SYSTEMATIC ANATOMY

OF THE DICQTYLEDONS
A HANDBOOK FOR -LABORATORIES
OF PURE AND APPLIED BOTANY

BY

DR: HANS SOLEREDER

PROFESSOR OF BOTANY IN THE 'UNIVERSITY OF ERLANGEN

TRANSLATED BY
L. A.. BOODLE, F.L.S.
ROVAL BOTANIC GARDENS) KEW

AND

F. E. FRITSCH, D.Se., PH.D., F.L.S.

ASSIstANT PROFESSOR OF BOTANY, UNIVERSITY OF LONDON, UNIVERSI1'V COLLEGE
AND LECTURER IN BOTANY_, EAST LONDON COLLEGE, UNIVERSITY OF LONDON

REVISED BY
D. H. SCOTT, M.A., LL.D.) PH.D., F.R.S.
LATE HONORARY KEEPER OF THE ]ODRELL LABORATORY,
ROYAL BOTANIC GAltDENS, KEW

'VOLa ~I~I
MONOCHLAMYDEAE, ADDENDA, CONCLUDING REMARKS
WITH 36 FIGURES IN THE TEXT

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First Printed 1908

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 CONTENTS OF VOL. II
ANATOMICAL CHARACTERS OF THE DlCOlYLEDONOUS ORDERS (continued):-

MONOCHLAMYDEAE PAGE ADDENDA

Nyctagineae. 645 POLYPETALAE PAGE
Illecebraceae .
Amarantaceae
649 Ranunculaceae 80s
651 Dilleniaceae . 807
Chenopodiaceae 655 Cal ycanthaceae 807
Basellaceae . 663 Magnoliaceae . 807
Phytolaccaceae 664 Trochodendraceae . 80g
Batideae . 668 Anonaceae . 810
Polygonaceae 669 Menispermaceae 812
Podostemaceae 674 Berberideae . 81 9
N epenthaceae 676 Nymphaeaceae 822
Cytinaceae . 680 Sarraceniaceae 82 3
Aristolochiaceae 682 Papaveraceae 824
Pi peraceae . 688 Fumariaceae. 82 4
Chloran thaceae 695 Cruciferae . 824
Myristicaceae 6g6 Capparideae . .. 82 5
Monirniaceae . 699 Resedaceae 826
Laurineae . 702 Cistineae 827
Hernandiaceae ·f 70 7 Violarieae 827
Go mortegaceae 709 Canellaceae 828
Proteaceae. .' . 709 Bixineae . 828
Thymelaeaceae . . 715 Pittosporeae . 83 1
Octolepis and Gonystylus 721 Polygaleae' . 83 1
Penaeaceae . 722 Vochysiacea.e 83 1
Geissoloma 7Z4 ' Caryophylleae 83 1
Elaeagnaceae 7Z4 Portulaceae . 833
Loran thaceae ~ 7Z6 Tam ariscine ae 833
Santala"eae . 730 Hypericineae 834
Myrodend"on 733 Guttiierae . 83 6
Champereia 136 Ternstroemiaceae ~ 837
GnWbia .. 737 Strasburgeria • 839
Balanophoreae . 73 8 Di pteroc .1l"peae 840
Euphorbiaceae . 139 A ndstrodadus 84 1
Daphniphyllaceae 760 Lophira. 84 1
Buxaceae
. 761 Monotes. 84 2
Balanopseae . 763 Chlaenaceae 84 2
Urticaceae . 76 4 Malvaceae. • 84 2
Ulmaceae, . 764. Triplochitonaceae . 843
Can nabine 769 Sterculiaceae . 844
Moraceae 77 0 Tiliaceae . . 846
Urticeae. 77S Rhaptopetalaceae . 846
Thelygoneae 779 Lineae. . .' 847
Platanaceae . 779 H umiriaceae . 849
Lei tnerieae . 782 Malpighiaceae 849
J ugl.andeae . 783 Zygophylleae 85 0
Myncaceae . 785 Geraniaceae . 85 1
Casuarineae . 7 86 Rutaceae . 854
Cupuliferae . 79 1 Simarubaceae 8S7
Salicineae . 797 Koebe"Unt'a 861
Lacistemaceae 799 Ochnaceae. . . . 862
Empetraceae 800 Ochnaceae, Van Tieghem 86,
Ceratophylleae 801 L\lxem burgiaceae 86;
 CONTENTS OF VOLUME II
IV. HAIRV CoVERING (continued) PAGJ!:
§ 36. Large glandular mechanisms (nectaries) . . • ([30
37. Chalk· andsa.lt·glandsof the Frankemaceae. Tamariscineae,
and Plumbagineae • • . . . . . I 133
38. Special forms of external glands occurring in insectivorous
plants. . . • . • . . . 1133
39. Cork-warts on leaves. 033
V. NORMAL STRUCTURE OF THll AXIS 1133
, 40. Medullary tissue . . . • . 1133
41. Appearance of the transverse section of the stem iD
herbaceous plants. . . . . . • 1134
2 Systematic importance of tbe structure o{ the wood in the
4- ' woody· Dicotyledons . . . . . . 1135
43· Wood-vessels. . 1136
44· Wood-p'rosencbyma . II4 I
45· Wood-parenchyma.. . . . . . . 1143
46 • Medullary rays of the wood . . • . . I14J
47· Special elements found in the normally 'developed xylem-
mass . . . . . . . . 1144
4 8• Axial wood and wood showing a tier..like structure 1145
49. Periderm with special reference to the cork . 1146
50. Aerencbyma . ' 1150
SJ. • Primary cortex • . • . • II So
52. Pericycle. . . . . . • . lJS2
53· Bast-groups and especially the secondary bast JJ54

VI. ANOllfALOUS STRUCTURE OF THE AXIS • • • • •

§ 54.Contracted vascular systems of submerged plants and
certain other Dicotyledons, and Dissociation of the ring
of vascular bundles • • "55
55. Axes showing poJystelic structure . I I 56
56. Medunary and cortical vascular bundles liS?
57. Intraxylary phloem. .. ... I159
58. Interxylary phJoem. ' . • . . . . U61
59. Successive development of secondary groups of wood and
bast . . . . • . 1163
60. Compound and divided xylem.ma.sses 1I6S
61. Unequal thickening of the xylem-mass u66
62. Cleava.ge of the xylem-mass II 6,
VII. STRUCTURE OF THE ROOT • • 1I67
§ 63. General ~tructure of the root . 1167
64. Anomalous structure of the root 1168

LITERATURE SUPPLEMENT

INDEX TO ORbERS AND ANOMALOUS GENERA (for Vols. 1 and II) 1I73

GLOSSARY • .. 1115
 NYCTAGINEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. The anomalous structure
of the stem (Fig. 155) and the oceurrence of raphides or styloids are character-
istic of the Order. The anomaly c_onsists in the appearance, in the pericycle,
of successive rings or strips of cambium, which produce secondary collateral
vascu1ar bundles and conjunctive tissue on their inner side. The conjunctive
tissue is either prosenchymatous or parenchymatous; its structure, combined
with the arrangement of the secondary vascular bundles, gives variety to the
'appearance of the transverse section of the stem. ~e p1:osenchymatous
-conjunctive tissue bears simple pits like the wood·prosenchyma of the vascular
bundfes, from which it is difficult to distinguish, and is sometimes traversed
by typical medullary rays. The xylem-vessels -have simple perforations. The
outer portion of the pericycle generally contains isolated groups of sclerenchy-
matous jbres. The development of cork is superficial. In Pisonia, Ncea,
and Lemaster the stomatal apparatus follows the Rubiacecus type, whilst
in J.1firabilis and Boerhaavia each pair of guard-cells is surrounded by an_ in-
definite number of ?fdinary epidermal cells. The __hairy covering consists _of
clothing and glandUlar hans. The former include the uniseriate trichomes
of Bougainvillaea, which are composed of- short cells, and the stellate hairs J
of Leucaster, in which the stellate portion consists of a rather large number
of unicellular rays. The glandular hairs' (Fig. 154, A) generally con'3ist of
a row of cells, the enlarged terminal cell being in most cases e1lipsoidal Of
-clavate, more rarely spherical; the contents of this -cell vary, but in the dry
state they are usually brown (Pisania, Neea" Mirabilis) . . A type intermediate
between clothing and glandular hairs is presented by the branched trichomes (Fig.
154, B) composing the dense hairy covering on the lower side of the leaf of
Pisonia tome~tosa, Casar.; in these hairs the axis and _branches are uniseriate ;
the terminal cells of the branches especially are commonly differentiated
into long sacs filled..with brown contents. Internal secretory organ~ a~ only
found in Okenia and Boerhaaviat in which enlarged epidermal cells, filled with
reddish-brown contents, give rise to a -red striation of the Ieave~ and floral
organs., • Oxalate of lime (Fig. 154, C) is excreted chiefly in the form of raphides
and styloidS, clustered crystals and a kind of crystal-sand (the latter in Leucaster)
being also found. The deposition of crystalline granules, consisting of Hie
same salt, in the cell-walls. of the epidermis in the.leaf and stem (Fig. 154, A),
has been shown to take place in the genera MirabiZis, Oxybaphus, Nyclaginia,
AUionia, Boerhaavia, AcieisantMs, Okenia and Abronia.
Z. STRUCTURE OF THE LEAF. The researches published on the anatomy
of the Nyctagineae almost always deal solely with the interesting anomalous
structure of the a~s, whilst the leaf~structure ~as practically not been investi-
_gated. And yet it would be important for sys matic purposes to obtain some.
knowledge of tpe,"di~tfibuJion of the. two type of stomata which have been
recorded in the Order;-and also of the distribution of the trichomes (se~ above).
With the, object of determining the different features which have been
briefly mentioned above, I examined the structure of the leaf in Mirabilis
Jalapa, L., BougainviUaea spectabilis, Willd. (Tribe Mirabilieae), Pisonia nitida,
Mart." Neea compressa, Schmidt (Triqe Pisonieae), and Leucaster canijiorus,
Choisy (Tribe Leltcastereae). The leaves in these plants have bifacial s~ruc­
ture. The stomata are only found on the)ower side of the leaf. In M.T(lbilis,
and Bougainvillaea, as stated above, they are surrounded by a variable,number
:Jf ordinary epidermal cells~ exhibiting no special arrangement, whilst in the

J The stella.te ha.irs of :lllJraCetz have not yet been carefully examined.
 646 NYCTAGINEAE

remammg cases they are accompanied on either side by one or more suo·
sidiary cells placed parallel to the pore. The important points regarding the
• trichomes investigated
on the leaf, have also
been mentioned in the
general diagnosis. Stellate
hairs occur in Leucaster
and A ndradea; simple um-
seriate hairs, composed of
short cells, with a blunt
A terminal cell not greatly
exceeding the others in
length, are found in
BougainmOUaea spectabilis.
Glandular hairs occur in
flisonia, Neea and M ira-
bilis; they have a stalk
of variable length, com-
posed of a very variable
nurnber of cells, and a
unicellular, sphtrical, or
elongated ellipsoidal head
(Fig. 154, A). The leaf of
Pisonia tomentosa, Casar.,
as mentioned above, bears
branched hairs, which have
a ghindular function (see
above, and Fig. 154, B}.
The large epidermal cells
of Okenia and Boerhaavia
are filled with brownish-
red, tanniniferous con-
tents, and are differenti-
ated as secretory cells;
they were observed by
Heimerl. The same author
demonstrated the deposi-
tion of crystalline granules
of oxalate of lime in the
walls of the epidelmal cells
(Fig. 154, A) in the genera
mentioned above. It
should be noted that these
genera belong to the Sub-
tribes Boerhaavieae and
FIG. 154. AI..Tranwttse 5eCtioD throng!!. the epidermis of lJarlzttavia Abronieae of the Tribe
rlj>en.r, L.; D, Hra.Dcbed tricbome of .Asonia tomenhsa, Casar; c, Crys-. 1\1· bili' din t
talline elemenu in tbe spongy tissue of P. "ftitla, Mart.-Original. Ira eae, accor g 0
the classification of Ben-
tham and Hooker; in the Sub·tribes Bougainvillaeeae and Boldoeae, and
in the Tribes Pisonieae and Leucastereae 1 Heimerl sought for this feature
in vain. The incrustatioi'l, as far as js known, is almost always present in
all the species of a genus; it is only known to be absent in a few

1 Species of BfJUcuinvillaea, Tricyda, Boldoa, Collignonw, Reidunbadda, E'isDnia, .Neca,

Phruoplilum, Cepha[olrnllflttdra,'lwciuter and CryptocarJnI!> were examined.
 NYCTAGINEAE 647
species of Mirabilis (M. Jalapa, L., 1W.. longifiora, L., M. Oaxacae, Heim. and
lVI. • Wrigntiana, Decne.'). In most cases the deposition is met with in the
epidennis, both of the leaf and stem. The variations found in the individual
species depend on (a) the occurrence of the deposition in both the upper and lower
epidermis of the leaf, or only in the lower, (b) the presence of the deposition in
the outer~ inner~ and lateral walls of the epidermal cells, and (c) the diff~t
degrees of abundance of the excretion. The layer of granules in the outeFwall
is directly covered by the cuticle, and is bounded towards the cell-lumen by a
lamella devoid of granules. Deposition is rare in the late~ walls, but common
in the internal walls of the epidermal cells; in these c~es also the lamella of
the wall adjoining the lumen is devoid of granules. The incrustation Iilayalso.
extend to the walls of the hairs, whilst the stomata are invariably free from it.
In some cases the deposition is'so abundant, that the leaves and stems of certain
species appear grey or whitish, the stem of Boerhaavia elegans, Chojsy being
chalky white 1. IIi Oxybaphus a study of the development has shown that
the_ incrustation only appears at a relatively late stage, and originates in
the cell-wall it.!klf. Regarding the mode of excretion of oxalate of lime, the
following statements may be added. Bundles of raphides have been shown
to occur in species of Bougainvillaea, Cryptocarpus, Mirabilis and Oxybaphus,
raphides and styloids in species of N eea, Phaeoptilt:m and Pisonia, styloids alone
in Eggersia. In the leaf of Pisonia nitida each cell of the palisade-tissue contains
a ra~her,small clustered crystal, whilst styloids and bundles of typical raphides
or of short. acicular crystals more like styloids also occur in the mesophyU
•(Fig. IS4, C). In the sacs containing crystal-sand in the leaf of Leucaste,
cani/lorus there are either tetrahedral crystalline granules or rafuer small
acicular or prismatic crystals.
3. STRUCTURE OF THE AXIS. Anomalous structure of the stem appa-
rently occurs in all the woody species, and is also found in a certain number
of the herbaceous species. It has been recorded by various authors (Regnault,
Unger, Gronlund, Petersen, De Barry, ~adlkofer. Solereder, H. Schenck,
Houlbert) in numerous species of the genera Mirabilis, Oxybaphus, Boerhaavia,
Bougainvillaea, Pisonia, Eggersia, Phaeoptilum, N eea, Leucaster and Crypto~
carpus. The constant feature of the an'omaly consists in the presence of
more than one ring of vascular bundles in the transverse section of the axis
{Fig. 15SY. The course of development of the anomaly has been referred to'
above. The secondary vascular bundles exhibit a concentric or irregular
arrangement; they are embedded in conjunctive tissue, and together with
the latter form the' wood' of the Nyctagineae ;,,<-in most members of the Order
the conjunctive tissue, except the innermost portion (on this point see below),
consists chiefly of prosenchymatous cells, which are provided with simple pits
and usually have rather thick walls, though in rare cases the thickeniJ1_g is but
slight (Pisonia tragrans, Desf.). Adjoining the phloem-groups of the vascular
bundles there are frequently 2: few unlignified and thin-walled or lignified paren·
chymatous cells, and in certain members of the Order (in all the species of Pi-
sonia and J.Veea investigated by Houlbert and myself, except p. hirtella, H. B. K.)
narrow, radial strips of tissue traverse the conjunctive tissue, and in their com-
position correspond to the medullary rays in woods with normal structure. The
thin-walled conjunctive tissue is developed in greater abundance in a minority
of the species investigated. Thus in BougainviUaea spectabilis the secondary
bundles, in which the xylem consists of vessels and prosenchyma, are separated
by rather broad parenchymatous sttips of tissue, resembling medullary rays,

1 The dun surface of the stems and leaves in some species of PiSOll;a and Neea. due to a
deposition of wax, must not be confounded with thi.s feature (Heimerl).
 NYCTAGINEAE

when seen in a transverse section of the stem; .parenchymatous cQnjunctive

tissue is also insinuated between the secondary bundles in a tangential
direction. B./astuosa, Hrcq. pres~nts similar features, according to Houlbert.
In Boerhaavia plumbaginea, Cav. (Petersen) and B. arborea, Lag. the thin-
walled, parenchymatous, conjunctive tissue is also rather abundant.. In these
species the phloem-groups and the adjoining conjunctive elements combine to
form concentric anmdar or band-shaped strips of tissue. The innermost
portion of the conjunctive tissGe, formed by the secondary meristem at the
commencement of its cambial activity, is differentiated so as to resemble
a pith in many members of the Order. Owing to this fact the leaf-trace
bundles, and sometimes also the innermost secondary vascular bundles, often

FIG. ISs.. Transverse section througb the stem or Plsoni'a mgt'tCaNs, Sw.-Original.

appear to be medullary (Mirabilis and Oxybaphus, according to. De Bary).

Probably"the medullary vascular bundles, which I met with in all the species
of BougainviUaea, Crypto carp us, Pisonia and N eea named in my , Holzsiruk-
tur,' are likewise only apparently medullary; they are wanting in Leztcaster
caniflorus and (according to Petersen) in Boerhaavia plumbaginea.
Regarding the component parts of the vascular bundles the following
facts remain to be mentioned. The xylem contains: (a) pitted vessels with
simple perforations accompanied by reticulately perforated septa in the
species of Boerhaavia (Dickson, Petersen); (b) prosenchyma; (c) paren-
chyma; (d) in the leaf-trace bundles, spiral tracheae which can be unrolled.
The latter elements, however, sometimes also occur in vascular bundles which,
from their position, must be regarded as inner secondary bundles, as has been
shown by De Bary in the case of Mirabilis, by Petersen in Boerhaavia and
Bougainvillaea, and by myself in species of N eea and Pisonia. In the ordinary
 NYCTAGINEAE

cases, iI) which the conjunctive tissue is formed by prosenchyma, the xylem-..
portions of the vascular bundles have not a distinct boundary. Together
with the conjunctive tissue they form a conn'ected xylem-mass, in which the
phloem-groups of the vascular bundles are embedded like the islands of soft
'bast in Strychnos.
External to the parenchymatous pericycle, in which the anomalous growth
in thickness of the stem takes place, a zone of small, isolated groups of scleren-
chymatous fibres is present in the species of Bougainvillaea, Cryptocarpus,
Leucaster, Pisonia and N eea examined by me; these elements are wanting
in Mirabilis Jalapa. and OxybaPhus viscosles (Regnault). The cork arises
subepidermally i~ BougaiJ1,villaea spectabilis (according\to H. Schenck), and
in Pisonia and Boerhaavia (according to Petersen), whIlst it is formed i~ a
deeper cell-layer of the primary cortex in N eea parvi/lora (according to Petersen).
The anom31ous structure, as far as is known, extends also to the roots
(Mirabilis with parenchymatous conjunctive tissue and concentric arrange-
ment of the second.<J.ry vascular bundles, according to De Bary; Bougainv£llaea
spectabilis and Pisdnia nitida, according to Avetta).
Literature: Unger, Bau u. Wachst. d. Dicotyledonenst., Petersburg, I 84o.-Niigeli, Beitr. z .
.)riss. Bot., i. Heft, 1858, pp. 1l9-3I.-Regnault, Anat. d. quelques tiges d. Cyclosp., Ann. sc. nat.,
scr. .of, 1. xiv, 186o, PP' 144-9 and pI. ix.-Gronlund, Stamm. og. wen. anat. bygn. has Neea, etc..
Vidensk. Meddel. nat. For. Kjobenhavn 1872, 19 pp. (French resume) and Tab.-Finger, Anat.
de. von Mirabilis Jafapa, Diss., Bonn, 1813, 25 pp.-Moller, Holzanat." Denkschr. Wiener Akad.
1876, pp. 35-6 and :UI.-De Bary, Vergl. Anat. 18n, especially p. 607 et seq.-Dickson, in
Transact. and Proceed. bot. Soc. of Edinburgh, vol. xiv, 1880, p. 121 et scq.-Petel'Sl!ll, Nyctag.-
Streng. Histolog. etc., Bot. Tidsskrift, Bd. II, 1879-80, pp. 149-76, Tab. iv-v and French resume
pp. (J6)-(19).-Radlkofer, in Abh. naturw. Ver. Bremen 1883. p. 435 et seq.-Morot, Pericycle,
Ann. sc. nat., ser. 6"t. xx, 1885, especiallyp. 282 et seq.-Hernil, Tige des Dicotyled., Ann. sc. nat.,
ser. 7, t. ii, 1885, pp. 246-7.-Solereder, Holzstr., 1885, pp. 207-10.-Heimerl, Einlag. v. Calcium-
oxala.t etc., Sitz.-Ber. Wiener Akad., Ed. xciii, Abt. I, 1886, sep. copy, 16 pp. and I Tab.-[Avetta,
Anomalie di straltura nelle radici delle dicotyled., Ann. _dell 1st. bot. di Roma 1887, p. 10.)--
Heimerl, Anal. d. N., Denkschr. \Viener Akad., Ed. 53, Abt. 2,1887, pp. ~[-78 and 3 Tab.; only
trea.ts of structure of flowers. and fruit t-PeteIgetl, Strengelb)'go. has Eggersia, Hot. Tidsskr., Bd. 16,
1888, pp. 216-20, Tab. iv and }I'rench resume p. (9) and Bot. Centralbl. 1888, iii, pp. 27-28.-
Eiselen, Rhaphiden, Diss., Halle a. S., 1888, p. 13.-Koh1, Kalks. etc., 1889, p. 79.-Heimerl, in
NatiirL Pfianzenfam., iii. Teil, Abt. I b, 1889, p. 15.-Houlbert, ,struct. compo du bois sec. clans les'
Apetales, Paris, These, 1893, pp. 75-82, scp. copy from Arm. sc. nat., ser. 7. t. xvii.-H. Schencll,
Anat. d. Lianen, 1893, pp. 56-8 and 252, Tab. ii.

ILLECEBRACEAE:
In this Order, which is mainly composed of her:'aceous plants, the anatomy
has been little investigated. From the few facts known it appears that in
the structure of the axis the following features promise to afford a diagnosis
of the Order: the usually normal structure of the stem; the lack of true
medullary rays in the wood; the simple perforations of the vessels; the
bordered pitting of the wood-prosenchyma; and the superficial development
of cork. The stomata are generally. surrounded by three or more epidermal
cells; in Scleranthus· the Caryophylleous type is found. The hairy covering
consists of unicellular or uniseriate clothing hairs, which are mostly simple,
rarely (PoUichia) branched, whilst uniseriate glandular hairs with a unicellular
head are very rare (Dysphania). Internal secretory receptacles are not present.
Oxalate of lime occurs in the form of large clustered crystals, and in Gymno-
carp os also as crystal-sand. Anomalous structure- of the stem in the form
of successive rings of growth has been shown to occur in Pollichia campestris_
Sol. and species of Corrigiola.
The structure of the leaf has been examined by Volkens in a number
of desert-plants (Pteranthus 'dichotomus, Forsk.,' H erniaria hemistemon, J.
Gay, Paronychia desertorum, Boiss., GYl1l-1Z0Carpos decandrum, Forsk.), and
650 ILLECEBRACEAE

by me in Pollichia campestris, SoL, A nychia d£chotoma, Michx., Paronychia

capitata, Lam., Pteranthus echinatus, Desf. and Scleranthus perenn":s, L. The
leaf-structure is commonly centric (e.g. Pteranthus diclt-otomus and Gymno-
carpos decandrum). The stomata are usually found on both sides of the leaf,
and either lie in the same plane as the surface of the leaf, or (Pteranthus dicho-
tomus and Gymnocarpos decc.ndrum) are strongly depressed. In the species
which I examined, with the exception of Sclera nth us perennis, the stomata
are surrounded by three or more epidermal cells, showing no special ~()de
of arrangement; in Scleranthus the stomata are parallel to one another, the
pore being parallel to the median vein, and they possess two subsidiary cells,
placed transversely to the pore. The following features are adaptations to
a desert-climate: a deposition of wax on the leaf-surface in Gymnocarpos
decandrum; high epidermal cells in the leaf of Pteranthus dichotomus and
Herniaria hemistemon; bladder-like differentiation of the epidermal cells in
the neighbourhood of the margin and above the midrib of the leaf in Pteranthus
dichotomus; and central aqueous tiSSue in the leaf of Gymnocarpos decandrftm.
The clothing hairs are mostly simple and unicellular (Herniaria, Parony-
chia, Anychia), or uniseriate (Sc/ercmthus). The branched trichomes of Polti-
chia campestris are generally two-armed, the main axis and the anns consist-
ing of a single row of cells. In the floral 'region of Pteranthus eehina/us there
are uniseriate trichomes of varying length, composed of few or many cells,
and provided with a capitate swollen t~rminal cell; they practically have n~
glandular function. On the other hand the similarly constructed capitate
hairs in the species of Dysphania (e.g. D. myriocephala, Benth.) are glandular;
the species of this genus were described by Bentham, Flor. Austral., as slightly
glandular. Oxalate of lime is often found in the mesophyll in the form of
large clustered crystals (in Pteranthus diehotomus and Gymnocarpos deeandrum,
and also in the five species investigated by me); the primary cortex of Gymno-
carpos decandrum contains large cells, which in place of clustered crystals
enclose a compact mass of crystal-sand consisting of oxalate of lime.
The structure of the axis, like that of the leaf, has been little investi-
gated; it was examined by me in PoUichia campestris, Sol. and Gymnocarpos
decandrum, Forsk., by Regnault in Paronychia bonariensis, Anychia dichotoma
and Corrigio/a littoralis, and by Petersen with reference to the ~ricycle in
species of Illecebrum, Pollichia, Paronychia, Herniaria t C01'rigiola~ Gymno-
carp os, Pteranthus, Cometes and Scleranthus. In the structure of the wood_
the most noteworthy feature is the absence of true medullary rays in species
in which the xylem-portio~ of the vascular bundles unite to form a ring (P<,lli-
chia campestris, Gymnocarpos decandrum, Paronychia bonanensis, Anychia
dichotoma). In Pollichia and Gymnocarpos the diameter of the vessels is not
great ('02-'03 mm.); they have rather thick walls and simple perforations
(denticulate at the margin in Pollichia campestris). Spiral thickening of the
vessel-wall is found in Gymnocarpos decandrum. Typical bordered pitting
has been demonstrated in the wood-prosenchym~J not only in PoUichia and
Gymnocarpos, but also in the species examined by Regnault. A sclerenchyma-
tous peri cycle is of frequent occurrence in the cortex (in the species examined
by Petersen and Regnault); in species with a considerable amount of growth
in thickness. it becomes split up into small sclerenchymatous groups. The
deVelopment of cork takes place subepidermally in Paronychia bonariensis.
Anomalous structure of the axis, in the form of concentric rings of
vascular bundles, is found in Pollichia campestris, and according to Petersen in
Corrigiola littoralis and C. telephiifolia (also in the root of C. littoratis, according
to Lohrer). The second ring of vascular bundles in .PoUichia (and probably
also in Corrigiola) is produced by a secondary meristem, which arises in the
inner parenchymatous portion of the pencycle.
 1LLECEBRACEAE
Literature: Regnault, Anal d. qq. tiges a. Cyclosp•• Ann. sc. nat., s~r. 4, t xiv, 1860, PP.. 1 13-17
nnd pl. 7.-Solereder, Hvlzstr.• 1885, pp. :iII 0, 31I.-Lohrer, Warzem, In Wigand, Bot. Hefte, "ii.
1887, pp. %6-7 and Tab. ii.-Volkens, Aegypt.-arab. Wiiste, IS87, pp. 103-5 and Tab. vii, x and xv.
-Petersen, Mom. til Caryophyll. anat., Bot. Tidsskrift, Bd 16' VP. 187-203, TavIe 5 and French
reswne p. (8).

I. REVIEW OF THE
AMARANTACE
ANATOMICAL
J
FEATURES.
.
With few exceptions
(Achatocarpus, Cladothrix) the woody members of this Order are distinguished
by the same anomalous structure of the axis as occurs in the related N ycta-
gineae and Chenopodiaceae (Fig. 157). The vascular bundles scattered in
the conjunctive tissue exhibit concentric or irregular arrangement in a trans-
verse sectio~ of the stem, and the conjunctive tissue either consists of pros-
enchyma with siJjlple pits or of unlignified parenchyma; when prosenchymatous,
medullary rays are sometimes ~evelcped in it. The perforations oi the vessels
are simple in all cases. The formation 9£ periderm takes place super.ficially
in Amarantus. The stomata do not possess any special subsidiary cells. In-
ternal secretory organs are wanting. The hairy covering is for the most
part formed by ordinary unicellular or uniseriate trichomes, and by capitate
hairs with a uniseriate stalk and a unicellular head. The following are special
forms of hait: (Fig. 156) 1: the trichomes of Ptilotus, with two arms or rarely
more, both the stalk and the arms being composed of a row of cells; the
two-armed trichomes of Dicraurus, in which the terminal cell has equal arms ;
the stellate hairs of species of Iresine with a many·rayed terminal cell,
occasionally passing by transitional forms into two-armed hairs with arms of
unequal length; finally the cande1abra-hairs of Alternanthera, Cladothrix,
'- and T elan~lwra, in which each tier is composed of a many-rayed cell. TIle
walls of the various forms of hair (with the exception of the capitate hairs)
are in many ca<;es raised into numerous small papillae.
2. STRUCTURE OF THE LEAF. The leaf-structure has been little investigated.
, The species which I examined, representing the three tribes (Amarantus Blitum, L.
and A. polygonoides, L., Celosia tfig')lna, L., Ge»nphrena deSt1torum, Mart.),
have stomata on both sides of the leaf. The stomata ar~ sunounded byithree
r;>r more ordinary epidermal cells. The size of the epidermal cells and the
nature DO their lateral margins vary in the species mentioned above. Celosia
trigyna is distinguished by papillose or bladder-like protrusion of the epidennal
cells on the lower side of the leaf oppo~ite the vascular bundles of the veins.
In Amatalltus Blitum, A. polygonoides and other species, -Gomphrena deser-
torum CL'1d other species, Alternanihera muscoides/Benth. et Hook., and Philo·
xcrus vermicularis, R. Br., th~.vascular bundles of the smaller and larger veins\
including their ultimate fcUllifications, are surrounded by a characteristic
sheath of large and almost cubicc4 parenchymatous ceUs (Fig. 156, A-B);
this is a feature which only rarely occurs amongst Dicotyledons, although
common t=J.ll1ongst Wx>nocotyledons. According to Johow and Wanning,
Philoxerus vmnicularis-possesses· a decidedly peculiar type of leaf-structure,
viz. rather massive aqueous tissue on the lower side of the leaf; stomata in
the upper epidermis only; assimilatory tissc.e only on the upper side; and
lastly. vascular bundles (of the veins) with a parenchymatous sheath of cells
with wide lumina, and radial arrangement of the adjoining cells of the palisade-
tissue.
Oxalate of lime is present in the leaf in the form of clustered crystals
or crystal-sand. Solitary crystals are rare (axis of Alternanthera ·procumbens,
according to Nemnich). CrystalAsand and clustered crystals sometimes occur
1 The stell,te hairs, mentioned by systematists as investing certain species of Tridu'uium, and
the hairy covering of Caluffl",a. described by them as furfuraceous. have still to be investigated.
 AMARANTACEAE

side by side in the leaf, as in Amaranltes BI#um and A. polygonoides, where'

large clustered crystals are found in the mesophylI, whilst sand occurs in
the veins; in other cases clustered crystals only are found in the leaf (Gomphrt.na
deserrorum), or sand only (Celosia trigyna); in Celosia trigyna the sand in the
cells of the mesopbyll consists of coarse granules or fragments, whilst that
in the veins is more finely granular. Crystal·sand is the characteristic form
(If excretion in the Order. Besides occurring in the leaf of A marantus and

FIG. 156. A-B, Sheath rotlnu the \'ascular lJundles in the veins of the leaf of Amal"tlH/iIs B/,./""" L. : A, ill
trallS'erse llee:tion i B, in suTfaee-\·iew. C-F, Forms of trichome. of the Amarantaceae: c, AIt"f14 lanaJ4, Juss. ;
0, CItu/otA".;% la""rinD.StI, Nutt.; E, !nsf,1e SeluJl/'"eri, \Vats.; F, D;et'auru4 leploddus, Hook. f.-Onginal.

Celosia, it has been shown to occur in the axis in species of the following
genera by Regnault, Arcangeli, Nemnich and myself: Deeringia, Celosia, Bosia,
Chamissoa, AUmania, Amarantus, Acnida, Cyathula, Pupalia. Crystalloids
have been xr.et with by Zimmermann in the chloroplasts of the leaf-tissue
in A chyranthes VerschaDeltii, Alogiphanes brasiliensis, and a plant described
as 'Aeroa SanguisOfba.~
Ordinary unicellular or uniseriate trichomes, which give- rise to a crisp,
woolly, or silky hairy covering, are widely distributed amongst the Amaran-
taceae. Capitate hairs with a unicellular or uniseriate stalk and a unicellular
 AMARANTACEAE

svherical or ellipsoidal head are found on the veins on the lower side of the
leaf in A marantus, and have also been 0 bserved by Nemnich in species of
Achyranthes, Acnida, Allmania, Alternanthera and Gomphrena. There are
numerous special forms of clothing hairs amongst the Amarantaceae; of
these I may mention the following examples derived from my own investigations.
Ptilotus latifolius, R. Br. has two-armed or many-armed trichomes, in which
the stalk and the frequently long arms are composed of a row of .cells with
thin walls; the lowest cells of the stalk are short and have yellow walls. Charac-
teristic uniseriate hairs, composed of a few short, basal cells with yellow walls,
and several longer cells, of which the terminal one is pointed, are found i!
Aerva lanata, Juss. (Fig. 156, C), but not in Aerva MonSfJnia, Mait., wher
ordinary uniseriate hairs OCCUI. The characteristic features presented b
the trichomes of Aerva lanata. are: (a) the presence of numerous small knobs I
on the longitudinal walls of the longer cells; these knobs are· not solid, but
constitute papillose protuberances of the wall of .the hair; and (b) the nature
of the transverse walJs in the upper portion of the hair; the surface of these
walls is not plane, the margin being produced into papillae, sq that the
cells of the-hair become firmly dovetailed with one another. If we 'imagine
some of the pap~llae of the long cells of a hair in Apva la1tata as growing out
into ray-cells at certain points, we should have d.ndelabra-hairs like those
of Alternanthera, Cladothrix (Fig. I56, D) and Telanthera. The following
points of difference are shown by these hairs. In T. /rutescens, Moq., the
upper cells of the candelabra-hairs bear whorls of pointed branches, each
whorl consisting of outgrowths from one of the cells immediately above the
lower transverse wall, whilst in Cladothrix lanuginosa, Nutt. the branches radiate
from the middle portion of the cell, the cells in question being cylindrical;
in Telanthera frutescens we meet with the same small conical protuberances
as in Aerva lanata, whilst in Cladothrix lanuginosa the nature of the trans-
verse walls in the upper portion of the hair is the same as in A erva lanata.
According to Schleiden and De Bary, the candelabra-hairs of Alternanthera
spinosa resemble tlJ.ose of Tela nth era /rutescens. J:he stellate hairs of l,esine
Pringlei, Wats. may be regar4ed as a reduced form of candelabra-hair; they
possess a uniseriate stalk and a terminal cell with three, four, or' more rays.
The hairy covering of a second species of Iresine (I. Schatfneri, Wats.) consists
of transitional forms, some of which lead up to the trichomes of Aerva lanata,
others to the two· armed hairs of Dicraurus. The commonest forms are uni-
seriate trichomes, in which the upper portion is-P!ovided with small papillose
protuberances, as in Aerva lal1ata, but the base of the terminal cell bears out-
growths in the form- of a number of short rays. The remaining trichomes
of Iresine SchaDneri are tW9-armed hairs with arms of unequal length; the
terminal cell has thick walls, and exhibits small papillae all over its longi-
tudinal walls, but bears still larger protuberances like rays just above the
point of insertion of the stalk (Fig. 156, E). From hairs of this type it is not
a great step to the two-armed trichomes of Dicraurus leptocladus, Hook. fil.
(Fig. I56, F); here. the.. terminal cell has equal arms, and its walls are likewise
provided with small papillae.
The petiole contains an arc of isolated vascular bundles in the character-
istic region in the species of Celosia, Amarantus and Gomphrena examined
by Petit.
3. STRUCTURE OF THE AXIS. The anomalous structure of the stem I
1 Unicellular or nniseriate hairs with similar papillae are also figured by Martius, Nov. Gen. et
Spec. plant. Brasi1., vol. ii, Tab. 127 et seq., in species of Hebalulu, Moczphants and Telantltera,
and by H. Schenck in Ha/JUteia; if I understand him rightly, Nemnich also observed them in species
of Acn)'rdnJ/us, Alternan/lura and Comphrena.
, The anomalous structure has also been met with in the root!; of the Amarantaceae.
 AMARANTACEAE

has been demonstrated by Sanio, Regnault, De Bary, Solereder, Morot, H.

Schenck,Houlbert and
N emnich in species of
the following genera :
Deeringia, Celosia,
H ermbstaedtia, Ro-
detia, Rosia, Chamis-
soa, A marantus, .lcni-
da, PUPalia, Psilotri-
chum, Aerva, Achy-
Tanthes, T elanthera,
Alternanlnera, Gam-
phrena, Froelichia, H e-
banthe. On the other
hand the anomaly is
wanting in the genus
A chatocarpus, which
has r~cently been
transferred from the
Amarantaceae to the
Phytolaccaceae by
Schinz and Autran,
and according to my
own observation a1so
in the woody Clado-
thrix lanuginosa, the
systematic position of
which amongst the
Amarantaceae had
never yet been doubt..
ed. The mode of
origin of the anomaly
is the same as in
the N yctagiIRae; the
secondary meristems
develop in the paren-
chymatous pericycle,
commonly on the
inner side of small
groups of sclerenchy-
matousfibres. In those
cases in which there
are successive meri-
stematic rings, each
producing a ring of
vascular bundles, the
latter are arranged in
concentric circles. On
the other hand, when
there are only strips
of meristem, aris-
ing irregularly, here
FIG. 1,0;7· Tran_ 8K'tlo!"s, through the axis of: A, &sill yervil fIiIWIJ, L. ; and ther~ and each
.. An'VIJ .stllflrUnI, WalI:-eripa.!, d I - 'ngl
eve Opl_JJg ~ a qi/AJJ. e
vascular bundle, whilst to the right and left mepstematic stripS of earlier
 AMARANTACEAE

origin are still active, the vascular bundles are irregularly arranged in the
transverse section. It may be noted that in the first case (Fig. 157, AI)
both the radial conjunctive tissue (simulating medullary rays), and the
tissue occurring tangentially between the vascular bundles, is paren-
chymatous, whilst in the -second case (Fig. 157, B t) the conjunctive tissue
is prosenchyriiatous, groups of unlignified and lignified parenchyma being
present only in' connexion with the phloem-groups. Transitions between
the two types are occasionally found (see Hebanthe pulverulenta, Mart. in
H. Schenck's work). The prosenchymatous conjunctive tissue bears simple
pits like the prosenchyma of the vascular bundles themselves. Medullary
rays sometimes occur in the prosenchymatous conjunctive tissue. The vessels,
which attain a diameter of '2 mm. in the twining species of Hebanthe, have
simple perforations. De Baij mentions the occurrence of apparently medul-
lary vascular bundles S (with regard to this term see Nyctagineae) in species
of Celosia, Achyranthes and Amarantus; in the species of the latter genus
they a:re accompanied by true medullary vascular bundles. The angles,
which are sometimes present on the stems of the Amarantaceae, consist of
~ollenchyma (Trichinium sericostachyum, Nees, &c.). According to Regnault
the development of cork takes place subepidermally in species of Am4rantus.
Literature: Regna.ult, Cyclospermees, Ann. sc. nat., ser. 4. t. xiv, 1860. pp. u7-33 a.nd pI. viii.
-Sanio, in Bot. Zeit. I864.-De Bary, Vergl. AnaL 1877. especially pp. 259 and 607 et seq.-
Jobow, in Pringsh~im, Bot. Jahrb., Ed. xv, 18S.., p. 309 Dote.-H. Schenck, Wandverdick. etc.,
DUs., Bonn, 188.., p. [.of.. and Tab.-Morot, Pc!:ricycle, Ann. sc. nat., ser•. 6, t. xx, 1885, especially
pp. 282-+-Herail, Tigedes Dicotylc!:d., Ann.sc. nat., ser. 7. t. ii, 1885, p. 146.-so1ereder, Holzstr.,'
1885, pp. 2I1-13.-Witte. Lianen, Diss., Frhlburg i. Br. u. Kiet 1886. pp. 13 and :14 et seq.-Petit,
Petiole, Mem. Soc. phys. et sc. nat. de Bordeaux, ser. 3, t. iii, 1881, pp. 239-41 and pI. ii.-
ArcaDgeli, Ossal calc. criptocrist., Naov. Giom. bot. Ital., vol. xxiii, 1891, p. 369.-Zimmennann,
Pfialuenzclle, 2. Heft, 1891, pp. 151, IS2.-H. Schenck, Anat. d. llanellI 1893, pp. -49-53 and
Tab. i.-Schim. in Natiirl. Pflanzenfam., iii. Teil, Abt. I a, 1893. pp. 93-3.-Schinz and Autran;
Ar:ilatocarpus, Bull. de l'Herbier Boissier 1893, pp. 1-14 and pI. i.-Houlbert. Boil sec. dans les
A~les, These, Paris, 1893, pp. ~.-Nemnich, Axe etc. d. A., Diss., Erlangen, 189 ..., 36 pp. and
1 Tab.-Herbst, Markstr., Bot. Centralbl. '1&)4, i, p. 295.-Schubert, Parenchymscbeidcn, Bot.
CentIalb1. 1897, iv, p. 63 et seq-:'"-Warmm[!. Halofyt Stud., K. Danske Vid. Selsk.. Skr. 1897, pp.
2l.f and 226.

CHENOPODIACEAE.
(SUBORDO i, CHENOPODIEAE, BENTH. ET HOOK.
I. REVIEW OF mE ANATOMICAL FEATURES. The most striking anato-
mical feature in this Order is the anomalous strutiure of the stem, occurring
in all Caenopodiaceae in which the growth in thickness is considerable. It
consists in the appearance of peticyclic rings or strips of cambium, which
"Iriginate and also lose their activity successively, forming secondary
bundles and conjunctive tissue. Two extreme types exhibited by the trans-
verse section of the stem may be distinguis~ed: the first shows cqncentric
zones of wood and.bast ; .in the second there' are vascular bundles embedded
in prosenchymatous conjunctive tissue, and arranged in various ways, either
·'The foUowiug species belong to this type: Celosia argmtea, Moq., Dosia )'dVa mtwa, L. and
CMuliissoa aJlissima, H. B. K., as described in my • Holzstruktur: HeblUlllu luJ~seriua, Mart.,
according to H. Schenck. and Rodelia Am/ursliana, Moq., according to Houlbert.
.. The following species belong to this type: according to my , Holzstruktnr' DeerinKia baccala,
Moq., HermosttUdtia Ca/fra, Moq., PujJalia lappacea•. Moq., Psilolric;'um cDrda/urn, Moq .• Auva
Sttmdens. Wall. and Telanlkera 1'amosUsima, Moq., and according to Witte /rt:Jine etatior, Rich.
I Ct/gsia' arrentea and Ckamissoa allissima also possess :l.pparently medullary vascular bundles
(see my 'Holzstruktur '). I will do no more than mention that there are (urther statements by
Nemnich OD this subject, as they do not appear to be quite reliable.
• The Suborder Baselleae of Bentham and Hooker is excluded from the following description,
and will be treated separately afth the rest of the Chcnopodiaceae.
 CHENOPODIACEAE

following a definite law Qf without regulanty. Other features Common to the

members oftheOrder are as fcHows: the vessels usuaHy have simple perforations
only (exception Axyris) ; both the wood-prosenchyma of the vascular bundles
and the prosenchymatouS' conjunctive tissue bear simple pits. True medul-
lary rays occur in the species showing the first type of transverse section,
but have not been observed in the pfosenchymatollS conjunctive tissue (in
the second type). A sclerenchymatous pericycle is' found in nearly all the

FIC. ISS. Trans\'erse section. throu.gh the leaves or; A, BtiSS;4 mu,i(4Io, All.; 8, Sa/scltr tonfr'folla, Forsk, ;
C, Alrlpltx Halimus, L.-AfterVolkens..

members of the Order; it rarely consists of a closed sclerenchymatous ring,

and is mostly composed of isolated groups of sclerenchymatous fibres. The
formation of cork varies, and may be either superficial or internal; in the latter
case it generally takes place in the pericyc1e. The following special anatomical
features are found in the stem: the peculiar structure of the stem in certain
Salsoleae ard Salicornieae (which have a peripheral network of vascular bundles
in the aqueous tissue t surrounding the usual vascular system of th~ axis) ;
the occurrence of apparently medullary vasculaL bundles in certain cases
(species of At,iplex, Beta and Chenopodium), and of true medullary bundles
i.n Acroglochin persicarioides, Moq.; finally the presence of spiral tracheids
 CHENOPODIACEAE

or spicular cells in the assimilatory parenchyma of the stem in certain species

of Arthrocnemum, Salicornia and Sarcobatus. The most noteworthy features
in the structure of the leaf are the absence of a definite type of stoma, and
of typical spongy tissue, which has not been observed in any species. In
spite of the xerophilous character of many species the cuticle rarely attains
a considerable thickness, nor have mucilaginous epidermal. cells been observed.
On the other hand aqueous tissue and a covering of wax on the surface of
the organs are present in certain species. The following anatomical features
of the leaf are of value for special diagnosis: the varied mode of con·
nexion between the assimilatory tissue and the vascular system of the veins
(in some cases by means of a characteristic sheath, Fig. 158); the peripheral
or central position of the aqueous tissue developed in the leaf (Fig. 158);
and the transverse arrangement of the stomata, not uncommonly shown by
narrow leaves (but"occurring also on the axis). No internal secretory recep·
tades are present. Oxalate of lime is excreted chiedy in the form of clustered
crystals and crystal.sand, rarely also in the form of solitary crystals. The
hairy covering is very diverse. Glandular hairs are rare (C amphorosma,
Chen<JPodium, Fig. 159, L-O), and always consist of a row of cells, of which
the terminal cell or the uppermost cells are glandular. Water-storing bladder·
like hairs (Fig. 159, C-F) are widely distributed; they have a unicellular
or uniseriate stalk and a unIcellular head, which varies in shape, being spherical,
or sac-like, or sometimes difiereJ}tiated like the terminal cell of a two·armed
hair. The following types of clothing hairs are found: ordinary uniseriate
trichomes; uniseriate trichomes with short basal cells and one or more
longer cells, the latter' being studded with cellulose-papillae (Camphorosmeae
and Salsoleae of Vol kens' classification, Fig. 159, A); two-armed hairs having
a terminal cell with equal arms, and likewise provided with cellulase-·papillae
(Petrosimonia, Fig. 159, B); stellate hairs in which all the rays are branches ...
of It single cell (Grayia, Eurotia, Fig. 159, G-H); stellate haIrs with several
uni- or multicellular rays (Axyris, Fig. 159, J); finally multicellular cande·
labra-hairs (Corispermum. 'Fig. 159, K, Agriophyllum).
2. STRUCTURE OF THE LEAF. The following description is mainly basedl
on Volkens' statements. ., ,
The leaf-structure of the Chenopodiaceae shows a number of anatomical
characters, which are correlated with the dry hab!tat of the members of the
Order. I

The mesophyll is in many cases differentiated into assimilatory and aqueous

tISSue. The assimilatory tissue is essentially composed of _palisade·cells.
Typical spongy tissue is absent in all the species examined by Volkens. Accord·
ing to Vol kens the aqueous tissue shows three types of differentiation. In
the first case (Rhagodia Billardieri, R. Br.) it forms several layers of palisade-
like cells with thin walls beneath the upper epidermis of the leaf; it has not
been determined whether these cells belong to the integumental tissue or
not. In the second case (in, "species of the genera Bassia, Kochia, Chenolea,
Panderia, l(irilowia and AtripZex) it forms the bulk of the mesophyll, whilst
the palisade-parenchyma is relegated to the middle layers of the mesophyll
(Atriplex Halimus, L., Fig. 158, C), or may even be confined to the immediate
neighbourhood of the vascular bundles of the. veins, the bundles then being
always provided with special sheaths (Bassia m'ttricata, AIl., Fig. 158, A).
In the third case, which is met with in the usually terete leaves of the repre-
sentatives of the Salicornieae, Suaedeae and Salsoleae, the chief part of the
internal tissue of the leaf is composed of centrally placed aqueous tissue, which
encloses the median vein together with its branches, lying at the periphery
of the aqueous tissue; the latter is often (in species of Halogeton, Salsola
and Traganum) surroUllded by a special starch-sheath, this betng followed
SOLnaDEJit U U
 CHENOPODIACEAE

by alayer- of palisadepparench~, and the latter by the epidermis (Fig. IS8, B).
Salts of calcium or other mineral salts are conunonly present, dissolved in the
contents of the aqueous ceUs.
The vascular bundles of the veins ()f the leaf are surrounded by special
sheaths in some members of the Order (in the CaInphorosmeae, Salsoleae,
and many species of AtriPlex, but not iIi the Chenopodeae, Corispermeae,
Polycnemeae, Salicornieae, Suaedeae and most Atripliceae, according to
Volkens 1); the cells of these sheaths are distinguished by their size and shape..
by the thickening of their walls, and by their contents (often including large
chloroplasts). In flat leaves (e.g. in Bassia mf4ricata, All., Fig. 158, A) these
sheaths are found in direct connexion with the vascular bundles, and even in
the finest ramifiCatio'ns of the veins they enclose the bundles" being either circular
or hippocreyiform in transverse section; in Inore or less terete leaves (e.g.
in. species of Halogeton, Salsola and T'aganum, Fig. 158, B) the sheath is
external to the strongly developed aqueous tis.~ue (see above), which encloseS
the fibrovascular system. '
The epideimIa of 'the leaf~ so far as is known, consists of a single layer
of celJs. The outer wall rarely has a thick cuticle (epidermis of the leaf of Agrio.
pltyllum, and that of the branch of Noaea, according to Volkens). Gelatiniza..
tion of the epidermis does not occur, but a granular coating of wax is some-
times found~ Stomata are present throughout the epidermis of the leaf
in the species I in which I examined the leaf-structure, and this is the case
even in plants with fiat leaves.. In the narrow, terete, succulent leaves
it is not a rare OCCWTence for the pairs of guard-cells to be parallel to
one another, their pores being' arranged transversely to the median vein of
the leaf .. A similar transverse arrangement of the stomata with reference
to the 10ngitudina1' axis is also occasionally met with on the stem '. In the
xerophilous species the stomata are commonly depressed below the surface.
Strictly speaking, special subsidiary cells have not been observed in this Order,
but in those cases in which the pairs of guard..cells show tAe abnonnal arrange..
ment mentioned above, and in 'Yhich the ordinary epidermal cells of the leaf
are also more or lesS' distinctly elongated transversely to the midrib" there
appear to be subsidiary cells placed parallel to'the pore (on the leaf of Cam-
plwrosma monsPeliae:u1n, Salsola Soda' and Suaeda /,uticosa, and on the axis
of Salicornia). It may be added that the narrow leaves of Poly&nemum IlrvetW,
L. also possess stomata showing a parallel arrangemept, but in this case the
pore ~ parallel to the median vein of the leaf.
The ba.fry covering p-esents very diverse fonns, Some of which are charac-
teristic of certain genera or groups of genera.. Unicellular hairs are not present,
but uniseriate hairs are widely distributed. Those on the leaf of Chenopodium
Botrys, L" for example, consist of a rather laJ:'ge number of thin·walled cells,
fCndering the hair somewhat articulated. The uniseriate, simple hairs,
which were pointed out by VoJkens in species of Bassia and KiriJowia
(Fig. 159, A) are especially characte_!istic; 1 also observed them in Cam-
phorosma and Echinopsilon, and according to Volkens they are widely
1 The tribes referred to in connexion with Volkens' investigations ate taken as defined in his
cla.ssl.ication in the Natilrliche Pfluzenfamilien.
I Th.5e were: C~iu/'l'l g!auctlm. L., Aln'p/,ex /taJlala, L, Camplumt.rma nzomjdliacNm, L.,
CwUJlrtnum k)'ssopifolium. 1..., Polycnemuln tl1"fJfflSe, L., llcn.;1IIJp:iilJn h.xs$opifoIUu, :&Ioq., SUIUda
./rrII{iosa, Fonk., s'iUstHa S«I4, L
S The aJDngem.ent o( the stomata as described above has been observed: I. Ob the l~ve& of
Campkotwma numspeliafu11l, L. t · Ec"i~psikn 1z7tSDjifO/il4S, Moq" lfeJ4gtton_a!P./.f/.ur(Jit4s, Moq.,
SalstJla Kali, L. S. kmii/o/ia, Forsk."1 S. SMa, L., SlMUtia.fruticosa, Forsk., S. marlHma, Dum. and
Tn:Ctlltum nudah"" Del., accordiJl2 to Brick, Vol1cens, Warming and my own observations; IL on
the stem of Sutw./4 f~lJSa (according to GeorghietT) and of the StdittWni4s (~g to Duval-
Jouve)
 CHENOPODIACEAE
distributed in the Camphorosmeae and SaIsole'ae. The hair is either seated
on one prominent epidermal c~ll or between two such celIs; it is composed of
a basal portion consisting of one Of a few cells, which have thin walls and, are
also often distinguished by their contents and of a long pointed terminal portion
p

consisting of one or a few cells, the walls of which are thick and more or less
densely covered with small solid papillae. Another special form of trichome
is seen in the two~armed hairs observed by Volkens in PetrosittWnia (Fig.
159, B); the stalk in this case is fonned by a few short cells, whilst the ter-
minal cell is cove~ed with papillae. Branched trichomes, which are of a similar

~ ·~·:~~X[·;::··z::;P

FIG. I~ Trichomes of the Chenopodia.ceae: A, KYi{Qwfa tr'lmtlJa, Bge.: B.. f'UlWi.

1'If{)fll'a of,j,nca, B~.; C AJripux leUcocfada &~; D-B, Gra)1ia polyraloide;B, HOO~. et
Am.; !"l CAtnopoJlu",. •"., L.: C? GrtJ'ta"j;l)1r~'tUs, Hook. etAm .. ; H~ Euroha. &n'tJIoidu,
C. A. Mey. ; J. A:qr,of IlI114r'anJol'ikr, L.; ~ Co","~u'" hyssoPi/o'.um, L.; 1., CAeno-
podium BtJlrys, L,; JI.o(), C, Illnrbl'#iddu, L,-A-X after Volkens, we :remainde:r original.

nature, and resemb1e stellate or candelabra hairs, have been met with by
Volkens in two groups of allied genera (in Grayia, Eurotia, Ceratocarpus and
Axyris, and in Corisp'ermu'!!!,.and 4griophyllum). The stellate hairs of Grayia
polygaloides. Hook. et Am. {Fig. IS9, G) and Eurotia ceratoe'des, C. A. Mey.
(Fig. 159, H) have a short stalk of a few cells, and a stellately. branched ter-
minal cell, whilst those of Axyris amarantoides, L. (Fig, 159, J) consist of a
long uniseriate hair bearing a few short ray-cells basally. The candelabra
hairs of COTispermum Jtyssopilolium, L. (Fig. 159, K) and Agriophyllum arena-
rium, Bieb. have a uniseriate axis and unicellular rays.
The trichomes, to which Volkens gave the name of vesicular hairs, are
especially characteristic of the Order, and give rise to th~ farinose surface
described by systematic botanists. According to Volkens they are found in
many species of Chenopodium, Rhagodia, Mo nolep is, A triplex, Exomis and
UU2
660 CHENOPODIACEAE

Salsola, and they also occur in Obione (Warming), Teloxys and Axyris (Geor-
ghieff). The bladder~like hairs are of the capitate type, but have no secretory
function. They consist of a unicellular or uniseriate stalk of variable length,
and of a unicellular head, which is thin-walled, and stores up water. The
head is mostly spherical in shape (Atriplex leucoclada, Boiss., Fig. I59, C, and
Cl}~nopodium glaucum, L.), but occasionally it is saccate. In the latter
c~ the longitudinal walls of the head are sometimes provided with short
protrusions (Fig. 159, E), thus constituting a transition to the stellate" hairs
of Grayia (see above); or the head is seated on the stalk in the same manner
as the terminal portion of a one- or two~armed hair (Fig. 159, D and F). Bio-
logically the bladder~like haits serve for the storage of water, as Volkens has
shown. When the waler contained in them is used up, they collapse and
form a membranous covering on parts of the plant where they have, been
present in large numbers.
True glandular hairs, secreting oil, have hitherto only been observed in
Camphorosma and in species of Chenopodium belonging to the sections Ambrina
and Botrydium. They are always uniseriate. In Camphorosma all the cells
of the filamentous glandular hairs are secretory, with the exception of a few
at the base; in Chenopodium Botrys (Fig. 159, L) the uppermost cells form
an ellipsoidal glandular head, only slightly marked off from the stalk,
which varies in length; in C. ambrosioides, L. (Fig. '159, M-O) there is
an approach to the bladder-like type of hair, since the terminal cell is espe-
cially strongly developed (being sac-like, sometimes with two unequal arms),
and is the principal or only seat of the secretory function.
Internal secretory receptacles have not been observed; but oxalate of
lime is commonly excreted, and chiefly in ·the .form of Clustered crystals or
crystal~sand. Octahedral or prismatic crystals," which apparently belong to
the tetragonal system (in the leaf of EChinopsilon hyssopifolius and Cam-
phorosma monspeliacum), are rare, and so are monoclinic crystals (Volkens).
Among the species in which I examined the leaf-structure, clustered crystals
only are present in Atriplex hastata, Corispermum hyssoPilolium, Polycnemum-
arvense, Echinopsilon hyssopitolius and Salsola Soda; in Chenopodium glaucum
and Camphorosma monspeliacum there are cells with clustered crystals, others
containing coarsely granular crystal~sand besides clustered crystals,jl.nd others
with COarse crystal-sand alone; Chenopodium Botrys has cells with coarsely
granular crystal-sand, while in Suaeda Iruticosa it is more finely granular. Crystal-
sand has also been observed in the stem in s~cies of the genera Ambrina,
Anabasis, Camphorosma, Echinopsilon, E%omis, Haloxylon, Salsola and Traga-
num as stated in my I Holzstruktur: In the succulent leaves of Salsola longi-
folia, Forsk., Halogeton alopecuroides, Moq., and Traganum nudatum, Del.,
the clustered crystals occupy; according to Volkens, a peculiar position. In
these species a loose layer of colourless, roundish cells is intercalated between
the epidermis of the leaf and the palisade-tissue; the majority of these cells,
or all Qf them, contain each a single clustered crystal; Volkens regards this
sheath of crystals as a protective arrangement against the attacks of snails.
In the species of the genera Atriplex, Blitum and Chenopodium examined
by Petit, the fibrovasclllar system of the petiole is formed by isolated vascular
bund1e~, which are arranged in an arc or ring.
3. STRUCTURE OF THE AXIS. Those Chenopodiaceae which exhibit con-
siderable growth in thickness are characterized by the same anomalous struc-
ture of the axis, as occurs in the N yctagineae and Amarantaceae. The
following statements regarding the course of development of this anomaly
are quoted from Morot's work: secondary rings or arcs of meristem (the
latter anastomosing reticulately) arise in centrifugal succession in the peri-
cycle (internally to the bast-fibres, where these are present) and produce
 CHENOPODJACEAE 661

secondary vascular bunclles as well as conjunctive tissue of varying structure.

The xylem-portions of these secondary vascular bundles always arise on
the inner, the bast-portions on the outer side of the secondary meristem.
The appearance of a transverse section of the axis differs, according to
the nature of the conjunctive tissue and that of the meristem. In the one
,case (e.g. in Camphorosma and Echinopsilon) rings of woody tissue alternate,
in the radial direction, with rings of thin-walled tissue; the former are
. traversed by lignifi~d or thin-walled medullary rays of varying breadth~
the broader rays sometimes separating the xylem~groups of the vascular
bundles of the same ring from one another; the other rings of tissue consist
of the phloem-portions of the vascular bundles together with thin-walled,
parenchymatous conjunctive tissue, which is present in variable amount.
In the second case, which is connected with that first described by means of
intermediate types, a,nd occurs in the majority of the Chenopodiaceae, the
vascular bundles are embedded in a prosenchymatous interfascicular tissue,
and exhibit a concentric, spiral, or irregular arrangement. In this case the
xylempgroups of the vascular bundles coalesce wHh. the conjunctive tissue;
they are only distinguishable owing to the arrangement of their vessels in
groups opposite the bast~portions. The latter vary in size, and are some·
times (Halostachya caspia, C. A. Mey.) remarkably small; a group of lignified
or unlignified parenchymatous conjunctive tissue is commonly associated
with them.
In a transverse section of the stem medullary vascular bundles also appear
in certain members of the Order, but these are rarely (in Acroglochin persi~
carioides, Moq., according to Georghieff) true medullary bundles. In most
cases 1 their "development shows that they are the normal Jeaf~trace bundlesf
and only apparently medullary; for the first secondary meristem produces
pith-like tissue external to' them before proceeding to the formation of the
secondary bundles.
The leaf,trace bundles sqmetimes possess considerable growth in thick·
ness, and thus delay the appea"ance of the anomalous growth (Camphorosma
and Echinopsilon, as also Blitum virgatum, Chenopodium murale and C. hybri-
dum~ according to De Bary, Kochia prostrata according to Georghieff)./ It
may be added here that in Halostachys caspia, C. A. Mey., 5-8 zones of growth
correspond to one period of vegetation, and consequently a decrease in the
size of the lumina of th<! vessels from within outwards may be recognized in
the region of these five zones, as in an annual ring.
The following statements regarding the distr~bution of the anomaly in
the Order are derived from the descriptions given by the various investigators
(Gernet, Regnault, De Bary, Morot, Solereder, Vol kens. Georghieff). Pro-
bably the anomaly occurs in all species having considerable growth in thick-
ness; at any rate no well established exception 2 in this respect is yet
known. Only those genera and species which exhibit little or no growth
in thickness have stems with normal structure. According to VoJkens,
Oreobliton, A phanisma- ,and M onolepis are examples: In Blitttm Bonus
Henricus, Beta trigyna and Hablt'tzia tamnoides the stem is normal in
structure only in its upper portion according to Georghieff. In the
following genera the anomaly has been described in the literature (the
sequence is that of Bentham and Hooker's classification): Acroglochin, Ha~
bUtzia, Rhagodia, .r.,.fonolepis, Chenopodiwn, Teloxys, Beta, Spinacia, Exomis,

1 In ma.ny herbaceous species of A fripl~, Beta, CllellopcdiulII and Obient; according to Georghieff
(see also De Bary). ,
, The two exceptions which Georghieff mentions (CamplzorcsJJla mcmpeliacum, L. and Gyay/(t
'SuI/uY/auai, Hook. et Am.') are non~existent; cf. Volkens',loc. cit.
 CHENOPODIACEAE
Atriplex, Grayia, EUTotia, Axyris, CamphOTOSma, Corispermum, Agriophyllum,
Chenolea, Kochia, Halostachys, Halocnemon, Arlhrocnemum, Salicornia, Suaeda p

Traganum, Cornulac.a, Haloxylon, Salsola, Noaea. Anabasis, Halogeton and

Sarcobatus.
The followmg facts may be noted regarding the detailed anatomical
structure of the axis. The stems of the herbaceous forms very commonly
possess subepidermal bundles of ·collenchyma, which project as nbs. The
portions of the epidermis adjoining the collenchyrnatous bundles externally
often consist of elongated cells (e.g. in Salsola Kali, L.), whilst the ·intervening
epidermal cells are approximately isodiametric in surface·view. The outer·
portion of the primary cortex sometimes (e.g. in species of Corispermum,
Salsola, Suaeda, &c.) contains palisade-tissue. In most cases the outer limit
of the pericycIe is formed by groups of sclerenchymatous fibres, so that the
occurrence of a sclerenchymatous pericycle may be considered as an ordinal
character· in the Chenopodiaceae, as is held by Georghieff. The sclerenchyma,
however, rarely attains a strong development; thus in Blitum Bonus Henricus,
for instance, only weak strands of collenchyma are present. According to
Georghieff a.nd Volkens the place of origin of the cork varies. In Camphorosma
and its immediate allies the subepidermal cell-layer becomes the phellogen
(Volkens); in Kochia prostrata, Schrad. the cork arises in the primary cortex
(Georghieff); finally in the Atripliceae, Salicomieae, Suaedeae and Salsoleae 1,.
according to Volkens, the cork cambium is constituted by ala yer of the pericycle,
which lies immediately to the interior of the groups of sclerenchyrnatous fibres,
where these occur. The prosenchymatous conjunctive tissue usually ha'i thick
walls, and always bears simple pits, and its cells may be arranged.in radial
series, or irregularly; medullary rays, such as occur in the Nyctagineous
genus Pisonia, have not been observed in this tissue, which in many cases
cannot be distinguished from the wood-prosenchyma of the vascular bundles,
since the elements of the latter tissue also have simple pits. True spiral
tracheae (primary vessels) only occur in the innermost vascular bundles,
i. e. in the leaf-traces. The pitted vessels of the wood generally possess
a diameter of '015-'°45 mm.; the twining species Hablitzia tamnoides has
vessels with wide lumina. The perforations of the vessels are as a rule
simple. Scalariform perforatiQ11s, 'with very oblique, almost l():lgitudinal
bars,' have only been recorded in Axyris amarantoides by Georghieff. Spiral
thickening of the vessel-wall is very widely distributed, and sometimes occurs
in all the pitted vessels, but in most cases only in those with narrow lumina;
it has been observed in certain species of Atriplex, Camphorosma, Eurotia,
H alocne1tton, H alostachys, H aloxylon, Kochia, N oaea, Rhagodia and Suaeda.
(for details see my , Holzstruktur and Georghieff's work).
J

Certain genera of the tribes Salicornieae and Salsoleae require a special

description. In the stem, as in the leaves of these genera (see above), stronsly
developed aqueous tissue is present; it surrounds the vascular system, and
is in its turn enclosed by a sheath of palisade-tissue, and this again by the
epidennis; the latter is sometimes (as in Anabasis aphylla, L., Brachylepis
eriopoda, Schrenk, &c.) strengthened by hypoderm. The aqueous tissue is
enveloped by a characteristic network of vascular bundles closed on all sides;
the xylem-groups of these bundles point inwards in the normal way in the
Salicornieae investigated by Dangeard, but are directed outwards in the Sal-
soleae examined by the same author. It may also be noted that the same
characteristic sheath of collecting-cells as occurs in the leaves has been observed
between the palisade and aqueous tissue of the stem but in the Salsoleae only.

I Again these groups are to be taken as defined in Volkens' revision of tite Chenopodiaceae.
 CHENOPODIACEAE
The features in question have been recorded in the following cases by Dal1geard :
amongst the Salicornieae in species ~f HaJostachys, Halocnemon, Arlhrocnemum and
Salicornia; amon~st the Salsoleae in species of O/a1.·ston, Noaea and Anabasis (with
Brachylepis); cortiCal vascular bundles with inverse orientation were also found in
species of H O1ani-no'Via, Caroxy Ion , Girgensohnia, H alimocnemis and H alanthium.
See also Volkens' work, with regard to species of Cornulaca, Haloxylonand Anabasis.
The stem of Salsola Kali, L. exhibits the layer of palisade, the characteristic sheath
and the aqueous tissue, but the network of bundles is wanting (Brick).
In certain species of Salicornia and Arthrocnemum the palisade-tissue
of the stem contains sac-like tracheids with wide lumina and strengthened
by means of a spiral band;' the longitudinal axes of these elements lie radially.
They are found in SalicOf'nia fruticosa, S. patula, S. peruviana, S. sarmemosa,
S. virginica and Arthrocnemum ambiguum, whilst they are only slightly de·
veloped in S. Emerici, and are wanting in S. herbacea, Arthrocnemum arbuscula,
A. capsicum and A. indicum (Duval-] ouve, Dangeard). In Salicorn£a macro-
stachya their place is taken by spicular cells, which are elongated in the same
direCtion, and slightly branched, especially at their apices. Similar' stereides'
occur in Arthrocnemum and Sarcobatus, according to Volkens.
The r09t exhibits the same anomalous structure as the stem. For syste-
matic'purposes it is an important uet that the anomaly may be demonstrated
in the root of species in which it does not appear in the stem, owing to the
small amount of growth in thickness in the latter (Blitum Bonus Henricus,
Beta trigyna, Hablitzia tamnoides, according to Georghieff).
Literature: Unger, Ban u. Wachst. d. Dicotyledonenst., St. Petersburg. 1840, pp. 104-8, Tab.
xiii-xiv.-Basiner, Kirghisensteppe, in Baer u. Helmersen, Beitr. z. Kenntn. d. russ. R., Bd. xv, .1848,
p. 93.-Schacht. Pftanzenz., 1852, p. 283 and Tab. xv.-Gemet, Holzk. einiger Ch., Bull Soc. imp.
Natural. de MoscoD, t. xxxii, 1859, pp. 164-88 and Tab. ii.-Regnanlt, Cyclospennees,Ano. !c. nat.,
sere 4, t. xiv, 1860, pp. 133-9 and pI. viii.-Sanio, in Bot. Zeit. 1863, p. 410 and 186,.., p. 2:16.-
A. Weiss, Pflanzenhaare, 1867, }O~ig. 198.-Duval-Jouve, SaHoomiees. Bull. Soc. bot. de France .[868,
pp. 132-40 and pl. I.-Van Tieghem, Symm. de struct., Ann. sc. nat., ser. 5. t. xiii, r871.-Martinet,
Org. de secret., Ann. sc. nat., set. 5, t. xiv, 1872, p. :139 and pt S.-De Buy, Vergl. Anat., 1877.-
Droysen, Anat. etc. d. Zuckerriibe. Diss., Halle a. S., 1877,37 pp.-Areschoug, Blad. anat., Minnes-
skrift Lund 18i8, P.ll1 et seq. and Tab. ix.-De Vries, Wachstumsgescb. d. ZuckerrUbe,Landwirtscb.
Jabrb. v. Thiel. Bd. viii, 1879. p. 417 et s.eq. and Ta.b. vi.-Hultberg, Salictwnia, Acta Lund., vol.
xviii, 1881-Z, 51 pp. a!ld 5 Tab.-Batalin, Salkornia !urlJacea, Bull. oongres de bot: et d'hort. a
St. Petersbourg~ 1884. pp. 219-33; Just 1884, I, p. 313.-Pichi, Beta vulgaris, var. sadu;vi/tnJ,
Nuov. GiOOl. boL !tal.. vol. xvi, 1884, Pp. :6l-81.-Volkens, Standort u. anat. Ban, Jabrb. Berlin.
ser.
Gatt. 1884, p. 37 et seq.----:-Morot, Pericycle, Ann. Be. nat., 6, t. xx; 1885, especially p. '77 et seq.
-Herail, Tige des Diootyl., AIUl. sC. nat., sere 7, t.:ii, 1885, pp. 34S-6.-So1ereder, Holzstr., 188S,
pp. U7-19.-Georghieft', Vergl. Anat. d. Ch., Bot. Centralbl.l887, ii, p. I I 7 et seq. and Tab. iv-vii and
1887, iii, p. 33 et seq.-Lohrer, Wurzel, in Wigand, Bot. Hefte. ii, 1887. ptl. 30-3 and Tab. ii.-Petit,
Petiole, Mem. Soc. sc. nat. et phys. de Bordeaux, ser. 3. t. iii, ~87J pp. 236-9 and pI. ii.-Volkens,
~pt.-arab. Wilste, 1881, pp. 138-4:1 and Tab. xi, xii and xv.-Brick, Bait. Strandpfl., Schrift.
natorf. Gesellseb. in Danzig, Bd.. xvii, I. Heft, 1888, pp. 136-43 and Tab. ii.-Dangeard, Stroct. des
Salicomieae et des' Salsolaceae, Bull. Soc. Linn. de Normandie, ser. 4, t. ii, Caen, 1888, pp. 88-gS;
see also Boll. Soc. bot. de France 1888, pp. 157 and 191 et ~.-Van Tieghem, in Ann. &c. nat.,
sere 7, t. viii, 1888. p. 7o.-Warming, Fra Vesterbavskystens Mstrskegne, Vidensk. Meddelels. batnrh.
Foren. i Kbbvn. 1890. pp. 214 et seq., 121 and 32s.-Arcangeli, Stratt. delle foglie dell' Atriplu,
Nuov. Giom. bot., vol. xxii, 1890, pp. 426-30.-Benecke, Nebenz. etc., Bot. Zeit. 1892, p. 51+-
Volkees, in Natiirl. Pftiiiienfam., iii. Teil, Abt. I a, 1892, pp. 39-4+-H. Schenck, Anat. d. Lianen,
1893, pp. 48-9 and 35l.-Houlbert, Bois dans les Apeta1es, These, Paris, 1893. pp. 7o-I.-Herbst,
Markstr., Bot. Centralbl. 1894. i, p. 295.-Russell, Climat mM., Ann. se. nat., sere 8, t. i, 1895.
p. 336 et aeq.-Schnbert, Parenchymscheiden, Bot. Centralbl. 1891, iv, p. 63 et seq.-Warming,
Halofyt Stud., K. Danske Vid. Selsk. Skr. 1897, pp. 190 et seq., especially 315 and 229 et seq.-
[Fron, Racine des SUlUda et des Sa/sola, Compt. rend., t. cxxv, 1897, p. 366 and Struct. spiratee des
rac. de cert. Ch., loco cit., t. cxxvU, 1898, p. 563.]

BASELLACEAE.
The limits of this small Order, which was established by Moquin-Tandon
in DC. Prodr. t vol. xiii 2, coincides with those of Subordo II Baselleae in
the classification of the Chenopodiaceae in Bentham and Hooker"s Genera
 BASELLACEAE
Plantarum. The elevation of this Suborder to the rank of an Order and its
separation from the Chenopodiaceae is also warranted on anatomical grounds.
Neither the stem nor the root of the Basellaceae exhibits the anomaly found
in the Chenopodiaceae, whilst on the other hand intraxylary soft bast occurs.
Other noteworthy features for the diagnosis of the Order are the structure
of the stomatal apparatus (which belongs to the Rubiaceous type) and the
sclerenchymatous peri cycle in the stem. Oxalate of lime is excreted in the
form of clustered crystals, and more rare1y in that of solitary tetragonal crystals.
Mucilaginous sap, which can be drawn out into threads, has been observed in
the leaves and cortex of certain members of the Order; it is due to the
occurrence of spherical mucilage-cells.
I have made a detailed examination of the leaf·structure in Basella rzebra,
L., Boussingaultia baselloides, H. B. K. and A nredera spicata, Pers. Trichomes
are not present. Both sides of the leaf are provided with stomata, which are
accompanied by two or more subsidiary cells, arranged parallel to the pore.
Large, spherical mucilage-cells, in which the mucilage apparently originates
from the membrane and does not belong to the cell-contents, are only found
in the mesophyll in the species of Anredera and Bouss£ngaultia. Clustered
crystals of oxalate of lime are present in the leaf.-tissue of the three species
investigated; in Baselta rubra there are also prismatic or octahedral crystals
of the same salt, and in the dried leaf of A nredera spicata yellowish sphaero-
crystaIIine masses of unknown chemical composition.
The petiole in Basella rubra contains two large, nearly median vascular
bundles, adjoined on either side by smaller bundles; in BottssingauUia basel-
loides there is an arc of wood and bast (Petit).
The structure of the axis has been investigated by Morot in Basella rzebra,
Boussingaultia baselloides and Ullucus tuberosus, and by me in Basella rubra
and A nredera spicata. In all these species there is a sclerenchymatous peri-
cycle, which is weakly developed in Ull'ltcits tuberosus, but strongly developed
and continuous in A nredera spicata; in Boussingaultia baselloides it is broken
up. The vascular bundles are of different sizes, smaller strands being inter-
calated between the larger; the bundles are isolated and arranged in a ring.
The larger bundles possess intraxylary soft bast, which is developed to a
variable extent and only arises secondarily, according to Morot. Thetdiameter
of the vessels is large (reaching '2 mm. or more) in Anredera, and probably
also in the other twining species; the perforations are simple. The formation
of cork takes place in the outermost cell-layers of the primary cortex in Bous,,:
singaultia baselloides.
Literature: Strasburger, Spaltoffit., Pringshcirn Jahrb. 1866-1, p. 316 and Tab. nxviii.-.Morot,
Anat. d. B., Bull. Soc. bot. de France 1884. pp. I04-7.-Georghieff, in Bot. Centralbl. 188" ii, pp.
371-J..-Petit, Petiole, Mem. Soc. sc. nat. et phys. de 130rdeaux, Set. ,3, t. iii, 188" p. 238 and pl. i.
-Volkens, in Natiirl. Pflanzenfam .• iii. Teil, Abt. I a, 1893, p. 125.

PHYTOLACCACEAE.
1. REVIEW OF THE ANATOMICAL FEATURES. In its anatomy this Order
only possesses a few distinctive characters common to all its members. Such
are the absence of external glands and of special internal secretory recep-
tacles, the simple perforations of the vessels and the superficial development
of the periderm. On the other hand the type of stoma and the nature of
the pitting in the wood~prosenchyma vary within the limits of the Order.
The Rubiaceous type is well marked only in certain genera (Ledenbergia,
llfonococcus, &c.); in other cases the stoma is surrounded by a relatively
large number of epidermal cel1s- exhibiting no special arrangement. The
 PHYTOLACCACEAE 665

wood-prosenchyma bears simple pits in the Rivineae (excl. Microtea) and

Euphytolacceae, but in the Gyrostemoneae, Microtea and the anomalous
genus Stegnosperma, the pits in this tissue have narrow though distinct borders.
The pericyde is sclerenchymatous, containing a composite and continuous
sc1erenchymatous sheath (e.g. in Codonocarpus, Gallesia, Phyto/acca, Seguieria),
or isolated groups of bast-fibres, or isolated fibres. Secondary hard bast has
not been observed. In' a number of genera belonging to the tribes Rivineae
and Euphytolacceae the structure of the stem or root is anomalous (stem
and root of Phytolacca, stem of Ercilla, Gallesia and Seguieria, root and (?)
stem of Anisomeria) j the anomaly consists in the formation of successive
secondary rings of vascular bundles in the pericycle. Oxalate of lime never
occurs in the form of the ordinary soJitary or clustered crystals. The Rivineae
(excluding Microtea) are distinguished by possessing styloids; in the Euphy-
tolacceae and the anomalous genus AgdesUs raphide-sacs are characteristlc,
and in the G)Tostemoneae the absence of oxalate of lime; the anomalous genus
Stegnosperma possesses sphaerites. Raphides and styloids not uncommonly
give rise to pellucid dots in the leaf. The hairy covering consists of, simple
uniseriate hairs. The following special anatomical features of the Order
may be shortly referred to at this point: the occurrence of hypoderm
on the upper side of the leaf (Galles£a Gorazema, Moq.); gelatinization of
epidermal cells (species of Codonocarpus and Gyrostemon) and more rarely of
internal cells in the tissue of the leaf; medullary vascular bundles in the
stem (Phylo/acea dioica, L.}; and sacs, crowded with numerous crystals of
varied shape (species of Gallesia and ]l.lonococcus).
2. STRUCTURE OF THE LEAF 1. This is bifacial in most members of
the Order, more rarely centric (in Gallesia Gorazema, Moq., Codonocarpus aus-
tralis, Cunn., Gyrostemon ramulosus, Desf., and according to Schulze Didy-
motheca thesioides). When the leaf is centric the mesophyll either consists
of palisade-tissue throughout (Gallesia Gorazema), or the middle of the leaf
contains aqueous tissue (Gyrostemon ramtdosus). The epidermal cells have
straight or undulated lateral walls. Gelatinization of the epidermis is men-
tioned by Blenk as occurring in Codonocarpus australis and Gyrostemon Yatnu·
losus; cells with mucilaginous membranes are also found in a subepidermal
position in the larger veins of the leaf in Codonocarpus australis, and in the
central aqueous tissue of Gyrostemon ratnulosus. In the species of Petiveria
the epidermal cells contain chlorophyll .. (Schulze). A hypoderm, composed
of two layers of rather large cells, is developed on the upper side of the leaf
in Gallesia Gorazema. The stomata are either confined to the lower side of
the leaf, or {in Phytolacca dioica. L., and according to Schulze es~~cially in
species of ~licrotea, CodO?'Wcarplls and Didymotheca) occur on both surfaces.
According to my own observations those of Codonocarpus arid Phytoiacca are
always surrounded by a number of ordinary epidermal cells exhibiting no
special arrangement, whilst in Rivina and Gallesia stomata belonging to the
Rubiaceous type ~.l~o occur, the latter being the only type present in
Ledenbergia. and -"!v[onococc'lis. Schulze describes the Rubiaceous type as
occurring in Pet£veria, MonococcttS and Gallesia, and for the most part in
Ledenbergia and Mohlana. In the larger veins there is generally no scleren-
cbyma accompanying the vascular bundles, but in those of Gallesia G01'azema
a sclerenchymatous ring is developed.

J The strul;ture of the leaf and nxis has receDtly been examined in detail by Schulze. In his
work the following genera. were i/lvestigated: lU'llina. Ledmhr~'a, .•V~hlanaJ Petivcria, ,.Vul'~ta,
ftfonQtoccus, Seguieria and Gallesia (Tribe Rivineae); Phytolacca, ErciJ/Q. and Anisomena (Tribe
Euphytolacceae); Didymol/uca, CodoHocarpus nod Gyrostemon (Tribe Gyrastemoneae) i and the
anomalous genera SteglloS#1"I1lll and Agdestis.
666 PHYTOLACCACEAE

The mode of excretion of oxalate of lime differs in the different tribes.

The genera Rivina, M ohiana, Petiveria and Seguieria (according to Blenk).
Gallesia (according to Radlkofer), Ledenbergia and Monococcus (according to
Schulze),and ViUamilla (V. octandra, Hook. f.) all belonging to the Tribe Rivineae,
possess styloids (sometimes having the shape of a swallow's tail); in Microtea,
which is included in the same tribe by Bentham and Hooker, oxalate of lime
appears to be wanting (according to my own investigation of the axis and leaf
in M. maypurensis, Don, aud according to Schulze). The genera Phytolacca,
Ercilla and A nisomeria, belonging to the Tribe Euphytolacceae, contain
raphides, according to Blenk; the same form of crystal also occurs in the
d.oomalous monotypic genus Agdestis, according to Schulze. In the Gyro-
stemoneae (Didymotheca, Codonoca rp us, Gyrostemon) oxalate of lime has not
been observed. The anomalous genus Stegnosperma is characterized by
possessing large sphaerites of oxalate of lime (Schulze). It may be added that
the styloids and bundles of raphides occur both in the mesophyll and in the
parenchymatous tissues and bast o£ .. the axis; tho~e in the mesophyll are
arranged either parallel or at right angles to the leaf-surface,
and in the latter case commonly give rise to transparent dots.
The styloids often attain very considerable dimensions. In
Gallesia Gorazema and Monococcus echinophorus, F. v. Milll., the
styloids are accompanied by special crystal-sacs in the meso-
phyll and veins of the leaf (according to my own observa-
tions), and in the former species similar sacs occur in the
primary cortex of the axii; they are filled with numerous,
rather small crystalline bodies, the shape of which varies, being
either shortly prismatic or rounded (Fig. r60). In the same
way small crystals, resembling styloids, are found in consider·
able numbers side by side with well-developed styloids in
certain cells of the bast of Rivina humitis, L., and in the leaf of
V illamilla octandra. Zimmermann states that nuclear crystal-
loids occur in the cells of the mesophyll in Rivina humitis and
Ledenbergia rosea-aenea, and in the former species in the epi-
s!'~:i~~e ;!rn~ta~r dermis of the leaf also. Kruch has recorded crystalloids,
the leaf of Galluia occurring free in the cell-sap, in the apical regioncOf the le~f
&.or;;::.rlJ • Moq.- in Rhytolacca abyssinica, and according to the same author
peculiar spherical bodies containing proteid and varying as to
the nature of theiisurface, occur in cells of the apex of the leaf in P. icosandra.
Special internal secretory ~eceptac1es 1 are absent.
The hairy covering consists only of simple uniseriate hairs; glandular
hairs are not present.
In Phytolacca decandra, L., a transverse section of the petiole m the charac-
teristic region shows an arc of three isolated vascular bundles, with smaller
bundles adjoining them towards the margins of the petiole.
3. STRUCTURE OF THE AXIS. Some of the genera belonging to the Tribes
Rivineae and Euphytolacceae are distinguished by having an anomalous
stem-structure, which consists in the appearance of successive rings of vascular
bundles in the pericycle. This anomaly has long been known to occur in
species of Phytolacca (P. abyssinica, Hoffm., P. acinosa, P. decandra, L. t P.
dioica, L., P. icosandra, L.), where it is found not only in the stem, but also
in the root (Avetta). Kruger also observed it in a-piece-.Qf-the branch of

1 Scbul:r:e correctly describes brown, tanniniferons cells as occurring in the mesophyll of Galles;,,:
GIJ'nZZOfIa; bnt his statement regarding the occnrrence of snbeficlermal secretory cells (=the gela-
tinized portions of the membrane of mucilaginous epidenna celli) in the leaf of CDdonocarpus
australis and in a plant with linear leaves, described as 'Cyros/muJn aust1'l1Iis,' is mcorrect.
 PHYTOLACCACEAE

ErciUa I)olubilis 2 cm. in thickness, and I have met with it in Seguieria /loti-
bunda, Benth. and S. longilolia, Benth., to which on Schulze's authority We
~ay add S. americana l. According to Schulze this anomaly is also found
in GaUesia Goraz&ma, Moq. and A nisomeria drastica, Moq.; I can confinn
these statements in the case of the stem of the former species, and at any
rate as regards the root of the latter. Transverse sections of sufficiently th~ck
branches of the above-mentioned species of Phytolacca, ErciUa, Seguieri(l and
Gallesia show al~ernating concentrIc rings of wood and bast. The inner-
most ring, which alone contains -primary spiral tracheae, consists of the leaf-
traces. In the young branen a parenchymatous pericyc1e adjoins the bast-groups
of the leaf· traces, and this is followed by a compOsite and continuous riIll of
sclerenchyma, which bounds the primary cortex on the inner side. The paren-
chymatous pericyc1e is the seat of the new formations, and in it there arises
a meristem, which before giving rise to a secondary ring of vascular bundies,
produces parenchyma on its inner side, part of this tissue becoming lignified.
Hence each secondary ring commences with a zone of parenchymatous tissue,
the amount of which varies. The medullary rays between the vascular bundles
of each ring are broad in most cases, and are generally lignified; some-
times (Gallesia Gorazema) even the rays traversing the zones of bast are ligni-
fied, at least in the internal rings of growth. The greater part of the napiform
root of Am'someria drastica consists of, large parenchymatoUs cells containing
raphides, as I found on examining the slices of the root which accompany
Lechler's herbarium-material; the centre of the root is occupied by the primary
vascular tissue, but this is split up by secondary dilatation; in the ground-
tissue adjoining this central mass of tissue one finds several concentric Circles
of iso1ated vascular bundles, in wl:ich each xylem-group is marked by an
elongated row of vessels with rather narrow lumina, the groups of bast being
clearly developed 2. It may also be noted that Phytolacca dioica has medul-
lary vascular bundles, the structure-of which is concentric according to Nag-eli
and Douliot, the phloem wing central. I t remains to be determined by future
investigation whether the other genera of the tribes Rivineae and Euphyto-
lacceae will not also prove to have anomalous structure in the axis, when suffi-
..ciently old material is examined. From Regnault's description, which howeyer
J5 not qui!.:) clear, it appears that Rivina humilis, L. possesses the same anomaly
as is found in Phytolacca. Schulze observed the commen~ment of anoptalous
growth in the root of Petiveria. In Ledenbergia rOSea-al4tM, on the other
hand, anomalous growth in thickness seems to'me to be out of the question,
since I found branches almost 2 em. in diameter exhibiting normal structure.
The structure of the wood has been examined by me in species of Rivina,
M icrotea, S eguieria, Gallesia, Ph y tolacca, E 'fcilta, A nisomeria, Codonocarpus
and Gyrostemon, and also by Schulze. The size of the lumina and the mode
)f arrangement of the vessels vary (maximum diameter = '03-'15 mm.);
the size of the borders of the pits in the vessels is also variable (diameter reach-
ing ·009 mm. in Phytolacca). Spiral thickening of the pitted vessels has been
observed in.A nisomcria dyastica. The perforations of the vessels are simple
in all cases. Where'the vessel-wall is in contact with parenchyma, it either
bears bordered pits only (Seguieria, Codonocarpus, Gyrostemon), or bordered
and simple pits (Gallesia, Phytolacca). Broad medullary rays are found (for
example) in GaUesia and Phytolacca; in Seguieria and Codonocarpus the rays
are from one to three cells in breadth; in Anisomeria distinct ray-parenchyma
1 The anomalous structure of this plant is incorrectly interpreted inMoller's work (loc. cit. 1876).
2 }~urtber investigation is re'1uired to determine whether tbis anomaly also occurs in the stem. of
Anisomeria arastica, as stated by Schulze, or whether woody tissue, some of which i!I unlignified, has
not been mistaken for phloem ~ II did, not, find the: J nnomaly in herbarium-material. even in thick
hranches.! ;.luU 10 ,jnl.Gx-O .r I', 'I I
668 PHYTOLACCACEAE
is not present. The wood.prosenchyma is characterized by having simple
pits in the Rivineae (excluding ]'.<[icrotea) and Euphytolacceae so far as investi-
gated, but in the Gyrostemoneae, ftlicrotea, and according to Schulze also in
Stegnosperma it is provided with small but distinct bordered pits. The
wood·fibres of Rivina humt"lis are septate in some places.
The following statements regarding the structure of the cortex are based
on my own examination of Rivt'na humih's, Ledenbergia rosea-aenea, Seg'ttt'eria
longi/olia, Gallesia Gorazema, Phytolacca dioica and Codonocarpus australis,
and on Schulze's description. In Rivina humilis and Ledenbergia ros'ea-aellea
the pericycle contains small isolated bundles of sc1erenchymatous fibres, while
in the rest of the species examined by me there is a composite and continuous
ring of sclerenchyma; Schu1ze states that bast.. fibres occur in the pericycle
in all cases, the fibres being some6mes accompanied by stone cells. Secondary
hard bast has not been observed. In Phytolacca dioica the primary cortex
shows a strong development of collenchyma in its outer portion, whilst' in
Codonocarpus australis it contains a few sclerosed. parenchymatous cells;
in Seg'ttieria longifolia the cortex is bounded by Ian epidermis composed of
cells elongated like palisade-tissue. The formation of cork takes place in the
subepidermal cell-layer (Ledenbergia, Phytolacca and Codonocarpus, accord-
ing to my own observation; Pett'veria, Seguieria, ,Gallesia, Phytolacca and
Gyrostemon, according to Schulze).
Literature: Nag-eli, Beitr. z. ",iss. Bot., Heft I, 1858, pp. 26 and 118.-Regnault, Cyclosp.,
Ann. sc. na"t., ser. 4, t. xiv, 1860, pp. 139-44 and pI. 9.-MolIer, HoIzanat .• Denkschr. VI'iener Akad.
18 76, pp. 75 and 37I.-De Hary, Vergl. Anat., 18n.-KrUger. Anom. Hobbild., Diss., Leip~ig,
J884, p. :n.-Blenk, DUTchs. P., Flora 1884, p. 375 et seq. and sep. copy, pp. 87-9·-Doultot,
Faisc. med. du Phyto/acca d;oica, Bull. Soc. bot. de :France 188S, pp. 391-::I.-Morot, Peri cycle, Ann.
sc. nat., SeT. 6, t. xx, 1885. pp. 275-7.-Herail, Tige des' Dicotyl., Ann. sc. nat., ser. 7, t. ii, 1885,
pp. 243-5.-So1ereder, Holl.Str., 1885. pp. 217-19.-RaCllkofer, Dnrchs. P., Sitz.-Ber. Munch.
Akad. J 886, pp. ~26-7.-[Avetta, Anomalie di struttura nelle radici etc., Ann. dell' 1st. bot. di Roma
1887·J-Petit, Petiole, Mem. Soc. sc. nat. et phys. de Bordeaux, ser. 3, t. iii, 1887, p. 2iI.-Eiselen,
J{haph., Di.s., Halle a. S., 1888, p. 16.-Heimerl, in Natiirl. Pffanzenfam., iii. Teil, Abt. I b, 1889,
p. 2.~Ross, Periderma, Malpighia, vol. iv, 1890-1, pp. 98-1oo.-Zimmermann. Pfianzen2:elle,
2. Heft. 1891, p. 137.-Houlbert, Bois sec. dans les Apetales, These, Paris, 1893, pp. 71-4.-
H. Schenck, Anat. d. Lianen, 1893. pp. 53-5 and 252, Tab. ii.-Herbst, Markstr., Bot. Centralbl.
1894. i, p. 295 et seq.-[Kruch, Ricerche anat. ed istolog. sulIa Phyt()/aaa d£'oica, Ann. del R. 1st.
bot. di Roma 1894. pp. 124-54, 3 Tab.]-C. Schulze, Anat. fian d. lnJu. d. Axe in der Fam. d. Ph.
ttc., Diss., Erlangen, 56 pp. and 1 Tab.-Kruch'fCristalloidi della Pkyto/acca ahpsi,uca, Atti della
reale Accad. dei Lincei, vol. v, 1896, pp. 364-6 and Sferoic1i e cristalloidi di alcune Fitolacche,
Annuar. R. ht. bot. di Roma, vol. vii, r897. I l pp. nnd Tab.

BATIDEAE.
This small Order, the systematic position of which is still uncertain,
includes only the monotypic genus Balis with B. maritima, L.
The axis of this plant is normal in structure. In transverse section
the mass of wood exhibits broad medullary rays, and rounded vessels
with thick walls and of no great diameter (reaching '03 mm.). The vessels
have simple perforations. In contact with parenchyma of the medul1ary
rays, the vessel-wall bears botdered pits. The wood-prosenchyma has thick
walls, bearing small, but distinctly bordered pits. The wood.parenchyma is
confined to the neighbourhood of the vessels. Regarding the structure of
the cortex it may be mentioned that isolated, massive groups of primary bast-
fibres 'are developed in the pericycle. In the secondary bast also scattered
sc1erenchymatous fibres with yellow walls are found. The formation of
cork takes place in the pericycle on the inner side of the bundles of primary
bast-fibres. In addition to the cork a large mass of phelloderm with abundant
intercellular spaces is formed by the phellogen. Oxalate of lime is present
 BATIDEAE

in the pith, primary cortex and bast, in the form of solitary crystals, which
in some cases have a corroded appearance.,
An important feature in the structure of the leaf lies in the fact that the
stoma is enclosed by two semilunar subsidiary ceBs arranged parallel to the
pore. The majority of the stomata are orientated so that their pores are
transverse with regard to the longitudinal axis of the narrow leaf. Trichomes
are not present. Oxalate of lime appears in the form of clustered crystals~
crystal-conglomerates, and also solitary crystals.
Literature: SoIereder, Holzstr., 1885, p. 22o.-Warming, in Vidensk. Meddel. naturh. For. i
Kbhvn., 1890, p. 23;1.-Dammer, Batis maritima, Ber. deutsch. bot. GeseIlscb. 189:l, pp. 643-4 and
in ~aliirl. .Pfianzenfam., iii. Teil •. Abt. 1 a, 1893, p. I}o.-Houl~rt, Bois sec. dans les Apetales,
These, PariS, 1893, p. '75.-Wannmg, Halofyt Stud., K. Danske Vld. Se\sk. Skr. 1897. p. H3.

POLYGONACEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. The following features
may be pointed out as characteristic of the Order: the absence of a special
type of stoma, the simple perforations of the vessels, and the simple
pits borne by the wood-prosenchyma. The peri cycle shows different types
of differentiation; the formation of cork is superficial or internal. Oxalate
oi lime forms clustered, or ordinary solitary crystals. The following types
of internal secretory organs are found,: cells with resinous contents (species
of Polygon1lm); branched secretory cells with brown contents (cortex of the
root in species of Calligonum); and elongated tannin-sacs (stem in species
of Polygonum and Fagopyrum). The hairy covering consists chiefly of simple
unicellular trichomes and glandular hairs,.in which the head is small, or
peltate and of larger size, being either unicellular or divided by vertical walls.
Anomalous structure of the axis occurs in some cases, the .following types
being represented: development of secondary vascular bundles in the peri-
cycle (Antigonon); occurrence of medullary vascular bundles (Rumex, Rheum),
with wood and bast inversely orientated, or with concentric structure "(the
phloem being central); presence of intra-xylary phloeIp. (Emex); development
of bundle~ of soft bast in the interfascicular tissue of the phloem (Polygonum,
Fagopyrum); or occurrence of cortical vascular bundles (Calligonum). The
followin_g special anatomical features can be employed in detailed diagnosis:
mucilaginous epidermal cells in the leaf; subepidermal strands of sclerenchy-
matous fibres and aqueous tissue in the same posil:ion in the leaf and stem;
hypoderm in the leaf; strands of collenchyma; the development of a: character-
istic sheath in the stem, and so on.
2. STRUCTURE OF THE LEAF. Since only scanty information has hitherto
been obt~ined regarding the structure of the leaf in the Polygonaceae, it was
necessary for me to make a rough survey by investigating a few representatives
of the five tribes (Eriogonum angulosiiin, Benth., Nemacaulis Nuttallii, Benth.,
Polygonum Bistorta, L., Rumex scutatus, L., Coccoloba ovata, Benth., C. longe-
pendula, Mart., Triplaris Pachau, Mart.). On the basis of these and
other investigations the following facts may be pointed out. In Coccoloba
o'IJata, Polygonum Bistorta, the land-form of P. amphilrium, L., in P.
equisetiJorme, Sibth. et Sm. and P. aviculare, the epidermis of the leaf
contain.s cells in which the inner membranes are mucilaginoUs. According
to Johow hypoderm occurs in some species of Coccoloba, such as G. nviteYa.
The stomata are always surrounded by several epidermal cells, wbich
are not differentiated by their shape except in Coccqloba and Triplaris.
They either occur on both sides of the leaf (Eriogonum angulosum,
Nemacaulis Nuttallii, Rumex scut«tt(s), there being very few on the upper
 POLYGONACEAE

surface in some of
these cases (Polygonum Bistorla, Coccoloba OfJata), Or they
are confined to the lower side (Triptaris Pachau); in the floating leaves of
the water-fonn of Polygonum amphibium they are of course only found on
the upper side. The leaf-structure maybe centric or hifacial. The leaves of Poly-
gonurn equiseti/orme, a desert-plant, exhibit a sheath of palisade-tissue beneath
the epidermis, and colourless aqueous tissue in the interior (Volkens). Accord-
ing to Dammer the vascular bundles of the veins (especially those of the median
vein) are accompanied in Coccoloba by a varying amount of sclerenchyma;
in species of Polygonum and Rheum the sclerenchyma· is replaced by collen-
ch~a. The occurrence of subepidermal strands of sclerenchymatous fibres in'
the leaf of Polygonum equiseti/orme des~rves special notice; they are separated
from the assimilatory tissue by a layer of aqueous cells, which evidently serves
to connect the palisade parenchyma (beneath these strands) with the water-
storing epidermis. It may be added that similar sclerench~atous strands
are also found in the stem of Polygonum aviculare, L. (Grevillius) and Calli-
gonurn comosum, VHerit. (Volkens). \'
The haJry covering consists of clothing and glandular hairs. The cloth-
ing hairs are generally long simple unitellular trichomes (E,iogtmum, Nema ..
caulis). In Polygonum amphibium there are also conical shaggy hairs 011 the
leaves of the land-form; the cells of these hairs exhibit a multiseriate arrange·
rnent, and their tips project as papillae. In the glandular hairs the head is
always unicellular, 01: divided by one or more vertical walls. In Eriogonum
angulos1,m the glandular hairs are composed of a cylindrical or conical basal cell,
a neck-cell, and a unicellular or bicellular head; in Poly~onum Bistcna and
Rumex scutatus there are sessile glands, the heads of whIch consist of from-
two to four cells, though sometimes unicellular in Rumex scutatus. Larger peltate
glands are present in Coccoloba and Triplar£s. They consist of a short stalk,
composed of two cells lying side by side, and a shield with an entire margin;
in Coccoloba ovata and C. longependula the shield is mainly divided by radial
vertical walls, only a certain number Qf which reach the centre, whilst in T,i-
plans Pachau the shield is subdivided by division.walls, running in various
directions, into a number of cells, which are four-sided or polygonal in surface··
view. External glands, similar to those of Polygonum and Rumex described
above, give rise to a mucilaginous gununy excretion on the buds in Jtplygonum,
Rheum and Rumex; but here the glandular structure of the wall of the trichome
is continued from the point of insertion of the latter into the smooth epidermis;
there are also mult iseri ate, elongated glandular shaggy hairs of more com·
plicated stnlcture, and hav.ing the form of a lamella Of band.
The nectaries found on the leaf· cushions of Polygonum cuspidatum, Willd.,
and of certain species of Muehlenbeckia, form depressions, in which shortly
stalked peltate hairs are densely crowded together {Delpino and Morini);
in these hairs the shield consists of prismatic cells, which are sometimes tran~~
versely divided.
The internal secretory receptacles include, in the first place, cells (or
cavities ?), 1illed with yellowish contents, which are soluble in alcohol. Such
secretory cells were found by Bokorny beneath the epidermis of the leaf in Poly-
gonUffl acre, H. B. K., and occur also in P. punctatum, Ell. ; they give rise to trans-
parent dots in the leaf. Other forms are the branched secretory cells observed by
Rindowsky in the cortex of the root in species of Calliganutn, and the long
tannin-sacs with brown, gelatinous contents, found by E. Schmidt -in the stem
of certain species 1 of Polygonum; the former have thin walls and are tilled

I" Viz. P. amp""!Jium, P. Ii7d~piper, P. /apatlzifo/z"u"" P. brientale, P. Ptt'sicaria, P. tinttCriU"'i

P., fTirgj"'''UIIl j F~m C)'mDsum; Meissn., F. emarginatunI, Mch., F. uculmtu1fl, Mch.•
F. /tllat"Ullm, Gacrtn.
 POLYGONACEAE

with a brown, tanniniferous gum-resin. The tannin-sacs resemble the well-

known sacs of Sambucus nigra. They at"ise by the elongation of single cells,
and attain a length of 12 cm. or more, so that they traverse entire internodes ;
in transverse section there is little to distinguish them from the neighbour-
in~ cells. They occur in the pith and bast, and sometimes also in the
pnmary cortex and are not found in the subterranean shoots, or only
in small numbers; they are not presen t in the roots. The brown
contents, seen in the tannin-sacs in older stages, sometimes also appear in
ordinary parenchymatous cells, which contain abundant tannin from the first.
Here we may also mention the bright yellow or red contents in the cells of
the medullary rays of the rhizome of rhubarb; they include the active chry-
sophanic acid.
Oxalate of Ume is for the most part developed in the fonn of clustered
crystals, but ordinary solitary crystals have also been observed. 1
The str~cture of the petiole, which has been investigated by C. de Candolle, Plitt
and Petit, is uniform only in so far as the vascular bundles have an isolated course.
In Muehlenbeckia complexa and M. varians the characteristic region exhibits two
vascular bundles with their xylem-groups pointing towards one another. In Rumex
AcetoseU'a'this part of the petiole contains an arc of isolated bundles, which is con-
tinued in~o the petiolar wings. In species of Polygonum, owing to the presence of one
or more larger vascular bundles in a median position, the arc is converted into a closed
ring; this is also the case in A ntigonon leptopus. In the petioles of certain species
of Rumex and Rheum there are numerous medullary vascular strands within a more
or less distinct ring of isolated bundles -; where the former become very numerous
(Rheum), the vascular bundles of the petiole appear quite irregularly scattered in
a transverse section. ~
The structure of the ochreate stipu!es has been examined by Grevillius and
O. Schultz in species of Rumex, Rheum, Polygonum and FagopYl'Um. The ochreae
exhibit mechanical strengthening in a certain number of the species only; it
may take the fonn of a thickened epidermis, the development of colfenchyma on the
inner (P. aviculare and P. Rafs") or outer side (P. ds'varZ:catum) of the ochreae
collenchymatous difierentiation of the whole of the ground.tissue (P. Hydyopipe"
Rheum), the occurrence of rings or crescents of ·sclerenchyma accompanying the fibro-
vascular system, or the differentiation of subepidermal strands of sclerenchymatous
fibres (P. amplexicauie, P. Bisrorta, &c.). A number of these anatomica~: features
may occur side by side in the same species. Palisade-tissue has not been' observed
in the ochrQe. Stomata occur, but are rare.
3. STRUCTURE OF THE AXIS. The following statements regarding the
structure of the wood are based on the investigation of woody species of the
genera. Eriogonum, Chorizanthe, Calligonum, Muehlenbeckia, Coccolaba, Tri-
plaris and Ruprechtia. The arrangement of the vessels and the size of their
lumina vary (maximum diameter = -024-.<>7 mm.). The perforations are
simple; the wall of the vessel bears bordered pits, where it is in contact with
parenchyma •of the medullary rays. Spiral thickening of the vessel·wall
occurs in Eriogonum lasciculatum, Benth., Chorizanthe paniculata, Benth. and
Triplaris Pachau, Mart. The medullary rays are generally from one to three
cells in breadth 2, ·arid according to Houlbert they are specially numerous;
in Chorizanthe paniculata distinct medullary ray· tissue is not present. The
wood-prosenchyma may be described as having simple pits; it is septate
by means of thin transverse \Yalls in species of Muehlenbeckia, Coccoloba,
Triplaris and Ruprechtia. Chambered parenchyma containing crystals is
found in Coccoloba.
The structure of the cortex has hitherto been little investigated. In

1 Walliczek's statement (Pringsheim Jahrh., Bd. xxv, 1893, p. 313) that raphi.des occur in the
tissue of the petiole. of R luum is incorrect. The petiole only contains large cl1litered crystals.
2 The herbaceous species are divergent in this respect (see Herbst, loco cit.).
 POLYGONACEAJ:.

many cases, and especially in the herbaceous species, the primary cortex con-
tains strongly developed, subepidermai strands of collenchyma, which project
in the form of ribs. The collenchyma is occasionally replaced by subepidermal
strands of sclerenchymatous fibres, which have been mentioned above in
the section dealing with the structure of the leaf; in other cases (A ntigonon
leptopus, Hook. et Arn., certain species of Rumex, such as R. Acetosa, &c.)
small vascular strands lie in the comers of the stem, and together with the larger
bundles corresponding to the furrows constitute the normal vascular ring.
An endodermis is sometimes developed in the herbaceous species. The peri-
cycle frequently exhibits sc1erenchymatous differentiation, either in the form
of a continuous ring (Antigonon leptopus, Polygonum pro parte), or of isolated
bundles of hard bast (Polygonum, pro parte). The development of cork takes
pJace subepiderrnally (Polygonum), or in the peri cycle (Atraphaxis spinosa~
L., according to Avetta).
Anomalous structure of the axis is found in some members of the Order.
Under this heading we may first mention the somewhat irregular formation
of the xylem-mass brought about during growth in thickness in Atraphaxis
spinosa, L. (xyl~m-mass irregularly lobed in the stem and root), according
to AVetta, and in Coccoloba striata, Benth. and C. ochreolata, Wedd. (flattened
or angular stems), according ·to H. Schenck. A true anomaly is described
by Avetta in the stem and root of
Antigonon leptopus, Hook. et Arn. In
transverse section the five-angled stem
of this species shows a 5-rayed pith;
the latter is surrounded by a vascular
rjng composed of .five larger and five
smaller· bundles alternating with one
another, the smaller strands lying ih
the corners; outside the bundles there
FIG. 161. Portion of a transverse section of the stem . • l' . f I h
or R"~.x cr;,rpUS, L.-Aftet' Herail. IS a pencyc lC qng 0 sc erenc yma.
In the parenchymatous portion of the
pericycle a secondary meristem produces five secondary vascular bundles, which
round off the outline of the stem. Another anomaly, first discovered by Sanio
in Rumex crispus, consists in the presence of medullary vascular bundles in
certain species of Rumex and Rheum 1. These hun dies are generally collateral,
and show inverse orientation of wood and bast, but in a few species (Rumex
cordifolius, Horn., R. domesticus, Hartm., and R. orientalis, Bernh., according to
Russow, Bergendal·and Mobius) they are concentric with central phloem.
The structure in question has been examined most carefully in Rumex cfisPl'S
(Fig.· 16 I) by Herail. The appearance of a transverse section through the rnature
stem of this plant is very striking. A common sc1erenchymatous sheath surrounds
each outer (normal) vascular bundle. together wlth the inverselY orientated strand
situated on its inner side. At first the peripheral bundles alone 3.re present, each of
them being surrounded by a sclerenchymatous sheath at an early stage; subse-
quently groups of soft bast arise at the margin of the pith, and these soon become
enveloped by sc1erenchyma; at a still later stage cambium appears at the outer limit
of each group of soft bast. and gives rise to the xylem of the medullary vascula.r
bundle externally, whilst.it adds to the soft bast internally; finally a small island of
sclerenchyma1;ous fi~res makes its appearance in the rmddle of the phloem. The
ste~ of Rumex cordI/a/ius has a still more complicated structure (Mobius). In this
speC1t'~s, in addition to other peculiarities (such as concentric vascular strands, &c.),
we find not only double vascular bundles, as in R. crispus, but even triple bundles.

I These are: Rumex confirtus, Willd., R. co'rdifolius, Hom., R. (ris/us, L., R. aonzts#Cfts,
Hartm., R. Hydro/apathll1n, Hud!i., R. I017g£fo/ius, R. ma:dmus, R. orienlalis, R. Patientia, R.
Imdu/qtus and Rheum .Gibes (Petersen, Rerail, Mobius).
 POL YGONA CEAE

owing to the appearance of a third bundle with inwardly directed xylem on the
inner side of the inversely orientated medullary vascular strand.
Having described the occurrence of medullary vascular bundles, we may
next consider the presence of intraxylary soft bast in Emex; the peculiar
streaky structure of Radix Rhei; the appearance of bundles of soft bast,
arising independently in the interfascicular tissue of the phloem, in numerous
species of Polygont{'m and F agopyrum; and the occurrence of cortical vascular
bundles in Calligomem comosum, L'Herit.
Intraxylary soft bast has been demonstrated by Petersen in Emex sp£nosa and
Centropod£um; it only occurs epposite a certain number of the vascular bundles;
according to Mobius, It appears also to be present in species of Rheum and Polygonum.
The streaky structure of Radix Rhei has been examined in detail by Schmitz and
Dutailly. In the tuberous branches of the rhizome of the officinal rhubarb-plant
(Rheum offic£nale, Baill.) the collateral leaf-trace strands form a normal vascular ring
enclosing a pith, which is continually undergoing increase in size. In a young branch
of the rhizome the pith is traversed by a complete network of anastomosing strands

FIG. 162. Trans\'el"le .ection through the axis of Calltjronum COmtMum, L'Herit.-Original.

of soft bast, which are arranged in transverse zones, followingJ closely upon
one another and corresponding to the nodes; these' strands unite the leaf-
traces, and are also connected with one another by bundles, which traverse the
internodes either in a vertical or oblique direction, and mostly run in the neighbour-
hood of the xylem-ring;-, Around each of these strands of soft bast a cambial ring.
developed at an early stage, produces rays of soft bast on its inner, and rays of wood
with abundant parenchyma on its outer side. whilst between the rays of wood and
bast it gives rise to medullary ray-tissue, which becomes filled with red colouring-
matter. Thus the radiate rings, or streaks, are medullary vascular bundles with
peripheral xylem. Besides occurring in Rheum ofticinale. they are found in R. Emodi,
R. Rhaponticum and R. palmaium. but they are not present In R. f'ugosum and other
species. The groups of soft bast found in species of Polygonum and FagoP'Yf'um have
been examined in detail by E. Schmidt j they arise between the phloem-groups of
the stem, and become connected with the vascular bundles at the nodes; with regard
to their relation to growth in thickness see Schmidt, loco cit. The cortical vascular
bundles of Calligonum comosum. L'Herit. (a desert-plant with advanced reduction
of the foliar organs). and of other species, are associated with a series of other notc-
SOLI:ItI:DI:R X X
 POL YGONACEAE
worthy structural features, such as the. differentiation Qf paliSade-tissue in the
p~ cortex, the. development of a characteristic sheath' of collecting cells below
the palisade layer (similar to that in the Salsoleae), the development of aqueous tissue
in tile cortex, and so on (for details see Fig. 162).
From the xerophildUs structure of Calligonum we may pass to the anatomy of
the phyiloclades of Polyg<mum platydadum, F. v. Miill. According to Pick, the cortex
contains two or three layers of palisade-tissue, limited internally by a zone of sc1eren-
chymatous fibres. The sclerenchy:matous tissue bears prominences reaching as far as
the epidermis,. and thus has the form of a wreath, !n which the elongated arcs are
concave towards the exterior and enclose,the a5simi~tory tissue, whilst the vascular
bundles lie in the small internal concavities of the wreath.
Literature: Salljo, in Bot. ~it. 1865, P. J ~9--HaJlstein, Harz- u. Schleimabs:., Bot. ZeiL J868.
p.699 et seq. and Tab. xi.-8chmitz, Masem von Radix Rhei, Bot. Zeit. 1875, pp. 260 and 276 et
seq.-Rindow3ky, Ristol. d. Gatt. Calligon~m, Mitt. Univ. Kiew 1875, :14 pp. and I Tab., Russian;
abstr. by Batalin in Just I875,PP. 394-6.-Moller, Holzanat., Denkschr. Wien. Akad. 18 76. pp. 34-5
and 330.-De Bary, Vergl. Anat., t8n, especially p. 6o".-Areschoug, BIad. anat., MinIlesskr. Lund
1878, pp. 67 and Il5 et seq.-)j:. Schmidt, Anat. veget.Org. v. Po'!grmum u. Fagobrum, Diss.,.
Bonn, 1879,38 pp.-Dutailly, Elements nouv. dans les tiges etc., These, Paris, 1879, p. 7" et seq.
and pl viii_C. de Candolle, Anat. compo d. feuilles, Mem. Soc. phys. et d'bist. nat. de Geneve,
t. xxvi, :I, 1879, p. +ll.-Pick, Armlaub. Pit, Diss., Bonn, 1879, p. 9.-Bokomy, Durchs. P., Flora
188:1, p. 371 and sep. copy, p. 25.-Petenen, in Engler, Bot. Jahrh., Bd. iii, 1882, pp. 377-9.-
Jobow, in Pringsheim Jahrb.• Ed. xv, 188 .., p. 309.-Volkens, Standort etc., Jahrb. Berliner
Gart. 1884, pp. 6 and 34 et seq.-HeraiJ, Tige des Dicotyl., Ann. se. nat., ser. 7, t. ii, 1885. pp.
:a83-7.-So1ereder, Holzstr., 1885, pp. nO-I.-Morini;: Nett. estranuz., Mem•. Accad. Bologna 1886,
p. 360 and tav. iv-v.-Plitt, Blattstiel, Diss., Marburg, 1886, p. 4o.-Grevillius, Stipelscheide etc.,
Bot. Centtalbl. 1887, ii, p. "54 et seq.-Lohrer, Wurzel, Wigand, Bot. Hefte, ii, 188 7, pp. 32-3.-
Mobius, KODzentr. Gefassb., Ber. deutsch. bot. Gesellsch. t887, pp. 16-18 and Tab. ii.-Petit,
Petiole, M6n. Soc. sc. phys. et nat. de Bordeaux, set. 3, t. iii, 1887, pp. 233-6 and pl. ii.-Volkens;
Aegypt.·arab. Waste, 1887, pp. 142-3 and Tab. ix.-Avetta, SuI fnsto e salla radice dell' Atrapkaxis
rpiwfa, Ann. del R. 1st. bot. di Roma, vol. iii. FIUiC. 2, J888, pp. 141-7 and Tab. xvi; abstr. in Jast
1888, i, p. 130 and Anat. ed iatol. della radice e del fusto dell' .:Antt."pnrJn leplojlus, loco cit., pp.
148-56 and Tab. xvii-xviii; abstr. in Jast, loe. cit., p. 728.-Delpino, in Mem. Accad. Bologna 1888,
p. 6:17.-Grevillius, Baa d. Stammes bei Polyg. aviculare, Bot. Centralbl. 1888, iv, p. 316 et seq.-
O. Schultz, Vergl. Anat. d. Ne't!eublattgeb., Flora. 1888, pp. IlI-4 and Tab. i.-Petit, Petiole. Actes
Soc. Unn. de Bordeaux, t. 43, 1889, p. 16 and pl. i.-Ross, Tessuto assim. e perid., Nnov. Giom.
bot. ltal., vol. xxi, l8S? p~. 229 and :I,36.-Dammer, in ~atiirl. Pflanzenfam., iii. Tell, Abt. la" 1891,
pp. 2-4.-A. Wagner, m SltZ••Ber. WIener Akad., Bd. CJ, I. Abt., I89~, p. 5IS.-Houlbert, BoIS sec.
dans les Apetales, These, Paris, 1893, pp. 91-7.-Koning!!;berger, Anat. Eigentumlichk. einig.
Rluum·A., Bot. Zeit. 1893, Abt. i, pp. 85-8 and Tab. iv.-H. Schenck, Anat. d. LiaDen, 1893, pp.
47-8 and Tab. i.-Herbst, Markstr., Bot. Centralbl. 1894J i, p. 294 et seq.-SchUling, Schleimbild.
d. Wasserpfl., Flora 1894. pp. 3:J8-31.-Russell, Climat med., Ann. sc. nat., se~. 8, t. i, 189S,
pp. 317 et seq. and 349.-Diels, Neuseel., Engler, Jahrb., Ed. xxii, J89<), p. l49.--Parmentier, in
Ann. sc. nat., ser. 8, t. ii, 1896. p. 3J.-WittIin, Oxalattaschen, Bot, Centralbl. 1896, iii.-Gillot
et Panntntier, .Rumex paJustris, BnlJ. Soc. bot. de France 1897, p. 3:1'; et .seq.-Wanningl Halofyt
Stud., K. Danske Vid. Selsk. Skr. 1897, p. 203.-[Montemartini, Fusto del Polfg. Sitboldii, Malpigbia.
1898, pp. 78-80 and tav. iii.-H.temmerle, Polyg. cuspidatum, Diss., Gottingen, 1898.]

PODOSTEMACEAE.
In external appearance the Podostemaceae remind one of Lichens, thalloid
Liverworts, Jungermanniaceae and Algae rather than of ,Phanerogams.- In
accordance with the object of this book, their anatomical featUres will
be quite shortly considered. In some cases, we find resemblances to Algae
in the internal structure as well as in the external characters. The
shoots are composed of fairly uniform tissue, which is very frequently
collenchymatous, especially in the neighbourhood of the vascular bundles.
Stomata are entirely absent. The epidermal cells sometimes contain chIoro·
phyll. The vascular bundles are never strongly developed; the xylem con·
tains a few annular and spiral tracheids, but sometimes even these are absent
(root of Mniopsis Weddelliana, Tu!.), or they become abortive at an early stage
and replaced by an intercellular space (stem of T"isticha hypnoides, Spreng.).
In the phloem the sieve-tubes are not always distinct, but in many species
 PODOSTEMACEAE

they are readily seen and together with their companion cells form cha.rac..
teristic groups of cells, An endodermis is not developed, ~ tJ:ansverse s~on
the stem shows ,more or less distinct dorsiventrality, depending on the position
of the. vascular bundles and the differentiation of the ground-tissue. As a rule
there are only a few isolated vascular bundles in the stem; they are placed. side
by side, so that there is no vascular ring. .In Hydrostachys imbricaja, Juss., on
the other hand, a ring of isolated vascular'bundles occurs in the axis of the
inflorescence, while additionai medUllary and cortical strands are also present.
In the structure of the root the dorsiventrality is still more distinctly ~k~
than in the axis. The vascular system is nearer to the ventral than tt> the
dorsal side of the root, and consists of two vascular bundles, which are placed
side by side, and fused together; the two xylem-groups
t;;! ~ ~

f
are situated ventrally, but may be absent.
Trichomes have on1i been met with on the leaf in
a few members of the Order; they occur in the form
of short, tubular protrusions of the epidermal cells, and
~:
.
.•
t
have a secretory function. Clustered crystals of oxalate ~ I
of lime have also only ~n obserVed in a few case~ ~:n9
(Hydrostachys and Lawia). Intercellular spaces, re- ~ ."
sembling resin..canals (without any secretion visible in
alcohol material), occur in the root of Weddellina
squamulosa, Tul. On the other hand peculiar siliceous
>

concretions are very widely distributed; they were

already observed by Tulasne, but were only correctly
interpreted for the first time (almost simultaneously)
by Cario and Warming,/
The silica-bodies are found in many s~es, and often
occur in great abundance; they are pnnClpally to be met
with in the peripheral partS of the root, stem and leaf,
and especially in the epidermis (even in the trichomes).
When the silica-bodies are abundant, the organs above
referred to possess. as it were, a coat of armour,
which is re~arded as a. protective arrangement aga.in.st
mechanical ~juriesJ the attacks of animals, and desiccation.
Thus, according to Warming, the silica-bodies are so
abundant in Podostemon Galvonis that the plant has the
same appearance in the dry state as when fresh; on the
other hand. plants of P. subulatus, a species which is a~
devoid of silica, become greatly shrunk in drying. In those FiG. 163. Silica.bodiea of
plants, however~ which have a complete subepidermal sUi- Trl.rJkA" A~, Spren,.-
ceous armour, the latter does not form an absolutely closed After Carlo.
sheath. but exhibits various' passage-ways: The appear-
ance of the silica-bodies varies in the same species. In most cases they fill the entire
lumen of the cell. Sometimes they are absolutely homogeneous and vitreous, but
frequen~y it is only the. outer _p~ of the silica-body that has ~ ".omogeneous
structure, the intenor appearing dull owing to the presence of a quantity of small
gas-vacuofes. The shape of the silica-bodies is also very varied, even:jn the same
species, as is shown sufficiently clearly by some of Cario's figures reproduced here
(Fig. 163). The surface of the silica-bodies is either smooth, or shows spi...al or
annular ridges, or is corroded in a very peculiar manner so as to exhibit a varying
number of depressions or excavations. Wanning and Kohl regard the silica-bOdies
in question as structures formed in the celJ-cavity, and hence homologous with the
well-known stegmata of Palms and other Monocotyledons; the same observers, attri-
bute their varied shape to the influence of other contents, component parts at .the
ceU, such as the nucleus, chlorophyll-bodies and starch-grains. According to Straa-
burger's investigations on Mniof>s-as Weddelliana. we are not in this case dealing 'With
a deposition of silica in the Living protoplasm, but rather with a progressive trans-
formation of the superficial portion of the protoplasm from without inwards.
XX2
 PODOSTEMACEAE

Literature: TulAsne, P. Manoge., Arch. Mus. d'hist. nat., t. vi, 18SZ, pp. 4 and 10 et seq.-
Cario, T1"'isticha nyjncides, Bot. Zeit. 1881, p. 25 et seq. and Tab. i.-Warming, Riselsyredannelser
hos P., Vidensk. Meddelels. naturh. For. i Kjobnhavn, 1881, pp. 89-92, with a French resUnte.-
Strasburger, Zellhiiute, ~ena, 1882, pp. ~34-5.~-WarmingJ Familien P., Vidensk. Selsk. Skr., 4- Abh .•
Danish with a French resume I: 1881) 34 pp.and 6 Tab.; II: 1882,88 pp.and 9 Tab.; III: 1888,
72 pp. and 12 Tab.; IV: 189[, 47 pp.; V : I899-Kohl, Kalks. u. Kiesels., 1889, pp. 24-9-67 and
Tab. v.-Warming, in Nattirl. Pf1anzenfam., iii. Teil, Abt. :z a, 1890, pp. 3 and 8-9 a.nd Hydroslachys,
Bull. Acad. roy. Danaise des sc. et Jettres 1891, sep. copy, 7 pp.-Wiichter, lVeddel/z'na sqUQfflu/osa,
Beitr. z. K. einig. W~rpfl., Diss., Rostock, .897, pp. 18-33, sep. copy from Flora 1897.

NEPENTHACEAE.
This Order comprises the single genus Nepenthes, widely known on acc()unt
of its peculiar ascidiform leaves or pitchers, which are adapted for the,capture
of animals, and· actually digest animal substances. The genus has been re-
peatedly investigated owing to its interesting foliar organs, and its anatomy
is consequently well known. , I

The leaf of Nepenthes consists of three parts,; there is a lamina, which

is produced into a cylindrical portion usually serving as a tendril, and ter-
minates in a pitcher, provided with a lid.
a
. . The lamina has the type of structure usual for 1eaf, as my own observa-
tions showed. The mesophyll is in some cases (N. gracilis, Korth. and N.
Boschiana, Korth.) distinctly bifacial. being differentiated into palisade and
spongy tissue, of which the former consists of a few layers of short, broad
cells; in other $pecies, such as N. ampullaria, Jack and N. albomarginata,
Lobb., the palisade-tissue is scarcely di~tinguishable, The lateral margins
of the epidermal cells are straight in most of the species investigated; in
N. ampullaria only they are curved on the lower side of the lamina. In the
epidermis of the lamina stomata occur on the lower side only; they afe also
present on both surfaces of the lid, and on the outer side of the pitcher itself.
The stomata are surrounded by several epidermal cells; those of the lamina
are arranged approximately parallel to oIle another, and with their pores
parallel to the mldrib. The integumental tissue of the lamina is supplemented
by a water-storing hypoderm. The latter is developed more especially beneath
the upper epidermis, where it usually forms a continuous layer ~f cells with
large lumina, considerably exceeding the epidermal cells in diameter (N.
gracilis, N. ampulla ria and N. albomarginata); in N. gracilis the cells are
elongated transversely to the median vein in surface-view, while in other tases
they are approximately isodiametric. In N. Boschiana there is a lay~r of
small cells lying immediately beneath the epidermis, and followed by the large-
celled hypoderm, On the lower side of the lamina the lowest layer or la.yers
of cells of the mesophyll also assume the character of a hypoderm. The
vascular bund1~ of the veins are provided with mechanical tissue. The
course of the veins is peculiar. There are, firstly, two or more lateral
veins, running parallel to the principal vein, while additiollal systems
of delicate, much branched, and anastomosing veins arise at short intervals
from the midrib and run near the upper side of the leaf. The meso-
phyll of the lamina, and also the ground-tissue of the lid, pitcher and axis,
are specially distinguished by the presence of spiral cells (Fig. 164, A-B),
serving for the ~torage of water. These cells are mostly tubular, closed on
all sides, and devoid of contents; they are '09-3'33 mm. in length and
-°45-13 mm. in breadth; their walls are stiffened by a spiral ban~ composed
of three or more partial bands. Both the wall and the spiral band consist of
cellulose. ..
Usually the inner wall of the pitcher may be divided into two sharply
,separated zones, the slippery and the glandular zone, which one can distinguish
 NEPENTHACEAE
on examining the internal surface with the naked eye; sQmetimes they are
even rec~gnizable from the outside. The slippery zone occupies the upper,
the glandular zone the lower portion of the pitcher. The relative areas occu·
pied by these two zones on the inner wall of the pitcher vary. In some
cases the line of demarcation between them is only a few millimetres
beneath the mouth of the pitcher (N. villosa, Hook. f., N. Edwardsiana, Low,
according to Wunschmann), but more commonly it lies in the middle of the
pitcher. The slippery zone is rarely completely absent (N. bicalcarata, Hook.,
N. madagascariensis, Poir., N. Raiah, Hook. f., and in some cases in N. ampul·
laria, Jack and N. RatJiesiana, Jack, according to Wunschmann). This
zone is specially distinguished by the absence of glands. Its surface is
not wetted by water, as it is covered with a granular layer of wax, which
sometimes has a blueish or reddish colour. In the epidermis of this layer,
besides cells of the ordinary type there are others which are semilunar and
resemble guard-cells; they project somewhat like papillae, and have their
concave sides directed towards the base of the pitcher (Fig. 164, Cl)._ The
glandular zone bears numerous glands of characteristic structure (digestive·
glands). These (Fig. 164, D-F) are wart-like bodies and may be seen even with
a lens to lie in pockets, which are open towards the base of the pitcher; the
pockets are produced by elongation of the row of epidermal cells bordering on
the upper margiIl of the gland so as to form a roof! with a sharp edge,
covering about one half, or sometimes -the whole of the gland, or in other
cases only a small portion of it. The glands themselves are placentiform
masses, consisting of either two or three horizontallayel's oi cells, of which the
uppermost is differentiated like palisade-tissue, and appears as a finely polygonal
network in surface-view; the gland is separated from the internal tissue of
the wall of the pitcher by one or more layers of cells, whkh when multi-
seriate are arranged like cork-cells, and have some of their walls suberized
(Fig. I64. F). Another important fact is that tracheal strands, springing
from branches of the veins, invariably terminate beneath the glands.
Glands of a similar structure to those described above, though sometimes
very different in appearance, are also found on other parts of the veg~tative
~organs, viz. on the stem, the petiole, the midrib of the leaf, the lower and
in rare Cc,;iies also the upper side of the lamina, as well ~ on the tendril,
the outer side of the pitcher, the inner and outer surface di the lid, and the
margin of the collar of the pitcher. These glands, however, excrete honey,
and have the function of attracting insects. Their structure has been examined
especially by Macfarlane, and in all ca:ses agrees ~h that of the digestive-
'glands in the following respects: the surface of the glands is formed by two
or three layers of secretory cells, the uppermost of which is composed of palisade-
like cells; the secretory portion is limited internally by one or more layers
of cells, some of whic.h are suberized; tracheal terminations are invariably
found beneath the gland. The glands, which occur on the inner side of the
lid in vari~ble'numbers, (numerous in N. Boschiana, few in N. albomarginata),
agree closely with the digestive glands in their flattened placentifonn shape,
and in being composed of three horizontal layers of cells; they lie in small
pits, which are overal'ched on all sides Gr 'on one side only (the upper) by a
projecting flap of tissue formed by the adjacent epiderI?:1al cells. The marginal
ftlands of the collar are likewise convex muiticellular structures, but they have
an elongated form, and are sunk in pits of a corresponding shape and often of
considerable depth; in these glands the layers of glandular cells cover the

1 These cells have nothing to do with the deposition ot wax. Macfarlue found transitions
between them and normal stomata, and eupposes that they take part in the excretion of water in the
linterior of the pitcher.
 NEPENTHACEAE
entire convex surface, apart from the insertion· of the gland. The glands,
described by Macfarlane as occurring on the other portions of the ascidium enume-
rated above and on the stem in various species, are essentially different. In

FIG. 164. A-s. Spiral ceUs "-om the leal of

N. al/xJma"'I!",.,ala, Webb. C, Epidermis or the
slippery zone. nAF, Digestive glands 0( N. ullJo..
marpNata : D, in surCa<:e-yiew; It-F in a IleCtiOI1
cut paralJd to tbelongit~uJjlJaJa:ds;Jtbe pitchn.
G, HOlley.gland from the stem of N_ F"~JIa_
phoYi~.-A., B, D-p Original, c, after GOebel,
G, after Macfarlane.

them the layers of glandular cells form the wall of a depression In the surface
of the organ, the shape of the depression varying in different cases. The
depressions are either shallow and widely open (leaf-glands of N. hybrida):.
o~ have the fonn of deep and narrow canals (stemAglands of N. Phyllampho,a,
FIg. I64, G), or of larger cavities, provided with numerous recesses and opening
 NEPENT1IACEAE

to the exterior by means of a narrow ostiole (petiolar glands of N. bica~cafata ;

glands occurring on the inner side of the lid of the pitcher in N. laevis 1,
N. Low#, and N. Pervillei).
In the axis (N. Phyllamph(Jra, Willd., N. Boschiana, Korth.) the normal
vascular ring is supplemented by numerous cortical vascular bundles, which
form a special annular zone in the primary cortex. According to Zacharias

I
E
c

F
FIG. 165' Hairy cGverill.\t Q£ the Nepenthaceae: A-B, Bxternal
glands, and D-F Clothing baits of N. sllxmuzrriMla, Lobb. C,
Trichome from the floral region of N. dutt11(J/()ritJ., L.-Original.

(N. Phyllamphora) 10-12 larger, and a con-

siderable number of smaller leaf-trace bundles
pass into the stem from each leaf; the lar~e
bundles first run for a varying distance In
t:abrimary cortex, and then enter the vascular ring, whilst the
s er bundles remain in the primary cortex. A transverse
section of the axis shows the following differentiation. The
pith, which contains spiral cells, elongated in the vertical direc-
tion, is surrounded by a medullary sheath of elongated
prosenchymatous or parenchymatous elements, provided with slit-like
pits and subdivided by delicate transverse walls. Outside the medullary
sheath lies the ring of vascular bundles. The pericycle contains hard bast
accompanied by numerous spiral cells, which are elongated in the vertical
direction; adjoining these elements is the endodermis. primary cortex The
1 The glands described above are not present in .N. graci/is, with which No 1aiviJ is frequently
confounded.
 680 NEPENTHACEAE
commences with a zone of large parenchymatous cells, amongst which short
spiral elements occur; the parenchymatous cells themselves sometimes (N.
Boschiana) possess spiral thickening, or spirally arranged slit-like pits. Outside
the parenchymatous zone is a ring of mechanical tissue exhibiting the same
structure as the medullary sheath and having the cortical vascular bundles
embedded in it; external to this is a small-celled tissue containing chlorophyll
and including spiral cells. The dev~lopment of cork takes place imm~diately
beneath the endodermis. In the structure of the wood we may mention that
the vessels attain a diameter of '07-'09 mm., and have simple perforations;
the medullary rays are narrOW and the wood-prosenchyma is provided with
typical bordered pits.
Oxalate of lime occurs in the form of clustered crystals. In addition to
these, sphaerocrystalline masses, the nature of which is not known, have been
observed in alcohol-material (Zacharias). Internal secretory receptacles are
not present.
The hairy covering, apart from the digestive and honey-glands described
above, is constituted by pelt ate glandular hairs and by clothing hairs, as my
own investigations proved. The former consist of (a) a basal cell; (b) a rather
long stalk with fairly thick walls, and divided by horizontal septa, the upper-
most tier being bicellular 'owing to the presence <>f a vertical wall; and (c) a
deciduous peltate head, which sometimes (N. gracih's, according to Haberlandt)
has a four-rayed form, and is divided into a number of cells (four or
more) by variously orientated vertical walls (Fig. 165, A-B). The clothing
hairs are of the fOllowing types: I. simple uniseriate trichomes, which are occa-
sionally (.in the floral region of N. destillatoria, Fig. 165, C) characterized by oblique
transverse walls and lateral protrusion of certain cells of the hair; 2. trichomes
of more complicated structure appearing to the naked eye like tufted Ot stellate
hairs (Fig. I65, D-E) or bristles (Fig. 165, F); some of these may be regarded
as derived from uniseriate hairs of sympodial structure, whilst others are com-
binations of similar trichomes with stellate hairs; and 3. lastly, stellate hairs
with a unic:ellular stalk and several ray-cells.
-Yteratllre;- ~abschD14nn) Neptnlne[) Diss., Berlin, rBtJ, 46 pp. (here and in DC. !'rodr, xvii,
p. 9r the older anatomical literatute)._':_Zacharias, Stamm d. Gatt . .Nepenthes, Diss., Strassbur~.
1817, ,~ pp. and 3 Tab.-Kny and Zimmermann, Spikularzellen von .A"ejenl/zes, Ber. ,--leutsch. bot.
Gcsellsch. 1885. pp. H3-8.-So1ereder. Hoizstt'., 1885, PP' 2l1-2.-Heckel and Chareyre, Ascidies
etc., Compt. rend., c. ci a, 1885. p. sSr.-Goehel, malogo Schild., ii. Teil, Lief. (, r89c, pp. cor-co
and Tab. xxii.-Wnnschmann, in Natiirl. Pfianzenfam., iii. Teil, Abt. 2, 1891, pp. 25G-7.-Mac.
{arlane, Pitchered plants, Ann. of Bot., vol. "tii. 1893, especiaUy pp. 420-40 and pI. xix-:ui.-
H. Schenck, Anat. d. Lianen, I893~ p. 7l.-Haberlandt, Trop. Lanbbl. ii, Sitz.~Ber. Wiener Akad.,
Ed. civ, Abt. r, (8951 p. 97 and Tab. iv.

CYTINACEAE.
This Order, as is weU known, comprises parasitic herbaceous plants, which
are for the most part leafless, and in other respects also show considerable
reduction of the vegetative organs.
This reduction is most pronounced in the tribe Rafflesieae, where the vege-
tative organs have the form of a thallus, embedded in the tissues of the host-plant;
according to Solms·Laubach) this structure is mycelial and 'devoid of vascular
tissue in the simplest cases (RafJIesia, Brugmans~'a, Pllostyles /faussknechtii,
Boiss.), whilst vascular bundles oCCur in the more massive thallus of Pilostyles
aetkiopica, Welw. and Cy#nus Hypocys#s, L. Unlike the RafHesieae the
members of the second tribe, the Hydnoreae, consisting of the two genera
Hydnora and Prosopan che, :possess I rhizoid-shoots.' The structure of these
rhizoids has been examined In detail by Schimper in Prosopanche Burmeisteri,
 CYTINACEAE 681
De Bary, Hydnora a/ricana, Thunb., and Hydnora abyssinica, A. Br.; in the
first two species the rhizoids are angular in cross-section, in the last species
they are cylindrical. In all three species the periphery of 'the transverse
section shows a thick layer of cork-cells. These surround a pare:achymatoUs
cylinder containing the vascular bundles; the cells of the parenchyma are
filled with brown, tanniniferous, gelatinous contents, which sometimes also
include starch and tetragonal crystals of oxalate of lime. The tissue
forming the innermost portion of the rhizoids consists of a strand of elongated
cells (pith), which in Prosopanche are extremely long and fibrous, while in the
remaining genera they are somewhat elongated and prismatic. The arrange-
ment o~ the vascular bundles differs in the three species investigated. In
Hydnora atricana a transverse section shows the usual ring of normally orientated
vascul..- bundles (xylem inwards, phloem outwards), the ring having five
projections corresponding to the usually pentangular shape of the rhizoid;
a few vascular bundles are also found scattered in the ground-tissue external
to the ring. In Prosopanche Burmeisteri, in which the rhizoids are likewise
4-5 angled.!. the distribution of the va!:icular bundles in the parenchymatous
ground-tissue is somewhat modi.4ied. In this plant the vascular ring is broken
up into (a) a central zone of four or five normally orientated vascular bundles,
which are placed singly opposite the concave lateral surfaces, and (b) four
or five peripheral groups of bundles; each of these groups lies in one of the
corners of the stem and is composed of two radial rows of bundles; the xylem-
groups in the bundles of one row are directed towards those of the other.
In Hydno,a abyssinica, finally, a transverse section of the rhizoid shows several
rings of normally orientated vascular bundles. Regarding the structure of
the vascular bundles, I may mention that distinct sieve-tubes and simple
perforations in the vessels have been observed in Prosopanche, and that the
bundles sometimes show slight growth in thickness. We may also note the
occurrence of peculiar secretory receptacles '"in the rhizoi4-shoots of ProsQ-
panche Burmeisteri; Schimper terms them receptacles of gelatinous substance,
and De Bary found them also in the pe'duncle of the same plant~ They fo.rm
cylindrical strands of large, loosely oonneeted alUS ,fJith ~l!.tiri{,)Q..C:; blntSlts,
the strands sometimes attaining a diameter.of 2 mm.; in 11lan,".f.ct.ses the
more centr~ cells sooner or later become disorganized, and replaced by an iJfteJ.'
cellular canal containing gelatine.' The gelatine-receptacles' are restricted
t I

to the r3.dii connecting the angles of the'stem (in transverse section) with the
centre; on each of these radii there are either two or three receptacles, which
decrease in size centrifugally.
Stomata appear not to be present in any member of the Order, not even
on the floral organs. Glandular hairs are figured by Chatin (Taf. xcii, Fig. 3)
on the bracts of Cytinus Hypocystis; a glandular head, composed of three
or four c~lIs separated by vertical walls, is se'a~ed on a conical multicellular
pedestal.
Literature:-Chatin-;-AnaL comp., PI. parasit., pI. xc bis-xcii bis and cii-cvii (without text).-
De Bary, Prosojancne, Abh. naturf. Gesellsch. Halle, Bd. x, 1868, pp. 241-69 and ~ Tab.-Solms-
Laubacll, Pilostyles Haussknechtz"i, Bot. Zeit. r874, pp. 49 and 65 et seq. and Tab. i and Iiaut. d.
,Loranth. u. Thallus der Rafflesiac. etc., Abh. naturl. Gesellsch. Halle, Bd. xiii. 1874, pp. 25~7
'and Tab. xxv-xxvi.-Schimper, Prosopancne, Abh. naturf. Gesellsch. Hal~: Bd. _n, 1882, pp. 21-47
and Tab. ii-iii.-Solms-Laubach, in Natiirl. Pftanzenfam., iii. Teil, Abt. I, 1889, pp. 275 and 28+-
Peirce. Haustoria, Ann. of Hot .• vol. "ii, 1893, p. 318 et seq. and pI. xiv-xv.
 682

ARISTOLOCHIACEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. This Order is characterized
by the following features: (a) the presence of secretory cells with oily
contents (Fig. 166, A-C), probably occurring in all the species, and sometimes
giving rise to transparent dots in the leaf, where they show a tendency
to be restricted to the epidermal tissue; (b) the lack of a special type of
stoma; and (c) the absence of glandular hairs. .In the woody species
other characters may be added, viz. the broad primary medullary rays,
the simple perforations of the vessels, the bordered pitting of the wood-prosen-
chyma and the absence of secondary hard ba.st. The development of cork,
as far as it is known, takes place superficially; in the woody species the peri-
cycle contains a continuous, or interrupted, composite ring of sclerenchyrna. In
addition to the'secretory cells, spherical cells, filled with brown, tanniniferous
contents, are present in certain species of Aristolochia and Holostylis. Oxalate
of lime is generally excreted in the form of small prismatic or clustered
I

crystals, very rarely as ordinary large solitary crystals. Grou~ of cells with
silicified walls (Fig. 166, D-G) are common in Holostylis and the species of
Aristolochia, while cells with siliceous contents (Fig. 166, H - J) are present
in all species of Apama and Thottea. The hairy covering consists of uniseriate
trichomes of varied structure; in species of Aristolochia, H olostylis, Thottea
and Apama these trichomes assume a special form, viz. bracket-hairs (Fig.
166, K), in which the terminal cen is bent like a hook; another special type
is that of the trichomes of Saruma, where the terminal cell encloses
fine crystal-sand composed of oxalate of .lime. ADomaIous structure of the
axis has only been met with in Aristolochia triangularis, Cham. (Fig. 167, B)
(splitting up of the original vascular ring, and secondary formation of b1.mdles
of wood and bast at the inner margin of the resulting segments). For the
purposes of finer .anatomical diagnosis the following characters have special
systematic importance: the details of the distribution, or the absence
(Aristclochia, s~tion Siphisia) of secretory cells in the leaf; the occurrence of
hypodermal tissue in the leaf lspecies Of Aristolochia and Thottea); and the
paJftllose differentiation of the· upper (species of Asarum) or lowe~ (species of
A.ristolochia and Thottea) epidermis.
2. STRUCTURE OF TIlE LEAF. This has been thoroughly investigated 1.
The leaf is generally bifacial, but in some cases the mesophyll consists of homo-
geneous muriform parep.,thyma. Centric leaf-structure with palisade-tissue
developed on both sides is rare (Aristolochia auricularia, Boiss., &c.). The
stomata have no special subsidiary cells, and in the great majority of species
occur only on the lower surface of the leaf. In certain species of Asarum and
Aristclochia and in Holostylis they are met with on both sides of the leaf, whilst
in Apama and Thottea they have been observed on the lower surface only.
It is a noteworthy fact that the cuticle is not striated in any of the Aristolochi·
aceae, but it frequentl y has a granular structure. Gelatinization of the epidermis
of the leaf has not been observed. Development of hypoderm is rare. Thottea
grandiflor(l., Rottb. is distinguished from other species of the same genus by
possessing a hypoderm situated beneath the upper epidermis of the leaf and
composed of a single layer of thick-walled cells with undulated lateral walls ;
hypoderm is also found on the upper side of the leaf in Aristolochia oblongata,
Jacq., and less completely differentiated in A. sericea, Benth. Papillose
development of the epidermis of the leaf has been demonstrated in species of
Asarum,Aristolochia and Thottea. In the species of Asarum in which this feature

1 See Solereder, in EPgler. Bot. Jahrb. I~89, loe, cit.

 ARISTOLOCHIACEAE 683

is found (A. Blumei, Duch., A. canadense, L., A. Thunbergii, A. Br. and A. va,ie-
gatum, A. Br. et Bouch.) the papillae occur on the cells of the upper epidermis;
in the other cases they are developed on the lower epidermal cells (Thcitea
dependens, Klotzsch, T. grandi/lora, Rottb., T. tricornis, Maingay; ATisto-
lochia albida, Duch., A. Clematitis, ·L., A. contorta, Bge., A. cymbifera, Mart.
et Zucc., 4. cynanchifolia, Mart. et Zucc., A. de'Mlis, Sieb. et Zucc., A. klan_s,
Willd., A. nervosa, Duch., A. ringens, Vahl and A. Uhdeana, Duch.). In Aristo-
lochia Tingens and certain other species the formation of papillae is restricted
to the veins, and occurs especially where several of the smaller veins meet;
in A. contorta the papillae are sometimes very long, resembling hairs, and some
of them may even become bicellular owing to the formation of transverse
walls. The vascular system of the smaller veins is invariably embedded;
in many species it is accompanied by varying amounts of sclerenchyma, whilst
in other species the sclerenchyma is entirely wanting.
The secretory cells (Fig. 166}, which in many species give rise to trans-
pareut dots in the leaf, require a more detailed description. They occur
not (ffily in the leaves, but also in all the other organs, such as stern, rhizome,
root, flower, fruit and seed. The secretory cells in the lamina of the leaf
have been especially examined. They are found in the lamina in nearly all
the members of the Order, being absent only in a few very closely related species
of Aristolochia, belonging to the section Siphisia (namely A. Kaempfet"i, Willd.,
A. platani/olia, Duch., A. Serpentaria, L., A. SiPho, L'Herit. and A. tomen-
tosa, Sims.). In these species, however, the secretory cells are met with in
other organs of the plant, so that their general occurrence ·may be regarded
as a specially valuable and Cdnstant character of the Order. The following
statements may -be made regarding the distribution of the secretory cells in
the tissues of the lamina in the various genera of the Aristolochiaceae,
to which I am able to add the new genus Saruma, Olivo on the basis of a
recent investigation. There is a general tendency in this Order to the develop-
ment of secretory cells in the integumental tissue. In the genus Asarum
~hey are invariably found in the epidermis, and in certain species they occur
In the mesophyll as well; the monotypic genus Saruma possesses secfetory cells
in the lower epidermis and in the mesophyll; in Thottea they occur only ~ the
mesophyll ~ in Apama in the mesophyll-in certain species in the epidennis
also; in the genera HolostyUs and Aristolochia exclusively in the integumental
ti4)sue: generally in the epidennis, rarely in the hypodernl on the upper side
of the leaf (A~ oblongata, Jacq.), or as basal cells to the uniseriate hairs (A.
sericea, Benth., Fig. 166, A, A. Griffithii, Hook. f. et.-Th. and A. saccata, Wall.,
in all of which independent secretory cells are wanting). The secretory cells
occurring in the epidermis of the leaf are either present on both sides or only
on the lower side; they are never found in the upper epidermis only; their
varied mode of occurrence may be employed systematically in the case of the
species of Asarum and Aristoloc/eia, and the same applies to the distribution
(r~ferred to above) of -the secretory cells in both mesophyll and epidermis,
or in only- one of these two tissues. The shape of the secretory cells is mostly
spherical or ellipsoidal; branched forms are of rare occurrence (Aristolochia
nervosa, Duch.). The size of these elements varies, their diameter being between
-075 and -012 mm. Their walls are usually suberized. The contents consist
of drops of a yellowish or whitish, or occasionally reddish substance, and include
ethereal oil. They are not of u!liform chemical composition; in A. Lindeniana
Duch. var. plagiophylla, Griseb. the contents of the secretory cells become
coloured indigo-blue by Eau de ] avene, this being due to the conversion of
some substance present in the secretion into indigo. The secretory cells in the
epidermis require special description, since frequently only a small portion of
each of these cells reaches the free surface of the leaf, while they often penetrate
 ARISTOLOCHIACEAE

deeply into the mesophyll (Fig. 166, B); consequently, if one examines a trans·
Verse section of the leaf in a superficial manner, they appear to belong to the

FlO. 166. A., Hair of A yist%du'a SIY,'UIZ, Benth. with A basal secretory ceU. a. Secretory cells.in the lower-
(!l?id~rmiiJ of the Jeaf {If A. OYfJchyur(J, Duch. r.:, Transverse section of leaf of A. h'~CkbJtont4, Gris~!:I. D-P,
SUiciJied gToup of cells from tae leaf f;)f A. lomIHJI1SIZ, Sims. : D, in section: :a, sit:cified group of epider1llal cells;
F, RTO'lP cfsubjacent pa1:aade- c ells in surface-view. G, Silici6~ group of cells in the mesopbyll of A. aCUJJj,,/iIJ,
.D,i;i:b. B-1< Si)jC~11S cells in the leaf of TkJlhIJ de/)lfnt/en.1, Hrotz.5Cb: H, in surface-view; J, in $ectjun. X.
Bracket-hatf of Ads/aloe"''' «mu",/o.sa, Sims.-Original..

mesophyl1, though this is really not the case. The small portion of the epi ..
dermal secretory cell which appears at the surface, occasionally lies at the
 ARISTOLOCHIACEAE 685

bottom of a small pit-like depression in the surface of the Jeaf (especiaIJy in

A. trichostoma, Griseb., Fig. 166, C, and A. spathulata, Duch.). Not uncommonly
the external surface of the secretory cell exhibits a centrally placed dot, which
has been shown to be due to a small, circular, thinner portion of the outer wall
(Fig. 166, B). The secretory cells of the axis occur not only in the epidermis,
but also in the pith, primary cortex and medullary rays. Spherical cells
filled with brown, tanniniferous contents must not be confounded with the secre-
tory cells; they are to be found in the neighbourhood of the veins in the leaves
of "Aristolochia GaleoUii, Duch., A. passi/loraetolia, A. Rich., A. veraguensis,
Duch. and Holostylis reniform£s, Duch.
Groups of cells with silicified walls are very widely distributed in this
Order, having been observed in Hoiostylis and in 94 species of Aristolochia.
The silicification usually involves a group of ceIls belonging to the upper epi-
dermis of the leaf, and the subjacent cells of the palisade"tissue; the con-
tiguous walls of these two layers are silicified, more or less strongly thickened,
and sometimes stratified (Fig. 166, D-F). Silicified groups of cells are of rarer
occurrence (Aristolochia acutitolia, Duch., Fig. 166, G) in the interior of the
leaf; they form spherical or hemispherical groups, in which those portions of
the walls abutting on one another at the centre as well as the radial walls are
thickened and silicified. The silicified groups of cells are generally visible
even with a lens as white or lustrous elevations on the upper surface of the leaf,
and resemble pustules; occasionally they give rise to transparent dots in the
leaf. Silica also occurs under another form in this Order, viz. as plugs
fi1ling the cavities of the cel1s; these were present in all species of the genera
Thottea and A pama, which I examined (Fig. 166, H - J). The siliceous cells 1 of
Tltattea and Apama are found in the leaf for the most part immediately beneath
the upper and lower epidermis, and occasionally in the middle of the mesophy1J
as well; in Thottea grandijiora, Rattb. they also occur in the hypoderm. They
ha ve also been met with in the primary cortex of Thottea grandi!lora, T. tl'icornis,
Maing. and Apama sil-iquosa, Lam.,. either in a subepidermal position or at
a greater depth in the tissue.
Oxalate of lime is excreted chiefly in the form of small prismatic or acicular
crystals, and as clustered crystals ;- large ordinary solitary crystals are very rare
(in the Df'i.ghbourhood of the vascular bundles of the veins in Aristolochia
reticulata, Nutt.). In Asarum, Saruma, Thottea, Holostylis and most of the
species of Apama the small crystals only have been observed. They are
commonly accompanied by clustered crystals of varied size in the species of
Aristolochia, and small clustered crystals are also f~)Und in the palisade-tissue
of Apama corymbosa (Griff.).
Regarding the hairy covering, two features have been sufficiently indicated
_above as important: the absence of glandular hairs and the occurrence of
bracket-hairs. The ordinary form of clothirig hairs in this Order is that
of simple, uniseriate trichomes, which exhibit a number of differences in the
thickness_of their walls and in the number and length of the component cells.
The following· forms-of these hairs require special mention: the narrow whip-
shaped trichomes of certain Aristolochias (A. barbata, Jacq., A. costaricensis,
Duch., &c.); the trichomes found in the species of Apama and distinguished by
having several relatively short basal cells and a few long terminal cells; and
the curious trichomes of Saruma Henryi, Oliv., which are rather thin-walled,
and have an elongated terminal cell with doubly refractive~ finely granular
contents 2. The above-mentioned bracket-hairs generally consist of a fairly
1 In my paper on the anatomy of the Aristolochiaceae these cells were described as secretory sacs.
2 As proved by their chemical behaviour towards acetic, hydrochloric and sulphuric acids, these
contents include fine crystal-sand composed of oxalate of lime, nnd this constitutes the doubly refract-
ing portion.
686 ARISTOLOCHIACEAE
high, unicellular, dome-shaped pedestal, which is seated on the epiderotis,
a short neck-cell, and a terminal cell, bent like a hook, the, tip being in most
cases solid and silicified (Fig. 166, K). When the pedestal is high, it is generally
composed of a rather large number of cells. In this case it consists of tiers,
which are either unicellular throughout (A. eriantka, Mart. et Zucc.), or multi·
cellular in the lower portion of the pedestal (A. pubescens, Willd.); or in other
cases there is a double pedestal (A. auricularia, Boiss.) consisting of a broad
base, the tiers of which are multicellular, surmounted by a long uniseriate
pedestal bearing the terminal cell. In the bracket-hairs of certain species
of Aristolochia (A. auricularia, Boiss., A. Chamissonis, Duch., A. cretica, Lam.,

FIG. J67. A. Tran3v~J"Se section thrOligh the youDg, and

a, throllg:h the older and anomalou.s a:.r:i!l of AmID/ocilla
Idan~14/a"i$, Cham.-After H. Schenck.

A. hirta, L., A. nervosa, Duch.) the hooked terminal cells are replaced by others
which are pointed, or only show a tendency to assume the shape of a hook
(' undeveloped bra'cket·hairs ').
The petiole has be€'u invcstiga ted in A sarom canadense and in a few A 1't:st%c/J·ias
by Petit, and in these cases three vascular bundles pass in to it. InA sarum canaden$e
they run through the entire petiole; in Aristolochia. on the other hand, their number
is increased by branching~ so that the characteristic region exhibits an arc of a rela.-
tively large number of approximated or isolated vascular bundles.
3. STRUCTURE OF THE AXIS. The structure of the axis is generally normal
in this Order. The broad primary med:illary rays, which are composed of
lignified or unlignified cells, are especially characteristic; they separate the
vascular bundles from one another in the Woody species of the genera Aristo-
lochia, Apama and Thottea, as well as in the rhizome of Asarum. In the woody
species of A ristQlochia they extend for long distances through the stem, and
are only traversed by stIands of woody tissue at a few points (Strasburger).
 ARISTOLOCHIACEAE

In some cases (e.g. the woody Aristolochias), in the course of growth in thickness,
broad secondary medullary rays are successively developed in the vascular
bundles themselves, occasioning a dichotomous fission of the plates of wood and
bast composing the vascular bundles; in this way we obtain a characteristic
appearance in transverse section, which H; Schenck has termed the Aristo-
lochia-type. This is specially distinct in young stems of A. triangularis, Cham.
(Fig. 167, A), in which the medullary rays are no narrower than the rays of
wood betwee~ them; the same structure is less typically differentiated in
A. Sipho, A. tomentosa and other species, in which the wood has a denser
structure.
The pith commonly consists of ·unlign:Uied cells in Aristolochia, of lignified
cells in Thottea and Apama: . .
The following facts may be mentioned regarding the structure of the wood,
more especially that of the woody species of Aristolochia, Thottea and Apama.
The vessels attain a considerable diameter (as much as ·14mm.) in the twining
species; their perforations are simple throughout (even in Asarum). The
wood-parenchyma is usually scantily developed, but is more abundant in
Apama siliquosa, Lam. and Thottea grandif/ora, Rottb., where it occurs in rows
accomp.anying the tracheae. The walls of the wood-prosenchyma bear large
bordered pits in Aristolochia, small but distinct bord.ered pits in Apama and
Thottea. Spiral thickening has been observed in the narrower vessels and in
the prosenchyma (with bordered pits) of 4. tomentosa, L. In the rhizome, of
Asarum the xylem contains no prosenchyma, and is composed of unlignified
parenchyma and vessels only.
The first feature to be described in the structure of the cortex is the origin
of the cork; this is only known in Aristolochia, where the cork arises super-
ficially, mostly in the subepidermal layer of cells. The cork of Aristolochia is
composed of alternating layers of thin-walled cubical cork-cells, and cells with
cellulose walls. Asarum europaeum develops no cork; the outer layers of the
cortex simply become suberized and subsequently peel off. Stone-cells are
sometimes fmind in the primary cortex. In Aristolochi~, Apama and Thottea
the outer portion' of the pericycle contains a closed or locally interrupted ~om­
posite sclerenchymatous ring, the fibrous cells of which h:.tve rather wide lurilina,
and are to~ome extent septate by means of thin transverse walls; the inner
parenchymatous portion of the pericycle. is often strongly developed. In
the rhizomes of Aristolochia and Asar'um the §iclerench~atous ring is replaced
by an endodermis. The secondary bast never contams bast-fibres, though
stone-cells may be present. The sieve-tubes often ljave wide lumina, and the
sieve-plates are commonly provided with coarse pores.
Anomalous structure of the axis is onlyl found in A1istolochia triangu-
laris, Cham., according to H. Schenck (Fig. 167, B). It first appears in stems of
this species 3'5 cm. in thickness. By-t;ecolldary dilatation of the pith the
vascular ring first becomes divided' up into ~eight fan-shaped .xylem-segments
corresponding to its eight component bundIes; 'narrow secondary plates of.
wood and bast-are formed, spreading out like a fan, from the sides of the
xylem-segments; and extending round their inner angles, these plates being
derived from a cambium, which appears in immediate contact with the original
segments of the wood.'
Literature: MohI, Ban u. Winden d:Ranken- u. Schlingptl., Tiibingen, 1827, p. 97 and Tab. xi.

1 The stem of the plant, described as t-lrl.rfrVQ£4ia 6ilaba' by Scbleiden and De Barr, !lnd stated
to possess successive rings of growth, is ri6~al in structure (see Solereder, 1.889). So also is the
axis of Bragrzntia Wallichii (Apama siNqUIJsa) j the stem described under this specific name by
Masters is provided with successive rings of growth of the Menispermaceous. type, but belongs to
a. species of Gnttum (see SoleTeder, 1894).
688 ARISTOLOCHIACEAE
-Decaisne, Lardizabalees, Arch. Mus. d'hist. nat., t. i, 18.39, p. 143 et seq.-Griffith, in Transact.
Linn. Soc. of London, vol. xix, 18..rz, p. 334.-Lindley, Veg. Ringd., 18... 6, p. 79:i.-Schleiden,
Gnmdz. d. wiss. Bot., ii. 1850, p. 167 and Fig. 15:1.-Dnchartre, Veget. et struct. anat. des A .•
Compt. rend., t. xxxviii, 1854. p. II4:1 et seq.-Vaupell, Peripher. Wachst. d. Gena.ssb., Leipzig,
18Ss.-Masters, Struct. affin. and distrib. of the genus Aristoloc1u'a etc., Jouro. Linn. Soc., vol. xiv)
1875, p. -487 et seq.-De Bary, Vergl. Anat., l877.-Zacharias, in Bot. Zeit. 1879. p. 633.-Moller,
Rind~anat., 1882, PP'. 12~-5·-Me~tovich, Mark, Klausenburg',,188S, Hungarian; abstr. in Just
1885. J, p. 789.-Henul, Tlge des Dlcot., Ann. sc. nat., s~r. 7, t. 11. 1885, pp. 248-51 and pI. 17.-
Solereder. Holzstr., 1885, pp. 2l:1-3.-Douliot, Periderme, Ann. sc. nat., SeT. 7, t. x, 1889, pp.
33:1-3.-Petit, Petiole, Act. Soc. Linn. de Bordeaux) t. 43. 1889, p. 19 and pl. i.:-Soleredet,
VergL Anat. d. A., Engler, Bot. Jahrb., Ed. x, 1889, pp. 410--5:14 and Tab. xii-xiv and in
Nattir1. Pflanzenfam., iii. Teil, Abt. I, 1889, p. 266-i.-Plancbon, Les Aristiloches, Montl)ellier'
(Hamelin Freres), 1891, l66 pp.-Strasburger, Leitungsbahnen, 1891, pp. 2S6-66.-H. Schenck,
Anat. d. Lianen, 1893, pp. 154-5 and Tab. viii.-Solereder, in Bull. de l'Herbier Boissier, t. ii, 1894,
pp. 38i-6.-(Bastin, Stroct. of Asarum canadense, Aroeric. Journ. of Pbarm. 1894 (see abo the
pharmacognostic works with regard to the-mostly obsolete rhizome :md root of species of Ari'sto a

lothia and A'sarum).-Schwabach, Mech. Ring, Bot. Centralbl. 1898, iv, p. 35 .. ~t seq.]

PIPERACEAE 1. t

I. REVIEW OF THE ANATOMICAL FEATURES. With the exception of the

genus Symbryon, the members of this Order are distinguished by the possession.
of secretory cells, which very- commonly give rise to a transparent dotting of
the leaves; besides these, lysigenous mucilage canals occur in the pith of the
stem of Piper. The stomata are only found on the under side of the leaf, and
are invariably surrounded by a considerable number of epidermal cells, which
are sometimes arranged in a rosette. Both scalariform and simple perforations
have been observed in the vessels. The wood-prosenchyma. when present,
bears simple pits. Where formation of ,:cork takes place it is superticiaI.
Four types may be distinguished in the arrangement of the vascular bundles
in the stem (Fig. 169): 1. The Saurureae (Sattrurus, H outtuynia and A nemiopsis)
have a normal ring of vascular bundles. II. The stem of Verhuellia only contains
a single vascular strand, which is concentric in structure. III. The species
of Piper (incl. Heckeria and Macropiper) have medullary vascular strands
in addition to a peripheral ring of bundles; -the peripheral ring is charact€~rized
by secondary growth by means of a cambium, and by broad primary medullary
rays, and is separated from the pith by a sclerenchymatous ring; the medullary,
bundles in most cases only have slight growth in thickness, and are arranged
in one or more circles. IV. In Peperom",-a the vascular bundles are scattered
in the ground-tissue and only exhibit slight growth in thickness. The hairy
covering is constituted by (a) simple, uniseriate hairs; (b) small structures,
which :tre composed of a limifed number of cells and r~b.1e glandular hairs
(Saururus, Peperomia, Piper); and (c) unicellular pearl~glanch (Piper); a
dense hairy covering is not present in mast cases. Oxalate of lim~ is generally
excreted in the form of small acicular crystals, or of small, sometimes minute,
sand-like crystalline bodies of varying shape, or as 'Clustered crystals; in
Symbryon clustered crystals are found in the epidermli of the leaf, whilst
ordinary large solitary crystals occur in the mesophyU." The Jollowing ana-
'tomical features are of value fo\. special diagnosis in thIS Order: the occurrence
of a hypoderm composed of a varying (often large) number of layers on the
upper side of the leaf (Piper, Saururus, Anemiopsis, and especially' Peperomia);
silicmcatjon of epidermal cells in the leaf (species of Pipet); papIllose differen·
tiation of the epidermis of the leaf (Peperomia marmorata); varying diffe-
rentiation of the peri cycle and of the layer of collenchyma in the primary cortex
of the stem in the species of Piper, and so on.

I For the genus L(ld~, which is included amongst the Piperaceae by Bentham and Hooker,
see Lactoridaceae, p. 39.
 PIPERACEAE
2. STRUCTURE OF THE LEAF. The 1eaves of the species investigated are
bifacial in structure in all cases. The stomata (A nemiopsis, Peperomia, Piper,
Saururus) are found exclusively on the lower side of the leaf, and are surrounded
by several ordinary epidermal cells, or by a rosette of subsidiary cells arranged
according to the Cruciferous type (species of Piper and Peperomia, according to
Benecke). Beneath the upper epidermis of the leaf a hypoderm of one
or more layers has been observed in all the species of Peperomia hitherto in·
vestigated (by Treviranus, Payen, Pfitzer, Beinling and Haberlandt), in Chavica
maculata (Treviranus), in Artanthe colubrina, Miq. (Payen) and also in Saururus
cernuus, L., Piper Khasianum, C. DC., P. Zu,ccarinii, C. DC.l and Anemiopsis
calilornica, Hook. et Arn. (according to my own observation).

lU 1 ! I! II
1-1- 1=!-f- l- I- ~ I- I-f-
~
l-
I- l- I--
l- I- l- l-
I- I-- f- f- I--
f- I- f- f-
t-- ~
l-
f- f- i- f- l-
f- I--f- f- I-- l-
f- f- f- I-
t- f- ~ f-
l-
t- f-
f-f- l- I-
t- f- f-- ~ l- f-
l- I-t- l- I-
f- f- I--
I- 1-1- I- ~ l-
I- f-I- l- I--
l- l- f- I-
l-
I--
t-
l- I- I-- l- l- I--
I-
l- f- l- I- l-
l- f- l- I-
I- I-f- f- ~ ~
~
l- I- f- f- l- I-
l- I- I-
I--
'- l-
I- 1-1- I-- l - f- I-- f- l - l - I-
I-- I- I- I--
I-
l-
i-
l-
r- I-
r-f-
I-f-
I--
-- - - -
l-

I-- l-
- l-
I--
I--
I-- l- I- I-- - I- t-
r-

~
IXI x::I I 1~ ,)
YT ;, 1
I :IT 11"1 IT

B
FIG, 168. A, TransveT'le section of the leaf of PtperO'f1"'(J incanD, Dietr. B, Traosvet'!le section or the I...,.l or
P. peruldaefo/ia, H.B.K. C-D, Hydathodes of P. incDnIJ. E, Pearl-gland of Arlan/"~ Sp.-B after }'Guer,
B after Nestler, the remainder onginalL

The hypoderm of the genus Pepeyomia has been examined in some dttail by
Ffitzer. The num~• •1 component layers varies, and is often very considerable;
in P. aritolia, Miq., the hypoderm consists of a single layer of cells; in P. blanda,
H.B.K., of 1-3: in P. imana, Dietr. (Fig. 168. A) of 6-7 or more; in P. peyeskiae-
folia, H.B.k. (Fig. 168, B) of 14-15 layers. Owing to the large number of layers
and the size (which is ofteJ;l considerable) of the cells in the inner layers, the epidermis
and the subj acent hypoqerm attain such dimensions that the two together in P. incana
are thicker than 1:}le whole of the remaining portion of the fleshy leaf, whilst in
P. magnoliaeloli~'Dietr~ and P. yubella, Hook. they exceed it several times in thick-
ness, in P. pveshiae/olia seven times. According to Pfitzer, the hypoderm in Pepe-
yomia is developed from a single-layered epidermis. In some cases, viz. when cell-
division and growth take place equally in the individual layers of the entire integu-
mental tissue, this mode of origin can still be recognized in the mature leaf, the cells
of the epidermis and those of the hypodermal layers being arranged in rows at right
angles to the surface of the leaf as seen in a transverse section (P. ari/olia). In
P. per8skiaejolia all of the numerous hypodermal cells' coincide vertically, whilst the

1 It may be mentioned here that the conjecture expressed in the Kew Index that Piptr Zuccariuii
might be a species of Pothos has become untenable since the finding of the type-specimen in the
Herb. Monac.
SOLlUIEDER vy
 PIPERACEAE
epidermis consists of smaller cells in consequence of secondary vertical divisions
having taken place. In other cases (P. incana) this arrangement is no longer
recognizable; the outer portion of the hypoderm consists of cells with small lumina,
the inner of cells with larger lumina. The hypodermal cells, in agreement with
their function as water-reservoirs, have contents consisting of watery cell-sap,
rarely including a few chlorophyll-gcains. The cell-walls are mostly thin, but in
P. incana they are thick in the mner layers of the hvpoderm; they are distinctly
collenchymatous in P. magnoliaefolia.
In those species in which there is no hypoderm, the single-layered epidermis
of the leaf often consists of large cells. The spongy tissue also takes part in
the storage of water in the Peperomias, and consequently contains little chloro-
phyll. Sclerenchyma has bee,n observed accompanying the vascular bundles
of the veins in the genus Symbryon only. The following are, special features
in the structure of the leaf: silicification of the epidermis (Piper COlU4
brinum, Link, P. elongatum, C. DC. and P. Betle, L. according to Engler,
P. hirsutum, Sw. and P. Hostmannianum, C. 'DC. according to my own obselva 4

tions); papillose differentiation of the epidermal cells on both sides of the

leaf in Peperomia marmorata (Beinling); spots with a silvery sheen onthe upper
side of the leaf in Peperomia 'argyracea, Hort. Bonn.' (=? P.- argyrea, Hort~),
these being due, according to Pfitzer, to a similar cause to those of the Begonias,
i.e. to intercellular spaces, which 'starting from the palisade-tissue penetrate
between the inner cells of the hypoderm and extend to about the middle of this
tissue; finally, the corky excrescences on the leaves of Peperomia maculosa
and P. obwsi/olia (E. Bachmann).
The hairy covering is dense in rare cases only. It is composed of clothing
hairs," small glands and pearl-glands. The clothing hairs are uniseriate, of
varying length, and have wide lumina. Small glands have been observed in
species of Saururus, Peperomia (Fig. 168, C-D) and Piper; they consist of a
basal cell, belonging to the epidermis, a short'stalk-cell, and a hemispherical,
spherical or sac-like terminal cell, which in the last case frequently lies upon
the surface of the epidermis. In the trichomes in question either no secretion
at all is produced, or only a small amount is formed beneath the cuticle of
the terminal cell. As Haberlandt suggests, they probably in all cases serve for
the excretion 0 f water (as h ydatho des ), ' or at least for the absorPtion of water.
The pearl-glands 1 (Fig. 168, E), first observed by Meyen and subsequently
examined more thoroughly especially by Penzig and Nestler, only appear
temporarily and under certain conditions; they often have the form of
large glistening spherica! bodies (1 mm. or more in diameter) occurring in
moderate numbers on all the aerial parts of the plant, while they are more
numerous on the stem and petiole. They consist of a single large cell, the
narrow base of which is inserted in the epidermis. Owing to the abundance of
plastk nutritive substances (proteid and fatty oil) which they contain, the readily
deciduous pearl-glands are regarded biologically as small food-bodies.
The usual form of excretion of oxalate of lime in this Order (Saururus,
Peperomia, Piper with Macropiper, and Heckeria) is constituted by small
acicular crystals, sometimes (Piper peltatum, L.) resembling raphides, and small
crystalline bodies of a difiert!nt shape, sometimes even so finely divjded as
to be like sand~ In addition to these crystalline elements, which occur in all parts
of the plant, and are frequently found in extraordinary abundance, clustered
crystals are prfsent in the axis;(pith and primary cortex} of Saururus cernuus,
L., and in the mesophyl1 ,of the peperam,&·as. The genus Symbryo,! differs

1 The pearl-glands are described: by Meyen in Piper spurium; by De Bary in P. ttigrum,

Enduo, glaucescffls and Artanllu e/()1tg'ata; by Nestler ill A,ta1t/ne I cordifolia,' Miq. (= 1 A.
comifoHa, Miq.).
 PJPERACEAE

from t~e other members of the Order in the features presented by its crystals :
the vems of the leaf of S. tetrastachyum, Griseb. contain ordinary large solitary
crystals, which have not been observed elsewhere in the Order, whilst both the
upper and lower epidermis include small cells mostly united in groups, and
filled with small clustered crystals.
Two kinds of internal secretory organs are found in the Pipefaceae, VIZ. :
(a) secretory cells, occurring in the leaf and axis, and, in the former~ giving
rise to transparent or opaque dots, ac"ording to the nature of the secretion,
and (b) mucilage-canals, which only occur in 1:he axis. The secretory cells
have been shown by Bokomy to occur in the leaf of numerous species of Sau-
rurus, Piper and Pep e1'omia , and according to my· own observations they are
also present in the monotypic g~nus Zippetia and in Anemiopsis (A. cali/ornica,
Hook.); they are wanting in Symbryon only 1. In the leaf the secretory cells
appear in the various tissues, e.g: in Piper Hostmannianum, C. DC. and P.
hirsutum, Sw. in the mesophyll only, in Saururus cernuus, L. in the mesophyll
and in the lower epidermis of the leaf, in A nemiopsis cali/ornica, Hook. in the
mesophyll and in both the upper and lower epidermis. In the axie; they occur
in the pith, bast and primary cortex. The secretory cells mostly have a spherical
shape. Their walls ate suberized, and their contents are generally clear, more
rarely (in the' folia nigro-punctata' of Peperotnia melanostigma, Miq., p, nigro-
punctata, Miq. and P. hirsutum, Sw.) of a brown colout. Tll~re are other elements
which must not be confounded with these spherical brown cells which replace
the clear secretory cells, viz. cells filled with brown contents Lut not differing
from the ordinary cells of, the mesophyll in shape; such cells may be observed,
for example, in the dried leaf of Piper cernu'!Jm, VeIl. and Sau1urus cernuus, L. ;
in the living leaf their contents (at least in Saururus) are dear and strongly
refractive. Mucilage~canals have only been observed in species of Piper
(Arlanthe Zacuapana, Cham. ct Schlecht.'=Piper titiaejolium, Cham. et
j

Schl. according to Debray, Piper fluminense, C. DC. according-to H. Schenck,

and P. Carpunya, R. et P., P. Khasianum, C. DC., P. nepalense, Miq., P.
nigrum, L. and P. Zuccarinii, C. DC. according to my own investigation).
Either one or several of them occur in the pith; they have wide lumina, and
without doubt have a lysigerious origin.
3. STRuCTURE OF THE AXIS. In the structure of the axis, which has
been examined especially by C. de Candolle, ]. E. \Veiss and Debray, we may
distinguish four types, requiring separate description.
The":Hrst type is found in the genera Saururus, Houttuynia and Anemiopsis,
which are members of the tribe Saurureae ; in this, type a normal ring of vascular
bundles is present. The bundles lie isolated in Jhe parenchymatous ground-
tissue of the stem, and are as a rule about '. equidistant from the centre;
but in the case of Saururus those bundles which are the next to bend out
into the leaves, project a little further towards the exterior of the transverse
section.
As an example of this type the stem-structure of Sauyurus cernuus~ L. may be
described in somewhat greater detail. In this plant, which grows in man;hy locali-
ties. the pith, containing numerous rather large intercellular spaces, is surrounded
by a ring of vascular bundles; these bundles may be shortly described as embedded
in a ring of sclerenchymatous fibres. Thus each vascular bundle is surrounded by
a ring of sclerenchyma, thick~r on the outer than on the inner side of the bundle,
and the outer parts of these sclerenchymatous rings are connected with one anoth.er
by narrow bridges of the same tissue. The xylem in the vascular bundles of Sauruffi_s
consists of primary tracheae, pitted vessels and lignified parenchyma; the pitted

1 Secretory cells are present in the three species of Pip" (P. aurilllm, H.B.K., P. arnuum,
VeIl. and P. Enckea, C. DC.), in which Bokomy states that they a~ absent.
Vy2
 PIPERACEAE

vessels a.ttain a diameter of '075 mm., and exhibit sca.larlform perforations with many
bars; they bear scalariform bordered pits wher~ they are in contact with other
vessels, and relatively large simple pits in contact ~tll parenchyma. Certain por·
pons of the primary cortex show rather large in.te7CeHlllar spaces..
V erhuellia belen~s
to the secOl14 type. Although it is not aquatic,
this genus, like certam dicotyledoJilous water-plants, possesses only an axile
strand, sympodial in structure and consisting of weak and ver8 closely approxi-

.
~ ~
~
~ . @
@ €lI
~
~ ~ ~
~

• ~
e
~
\j
~ FIG. 169. A-B, p,~" CtJrIJil1J;,a,
R. et P.: A, Transverse section of the

~.
young stem; H, Portion of a \ransveree
.. section of the stem after tbe com-
~ ~,
,
menc:ement (If gT'O'o\th in thickness •
~ C. Transverse section of the t_aang
stem of P. fiumhun.u, C. D n,
Tran!ilVerge section of the st.ern or R!-
fi t"imJia incana. Dietr.-C after H.
c:henck, the remainder orig inaJ.

mated leaf·traces, which are fused longitudinally. The axile strand appears
to be concentric, according to Schmitz's brief statements.
The third type (Fig. 169, A -B) is consti tuted by the genus Piper in the w.ider
sense, including Heckeria and Macropiper. In this case two kinds of vascular
bundles are present. The one kind forms a peripheral circle, in which the bundles
are distinct from one another in the young stem (Fig. 169, A), and are applied to
a scIerenchymatous zone limiting the pith externally and generally having an
undulated course; after the commencement of activity of the cambial ring
(Fig. I~, B) these bundles grow in thickness and become separated from one
 PIPERACEAE

another by broad primary medullary rays. ,Those of the second kind are
medullary vascular bundles, which lie isolated in the pith and are arranged
more or less <_iistinctly in one or more rings; as a rule they only exhibit
a limited growth in thickness by means of sirips of cambium of the same
breadth as the'vascular bundles.
The following statements regarding the detailed £tructure of the stem
of Piper may be quoted (chiefly from J. E. Weiss). The formation of cork takes
place in the epidermis (P. Carpunya, R. et P., Chavica Roxburghii, Miq.), or in
the outermost cell-layer of the primary cortex (Artanthe cordi/oUa,' Miq.,1 f

Piper bullatum, VahI, P. geniculaturn, Sw.). The primary caortex contains

either a collcnchymatous ring or isolated bundles of collenchyma. In certain
species (Piper bullatum, P. genicula tum, P. peltatum, L.) the collenchyma retains
its character, even in the older branches; in other.s sclerenchymatous fibres in
varying numbers develop in the collenchy:r;na (Artanthe cornifolia" Ckavica
Betle, L., C. Roxburghii, Piper Bredemeyeri, Jacq.), or the whole of the collen-
chyma becomes transformed into a ring of sclerenchymatous fibres (Piper
Carpunya, P. rivinoides, Kth.), which subsequently becomes ruptured in the
course of growth in thickness. In the older branches of Piper nigrum, L. I found
isolated bundles of sclerenchymatous fibres, which were to some extent united
by sclerosed parenchyma. The innermost portion of the primary cortex is
formed by an endodermis, which either surrounds the entire peripheral ring
of vascular bundles (Artanthe cornijolia, Chavica Betle, &c.), or is only deveioped
in contact with the bast of the bundles (Piper Carpunya'. Towards
the pith the peripheral ring of bundles rests against the inner scleren-
chymatous sheath mentioned above; this sheath is either entirely composed
of sclerenchymatous fibres, or is parenchymatous opposite the primary medullary
rays; it is continuous in the young branch, but becomes split up into fragments
after the commencement of growth in thickness. Corresponding to this inner
sheath there is frequently a sclerenchymatous layer at the outer margin of
the groups of bast; it is either developed in the form of semilunar bundles of
fibres united into a ring by groups of sclerosed parenchymatous cells (Piper
nigrum), or it is represented by semilunar groups of hard bast alone (A1tanthe
corni/olia, Chavica Betle, &c.) or by a few bast-fibres (Piper Carpunya, &c.),
whilst in Piper bulla tum, for example, it is absent. The growth of the peripheral
zone of vascular bundles takes place by means ,9£ a cambial ring, which produces
primary medullary rays, 10-16 cells in breadth, between the bundles;
these medullary rays traverse the internodes in the form of longitudinal
plates and are composed of cells only slightly elongated in the vertical
direction. The xylem-portions of the vascular bundles, apart from the
primary tracheae, which are enclosed in thin-walled parenchyma, consist
chiefly of the following elements: (a) pitted vessels, having mostly simplet
more rarely (according to Debray) also scalariform perforations with few bars;
(b) wood-prosenchyma bearing simple pits, and sometimes septate by means
of thin transverse walls~;""and (c) secondary, generally broad, medullary rays
(of the Aristolochia-type). #-lhe medullary vascular bundles are embedded
in a thin-walled pith and are of the nature of leaf-traces; they are collateral
in structure, and their wood and bast are generally normally orientated. They
are either arranged in a single, more. or/less regular ring (Piper Byedemeyeri
Chavica Bette, &c.), or in two (Piper gcniculatum) or even more rings (P. bullatum
with about 25 vascular bundles in the pith). The xylem in these bundles
has the same composition as that of the peripheral bundles; groups of sder~n·
chymatous fibres are frequently developed at the inner margin of the xylem

1 No doubt Arlonthe ~ornifo/ia, Miq. is the plant referred to. The incorrect Qamc 'A. cmi-
folia' seems to be widely spread in gardens (cf. footnote, p. 690).
 P/PERACEAE

and at the outer margin of the bast. The growth in thickness of these bundles
is often very considerable (especially in P. geniculatum and Chavica /rustrata,
Miq.).
According to H. Schenck, the stem-structure of Piper fluminensc. C. DC. deviates
somewhat from that of the other species of the genus. The young stem (Fig. 159, C)
bas, as its essentiaJ feature, two rings of vascular bundles, of which, curiously enQugh,
the bundles composing the peripheral ring and resting against the sClerenchyma.
are very small, while the bundles forming the medullary ring are much larger.
In the older stem (5 cm. in diameter) the bundles of both rings are found to have
grown in thickness individually. even the peripheral bundles growing by means of
cambial strips, which do not unite to form a, ring.
The genus Peperomia belongs to the fourth type, in which the vascular
bundles are scattered in the ground-tissue (Fig. r69, D). Regarding the detailed
structure of the stem of Peperomia, the following statements may be made on
the authority of. J. E. Weiss and from my own investigation of P. carthagi-
nensis, C. DC. The epidermis is simple (P. rubella), or consists of two layers
(P. va1'iegata~ R. et P ..&c.). The cork arises in the epidermis; when tbe latter
consists of two layers, the cork develops in the outer layer (P. carthaginensis,
P. variegata). Collenchymatous tissue is very strongly developed at a depth
of a few layers of cells" beneath the epidermis. This is followed internally
by parenchymatous ground-tissue, in which the vascular bundles are embedded.
The number and mode of· arrangement of the bundles varies. Their distribu-
tion is sometimes rather irregular, in othel! cases one can distinguish two or
more rings of bundles ~ -when two of these are present, the bundles composing
- them alternate in the'transverse section. Tp.e bundles are generally collateral
in structure, and their wood and bast is normally orientated. They are either
of equal size, or the outer (P. incana, Dietr.) or inner bundles (P. brachy-
phylla, Dietr.) are the larger; in P. verticillata, Miq. there is a very large central
vascular bundle, which has concentric structure (with central xylem) and
may perhaps have arisen by the fusion of several bundles. The bundles are
commonly provided with an endodermis, which either s~rounds the whole
bundle (P. obtusi/olia, Dietr., &c.) or only-forms a semicircle on the outer side
of the bast (P. urocarpa, Fisch. et Mey., &c.); in other species, such as
P. amplexi/olia, Dietr., &c., the endodermis is absent. There are 0.0 scleren-
chymatous elements in the vascular b~ndles. The bast is sometimes collen-
chymatous (P. variegata, R. et P.); the xylem consists of unlignified paren-
chyma and vesselS>. There is only slight growth in thickness.
Literature: Treviranus, in Verm. Schr. iv, ]S:u, p. II and Phys. d. Gew. i, ,]835, p. 449.-
Meyen, Sekretionsorg., 183i, p. 47.-Sanio, in Eot. Zeit. 1862. p. 213 and 1864, p. :J7.-C. de
Candolle. Mem. s. 1. fam. d. P., Mem. Soc. de phys. et d'hist. nat. Geneve, t. xviii, 1, 1866,32 pp.
and 6 Tab. and in DC. Prodr. xvi,], 1869, p. 235 6~ et seq.-Pfit:ter, Mehrsch. Epid. etc., Pringsheim
Jahrb., Bd. viii, 1871-2, pp. 26-31 and Tab. vi.-Schmitz, Fibrovasalsyst. d. P., Diss., 1871 and
Vuilut:llia, Flora 18i2,t P~_408'-l' E. Weiss, Wachstumsverh. etc. d. P., Flora 1876, p. 311I et seq.
and Tab. xi-xii.-De .cary, Verg. Anat., 1877; here on p. 260, footnote 2, further older litera·
ture.-E. Bachmann, Korkwuch., Pringsheim Jahrb., Bd. xii, 1879-81, p. 203 et aeq.-Beinling,
Anat. d. Laubbl. u. Wurz. .-on Peptromia, Cohn, &itt. z. Biol., Ed. iii, Heft I, 1879, pp. 32-7 and
Tab. iv.-Zacnarias, in Bot. Zeit. z879, p. 61oif..-Bokorny, Dllrchs. P., Flora. 188z, P.565 et seq.
and sep. copy, pp. :U-3.-Morot, Pericycle, Ann. lie. nat., set. 6, t. xx, 1885, p. 2C)8.-So1ereder,
Holzstr., 1885, pp. 223-4·-Debray, Caract. anat. et parcours des fai~. fibro.vase. d. P., Paris
(0. Doin), 1886, 107 pp. and 16 pI.; abstr. by Sanio, in BoL CentIalbl. 1886, ij, pp. J36-60.-
Solereder. Sekretz. bei den P., Engler, Bot. Jahrb., Bd. x, 1887, pp. 509-II.-Beneclce, in Bot. Zeit.
18glil, p. 555 et seq.-Houlbert, Bois sec. dans Ies Apetales, These, Paris, 1893, PP.5S-62.-Nestler,
Perldriisen von Artanthe (Drdzfolia, Miq., Oesterr. bot. Zeitschr. 1893, pp. 332 and 386 et seq.,
Tab. xvi.-Penzig, Perldriisen, Report of Congress, Genoa, 1893, p. 237 et seq. and Tab. XV.-
H. Schenck, Anat. d. Lianen, 1893, pp. 42-5 and Tab. i.-Engler, in Natiirl. Pflanzenfam., iii. Teil,
Abt. I (1894), PP' 1 and 4-s.-Haberlandt, Trop. Laubbl. ii, Sitz .• Ber. Wiener Akad., Bd. ciii,
Aht. r. 1894. p. 5u et seq. a.o.d Tab. iii.-Virchow, Blatttahnet.Archiv d. Pharm. 1896, scp. coPl~
p. 39.-Uaderbolm, Anat. stud. ofver sydamerik. Peperomier, Diss., Upsala, 1898, 99 PP') :I Tab.J

,""'" .- .
 695

CHLORANTHACEAE.
This Order is distinguished anatomically by the following characters;
the possession of secretory cells; the exceptionally numerous bars in the per-
forations of the xylem-vessels; normal stem-structure; the absence of a hairy
covering and the lack of a special type of stoma. The two genera Chlo,an-
thus and Hedyosmum (the third genus Ascarina has not been investigated)
may be distinguished by the pitting. of the wood-prosenchyma, which is
typically bordered in Chloranthus, but simple in H edyosmum, as well as by the
occurrence (Hedyosmum) or absence (Chloranthus) of mucilage-canals at the
margin of the pith.
The secretory cells were first observed by Blenk, according to whom they
frequently give rise" to transparent dots in the leaf. In',the leaf, as far as I am
aware, they are only present in the mesophyll, never in the epidermis. They
are also found in the cortex and pith of the axis. They are spherical in shape
and have a suberized membrane. They have been shown to occur in the two
genera Chloranthus and Hedyosmum and will probably also be found in
Ascarina, since the plants of this genus are described as aromatic.
Secretory organs of a different kind, viz. the mucilage-canals 1, referred to
above, occur only in the species of Hedyosmttm. I have met with these canals
at the margin of the pith in H. brasiliense, Mart., H. glabratum, H.B.K. and
H. racemosum, Don, and in a similar position in the petiole of H. arborescens,
Sw.; they appear in the form of wide iIltercellular spaces, which are elliptical
in transverse section and are apparently of schizogenous origin. Their contents
consist of mut:ilage, which in H. brasiliense encloses sphaerqcrystalline masses;
the latter are insoluble in hot water, alcohol and ether, but readily soluble
in hydrochloric acid.
I have examined the structure of the leaf in H edyosmum brasiliense, Mart.,
H. arborescens, Sw. and Chloranthus bra chystachys, Bl. In these species it is
bifacial, though typical palisade-parenchyma composed of long cells does not
occur. The latter is replaced by short-celled arm-palisade (of Haberlandt).
The spongy tissue has large intercellular spaces. The stomata are only found
on the lower side of the leaf, and no definite type of stoma is evident. ;But
in H. arborescens the neighbouring cells are marked by their arrangement in a
rosette, arld in Chloranthus brackystachys ODe or two subsidiary cells, parallel
to the pore, may be seen here and there. In H. arborescens there is a single
layer of hypoderm beneath the epidermis on the upper side of the leaf. The
vascular bundles of the veins are provided with a sclerenchymatous sheath.
The structure of the wood has been examined in H edyosmutn brasiliense,
Mart., H. arborescens, Sw:, H. racemosum, Don, Chloranthus brachystachys, Bl. and
C. officinalis, Bl. The medullary rays of the wood are o~asionally very broad
(e.g. in H. brasiliense), in other cases less so ;- their cells are always considerably
elongated in the vertical direction. The diameter of the vessels is not great
(reaching ·045 mm.); in branches of Chloranth~ts brachystachys from herbarium..
material the pitted vessels are confined to the neighbourhood of the
primary wood._· Scalariform perforations, with very numerous bars, are
highly characteristic of the Chloranthaceae; in H. brasiliense and H. racemosum
as many as IOO bars may be counted on the same perforation. In contact
with parenchyma the walls of the vessels are furnished with bordered pits.
The wood-prosenchyma bears typical, large bordered pits in Chloranthus, whilst
in Hedyosmum the elements of this tissue have simple pits, and are frequently
provided with delicate transverse walls.
In order to be able to add some information regarding the structure of
J Engler's statement, that the species of Hedyosmum are characterized by gelatinization of their
medullary tissue, is inaccurate.
 CHLORANTHAC£AE
the cortex, I examined the'latter in Chloranthus brachystachys and Hedyosmum
brasiliense. No formation of cork was observed. In Chloranthus brachystachys
the outer limit of the bast is formed by a composite and continuous scleren~
chymatous ring composed of groups of bast-fibres and of sclerosed parenchy-
matous cells with wide lumina; in H. brasiliense isolated groups of scleren-
chymatous fibres alone occur in this region. In this species the Quter portion
of the primary cortex is differentiated as typical collenchyma, and the epidermis
is subpapillose.
The occurrence of silicified groups of cells in the leaf of H; brasiliense
and H. a'fborescens deserves special notice. The silicification involves a group
of epidermal cells and the mesophylJ-ceJls lying beneath them. Similar silicified
cells have also been observed in the cortex of Hedyosmum brasiliense in the neigh-
bourhood of the primary groups of hard bast. Crystalline elements were
entirely ab=ent in the leaves examined. In the pith and the inner. portion
of the primary cortical parenchyma of H. brasiliense numerous small clustered
crystals were observed, but nowhere else.
I
Literature! Moller, Holzanat., Denkschr, Wiener Akad. 1876, pp. 19 and 3fs.-Blenk, Durchll.
P., Flora 1884, p. 37~ and sep. copy, p. 84.-Solereder, Holzstr., 1885, pp. 124-s.-Engler, in
Natiirl. Pflanzenfam., iii. Teil, Abt. 1, p. r:.:.

MYRISTICACEAE.
The following anatomical features are characteristic of this Order: secretory
cells; the excretion of oxalate of lime in the form of small acicUlar crystals,
which are accompanied by clustered crystals; the absence of glandular hairs (?),
and the occurrence of extremely characteristic trichomes, resembling stellate
hairs and composed of one· or two·armed cells; stomata with two subsidiary
cells, arranged parallel to the pore; the tendency to the formation of scalariform
perforations in the vessels; the simple pitting of the wood-prosenchyma; the
superficial formation of cork; the occurrence of isolated groups of hard bast
in the pericycle, and the development of secondary hard bast.
The secretory cells, which like those of the preceding Orders frequently
give rise to pellucid dots in the leaf, have been shown by Blenk to occur in the
leaf of a large number of species. They a.re, however, also found :.n. the axis,
being situated in the pith, bast and primary cortex. The secretory cells of
the leaf never occur in the epidermis, but are restricted to the rnesophyll ;
where they occur in the palisade·parenchyma, they are mostly found onli in
the inner layer of this tissue, though exceptionally (Iryanthera ma crop hylla,
Warb.) they may lie immediately beneath the upper epidermis. The secretory
cells are spherical in shape; their contents are semi-fluid in some cases, whilst
they are solid, crystalline and doubly refractive in others. The colour
of the secretion may be either yellow or reddish, in a few cases deep brown,
or almost black. In some species the secretion had entirely or partially
disappeared in herbarium material. When treated rnicrochemically, the walls
of the secretory cells in the leaf of Myristica o{ficinalis, Mart. show distinct
. differentiation into an outer suberized and an inner cellulose lamella. In
M. Horsfieldii, Bl. (= Horsfieldia iryaghedhi, \Varb.) and M. Vrieseana, Miq.
(= H. Irya, Warb.), according to Blenk, the membranes of the secretory cells
in the leaf are strongly thickened and gelatinized; in water they sweJl up to
such an extent that thry appear to dissolve, and in this way the resinous contents
are forced out of the secretory cells 1.
~------------~~----~--~--~
1 For information regarding the secretory cells occurring in otLer parts of the plant $te Voigt,
Ban u. Entwickl. d. Samens u. SAmenmantels von .Myrislica f"ag1'ans, Diss., Gottingen, 1885;
Tschirch, Inbaltsst. des Arillus vo!). lIfyristica jmgrans, Ber. deatsch. bot. Gesellscb. 1888, p. 138
et seq. and Warburg, Monogr., loco cit.
 .lIlYRIS T ICA CEA E 6c)7

In many members of the Or~r, besides the secretory cells, therEt are
elongated sacs having brown or red tanniniferous contents and occurring at the
margin of the pith and in the secondary l*lst of the branch. as well as in 'tofTe-
sponding positions in the veins of the leaf.; these elements are generally dis-
tinguished from the neighbouring cells by baving larger lumina, and they
recall the well-known tannin-sacs. In the living plant their contents are
aqueous or bright yellow, but become red on exposure to the air.
I hav~ examined the leaf-structure especially in Virola otficinalis, Warb.~
and a few statements on this subject have also been made by Blenk, Prantl
and Warburg.- In the species just mentioned the leaf has bifacial structure, and
the palisade-tissue consists of several.layers of long cells. The epidermal cells
on both sides of the leaf have small polygonal outlines. Hypoderm is mentioned
by Warburg as occurring on the upper side of the leaf in Dialyanthera. Accord-
ing to Blenk, the lower epidermis of the leaf commonly exhibits a more or less
papillose differentiation. According to my own observations, distinct papillae
are found in Virola offici'nalis, whilst in V. sebifera, AubI. the lower epidermal
cells ant merely arched outwards. In V. otJicinalis and V. sebifera the stomata
are confined to the lower surface of the leaf; in both 5pecies a single subsidiary
cell is placed parallel to the pore on either side of the pair of guard-cells. In
V. otJicinalis the smaller veins show a tendency to be verticaily -transcurrent
by means of unlignified tissue; no sclerenchyma is developed in connexion
with them, and the same applies to the larger veins. Warburg describes
sclerenchymatous fibres, running irregularly through the mesophyll, in the genus
Gymna cranthera, and branched spiclliar cells, frequently of stellate form, in
species of Iryanthera, especially I. macrophylla, Warb.
The following statements regarding the structure of the wood are based
on Warburg's work and on my investigation of Virola officinalis and V. sebife1·a.
The medullary rays are narrow, being from one to two cells in breadth. 'The
vessels are for the most part arranged radially in the transverse section of the
axis, and attain a diameter of ·o6-·r mnl. The perforations of the vessels are
chiefly scalariform with r-8 bars, but, besides these, simple, elliptical or
circular perforations are also present. In contact with parenchyma the walls
of the vessels show all transitions from large simple pits to bordered pits.
The wood~arenchyma is scantily developed; the wood-prosenchyma bears
simple pits and is locally septate by means of delicate transverse walls.
The structure of the cortex has been examined by Moller in rather old
portions of the bark of V. sebi/era, and by me in branches from herbarium-
material of V. otJicinalis. The cork arises superficially in V. otJicinalis, viz.
in the second layer of the primary cortex, and includes cens sclerosed on one
side. In the same species the outer limit of the bast is {)ccupied by isolated
groups of primary bast-fibres. In the inner portion of the primary cortex
there are groups of lignified cells corresponding in position to the interspaces
between the groups of bast-fibres just mentioned. The occurrence of secondary
hard bas~js.especiaIly-.noteworthy, and could be determined even in herbarium-I
material of. V. ctJicinalis. In older specimens of the cortex (V. sebifera). the
secondary groups of bast-fibres combined with stone-cells give rise to a concen-
tric stratification of the bast. Moller states that in the sieve-tubes of V. sebifera
the sieve-plates are not found on the end-walls of the segments, but on their
longitudinal walls; it remains to be determined whether this is the case in
all the members of this Order. Tannin is present in some abundance in the
cortex.
Oxalate of lime is excreted chiefly in the form of small acicular crystals,
and these occur in abundance in the cortex of the stem and in the veins of
the leaf. In addition to these, clustered crystals are present in the leaf in
many species, being mostly enclosed in rat~er large subepidermal cells.
6g8 MYRISTICACEAE

The hairy covering of the Myristicaceae (Fig. 170) is of a very peculiar

type, the hairs being generally described by systematists as of the stellate
form; Warburg has recently made a careful examination of them. Although
the various forms of hairs which occur in the individual species, or even on
the different organs of the same plant, show considerable diversity when ex-
amined in detail, they may all be regarded as modifications of a single principal
type. To express it shortly, the hairs of all the Myristicaceae are uniseriate
trichomes sympodially branched. They may be classified in two categories,
which are connected by transitional forms. The first of these (Fig. 170, A-C)
is represented by hairs consisting of cells of the one-armed form; hairs
of this type originate from a uniseriate trichome, through the component

FIG. 170- Hs.ifY coveriog of the Myristic,!-cue: A, Kntma H~Ju'''(l_, Warb.

(lear). B-C, Vlrola. sehiftra Aubl. (leaf): !I, Trichome seen f~m the Iillde; c, &E.eD f:om
above. D-B, Dia/yan/lura 'Oloha, Warb. (inflorescence): D, Trkhc.me from the ~~;
E, rrom above. F, My","sHca C()()AU, Warb. (leaf): trichome from above. G, Mj.,.,slietJ
vilJosa, Warb. (Ieaf).-Aner Warburg.

cells forming lateral protrusions, like branches or rays, these l

being produced at the upper end, or m<;>re ruely at the
middle or base of the cells. Various modifications of this
type are brought about as follows: protrusions may. be
formed by all the cells of the trichome, or only in the
apical region of the latter, so that the hairs may possess a
stalk, variously differentiated, or may have none; again the
portions of the cells combining to form the main 'stem J
of the trichome may be short or long; and lastly, the protrusions vary in
length and show all transitions from papillose protuberances to ray-
like processes, and the latter may radiate in the most varied directions.
In this way a series of forms is obtained, from trichomes resembling stellate
hairs to those of an abietiform type. The second category (Fig. 170,
.D-G) is composed of trichomes the cells of which are two-armed. In
the simplest case we have a bue, simple, two-armed hair, which ic;
unicellular and is mostly seat~d on a low stalk-cell. More commonly one
finds a pair of two-armed trichomes one above the other; in such hairs
the two segments do not generally lie in one vertical plane, but intersect
one another (Fig. 170, D-E). Further, it is not rare to find trichomes, which,
apart from one or a few short basal cells, consist of a still larger number
(as many as six) of two-armed celis, all intersecting at various angles in such
a manner that the trichomes have the appearance of stellate hairs, when seen
from above (Fig. 170, F). In other cases the number of two-anned celIs is
 MYRISTICACEAE

much grea!er, and the trichomes ap~c:r like shaggy ~airs to_ the ?aked eye;
in these halTs the two·armed cells all he In the same vertIcal plane (FIg. I70, G 1).
It may be added that Warburg also met with simple, unicellular hairs as an
exceptional occurrence. Glandular hairs appear to be absent II.
Literature: Moller, Holtanat., Denkschr. Wiener Akad. 1876, pp. ~ and 36s.-Moller, Rinden-
.ahat., 188l, pp. :ua-s.-B1enk, Durchs. P., nora 1884, p. 372 and sep. copy, pp. 84-5.-So1ereder,
Holzstr., 1885, pp. ll5-6.-[Thouvenin, LOcal. do tannin dans les M. and Struct. des M., Bull. Soc.
sc. Nancy I887.J-Prantl, in NatUrl. Ptlanunfam., iii. Teil, Abt. 2, p. 4o.-Warburg, Haarbild. d.
M., Ber. deutsch. bot. Gesellsch. 1895, pp. (78)-(8a) aud Tab. xxix j in Naturl. Pflanzenfam.,~achtr.
u. Reg. zn Teil ii-iv, 1891, p. 162; and Monogr. d. M., Nova Acta, Halle, 1891.

MONIMIACEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. The following characters
are those of most value for the anatomical diagnosis of the Order: the presence
of secretory cells, which frequently give rise to transparent dots in the, leaf;
the absence of mucilage· cells and glandular hairs; the occurrence in the vessels
of scalariform perforations, in addition to which simple perforations may also
be present; the development of-a composite and continuous sc1erenchymatous
ring in the pericyde, induding stone-cells thickened in the shape of a horse-shoe
(exceptions: Conuleum and many species of Siparuna); the excretion of oxalate
of lime in the form of small acicular crystals, or small cubical or rhombohedral
crystals, several of which invariably occur together in the same cell. With
reference to the leaf-structure, it may be added that the leaves are bifacial, that
hypoderm is developed beneath the upper epidermis in most species, and that
the vascular bundles of the veins are usually surrounded by a sderenchymatous
ring. The type of stoma is not uniform, the guard· cells being either surrounded
by several epidermal cells, or accompanied by subsidiary cells, which lie parallel
to the pore. The prosenchyma which forms the groundwork of the wood bears
either bordered or simple pits; septate prosenchyma bearing simple pits has been
observed in all the species investigated. The medullary rays are strikingly
broad in the Monimieae, but narrow in the Atherospermeae. The formation
~f cork takes place superficially. The hairy covering consists of simple uni-
cellular trimome~, two-armed unicellular hairs (Mollinedia, Matthaea, Athero-'
spenna), tufted hairs (Peumus, Siparuna), stellate hairs (Monimia, Palme1ia,
lIorlonia, species of Siparuna, Fig. I7I), and peltate hairs (Conuleum, species
of f.lonimia and Siparuna). The following special features, found in certain
species of Siparuna, may also be mentioned: the presence of tannin·sacs
with wide lumina in the pith and bast; the occurrence of sderenchymatous
fibres in the mesophyll, and of an epidermis, two or more layers thick, in the leaf.
2. STRUCTU~ OF THE LEAF. This has been investigated in detail by
Hobein '. In all the species the leaf-structure is bifacial; the palisade-tissue
mostly consists of a single layer, rarely of two or more layers of cells; which vary
in length, whilst the spongy tissue"is loose and frequently characterized by the
presence of large intercellular spaces. The epidermal ceUs on both sides of
I the leaf are generally small, and usually have straight lateral walls, strongly
undulated walls having only been observed in Doryphora. Gelatinization of
the epidermis of the leaf does not occur, but the development of a hypoderm

1See also tab. 40 et seq. in Flor. brasil. v, I.

SI was unable to find the' glandular papillae, consisting of a short stalk-eell and. a second ~11
of only slightly greater size fonning the head,' as mentioned by Wllrburg in M~ristica arg'tnlea,
Warb.
a The following genera. were examined: (a) Tribe Monimiea.e: Monimia, TanllMurissiI,
Palmerla,Mo/1inedia,Kibara, Mallktua,H~d)'ca"'iI, PeumUJ,HiWlonia; {b)Tri~ Aiherospenneae:
Conuleum, Sipanma, Atlzel'osperma, DDY')'pnora, Laurelt'a, Dapnnandra.
 MONIMIACEAE

is
on the upper side of the leaf a phenomenon of wide occurrence; among the
genera and species examined by Hobein hypoderm is absent in the fol1ow~
ing cases only: Atherosperma, Daphnandra, Matthaea and certain species of
Siparuna; according to Poisson, it is present in Hennecartia. The hypoderm
consists either of a singl e layer (e. g. in M ollinedia) or of several (e. g. in
Monimia ovaUlolia, P. Th.). Its cells are usually larger than the epidermal
cells in surface-view Rnd frequently (Monimia, Palmeria, Mollinedia, Tam-
bourissa) also exceed the latter in height in the transverse sectiofl. In some
cases the walls of the hypodermal cells are thin (e.g. in Hedycarya), but often
they are strongly thickened and pitted, and in M onimia ovalifolia they are
lignified as well. In those species of Siparuna which have no hypoderm, the
epidermis on the upper side of the leaf is simple (S. chiridota, A. DC., &c.), or
consists of two or more layers, either throughout or only at certain points (in
the majority of the species of Siparuna); in the latter case the appearance
of horizontal division-walls in the epidermal cells is often confined -to the
neighbourhood of the trichomes. On the lower side of the leaf hypoderm has
only been met with in Laurelia sempervirens, Tul. The stomata are almost
exclusively confined to the lower_side of the leaf; only in species of-SiParu1~a
have they been observed on the upper side, where they are restricted to the
neighbourhood of the veins. The guard-cells are in most cases surrounded
by several ordinary epidermal ceIJs; Hobein states that in Mollinedia, Kibara,
MaUhaea and Hedycarya there are four subsidiary cells, two of which are
placed laterally with reference to the guard-cells; he describes two lateral
subsidiary cells in Conuleum also. The vascular systems of the larger veins
are as a rule surrounded by a ring of hard bast. The only exception is
found in a few species of Siparuna, in which the hard bast is absent or only
slightly developed. In S. mollicoma, A. DC. and S. mollis, A. DC. sclerenchy...
matous fibres branch off from the sclerenchyma of the veins and run freely
in the mesophyll, sometimes extending as far as the epidermis of the leaf.
Bokorny and Hobein met with secretory cells in the leal in all the members
of the Order investigated, and according to my own observations they are also
present in the leaf of Hennecartia. The position in which they occur in the
leaf is generally the mesophyll, and they frequently give rise to transparent
dots in this tissue. Besides occurring in the mesophyll, howe~r, they are
also present in the epidermis of the leaf in certain species of Tambourissa,
Mollinedia, Conuleum and S'paruna, and in Daphnandra, in the hypoderm
in Palmeria; in M onimia, on the other hand, they are confined to the hypoderm
and are only present in small numbers. In the axis the secretory cells are
found in the pith, bast and primary cortex, and rarely (Hedycarya arborea,
Forst.) in the medullary rays of the wood. The secretory elements are spherical
in shape and their contents are for the most part clear, more rarely (species
of Siparuna) coloured brown by tannin; their diameter varies from '02 to
'06 mm, Mucilage-cells are not present in the Monimiaceae, this being a point
of difference from the allied Laurineae.
Oxalate of lime is mostly present in the leaf and axis in the form of small
acicular crystals, large numbers of which occur together in the same cell; they
frequently OCCU!?y the whole of the mesophyll. In the "genera belonging to
the tribe Monimleae, besides the crystalline elements just referred to, there are
smalJ, cubical or rhombohedral crystaJs (attainjng the greatest relative dimensions
in M oUinedia), several occurring in the same cell; these crystals are found in
the neighbourhood of the vascular bundle3 of the leaf and in the medullary
rays of the axis. Ordinary large solitary and clustered crystals have not
been observed in this Order.
The hairy covering consists of clothing hairs only. In most cases they
are ordinary unicellular trichomes, usually with strongly thickened walls;
 MONIMIACEAE 79 1 -

in Kibara the subsidiary ce!1s_cof the hairs are also considerably thickened.
The unicellular hairs of Doryphora, Daphnandra and Laurelia show a tendency
to become tufted, two or three of them being frequently sunk in the epidermis
side by side. Mollinediarepanda, R. et P. andll.f. tri/lora, Tul. exhibit transitions
between ordinary unicellular trichomes and short two-armed unicellular hairs
with thick or thin walls, such as occur in other species of M ollinedia, in M althaea
j and Atherosperma. Large tufted hairs, often seated on prominent elevations
of the leaf-surface, are found in Peumus and Siparuna; their sclerenchymatous
cells are concrescent d the base so as to produce a fbot, which is often deeply
sunk in the tissue of thf leaf. Structures closely allied to the tufted hairs are
the stellate hairs, in which the ray--cellS" are- spread out. in a plane parallel to
the surface of the organ; these hairs have been observed in certain species
of Siparuna, in Itlonimia, Palmeria and Hortonia. Pelt ate hairs have been
recorded L"1 species of Monimia, Siparuna and Conuleum, and according to
O. Bachmann and Hobein they r~re distinguished by the fact that their ray-
cells have thin walls are only partially concrescent, and do not meet at the
f

centre of the shield, but along a median line (Fig. 171).

l 3· STRUCTURE OF THE AXIS. I have examined the structure of the wood
in Monimia ovalifolia, P. Th., ]a,[ollinedia Selloi, A. DC., Hedycarya arborea,
Forst., Peumus Boldus, Mol., A therosperma moscha~
tum, Labil1., Doryphora Sassafras, Endl. and Daph-
nandra micrantha, Benth.; Hobein only dealt with
the tissue of the medullary rays. According to him
the medullary rays of the wood are broad in an
Monimieae, being visible to the naked eye, or at
any rate with a lens, while in the Atherospermeae
they are narrow (from one to three cells in breadth).
The'diatneter of the vessels is small (not exceeding
'°36 mm.). The wall of one vessel in contact with
that of another sometimes bears scalariIorm
bordered pits, whilst in contact with parenchyma
it generally has very large elliptical simple pits, rIG. 171. Peltate hair of Si'parzma
. b d d' II I P B ld crillaio • •'\. DC.-After O. Bachmann.
WIth or ere PIts as we. n eumus 0 us I
a delicate ~piral thickening of the wall is also present. The perforations of the
vessels are generally exclusively sca1ariform, and often (e.g. in Atherosperma)
have very numerous bars; in M onimia ovalifolia, besides perforations with
I-IZ bars, there are others which are simple and have an elongated elliptical
outIL'le;. Peumus Boldus has for the most part simple circular or elliptical
perforations, those of the ~calariform type occurring only in the neighbourhood
of the primary wood. The prosenchymatous groundwork of the wood bears
simple pits in Monimia, Hedycar'}'a, Peumus and Daphnandra, small and often
indis tinct bordere'd pits in M ollinedia, and typical bordered pits in A therosperma
and Doryphora; septate wood-fibres with simple pits have been observed in
all species.
AccordiDg.to Hobein, the most important character of. the cortex. is the
composite and continuous sclerenchymatous ring in the pericycle. This ring
is partly composed (perhaps in' all cases, at any rate in the seVen species in
which I examined the structure of the wood) of hippocrepiform stone-cells,
in which the outer tangential walls are not sclerosed. The sclerotic ring is
wanting in Conuleum and in a large number of species of Siparuna, in which
only isolated groups of bast-fibres are present. The development of cork
takes place superficially, in a subepidermal position in Siparuna limoniodo1a,
A. DC., and beneath the hypoderm in Peumus Baldus, Mol. and Monimia
ovalifolia, P. Th. Tne primary cortex is not strongly developed, and consists,
in most casesJ of cells which have thin walls or are slightly collenchymatous;
 MONIMIACEAE

~n'L~uTelia laIge iiltercell_ul~ ~paces'?ccur:be~ween the cells;, The collench~~

lS.1morel strongly ~develope(bml.specles·'of' SIPa'u.na~' \. In imost, of, the.Jgenera
(except\M attiuua,tPeunius, ,Laurelja and 1Daphnandra) sclerosis of. the lprimary
corteXJ sets linr lat'l an, early: stage~ 'and in .. SiPa,una~ limoniodoTa, . A.,fDC~ and
S:negl~c~a;rA;iDC! ~~ !eads_ to the fo~~ation of a rin~ of stone-~ells. _"Acoording
to ;Hobem~ Jthe"secondary oast. contains J bast.;ftbres' m. Conuleum; and rod-cells
in' M onimia~"HedycaTya;,-Peumus and' Conuleum ; Moller also mentions the bccur~
rencelofrrod-cells indhkk'pieces' ofrthe :cortex, of[IAtherosperma moschatum,
LabilI: i' It;remains'lto,-mention the presence. of peculiar, tannin-sacs;',which
correspond dn "every 'respect ·With.'the ,well-known' sacs of ,SambucUs"nigta;
l.tliave"floticed,Jhese structures in St:paruna h'moniodara,rA. 'DC.;and'S.; g·uia-
ninsis, Aubl.:at.the margin of the pith; and' in the]bast:, l("~ T ' . ' LJ:' r;jj ;j.

"(,\ I ,.~. ...r f" I .,.--i~. , "".. ' H . '\ .~ ... \ ,l't'tt1; '-1 r: " ... ,,("' .....
.,~!~~at~~e : ~?~?my I'}?~.chs. P;, ,Flora 188 .. , 'P; '366'"et Seq'. arid .sep. ~opY. pp: '~3_:'5:-:-~01)er.,'
Rlndenanat,Il88a j pp. 99-1 o~-S~lereder, Holzstr.,'1885, 'pp; ·l26.:..7 •. . .:. . POisson. Hennecarlta, I'aris,
1885, 6: pp; and I .pL-O~iBachm8.nD; Schildh.,· Flora' 1886~Jsep •.'copy, p., 16Iand_Tab.~;riii.:"':"
Radlkofer, in 'Sit~~~Ber.,lM~cb. Akad. ,1886, p. 327 .----:Hobeio, ~Anat. Charakt.• d. M. etc., Engle~,
Bot. Jahrb., Bd. x, 1888, pp. 51-73.-Douliot, Perlderme, Ann. lie; nat., ser. 7, t. X,. 1889. p. 33•.-
~~~~e, i~ ~gler, Bot. J.ab~b,~ ~d~.~i, f~89i;·,);. '37"~d Chilen:.'H,~lig~., -.~rlDgsheim J~~b.~~Bd:
x~,_. !~9(,' p. B8.-[Pe!~JDs; ,.yPltr. 7:. 1<:'. ,~. M .. Fn",l"r, Hnt- TAhrh. Ha. xxv;~ !898, Heft '4-~.J,

',',LAURINEAE 1,
lL . ~ ~: ." . . . . ' .. , .. .•
I;-- KEVlEW OF/THE ANATOMICAL FEATURES. The Laurineae, like the allied
Monimiaceae, 'aie characterized bi the-"constant presence',jof oil-cellS:,'~ The
ni~dlag~~~~s _wpich,,~cur hi- m~ny, genera. of .Laurin~a~, t~~~gh' .ri~fin ·.:~l
tbe:specles;,0,,1- ~ tbe.!!e' genera, consb tu te ·a ._ye!y' fP~!a_cJ.e~lstic Je~ tl;1I'e}.not ;:.~01P.l ~
in ~~Monimiaceae.::: .,pther(jmpqrt~~t characte!s ,are.'las .f.olloWS:; -the_. ~ul?i
sidi3:!y~cell,s, ~f the< stomata are ~ placed :parallel·to"Jhe_, pqr:e.:.O!' the. gUa~?:~e~ J;
the smallevveins ~of\ the leaf. are yeJ;ti~allYJ ~ranscurrent ; 'l.the,tmedullary, rays
of theW:0od 'ru:e 'haft-ow; t4~r~jsi~~ tendencY·~to t~e)o~_ni_atiOD:l9!i~C3l.¥ifq~,~
p~rforatlOns, which, however,- ·never; -h3;ve ;Xery "n~etO~, .!>.¥,S ';:l,~·,~~~t~.!
!1th,~p~r~!lcl,t~l t.he .~L~!!s of. t~e)"y;e~e~ ...~~_3:r! 1.h9~dered I.or Sl~P~~, P!~, 1 Q.!~e_.g
of large sIZe; . the wood.:prosenchyma bears SImple ~Its and 1S,S9:m~!~~ ~~pt~~~;
ba:st~fibres may be~rather abund8:nt, or only present~ smaiJ I).Embe~~ 1Jl t~~ ~~~~n.;
dary ~ast ;.there,is ~tendencYJo_.!or.m a_cQ~posi~~ ap.d ~ont~uous sC?er~nc~~:,
tous rmg, [mcludmg stone~c~lls ~~lth\.U:,shaped,thickenmgs ; r .~he Aey.elop~e~~ &~
cork( is fsuperfi~ial; glan?ular hairs are absent,! and,,~i~p~~_, -unic~llul}l! h~~
areLt!le':only forms ofJ_tnchome.cp:resen!; }.9~a).a.t~qf;lun~~1:S s~crete!l ~o:~~!YI~
the.form of~mall crystals, which are f~s,Iform) aClcular,.9! <;If ~.t~~,~. s~~p'~'~o:~e
p3.!asi!ic .genus. Cassyth~ Jlike~ise J has. ; s~cre~ory'( cells, IIi~~J.!~¢~~~!!~!~.dJ ~~~}
aClc~arJ_crysta1s, ~ bUt.~!lS r'speclallyi' d1Stl~gUJ.s,h.~d •. bYJP~e~~g'~ a, ~ylep,~r!1.lg
devol~ 'of.,'medullary ·rays.l,o(The, ar!atoIDlcal \st~ctureLo~ Ith~ ,.m.ffiy!<)'u.~~ g~!l.c:r~
and) speCIes shows •little:. diyersity. 11The folloY'!Dg ..~p~q,!.!' f~atur~s i ~~y~e, ~~~~
under observation: development of hypoderm beneath the upper r~P~4~[~
off the .leaf I (species} ohA ydendron, JB eil~chm~edia, ',CrY1?!g~a,rylf. 1~~?r !W:!!~,,!!~la) ;
scleroslSTof lSolated,,~ells lori groups:of J~ells.!~n_Jh~ ;paJlS_C!.4e:t~ue! (~ ~p'~~e~~BJ
Ocotea) ; I<transversel orientation. ot st~l!lata ,on the.,ste!ll,Fo~ ~q-,ssyth,~:\ vl:t1.G T G,(
.aH<K2.,STRUCTURE oF.lmErLEAF:bcAdetailed investigatIon. of the leaf-structure
.. ....~....... I ' • • - .... _ . " . ... ~ ... l ............. J . . . . . . . . . ..

has: not .yet, been~ madet JJn -the few SpeCl~~ jwl!!~h I,~ayer ~x3:nul'!~d_1(L~,14T~~
1iabilis,fiL.,~ Ocotea, api/era,) ~arL, Litsea iNeesiana, JHemsley, ,!?~t:~~a graJi§.~i~,
Gaertm f:)' the~ leaf·structure is.,bifacial,: and:.the,J~~o~ata:.¥~ ~~YA9.!!nd!.~n
the lower su'rface.1:.,It is_ch~aracter!sticLof.th~ s!QIp.~t~tha~ !;hey:~x~.ac~~~p?-nl~9
f)u«m"l~ Dr..: r)1.1 , .. ".v , ...... ~ ,.., ,~~~ ,.~... . .,.; ...'1:; j b'(J(~ JO. A
t~Ja..l.::...We will follow the example"of Pax hy,dealing:,witb. t~~' ~u.s,H~",:andia ~ogethef.. ,~~th, ~~
~IGyrocarpus,. /1/igerar._.and~SpartJllant"eliuml· (of.Jhe Giioca~e) a.s· a separate Order,
fi: '
man dlaceae, r II .•
lO oWlng
th·e -L" n'un"neae.H , . u u · ' .~...,
<&- • L' :,}ii~"" Gl) ~/I .t.l J t.:'· ~;:} I r:,.( ;~11 111
• .
 ~'AURINEAE

by subsidiary cell~ne'on either 'side and .parallel to the pore; the subsidiary
cell~ )are noOeasy"to.l.t~cOgruze (Lit~~a!Nee~ia~;~P~rsea rgratissima), when : bo~h
guard.cells and SubsIdIary cells·:are~,depressed.~()· Cassytha,' also (C:\amerJcana,
Nees) possesses' 'stornata'-' with r subsidiary cells ',\paialle!l to )~the \: pore; tdt\', is
noteworthy~t however, 'thair.r:the stomatalJboth'don ,the!'stern:.( and:lIon ~the
rudimeiitarY·lt~aves 1:: lie'~transversely _to Jthe 110ngitudinalJ axis ~of1:the:lorgan:
In. many members 'oft the: Order [the) lower.' side:1oflfthedeaf,jhasla glaucous
bluish bloom' .'dueTto' 'an) excretion ['0£' WaX.-1If :AccorCling" to~:Pax~" hypodermr: is
formed on the upper.iside,iof ,fthe-leaflin"!C1yptocarya,'J:Aydentiron, 1Ravtmsara
an4.1n certain species ~of Beilsch~iedia: ;Accordingl '"to uMez,fsome 11 species bf
Ocotea(O, KJ.cmditolia, Mez, O. opiferti', Mare; O. Kunthiana,'Miz and O:'''Rusbyana,
Mez) show'the:followmg special ~peculiaIity~' individil31 cells ·ofltlie-palisade-
ti~u~ ,or 'small groups' of these' cells .'are -'transformedr mto. similarly: elongated
ston~-cells~, r ..1These ~ 'scl ere nch yina
t«?US)' elements:1 are' 1viSible' ~ to Il the .Inaked "eye
as .distinct i dots', 6n •theT§urface> of [the! leaf:1.bJ Mez: further states, that.l in .: other
species of the' same:genus.J(OcJtealalriijoliai'lMez,';Or-Bojo;llL B~jKr: q!!dispersa,
Mez and O.c·punctulata,IMez).the surface Of the leaf bears fine black dotsra;point
which"- requires reirivestigationT and,- 'explanation': (cork-\varts. ?). -; rI I was'iun~ble
to,obt!lin material ,for this; purpose. , IThe~veins 'of medium siZe arc.;~itically
transcurrent 'on"both sides by'means' of' more 'onless 1differentiated sclerenchy-
matous 'cells-Y (1)oth iIi the' 'species mentioIled~above '.and,ill specieS of~JP~ebe
and Cinnamomum, according/to Perrot);· ::!!t~,H ::.mI,kit· NUfi(_ 2..;;.;:.bJ I(!Om .{ll
IThe :secretory:elements'.found liri.·,tnetLauHneacjrequire I a:.more detailed
description: ',Jbey; ;are~ of: ~ fwo' ,kinds, ") ,viz;"' (I): 'secretory cells -fWith rf oily
contents,. 'and TJ (2) Jmucilage-~cells:' _:,' Bokorny's:-:investigatiorlS, ~stipplemented
by~ thos~;,of '~obein; ha v~)shown' that oil:_c~lls r'?ccur'/~m,~he deayes ,of; illlth~
plants' mV,e~Vgate~ (speCIes ,of Cryptocarya, ~:~tnlschm",ed",a, Dehatis..a;rAyden~
,dTon, Aiouea, 'E1'idiandra;.. A.Crodiclidium, ;Ctn1tamomum;""·1l-1achilus,t... Persea~
,AlseddaPhne~ t Phoebe, Ocotea,·Mespilodaphne,' Dic'jipeltium; N eCtcindra; Sassafras,
A ctinodaphrte:-,Dodecaaen1.a;'Litsea; Lindera,' ,Laurus ;-: andraccording to: recent
pbservations of my_'own' in' Umbellula'fia also); J i.These secretory cells are moreover
'preSent inJtne' parasitic gciJ.us 'Cassytha (C. anieric~na,:NeOes ),'. w liere~ Hacke~berg
and I observed them irithe pith' and leaf-tissue respectIvely: The secretory cells~
like the mt:cilage·cells to be.'dcscribed i below,ilfrequently:give rise, toutrans-:
parent dots1 in the leaf; they"are found both'jn-the palisade-and spongy,tissue;
and,in rare , caSes (UmbeliUlaria -ctilijornica;,:Nutt!):in:the lower. epidermis also;
They are generallyfspherical lin 'shape"'; those1:situated<L_1.othe'palisade?form
'the om y' exceptions,'since 1 they appear as enlarged' sac-like rcells -of:. t'1is tissu~~ <

The walls of the secretqry c~lls are suberized, and their contents'are",homo'::'
geneous and in -most cases yellowish. In'every case~ by..suitable treatment 'of the
sections, ~the presence l of·' the" wall of the, secretory' cell' rbe ldemonstrated;; can
the resin-lacunae, stated oy[Chatiti ;to 1oc'cur f in",.the'1Latirineael:do !not- exist.;
~In :the 'epl_dermar'secretory 'cellsltofl Umbellularia,.lthe'twhole:!,of> the/outer
,wall, does not"re~n th~~exterilal ;surface,':bbtI'oruy~a" circular'pDrtion,~arid[at
the centre of the latter there is a small dot-like:'arealof 'ltb.inner~<:e!l-\\"all.
IIi 'the,;branch [the secretory; 'cells' 'are found::iti, theTpithland.> cortex;. and
,a1sc;' iIi the' medullary lniys' and the parenchyma of; the' .:wo'cjd~j c!'In :"the Jpitli
'and primary cortex~they' areL.approximatcly' isodiametric, :while£thbse'jn, the
bast are elongated in the vertical direction. The occurrence of these'seCTI.!tory
,cells. in t!le_~ood "{),f. !h,~ ~'9o<?t._in: §~ss..ajr~~ ~~ !0I?-g l?~,~n! ~9~wE?, ~!~j., o? this
depends,· the, ,medlcmal.,; use of" this ,part. of ,the •. plant. , 'J Accor,ding tlto,.H6hnel,
Felix and Knoblauch,. hcweyer, Jhe_~ecrElto!y~lls..E~ also m the _present wooa.
~f the,.stern, i~" ~any ~e~b!~ ~f~!?~s2.t:.~~r(.~. r;~~~b!~~;~~~~~aWt~IL1t~;~ in
1
\. D,

Felix
. ,

~.lso
••

met"with' them
,

in
; , r'" ","'I' .. " • • ~~. ," (>"~ • "! 'n' '..... '"
fossil 'Wo6d~ belonging' to members of the·Uddneae . . . ' ' l d •
rt· i,,·.,."". . .~..- "u!
•Co" "",lJ ,J. '-V ".~'~..;;, ~.u.l __ ..... ~;..;JuJ· U.i -;Jc.,a..;.J-C,l.114 k~ CiiU:;;;"';,JijJa r;1um, ~
 LAURINEAE

the medullary rays or parenchyma of the wood in the stem of species of the
genera Cryptocarya, Beilschmiedia (Hufelandia), Aydendron, Aerodiclidium,
Cinnamomum, Persea, Oreodaphne, Dicypellium, N ectandra, Sassafras,
Litsea, Umbellularia and Laurus. I may add that occasionally the secre-
tory cells may even be observed in the medullary rays of the wood in
branches from herbarium-material (e.g. in Nectandra angusti/olia, Nees et
Mart.). Secretory elements of the second kind, i.e. the mucilage-cells, are
not so widely distributed as the oil-cells, for they only occur in certair~
species of certain genera. Bokomy met with them in the leaf in species
of the genera Beilschmiedia, Dehaasia, Aydendron, Machilus, .Persea, Alsec ..
daphne, Phoebe, Ocotea, Mespilodaphne, Neetandra, Sassafras, Actinodaphne,
Litsea and Lindera, whilst Radlkofer found them in ACfOdiclidium; ac-
cording to Ballier and my own observations, they also occur in the leaves of
Cassytha americana. With the exception of this 1ast species, in which the
entire mesophyll aboun ds in mucilage-cells with wide lumina, these cells generally
occur only in the palisade~tissue in the leaf. In the axis the mucilage~cells
are invariably (?) present in the form of cells, elongated in the vertical direction;
they occur chiefly in the bast (e. g. in cinnamon-barks, in Litsea Neesiana;
Hemsley and Persea gratissima, Gaertn. f.), more rarely in the primary cortex
also 1 (e.g. in Cassytha american.a, "Nees, where they were observed by Hacken-
berg, but inconectly interpreted, and in Persea gratissinta). The mucilage-cells
in most cases show distinct stratification of the mucilage and a very small
lumen ;' hence it may be concluded that the mucilage arises by metamorphosis
of the cell-wall. Before leav.ing the subject of the secretory organs we rilaY
point out that Hohnel's statement as to the occurrence of resin-canals in the
medullary rays of the wood in a species of Oreodaphne (Laurier de Montagne
from Martinique) is without doubt incorrect; and the same is to be said of
Plitt's statement regarding·the occurrence of oil-canals in the petiole.
Oxalate of lime occurs chiefly in the form of small acicular or spindle~shaped
crystals; crystals of somewhat larger 'size are rarc, being i'ound in the neigh-
bourhood of the vascular bundles (e.g. in the cortex of the branch of Endiandra
virens, F. v. Miill., in the cells of the sc1erenchyma-ring, which exhibit U-shaped
thickenings). The crystals first -mentioned occur both in the leaf and axis t
several being invariably found in the same cell. Clustered a~d ordinary
large solitary crystals are entirely a~sent. Pe~rot, it is true, mentions
the occurrence of clustered crystals ill the pnmary cortex of lVectandra
angusti/olia; his material, however, as shown by reinvestigation of the same
species, was incorrectly determined, and does not belong to any member of
the 'Laurineae.
The hairy covering is very uniform. Glandular hairs are not present, nor
are there any special forms of trichomes. All the hairs found in the Laurineae
are simple and unicellular, and are often sclerenchymatous.
According to Plitt's investigations the fibro·vascular system of the petiole
consists of an arc of vascular bundles (in species of Cinnamomum, Cryptocarya,
Lam'us, Persea and Tetrantkera).
3. STRUCTURE OF THE AXIS. The Laurineae on the whole also show uni-
form features in the structure of the axis. The genus Cassytha alone constitutes
an exception, and may therefore be described separately in the following
paragraph.
In CJ,ssyika amer£co.na, Nees the pith consists of lignified. but thin-walled cells;
it is surrounded by a xylem·ring. which shows two characteristic features: firstly,

1 Hahnel also observed spindle-shaped mucilage-cells in n wood, used in Southern China for
the prepan.tinD of an adhesive gum under the nam~ of Pan-FIl, and said to belong to the Lamineae;
the mucilage--cells in this case are fonnd either singly or in groups.
 LAURINEAE
the absence of medullary rays, and. secondly, the presence of slight ridges alternating
witlt furrows at the periphery of the xylem, the bundles of soft bast of the vascular
ring being inserted in the furrows. The inner portion of the xylem-ring, with the
exception of the small primary groups of tracheae, consists of vessels with very wide
lumina (diameter = ·12 mm. or more) and of wood· parenchyma, whilst the peripheral
portion is dense, and is composed of vessels with very smail lumina, wood-prosen-
chyma and wood-parenchyma. The vessels of Cassytha have simple perforations
throughout; their walls are provided with very large, circular. bordered pits. The
wood-prosenchyma bears simple pits. In the pericycle one finds isolated groups of
bast-fibres with white walls. Externally this is followed by the primary cortex
containing chlorophylll, mucilage-cells and abundant small acicular crystals, and
bounded by the epidermis, which has transversely placed stomata, a!ready described
~bove 2.

The features of the axis common to the ether members of the Order have
been summarized above in the general review of the anatomical characters.
The structure of the wood has been examined chi·~fly by Knoblauch, who
had abundant material 1.
The vessels have lumina of medium size (maximum diameter = ·03-
·075 mm.)~ usually fairly constant throughout the same annual ring. Sassafras
alone is distinguished by the fact that the vessels of the spring-wood have very
wide lumina, while those of the autumn-wood have narrow lumina. In m0:5t
of the species the perforations of the vessels are both simple and scalariform.
Scalariform perforations occurring alone were observed by Knoblauch in. Hufe-
landia penclula, Nees, but in no other cases. The scalariform p~rforations
mostly have few bars, the largest number observed being seventeen (in Crypto-
carya Wightiana, Thw.). Knohlauch also mentions a number of cases in
which he observed simple perforations only (species of Beilschmiedia, Acroiii-
clidium, Oreodaphne pro parte, Dicypellium, N ectandra, Actinodaphne, Litsea
pro parte, Umbellularia, Laurus pro parte). Further investigation is necessary
to determine whether scalarifonn perforations do not really occur in these
species in the neighbourhood of the primary wood, as is to be expected
from analogy (e.g; with Endiandra virens, F. v. Mull., which I investigated).
The structure' of the wall of the vessel in contact with parenchyma of the
medullary rays is very characteristic. It shows b'ordered pits with transitions
to simple pjts; the simple pits are sometimes large and elliptically elongated,
a group of them reminding one of a scalariform perforation. Spiral striation
of the wall.:; of the vessels is mentioned by J. Moller as occurring in 'species
of Tetranthera and Camphora. The medullary rays of the wood are mostly 1-3,
rarely as much as five cens in breadth, and the ray-cells vary in height. The
wood-parenchyma may be little developed or fairly abundant. Tangential
bands of wood-parenchyma ar.:: state.d by Knoblauch to occur in Beu-
schmiedia R_oxburghiana, Nees, Actinodaphne elegans, Thw. and Litsea dealbata,
Nees, and by Reiche in Cryptocarya Peumus, Nees. T'ile' wood-prosenchyma
has walls of varying thickness; in the young parts of the wood the gelatinous
layer is occ.asionally differentiated (e.g. in ~ersea inciica, Spreng. or Oreodaphne
bullala, Nees). The"pits of tne wood-prosenchyma are simple; t.he lumina are
septate by means of delicate transverse walls in species of Beilschmiedia, Ayden-
-_----_._. - -.~. -'--_~----,-

1 ihus CD.Jsytna. is not devoid of chlorophyll, as stated in mar.y works (Kerner, Pflanzenlc:bcn.
i, p. IS8, &c.). 1<-0:r details ~ Hackenberg, loco cit.
2 Aml.tnmical investigation affords an eesy means of distinguishing Cassytha from Cuscuta, whir.h
is similar in habit; in Cau)'t"a alone a pith is present, and a. mass of wood showing- a slightly stet·
late transverse section.
S Knobla.uch im-estigated the wood in the stem of thi.rty-three species of the following genera:
C?,pt()I;arya, Beilschmz'tdia (inel. HuftlanJia), Ayde1u1rtm, AcrodididillJll, Cinna~mum, Madzilus.
Plrsca, 01-eotiap.lllu, Dir)'pllium, Nect(J.~dra, Sa.l$aj,.as, ActilU1daphne, L#sea, Telranlht.,.", U",bel-
lulan·a. Lindera and Lc.ut'tu.
~~wu Zz
 L~URINEAE

d,on/,:A crodiCZidium,1 Cinnamomum; ,.,1achilusi .persea;: Oreoaaphne,tDicypellium!

Nectandfit'Sassa!1as"'Litsea''aiidiUnibellidaria:t~' -' '-;'1 :r" t: (!':·I()rr~ d:trll
'~:': r nle:';irUcture of ;the' pftn,has)jeen ExammecLm :delill ~o'n(·' in'l tne fcase('o~
.Cin·nlf,m.qrn.~m.. ~~~Yl.an.~~t{11jJ .~y;~¥..eIlioVic4; r." The,. pith ;~l ~this,_ ~fmL ~)~c~~~~
r

geneous.::, .It )15 i,difIeren tia ted ill to a -:penpheral 'portIon, ,composed (,of 1 small
active-cells with,thick'walls, andla central portion 'COnsistinglOf, emptYl.cellS
with thin walls., :I ," '~t-: " J ,S" I " • -~. " I ,_"I ,_,~r ~ :t!J0.[f~1I0'1Ib
, The development ()f cork takes'place at a: relatively late "stage-:' 'The,fcork
ames' su ~erticiall· eiilier'liii the"e 'idermis"'wliich usWill hg'a thiCk oufer wall
l

(Laii~us'~bilis,-t\lna;Cinnamonium dUZC;!.:Nees,'acco'rlmg 'to 'Moller); orjm 'the

outenllost cen~layer-of the'primary"cortex (Teirantliera iaponica~ Spreng.;a~4r~;
ing to }. E. Weiss). According to Moller, the cells of the cork either have
thiri' waIlS.l (species of l Persea,- Litsea: S assa!;(is), . "the !uuler 'tangeritialT walls or
are sClerosed' (spec~es '~ff9innamo1nunlrJD~cypellium,1:OreodaPhi(e; 'T~ai#.lie,a)'.
'11\ "Tetiiiitthera (iaponica,; ~according 'torWciss;:SCIerosea: ~or k -cellS": with rstrongly
thickened outer,walls alternate with thin-walled cells, which<are'elol1gated_in~the
radial·diiection·~nd[rapparently'-riot:-suberized!' .t.Stone-ceIIs"sol1ieti.nl;eS ra'ppear
iii thtfprimary'cortex of older branches.t .lThe struCture'of thcpericyc1e is very
dW-actensHc. \. 'The '-isola ted gT9UPS of jpriri:lai-y" bast-fibresl,'are Wfited tif' f9tm
&'closed oroillY'lqcally,interrupte.d ~sclerenchymatoU5' ri.'1g by~mea)is-of 's~one:
cells; some ~of·the latter have thin outer'tangenti<iFwal1s; 'buVsclerosed-inner
J

tangential and'radlaI.walls, so 'that they -exhibit' U-shaped l thickenin~cin' tr:ans:'

verse'sectionS' of the'br~lliGh~ A pericycllc'ring of this kind has'beeri obs~rved
b~ \Moller' in"s~cies ~ ~f 1Crypto~arya; (Beitschm,ie1ia~. Eitdli~keri",. A C1'!Jd~~lidium,
'(Jinnamomum~:' PersMi Phoebe,' Oco~ea, Agat/Wpnyllu'm; Dlcypelhum;, N ectand,a,
Sassatras~. Li!sea,·'Linde~a '~d Lau1'us. ;' 'also' by 'Perrot in DaPh~idi't!tn~ I aiid 'by
I

me'in Neetandra ~a'ngustifolia;"Nees' et Mart: (contrary-to the'statenient\~made

by:Perrot,. wboseL·m.aterihl certainly- 'did not belong ·to any. member:,of Ith~
'Laurineae) and Endiandra virens}'Fil.v~~Miill. The secondary bastih most Cases
~9ntains spindle-shaped=-ftbres o~[rotinded qitanrangUlar- 'sect~on~ and ,having
nruTOW lumina'; 'these 1j.bres·rarelY (Aerodiclidiuni;'Beilschmiedia, Ocotea) app~~r
in large 'nufnbers,\ formirig[ bundles;- usually there ·aie 'only··few. present. L

Stone:cells sometimes 'accoml'an"y .the ,:fibres', ~some' 'of ,them' being' thickene.a 'on
one side'only,_,like:t~ose of,the pericycle';' the stone cells arerdev~oped in'ex~
ceptional, abuiidaiice ,m;the -older-' cortex of certain' ~pedes" (species 1of' C,ypto-
caryii and Tetrant"~ra)~ l ' ;J _ ... .', ~J:to
•. I • ... , _.. ,
~. ._ _ . _.. . :I. .... ~ ..-t..I ~', "010' ... ~, JL'"
,....~.""
.... .."
'C~,-r"l'l
oJ 1. ,
.•• Liter,atlUt;,:, C,~tin,- ~D;at. .comJ>., ~~s,!egetaux, pI. iv-y, and G~d., (ol..,Ann. ;8C, ~~., ~::1,6,
t. 11, ,l8n;, pp~ :I 1 4-16, and pI. ,I 5.-Moller, Holzanat., 'DenkSt;hr:Wlener Akad. 1876. pp. 36-9 and
'3Z~ ct i;eq.-Zachaiias; in' Bot. 'Zeit. '1879, p: 6l6~:"":"Hohn'el, stkret!orisorg~, Sit%.I~Ber;'Wiener :Akad!,
Bd.~lxu.iv, Abt~ J~ 1881, P: !;96.-Bokomy, Durchs: P., :Flora. 1883, p_- 359".et' seq .. a.nd~.sep.. copy,
pp.,16-::32.-Hohnel, Gefassi. Holzer,mit Harig., BQt•• Zeit. 1882, pp. J6.;-6.~Moller.. RindenanaL,
~.~8l. pp:"IQ3-13.:-!elix, ~os3ile,Hol~er. Zeitschr...d.eutsch. geol. Ge~llsc:~ •• ,J~d. xxxv. ~~83, p.,61
-and Holzopale,'Mltt Jahrb. ung. geol. Anst., Bd. VII, 1883, PP.'27-8.~Mentovlch, Mark, Klau&en.
burg~ '1885 ;,!:-a~ti::in JUSt'11885'. i, 'P':..J78~:;-solerederi Holzstr:, 1885. <p'. 1~i7;':::"Plitt;· Bla'hmel.
Diss., Marbur~;' I 886/pp:'-35-6•..:::..Radlko{t>r,tDurcbs.. P., Sitz:-Ber;. Miinch~; llad.; 1886; P: 327.:!.
c

:Knobla.llch. Anat. d. BoIzes d. L., Flora,I888,pp. 339--too and Tab.·,vi~.-HobeiI!, Anat..Charakt.

d. L., Engler, Bot .• Jabrb., Rd. x, J889J'P" 74.-Hackt:nberg. ,Cassytlta 4f1Uricana;rVerh. naturhist.
Ver: d. preuss. 'Rheinlude etc. I889,-PP' 98~I38:':"":Me:i;'I ... americ;, ]ahrb. Berli.:ier Gart.;Rd. v,
1889, sep. copy, pp. i99-500.--Lalanne, Feuilles persist., Act. Soc. Linn. rle Bordeaux, ser. 5. t. iv,
1890, p. 66 and pI. iv.-J. E. Weiss, Korkbild" Dellkschr. Regensb. bot. Geseilseh. ~890. sep. oopy,
p: ~!i."-Pu. in N at~r1. :Pflanzenfam.,· iii:' 'Teil,~ Abt; ,~, r891 .' pp.; 106-7 .-:-1::e:rot;· Et. \ bist.' des ~ I
fbese. Lons-le-Saumer, 1891, 6:1 pp.-H. Hal her; G hed .. d. Con ,!,:olv., Engler" Bot. Jabrb.~Bd. XVI,
1893~ p; 540,~Reicht')'~hi!en. Holzptl., Prin~sheimj Jahrb":, Bd. >.xx~ (897, p:"S6 se'q:""JtMiiller, et
Lignum Aloes~ Phs.!'Dl8.Cent; Post I 897.~(With' regard' to- the cinnamon-barks' see" the "l:.barma·
c~n"He wnrllc ."if R~T"', Vaal TqC'hirMl. Ilt.c._ ,J:
 HEl{NANDIACEAE; ,-
~ .:: .. "J:T. "1 '.

According: to, Pax;. this Order ~is' composed, ofr the three I genera' GYTocarpus;
SpaTattanthelium'and.Illigeral(of., the' Gyrocarpeae); and the~genus HeTnandia!;
While the mode of dehiscence of, the an thers and other exomorphic 'fea tures show
the close conrie~(ion between this Order. and the Laurineae; thiS~ is 'also indicated
by~th~ anatomic~l'chara<:.terS~(A,The most important of~these¥are the possession
ot-secretory cellS\ and the: excretion 'of oxalate_of,Jimeimctnb".J.orm",of(sinall;
acicular. crystals~~ In these tW9 ailatomiGaI characters, and also in,the absence
\of in traxyJ.a:ry ;:soft l Deist, the genera of:. Gyrocarpeae ,~ffer ',Widely., from Jt.he
,Combretaceae, with .which they. ace associated. by BenthamJ'and,.HoOker, !lIld
others. The cystoliths of Gyrocarpus and SParattanthelium constitute a'special
ana'tomical feature. tliough~not;forind in'!all members of,the Order<~. ') " i.l
, l'he leaves of;the'Hernandiaceae i have bifacial structure, ,the .paliSade-.tiSsue
most1usually consisting of a·single layer. The, stomata. are' found only on the
lower side of' the le~if. In Gyrocarpus and
SParattanthelium they are surrounded
by a'number of ordinary epidermalI'cells;,while:in IUigera (excludingd.r;obtusa,
M~issn. 3) and H ernandia they~ are accompanied by subsidiary cells, arranged
pcitallelto the· pore; ',Hypoderm, consisting of from on.e.to two layers, has
been' observed 'on 'the upper: side of the le~f in Gyrocarpus ,'acuininatus, Meissn.,
G.. "asiaticus, -Willd.J vaT.' y, G.-rugosus, R:-]'Br., and Illigcra' appendiculata, BI.;
o~er_l species: of Gyro crirp us , and I ll,igera, and:' also: s~cies 'of, Sparattantheli'!lm
show'division bylmeans of horizontal walls:in certairi celIs:of the_upper epi·
dermis of the leaf. ,In the anomalous species Illigera ,.Dbtusa Jhe 10wer~ epidermal
celli of'the leaf, are produced· into coronate' papillae, Which are· connected ··with
one 'another by~ridges' of cellulose; The vascUlar ,hundles lof tlie veins -are
accompanied by sclerenchyma in the investigated species of SPa;attanthelium,
nligera~and,Hernimdia;. - _ '," . .' ' .
The hairy covering usually consists ~only of 'clothing' hairs: ~The latjer
'in Gyrocarpus~· Sparattanthelium and Illigera are ordinary unicellular scleren-
chymatou~ i!icliomes; aCcompanie~, bYlf,!1iSelln1,ar 1?ra~~~~ih~iis iI\,c~~!ain sI?:~.ies
of the~ three genera: II "i: In ,Hernandsa sonora; '- fa ther '-short,~lunicell\ilai hairs J Wl th
,wide 'luminal are' 'present in the floral region; the anomalous species Illige'ra
obtusa- has' drdiiiarY.Junicellular ,trichomes'. withdransitions·',to.,typically JWQ:
armed hairs, ;the: anns. being' of. equal ,length. 1 The glandular~hairs IlligeT(J of
(exc!."I ... obtusa)~consist~ ofl a~ short.,~ni.cellu.lar! stalkrand r a ,two:celled{ head,
,shaped likejthe teleutospore of ~'tP.uccsnta (Flg.1I7Z, ·A): , .J.: r 'I0~~::: 50:'1 11,' br~'
Oxalate of·,time, both in the axis and,leaf;·, islaImost.exclusive1y~excreted
in the form of small/acicular crystals ,(except in Illigera obtusa, where,Ol:dinary
large solitary crystals: occur, in tl!eineighbourhood'lof .t~e'\vascular.~bun_dles).
In ~the' leaf the ',acicular crystals! are found· both. in fthe ;..vascular.) bundles~arld
mesophyll,-and :als? in,the uI?per andtlo\,\'er epide~~, where~they are ab~dant
and are-'accomparued, by, qmte, small. octohedral sohtary'! crystals; Jhey",~~~:­
times :'o~ur.L eve~~in the~:g_uard:cells,Land in- !l!igera1j also~:in ,:the"headS:1 0f
the glandular ,harrs:; lBy Ahe, presence) oft cys~o1iths therJ~QI closely; t:elate:<)
genera Gyrocarpus \,land ;Sparattantheliuml,arer disting?ished :t.fromt t1;te,t ~tp.~r
members ·of Ithe Order.cj :.The cystoliths of ,the.lleaf) areJgenerally,'~confi.i::1ed"to
·the;integumental,·tissue; :they:are: rarely found' alsolip3,t~eJ_sQftLb~tjlQt:th_e
'vems, (SParattanthelium amazonllm,- Mart.');: pr ,in 'lthe l ,tissue. surround_ing £.t!t~
_ ~""T0 ... • - '"

l' j, ' . ' - ~ " . . ~ ,. _., c, h i . ... , ' ,o'-{ (TTl, -.1 (! ,'{01::.ri:JI.slJ
See ootnotes on pp. 3~3 and 70 '].. 'J ' -t- ......... "'f'- • ..,..
• " -,t ..... - ..,. _::J"'. ,
J Most ioiportant'literature ':' Solereder~ 1889;' itl addition to' the', sptCies of Gyrocarpeae; men-
tioned there, the structure of the axis and leaf was also investigated iIi Hermmdia sonora; L, 18 91,6
• IlHg-era oo/usa, Meissn. differs (rom the other species of J/Jigua not only in this respect but
also in the nature of the fruit and in other anatomical characters (see below), and should at auf rate
be excluded from 'the
genus llligera (Ct. 'Flora'Brit: Ind. ii. '1819; p. ~61).: "..,_" :1 1I.. ..J_'" ";.~ ~ 0
• NO" ~1\i _'\ I.a. or gn... I u" I~ ~.

ZZ2
708 HERNANDIACEAE

vascular bundles (Gyro carpus asiaticus, Willet.1., or even in the mesophyll

(G. rugosus, R. Br.). The L'1.tegumental cystoliths. occur in the hypoderm in
those species in which a hypoderm is present; in o'ther cases they lie in epi-
dermal cells the shape of which is adapted to that of the cystoliths; in the
latter case only a small portion of the epidermal cell reaches the surface of the
leaf, but it is to this part of the wall that the stalk of the cystolith is attached.
The form of the cystolith differs in the two genera. Gyrocarp'lfS has spherical
or ellipsoidal cystoliths, whilst those of Sparattanthelium (Fig. 172, B) are
branched, and usually have six arms, radiating from a common centre, their
arrangement coinciding with three intersecting axes. The cystoliths mayor
may not be calcified; their skeleton gives the cellulose-reaction. The calcified
cystoliths are generally visible even with the naked eye'as granular dots (Gyro-
carpus), or small striae or stars (SParattanthelium) on the surface of the leaf.
The cystoliths may a150 occur in the axis, e.g. in the primary cortex of C. asiaticus.
Secretory cells with oily contents are present in all the lllembers of the
Order, and in Hernandia 1 mucilage-cells occur as well. The secretory cell5
sometimes give rise to transparent dots in the leaf; they are found. in the
epidermis of the leaf (only in
certain species of SParattanthe-
limn, Illigera and Hernandia), in
B the palisade (only in I1Ugera and
Hernandia), or spongy tissues)
and in the ground-tissue of the
veins; in the axis the secretory
cells are found in the pith, bast
and primary cortex. Tne se-
cretory cells of the 1eaf are
spherical, with the exception of
those occurring in the palisade-
tissue, which have the form of
FIG. J7:1. A, Glandillar hairs of IlIijfof1"o C(J'7:adofnio,
Meisso. B, Transverse section through the up~r portion of the enlarged cells of this tissue.
lamina or the leaf of Spa".aJ14nthdium BotomdontfIJ, Mart.- The structure of the axis has
Ori~inal.
been examined in Gyrocarpus
asiaticus, Willd., Sparattanthelium T-upinambazum, Martius, Illigeta Khasiana,
Clarke, and H ernandia sonora, L. The forma tion of cork takes place super-
ficially, viz. in the outermost cell-layer of the primary cortex in Hernandia sonora,
and in the second layer of the cortex in Gyrocarpus asiaticus. In the two species
mentioned the cork consists of cells with wide lumina and thin walls. The outer
portion of the primary cortex is composed of collenchymatous tissue; in
SParattanthelium Tupinambazum the inner portion consists of parenchymatous
cells with wide lumina and lignified walls. In the four species mentioned above
the pericycle contains strongly developed bundles of various types of scleren·
chymatous fibres; only in Gyrocarpus are these fibres united to form a con-
tinuous sclerenchymatous ring by means of stone-cells with U-shaped thicken-
ings, as in the Laurineae. The bast requires no special mention. The v; ood
is soft, and consists of (a) vessels with fairly wide lumina, (b) narrow rnt:dullary
rays, and (c) wood-prosenchyma with relatively wide lumina, walls of no great
thICkness, and simple pits. Where the vessels abut on one another they have
relatively large bordered pits, and in contact with parenchyma they bear
transitions from bordered pits to large simple pits. The perforations of the
vessels are for the most part exclusively simple; in Hernandia some of them
are scalariform, but with few bars.

I This statement is foulldell on an .investigation of II. sOJJora; Bo.kotny only mentions resin-cells
as occurring in Hernaltdia.
 HERNANDIACEAE
Literature: Moller, Hobanat., ~kschr. Wiener Akad. 1876, pr. 41 and 337.-Bokomy,
Dnrchs. P., Flora I88.2, p. 359 et seq. and sep. copy, p. 16 et seq.-Solereder. Anat. u. Syst. d.
Combret., Bot. Centralbl. 1885, iii, p. 161 et seq., Holzstr., J885, pp. J21-9 and 227 and Blattspl'.
bei den GYl'ocarp., Engler, Bot. Jahrb., Rd. x, ItS89, pp. 511-20 and Tab. xiv.-Kohl, Kallts. etc.,
1889, p. I34.-P ax, in Nabirl. Ptlanzenfam., iii. Teil, Abt. 2, 1889, pp. 126-9. .

GOMORTEGACEAE.
Gomortega nitida, Ruiz et Pav., which is the only representative of this Order,
agrees with tlie Laurineae and Monimiaceae in possessin~ secretory cells, and
in the presence of a composite sclerenchymatous ring incluaing stone-cells with
horseshoe-like thickenings in the pericycle.
The leathery leaf is typically bifacial, the palisade-tissue consisting of
several layers, while the spongy tissue is lacunar. The epidermal cells on
both sides of the leaf have straight lateral walls. Beneath the upper epidermis
of the leaf a tHick-walled hypoderm is developed,. consisting of from one to two
layers~ the cells of which are polygonal in surface-view and considerably larger
than the epiderml.l cells. The stomata are found only on the lower side of the
leaf, and are accompanied on either side by a single subsidiary cell placed
parallel to the pore; a secondary division occasionally takes place in the
subsidiary cells in a direction at right angles to the pore. The vascular bundles
of the veins are provided with a strongly developed sheath of sclerenchyma.
The following statements may be· made regarding the structure of the
stem. The xylem consists of (a) wood-prosenchyma with bordered pits;
(b) narrow medullary rays, the cells of which are somewhat elongated in the
vertical and radial directions; (c) vessels with small lumina (maximum diameter
= '03 mm.), exclusively scalariform perforations (mostly having numerous bars),
and walls bearing relatively large simple pits in contact with parenchyma of
the medullary rays; and (d) a small amount of wood-parenchyma. The peri-
cycle contains a composite and fairly continuous ring of sclerenchyma, composed
of groups of bast-fibres, and of stone-cells sclerosed on all sides or on one side
only. According to Reiche, 'sclerenchymatous cells resembling idioblasts' occur
in the older portions of the secondary bast.
The secretory cells are filled with a yellow resinous secretion; their shape
is approximately spherical in the pith and primary cor.tex, and in the palisade'
and spongy tissues and the hypoderm of tIle leaf, while those in the soft bast are
elongated in the vertical direction. Oxalate of lime is only present in small
quantities; and occurs in the medullary rays of the bast, and in the neighbour-
hood of the veins of the leaf, in the form of small acicular or prismatic crystals,
a number of them being found in the same cell. Trichomes are absent.
Literature: Reiche, Comorlqra, Ber. dentsch. bot. Gesellsch. 1896, p. 229.-Harms, in NatUrl.
Pflanzenfam., Nachtr. u. Reg. zu Teil ii-iv, 1897, p. 173.

PROTEACEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. Existing investigations point
to the following anatomical characters as common to the Proteaceae: (a) in
the structure of the branch, the simple perforations of the vessels; the bor-
dered pits on the thick walls of the \\'ood-prosenchyma; the superficial develop-
ment of cork; and the occurrence of secondary hard bast; (b) in the struct;ure
of the leaf, the nature of the stomatal apparatus, the guard-cells being accom-
panied by subsidiary cells placed parallel to the pore. In most cases the wood
in transverse section shows vessels with relatively small 'lumina and arranged
tangentially; the vessels are embedded in tangential bands of wood-paren-
 710 PROTEACEAE

chyma, and the medullary rays are broad. Internal secretory receptacles,.
viz. secretory cavities with red or reddish-brown contents, have only been
observed in the genera A denanthos and Frankland1;a; in Franklandia (Fig.
173, C) the secretory space is traversed in a remarkable m:anner by a network
of narrow cells, which are inserted on the papillose epithelial cells. Gelatini-
zation of the epidermis of {he leaf has not been observed in any species. Oxalate
of lime is present in the form of clustered and solitary crystals. The hairy
coverin~ (Fig. 173, E-F) consists of simple trichomes composed of on~ or a few
cells; m the genera Grevillea and Hakea bicellular two-armed hairs occur;
short hairs with a terminal cell shaped like an ascus (glandub.r hairs ?) have.
been observed ~ Lamherh:a. An investj~ation of, th7 leaf prese.nts a large
number of speCIal anatomIcal features, VlZ. the blfacIal or centr~c structure
of the leaf; the numerous details connected with the differentiation of the
middle layer (see Fig. I73); the occurrence of mechanical cells of va.rious
shapes in the mesophyll in numerous members of the Order; the vertical
transcurrence of, the veins in some cases by mea~s of sderenchyma; the de-
velopment of hypodermal tissue (Banks,'a, Dryandra, Franklandia); the varied
position of the stomata, which either lie on a level with the epidermis or are
depressed; the occurrence of stomata in small pits on the leaf-surface (Banksia,
Dryandra).
Z. STRUCTURE OF rHE LEAF. In the leaves, which are mostly leathery
and narrow, the anatomical structure is the expression of an adaptation to
the dry season, which the plants have to withstand, more particularly those
growing in the subtropical regions of South Africa and New Holland. The
mesophyll of the leaves has been especially ,examined by.Bengt Jonsson, while
MoW, Strasburger, Tschirch and others have investigated the peculiar position
of the stomata in many species.
The leaf-structure is either bifacial or centric (Fig. 173, A-C). In the
latter case, which is quite general where the leaf is very narrow or acicular, an
envelope of one or more layers of palisade-cells sunounds a medullary tissue
with little or no chlorophyll; in some cases the medullary tissue probably serves
for water-storage, whilst in others (Franklandia fucifolia, R. Br.) it has thick
walls, and stores up starch. Mechanical cells frequently occur as special ele-
ments belonging to the mesophyll; according to Jonsson there ara essentially
three forms of these elements. The first are prop-cells, viz. sc1erenchymatous
elements, elongated like palisade, belonging to the palisade-parenchyma an~
having short, root-like branches at both ends (species of Adenantkos, Grevillea,
Hakea, Is opogon, Molloya, Petrophila, Roupala and Stenocarpus). In the
medullary tissue of the leaves of some species of /sopog(Fn (1. petropkiloides,
R. Br., I. cornigerus, Lind!. and I. spathulatus, R. Br.) these mechanical ele-
ments are replaced by stellately branched sc1erenchyma-cells with thin rays
resembling the arms of an Ophiurid (Fig, 173, B). In a third group of species,
only ordinary sclerenchyma-cells are present (species of Adenanthos, Bellendena,
Hakea Isopogon, Leucospermum, Nivenia, Sorocephalus·and Xylomelum); they
are usually unbranched, or in other cases more or less branched. In the
acicular leaves of I sopogon adenanthoides, Meissn. the whole of the medullary
tissue is converted into sclerenchyma. with the exception of a narrow peripheral
zone containing solitary crystals; finally, in the bifacialleaf of Grevillea Hiltiana,
F. v. Milll. the sclerenchymatous fibres accompanying the vascular bundles
branch off from the latter and traverse the palisade-tissue in a direction at right
angles to the surface of the leaf, ultimately spreading out in considerable num·
bers between the palisade-tissue and the upper epidermis. The vascular
bundles of the leaf are always accompanied by· sclerenchyma, but the latter
varies in amount. Sometimes, even in the smaller veins of flat leaves, it forms
vertically transcurrent plates reaching as far as the epidermis on either side
 PROTEACEAE 7II
(species of Banksia, Synaphea and DrYandra). In flat leaves the arrangement
of the vascular bundles follows the normal type; in very narrow or acicular
leaves having a considerable number of bundles there is either one larger bundle
at the centre of the medullary. tissue, while the remainder are situated near the
periphery of this tissue, or both large and small bundles are scattered irregu-
larly in it, or all or/them lie at the periphery. In .l/akea sulcata, R. Br.
the larger vascular, bundles of the leaf, which are arranged in a ring at the
margin of the pith~ resemble the bundles in the fiat leaves of certain species
of the genera Banksia" &c., in having strongly developed masses of sc1eren-
chyma opposite and external to their xylem-groups; these travers~ the
palisade-tissue: and reach as far as the epidermis. .
There is little to be said regarding special contents in the mesophyll.
Crystals of oxalate of lime appear to be, on the whole, of rather rare occurrence.
Among the different species figured by Jonssen, he only shows solitary crystals
in the peripheral part of the medulla of the leaf of I sopogon adenanthoides, and
in the epidermis of the leaf in H akea marginata j De Bary men tions the occurrence
of solitary crystals in the epidermis of the leaf of Hakea sa/igna; in a casual
examination of Adenanthos obovata, Labill. and Franklandia fuC'ifolia, R. Br.
I foun~ in the medullary tissue of the leaf, abundant clustered crystals of
varied structure, as well as comparatively small numbers of solitary crystals.
Moller states that, in investigating the I bark J of species belonging to the genera
Banksia, Hakea, Leucadendron and Leucospermum, he observed clustered
crystals in the primary cortex in Banksia and Leucadendron only; in all other
cases (even in the bast) he found no crystals.
Another pQint of special interest is Jonsson's statemen~ on the occurrence
of' glandel ' in the leaf of Franklandia tucifol£a and some species of Adenanihos
CA. apicu/ata, R. Er., A. barbigera, Lind!., A. obovata, Labill., A. sericea, Labill.).
I have investigated Franklandia lucitolia, R. Br. and Adenanthos obovata,
Labill. with reference to this point, and I am able to state that these glands
are secretory cavities, lined by a delicate one-layered epithelium, and .filled
with brown contents. The secretory cavities of Adenanthos obovata are fairly
large spherical spaces in the palisade-tissue. Those of Franklandia lucilolia
(Fig. I73, C) are far larger, and are even visible to the naked eye as pustules on
t.he peculiar, dichotomously divided leaves, which look more like branches;
a transverse section shows that the secretory/cavities not only traverse the
palisade-parenchyma, but also penetrate deeply into the medulla of the centric
leaf. Closer eX!lmlnation of the secretory cavities of Franklandia shows that
the s'ecretory space, which is filled with reddish~brown, crystalline~ doubly
refractive contents, is traversed by a network of narrow~ elongated, thin-walled
cells~ connected with papillose protrusions of the epi'thelium. These extremely
peculiar secretory cavities of Franklandia may be recommended for develop-
mental investigation in fresh material. In some respects they are probably
comparable to the well-known intramural glands of Psoralea, and the secretory
cavities discovered by Kopff in certain species of Lonchocarpus. A third feature
connected with the presence of special contents in the mesophyll is the occur-
rence of a sheath of cells surrounding the medulla and the vascular system in
the centric leaf of A ulax umbellata, R. Br.; the cells of this sheath are elongated
parallel to the surface of the leaf, and are filled with brown contents.
In describing the integumental tissue the subjects deserving more thorough
treatment are: the development of hypDderm, the stomata, and the trichomes.
There are, however, other features worth mentioning, and we will take these
first. Gelatinization of the epidermis of the leaf, contrary to what one would
expect, has not been observed in any member of the Order; silicification of the
walls of the epidermal cdls occasionally takes place (species of Hakea). The
outer wall of the epidermal ceBs is considerably thickened in most cases in this
71.2 PROTEACEAE

Order, and so is the cuticle. In some Proteaceae, at the line' of junction

of the cuticle and the cellulose portion of the outer wall, there.are, accord-
ing to Nageli and De Bary, blunt cuticular processes with intervening slits
into which the cellulose portion of the wall penetrates; in a surface-view
of the cell, these slits appear as striae with irregular radial arrangement
around a central point, or in elongated cells arranged about two excentric
points; sometimes also they branch and anastomose~ Tschirch mentions
the occurrence of a waxy covering on the surface of the leaf "in Protea
melli/era, In EmbothriU1n coccineum. Forst. the epidecnal cells are provided
with papillae, in the formation of which the external wall is principally
concerned. The hypoderm consists of one or more layers; according to
JDnsson's drawings the cells of this ·tissue q_re generally not much larger
than those of the epidermis, and in Dryand1'a floribunda, R. Br. they re-
semble fibres. Hypoderm is present in the bifacialleaves of numerous species
of Banksia and Dl'yandra, as well as i.'1 the centtic leaf of Franklandia futi/olia,
R. Br. (Fig. 173, C). With regard to the. stomata, the most important feature,
at any rate for systematic purposes, is that the pair of guard~cells, in the cases
more carefully examined by Mohl, Strasburger and De Bary (species of Gre-
villea, Hakea, LMtcadendron, .Zlfimetes, Persoonia and Protea, and also in Aden-
anthos obovtlta, Labill. and Lambertia multiflora, LindL, as I learnt from a casual
observation), are accompanied on either side by one or two subsidiary cells,
arranged parallel to the pore; these subSidiary tells are cut off secondarily from
the cells adjacent to the guard-cells or from tbe mother·cell of the stoma, as
Strasburger has shown in species of Grevillea and Hake«. It remains for future
investigation to determine whether this type of stoma occurs in all Proteaceae;
the demonstration of the subsidiary cellS is not always an entirely simple
matter, since the whole of the stomatal apparatus (i.e.· the guard·cells with the
subsidiary cells) is frequently depressed. The mode of insertion of the stomatal
apparatus in the epidermis has much less systematic importance. In the great
majority of species the guard·cells are situated at the same level as the adjacent
epidermal celis, or somewhat higher; in the latter case the guard·cells are
frequently distinguished by the marktd development (strong in Leucadendron
decorum, less so in Grevillea Hilliana) of their outer cuticular ridges, amounting
to the formation of a rampart of cuticle. In the species of Banksia and. Drya1tdra
(according to Mohl) and Lambertia (according to Engler 1) the pairs of guard-
cells have the same structure as that just described; in these cases several
stomata occur together in ampu]]iform depressions in the surface of the leaf,
the depressions being lined with delicate felted hairs. In other species (e.g.
Petrophila rigida, Protea melli/era, Roupala brasiliensis, A ulax umbellata, accord-
ing to ~schirch) the guard-r:el1s and epidermal cells lie at the same level and
both have their external walls considerably thickened and to an equal extent.;
consequently the cuticular- ridge is raised, and the vestibule correspondingly
elongated. Finally, in other spedes the guard·ce]]s wHh the subsidiary ceJJs
belonging to them are depressed to a greater or less depth below the level of the
epidermis of the leaf, and the epidermal cells, which adjoin the stomatal ap-
paratus. form the wall of a variously shaped outer respiratory cavity, the latter
being ampulliform (Franklandia fucijolia, Fig. 173, D), cylindrical (Stirlingia
teretifolia), funnel·shaped (species of Hakea, e. g. Hakea saligna) or double
funnel-shaped (Hakea cyclocafPa). Naturally the stomata may occur on both
sides of the leaf (principally in centric leaves). or they may be present on the
lower surface only. According to Mohl, they are mostly (WIth the exception of

I In Lamberlia. imrlllis. R. Ur. and L. lIlultijloyo, LinJl. I did not find this foenture; [lor Ji,l
JOnsson in the species of Lambert/a e~amined by him.
 PROTEACEAE
I

those occumng In small pits) placed parallel to one another, as in the Mono-
cotyledons, and at the same time paral1el to the length of the leaf.
The hairy covering consists as a rule of simple, unicellular hairs only,
though they vary in other respects. In most cases the hairs have thick walls
and narrow lumina, and are stiff; thin, wavy hairs, torming a dense felt, occur
in Ba1lks1'a and Dr.vandra; also in Grevillea Pinaster, Meissn., according to

FJG. 1{~3. Traos\'erse section of

the lea : A, of HtJ"e,~ is u/cala ,
R .. Br.: fl. of Isop0'f.0n petruphi-,
''!Iaes, R. Br.: C# 0 FrtJllkla"dia
jUcito/ia, R. Br. with a secretory
cavity. D, Transverse section
through the ~tomatal a.pIlliratus of
Frallldandti: fuct/olia. E, Tri-
ch~s of LamlMrlia l1unms.
R. Br. F, Two-annerl hair of Cre·
'Vil/ea [;ntaris, R. Br.-A:'H aftl'"r~
Jonsson, the remainder original.

Engler. The simple hairs, moreover, are not always unicellular; in Lambe~,ti£l.
inermis, R. Br. (Fig 173, E), for example, they consist of quite a short basal
cell, inserted on the epidermis, and a long terminal cell with thick walls and
narrow lumina. Hairs of a special form appear in the two genera Grevillea
and Hakea (according to Moller, Jonsson and Engler). They a.re bicellular,
two-armed hairs (Fig. 173, F), which are probably to be found in all species
of these two genera. at any rate on some part of the plant. They consjst of
 PROTEACEAE

a basal and a terminal cell; the former is often long, and sometimes almost
without a lumen owing to the strong thickening of the wall; the terminal cell
has equal arms, may be thick- or thin-walled~ and is pointed at both ends; the
middle of the cell is inserted on the basal cell, and its arrns either form a wide
angle or are directed upwards dichotomously at an angle of 60°-<)0°. Glandular
hairs appear to be absent; an investigation of living material is required to
determine whether the hairs, observed by me on the lower s.urface of the leaf in
Lambertia inermis, R. Br. (Fig. 173, E), and having an ascus-shaped termina1
ceU and a short basal cell sea ted on the epidermis, are external glands.
To conclude the section on the structure of the leaf a tabulated review of the
more special results obtained by Jonsson is added. Jonsson's types are retained
in this review; they are not systematic groups, and transitions between them occur;
as regards the species investigated the original work must be consulted.
I. Hakea-type. Leaf-structure centric; sclerenchym<l.tous rod-cells in the pali-
sade-tissue: species of Adenanthos. GywiUea. Hakea, Isopogo'n, Molloya, Petfophila,
Roupala and Stertocarpus•
.1I. lsopogon-type I. Leaf-structure centric; ophiurid-like spicular cells:
~pecles of I sopogon. .
III. I sopogon-type II. Leaf-structure centric; otdmary scIerenc.:hymatous
cells: species of Adenanthos, BeJlendena. Hakea, Isopogon, LeucospCJ'1num, /Vivenia.
Sorocephalus and Xylomelum. ,
IV. 'Pe-rsoonia-type. Leaf-structure centric; scletenchyma of the vascular
bundles not vertically transcurrent: species of A denanthos, A ulax, Conospermum,
Emb.!Jt"ri~m. Leucadendron, M,,'metes, Nit,em'a. Persoonia~ Petrophi/a, Protea. Serruria.
Spatat/a and Stirlingia. .'
V. Synaphea-type. Leaf-structure centric; sc1erertchyma of the vascular
bundles vertically transcurrent: species of Hakea and Synaphea.
VI. Banksia-type. Leaf-structure bifacial; sclerenchyma of the veins verti-
cally tra.nscurrent; nypoderm of 1-3 layers: species of Banksia and DryanJ..ra.
VII. Gr8IJiilea-type. Leaf-structure bifacial; sclerenchyma of the veins not
vertically transcurrent; no hypoderm: species of A nadcnia, Brabejum, Grevillea;
Helicia, Lambertt'a, Lomalia. Orz'tes.
VIII. Franklandia fuci/olia, R. Br.: Leaf-structure centric; hypoderm.
IX. Aulax umbeilata, R. Br.! Leaf..gtructure centric; between the meuulla of
the leaf a,nd the palisade-tissue a characteristic layer of cells with brown contents.
3. STRUCTURE OF TIlE AXIS. The structure of the wood has been examined
by me in representatives of aU the tribes \ and has also been in~tjgated by
Houlbert. The vessels have a maximum-diameter of '024-'05 mm., and in many
cases (in the species of Banksia, Gr&villea, Helicia and Lomatia examined by
me, and according to Houlbert in Banksia, Dryand1'a, Emboth,.ium, C,e'IJillca,
Guevina, Hakea, Knightia, Macadamia, Oritt.s, Roupala, Ste1'wcarpus and Xylo·
melum, but not in Brabejt£1n, Isopogon, Persoonia and Protea) they have a charac~
teristic arrangement in the tangential direction, being at the same time em~
bedded in zones of wood"parenchyma similarl1 situated. The perforations of
the vesselc; are exclusively simple. The vessel-wall is provided with bordered
pits where it is in contact with parenchyma. Spiral thickening of the walls. of
the vessels has been observed in Dryandra formosa, R. Br., Grevillea Baueri,
R, Br. and Persoonia acerOSil, Sieb. The medullary rays are usually broad; they
are narrow, 1-3 cells thick, only in the species of Franklandia, Persoonia and
Symphyonema investigated by me, The wood-prosenchyma is invariably
thick-walled, occasionally (Hakea s'ttaveolens, according to De Bary) provided
with a gelatinous layer, and always bears distinct, though sometimes small
bordered pits. On the inner side of the primary vessels groups of scleren-
chymatous fibres are developed in many cases (in all the species investigated
by me, excepting those of Symphyonema and Persoonia; see also Vesque, Ioc. cit.
and Baillon, Hist. d.,.pl., t. ii, p, 406).
I Species of Pro/ea, A delUlnth os, S)'llaphea, Conosjermum, .Frankltllldia, SYlllphJi01le1lla,
PeJ'soollia, Hdida, Grevillc(l, Lomiltia. Em/rol/lrium, Ballksr'a and Drj'andra.
 PROTEACEAE 715
Regarding the structure of the cortex our information is still scanty;
Moller's investigations extend to species of Banksia, Hakea, Leucadend'lon and
Leucospermum. The formation of cork takes place subepidermally (species of
Banksia, Grevillea and Hakea, according to Sanio, Molle~. Doullot and J. E.
Weiss). The differentiated cork consists of cells with thin walls and wide
lumina, or of somewhat thick-walled cells. The primary cortex sometimes
contains stone-celIs; in -the pericyde there are isolated groups of bast-fibres.
In Banksia, Le~~cospermum and Leucadend'lon the secondary bast is charac-
terized by the following features: the tangential arrangement of compact
bundles of hard bast, in addition to which there are smaller isolated groups
of hard bast and sclerenchymatous parenchyma with elements of varying
shape; the broad medullary rays, which become sclerosed independently of
the hard bast; and the sieve-tubes, the clements of which are devoid of sieve-
fields only for a short distance in the middle of their length.
Literature; .~ohl, Spaltoff. d. Pr., Verb. Leopold. Akad., Bd. viii, " Abt., 18 33, pp. 789-804
and Tab. lx-lxi, aDd Yermo Schr., 1845. p. 245 et seq.-Strasburger, Spaltoft'n., Pringsheim 'Ja.hrb.,
f..
Bd. v, 1866-7, pp. 3~8-,9. and Tab. xli.-Vesque, in Ann. sc. nat., ser. 6, t. ii, 18 75, 145.-Moller,
Hobanat., Denkschr. "\\-iener Akad. 1876, pp ..403-4 and 338 et seq.-De Bary, Verg . Anat., 1877.-
Areschoug, Blad. &nat., Minnesskr. Lund, 187!!> p. uS et seq.-Bengt Jonsson, Bidr. till kannedoDl.
om. blad. anat.. byggo. hos fr., Acta Univ. Lund, vol. XV, 1878-9, 49 pp. and 3 Tab.; see also
Jast 1880, i, pp. 1I3-14.-Tschirch, Assimilationsorg., Linnaea, Bd. 43, 1880-83, p. 139 et seq. and
Tab. H.-Moner, Rindenanat., 1883, pp. 119-304.--S01ereder, Holzstr., 1885, pp. 328-30.-0. Bach·
mann, Sehildh., Flora 1886,· sep. copy, p. I6.-Douliot, in Ann. se. nat., ser. 7, t. x, 1889, pp. 331-'.
-J. E. Weiss. Korkbild., Denkschr. Regensb. bot. GeseUsch. 1890, sep. copy, p. 5S.-Engler, in
NatiirJ. Pfianzenfam" iii. Teil, Abt. I (189....), pp. 12D-2.-Houlbert, Bois sec. dans les Apetales,
These. Paris, 1893, pp. 13-43 and pI. i-H.-Reiche, in Engler, ]ahrb., Bd. xxi, 1895, [. 37 and
Cbileo. Holzgew., Priogsheirn Jahrb., Bd. xxx, 1897, p. 92.-[Tassi, Le Proteacee, .Bul • del Lab.
ed Orto bot. della Univ. di Siena 1898, pp. 67- 1 34, 13 tnv.]

THYMELAEACEAE.
I. REVIEW OF THE ANATOMICAL FEATURES. This Order is characterized
in a most excellent manner by the structure of the stem. In all the genera, with
the single exception of Drapetes, there is intraxylary phloem 1, accompanied
by more os less abundant hard bast-fibres, and the external bast is likewise
distinguished by the presence of numerous baJt-fibres. The following features'
may also be described as general ana tomical characters of the Order: the super-
ficial development of the cork, the simple perforations of the vessels, wood-pros-
enchyma with bordered pits, narrow medullary rays in the wood, the absence
of external and internal glands, and the lack of a special type of stoma. Oxalate
of lime is deposited in very different forms. namely as ordinary solitary crystals,
styloids, clustered crystals and crystal-sand; these forms have no great
systematic importance. The trichomes are simple and unicellular; hairs of
a special form, viz. two-armed hairs, have only been observed in Daphnopsis.
Interxylary~_in addition to the intraxylary phloem is present in six genera,
namely Linostoma, Lophostoma, Synaptolepis, Aquilaria, Gyrinops and Gyri~
nopsis; it is given off internally by the cambium. The following are special
anatomical features, which are of value for specific or generic diagnosis: the
gelatinization of the epidermis of the leaf and stem, occurring in very many
cases; papillose differentiation of the epidermis of the leaf (species of Daphne) ;
the occurrence of stomata exclusive1y on the upper surface of the leaf (species
of Passerina); the enclosure of the individual stomata in receptacles form.ed
by the papillose elevation of the neighbouring cells (species of Edgeworfhia t
Enkleia, Lasiosiphon, Linostoma, Lophostoma and Synaptolepis); scleren ..
1 In Afjuilaria Agalloeha, Roxb. the internal groups of soft bast become f;l'ansformed by
>«onda.ry changes into inversely orienta.ted med\lUary va:>eular bundles (see below).
716 THYMELAEACEAE

chymatous fibres in the mesophyll (species of Daphne, Daphnopsis; Enkleia,

Lasiosiphon, Lophostoma, Peddiea, Stephanodaphne); finally, the occurrence or
absence of intraxylary phloem in the midrib of the leaf and in the petiole, and
also the epidermal or subepidermal origin of the cork (according to Van
Tieghem).
2. STRUCTURE OF THE LEAF. The structure of the leaf and axis is well
known from the investigations of Van Tieg!Iem and Supprian.
The, leaf· structure is bifacial in most cases. The palisade;-tissue gene a

rally consists of short cells; the spongy tissue has large or small in tercellular
spaces. Centric leafAstructure with development of palisade-tissue on both sides
of the leaf has been demonstrated in species of Pimelea, Thymelaea, Stellera
and Diarthron; a mesophyll composed of isodiametric cells has been found in
species of Drapetes. In certain species the mesophyll is traversed quite in·egu-
larly by sclerenchymato1!s fibres (Daphnopsis Guacacoa, Wright ed. Griseb.
according to Radlkofer; species of Enkleia, Daphne section Eriosolena, Lopho-
stoma, Peddiea and Stephanodaphne according Ito Van Tieghem; Daphne pen-
dula, Sm., D. Wallichii, Meissn., Lasiosiphon scandens, Endl., Peddiea Fischeri.
Engl., P. parvi/lora, Hook. f.~ and Stephanodaphne cremostachya, Baill.
according to Supprian).
In the leaf the cells of the epidermis are generally low; their lateral walls
are usually straight on the upper side of the leaf, undulated or straight on the
lower side. Daphnopsis Humboldtii, l\feissn., Drrlpetes DietJenbachii, Hook.
and Thymelaea hirsuta, Endl. possess epidermal cells of greater height; in
Dais cotinijolia, L. this is only the case above the median vein, while on either
side of the latter the epidermal cells graq.ually decrease in height. Epidermal
cells having their walls arched outwards are found in species of Linostoma;
papillose epidermal cells occur on the lower side of the leaf in Daphne' com-
posita and D. involucrata (Van Tieghem). The thickness of the outer waIl
I

varies in relation to climate and habitat. The cuticle is usually smooth; excre-
tion of wax is rare, and the amount of it is never considerable (tagetta, Pimelea). •
Gelatinization of the inner membranes of epidermal cells, on the other hand,
is very common. Mucilaginous epidermal cells in the leaf have been observed
by Radlkofer, Bokorny, A. W'1gner, Van Tieghem and Supprian in species
of the following genera: Arthrosolen, Chymococca, Cryptadenia. Dc.phne, Diar-
thron, Dicranolepis, Edgeworthia, Gnidia, Lachnaea. Lagetta, Lasiadenia, Lasio-
siphon, Leucosmia, Linodendron, Linostoma, Lophostoma, Ovidia, Passerina 1,
Peddiea, Phaleria, P£melea, Stellefa, Struthiola, Synaptolepis, Thymelaea, and
Wikstroemia 2; in some cases they give rise to transparent dots in the leaf.
The gelatinization of the internal membranes is often considerable; sometimes
a few unchanged cellulose lamellae still remain in the gelatinized membrane
(Arthrosolen gymnostachys, C. A. Mey. and A. somalensis, Fig. 174, A, according
to Supprian and Van Tieghem respectively). It may be added that, according
to Van Tieghem, cells with mucilaginous inner membranes sometimes occur
also in the ppidermis of the stem (Arthrosolen, Diarthron, species of Gnidia
:iection Phidia, species of Stellera section Dendrostellera, species of Thymelaea
section Lygia), or in the subepidermal layer of cells in the stem (species of
Phaleria). The structure of the epidermis of the leaf in Phaler£a coccinea, Baili.
and P. octandra, Baill. deserves special notice; in the first of these species
epidermal cells with their walls uniformly and strongly thickened occur scattered
a.mongst the ordinary epidermal cells on both sides of the leaf; in the second
I De Bary's statement regarding division of the epidermal cells by means of horizontal walls in

P. ericoi'des (p. 35) is erroneous, and referable to the gelatinization referred to above.
2 Van Tiegbem (lac. cit.) goes too far in ascribing generic valne to the gelatinizaticn, without
having examined a sufficient number of species; in general the occurrence of mucilaginous epidermal
cells is only a specific charncter.
 THYMELAEACEAE

species these cells are only present on the upper side of the leaf. The stomata
are generally found either on both surfaces or only on the lower. In Passerina
ericoides~ P. !ili/ormis and P. hirsuta, the leaves, which are adpressed to the
branch, bear stomata on the upper side only; the tissues of the leaf have cor-
responding positions~ the palisade-tissue being situated on the lower (external),
and the spongy tissue on the upper side (Caruel). The stomata, as far as
I have ascertained, possess no special subsidiary cells (Daphne La2treoia, Aqui-
{aria Agal/ocha). They may either lie at the same level as the epidermis, or
may be depressed, or rarely (Passerina) somewhat raised. Peculiar stomata
(Fig. 174, B-C), viz. such as occur singly at the base of flask-shaped receptacles,
of which the wall is formed by the elongated cells (six to ten in number) adjoin-
ing the guard-cells, are present in Enkleia malaccensis, Griff. (according to
Van Tieghem)~ Linodendron (accorcling to Radlkofer), Linostoma decandr2lnt,

FIG. '7~' A, M ucllaginous epi,iennal cell~ in the leaf of A rllll'~.

SO/~IJ som(l/~lIsis. II-C, Stomata enclosed In receptacles: II III
D transverse section' c surface·view. D, Trans\'erse section of tbf'
If'af of lJicranoiepis 'RenthamiallQ with styloids.-A and 0 aft~r
Van Tieghem, B--{; after Supprian.

Wall., Lasios i phon scandens, End!., Lophostoma

caiophylloides, Meissn., Synaptolepis Kirkii,
Olivo and Edgeworthia Gardner":, Meissn. (ac-
cording to Supprian). I
The vascular bundles of the vems of the
leaf are invariably accompanied by bast-fibres
(Supprian). Bundles of hard bast occur at the
margin of the leaf in species of Daphnopsis,
Dicranolepis, Passerina and Synaptolepis.
Neither internal secretory organs nor
external glands are present. The hairy
covering in all cases consists of unicellular trichomes, according to Supp:rian.
A special form of these is constituted by the two-armed hairs, which. according
to Radlkofer, occur only in Daphnopsis Guacacoa, Wright, while in D. cuneata,
Radlk. none but ordinary unicellular trichomes are present; in D. angusti/olia,
Wright _e~:l..Griseb.t however, the trichomes of the floral region show a tendency
to the formation of two-armed hairs.
Oxalate of lime occurs in the form of ordinary rhombohedral solitary
crystals, prismatic crystals of more or less pronounced styloid-like shape
(Fig. I74, D), clustered crystals, and typical crystal-sand. No very great
systematic value should be attached to the various forms of excretion in this
Order, which is contrary to the results obtained in other Orders; this conclu-
sion agrees with the statements of Gilg and Supprian, but differs from those of
Van Tieghem; I convinced myself of this point during an investigation (under-
taken some years ago, but not published) of the structure of the axis in most
of the genera of this Order.
I t is not a rare occurrence for ordinary soli tary crystals to be connected with
 THYMELAEACEAE
typical styloids by transitions in the same species. Clustered crystals, styloids and
solitary crystals occur embedded in crystal-sand. Several of these forms of crystals,
even the most characteristic types, viz. styloids and crystal-sand, are sometimes
found within the limits of a genus in closely related species, or even side by side in
the same species. Under these circumstances the employment of the features pre-
sented by the crystals for systematic purposes is rendered very difficult. For the
actual determination of their systematic value far more extensive investigations
than have hitherto been carried out would be necessary; such work would repay
none but the monographer, who employs endomorphic to supplement exomorphic
characters when distinguishing different genera and species. For the purpose of
this book the following table may suffice; it combines Van Tieghem's statements
with the results of my own investigation, referred to above. In order to avoid false
conclusions when usmg this tabulated review: it must be pointed out that several
forms of crystals sometimes occur side by side in the same species:
I. Clustered crystals have been observed in species of Pimelea, Schoenobiblos.
Daphne, Ovidia, Dirca, Daphnopsis, Lasiadenia, DaiI4 Edgeworthia, A t'throsolen,
GorxlaJtiaj FU'lJife'l'a, Lagetta, Cryptadenia, Struthiola, Gn£d~'a, Lasz"osz-phon, L~'nostoma,
Lophoslcma, Enkleia, Synaptolepis, Stephanodaphne; Peddiea, Leucosmia, Phaleria,
Pseudais.
II. Ordinary" solitary crystals in species of Schoenobiblos, Difca, Daphnopsis,
Lasiadem'a, Da.is.,...A'Ytn,pso/en, GaodalNa, Lagetta, Last'osz·pl&on .. Peddt'ea, Phalen'a.
III. Columnar crystals and typical styloids with intermediate forms in species of
Pime/ea, Daphne, Ovidia, Dirca, Daphnapsis, Lasiadenia, Dais, A rlhrosolen, Goodallia,
Funilera, Lagetta, Cryptadenia, Struthiola, Gnidia. Lasiosiphon, Linostoma, Lopho-
stoma. D~'c1'anolePis, Lt"nodendron. Stepkanodapkne; Pedd~"ea, Leucosmia, Phalel'ia,
Pseudais; Aquilaria, Gyrinops, Gyrinopsis.
IV. Bundles of narrow acicular or prismatic crystals in species of Dirca and
Pime/ea.
V. Crystal-sand in species of Ovidia, Daz's, FaSS8YZ'na, Chymococca, Funz'/81'a,
Cryptadenia, Lachnaea, Gnidia, Synaptolepis: Gjlrinops (namely Lachnolepis mo/uc-
cana, Miq.).
VI. Crystal-sand, enclosing an ordinary solitary crystal or a prismatic crysta1,
in species of Sc.hoenobiblos, Daphnopsis. Dais, Lagetta, Phaleria, Pseudais.
VII. Crystal-sand, enclosing clustered crystals, in species of Ovidia and Phaleria.
VIII. According to Van Tieghem and Supprian, no crystals are present in species
of Daphne, Thymelaea, Wiksiroemia, StelJera, Diarthron, Drapetes.
Before leaving this subject it may be added that in the tissues of young
organs of Daphne Laureola, L. peculiar sphaerocrystalline bodie.appear on
the addition of alcohol; they have been investigated by Hansen and Baccarini.
3. STRUCTURE OF THE AXIS. The most important feature is the intra·
xylary phloem, which has been demonstrated by Petersen, Solereder, Supprian,
Van Tieghem and Gilg in all the genera with the exception of Drapetes. Supprian
and Van Tieghem between them have made investigations with regard to this
feature in species of almost all the genera of Thymelaeaceae given by Durand,
as well as Linodendron, Lophostoma and Enkleia; the only genera in which
they had no material were Schoenobiblos and GoodaUia. For these two genera
also I am able to record the occurrence, of internal soft bast, having myself
examined Schoenobiblos daphnoides, Mart. (Herb. Monac.) and Goodallia guia-
nensis, Benth. (Herb. DC.); this tissue is also present in the new genus
Englerodendron, Gilg. A few remarks may be made on the exceptional case
of Drapetes; the species (of which D. Dieffenbach":":, D. erico":des, D. Lyallii and
D. muscosus have been investigated) have a moss-like habit; on the evidence
of exomorphic characters Drapetes is a true member of the Thymelaeaceae,
in spite of the absence of intraxylary phloem and of other anatomical charac-
ters, which are otherwise of general occurrence in this Order, and will be referred
to later. This is not the place for entering into theoretical speCUlations as to
how far the absence of these anatomical features in -Drapetes may be explained
by mode of life and habit.
The intraxylary soft bast is developed in varied abundance. Very fre·
 THYMELAEACEAE

quently bast-fibres are present at the inner margin of this tissue, and they may
also be embedded in it; in the latter case the bast-fibres often have a similar
distribution to those in the outer bast. Not uncommonly an increase of the
intraxylary phloem takes place by means of a cambial ring appearing at the
inner margin of the xylem-ring, e.g. in species of Daphne and Aquilaria.
In Aquilaria Agallocha, according to Van Tieghem, this cambium not only
produces phloem internally, but also woody tissue externally, so that in
this plant we have secondary formation of inversely orientated medullary
vascular bundles.
The intraxylary phloem, unlike the same tissue in other Orders in which it occurs.
is to bejound in the petiole and the midrib of the leaf only in a certain proportion
of the genera (Lamounette and Van Tieghem). According to Van Tieghem. the
presence or absence of this tissue in the petiole and midrib has system~tic value
for genera and sections of genera, as the following review shows.
Internal 50ft bast (' tubes peridesmiques ') is not present, according to- Van
Tieghem, in the vascular system of the petiole and midrib of the leaf in the foHowing
genera: Pimelea sections Eupimelea, Thecanthes and Gymnococca, Schoenob~·blos,
Daphne (excl. section Eriosolena), Ovidia, Dirca, Thymelaea, Dais, Stellera, A'I'thro-
solen, Dia'l'thron. Passerina, Chymococca. Cryptadenia, Lachnaea, Drapetes, Strutht"ola,
Gnidia, Lasiosiphon, Linostoma. Dicranolepis .. Pseudais, Peildiea. Internal phloem
(' tubes peridesmiques') is present, on the other hand, in' the following genera
and sections respectively: Daphne section E,.iosolena, Daphnopsis, Lasiadenia,
Edgeworthia, Wikstroemia, Funitera, Lagetta, Lophostoma. Enkleia, Synaptolepis.
Stephanodaphne, Linodendron; Leucosmia, Phaleria " Aquilaria, Gyrinops. Gyrin-
opsis.
The pith consists of lignified and unlignified cells. Stone-cells are found
in the species of Drapetes, Lasiosiphon, Li1tostoma, Lophostoma, P£melea, Steller a
and Synaptolepis.
The structure of the wood has been examined in almost all the genera of
the Order by Supprian, and by me in the investigations mentioned above.
On this subject -the following statem'ents may be cited. The vessels in the
secondary wood generally form groups of variable size, or rarely (Thymelaea
villosa, Endl.) radial rows. Their maximum diameter varies between ·02 and
·07 mm. They have exclusively simple perforations, and there are/bordered
pits on their walls, even on those in ,contact with parenchyma. Spiral thickening
of the walls of the vessels has been observed in species of Arthrosolen, Dais,
Daphne, Daphnopsis, pirca, Lachnaea, Ovidia, Passerina, Pimelea, Stellera and
Wikstroemia. The wood-parenchyma is usually scantily developed, but is
somewhat more abundant, and takes the form of tangential bands .in Lagetta
lintearia, Lam., Dirca palustris, L. and Pimelea incana, R. Br., according to
Houlbert. The wood-prosenchyma in most cases has wide lumina and bor-
dered pits. The borders of the pits may be small or large, but even when small
'they are distinct in section. The following constitute exceptions as regards
the pitting of the wood-prosenchyma: Diartkron vesiculosum, C. A. Mey., in
which the wood-prosenchyma may be described as having simple pits, Edge~
worthia chrysanthat'Lindl. witli wood-fibres bearing simple (roundish) and
bordered pits, and Lasiadenia rupestris, Benth. with short prosenchymatous
cells having wide lumina and relatively thin walls, which are provided with
simple' roundish pits. The medullary rays consist of from one to two rows of
~ells, rarely as many as four.
Interxylary phloem has been observed in six genera, namely Linostoma,
Lophostoma, Synaptolepis, Aquilaria 1 (inel. Aquilariella and Lachnolepis),
Gyrinops and Gyrinopsis. This feature was demonstrated by me (1885 and
- - - - - - - _ . ----- --_-
The inlcrxylary phloem of Aqmwria is wrongly interpreted by Moller (1876).
720 THYMELAEACEAE

1890) m Linostoma decandrmn, \Vall. \ Lophostoma calophylloides, Meissn.~

L. ovatum, Mei~sn., Aquilaria Agallocha, Roxb., A. grandi/lorum, Benth.,
A. malaccensis, Lam., A. microcarpa, Baill., A. 0phispermum, Poir. (?), Gyrinops
WaUa, Gaertn. and Gyrinopsis Cumingiana, Decne., by Van Tieghem in Synap-
tolepis Kirkii, Oliv., Aquilaria Beccariana, V. T., Aqltilariella borneensis, V. T.
and Lachnolepis moluccana, 1\fiq. The origin of this tissue has been investigated
in Linostoma, Aquilarz'a, Gyrinops and Gyrinopsis, and in these it is produced
internally by the cambium (Thouvenin, Van Tieghem). It is worthy of remark
that in some of the species mentioned (in Aquilaria Agallocha, A. malaccensis,
and A. microcarpa, Gyrinops Walta; and also in Lophostoma calophylloides,
according to Van Tieghern) bast-fibres occur jn the jslands of soft bast ill the
wood; this is doubtless connected with the abundant development of the hard
bast in this Order, for islands of soft bast in the wood do not require any me-
chanical elements. 'I.

In the structure of the cortex the development of the cork is speCially

characteristic. The cork-cambium arises either in the epide~mis ~tseif, or in
the outermost cell-layer of the primary cortex. Van Tieghem ascribes con-
siderable systematic importance to these two modes of origin, as is evident from
the following summary; it remains to be determined whether this view is really
correct. Tbe cork-cells are. mostly fiat, rarefy (Lachnaea and Cryptaden£a,
according to Van Tieghem) they have rather wide lumina. They h~\Ve uni-
formly thickened membranes.
The development of cork in the epidermis has been observed by Van Tieghem
in the genera Daphne (incl. section Eriosolena), Lasiadenia, Dais, EdgewortMa,
Wikstroemia. Stellera (section Chamaestetlera), Arthrosolen laxus (= Rhytidos olen ,
Van Tiegh.), Lagetta. Drapetes pro parte, Gnidia,.pro parte (Gnidiopsis, Van Tiegh.).
Linostoma, Lophostoma, Enkleia, Dicranolepis,; Synaptolepis, Linodend1'On.. Le~,­
cosmia, Pha/eria, Pseudais .. Aguilaria pro parte, Gyrinops, Gyrinopsis.
The following ha.ve subepidermal formation of cork, according to Van Tieghem :
Pinzelea, Ovidia, Dirca. Thymelaea, Daphnopsis, Stellera (section Dendrostelle'l'a),
Arth'l'osolen, Diarthron, Passerina. Chymotocca, Funifera, Cr'Y'ptadenia, Lachnaea,
Drapetes PJ'O parle (Daphnobryon eYiccides), Styuthiola, Gnidia pro parte, Lasiosiphon.
Stephanodaphne,' Peddiea; Aquilaria pro parte (Aquilariella. Van Tiegh.).
The primary cortex contains strongly developed collenchymatous tissue
in Dil'ca (Supprian), palisade-tissue in species of Gnid£a and Thymelaea (Van
Tieghem). In a transverse section the groups of bast belonging to the vascular
bundles usually become narrower outwards in the form of a wedge, as in the
Iime~ the primary medullary rays between them being correspondingly widened.
Primary hard bast occurs in all, secondary hard bast in almost all cases (the
only exception known being Drapttes). The bast-fibres are generally developed
in abundance, and are visible even to the naked eye on breaking a young branch.
for they then project as silky fibres at the broken surface. Their arrange-
ment in the secondary bast varies. They are either scattered, or form den-
dritic figures in the transverse section of the branch, or they give rise to
a concentric stratification into hard and soft bast. As regards the structure of
the bast-fibres, it may be mentioned that the degree of thickening of the walls
varies, the walls may be lignified or unlignified, frequently no pits are present.
and septation of the lumina never occurs. Peculiar swellings in the bast-
fibres have been observed by Wiesner (Rohstoffe) in Lasiosipho1t speciosus~
Decne., and by Supprian in Daphnopsis Bonplandii, l\{eissn.) while the fibres
were found by Supprian to have an undulated course inlPeddiea Fischeri, Engl.

I Lt'nostgma scandem, Kurz., :l species which has often heen transferred (Syn.: Enl.:/da malac·
remis, Griff., Last'osiplulIl scandens, Endl. &c.), does not possesS' these islands of soft bast, at least
not in branches from herbarium-material.
 THYMELAEACEAE 721
Appendix: On the anomalous genera Octolepis and Gonystylus.
Gonystylus has been split into three genera (Gonystylus,Asc" rum and Am.yxa)
by Van Tieghem, but this subdivision is unwarranted. The two genera Octolepis
and Gonystylus agree with the Thymelaeaceae in the structure of the bast, the
superficial development of cork, the occurrence of simple, unicellular hairs, and
the absence of glandular hairs, but are essentially distinguished from them
by the absence of the intraxylary phloem and the presence of cells containing
mucilage in the parenchymatous tissues of the leaf and axis. The genus
Gonystylus is especially characterized by secretory cavities, which are apparently
lysigenous, and sometimes give rise to transparent dots in the leaf.
The genus Gonystylus, in which I was the first to observe secretory cavities and
the absence of internal soft bast, I.as recently been raised to the rank of a separate
Order, the Gonystylaceae, and placed near the Tiliaceae. Radlkofer and Van
Tieghem have likewise investigated it. Van Tieghem also had the opportunity of
-examining Octolepis Casearia, Olivo Of Gonystylus he had the following material
at his disposal: G. Miquelianus, Teysm. et Binn., G. affinis, Radlk. (5yn.: G. Becca-
f'ianus, Van Tiegh.), G. borneensis (Syn.: Asclerum boyneense, Van Tiegh.) and
G. Pluyicornis, Radlk. (Syn.: Amyxa kutcinensis, Van Tiegh.).
The following statements may be added regarding the two genera, taking
Gonystylus first. The leaf-structure is bifacial. Stomata are only found on the
lower side of the leaf; in G. affinis they are surrounded by a rosette of somewhat
smaller epidermal cells. The epidermis of the leathery leaves of G. Aliquelianus
and G. affinis has a remarkable structure; its cells are elongated towards the meso-
phyll in the form of conical proliferations; some of these cells are sclerosed ontall
sides, the remainder have mucilaginous inner walls. In G. Dorneensis the mucila-
ginous epidermal cells alone are present; in G. pluricornls both mucilaginous and
sclerosed epidermal cells are ausent. Secretory cavities and mucilage-cells are found
in the mcsophyll of all the species. Oxalate of lime is excreted in the form of clustered
crystals, and, in the axis, also in the form of solitary crystals according to Moller.
As regards the structure of the wood, it may be mentioned that the perforations of
the vessels are exclusively simple, and that the wood-prosenchyma, which has wide
lumina, is provided with small bordered pits. The development of periderm takes
place superficially, in the epidermis or subepidermally (Van Tieghem). In the axis
the secretory cavities are found in the primary cortex, the mucilage-cells in the pith
and primary cortex. I
Octolepis;possesses mucilage-cells only which are found in the pith and primary
-cortex, as well as in the mesophyll, and has no secretory cavities. The cork-cambium
arises in the epidermis of the branch in this plant. Oxalate of lime is found in
the form of clustered crystals. /'
Literature: Caruel, Foglie della i'asserilla hirsuta, Nuov. Giom. bot., vol. i, 1869, pp. 19-4-5
(there cited: Pasquale, Eterofilla, Diss., Napoli, 1 867).-Radlkofer, Monogr. Serjania, 1875, p. 103.
-Moller, Holzanat., Denkscbr. Wiener Akad. 1876, pp. 39-42 and 335 et seq.-De Bary, Vergl.
Anat., 18n.-Mol!er, Rindenanat., 1882, pp. II~-16.-Petel'sen, in Engler, Bot. Jahrb., Ed. iii, 1882,
pp. 5 64-:;.-Bokorny, Durchs. P., Flora 1882, p. 359 and sep. copy, p. 16.-Radlkofer, in Sitz.-
Ber. Miinch. Akad. 1884, p. 487 et ~q.-Solerede-r, Holzstr., 1885, pp. 230-3.-Radlkofer, Dnrchs.
Punkte auf Blattern Sitz.·Ber. Munch. Akad. 1886, pp. 328-30.-Baccarini, Sferocristalli, Malpighia t
vol. ii, 1888--1), pp. 13-r:;.-Lamounette, Liber iuterne, Ann. sc. nat., sch. 7, t. xi, 18 9 0 , pp. 274-5.-
Solereder, in Ber. deut"eJ,_ bot. Gesellsch. 1890, p. (98) note.-Cohn, La./ietta linttaria, Jahresber.
schles. Gesellsch: f. vaterl. Kultur 1892, Bot: Sekt., p. 65.-Thouvenin, Stnlct. des Aquilal la, Joum.
de Bot., t. vi, 1892, pp. 2I2-15.-A. Wagner, in Sitz.·Ber. Wiener Akad., Bd. ci, Abt. I, 1892,
p. 51 5.- Van Tieghem, Strnct. des Afjllilana t Joum. de Bot. 1892, pp. 2 17-19; Struct. et aft des
Th. etc., Ann. sc. nat., ser. 7, t. xvii, 1893, pp. 185-294 and Tab. ix; see also Bull. Soc. bot. de France
1893. pp. 65-78.-Houlbert, Bois sec. dans Ies Apetales, These, Paris, 1893, pp. 83-9I.-Supprian,
Britr. z. Kenntn. d. Th. etc., Diss., Berlin, 1894, 52 pp. and I Tab., also Eugler, Bot. Jahrb., lXl.
xviii.-Gilg, Verwandtschaftsverh. d. Thymelaeales etc., Engler, Bot. Jahrb., Bd. xviii, 1894, pp.
::;,12-54, and in Natiirl. Pflanzenfam., iii. Teil t Abt. 6a, 1894, p. 2I7.-Gilg, in Natiirl. Pflanzenfam.,
Nachtr. u. Reg., Teile ii-iv, 1897, p. 23T.-Kuhla, Phelloderm, Bot. Centralbl. 1897, iii, p. 199.-
J. Mi.jller, Lignum Aloe:;, Phann. Post 1897.

501F.REDIUI 3 :\
 722

PENAEACEAE.
This small Order, which is generally placed in the neighbourhood of the
Thymelaeaceae, agrees anatomically with the latter in possessing intraxylary
soft 'bast, simple perforations in the vessels, and wood-prosenchyrna with
typical bordered pits, as well as in the lack of internal and external glands.
It is distinguished from the Thymelaeaceae by the absence of hard bast-fibres,
which are often developed in such abundance in that Order. The stomata,.
like those of the Thyrnelaeaceae, have no special subsidiary cells. Oxalate of
lime is excreted exclusively in the form of clustered crystals. The thick leaves
contain fibrous cells, which run irregularly in the mesophyll, and are of two
kinds, being ei~her sc1erenchyrnatous fibres having thick walls and narrow
lumina, and thus serving for mechanical purposes, or relatively thin:-walled
fibres, stiffened by means of a spiral band, and apparently constituting a system
for water~supply; one, at least, of these two forms of fibrous cells is present·
in every member of the Order which has been examined.
_The structure of the leaf and axis is known in detail from the investigations
of Van Tieghem and Supprian 1. The leaf. structure is either bifacial (species
of Glischrocolla, Endonemn, and Penaea, with a tendency to centric structure
in some cases), or centric with palisade-tissue on both sides (species of Sarcocolla,
Brachysiphon, Stytapterus). The palisade-tissue consists of short cells; the
spongy tissue is composed of small cells and is never very lacunar. The epi-
dermis of the leaf invariably consists of one layer, and the cells are frequently
filled with brown, tanniniferous contents. The outer wall of the epidermal cells.
is thick; gelatinization of the inner wall has not been observed in any species.
belonging to this Order. The cuticle shows ret~culate striation in some cases,.
e.g. Endonema Thunbergii, while in Sarcocolla fucata it has a granular structure
(according to my own observations). In Endonema retzioides the cuticle fonns
well-marked external pegs, situated centrally to the surface of the outer walls of
the epidermal cells (Supprian). The stomata are either found on both surfaces
of the leaf (in species of Sarcocotla, Brachys£phon, Stylapterus, as well as in
Penaea Cneorum and P. acutitolia), or only on the lower surface (in most species
of Penaea, and in Gtischrocolla and Endonema). As I have been able to COD4
vince myself in Penaea myrtoides, Sarcocolla fucata and Endoneni'a Thunbergii,
the stomata are not accompanied by any special subsidiary cells, but are sur-
rounded by a varying number of ordinary epidermal cells. We must not omit to
mention the peculiar peg-like processes which project into the cavities of the cells
adjoining the stomata in Sarcocolla fuca/a, and especially inPenaea fflyrto-ides(Fig.
175, A) ; they spring from the vertical cell-walls bordering on the uppermost
portion of the respiratory cavity; their function has not been determined. The
occurrence in the mesophyll of the two forms of fibrous cells (Fig. 175, B),
referred to above, is very characteristic of the Order. Both forms are branched
and run irregularly through the mesophyU. On reaching the upper and lower
epidermis the sclerenchymatous fibres very frequently spread out in con-
siderable numbers, parallel to the surface of the leaf, between the epidermis
and palisade-parenchyma. The spirally thickened fibrous cells, on the other
hand, mostly become enlarged and terminate in contact with the epidermis.

1 The following is an enttmeration of the species investigated by Van Tieghem, the nomenclature
being that of DC. Prodr.: Penaea acutifolia, P. CneOrrll1l. P. 11fflcronata, P. tII)'rloides, P. DVala;
Sty/ajterns fruticu/osus; Say(ocolla formosa, s. fuca/a, s. squamosa .. Bracnysiphon acutus, B.
imhricatus, B. speciosus,. Eut/mema rel::.ioidts, E. Tlzunbe'"ffii " Gtiscn"ocolla Lessertiana. The
species examineu by Supprian are included in this list. I have myself made a casual examination
of the structure of the wood in all the species just cited with the exception of S. squamosa, and also
in Pmaea myrtijolia, Sly/apterus oar6a/lls and S. e,.irot"dcs, Sar,1)colla ",;"or and Brachysiplltm
I'ricaejo/ius.
 PENAEACEAE

Van Tieghem sta tes that both kinds of fibrous cells occur in the species of Penaea,
Sarcocolla and Endonema, those with spiral thickening alone are found in
Glischrocolla and Stylapterus. and the sclerenchymatous type alone in Brachy-
siphon. The only crystalline elements occurring in the leaf are clustered
crystals. Trichomes are very rare in the Order; such as are present being
short, simple, unicellular hairs (stem of Penaea mucronata).
In the structure of the axis, the first feature to be described is the intra-
xylary phloem, which I was the first tQ.demonstrate in this Order (in 1885). None
of the species are without .intraxr'lary phloem, which, together with the pith,
generally forms a mass of tissue 0 rhombic shape in a transverse section of the
branch. Whilst the internal phloem in the allied Order Thymelaeaceae is
almost always provided with hard bast-fibres, these elements are not present
in the Penaeaceae. Only in rare cases (Penaea mucronata, L. {3 microphyUa,
Eckl.) are a few sclerenchymatous rod·cells developed at its inner margin. In
all cases the structure of the wood shows the characters which I previously
indicated as markmg the Penaeaceae, though the material then examined was
very limited. The medullary rays are invariably narrow (1-2 seriate),· and

FIG. 17$. A Surface-\'iew of the epidermis on the lower side of the leaf of Pmaea mY"/lJidu L. [, seen
from within. D,transverse section through the leaf of Penaea ".tterona/a.-A Original, D after Van Tieghem.

consist of cells, which are sometimes considerably elongated in the vertical

direction. The vessels have small lumina (maximum diameter = '022-'04 mni.),
and are either isolated or arranged in groups. The perforations are exclusively
simple, and the walls of the vessels bear bordered pits, even where they are in
contact with parenchyma. The wood-parenchyma is scantily developed. The
wood.prosenchyma has typical bordered pits on its walls, which are usually rather
thick. The cells of the pith have, for the mo~t part, unlignified walls, and are
frequently collenchymatous; scattered stone· cells in the pith are not uncommon.
The primary cortex usually shows collenchymatous differentiation, and, like
the pit~t now '.ln~ then includes stone-cells. !he inner liI?it of the primary
cortex IS sometImes (Penaea mucronata, accordmg to Van Tleghem) formed 'by
a large-celled endodermis provided with Caspary's dots. Bast~fibres are not
present in,the pericyc1e nor, in the secondary bast. Rod·cells are sometimes
developed in place of them, e.g. in the pericycle of Glischrocolla Lesserliana or
in the secondary bast of Brachysiphon acutus. The bast, w~ich is often collen-
chymatolls, generally includes chambered fibres with clustered crystals. The
development of cork takes place in the pericycle I in Penaea mucronata, according
to Van Tieghem's and my own observations. The cork is generally composed of
two kinds of cells, which sometimes form alternating layers, viz. cells with
thin walls and wide lumina, and others with thickened walls and having wide
lumina or a tabular form. We may add that in Endonema retzioides and

1 Supprian's statement, that the formlltion of cork takes place in the subepidennallayer of cells
. in I all species,' ooe5 not agree with this.
3 A2
72 4 PENAEACEAE

E. Thunbergii small vascular bundles run in the four corners of the stem and
bend out into the leaf at the node (Van Tieghem).
Appendix: On the anomalous genus Geissoloma, Lind!.
The monotypic genus Geisso!oma (with G. marginatum, Kth.), placed by Bentham
and Hooker at the end of the Penaeaceae as a genus anomaluffi, is probably best
regarded as a separate Order in accordance with the views expressed by Sonder and
A. de candolle, since it is essentially distinguished from the Penaeaceae by its endo-
morphic as well as its exomorphic characters. Thus Geissolomf4 has no bicollateral
vascular bundles, but has scalariform perforations with many bars in the vesseis, and
a composite and continuous sclerenchymatou5 ring in the peri cycle of the axis. In
the pitting of the wood-prosenchyma, which is bordered, this genus agrees with the
Penaeaceae, as also in the lack of external and internal glands. On the other hand
the sclerenchyma-fibres present in the Penaeaceae are entirely absent in the leaf of
Geissoloma. Oxalate of lime is excreted in the form of solitary and clustered crystals.
The trichomes, which occur on the stem, are simple and unicellular.
Van Tieghem and Supprian investigated the leaf and axis of the genus Geissoloma;
I had only an opportumty of examining the axis. 'The following statements re-
garding the anatomy of this genus may be added to those given above. The leaf
has bifacial structure. The stomata are only found on the lower surface of the leaf.
Both the upper and lower epidermis include numerous mucilaginous cells. The
epidermal cells on the upper side of the leaf have strongly thickened outer walls and
well-marked cuticular pegs. The peculiar thickened and raised margin of the leaf is,
formed by higher epidermal cells, thickened on all sides. In the mesophyll clust.ered
crystals are present. according to Van Tieghem. The pith consists of lignified and
pitted cells. The medullary rays are 1-3 cells thick, and are formed by cells which are
more or less strongly elongated in the vertical direction. The vessels are scattered,
and have small lumina (mean diameter = ·o25:mm.); their walls for the most part
bear large simple pi ts where they are in contact with medullary ray-parenchyma. In
the primary cortex. stone-cells with wide lumina and clustered, and solitary crystals
have been observed; in the bast solitary crystals are found. The development of
cork takes place in the outermost cell-layer of the primary cortex; the cork-cells
have fairly wide lumina.
Literature: Solereder, Holzstr., 1885. p. 23J.-Van Tieghem, Sur les Thymelaeacees et l,es P.,
Ann. $(:. nat.,ser. 7, t. xvii, 1893, pp. 277-88.-Supprian, Kenntn. d. Thymelaeac. u. P., Diss.,
Berlin, 1894, pp. 25-9. also, in Engler, Bot. Jahrb., Bd. xviii.-Gilg, in Nattirl. PBan zenfam., iii.
Teil, Abt. 6 a, 1894, pp. 206 and 209.

ELAEAGNACEAE.
This Order, which is generally placed near the Thymelaeaceae, is charac-
terized by the following features: the absence of intraxylary soft bast i the
presence of simple perforations in the vessels, and of wood-prosenchyma bearing
border,=d pits; the superficial development of the cork; the occurrence of groups of
sclerenchymatous fibres in the pericyc1e and secondary bast; the lack of a special
type of stoma; the nature of the hairy covering, which· consists of pelt ate and
stellate hairs; and the excretion of oxalate of lime exclusively in the form of
small acicular crystals.
The structure of the leaf and axis has been examined in species of all three
genera. The leaves are either typically bifacial (Shepherdia canadensis, N utt.). or
tend towards centric structure, the lowest cell-layer of the spongy tissue being, in
the latter case, differentiated more or less like palisade (Hippophaif rhamnoides,
L., Elaeagnus angustijolia, L.). Hypoderm has been met with on"the upper side
of the leaf in Elaeagnus reflex a (Lalanne). The stomata. which are sWTounded
by a varying number of ordinary epidermal cells, are only found on the lower
surface of the leaf. The vascular bundles of the veins are not accompanied by
sc1erenchyilla. The covering of stellate and peltate hairs is found in all the
 ELAEAGNACEAE
species. The peltate hairs con'sist of a large number of narrow ray·cells, which
have relatively thin walls and wide lumina, and do not all reach the centre of the
shield; the ends of the ray~cells are free at the margin of the shield and taper to
a point. In Elaeagnus orientalis, L. and E. pungens, Thunb. an additional tuft of
rays is inserted centrally on the upper side of the shield (0. Bachmann). Thestalk
of the peltate and stellate hairs (these two types being connected by transitional
forms) is of very varied length. The long stalks of the hairs of Shepherdia
canadensis are multiseriate, and consist of rather long cells, replaced immediately
beneath the terminal portion of the hair by short cells with yellow walls; in
Hippophae' rhamnoides (Fig. 176) and Elaeagnus angustifolia, on the other hand,
the halrs have a short stalk, composed of a: single layer of short cells with thick,
yellow walls, and giving rise to a false lower scale, when seen in a surface~view of
the hair. The mode of excretion of oxalate of lime in this Order is very charac~
teristic. Clustered or solitary crystals have not been observed either in the
leaf or in the axis. In all cases only small acicular crystals are present; they
maybe long or short,in the latter case sometimes almost resembling crystal-sand;
several of them invariably occur in the same cell. In the leaf they are found
in the epidermis and mesophyll, while
in the axis they occur in the pith and
primary cortex.
Some further information regard-
ing the structure of the wood and
cortex may be added. The medul-
lary rays of the wood are from one
to two cells broad in H ippophal and
Shepherdia; as much as four cells in
breadth in Elaeagnus. The vessels
attain _diameters of '15 mm. (Elae-
agnus angustijolia) and '075 mm.
(Hippophac· rhamnoides), and are pro-
vided with bordered pits, even where
they are in contact with parenchyma
of the medullary rays. The wood~
parenchynu is scantily developed j
the wood-prosenchyma 1 is invariably seen }<;:~~f~io!.~totr~~i~~tf l1i/'l'(Jphae' rhall""oides, L.
covered with distinct bordered pits, ./
and has more or less thickened, walls. The cork arises in the epidermis,in
Elaeagnus angustifolia, and subepidermally in Hippophal rhamnoides (Douliot);
t it consists of somewhat flattened cells with wide lumina and thin walls. The
primary cortex is composed of loosely united cells, and even:in late stages
shows no signs of sclerosis. With regard to the structure of the b~t, Moiler
points out that in Hippophae· the bundles of secondary hard bast are Irregularly
arranged, while the medullary rays are narrow (one or two cells broad), and at
some points become sclerosed between the bundles of hard bast; this differs
from whafis- found~in Elaeagnus, where the hard bast, owing to its regular
arrangement, gives rise to stratification of the secondary bast, while the .me"'
du11ary rays are mostly broad and remain thin-walled.
In £taeagnus and H ippop"'ae~ according to Men tovich, the pith is hetero-
geneous, and consists of a peripheral portion, in which the cells are smaller,
have thicker walls, and are actIve, and a central cylindrical portion, in 'which
the cells contain no starch, and, after the lapse of a·year,.are,empty.

I In the .ood-prolencbyml. t)( H. ,.Aa1m,oider (\yhich here has bordereQ pits). Sanio observed

occalion .. l tbicktninfl of the walls, projecting into the cavities in the form of blunt cylindrical pegl, or
bums, extend in, trlllJl't'ersely from one ,ide to the other, as in the tracheideiO of Pinus si/wI/n·s, !ce.
 ELAEAGNACEAE'
Literature: Rauter; Trlchomgeb., Denkscbr. Wiener Akad .. J871, .ep. copy, pp. 7-S and Tab.
i-H.-Moller. Holzanat., Dmkscbr. Wiener,Akad. 1876, p. 4" and 338.-De Buy, VeTgl. Anat.,
J8n.-Moller. Rindenanat., 1882, pp. iI6--19.-Menlovich, MIUk., KlauSenburg, 1885; see'just
1885, i. p. 788.-So1ereder, Holzstr.; 1885, p. :13 ..... -0. Bacbmann, Schildb., Flora 1886, &tp. copy,
p. 17.-Douliot, in Ann. sc. ~at.. ser. 7. t.,x, 1889, p. 333.-Lalanne, Feuilles pedist., Actes Soc.
Linn. Bordeaux, sere 5, t. iv, 1890, p. 101 and PI. vi.-:Gilg, in Natiirl. Ptlan%enfam., iv. Teil. Abt.
3 a, 189-4, pp. s..6-7·

LORANTHACEAE.
1. REVIEW OF THE ANATOMICAL- FEATURES, 'According'to existing' inveSti':, ,
gat ions the following anatomical 'characters should, be kept" in view for the dia·
gnosis of the Order ~ the'type of stoma, the subsidiary cells being placed parallel
to the pore; the 'position of the stomata~ on the 'branch, the stomata 'being!'
arranged transversely or somewhat obliq'uely ·to .the longitudinal axis;, the'
simple perforations of the vessels; the~ isolated groups of bast-fibres' in ,the
pericycle; the absence of secondary 'hard bast';' the superficial developmept of
cork; the absence of glandular hairs: The medullary raYs of the wood vary in'
breadth; the wood-prosenchyma generally (exception Viscum) bears bordered
pits, Oxalate of lime is excreted in the form of ordinary solitary and clustered'
crystals, The hairy covering (Fig. 177. B-E) in Loranthus consists of (a)
uniseriate trichomes, in which each of the cells, with the exception of the'ter·
minal cell, is produced at its upper extremity into' a lateral protrusion, and (b)
candelabra-hairs, the tiers of which are unicellular; ·from analogy the stellate
and' pelt ate hairs, mentioned by Engler as occurring in species of. Loranthus,
probably have a unicellular ray-portion and shield respectively, The follo~ng.
special anatomical features have been shown,to occur in this Order: cork-\J,iai'ts'
on the lower side of'the leaf in Loranthus punctatus, R. etP.; branched for'
unbranched stone-cells~occurring in the tissue of the leaf and cortex in many'
members ot~he Order)aiid enclosing solitary crystals (Fig. I77, A); frequently,
swollen tracheae in Ithe terminations of the veins; silicified groups ofcells in\.the
leaf·tissue in certain species; lysigenous mucilage-canals in the pith and bast of
the axis in Nuytsia; group~\'of sclerenchymatuus fibres at the inner margin of'
th~ larger vascular bundles in some Loranthaceae; anomalous structure of ',the'
axis in Nuytsia floribunda, R. Br., consisting in the occurrence oP'soft bast-
tissue in the xylem. ' , .,
2. STRUCTURE OF THE LEAF. The leaf·structure -varies; it is sometimes
typically bifacial, e.g:in Loranthus punctatus, R. e't P., 50metim~~ tyPically centric
with palisade-tissue on both sides of the leaf, whilst the inesophylr of T'upeia
pubigera, Miq. or Loranthtts europaeus, Jacq. is composed. solely of isodiametric
cells. In V iscum album. L., the leaves of which are biennial, {tie mesoiJhyll
consists of isodiametric cells duririg the' first'year { in the second'year, however,
the layers of cells lying beneath the upper and lower epidermis become elqngited_
like a palisade, and a simultaneous increase in 'the number of chloroplasts takes
place. 50 that the leaf-structure becomes centric. The- stomata, as far' as IS
known ·(in :Arceuthobiwn, Antidaphne, LePidoc~ras, Nuytsia, Tupeia. Viscut!i)~'
. are accompanied by subsidiary cells, placed, parallel to the pore, only a. single
subsidiary cell being, as a rule, found on either-side of the ~pair of guard-cells t;
in rare cases there are severa}. . In leaves wHh cen'tric or'hornogeneous structure
the stomata occur on hoth surfaces, being present iii smaller numbers the upper' ori
side; in the bifacial leaves of Loranthus: punctatus' they are only 'found the on
lower side, but here they occur in large numbers: The arrangement of Hie'
stomata with reference to one another is usually irregular; in Nuytsia floribttnda
alone the stomata are placed transversely to the longitudinal axis of the leaf.
On the branches of Antidaphlu, Arceuthobium, Lepidoceras;' Lor~nthus, 'Nuytsia
and Viscu11J. the stomata exhibit a.-~imilar orientation, i.e.,·they-·a:re"'arranged
 LORANTHACEAE

transversely to the longitudinal axis of the branch (Chatin). . The outer walls
of the cells in the epidermis of the leaf ~ well as in that of the branch are dis-
tinguished by their considerable thickness in all the species investigated. I
have not obsetved gelatinization of the epidermis of th~ leaf in any case, not even
in Nuytsia {ldribunda,. where it is stated to occur by Van Tiegbem. The
mechanical system in .the veins of the leaf is developed in various ways. In
Lorantnus punctatus it consists of strongly developed arcs of sclcrenchymatous
fibres, whilst in V iscum album, Loranthus europaeus, Tupeia pubigera, &c., an
arc of collenchyma is associated with the vascular system of the larger veins;
tbis collenchyma may in some cases (Viscum album) subsequently become
sclerosed. The occurrence of enlarged terminal tracheides (e.g. in Viscum album,
Lora nth us europaeus, &c.), or of independent storage-tracheides in the mesophyU
(Loranthus punctatus) is very common. Similar water-reservoirs are sometimes
found in the form of special systems of tracheae, accqmpanying the vascular
bundles of the,veins, e.g. in Nuytsia floribunda and species of Loranthus belonging
to the section Gaiadendron, according to Van Tieghem. ~
Special features in the mesopb"yU are silicified gn;mps of cells' and groups of
stone-cells with crystalline inclusions; the latter have also been mef.with in the
cortex. The silicified groups of cells in Loranthus europaeus were erroneously
described by Marktanner-Tumeretscher as groups of mucilage cells 1, and their
true nature has only recently been recognized by Ravn; they consist of a vary-
ing number of cells, which when numerous are often united in the form of a sphere;
the contiguous walls of these cells are considerably thickened, stratified and
silicified, or they possess silicified protuberances rese~bling cystoliths, which
project convexly into the lumina of the cells and are sometimes stratified.
These silicified cell-groups occur principally in connexion with the vascular
bwulles, and have been observed in the following species, according to Ravn :
LOTanthus europaeus, Phoradendron emarginatum, Mart., P. rubrum, Griseb.,
Stluhyphyllum Fendleri, V. Tiegh., Tupeia antarctica, Ch. et Schl., Viscum album
and V. articulatum, Bunn. Groups of stone·cells appear to be widely distributed
in this Order, so far as one can judge from Van Tieghem's investigations, which
are n.ot yet c~mpleted. They are ~ound, for example, in the ~issue of/the leaf,
and m the pIth and cortex of Loranthus europaeus (Mentovlch, Matktanner-
Turoeretscr....er), and, according to my own observations, in the leaves of L./er·
rugineus, Roxb. (Fig. 177, A) and L.' punctatus; R. et P., and in the cortex of
Tupeia pubigera. These stone-cells are m9stly branched, and ultimately become
thickened to such an extent that only one or more peripheral portions of the
lumep remain; these are filled by solitary crystals. In the leaf, stone-celIs not
only accompany the vascular bundles of the veins, but, are also tOWld free in
the mesophyll (Loranthus lerrugineus).
The simplest forms of trichomes are uniseriate clothing hairs, in which the
upper ends of the cells are produced into lateral processes directed towards the
apex of the hair (Loranthus I'ulescens, DC., Fig_ 177~ B). Candelabra-hairs with
unicellular tiers are fOWld in the species 'of Loranthus belonging to the section
Cicldanthus (e.g. L. ,/errugiifezts, L. Scurulla, L. var. obtecta, Kurz, L. lepidotus,
Bl., &c.), and, according to Engler, also in Notothixos. These hairs (Fig. 177,
C-D) consist of a row of short, and somewhat ventricose, thin-walled cells, in
which the wall is protruded, in a horizontal plane at the middle of each cell, into
four or more rays; the horizontal walls, separating the individual tiers of the
trichome, are pitted. The trichomes found in the floral region of Loranthus
rUfescens. DC. (Fig. 177, E) have a structure similar to that of the candelabra-
---_. __
hairs. The pelt ate and stellate hairs, mentioned by Engler as occurring in
-
I Marktanner-Tnrneretscher's incorrect interpretation rea.ppears in Haberla.ndt's Phys. Pflanz~n.
anal., 2nd ed., 18¢. p. 353 and fig ...,6.
 LORANTHACEAE
species of Loranthus (the former, for example in L. Soyauxii, Engl., the latter
y

in L. taborensis, Engl.), may be assumed from analogy to possess a unicellular

shield and ray-portion respectively.
The only internal secretory organs found in this Order are mucilage-canals,
which occur in Nuytsia /loribunda, R. Br. They appear in the pith, and in
later stages in the bast; the pith contains a central mucilage-canal, and others

FIG. t1T. A, Portion of .a

trulvrrse ~ction of the leaf
of Ltlf"wnl}JU$ f,rrur-·"'II$.
RO'lb. a-a, Trichome.: Band
I £. L. -f~s, -DC.; c and
v, L.ft'J1'Uri",..,.-Orj~naJ.

which are peripheral, and situated opposite the larger vascular bundles. The
peripheral canals pass out in to the midrib of the leaf with the -vascular bundles,
retaining their position on the upper side of the latter; they do not appear to
undergo any further branching. but become _considerably swollen at certain
points. The mucilage-canals of Nuytsia are provided with an epithelium of
several layers, but are nevertheless of lysigenous origin; in the leaf the middle
 L(JRANTliACEAE

lamellae of the gelatinized cells, which form the canal, can still be clearly
recognized.
3. STRUCTURE OF THE AXIS. The axis, excepting that of Nuytsia, has
normal structure. The formation of cork generally (Loranthus, NU)'tsia, Tupeia)
takes place superficially, whilst in V iscum, as is well known, no cork is produced.
The primary cortex is frequently (Nuytsia, Viscum) developed as assimilatory
tissue, and;its outer portion contains palisade-parenchyma in Nuytsia. Stone-
cells are not uncommonly present in the pr_jmary cortex (Loranthus, Tupeia, Vis-
cum), their lumina in some cases containing solitary crystals. Isolated groups of
hard bast are found in the per~cycle in the species of Antidaphne, Arceuthobium,
Gaiadendron, Lepidoceras, Loranthus, Nuytsia, Tupeia and Vis cum, which have
been investigated. No secondary hard bast is formed. The structure of the
wood has been examined in Viscum album, L., Loranthus pentapetalus, Roxb.,
L. europaeus, Jacq., Tupeia pubigera, Miq. and Nuytsia floribunda, R. Br. The
breadth of the medullary rays varies even within the genus Loranthus, L. euro-
paeus having very broad rays, while those of L. pentapetalus are only from one
to three cells in breadth. Engler mentions me~ullary rays which are only 1-3
cells thick as occurring in most specif'~ of Loranthus, Phthirusa, Struthanthus,
.?ltrygilanthus and Lepidoceras, and rnultiseriate medullary rays as present in
certain species of Viscum and Lorantht-1: As in the case of the breadth of the
medullary rays, variation also occurs in the arrangement of the vessels, the size
of their lumina, and their abundance. In V iscum album, for example, the vessels
are very numerous, have small lumina (maximum diameter == '035 mm.), and
reSemble tracheids; in Loranthus europaetls spirally thickened, pitted vessels
with narrow lumina are accompanied by others with somewhat wider lumina
(diameter reaching '045 mm.). The perforations of the vessels are invariably
simple. \Vood-parenchyma is rather abundantly developed. The wood-prosen-
chyma bears bordered pits in Loranthus, Tupeia and Nuytsia, while in Viscum
it is scantily developed and sclerenchymatous, and has simple pits. -At the
inner margins of the bundles of primary xylem massive groups of sclerenchy-
matous fibres, resembling hard bast, sometimes occur (Viscum album; Anti-
daphne viscoidea, according to Chatin; Nuytsia floribunda, according to Van
Tieghem -; Phoradendron, according to Engler). /
The pith consists ofcollenchymatous cells in Viscum album and in the young
branches of Loranthus europaeus; the cells become lignified in older branches of
Loranthus (Mentovich). /'
It nowremains to describe (a) the curious appearance presented by the stem
of Arceuthobium Oxycedr; in. transverse section (quoting from Solms-Laubach's
account), the peculiarities being probably shared by other species of the genus, and
(6) the anomalollS structure of the axis of Nuytsia floribunda, discovered by Van
Tieghem. A transverse section through an internode of Arceuthobium Oxycea,;
shows four vascular bundles, which lie in pairs opposite one another, one pair
being more strongly developed than the other. This structure is connected
with the course of the bundles (see loco cit.). The anomalous structure of
Nuytsia consists -in'- the occurrence of bands of soft bast in the xylem. Van
Tieghem, who examined young branches in the living condition, came to the
conclusion that the interxylary phloem of Nuytsia originates in the same way
as in Strychnos. Older portions of the stem apparently show the anomaly of
successive rings of growth; this is illustrated by a specimen which lies before
me; it is 10 cm. in diameter, and was presented to our Museum by F. v. Muller.
The xylem is traversed by concentric layers of thin-walled tissue, which consists
chiefly of large parenchymatous cells and includes mucilage-canals, as well as
small groups of soft bast; the latter project convexly on the outer side, and
correspond to the individual segments of the xylem-rings, these being limited
laterally by broad, lignified medullary rays; these bands of thin-walled tissue
 LORANTHACEAE

are moreover traversed by the broad lignified rays, which continue their course
outwards in the next xylem-ring. These facts are not opposed to Van Tieghem's
observations,especiallyif one bears in mind that the anomalies of successive rings
of growth in the pericycle, and of interxylary phloem of the type formd in
Strychnos, are very closely related; for in the first case extinction and extra-
fascicular renewal of the cambium,take place simultaneously at all points, While
in the second case they only occur at certain points in the transverse,section.
Literatnre: [Unger, Parasit. Pfl .• Ann. Wiener Mus., Bd. ii, 1'8",,0, p. 3', Tab. iiLJ-Solms-
Laubach, Parasit. Ph., Pringsheim ]ahrb., Bel. vi, 1867-8, pp. 603 and 6J5 et seq. and Tab. xxxix.-
De Bary, Verg1. Anat., 1877.-Hesselbartb, Anat. d. Holzes, Diss., Leipzig, 1879, p. 7I.-:-Mentovich.
L.-~inden (Hungarian), Magyar Novenyt. Lapok. vii, 188~h n. 7.... , pp. 17-'3; ~bstr. ~ Bot.
Centralbl. 1883, ii, pp. 74-5. and Just IS8,3, i, pp. 18O-I.-:-Solereder, Holzstr., 1885, pp. 234-5.-
Mentovich, Mark, Klausenburg, 1885; abstr. in Just l 1885, i, p. 788.-Marktanner-Tumeretscher.
Anat. Ban uns. L., Sitz.-Ber. Wiener Akad. 1885, u pp. and I Tab.-Van Tieghem, Nuytsia et,
Gaiadmtirtm, Bull. Soc. bot. de France 1893. pp. 317-28 j Nalkcia d Triartizron., loco cit .• 8~,
pp. 61-,J ; Rae. des L~ loe. cit. 1894-, p. I:n et seq.; see alSo loco cit., J895.-Engler, in Natiirl.
P&nzenfam., iii. Teil, Abt. I (I~"'.), pp. 158-9 and 1.. africa.nae, Engler, Bot• .]ahrb., Ed. xx, 1895,
'.0.
p. 77 et seq., Tab. i-iii, especially ii.-Diels, Nenseeland, Engler, Bot. Jahrb., Bd. xxii, 1896, p.
-Kolpin Ravn, Cystolithes rudim. siliciJ1es chez qu. 'L., Bot. Tidss1crift, Ed. :Il, i, 'I~7, pp. 53-8.

SANTALACEAE.
1. REVIEW OF THE ANATOMICAL FEATURES. The following anatomIcal
features may be pointed out as important diagnostic characters of this Order
(excluding the genera Champereia, Grubbia and.Myzodendron, which will be
discussed separately in an appendix to the Order): the superficial origin of the
cork; the uniform structure of the wood, viz. (a) vessels with simple perforations
throughout, and with the walls usually bearing bordered pits, even where they
are in contact with parenchyma of the medullary rays, and (b) wood-prosen-
chyma bearing bordered pits; the almost exclusive occurrence of parallel sub-
sidiary cells on either side of the pair of guard~ells; the fact that the smaUer
veins of the leaf are invariably embedded; the absence of gelatinization ~ the
epidermis of the leaf, and of internal and external glands; the frequent ,occur-
rence of groups of silicified cells, and the excretion of oxalate of lime in the. form
of clustered and ordinary solitary crystals. These, which are the mo9J: important
features, are supplemented by a number of others, affording characters for
generic and specific diagnosis. Such are: the frequent occurrence of isolated
bundles of primary hard bast in the pericycle, whilst a composite and con-
tinuous ring of sclerenchyma is rarely (Henslowia, and also in Champereia and
Grubbia) present; the very rare development of secondary hard bast (Santalum,
Ompluuomeria); the scantiness of the hairy covering, both as regards the actual
occurrence of trichomes and the number of forms of hairs present, the hairs
being mostly simple and ~icellular, rarely (species of Buckleya and Pyrula,ia)
simple and uniseriate. or,still more ,rarely (Exocarpus latitolius, R. Br.) tufted;
the enlarged terminal tracheids occurring in many members of the Order, and
the rows of tracheids found in the leaf-tissue in some genera (Quinchar.zalium,
Thesium); the papillose differentiation of the epidermis of the leaf j the occur-
rence of hypoderm in the leaf; the transverse position of the stomata on the
leaf and axis, and so on.
A few facts concerning the three anomalous genera referred to above may
be mentioned here. The genus Myzodendron is distiuguished by the special
structure of its xylem and by the occurrence of medullary vascular bundles
(Fig. 178) in certain species j another distinctive feature is the absenc~ of
parallel subsidiary cells accompanying the stomata. Champereia is charac-
terized by the occurrence of double cystoliths (Fig. 179). Grubbia differs
essentially froin the structure typical of the Santalaceae in having scalariform
 SANTALACEAE 73 1
perforations with numerous bars in its vessels, and in the absence of the parallel
subsidiary cells which accompany the guard-cells in the'Santalaceae.
2. STRUcrURE OF THE LEAF. The structure of the leaf and axis in this Order
have recently been made the subject of a detailed investigation by Behm; his
statements!are supplementary to the illustrations in Chatin's Atlas I.
The structure of ~the leaf varies, and is only of importance for specific dia-
gnosis; it'may be either bifacial or centric; in many cases the mesophyllconsists
of rmiform, approximately isodiametric parenchymatous cells. The structural
features of the epidermis are of greater systematic value. Gelatinization of the
epidermis has not been observed in the leaf of any member of the Order; nor
are the lateral,walls of the epidermal cells ever strongly 'undulated. The outer'
wall is often considerably thickened, and the cuticle is not uncommonly striated.
In Cervantesia tomeniosa, Ruiz; et Pay. the upper side of the leaf has specially
high epidermal 'cells, elongated like' a palisade, and having a thick outer wall ;
the'lumentis narrowed conically,towards the exterior, and contains a deposit of
fat-bodies. 'Papillose differentiation of the epidermis, or the development of a
hypoderm: are characteristic of a large numb~r of species. Behm mentions the.
occurrence of 'papillae in species of A rjona, Exocarpus, Leptomeria, Quinchamalium,
Santalum and Thesium;' their development extends for the most part only to
single epidermal cells, or to groups of such cells, which are either situated on the
surface10r confined to the margin of the leaf; papillose differentiation of the,
whole' of the lower epidermis appears only to have been observed in Santalum
album, L. According to Behm, 'the' development of hypoderm is found in
Henslowia '(H: granulata, Hook: fil. et Th., H. heterantha, Hook. fil. et Th.,
H. retusa, Bl., :H; varians~ BL), in Jodina rhombi/olia, Hook. et Am. and in
Santai'u.m aCllminatum, A. DC: (and also locally in S. Preissianum, Miq.). In
Henslowia arid Santalum acuminatum the hypoderm is composed of somewhat
collenchymatotis cells, which are larger than those of the epidermis; it consists
of two layers in Henslowia, of one layer in Santalum acuminatum. In Jodina
rhombi/olia the strongly collenchymatous hypoderm and the epidermis together
give one the impression of 'a many-layered epidermis, when seen in transverse'
section. The structure of the stomatal apparatus is, on the whole, uniform/;
each pair of guard-cells is generally accompanied on either side by one or mort'
subsidiary ctlls, placed parallel to: the pore. It should, however, be mentioned
that the type of stoma just described is ,sometimes less distinctly marked,
even in the same superficial section, owing t6 the fact that secondary
vertical walls, running at right angles to the pore, more or less frequently
arise in the subsidiary cells. Actual exceptions to the characteristic type
of stoma are, however, only afforded by species of Arjona and Quillchama-
lium~ iIi which the same superficial section shows stomata with parallel.sub-
sidiary cells accompanied by others, which' are surrounded by a relatively,
large number of adjacent cells exhibiting no special arrangement. The arrange-
ment of the stomata with reference to one another is also of value for the recog-
nition of certain g~!l~r~ and_ species., In some cases they are placed parallel to
one another and transversely "to. the midrib of the leaf i this arrangement i~
well marked, acco~ding to Behm, in species of Exocarpus, Leptomeria. Ompha-
comeria, Osyfis, Santalum and Thesium. and, according to ChaUn, in A II/hobo/res
and Chorethrum, Whilst; according to Behm, in other species of Osyris and .in
species of Comandra and Osyr£dicarpus the tendency towards transverse arrange-

J Behm examined numerous species of the genera QlIinc/zanralirtfll, Alj01ta, Thesium, usyridi-
carpus, Cervalltesia, jodt'lIa, Pyru/art'a, Aca1tt/tos)'l"is, Comamlra, Saltlalum, Myos(/Zi/os, h'tukltya,
Osyris, Omp" acom ert'a, llens/{)Wia, C/zorelrum, Lfplomeria, Exorar/,us; Chatin, besides 'de3.linJ,:
with species of certnin of the genera a.lready mentioned, has also figured the structural' (ealmes of
Narrodea, FuulnuS (Mida) nnd An/hobo/us.
732 SANTALACEAE

ment of the. guanl-cells is only seen here and there. In other cases, which are
restricted to the genus Ariona, the pairs of guard-cells are likewise arranged
parallel to one another, but at the same time they lie parallel to the midrib of
the leaf. In other cases again we have an irregular arrangemen.t of the pairs of
guard-cells, this being the most widely distributed type (in species of A can tho "
syris, Buckleya, Cervantesia, Exocarpus, H enslowia, Iodina, M yoschilos, Pyrularia,
Quinchamalium and Santalurn, according to Behm). A transverse arrangement
of the stomata, similar to that shown by the leaves, occurs also on the young
branches in many members of the Order; Chatm mentions this feature as being
more or less distinct in species of A nthobolus, Choretrum, Fusanus, H enslowia,
Myoschilos, Osyris and Thesium, whilst De Bary mentions its occurrence in_
Exocarpus and Santalurn, and Hieronymus in Iodina and Leptorneria. The
stomata either occur on both sides, or only on the lower surface of the leaf.
The guard-cells in'most cases lie on a level with the epidermis, but they ~ave
been observed to be deeply depressed in A1'iona, Iodin" and Osyris. The
vascular bundles of the veins, as stated abov(f; -are invariably embedded;
mechanical tissue may accompany them, or may be absent. The: terminal
tracheids, previously mentioned in the general diagnosis, are found in most ot
the genera; amongst the genera examined by Behm, they are only absent in
A canthosyris, Buckleya and Iodina. They form groups of two to five ele-
ments, and are developed in a specially typical manner in Henslowia. In
Quinchamalium and Thesi1tm, besides the terminal tracheids, there are others,
which are independent of the veins, and form-rows of elements running parallel
to the latter; these tracheids are more or less· elongated, and their walls
-either have bordered pits, or are spirally or reticulately thickened.
Regarding the occurrence of oxalateofUme, we may mention once more that
clustered and ordinary solitary crystals occur in both the leaf and axis. Ac-
cording to Behm, certain species of Cervantesia and Leptomeria have. specially
large clustered crystals in the leaf. In Santalum album, L. and Iodina rhombi-
folia, Hook. et Arn. solitary crystals are found in the epidermis of the leaf; in
the case of ] odina rhombi/alia they appear, on superficial observation, to lie in
the considerably thickened outer wall of the epidermis. Behm also notes the
presence of crystalline, in some cases sphaerocrystalline, doubly refractive
masses in, the epidermal cells of Thesium montanum. Ehrh., T. ebracteatum,
Ha:yne, and Santalurn acttminatum, A. DC. -; in .view of their ,solubility in
alcohol and ether, these masses are probably ora fatty nature. Bebm observed
grou ps of silicified cells in species of A canthosyris, Buckleya, Comandra, Exocarpus,
Osyris, Pyrularia 1, Quinchamalium, Santalum and Thesium. They are present
chiefly in the neighbourhood of the veins, and either consist of pairs of cells, or
of larger groups. The walls of the cells in question either exhibit silicified
protuberances (often hemispherical), or they are only locally thickened (like
a horseshoe. as seen in transverse sections of the leaf) and silicified.
The most important points concerning the hairy covering have already
been stated above. In most cases the simple unicellular hairs are small and
pointed. Uniseriate hairs are mentioned by Behm as occurring in Buckleya
lanceolata, Miq. (incorrectly named B. Quadriala. S. et Z. by Behm), and are
also present in Pyrularia. according to Chatin. The tufted hairs of Exocarpus
lati/olius, R. Br. consist-of short unicellular hairs having thick walls and-narrow
lumina, and sunk side by side in the epidermis.
3. STRUCTURE OF THE AXIS. The most important characters presented by

1 These groups of silicified cells were previously se~n by Bokorny in P. pu"ua, Michx., wh~e
they iive rise to tranllparent dots nt the margin of the lenf, bnt they were not correctly interpreted
by him.
 SANTALACEAE 733
the structure of the wood and cortex have already been pointed out in the
general review at the beginning of the Order.
The structure of the axis is normal. The xylem-portions of the vascular
ring as a rule form a closed mechanical ring, traversed by medullary rays which
are most1y narrow, though occasionally (in species of Acanthosyris, Osyridi-
carpus, Osyris, ] odina) rays 4-5 cells thick are present. Exceptions to this type
are presented by some herbaceous species, viz. Nanodea, in which, according to
Chatin, the transverse section of the stem exhibits four isolated vascular
bundles, arranged in a ring, and species of Thesium and Arjona, in which the
bundles. of the vascular ring are separated from one another by unlignified
tissue. The pbyllocladee of the leafless species or of those with reduced leaves
(Exocarpus phyllanthoides, Endl., Omphacomeria psilotoides, A. DC.) possess
a vascular ring, the bundles of which are almost compressed into one plane in
Exocarpus, whilst in Omphacotr..eria thre~ larger bundles, which form a ring
open at the sides, are seen in the middle of the main vascular ring; in other
respects the phylloclades have the structure of a centric leaf.
The pith consists either of lignified or unligr.ified cells. The vessels are
scattered in the transverse section, and have relatively small lumina (between
·018 and ·03 mm.). In contact with parenchyma of the medullary rays, the
walls of the vessels mostly bear bordered pits, though there are sometimes
simple pits as well; the latter are never large, and do not exceed the borders
of the bordered pits in size. Spiral striation or thickening of the walls of the
v~~·is' stated by Behm to be present in species of Exocarpus, Osyris, Santalum
"aDd Thesium. The development of cork takes place either in the epidermis
(A cantlwsyris and Comandra), or in the outermost cell-layer of the primary cortex
(Buckleya, Choretrum, Henslowia, Leptomeria, Myoschilus, Osyris, Santalum and
Thesium). In many of ·the Santalaceae showing reduction in the leaves, the
primary cortex is differentiated as typical assimilatory tissue, as in the phyllo-
clades mentioned above. In some cases the primary'cortex includes stone·cells
(species of Choretrum, Henslowia, Santalum and Thesium), and occasionally
there are also bundles of sclerenchymatous fibres (according to Behrn, in San·
talum, and in the shoots of Omphacomeria which resemble 'phylloclades~· in these
two cases the fibres are mainly subepidermal; according to Chatin, they also
occur in Leptomeria BiUardieri). The outer limit of the bast is in most cases
formed by isolated groups of primary bast-fibres, between which scattered stone-
cells may occur at some points. The stone-cells are present in such abundance
in Henswwia that a composite and continuous ring of sclerenchyma is formed.
Amongst the material investigated by Behm, the groups of primary bast·fibres
are only wanting in the species of Comandra; according to Chatin, however,
they are present in a species of this genus (C.livida), which was not examined by
Behm. Typical bast-fibres have only been observed in the secondary bast in
the case of Omphacomeria and Santalum (Behm); the secondary bast contains
stone-cells in species of Osyridicarpus and Henslowia (Behm), and peculiar
sderenchymatous· elements, resembling' both stone-cells and bast-fibres,' in
Exocarpus c'Upressiformis, Labill. (MeHler I).

APPENDIX.
I. On the genus Myzodendl'on.

The parasitic ~enus Myzodend,on. which has recently been separated from the
Santalaceae and raISed to the rank of an independent Order by HIeronymus in the
Nattirliche Pflanzenfamilien, agrees with the Santalaceae in many ofits anatomjcal
characters; thus it has: superficial development of cork; simple :perforations in ,the
vessels; no internal or external glands; silicified groups of cells; solitary and clustered
1 RegardiDg the haustoria of the Santalaceae see Solms·Laubach and De Bary, 11. ce.
73+ SANTALACEAE

crystals; isolated groups of hard bast in the pericycle, simple uni<;;ellular hairs; again,
in one species (M. {'1'near-e, Popp. et Endl.) systems of tracheids are preseatin the leaf,
similar to those of the species of Quinchamillium and Thesium, and another species
(M. quadriflO1'um, DC.) has crystalline masses in the epidermis of the leaf (and also
in· the ground-tissue of the axis) like those found in Thesium montanum, &c. There
are only two important anatomical distinctions between Myzodendt'on and the majo-
rity of the Santalaceae to be noted, viz.: I. The stomata are surrounded by a rather
large number of irregularly arranged epidermal cells. II. The wood-prosenchyma.,
i.e. when there is any development of mechanical tissue in the xylem, lS replaced by
sclerenchymatous cells bearing simple pits and resembling rod-cells.
Little remains to be said on the subject of the structure of the leaf. The meso-
phyll consists of uniform isodiametric cells. The stomata ar~ found on both surfaces
of the leaf, and are placed parallel to one another and to the midrib of the leaf. The
vascular bundles both of the large and small veins are embedded in the mesophyll ;
there is no sclerenchyma accompanying them. Silicified groups of cells have been
observed by.Behm only in M. imbricatum, POpp. et Endl., M.lineafe, popp. et EI;tdl.,
M. punctulatum, Soland. and M. quadfi{lorum, DC. .
The structure of the axis requires a more detailed description, and on the
characters .presented by it the investigated species can be divided into two groups.
corresponding to the two sections Eumyzodendron and Gymnophyton, established by
Hooker fit The species of the section Eumyzodendron (M. brachystachyum, DC.,
M. ,,"neare, popp. et Enell. = M. lineat'ilolium. DC., M. oblongilolium, DC., M~
quadriflofum, DC.) are characterized by the following features: (a) the vascular
bundles are separated from one another by radial strips of unIignified ground-tissue;
(b) the ground~mass of the wood consists of unlignified cells and is developed in
varying abundance in the different species, depending on.-the number of vessels and
groups of mechanical cells which are embedded in it; and (c) the secondary wood
contains two kinds of vessels, some being scalariform and others having spiral
thickenings. On the other hand the investigated species of the second sec.tion
(Gymnophyton), viz. M. tunctulatum, Soland. and M. imbricatum, popp. et Endl..
ha.ve the xylem-groups 0 the vascular bundles united to form a dosed mechanical
ring. composed of lignified cells only, and traversed neither by primary nor
secondary medullary rays. In certain species of the first section, though not in
aU (only in M. brachystachyum, }II. oblongi/olium, M. quadriflo'l'um. and also in M.
angulatum, Phil., according to Hieronymus) •. medullary vascular bundles occur.
They arc normally orientated, are present in varying abundance, and possess con-
siderable secondary growth in thickness. Not uncommonly they form a. second
(inner) ring of bundles ,Fig. 178, A). According to Behm's investigations the
medullary bundles are of secondary origin in ~w:. oblongifoli-um. Their development
appears to be connected with the formation of pseudoendogenous buds, and this
may possibly explain the fact that, acc<?rding to Behm, in one and the. same speci~s
the medullary bundles may be present In one branch, though absent III another of
equal thickness.
Of the more special anatomical features in the structure of the stem the
follOwing are noteworthy. M. punctulatum alone has numerous verrucose promi-
nences on the surface of the stem, giving the latter a punctate appearance; hence
the specific name. These prominences are due to large respiratory cavities, and
each bears a stoma at its apex. The formation of cork is superficial. In most
species the cork develops in the epidermis; in M. obJongiloJium (in the specimen
described by Behm as M. hetet'ophyllum. POpp.). in which the epidermal cells on the
young branch are produced into finger-shaped unicellular hairs I it·arises in the sub-
epidermal layer of cells. In M. b"achystachyum and M. oblongifolium the primary
cortex contains a more or less closed ring of stone-cells; in M. obioffg-itolium there are
also scattered groups of stone-cells. In M.l'ineafe stone-cells are distributed throu~h­
au t the ground -tissu e of the axis, and are a Iso found in the soft bast. In all the SpecIes
groups of hard bast are present at the outer limit of the phloem, but" are developed
to a varying extent. The xylem in all species of Myzodend"on is characterized by the
fact that the vessels have relatively small luminal are composed of short elements,
and exhibit simple perforations. In the species belonging to the section Eum1zoden.
dl'on, asalready mentioned, the secondary wood possesses two kinds of vessels, viz. (Q)
vessels with rather large lumina and provided with·scalariform bordered pits, and
(b) spirally thickened vessels with somewhat narrower lumina; in the species of the
section Gymnophyton, on the other hand. we only find vessels of th~ first kind. In
 SANTALACEAE 735
the species of both sections themechanicalelementsof the xylem consist of sclerenchy.
matous cells (' wood-prosenchyma ') resembling rod-cells; they have thick walls and
narrow lumina, and bear simple pits. It has already been stated above that. in
the species of the section Gymnophyton, these sclerenchymatous cells together with
the pitted vessels form a closed woody ring, devoid of medullary rays. In the
SpeCIes of the section Eumyzodendron, on the other hand, the ground.mass of the
wood in the individual vascular bundles consists of unlignified cells, the bundles
being separated from one another by strips of thin-walled tissue. In those species
in which material of sufficiently thick branches has been examined (M. oblongitolium,
M. bt'achystachyum and M. quadrifl(Wum, according to Behm, and M. linearifolium,
according to Hooker and Chatin) it has been found that the vessels and sometimes
also the wood·'Prosenchyma show a very characteristic distribution amo:l~t the
unlignifted cells In a transverse section of the branch. In .M. obion_ei/olium (fIg. 178)

FIGr.178. M.y,OtInulr'()n DiH()nriffJiium, DC.: A, Dia~matic ~p~sentation of a trans\-erse sr.ction

through the ois; 8, a single vascula.r bundle.-Original. -

each of the individual vascular bundles (including the medullary bundles, when
they have undergone considerable growth in thickness) contains groups of scalari-
form vessels, which are embedded in a small quantity of thin-walled tissue and
alternate in the radial direction with portions of tissue composed of spirally thickened
vessels~ a few thin·walled cells, c.nd t.l-te mechanical elements (wood-prosenchyma).
In the portions of tissue last mentioned, the arrangement of the elements is as
follows: the middle region is occupied by a radial series of the thin-walled cells;
amongst these the spirally thickened vessels are embedded, whilst a bundle of
short prosenchymatous cells lies on either side of the radial group. A similar
stratification of the secondary wood has also been observed in the other species of
the se~tion Eum"zodendt'on. mentioned above, except that the groups of mechanical
tissue are only dev~lol?ed at certain points, or are entirely absent. In some species
(M .linea1'e, 11,1. oblongstoli um) bundles of elongated sclerenchymatous cells, resembling
hard bast, are found on the inner side of the vascular bundles. The pith as a..rule
~
 SANTALACEAE

consists of unlignified cells with thin walls; in the investigated species of the
section Gymnophyton it is composed of prosenchymatous cells having thick walls.
For the structure of the haustorium of Myzodendron reference may be made to
Chatin, pI. !xxv.
2. On the genus Champereia.
Recent authors refer the genus Champeyeia to the Opilieae (which are provi4ed
with cystoliths, according to Edelhofi and Valeton); and like the Opilieae, Cham-
pe1'eiQ possesses typical cystoliths, a point to which attention was first drawn by Van
l·ieghem. T~ese cT-?toliths cannot. however. be regarded as decisive in determining
the systematic posItion of the genus. For Eehm bas shown that transitional forms
connect the cystoJiths with the cystolith-like. silicified protuberances. which occur
both in the Opilieae and in the typical Santalaceae.
The cystolith-like structures of Champereia may be divided into threP. types.
The first consists of true double cystoliths (Fig. I79. A and B). They were observed
by Behm in all the ma.terial which he examined; he found them to occur in the bast
of the branph. and also in the pith and the tissue. Qf the medullary rays. Instead of
two there may sometimes be three or four cystoliths together. The double cystoliths
lie in two adjacent cells, which are in contact wi th on~ another by their narrow sides,

FIG. 179. A, Double cystoliths (rom the bast of the axis in '(;1uz111/¥l'ei4
Gr;.fIi,nian.J, Pll!nch. (Gnffith, n. 4388, Herb. Mon2.c..), calcified; B. t~
same after de!;alcitkation with acetic acid. C, SeC'.ondarv metamorphosis
of the double crstolitb$ of CIu:_pereia sp. (Cuming, lI2Q. Herb. Monac.),
see description In text. D, Group of ceils with silicified cystolith-like bodie!l
from the leal·tissue (If CAo.mperda Gr,£1illli.:rna.-Otiginal.

the latter being placed horizontally in the pith and bast. and vertically in the medul- ,
lary rays. The stalks of the two cystoliths, which are mostly calcified. spring from
the common wall of contact. The second type is formed by cystolith-iikc structures,
which may be shortly described as cystoliths which are sessile, or devoid of a stalk,
and have a reduced skeleton; they have been observed in the primary cortex and in
the tissue of the leaf, giving rise to trans'parent dots in the latter. They form spherical
gr0";lJ>.S of 2~ c~lls. each of wh~ch c~ntams a body resembling a cystoht~. These cells
exhibit conSiderable local thlckenmgs on the walls of contact, and 10 each cell a
crystal of carbonate of lime. having a rather irregular outline and almost fiUing the
lumen of the cell, is inserted in the thickening of the wall like a tooth in the jaw.
These characteristic groups of cells. or rather the crystals they contain, together
give one tlie impression of a large clustered crystal, when examined under a low
magnification. The third type is formed by silicified protuberances having the shape
of a cystolith (Fig. 179. D). the lithocysts being arranged in a group. as in the second
type. and t~e group ~Ying.a more?r less sp.herical sh<1:pe. Behm. only m~t with this
third type m a certam portion of hiS materIal. where It occurred'm the tissue of the
leaf. For the transitions between the three types. observed by Behm, we may refer to
his treatise. An interesting metamorphosis of the twin cystoliths and their'litho-
.::ysts·(Fig. 179, C). however. must pot be omitted here; all stage§' were obserVed by
Behm in the bast of the material collected by Cuming (n. 1129. Herb. Monac.). ThiS
secondary mebmorphosis may'extend to both or only to one of the cystoHths with
 SANTALACEAE 737
the corresponding lithocysts, and is 1nitiated by the disappearance of carbonate of
lime from t..'J.e cystoliths. At the same time the cell-wall of the lithocyst becomes
sclerosed. A further stage is found in those cases in which sclerosis also affects the
outer portion of the cystolith, so that one might imagine there were two sclerosed cells,
the one inserted in the other (Fig. 179, C). Finally, in some cases sclerosis is carried
so far that the sclerosed wall of the lithocyst fuses with the sclerosed outer Y0rtion
of the bOGy of the cystolith to form a homogeneous membrane; in this fina stage,
therefore, we obtain a stone-cell, the lumen of which still shows the approximate
shape of the original cystolith.
The following points are noteworthy in the structure of the leaf and axi. of
Champercia. The leaf-structure is bifaclal. The stomata are only present on the
lower side of the leaf, and possess several subsidiary cells, placed parallel to the
pore. The vascular bundles of the smaller and larger veins of the leaf are embedded.
The xylem is composed of: (a) vessels with small lumina (diameter = 0033 mm.),
simple perforations, and bordered pitting in contact with parenchyma; (b) two- to
three-seriate medullary rays; (c) wood-prosenchyma with rather thick walls and
bordered pits; and (d) a small amount of wood~parenchyma. A composite and
continuous ring of sclerenchyma is found in the pericycle. The cork arises in the epi.
dermis, and consists of cells with thick waUs and narrow lumina.
3. On th~- genus Grubbia.
The genus G,ubbia has, on the one hand. been referred to the Santalaceae as
an anomalous genus, on the other hand it has been associated with other Orders,
such as Hamamelidaceae and Bruniaceae. and recently it has also been regarded as
a separate Order, the Grubbiaceae. It is essentially distinguishe<l fr,pm the Santa-
laceae anatomically by the occurrence of scalariform perforations i.P: the vessels, and
by the absence of parallel subsidiary cells in the stomatal apparatus. Neither have
silicified groups of cells, nor terminal tracheids, nor indepenoent rows of tracheids in
the leaf-tissue been observed in this genus. Points of resemblance between Grubbia
and the Santalaceae are the small lumina of the..vessels, the wood-prosenchyma with
bordered pits, and the superficial formation of the cork.
The following facts maybe added regarding the more detailed structure of the
axis and leaf. The leaf-structure is bifacial. The stomata are found only on the
lower surface of the leaf. a.nd are surrounded by a relatively large number of irrep-
larlyarranged epidermal cells. The wood consists of; Ca} vessels With smallluml!la
(diameter = -03 mm.) and scalariform perforations with numerous bars (40 or more),
and both simple and borde~ed pits on t~~~llsin contact with parenchyma of themedul-
lary-rays; (b) narrow, urusenate or blsenate medullary rays ; (c)wood-prosenchyma
with bordered pits; and (::I) a small amount of wood-parenchyma. A com~ite and
continuous ring of sclerenchyma is present at the outer limit of the bast lD Gfubbia
stricta. A. DC., while in C. t'osmatinitolia, Berg there are only isolated groups of hard
bast. According to Hieronymus the branch'in the species of the section 0phi,a is
characterized by a hypoderm, which is sharpll differentiated from the mtemal
tissue, whilst in the species of the section Styobuocaypus the hypoderm and cortical
tissue merge into one another. The ,cork arises subepidereiallyand is composed of
cells with rather thick walls. Oxalate of lime is found in the form of clustered and
solitary crystals. In the leaf of Cyubbia yosmarinilolia the clustered crystals occur
chiefly in septate cells of the palisade-tissue; while in G. styicta idiobla.sts,
situated immediately beneath the epidermis of the leaf, contain solitary or
more rarely clustered crystals. The hairy covering cqnsists of simple. unicellular
triehomes, which sometimes have verrucose thickenings.
- ... ~

Literature: Chatin, Anat. comp., PI. parasit., 1856-63, pI. lvi-lxxvi a and c::t.-Dalton~Hooker,
Mpodendr?It, Ann. sc. nat., ser. 3, t. v, 1846. p. 193 et seq. and pI. 5-9, and in Flora antarct., vol. ii,
18#-5, p. 289, Tab. cii-<:vii.-Pitrll, Phan. Paras., Bot. Zeit. 1861, p. 69.-Solms·Laubach, Paras.
Phan., Pringsheim Jahrb., Bd. vi, 1867-8, PP' 539-60 and Tab. xxxii-xxxiii, and abstr. of Clarke,
LOnLutha.ceae, Bot. Zeit. 18H, p. 145 et seq.-MoUer, Hobanat., Denkschr. Wiener Akacl. 1876,
p. 39 and 335.-De Dary, Vergl. Anat., 18n.-Moller, Rindenanat., ISh, pp. 1I;\-I4.-Bokomy,
Dnrchs. P., Flora 1882, p. 358 et 5eq. and sep. copy, pp. IS-16.-So1ereder, Holzstr., 1885, pp.
:3S-6.-Radlkofer, in Sitz.-Ber. Miinch. Akad. 1886, p. 330.-Hieronymus, in Natiirl. Pflanzenfam.,
iii. Teil, Abt. J, 1889, pp. 199-200, 20+-206 and ~:39.-Van Tieghem, Clzamjereia, Ann. sc. nat~J
set. '1. t. xvii, 1893, pp. 25S-6.-Houlbert, Bois 6('C. dans les Apetales, These, Paris, 1893, pp. 165-
,o.-Van Tieghem, .A"allogia, Bull. Soc. bot. de France 189+. p. 61 et se9..-Behm, Anat. Charakt.
d. S., Diss., Erla.n~D, 1895,56 pp., sep. copy from Bot. Centralbl. ]895, 11.-[Engler and Volkens,
Ostafr. Sandelholz, Notizbl. bot. Gart. n. Mns. Berlin 1897, n. 9.] .
SOLEIIEDER 3R
 BALANOPHOREAE.
The Balanophoreae, as is well known, are root-parasites devoid of chLoro-
phyll; th~y will only be treated here as far as is compatible with the object of
this book. For information regarding other features the literature cited must
be referred to.
The ground-tissue in these fleshy plants, which have a yellowish or reddish
colour, consists of large parenchymatous cells; these become smaller towards
the periphery both in the rhizome and in the peduncles. In the genera Balano-
phora, Langsdortfia and Thonningia, this tissue contains a substance 1 . first
observed by Goeppert, who gave it the name balanophorin; according' to
Poleck, it is a body having the appearance of wax, but of the nature of resin,
and essentially distinguished from wax by its stickiness and higher melting
point. In the other genera starch is found m place of the balanophorin. Other
special cell-contents in the ground-tissue are the clustered and solitary crystals,
met with in Balanophora alutacea by Goeppert, and the sphaero-crysialline
clusters consisting of oxalate of lime and observed il'l Lophophytum mirabile~
Schott et Endl. by Eichler. In certain species (Helosis guianensis, Rich.,
H. mexican-a, Liebm., Langsdortfia hypogaea, Mart., Lophophytum mirabile,
Schott et Endl., Rhopalocnemis, Scybalium iamaicense, Schott et Endl.) stone-
cells, which are either isola.ted or united into groups, occur in the ground-
tissue j the cells are sclerosed on all sides, or (Langsdortfia hypogaea) on one side
only. In the tuberous rhizome of LoPhophytum mira bile they become secon-
darily corroded owing to a process of solution, and ultimately become completely
I dissolved (Eichler).
! The epidermis consists of small cells, ana contains no stomata. In Balano-
, plwra involucrata, Hook .. f. et Th. the epidermis includes cells of a papillose
nature, occurring singly or united into groups (Hooker). Trichomes are rare
(LangsdortJia, Thonningia) on the vegetative organs; where present they are
unicellular, have thin walls and rather wide lumina, and a bulbous basal enlarge~
ment.
The vascular bundles are always collateral; the following facts may be
mentioned regarding their distribution n the rhizome and peduncle. Owing
to the irregular course and the branching of the vascular bundles, their arrange-
ment is irregular in the tuberous rhizomes (Helosis, Langsdortfia, Lathrophytum,
LfYj>/t.,Qphytum, Rhop alo cnemis, Scybalium) and in the axis of the inflorescence
(Cynomorium~ He-Iosis, Langsdottfia, LoPhophytum, Rhopalocnemis, Sa ...cophyte,
Scybalium). This irregular arrangement IS most marked in Scybalium /ungi/orme,
Schott et Endl. and LoPJwphytum mirabile, Schott et Endl., in which the number
of bundles in the transverse section amounts to 200-300, whilst in the axis of
the infiorescence of Helosis guianensis one can distinguish two more or less
distinct rin~s of vascular bundles, and in that of Langsdorffta hypogaea a main
ring with medullary and cortical bundles. The cylindrical rhizomes of Helosis
and Langsdorgia generally have only a single ring consisting of a few isolated
vascular bundles; at those points, however~ at which local tuberous thickenings
occur, the regular arrangement ceases. In the floral axis and cylindrical
rhizomes of Helosis guianensis the vascular bundles are provided with a layer
of sc1erenchyma at the margin of the wood and bast. Further, in the cylindrical
,rhizome...;; of Helosis the pith, situated internally to the vascular ring, consists
J

entirely (H. guianensis), or at least in its peripheral portion (H. mexicana), of

sclerenchymatous fibres-, aud the strips of ground-tissue, lying be~~een the
vascular bundles, also become sclerosed. The vessels of the Balanophoreae

1 The amount of balanophorin is so considerable that the shoots containing this substance are
used as torches by the natives.
 BALANOPHOREAE 739
exhibit reticulate or striate thickenings; their perforations are simple in Helosis
(E. Zimmermann). It is doubtful whether Eichler's statement (which is
repeated by Engler) regarding the occurrence of scalariform perfora.tions is
correct. Tracheae with spiral thickenings, which can be ~rolled, and annular
vessels are said to be absent in this Order; mechanical elements are not deV'e~
loped in the xylem. The sieve-tubes of H elosis have scalariform sieve-plates
at their ends and on the lateral walls (E. Zimmermann).
The scale-leaves of the Balanophoreae consist of homogeneous tissue, which
includes groups of stone-cells in Langsdortfia hypogaea and Lophophytum mirabile.
'The number of the vascular bundles which pass into the scale-leaves and there
undergo further branching, is three in Langsdorffia hypogaea and LoPhophytum
mirabile, and one in Scybalium fungiformc.
Literature: Unger, Kellatn. d. ·pRTasit. Pfl., Ann. Wiener Mus., Bd. ii, 1840, pp. u-60.-
Goeppert. Ban d. B. &c., Verh. his. Leopold .• Karol. Akad. d. Naturf., Bd. xviii, Suppl. I, 1841,
p. 239 et seq. and 3 tab •• and Rlwpalonu",is, loco cit., Bd. xxii, I, 1847, p. lJ7 et seq. and Tab.
:Q-:n.-Poleck, Wachsiihnl. Best. d. Baian()pn()ra, Verb. kais. Leopold.-KatoIin. Akad., Bd. uii, 1,
1847, p. 159 et seq.-Hooker, Struet. and aff. of B., Transact. Linn. Soc., vol. xxii, I, 1855-6, p. I.
et seq. and pI. i-xvi.-WeddelI, CJ"U)1nOn"um &()(cineum, etc., Arch. Mus. d'hist. nat., vol. lE, 1858-
61, pp. 369-308.-Chtin, Anat. compo d. v~get.: pI. parasit., pp. 93-101 and IC5.-So1~Laubacb.
Puasit. Phan., Pringsbeim Jahrb., Bd. vi, 1867-8, pp. 539-39.-Eichler, Latnroph)'Ium, Bot. Zeit.
1868, pp. 514-15, and B., in Martius Flor. bruil., vol. iv, l, 1869. p. 1 et seq. and tab. ii, vi, viii,
xii and xiii.-Beccari, in Nonv. Giorn. bot., vol. i, 1869< p. 65 et seq. and Tab. iii.-Solms-Laub2.ch,
Haustorium. Abh. naturf. Gesellsch. Hallc, Btl. xiii, 1877, p. 267 et seq. and Tab._xxv.-De Buy.
Vergl. Anat., I8n.-E. Zimmennann, He"'sis guianensis, Flora 1886, pp. 371-86 and 400-::11, Tab.
fl.-EDgier, in Natilrl. PBanzenfam., iii. Teil, Abt. I, 1889. pp. l#-7.-Peirce, Street. of haustoria
&c., Ann. of bot., 'Vol. vii, 18,93, p. 318 et seq.-(Pirotta e LQ!lgo, Stomi nel Cynomtirium, R. Accad.
d~ Lienei 1899. pp. 98-loo.J

EUPHORBIACEAE
(exel. Daphniphyllaceae and Buxaceae 1).
I. REVIEW OF THE ANATOMICAL FEATURES. No special anatomical features
are common to all the Euphorbiaceae. On the other hand there are numerous
anatomical characters, which may be of value in enabling one to recognize a
plant as a member of this Order, and also for purposes of more precise deter-
mination.
In the structure of the leaf we may specially note that the stomatal appara-
tus is not of uniform structure. The pairs of guard-cells possess subsidiary cells
of the type found in the Cruciferae or Rubiaceae-, or they are surrounded by a
nwnber of ordinary epidermal cells. In the structure of the axis the following
features may be mentioned. The cork in most cases develops subepidermally,
rarely in a deeper cell-layer of the primary cortex~ or in the epidermis. The
pericycle either contains no sclerenchyma, or isolated bundles of bast-fibres,
sometimes uniting to form a ring, or a composite sclerenchymatous ring, which
is completely continuous or nearly s()_. Groups of bast-fibres are frequently
present in the secondary'hast, but only very rarely give rise to stratification of
the latter. The primary and secondary bast·fibres are often characterized by
a well·marked stratification of their walls. The xylem consists of: (a) medul·
lary rays which are mostly narrow; (b) vessels, which either (in most cases) have
simple perforations only, or both simple and scalariform; or (very rarely) ex-
clusively scalariform perforations; in contact with parenchyma they either bear
bordered pits only, or the latter are intermingled with simple pits-of varying size ;
(c) wood·parenchyma. which is abundant in only_aJew members ofthe Order;
(d) finally, wood-prosenchyma with thick walls and narrow lwriina, or ra-ther

I The Daphniphyllaceae and Buxaceae will be treated separately in an. Appendilt at the end of
the Euphorbiaceae.
3B2
 EUPHORBIACEAE

thin walls and wider lumina; in the former case the wood·prosenchyma is some.:
times provided with a gelatinous layer, while in the second case it is septate; in
the majority of the Euphorbiaceae the walls of the wood-prosenchyma are.
furnished with simple pits, whJe in a small number of genera (of the
Tribes Caletieae, Phyllantheae, Hippomaneae and Acalypheae) they bear
bordered pits.
Oxalate of lime is mostly excreted in the form of ordinary solitary and
clustered crystals; in certain species it is entirely absent. The following special
forms of crystals are found: sphaerites (Acalypha, Claoxylon, Croton, Crotonopsis,
E,emoca,pus, Gelonium); stvloids (Bernardia, Colliguaya, Excoecaria, Pachy-
stroma, PhyUanthus); and rod-shaped crystals, bent like a VorW (Phyllanthus).
In certain Ampereae, Crotoneae, Acalypheae and Hippomaneae clustered and
solitary cryst::-Js occur in the epidermis of the leaf, while in many EuphorHiaceae
they are found in special idioblasts in the mesophyll, the latter sometini~
giving ·rise to transparent dots in the leaf. The internal secretory system
is constituted as follows: I. laticiferous cells (in Croton pro parte, some
Acalypheae, and many Hippomaneae and Euphorbieae) ;' I1.laticiferous vessels
(in H evea and M anihot only); III. rows of laticiferous sacs (only in -111 icrand,a) ;
IV. tanniniferous cells, sometimes differing from the surrounding cells in the size
of their lumba or in their length, and occurring singly or in rows j they are
found principally in the bac;t, where they appear as more or less distinct idioblasts
(widely distributed in all the Tribes); V. very much elongated secretory sacs,
which frequently have wide lumina and are mostly filled with brown contents
(in Alchornea (?), Givotia, Mallotus, Pausandra, Pogonoplwra); they must not
be confounded with the laticiferous cells; VI., secretory cells with oily or
resinous contents (Fig. 180, A-C), which show varied differentiation, and have
various shapes, and sometimes give rise to pellucid dots in the leaf (in Ricinus
and the Eucrotoneae, excepting some species of Croto"" belonging to the section
Ast,aea); VII. secretory lacunae, i.e. intercellular secretory spaces with curved
epithelial cells loosely connected with one another; they usually give ric;e to
transparent dots in the leaf (in Cluytia and Gelonium); VIII. mucilage· cavities,
or groups of mucilaginous cells (in certain Phyllantheae). Of all these secretory
organs, only those numbered I-III have geen shown to produce a milky juice.
The trichomes maybe classified under clothing hairs, external glands, and
stinging hairs (including the hairs containing clustered crystals). The ,usual
form of the clothing hairs is the simple, unicellular or uniseriate trichome, which
is frequently present. Special forms of such trichomes are the unicellular, t\yo-
armed hairs of Argyrothamnia and Pausandra, and the branched hairs of species.
of Phyllanthus and Mabea. The following types of clothing hairs (see Fig. ISo)
have a more complicated structure: stellate and tufted hairs (in the Ricmo-
carpeae, Phyllantheae, Crotoneae, Acalypheae and Hippomaneac); peltate
hairs (in Aextoxicon and certain Phyllantheae, Crotoneae and Acalypheae) ;
simple shaggy hairs (in certain Phyllantheae and Acalypheae); and candelabra-
hairs (in species of Croton, Fig. lBo, KJ. Regarding these hairs it may be added
that sc1erenchymatous cells belonging to the mesophyll are sometimes associated
with the complicated trichomes of the species of Croton in such a way that they
appear to form a component part of the hair; some of the peltate hairs (Fig. lBo,
L) are provided with a small lower scale: Ordinary glandular hairs are ,not
I

common; they have been observed in the tribes Ricinocarpeae, Phyllantheae,

Acalypheae (here more frequent than elsewhere) and Hippomaneae. Large,
vario11sly shaped glands (Fig. lBo, !vI-O)t provided with a secretory palisade-
like epidermis, are inuchmore widelydistributed; they occur on the petiole, on the
surface of the leaf, on the leaf·teeth, or on the stipules of many species, and some
of them. are transitory. The genera Cnesmone, Leptcwhachis and T,agia, be-
longing to the Acalypheae, and the genus Dalechampia have very characteristic
 EUPHORBIACEAE 741
stinging hairs (Fjg~ ~Bo" P-Q), the essential part of which is a cell containing
a subulate crystal .of· oxalate of lime, suspended from beams of cellulose;
I atropha has stinging hairs showing the same structure as those of the nettle.
Hairs or papillae with clustered crystals (Fig. lBo, R-S), i.e. papillose projections
of the epidermis, resembling stellate hairs or of spherical shape, and filled by a
clustered crystal of oxalate of lime, are found in the genera Acalypha, Argyro-
thamnia, Caperonia, Claoxylon, Fragariopsis and Plukenetia, belonging to the
Acalypheae. .
Anomalous structural features are found in the axis of certain Euphorbia-
ceae; these cooiist in the occurrence of' I. intraxylary phloem (in Croton excl.
species of the section Astraea, iiI Crotonopsis, Eremocarpus and Iulacroton, of
the Tribe Crotoneae ; in Alchornea, Conceveiba, Mallotus pro parte and Pera, of
the Tribe Acalypheae; in Dactylostemon, M abea, Sene/eldera and Sebastiania
pro parte, of the Tribe Hippomaneae); II. secondary bundles of wood and
bast in the parenchymatous pericycle in species of Dalechampiq, and Fragariapsis;
III. inrerxylary phloem in Dalechampia; IV. medullary vascular bundles in
RicintA.
Finally, the following anatomical features are of minor systematic value:
gelatinization of the epidermis of the leaf; papillose differentiation of the
epidermis on the lower side of the leaf; development of hypoderm on the upper
side of the leaf; cork·wai'is on the leaf (A manoa) ; sderenchymatous fibres and
elongated spiral tracheids (Pogonophora) in the 'mesophyll; crystalloids and
sphaero·crystals of Wldetermined nature, &c.
2. STRUCTURE OF THE LEAF. The following description of the structure of
the leaf and axis is mainly based on the various researches which have been
carried out in Professor Radlkofer's botanical laboratory during the last few
years 1. The whole of the literature cited was also taken into account.
The anatomical fe<1tures of the leaf are very varied; this is to be expected
when the great extent of the Order is taken into consideration, and is well shown
in the structure of the integumental tissue. The lateral walls of the epidermal
cells may be either straight or undula ted. The thickness of the ou ter walls varies,
and is often considerable. In some cases the inner walls are strongly thickened,
e.g. in some species of Croton (Frombling), and also in some Hippomaneae (spedes
of Actinosteman, Givotia and Seneleldcra, according to Herbert). Rothdauscher
/'
I In the description of the Euphorbiaceae 1 follow the cla.ssification of Miiller Arg. in DC.
PrOOr., especially since the systemntic.anatomical papers, referred to above, followed the same
system of classification. It will therefore be appropriate to enumerate the ten Tribes of·Milller's
system, and at the same time to give an alphabetical list o( the genera or species, which have been
eumined anatomically, in each Tribe: 1. Caletieae: Caletia hexandra, Miill; Arg. and Pseuda"tkus
pz'meltows, Spreng. were examined by me.-II. Ricinocarpeae: Berlya gummiferQ, Milii. Arg.,
Beyeria VI'scosa, MUll. Arg. and Rict'nocaryus pinifolius, Desf. were examined by me.-IlI.
Ampereae: I examined Amperea spartiordes, Brongn.-IV. Phyllantheae: Frombling .and Roth·
dauschel' examined species of the genera: A clephiia, Amanoa, A ndrach m, AnHdesma, Aporosa,
Baccaurea, Buchojjia, Breynia, Cyclostemlm, £!iscocarpus, Drypetes, Hemicyclia, Hieronyma,
Hymmocardia, Laclmo!iYlis, Milanthesopsis, Pelaiostigma, Phylla nth us , Pu/ranjiva, Richeria,
Sauro/us, Sa'lJia, Secunneca.-V. Bridelieae: Bridt!/ia s/ipularis j Bl. was examined by me.-VI.
Crotoneae: Frombling investigated species of the genera Cro/Im, Cro/01UJpst"S, Eremocarjus, Jul0.
crolon a.nd .JI.ficrandra.-VII. Acalypheae: Rittershansen examined species of the genera Acalypha,
Atin'a1ta, Agroslis/achys, A lcltornea , Aleurites, Argyrolltamnia, Benttl1'difl, Caperonia, Cepltalo-
cfflm, Claoxylon, Clddz'lm, Cnesmone, Coccoceras, Coelodz'scus, Concevdba, Crozopkora, Dysopsis,
Fragan'opsis, Hevea, HomonoY4, flhannesia, Leptorhachis, l'tlacaranga, Alallotus, lllercun"aHs,
P(lchystroma, Pera, Plukene/ia, Bicindla, Ricinus, Sarcoclinium, Symphyllia, Tragia and Trewia.
VIII. Hippomaneae: Herbert investigated species of the genera Addoc,.oton, Actinostemon, Adeno·
dine, Ackmpelti's, Carumbium, Chat/oearyus, Cluytia, CoditUum, Colliguaya, .Dactylos/emon,
Eris,nantlzus, Excoecaria, Celom'um, GivoHa, Hippomane, Dura, latropha, l't-fabea, Man ikol ,
Maprounea, Alkrodesmis, Os/odes, Paradenocline, Pausandra, Pogollophora, Sebastiam·a. Seneftldera,
Stillingia and Trigonos/emon.-IX. Dalechampieae: Dalechampia has been examined by Ritter5~
hausen.-X. Varions Euphorbieae by several authors,
742 EUPHORBIACEAE

mentions a specially strong sclerotical differentiation of the outer wall (If

the epidermis as occurring on the lower side of the leaf in species of A 1nanoa
and Discocarpus. According to the same author marginal pits are found in
species of Aporosa, Bisc/zoffia, Cyclostemon, Drypetes and Hemicyclia. Another
noteworthy anatomical feature of the epidermis is ,its gelatinization, which is
very extensive on the upper side of the leaf in the investigated species of the
Caletieae. While the lower epidermis only includes isolated cells with muci-
laginous inner walls, the corresponding walls of the ,upper epidermal cells coalesce
to form a common mucilaginous layer. Amongst the Phyllantheae, Rothdauscher
met with mucilaginous epidermal cells in certain species of Actephila, Andrachne,
Antidesma, Aporosa, Hieronyma, Hymenocardia, Richeria, and Securinega.
whilst Frombling found them in species of Petalostigma and Phyllanthus; in
the species examined by Rothdauscher, the mucilage-cells are occasionally of
large size and spherical in shape; their lower ends penetrate into the mesophyll~
and frequently only a small portion of the wall reaches the surface of the leaf.
In the Crotoneae and Hippomaneae gelatinization of, the epidermis of the leaf
has not been observed, but it occurs in the Acalypheae (species of Acalypha,
Claoxylon, H omonoya, Leptorhachis, Pera and Tragia, according to Rittershausen),
and in the species of Bridelia (Tribe Bridelieae), examined by me. The nature
of the markings on the cuticle is also very varied; of these we need only men-
tion the occurrence of thick cuticular crests in species of Mabea (Herbert),
and of very characteristic markings on the lower side of the leaf in Aleurites,
Hevea and Johannesia (Rittershausen). According to Herbert, groups of
silicified epidermal cells are found in species of Actinostemon, Manihot, Pau-
sandra, Sebastiania and Trigonostemon; according to Rittershausen, rosettes
of silicified cells containing cystolith-like bodies (like those of the Boragineae,
but only silicified not calcified) occur in the epidermis of the, leaf in the species
of Bernardia in the immediate neighbourhood of the trichomes. Papillose
development of the epidermis of the leaf is also very widely distributed.
Amongst the Phyllantheae it has been observed on the lower side of the leaf
in Amanoa oblongi/olia, Mull. Arg., Securinega Acidothamnus, 1\ltill. Arg. and
S. obovata, Mtill. Arg. by Rothdauscher, and in numerous species of Phyllanthus,
and in Melanthesopsis fruticosa, MUll. Arg. by FrombIing. According to
Rittershausen, a number of Acalypheae show the same feature, viz.: Alc/ro"nea
Martiana, Miill. Arg., A. Schomburgkii, Klotzsch; Conceveiba guianensis,
Aubl.; Hevea discolor, Miill. Arg.; Homonoya riparia, Lour.; Johannesia
princeps, VeIl. (only the subsidiary cells of the stomata); Macaranga gtJmmi-
flua, Miill. Arg., M. Hel/eri, MUll. Arg., M. 1'avanica, Milll. Arg. p genuina (in
the neighbourhood of the glandular hairs), M. indica, Wight, M. min utiflo ra,
Miill. Arg., M. PoPuli/olia, Mull. Arg., M. tomentosa, Wight; MallotUs' flO1'i-
bundus, Miill. Arg. ;-in }"facaranga triloba, Milll. Arg., ~fallotus ricinoides,
Milll. Arg. and Ricinus communis, L. the epidermal cells on the upper side of
the leaf are subpapillose. Finally, the lower epidermis of the leaf is entirely
or partially papillose in some Hippomaneae, according to Herbert, namely:
Carumbittm populneum, Mtill. Arg.; Cluytia daphnoides, Milll. Arg.; Mabea
angusti/olia, Benth. tl genuina, M. fistuli/era, Mart.; Manihot coerultscens,
Mull. Arg. f3 genuina, M. palmata, MUll. Arg. a ditJusa. M. pilosa, Pobl, M.
trip artita , Miill. Arg. f3 porrecta, }"f. utilisst'ma, Pohl; M aprounea guianensis.
Aubl.; Stillingia discolor, Champ. et Benth., S. nutans, Vahl and S. sebifera,
Michx.. In Euphorbia bttxifolia, Lam. the upper epidermis of the leaf is papil·
lose (Warming). The shape of the papillae varies, e.g. capitate in Mapf()Unea
guianensis, in other cases frequently long and finger-shaped. Transitions from
the latter type of papillae to simple, unicellular trichomes are sometimes
found. Such transitional forms have been observed by Frombling in PhyUan-
thus praetervisus, Mull. Arg. and by Herbert in Sebastiania daphnoides, Miill.
 EUPHORBIACEAE 743
Arg. a myrtiUoides. Another noteworthy feature is the occurrence oi hypoderm
on the upper side of the leaf. Amongst the Phyllanth~ae a single layer of
hypoderm is found, according to Rothdauscher, in Bischotfia javanica, Bl.~
Cyclostemon Cumingii, Baill., Hemicyclia andamanica, Kurz, and locally also
in Hymenocardia acida, Tul.; according to Frombling, in Phyllanthus indicus,
Miill. Arg., P. obovatus, Milll. Arg. and P. puberus, Mull. Ar$'" and locally also
in certain other species of Phyllanthus. Rittershausen mentions the occurrence
of a similar hypoderm in the following Acalypheae: Agrostistachys indica,
MUll. Arg., Aleurites moluccana, \Villd., Pachystroma ilicifolium, Milll. Arg.;
amongst the Hippomaneae Herbert fowd a hypoderm in all the investigated
species of Celonium and in Excoecaria Agallocha, Milll. Arg. The occurrence
of structures resembling cork-warts can be demonstrated on the lower side of
the leaf in Amanoa oblongifolia, MUll. Arg. (Phyllantheae). Other special
features of the integumental tissue, such as secretory cells (in Ricinus) and
crystals (in representatives of the Ampereae, Crotoneae, Acalypheae and
Hlppornaneae) will be fully treated in the sections dealing with those subjects.
Concerning the stomata, we may first mention that in most of the-Tribes
(excluding the Crotoneae) they are usually confined to the lower side of the
leaf, occurring more rarely on both surfaces. In the leaf of Euphorbia buxi-
folia~ Lam., in which the lower half of the mesophyll is occupied by aqueous
tissue, the stomata are found only on the upper side of the leaf (Warming),
In very many Euphorbiaceae the stomatal apparatus belongs to the Rubiaceous
type, the stomata being accompanied on elther side by one or more sub-
sidiary cells, which are placed parallel to the pore. In other cases, however,
the Cruciferous type occurs, or the pairs of guard-cells may be surrounded by
a relatively large number of cells, exhibiting no special arrangement. Regard-
ing the special distribution of the Rubiaceous type, &c., amongst the Euphor-
biaceae, the following facts may be mentioned. In the Arnpereae (Amperea)
the sto~ata are mostly surrounded by three epidermal cells. In the Ph,llin-
theae -(excl. Euphyllantheae) examined by Rothdauscher, the. Rubiaceous
type preponderates; Andrachne, Aporosa, Baccaurea, Lachnostylis and Richeria
constitute the sole exceptions, the Cruciferous type being more or less distinct
in these genera. In the Euphyllantheae Frombling found no unifornl' type
of stoma, for the cells adjacent to the guard-cells in this Tribe appear as sub-
sidiary cells either of the Cruciferous or Rubiaceous types, both types often
occurring side by side on the same leaf-surface. In the Acalypheae and
Dalechampieae, ac~ording to Rittershausen,/the Rubiaceous type is present
in all cases, and the same (exception: Manihot caerutescens, Miill, Arg. fJ
genuina) applies almost throughout the Hippomaneae, according to Herbert.
In the _Euphorbieae Benecke mentions the occurrence of the Rubiaceous type
in Mercun'aUs and Euphorbia. It is impossible to enter into details here
regarding the varied mode of insertion of the guard-cells, i.e, whether they
are depressed, not depressed, or elevated. Only one noteworthy feature will
be mentioned, that shown by the stomatal apparatus in Caletia and Pseudanthus,
or, to be more preciserby the cells-which adjoin the guard-cells and penetrate
beneath them. The walls of these cells, which run at right angles to the surface
of the leaf, and limit the outermost part of the respiratory cavity, appear,
when examined in a superficial section, to be folded in a precisely similar
manner to the well-known arm-palisade-cells (of Haberlandt) of the type with
numerous arms. The object of this arrangement, which has also been observed
in other Orders of plants, remains to be determined.
The leaf-structure may be either centric or bifacial; for details reference
must be made to the special papers cited. I have met with rolled leaves having
two furrows, one on either side of the median vein, and showing a corresponding
type of structure, in the species of Bertya and Ricinocarpus mentioned above.
 744 EUPHORBIACEAE

The following spe~tal features o~ the mesophyll ~re worth mentio!ting: t~~
occurrence of a kind of arm.palisade-parenchyma m PhyUan~hus Roxbul'gha..",
Miill. Arg.; the peculiar differentiation of the lowest cell-layer 01 the spongy
tissue in M icrodesmis caseariaetolia, Planch. ; and the occurrence of sc1erenchy.-
matoUs elements running freely in the mesoph}'ll, of spiral tracheids, and of
enlarged terminal tracheids. In the species of J\;!icrodesmis mentioned above,
the layer of cells referred to is strongly thickened on the side facing the
interior of the leaf. Sclerenchymatous fibres, running freely in the leaf, are
found chiefly in the Acalypheae, viz. according to Rittershausen in: Acalypha
diversifolia, Milll. Arg. a popoyaensis; Alchornea .,:lici/olia, Miill. Arg., A.
latilolia, Sw., A. Martiana, Milll. Arg. (according to loco cit., p. 38), A. Schom-
burgkii, Klotzsch; Bernardia axillaris, Milll. Arg. f3 genuina, B. celastrinea,
Miill. Arg., B. gambosa, Milll. Arg., B. scabra, Miill, Arg.; Conceveiba guianensis,
Aubl., C. trigonocarpa, Mull. Arg.; Mallotus Lawii, Miill. Arg.; Pera a'niso-
tricha, Milll. Arg., P. coccinea, Miill. Arg., P. distichophylla, Baill., P. heterodoxa.,
Miill. Arg.; Sarcoclinium longitotium, Wight ;-also according to the same
author amongst the Dalechampieae in Dalechampia atfinis, MUll. Arg., D.
bra,siliensis, Lam., D. capensi~, Spreng. f., D .. cap~onioi.~es, ~iill. Arg., D. /ici-
lolsa, Lam., D. Leandn, Baill., D. magnoltaefol"a, Miill. Arg., D. scan dens,
Miill. Arg. f3 tallax, D. semitriloba, Tausch, and D. triphylla, Mull. Arg. The
same kind of sclerenchymatous -fibres, frequently snowing well-marked strati~
fication of their walls, have also been observed by Herbert in the following
Hippomaneae: Actinostemon concolor, Miill. Arg.; Chaetocarpus castaneae-
carpus, Thw" C. Pohlii, Miill. Arg.; Erismanthus obliqu~,s, Milll. Arg.; Pau-
sandra Aforisiana, Radlk.; Sebastiania brasiliensis, Milll. Arg., S. multiramea,
Milll. Arg.; Trigonostemon lauritolius, Baill.. ;-and finally Rothdauscher met
with them in Actephila lati/oUa, Benth. (Phyllantheae). Here we may
mention the sclerenchymatous cells (rod-cells and sclerenchymatous fibres),
, which occur in connexion with the trichomes in many Crotoneae; they
will be dealt with more fully in the description of the hairs. The spiral
tracheids mentioned above occur only in the spongy tissue of Pogonophora
Schomburgkiana, Miers; they have a tubular shape, are provided with a very
delicate spiral band, and are about 40 J-t in breadth. Herbert has shown that
enlarged terminal tracheids occur in species of the genera Givotia, ..'~fabea,
Sebastiania and Stillingia (of the Hippomaneae). The special crystal-idio-
blasts of the mesophyll will be described in the section dealing with the crystal ..
line elements.
In the structure of the veins we may first point out that the vascular
bundles mayor may not be accompanied by sclerenchymatous tissue.
In some species of Euphorbia (E. bahiensis, Boiss. and E. buxilolia, Lam.)
the bundles are surrounded by a parenchymatous sheath of large, wiele cells
(Warming). The smaller veins are either embedded or vertically trans-
current. In the following members of the Order the latter is the case:
amongst the Phyllantheae the genera A manoa, Discocarpus, Hymenocaydia~
Lachnostylis, Richeria and Sa via, and some of the species of Aporosa~
H emicyclia and Securinega; amongst the Crotoneae some of the species of
Croton; amongst the Acalypheae Alchornea, Conceveiba, Macaranga, MaUotus.
Trewia, and Bernardia celastrinea, Mill!. Arg., and Hevea (here vertically trans-
current on the upper side only); amongst the Hippomaneae Hippomane and
species of M anihot and Stillingia.
Oxalate of lime is generally excreted in the form of clustered and solitary
crystals. Both forms may occur side by ,side in the same plant; in other
cases there are only clustered crystals, or (extremely rarely, in Pachystroma)
only solitary crystals, or oxalate of lime may be entirely absent. The following
special forms of crystals of oxalate of lime have been observed in this Order·~
 EUPHORBIACEAE 745-
,
sphaeritt:s, small rod-shaped crystals, often bent like a V or W, and styloids.
Amongst the Crotoneae Frombling met with sphaerites of oxalate of lime in
the mesopbyll in C,otonopsis, Eremocarpus, Croton ~aPitatus, Milll. Arg. and
C. monanthogynus) Michx.; of the AcalYl>heae Acalypha and Claoxylon like-
wise possesses sphaerites, which lie in hemlspherical, projecting epidermal cells
of the axis and leaf (Rittershausen); lastly, they are also found in the pith of
Gelonium, according to Herbert. The small rod-shaped crystals, which are bent
like a V or yv, and recall the analogous forms of crystals found in the Papillo-
naceae, have only been observed in the mcsophyll of some species of PhyUanthus.
Typically differentiated styloids are found in the bast in species of Phytlanthus,
in the leaf and axis in Pachystroma, and in the bast of Bernardia; styloids,
showing less typical differentiation, occur in the bast in Colliguaya and Excoe-
caria. The terminal cells of the stinging-hairs of Cnesmone, Dalechampia,
Leptorhachis and Tragia contain crystals resembling styloids; furthtr details
will be given below. The ordinary crystalline elements of oxalate of lime,
especially the clustered crystals, more rarely the solitary crystals, are con-
tained in special idioblasts, in certain members of the Order; these idioblasts
are distinguished either by their large or small size, sometimes also by their
shape. Very frequently they are found in the epidermis of the leaf, either
singly or arranged in small groups, and they are often characterized by th(!ir
sman size and their rounded outline in surface-view.
Amongst the Ampereae I met with roundish crystal-idioblasts in both the
upper and lower epidermis of Amperea spart£oides, Brongn.; the clustered crystals,
which they contain, are embedded in a thickening of the inner wall. Amongst
the Crotoneae clustered crystals in the epidermis of the leaf have only been observed
in Julocroton triquete'Y, MUll. Arg. On the other hand they are present in many
Acalypheae and Hippomaneae. Amongst the Acalypheae Agrostistachys, Aleurites,
Cleidion, Coelodiscus, Conceveiba pro parte. Homonoya and Trewia possess idio-
blasts, which contain clustered crystals, but otherv.'ise differ little from the rest of
the epidermal cells; in A calypha pro parte, A Ichornea, Claoxylon, Datechamp'ia
and Mallotus the epidermal cells containing the clustered crystals are of consider-
able size, and con~quently penetrate into the mesophyll; the idioblasts of Argyro-
thamnia and CapeYonia, which also contain clustered crystals, are distinguished
by having a smaller~ roundish lumen~ and by being arranged in small groups! In some
other Acalypheae, the idioblasts, containing clustered crystals, project like papillae
or hairs, but these will be dealt with in the description of the hairy covering.
Amongst.the Acalypheae solitary crystals in the epidermis of the leaf have hitherto
only been observed in Cnesmone iavanica, Bl. and in Dalechampia..§candens, Miill.
Arg. {3 ta#ax. Amongst the Hippomaneae small idioblasts, contJting clustered
crystals, have been shown to occur in the epidermis of the leaf in species of H,·ppo-
mane. M anihot. Ostodes and Pausand'Ya. Herbert, who investigated this Tribe,
was not able in all cases to determine with certainty whether the -large crystal-
idioblasts. which occur in other Hippomaneae, belong exclusively to the palisade-
tissue or are epidermal cells depressed to the level of the palisade; these idioblasts
either contain clustered crystals (Acidocroton. Jatropha pro parte, Givo#a) or
solitary crystals (Erismanthus, MicYodesmis, Pogonophora, Stillingia pro parte),
the latter_ qeing freq~_ep..!Jy inserted in a thickening of the inner wall.
Large crystal-idioblasts, which for the most part have a suberized wall,
and are occupied by clustered or solitary crystals, are equally common in the
mesophyll, and sometimes give rise to transparent dots in the leaf 1.
The mesophyll in all the investigated Crotoneae (Frombling) contains large
crystal-idioblasts with suberized walls; in the majority of the Crotoneae large

I Transparent dots in the leaf in the Euphorbiaceae are also caused by tecretcry cells and
lacunae (see belo\v), and, in certain specie!> of Excoeco.ria and Euphorbia, by special features of the
mesopbYll, viz. respiratory cavities on the upper side of the leaf, and the. meshes ~n tbe network ot
veins (Radlkofer).
 EUPHORBIACEAE

clustered crystals are found in these elements, while in Crotonopsis, Eremacarpus,

Croton capt"tatus, Miill. Arg. and C. fflonanthogy1Cus, Michx. the latter contain the
sphaerites mentioned above. In the Acalypheae also, as far as can be gathered
from Rittershausen's statements, this feature is not rare, being found for example
in Ricinella, in which Radlkofer first pointed ou~ the pellucid dots caused by the
tlustered crystals. According to my own observation I may add that large sacs,
filled with solitary crystals, are present in the palisade-parenchyma of l1ridetia
s#pulan's, Bl. (Tribe Bridelieae),
A third special feature presented by the crystalline elements ~ the leaf
is the occurrence of small clustered crystals in chambered cells of the pa1isade-
tissue; this feature is recorded by Rittershausen in species of Alchornea,
Cephalocroton and Hevea.
Having dealt with the oxalate of lime we may add a few words on the
occurrence of crystalloids, crystals of abietinic acid, sphaerites in alcohol
material of Euphorbias, and fatty bodies. Crystalloids have been obs~rved
by Fry in the stem of Euphorbia splendens, where they occur in the inner cell-
layers of the primary cortex, and also in the Ie,af (mainly in the mesophyll
above the vascular bundles of the veins); beautiful crystalloids are also found
in the latex of Jatropha (see p. 747). Pax mentions the oCCurrence of crystals
having the shape of a whets"tone and consisting of abietinic acid in the cortical
tissue of Euphorbia splendens. Sphaerites are met with in alcohol material
in the tissues of the Cactus-like Euphorbias, e.g. E. Caput Medusae, and have
been investigated by variQus authors (Leitgeb, Rodier, Schaarschmidt, Hansen
and Belzung). To judge from a microchemical examination carried out by
Hansen, they consist of calcium phosphate, but according to a macrochemical
investigation by Belzung, they are composed partly of calcium malophos-
phate, and partly of calcium malate. Fatty bodies are present in the mesophyll
in many members of the Order.
The secretory elements of the Euphorbiaceae require a detailed description,
especially as some of them were not quite correctly 1 interpreted by Pax (in
Engler, Bot. Jahrb. 1884, and Natiir!' Pflanzenfam.). The following types
of secretory elements are found in the Euphorbiaceae, and will be described
in order: I. Laticiferous cells; II. Laticiferous vessels; III. Rows of latici-
ferous sacs; IV. Tannin-sacs and rows of tannin-cells (' g~gliederte Milchsaft-
rohren' and f gegliederte Milchsaftschlauche ' of Pax); V. Elongated secretory
sacs with wide lumina, and usually with brown contents; VI. Secretory cells;
VII, Secretory lacunae; VIII. Mucilage-lacunae. I

1. The laticiferous cells are confined to the tribes Crotoneae, Acalypheae,

Hippomaneae and Euphorbieae. Amongst the Crotoneae they are only found
in some of the species of Croton 2 (Frombling); in the tribe Acalypheae only
1 It would be out of place here to enter into a detailed criticism of Pu's statements: two points
only mny be shortly referred to. The elements described by Pa.x as laticifcrOllS vessels (' gegliederte
Milchrohren ') or articulated sacs Cgegliederte Schlauche ') do not correspond to what a.re called
laticiferous \tessels (I gegliederte Milchrohren') in the plant· anatomy of the present day, but are only
rOWi of cells with distinct transverse waIfs and speciaJ contents (mostly brown a.nd tanniniferous) j more-
over the laticiferous nature of these contents in the living plant has 110t 1tt been proved either by Pax
or by other observers. In the second place the following statement on p. 5 in th~Natiirl. Pflanzen-
familien js only partially Correct: 'haafig werden die Querwande resorhiert und <lie Milchrobren
nehmem in bohem Grade das Aussehen ungegliederter Schliiuche an, so bei den Jatrophene, Manihoteae
und Crotoneae: This is shown by the fact thAt laticiferons vessels are only known in Hevea and
Manihot, while Jatropka, Croton, &.c., possess true laticiferons cells.-Similarly.in view of the wid~
distribution and general use of Moller's book on the anatomy of the Cl)rtex it seems advisable to refer
to the partly incorrect terminology of the secretory organs in the Eupborbiacene employed in this
book also. The laticiferous cells are termed l:lticiferous canals in the bllSt of A.ndira brasilimsis
( = JQliann,-sia prilueps), se~etory sacs in tbe cortex of Jatropha Curcas, and Iaticiferous sacs in the
bast of Baloghia lucida (= Codiaemn lucidum). _
a Namely Croton tu:knophy//us, Spreng., C. agrarius, Mull. Arg" C. argyranthtlnus, Michx.,
C. Cojil<.a1'l~, Benth., C. CaJ{ol'i/lQ, BellD., C. ullidijoliu.I, Bam., C. dlrj'.f«ladu.I, MUll. Arg.,
 EUPHORBIACEAE 747
in the genera Aleurites, Johannesia, Macaranga and Pachystroma (but according
to my own observations not in Alchornea and M allotus, where they are stated
to occur by Rittershausen); in the Tribe Hippomaneae in the genera Actino-
stemon, Adenocline, Adenopeltis, Codiaeum, Cotliguaya, Dactylostemon, Exco ecaria ,
Hippomane, Hura, Jatropha, Mabea, Maprounea, Ostodes, Para de nocline,
Sebastiania, Senejeldera, Stillingia and Trigonostemon (Herbert). In the Tribe
Euphorbieae the distribution of the latidferous cells has been little investi-
gated, and. hitherto these elements have only been demonstrated in Euphorbia,
in Mercurialis and Ricinus (llanstein), and Poinsettia (Pirotta and Marcatili).
Regarding their course of development, Hanstein and Chauveaud have shown
that they are differentiated at a very early stage in the plane of the cotyledonary
node. their initial cells bein& situated in the outermost cclI-layer of the central
cylinder; the initial cells elther compose the whole of this layer, or they are
arranged in it in the form of four arcs; in Croton a second ring of initial cells
is found in the middle of the primary cortex. The course of the laticiferous
tubes in the mature plant is as follows: in the axis they occur in the pith .,
bast, and primary cortex, whilst in the leaf they are found in the veins and
sometimes II also free in the mesophyll. The laticiferous tubes, which run
freely in the mesophyll, sometimes exhibit a very striking relation to the assimila-
tory system (Haberlandt), and not uncommonly extend as far as the epidermis
of the leaf. The wall of the laticiferous tubes is sometimes (e.g: in the fleshy
Euplwrhias) exceptionally thick, and is often provided with large pits. The
thick transverse walls, observed by De Bary and Dippel in the older latici·
ferous tubes of Euphorbia Lathyris, appear to be secondary formations. The
contents are generally milky in the living plant; in herbarium-material they
have a brown or grey colour. Dietz has shown that a number of crystal-
lizable organic substances are present in the coagulated latex of the Euphorbias ;
the latex also frequently contClins tannin. Treub found nuclei in the latid-
ferous tubes of the Euphorbias. The following special substances are found
in the contents of the laticiferous tubes: clustered crystals of oxalate of lime
(in Hura a-qd Mabea, according to Herbert); crystalloids having the form of
tetragonal flakes with rounded corners (in Jatropha, according to Trecul and
Herbert); finally, rod- or bone~shaped starch-grains (in Euphorbia splendens
and other species, and also in Hura crepitans, Milll. Arg.).

C. dliato--glantiulosus. Ortega, c. com()sus, Miill. Arg., C, Corlest'anus, H. B. K., C.d,·sc%r, Willd.,

C. exukrans. MUll. Arg., C._/lavens, Miill. Arg., C. ,Pr£onis, MUll. Aig., C.jruticulos'Us, Mijll. Arg.,
C. gracilipes, Baill., C. kumilis, L., C. inccrtus, Miill. Arg., C, Klotucl:a"£, Mull. Arg., C. /a.dzft()-
clad:Js, Mart., C.limarifolius, Mull. Arg., C. /inearis, Jacq., C. lobatus, Miill. Arg., C. Luntiianus,
!'rHin. Arg., C. morito/ius, Mull. Arg., C. origanifoUus, Miill. Arg., C. pallidus, Miill. Arg., C. pana-
menm, Miill. Arg, C. paraensi's, ""Mull. Arg., C. pauli"us, MiilL Arg., C. pedice1!atus, H. B. K.,
C. Pok!ianus, Mijll. Arg., C. pulegiodorus, Baill •• C. pungtns, MUll. Arg., C. rkamnifolius, Mull.
Arg., C. Sapotanus, Mull. Arg., C. scltrocalyx, Mull. Arg., C. semitJ~stilUSJ Mlill. Arg., C. Soliman,
Cham. et Schlecht., C. stipulaceus, II. D. K., C. sUDacultls, Mlill. Arg., C. subv!'llosus, Miill. Arg.,
c. tridentalus, Mart., C. Urucurana, BaH1., C. 1/a",'utn'anus, Baill., C. veprelorum, Miill. Arg.,
C. Wilsonii, Griseb., C. xalapensu, H. B. K.
1 The 11lticiCerons cells are never found soiely in the pith. Rittershausen's statements regarding
their occnrring e.s:tlusively in the pi~h in Alchornea, Altudtes and Mallatu! require ccrrectioD, as
shown by the following data: firstly, in Alch.ornea (according to my C'w'll exa.mination of A. cfWdata,
Milll. Arg.) laticiferous tubes are not present-elongated secretory sacs may possibly be present in
some of the species examined by R.ittershauscn, and, if so, were incorrectly interpreted by him;
secondly, the latieiferous cells of Aleurites (A. mo/uccana, Willd.) occur not only in the pith, but
also in the bast and primary cortex; and thirdly, the laticiferous cells, mentioned by Rittershausen
as occurring in the pith of Mal/o/us, are really secretory sacs (see under V) according to my own
examination of M. oreojMlus, Miill. Arg.
3 According to Rittershausen, amongst the Acalypheae in P~k)lstY~ma only; according to
Frombling, in almost all species of C,.otOIl; according to Herbert, quite generall y in the H!ppomaneae;
according to Marcatili and Pirotta, in EupluJrlJia and Poinsettia (c(. also Haberlandt, Groom and
Scott, II. ce.).
 EUPHORBIACEAE
II. The laticiferous vessels are onlyfou.nd in two genera, viz. Hevea (well-
known caoutchouc-plants) and Manihot, where they were first observed by
Scott, Calvert and Boodle. They occur chiefly in the bast of the vascular

FIG. tP.o (aj. A, Crat.eri(orm seeretory cell of Croitm mona,,'J,"r1"us, MichJ:. lI~.Pyriform ~cretorv cell
or i"/(}cr%n fususce"s, Baill. c, Secretory cell in the leaf of C.,.,Jlon e~"ilus. Mull. Arg-. D-F, Stellate
~r tufted hails with secretMy cells at their base: D, CYoltJn subacul_s. Mull. Arg.; It-F, C. pule£iodI)YU.I....Baill.
Q-t. Various trichomes: G, C. eXUOeYdnS, Mull. Arg.; H. C. a~I!"s;s, Haill.· I. /uiOt:rolon /U.lcuuns, naill.;
k, Crow" discolo,., W.; L, C. jfo"'ihu"du~ Spreng. M,·Patelliform gland of t.rJdndulosus, }Jlill. Arg., section.
H. Glandular mat_:gin of the leaf of C. ".eJrtUtzu, Milil. Arg. in section. 0, Morulose gland of C. dliatO-,flandu.
1tiIW, ou. P-Q. Stinging hairs of TraK"_iJ ciSSDi(Us, Mull. Arg. R--S, Hairs containing clll.'Itered crystals, from
Plllleefteli'a.-A-O. G-H and M-{) after FromLling, E-f", I-K and P-Q Original. L after Bachmann, R-S after
R itt~rshal1sen.

bundles in the axis and leaf; but they have also been met with in thelrimary
cortex in Hevea (Rittershausen), and at the margin of the pith an in the
 EUPHORBIACEAE 749
primary cortex m Manihot. In tne living plant their contents are milky in
HefJea, of the nature of a watery sap m Manihot. Nuclei have been
demonstrated in the laticiferous tubes of M anihot.

··•
I

•
·
I

FIG. l~ (6).

III. The rows of latic.iferous sacs are confined to the genus M1-cyandra
(Frombling). The cells forming these rows have rather wide lumina, show
various degrees of elongation, and in herbarium.material are filled with granular
 EUPHORBIACEAE

contents having a grey colour. In the axis, they are present in the pith, bast,
and primary cortexp while in the leaf they run independently of the veins~
being free in the mesophyU, where they form long chains of cells. The expres-
sion 'rows of laticiferous sacs' is justified by the fact that the species of
Micrandra are described as trees with latex, and that other secretory organs
are not present. _
IV. Elements very widely distributed in almost all the Tribes are cells
filled with brown, tanniniferous contents (in herbarium material); these
elements are often distinguished from the adjoining cells by the size of their
lumina, or by their greater elongation, and either occur singly or united in
groups; they are chiefly found in the bast, but are also present in the pith
and primary cortex of the branch. Pax terms these elements articulated sacs
(gegliederte SchHiuche), articulated laticiferous sacs (gegliederte Milchsaft-
schHiuche), or even (inadmissibly) articulated laticiferous tubes (gegliederte
Milchsaftrohren), and ascribes great systematic importance to them for the
subdivision of the Euphorbiaceae according to his scheme of classification.
According to more recent observers, and my own 'views also, Pax has certainly
gone too far·in this respect. The elements in question are frequently connected
with ordinary tanniniferous.cells by transitional forms, so that it is often difficult
to decide vlhether the tanniniferous cells of a given plant should be regarded
as Pax's elements or not. The contents of these elements, at any rate in
herbarium-material, give no more assistance than does their shape towards
making a clear diagnosis; moreover not a single case has hitherto been
recorded in which the contents have proved to be characteristic an~ of the
nature of latex, in the livjng plant.
V. The fifth kind of secretory organ is formed by elongated sacs, which
mostly have wide lumina, and are generally filled with brown contents in
herbarium-material. They are often very long, and are then easily confounded
with laticiferous cells, unless their development is investigated, especially as
the laticiferous cells often possess similar brown contents. In order that such
mistakes may be avoided, it maybe noted that the secretory sacs usually occur
solely in the pith. while this is never the case with the laticiferous cells, and
that they generally have a larger diameter (reaching 95 IL) than the latter.
In some cases (M allotus oreophilus, Milll. Arg.) the secretory sacs in a trans'
verse section of the branch are surrounded by a ring of cells, having small
lumina and resembling an epithelium. They occur in the pith, and sometimes
also in the bast, perkycle, and primary'cortex. They have been observed in
species of M aUotus and Alchornea (Tribe Acalypheae), and species of G1:votiai
Pausandra and Pogonophora (Tribe Hippomaneae, according to Herbert).
VI. Secretory cells, viz. idioblasts with oily or resinous contents, are only
known to occur in Ricinus, and in the genera Croton, ]ulocroton, Croto1'topsis
and Eremocarpus of the Crotoneae, that is to say in all the Eucrotoneae (with
the exception of a few species of Croton, belonging to the section Ast,aea).
In the Eucrotoneae they frequently give rise to transparent dots in the leaf.
The secretory cells of Ricinus communis, L. are large cells present in the epider-
mis on both sides of the leaf, and penetrating somewhat into the mesophyll,
whilst only a small portion of the wall of these cells reaches the surface of the
leaf; they are m-ore commonly met with on the under side of the lamina, where
they usually have an elongated shape; they contain a secretion, which is
yellowish-brown in the dried plant, but bright and strongly refractive in the
living plant. The following statements may be made regarding the secretory
cells of· the Euctotoneae. They are absent in only a few species of Croton
belonging to the section Astraea (Co' comosus, Miill.. Arg., C. Klotzschii, Milll.
Arg., C. lofJatus, Miill. Arg. and C. paul";nus, MOO. Arg.); a further peculiarity
of these species is the absence of the intraxylary soft bast characteristic of the
 EUPHORBIACEAE 75 I
Eucrotoneae, and for these reasons they should probably be excluded from
the latter Tribe. In herbarium~material the secretory cells of the Eucrotoneae
have bright~yellow translucent, oily contents readily soluble in alcohol and
ether; their walls are suberized; in many cases they emit an aromatic
scent, ,,:,hich is usually agreeable, and is recognizable even in herbarium-
material. They are found both in the leaf and axis. In the axis, they occur
in the epidermis, pith and primary cortex, and also in the bast; in the first
three tissues they are mostly spherical in shape, while the secretory cells in the
phloem are elongated and tubular. In the leaf, they are found in the epidermis
and mesophyll. In the latter tiss~e they have a spherical or long oval shape,
according as they belong to the spongy or palisade-tissue. The epidermal
secretory cells vary greatly in shape, and according to Frombling the following
forms may be distinguished: (1) secretory cells, situated on a level with the
epidermis, and only distinguished from the rest of the epidermal cells by their
spherical shape and their contents; (2) secretory cells projecting above the
epidermis as papillae; but again possessing a completely spherical shape;
(3) rapillose secretory cells, projecting more strongly, and either craterifonn
(Fig. 180, A) or pyriform (Fig. lBo, B); (4) depressed secretory cells, lying on
a lower level than the remaining epidermal cells; only a small portion of the
wall of these elements reaches the surface of the leaf, and thus limits part of
the pit-like depression situated above the secretory cell; the latter has a
spherical or long tubular shape and penetrates into the mesophyll (Fig. 180, C).
Regarding the distribution of the secretory'cells in the tissues of the leaf, the
following points remain to be mentioned. The secretory cells are found in the
epidermis of the leaf 1 in all species. They are never restricted to the upper
epidermis, for when present there they are also found in the lower epidermis.
They are of rare occurrence in the mesophyll (Croton amabilis, Miill. Arg.,
C. compressus, Lam., C. corylijolius, Lam., C. gratissimus, Burch., C. hemiar~
gyreus, Mull. Arg., C. Mubango, Miill. Arg. and C. reticulatus, Miill. Arg., and
also Julocroton fuscescens, Baill.). A specially noteworthy feature is the
presence of secretory cells in the epidermis of the":'pedestals of the stellate and
tufted hairs (Fig. lBo, D-F). It is impossible to establish a definite rul~ regard-
ing the distribution of the various forms of epidermal secretory cells, since
different types are frequently present side by side in the same plant. According
to Frombling, however, tubular secretory cells traversing the entire thickness
of the leaf, and those of crateriform shape, appear to be characteristic of definite
groups of species; this is seen to be the case if the list of species ~ enumerated
for this purpose in the foot-note below be compared with the classification
given by Muller Arg. It may be added that the crateriform secretory cells
are in most cases (exception Croton astroites, Milll. Arg.) found only on the

J Frombling's statement (on p. +9 of his treatise) that in Croton reticula/us secretory cells occur
only in the spongy tisstIe is a mistake, which may be corrected here.
, The following s~i~ posses_s crateriform secretory cells: Croton agoensis, BaHI., C. amabilis,
Miilt. Arg., C. as/roiles; -Miill. Arg., C. betulin us, Vahl, C. buxijolius, Miill. Arg., C. capitatus,
Milll. Arg., C. Calinganus, Miill. Arg., C. ceanothijo/ius, Baill., C. cneorifolius, MUll. Arg., C. C(l1ll-
fruS'US, Lam., C. Corc~psis, Ba.ill. t C. jlgri/>undus, Spreng., C. Gaudichaudi, Baill., C. gratissi-
mw, Burch., C. IumiargyreuI, Miill. Arg., C. longz'ntrvi1lS, Mull. Arg., C. man'timus, Walt., C.
mkans, Mull. Arg., C. mi'grans, Casar., C. Muban~, Miill. Arg., C. Palanostt"gm4, Klotzsch,
C. tenellus, Miill. Arg., C. verbenaefo/iUl, MUll. Arg., C. Widp-enianus, Mull. Arg. ; Julocrot"n
argenteust Fr. Didrichs., J. fuscesuns, Baill., J. stijularis, Miill. Arg., J. trifjueter, Mull. Arg. ;
Erem~ar'pus uti'gerus, Benth. Elongated secretory cells, which traverse a portion, or often the
whole of the mesophyll, have been observed in Croton Bentlzamianus, Miill. Arg., C. Betulasler,
Mull. Arg., C. arinIJ·dentalus, Miill. Arg., C. dicM/omus, Mull. Arg., C. erel1lophilus, Mull. Arg.,
C. g/utinosus, Mull. Arg., C. ,rrewiaefolius, Mull. Arg., C. Rja/marson"", Griseb., C. lucidus, L.,
C. Lundianus, Miill. Arg., C. 'lna/ourens;s, Miill. Arg., C. muscicapa, Miill. Arg., C. oxyphyllttt,
Mull. Arg., C. sc/tro,a/yx, Miill. Arg., C. sincorensis, Mart., C. timam/rIJides, Mull. Arg., C.
7.',"rgultosus, Mull. Arg.
75 2 EUPHORBJACEAE

lower side of the leaf, and only in species the leaves of which bear a dense
hairy covering. Regarding the general distribution of the forms of secretory
cells described above amongst the genera of the Eucrotoneae, the following
statements may be taken from Frombling's work. In Croton all the different
types occur, viz. secretory cells in the mesophyll (in a few of the species, see
above), and the various forms of secretory cells found in the. epidermis; in
Julocroton there are secretory cells in the mesophyU (Julocroton fuscescens,
Baill.), and those of crateriform shape in the epidermis; in Crotonopsis the
second and fourth types of epidermal secretory cells are present; finally in
Eremocarpus, crateriform secretory cells only.
VII. The secretory lacunae, observed by Herbert in almost all species
of the two genera Cluytia and Gelonium (exception: Cluytia alaternoides,
Mull. Arg.) belonging to the Hippomaneae, are unique; they frequently
give rise to well·marked transparent dots in the ,leaf, and their structure
recalls that of the intercellular secretory receptacles of Derris, Lonchocarpus
and other Leguminosae. The spongy tissue of ,the two genera mentioned
above contains rather large intercellular spaces, filled with a colourless,
or yellow, or brown secretion, and surrounded by secretory cells, which for
the most part have a serpentine form. Closed intercellular secretory spaces,
provided with an epithelium~ have not been observed~
VIII. Mucilage.lacunae, or rather groups of cells with mucilaginous walls,
were first met with by Vesque in the primary cortex of BischotJia javanica,
and subsequently by Rothd~uscher in other Phyllantheae also (viz. species of
A ntidesma, Baccaferea, Hieronyma, Hymenocardia and Securinega), and by
.Rittershausen in Adriana (Acalypheae).
Having described the secretory organs; we may now deal with the tri·
chomes, which may be divided in to three groups: I. Clothing hairs; I I. External
glands; III. Stinging hairs (including the hairs containing clustered crystals).
1. The clothing hairs include a large number of forms. In the simplest
case they are unbranched unicellular or uniseriate structures. A type of
clothing hairs found by Rothdauscher in the Phyllantheae consisted of simple
and unicellular or septate hairs, the latter being divided by a few walls;
according to my own investigation Bridelia stipularis, Bl. (Tribe Bridelieae)
possesses uniseriate trichomes; the Crotoneae, according to Frombling, have
unicellular hairs, but (excepting species of Croton belonging to the section
Astraea) these hairs never occur alone, but are always accompanied by other
more complicated trichomes (stellate and tufted hairs); according to Ritters-
hausen both forms of hairs are widely distributed in the Acalypheae, whilst
unicellular trichomes are present in Dalechampia (Tribe Dalechampieae);
finally Herbert also mentions simple unicellular, or rarely uniseriate trichomcs
as occurring in the Hippomaneae. The following special forms of the simple
clothing hairs are noteworthy: (a) unicellular two-armed trichomes, which
have thick or thin walls, and have narrow lumina in the former, and wider
lumina in the latter case; they occur in the genus Argyrothamnia (Acalypheae),
and in the genus Pausandra (Hippomaneae); (b) the trichomes of Sarcoclinium,
which are quite short, and have rather wide lumina and thick walls, and the
trichomes of Coelodiscus, which are likewise very short and have thick walls,
and are mostly bent like a hook at the apex; lastly (c) the branched multi-
cellular trichomes found in Phyllanthus reticulatus, MiilL Arg., and in the genus
M abea (Hippomaneae). The trichomes of Phyllanthus reticulatus are dichoto-
mously branched, the axis and branches consisting of a row of thin-walled cells.
The branched hairs of the species of Mabea are shaped like antlers; the main
axis and its branches are formed by a row of cells~ which have thick walls and
narrow lumina, the terminal cells of the branches being provided with narrow
lateral protrusions, an indication of branches of the second order. The following
 EUPHORBIACEAE 753
types of clothing hairs exhibiting a more complicated structure are found in
this Order: (a) tufted and stellate hairs with transitions to peltate hairs (the
former in the Ricinocarpeae, Phyllantheae, Crotoneae, Acalypheae and Hippo-
maneae, the latter in certain representatives of the Phyl1antheae, 'Crotoneae,
Acalypheae and in the doubtful genu~ Aextoxicon, which is referred to the
Eupho_rbiace,ae by Bentham and Hooker); (b) simple shaggy hairs, i.~. bi- or
multiseriate trichomes (in representatives of the Phyllantheae and Acalypheae) ;
and (c) candelabra-hairs with a multiseriate main axis (in Croton, Fig. 180, K).
We may add that the stellate, tufted and peltate hairs are often seated on
multiseriate stalks oi varying length.
In the following paragraphs the more complicated trichomes will be described
in detail under the individual Tribes. Amongst the RicillOcarpeae, I have observed
tufted hairs in Ricinoca'Ypus, Beyeria and Bertya, where they fill the furrows of the
~olled leaves. ~ ,The ray-cells in this case are eith.er sunk in the epidermis of the
leaf or are seated on a short, multiseriate stalk. Stellate hairs are also mentioned
by systematists as occurring in the genus Beyeriopsis, which belongs to the same
T~~ .
In the Phyllantheae, clothing hairs of complicated structure are not com·
mono According to Rothdauscher, Andraclme aspera, Spreng. possesses uniseriate
trichomes with a short, almost spherical. though apparently not secreto~, terminal
cell, and, accompanying these, similarly constructed shaggy hairs, which are bi-
seriate at the base, but either become uniseriate above, and terminate in a spherical
cell, or fork into two uniseriate branches, each of these having a spherical
terminal cell. Peltate hairs occur in the genus Ht"eronyma; the shield is composed
of a large number (up to 36) of thick-walled ray-cells. which ben4 downwards near
the centre of the shield, and thus form the stalk. the latter being sunk in the epi-
dermis; the peltate hairs of the genus Hymenocardia (which is likewise described
as scaly) are bladder-like and glandular, and will be described in greater detail
amongst the external glands. Stellate or tufted hairs appear to be very rare
among the Phyllanthea~; the only record I am aware of is a statement made by
Miiller Arg. in DC. Prodr. on the rare occurrence of stellate hairs in the genus
Baccaurea.
In the Crotoneae, the third Tribe to be considered, the forms of the trichomes
are very varied. Typical simple hairs are extremely rare in this Tribe, as remarked
above, and where they occur, they mostly appear as rudimentary forms/hf more
complicated trichomes (Fig. 180, G). Frombling, whose description I follow here.
divides the trichomes having a more complicated structure into two groups:
(a) ilairs with a distinctly developed central cell; and (b) hairs without a central
cell, or those in which a central cell. though-<:lriginally present, is not recognizable
in the mature condition. In those hairs, which have a distinctly developed central
cell, the latter is either not specially prominent, or forms a 'central ray' of vari-
able length and structure. The hairs with a central cell belong either' ~ the
peltate or stellate type; amongst the latter Frombling only includes those tri-
chomes in which the rays run in a flat or funnel-shaped plane, and are only connected
with one another basally. Numerous transitions' are to be found between the
stellate and peltate hairs. The stalk of the trichomes in question is short or of some
little length, and consists of several rows of cells; not uncommonly the component
cells are suberized at a certain level throughout the whole thickness of the stalk.
According -to Q_~ :Bacnmann and Radlkofer the occurrence of a small lower scale
is specially characteristic of the peltate hairs (Fig. 180, L) and of the forms of
stellate hairs most nearly related to them; below the actual shield in these cases
one finds a second central layer of smaller cells, which are mostly provided with
large pits. The composite trichomes with no central cell are divided by Frombling
into: (a) tufted hairs, in which the ray-cells are directed upwards, and are usually
sunk in the epidermis, or are attached to a more or less elevated portion of the
latter, i.e. to a kind of pedestal; (b) stalked star-shaped hairs, in which the ray-
cells are borne on a stalk (usually rather long), and extend irregularly in all direc-
tions; finally, (c) candelabra-hairs (Fig. 180, K), consisting of a cylindrical, multi-
seriate main axis, some of the superficial cells of the latter being produced into
rays, which are often arranged in tiers. Between these three forms of trichomes
transitions are found. Their stalks often possess suberized cell-areas like those
SOI.F.RItDEIt 3C
 754 EUPHORBIACEAE
in the stalks of the peltate arid stellate hairs. Epidermal secretory cells are fre·
quently 1 present in the pedestals of the tufted, star-shaped and candelabra-hairs
(Fig. 180, D-F). In many species of the Crotoneae parenchymatous or pros-
enchymatous sclerenchyma-cells are associated with all the forms of trichomes
mentioned above as occurring in this Tribe, with the exception of the candelabra-
hairs; these sclerenchymatous cells belong to the mesophyll, but are so intimately
connected with the hairs that they may be regarded as an integral part of the
latter. In sessile hairs the sclerenchyma-cells immediately adjoin the base of the
hair, whilst in the stalked hairs they traverse the stalk. The precise nature of the
sclerenchyma-cells varies according to the species. In the simplest case they are
slightly sclerosed cells of the palisade-tissue (e.g. in Croton exuberans, Miill. Arg.
(Fig. ISo, G), or C. ch a maedTyfolius , Griseb.). In other cases (e.g. C. cU'neatus,
Klotzsch) they are of the same nature, but are more strongly sclerosed. In C.
antisiphyliticus, Miill. Ar~.• for example, the sclerenchyma-cells penetrate further
into the mesophyll than m the caseS mentioned above; at some points they even
traverse nearly the entire thickness of the spongy tissue, thus coming into contact
with the epidermis on the other side. Not uncommonly the groups of scleren~
chymatous fibres, connected with two trichomes, situated opposite one another
on the upper and lower side of the leaf respectively, have their ends dovetailed,
so that they unite to form a sc1erenchymatous column, traversing the entire thick-
ness of the leaf perpendicularly (Fig. 180, I). It remains to mention the case,
illustrated for example by C. agolnsis, BaHt (Fig. 18o, H), in which the sclerenchy-
matous fibres, connected with the trichome, terminate in the mesophyll like roots.
Regarding the distribution of the sclerenchyma-cells in question, amongst the
Crotoneae it may be stated that they have been observed only in Croton and Julo-
croton; for further details the reader is referred to the foot-note 2 and to From-
bIing's statements. To conclude the subject of the trichomcs of compijcated type
found in the Crotoneae, it remains to mention that they occur in all the genera of
this Tribe except Micrandra. but that peltate hairs have been met with in Croton
and Crotonopsis only, and candelabra~hairs in- Croton only.
In the Acalypheae, according to Rittershausen. tufted, stellate and peltate
hairs are present, as well as shaggy hairs. The tufted hairs are the most widely
distributed (Acalypha. Alchornea, Argyrothamnia, Bernardia, Cephaloctoton, Coe-
lodiscus, Conceveiba, Crozophora, Mallotus and Pera). They generally consist of
a large number of unicellular trichomes, which are sunk side by side in the epider-
mis. In some cases (Crozophora, Mallot'Us) they possess multiseriate stalks of varying

1 CrotDn agrarim, Miill. Arg., C. Cascarilla, Benn., C. celtidifllius, Bail!., C. ~hrysocladus,

Miill. Arg., C. ciliato-glanduloms, Ortega, C. Cortesianus, H. B. K.. C. discolor, Willd., C. jlavens,
MUll. Arg., C. Frionis, Miill. Arg., C.jruticuloSUJ, Mull. Arg., C. g-racilipes, Baill., C. 4umilis, L.,
C. incerlus, Miill. Arg., C. laca'Nlldadus, Mart., C. linearis, Jacq., C. f1ll1rtpliu.r, Mlil1. Arg., C. 'qri-
rani/oHus, Miill. Arg., C. fa'lidus, Miill. Arg., C. panamensis, Miill. Arg., C. pa1aimir, Mull. Arg.,
C. Poh/ianus, Mull. Arg., C. pulegiodtJrus, Raill., C. puncens, Mull. Arg., C. rltamnifolius, Mull.
Arg., C. Sap-aeanus, Miill. Arg., C. semives#tus, MUll. Arg., C. Soliman, Cham. et S~hlei:ht.,
C. stipulaClus, H. B. K., C. subacutus, MUll. Arg., C. sub'lJi//osus, Miill. Arg., C. Urucurana, Bam.,
C. Vaulkieranus, Baill., C. vepretorum, Mull. Arg., C. xa/apensis, H. B. K. All these species have
laticiferous cells also (see p. 746, footnote l).
I In the following enumeration of the species exhibiting a distinct connexion of sclerenchyma
with the trichomes, the words' column,' , root-like' and' palisade-cells,' which are added in brackets,
indicate a columnar connexion of the sclerenchyma of two opposite trichomes, a root-like branching
of the sclerenchyma in the mesophy11, or a palisllde-like differentiation of the. sclerenchyma as the
use may be. It may be added that various morIes of differentiation of the sc1erenchyma often
occur side by side in the same species. The species baving sclerenchyma connected with their
trichomes are as follows: Croton ago€'nsis, Baill. (root-like), C. anlisipnyliticus, Miill. Arg., C.
arg;yranlhemu.r, Michx., C. asftrrimus, Benth. (palisade-cells), C. as/routs, Mull. Arg. (columns),
C. BenlAamianus, Miill. Arg., C. betulinus~ Vahl (root-like), C. BiO!Jergianus, MiiIl.·Arg., C. (aJi-
Ialus, Milli. Arg. (columns), C. caryopnyll'MS, Benth., C. Catinganus, Miill. Arg., C. cautiatus, Miill.
Arg., C. ceanothifolius, Baill., C. c/taet(J(a/yx, Mull. Arg" C. (1UOt'I/o/ius, Miill. Arg., C. (o"'jwessus,
Lam. (columns), C. Corclzoropsis, Baill. (ropt-like), C. cumatus, Klotzscb, C. jloribundus, Spreng.
(columns), C. glandulosus, MUll. Arg. (palisade-cdls), C. gratissimus. Burch. (palisade-cells), C.
lonlinervius, Miill. Arg. (root-like), C. marjlimu.r, Walt., C. miams, Miill. Arg., C. ms"grans,
Casar., C. menanlkogynus. Micbx. (columns), C. NUDan!O, Miill. 'Arg. (palisade-cells), C. Palantl·
stigma, Klotzsch, C. pedicel/atus, H. B. K., C. I~nel/us, Mull. Arg. ; Ju/ocroton argenteus, Fr.
Didrichs.,J./usctscens, Baill.,J. stipularis, MUll. Arg.,J, tn'fjuelet", Miill. Arg. (columns in all fonr
species).
 EUPHORBIACEAE 755
length. In some species of A lchornea the tufted hairs graduate into stellate hairs
by the spreading out of the ray-cells in a plane parallel to the epidermis. In the
stellate hairs of Aleurites the ray-cells have thin walls and wide lumina, and are
concrescent for half their length. Peltate hairs are described by Baillon·as occurring
in Crozophora (on the ovary only), and by O. Bachmann in species of Homonoya
and Pera. The peltate hairs of Homonoya have a four-celled basal plate, and
a shield of narrow thin-walled ray-cells, which are united so as to form a cup; the
peltate hairs of Pera are characterized by the presence of a smaIl lower scale.
Shaggy hairs graduating into spines have been met within Caperonia. The follow-
ing types of hairs require further investigation: the stellate hairy covering found
in the genera Adriana, Epiprinus, Manm"ophyton, Neoboutonia, Symphy/tia and
5 umbavia, and the scales. of Crotonogyne, Leucocroton and Pseudocroton (see 1\1 liller
Arg. in DC. Prodr., and Bentham and Hooker, Gen. plant.).
In the Hippomaneae. as in the Phyllantheae, the trichomes of the type with
which we are dealing are not common. Herbert records stellate hairs with a multi-
seriate stalk and many rays in only one species each of Givotia and Trigonostemot2,
whilst a stellate hairy covering is mentioned by systematists as occurring in the
genera Endospermum. Paracroton and Ricinodendro1l as well. O. Bachmann's
statement that peltate hairs occur in Pausandra is incorrect. The ttichomes
which Herbert met with in Jatropha urens, L .• must be classed as shaggy hairs.
They consist of (a) a multiseriate. columnar pedestal, which becomes slightly nar·
rower upwards, and apparently encloses an intercellular canal, and (b) a rod-shaped
terminal cell with thin walls.-In connexion with the Phyllantheae it may be men-
tioned that the peltate hairs of the anomalous genus Aextoxicum possess a small
lower scale.
II. Ordinary glandular hairs are not very common amongst the Euphor-
biaceae. In the Ricinocarpeae small glandular hairs are found in Bertya
gummitera, l\fiill. Arg. and Beyeria viscosa, Miill. Arg.; they are composed
of a short, unicellular stalk and a spherical unicellular head, and in both cases
give rise to a considerable deposit of secretion on the surface of the leaf; besides
these, club-shaped glands, consisting of a row of cells, also occur in the furrows on
the leaves of Bertya gummifera. Amongst the Phyllantheae, only glandular,
vesicular scales, sunk in small pits in the epidermis, have been recorded (in
Hymenocardia). The shield. is composed of radially arranged cells, and the
secretion collects beneath the cuticle. In the Crotoneae glandular hairs may be
said to be present only in so far as one may thus describe the epidermal secretory
cells projecting like papillae and mentioned above. Amongst the Ac'alypheae
glandular hairs are represented in somewhat greater abundance. External
glands of varying size, and having a uni· or biseriate stalk of varied length,
and a spherical or oval head of few or many cells, have been shown to occur
in species of Acalypha, Cephalocroton, Cnesmone, Ricinella and Tragia. In
Caperonia there are long-stalked glandular shaggy hairs, which are visible
to the naked eye; the ellipsoidal head encloses clustered crystals, and is
sheathed by a secretory epidermis composed of a few elongated prismatic
cells. The genera Coccoceras, Coelodisctls, Macaranga and Mallotus have large
spherical external glands, which are sunk in the surface of the leaf, and have
the structure of the well-known Kamala, derived from Mallotus philippin~nsis,
i.e. they exhibit abundant formation of secretion beneath the cuticle. Amongst
the Hippomaneae, glandular hairs have only been observed in Acidocroton
,adelioides, Griseb. They are small structures sunk in the surface of the leaf,
and consist of a stalk·cell and a head, the latter being divided by vertical wa11s
arranged crosswise; the four cells of the head project slightly as papillae.
Glandular organs of a different type are much commoner among the
Euphorbiaceae than the glandular hairs described above, namely rather .large
external glands (nectaries), which are visible to the naked eye, and have already
occupied the attention of systematists (Baillon, Millier Arg., Bentham and
Hooker I). Firstly they occur at the base of the petiole, where two or more of
1 See also Delpino, ~fem. ACC3d. Bologna, 1888, p. 30 et seq.
3C2
 EUPHORBIACEA£.

them are situated. Not uncommonly, however, they replace the stipLlles, or in
certain species they surmount them, the stipules in this case being laciniate
and filamentous (e.g. in Croton ciliato-glandulosHs~ Ortego or Jatropha multi-
{ida, L.). In other cases they are found at the base of the lamina, sometimes
occurring on a special lobe of the latter. Occasionally they are shifted higher
up on the lamina, two or more of them being pres en t on the lower side of the
leaf; they are rarely met with on the upper side, but frequently occur on
the leaf·teeth. For details of their distribution reference must _be made to
the monograph of Muller Arg.
The anatomical structure of these large glands has hitherto been little investi-
gated; the chief work is that of Frombling on the Crotoneae. This author classifies
the glands as patelliform and morulose. The shape of the patelliform glands
(Croton, Fig. 180, M) recalls that of the perlthecium of the DiscOInycetes; the
glands may be either stalked or sessile. The epidermis of the lower arched portion
has sclerenchymatous walls, and the superficial cells of the'somewhat raiied margin
of the upper disc-shaped portion have a similar structure. The discoid, depressed
surface of the gland is formed by an epidermis~ differentiated jike·a palisade, and
having a subcuticular deposit of secretion; beneath this is a second layer of palisade-
cells with thicker walls, followed by spongy tissue with cells containing clustered
crystals, and surroundiI?-g the termination of a vein. The pat~lliform glands on
the upper side of the leaf of Nle'crandra bra.;teosa, Benth. have an analogous structure.
but do not project as hairs. The glandular margin of the leaf in Croton refractus,
Miill. Arg. (Fig. 180, N) may be regarde4 as a transition to the morulose glands;
in C; f'e/1'actus the margin of the leaf is composed solely of a prominent arched group
of epidermal cells, elongated like a palisade. In other cases, where the curvature
of the surface of the gland is greater, spongy tissue, or even the termination of
a vascular bundle, enters into the compQsition of the glandular body, which thus
becomes club-shaped; in this way we obtain rnorulose glands (Fig. 180, 0), which
are either sessile or possess a multiseriate stalk traversed by a vascular bundle.
The secretion of these glands is often considerable and lasting; in some cases
however the glands are only functional during the early stages of the organs on
which they occur.
Morini examined the nectaries of R£cinus communis, L., Crozophora tinctoria, L.
and Homa/antnus popult/otia. Reinw.; a secretory palisade-like epidermis is found
in these species also.
III. Stinging hairs have been observed in the genera Cnesmone, Lepto-
rhachis and Tragia (Acalypheae}, in Dalechampia and in Jatropha. According to
Rittershausen the stinging hairs in these three genera of the Acalypheae and
in Dalechampia have a very characteristic type of structure, first noticed by
Criiger in Tragia, and subsequently by Kohl and Stahl. In their simplest
form (Fig. lBo, P) they consist of a simple unicellular hair, which frequently
exhibits a ventricose basal enlargement, whilst the upper pointed portion
contains a subulate crystal of oxalate of lime attached to the wall of the hair
by strands of cellulose; the upper end of the crystal is sharply pointed, and
the lower end is somewhat enlarged, while the very base is provided with two
or three small rounded teeth. These simple stinging hairs are accompanied
by others of more complicated structure (Fig. 180, Q) and consisting of two
parts, viz. (a) a fairly long pedestal, composed of five contiguous cells, of which
one lies in the middle, and differs from the four peripheral cells in having thin
walls and possibly containing a secretion; and (b) a terminal cell, having its
basal wall pitted, and its entire length occupied by a crystal showing the same
structure and the same mode of attachment as in the simple stinging hairs.
The stinging hairs of Jatropha urens. Miill. Arg., &c., agree in structure with
the well-known hairs in the nettle. They are unicellular, conical trichomes,
about 4 mm. in length, and are surrounded basally by a multicellular pedestal,
the upper part of which is differentiated like a cupule. At the apex of the
hair there is a head, which is seated obliquely on the end of the trichome;
 EUPHORBIACEAE 757
the head drops off readily owing to the presence of a thin area on the concave
side, and a thickened area on the convex side of the wan just below the head.
As structures allied to the stinging hairs we may now describe the hairs
containing clustered crystals (which were observed by Rittershausen in Pluke-
netia and Fragariopsis) and similar trichomes, which no doubt constitute
a protection against herbivorous animals. The hairs containing clustered
crystals (Fig. ISO, R-S) are epidermal cells, which are differentiated as papillae
or short hairs, and contain a clustered crystal of oxalate of lime; the latter
fills the lumen of the hair, its few (3-6) pointed spines being all directed out-
wards, and enclosed by the extremely thin outer wall in such a way that the
entire structure assumes the appearance of a stellate hair; beneath the cell
containing the clustered crystal there is usually a narrow cell, which penetrates
to a considerable depth in the mesophyll; this cell possibly contains an
irritant substance. Transitions to such hairs, containing clustered crystals,
occur in other Acalypheae. Caperonia and Argyrothamnia possess epidermal
cells containing clustered crystals, and having thin outer walls, which pro-
ject somewhat externally; in Acalypha and Claoxylon epidermal cells, which
project as hemispherical structures, and are filled with a sphaerite of oxalate
of lime, have been observed in the leaf and axis.
3. STRUCTURE OF THE AxIS. The structural features of ·the axis afe not
very uniform, as is to be expected in so large an Order.
The vascular bundles generally have a simple, collateral structure~ The
following genera possess intraxylary phloem: amongst the-Eucrotoneae, the
genera Croton, Crotonopsis, Eremocal'pus and jttloeroton, with the exception
of a few species of Croton, which belong to the section Astraea, and are peculiar
in other respects also (C. comosus, Mull. Arg., C. Klotzschii, Miill. Arg., C. lobatus,
Miill. Arg. and C. Paulinus, Miill. Arg.); amongst the Acalypheae, the genera I
Alchornea~ Conceveiba, Pera and Mallotus integri/olius, l\Hill. Arg. (but not
other species of lllallotus); and amongst the Rippomaneae, the genera Dactylo-
siemon, Mabea, Senefeldera, as well as some of the species'of SebasHan£a 2.
Some of the Euphorbiaceous lianes exhibit normal structure even in thick
stems (species of Bridelia, Croton, Dalechampia, Omphalea, Phyllanthus and
Tragia, according to Criiger and H. Schenck). Anomalous structure is only
known in Dalechampia and Fragariopsis, where it was discovered by H. Schenck.
In both genera (Daleckampia /ici/olia, Lam. in branches r'5 em. thick, Fragari-
apsis montana, Taub. and F. sea n dens, St. Hil.) the anomaly consists in the
appearance of secondary strands of wood and bast in the pericycle, whilst in
Dalechampia there is the additional anomaly of the development of interxylary
phloem. The latter has been demonstrated not only in D. fici/olia, but .also
in D. Leandri, Baill., D. pentaphylla, Lam. and D. triphylla, Lam. It is enclosed
in bands of thin· walled parenchyma, which are concentrica.lly arra.nged, and
is given off by the cambium jnternally. It may be' added that medullary
vascular bnndles have been met with in Ricinus (Sachs, Dutailly).
The following points may be mentioned regarding the structure of the
wood. The medullary rays are in most cases narrow; broader rays are found
in species of Aporosa, Dalechampia and Phylla Il thus. The mode of arrange-
ment of the vessels and the size of their lumina vary (maximum" diameter in
Phyllanthus reticulatfts, Miill. Arg. =·2 mm., in Hemicyclia sepiaria, Wight et
Arn. = -014 mm.). The perforations of the vessels may be either exclusively
Simple, or both simple and scalariform, or scalariform only, in one and the
same species. The bordered pits on the walls of the vessels are· sometimes

1 Pax's statement regarding the occurrence of interna.l soft bast in all tbe species of'AftJ//oluS and
in A/~urit~j' jsincorrect.
, See Herbert, loco cit. p. -49.
 EUPHORBIACEAE
of considerable size (diameter of border = ·004 mm. m Croton pedicellatus,
H. B. K., and '007 mm. in Jatropha divaricata, Sw.). Scalariform bordered
pits are occasionally (Phyllanthus) present on the common wall of two con·
tiguous vessels. In contact with parenchyma of the medullary rays the walls of
the vessels either bear bordered pits only, or bordered accompanied by simple
pits, the latter being sometimes of large siJ;e. The following paragraphs give
a synopsis of the distribution of the perforations, and of the structure of the
veSsel·wall in contact with parenchyma in tbe individual genera.
Perforations: I. Simple perforations only are present in: Caletieae: Calet£a.-
Ricinocarpeae: Beyeria, Ricinocarpus.-AIIlpereae: Amperea.-Phyllaritheae:
A ctephila pro parte, A manoa, A ndrachne, A n#desma pro parte, Breynia, Diseo-
carpus, Hymenocardia pro parte, Lachnostylis, Melanthesopsis, Petalostigma, Phyl-
lanthus, Sauropus pro parte, Savia, Secur,"nega pro parte.-"Crotoneae.-Acalypheae:
A calypha, A driana. A leurites, A rgyrothamnia, B C1'nardia, Caperonia, Cephaloeroton,
Clao~ylon, Cleidion, Coccoceras, Coelodiscus, Conceveiba. Crozophora," Homonoya,
J ohannesia, M acaranga, Pera. Plukenetia, Ricin ella, Ricinus, 5ymphyllia, TYagia,
Trewia.-Hippomaneae: almost all genera, ,except those given in II and III.-
Dalechampieae : Dalechampia.-Euphorbieae : Euphorbia. Pedilanthus.-Gen.
anomal. : Lophopyxis.-II. Scalariform perforations only in: Phyllantheae :
Actephila pro J?arte, Ap()1"osa pro parte, Bmcaurea, Cyclostemon pro parte, Hemi-
cyetia, Putran,iva.-Hippomaneae: Mierodesrnis,; Aextoxieum.-lII. Both scalari-
form and simple perforations have been observed in: P~llantheae: Actephila pro
parte, Antidesma pro patte, Aporosa pro parte, B":schotp,a, Cyclostemon pro parte,
Dl'ypetes, Hieronyma, Hymenocardia pro parte, Riehen"a, Saul'opus prolarte, Savia
pro parte.-Acalypheae: Agrostistachys, Alchornea, Cnesmone, Hevea, eptorhachis,
Mallotus, Mercul'iahs, Pac hystrom a, Sarcoclinium.-Hippomaneae: Adenocline.
Bennetia, HUYQ, Manihot, Ostodes, Paradenocline, Pausandra, Pogonophora.
Pitting of the vessel-wall in contact with parenchyma: I. Bordered pits
only: Caletieae: Caletia.-Ampereae: Amperea.-Phyllantheae: in most. cases,
excepting those genera quoted under II.-Crotoneae (see II).-Acalypheae: Agro-
stistachy s, A rgyrothamnia. Caperonia, Crozophora, M ercuriaiis (according to illtters-
hausen, p. 102). Sarcoclirdum.-Hippomancae: Acidocroton, A etinostemon , Adeno-
pellis, Bennetia. Colliguaya. Excoecan"a, Gelonium, Givotia, Hippomane, Mabea pro
parte. IIJicrodesmis, Pausandra, Sebastiania, Senejeldera, Stillingza, Trigonostemon.-
II. Simple pits often of brge size, sometimes accompanied by bordered pits and by
tr~nsitional forms between both kinds of pits: Ricinocarpeae: Ricinocarpfts.-
Phy llantheae: A n#desmo. A porosa, Baecaurea, Bischoffia, H ieronyma, H ymeno-
ca,.dia, RiehC7ia, 5ecurinega.-Bridelieae: Brt'delia.-Crotoneae: sometimes small
(never large) simple pits.-Acalypheae: AcalYPha, Adr£ana, Alchornea, Aleurites
(according to Rittershausen, p. 46), Bernardia. Cephalocroton, Cia oxylo1J , Cleid~"on,
Cnesmone. Coccoceras, Coelodiscus, Conceveiba, H eVea, H omonoya, J oha'll.f1esia,
Leptorhachis, M acaranga, M altotus, Pachystroma. Pera, Plukenetia. RicineUa,
Ricinus, Symphyltia, Tragia, Trewia.-Hippomaneae: Carumbium, Chaetocarpus,
Cluytia. Codiaeum, Dactylostemon, Erismanthus. Hura, Jatropha, lMabea pro parte,
1\1atlihot, llJ aprounea. Ostodes, Pogonophora.-Dalechampicac: Dalechampia.

The wood·parenchyma is generally scantily developed. It is found in

greater quantity amongst the Phyllantheae in species of Amanoa, Aporosa,
Baccaurea, Cyclostemon, Drypetes, Hemicycl1:a, Lachnostylis, Richeria, Sauropus
and Savia; also in Ridmts (Acalypheae); and in Adenocline, Givot£a, Hip-
pomane, Paradenocline and Pogonophora (Hippomaneae). The wood· prosenchyma
may have either thick or thin walls; in the former case it is sometimes provided
with a gelatinous layer, while in the latter case it is occasionally septate. The
pits of the prosenchyma are generally simple. \Vood-prosenchyma with
bordered pits has only been demonstrated in certain genera of the Tribes
Caletieae, Phyllantheae and Hippomaneae.
Septation of the wood·proscnchyma has been obsen:ed in: Phyllantheae:
Actephi/a, Andrachne, Antidesma, Bisclwtfia; Discocayplts. Hieronyma. Hymeno-
c(lydia, Sallropus.-Bridclieae: Bridel"a.-Acalypheac: Acalypha.
 EUPHORBIACEAE 759
The following genera have wood-prosenchyma with simple pits: Rici.nocarpeae:
Bey,ria, Ricinocarpu,s (very small bordered pits as well).-Phyllantheae: Actephila,
A manOQ, A ndrachne pro parte. A ntidesma, Baccaurea, Bischoffia, Breynia. Disco-
Garpus, Drypetes, Hemicyclia, Hymenocardia, Lachnostylis, Melanthesopsis. Petalo-
stigma, Phyllanthus, Richeria, Sauropus, Savt'a, Securinega.-Bridelieae: Bl'idelia.-
Crotoneae.-Acalypheae: Ac-alypha, Adriana, Ag1ostistachys, Alchoynea, Aleurites.
Argyrothamnia, Berna,dia, Caperonia, Cephalocl'oton, Claoxy(on, Coccoceras, Coelodis-
GUS, Crozophora, Hevea, Homonoya, Johannesia, Leptorhachis, Macaran~, Mallotus,
Pachyst,oma, Per~a. Plukenetia, Ricine/la, Ricinus, Sarcoclinium, Symphyllia, Tl'agia.
-Hippomaneae: Acidocroton, Actinostemon, Carumbium, Chaetocarpus pro parte,
Erismanthus, Excoecaria, Givotia, Hippomane, Hura, Mabea, Paradenocline, Pogono-
phora, Sebastiania, Seneleldera, Trigonostemon.-Dalechampieae: Dalechampia.-
Euphorbieae: Euphorbia, Pedilanthus.-Gen. anomal.: Lophopyxis.
Wood-prosenchyma with bordered pits is present in the following genera:
Caletieae: Caletia.-Phyllantheae: Hieronyma, most species of Andrachne,
Aporosa sphaet'ocarpa~ Miill. Arg.-Hippomaneae: Adenocline~ Adenopelt'is~ Ben-
netia~ Chaetocarpus pro parte, Codiaeum, Colliguaya, Cluytia (indistinct), Dactylo-
siemon, Gelonium, Jatrot?ha. Manihot (narrow borders). A1aprounea, Microdesmis,
Ostodes, Pausandra, St'tllingia.-Acalypheae: Conceveiba (according to Ritters-
hausen, p. 67).
The following genera have wood-prosenchyma with both simple and bordered
pits: Ampereae: Amperea.-Phyllantheae: CyclostemDn.-Acalypheae: Cleidion,
Cnesmo-ne. MeTCU'riaiis, Tt'cwia.
Those structural features in the cortex which are cornman to the Order
were mentioned in the general diagnosis. The development of cork takes place
in the outermost cell-layer of the primary cortex in a large nwnber of genera
belonging to the Tribes Ricinocarpeae, Phyllantheae, Bridelieae, Crotoneae,
Acalypheae. Hippomaneae and Dalechampieae (see the special papers). In
Actephila the cork develops in the epidermis; while in Baccaurea, Andrachne
Roemeriana, Miill. Arg. (but not the other species of Andrachne) and Amperea
sparlioides, Brongn. the cork is produced at a considerable depth in the priinary
cortex. The varied structure of the cork-cells can only be referred to; the
cells may be flat or provided with wide lumina, they- may have thick or thin
walls, and may be sclerosed on one side or on all sides. The outer portion.of
the primary cortex is frequently collenchymatous, and often contains stohe.
cells (e.g. in special abundance in Aporosa and Henticyclia); according to Pax
the stone-cells in Hyaenancht and Richeria are distinguished by being strongly
elongated in the vertical direction. Other points requiring special mention
are: (a) the occurrence of an apparently phellodennal ring of stone·cells in
!ohannesia; (b) the presence of a ring of stone-cells in the innermost part of
the primary cortex and in contact with the primary hard bast in Pogonophora ;
and (c) the peculiar cortical collenchyma found in species of Pera having
its small intercellular spaces filled with a peculiar, doubly _ refractive
substance (Rittershausen). In most cases the pericyde is formed by isolated
groups of bast~fibrest which in young branches frequently unite to form a ring,
or by a continuous and composite sclerenchymatous ring. In .H'lera there
is a broad parenchymatous pericycle with abundant tannin, situated on the
inner side of the" sclerenchyma-ring. The bast-fibres of the pericycle are in
many cases distinguished by an exceptionally well-marked stratification of
their walls, which are either white or yellow. In some cases (in Amperea
sparlioides, "Brongn. and probably also in a number of the herbaceous Euphor·
biaceae) the sclerenchymatous pericycle is entirely wanting.
A composite and completely or nearly completely continuous ring of scleren-
cbyma is stated to occur in: Phyllantheae: Amanoa. Apoyosa pro parte, Cyclo-
Siemon, Dyypetes, Hemicyclia, Hieyonyma pro parte.-Acalypheae: Alchofnea pro
parte. Cleidion, Conceveiba. Hevea, Homonoya. Macaranga pro parte, Pera, Sym~
p/~)·lIia.-Hippomaneae: AC#'1ostemon pro parte, Chaetocaypus pro parte, Cluyti'a
 EUPHORBIACEAE
(slightly interrupted), Dactylostemon, Gelonium, Maprounea, Microdesmis, Sebas~
tiania pro parte and Seneletdera (in Gelonium and Sene/eldefa the ring of scleren~
chyma is composed of stone-cells, and at its outer margin the groups of primary
bast-fibres are situated).-An interrupted and composite ring of sclerenchyma occurs
in: Phyllantheae: Actephila, Aporosa pro parte, D,'scocarpus, Hymenocard£a,
Hieronyma pro parte, Lachnostylis.
It has already been pointed out on p. 739 that typical stratification of
the bast in,.,to hard and soft bast appears to be of rare occurrence (B,idelia
stipularis, Bl.). In many members of the Order, belonging to the various
Tribes one finds small groups of bast-fibres Of isolated bast-fibres, the latter
sometimes exhibiting a reticulate arrangement in transverse section. The
bast-fibres> found in the secondary bast resemble those of the pericycle in fre-
quently possessing well~marked stratification of their walls. Two specially
noteworthy features may be mentioned: the bast in Securinega Acidothflmnus,
Miill. Arg. shows sclerosis of the parenchyma lying between the groups of
secondary bast.fibres, so that a composite and continuous sclerenchymatous
ring is formed; and the bast in the species of Ricinella (according to Radlkofer's
verbal statements) is distinguished by the fact that the innermost, annular
portion has a characteristic. reddish-brown or brown colour owing to the pre-
sence of a secretion. The crystals found in the bast '-(mostly solitary or clus-
tered crystals) are contained in chambered fibres.
The cells of the pith mayor may not be lignified. Stone-cells are not
uncommonly present in the pith. In Cluytia alaterno£des, Miill. Arg., the cells
of the pith are specially characterized by the curious fact that their transverse
walls are provided with large, circular or ellj.ptical perforations at the points
of junction with the vertical walls. :
Regarding the features presented by the crystals and secretory organs in
the axjs, see above, pp. 744 and 746 et seq.

DAPHNIPHYLLACEAE.
At this point we may briefly_ deal with the anatomical features of the genus
Daphniphyllum, which was raised to the rank of a separate Order by Muller Arg.
on account of its small embryo, but is included amon.sst the Phyllantheae by Ben-
tham and Hooker. I have examined the leaf and axlS of D. laurinum, Baill., and
the leaf of D. glaucescens, BI. There are no special anatomical characters separating
the genus Daphniphyllum from the Euphorbiaceae. The characteristic features
in the structure of the axis, viz. the scalariform perforations of the vessels, and the
bordered pitting of the wood-prosenchyma. are also found in the Euphorbiaceae;
the same applies to the special structural features of the leaf, viz. the presence of
subsidiary cells placed parallel to the pore of the guard-cells, and the occurrence
of small cells, containing clustered crystals, in both the upper and lower epidermis.
Trichomes and secretory elements are not present. The septation of the pith (with-
out sclerosis) in the two speCies mentioned above is specially noteworthy.
The follOwing statements may be added regarding the structure of the axis
an~ leaf. The wood possesses narrow, uni- to triseriate medullary rays, the cells of
whlch are somewhat. elongated in the vertical direction* The vessels of the wood
have small lumina (diameter reaching '036 mm.), perforations with very numer-
ous bars, and bordered pitting (especially scalariforrn bordered pits) in contact wifu
parenchYIlla of the medullary rays. The wood-parenchyma is only scantily de-
veloped. Isolated groups of sclerenchymatous fibres are present at the outer limit
of the bast. Clustered crystals occur in the bast and in the primary cortex. The
leaves of the two species mentioned above have bifacial structure. The stomata
are found only on the lower side of the leaf. The vascular bundles of the veins are
accompanied by sclerenchyma. In D. glaucescens the epiaermis on the lower side
of the leaf shows papillose differentiation.
 BUXACEAE.
The following statements are based on my own examination of the axis and
leaf in Simmondsia calijornica, Nutt. {described as Brocchia glauca in my (Holz-
struktur '), Sarcococca prunitofmis, Lindl., Buxus sempervirens, L. and Pachysandra
p,-otumbensf Michx.
As characters common to the whole group, we may point out that the wood-
prosenchyma invariab1y beaIS bordered pits, the vessels always have bordered pits
on the walls in contact with parenchyma, the stomata are never provided with
parallel subsidiary cells, and external glands are absent. The perforations of the
vessels are usually scalariform only, but are simple in Simmondsia. The place of
origin of the cork varies; in Buxus it is subepidermal, whilst in S'I:mmoncist:a the
cork arises in the parenchymatous pericycle, immediately internal to the pericyclic
groups of sClerenchymatous fibres. Oxalate of lime is excreted in the form of
clustered crystals, ordinary or styloid-like solitary crystals, or as crystal-sand.
Internal secretory organs are only present in the form of secretory cells (Pachy-
sandra, Simmondsi a), and never show typical differentiation. The following tea tures
require special mention: the anomalous structure of the axis in the monotypic genus
Simmondsia (appearance of successive rings of growth)1 and the cortical vascular
bundles in Buxus sempcrviycns.
Regarding the structure of the leafin the species mentioned above, the following
facts may be added. The leaf is bifacial in Buxus, Pachysandra and Sarcococca,
whilst in St·mmondsia the entire rnesophyll consists of palisade-tissue. The species
exhibiting bifacial structure have stomata only on the lower side of the leaf; in
Simmondsia, on the other hand, they are equally distributed on both surfaces. The
stomata of Simmondsia are somewhat depressed, and have no subsidiary cells.
In the remaining genera the guard-cells are surrounded by a rosette of more or less
distinct subsidiary cells. Buxus sempervirens is distinguished by the specially
strong development of crests on the guard-celis. The epidermis in the leaf of
Simmondsia shows some noteworthy' characters: its cells have a small polygonal
outline in surface-view, whilst in a transverse section of the leaf they have the
shape of palisade-cells, but with a conical tapering of the cell·lumen towards the
upper side, and a thick outer wall. Sc1erenchyma may (Buxus. Simmondsia) or may
not (Pachysandra) accompany the vascular bundles of the veins. In the leaves of
~uxus. Pachysandra and Sarcococca I found no oxalate of lime, while in Simmondsia
I observed that it occurred abundantly in the peripheral portions of the leaf in the
form of clustered crystals, and in the middle region in the form of solitary crystals.
some of which were differentiated like styloids. In the dried leaf of S£mmondsia the
palisade-parenchyma contains relatively large cells. filled with a yellowish secretion,
which at once acquires a brown colour with Eau de Javelle, owing to the presence
of tannin. Trichomes have been observed on the, leaves of Pachysandra and 5im-
mond~·a. The hairs of Pachysandra are simple, have pointed ends. and consist
of from two to three cells; those of Simmondsia are likewise uniseriate, but are
composed of a larger number of cells, which have thick walls, and mostly exhibit
a certain amount of articulation, whilst the last one or two cells are relatively short
and have thin walls (glandular ?).
In the structure of the axis the anomaly presented by Simmondsia califoynica
first requires a somewhat detailed description. Even branches from herbarium
material show successive rings of growth; these are arranged concentrically, and
are separated from one another by narrow rings of lignified parenchyma (con-
junctive tissue).' 'The· soft "bast in the-individual vascular rings does not form a
continuous annular "zone, but consists of isolated groups arranged in concentric
series, with parenchymatous conjunctive tissue penetrating between the grou~.
The secondary rings of bundles arise in the parenchymatous portion of the pen-
cycle, internal to the pericyclic groups of sclerenchymatous fibres.
A transverse section of the wood in the Buxaceae shows narrow medullary
rays, one Or two, rarely three cells in breadth. The vessels are generally isolated,
and their diameter is not great ('015-'°3 mm.). The bordered pits on the vessels
are small in Buxus and S~·mmondsia, but relatively large in Pachysandra and
Sarcococca. Bordered pits are also found on those walls of the vessels which
are in contact with tissue of the medullary rays. The scalariform perforations
have rather numerous bars (3() or more) in Pachysandl'a, Sal'cococca, and in Buxus
subco/umHaris, Mull. Arg., which belongs to the section T"icet'a; the bars are
 BUXACEAE
less numerous (not exceeding IS) in the investigated species of Buxus belonging
to the section Eubuxus (B. balealica, Willd., B. iaponica, Miill. Arg., B. sempct'-
virens, L. and B. Watlichiana, Baill.); S~·mmondsia has simple perforations only.
The size of the bordered ~its on the wood-prosenchyma agrees with tha.t of the pits
on the vessels. Spiral thickening is found on the walls of the vessels in Simmondsia
and on the walls of the wood-prosenchyma in Pachysandra. The wood.paren-
chymais in most cases scantily c1eveloped, being somewhat more abundant in Buxus
only.-Turning now to the structure of the cortex, we must first refer to the cor-
tical vascular bundles of Buxus sempeyvirens, which were mentioned above. Whether
they are present in other species of this genus remains to be determined. They are
found singly in the four corners of the branches, and according to J. E. Weiss each
bundle is a leaf-trace, which ends blindly in the primary cortex. In Pachysand~a
pf'ocumbens the primary cortex includes groups of sclerenchymatous cells, which
are either sclerosed on all sides or on one side only; the s.clerenchyma-cel1s of the
latter type enclose solitary crystals of oxalate of lime. In Simmondsia. Sal'COcocca
and Pachysandra the bast is limited towards the primary cortex by groups of
sc1erenchymatous fibres, but these are not present in Buxus sempervirens. I have
also observed isolated sclerenchymatous fibres in the secondary bast in Sa1'cococca.
The mode of development of the cork has already been mentioned above. Secretory
elements have been met with in the primary cortex bf Pachysandra pl'ocumbens
(herbarium~materia1), where they occur in the form of secretory cells arranged in
longitudinal rows. Oxalate of lime is found in the axis of S~·mmQndsia in the form of
clustered crystals; in Pachysand'fa the pith and primary cortex contain crystal-sand,
and the primary cortex has SOlitary crystals as well; in Buxus sempervirens solitary
crystals and small crystalline granules are found in the bast, occurring side by side
in the same cells; in Sarcococca prunijorm£s oxalate of lime is not present.

Literature: Cruger, in Bot. Zeit. 1855, p. 618 et seq. and'Tab. viiL-Bailloo, ft. gen. da groupe
des Eupb., Paris, 1858, pp. ~30-40.-Hanstein, MiIchsaftger. etc.;Berlin. 1864, pp. :H. 76 elc. and
Tab. x.-Dippel, Milchsaftgef., Rotterdam, 1865.-Trecul, Lacticiferes etc. dans les Euph. etc••
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lld. ill, (88~, p. ~So.-Schunerus) Milchs. von Euphorbia Latnyris, Abh. bot. Ver. Provo Branden-
burg, Bd. xxvi, 1882, p. 27.-Haberlandt. Phys. Anat. d. Milchr•• Sitz.-Ber. 'Viener Akad., Bd.
lxxxvii, Abt. I, 1883. pp. 51-69 and ~ Tab.-J. E. Weiss, Markst. Gefassbiindelsyst., Bot. Centralbl.
J883, iii, p.4-II and Tab. i.-Scott, Laticif. tissue of Maniltot GlazitflJii, LaUcif. tissue of Ht'ZIta
Spruceana, The quarterly Joum. of mictosc. sc., vol. xxiv, 1884, pp. 19J-~o6.-Pax. Anat. d. Eaph.
etc., Engler, Bot. Jahrb., Bd. v, 1884. pp. 384-4:1J ruld Tab. vi-vii.-Dingler, FlAchsprosse, i.
Plty/lan/hur, Miinchen, J88S, p. ~9 et seq.-Mentovich, Mark, Klausenburg, 1885, Hungarian; abstr.
in Just 1885.1, p. ,S,.-Scott, Articul. laticif. vessels in Hcvea, Journ. Linn. Soc., vo1. xxi, 1885, pp.
566-S73.-So1ereder, Holzstr., 1885. pp. 236-.f1.-0. Bachmann, Schildh., Flora 1886. sep. copy,
pp. 5-13 and Tab. vii.-Haberlandt, Brennhaare. Sitz.-Ber. ,"Viener Akad., Ed. xciii, Abt. I, 1886,
p. n3 et seq. and Tab.-Morini, Nett. estranuz., Mem. Accad. Bologna 1886, pp. 343,346 and 347.
tav. i-ii.-Pirotta e Marcatili. Vasi laticiferi ed it sisto assimil., Annuario R. 1st. bot. di Roma 1886,
n. 2; abstr. in Just 1886, i, p. 922 and Bot. Centralbl. 1886, ii, pp. :n~-13.-Radlkofer, Drnchs.
Pnnkte auf Blattem. Sitz.-Ber. Munch. Akad. 1886, pp. 330-8.-Volkens, Aegypt.-arab. WUste, t887,
p. 1 43.-Calvert and Boodle, Laticif. tissue in the pith of Man;"'" etc,. Ann. of bot., vol. i, 1887, pp.
55-62 and pI. v.-Stahl, PII. u. Schnecken, Jena, 1880, p. 7o.-Hansen, Spbaerokr., Arb. Wiirzburger
rnst .• Bd. iii, 1888, p. 93 et seq.-Groom, Laticif. tubes, Ann. of bot., vol. iii, 1889, pp. 161-3 and
pI. xi.-Kohl, Kalks. u. Kiesels., 1889, p. 164; here cited Johow, in Sitz.-Ber. neiderrhein. Gesellsch.
f. Natur- u. Heilkunde, 4. Joni I888.-Leitgeb, Spharite, Mitt. bot. lnst. Gnu, Heft 2, 188S.-Ross,
Tess. ass, e periderma, Nuov. Giorn. bot., vol. ui, 1889, pp. 236-7.--Scott, Laticif. ti:.sne in the
leaf, Ann. of bot., vol. iii, 1889, pp. 44S--6.-Lalanne, }<'euilles persist., Act. Soc. Linn. Bordeaux,
ser. S. t. iv, 1890. p. 57 and pI. iii.-Pax, in Natiirl. Pfl:mzenfam., iii. Teil, Abt. 5, 1890. PP' 3-5
t"t seq.-Rodier. Spherocristaux, Mem. Soc. se. phys. et.nat. de Bordeanx I890.-Chau\,eaud, App.
 BUXACEAE

lact. des Euph. etc., Ann. sc. nat., ser. 7, t. xiv, 1891, p. 25 and 138 et seq. and pI. 1-6.-Fry,
Aggreg. of proteid in the cells of Eupk(J1"oia splendem, Ann. of bot., vol. v, 1891, pp. ",15-18 and
pl. xxiv.-Benecke, Nebenz. d. Spaltoff"n .• Bot. Zeit. 189:l, pp. 589-91 and Tab. viii.-Rittershansen,
Anat.·syst. Unters. von BI. 11. Axe d. Acalypheen, Diss., Erlangeu, 1893, 123 pp. and I Tab.-
Belzung, Spherocristaux des Euph. cactif., Joum. de bot. t. vii, 1893. p. III and 261 et seq.-
H. Schenck, Anat. d. Lianen, 1893, pp. I",:l-7 and Tab. viii.-Engler, Icacineae. Sitz.-Ber. Berliner
Akad. i893, pp. 265-6 and Tab. (Luphopyxis).-Herbst, Markstr., Bot. Centralbl. 1894. i, pp. 328-9.
-Frombling, Anat.-syst. UnteJ'S. von 1:11. u. Axe d. Crotoneen u. Euphyllantheen, Diss., Munchen,
1896, 76 pp. and :l Tab.; sep. copy from Bot. CenlTalbl. 1896.-Rothdauscher, Anat.-syst. Unters.
von BI. u. Axe der Phyllantheen etc., Diss., Munchen, 1896, 89 pp.; sep. copy from Bot. Centralbl.
1 896.-Wittlin, Kalkoxalattascb., Bot. Centralbl. 1896, iii, p. 68.-Warming, Feuilles de l'Euphgrbia
/Juxifolia, Danske Vid. Selsk. Oversigt 1896; abstr. in Bot. Centralbl. 1897, ii, p. :.J79.-Herbert,
Anat. Unte:s- v. Bl. u. Axe d. Hippomaneen, Diss., Munchen. 1897,62 pp.-Reiche, Ch~~7Il. Holzpfl.,
Jahrb. f. Wl5S. Bot., Ed. xu, 1897, p. 9o.-Kuhla, Phelloderm; Bot. Centralbl. 1897, lll, p. 196.-
Moller, Lignum Aloes, Pharm. Post 1897.-Warming, Halofyt Stud., K. Danske Vid. Selsk. Skr.
1897, pp. 190 , 2H and 2:u.-[Chauveaud, Caract. anat. des EupMrbia Pep/us etc., Joum. de bot.
1897, p. 534.-Poulsen, IDe. cit.]

BALANOPSEAE.
There are scarcely any statements I in the literature regarding the anatomy
of this small Order, which consists of the single genus Balanops, and is regarded
as a Tribe of the Cupuliferae by Baillon. The following remarks are based on
my own examination of fragments of certain of Balansa's type·specimens
kindly sent to me by Prof. Urban (axis and leaf of Balanops Balansae, BailI.
and B. microstachya, Baill., and leaf of B. Pan cheri, BailI. and B. Vieillardi,
Baill.). The following features are characteristic of the genus: vessels with
extremely small lumina, and scalariform perforations having numerous bars;
a composite and continuous ring of sclerencnyma at the outer margin o£ the
bast; superficial development of cork; excretion of oxalate of lime in the
form of clustered and solitary crystals; and the absence of trichomes.
The smooth leathery leaves have bifacial structure. The epidermis of the
leaf is distinguished by having a considerably thickened outer wall and by
~he polygonal outline of its cells in surface-view. In two of the investigated
species, viz. B. Balansae and B. micro sta chya. a single layer of hypoderm is
present beneath the upper epidermis. The stomata occur only on the lower
side of the leaf; their guard. cells are provided with strongly developed cuti~
cular ridges, and are surrounded by s~veral epidermal cells. The vascular
bundles of the veins are embedded, and. are surrounded by a ring of scleren·
chyma. Enlarged terminal tracheides are sometimes found at the ends of
the vascular bundles. In B. Pancheri and B. Vieillardi clustered crystals are
rather abundant in the mesophyll.
According to Petit, the b~sal portion of the petiole in B. V ieillardi contains
three arc~shaped vascular bundles; these become closed higher up to form
three rings, but finally open out again.
The wood in the genus Balanops consists of: (a) vessels which have very
small, rounded-lumina (only attaining a diameter of '021 mm.), and are mostly
isolated; (b) wood.prosenchyma with thick walls and narrow lumina; and
(c) very narrow medullary rays. The vessels-, as stated above. have scalariform
perforations with numerous bars in all cases. In contact with parenchyma
of the medullary rays, the walls of the vessels bear rounded or slit-shaped
pits, which may be described a.s unbordered. In B. microstachya the wood·
prosenchyma has simple pits, but in B. Balansae the pits may be recognized
as distinctly bordered when seen in section. The cells of the medullary rays
are often considerably elongated in the vertical direction; they are sometimes
low, but never show considerable radial elongation.
I Petit, Petiole. Actes Soc. Linn. de Bordenllx, 1. xliii, 1889, ". 18 and pl. i.
 BALANOPSEAE

Regarding the structure of the cortex, the following statements may be

made. The cork arises sub epiderm ally (B. microstachya), and consists of
narrow, tabular cells with rather thick waIls. In B. microstachya a ring of
stone-cells is found adjoining the cork-cambium; this ring is perhaps of the
nature of a phelloderm. In B. Balansae numerous slightly sclerosed cells
occur in the primary cortical parenchyma. The outer limit of the bast is
formed by a composite and continuous sclerenchymatous ring, which also
includes crystal-cells (with solitary crystals). In the bast we occasionally find
solitary crystals, or structures resembling clustered crystals. True chambered
fibres containing solitary or clustered crystals are not present in the bast;
secondary bast-fibres are also absent, at least" in such material of the branch
as has been investigated.

URTICACEAE.
I

The most satisfactory method of dealing with the anatomical features of

this Order will be to describe them under different groups, viz. : 'I. Ulmaceae J

(to include the Tribes Ulmeae and Celtideae); • 2. Cannabineae J (the Tribe
Cannabineae); • 3. Moraceae' (the Tribes Moreae, Arctocarpeae and Cono-
cephaleae); and' 4. Urticeae' (the Tribe Urticeae). The Tribe Thelygoneae
wi.ll be considered in an appendix to these groups, since.it differs anatOIpically
from the rest of the Urticaceae in possessing bundles of 'rap hides.
At this point we may give a brief review of the most important anatomical
features. The occurrence of silicified or cal<;ified cell~walls, and of similarly
constituted cystoliths or cystolith-like structures is very general. In some
cases the cystoliths occur independently of the trichomes, this type being found
chiefly in epidermal cells (independent cystoliths); in other cases they art~
met with in the hairs (hair-cystoliths). Independent cystoliths have in the
first place been found in certain Ulmaceae and Moraceae, but attain their
widest distribution in the Urticeae. A second anatomical feature, viz .. the
occurrence of laticiferous cells, is confined to Cannabis and representatives
of·the Moraceae. The following important anatomi~al characters af(~ common
to all Urticaceae: the superficial development of the cork in most cases;
the considerable length of the bast-fibres 1; the simple perforations of the yessels;
the usually simple pits on the wood~prosenchyma; the lack of a uniform type
of stoma; the excretion of oxalate of lime in the form of ordinary solitary
and clustered crystals (except in Thelygonum, see above). The hairy covering
consists of simple, mostly unicellular clothing hairs, and glandular hairs of
varied structure. Special forms of hairs are: the bracket-hairs, occurring in
some genera; the two-armed climbing hairs of Humulus; the stinging hairs
of certain Urticeae; and the pearl-glands. In addition to laticiferous tubes
and external glands, the secretory system consists of resin-cells.and mucilage~
receptacles (mucilaginous cells and mucilage-canals).

I. ULMACEAE.

The Ulmaceae agree with the rest of the Urticaceae in certain features pre-
sented by the structure of the axis (viz. the superficial development of cork,
the almost exclusively simple perforations in the vessels, and the nearly universal
occurrence of simple pits on the wood-prosenchyma); also in the frequent occur~
rence of calcification and silicification, and more especially of cystolith-like

1 These long bast.fibres are multiDl1cleate In l/rlica dioi,Q, and Cannabis saliva (Treub and
Kallen).
 ULMACEAE
structures. The latter are either independent cystoliths, or hair-cystoliths, or
cystotyles. As a point of distinction from the Cannabineae and Moraceae,
the absence of laticiferous cells in the Ulmaceae deserves notice. The only
internal secretory elements which have been observed are ~nucilage-cells (in
the bast of species of Ulmus, and in the mesophyll of species of Celtis, Giron-
niera and PhyllostylQIL), if we exclude the gelatinization of the epidermis of
the leaf found in many species. The type of stoma is not uniform; similarly
the pericycle shows varied differentiation. Oxalate of lime occurs in the form
of clustered and solitary crystals, the latter sometimes resembling styloids in
shape as in the neighbourhood of the primary hard bast of Celtis. The hairy
covering consists of clothing and glandular hairs, both of which show little
diversity of form, the clothing hairs being mostly unicellular. Other special
features in the structure of the leaf are: papillose differentiation of the lower
epidermis (species of Celtis); and development of hypoderm on the upper side
of the leaf (species of Aphananthe, Celtis, Gironniera, Holoptelea, Parasponia.
Trema).
The leaf·structure of the Ulmaceae has been investigated especially by
Priemer 1. The following statements may be cited regarding the integu4
mentary tissue of the leaf. The lateral margins of the epidermal cells are either
straight or undulated. Certain species of Ulmus and Celtis are distinguished
by the development of strong cuticular ridges, especially on the lower side of
the leaf. Papillae are found on the lower side of the leaf in certain species
of Celtis 2. All the investigated species of Ulmus, Phyllostylon, Planera,
Z elkova incl. H emiptelea, A mpelocera, Celtis, Parasponia, Gironn'iera, Chaetacme,
and some of the species of Trema exhibit gelatinization of the epidermis of
the leaf. Another feature is the' two-layered epidermis' found in species of
Aphananthe, Celtis, Gironniera, Holoptelea, Trema and Parasponia s, the lower
layer of cells usually differing from the upper. The stomata are generally
present only on the lower _side of the leaf, 'but in Trema aspera, Bl. and some
species of Celtis they also occur on the upper side. The guard-cells usually
lie on a level with the epidermis, but l in the species with a papillose epidermis,
and in H emiptelea Davidii, PI. they are depressed, while in Trema amboinensis, /
Bl. and other species they are elevated. With regard to the subsidiary celis,
Priemer makes the following rather vague statement: 'The stomata are
invariably enclosed by accessory subsidiary cells' _; in Ulmus campestris, L.
I found the stomata surrounded irregularly by a number of ordinary epidermal
cells, while in Celtis tetrandra, Roxb. one of the neighbouring cells on either
side of the pore is placed parallel to the latter. The leaf· structure is mostly
bifacial, but is subcentric in some species of Celtis. The palisade· tissue consists
of one or more layers, the number being sometimes constant for certain species,
or even genera. In some species of Celtis the lateral walls of the palisade· cells
are finely undulated. The spongy tissue either contains large lacunae (Planera,
Ulmus, Zelko va) or is dense. In certain species of Celtis, Gironniera and
Phyllostylon 4, mucilag~.:cells~-occur in the mesophyll~. The vascular bundles

1 Priemer's work deals with the following genera: (I) Ulmeae: Ulmus, Holople/to, Pkyllosty/o1Z.
Plantrai (2) Celtideae: Ztlk()va incl. Hem£pldea, Ampelocera. Cellis, Trema, Parasponia, Apha-
nantne, Gironniera. Chaelacme and eel/ida/sir (Alerlensia pubescens, Ktb., incorrectly na.med
Merlensia citn"foHa or Celtidopsis cilrifolia by Priemer). Priemer's nomenclature of the species is
on the whole retained here. although in his work the $ame species sometimes figures under several
synonyms j I bave only corrected the numerous misprints.
2 See Priemer, lac. cit.• p. 462.
:t. See Priemer. loco cit., p. 424, foot·note I.
t Celtis lUuleata, Sw., C. dicholoma. Rniz, C. j'amaicemis, Pl., C. lall/olia, Pl., C. mauri/iana,
Pl., C. panicu/a/a, Pl., C. ri"gescem, Pl.; Cironniero ce/tidijo/ia, Gaud.; Pltyllostylon ''''ariHens,\
Capan., P. rhamnoidu. Taub.
~ 1 do not know what Priemer means by the I (ree sclerenchymatous elements' of some species
 ULII/ACEAE

of the veins are either accompanied by sc1erenchyma only (Ulmus, Planera,

Zelkova, Celtis pro parte, Gironniera, A phananthe pro parte) or by a sheath of
large parenchymatous cells devoid of chlorophyll (Hemiptelea, Phyllostylon,
Celtis pro parte, Trema, Parasponia, Aphananthe pro parte); in other cases
neither sclerenchyma nor a parenchymatous sheath is present (Celtis cinna·
momea, Lindl., Ampelocera, Apltananthe philippinensis, Pl.). Vertically trans-
current veins are found in the genera Gironniera, Hemiptelea, Parasponia, Planera,
Trema, Ulm-tIS, Zelkova, Celtis pro parte, and Phyllostylon rhamnoides, Taub.
Oxalate of lime is present in the form of ordinary solitary and clus'tered
crystals. Both forms of crystals are found in the leaf in the genera A pha-
nanthe, Celtis, Gironniera, H oloptetea 1, Phyllostylon and Ulmu'i, the clustered
crystals occurring in the mesophyll, the solitary crystals accompanying the
vascular bundles of the veins. Solitary crystals alone have been observed in
the leaf in Ampelocera (only in the petiole) and Zelko va, clustered crystal~
alone in Hemiptelea, Parasponia, Planera and Trema, while in Ampelocera
neither clustered nor solitary crystals are present.
The hairy covering consists of clothing and glandular hairs. The former
have pointed apices, and are mostly unicellular, or as an exception (C. Krauss-
iana, Bernh.) bicellular. The walls of the hairs are mostly thick, and are
frequently calcified or silicified; the outer surface is sometimes (species of
Aphananthe, Celtis, Girom~iera, Trema) furnished with calcified warts, and not
uncommonly a crystolith-like body springs from the lateral wall. Special forms of
trichomes are: the long and narrow clothing hairs of Celtis boliviensis, PI.,
C. Tala, Gill. and Zelkova cretica, Spach; the triChomes of Ampelocera Ruizii,
Klotzsch, which have thin walls, and are broadest at the middle, not at the
base; and the hairs of certain species of Trema,i (such as T. amboinensis, Bl.),
which likewise have thin walls, sometimes contain cystoliths, and form a felt·
like covering on the lower side of the leaf. Glandular hairs are present in all,
the genera examined by Priemer. They generally have a unicellular or uni-
seriate stalk of variable length, and a head composed of one or several cells.
The shape of the glandular hairs varies even within the limits of a genus. In
the simplest case (species of Celtis and Trema) the glandular hairs consist of a row
of cells with a more or less distinct demarcation of the terminal cell. In other
species of Trema the hairs are composed of a double row of cells. Ulmus, Planera,
Phyllostylon, Aphananthe and Hemiptelea have club-shaped glandular hairs con-
sisting of about three cells lying in a row, the terminal cell being occasionally
somewhat swollen. In A mpelocet'a there are glandular hairs with a unicellular stalk.
and a club-shaped head divided by horizontal and vertical walls; in Gironniera
there are hammer-shaped external glands with a unicellular stalk1 and a head
divided by vertical walls only, whilst in certain species of Celtis the glandular hairs
are derived from uniseriate hairs by enlargement of the distal cells which further
undergo vertical divisions.
In some cases the marginal teeth of the young leaves also produce a secretion,
e.g. in Ulmus campestris. according to Reinke. These teeth include the termination
of a vein; mucilage is found in the epidermis and the subjacent tissue, and a few
stomata OCcur above the end of the vein
Silicification and calcification are common phenomena, as stated above.

of Celtis, described on p. 438 ; possibly the enlarged terminal tracheides are referred to. In the hvo
species which I had at my disposal for checking this observation (c. paniculata, 1>1. a.nd C. telra,tdra,
Roxb.) I fonnd nothing to which such a term would be applicable. -
1 In the general part of Priemer's work PhyllQstylon is not included amongst the genera enume-
rated as having both solitary and clustered crystals in the leaf; in place of this genus, however,
Hemilteie4 is quoted, involving a contradiction of the statements in the special portion of the same
treatise. In the general part H%plelta is given amongst the genera-which have clustered crystals
only, while, according to the special part, solitary crystals are present in the petiole of Holopttlea.
Such contradictions are unfortunately not uncommon in Priemer's work.
 ULMACEAE

In some cases the silicification, besides affecting the walls of the hairs, frequently
extends to the outer walls of the epidermal cells and the adjacent lateral walls.
In other cases 'the silicified parts are protuberances of the cell-wall, the shape
of which may graduate into that of cystoliths. These protuberances occur
not·only in the integumental tissue~ but also in the mesophyll. The silicifica-
tion is specially strong in Celtis, Chaetacme, Hemiptelea, Holoptelea and
Trema. In many cases calcification accompanies the silicification. The
calcification of the walls of the hairs has already been mentioned above. We
may follow Priemer in dividing the calcified cystolith-like structures into
cystotyles and cystoliths. According to Priemer the cystotyles comprise
all encrusted . . structures which have no stalk and no stratification, while
the cystoliths are those which are provided with a stalk, or are stratified,
or exhibit both these features. The cystolith-like structures are almost always
confined to the integumentary tissue; they are restricted to the mesophyll in
Celtidopsis only. Cystoliths have been observed in the genera A mpelocera,
Celtis, Chaetae me, Hemiptelea, Holoptelea, Parasponia, Phyllostylon'and Trema,
and cystotyles in: all the investigated genera. The cystoliths are either inde-
pendent, or occur in the hairs. The independen t cystoliths are found in epidermal
cells, which are enlarged towards the mesophyll, only a small portion of the
wall of the cell in most cases reaching the surface of the leaf; it is to this por-
tion of the wall that the stalk of the cystolith is attached. Cystoliths of the
second type, viz. hair-cystoliths, occur in the trichomes. The lithocysts con-
taining the independent cystoliths usually lie with their outer wall on a level
with the epidermis~ and rarely (Celtis brasiliensis, Pl.) occur in a superficial depres-
sion. The independent cystoliths are as a rule botryoidal, cylindrical, or
spherical in shape, all these different forms being found in the genus Celtis.
Among rarer forms are reniform or arc-shap_ed cystoliths (Celtis pubescens,
H.B.K., &c.), or mushroom-like cystoliths (Chaetacme aristata, Pl.). The
cystoJiths in the hairs are pear-shaped and are seated on the lateral wall,
y

near the base of the hair. The occurrence of cystoliths in mucilaginous epi-
dermal cells is specially noteworthy (species of Chaetacme and Trema),
In dealing with the systematic value of these structures Priemer distinguishes
the following types of cystoliths, employing the terms introduced by Mez in his
stUdy of the cystoliths of the Cordiaceae: I. Independent cystoliths, i.e. such as
OCCur independently of the hairs; at the most the outer wall of the lithocyst ter-
minates in a small point. A mpelocera (containirtg very little lime and only occurring
in the upper epidermis of the leaf); Celtis (except C. Kraussiana, Bernh.; some
of the species also have types II and III); Chae/acme (mushroom-shaped, mostly
in mucilaginous epidermal cells. and only on the upper side); Holoptelea (only in
the lower epidermis); Phylloslylon (accompanying type II); Trema (in mucilaginous
epidermal cells in a few of the species; accompanied by type 11, and in some of
the species by III also); Celtidopsis (only in the mesophyll). II. Hair-cystoliths:
Celtis pro parte (accompanied by type I); Phyllostylon (accompanied by I); T,ema
(accompanied by I, and in some of the species also by III). III. Spherical cysto-
liths, i.e. hair~cystolith~,_ surrounded by epidermal cells containing cystotyles, the
whole group forming small false scales: Celtis Kraussiana, Bernh. and other species
of Celtis (the latter also-having types I and II); Hemiptelea; Parasponia (accom-
panied by II); Tt'ema (accompanied by I and II). IV. Smaller cystoliths, oiten
poor in carbonate of lime and rich in silica, and seated in groups of 2-5 in the corners
of adjacent epidermal cells: in all the genera and of no special systematic value.
According to Priemer the base of the petiole contains either an arc-shaped
vascular bundle (Celtis, Trema, Zelkova, Holoptelea, Aphananthe), or a ring of
wood and bast (Ulmus, Planera, Phyllostylon, Hem£ptelea, ChaeJacme), or three
vascular bundles (Gironniera). According to Petit, three vascular bundles
enter the petiole in species of Celtis, Ulmus and Planera.
The structure of the axis has been specially examined by me with reference
 768 ULMACEAE

to the nature 91 the wood in Ulmus campestris, L., Holoptelea integrifolia, Pl.,
Planera aquatic4, Gmel., Celtis occidentalis, L. and Aphananthe aspera, Pl., and
by Moller with referenGe to the cortex in species of Ulmus and Celtis. In Ulmus
the medullary rays of the wood reach a breadth of seven cells even in material
from branches, the cells being much elongated radially, and exclusively in that
direction; in the remaining genera the medullary rays are narrower, reaching
a breadth of three cells, and some of the cells are elongated in the vertical,
S01ll£ in the radial direction. The vessels vary as to the size of their lumina
and the mode of their arrangement. In Holoptelea, the vessels, which are mostly
isolated, more rarely arranged in radial series of 2-3, attain a diameter of
'09 mm.; in Planera the majority of the vessels form radial rows, and their
diameter only reaches '04 mm.; in Ulmus the diameter of the vessels
in the spring.wood reaches 'IS mm., and they are isolated, while the vessels
of the autumn-wood have smaller lumina and form tangential groups. On
the common walls between contiguous vessels rather large bordered pits occur
(diameter of border in Ulmus as much as '009 mm., in other cases '004 mm.).
In contact with parenchyma of the medullary rays, the walls of the vessels are
also mostly provided with distinct bordered pits. Spiral thickening of the
vessel-wall has been observed in Ulmus campestris (according to Hesselbarth
in U. pedunc~data also), and in Aphananthe aspera. The perforations of the
vessels are simple, and round or elliptical. Scalariform perforations with 1-6
bars have only been met with in Planera aquatica, where they occur in the
neighbourhood of the primary wood, elongated elliptical perforations being
also present. The wood·parenchyma is sometimes (Celtis, Hotoptelea) developed
in considerable quantity. In Planera the wood·prosenchyma is provided with
_pits having a small border {similar to th~se of Betula). but in other cases it bears
simple pits. Thewood·fibres frequently have relatively wide lumina and rather
thick walls. In some cases (species of Celtis. Holoptelea and Ulmus) the size
of the lumen is diminished by the presence of a gelatinous layer. It remains
to mention that in the heart·wood and pith of certain species of Ulmus and
Celtis it has been demonstrated that carbonate of lime forms a component part
of the cell·contents (Molisch and Priemer).
The outermost cell-layer of the cortex is sometimes (CelUs, Ulmus) silici-
fied. According to Moller, the development of cork takes place subepidermally.
The cork consists of cells with wide lumina, and in Celtis contains stone-cells.
As is well known, Ulmus suberosa, Mch. is characterized by a massive develop-
ment of the superficial periderm. The pericycle contains isolated bundles of
bast·fibres in Ulmus, and a composite and continuous ring of sclerenchyma in
Celtis. In both genera the secondary bast includes groups of long, smooth
bast· fibres, which occasionally give rise to a stratification of the bast. In the
older cortex of Celtis the bast·parenchYma becomes sclerosed. The sieve·tubes
have simple sieve·plates with coarse meshes. The medullary rays of the bast
contain no crystals, but solitary and clustered crystals are fairly common in
other parts of the cortex; in Ulmus the solitary crystals in the neighbourhood
of the primary bast are rod-shaped (resembling styloids). Of secretory ele-
ments, mucilage-cells are found in the bast of Ulmus; they have large rounded
, lumina, and are somewhat elongated in the vertical direction: according to
Moller they vary in abundance, and are quite absent in some specimens.
According to Gris, the pith in Planera is entirely composed of active cells
with thick walls, whilst in Ulmus and Celtis the central portion consists of
rather large, empty cells with thin walls~ and the peripheral. portion is formed
oy relatively smaU active cells with thick walls '.

I It is not easy to understand how Gris comes to the conclusion, from these' data, that Cdtis
~ho\\ld be separated Crom Ullllf(s and P/a1tera as an independent group.
 2. CANNABINEAE.

Cannabis and Humulus, like the other Urticaceae, exhibit the following
features: simple perforations in the vessels; wood·prosenchyma with simple
pits; secondary hard bast; long bast·fibres (as much as 22 mm. in length in
Cannabis); cystolith·like structures; unicellular clothing hairs often ~on·
taining cystoliths; and glandular hairs, the heads of which are mostly divided
by vertical walls only. Laticiferous cells have only been observed in Humulus.
The stomata have no special neighbouring cells. Oxalate of lime takes the
form of clustered crystals.
To this brief diagnosis we may add some information regarding the lati·
ciferous tubes, the hairy covering, the cystoliths,' &c., and the structure of the
petiole. The laticiferous tubes of Humulus are found in the parenchymatous
pericycle, on the inner side of the primary groups of hard bast. According to
Holzner the laticiferous tubes do not appear to be present in the embryo.
Further investigation is required to determine whether these elements occur

A~~Q D 1. D 2.
A283

FIG. 18.. A-F, Humulus Lutultu, L.: A-D, Glando.lar hairs; E, Lupulin.
~land~ ; F, Climbint hairs. G, Transverse section throu~h the leaf of Ca1tnabis
sali1ltJ, L.-A-D and F after Holzner, E after Kny, G Origmal.

in Cannabt's, or are really absent, as is apparently the case I. The clothing

hairs are mostly unicellular, more rarely (Humulus) uniseriate; they vary in
length. As in other Orders possessing cystoIiths, we find an antagonistic
relation between the development of hairs on the one hand, and of cystoliths
on the .<?_ther. (Fig. r8I, G). The large cystoliths on the upper side of the leaf
of Cannabis sativa; L. are contained in trichomes, the basal portion of which
is strongly swollen, and penetrates into the mesophyll, while the actual body
of the hair is often considerably reduced; on the other hand the longer hairs
on the lower side of the leaf only include small cystoliths, and in these the
stalk of the cystolith may be said to be formed by the body of the hair, which
has become solid owing to calcification and silicification; in other cases these
Chauveaud's statement (loc. cit., p. 119) that laticiferous tubes occnr' in CannalJis is based on
the fact that Engler ascribes them quite generally to the members of his Order Moraceae (including
the Cannabineae). According to Chauveaud laticiferous tubes are m,t present in the embryo of
Cnnnabb.
SOLEREDER 3 D
770 CANNABINEAE

long hairs only possess a calcified apex. In the cells adjoining the large hair-
cystoliths, cystotyles sometimes occur. The climbing-hairs found in the hop
(Fig. r8I, F) are a special form of clothing hair. They are unicellular mal-
pighian t~irs with equal or unequal arms, and a strongly silicified wall, and
are either directly inserted in the epidermis or seated at the apex of a multi-
cellular pedestal. These hairs are connected with the ordinary clothing hairs
by transitional forms. The glandular hairs of Humulus exhibit a series of
fonns, which have been described in detail by Holzner (Fig. r8r, A-E) ;, the
following types are found: (a) stalked capitate external glands, with a uni-
cellular, Of uniseriate and then locally (i.e. in the upper part) biseriate stalk, and
a spherical head, which is composed of four or more cells separated by vertical
walls, or is divided into a still larger number of cells by both hor~ontal and
vertical walls; (b) discoid glands with a unicellular stalk and a disc-shaped
head consisting of a single layer of cells; (c) finally glands with a crateriform,
head, composed of a single layer of cells, and exhibiting an abundant accumula-
tion of secretion beneath the cuticle. Cannabis has discoid glands similar
to those of Humulus, and also small external glands with a unicellular stalk,
and a head which is either unicellular or bicellular, owing to the presence of
a vertical wall. The teeth on the young leaf of Humulus are also glandular,
being provided v-ith a secretory palisade-like epidermis~ The characteristic
region of the petiole contains an arc-shaped fibrovascular system in Cannabis
and Humulus; in the latter genus the seven component vascular bundles
which enter the petiole, can be distinctly recognized in the arc of vascular tissue.

3. MORACEAE.
The Moraceae are spedally distinguished by the presence of laticiferous
cells. Certain other features they have in common with the rest of the Urti-
caceae; these are: the simple perforations in the vessels; the simple pitting of
the wood-prosenchyma; the superficial development of the cork; the presence
of cystolith-like structures j and the frequent occurrence of very long bast-
fibres. The stomata usually have no special subsidiary cells. The hairy
covering consists of: (a) simple unicellular hairs, which are sometimes (A"to-
carpus, D01stenia, Broussonetia, Cecropia) bent like a hook at the apex; (b)
uniseriate hairs (Ficus pro parte, Cecropia); and (c) glandular hairs of varied
structure (in some cases pearl-glands). Oxalate of lime is excreted in the form
of solitary and clustered crystals. The following internal secretory organs
are present: resin-cells (Artocarpus), tanniniferous idioblasts (Ficus), and
mucilage-canals (Cecropia, Conocephalus). Special features in the leaf are the
occurrence of hypoderm (species of F'icus and Conocephalus), and of clustered
crystals in the epidermis (species of Antiaris, Artocarpus, Ficus, Maclura).
A detailed investigation of the structure of the leaf has not yet been made.
In the representatives of the three Tribes examined by me (Morus alba, L.,
Artocarpus incisa, L. f. and Conocephalus suaveolens, Bl.) the stomata are
surrounded in the first two species 1 by a number of epidermal cells irre-
gularlyarranged, in Conocephalus suaveolens I by three subsidiary cells. Gela-
tinization of the epidermis of the leaf has been observed in M orus alba, and
by Richter in Antiaris toxiclJria, Lesch. and species of Cecropia. Hypoderm
has been found in Conocephalus suaveolens, species of Cecropia and numerous
species of Ficus (see De Bary and Mobius); it occurs either on both sides or
1 The same is the case in AlIlian"s. according to A. Richter, and in Fuus, according to Stras-
burger. In Ficus the cells immediately surrounding the pairs of guard-cells are cut off by secondArY
divisions from the cells adjoining the mother-cell of the guard-cells '(Strasburger, in Pringsheim's
Jahrb., Bd. v, 1866-7, p. 332 and Tab. xli).
2 A Iso in /)orstenia, according to Benecke.
 /I-/ORACEAE 77 1
only on the upper side of the leaf. In 'Brosimopsis lactescens~ Sp.-Moore and
Ficus Carica the upper epidermis consists locally of two layers. The genus
Cecropia has vertically transcurrent veins (Richter). A noteworthy feature
is the occurrence of clustered crystals in the integumentary tissue. These
crystals occupy small epidermal cells in A ntiaTis sacciodora, Dalz. and A rio ..
carpus communis, Forst., according to A. Richter, and in Madura tricuspidata
and Ficus indica,' -according to Mobius; in the first three !pecies these ('ells
I

form small groups. In Ficus elastica and F. macrophyUa clustered crystals

are found in the hypoderm (Mobius). It remains to mention the peculiar
small pits, warts or spots (forming hydathodes), which are found in many
species of Ficus, in Conocephalus ovatus and CecroPia Schiedeana; they are
sItuated on the upper side of the leaf, and generally occur above the points
of junction of the veins, or more rarely are restricted to the margin of the
leaf. They are due to the presence of a group of cells resembling an epi-
thema and containing the termination of a group of tracheae belonging to the
vascular system; above the epithema there is a group of water·pores (De
Bary,. Haberlandt, Mobius).
The distribution of the laticiferous tubes has hitherto been insufficiently
investigated from an anatbmical point of view. By no means all the Moraceae
are stated by systematists to have latex, but that does not exclude the possi.
bilityof laticiferous tubes occurring throughout the Tribe, though in some cases
with contents which are not milky. In Bentham and Hooker's Gen. Plant.,
and in DC. Prodr. the following genera are described as lactescent: amongst the
Moreae-Bleekrodia, Taxotrophis, M aillardia, Broussonetia, Allaeanthus, Bagassa,
Chlorophora, Maclura, Pachytrophe (succo opalino), Paratrophis, Pseudomorus,
Morus, Ampalis (succo opalino), Trophis, Sloetia and Dorstenia; amongst the
Artocarpeae--Ficus, Brosimum, Lanessania, Antiaris, Olmedia, CastiUoa,
Perebea, N oyera, A ,tocarpus, Sahagunia, Clarisia, Balanostrebus and SOTocea ;
and amongst the ConocephaIeae-Cecropia and Coussapoa. According to Poppig.
Trymatococcus (Morea) has-no latex; the genera Pseudolmedia, Helicostylis and
Batocarpus (Artocarpeae) are described as (?) lactescent. According to Treeul,
6

laticiferous tubes are not present in Conocephalus suaveolens, but they have /
been shown to occur in the recently established Artocarpeous genus B,osim·
opsis, Sp.-Moore. According to Schmalhausen and Chauveau~ the initials
of the laticiferous system in the Moraceae are found in the nodal plane of the
embryo; they form two groups, each composed of 4~5 cells, which are situated
peripherally in the central cylinder at points corresponding to the depressions
between the two cotyledons. In the mature plant they are present both
in the axis and leaf; those in the axis occur in the pith, primary cortex, peri·
cycle and bast. The medullary laticiferous tubes are, often connected with
those in the cortex through the medullary rays. Those occurring in the secon·
dary bast are not of secondary origin, but are merely branches of the cortical
laticiferous system. The course of the laticiferous tubes in the leaf has only
been examined in--detail in ~Ficus; we may follow Pirotta and Marcatili, as
well as Groom and Scott~ in distinguishing two cases. In some of the species
of Ficus (e.g. F. laurifolia) the laticiferous tubes are only found accompanying
the vascular bundles of the veins; in other species (e.g. F. elastica, F. religiosa,
&c.) the main branches of the laticiferous tubes do not leave the vascular
bundles of the veins, but they send out branches into the mesopbyll, and these
sometimes traverse the hypodermal aqueous tissue as far as the epidermis, or
even (F. retusa) reach the cuticle. In Ficus Carica, F. elastica, Broussonetia
papyri/era, Maclura aurantiaca and MOTus nigra the contents of the latici·
ferous tubes include large grains, the nature of which has not been determined;
these grains frequently show stratification, which was first observed by CarueJ.
The laticiferous tubes also contain nuclei, according to Treub.
~n2
 77 2 MORACEAE
The spherical secretory cells with resinous contents mentioned above
have been observed by A. Richter in the spongy tissue of the leaf in Artocaypw.
communis, Forst. and A. echinata, Roxb. (but not in A. integri/oiia, L.). The
mucilage~canals are probably of lysigenous origin, and are found in the pith in
species of Cecropia and in Conocephalus suaveolens (see also Treeul), The tanni.
niferous idioblast~ which were described by Mobius as mucilage-cells, have
contents which are strongly refractive in the living, and ,reddish-brown in the
dry plant; these elements are present in the palisade-tissue (as enla.rged cells
of this tissue), and in the neighbourhood of the vascular system of the veins
in Ficus austraUs (=F. rubiginosa).
Calcification and silicification of the cell-wall are two common phenomena
in the Moraceae, and often accompany One another. The silicification affects
cells of the different tissues, the hairs and cystoliths. The following cases of
silici1ication may be mentioned as examples: the walls of the epidermai cells in
species of Ficus and M orus are silicified; so also are the walls of the outermost
cell-layer of the primary cortex in species of Artocarpus, Brosimum, Coussapoa,
Ficus, M OTUS, Pourouma, Sorocea, Trophis and Trymatococcus, and the walls
of the hairs in Ficus; according to Kohl, the walls of the papillose epidermal
cells and of the trichomes in Dorstenia nervosa, Desv. are silicified to such an
extent that the lumina of the cells almost entirely disappear. In the case of
the cystoliths, silicification sometimes (Fictls Sycomorus, a<:cording to Miliarikis)
affects the lithocysts, and very frequently the stalk and the adjacent portion
of the nucleus of the cystolith. In F. Sycomorus moreover, according to
Miliarikis, a siliceous shell may be formed round tbe head of the cystolit~
thus enclosing it, or the entire cystolith may be impregnated with silica.
The cystoliths, which have been chiefly examined in the leaf, are inde-
pendent in some cases, whiJst in others they are hair-cystoliths. Their exact
distribution has not yet been determined. In Kohl's treatise, in which the state-
ments of earlier observers are combined with his own observations, cystoliths
are described as occurring in F':cus, it{orus, Broussonetia and Chloropkora
(Kohl's Maclura tinctoria=Chlorophora tinctoria, Gaud., non fvlaclura, NutL);
to these I am able to add Conocephalus. According to Payen, cystoliths are
absent in Dorstenia. The cystoliths found in the leaf, when not present in
the hairs, are generally 1 situated in epidermal cells, which often penetrate
deeply into the mesophyl1, while the portion of their wall reaching the
surface of the leaf varies in area. In most caSes the cystoliths occur singly;
more rarely several of them are found in the same cell; within the limi ts of
the same genus (Ficus) they may predominate sometimes on the upper, some-
times on the lower side of the leaf. Only the independent cystoliths show
typical differentiation; they are provided with a distinct stalk attached to
the outer wall, exhibit distinct stratification, and sometimes also show radial
strands, whilst in the hair-cystoliths these structural features are more or less
reduced.
The cystoliths have been most thoroughly investigated in the genus Fict.!-s by
KoW and Mobius, and their results, which are based on the examination of a rela-
tively small number of species, are sufficient to show what valuable data will be
obtamed for specific diagnosis when further investigations are carried out. The
cystoliths of F£cus elastica (Fig. J 82, A -B) are those best known; they are ellipsoidal
in shape, with a verrucose surface, and occupy large idioblasts, adapted to the
sha pe of the cystoliths and occurring in the integumentary tissue of the leaf, e~pecially
on the upper side. They are strongly impregnated with carbonate of lime, and
to some extent (chiefly in the stalk) with silica as well; in section (as may be
specially Clearly seen after decalcification) they e.xhibit very fine stratification

1 In no case do they occur in the cells of the assimilatory tissne, but they are s.ometimes present
in the parenchyma of the veins of the leaf.
 MORACEAE 773
about the end of the stalk as a centre, and an additional system 01 delicate, frequently
branched strands, radiating from the same point and terminating in the warts.

FIG. 182. A-B, Cystolith. of Fieu$ elastica, Rm:b.

in 'the hypodermal ti8SUe on the upper side of the
leaf: B, after decalcification with hfdrochlorjc acid.
~J Cystoliths and cystolith·hairs on the lower side of
Ule leaf of Ficus Ca".ica, L. D-E, Broussontl,a
Ialyriftr(J1 Vent.: D, Transve:rse section of the leaf
with cysto ith-hairs, bracket-hairs and glandular
hairs i E, Cystolith. hairs. on the: upper side: of the
younr leaf.-Original.

In F. Cooperi, especially io the lower

c,1?idermis, there are spherical cysto.
liths in which the stalk is short and
thick, and the surface bears broad flat
warts. The cystoliths 00 the upper
side of the leaf in F. 1'eligiosa are
characterized by the fact that their
lithocysts do not differ in size from
the remaining epidermal cells, and are
arranged in groups. In F. Carica the
cells containing the cystoliths show a
tendency to develop'" into· trichomes
(F)g. 182. C).- In the leaf of this
species we find the following structures:
(a) large, distinctly stalked cystoliths C
occurring on the lower side of the leaf
in enlarged papillose epidermal cells
provided with a small solid hair-like
tip; (b) hair-cystoliths; and (c) hairs,
of which the upper portion is filled with
calcified caps of cellulose. In a species
which Kohl describes as F. ceyasijeya.
he found (besides other types) large cystoliths of an irregular tuberous shape,
having neither warts nor strands. For further examples see Kohl and Mobius_
774 MORACEAE
According to Bokorny, the cystoliths of F. cordata, Thunb., which give rise to
transparent dots in the leaf, are not calcified, but appear to be suberized, to judge
by the reaction they give with iodine and sulphuric acid.-The cystoliths found in
the species of Morus are hemispherical or spherical, and are provided with radiating
fibres. Their lithocysts are situated on the upper side of the leaf, and are of large
size.-In Broussonetia .papyrifera, Vent. (Fig. r82, D-E) the large lithocysts on the
upper side of the leaf show papillose differentiation: on the lower side hair-cysto-
liths are found. The large cystoliths on the upper side are sometimes produced
by the fusion of two or three cystolith-rudiments.

The most essential points regarding the hairy covering have already been
mentioned above. Unicellular bracket-hairs are present in Dorstenia, according
to Kohl; in Artocarpus and Cecropia, according to A. Richter; and in Brousso-
netia, according to my own observation. The glandular hairs found in Antiaris~
B10ussonetia and Morus have a unicellular stalk and a more or less distinctly
delimited ellipsoidal head, which is multicellular with transverse and longitu-
dinal division~wal1s. Similar glandular hairs appear to occur also in Cecropia
(Richter) and in some of the species of Ficus, \ at least on the young leaves
(Meyer and Mobius); in F. rubig~'nosa glandular hairs have been found, con~
sisting of a row of three cells, of which the terminal cell is long and tubular;
, F. indica' has glandular hairs with a bicellular stalk and an elongated head,
divided into two cells by a vertical wall (Mobius). The external glands of
Arlocarpus have a unicellular stalk and a disc-shaped head, divided· by vertical
walls; according to Haberiandt. these glands are hydathodes. The large
pearl-glands of Cecropia and Pouro,uma. which were known to Meyen,' remain
to be described; it has been shown that in Cecropia they have a biological
function 'as food~bodies. Several square centimetres on the lower surface of the
base of the petiole'in Cecropia and Pourouma are coated by a velvety brown
covering of mniseriate hairs, amongst which the pearl·glands are situated;
these glands look like insects' eggs; they fall off readily, but are continually
r~~aced by others. They are ovoid or pear~shaped~ and consist of numerous
c , which in Cecropia are rich in proteid· matter and fatty oil, according to
Schimper; in this genus a stoma is situated at the apex of the gland. Glandular
leaf-teeth having a similar structure to those of Ulmus (see above) are found
for instance in Morus (Reinke).
According to Petit, the differentiation of the fibrovascular system ~n the
petiole is not of a uniform type. In ]}[orus alba the main system forms an arc~
in species of Ficus and in Castilloa elastica a slightly intenupted ring, in Brous ..
sonetia papyri/era, Maclura a~rantiaca and Artocarpus integrifolia a ring ~f
5-7 isolated vascular bundles, while in Cecropia peltata the petiole contains
a still. larger number of vascular bundles (about 30). For details see Petit,.
loco cit.
The following statements may be made regarding the structure of the
wood; they are based on my own examination of species of the genera Sloetia,
Morus, Conocephalus, Artocarpus and Ficus, and also on the statements of
MOller, Hesselbarth, Houlbert and A. Richter. The medullary rays vary in-
. breadth; in Conocephalus suaveolens, Bl. and Fictts indica, L., they attain
a breadth of seven cells even in herbarium-material. The size of the lumina
of the vessels varies greatly; the perforations are simple. In contact with
parenchyma the walls of the vessels as a rule bear bordered pits with tran~itioJ1S.
to large simple pits, but in Slodia St'deroxylon, Teysm. et Binn. there are
bordered pits only. Spiral thickening of the vessel-wall has been met with
in species of Morus (chiefly in the vessels with narrow lumina), in Broussonetia
papyri/era, Vent., and Maclura aurantiaca, Nutt. The wood·prosenchym~
always bears simple pits and may have either wide or narrow lumina. Accord·
ing to De Bary. a gelatinous thickening-layer is p-resent in the wood·fibres in.
 MORACEAE 775
species of Mol'us, Broussonetia and Ficus; septation of the wood-fibres is found
in species of Ficus. In many species of Ficus, in Cecropia obtusa, Tree. and
in species of Artocarpus, Brosimum and Streblus the wood-parenchyma forms
tangential bands; this is also the case, though less distinct, in Antial'is and
Bagassa.
The pith is homogeneous in Ficus (Mentovich).
The structure of the cortex has been specially examined by Moller in
species of M orus, M aclura, Ficus, Broussonetia, Artocarpus and Cecropia, and
by A. Richter in Antiaris. In these genera the cork arises in the outermost
celI..layer of the cortex. The collenchyma of the primary cortex is developed
in various ways. Artocarpus and Cecropia are ,characterized by extensive
sclerosis of the primary corte~ accompanied by a certain amount of thicken~
ing. A ring of stone~cells, occurring in the primary cortex, is distinctive of
M OTUS alba, L., according to Moller, and Chlorophora tinctoria, Gaud., Cudrania
iavanensis, Tree. and Streblus asper, Lour., according to Engler. The pericycle
includes isolated groups of bast-fibres, the indiyidual fibres being often broad
and flattened like a strap. The secondary bast contains bast~fibres, which
vary in abundance and arrangement j in some cases (Morus alba, Streblus
asper, according to Engler) the secondary bast·fibres, which are always
distinguished by their considerable length, give rise to stratification of the
bast. Some of the medullary rays of the bast are broad; the sieve~tubes
have simple sieve-plates with coarse -meshes.

4. URTICEAE.

The Urticeae are characterized by the absence of laticiferous tubes and

the frequent occurrence of independent cystoliths. Only one genus (Neraudia)
is stated by Gaudichaud to exude a milky juice, but this genus has not yet
been investigated anatomically I. The Urticeae have the following features
in common with the rest of the Urticaceae: the lack of a uniform type of stoma;
the usually superficial development of the cork; the simple perforations of
the vessels; the simple pitting of the wood-prosenchyma; the long bast·fibres ;
and the excretion of oxalate of lime in the form of clustered and solitary crys~
tals S, The hairy covering consists of -simple unicellular clothing hairs, and
small glandular hairs. As special forms~of the clothing hairs we may mention
the bracket~hairs (Boehmeria, according to Demeter, Forskohlea, according to
Weddell), and the stinging hairs occurring in some genera, whilst the pearl·
glands found in certain species of Urtica constitute a special form of glandular
hair. The internal secretory elements 3 known to occur are lysigenous mucilage~
canals in the pith and cortex of Boehmeria platyphylla, Don et Ham. and
Pipturus argenteus, Hort. (Moller, Engler).
The. leaf. structure has hitherto been little investigated. The stomata,
which frequently occur on both sides of the leaf, have no special subsidiary
cells in Urtica-and "Some-species of Pilea, while in other species of Pilea, and
in the species of Boehme-ria, Elatostema and Pellionia examined by Benecke
there are three subsidiary cells. The thick fleshy leaves of Pilea serpyllifolia,

1 Chauveaud's statement (loc. cit., p. 119) regarding the occurrence of laticiferous tnbes in the
ma.ture plant in Urtica dioua is incorrect, and is probably based on a remark made by Treub (Arch.
need., 1879), who mentions 'laticiferes' in Urlica. According to Gravis (loc. cit., p. 19) the
elements referred to are merely bast· fibres having rather wide lumina in which contents are present.
t Schaarschmidt describes the occurrence of sphaerocrystals of a substanCe allied to besperidin
in alcohol material of U,.tita majDY; accOIding to Kallen nuclear crystalloids occur in the bristle-
hairs of Urlica dioica.
3 The tannin-cells described by Zopf (Bibl. bot., Heft :1,1886, p. 23 and Tab. iii) in the stem of
Pa,.ielaria diffuSlI Are not distinct idiobillosts.
 URTICEAE
Wedd. have stomata only on the upper side, where they occur in large numbers;
on the lower side the margin of the leaf bears hydathodes consisting of a group
of water-pores with a subjacent epithema provided with the termination of
a vein. Hydathodes having the same type of structure are also found scattered
on the lower side of the leaf in Pilea elegans, and are similarly distributed on
the upper side of the leaf in Urtica urens and U. macrophytla, ThWlb.; on older
leaves of the last species the hydathodes are covered by small iridescent scales,
composed of an organic substance, silica, and carbonate of lime. The lower
epidermis of the leaf in Pilea serpyllifolia consists of water-storing cells of
great height and with wide lumina. Hypoderm is stated by Pfitzer to occur
in 'Urlica crassifolia.'
The cystoliths of the Urticeae have been examined by WeddelL They are
independent, Le. they do not occur in connexion with the trichomes. As a'rule
they belong to the integum~ntary tissue" but they also occur in the pith, bast
and primary cortex of the branch. They are usually spherical, ellipsoidal or
fusiform (Fig. 183, A); other forms of rare occurrence are stellately branched
(Pika stelluligera. \Vedd.). In those species which have fusiform cystoliths,
others of irregular form are sometimes present in smaller numbers (e.g. bent

c D

FIG. 183, A, Upper epidermis of the leaf of Pika .zerpyllf/()/i,z, Wedd, with a cxstoJith. B-D, Stinging bair of
Url;ca diuica, L,: D, ll~r p4>rtion of the same; the 'haded part of the wall is clilciued, the un shaded part sili·
cified.i.Cz Tip of the iitinging hair before, and D after the head has been broken Off.-A Original, B after Kohl, C-D
after l1aoerlandt.

like a horseshoe, &c.). The stalk of the cystolith is only slightly developed,
or appears to be absent; in many cases it probably becomes resorbed. In
the fusiform cystoliths, which lie parallel to the surfaE:e of the organ, the
stalk is attached to the middle of the cystolith. The latter sometimes shows
concentric stratification and striation, and generally has a siliceous skeleton.
In certain species of Pilea the body of the cystolith has been shown to be
enveloped in a siliceous shell (Miliarikis), but this feature is not of systematic
value.
In the dried leaf the cystoliths are generally visible, according to their
different shapes, either as dots (cystolithi punctiformes) or as elongated,
tubular or linear markings (cystolithi oblongi, fusiformes, !ineares), or rarely
(viz. in Pilea stelluligera mentioned above) as star-shaped bodies (cystolithi
stellati). These and other features, which are likewise visible with a lens
(e.g. the occurrence of the cystoliths on both sides, or only on one side of the
leaf, the position of the cystoliths with reference to the veins, &c.), were employed
for systematic purposes by Weddell in his monograph of the Urticeae; in the
first place he showed that independent cystoliths occur in almost all the genera
included in his enumeration in De Candolle's Prodromus (excepting only Laurea,
Ville'brunea, Debregeas";a, Roussel£a, Forskohlea and Dt'dymodoxa; of these
 URTICEAE 777
Laurea is stated to possess no cystoliths, whilst the remaining genera have not
yet been examined microscopically). The various forms of cystoliths are
distributed in such a way amongst the different genera of Urticeae that the
members of the sub·tribes Urereae, Boehmerieae, Parietarieae and Forskohleae
have chiefly cystolithi punctiformes, while those of the sub-tribe Procrideae
have for the most part cystolithi lineares.
Cystolithi punctiformes only are present in the following genera (arranged in the
same order as in Weddell's work in DC. Prodr.): Obetia, Laportea (cystoliths small).
Scepocarpus (cyst. very small), Girardint"a. Boehmeria 1, Chamabaz'nia, Poulo/ria,
i.l1emorialis, CyPhotoPhus, Touchardia, Sarcochlamys, Neraudz'a, Pipturus, Leucosyke,
Maoutia, Phenax~ Parietaria, Gesnouim'a, Hemistyiis, Droguetia and Distemon.
Cystolithi oblongi, lineares, fusiformes, &c., are found exclusively in the genera
Nanocnide, Flewyya, Achudemia, Lecanthus, Pellion£a, Eta/ostema, Procris (cyst. very
small), Helxine and Myriocarpa; in Myrz"ocarpa the linear cystoliths radiate from
the bases of the hairs and thus produce a characteristic marking on the upper side
of the leaf. Finally cystoliths of both forms are found within the limits of the
following genera: Urt£ca. Hesperocnide, Urera, Gyrotaenia, Pilea (cyst. sometimes
also star-shaped in this genus), Poikilospermum and AustraUna.
Regarding the hairy covering, the following statements may be added.
The glandular hairs of Boehmeria and Urtica possess a unicellular stalk, and
a head composed of a few cells. The pearl-gla"nds found in Urtica macrophylla,
U. macrostachya and U. penduliflora have a much more complicated structure,
according to Meyen and Penzig. They are spherical emergences made up of
an epidermis composed of small tabular cells with no stomata, and of a central
mass consisting of a few large polygonal cells containing abundant protoplasm
and no chlorophyll, but remarkably large quantities of a fatty body; the
epidermal cells may sometimes grow out locally into hairs. Stinging hairs are
described by Weddell as occurring in Urtica, Hesperocnide, Fleurya, Laportea,
Urera, Scepocarpus and Girardinia. Those of Urtt"ca and Laportea have been
examined in detail by Haberlandt. Their structure, which is well known, is as
follows: they are rather long, conical unicellular hairs, the apex bearing a small
spherical or ovoid head, which is obliquely inserted, and readily breaks off, thus
leading to the emission of the contents of the stinging hair. The hair is fre- ;
quen tly seated on a multicellular pedestal, which surrounds its base like a cup, and
is partly formed by the periblem. The nature of the wall of the stinging hairs
is peculiar. The head and the neighbouring portions of the wall of the hair
are silicified; in the latter (Fig. 183, B) the amount of silicification gradually
decreases downwards, i.e. towards the base of the hair, this decrease in silici-
fication being first shown by the innermost portions of the wall (i.e. those
nearest the lumen of the hair); simultaneously with the decrease in silicifica-
tion calcification begins, and ultimately near the base oLthe hair silicification
:is entirely replaced by the calcification. Regarding the 'unequal thickening
of the wall of the head, and the characteristic line of fracture thus produced,
&c., see Haberlandt and Fig. 183, C-D. Since the work of Gorup-Bessanez
the irritant causing-the stinging sensation has usually been stated to be formic
acid; this, however, :is incorrect, and it is probably a substance related to the
ferments.

1 This opportunity may be taken of correcting an incorrect statement which bas passed from one
book to another. I refer to the fusifonn cystoliths of a plant described as u,t£ca macrophylla. which
De Barr figures in his 'Vergleichende Anatomie,' p. IU, Fig. 45; these cystoliths are reproduced
by Engler in his revision of the Urticaceae (in the 'Natiirl. Pfianzenfam.') under the synonymous
name BoeJmzeria platyphylla, Don, and are also mentioned in Kohl's work (lac. cit., p. 127). The
fusiform cystoliths in question do not belong to Bodlmer~"a platyjlzylla, which like otber~ species of
Boehmeria has' cystolitbi punctiCormes' only; the material investigated by De Bary was incorrectly
~etermined.
 URTICEAE

The petiole contains isolated vascular bundles in the species in'Vestigated

belonging to the genera Urlica, Parietaria and Boehmeria (Petit).
The structure of the wood and cortex have hitherto been little investigated.
In the woody species the medullary rays of the wood are broad (Urera sinuata,
Wedd., Laportea canadensis, Gaud.), or larrow· (Gesnouinia arborea, Gaud.,
according to Engler). The herbaceous species usually have broad medullary
rays,.. but in other cases (Urtica dioica) medullary rays are not present. _Demeter
and Herbst state that in species of Boehmeria the medullary rays are of excep-
tional height.
The vessels in Urera and Laportea are mostly isolated, and have rather
wide lumina (diameter reaching '08 mm.). In contact with other vessels they
bear relatively large bordered pits (diameter of border = '004-'007 mm.), while
in contact with parenchyma transHions to simple pits are present. The per-
forations of the vessels are simple (even in Boehmeria and Urtica), and the
wood-prosenchyma bears simple pits. The xylem of Urtica d";oica includes-
groups of un lignified parenchyma.
The cork originates in the subepidermal layer of cells in Pouzolzia and
Boehmeria (Moller), but at a considerable depth in the primary cortex in
Urlica dioica (Gravis). The peri cycle contains isolated groups of bast-fibres
in Pouzolzia, Boehmeria and Ur#ca. The bast-fibres in Urtica attain a 1ength
of 77 mm.; their walls are silicified (Wicke).
Literature: Meyen, Sekretionsorg., Berlin, 1837, p. 46, and ~lldl1ngsgesch. etc .. MUller, Arch. d.
Pharm. 1839, p. aSS et seq.-Payen, DeveI. des veget., Mem. pres. pat div. sav. al'Acad. roy. des
sc. de l'lnst. de France, t. ix, 1846, p. 79 et seq.-Weddell. Cystolithes etc. des U., Ann. sc. nat.,
ser. 4. t. ii, I8S'''' pp. l67-72 and pl. I8.-Schacht, Traub~korp. im Bl. v. U. etc., Abh. Senckenberg.
Gesellsch., :&1. i, 1854-5, pp. 133-53 :lnd Tab. vii.-Weddell, Monogr. des U. t Arch. Mus. d'hist.
nat., t. ix, 1856-7, pp. JO-rI et seq. (also in DC. Prodr. xvi, I, J869).-Wicket in Bot. Zeit. 186f t
p. 91 et seq.-Hanstein, Milchsaftgef., Berlin, 1864, p. 75 et seq.-Dippel, Milcasaftgef., Rotterdam,
186s.-Carnel, Sac laiteux du figuier. Bull. Soc. bot. Fra.nce 1865, p. 273.-Strasburger, in Pringsheim
Jabrb., Ed. V, 1866-7, p. 330 and Tab. xli.-Gris, Mcklle. NOllv. Arch. Mns. d'hist. nat., t. vi, 1870,
Pp,- ~64-5 and pl. 16.-Rauter, Tricbomgeb., Denkschr. Wiener Akad. 1872, pp. 24-30 and Tab.
vli-h.-Davh1, Milcbz., Breslau, 1872, p. 38 et $eq.-Martjnet, Org. de secreL, Ann. sc. nat., ser. 5,
t. xiv, 187:z, p. 178 et seq. and pi. 17.-Bot'rlage, Bydrage tot de Kennis der Houtanatolllie, Leiden,
18i5.-Delbrouck, Pflanzenstacb., Hanstein, Bot. Abh., Bd. ii, 1875, pp. H-5.-Moller, Holzanat.,
Denkschr. Wiener Akad. 1816, pp. 25-32 and 321.-Reinke, Sekretionsorg., Pringsheitn Jahrb.,
Rd. x, 1876, pp. ISi-S.-De Bary, Vergl. Anat. 1877.-K. Richter, Cyst., Sitz.·Ber. Wiener Akad.
1871, sep. copy, 34 pp. and 2 Tab.-Schmalbausen, Milchsaftbeh., Mem. Acad. des sc. de St. Peters--
bonrg, ser. 7, t. xxiv, n. 2, 1877, pp. lS-20.-C. de Candolle, Anat. comp. des feuilles, Mem. Soc.
phys. et hist. nat. de Geneve, t. xxvi, 2,1879, p. 443 and 447.-[Fougairson, Rech. auat. s. Ie groupe
des U., These, Toulouse, I 879.]-Hesselbarth, Vergl. Mat. d. Holzes, Diss., Berlin, 1819, pp. 35-
s6.-Treub, in Arch. neerland., t. xv, 1880.-Demeter, Rosanoff'sche Kr. bei U., Magyar Novenyt.
La pok '" 188 I, p. i\ 2 , Hungarian; a bstr. in Just 1881, i, p. 436 et seq. and Bot. Centralbl. 188 I, ii,
p. 34I.-Derneter, Hist. d. U., Klausenbnrg, 1881, 43 pp. and 2 Tab., Hungarian; abstr. in Just
[88[, it pp. '436-9 and in Bot. Centralbl. 188r, iii, p. ;p8.-Molisch, KoMens. Kalk im Stamme etc.,
~itz.-Ber. Wiener Akad .• Bd. lxxxiv, AM. r, 1831, p. 1 et :;;eq. and Tab. i.-Schaarschmidt, Spharokr.,
Magyar N<)\'eIlyt. Lapok 1881, pp. 134-8, abstr. in Bot. Centralbl. 188~j i, p. 47.-Bokomy, Dan:hs.
P .• Flora. 1882, p. 356 et seq. and sep. copy, p. , 3 et seq.-KaUen, Protoplasrna in Gew. 'V. Vmca
etc., Flora 1882, pp. 65, 81 and 97 et seq., Tab. iii.-MoUer, Rindenanat., 1882, pp. ~5.-Miliarikjs,
VerkieseJ.leb. ElementaroTg., Diss., Wiirzburg, 188,4, p. 28.-Chareyre, Cyslolithes, Revue sc. nat,
Montpellier, ser. 3, t. ill, 18S..,.. pp. 523-602; abstr. in Bull. Soc. bot. 1894, p. (94).-Gravis, Urii,,,,
dioica, Mem. cour. des say. ctrang. publ. par l'Aca-d. roy. de Belgiqne, t. xlvii, 1884, 2,56 pp. and
l3 pl.-Mentovich, Mark, KlausenbllTg, 188,5, Hungarian; abstr. in Just 188.;, i, p. 789.-Pitotta e
Marcatili, Vasi latie. ed i1 sisto assim., Ann. 1st. bot. di Roma, vol. ii. 188,5, ~ pp.; abstr. in Ju!Ot
J 885, t, p. 793 and Bot. Celltralbl. 1886, ii, p. 21l.-So1ereder, Holzstr., 188S, pp. 241-3.-
Haberlandt, Anat. u. Phys. d. pfl. Btennh., Sitz.·Bet. Wiener Akad., Bd. xciii, ·Abt. I, 1586, p. !:iI3
et seq.-Lohrer, Wurzel, Wigand, Bot. Hefte, ii, 188" p. 33 et 5e't--Petit, Petiole, Mem. Soc. sc.
phys. et nat. de Bon]eaux. scr. 3, t. iii, 1887, PP. 227-33 and p1. ii.-Volkens, Aegypt.-arab. Wtiste,
18 8 7, p. 14+-Schhnper, Pfi. u. Ameisen, jena, 1888, p. {O et seq. and Tab. ii.-[Hriosi e Tognini,
Anat. compo della Cannab.• Atti 1st. bot. llniversita di Pavia, ser. 2, vol. ii, 1889.]-Dehmel,
Milchsaftbeh., Diss.• ErIangen, 1889, p. 29.-DouIiot, in Ann. sc. nat., Set. 7, t. x, 1889, pp. 329-30.
-Groom, Latic. tubes, Ann. of bot., vol. iii, 1889, pp. 163-4.-Marcatili, f'asci midoll. fogl, dei
Ficus, Malpighia, iii, 1889, pp. I29-33.-Kohl, Kalks. etc., 1889. pp. Il3, Il6, t23, I36, 231 etc.
 URTICEAE 779
and Tab. iv and viii.-Scott, Latic. tissue in the leaf, Ann. of bot., vol. iii, 1889, pp....6-7.-
Giesenbagen, Cyst. v. Ficus elasHca, Flora 1890, Diss., M:uburg; see also Zimmermann and
Giesenhagen, in Ber. deutsch. bot. Gesellscb. 1891, p. 17 and 74.-Gibson and Warham, Stinging
hai~ of Urtica dioua! Proceed. Liverpool BioI. Soc,! vol. iv, pp. 9 1 -4 and fl.. I.-Lalanne, ~euilles
persist., Act. Soc. LtnD. de Bordeaux, ser. 5, t. IV, 1890, p. 63 and p. tv.-Ross, Penderma,
Malpighia iv, 1890, pp. 54-5.-J. E. Weiss, Korkbild., Denkscbr. Regensburg. bot Gesellsch. 1890.
sep. copy, pp. 54-5.-Chauveaud, Appareil laticifere, Ann. sc. nat., lIer. " t. xiv, 1891, especially
pp. IJZ-I9 and pl. 7-8.~Strasburger, Leitungsbahnen, Jena, 1891, pp. 3OG-7.-Zimmermann,
Pflanzenzelle, 2. Heft, 1891, p. I,38.-Benecke, in Bot. Zeit. 189', p. 545 and 553 et seq.-Holzner
and Lenner, Hop(en, Zeitschr. ges. Brauwesen, Jahrg. 1892-5, with many pla(es.-Houlbert, Bois
sec. dans les Apetales, These, Paris, 1893, pp. 103-30.-Penzig, Perldrifsen etc., Report of Congress.
Genoa, 1893, p. 237 and 241 et seq. and Tab. xv.-Priemer, Anat. Verh. d. LaubbI. d. Ulmac.
(cinschi. Celtid.) etc., Engler, Bot. Jahrb., Ed. xvii, 1893, pp. 419-75 and Tab. x-xi; see also Sitz.-
Her. schles. bot. Gesellscb. 1892.-H. Schenck, Anat. d. Lianen, 189.l. pp. 45-7.-Engler, in Naturl.
Pflanzenfam., iii. Teil, I. Hiilfte, Abt. I, (1894). pp. 60, 67 and 100 et seq.-Haberlandt. Trap.
LaubbL j and ii, Sitz.·Ber. Wiener Akad., Ed. ciii, Abt. I, 1894, p. 532 et seq. and Tab. iii and
Bd, eiv, Abt. I, 1895, p. 58 et seq. and Tab. i-ii,-Herbst, Markstr., Bot. Centralbl. 1894, i, p. l89
et seq.-Chimani. Milchrobren, Diss., Bern, 1895. pp. 38-41. also in Bot. Centralbl. 1895, i.-Kny,
LupulindriiseD, Text z. bot. W~ndtaf. xci, 1895. pp. 5-8.-A. Richter, Z. Anat. d. Artocarp., Mathe-
matikai es termeszetudomanyi Ertesito, Budapest, 1895, pp. l88-98, and Cudrania, Plecospermum u.
Cardiogym, Termeszetrajzi Fiizetek 1895, pp. 294-307 and 2 Tab.-Spencer Moore, Matto Grosso
Expedition, Tra.nsact. Linn. Soc., vol. lV, part 3, 1895, p. 47+ and pI. 39.-Pfitzer and Ad. Meyer,
Bluten- u. Frucbtst. von Artocarpus, Ber. deutsch. bot. Gesellscb. 1896, pp. 52-3.-A. Richter,
AnJiaris u. Artocarjus, Math. u. naturw. Ber. aus Ungaro, Rd. xiii, 1896, pp. 128-143 and Tab. iii.
-Vircbow, Blattzahne, Archiv d. Pharm. 1896, sep.- copy, p. 53.-Wittlin, Kalkoxalattasch., Bot.
Centralbl. 1896, iii, p. 67.-[Yasa.da, Cystoliths, Bot. Magazine Tokyo 1896.]-Kubla, Phelloderm,
Bot. Centralbl. 1897, iii, p. 164.-Mobius. Anat. d. Ficus-BI., Ber. Senckenberg. Gesellscb. 1897.
pp. II7-38 and Tab. ii-iii.-A. Richter, BlaUstr. v. Cecrojia, Bibl. bot., 1-4. Heft, I898.-[Biittncr,
Cortex Murure, Diss., Ert.mgen, 1896, 31 pp.-Briosi e Tognini, Anat. della Canopa, Parte sec.:
Org. veget.• Atti dell' 1st. bot. della Univ. di Pavia, ser. l, vol. iv, p. 175 et seq., 26 tav., see Bot.
Centralbl., Beiblatt, Bd. viii, 1898, p. 27.-Engler, Moraceae, in Engler, Monogr. afro Pfi.·Fam. u.
•Gatt., Leipzig, 1898, p. 8 and 38 et seq.]

•
ApPENDIX: TRIBE VIII, THELYGONEAE.
It is best to separate the monotypic genus Thelygonum from the Urticaceae
as a distinct Order, as was done by Poulsen. This genus is essentially distinguished
from the Urticaceae in its anatomy by the presence of bundles of raphides.
The structure of the leaf is bifacial. On both sides of the leaf the epidermis
bears stomata, but on the upper side they are only present in small numbers. The
stomata are accompanied on either side by one or more subsidiary cells, placed
parallel to the pore. The upper epidermis of the leaf contains chlorophyll and
tannin, and is composed of c.ells, which are not so strongly undulated as those of
the lower epidermis. At the upper end of the petiole the fibrovascular system
consists of isolated vascular bundles (Petit).
Poulsen makes the following statements regarding the structure of the stem.
The epidermis consists of small cells and bears unicellular'hairs. The cortex con-
tains no collenchyma, and is separated by a distinct endodermis from the pericycle
and the vascular ring. ,
In addition to the unicellular trichomes just mentioned, colleters with a secretory
palisade-like epidermis are met with on the young or ans cf Thelygonum (Balicka·
I wanowska).
Literature: Petit, Peti61e;Act. SOC~Linri. de Bordeaux, t. 43, 1889, p. 16.-Poulsen, in Natlirl.
Pfianzellfam., iii. Teil, Abt. i a:' 1893, p. u3.-Balicka·lwanowska, Thelygonum Cynoorambe, Flora
1897, pp. 365-6.

PLATANACEAE.
A whole series of anatomical features characterize this Order, which con~
sists of the single genus Platanus. Of these we may mention :. the simultaneous
occurrence of simple and scalariform perforations, the latter with 1-12 bars;
the broad primary medullary rays in the xylem and bast of the vascular'ring;
the bordered pitting of the wood-prosenchyma; the occurrence of sclerotic
 PLATANACEAE

parenchyma in place of secondary hard bast; the subepidermal ongm of

the cork; the absence of a special type of stoma; and the characteristic
hairy covering, composed of candelabra· hairs, and accompanied by glandular-
hairs with unicellular glandular heads. Oxalate of lime occurs in the form
of clustered and solitary crystals. Internal glands are absent.
The following statements may be made regarding the anatomy of the leaf.
In the investigated species (e.g. P. occidentalis, L., p, mexicana, Moric., &c.)
the leaves have centric structure. On both sides there is a single layer of 'pali-
sade·tissue~ the cells of which are considerably less elongated on the lower
than on the upper side of the leaf. The vas:;ular bundles of the larger veins
are provided with a ring of sclerenchyma in P. occidentalis, L.; the smaller
veins are vertically transcurrent on both sides by means of lignified tissue,
having rather wide lumina, and extending as far as the epidermis. The guard·
cells of the stomata lie on a somewhat higher level than the adjacent cells,
the stoma being surrounded by a rather large number of these cells showing
no special arrangement. In P. occidentalis stomata 'are also found in small
numbers on the upper side of the leaf. A number of water·pores have been
observed on the tips of the leaf·teeth
in P. occidentalis. Oxalate of lime is
excreted in the veins of the leaf, chiefly
in the form of clustered crystals. The
dense wooUy covering of hairs, which
is found on the, young organs, but
subsequently falls off (persistent on the
lower side of the leaf in the South
American species), is of a very peculiar
type. It consists of candelabra·hairs,
in which the main axis is formed by
a row of cells, short at the base, but
longer in the upper part of the tri-
chome; at the points at which the
transverse walls of this main axis are
situated, whorls of 3-5 unicellular rays
.- are inserted. The form of hair just
Fm. 184·-Candelabra-ha;r of PltJla"u.roccichntaUs, described is the normal type, but
L.-Original.
numerous modifications occur; reduc-
tion in the branching is very common, especially in the hairs on the sepals and
on the ovary, where one sometimes finds a certain number of simple uniseriate
trichomes, composed of long cells and only provided with an occasional lateral
branch. The peculiar hairy covering is also noteworthy from a hygienic
standpoint, since, owing to the large numbers of these hairs falling off at the
same time, they have an annoying inflammatory effect on the mucous mem-
branes (for details see Morren and Drude). Besides these clothing hairs
glandular hairs are also present. According to my investigations the glandular
hairs situated on the surface of the leaf in P. orientalis, L. and P. occidenJalis, L.
consist of a short stalk~cell seated on the epidermis, and a spherical terminal
cell. Niedenzu figures similar capitate glands with a rather long, uniseriate
stalk in P. ocddentalis var. kispan£ca, Wesmael. According to Hanstein
the glandular hairs are sometimes (P. acerijolia) combined with the clothing
hairs. some of the branches of the latter bearing glandular heads.
TIle structure of the petiole has been examined in detail by Petit in Plata-
nus occidentalis. The course of the petiolar vascular system is very com-
plicated; this is chiefly due to the well-known fact that the basal portion of
the petiole has an internal conical cavity, and encloses the axillary bud. For
details the reader is referred to the statements of Petit and Niedenzu, but the
 PLATANACEAE

following points may be mentioned here. A transverse section through the

base of the petiole shows the central wide cavity (enclosing the axillary bud),
while the wall of the latter contains a rather large number of vascular systems,
some of which consist of rings of vascular bundles. At the apex of the petiole
the fibrovascular system is composed of three arc-shaped or circular systems,
placed one above the other. According to De Bary, only some of the vascular
strands found in the stipules (viz. those in the two larger veins) are branches
of the leaf-trace; the remaining vascular bundles of the stipules pass inde·
pendently into the ring of bundles in the axis.
To the statements made above regarding the structure of the wood, we
may add that the vessels have lumina of an average size of '03-'04 mm., and
that the wood-parenchyma is scantily developed.
The pith is homogeneous and consists of active cells with thick walls
(Gris 1).
We may now deal with the structure of the cortex. The pericycle can·
tains massive groups of primary bast-fibres, united by stone-cells (having
rather wide lumina) to form a composite and continuous ring of sclercnchyma,
wrJch is developed even in branches from herbarium-material (P. occidental1:s
and P. orientalis); the primary medullary rays of the cortex, which separate
the groups of soft bast belonging to the individual vascular bundles from one
another, also become sclerose~ even in thin branches; in the latter sclerosis is
also to be seen in the primary cortical parenchyma. Subsequently sclerosis
becomes still more general in the tissues of the cortex, and especially in those
of the bast. The secondary cortex then exhibits stratification, broad layers
of sclerotic parenchyma alt~rnating with narrow bands of thick-walled, but
unlignified parenchymatous cells, and narrow strands of sieve tubes (Moller).
The sclerohc medullary rays of the cortex also penetrate in a characteristic
manner into the medullary rays of the wood as short vertical ridges. 'The
stone·cells· never have greatly thickened walls or narrow lumina, but are
distinguished by the relatively slight thickening' of their walls. The cork has
been shown to originate in the outermost cell-layer of the cortex in P. ort'entalis
and P. occidentalis; its cells have thick walls and rather wide lumina. As is
well known the formation of scale-bark subsequently takes place. The layer
of cork cutting off a scale of hark is only a few layers of cells in thickness;
its outer portion, consists of thin-walled cells, its inner portion of cells with
thick yellow walls; by the tearing of the thin-walled cells, the exfoliation of
the scale of bark is brought about, while the thick-walled layer remains as
a smooth covering for the living cortical tissue, until the next exfoliation takes
place. Oxalate of lime is found in the cortex, chiefly in the form of solitary
crystals.
Literature: Morren, Duvet du' Platane, Bull. Acad. roy. de Bruxelles, t. iv, 1~37, 8 pp. and.
Tab.-Hanstein, Harz- u. Schleimabs .• Dot. Zeit. 1868, pp. 749-51 and Tab. xii.-Gl'is, Moelle,
Non.,. Arch. Mus. d'bist. nat., t. vi, 1870, p. 267 and pl. xv.-Moller, ~olu.nat., Denkschr. Wiener
A.kad. 1876, p. 33 and~329 ..-~ Bary, Vergl. Anat., IS77.-Hesselbarth, Vergl. Anat. d. Holzes.
Diss., Leipzig, 1879. pp. S7-8.-Moller, Rindenanat., 188:3, pp. 8S-7.-So1ereder, Holzw., 1885.
p. 243.-Pdit, Petiole, Mtm. Soc. sc. phys. de Bordeaux, ser. 3. t. iii, 1887, pp.·2SO-l and pi. iii.-
Drude, Haarfib d. Platanenbl. etc., Gartenflora 1889 •. p. 393.-Ross, Periderma, Matpigbia, yol. i".
18900-1, p. JOl.-Niedenzn, in Natiirl. Pfianzenfam .• iii. Teil, Aht. 2 a, 1891, pp. I37-8.-Houlhert.
Bois lee. dans Ies Apetales, These, Paris, 1893, pp. IS4-6.-Virchow. Blattzahne, Arch. d .. Pharm.
1896, p. 62.-K uhla, Phelloderm, &t. Ccntralbl. 1897, iii, p. 165.

1 1 did not find the concentric bundles, stated by Niedenzu to occur at the peri_phery of the
pith in P. occUkn/alu and P. orimtalis. Possillly the strnctnres interpreted as concentnc medullary
Lundles by Niedenzn may be the proto:r.ylem·groups of the vascular bunnies together with tbe gsoup ..
or sclerenchyma situated opposite them on the inner side.
 LEITNERIEAE.
This Order consists of the single genus Leitneria, which in its anatomical
features recalls the Dipterocarpeae and to some extent the BaIsamifiuae also;
it has likewise been regarded as related to these two Orders on account of its
exomorphic features. Leitneria, like the Balsamifiuae and Dipterocarpeae,
has secretory canals situated just at the margin of thepith; the structure of
the secondary cortex is the same as in the Dipterocarpeae, viz. broad primary
cortical medullary rays, which are enlarged outwards in the form of wedges,
while the intervening portions of bast become correspondingly narrowed ex-
ternally, and exhibit distinct stratification into hard and soft blst. The details
of the anatomical features of Leitneria /loridana, Chapm. are as follows:
·In the in terior of the branch there is a pith composed of isodiarnetric
polygonal cells. At the margin of the pith the cells become smaller, and it is
in this small-cell~d tissue that the balsam·canals run; they are situated in
such close proximity to the primary xylem-groups of the vascular ring that
Van Tieghem and Lecomte, who were the first to observe them, originally
regarded them as belonging to the primary wood. A transverse section
of the branch shows one or two dozen of these secretory canals, each
lined by a single layer of epithelial cells, which project somewhat as papillae.
The resin is yellow and brittle; it is insoluble in water, but readily soluble
in alcohol. The wood is distinguished by its exceptional lightness and its
spongy character, thus recalling the well-known Tupelo-wood; Leitneria
inhabits the same marshy habitat as the plants from which this wood is derived
(species of Nyssa). The medullary rays of the wood are narrow, being either
oneor two cells in breadth; none of the cells of the medullary rays are elongated
to any considerable extent in the vertical direction. The vessels attain a dia-
meter of '05-'09 mm., and have simple perforations; in contact with one
another they bear bordered pits, but in contact with parenchyma of the medul-
lary rays they have large simple pits; these together sometimes resemble
scalariform perforations. The wood-parenchyma is not abundant, and is
only present in the neighbourhood of the vessels. The vessels are accompanied
by tracheides with spirally thickened walls, whilst the ground-mass of the wood
is 'composed of wood-fibres with wide lumina and simple pits. There are no
secretory canals in the wood. The cortex is rich in tannin, and, like the wood~
is devoid of resin-canals. In the secondary cortex, as stated above, the groups.
of bast be]onging to the vascular ring become narrower outwards in the form
of wedges, while the primary medullary rays between them become correspond-
ingly broader outwards. The groups of phloem a.re stratified into hard and
50ft bast. The hard bast in Lc,:tneria is composed of hast-fibres with very
wide lumina, exceeding those of the elements of the bast-parenchrnya; the
pericyclic hard bast alone is composed of cells with thicker walls. The outer
portion of the primary cortex is collenchymatous. Cortical vascular bundles
are not present (a point of distinction from the Dipterocarpeae). The cork
.arises immediately beneath the epidermis, and consists of low cells.
The leaf of Leitneria floridana has bifacial structure. The stomata are
not depressed. Beneath the upper epidermis of the leaf there is a hypoderm of
several layers, containing clustered crystals; the latter, it may be added, are
also met with in the pith and bast of the branch, and in the petiole. Two kinds
of trichomes occur: viz. simple uniseriate hairs with an often bulbous base,
two of these hairs being occasionally sunk in the epidermis side by side; -and
club-shaped glandular hairs, in which the biseriate stalk is only slightly marked
off:from the multicellular clavate head.
Three vascular bundles pass out from the branch into each leaf. They
soon unite to form a ring, the secretory c.anals here also being situated at
 LEITNERIEAE

the margin of the pith thus formed. In the leaf the secretory canals run in
a similar position (on the upper side of the xylem), and are continued into
the finest veins. There are no balsam-canals in the root.
In concluding the description of this Order we may add the b!lowing remark.
The anomalous genus Didymeles, which is indigenous in Madagascar, and is referred
to the Leitnerieae by Baillon, has no secretory canals, according to Lecomte and
Van Tieghem; in the structure of its leaf (occurrence of sclerenchymatous fibres
in the parenchyma of the petiole and in the mesophyll) Didymeles may perhaps be
classed with the Ternstroemiaceae.
Literature: Van Tieghem et Lecomte) Struct. et aff. du Leitneria, Bull. Soc. bot. de France
]886, pp. 18r-4.-Van Tieghem, in JOllm. de bot. 189], pp. 387-8.-Heim, Leit1teyia, Assoc.
frao t . p. l'avancem. des sc., Marseille, 1891, pp. 233-4. and Rech. s. les Dipte~oc:lrp" These, 189::1,
p. 175 and pI. xi.-Trelease, Leitneria jioridana, Report Missouri bot. Gard. 1894, 36 pp. and
pI. SO-H.-Engler, in Nattir!. Pftanzellfam., Nachtr. u. Reg. zu Teil ii-iv, 18 97, p. 117.

]UGLANDEAE.
This Order is well characterized anatomically. One of the most distinctive
features is the occurrence of peltate glands in all the species; the glands
vary in size. The absence of resin-canals distinguishes this Order from
the Anacardiaceae, with which an affinity has repeatedly been suggested.
The following features are noteworthy: in the structure of the leaf, the
stomata are surrounded by several ordinary epidermal <;:ells exhibiting no
special arrangement; in the structure of the wood, the medullary rays are
not broad, the perforations of the vessels are mostly simple, and the wood-
parenchyma is rather abundant; in the structure of the cortex, the develop-
ment of periderm takes place superficially, isolated groups of primary 'hard
bast are usually present in the pericycle (Carya, Engelhardtia,' Platycarya and
Pterocarya), or a composite sclerenchymatous ring occurs in the same position
(]ugl,ans), and secondary hard bast is abundant. Septation of the pith {without
sclerosis), setting in at an early stage, is characteristic of the two very closely
allied genera ] uglans and Pterocarya. The hairy covering, apart from the
pelt ate glands already mentioned,-consists of simple unicellular hairs, a number
of which are occasionally sunk side by side in the epidermis, thus forming
tufted hairs. Oxalate of lim~ is found chiefly in the form of clustered crystals,
but ordinary solitary crystals also OCCUf.
, I made a general comparative examination of the structure of the leaf
and the hairy covering in ]uglans regia, L., Carya tomentosa, Nutt., Ellgelhardtia
spicata, Bl., Platycarya strobilacea, S. et Z. and Pterocarya sorbi/oUa, S. et Z.
To the statements made above regarding the stomata we may add that
stomata of two sizes frequently occur on the same leaf-surface. and that,
in the_species_ whicb._I examined,. the stomata are restricted to the lower
side. In most- cases oxalate of lime is only present in the leaf in the form
of clustered crystals; the veins in Platycarya strobilacea alone contain solitary
crystals besides numerous clustered crystals. In the mesophyll of Platycarya
strobilacea large clustered crystals are present; and according to Radlkofer.
the same is the case in Carya porct"na, Nutt., where the crystals give rise to
transparent dots. In all the species the peltate glands attain a moderate
size; the short, unicellular or uniseriate stalk consists of cens with yellow
walls; the shield has an entire margin and is composed of a varying number
of ray-cells, which have thin walls, and become somewhat broader externally,
in accordance with the form of the, shield. In Carya tomentosa the large peltate
glands are. accompanied by others of smaller size but having the same structure;
 JUGLANDEAE

Juglans regia and Pterocarya sorbifolia, besides having large glands, have others
with a very small shield, divided into four cells by two vertical walls arranged
as an orthogonal cross 1. The glandular hairs in Engelhardtia spicata are
accompanied by simple unicellular hairs, and in Pterocarya sorbi/oh"a by (a)
simple straight unicellular hairs, which have pointed apices, and a bulbous
swollen base, and are provided with subsidiary cells, and (b) long narrow,
somewhat curly unicellular trichomes with thick walls, a few of these hairs
being sometimes sunk in the epidermis side by side; finally in Carya tomentosa,
besides glandular hairs there are tufted hairs, composed of a group of stiff,
unicellular, sclerenchymatous trichomes.
According to Petit, the basal portion of the petiole in J uglans, Carya,
Pterocarya and Platycarya contains a closed fibrovascular system, triangular
in. outline. Higher up in the petiole vascular bundles branch off from this
system, and in the characteristic region form a straight row of inversely orien~­
tated bundles, i.e. with bast above (Juglans), or one or more rings of wood
and bast, on the upper side of the annular closed principal system. No scleren-
chyma accompanies the fibrovascular system. Accordmg to C. de Candolle
the petiole of Engelhardtia- has a structure similar to that of
Juglans, &c.
The structure of the wood has been examined by me in
numerous species of the five genera forming the Order. The
medullary rays are narrow) from one to three cells in· breadth;
solitary or clustered crystals are sometimes (Carya, Engel-
hardtia) found in the medullary rays, and are contained in
ellipsoidal and swollen, or in transversely septate cells. The
vessels as a rule have simple perforations. Scalariform perfora-
tions with a· rather small number of bars have only been
observed in the primary wood in Carya aquatica, Nutt., and in
relatively small numbers in the secondary wood in all investi-
gated species of Engelhardtia. In contact with parenchyma
FIG. 185. Brancb the vessels bear simple pits, sometimes with transitions to
of Pt#roca~YG CtlU· bordered pits. The genus Carya is specially characterized by
C/u;r:4, c. A. Mey., •
Cllt SO.IlS\o show ~ne the thIck walls of the pitted vessels (principally in the autumn
~~~;~~al.rthep'th. wood), and the genus Platycarya by the spiral thickening of
the walls of the pitted vessels and tracheides. Wood-paren-
chyma is abundant. The wood·prosenchyma bears simple pits in Carya, while
in the remaining genera the pits have a distinct border, which is smaller th~m
the pore.
The structure of the cortex has been examined by Moller in J uglans regia,
L., J. nigra, L., Carya alba, Nutt., C. amara, Nutt., and by me in Engelhardtia
parvifolia, C. DC., Platycarya strobilacea, S. et Z. and Pterocarya rhoi/olia, S.
et Z. The formation of cork takes place in the subepidermal cell·layer in
Carya, J2~glans, Pterocarya and Engelhardtia. The cells of the cork either have
thin waIls and r~ther wide lumina, or they have thick walls and are flat. The
primary cortex is frequently collenchymatous. and often (e.g. PtlJrocarya
rhoi/olia) contains a very considerable numher of clustered crystals, which are
visible even with the naked eye, since they give a white colour to the transverse
section. In all cases the pericycle contains massive groups of bas t· fibres,
which in Juglans are united by a small number of stone cells so as to form
a ring. The medullary rays of the bast show considerable broadening towards

1 The teeth on the margin of the leaf also have a glandular nature in some cases (Jug/am
a-
cinerea and Pteroca"ya). They include the termination of vein, muCilage-containing cells, and
clustered crystals; aud the apex of the tooth bears a few stomata (see Reinke. in Pringsbeim's Jahrb.,
Bd. x, 1876, p. 167).
 JUGLANDEAE

the exterior in Engelhardtia only, and in this genus the appearance of the bast
in transverse section recalls that of the lime. Secondary hard bast is present
in all cases, and sometimes gives rise to distinct stratification of the bast. The
soft bast contains numerous clustered crystals; more rarely (Juglam, Carya,
Engelhardtia) solitary crystals are also present, the latter showing a somewhat
prismatic elongation in the bast of Carya amara. Regarding the structure at
the pith it has already been pointed out above that septation of the pith without
sclerosis occurs in Juglans and Pterocarya (Fig. 185) only. According to Gris.
the diaphragms consist of cells which are generally empty or contain clustered
crystals, while the peripheral portion of the pith is formed by active cells with
thick walls. The non-septate pith of Carya amara is also heterogeneous.
Literature: Gris, Moelle, Nouv. Arch. Mus. d'hist. nat., t. vi, 1870, pp. :l77-8 and pI. xviii.-
Moller, Holzanat., Denkschr. Wiener Akad. 1876, pp. 94-6 and 390.-C. de Candolle. Anat. compo
de la (eaiIle, Mem. Soc. phys. et bist. nat. de Geneve 1879, p. 477 et seq.-Moller, Rindenanat., 1882.
pp. 308-12.-So1ereder, Holzstr., 1885, pp. 243-6.-0. Bachmann, Schildh., Flora 1886, sep. copy,
pp. 17-18.-Radlkofer, in Sitz.-Ber. Munch. Akad. 1886, pp. 338-9.-Petit, Petiole, Mem. Soc. sc.
phys. et nat. de Bordeaux, ser. iii, t. 3, 1887, pp. 249-50 and pl ii; and Act. Soc. Linn. c:le
Bordeaux, t. 43. 1889, p. U and pI. ii-J. E. Weiss, Korkbild., Denkscbr. Regensburg. bot-
Gesellsch. 1890, p. ;'i4.-Houlbert, Bois sec. dans Ies Apetales, These, Pari!', 1893, pp. 156-64.-
Engler. in Natiirl. Pftanzenfam., iii. Teil, Abt. I (1894), p. 21.-F. Schneider, Esche, Forstl..natllrw.
Zeitschr. 1896, p. 421 et seq.-Kuhla, Pbelloderm, Bot. Centralbl. 1897, iii, p. llS.

MYRICACEAE.
The characteristic anatomical features of this Order, which includes the
single genus Myrica, are as follows: large pelt ate glands; vertical trans e

currence of the smaller veins in the leaf; absence of a characteristic stomatal

apparatus; narrow medullary rays in the wood; tendency to form scalarifonn
perforations in the vesse]s~ which never have specially wide lumina; wood·
prosenchyma with bordered pits j tendency to form a composite and con-
tinuous ring of scIerenchyma in the pericycle 1 j superficial formation of cork.
Oxalate of lime occurs in the form of clustered and solitary crystals. Besides
the peltate glands simple unicellular hairs are present.
The structure of the leaf in the species of Myrica has hitherto been little
investigated. In Myrica Gale, L. I found it to be almost centric, owing to the
palisade-like differentiation of the lowest layer of the spongy tissue; in M.
microcarpa, Benth. the leaf-structure is bifacial. /The stomata aTe found only
on the lower sUrface of the leaf, and are surrounded by a number of adjacent
cells. In M. Gale the cells of the lower epidermis are papillose, especially the
cells adjoining the stomata. The vascular systems in the midrib are provided
with arcs of sclerenchyma; in the smaller veins the vascular system is con-
nected with both the upper and lower epidermis by special layers of elongated
cells with wide lumina. In Myrica Gale clustered crystals are found in the
veins of the leaf, in M. microcarpa clustered and solitary cry~tals, and also
cells containing solitary crystals accompanied by small crystalline granules.
The peltate glands-consist·ofa uni-'or biSeriate stalk (the length of which varies
according to the degree of depression of the gland in the surface of the leaf),
and a shield, in which some of the ray-cells often do not reach the centre; the
secretion is deposited beneath the cuticle. In M. Gale there are simple uni-
cellular sclerenchymatous hairs in addition to the peltate glands.
The structure of the wood has been examined in 20 species. The maxi ..
mum diameter of the vessels varies between '02 and ·05 mm. In most species
the vessels are isolated, and exhibit radial arrangement in M. Gale only. The
perforations of the vessels, as already stated, show a tendency towards scalari-
1 Engler's statement, that bast.fibres and stone·cells are absent in the Myricac<:ae, is incorrect,
and is due to insufficient study of .Muller's work.
~OLUED£II 3 E
 MYRICACEAE

form differentiation in all cases. In some species (M. F(J.ya, Ait., M. Gale, L.~
AI. in tegri/olia , Roxb., }If. iavanica, Bl., AI. mexicana, \VilId., M. ,ubra, S. et
Z.) only scalariform perforations with 1-15 bars have been obser_ved. All
the other investigated species have simple (mostly elliptical, more rarely
circular) perforations accompanied by others of the scalariform type; in these
cases either the simple type is prevalent (e.g. in M. aethiopica, L. and M. Bur-
mannii, E. Mey.), the scalariform perforations being almost confined to the
neighbourhood of the primary wood, Of the scalariform type is prevalent
(e.g. in M. altera, C. DC. and M. cordi/olia, L.). The walls of the vessels are
provided with bordered pits in contact with parenchyma. Wood-parenchyma
is scantily developed. The medullary rays of the wood are narrow, at the
most 4-seriate; the cells vary in height. The w0od-prosenchyma has elements
with lumina of varying size, and bears bordered pits on its walls.
In the structure of the cortex we may first mention that the cork originates
immediately beneath the epidermis in M. Gale and M. sapida, Wall, according
to my own observations, and in M. cali/ornica, Cham. et Schlecht. according
to Moller. In the three species just mentioned the cells of the cork are sclerosed
on one side only, viz. on the inner tangential walls. In M. sapida the primary
cortex includes stone-cells, which have thick walls and narrow lumina, and
are sometimes slightly branched. The pericycle <;ontains isolated bundles of
primary bast-fibres with intermediate stone-cells, so that either an interrupted
(M. Gale) or a continuous (M. sapida and M. cali/arnica) composite scleren·
chymatous ring is formed. In the bast of M. sapida there are elongated pitted
cells, resembling sclerenchymatous fibres, and either occWTing singly or
arranged in small groups. The ends of the sieve· tubes bear several roundish
sieve-fields arranged in a longitudinal ro~. Oxalate of lime is found in the
axis in the form of solitary and clustered crystals.. It rClnains to mention
that H6hnel's statement regarding the occurrence of schizogenous resin-canals
in the cortex of ftf. sapida, \Vall. is incorrect. As will be seen from the descrip-
tion given above, I have examined this species and have found resin~canals to
be absent, as in other membCf'S of the Order..
According to Gris, the pith in Afyrica Gale consists of active cells with thick
walls.
Literature: Gris, Moelle, N()uv. Arch. Mus. d'bist. nat., t. vi, 1870, p. l84.-De Bary. Vergl.
Anat., I8n.-Hobnel, Holzer mit Hang., Bot. Zeit. 1882, pp. J66-7.-Moller, Rindenanat., 188:l,
pro 48-9.-So1ereder, Holzstr., 18SS, Jlp. 247-8.-Engler. in Natiirl. l'flanzenfam., iii. Teil, Abt. [
(1894), p. 'J7·

CASUARINEAE.
It is well known that the Casuarineae, which comprise the single genus
Cas!uzrina, are plants resembling horse-tails (Equisetum) in habit. As a result
of the reduction of the foliar organs, abundant palisade-parenchyma is
developed in the cortex of the young branches. The following additional
features characterize the Casuarineae anatomically: the occurrence of
cortical vascular bundles, which alternate with the bundles of the central
cylinder, and, after running through the length of one internode in the cortex,
pass into the central cylinder at the next node (Fig. 186 A); the structure of
the wood, viz. (a) vessels having small lumina and simple or scalarifonn
perforations, and bearing bordered pits in. contact with parenchyma.
(b) tangential bands of wood-parenchyma, (c) wood-prosenchyma with
bo!dered pits, and (d) broad medullary rays; the peculiar origin of the peri-
derm (see below), which is connected with the nature of the surface of the
young branches; stomata, placed transversely to the longitudinal axis of
the branch, and provided with parallel subsidiary cells (Fig. 186, B); finally,
t he occurrence of rows of tracheides on either side of the cortical vascular
 CAS UA RINEAE

bundles (Fig. 186, C The hairy covering (Fig. I86, D-E) consists of simple
or branched trichomes. Neith~r internal nor external glands are present.
Oxalate of lime occurs in the form of clustered and ordinary solitary crystals.
In the following paragraphs we will first deal with the structure of the young
branches, some of which in certain members of the Order exhibit no growth in
thickness. and are subsequently thrown off like leaves; a description of the structure
of the thick branches and stems will follow. -
It should be borne in mind that the young branches of Casuarina consist of
a row of joints or internodes, each of which is prolonged above into a short sheath,
surrounding the base of the next higher internode, and terminating in 4-20 teeth.
Coinciding with each of these teeth is a more or less strongly developed longitudinal
rib, which runs down the sheath and the internode below it. Between the ribs
there are corresponding longitudinal furrows of varying del'th. The teeth of the
sheatqs alternate at successive nodes, and the same relation IS exhibited by the ribs
and furrows in the successive internodes.
First we will describe the structure of the young branch in C. equiseti/olia, L.
(Fig. 186. A); this description will be followed by a consideration of the points of
difference presented by the other species. The internode in C. equisetifolia is
distinguished by having furrows of considerable depth separated by proportionately
high ribs; in a transverse section the ribs appear broad, the deep furrows very
narrow. At the inner margin of the ribs, with which they correspond in position,
the cortical vascular bundles are situated, while the bundles of the central cylinder,
which alternate with the cortical bundles, lie opposite the furrows. We will now
consider the structure of the transverse section, taking the tissues from without
in-wards. The nature of the epidermis varies according as it borders directly on the
exterior, or forms the lateral surfaces of the longitudinal furrows. The peripheral
portions of the epidermis, which form the external limit of the ribs, consist
centrally of somewhat isodiametric cells polygonal in surface-view, but towards the
furrows the cells become slightly elongated in the vertical direction. Stomata
are not found in these portions at the epidermis, in which the outer wall is con-
siderably thickened, and encloses small roundish doubly refractive bodies (not
oxalate of lime according to my investigations, Fig. 186. A and B). The stomata
are confined to those parts of the epidermis which line the longitudinal furrows.
They are arranged (Fig. 186, B) in rows, and are placed transversely to the longi-
tudinal axis of the branch. The stomata are provided with subsidiary cells lying
parallel to the pore, and are situated between epidermal cells which are/likewise
elongated transversely to the longitudinal axis of the branch. Hairs spring from
the base of the furrows (Fig. 186, D-E); in C. equiset£/oUa they are of two kinds.
viz.: (a) simple trichomes, consisting of tw:o short thin-walled basal cells, and
a longer terminal cell with thicker walls (Fig. 186. E); and (b) branched trichomes
of sympodial structure, having the form shown in Fig. 186. D. In other species
or varieties the hairs have a different structure. Thus in C. equisetilolia var.
incana Poisson figures hairs with two relatively short, superposed basal cells,
above which, dichotomously inserted, are two longer cells forming an acute angle
with one another; in some cases the dichotomous branching is repeated a second
t.ime. In the ribs of C. equiseti/olia hypodermal parenchyma lies immediately
beneath the epidermis, and is followed by one or more layers of hypoderm com-
posed of sclerenchymatous fibres, which are elongated in the vertical direction, and
are thus cut. transversely in-a transverse section of the branch. From the middle
of this hypoderm a ridge of similarly arranged sclerenchymatous elements extends
radially towards the bast of the cortical vascular bundle. The remaining portion
of the rib is occupied by a number of layers of palisade parenchyma. the cells of
which are elongated in a radial direction.
In those species which have branches destitute of furrows, or with only slight
indications of them, the structure of the outer portion of the young branch is
different. A few examples taken from Poisson's work may suffice. In C.
Chatnaecyparis, Poiss. the transverse section of the branch has a quadran~lar
outline. the four comers being produced into slight wings, and each contAlniug
subepidermal sclerenchyma, and a sclerenchymatous ridge extending towards 'the
cortical vascular bundle; on the whole of the remaining circumferential portions
of the branch, palisade. parenchyma is developed beneath the epidermis. except at
four points. situated in the middle of the four lateral surfaces. where the internal
3 E2
 CASUARINEAE

FIG. 186. CasutJyina equi~li./oli(J, L. A, TranS\'erse Sf'ction through a young stem (the crystals of OJ:alatt::
of lime have been omitted (or the sake of clearness i the bodies enclosed in tht" outer wall of the f'pidermis art·
shown Bi black dots). D, Epidermis (TOm the lateral surface of a (11I1"OW of the branch i on the left, a portion of
the epidermis, situated at tlie mouth of the furrow i it is de,'oid of stomata, and has warty prominence!! (drawn
as circJ\"S), each of which corresponds to a body deposited in thr outer wall (see A.); on the right, a portioA
of the epidermis 'yin~ d~per in the furrow; thi. showli' the characteristic Sltomata. c, A row Df st(]rage-tracheid ..~
in a longitudinal seelio. i rows like this rUIl on either .ide of the cortieal vascular hundles (cr. A). ll~a, Forms (It
hairs found in the furrows.-Original.
 CASUARINEAE

cortical parenchyma extends as far as the epidermis, whicb bears hairs at these
points; stomata are developed on the lateral surfaces wherever palisade-parenchyma
borders directly on the epidermis. In C. leucodon, Poiss. the transverse section of
the branch is quite similar, except that the four corners are rounded off; hairs an~
eutirely absent, and a subepidermal band of sclerenchyma is found in the .middle
of each of the lateral surfaces. Other species,. such as C. nodifiorG, Forst.,
C. angula'ta, Poiss. and C. Deplancheana, Poiss. var. genuina are essentially
distinguished from C. Chamaecyparis and C. leucodon by the absence of the
sclerenchyniatous ridges connecting the subepidermal sclerenchyma with the
cortical vascular bundles. In other respects the appearance of the trans"erse
section'tn the first two of these species recalls that of C. Ch a maecypa ris , whilst in
C. Depl~ncheana var. genuina it is more like that of C. kucodon. In C. Deplancneana,
Poiss. var. crassidens the distribution of sclerenchyma and palisade-tissue in the
branches is very peculiar. The branches have a rounded quadrangular outline in
transverse section. The internal cortical parenchyma only extends up to the
epidermis at four points, corresponding to the middle regions of the four lateral
surfaces; everywhere else palisade-tissue is present beneath the epidermis, but is
traversed by numerous radial sclerenchymatous ridges. Finally, Lecomte mentions
that there is a continuous sheath of palisade-tissue beneath the epidermis in
C. Rumphiana. Miq. The anatomical features, which have just been described, are
of great value for the more special diagnosis of the Casuarineae, and a detailed
examination of them from sp'ecies to species is desirable; the results thus obtained
will however only be of use If the exomorphic features are taken into consideration
at the same time; not until this has been done will it be possible, for example, to
determine whether the variety of C. Deplancheana characterized by the peculiar
arrangement of the sclerenchyma is not best regarded as a distinct species, and so
on. It may be added that Lecomte describes the occurrence of sclerenchymatous
ridges in the ribs or corners of the branch in the following species: C. Chamaecyparis,
Poiss., C. Cunninghamiana. Miq., C. Decaisneana, F. Mull., C. Deplancneana, Miq.
var. intermedia, Poiss., C. equiseti/ol£a, L. var. incana, A. Cunn., C.leucodon, Poiss.,
C. oxyclada, Miq., C. quadrivalvis, Lahill.; but not in C. angulata, Poiss .• C. decussata,
Benth., C. micYostachya, Miq., C. nana, Sieb., C. nodiflora, Forst. f., C. sUmatrana,
Jungh., C. thuyoides, Mig., C. torulosa, Ait.
The inner portion of the young branch in C. equisetifolia (d. Fig. 186, A) up to
the outer boundary of the central vascular cylinder consists of unlignified parenchyma
with thin walls. This contains the cortical vascular bundles, which lie directly
opposite the ribs, and are equal to them in number. They consist of a weakly
developed mass of xylem situated on the inner side, and a phloem-group, directed
outwards, and supported at its outer margin by a bundle of sclerenchymatous
fibres. On the right and left of the phloem-groups rows of tracheides (Fig. 186, A
and C) are situated. These tracheides are either of irregular shape, or almost
isodiametric, or elongated like fibres, and their lignified walls bear simple pits· of
various" forms. This system of water-storing elements unites the wood of the'
cortical vascular bundles with the palisade-tissue at certain ,points, and eviden~y
serves for the supply of water to the latter tissue. The parenchymatous cortex IS
bounded towards the central cylinder by an endodermis. The vascular bundles of
the central cylinder are separated by rather broad medullary rays, and at the outer
margin of the masses of bast there are groups of· sclerenchymatous fibres; these
groups vary in size.
The structure of the leaf,sheath on the whole agrees with that of the outer
portion of the cortex i~_ the young branch. For information on this point, and also
on the peculiar transverse bundles of fibres, which are present in the commissural
portions of the sheath, and prevent tearing, see C. MUller and Morini.
The occurrence of cortical bundles is connected with tht~pa.th followed by the
vascular strands. The latter begin their course "in the teeth of the leaf-sheaths, and run.
downwards in the ribs of the sheath j- at the node they pass into the internode, a:.~d
continue their course in the latter. running in the ribs, which are merely continua- r

tions of those on the sheath. At the next node they enter the inner ring of DundleS.
but first fuse with_ the' leaf-traces' which formed the inner vascular ring in the
prec~ding internode. , To accomplish this fusion each of the inner vascular bundles
forks at the node into two short branches, each of which fuses with the nearest
cortical bundle' entering' the inner ring. .
The peculiar origin of the cork in the young branch also deserves notice; it has
 CASUARINEAE
been investigated by Sanio, Low, Ross, Lecomte, and others. The phellogen first
appea~s in the fur~ows, where it is subepiderm~. In the ribs. it dey~l0I'.s on ~c
inner side of the palisade-parenchyma, and accordmg to Lecomte Its pOSItion IS outside
the cortical vascular bundles in the basal portion of the internode, whilst further up
it cuts through the middle of the cortical bundles, and at a still higher J~el.it lies
on the inner side of"these bundles. In C. tOf'ulosa according to Sanio the cork,
formed beneath the furrows, consists of cells which have their inner tangential
walls strongly thickened, whilst on the inner side of the ribs there are at the most
only indications of one-sided thickening in the cork-cells.
I t now remains to consider the structure of the thicker branches and of the main
stem. Numerous stone-cells are developed in the parenchymatous outer cortex as
the bra.nch increases in age; large numbers of stone-cells also arise between the
groups of primary bast-fibres belonging to the vascular ring. and connect them so
as to form a composite and continuous ring of sc1erenchyma (C. qua.drivalvis,
Labill. according to MOller). In the young branches of C. eq'Uiseti/olia only
clustered crystals are found, occurring in the palisade·tissue, in the groups of bast.
and in the parenchymatous outer cortex, but in older stems numeliOus solitary
~rystals are also present in the outer cortex. According to MOller's in$'estigation~
the bast in C; equisetifolia, L. contains abundant sclerenchyma in, the form of
groups of bast·fibres and stone-cells, these being accompanied by numerous
4Chambered fibres with solitary crystals. The inner portions of the primary corlical
medullary rays also undergo considerable sclerosis, and in older stems penetrate
the wood in the fonn of sclerenchymatous ridges. The sieve-tubes have sieve-fields
showing scalariform arrangement. The following statements may be made re-
garding the structure of the wood. The vessels never have a large diameter
(maximum diameter between '015 and ·06 mm.), and are scattered in a trans-
verse section. They have simple, mostly elliptical perforations, which, in all the
species (14 in number) examined by Poisson and myself, are accompanied by
scalariform perforations; the latter are at any rate present in the neighbourhood
of the primary wood, and often also further out in the secondary wood. In some
cases peculiar forms of scalariform perforations (some of them distorted forms) are
found; they have been described in detail by Boodle and Worsdell. The walls of
the vessels. even where they are in contact with parenchyma. are provided with
bordered Vits. Spiral or reticulate thickening of the pitted vessels is not uncommon
(C. laumil,s, Link, C. mz·crostachya, Miq .• C. nana, Sieb .. C. paludosa. Sieb., C. stricta,
Ait.). Some of the medullary rays of the wood are distinguished by their breadth.
In the herbarium-material of some species (C. equisetifolia, L.. C. Hugeliana, Miq.,
C. microstachya, Miq.) I did not meet with specially broad medullary rays, but this
does not neceSsarily mean that broad medullary rays are absent in these species, for
it has been shown that the rays sometimes only broaden out in their outer portions,
where they may ultimately attain a breadth of 24 cells or more. According to
Goeppert the formation of new vascular bundles sometimes takes place in the broad
medullary rays of the older wood in the same way as in Clematis. The wood·
parenchyma forms tangential bands in the transverse section, and is very strongly
developed. The wood-prosenchyma has thick walis, is sometimes distinguished by
the presence of a gelatinous layer, and always bears bordered pits, though in
varying numbers. In those species in which the pitted vessels were stated above
to be spirally thickened, a similar thickening is also found on some of the elements
of the wood-prosenchyma (except in the case of C. stricta).
Literature: Goeppert, Anat. Ban d. C., Linnaea, 1841, pp. 747-56, and Tab. iv.-Stache, De
Casuarinis. etc., Vratisl., 18S5.-Sanio, Kork, Pringsheim Jahrb., Bd. ii, 1860, pp. 103-5 and Tab.
xiii, and Holzk., Bot. Zeit., 1863.-E. LOw, De C. caulis foliique evolutione et structnra, Diss.,
Berolini. 1865,54 pp.-Poisson, Casuan'na, Nauv. Arch. Mus. d'hist. nat., t. x, 181+. pp. 59-III
and )11. iv-vii.-Moller, Holzanat., Denkschr. Wiener Akad. 1876, pp. 19-20 and 315 and Tab. i.-
De Bary, Vergl. Anat. I877.-Moller, Rindenanat., 1882, pp . .o4S-7.-S01ereder, Holzstr., J88~,
pp. l.of.8-So.-Schnbe. Blattarme Pit, Breslau, 1885, p. ~.of. and Tab. ii.-Lecomte, Anat. de la tige
et de Ia feume des C., Bull. Soc. bot. de France 1886, pp. 311-J7.-C. Miiller, in Pringsheim Jahrb.,
Bd. xix, 1888, pp. ·571-.~.-Ross, Tessuto assim. e periderm a, Nuov. Ciom. bot. Ital., vol. xxi, 1889,
pp. l4I-4 and tav. ii; see also Ber. deutsch. bot. Gesellsch. 1886, pp. 367 et seq.-Engler, in
Naturl. Pflanzenfam., iii. Teil. Abt. I (1894), pp. 16-18.-Boodle and Worsdell, Compo anat. of th~
C. etc., Ann. of bot., vol. viii, 1~9", pp. 231-6.. and pI. XV-x\·i.-[Morini, Anat. del canle e della
ioglia delle C .• Mem. della R. Accad. delle scienze dell' 1st. di Bologna, str. 5, t. iv, 189+.]-Morini,
.'\rca ,;oJ)D.ettiva della gua.ina fogI. delle C .• Malpighia 1896, pp. l04-19 and tav. ix.
 79 1

CUPULIFERAE.
I. RE'fIEW OF THE ANATOMICAL FEATURES. The following anatomical
feature's,3lt common to the members of this Order: a tendency to form
scalarifonn perforations in the vessels; wood-prosenchyma with bordered pits
(with thel single exception of Nothofagus); a composite and continuous, and
in most cases permanently continuous, ring of sclerenchyma in the pericyc1e;
subepidermal development of cork; stomata without special subsidiary cells;
the occUl'rence of glandular hairs; the vertical transcurrence of the medium-
sized vems in the leaf. A number of structural features in the axis are of
value for the 4istinction of genera or groups of genera. Some of the medullary
rays of the wood are very broad in the species of Fagus belonging to the
section Eufagus, and in Quercus, but narrow, 1-4 cells thick, in the remaining
mem_rs ofl the Order. The Tribes Betuleae (Betula, Alnus) and Coryleae
(Carpinus, O!1rya, Corylus) are characterized by the radial arrangement of the
vessels in the transverse section of the branch; in the Tribe Quercineae
{Quercus, Castanopsis, Castanea, Fagus) this feature is more or less indistinct.
There are only two genera, Betula and Alnus, which bear exclusively bordered
pits on the walls of the vessels bordering on parenchyma of the medullary rays;
in the remaining genera these portions of the walls are provided with large
simple pits and bordered pits. Sealariform perforations are the only type
present in the vessels in Betula. Alnus. Corylus, and Distegocarpus Carpinus,
Sieb. et. Zucc., while in Ostrya, Fagus, Castanea, Castanopsis, and Quercus both
. simple and scalariform perforations are found. In most of the CupuIiferae
(species of Carpinus, Ostrya, Corylus, Quercus, Casta·nea) the secondary bast
contains groups-of bast-fibres; in Betula, Alnus, and Fagus they are replaced
by groups of stone-cells. Hypoderm haS only been observed jn the leaf in
certain species of Alnus. There are no special internal secretory elements,
'but many species are characterized by the presence of a mucilaginous epidermis
in the leaf. Oxalate of lime is excreted in the form of clustered and soli t~ry
.crystals; large, well-differentiated solitary crystals, occupying correspondingly
'large idioblasts in the mesophyll, give rise to transparent dots in the species
-of Carpinus (ind. Distegocarpus) and Ostrya. ",The hairy covering is formed
by clothing and glandular hairs (Fig. 187). The clothing hairs are: (a) simple
unicellular. or uniseriate (with thin transverse walls), generally sc1erenchyma-
tous triC~omes; (b) tufted hairs (species of Castanea and Quercus); and
(c) peltate hairs (species of Castanea and Quercus). The glandular hairs are
of various shapes; glandular scales are found in the species of Betula and
Alnus~ and also in Notholagus.
2. STRUCTURE OF THE LEAF. The leaf-structure has been examined in
detail by Boubier in the Betuleae and Coryleae; my own investigations
extended to Betula alba~"L., Alnus glutinosa, Willd., Carpinus Betulus, L.,
Ostrya carpinijolia, Seop., Corylus Avellana, L., Quercus Farnetto, Ten.,
Q. [lex, L., Castanopsis indica, A. DC., Castanea vulgaris, Lam., Fagus (Eutagus)
sylvatica, L., and F. (Nothofa~us) antarctica, Forst. The leaf-structure varies;
it is either distinctly bifacial (e. g. in Fagus antarctica) or typically centric,
the mesophyll consisting of palisade-parenchyma throughout its entire thick-
ness (e. g. in Quercus ]lex) or only subcentric, the lowermost layer of the
mesophyll being differentiated as palisade. Boubier found the mesophyll
to have bifacial structure in all the Betuleae and Coryleae examined by him,
with the single exception of Ostrya carpinifolia. The epidermal cells have
straight or undulated lateral walls. Hypoderm has been mf:t with on the
upper side of the leaf in certain species of Alnus (Alnus acuminata. A. elliptica,
792 CUPULIFERAE

A. firma, A. glutinosa 1, A. incana, A. jorullensis, A. nepalensis, A. obloltgifolia~

A. pubescens, and A. rhombi/olia); it consists of a single layer, the compofieat
cells of which vary in size. The occurrence of a mucilaginous epidermis in
the leaf is a far more frequent phenomenon. I t has been ob$.etved: in all
the species of Betula examined by Boubier (including B./ruticosa, collkary to
Radlkofer's statement); in the genus Alnus, in A. cordi/olia, A. iapantca,
A. maritima, A.· orientalis, A. rubra, A. serrulata, and A. viridis (l3o.l).bie.r); in
Corylus americana and C. rostrata, and in Carpinus duinensis and C,'j!ponica
(according to Boubier); but also in Corylus A vellana, Quercus RMnetJo~
Q. pedunculata, Q. robuT, Castanea vulgaris, and Fagus antarctica. 5.ijnilarIy,
some of the hypodermal cells in the species of Alnus cited above (except 4

A. /irma, and thus including A. glutinosa) also have mucilagin~)Us inner

membranes. Boubier mentions the formation of papillae in the~lbwer'epidermis .
of the leaf in Betula corylifolia, B. nigra, Alnus nepalensis, A. oblongi/olia, 'and
A. pubescens, the papillae being only slightly differentiated in tli~ ~peqes 4)f
Alnus. The stomata are restricted to the lower side of the leaf in almost
all the investigated species (exception: Alnus orientalis), and.. are always
surrounded by several irregularly arranged epidermal cells. The larger veins
of the leaf are provided with arcs of sclerenchyma; those of medium size are
distinguished by the fact that they are vertically transcurrent, extending as
far as the epidermis on both sides of the leaf by means of variously difieren-
Hated connecting tissue.
Oxalate of lime occurs in the leaf and axis in the form of clustered and
solitary crystals, both forms being generally present in the same species, but
exhibiting different distribution; the latter feature can only be employed for
special diagnosis. Sanio (Monatsber. def Berliner Akad. 1857) mentions the
occurrence of crystal-sand in the cortex in Alnus glutinosa and Betula alba;
according to my own investigations, however, this is incorrect. In all the
species of Carpinus (see Blenk) and in Ostrya (according to Boubier) the leaf-
tissue includes idioblasts, which are filled with solitary crystals, and in both
genera give rise to pellucid dots in the leaf.
The clothing hairs are simple unicellular or uniseriate trichomes, or stellate
or pelt ate hairs. Simple unicellular hairs have been met with in species of
Betula, Alnus, Carpinus, Ostrya, Corylus, Castanopsis, and Fagus; simple
uniseriate hairs i~ species of Alnus, Ostrya, and Corylus. In Corylus Avellana
and Ostrya carpinifolia the wal1s of the hairs exhibit a curious striation, which
is connected with the internal structure of the wall. Stellate hairs have been
shown to occur in Castanea vulgaris, Quercus Farnetto, and Q. Ilex. They
consist of a tuft of sclerenchymatous cells (hairs), the whole group being either
inserted directly in the epidermis, or seated on the apex of a short multi-
seriate pedestal developed from a group of epidermal cells. The peltate hairs
(Fig. r87, A), which I examined in Castanopsis chrysophylla, A. DC, are allied
to the stellate hairs; according to PrantI, they occur not only in species of
Castanopst's but also in the species of Qu.erctts belonging to the section
Pasania. The peltate (glandular?) hairs of Castanopsis chrysophylla, are
filled with yellow contents and consist of a low basal cell and a medium-sized
shield with a slightly undulated margin; the ray-cells, only some of which
reach the centre of the shield, have thin walls and exhibit secondary division-
walls. Glandular hairs of various forms have been observed in the genera
Betula, Alnus, Carpinus, Ostrya, Corylus, Quercus, Castanea, and Fagus. The
two genera of the Betuleae (Alnus and Betula} are characteriz~d by having
glandular scales (Fig. 187, B-D). These possess (a) a short but broad stalk, which
1 "'alliczek's statement to the contrary (loc. cit.. p. 336). viz. that Alnus glutiJlosa has a"{)ne~
layered, partly mucilaginou! epidermis, i:; incorrect.
 CUPULIFERAE· 793
l'sually projects into the shield, and is composed of several layers of low,
suberized cells ; and (b) a shield, thecellso£ which appear elongated like palisade-
tissue in sections of the gland, and are mostly polygonal in surface-view. Similar
glamular scales occur in Fagus a'Htarctica, and probably also in other species
of the sectwn Nothofagus.. The external glands found in the species of Corylus
(Fig. 187, E-F) have a clavate shape; their basal portion is formed by a few
rows of low, apparently suberized cells, whilst the upper, more or less swollen
portion consists of a few thin-walled cells. According to Boubier, Ostrya
virginiccl (Fig. r87, G), Carpinus cordata, and C. Tschonoskii have glandular
hairs with a short uniseriate stalk composed of a few low cells, and a discoid
flattened head consisting of a rather larger number of cells; Carpinus Betulus
(Fig. 187, H), C. caroliniana, Ostrya carpini/olia, and Castanea vulgaris possess
glandular hairs with a rather short, uniseriate stalk composed of a few cells,
and a spherical or ellipsoidal head consisting of several cells exhibiting no

.~

FiG. 187. A CasttJnopsi's cnrystJ/J/'1U4, A. DC. B-D, Feltate glahd~ of BetuitJ alba, .'L.: H, in transverse
section; c, ~lal1dular disc ~11 at a fow, and D, at a high fOc1lS. K-F, Cory/us Ave/lana. G, Ostrya viy,einica.
H, Cayeinrl8.Betulus.
, I, Q~YCUS Fanutlo.-A-D and I original, E-ll after Bonbier•

special arrangement. Glande; of the latter kind are also to be met with,
though rarely, in Quercus Farnetto, where they accompany tubular. uniseriate
external glands, which are curved in various ways (Fig. r87, I). Lastly, we
may mention the rather_larg~ glands observed in Coyylus americana, C. !erox,
and C. Avellana (hi the last of these species only on the petiole) ; they have
a multiseriate stalk and a flattened spherical head composed of numerous
cells. Having described the glandular hairs, we may make a few remarks on
the glandular nature of the leaf-teeth, this subject having been investigated
in species of Alnus, Betula, Carpinus, and Corylus. According to Reinke, the
glandular character in Alnus cordata and Betula alba is due to glandular
shaggy hairs with a structure similar to that .of the glandular scales (i. e. they
consist of a small amount of ground~tissue and a palisade-like epidermis),
whilst in Carpinus Betulus and Corylus A vellana it is caused by glandular
hairs similar in structure to those on the foliage leaves. According to
Hanstein, identical, or at any rate similar, glandular scales or hairs are also
met with in Alnus, Corylus, Carpinus, and Ostrya as a protection to the buds.
794 CUPULIFERAE

The structure of the petiole has been examined by C. de Candolle and

Petit, and more especially in the BetuJeae and Coryleae by Boubier, and in
Quercus by Bossebreuf and Pereira. Without discussi.ng points of detail,
which are of importance only for the diagnosis of species, we may note the
following features as being useful in generic diagnosis; for other _peints the
reader is referred to the works cited below. In all the genera of the \Juercineae
(Quercus" Castanea, Castanopsis, Fagus) the characteristic region of the petiole
contains a closed annular fibrovascular system; according to Praml, five
vascular bundles pass out into the petiole in these genera. Th6 genera
Alnus, Carpinus, and Corylus agree in the structure of the characteristic
region, the fibrovascular system forming in most cases a closed or almost
dosed ring of wood and bast, above which lies an arc of wood and bast (with
the xylem directed upwards); the number of bundles in the initial-region is
three in Corylus, and usually three in Alnus, though from five to seven may
be present. In the structure of the characteristic region the genera Ostrya,
Ostryopsis, and Betula agree in as much as the vascular system in transverse
section is open on the upper side, and has a lyrate or arc-like shape; there are
three vascular bundles in the initial region in these genera.
3. STRUCTURE OF THE Ax~s. The structure of the axis in the Cupuliferae
has been thoroughly investigated. The following description is chiefly based,
on Moller's' Rindenanatomie' and the statemehts in my 'Holzstruktur 11,
The origin of the cork is the same in all cases, to judge from the concor-
dant results of investigations carried out by MOller, J. E. Weiss, Douliot, and
others, who examined numerous species of the genera Betula, Alnus, Carpinus il

Ostrya, Corylus, Quercus, Castanea, and Fagus; the phellogen always arises in
the outermost cell-layer of the primary cortex. The cells of the cork vary
in their nature. The well-known white exfoliating cork (not bark) of the birch
is_ distinguished by the presence of linear (transversely placed) lenticels; it is
dearly differentiated from the fifth year onwards, and consists of alternating
layers of tabular thick-walled cells and scarcely flattened thin-walled cells, the
latter being filled with minute white granules (betulin 2). In Alnus the cells
of the cork, which is persistent for a long time in this genus also, arc rather
low and almost tabular. Amongst the Coryleae, Ostrya (0. virginica, Willd.)
and Corylus (C. Colurna, L.) show a stratification of the cork similar to that
found in Betula, layers of cells with wide lumina alternating with layers of
tabular cells; in Carpinus Betulus, L., the cells of the ..:ork are tabular and
have thick walls. The species of Quercus have for the most part flat cork--cells
with thick walls, while in Fagus sylvatica the cells have thin walls and are
tabular, and in Castanea-vrtlgaris they have thick walls, and are maia-ately
flattened. The outer portion of the primary cortex is distinctly collenchyma·
tous in all the members of the Order. Both clustered and solitary crystals
may occur in the primary cortex. In Alwlts glutinosa and Ostrya t'irgz"nica
large intercellular spaces are formed in the inner portion of the primary cortex
in consequence of the tangential elongation of the cells and the separation of
the walls in the tangential direction. The pericyc1e in axes not exceeding
a certain thickness contains a composite and continuous ring of sclerenchyma
in all investigated species. In most members of the Order this ring is retamed
during growth ill thickness, and only in Castanea vulgaris and Betula alba does

1 The latter give the results of an investigatioll, which extended to all the species 01 all the genera of
Cnptlliferae rep,esented in the Herbarium Monacense in the year J 884 (excluding Ostryopsis).
~ With reftrence to betulin. besides other works, see Hohne1, Kork. etc., Sitz.-Ber. Wiener Abd ..
tkl. hlK'1, Abt. I, 1877. and Merklin, in Melanges biologiques St. Pde.l6b., iv, 1865. p. 565 et seq.
According to Bonbier. betlllin does not oeem in all the species of B,tula, being absent in JI.
ft/uy;lltii and B.''':fra.
 CUPULIFERAE 795
it undergo local rupture' sooner or later. Groups of secondary bast-fibres
have only been observed in the genera Carpinus, Ost7ya, Cory/us, Quercus, aud
Castanea, not in Alnus, Betula, and Fagus. They often give rise to a distinct
stratification of the secondary bast. In Alnus, Betula, and Fagus groups of
stone-rells are developed in place of the secondary bast·fibres, while in the
secondary bast of Quercus similar groups of stone-cells accompany the groups
of bast-fibres .. A very peculiar feature is the sclerosis of the inner portions of
the prim~ry medullary rays in the cortex of Fagus sylvatica and certain species
of oak, :3nch as Quercus Suber; the sclerosed groups of cells, as in Platanus,
form crest-like projections on the inner side of the cortex, penetrating the
med~lary rays of the wood, and thus establishing a fir~ dovetailing between
the wood and the cortex.
The characteristic features in the structure of the wood have already been
summarized above, but the following special points in the. individual Tribes'
and genera may be mentioned. The genera Alnus and Betula (Betuleae)
agree in almost all respects in the structure of the wood; they have: (a) narrow
medullary rayS', which are one or two, or at the most four cells in breadth,
the cells being mostly elongated in the radial direction, and containing
clustered crystals; (b) vessels, which have rather small lumina (diameter
reaching '04 rom.), and are arranged in radial rows, bear bordered pits in
contact with parenchyma of the medullary rays, and have exclusively
scalariform perforations with 5-30 bars; (cb...a small amount of wood-
parenchyma; and (d) wood-prosenchyma with W;t\,}lumina,and walls furnished
with small but distinct bordered pits, the borders' being smaller than the pits,
which are not numerous. The different size of the bordered pits on the
common walls of contact of the vessels servt's to distinguish the two genera;
for the diameter of the border is only ·Q017 mm. in Betula, but .003-.004 mm.
in Alnus. The Coryleae (Carpinus, incl. Distegocarpus, Ostrya, and Cory/us)
show certain points of agreement with the Betuleae, viz. medullary rays,
which are one or two, or at the most three cells in breadth; wood.prosen-
chyma with a constant type of structure (characterized by wide lumina,
and small but not numerous bordered pits); and the always prevalent
radial arrangement of the vessels, which have small lumina (maximum
diameter = ·04 mm.). In other respects, however, there are a number of
differences. The walls of contact between two vessels bear densely packed
and rather large bordered pits, the borders being sometimes hexagonal owing
to mutual contact. \Vhere they border on the medullary rays the walls of
the veS!iels are furnished with almost simple pits. Spiral thickening of the
pitted vessels has been observed in Carpinus Betulus, L., C. caro/iniana, Walt.,
C. duinensis, Scop., C. viminea, Lindl., Distegocarpus laxif/ora, Sieb. et Zucc.,
D. Carpinus, Sieb. et Zucc., Ostrya carpimfolia, Scop., O. virgin":ca, Willd., and
Corylus Colurna, L. I:l all the investigated species of Carpinus and Ostrya,
as well as in Distegocarpus laxijlora, the perforations of the vessels are for the
most part simple and elliptical; only in the neighbourhood of the primary
wood, and more rarely also at sO!TIe points in the secondary wood, scalariform
perforations with few bars are present. Distegocarpus Carpinus and all the
investigated species at Cory/us, on the other hand, have exclusivdy
scalariform perforations with a varying number of bars (4-r6). Wood-
parenchyma is developed in greater abundance in the Coryleae .than in the
Betuleae, and sometimes forms tangential bands. Amongst the Quercineae
the genus Fagus shows considerable diversity in the structure of the wood,
and the differences agree with the splitting of the genus into two genera,
Notlwfagus and Eulag~ attempted Itly certaw authors. Thus the s_I)@eit:!s
of Eujagus po~sess very broad medullary rays, which attairt a. breadth of eigl)t
celJs even in ·,the ~vood of the second year, whilst in the 'specits of Notho-
 7if> CUPULIFERAE

fagus, which are indigenous in the antarctic regioris of America and Australia, the
r dYS are only f'<>m one to two cells broad; besides this difference we have the fact
that the wood·fibres have bordered pits in EtI.fagus only, while in N othojagus they
bear simple pits, and are sometimes septate. In all the species of Fagus the
vessels have smaller lumina than in the remaining genera of Querciueae; they
alsp exhibit a more or less distinct radial arrangement. The perforations
of the vessels axe for the most part simple and elliptical, often elongated..
elliptical, in outline; in all the species, however, scalariform perforations have
also been observed; they have rather a small number of bars, and occur in
the neighbourhQod of the !,Jrimary wood, sometimes also in the secondary
wood. The walls of con tact between the vessels bear numeroUS bordered
pits, which occasionally show a scalariform arrangement; in contact with
parenchyma of the medullary rays simple pits are present on the walls
of the vessels. Spiral thickening of the walls is found in Pagus australis,
POpp., and F. pro cera, P<5pp. Molisch met with deposits of carbonate
.of lime in the vessels of the heart·wood in Fagus sylvatica, and the same
feature is found in Betula alba. Wood-parenchyma is scantily developed
in Fagus. The structure of the wood in the two genera, Castanea and
Castanopsis, belonging to the Quercineae, shows close agreement. Only
narrow medullary rays, 1-3 cells broad, extend outwards from the {our-
rayed pith. The vessels are scattered in the transverse section of the
branch, the radial arrangement of the vessels seen in most Cupuliferae being
suppressed in this case. The perforations of the vessels are chiefly simple,
and circular or elliptical; but, as in the other members of the Order having
simple perforations, those of the scalariform type are to be found in the
neighb&urhood of the primary wood, and in some cases in the secOIldary wood
also. In contact with parenchyma of the medullary rays the walls of the
vessels bear simple pits. The wood-prosenchyma has bordered pits in Castanea
and Castanopsis; the wo?d-parenchyma, which sometimes contains solitary
crystals, is developed in abundance. The genus Quercus agrees with Castanea
and Castanopsis in the following features: (a) the tendency to form scalariform
perforations in the vessels; (b) the occurrence of simple and bordered pits on
the walls of the vessels in contact with parenchyma of the medullary rays;
(c) the bordered pitting of the wood-prosenchyma; (d) the abundant develop-
ment of wood-parenchyma, which frequently contains crystals; cmd (e) the
suppression of the radial arrangement of the vessels; it is distinguished from
these genera, however, by the presence in the wood of broad medullary rays,
extending outwards from a pith· which is five.rayed in almost ~l] cases
(exceptions: Quercus lappacea, Roxb., and Q. macrolepis, Kotschy). Amongst
the perforations of the vessels in the secondary wood of Quercus, t~lOse of
the simple, elliptical or circular type invariably preponderate, but in all the
species which I investigated (sixty-seven in number), and likewise in those
examined by Abromeit, scalariform perforations were always present, .if not
in the secondary wood, at any rate in the neighbourhood of the primary wood ..
With regard to the grouping of the species of Quercus on the basis of the
.anammyof the wood, see Abromeit.
The structure of the plth has been examined by Gris. Alnus and Betula,
Carpinus and Ostrya, Quercus, Fagus, and Castanea have a homogeneous'yith,
whilst in Corylus alone it is heterogeneouS. In Corylus the periphery 0 the
pith is composed of active cells with rather thick walls, whilst, the ~entraI
portion consists of empty cells with rather wide lumina and thin walls.
Regarding an abnormal occurrence of medullary vascular bundles with central
phloem in Alnus glutinoS4 see Ktinkele.
Literature: Frank, in Bot. Zeit. 1864. p. 377 et seq.- .Hanstein, Harz· u. Schleim#bs., Hot. Zeit.
1868, pp. 72l and 7,6 ,et seq. and Tab. xi-xii.-Gris, Moelle, NOllV. Arch. Mus. d'hist. nat., t ..vi,
 CUPULIFERAE 797
l870, pp. 279- 84 and pI. xviii-xix.-Radikofer, in MOi}ogr. Su:/ania, 1875, p. !04.-Moller,
Holzanat., Denkscbr. Wiener Akad. J876, pp. <10-25 and 316.-Reinke, Sekretio.sorg., Pringsheim
J&brb., Rd. x, 1876, pp. 139-40.-De Bary, Vergl. Anat. J877.-C. de Candolle, Anat. compo des
feuilles, Mem. Soc. phys. et d'hist. nat. de Geneve 1819, P..... 3 et seq.-He!lSelbartb, Vergl. Anat.d.
Holtes, Diss., Leipzig, I8i9, pp. 15-34.-Molisch, Kohlens. Kalk, Sitz.-Ber. Wiener Akacl.,
Bd. lxxxiv, Abt. I, t881, p. 18.-Mollel, Rindenanat., 1882, Pp.49-68 (see here the older literature on
the -.Datum), of the cortex in the C.)-Abromeit, Anat. d. Eichenbolzes, Diss., Konigsberg, 1884. scp.
copy from PringsheimJahrb., Rd. xv, 1884, pp. z09-81.-Blenk, Durcbs,P., Flora 1884, p. 371 and Scp.
copy, pp. 83-+-Soiereder. Holzstr., 1885, pp. 2!jO-9.-Kny, Hola v. Quercus suber., Text xu bot.
Wandtaf.)xxiv-lxxvi, 1886, pp. 303-3~:-Pt:tit, Petiole, Me~. Soc. sc. phys. et na~. de Bordet\lx,
5/:r. 3, t. llJ, 1887, pp. 2..f-2-6 and pI. 11; and Actes Soc. Linn. de .Bordeaux, t. ft.J 889, p. J 7.-
Hartig and Weber, Holz d. Rotbuche, Berlin, 1888, p. 20 et seq.-Douliot, ill APD. !i.t. nat., Ii(~r. 7,
t. x, 1889, p. 332.-Lalanne, Feuilles persi!i.t., Act. Soc. Linn. Eordeaux, ser. 5, t. iv, 1890, P. 11 J
and pI. vii.-J. E. Weiss, Korkbild .. ·Denkschr. Regensb. bot. Gesellsch. 1890. sep. copy, pp. ~:.I-4.
-Ros&, Periderma, Malpighia, vol. iv, 1890, p. 183.-Arcangeli, in Nuov. Giom. bot. Ital., vol. xxiii.
I SIp • p. 37o.-StrasDuTger, Leitllngsbahnen, 1891, _pp• .:I 66-76.-Houl bert, Bais Sec. <lans les
A petales, These, Paris, 1893, pp. 132-54.-W alliczek, Mem branschleime, Pringsheim Jahrb., Bd. xxv.
1893, p. 209 et seq.-Prantl, in Natiir!. Pflanzenfam., iii. Teil. Abt. 1 (189....). pp. 38 and 47 et leq.
-Bosseboeuf, Petiole riu Quercus, Bull. Soc. bot. de .France 1896, pp. 26O-S.-Boubier, Anat. SYlt.
de Betulacees~Corylacees, Malpigbia 18.26, pp. 349-436.-Kra·~ '!' 5yst. d. Buchen. Anil. Wiener
Ho{museum, Bd. xi, 1896, pp. r6r-2.-lPereira da Fonseca, Estudo compo da cstructura do peciolo
de alg. cipee. de Quercus. Bolet. da 50ciedade Broteriana Coimbra, xiii, 1896, pp. 48-59 and Tab.]
-Reiche, Chilen. HolzpA., Jahrb. f. win. Bot.. Bd. xxx, 1897. P.96.-Virchow, Blattzahne, Arch.
d. Phann. 1896. p. 62.-Kiinkele, Strangbild. im Marke von Alnus C'ulinosa, Bot. Centralbl. 1897.
iii. pp. 1-6 and Tab. i.-Kuhla, Pbelloderm, Bot. Centralbl. 1897, iii, pp. 118 and 161.

SALICINEAE.
The two genera of this Order, Salix and Populus, have the following
anatomical characters in common: in the structure of the axis-superficial
development of the periderm, isolated bundles of bast-fibres in the pericycle,
stratification of the phloem into hard and soft bast, narrow medullary rays in
the wood, simple perforations in :tt~essels, simple pits on the walls of the
vessels where they are in contact ~th parenchyma of the medullary rays,
wood-prosenchyma with simple pits and sometimes septate, and scantily
developed wood-parenchyma; in the structure of the leaf-the tendency of
the stomatal apparatus to differentiate in accordance with the Rubiaceous
type, and the absence of glandular hairs. Oxalate of lime is excreted in the
leaf and axis in the form of clustered and solitary crystals. Internal secretory
organs are absent. The two genera of.~the Salicineae may be distinguished
anatomically by the place of origin of the cork, in Salix it is the epidermis
itself, in Populus. the outermost cell-layer of the primary cortex.
I hc:rve examined the structure of the leaf in Populus nigra, L. and Salix
alba. L. In both species the epidermis bears simple unicellular hairs, and
stomata are prerent on the two sides of the leaf. In Salix alba the stomata arE'
accompanied on either side by one or more subsidiary cells, placed parallel to
the pore; whilst in Populus nigra this type of, stoma is not always so well
mark~d.~_The ,epidermis jn many cases includes cells with mucilaginous
inner walls, thus according to Radlkofer in Salix acuminata, S. aJba, S. amdJa,
S. Caprea, .5'. cinerea, S. daphnoides, S. tragilis, S. grandi/alia, S. incana~ and
S. ni~cans, but not in S. amygdalina, S. ,ub,a, S. viminalis, Populus alba,
P. nsgra, and P. tremula; occasionally (5. alba according to Walhczek) both
the inner and outer wall of the same epidermal cell may be mucHaginous
According to Rothert's abstract of Dobrowlianskij's work, hypoderm occurs
beneath the upper epidermis in some members of the Order, and, in certain
species of SaZ,x belonging to the section Rugosae, the epidermal cells here
and there show septation into two or three cells; the latter statement mny
possibly be due to an incorrect interpretation of the mucilaginous epidermal
cells. According to Dobrowlianskij, the mesophyll varies in its structure.
Very commonly the lowest ceU-Iayer of this tissue contains little chioro-
 798 .,<"" I1LICINEAE·

phyll, and is differentiated like a hypoderm; accord.ing to my own observation

this is also the case in Populus nigra and Salix alba. In these two species the
remainder of the mesophyll consists. of palisade~tissue; in other species dense
or· loose spongy tissue is presen t. Glandular hairs are not present in this
Order~ In place of them the leaf·teeth· in certain species of Salix and
Populus have a vesicular glandular· structure, the glandular tissue being
formed by th~ epidermis, the cens of which are transformed into long, narrow,
radially arranged prisms I.
In Salix and Papulus three vascular bundles always pass into the petiole,
and in both genera these bundles show a tendency to.. become rolled up into
a circular form during their further course thr:ough the petiole. In the
characteristic region we meet with a ring of wood and bast. in Salix, and
several superposed rings -of vascular bundles in Populus.. For details see
C. de Candolle, ,Petit, and especially Komaroff. ~
.1 ha ve-: examined the structure of the wood in Populus tremula, L.,
P. nigra, L., Salix alba, L., and S. purpurea, L. The medullary rays. are
narrow, oeing from one to two cells in breadth. According to Schulz, the
medullary, rays of the poplars consist entirely of cells of equal height, whilst
those of the willows contain two kinds of cells, the one elongated in the
verti~, the other ill the radial direction. The vessels are scattered in the
transverse section or are arranged in radial rows, several of them fonning
a row; their diameter attains '066 mm. The perforations are simple. In·
contact with one another the vessels bear rather large bordered pits (diameter
of the polygonal borders = ,oo4-·oo6.mm.), but in contact wjth parenc,hyma
of the medullary rays thfre are simple pits of the size of the border.in the
bordered pits. Wood .. parenchyma is. scantily deve]o~d. The wood ..
prosenchyma, which has fairly thick walls and rather wtde lumina, bears
simple pits:. and is sometimes (S. alba) septate.
The following statements regarding the structure of the cortex are chiefly
based on 'Moller's work. According tl) Sanio, Moller, J. E. Weiss, and others.
the cork arises in the epidermis in the numerous investigated species of Salix,
but in the species of Populus it always develop~ in the sub·epidennal layer of
·cells. In Populus the cork-cambium first gives rise to several rows of cubical
cel1s with wide lumina, and flattened cork-cells are not formed until later; in
SaUx a thin stratum of cork _is. formed each year consisting of a few layers of
cells, which have their outer tangential walls sclero~ed. The outer fortion ~f
the primary cortex is coIl en chymatous, and in certain species 0 . Populus c

includes stone-cells, which sometimes (P. alba, L.) unite to form a ring.
Isolated groups of bast-fibres are developed in the pericycle, Groups of hard
bast are always present in the secondary bast, and they are often so arranged
as to ~usestratification ofthephloem; in some cases (P. alba, L., P.fryramidalis,
Roz.) groups of stone~celJs occur a_s well. Oxalate of lime is found in the bast
in tl1f1 form of solitary and Clustered crystals. th~ for~et being contained in
chemoored crystal.fibres. which sheathe the secondary bundles of hard bast;
they 'ilI"e enclosed in the cell-wall. The medullary rays of the phloem
tiavelsa·tbe second~.ry hard bast without becoming sclerosed.
The pith is heterogeneous according to Mentovich, the periphery being
formed by active cells, the centr3.1 portion. by empty cells. In thos~ portions
of the stem of Populus alba and Salix amygdatina. In which the formation of
heart"wood h~s already set" in~ the pit~.contains deposits of carbonate of lime
(l\Iol~ch).

'l Th( buds of the poplar; which arc covered with abundant ba~sam, exhibit a limilar stntdure j
thl: iIlternal bud.&eales have glandular surfaces, conai&ting o( epidermal cells tlongated like palisade,
~nd coyered by a thin cuticle (see Hanstem, lac. cit., and Areichoug, Acta Land. IS70, ta.b. iv, 4-0)'
 SAL 1CINJ:.AE

Literature: Chalon, Tiges lign. dico~yl., ~ Mem., 1868, pp. 49-s3.-Haosteill, Harz- n.
Schleitnabs., Bot. Zeit. 186R, p. iS6.-Radlkofer, Monogr. Serjania, 1875, p. 103.- Moller, Holzanat.,
Denkschr. Wiener Akad. 1876, pp. 33-4 and 3:Z9--Reinke, SeKtetit;msorg., Pringsheim Jahrb., Bel. x,
1876, p. 168.-Areschong, BIad. anat., Minnesskr. Lund 1878, p. ~7 et seq.-De Bary, Vergl. Anat •.
18i7.-C. de Candolle, in Mem. Soc. phys. et hist. nat. de Geneve 1879, p. 446 et seq.-Molisch;
Kohlens. Kalk, Sitz.-'Ber. Wiener Akad., Bd. lxxxiv, Abt. I, 1881, p. 7 et seq.-Moller, Rindenanat.,
1882, pp. 89-95.-Sthu!z, Markstrahlgew., Diss., Berlin, 1882, pp. 18-19; als~ Jahrb. Berliner bot.
Gart., Bd. ii. -Mentovicb, Mark, Klausenhurg, 1885, Hungarian; abstr. Just 188.5, i. p. ,87.-
£Qlue<ier, Holtstr., 1885, p. ~S9.-Petit, Petiole, Mem. Soc. sc. phys. et" nat. de Bordeaux, ser. 3,
t. iii, 1887, F. 246 aod pI. ii-iii.-Dobrowlia.'l5kij, Verg1. Anat. d. Bl. d. S., Arb. St. Petersb. natun.
Gesellsch., Bd. xix, 1888 (Russian) pp. J6r-70 ; abstt. by Rothert, in Bot. Centralbl. 1889. ii, p. 487.
-Douliot, Periderme, Ann, sc. nat., ser. 7, t. x, 1889, pp. 330-1.- J. E. Weiss, Korkbild., Denkschr.
Regensburg. bot. Gesellsch. 1'890, sep. copy, p. 53.-ROSSt Peri derma. Malpighia, vol. iv, 1890, p. 104.
-Strasburger, Leitungsbahnen, z891, pp. 207-13.-Houlbert, Bois sec. dans les A~tales, Tb.ese,
Paris, 1893, pp~ 130-2.- Walliczek, Membranscbleime, Pringsheim Jahtb., Bd. xxv" 1893, p. 238.-
Pax, in Natiirl. Ptlanzenfam., iii. Teil, Abt. 1 (1894), pp. 30-J.-Lazniewski, Bioi. d. Alpenpfl .•
Diss., Munchen, 1896, p. 40 et seq. ; sep. copy from Flora 1896.-Virchow, Blattziibne, Arch. d.
Phann. 1896, sep. copy, PP' 51 and 6I.-Komaroff, Struct. fol., Bull. Herb. Boissier l897, pp. '~26-
46.-Kubla, Phelloderm, Dot. Centralbl. 1897. Hi, p. II6.-[Theorin, Om bladt. glandlema hos en
del Salices, Stockholm, 188:a.-·So~taric, Bau d. Stam::nes d. S., absfr. in Oest. bot. Zeitschr. 1899,
P.·1I7 (a.ppears in Sitz.-Ber. Wiener Akad.).] .

LACISTEMACEAE. "
This small Order, which consists of the singl~' g~nus Lacistema, is of
uncertain systematic position, having been.regarded by s~tematists as allied
to the Piperaceae, Chloranthaceae, Urticaceae, and Samydaceae; it has neither
the resin-cells of the Piperaceae and Chloranthaceae, nor the secretory cavities
of the Samydaceae, nor the -cystoliths of the Urticaceae. Its anatomical
characteristics are; vessels with rather small lumina, and scalarifonn perfora-
tions; narrow medullary rays in the wood; wood-prosenchyma with few but
distinctly bordered pits; a composite and continuous ring of sclerenchyma in
the pcricycle; superficial development of cork; and the absence of internal
and external glands. Oxalate of lime is excreted in the form of clustered and
solitary crystals. The trichomes are simple. Tannin is present in a~undance
in all parts of the plant. /
I have examined the structure of the leaf in Lacistema intermedium,
Schnizl., and L. pubescens, Mart. The leaves in these two species have typical
bifacial structure, the spongy tissue being very lacunar. The stomata are
restricted to the lower side of the leaf, and are surrounded by three or more
epidermal cells, which in some cases show a certain degree of differentiation
as subsidiary cells. The vascular bundles of th~ larger veins are accompanied
by sclerenchyma. Clustered crystals are found in the mesophyll, and in the
veins. The hairs are simple; they have rather thick walls, and are unicellular,
or septate owing to the presence of delicate transverse walls; the basal
portion of the hair is provided with father large pits, elongated in the direction
of the length of the hair.
I have. examined-. the structure of the 'axis in L. pubescens, Mart., and
L. myricoides, Sw. The following statements may be added to the dataglven
above on this subject. The medullary ra:'s of the wood are from one to two
cells in breadth, and are chiefly composed of cells, which are elongated in the
vertical direction. The vessels are quadrangular in transverse section, varying
numbers of them being arranged in radial rows. The bordered pits on the
walls of the vessels are small (diameter of border -== ·002·-'003 mm.), and are
also found on the walls in contact with wood-parenchyma, or parenchyma of
the medullary rays. The scalariform perforations have a varying number of
bars; in L. myricoide's, where the bars are rather broad, a..c; many as twenty
may be present. Wood-parenchyma is somewhat abundant. A few stone-
cells sometimes occur·in the primary cortex. The cork in L. pubescens arises
800 LA CIS TEMA CEA E

sub-epidermally, and includes cells having their wall? more strongly thickened
on one side (viz. the inner \.angeJ1tial and the contiguous portions of the
radial walls). Oxalate of lime is found in the medullary rays of the wood in
the -form of ordinary solitary crysWs.
Literatnre: Bokorny, DUTCh•. P., Flora, 1883, p. 371, aep. copy, p. 3s.-So1ereder, Holzstr.,
188S, pp. 359-6o.-Engler, in Natilrl. Pflanzenfam., iii. Teil, Abt. I (1894), p. 15.

EMPETRACEAE.
This small Order, the members of which are distinguished by ericoid
.habit and by possessing rolled leaves, shows great uniformity in the anatomical
structure of the vegetative organs. In illustration of this statement the
following characters may:be pointed out: the apparently bifacial structure
of the leaf; the mucilaginous epidermis of the leaf; the absence of a special
type of stoma; the very small lumina of the vessels; the tendency to form
scalariform perforations in the vessels; the narrow medullary rays of the
wood; wood-prosenchyma with bordered pits'; the scanty development or

FIG. 188. Transverse section th ..ough the leaf of Emj>eJru11t niitrum, L.-OriJil'inaL

complete absence of sclerenchyma in the cortex; the origin of the cork in the
pericycle ; the excretion of oxalate of lime in the form of clustered crystals
or sphaerites; and the hairy covering, which is formed by simple unkellular
trichomes, and short glandular" hairs having a stalk composed of one or a few
cells, and an ellip~oidal, uni- or multicellular head.
The structure of the leaf has been examined in all the four species of the'
Order (Empetrum nigrum, L. Fig. r88, Corema album, Don, C. Conradii, Torr.,
and eeTat'Ola ericoides, Michx.). T!,1e leaves of all the species belong to the
category of rolled leaves, being deeply furrowed on the lower side. This
furro'\l1 does not, as would appear at first sight, originate by the bending back
of the mlll'gins of the leaf, but according to Gibelli and Gruber it is due to
more rapid growth of the portions of the lower side of the leaf situated on
either side of the midrib; in consequence of this, a groove first arises, but
this subsequently becomes closed owing to the lateral extension of the'more
rapidly growing portions, and thus forms a central cavity. The epidermis,
which limits the rolled leaf externally, is distinguished by having a strongly
thickened outer wall, by the absence of stomata, and by the gelatinization
of the inner cell-walls. ThE' gelatinization has been correctly interpreted by
Gruber and others only in the case of Empetrum 1, but according to my own
1 The gelatinization is found both in the lowland, and in arctic and :llpine specimens (see
Wagner, Sitz.-Her. Wiener AJ,ad., Bd. ci, Abt. I, 1892; Warming, Om Groc:nlands Vegetation,
1888; and Boergesen, Journal de Botaniqnc, 1895, p. 3).
 EMPETRACEAE 801
investigations it also occurs in the other three species, in which some of the
mucilaginous inner walls (Corema album and Ceratiola) have a conical form
and penetrate the leaf-tissue, while in some cases they also exhibit a number of
unaltered cellulose-lamellae (e. g. in Corema Conradii). The epidermis lining
the central cavity of the rolled leaf has thinner outer walls, and includes the
stomata, the latter, according to Gibelli, being surrounded by several ordinary
epidermal cells in Empetrum. The inner epidermis is also furnished with
trichomes, which consist of glandular hairs and simple unicellular hairs. The
former have a short unicellular or uniseriate stalk (in the latter case composed
of a few cells), and an ellipsoidal head, which is unicellular, or divided into
several cells, the diVIsion-walls being chiefly horizontal, one or two vertical
walls being also present. The simple, unicellular hairs are so abundant in
Corema album and Ceratiola that they form a tangled mass filling the entire
cavity of the rolled leaf. In Empetrum and Corema Conradii they are chiefly
present at the margin of the leaf, and those on the two margins become
interlocked, thus.firmly closing the aperture of the central cavity. Beneath
the outer epidermis of the rolled leaf there is more or less typical palisade-
parenchyma, whilst spongy tissue is situated above the epidermis, lining the
central hollow, so that, if the course of development be disregarded, the leaf
may be described as bifacial. There are no sclerenchymatous elements
accompanying the vascular bundles of the veins. Internal secretory elements
are likewise absent. Oxalate of lime is found in the leaf of Empetrum in the
form of clustered crystals, and of beautiful spbaerites with radial striation and
concentric stratification; in the leaf-tissue of Corema album oxalate of lime
only occurs in the form of clustered crystals.
I have examined the axis in Empetrum, Corema album, and Ce1atiola
ericoides. In transverse section the wood (and this also applies to Corema
Conradii) shows scaUerer! vessels with small lumina (maximum diameter
= '024 - '03 mm.), and narrow I -2-seriate medullary rays, the cells of which
are more or less elongated in the vertical direction. The perforations of the
vessels c.re scalariform (with as many as ten bars) in Empetrum, whilst in
Corema and Ceratiola they are, for the most part, simple and elliptical, more
rarely scalariform. In contact with parenchyma of the medullary rays the
walls of the vessels bear slit-shaped bordered or unbordered (?) pits. Wood-
p2.renchyma is scantily devebped; the wood - prosenchyma has distinct
bordered pits. The following points are noteworthy in the cortex, etc. The
epidermis, like the outer epidermis of tl).e rolled leaf, has a very thick external
wall. Sclerenchymatous fibres are rarely met with at the outer limit of the
bast, being isolated in Empetrum according to GibeIli, and forming bundles at
some points· in Ceratiola. The first layer of periderm arises in the pericycIe.
In Corema album the cells of the cork are strongly thickened on one side (viz.
on the inner, tangential wall). Sclerenchymatous elements are entirely absent
in the secondary bast. In Ceratiola the subepidermal cell-layer of the cortex
has its inner tangential walls sclerosed, so that the cells appear thickened in
the form, of a ~horsesh6e in transverse sections of the branch.
"iii,
literature": Gibelli, Strntt. delle fogJie delle E., Naov. Giom. bot. Ital., vol. 18 76, pp. 49-
60 and tav. v-vi.-Graber, Anat. 11. Entw. d. Bl. von E11lpetrum. Diss., Konigsberg, 1881, 38 pp.-
Solercder, Holzstr., 1882, p. 260.-Pax, in Natlirl. Pflanzenfam., iii. Teil, Abt. 5, 1891, P. 124.-
[Mac Ewau, Compo anat. of Cortma, etc., Bull. Torrey bot. Club, vol. xxi, 1894, pp. 277-85.)

CERATOPHYLLEAE.
There is litt]c to be said regarding the anatomy of this interesting Order,
which includes the single genus CeratophyUum with about three species; it is
well known that the members of this Order are submerged plants, having no
SOLERl:DER 3 F
 CERA TOPHYLLEAE

land.forms, producing no roots, and flowering and fruiting beneat4 the

surface of the water. Fe::ttures of special note are the complete absen(..'C of
yessels and of stomata.
The axis, which, like the leaf, has been specially examined in C. deme,.
stem, L. by H. Schenck, has an epidermis con taining chlorophyll, and devoid
of stomata. Beneath the epidermis is the primary cortex, in which the outer-
most cell·layers are collenchymatous, while the inner portion contains a ring
of air·spaces separated from one another by radial plates, one layer of cells in
thIckness. Numerous tannin-cells with suberized walls are present throughout
the primary cort x. The vascular system of the stem is delimited from the
primary cortex by a typical endoderm is containing starch, and provided with
Caspary's dots. The centre of the vascular strand is occupied by an axile air-
canal, arising by the resorption of a small group of elements, but according to
Caspary, Sanio, and Schleiden this group does not consist of tracheae _~as is

FIG. I Rq, Transverse st'ction t h rou,h the fibro-vlU(':ular Iyst~m of CeY(J/opIJyllt41" tle",ersu",. L. -after
H. Schenck.

the case in Aldrovanda. for example), but of procambial cells with narrow
lumina and delicate walls. The axile canal -is surrounded by ail annular zone
of coil enchymatous, parenchymatous cells, which contain starch and are
elongated in the vertical direction; as shown by H. Schenck, they are to he'
regarded as xylem (wood·parenchyma), though they include no vessels.
From the xylem·portion a ring of phloem extends as far as the endo·
dermis; this phloem consists of bast-parenchyma and sieve-tubes; the latter
have wide lumina, are provided with companion cells, and are arranged in two
rings in the transverse section. Vessels and lignification are thus entirely
absent in the stem of Ceratophyllum.
According to H. Schenck, the segments of the leaf in Ceratophyllum
demersunz are elliptical or circular in transverse section, and are traversed by
a vascular bundle, which runs nearer to the upper side, contains no vessels,
and is surrotL"lded by a parenchyrp.atous sheath. The epidermis of the leaf is
devoid of stomata, and consists of low cells, which are polygonal in surface-view,
and contain chlorophyll. The latter is less abung.arit in the parenchyma vf
the leaf than in the epidermis. The mesophyll is not particularly strongly
developed, and is traversed longitudinally by air·canals; of these three on
 CERATOPHYLLEAE
the lower side are large, and are separated from one a.nother by septa of
a single layer of cells, while on the upper side there are two much smaller
canals. The parenchyma of the leaf, like that of the cortex, contains tannin-
cells. The apices of the segments of the leaf in CeratophyUum, as in those
of MyriopltyUum., are occupied by a long, shaggy, multiseriate glandular
structure, the oily contents of which are rose-coloured in young stages and
subsequently become rich in tannin.; On either side of each of these gland~
there is a long, pointed unicellulaI\ hair. I t has not yet been determined
whether these peculiar glands, whi~h !)ubsequentJy disappear, serve for the
protection of the young leaf-tips against parasites.
Regarding the relation of the axile strand in the stem to the leaf-trace
strands, we may state on the authority of De Bary that~Ceratoph'YUum is
one of those plants in which the strands found in the stem are cauline, and
have apical growth, whilst at the nodes branches pass off from the cauline
strands into the leaves.
Literature: Schleiden, Kenntnis der C., Linnaea 1837. pp. 53~2.-Caspary, in Monatlber.
Berliner Akad., 1862. p. 466.-Sanio, Endog. Gefassbilndelbitd., Bot. Zeit. 186~ p. H3.-Botodin.
Blattsp. einig. WasserptJ., Bot. Zeit. 18io, p. 8so.-Magnus, Bemerk. z. Aues. v. Boro<iin, Bot. Zeit.
1871, p. 481: bere cited: Mercklin, Blattgest., Jena, 18.,.6, pp. ,.~-, and Tab. \; see also Hegelm.aier,
in Bot. Zeit. r871, p. SOI.-De BarYt Vergl. Anat. 187i.-Areschoug, Blad. anat., Minnesskr.,
Lund. 1878, p. 1« et seq.-Klercker, Anat. et devel. de CeralojJltyllum, Bihang K .. Svenska VeL-
Akad. Handlingat, vol. ix, n. 10, 1885, 23 pp. and 3 Tab.: see also Bot. Centralbl. 1885. i. p. 157
et scq.-H. &:henck, Vergl. Anat. d. submers. Gew., Kusel, 1886, pp. IO-JI and 38, Tab. i aDd vi.
-Duta.iUy, Glandes fol. G.es 'C,rat(lp4yllum, etc., Assoc. {fan,;. p. l'&vanccm. des K., Pads, l~l.
p. 32o.-Engler, in Natflrl: P6anzenfam., iii. Teil, Abt. 2 (189+), p. fl.-Schilling, Schieilllbiid. d.
Was$erpft., Flora 189.., p. 3 2 7.
 ADDENDA
RANUNCULACEAE (pp. 14-20).
I. T~E is little to add to.the -REVIEW OF ANATOMICAL FEArtrRES glV"en
ii! the earlier part of this book. Clustered crystals of oxalate of lime have..
also been observed in the petiole in species of ThalictlUtn, and in tbf leaf-
sheath in Jtnemone iaponica, S. et Z.; arm-palisade-cells have also bee.Q. re-
conkd in s~ecie$ of Adonis, Clematis, Delphinium and Nigella. The structure
of the. stem has been shown to be astelic in species of Anemone, Caltha, E1'ant"is~
OXYlUllPhis and Ranunculus (incl. Ficaria) .
.~~ STRUCTURE OF THE LEAF. Numerous details regarding tke natw-e
of the epidermis and the structure of the leaf are to be found in Goffart"s
work I. . The stomata are confined to the lower side of the leaf in Ac(aea,
Anemone pro parte, Aquilegia, Delphinium and Trollius, but OC(:ur on'· bot~
sides in 4 ~.is, A nemone pro parte, Caltha, Celatocej>halus, I sO/>'Y'lfm, M yosut'us
and NigellJl; There is no arm-palisade-parenchyrna (see also above) in Hepatica
and Puhatl11a, while in Aconitum columbianum, according to Schwartz:-Clemen~,
it is o;nly prESent in the shade·leaV'es. The clustered crystals ot oxalate ef lime,
fqtftd In Anemone and Thalict1um, have already been referred to above.
Sphaerocrystalline masses of unknown chemical composition have been o&served.
in Rtmunculus aco7titifolius, L., R. platanifolius, L. and R. ",,00m, Crtz.;
according to Schaffnit, similar deposits are present in the floral organs in species
of Ceratocephalus and Thalict,um. The 'small club·shaped (see Fig. I, J) or
conical (see Fig. I, G) trichomes have recently been· shown to occur also in
Species of Adonis, Anenwne, Aquilegia, Delphinium, Isopyrum, Nigella and
Trollius, whilst longer tubular hairs with a basal ventricose swelling have been
recorded for species of Delphinium and HellebOfus. For information as to
the number and course of the vascular bundles-in the petiole, see especially
Goffart and Ziegler ;' in Paeonia the bundles are less isolated than ih the other
genera.
. 3. STRUCTURE OF THE AxIS. According to Ziegler, anomalous structure
of the stem is also found: (a) in Delphinium Requieni, in which numerous
medullary vascular bundl~ are present in the lower part of the stem; and
(b) in Paeonia o{ficinalis, in which a single cortical concentric vascUlar b~ .
WIth central phloem is found in certain parts. According, to the same author;
TlwIlictrunt P'Y"hoc~!pum. d_iffers from other species of its genus in having ·no
medullary bundles.
NOTE.-Accordin~ to Stapf, an anomalous structure of the tuberous roots of
AtiOJIilum is characteristic of fhe Indian ~iest belonging to the sections Antiot'''
(A. rotunditolium, Kar. et Kir., A. heteroph-yllum, Wall, A. navicula,.-e, Stapf. A. pal~
tMlum, Hook.. t, and probably also A. lloolleri, Stapf) and Dtino"hilutn (A .. dlituW-
,.~izum. Stapf and A. Bal/OUfis, Stapf). and, accordine to If1l).isch, Goris and A. Meyer,
is also found in A. Anthcwa. ' In these species the parenchymatqus ground~tissue
of the central cylinder includes several·vascular strand". for the most part cori.~

I In this treatise the following genera z::.dea.lt with t -tI(lmt"ttU1l, AcllUa, Al/Mis, Ane""N,
1J".~' CtlitAa, Cb-tII«el halus, Cimi(~";I"
i Delpl'inlw",- Ertlnlhis, Rel/tHrus, IsD./J7rJilt'l,
"",."s,lli.erU", Ph TJd., PanunNl/u.ft "",lietrum., 'frW/ius. .
806 ADDENDA--RANUNCULACEAE
centric in structure, and each having a small pith of its own, and growing in thickness
by means of a partial cambium; these strands are arranged in a circle in the trans·
verse section, the outline of each strand being annular or stellate in the Anthora-
type, but tangentially elongated (band·shaped) in the Dein01'1'hizum-type. Both
on the outer and inner sides of this vascular system there are numerous phloem-
bundles situated in the parenchymatous ground-tissue, and right at the centre there
is a small indistinct pith. The mode of deVelopment of this anomaly bas been
determined by A. Meyer in A. heterophyllum. In the pith of the tuberous root,
that is to say on the inner side of the normal vascular system, which grows in thick-
fleaS by means of a normal cambium, a secondary meristem of circular outline in
transverse section is developed; this meristem produces parenchyma and phloem-
.bundles on its inner side, and parenchytna, sometimes together with tracheae, on its
outer side. This secondary cambial ring next becomes united at one 'point with the
external (normal) cambium, and consequently in a transverse section it appears
as though only one (the outer normal) cambium were present, though strongly-
jnvaginated at one point into the pith. In the further course of development
the two cambia, i. e. the secondary meristem and the normal ~bium, fuse at
other points also, and in this way the concentric bundles mentioned above originate.
each being surrounded by a partial cambium. It will be well to. mention that
according to Hartwich and Neuber the anomalous structure just described may
sometimes also be met with in A. Napellus; in the tuberous roots of this species,
moreover, other anomalies occur locally. but for these the reader must be referred
to the literature, especially the wflrks of A. Meyer, Ha,rtwich. and Neuber. Re-
garding-.the retiform rhizomes and roots of A. Lycoctonum see ibid. and Jost, loc. cit.
For the structure of the root in the Ranunculaceae generally see also Lenfant,
Mansion, Sterckx, n. ce. The bulbils in the axils of the leaves of Ranunculus Ficaf'ia
are, we may add,. formed mainly by a negatively geotropic root-structure, as s)lown
by their anatomy.
Literature: Irmisch, AconitulIJ .Noptlilts, Zeitschr. d. ges. Naturwiss., IS.54, pp. 181~3 aod
Tab. iii-v.-Wasowicz, Acon;tu", luterophyl/um, Archivd. Pharm., 1879. i, p. :117 et seq.-A. Meyer,
A(lm;/u1n Napellus, etc., Archiv d. Pbarm., 18S1, ii. pp. 240-j6.-Langgaard, Japan. d. chin.
Aconitknollen, Archiv d. Pharm., 1881. i. p. 161 et seq.-Costantin, Tiges aero et sout., AI •. sc.
,.ix.
nat., ser. 6, t. xvi, 1883, p. 88 et seq.-Costantin, Tiges d. pl. aqu., Ann. SC. nat., ser. 6, t. 1884.
p. :187 et seq. and pI. 16.-Jost, Zerkliiftung einiger Rhiz. etc., Bot. Zeit•• 1890, p. 485 et ~q. and
Tab. vi.-fMiczinsld, Anat. des Anemones hybrides, Compt. rend. Acad. Cracovie, xxiv, 1891,
pp. 105-36 and pl. ii-iil.]-[Togoini. Stomi, Atti ]st. bot. Pavia, 1894 j abstr. in Just, 189-4.
I, p. 466. and Bot. Centralbl. Beib., ]S94-, p. 4:13.]-[Eastin, Struct. of Cimicifuga, Americ.
Journ. Phanp. 1895, pp. 121-8. ]-Borgesen, Arkt. pI. bladbygn., Bot. Tidsskrift, 1895. p. 219 et se<l.
-(Nihonl, Et ...nat. des Ranunc., Mem. cour. et memo d. savants etrang. Acad. d. Belgique, 1895.)-
Ziegler. Gefassbundel im Stengel der Ranunc., Diss., Erlangen, J 895. 41 pp.-MacDougal, /!o/,yrtt",
«riden/ale. Bot. Gaz. xxi, 1896, pp. 28O-3.-Janczewski, Ane11ltJm, Revue de bot. 1897, p. 337 et
seq. :lnd pl. 18-19 (racine). and lac. cit. 1898, pp. 433 and 507 et seq. and pI. 16-19 (tige).--
LenEant, Delphinium. Arch. de l'Institut bot. de rUniv. de Liege, i, 1897.70 pp., pl i-xi.-Mansion,
TluJlictrum jlavum, ibid., 1°4 pp., pI. i-xiv.-Sterckx, Clematidees, ibid., 88 pp., pI. :or.;v-.xxix.-
Spanjer, Wasserapparate, Bot. Zeit. 189S, i. p. 'i4.-Weinrowsky, ScheitelOffn. bei Wasserptl., Diss.,
BerliD, ]898, p. 32.-Minden, Wassersec. Org.... Bibl. bot., Heft 46, 1899. pp. :13 and p.-[C611in,
HyJrastis canatimsis, Jouro. de Pharm. et de Chimie 1900, p; 309; abstr. in JUS1.I90o, ii, p.·16.1-
Kliem, Veget. RegeneratioDSorg., Diss., Erlangen, J9OO. p. ,p.-[Rer.voll, Nogle arkt. rAIlunklers
morfologi og anatomi. Nyt. Magaz. for Naturvidenskab.. 1900, 25 pp., pt. xii-xiv.]-Schldchert,
Xeropbyten der Muschelkalkh. bei Jena.. Naturwiss. Wochenschr. 1900, p .•449 (Anenume).-Thomas,
Feuilles sout., These, Paris, 19oo.-Goffart, Anat. de_s feui1les dans les Rannnc., Arch. de Plnstitut
bot. de l'Univ. de Liege, iii, 19°1, H~O pp. and 14 pL, especially p. 150 et seq.-[Goris, Accm·t.m,
Bull. Sc. pharmacol. 19°1. no.... ; abstr. in Just, 1901. ii, p. 37.]-Petersen, Vedanatomi, 1901, p. 41.
-Bouygues, Petiole, These, Paris, 190:1, p. to.-Mardner, Phan.-Vegetat. der Kerguelen, Diss.,
Ba4el. 190 3 , pp. 28-31 (Ranu1U"tllus).-Knothe. Unbenetzbare Blatter. Din•• Heidelberg, 1902, p. 16.
-[Tschircb, HtllelHwus-Drogen. Schweizer. Wochenschr. f. Pharm. 1903. no. 35; abstr. in lust,
J902. ii. p. 68.)-[Vanderlinden. Alcal. et glycosides dans la famille des RanUDc., Recueil de 1 Jost.
bot. de Bruxelles, v, 190:1, pp. )~5-i8; abstr. in Bot. Centralbl., xcii. p. ]67.]-[Finlayson, Stem-
structure of some leafless plants, Transact. and Proc. New Zealand Inst. 1903, p. 360et seq. (Clemali"s
afolJ'at.).l-Scboote,,stelartheorie, Jena and Groningen, 1903, pp. IU-I.5.-Theorin, Vaxttrichom.,
ArkivfOr Bot. i, 1903, p. 171.-[Futo, Hlpatiu~t,.anssilvani('(I, Diss., Klansenburg, 19°4, Hungarian.]
-Freidenfeldt, Anat. Bau der Wurzel, Bibl. bot., Heft 61, 1904, pp....S-54.-Neuber, VergI. Anat.
d. Wuncln. Diss.. Bern, 1904, pp. 5-53 (Ac~tiJum, Belle6orus).-Schaffnit. Nekt. der Ranunc.,
Diss., Erlangen, 190....-Siissenguth. Behaarongsvetb. der Wtirzburger Muschelkalkpfl., Dis50., Wiirz-
burg, 1 go.... pp. 17-18.-Theorin, Vaxttricbow., Arldv for Bot., iii, no. 5. 1904, p. 17.-[Clark
 ADDENDA--RA NUNCULA CEAE
CotYledon- and leaf-structure in certain Ranunc., Torreya, 19°5, pp. 16..-6.]-Sarton, Recb. e:l~r.
sur l'anat. des pl. affines. Ann. sc. nat., ser. 9, t. ii, '1905, pp. 1-35 (Anmume, Ranunculus, T/uz·
lictrum).-Scbwartz-Clements, Relat. of leaf.structure to pbys. factors, Trans. Amenc. Microsco_p.
Soc. 1905, p. 59, pI. vii, 3--[Senft, Medizin. verw. PfL aUi der Fam. d. Ranunc.. Pharm. Pruu.
1905, DO. 1 et seq. ; abstr. in Bot. Centralb1., cH, p. 268.]-Stapf, Aconites of IndiA, Annals Bot.
Gard. Calcutta, x, 2, 1905, pp. 12~-30.-Geneau de Lamarm~re, Membr. cut. d. pI. aqu., Revue de
bot. 19Q6, p. 289 et seq.-Piccioli, Legnami,. ~ull. Sien~ 1906, p. 132. [For additional literature
see p. II71.]

DILLENIACEAE (pp. 20-24).

2. STRUCTURE OF TIlE LEAF. In certain species of lVormia a hypoderm
composed of large cells is found beneath the upper ~pide.rmis, while in Dillem:a
indica, L. there is a single layer of hypoderm.
3. STRUCTURE OF THE AXIS. The wood in W ormia /erruginea, Baill.
consists of : (a) broad medullary rays reaching a breadth of ten cells; (b) vessels
with exclusively scalariform perforations (with as many as forty bars), and
bearing bordered pits in contact with wood-parenchyma;' and (c) wood-prosen-
chyma having. bordered pits (Ursprung). A composite and continuous ring
of sclerenchyma in the pericycle is also present in the young axis· of Dillenia
ochreata (= W ormia ochreata) and Doliocarpus semidentatus (Pitard).
Literature: Ursprung, Anat. u. Jahresringbild. trop. IIoizarten, Diss., Basel, 1900, pp. 10-13
(Wor1lliafirrug":nea).-Pitard, Pericycle, These, Bordeaux, 1901, p. Sa.-Fabricius, Laubbrattanat.,
etc., Beih. Bot. Centralbl., xii, 1902, pp. 310, 3II.-Bargagli.Petrucci, Legnami, Malpighia, 1902,
p. 333 (Dillenia).-Areschoug, Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handling. 39, n. 2, 1905,
pp. H-15 and Tab. xxiv-v (Dillenia), and pp. 105-6 (Tetracera).-[Moll and Janssonius, Mikro-
graphie d. Holzes, Leiden, 1906, pp. 65-80 (/Formia, Dillmia).]

CALYCANTHACEAE (pp. 25-27).

1. The statement in the REVIEW OF ANATOMICAL FEATURES regarding
the absence of a sclerenchymatous ring in the pericycle requires some modifica-
tion, since according to Van Tieghem (see also Pitard, loco cit.) the lower inter·
nodes in Chimonanthtts contain a composite and continuous sclerenchymatous
ring composed of groups of bast·fibres and stone-cells with U-shaped l thicken-
ings; in Calycanthus, on the other hand, there are invariably isolated groups
of bast-fibres. The stone-cells with U-shaped thickenings afford an important
diagnostic character of the Order, since they also occur in Calycanthu,s.
3. STRUCTURE OF THE AXIS. According to Van Tieghem the anomalous
structure of the axis (occurrence of four cortical vascular bundles with inversely
orientated wood and bast) is not the same in the two. genera belonging to the
Order. In Chimonanthus the normal vascular ring (stele) has a quadrangular
outline, and the primary cortex is normal; the cortical vascular bundles lie
in the parenchymatous per~cycle on the inner side of strongly developed groups
of pericyclic fibres; the cortical 'bundles' actually consist of two strands.
In Calycanthus, on the other hand, the stele is cylindrical, but the primary
cortex has thickened angles; the vascular bundles, which here consist only of
a single strand, lie in the inner portion of the primary cortex.
Literature: Lloyd ~~illiamsJ Sieve tubes of Cal)'<mll/zus occidmtalis, Ann. of Bot., ,·iii, 189+,
pp. 367-7o.-Biennann. Olzellen. Diss.• Bern. 1898. pp. 58, 59.-Pitard, Pericycle. These, Bordeaux,
~901, p. 8a.-Van Tieghem, Tige des Calycanth.• Ann. sc. nat., SeT. 8, t; xix, 1904. pp. 305-jO;
also Bull. Mus. d'hist. nat. 1904, p. 68 'et seq . -Van Tieghem, Meristetes corticales, Ann. SC. nat.,
ser'9, t. i, 1905. p. 40.
MAGNOLIACEAE (pp. 27-31).
I. In the REVIEW OF A~ATOMICAL FEATURES the following additions or
modifications are necessary. The clothing hairs are mostly multicellular and
uniseriate; in some cases (species of JJagnolia and Miche.lia) they are uni-
 A DDENDA-MAGNOLIAGEAE
cellular. Tufted hairs with 1-2·celled rays are. found in Magnolia /asciculizta~
Parm. The statement as to the absence of a sclerenchymatous ring in the
pericycle is incorrect, for not uncommonly there is a' continuous or interrupted
composite ring of sclerenchyma in place of the isolated groups of bast-fibres.
The genus Zygogynum agrees with Drimys in having no vessels, and in the
structure of the wood reseml1lipg that of the Coniferae.
2. STRUCTURE OF THE LEAF. Papillose differ~ntiation of the lower epi-.
dermis is also found in Magnolia le"u~inea, Parm. and in Drimys retorta, Miers,
but not in D. granatetfsis, Mut., as was JOcorrectly stated on p. 28. A continuous
one-layered hypoderm on the upper side of the leaf has also been demonstrated
in Magnolia CampbeUi, Hook. f. et Thoms., M. /asciculata, Parm., M: ovata.
Parm., M. xerophila, Parm., Talauma inflata, Parm., Michelia calcuttensis,
Parm., M. Kisopa, Buch.-Ham., M. montana, B1., M. ovalitolia, Wight, PrIan-
glietia insignis, B1., M. pilosa, Parm., Drimys brasiliensis, Miers and D. grana-
tens is, Mu t.
Secretory cells are also present in the mesophyll and in the cortex in
Zygogynum. According to Parmentier, the secretory organs, described as
mucilaginous cells by Blenk, (Lacunes a gomme' by Vesque, resiniferons intet.:.
cellular spaces by D'Ippolito, ~d c Cana~x gummiferes by Parme!ltier, are I

found qwte generally In the· Schizandreae In the bast of the larger vems of the
leaf.
Branched sc1erenchyma-cells also occur in the mesophyll, though this ~
exceptional, e.g. in the spongy tissue in Drimys rdorta, Miers. In certain
SpeCies of Zygogynum stone-cells have been observed in the mesophyU, whilst
in certain species of nrimys and Zygogyhum there are zones of more strongly
thickened and lignified mesophyll·cells. .
Special mention should also be made of the occurrence of clustered crystals
in the epidermal cells in a new species named Magnolia echinina by Parmentier.
3. STRUCTURE OF THE AXIS. With reference to the structure of the pith,
we may add that incomplete diaphragms composed of stone-cells occur in
Drimys HOfIJeana, F. v. Miill., and that stone-cells, isolated or in groups, are
not uncommonly found in the Wintereae.
Wood resembling that of the Conifers in structure has recently been
demonstrated by Van Tieghem in almost all the known species of Drimys I
(with the single exception of D. uniflora, Turcz, which he was unable to
examine), and by me and Van Tieghem in Zygogynum, A certain number
of the species are further characterized by the presence of scalariform bordered
P~ts on t.he tracheids ~f the spring~wood: The excepti~ns quoted by PalElentier,
VIZ. Dnmys Mallen and D. vasculans, both of whIch possess vess& itt. the
wood, are certainly not members of the genus Drimys.
In the development of the cork the phellogen, according to Parmentier"
arises quite generally in the subepidermal layer of cells in species of Magnolia,
TlIlauma, Michelia, Manglietia, Liriodendron, Illicium, Drimys and Schizandl'a.
According to Van Tieghem and Douliot, however, the cork in Drimys develop~
in the epidermis (and not subepidermally). The pericycle contains eitherj

I It tnay be tnentioned here that owing to the peculiar structure of the wood Van Tieghem
unites the genua Drimy&, which he moreover subdivides into five genera (Dn'mys, Wintera, Buhbia,
Bell;otum, and EXl1sjJet'mum), with ZYgrJgynum, as well as Trodrodendron and Tetracentron (see
tM note undL'1- 'Irochodendraceae) to form .. special taxonomic group, the HomoxylCes, as oppo~
to the rest,ofth~ Dicoo/ledons (Heteroxylee.). . .
It may a~ be POlnted out that D1.ppolito'. statement to the effect that tracheids onlyaod no
vessell are present, at any rate in the later growth of the wood, in species of Illilium, LiritJdenaron,
AfagnoJi'a, AIt't:It~/ta, and Talaul1Ia, is without douneincorrect. 1 have recently examined the w~
from tbick pieces of the stern of LiriOiUndrtm tutipi/era and Magllolia grandi/lo,a,.and have met
with vessels baving ch.racteristic scalarifonn perforations.
 ADDENDA. -MAGNOLIACEA E

a composite and continuous sclerenchymatous rin~ (Magnolia pro parte,

Talauma, Michelia pro parte, Drimys pro parte), or ISolated groups of bast-
fibres (Magnolia pro parte, Mi(helia pro parte, Manglietia, Liriodendron,
IUicium, DTimys pro parte, Schizandra, Kadsura, Zygogynum), between which
there are sometimes groups of stone-cells, so that.an interrupted and composite
mechanical ring is formed. For fttrther details regarding the differentiation
of the sclerenchymatous riIJg in MtJgnolia and Talauma see also Pitard, loco cit.
Secondary hard bast showing varied differentiation is a1so present in species
of Talauma, Michelia, Manglielia, Illicium and Drimys, while in the Schizan-
dreae the bast includes only the crystal-~clerenchyma, described on p. 29;
similar elements may also occur in the primary cortex and in the pith 1.
LiteratuTC: Wijnaendts Fnmcken, Sc1ereiden, Dis.. , 'Utrecbt, 1890, pp. 36-40.-Brandt, Wenig
bek. Rinden, Diss., Dorpat, 1894, p. :n et seq.-[Matsuda, Anatomy of Magnol., loum. CoIl. Se.
Imp. Univ. Japan, vi, 1894, p. Il5 et seq.]-Parmentier, Hist. d. Magnol., in Giard, Bull. scientif.
de la France et de Ia Belgique l1. 1896, pp. 159-337 and pl. viii-xi.-Biermann, Olzellen, Diss.,
Bern. 1898, pp.....-7.-So1ereder, Cercidipkyllum, Ber. deutsch. bot. Gesellsch. 1899. pp. 396.397.-
Van Tieghem, Homoxylees, Journ. de bot. 1<)00, separate copy, 68 pp.-D'Ippolito, Canle delle
Magnol., Malpighia, xv, 1901, pp. 438-60.-Pitard, Pericycle, These. Bordeaux, 1901, pp. 103, 104.
-BaJ"gagli·Petrucci. Legnami, Malpigbia. 190:3, p. 19.f (Talauma).-Simon, Wintergriine Gew.,
Ber. deutsch. bot. Gesellsch. 1902, p. :3.fI.-Strasburger, Drimys-Winleri, Flora, 1905. Erganz .• Bd.,
pp. :a 1 i, 21 8.-[Moll and Jamsonius, Milcrographi"e des Hobes, Leiden, 1906, pp. 80-106 (Talaunza,
Magw/ia, Manglietia, .J-.ft'chelia).]-Piccioli, .Legnami, Bull. Siena, 1906, pp. 127 and 14-8.-
[Hol~D, Einfluss d. Klimas, 1907, p. I17 (Micbelia).]

TROCHODENDRACEAE (pp. 3I-34)'.

I. To the statements made in the REVIEW OF ANATOMICAL FEATURES the
following additions are necessary. In Euptelea simple uniseriate clothing hairs
generally occur on the young leaf; these hairs consist of: (a) a baSal cell, or
several short basal cells, showing local biseriate arrangement; and (b) one or
several, more elongated, terminal cells with wide lumina. Arm-palisade
parenchyma is present in Euptelea polyandra, S. et Z. and E. pleiosperma,
Hook. f. et Th. I
2. STRUCTURE OF THE LEAF. The lower epidermis in the leaf of Euptelea
pleiosperma, like that of certain other members of the Order, shows papillose
differentiation.
The caou tchouc-ce1ls in the leaf of Eucommia are generally found in the veins
(in the ba.t of the vascular bundles, and in the ground-tissue), but they sometimes
also pejllet'Ste into the mesophyll. where they ultimately terminate with swollen
ends. In the root of Eutmnmia these caoutchouc-cells are present in the bast
and in the neighbouring pericyclic region.
3. SritUCTURE OF THE AxIS. Subepidermal development of the cork is
also found ~ Trochodendron and Tetracentron .
. Uttnture: Parmentier, in Gia.rd, Bull. scient. de la France et de la Belgique 27, 1896, p. 318
et~."";'&lereder, Ctrddipk.7llum, Her. deutsch.. bot. GeseUsch. 1899, pp. 400, .f0I.-Van Tieghem,
DomoxyUa, loam. de bot. 1900, separate copy, pp. 5 and 63.-Barthelat, Laticiferes de l'Eucom11lia,
Joa.ro. de boL 1900, pp. 55-9·

1 Pmot'.ltatement (Tissu erible, These. Paris, 1899. p. 1 .... ) as to the occurrence Q[ secondary
bud1n of wood and but In the primary cortex of the ScWzandreae is incorrect.
• In retaining thil Order 1 am guided solely by practical conaiderationB. Recent ipestlgatioD.l
(see Solereder, lOCo cit.. and Hallier, in Beih. Bot. Centralbl. 1903, p. 2.7 et seq.) have shown that
the Emera Cwc;a"I";,llum, Ruple/ea, and Eucomm;(J must be transferred to the Hamamelidac:eae,
and Tetmcenbrtll to the Magnoliaceae. Harms' recent interpretation of the tIoral features of
Cercidip;'yllum (Erganz.-Heft ii to Telleo ii-iv of the NatUrl. Ptlanzen(am. 1906) in no way alten
my views regarding this genus.
HIO ADDENDA

ANONACEAE (pp. 34-39).

I. The REVIEW OF ANATOMICAL FEATURES' in the first -place ·requires
amplification as regards the hairy covering. Short, simple, unicellular tri~
chomes occur (Polyaltkia hypoleuca, Hook. f. et Th.), but are rare. Stel1ate
or tufted hairs have also been recorded in Ellipeiq. and pelt ate hairs also in
M eiocarpidium. SpiCular cells traversing the leaf in the vertical direction are
a1so found in Heterapetalum brasiUense, Benth., whilst sc1erenchymatous fibres
running freely in the mesopbyll are also present in certain species of Asteranthe,
Heteropetalum, Sageraea, 'and Uvaria 1. Lastly, we may mention here the
occurrence of brown middle layers in the dry leaf in species of Popowia and
MezeUia.
2. STRUCTURE OF THE LEAF. The reinvestigation of the anatomy of, the
leaf in numerous African, American, and Asiatic species by Beyer t has con~
siderably increased our knowledge of the structure of the leaf in this Order.
With regard to this subject and the special types of differentiation of
the epidermis, Beyer's treatise should be consulted. The following features
presented by the in tegumentary tissue are of systematic importance: formation
of papillae on the lower epidermis in conjunction with the development of
cutlcular ridges connecting the papillae with one another occurs in Anona
glauca, Schum. et Thonn., Cleistopholis glauca, Pierre, C. Staudtii, Engl. et
Diels, an.d Enantia Kummeriae, Eng!. et Diels; a multiseriate (mostly two-
layered) epidermis is present in certain species of Anona, Cleistopholis, EUipeia,
Miliusa, Mitrephora, Packypodanthium, and Xylopia 3; palisade-like elonga~
tion of the epidermal cells is also found in U varia gabonensis, Eng!.. et Diels;
finally gelatinization of the epidermis has only been observed in Heteropetalttm
h~il"ense, Benth. According to Beyer (loc. cit., p. 545), all the species which
he examined appear to have stomata with, subsidiary cells parallel to the pore.
According to Beyer, mechanical elements are abundant in the veins;
vertical transcurrence of the latter by means of hard bast is also found in the
genera Anona, Ellipeia, Hexalobus, Is%ma, Miliusa, "Afitrephora, Polycerato-
carpus, Sagefaea~ Uvaria and Xylopia_. .
Beyerfound that the secretory cells were present throughout. Stenanthera
gabonensis, Engl. et Diels is especially characterized by having two superposed
layers of secretory cells in the spongy tissue. In Xylopia tenu.i/olia the walls
of the secretory cells exhibit stratification. The branched spicular cells and
the spicular fibres found in the mesophyll have already been mentioned above
(see under I).
Beyerreinvestigated the crystals of oxalate oC lime, which afford important
characters for systematic purposes. According to him also the occurrence
of solitary and clustered crystals of oxalate of lime in the epidermis of the
leaf is an important ordinal character. In other respects Beyer'did not find
in all cases that the older statements of Borodin were quite correct; it will
therefore be best to reproduce in the following paragraphs Beye:'s review of
- - - - - - -----------
1 The species in which Beyer has recently met with spicular fibres are a.s follows: A,uma
rfauca, Schum. et Thonn., A. sen~galm$is, PeI'S.; Asleran/lu As/erias, Eng}. et Diels; Gumleria
australis, St. Hil., G. Sellowiana, Schlecht., G. 'lJillosissima, St. Hil.; lIettroptfalum ImJSilumt,
Bentb.; Sag-eraea t/lijtic(J, Hook. {. et Th.; Uvaria Aftel#, U. ollgo/ellsis, Welw., v: cffffra.
E. Mey., l'!. ceylanica, Lo, U. IJ£nklagei, Engl. et Diel5, V. J,.·irllii, Olivo -' _
~ Beyer's investigations extend to numerons species belonging to (orty genera.
3 TheE<: aJ:e: Anona g'lauco, A. Klaz'nei, Pierre, A. paluslris, L, A. senega/tnsis, Peri., A.
Sfju411JOSa, L.,; Cfttslopholis a/oida, Eng!. et Diels (with a typical two-layered epidermis according
to Beyer's figure); Ellipeia I~pt()poda, King; Miliusa indica, Lesch.; Alilrtphora Maillgayi, King;
Ptu4ypodallihium confine, Engl. et Diels; XylojJJ'a ol/hiop;ca, A. Rich., X. a/riaJna, Olivo
 ADDENDA--ANONACEAE 8u
the nature and mode of occurrence of the crystals in the upper epidermis of the
leaf:
(a) All the cells of the epidermis, or only isolated cells containing each a single
clustered crystal :-A naxagorctl, A nana. A nonidt"um, A rtabotrys, A steranthe, Cle':s/o-
pholis, Duguetia, Eupomatia, Goniothalamus, Heteropetalum, Hexalobus. Meiocal'-
piditl-m, Monodora, PachypodantMum, Potyalthia, PopoW1'a, Rollinia, Unona, Uvaria,
Xylopia. (Exceptions: Monodora Preuss':':, Engl. et DieIs-clustered and solitary
crystals; Popowia elegans,' Engl. et Diels-clustercd crystals in upper. solitary
crystals in lower epidermis; Xylopia a/rt'cana, OJiv.---occasional sOlitary crystals
in both upper and lo",cr epidermis.)
(aa) Clustered and s<?litary crystals :-Bocagea (with round crystal-cells),
Ellipeia, Guatteria.
(b) Cells of the epidermis (usually all of them) containing each a solitary
crystal :-Enantia. Melodorum. Miliusa, Sageraea (not all th~ cells in this case).
Uvariopsis.
(bb) Solitary crystals chiefly present in the epidermal cells of the veins, also
in scattered cells in the rest of the epidermis :-Alphonsea, Ephedranlhus, Mitye-
phOJ'a.
(c) Crystals not observed :-Cananga, I soloma, M ezettia, Oxymitl'4, Poly-
ceratocarpus, Stenantheya.
In investigating the hairy covering Beyer met with trichomes of a type
new to the Order, viz. the unicellular' clothing hairs mentioned above, (see
sectioll 1). According to the same authority uniseriate clothing 'hairs are
widely distributed in the Order (Anona, Asteranthe, Cananga. Enantia, Ephe-
dra nth us, Goniothalamus, Guatteria, Heteyopetalum, Hexalobus, Me/IfJ do rum.
Miliusa, Oxymitra, Piptostigma, Popowia.. R(lllinia, Unona, Uvaria (Asimina
and Sect. Uvariodendron), Xylopia). They consist of two or three cells, of which
the terminal cell is invariably longer than the others, and sometimes (Anona,
Rollinia) of remarkable ~ength. Stellate or tufted hairs also occur in spe.cies
of Ellipeia, whilst peltate hairs 1 are ah;o found in M eiocarpidium lepidotz_,
Engl. et Diels. The rays of the tufted hairs in Uvaria lurida, Hook. f. et Th.
(sphalm. Ilucida' in Beyer's paper) are multicellular and uniseriateowingto th~
presence of one or inore division-walls, a point which I am able to confirm.
3. STRUCTURE OF THE AXIS. According to recent investigations ,stone·
cells are very commonly present in the pith; they are but. rarely absent
(Asimina and Monodora besides Eupomatia). \Vell-developed diaphragms
composed of stone-cells are also found, for ex~mple, in species of Goniothalamus,
Guatteria, H exalo bus, M elodorum, Popowia, and Stenanthera.
With regard to the structure of the wood and cortex., Beyer confirms the
anatomical characters which I pointed out as important in the diagnosis
of the Order, for the material which he investigated. Stone-cells are of occa-
sional occurrence in the bast, but never show such a regular arrangement as
in the cortex of Guatteria villosissima, St. Hi1., described by Moller. Beyer
demonstrated superficial development of the cork also in Cleistopholis and
Uvariopsis. .
We may add-thabthe species·of-lJlelodorltm examined by Beyer agree with
the species described by Schenck in having normal structure in the stem.
Literature: BOrgesen og Paulsen, Veget. dansk·vestind. Der, Bot. Tidsskrift, xxii, 1898-9.
pp. S0, 51 (AnblUl palllslns).-Pitard, Peri cycle, These, Bordeaux, 1901, p. ] o+-Arescbong,
Mangroveptl., Bibl. bot •• Heft 56, 190::1, p. 63 and Tab. vii.-Bargagli-Petrucci, Legnami, Malpighia,
1902, p. 295 (Xylopia).-H. Beyer, Anatomie der AnoM~:eae, insLes. d. afrikan., in Engler, Bot.

J Beyer doubts the correctness of O. Bachmann', statement as to the occurrence of peftate h~irs
in AlUma fuifuracea, St. Hit, or the correct determination of the plant examined by Bachmann;
but in this he is wrong. In Martius, Fl. brasil., xiii, I, p. 8, we find the follo"in~ statement with
reference to this species: 'folia .•• superne parcins lepidota, pHis stellatis.' Whether the species
belongs to the genus AnolltJ is • different qnestion.
 ADDENDA--ANONACEAE
Jahrb.. xxxi, 190:1) pp. 516-55; see also Engfer, M onogr. arrikau. Pflnnzenfam. : vi. Anooaceae, 19() I.
-Areschoug, Trap. rut. bladbyggn., Sv. Vet. Akad. Ha.ndl •. ~! D. z, '90S, pp. u6-.;o (Antma,
Arlabo/rys), pp. 14:1--4 and Tab. xxi-xxii (G(Jltio//za/amus)) and pp. 156-8 (C)'a/hOt"aJyx).-[For
additional literature see p. Il69.]

MENISPERMACEAE (Pp.39-43).
I. In the REVIEW OF THE ANATOMICAL FEATURES the following correc-
tions or additions are necessary. A uniform and characteristic type of stoma
is not present. In the axis the pericycle sometimes (Cissampelos /ascicuJata,
Benth.) contains no sclerenchymatous elements. Secretory sacs of varying
length, besides occurring in Anami,ta, &c., have been recorded in the genera
Alberlisia, Antizoma, Calycocarpum (?), Chasmanthera, Cocctdus, Desmonema,
Disciphania, Fa'UJcettia, Fibraurea, Hypsipodes, Kolobopetalum, ParaOaena,
Pericampylus (?), Syntriandrium, Tiliacora, Tinomiscimn and Tridisia, whilst
open intercellular secretory spaces have been shown to be present in certain
species of A nomospermum, M iersiophyton and Tinospora. Anomalous struc-
ture of the axis (VIZ. the occurrence of secondary vascular bundles in various
regions) has recently been stated to occur also in species of Chasmanthera,
Menispermum, and Stephan·ia. Oxalate of lime is deposited in the following
special forms: large ordinary, solitary crystals in the veins of the leaf in
Burasaia, Chasmanthera, Chlaenandra, Fawcettia, Hypsipodes, Kolob~etalum,
Leichhardtia, Syntriandrium, Tinospora ; rather large crystals resembling styloids
and situated in the leaf in pairs of epidermal cells and in the palisade-tissue
in Leichhardtia clamboides, F. v. Mull.; large clustered crystals in species of
Clwnd,odend,on and Syrrhonema, smaller (:rystals of this type in species of
Macrococculus and Pericampylus. The trichomes of the Menispermaceae are.
either typical clothing hairs or hairs of a glandular nature. No member of the.
Ord~ has been found to have exclusively unicellular clothing hairs. The
typical clothing hairs .for the most part consist of two cells, a short basal cell and'
a long terminal cell. Uniseriate dothing hairs composed of more than two'
cells have been observed onl y in species of A namirta, A rcangelisia, Calycocarpum,
Chasmanthera, Coscinium, Disciphania, Heptacyclum, Menispermum. (together:
with two-celled hairs), Paraoaena, StephanM, T";nospora (here ac.companied by
unicellular hairs). Besides the glandular shaggy hairs of lateorhiza we may
mention the following types of glandular hairs: the small unicellular, ellip-
soidal or club-shaped trichomes present in species of Kolobopetalum, M ierszo- '
phyton and Tinospora; and the longer'uniseriate trichomes found in species of
Calycocarpum and Par'!-oaena. The unicellular hydathodes, first recorded by
Haberlandt in Anamirta, are also found in Arcangelisia. Peculiar small rosettes
of silicified cells occur in the epidermis of the leaf jn Coscinium Blumeanum,
Miers. We may add the following special features presented by the leaf
in certain species to the enumeration given in the earlier portion of this work:
the papillose or sclerotic differentiation of epidermal celIs; the very rare
occurrence of a hypoderm or of a locally tWO-layered epidermis; the very
rare arrangement of the stomata in groups; the occurrence in the mesophyll
of arm-palisade tissue (scarcely typically differentiated)~ and of spicular cells
differentiated as hairs or assuming some other form; the presence of tubular
cells filled with siliceous substance at the ends of the veins, &c.
Z. STRUCTURE OF THE LEAF 1. Recent investigations have shown that

1 Re<:ent researches on the strncture of the lea.f, the results of which lie before me in manuscript,
have been canied out under my guidance by K. Krafft, who continued Aner's uncompleted investiga-
tions (see the earJier part of this work, p. 40, footnote). Krafft's researches extend to lhe follow-
i.ng genera, which are enumerated according to Engler and Prantl's system: I. Cocculeae: il1eni-
spwmum, Pericampylm, Sarcope/alum, Cocculus, lJip/oelisia, Sltpnallia, AnliuJnUJ, Cissampeks,
Perajhol'(I., eye/ea, TiliactJra, Abuta j 11. Tinosporeae: HUSlmamlUt, \ .Iatan-hiso, Ti1U)sp~ra,
 ADDENDA-},IEN1SPERMACEAE

bifacial structure 01 the leaf·!S tar more WIdely distributed than centnc. The
palisade and spongy tissue show a very varied type of differentiation. Krafft
records the presence of mar~nal pits ill the epidermis in. species of Abuta,
Adeliopsis, Anomospe,mum, Arcangelisia, Bania, Cocculus, Fawcettia, Hepta-
cyclum, Husemannia, Hyperbaena, Limacia, Mac,ococculus, Pachygone and
Pycnarrhena, where he. found them in nearly all cases on both sides of the leaf,
while in A deliopsis they are confined to the upper epidermis. In the course of the
recent investigations gelatinization of isolated epidermal cells has not been
observed in any species. But subepidermal ~yers of mucilage similar to those
described by Auer·inAnomospermum reticulatum are rather widely distributed;
each patch of mucilage is derived from the inner walls of a group of upper epi·
dermal cells and the adjacent w~ of the uppermost layer of palisade-cells; this
feature is found in A deliopsis, A nomospermum pro parte, Cissampelos pro parte,
Cyclea pro parte, Limacia pro parte, and Stephania pro parte. Epidermal cells
of exceptional size are stated by Krafft to occur in Antszoma and Cocculus pro
part,. especially on the upper side of the leaf; the same author mentions the
presence of sclerotic epidermal cells situated over the veins on both sides of
the leaf in,Anomospermum reticulatum, Eichl., Hyperbaena lauri/olia, Urb. and
Tridisia 1l1'Ucoubensis, Baill. Papillose differentiation of the epidermis in the
leaf has also been observed: (a) on both sides in A ntizoma calcari/era, Miers,
A. lycioi4es, Miers, Cissampelos c.apensis, Thunb., C. Pareira, L. (not in all the
forms of this polymorphic species), Cocculus Leaeba, DC. and Stephania rotunda,
Lour. (not all the cells papillose in this case); (b) on the lower side ,only in
Cissampelos fluminensis, Eichl. (slightly papillose), C. ~laberrima, St. Hil., C.
Pareira pro parte (see above), Diplodisf,a macrocarpa, MIers, Hypsipodes subcor-
datus, Miq., Jateorhiza Columba, Miers, Menispermum dauricum, DC. (isolated
cells above the veins), Miersiophyton kamerunense, Engl. (isolated cells above
the veins), Peraphora robusta, Miers, Stephania abyssinica, Walp., S. di$color,
Spreng., S. elegans, Hook. f. et Th., S. hernandifolia, Walp.and S. rofunda,
Lour. Curiously enough there is no new record of a species with hypoderm; in
Parabaena sagiUata, Miers, the upper epidermis consists locally of two layers.
A special type of differentiation is exhibited by the upper epidermis ~ Desmo-
nema pallide-auTantiaca, Engl. et Gilg, both the outer and inner walls being
very strongly thickened and partly converted into mucilage, and by the upper
epidermis in ChlaenandTa ovata, Miq., in which the strongly thickened outer
walls project convexly into the lumina·-'Of the cells. Finally, a distinctly
peculiar feature of the epidermis is presented by the small groups of silicified
cells in Coscinium Blumeanum, Miers. They are situated in the upper epi-
dermis and lie immediately above the sclerenchyma of the smaller vertic~y
transcurrent veins. They consist of a small central cell, which is rounded In
surface-view and might possibly be interpreted,as the rudiment of a hair, and
of a rosette of cells with unevenly thickened wal1s~ which in surface~view have
the same appearance as the well· known cystolith~like bodies in the subsidiary
cells of- the trichomes of Lithosperm·um and other Boragineae. In some cases
two of these groups of silicified cells arc united. Regarding their function
nothing is known.
In very many Menispermaceae the stomata are surrounded by ordinary
neighbouring cells. A definite ,and uniform type of stoma is not present. Not

,/i"awceltia, .Afiersiopll_Y/olI, C!tasmallflura, IIypsipodes, Deslllt»Uma, Parabama, Ducip!iania, Kolo/Jo-

pdiJIUIII, SYII,Jr;4I1drium, Ca/ycocarpum, AIIa.mi,./a, Coscilzium, ArcaltKtlisia, Cltlll4nanrira; HI.
Limacieae: Allol}lospermum, Limacia i IV. Pachygoneae: T.,.t'(/isia, Hejlacyc!u"" PycnarrMml,
,1t/tUnKOCCU/UI, PleoDlIe, Sdadotamia, Alh,,,lisia, PachYP',e, Hyje,,/JaeluI, C/uJndro(/elUlnJn, De-
100u/m, CtZrf'"QJlia, Adeli(}/JJls, Ballia, S)lrrMIt~ma, L.~i'chhlZrdlia. Additional recent statementl 011
the structure of the leaf and axis are further also to be fouod in Maheu's p3pcrs, 11. ce.
 ADDENDA-MEN1SPERMACEAE

uncommonly, however, one observes a tendency for subsidiary cells to develop

in connexion with all or some of the stomata; atcording to Krafft this is the
case in: Albertise'a (subsidiary cells arranged like a rosette), Anamirta, Ban·ia
(rosette-like subsidiary cells). Burasaia pro parte (4-6 neigbbouring cells, of
which one is placed parallel to the pore on either side), Carronia (rosette-like
subsidiary cells), Chlaenandra (neighbouring celS"with thin walls, so that they
appear as subsidiary cells), Cocclllus, Cyclea pro parte, Fawcettt'a, Heptacyclum,
Husemannia (rosette-like subsidiary cells), Hyperbaena (subsidiary cells rosette-
like in H. laurifolia, Drb.), Leichhardtia (4-6 neighbouring cells, one being
placed on either side of and parallel to the pore), Limacia pro parte, M acro-
cocculus (subsidiary cel1s rosette-like), Paraoaena, Pleogyne, Pycnarrhena (narrow
subsidiary cells), Sa."copetalum. In Macrococcu/us p(Jmiterus, Becc. severa}
pairs of guard-cells with their subsidiary cells together form stomatal groups,
separated from one another by rows of ordinary epidermal cells, without
stomata and differently shaped. The only additional record of the occurrence
of stomata on the upper side of the leaf is in Antizoma pro parte.
The' previous statements regarding the structure of the mesophyll
require the following additions. The. thickenings and swellings confined
to certain portions of the wall in cells of the palisade and spongy
tissues, first recorded b-y- Auer in Stephania kerna",d"'lolt'a, are stated by Krafft
to occur in Adeliopsis, Carronia, Cissampdos, CoccuJus pro parte, Cyclul,
Limacia pro parte, Pachygone, Pericampylus, Stephania ind Synhonema.
The palisade-tissue in Disciphania lobata, Eichl. is composed of a single
layer of short cells, and shows slight differentiation as arm-palisade-tissue. In
certain species a part or the whole of the spongy tissue has thick walls, which
are then in most cases lignified; this feature is found in Alberlisia papuana,
Bece., Anamirta Cocculus, Wight et Arn., Bania thyrsifiora, Becc., Husemannia
protensa, F. v. Mull. and Parahaena sagt'ttata, Miers. Mechanical elements are
also frequently developed in the mesophyll. The following types are found:
isolated palisade-cells resembling idioblasts, and strongly thickened, but with
wide lumina, in Coscinium Blumeanum, Miers; isolated sclerosed cells belong-
ing to the spongy tissue or entire layers of such cells near the lower epidermis
in Abuta conca/or, Poepp. et Endl.; branched -stone~cells in the neighbourhood
of the vascular bundles of the larger veins in Chlaenandra ovata, Miq.; slightly
or considerably developed branches of the sclerenchyma of the veins, though
not typically fibrous, in the mesophyll in Abuta concolor, Poepp. et End!.,
CIaond1ltbdendron platypnyllum (Miers), and Detandra paraens£s (Eichl.); spicular
fibres, which branch off from the sclerenchyma of the veins into the mesophyll,
in Anamirta Cocculus, Wight et Arn. and Arcangelisia lemniscata~ Becc.;
girder-shaped spicular cells, which traverse the entire thickness of the leaf-tissue
from one epidermis to the other, in Heptacyclttm Zenkeri, Eng!. (where they
have wide lumina, and are ~ac-like. often dichotomously divided, and give
off root-like braoches beneath the epidermis), Anamirta Cocculus (like fibres and
unbranched), Burasaia gracilis, Decne., B. congesta, Decne. (branched with
transitions to hair-like differentiation), and probably also in Tinomiscium
reticulatu1n, Miers (' piliers' according to Maheu); girder-like spicular cells~
which essentially be10ng to the palisade.tissue onlYi branch beneath the
upper epidermis, and give off rays into the spongy tis~ue, while they show
transitions to hair-like differentiation, in Adeliopsis decumbens, Benth. (sub-
epidermal rays penetrating the subepidermal layer of mucilage), Burasaia
madagascaT1'enst's, DC., and Limacia cuspidata, Hook. f. et Th. (Cuming,
No. 1252, Philippines, subepidermal rays penetrating the subepidermal layer
of mucilage); lastly, spicular cells differentiated like hairs in the spongy tissue
in Adeliopsis decumbens (star-shaped), Anomospe,mum iapurense, EichI. (with
few rays), Limada cuspidata and L. microphylla, Miq. (sta.r-shaped).
 ADDENDA-MENISPERMACEAE 815
,U{)
The vascular bundles oT the veins are in most cases accompanied by well ..
developed sclerenchymatous tissue. According to Krafft, exceptions to this
rule are found, especially. in A ntizoma, Cissampelos pro parte, Desmonema,
Dipl"clisia, Disciphania, Miersiophyton, Stephania pro parte and Tinospora
pro parte; in these the sclerenchyma is absent. In very many genera and
species the lateral veins of tH, second order, and even some of the next smaller
size, tend to be vertically transcurrent by means of special parenchymatous or
sc1erenchymatous tissue accompanying them. According to Krafft, the follow ..
jng genera and species may be especially mentioned as examples of this:
DiPloclisia and Menispermum (veins vertically transcurrent by means of tissue,
which is not sclerosed), Heptacyclum, Hyperbaena, Limacia microphylla, Miq.,
M acrococculus, Peraphora, Pericampylus, Pycnarrhena, Sarcopetalum and
T,iclisia (veins vertically transcurrent by means of a sclerenchymatous
ring). Abuta, Anomospermum reticula tum, Eichl., Coscinium, Limacia velutina,
Miers and Pachygone (veins vertically transcurrent by means of regular plates
of sclerenchyma). The structure of the smaller veins of Chondrodendron
platyphyllum, Miers is also noteworthy; these veins, which project on the
lower sid~ are vertically transcurrent on the upper by means of sclerenchyma·
tous plates, whilst on the lower side a group of palisade-cells fonning a single
layer adjoins the vascular bundle.
The forms in which oxalate of lime i~ deposited are very numerous, as
may be gathered from the previous description. Specially common types
are the crystals, varying in size, of prismatic, acicular, rhombohedral or
other sh~pes, with transitions to styloid~like crystals or crystal-sand; other
common forms are small or sometimes larger clustered crystals, and ordinary
large solitary crystals. In most cases oxalate of lime is excreted in abundance,
being present not only in the mesophyll and veins, but often also in the
epidermis. The only cases in which Krafft found no oxalate of lime in the leaf
were Abuta Grisebachii, Tr. et Pl., A. Imene, Eiehl., Miersiophyton kamertlnense~
Engl. and Sciadotaenia amazollica, Eichl. Only tht typical and principal
forms of crystals can be employed for systematic purposes, since the others are
frequently connected by transitional forms, so that their delimitation bee'omes
difficult. In the first place the occurrence of the following types may be
mentioned as specially important; ordinary large solitary crystals are "found
in the veins in Burasaia pro parte, Chasmanthera, Chlaenandra, Fa wcettia ,
Hypsipodes, Kotobopetatum, Leichhardtia, Syntriandrittm and Tinospora;
somewhat smaller rhombohedral crystals are present in Abuta prQ parte,
A nomospermum, Burasaia pro parte, Heptacyclum, Husemannia, Hyperbaena
pro parte, Limacia, Atacrococculus, Pachygonc, ,Pleogyne, Pycnarrhena and
Tiliacora pro parte; rather large, styloid-shaped or prismatic crystals, which
under a low magnification give the appearance of the veins being striulated, occur
in Cissampelos pro parte, Cyclea pro parte, Diploclisia macro carp a, Miers,
Jateorhiza Columba, :Miers, Pericampylus· incanus, Miers and Stephania; rather
large styloid-like;.mOstlygeniculate, hemitropic crystals are found in numerous
cells of the palisade-tissue in Leichhardtia clambo1'des, F. v. Miill.; large clustered
crystals are present in the palisade-tissue in the immediate neighbourhood of
the vascuIa'r bundles of the veins in Chondrodendron platyphyllum, M;iers and
Syrrhonema jasciculatum, Miers, smaller clustered crystals in the palisade.. tissue
of the veins in Pericampylus incanus, and small clustered crystals or aggre ..
gates resembling clustered crystals singly in the palisade..cells of Macrococculus
pomiJerus. The epidermis, as mentioned above, frequently contains various
forms of oxalate of lime. In Desmonema pallidt-attrantiaca, Eng!. et Gilg and
Tiliacora Warneckei, Engl., in particular, and also in Anamiria Cocculus the
epidermis on both sides is differentiated as a regular crystal-containing armour ;
the cells in this case contain chiefly large rhombohedral crystals. Finally, we
 816 ADDENDA-MENISPERMACEAE
(3 -e.:,
may mention the paired crystal-cells, found in- the upper epidennis of Leich-
hardtia clamboides; these, like the paired crystal-cellS of Canavalia villosa (see
Systematic Anatomy, p. 265, Fig. 58, e), as a rule contain hemitropic crystals
resembling styloids. Krafft met with sphaerocrystalline masses composed of
an unknown chemical substance in the lower epidermis of Sarcopetalum Harvey-
anum, F. v. Miill. In Arcangelisia lemniscata tubular cells filled with a siliceous
substance are found at the ends of the smaller veins.
Several additions may be made to the earlier statements on the' secretory
receptacles of the Menisperrnaceae, which referred for the most part to those
occurring in the axis. In the first place we may note that, besides secretory
cells of varying length and with diverse contents, there are also open inter-
cellular secretory receptacles. The latter are situated singly in the nleshes
of the veins, and are often enclosed by arc-shaped cells of the spongy ,tissue:
Krafft met with them in the genus Ti1tOSpora (but only in T. Bakis, Miers),
and also in A nomospermum iapurense, Eichl. and M ierst'ophyton kamerunense,
Eng!. The nature of the secretion varies. As regards the distribution of the
secretory cells, we may in the first place enumerate those genera in which Krafft
met with no secretory cells in the leaf, or only found them in a certain number
of species; these are: Abuta, Adeliopsis, Anomospermum, Arcangelisia, Bania,
Calycocarpum, Carronia, Chlaenandra, Chon dro dendron , Cocculus pro parte,
Coscinitem, Cyclea, Detandra, Heptacyclum. Husemannia, Hyperbaena, Leich-
hardtia, Limacia pro parte, Macrococcultts, Afutispermum, Miersiophyton,
Pachygone, Peraphora, Pericampylus, Pleogyne, Pycnarrhena, Sarcopetalum,
Sciadotaenia, Stephania and Syrrhonema. The secretory cells in' the leaf are
frequently restricted to the vems, being for the most part associated with the
sc1erenchyma of the latter. and sometimes even embedded in it; they then
usually have an elongated shape, like the secretory cells of the axis. It is not so
common to find the secretoryceUs free in the mesophyll, where they occur in the
form of rather short sacs; when this is the case the veins also generally contain
short secretory sacs only, more rarely sacs of greater length. The contents of the
secretory sacs vary greatly. In herbarium-material the long sacs in the veins
mostly have yellowish contents, which after treatment with eau de Javelle
resemble latex; sometimes, however, they are brown. In the short sacs the
secretion generally has a reddish-brown colour, but in some cases it resembles
gum, e.g. in A ntizoma. Hallier informs me that Fibraurea possesses white latex ..
Maheu ~s recently discovered contents resembling caoutchouc, and capable
of beirJIJ 6t'awn out into threads when parts of the branches or leaves are broken
across, m the secretory sacs of Tinomiscium javanicum, Miers, T. petiolare,
Miers, and T. phytocrenoides, Kurz; these sacs are l:20-150 f" long, and occur
. in the stem at the margin of the pith, and on the outer side of the arcs of
peri cyclic hard bast, and in the leaf especially in the neighbourhood of the veins.
Poulsen met with mucilaginous contents in the aerial roots of Tinospora crispa,
Miers.
Elongated secretory sacs with yellOwish or brownish contents have been shown
by Krafft to occur in the veins of the leaf in the following genera: Albenisia,
.A namirta, BUYQsaia, Chasmanthera, Cissampelos pro parte, Cocculus pro parte
(in most cases), Desmonema. Disciphania, Fawcett-ia, Hypslpodes, jate01'lli-za, Ko'obo~
j>etalum. Limacia prQ parte, Pal'abaena, Syntriandrium, Tilio.coya, Tinospora;
short secretory sacs, which mostly have reddish~brown contents, and are sometimes
recognizable in transrnitted light by. using a lens, were found to occur independently
in the mesophyll, and, where expressly stated, also in the veins in: Antizoma
calcarijeya. Miers, A. ly c£o ides, Miers (also in the veins), Cissampelos PareiYa pro
parte, C. jasciculata, Benth., Diploclisia mac1'ocarpa. Miers. (also in the veins).
Tiliacora racemosa, Colebr., Triclisia IOtfCottbensis, BailI. (also in the veins, and 1fl
the mesophyll sometimes in rows).
The earlier statements and those of Krafft agree with regard to the occurrence
 ADDENDA-IIENISPERMACEAE
of elongated secretory sacs inthe petiole and axis in species of Anamirla, BUf'asaiiJ,
Cissampelos. Diplocluia. jateOf'Aiza, Limac;a and TinospOf'tJ. Maheu has recently
demonstrated secretory sacs also in Filwau,ea d.l()l'oIeuca (= F. tinclm-ia, Lour.
ex syn.), Calycoca,pum sp., and PericamPylus sp., as well as in additional species
of Burasaia, Cissampelos, and' Codulus; m view of Krafft's results Maheu's state·
ments with reference to Cal~ca,pum and Pe,icampylus require re-examination
and confirmation.
Owing to recent investigations our knowledge of the hairy covering of the
Menispermaceae is much more complete than formerly. The trichomes may
be classified under typical clothing hairs and glandular hairs, but the prec~e
function of the latter remains to be determined in living material. A specially
noteworthy feature of the typical clothing hairs is that they are not exclusively
unicellular in: any member of the Order. Krafft, moreover, only records these
unicellular hairs in Tinospora Bakis, where they represent a reduced form
of the multicellular clothing hairs. Blottiere's earlier statement regarding the
presence of unicellular clothing hairs in Abuta ,"'eseem is incorrect and may
be disregarded. The most widely distributed form of clothing hair is represented
by bicellular trichomes with a short basal cell and a longer terminal cell of
varying length. Points of difference are found in the structure of the basal
cell, the upper end of which frequently passes over into a stalk-like process
of varying leng~h and either solid or provided with a lumen, as well as in the
nature of the 'Wall and lumen of the terminal cell. These bicellular dothing
hairs have beert recorded in the following genera: Abuta, Adeliopsis, Alberlisia,
Antizoma, Bania, Carronia, Chondrodendron (with a specially long terminal cell),
Cissampelos, Cocculus, Oyclea, ,Detandra (with a specially long terminal cell),
H usemannia, H yperbaena, Limacia, M acrococcu[us, M enispermum (accompanied
by three· celled hairs, see below), Pachygone, Peraphora, Pericampylus, Pleogyne,
Pycna"hena, Sciadotaenia,-Sy"honema, Tiliacora. The two following forms of
hairs may be interpreted as modifications of the bicellular trichomes: (a) those
of A nomospermum reticulatum, Eichl., in which the basal cell Undergoes subse-
quent division by thin vertical walls; and (b) the three-celled trichomes accom-
panying others composed of two cells in the species of M enispennum; here
a third ceUis intercalated between the basal and terminal cell, 'and is/delimited
from the latter by an oblique wall. The trichomes of M enispermum constitute
a transition to the uniseriate clothing hairs, which are composed of more than
two cells, and exhibit various types of differentiation. The uniseriat~ hairs
are found in the following species : A namirta Cocculus (3-6-celled, wtth thick
division-walls; especially on the do matia) , Arcangelisia lemniscata, Becc.
(3-8·celled, with both thick and thin division-walls; only on the domatia),
CaJ'Ycocarpum Lyonii, Nutt. (very long, composed of 4-7 cells, and with thin
division-walls), Chasmanthera dependens, Hochst. and C. strigosa, Baill. (2-6·
celled, those of C. strigosa with subsidiary cells, each of which contains a large
solitary crystal of oxalate of lime); -Disciphania lobata, Eichl. (3-?,-celled).
Heptacyclum Zenkeri, Engl. (3-?,-celled, recurved at the base), Pa,abaena
~. Miers (2-s-celled, with thick division-walls; see under the forms
of glandular hairs), Stephania discolor, Spreng. and S. hernandifoUa, Walp.
(5-Io-celled, with thin walls, a number of cells being sometimes placed ~ide by
side at the base of the hair), Tinospora Bakis, Miers (2-4-celled, conical, occa-
sionally biseriate at the base). Lastly, to this type of hair we may also refer
the characteristic 5-8-celled, uniseriate clothing hairs of Coscinium Blumeanum,
Miers, these being cpmposed of several short basal cells exhibiting uniseriate
arrangement, and a very long flagelliform terminal cell; in a certain sense
these again form a transition to bicellular clothing hairs with a short basal cell.
Amongst the glandular forms of hairs we may firstly include the small
unicellular club-shaped or ellipsoiqal trichomes (possibly hydathodes), which
IIOUIEU. 3 G
818 ADDENDA-MENISPERMACEAE

either occur singly on the leaf~surface, especially in the neighbourhood of the

veins (and sometimes sunk singly in small pits, e:g. in Tinospora aurictdata,
Engl.), or lie in groups at the base of the lamina in the angles formed hy the
principal veins on the lower side of the leaf, Qeing sometimes situated in small
pit-like depressions (domatia ?). These unicellular club-shaped trichomes
have only been recorded in the following ~pecies: Kolobopetalum auriculatu,n,
Engl., Miersiophyton kamerunen~e, Engl., Tinosj>ora auriculata, Engl., T. Bakis,
Miers, T. cordi/olia, Miers and T. reticulata, Miers. With these unicellular
dub-shaped trichomes we may class the glandular shaggy hairs of Jateothiza
Columba, represented in Fig. 7, A in this work. Finally, we may mention here
the uniseriate trichomes of Catycocarpum Lyonii (narrowed basally, and. thin-
walled throughout), and Parabaena sagittata (3-celled, dub-shaped, with
a specially thin-walled terminal cell), which are apparently glandular and may
be derived from the typic~l clothing hairs found in these species.
Krafft met with unicellular hydathodes, having the same structure as
those of Anamirta Cocculus (according to Haberlandt), in the upper and lower
epidermis of the .veins in the leaf of Arcangelisia lemniscata. He made the
interesting observation that in both species the walls of the hydathodes,
originally composed of cellulose, became subsequently suberized, and that
additional thickening layers, which. are lignified and also 'Pitted, are deposited
on the suberized wall, naturally involving the loss of function of the hydathodes.
Typical domatia in the form of sman pits are found, in the angles' formed
by the veins of the second order in A namirta Cocculus and A rC4ngelisia lemnis-
cata. The entrance to the pits is clothed with hairs.
3. STRUCTURE OF THE AXIS. According to Maheu, the development of
cork takes place in various positions: either ;in the epidermis (Tinomisciu,,,
petiolare), or in the subepidermal layer (Menispermutn), or first in the primary
cortex and then in the pericyde CCissampelos obovata, DC.'). In Meni..
spermum canadense and species of Abuta, Cocculus and Pericampylus Damm
observed the formation of 'a cuticular epithelium,' which sooner or later
becomes replaced by cork.
The following additional forms are stated by Maheu to exhibit an
anomalous structure in the axis: Chasmanthera, Cissampelos hexandra
(= Stephania discolor, Spr. ex syn.), Cocculus Thunbe1'gii and M enispe1mum.
According to Maheu, the origin of the anomalous growth is not always the
same.
In the first case (in an unnamed species of Menispel'mum) the secondary vascular
bundles originate, according to Maheu, in the primary cortex; in the second case,
which is the most widely distributed (Pal'eira brava = Chondrodendron tomentosum.
Ruiz et Pay. ex syn. and CoccuJus laurifoi'ills, DC.; see also Gerard with reference
to A bula f'ufescens and Chondrodendl'on platyphyllum) they arise in a secondary
tissue, formed by division of the endodermts; in the third case (Cocculus Leaeba, DC.,
and Cissampelos Pareif'a) they appear in the pericycle; and in the fourth al1d last
case (Abuta Selloana, Anomospermum gtand~'folium) Cocctd«s platyphyllus = ChondfO-
dendron platyphyllum. ex syn., Cissampelo$ Mauys'tiana, Wall. = Pet'icamf>ylus
incanus, Miers ex syn.) immediately external to the soft bast of the normal ring
of bundles. Vlith reference to Cocculus Th",nbtfgii and Cissampelos hexandfa
I quote Maheu's own words: 'Ie cambium peut ~tre inactif. par places ou pro-
gressivement sur toute sa surface; il apparatt alors en dehors de lui une nouvelle
assise generatrice.·
Literature: Gerard, Form. &nom. des Menisperm., Comptes renduI, Paris, cHi, 1886, pp. 1027.
I028.-(Heckel et Schlagdenhautren. Bakis et Sangol, Ann. lnst. col. Marseille, iii, 18 95, p. ~9
d seq.]-Schwabaeh, in Bot. Centralbl. 1898, iv, p. 359.-Spanjer, Wasserapparate, Bot. Zeit. 1898,
i, p. 6I.-Pitard, Pericycle, These. Bordeaux, 1901, pp. SO-S3.-Damm, MeJujiihr. Epld., Beih. Bot.
Centralbl., xi, 1902, especially pp. 236-9.-Mabeu, Recb. anat. lur lea Menbperm., lonrn. de bot.
1902, pp. 369-78; !;ee also Congres internat. de Pharm., sep. copy, 7 pp. (received xii, 19°-4).-
Foul sell, Luftr.,dderne hos Tin()sptwa ais/a, Vidensk. Meddelels. Kjfl~nhavn, 1902, pp. 235-7.-
 ADDENDA-MENISPERMACEAE
Morini, Stud. anat. del caule delle Menisperm., Mem. Acead. Bologna, 190.... -AreschoUf, Trop.
vaxt. bladbyggn., Sv. Vet. Akad. Hand]. 39, D. 2, 1905, pp. 21, U and T:r.b. xxiv (Pen·ca1llJyltU).-
a
Maheu, Laticiferes caoutchouc: Tinomiscitlm, Comptes rend us, Paris, • d~c. 1905, sep. copy, II plJ.
-[For additionalliteratnre, see p. 1171.) ,

BERBERIDEAE (pp. 44-47).

I. In the ,REVIEW OF THE ANATOMICAL FEATURES the following changes
and additions are riecessary. Glandular hairs with a uniseriate stalk and an
ellipsoidal multicellular head occur in Epimedium. Differentiation of papillae
on the epidermal cells of the leaf (occasionally on those of the upper side as
well) is also found in numerous species of Berberis and in certain species of
Epimedium and Berberidopsis. In certain species of Berberis and M ahonia
a parenchymatous hypoderm is situated beneath the upper epidermis of the
leaf, and in very many species of these two genera there is a hypodenn, com-
posed of sclerenchyrnatous fibres, in the same position. The tendency of the
vascular bundles in the subaerial stem to show an arrangement, like that
typical of Monocotyledonst is at least indicated in other herbaceous members
of the Order besides those previously mentioned. The nature of the peri cycle
varies. The' cork either arises on the inner side of the groups of pericyclic
fibres, or in the primary cortex, sometimes in a subepidermal position.
z. STRUCT-URE OF THE LEAF. The ,structure of the leaf in the genera
belongir.g to the tribe Berbereae is now well known, owing to the investigations
of Citerne, Kohne, Fedde and C. K. Schneider. The epidermis furnishes
numerous anatomical characters suitable for specific diagnosis, e.g. besides
the shape of the cells, firstly the nature of the cell-wall, and the frequent occur-
rence of papillae 1 in Berberis and M ahonia. Cuticular beads are also found
1 According to Citernc: and otheu, the formation of papillae on the lower side of the leaf hal
been demoQstrated in the following species: Btrhris a,fapaltllJis, B. tlrmata, B. asial,'(a, B.
~radl)'lJot",a, B. carinata, B. ,Ailmsis, B. 'I)ncinna, B. (orym/Josa, B. das),stlUhya, B. dmsijt~ra,
Kohne, B. tliapnana, B. dicl),opltylla, B. diff'lUa, B. tmpetn"folia. B. Jertlz, B. jerruginea, B.
IhxUf)stJ, B. Gayi, B. rlauca, B. Goudotii, B. GristbacAii, B. gui/adtt, fl. lJettrtljada var.lajJi//4sd,
B. Itorrida, B. lau,ina, B. Iycium, B. ftUUrtlstpala, B. tlt1atd, B. palltns, B. papilli/erd, Kohne,
B. picktncmsil, B. pyrocarja, Kohne, B. rejJem, Lindl., B. t'igidifolia, B. sas,:cola, B. serra/lI-
amtala, B. sinellsis, B. tl)/imtnsis. B. Tohonos~iana, B. UmDe/lattl, Wall., B. virtsullS, Hook. f' I
B. IVtddtllii, B. ~uttllanensu; Malumia ancustifolia (Harlw,), M. Fytmontii (Torr.), M. lJae"us!tJ-
carpa (Woot.), M. incerta, Fedde, M. pinna/a (Lag;), AI. pumUa (Greene), M. relens, Don, M.
lri/o/ioJata (Morie.); Elim/dium Davuti, E. sintlUis j BerbeJidopsis coraJlina; papillae arc
present OD the upper side of the leaf in: Ber/Jeris tutinacantha, B. acuminala, B. ag-apatemis, B.
asiatica, B. crispa, B. ferru,rinta, B. fluuosa, B. lazijltJra, B. /evil, B. /ycium, B. Weddellii i
~fa1wnia Andrieuxii (Hook. et Am.), M. Fremtmlii, M. htumatoearja, M. pumila, M. Jubintegri-
fa/;;', Fedde, M. tnfoNo/ala. The papillae found in Btrlltns gui/adu, B. ()fJQta. and B. toliinensis
are of a specially curious type; they are united into groups' and are situated at the margins of the
ia.dividua.l epidermal cells.
We may ennmerate here separately from the foregoing list those species of Berbtris (EuOel'luris)
in which C. K. Schneider ment.ions the Occurrence of a papillose epidermis in the leaf. Pollpillae are
present (they we~e only !a~en into account by Schneider when they were evident in a surface-view of
th~ epidermis)on the:lower side of the leaf in: Berberis afg-ham'ca, Schn., B. ag'ajJalemis, Lecbl.,
B. armata, Cit., B. aurakuactnsis, Lem., B. asiatica, Roxb., B. brach),o(J(/ria, Gay, B. butneHat-
jolia, Schn., B. Duxt'fo/ia, Lam. var. papillosa, Schn" B. diltnsi's. Gill .. B. dtil1lDomtis. Scho.,
B. chitria, LiDdl. var. sikkimensis, Schn. (slight pApillae), B. co/zcinlta, Hook. f., B. (o'Ym6asa,
Hook. et Am., B. (u!leala, DC., B. das,Ystacnya, Maxim., B. diaphana, Ma.xim., B. t[tgllllS, Scho.,
B. empelrifolia, Lam., B. fle.xucsa, R. et P., 8. ForskaliaNJ, Schn. (not at all points), B. rtzl'Awa-
UtUu, Schn. (more or leu marked), B. rranaijl",.tI, Tura., B. Hallii. Hieron., ~. Jatu'uwma,
Schn. (not always distinct), B. i,rMr'a/(J, Sehn., B. inlerwrima, Bge. var. triW(JlUnsis.i:Sclm. and
VaT. typic., Sc:hP.., B. kase/'.rarica, Rupr., B. KiiAneana, Schn .• B. M4ma(ltUnJU-, Scho., /1. lati/D/itJ,
R. et P., B. litllarifolia, Phil., B. Julea, R. et P., B. lye;I4_, Royle, B. madlJJepala, Hook. f. et Th"
B. M".i't%ii, Hicroo., B. niK"'c4ttS, O. Ktt.e., B. n_m",ularia, Hge. vat. pyr(J£arpa, Schn., B.
pal/eftS, Francho, B. Pearcei, Phil., B. PtliJialla, Schn., B. fJII)'l/ocanllta, Rusby, B. ~/,antlla,
Hems1., B . .fJi~, TutcZo, B. ,.ari~ra. Lecb1., B. I"uti1Urvia, Rusby, B. Soulltana, Schn .•
B. Tlw",stlJe.anII. Schn., B. TIJuII6tYril, DC. (in part), B. li,"ltlrils, Lesch., B. In~na, Kze., un~
'=tG2
 Al)DENDA-BERBERIDEAE

in· Berberis ma'ginata, whilst characteristic thickenings of the outer wall, in.
the form of a. network of ridges, are present in those species of Mahonia, which
Fedde includes in thtt Aquifoliatae and Paniculatae. In certain species ,of
Berberis and Mahonia 1 a parenchymatous hypoderm, mostly with thin walls,
occurs beneath the upper epidermis; in species of Berberis and Malwnia t with
thick evergreen leave:; a still more widely distributed feature is a hypoderm
composed of sclerenchymatous fibres and situated likewise beneath the upper ,
epidermis s. The stomata are found op the upper as well as on the lower side
of the leaf only in certain species of Berberis and Leontice '. Subsidiary cells
can only be SaId to be present in BerbeTidopsis, where each pair of guard.cells
is surrounded by a multiseriate ring of rather small epidermal. cells. Citeme
mentions the occurrence of lobe-like appendages at the ends of the guard·cells
in CauJophyllum, Epimedium David; and E. sinense. For the wateI'-pores of
PodophyUum see Spanjer, loco cit. In certain species of Berberis and-Mahonia'
one or more layers of the spongy tissue have thicker walls and scanty-chiaro..
t~ll,· so that they function as mechanical ,tissue; in some cases they are
. erentiated like a hypoderm.
As regards the veins of the leaf we may mention that Citeme attributes
generic importance to the number of vascular bundles in !he priricipal
veins {one bundle in CaulophYllum, JeUersonia; Leontic8; sev~al in Achlys,
Berberis incl. Mahonia, Diphylleia, Epimedium, Nandina, rodophyUum),
and to the occurrence of groups of sclerenchymatous fibres sftuated at the
margin of the leaf and sometimes developed in connexlon with a marginal
vascular bundle (only in Berberis incl. Mahonia, Epimedium, _Nandina).
The groups of sc1erenchymatous fibres, accompanying the vascular bundles of
i

named species belonP.'?g to the section TruxiUenscs, B. tJfltlullafa, Wall. (rather distinct), B. rNVii,
114ra, Schn., B. v~JCillata, Turcz., B. virescens, Hook. C' I B. virga/a, R. et P.,.JJ. 'lIiIe'lJina, Hl.eroG."
B. WartIt"ana, Schn., B. WfglJtiana, Schn. (in most cases), B'7unnanensu, Franch. .
1 These sp«ies are: BerberiJ appaJtnsir, B. armata, B. conferia, B • .Jle~uosa, B. Jzqrri(ua pro
parte, B.loxhtSiJ, B. multijlDra, B. janiculata, B. Wa/licAiana pro parte, B. Weddell;;~' Ma4iniG
HtuhDe.(ii (Bentb.).
I The following species: Berberis tutl'nacanl/zd, Mart., B. brac/Jy60trya, B. cannattl., B.
~"ilntd$, B. Clauuenii, B. t:t1rWed, B • .[)arrqinit~ B. Darwini-; x emjetriNlia, B. diffNStl, B.
I1!'pelri/~lia, B. feroz, B. ftrrurinea, B. Grirebachii. B. Jzdert'lhylla, B. Iwm'da, B. ilic;'jlJliIJ;
B. inngnis, B. laurina, B. levis, B. litoralis, B. NeCtriana, TiSchI., B. Potanini, B. JruitUlStI..
B. rJltcifolill, B • .ran·cola, B. serrato-dentala, B. spinu/IJra, 8. Valdiviana, B. 'llJ'rga/a; Ma"m;a
tur)'6racteal4, Fedde. M. Fortunei, Lindl., M. F~tmo;'tii (Torr.), M. luumalocarpa (WooL).
M. ./ojtmico, Thunb., M. _poIemis, DC., M. n4nwa, NDtt., M. #Yl)aonl4, Fedde, N. triftl1io1~
(Moric.), these being the species belonging to the sections which Fedde names Longibracteatae and
Horridae.
i C. K. Schneider mentions the occurrence of hypoderm in the following species withont giving
further details: Ber6tris tklinacanllza, Mart. (almost invariably), B. acala/emu, Lechl., O.
/Jarantiana, Vid., B. /wack_y6odria, Gay, B. dli/msis. Gill., B. conf~rla, Klh., B. (uneala, DC.,
B. Danuinii, Hook., B. divaricattJ, Rushy, B. empelrifolia, Lam., B. Gn'jfiJkiana, Scbn., B. HaJlii,
Hieron., B. ilicifoli'o, FOISt., B. i'mienis, Hoole. f. et Th., 8.lauri'na, BjTIbg., B. iimarif()/itI, PbjJ.,
D. liltmzlisl phil., B. loxensis, Benth., B. lutelZ, R. et P., B. paniculala, Juss., B. Pt(J1"ca, Phil.,
B. pedt'nala, Hieron., B. IA~llacanJ/za, Rusby, B. tintit"licensis, Hieron., B. pn"kpotla, Tura:.,
B. rarijllWa, ~cb1., B. t'ecl,nerrn~, Rusby, B. ruscijol.a_, ~rn., B: S(/Jwerin~! SchD., lJ. Sel/tnuianIJ.,
Schn., B. SflUluana, Scho., B. In,pu, Knnze, B. flaldJfJUlna, Phll., B. 1JtJnijlna, Scho., B. verl,·
tillaJa. Turcz., B. virrata, R: et :P., B. WallicAiana. DC., B. JYarsct'ftJi&zii, Hieron., B. Wawrana,
Selm., B. Wellsteiniana, Schn.
• fl~rdiDt: to C. K. SchneJdft, they ~e present in luge ~uzm~ on the opper. side m tbe
Collow~~. I~es only: B~ ___ ns. P.resl, B. ""fStral,.,. Mons var; _Hackelian.a., Scho.,
B. Blnlllen, Scha' f B. crM,!fZ'tuI. DC., B. cntua, L., D. JnlegernmtJ, Bge., B. Aasclzganca, Rupr.
• 1'he- following spec:ieJ: BwlJeril tUi4Ii&IJ. B.lwac/,y60trya, B. carina/a, B. ~lIilmsi.t, B. Clauj·
sen;;, .B,. UIIfnjlJ, B. c~~a.. B. a."ift!s"!, B. ferox, B .. Gtnldolii, B. iliciJolia. B. i,"!gnis l fl· ~/,
B. jwfIlMJd, n. IUIeD/d, B.- ValJi7!UllM; MaAlJ1JIlIJ eurylwadeoJa, M. FlJrluJU., M. ja/MJetl,.
M. IUjIG/e1uU, M. MItTHISII, M. po!yotl#NltI. i. e. species belonging to the section LongibrKte&t&e of
Fedele.
 ADDENDA--BERBERIDE&E 821

the smaller veins are often strongly developed, and in many species of Ber'hetis
and Malwnia cause the veins to be vertically transcurrent.
Regarding the mode of deposition of oxalate of 4iime we may add the
following information. The ,clustered crystals occasionally (e.g. in the pith
of M ahonia lanceolala a..""ld M. pinnata) exhibit a sphaerocrystalline structure.
The prismatic crystals of unknown chemical c.ompositio~discovered by Vesque
have also been observed by Fedde in 1.\1ahonia nepalensis and by Citeme in
Berberis cretica, B. Grisebachii and B, levis.
The glandular hairs of Epimedium have .already beeh mentioned aoove.
Uniseriate hairs, composed of strongly thickened cells (and likewise observed
by Citerne), are present in Epimedium acuminatum· and E. sinense; they
-consist of a few short stalk-cells and a long terminal cell, which is specially
thick-walled and pitted. Structures resembling lenticels are nresent in larpe q
numbers in Berberis Feddeana, Schn., where they cause broWn dotsl·.o£~"tn~r
iower side of the leaf; in other species of Berberis (e.g. B. canaaetiSis,HtlilL):>
they are not abundant, nor are they always present .J ,joH \\'"·~"'~b~~
'
, F or t h e structure 0 f t h e 1eaf -spmes C't "J..rRn~1"jI,i) T":.rh..ft1W
see M"IttmanD:~'d,~t~rf~~,~llu'~~"[~~~C!I
1ler, n. ce. 'c1 M at .1;m~J\tI'l\ .1111;) ,{) :>jr!qJl1~
3. STRUCTURE OF THE AXIS. According to Citerne; the,herQaceous mem-fI
bers ~f the OJ_'der (Achlys, CaulophyUu.m, Diphylleia~"JJ~p..~fne~f1!?t!,i 1}!letsoji!q_~?
Leonttce, Pod.ophyllum) have a normal rIng of vascular bundles ~·tne~suotet~~
ranean axis, bke that in the woody species, In Achlys,··.Dtj/"Yilfra~-'apd',~.fjaq;J
phyllum, besides the normal vascular bundles there are others sItuated in the
primary cortex and composed mainly of sclerenchyma; these cortical bundles
-originate in the scale~leaves clothing the rhizome, and, according to Tischler,
they finally pass into the normal ring of bundles. On the other hap.~l in.. the
subaerial portions of the axis among the herbaceous species there is a t~..Q,q~n~y~
for the vascular bundles to show a scattered arrangement, or to be groupedAlJ..J
two or more rings 1. This anomaly' is specially pronounced in- DiphyUe~'!,d
Leontice and Podophyllum; in Epimed~'um, according to Citeme, there' ar~o
two rings of bundles, whereas, according to Tischler, the presence of two rings .
.can only be inferred; in Achlys Tischler records two rings of bundles, lor a
'Single ring of dovetailing bundles. In the same way the other genera (including
Ranzania according to Tischler) have for the most part vascular bundles of
two sizes, and here also two rings of bundles- may be. supposed to be present.
A typical scattered arrangement of the bundles or their disposition in two
'Or more rings is only found when the number of vascular bundles is sufficiently
larger In the case of Podophyllum Tischler has shown that the inner vascular
bundles are of the nature of leaf-traces. We may add that th~ soft bast in
PodophyllHm, like that of the Monocotyledons, contains only sievertubes and
companion·ceils, and no bast·parenchyma. ,
Our knowledge of the structure of the cortex in the Berbereae has also
been extended. In most cases the pericycle comprises a continuous or inter~
rupted ring of. bast-fibres, or isolated groups of bast·fibres. Pericyc1ic bast-
:fibres (which we may note are frequently septate by means of thi."1, transverse
walls) are only wanting in the subterranean axes of the herbaceous forms
,(with the exception of Epitnedium) and in the subaerial stem of Leont£ce altaica.
In the subaerial axes of Epimedium. and of Nandina, Mahonia Fortunei and
J.f. iaponica, two_zones of bast·fibres are found in the pericycle (Citeme); in
t M ahonia Ehrenbergii, Kunze. the pericycle contains a compoSIte and continuous
Ting of sc1erenchyma (Fedde), as in the Lardizabaleae. In Berberis and

1 Perrot's statement (Tissll cribIe, These. Paris, 1899, p. 14-1) that secondary bundles of wood
and bast occur in the primary cortex of the Lardiza.baleae is incorrect.
 ADDENDA-BERBERIDEAE

M ahonia the bast occasionally includes bast-fibres, which are sometimes

septate or merge into shdtt sclerenchymatous elements. The cork originates
in various positions; either in a layer of the primary cortex (in some cases
the subepidermal layer), or the pericycle. The former has been shown to be
the case in: Berberis ulicina; those species of M ahonia of the section Pani-
culatae, which have been investigated with regard to this character; the
subterranean portions of the axis in Caulophyllum, Diphylle'f,'a, /eOerson'ia
(subepidermal), Leontice, Podophyllum (subepidermal); and Berberidopsis. The
cork arises in the pericycle in most species of Berberis, in M ahonia pro parte~
and in the rhizomes of Achlys and Epimedium.
For the structure of the root in the Berbereae see also Citerne, lac. cit.
Literature: Nanke, Axen bei dikotyl. HolzpR., Diss., Konigsberg, 1886, p. 35.-Mittmann,.
Pfla.nzenstacb'1 Verb. bot-.Ver. Brandenburg, 1889, p .• 54-.-Citerne, HerberlMes et Erytbrospermees,.
These, Paris. 189:J, 161 pp. and 8 pl.-Lothelicr, Epines, These; Paris, I 893.-Schwabach, Bot.
Centralbl. 1898, iv, p. 359.-Spanjer, Wasserappar.l.te, Bot. Zeit., 1898, i, p. 55.-Ho1m,
P~"y/lum. 13ot. Gazette, 1899, pp. 429-31.-Kohne, Anatom. Merkmale bei Berheris.Arten, in.
Wittmack, G a rtenfl ora, 1899. pp. 19, 3, 9 and 68 et seq. j see also Mitteil. deutsch. dendrolog. Gesellsch.
z899, p. 54.-Roedler. Auimilator. Gewebesyst., Disi., Freiburg i. Schw.z899, p. 3T.-Feddc, Mono-
~raphie d. Gatt. Makonia, in Engler, Bot. Jahrb., xxxi, 19°1, pp. 3Q-133.-Pitard, Pen cycle , Th<:!e,
Bordeaux, 1901, pp. 49 and·S3·-Petersen, Vedanatomi, 1901, p. 43.-Simon. in Bet. deutscll. bot.
Gestllsch. 190', p. 238.-Tjschler, Berberidaceen n. PodophyJIaceen, in Engler, Bot. Jahrb. • .xud ..
1903, p. 596 et seq.-Col, Faisceaux, Ann. $C. ont., ser. 8, t. xx, 1904, p. 6\6.-C. K. Schneider.
Btr6eris (EuofftJen's), Bull. de l'Herbier Boissier, s~r. '1, V, 1905. pp. 33 et seq.-Piccioti, Legnami~
Bull. Siena., 1906, p. 169.-[For additional literature, see p. 1169']

NY11PHAEACEAE (Pp.47-Sr).
I

2. STRUCTURE OF THE LEAF. It is characteristic of the stomata of Nym-

phaea, and also of those of Euryale, Nuphar and Victoria, that the closing of the
pore is not effected by contact of the arched ventral walls of the guard-cells,
but by more or less complete approximation of the external cuticular ridges
of the latter (Haberlandt). The branched sclerenchymatous cells (internal
hairs, trichoblasts) found in the genera Nymphaea, Nuphal', Victoria and
Euryale have recently been thoroughly investigated by GUrtler. Their form
varies greatly, even within the limits of the same species. Besides the stellate
IdiobJasts and those of girder-like form showing one-sided d:evelapment and
resembling hairs, other specially noteworthy forms are the H-shaped elements
resembling the spicular cells of the Aroideae and occurring in the peripheral
portions of the petiole and peduncle of Nymphaea. The crystals of oJ;'3.1ate
bf lime on the idioblasts of Nymphaea are sometimes reduced in number;
according to GUrtler, p. 19 (but not quite in accordance with p. 42), similar
crystals apparently occur in Victoria regia also; in Nymphaea and Nuphar
the walls of the tnchoblasts bear pits where they are in contact with the neigh-
bouring Cp.Us. According to Gurtler, the trichoblasts have a purely mechanical
function.
Other structures which may, according to Gurtler, be classed with these
unicellular internal hairs are multicellular intercellular structures resembling
shaggy hairs; they consist of several rows of cells with thick pitted walls,
attain a length of '02-'1 mm., and are terminated by a stellate cell enclosing
a large clustered crystal. These hairs are either unbrancbed or branched,
and in the latter case each branch terminates in a cell with a dustered crystal.
They occur in N elumbittm speciosttm, where they accompany the hair· like
cells containing clustered crystals previously mentioned, and are situated in
the intercellular spaces in the neighhourhood of the diaphragms, which in
this species are specially lacunar; they Serve as a mechanical support to these
diaphragms. Regarded developmentally they are emergences, since they
l.rise from the two outermost cell-layers of the wall of the intercellular canal.
 ADDEND~--NYMPHAEACEAE

According to GUrtler, structures similar to those forms of internal hairs

described as false diaphragms by Treeul (see the previous statements on p. 49)
are also found in Nymphaea capensis and other species, as well as in Euryale
/trox. The hair-like structures found in Brasenia, mentioned and figured in
the earlier part of this work (after Schrenk), are, according to Gurtler. probably
of a pathological nature, since they have not been observed under normal
conditions. Intercellular hairs of a pathological nature are moreover quite
'common in the Nymphaeaceae; for this point, see Gurtler, lac. cit., p. 54-
et seq.
\Vith reference to the systematic value of the air-canals in the petiole
see Conard, loco cit.
3. STRUCTURE OF THE AXIS. According to Van Tieghem and Schoute,
astely is of almost universal occurrence in the rhizomes of the Nymphaeaceae.
NelumbiHm alone has a general endodermis around the central cylinder.
According to Gwynne-Vaughan, polystely (with twelve or more' root-bearing
steles ') is also found in the rhizome of Victoria regia and certain species of
Nymphaea (locally beneath the points of insertion of the leaves).
NOTE.-We may mention that, according to Hewitt, the Nymphaeaccae are
the only Order of Dicotyledons in which the root-hairs arise from special cells,
noticeable as' idioblasts even during the differentiation of the epidermis of the
root, this being analogous to what has been observed in very many Monocotyledons.
Literature: Costantin, Tiges d. pI. aquat., Ann. sc. nat., ler. 6, t. xix, 1 S84, p. 287 et seq. and
pI. 17.-lIaberlandt, Spaltoffn. d. Schwimmpfl., Flora, I 8Sj, p. 102.-Mittmann. PflanzenstachelD,
Verh. bot. Ver. Brandenburg, 1889, p. 63.-Wijnaendts Francken, Sc1ereiden, Diss., Utrecht, 1890,
pp. 32-S.-Gwynne-Vaughan, Morph. and nnat. of Nymphaeaceae, Transact. Linn. Soc. Bot., 1897,
pp. 287-99 and pI. H, 22.-Wollenweber, Anat. d. Schwimmbl., Diss., Freibu.rg i. Br., 1897, pp.
l3-6 and 32, 33.-Weinrowsky, ScheitelOffn. bei \Va.sserpfl., Diss., Berlin, 1898, p. 3S.-Thomas,
Fenilles sout., These, Paris, 19oo.-Masters, Air-canals in the stalks of Nymphaeaceae, Joum. Roy.
Hortie. Soc., xxvi, 1901-:1, pp. 840-3.-Knothe, Unbenettbare BL, Diss., Heidelberg, 1901, p. 16.-
Schoute, Stelartheorie, J903, p. Ils.-Freidenfeldt, Anat. Bau d. Wurzel, Bibl. bot., H. 61, 1904,
p. 5s.-uavitt, Trichomes of the root, Proceed. Boston Soc. Nat. !-Jist., xxxi, 1904. p. 300.-Pilzetti,
Local. dell' alcaloide net Nuphar, etc., Malpighia, 1904, pp. lo6-9,-Conard, Waterlilies, Washington,
1905, pp. 'l7-17.-Giirtier, Intercellulare Haarbild_, Diss., Berlin, 1905, 91 pp.-[Montemartini,
Sisto mecc. della Victoria regia, Atti 1st. bot. Pavia, 1905, 5 pp. and 3 tab.]-[Chiffiot, Anat. camp.
des Barclaya langifalia et B. Mol/leyi, Dull. Soc. sc. nat. Sa8ne-et-Loire, 1906.]-G~nean de
Lamarliere, Membr. cut. des pI. aqu., Revue g~D. de bot., 1906, p. 289 et seq.

SARRACENIACEAE (pp. 5I-54).

Fenner's 1 recent investigations include Sarracem'a flava, L., in which they
deal with the nectarial glands found on the under side of the lid and around the
entrance to the pitcher, the development of the hairs in the • eel-trap-zone,' and
the fourth or lowermost zone of the pitcher, which is regarded as the absorptive
region. The.nectarial glands are of epidermal origin, having a patelliform depres.
sion at the apex and resembling the glands of the pitcher in structure. The absorp.
tive zone has an epidermis of peculiar structure, the cells having undergone division,
though the"~d~.!Ighter-cells -are only bounded by incomplete walls in the form of
ridges. Without further investigation I cannot accept Fenner's interpretation
that we are dealing with simple epidermal cells, which have become segmented
into a number of ' niches' by means of ridges of cellulose and include • several
nuclei:

1 Fenner, Anat' t En tw ickl nngsgesch. u. BioI. d. La ubbl. u. Drusen einiger Insectivoren, Diss.,
Zurich, 1904, pp. 21-7 and Tah. ix and xxi (also Flora., 1904).-See also: Freidcnfeldt, Anat. Ban
d. Wurzel, Bibl. bot., H. 61, 1904, p. 6.~.-[Macfarlane, Sarractnia Calesbaei. Contrib. Bot. Lab.
Philadelphia, ii, 190", p. 426 et seq.)-Forrest Shreve, Sarrarfflia Furpur~a, Dot. Gaz., :xlii, I~.
pp. 118-:13·
 ADDENDA

PAPAVERACEAE (pp. S4-~6).

3. STRUCTjJRE OF THE AXIS. Astely is found in Chelidonium ma1U$
(&ilouteJ.
Literature: E. Schmidt, ?\fi!chr., Bot. Zeit., 1882, p. H:;.-[Tognini, Stomi, Atti 1st. bot.
Pavia, 1894.J-Boert:esen. Arkt. pI. bladbygn., Bot. Tidsskr., xix, 1895, p. 1II9 et se<:j.-8chubert,
Parenchymscheiden, Bot. Centralbl., 1897, iii, p. 4i4.-Millden, Wassersec. Org., Bibl. bot., H. +6,
1899, p. 34.-Tbomas, Feuilles sout., These,l'aris, 1900.-Molisch, Milchsaft u. Schleimsaft, 1901,
p. 71 et seq.-Knotbe, Unbenetzb. m., Diss., Heidelberg, 1902, p. l .....-Schoute, SteUirt1]eorie, 1903,
p. II6.- Theolin, Vaxttrichon1., Arkiv {or Bot., i, 19°3, p. 17f.-Kniep, Bedeut. d. Milchsaftes,
:Florn, 1905, p. 176 et seq.-Mayus, Milchr. in Qen m., Beih. bot. Centralbl., xviii, Abt. I, 19()S.
pp. '76-8.-Sarton, Rech. expo sur l'anat. d~s pI. affines, Ann.·sc. nat.. sCr. 9. t. ii, ]905, pp. 25-3"
(Cnelidor.it(m).-[Faltis, Opinmalkaloide, Pharm. Post., 1906, n. ~p, 32; abstr. in Bot. Centralbl.,
civ, p. 61.]

FUMARIACEAE (pp. 56-58).

Zsak Zoltan has recently met with 'unicellular fir..ger-shapcd trichomes in
specimens of Corydalis cava and C. solida, two species which he regards as varieties.
Literature: Costantin, Tige:s aeroet SQnt., Ann, sc. nat., ser.6, t. xvi, I88~, p. 96 et ~.-Jost,
Zerkliift. einiger Rhiz. n. Wurz" Bot. Zeit., 1890, p. 469 et seq. (Co1j'tklis).- Thomas, Feuilles !iOtlt.,
These, P:ll"lS, J90o.-Knothe, UnbeJletzb. Bt, Diss., Heidelberg, J902, p. 15.-Zsak ZoJtan, in Magyar
Bot. Lapok, 19°4. 4- pp.-[Haberlandt, Sinnesorg., ii Aufi., 1906.j

CRUCIFERAE (pp. 58-67).

2. Schweidler's recent investigations ~n the my rosin-cells found in the
foliage-leaves of the Cruciferae contain the following statements, which w€ may
add to the information given on pp. 60-62. The earlier statement (p. 60) regard-
ing the absence of chlorophyll~grains in the myrosin-cells i;; incorrect, since
Schweidler found chloroplasts in the secretory cells of the assimilatory tissue
in those species which hI';: subjected t.v a detailed examinahon. According to
Schweidler, the local distribution and the differentIation of the idioblasts in the
sepals, petals and the valves of the pod are essentially the same as In the foliage
le.aves. The presence or ~bsence of idioblasts is probably a character of generic
value. In those cases.: in which no myrosin-cell.;; have been found in certain
species of a genus generally possessing .~hese elements, this is probably due
to a reduction of the secretory cells, which has hitherto rendered it impossible to
recognize them. CJoselyrelated species possess albumi1;.ous cells agreeing in their
..)ocal distribution. size, shape, &c. Schweidler distinguishes between 4 idioblasts
. of the vascular bundles,' which are mostly prosenchymatous, and' ldioblasts
of the mesophyll/ which are parenchymatous elements containing chlorophyll
a11d showing little or no difference from the ordinary cells of the IDesophyll; he
subdivides the idioblasts of the. vascular bundles into 4 idioblasts of the phloem-
sheath J and • idioblasts of the parenchyma~sheath.' According to the same
authority another point of systematic im,ortance is t6 be found in the prcs~nce
or absence of spetial albuminous contents in the guard~cells of the stomata
in those ~pecies, which have no idioblasts.
Th~ genus Arabis has been made the s.ubject of detain:;d _inve~tigation by
Schweidler. his results being as fellows. In the species of the section Turrf,'tz's, L.
th~ idioblasts a.re present in the phloem-sheath only. in the species of the section
Cardamt'nopsis, Boi~s. they occur both in the phlot:ro-sheath and iIi..the pCI..L'enchyma-
sheath~ in those of the sections Pseudarabt's, C. A. Mey. and Turritella, C. A. Mey. the
idiobiasts are situated in the mesophyll, and in these of the section ElI,af'tibis there
are only special albuminous content.:. in the guard-cells, whilst in some few species
no idioblasts were recogni.zable, In consequenc.e of these re~ults (l.l)d :!rom a con-
sideratior.. of ether char'act~rs Schweidler comes to the conclusion that the genc.s
 ADDENDA--CRUCIFERAE
.Arabis must be provisionally restricted to the sections Eflarabi~, Pseudarabis and
Turritelia, whilst the section Cardaminopsis should be included in Cardamine, and
Turt"itis raised once more to the ra.nk of an independent genus.
The same featmes are moreover of va\ue in connexion with the subdivisiill
of the Cruciferae, if one studies the affinities of the genera having ,idioblasts in 't1ie
mesophyll only (Exo-idioblastae), or idioblasts in the vascular bundles only (Endo-
idioblasu.. ',or both kinds of idlOblasts (Hetero-idioblastae). .
The Exo-idioblastae include the following genera: L1mat'ia, Vesicar£a, Schive-
rec1~ia. Peltaria, Petrocc;,llis, Draba, Cochlea ria (Alyssineae); T}tlaspi, Teesdalia
(Thlasp1deae); Anastatica (Anastaticeae); Cakile, Chorispora (Cakilineae); Gold-
bachia (Anchonieae); Brassica, Sinapis, Moricandia, Dz·plotaxis, Eruca (Brassiceae) ;
Vella, Carrichtera, Succovia (VeUeae); Crambe, Rapistrum, Raphanus (Raphaneae) i
Bunias (Buniadeae); Braya, Arabis sens. strict_ (Arabideae); Al/.1,·ar£a, Conringia
{Sisymbrieae). The Endo-idioblastae include: Cheiranthus, Nasturtium, Barbarea.
TU'Y1'itis, Arabis Sect. Cardami'nopsis, Caraamine, Del1,taria (Arabideae); Maleolmia,
Hesperis, Sisymbrium, Erysimum (Sisymbrieae); Camelina (Camelineae); Sene.
biera, Lepidium, Aethionema (Lepidineae); Capse!la, Biscutella (Thlaspideae);
Heliophila (Heliophileae). The Hetero-idioblastae include: l.beris; Ltpidium
D,aba; lsalis Myagrum (Isatideae).
Mardner also mentions the occurrence of myr03in-cdls in Pringlea anti~cor·
tutina, R, Br. '
According to Bouygues, the petiole of the Cruciferae is distinguished by
the presence of 'faisceaux rayonnes.' which have a characteristic mode of
development. The vascular bundles of the petiole are arranged in a rjng,
and are embedded in a parenchymatous tissue, the cells of which differ from
the parenchyma of the ground-tissue in having smaller lumina and thicker
walls. The entire ring of vascular bundles, together with the small-celled
parenchyma, develops from a aingle procambial strand, in which secondary
procambial bundles become differentiated and give rise to the individual
vascular bundles.
Regarding the anomalous strands of soft bast in the rhizome and root of
Cochlearia Armo,acia, see also Viret. loco cit.
LiteratuT~: Costantin, Tiges aero et 8oUt., Ann. se. 'nat., ser. 6, t. xvi, 1883, p. 98 et seq.-
COStantin, Tiges aero et souL d. pI. aql1at., Ann. sc. nat., Ilt~T. 6, t. xix, I SS4, p. :287 et seq." and pI.
1".-I6.-[Tognini, Stomi, Atti 1st. bot. P:..via, I 894.]-Boergesen, Alkt. rl. bladbygn., Bet.
Tidsskr., xix, 189.s~ p. 2J9 et seq.-Schubert, Parenchymscheiden, Bot. Centrillb . 189" iii, p. 475.-
Le~ring. Interxyl. Leptom. Diss., Berlin, 1899. p. :28.-Anheisser, Arunkoide Blattspreite, Diss••
Jena, 1900, pp.' 18, 19.-Paulesco, Stnlct. anat. des hybrides, These, Geneve, 1900, p. 63 (Den/aria).
-Scbleichert, Xerophyt::n bei Jena, Naturwiss. Wochebschr., 1900, P.446 (Thlaspi).-Thomas:
Feui!les sout., These, Paris, 1900.-Veba, Anat. d. Axen von Alyssum calycinum, Oesterreich. hot.
7..eitschr. J901. pp. 225-33.-Bo'Cygues, Petiole, Thbe, Paris, I(j02, p. 31 et seq.-Clanditz, Blattal'lat•
.canar~ Gew., Diss., Ba!>el, 1902, pp. 57. 58 (Chei,.anthus l).-Knothe, Vnbenetzb. Bl., Diss., Heidel·
berg, J90l, p. Is.-Mardner, Phan.- Veg. der Kerguelen, Di.ss., Basel, 1902, pp. 8-14 (Prir.gle?J).-
[Armari, Pi:lDte della reg. medit., Ann. oi bot., i, 1903, p. 17 et seq. (.lberis).]- Theann, Vaxttrichom.\
Arkiv for Bot., i, 19()3, p. 170 and iv, n. 18, 1905, pp. 17, 18.-[Boodle, Wallflower, New Phytol.a
iii, 1904, p. 39; abstr. in Hot. Centralbl., xcv, p. S04.J-Chrysler, Strand plmts, Bot. Gazette, xxx\ii,
19°4, p. 461 et seq. (Cakile).-Slissenguth, Bt:h.-Verh. d. Wiir:1:b. Muschelknlkpti.., Diss., Wiirzburg,
1904. PJl.. 19, ~~.:--:-Viret. Liais~n$ ?u phloeme rned. ~'. lIi.stifut bot. Geneve; 1904, pp. 36-4~'
-Schw~idlet, Elwelssi:ellen d~ ,Cruclf., Ber.J.leutsch. bOt. Gesellscb' J 19'J5, pp. 274-85 and TaC. XII.
-[For additionalliterature,"gee p. Jl70.]

CAPPARIDEAE (pp. 67-77).

I. Under the general REVIEW OF ANATOMICAL FEATURES we may adp
that crystalloids are occasionally present (in cells of the ground-tissue and
epidermis of the petiole of Boscia) in the form of smail rhombohedral; octahedral
or roundeti crystalline bod.i~s.
l The statement that gl:ddular hairs a:-e present in C/l(£'ranthus s<'tJjarius appenrs questionable
tomt'.
 ADDENDA--CAPPARIDEAE
2. STRUCTURE OF THE LEAF. In connexion with a monographic revision
of Boscia Pestalozzi made a thorough examination of the structure of the
leaf in this genus. The most characteristic feature of the genus is the occur-
rence in the mesopbyU of sclerencbymatous cells elongated in the same direc-
tion as the palisade-tissue; these elements are situated beneath the epidermis
on both sides of the leaf, and where they come in contact with the epidermis
they exhibit either a bulbous (B. angustifolia, Rich.) or a somewhat T-shaped
(B. rotundijolia, Pax, &c.) enlargement, or a more or less abundant ramification.
The thickness of their walls varies greatly. In some cases these elements,
]ike the sclerenchymatous cells of certain species of Capparis, may even pene-
trate between the epidermal cells, and thus form part of the surface of the
leaf. The structure of the leaf in Boscia is bifacial or centric; the stomata
are either restricted to the lower surface or are found on both sides of the
leaf. No hypoderm is present. On the other hand storage tracheids (de~cribed
by Pestalozzi as 'sclereids with a lobed form ') occur in the spongy tissue.
The vascular bundles of the smaller veins are ,provided with hard bast and
are embedded. The hairy covering consists only of unicellular clothing hairs
of varied length; in B. corymbosa, Gilg. these hairs appear like finger-shaped
papillae. For the remaining features of the leaf-structure see the work cited
below. Large solitary crystals of oxalate of lime·(cf. Syst. Anat., p. 7I) also
occur in the axis of Cadatm; they are situated in the stone-cells of the pericycIe.
The thorns of Capparis spinosa, which are interpreted as stipules, contain
neither vascular bundles nor terminations of bundles (Lothelier).
3. STRUCTURE OF THE AXIS. The cork in Cadaba glandulosa, Forsk.· also
arises subepidermally. In this species the primary cortex contains groups of
stone-cells, and the pericycle exhibits a composite and continuous ring of
sclerenchyma.
, Litl!rature: Wijnaendts. Francken. Sc1ereiden, Diss .• Utrecht, 1890, pp. 50, 5I.-Lotbelier,
Epines, These, Paris, IS93. pp. 35 and 46.-Pesta.lozzi, Bosda, Bull. de l'Herbier Boissier, 189 8 ,
App. iii; also Diss., Ziiriq,b, 152 pp. !lnd 13 pl.-Vrs-pung, Cadaba glandulosa, Ber. deutsch. bot.
Ges., 1901, pp. 501-8 and Tab. xxix.-[Moll and Janssonius, Mikrographie d. Holzes, Leiden, 1906,
pp. 175-92 (Capparis, Cmtacz'a).)-lHoltermann, Einfluss d. Klimas, 1907, p. 106 (Capparis,
Naerua).]

RESEDACEAE (pp. 77-79).

l\Iorstatt's recent investi~ations deal with only a few species of Reseda.
The discovery of oxalate of hme in the form of small crystals in the pith of
Reseda odorata is specially noteworthy.
Other points of importance are as follows. Large vesicular papillae with
transitions to simple unicellular blunt hairs are borne by the epidermis of the
stem in R. lutea; on the leaf the papillae are- restricted to the margin. The
pericycle in the stem of Reseda includes groups of bast-fibres. The structure
of the wood in Reseda exhibits the same features as in Ochradenus.
The basal appendages (' denticuli basilares ') of the leaves of Reseda
exhibit various types of structure; in R. lutea they have a distinct pal~sade­
like epidermis, which secretes mucilage; in no case do they contain a vascular
bundle.
Literature: J. Milller, Monogr. de la famille des Resed., ZUrich, 1857, p. I I ct seq. and Tab.
i-iii.-[Tognini, Stomi, Atti. 1st. bot. Pavia, 1894.]-Morstatt, Beitr. z. Kenntn. d. Resed., Diss .•
Heidel~rg. J903. 64 pp.-Col, Fa.isceaux, Anll. sc. n:lt.;ser. 8, t. xx, 1904, p. 1"9.-SUsseoguth.
Rebaarungsverb. d. Wiirzh. MuschelkalkpR., Diss., Wi.irzburg, 1904. p. lO.
 ADDENDA 82']

CISTINEAE (pp. 79-82).

Withrefeq:nce to the hairy covering (p. 80) we may add, on Siissenguth's
authority, that tufted hairs like those found in Cistus creticus (Syst. Anat.,
Fig. 21, B) also occur in H elianthemum nanum, but not in H. vulgare and H.
Polifolium.
Regarding the structure of the axis (d. p. 82) the following data may be
quoted from Piccioli's investigations, which deal with the four genera of the
Order. In the wood, medullary rays are not present in Lechea, whilst in the
remaining genera they are narrow and mostly uniseriate. As a rule the bulk
of the wood is composed of prosenchyma with bordered pits; in the species
of Helian!hemttm belonging to the section Eriocarpum, however, the tracheids
are replaced by mechanical elements. \Vood·parenchyma only occurs in
relatively small amount. The region in which the first layer of cork is pro-
duced varies, being the epidermis in Cistus and Helianthemum § Lecheoides,
the subepidermal layer of cells in Lechea, H elianthemum § H alimium and
E1thelianthemum, a more or less deeply situated layer of the primary cortex
in Helianthemum § Macularia, Brachypetalum pro parte, Eriocarpum, Fumana·
pro parte and Pseudocistus, the endodermis in § Brachypetalum pro parte, and·
lastly the cell-layer situated on the inner side of the bast-fibres of the pericycle
in Budsonia and § Fumana pro parte. The endodermis is seldom distinctly
differentiated; it constitutes aqueous tissue in Helianthemum politolitem and
H. salicifolium. The outer portion of the primary cortex is often collen-
chymatous. The pericycle in species of all the genera includes a ring of fibres,
or isolated groups of bast-fibres; in certain species of Helianthemum the bast-
fibres are scanty or absent. In H. gutta!ttm the pith is differentiated as aqueous
tissue. It remains to mention that in all the genera oxalate of lime occurs.
in the form of clustered crystals; and that additional solitary crystals have
only been observed in different parts of the axis in the species of Helianthemum
belonging to the section Eriocarpum (H. kahiricum and H. sessiliflorum).
Literature 1 : Gauchery, Hybrldes dans Je genre Cls!us, Assoc. (rant. Besanyon, IS93, i, p. 238,
and ii, pp. 534-4I.-Schubert, Parenchymscheiden, Bot. Centralbl. 1897, iv, p. 16.-Paulesco, Street.
anat. des hybrides, These, Geneve, 1900, p. 75 (Cistus).-Clauditz, Blattanatomie canar. Gew.,
Diss., Basel., 1901, pp. 12, 13 (Ct·stus).-Gauchery, in Assoc. fran y• Ajaccio, 1901, ii, ed. 1902 , pp.
4°8-13 (Htlian/hemum "alimifolz'um x Cls/UJ sal1;ifo/ius).-Petersen, Vedanatomi, '90J, p. 43.-
Pitard, Pericycle, These, Bordeaux, 1901, p. I02.-Grosser, Cistaceae, in Pflanzenreich, H. 1-4, 1903,
pp. 3-4.-Piccioli, Legno e carteccia delle Cistin., Nuov. Giarn. bot. ItaL, N. S., xi, 1904" pp. 47:1-
50.f.-Siissenguth, Hehaarnngsverh. d. \VUrzb. Muschelkalkptt, Diss., Wtirzburg, '9°4, pp. 20-22.-
Sarton, Anat. d. pI. affines, Ann. sc. nat.. ser. 9. t. ii. 1905, pp. 39-43 (Heliant}umum).-Piccioli,
!.egn ami, Bull. Siena, 19Q6. p. 167.-l For additional Iiterll:ture, see p. 1169.]

VIOLARIEAE (pp. ,82-86).

SECTION 2. _ An- undetermined Peruvian species of Viola (Viola 62) inve:,ti.
gated by Weberbauer has a papillose epidermis on the upper side of the lea.f;
on the same side there is also a strongly projecting network formed by strands
of tissue containing 2-3 layers of palisade-cells showing a fan-like-Iarrangement.
With reference to the teeth on the margin of the leaf (p. 85) we may ddd
that they secrete lime in some species of Viola (e.g. V. scan dens, Willd., accord-
ing to Triana et Planchon, Prodr. Flor. Nov. Granat., Ann. sc. nat., ser. 4,
t. xvii, 1862, p. 121, and V. Lindeniana, Turcz., Tonduz, n. 2123, Costarica).

I Gerber's paper (Et. anat., phys. et bioi. sur les Cistes de Pro\'ence, Annuaire Fac. Sc. Marseille,
1899) only deals with the reproductive organs.
 ADDENDA-VIOLARIEAE

SECTION 3. According to Pitard, a composite and continuous ring of

sclerenchyma in the pericycle is also developed in 'I onidium salidfolium'
and species of Alsodeia.
Literature: G. Kraus, Inulin !lei Viol., Sitt.-13er. naturf. Gesellsch., Halle, 1880, p. 6 j ~ also
Fames, in Pharm. Journ. and Transact., 1884, p. 515, and Beauvisage, in Bull. trimestr. Soc. bot. de
Lyon, J888, pp. 12 and 39.-Costantin. Tiges aero et sout., Ann. sC. nat., SeT.6, t. xvi, 1883. p. 10:3
et seq.-Schubert, Parenchymscheiden, Bot. Centralbl. 1897, iv, p. 17.-[Hartwich, Falschc
Ipecacuanhawurz., Schweize~. \Vochenschr. f. Chemie etc., 1899, n . .f8; abstr. in Just, 1900, ii,
p. :18, Inulin n.-Paulesco, Struct. anat. des hybrides, These, Geneve, 1900, p. 70 (Viola).-Pitard,
Pericycle, These, Bord~nx, 1901, p. 70.-Clauditz, Blattanat. canar. Gew" Diss., Basel, 1902.
pp. 55. s6.-Freidenfeldt, Anat. Ban d. Wurzel, Bibl. bot., H. 61, 1904, pp. 61-3.-Sarto:l, Rech.
expo sur l'anat. Ges pl. affines, Ann. IC. nat., ser. 9, t. ii, 19°5, p. 86 (Piola).-Siissenguth,
Behaamngsverh. der Wurzb. Muscbelkalkptl., Diss., Wunburg, 19°4, p. l:il.-l'beorin, Vaxttri~h<;\III.•
Arkiv for Bolo, iii, n. 5, 1904, p. 18; see also loco cit., iv, n. 18, J905, p. I 4.- Weberbauer~ V~get.
der Hocbanden Pems, in Engler, Bot. Jahrb., uxvii. 1905, p. 88.-[MoIl and Janssoninj, Mikro-
gr;)phie d. Holzes, Leiden, i, 1906, pp. 193-7 (A!sodeia).] ,

CANELLACEAE. (pp. 86-87).

2. STRUCTURE OF THE LEAF. In Canella alba, P. Browne, C. obtusi/olia,
Miers, and Cinnamodendron macranthu11t, Baill. the mesophyll contains no
typical palisade-tissue. According to Parmentier, clustered crystals are found
in the epidermis, where they occur in every cell in Canella alba, in ~ach cell
of the upper epidermis in Cinnamodendron macranthum~ and in small
epidermal cells in Cinnamosma /ragrans. In Canella ·obtusifolia and Cinnamo-
dendron macranthum the stomata are only present on the lower side of the leaf,
and are accompanied by subsidiary cells placed parallel to the pore. In -all
three genera the vascular bundles of the veins are strengthened both above
and bel~w by groups of fibres. Iri Cinnamodendron also three vascular bundles
-pass iI;to the leaf.
3. STRUCTURE OF THE AXIS. Regarding the structure of the cortex we may
4dd that In Canella obtusifolia, Cirmamodendron macranthtem and Cinnamosma
~',ugrans also the cork develops in the subepidermal layer of cells. In CaneUa
only are-. the U-sbaped sclerosed cells present in the phelloderm j this tissue is
not icl~~eJ in Cinnamodendron macranthum, and in Cinnamosma /ragrans
&Sclerosis 1s only local. In all three genera the pericycle includes bundles of
fibres. ~ich in some cases only develop at a later stage.
Lite~tute: [Greenish, Canelfa bark, Pbarm. Joum. and Transact., xxiv, 18 93-4, pp. 793-7.)-
Parmentier, in Giard, Bull. scient. de la France et de la Belgique, xxvii, 1895-6. pp. 3I5-18.-Bler-
mann, Oelzellen. Diss., Bern, 1898, pp. 29. 3o.-Van Tieghem, Canellacees, Journ. de Lot., 1899, pp.
~66-76.-[Courchet, Cinnamosma, Ann. lnst. Marseille, 1906.]

BIXINEAE (pp. 87-(1).

I. To the REVIEW OF ANATOMICAL FEATURES we may make the following
additions. In the four genera of the Cochlospermeae the structure of the bast
is like that of the lime and shows the same character in the primary cortical
medullary rays, i.e. they broaden outwards like a wedge; the development
of cork besides being subepidermal may also be _epidermal and (Aphloia) peri~
cyclic; groups of soall epidermal cells containing crystals are also found in
Camptostylus~ Erythrospermum, [toa, Rawsonia, and Scottellia; mucilaginous
epidermal cells in the leaf are also met with in the genera Amoreuxia and Mar-
quesJ:a; mllcilage·cells (not mucilage·canals) occur according to Van Tieghem in
Sphae1'()sepalum; simple unicellular clothing hairs provided with blunt pro~
tuberances are found in Hoplestigma; tufted hairs also in Marquesia; peltate
hairs also in Camptostylus and Cochlospermum; glandular hairs, differing in
structure from those of Oncoba and Poggea, in Hoplestigma and !Jarquesia;
 ADDENDA--BIXINEAE
wart·like emergences on the petiole in Scolopia; hypoderm in the leaf also in
species of Itoa and Scottellia; spicular fibres in the mesophyll also in Centro·
placus paniculatus, Gilg (where their lumina are often filled with silica) and
Erythrospermum amplexicaule, DC.
2. STRUCTURE OF THE LEAF. The spicular fibres of Centraplacfts and
E1ythrospermum mentioned above were observed by Brandlein and Citerne
respectively. The sc1erenchymatous fibres of Centroplacus paniculatus branch
off from the sc1erenchyma of the veins, and sometimes show ramification
themselves; they are specially distinguished by the frequent presence of
amorphous silica filling their wide lumina. Mucilaginous epidermal cells
also occur in A moreuxia palmatifida, Mo~. et Sesse, M arquesia macrura, Gilg,
Neumannia deltoides, Warb., N. minima, Warb. and N. theaejormis, Rich.
(speci~ of Aphloia). A hypoderm of one or several layers is found beneath the
upper epidermis in Erythrospermum amplexicaule, DC., E. amplifolium, E. coro·
narium, E. laxi/lorum, E. phytola,fcoides, E. pyrijoli'Hm, E. tetrasepalon, E. verti·
tiUatum, Lam., Itoa orientalis, Hemsl. (locally), and Scottellia macropus, Gilg
et Dinkl., while a single hypodcfmallayer is situated above the lower epidermis
in Erythrospermum amplexicaule and E. verticillatum (Brandlein, CHerne and
Van Tieghem). The small crystal.idioblasts in the epidermis are recorded
by Citerne in Erythrospermum (with clustered crystals) and by Brandlein in
Camptostylus caudatus, Gilg (with clustered and solitary crystals), Itoa
orientalis (dust. cryst.), Rawsonia Schlechteri, Gilg (clust. cryst.), and Scot·
tellia macropus (elust. and solit. cryst.). A stomatal apparatus of a
distinctly Cruciferous type is found also in Centroplac'Lts and Rawsonia.
whilst the Rubiaceous type is clearly developed also in Carrierea, Itoa and
Trichostephanus; in the remaining genera investigated by Brandlein only
ordinary neighbouring cells are present. The stomata of Sphaerosepalum
alternilolium, Bak. occur on both sides (If the leaf, and are provided with
subsidiary cells (Van Tieghem).
In Bixa, Erythrospermum and Sphaerosepalum the vascular bundles ui
the veins are accompanied by sclerenchymatous fibres, but this is not the
case in A moreuxia and Cochlospermum. One or three vascular bundles pa~
out into the leaf. In the genera Bixa, Cochlospermum (Mttximilianea). A1#O·
reuxia and Sphaerosepalum (which was formerly included among the Cruttiferae)
-all four members of the Cochlospermeae-the three bundles, which enter th,.
petiole, sooner or later unite to form a rfng, and in Bixa the latter encloses
an arc of wood and bast with the xylem directed upwards. In Bixa the
vascular system of the midrib is the same as that of the petiole (this is contrary
to the earlier statement on p. 89). In the Flacourtieae, on the other hand,
there is either one vascular bundle (Dovyaz",'s, Flacourtia, Ludia, Seolopia,
Xylosma, &c.), or three (Azara, Idesia, Tisonia, Trimeria, &c.) which unite
to form an arc open on its upper side; in this tribe the vascular system nowhere
forms a ring. In the genus Aphloia (Neumannia), which in its anatomy departs
in some respect~Jsee -the development of the cork) from the Flacourtieae, three
vascular bunClles (one large and two small) pass out from the axis into the
petiole. In the same way three vascular bundles also occur in ErythrocMton,
but a modification is introduced inasmuch as the bundles in this genus
ultimately become concentric in structure.
In connexion with the subject of oxalate of lime (p. 89) we may mention
that Fabricius' recent statement as to the occurrence of cystoliths in a member
of this Order (Aphloia madagascariensis) is incorrect. The plant examined by
Fabricius does not belong to the Bixineae, but was a specimen of A'rtocarpus
integrijolia, L.f. (Urticaceae).
The most important addition to the previous statements on internal
secretory organs is as follows. In Sphaerosepalum alternifolium, Bak., Van
 ADDENDA-BIXINEAE

Tieghem only met with mucilage-cel,ls in the primary cortex of the branch, in
the parenchyma of the petiole, and in the mesophyll; thi~ is not quite in
agreement with \Varhurg's earlier observations on the occurrence of mucilage-
canals.
The secretory cells are also present in Amoreuxia palmattfida, MoC;. et
Sesse and A. ~mipora, Van Tieghem. They are quite generally distribut~d
in the outer portion of the primary cortex of the branch, in the parenchymatous
ground-tissue of the petiole, and in the mesophyll. The mucilage-can,als actually
exhibit lysigenous development, and are also found in the petiole and in the
larger veins of the leaf; in the primary cortex they o.ccupy the inner portion.
A central, medullary mucilage-canal has only been recorded in the axis of
Bixa Orellana and A moreuxia unipora.
To the section dealing with the hairy covering w~ may firstly add that
simple unicellular clothing hairs have been met with in Carrierea, Hoplestigma
(where they bear the protuberances mentioned above), ltoa and Marquesia;
in the last of these genera the simple hairs are accompanied by tufted hairs
(Brandlein~. Peltate hairs are described by Van Tieghem in Cochlospermum,
and by Bdindlein in Camptostylus (in this genus they have a short stalk and
thin-walled ray-cells). The glandular trichomes of Hoplestigma Pierreanum,
GiIg~ which were likewise observed by Brandlein, ,are uniseriate and in some
cases of great length; they consist of a few basal cells with relatively thick
walls, followed by several longer cells with thin walls, and a rounded terminal
cell, which is not sharply marked off, and has thin walls. According to the
same authority, the glandular hairs of Marquesia macrura have a short stalk
and an obliquely inserted head, the latter being multicellular and having both
horizontal and vertical division-walls. According to Areschoug, the wart-
like emergences~ mentioned above, found on the petiole in Scolopia and having
the function of hydathodes, include an epithema and the termination of
a vascular bundle.
3. STRUCTURE OF THE AXIS. The structure of the cortex has recently
been investigated in detail by Van Tieghem, more particularly in the genera
of the Cochlospermeae, as well as in Aphloia (Neumannia) and Erythrospcrmum.
In Bixa, Amorcuxt'a t Cochlospermum, Sphaeroslpalum and Erythrospermum the
pericycle contains isolated bundles of bast·fibres, whilst in the Flacourtieae,
according to Van Tieghem t though not quite in accordance with the earlier
statements of Harms (see p. 87), a composite and continuous ring of scleren-
chyma is quite generally developed. A similar ring is found in the pericycle
in the species of A phloia, and according to Pitard also in X ylosma nitidum
(where it includes cells with U-shaped thickening), and in a plant described as
Rumea coriacea (= .)(ylosma nitidum ex syn.!) (with ordinary stone-cells).
A distinct endodermis composed of relatively large cells with Caspary's
dots on the radial walls is present in Aphloia only. The bast has a characteristic
structure in four genera of the Cochlospermeae, viz. Bixa, Amoreuxia, Coch/o-
spermum and Sphaerosepalum. The groups of phloem are narrowed outwards
in the form of a wedge, while the primary medullary rays become correspond-
ingly broader in the same direction; moreover the groups of bast show the
same characteristic stratification into hard and soft bast as is seen in the lime.
A similar broadening of the medullary rays outwards, and a narrowing of the
phloem-groups between the rays is also more or less marked in Erythrospermum,
but no secondary hard bast is present. In Aphloia sclerosis of the bast only
occurs in the older branches. Bast-fibres occurring singly or in groups are
formed from the thin-walled tissue in the outer portion of the phloem, whilst
the tissue of the primary medullary rays between the phloem-groups undergoes
sclerosis as far as the boundary of the xylem. Consequently the newly-formed
soft bast appears in a transverse section in the form of nests of tissue corre-
 ADDENDA--BIXINEAE
sponding to the individual vascular bundles. In the typical Flacourtieae,
according to Van Tieghem, there is neither stratification of the bast into hard
and soft bast, nor any wedge-like broadening of the primary cortical medullary
rays outwards. The cork ari~s in the epidermis in B~'xa a'P.d Amoreuxia, in
the outer cell-layer of the primary cortex in Cochlospermum, Sphaerosepalum
and Erythrospermum, but on the inner side of the pericyclic parenchyma in
Aphloia. The cells of the cork are flat and have thin walls in Bixa; they are
flat and somewhat thickened in Erythrospermum, fiat and provided with
slightly thickened tangential walls in Aphloia, and so on.
Literature: Citerne~ l3erberldees et Erythrospermees, These, Paris, 1891, pp. 104 and 127.-
Briquet, Hydathodes fo1. des Scolopia, Bull. Herb. Boissier, 1898, pp. 503, 504.-Van Tieghem,
Neumannie, Jauro. de bot., 1899, pp. 361-7.-Van Tieghem, ~ixacees, Cochlospermacees et
Spherosepalacees, Journ. de bot., 1900, pp. 31":S4.-Van Tieghem, Erythrospenne, Joum. de bot.,
1900, pp. 1 2 5-9.-Pi tard, Peri cycle, These, Bordeaux, 1901, p. 68. -Areschoug, Maogrovepfl., Bibl.
bot., H~ 56, 190~, pp. 61-3, and Tab. vi.-Fabricius, Laubblattanat., Beih. Bot. Centralbl., xii, 1901,
pp. 3J7, 318.-Solereder, Zwei Bericht., Bull. Herbier Boissier, 1903, p. 318 et seq.-[Rippa,
Nuovo genere etc. (Licopolia), Bull. Orto bot. Napoli, ii, 1904, p. 69 et seq.]-Areschoug, Trop.
vaxt. bladbyggn., Sv. Vet. Akad. Handl. 39, n. 2, 1905, pp. 103-5 ETaraktogmos), and pp. Il6-18
(Ryjarosa).-Brandlein, Syst.-anat. Untersuch. d. Bl. der Samydaceen, Benth.-Hook., Diss.,
Erlangen, :906, Manuscript (Camptostylus, Carr£ena, Cenlroplacus. IIojlestigma. floa, Marquesia,
Rawsoltia, Scottellia, Trichostepham(s).-[For additional literature, see p. 1169.J

PITTOSPOREAE -(pp. 91-94).

Literature: Pitard, Peri cycle, These, Bordeaux, 19°1, p. 4'J.-Hooper, Supposed Beilsdwziedta-
bark, Pbarm. Joum., 1904-, p. 361 et seq. (According to ~he, results of my own investigation this
bark belongs to Pittosporu11l.)-[For additional literature, see p. 1I7I.]

POLYGALEAE (pp. 96-roo).

Regarding the anatomy of the saprophytic genus EpirrM2anthes we may note
that the scale-leaves have no stpmata. unicellular clothing hairs are present, and
the vascular cylinder of the stem is enveloped by a ring of fibres.
Literature: Knoblauch, Oekolog. Anat. etc., Habilitat. Schr., Tiibingen, 1896, p. 13 It
seq.-
Penzig, Epirl'ltizanthcs, Ann. Jard. Huitenzorg, xvii, 1901, pp. 141-70, and Tab. xx-xxvi.-[For
additional literature, see. p. lJ7I.] -

VOCHYSIACEAE (pp. 100-104).

3. STRUCTURE OF THE AXIS. Leisering thinks it probable that the inter·
xylary phloem, which is also found in Erisma nt'tidum, DC, is developed by
subsequent differentiation of sieve·tubes from parenchyma given off internal1y
by the cambium.
Literature: Johannson, Noch wenig bek. Rinden, Diss., Dorpat, I89T, pp. 15-17.-Leiserinc,
Interxylares Le'p,!_~~, Diss~.t.-~~lin, ~~99, PP_,__45, 4 6 .

CARYOPHYLLEAE (pp. r07-III).

I. The REVIEW OF ANATOMICAL FEATURES requires the following additional
remarks. The structure of the stomatal apparatus is not the same in all the
members of the Order, the Caryophylleou5 type with two cells placed trans-
versely to the pore being rare (Loe{lt'ngia) in the representatives of the tribe
Polycarpeae. Solitary crystals of oxalate of lime probably only occur in
exceptional cases (axis of Polycarpaea filifolia, Webb). Clothing hairs of a type
not previously mentioned are branched sympodial trichomes with a varying
number of rays (species _of Cerastt"wn, Polycarpaea, PolYfarp01L and Stipulicida).
 ADDENDA-CARYOPHYLLEAE

The occu_rrence of secondary zones of growth has also been recently demon-
strated in the axis of numerous species of Polycarpaea, as well as in the root
of certain species of Cerdia, Ortegia, Polycarpon, Pycnophyllum,. Spergula,
SPergularia and Stipulicida.
2. STRUCTURE OF ¥IE LEAF. A not uncommon feature in the Polycarpeae
(Loef/ingia, Ortegia, PMycarpaea, Polycarpon) is the presence of longitudinal
rows of epidermal cells with vesicular papillose protuberances; they occur on
the midribs and in .the neighbourhood of the margin of the leaf. Similar
longitudinal rows of papillae are found on the epidermis of the stem in species.
of Loeflingia, Ortegia, Polycarpon and Stipulicida (] osting). Amongst the
representatives of the Caryophylleae (especially the members of the Poly-
carpeae) investigated by Josting, stomata of the CaryophyUeous type are only
present in SPergularia and Loe/lingia, but not in SPergula nor in numerous
other Polycarpeae examined in addition to Loejlingia (see also Liiders, 10c. cit.).
Colobanthus kergzeelensis, Hook. f. has a one·layered hypoderm above the'lower
epidermis (Mardner). On the occurrence of water-pores, see Spanjer, loc. cit.
Neither unicellular clothing hairs, nor even' unicellular trichomes of any
kind occur in this Order. Josting's statement as to their occurrence in Poly-
c~rpaea is certainly incorrect in the case of P. TeneriUae, Lam., and probably
also P. candida, Webb et Berth. The branched multicellular hairs are more
or less distinctly sympodial in structure; they have the form of a candelabra
or stellate hair (Cerastium mollissimum, Poir., Polycarpon Loejl£ngiae, Benth.
et Hook.), or one with two or more arms (Cerasti-um dicrotrichttm, Fenzl, and
species of Polycarpaea, including P. Teneriffae). Trichomes similar to those
found in Polycarpaea appear to be present also in Stipulicida, according to
Lliders. The uniseriate glandular hairs with a unicellular head are also found
in Loe/lingia, SPergula and SPergularia (on the inflorescence).
3. STRUCTURE OF THE AXIS. The structure of the stem has recently been
investigated in the Sileneae and Alsineae, chiefly by \V. Meyer, and in the
Polycarpeae, especially by Jesting. The statements of the earlier authors
have in some cases been confirmed by these researches, in other cases extended.
\Vith reference to the structure and position of the mechanical ring and its
occasional absence, see the papers cited.
The development of cork in Polycarpaea also takes place immediately
on the inner side of the peri cyclic strengthening ring.
In the following additional cases anomalous zones of growth have been
observed: by me, in the axis of Polycarpaea aristata, Chr. Sm., P. carnosa,
Chr. Sm., P. /iIi/armis, Webb, P. latifolia, Poir., P. microphylla, Cav., and
P. Smithii, Link; by J osting, in the root of Ortegia Mspanica, L., Polycarpaea
TeneriUae, Lam., Polycarpon peploides, DC., Spergula arvensis, L., S. fl.forisonii.
Boreau and SPergularia rubra, Presl: by Liiders, in the root of unnamed
species of Cerdia, Pycnophyllum and Stipulicida.
The development of the secondary zones is probably 1 in all cases extra-
fascicular.
Literature: Costantin, Tiges aero et 50Ut., Ann. sC. nat., ser. 6, t. xvi, 1883, p. 80 et seq.-
[Dam anti, Nettarii estrann:r:. della Silmt fuscata, Giom. Soc. d':tcc1imaz. ct age. in Sicilia, I~8S,
p. 101 ; according to Just, 1885, i, p. 745.]-BOrgesen, Arkt. pI. bJadbygn., Bot. Tidsskrift, xix,
1895, p. 219 et seq.- Schubert, Parenchymscheiden, Bot. Centralbl. 1897, iii, pp. 472-4.-Guffroy.
Dianthus, Bull. Soc. bot. de France, 1898, p. 343.-Spanjer, Wasserapparate, Bot. :Zeit., 1898, p. 53.
-[Clements, Histog. of the Caryophyllales, I, Transact. Amer. l\1icroscop. Soc., xx, 1899, pp. 67-
164 and pI. viii-xxv i see also Contribs. from the bot. Lab., Univ. of Nebraska.]-W. Meyer, Vergl.
Anat. d. CaryopbyU. u. Primulaceen, Diss., Gottingen, 1899, 74 pp.-W. Meyer, Einfluss etc., Bot.
CentralbI. 1899, iii, p. 337 et seq.-Keamy, in Contribs. U.S. Nat. Herb., v, S, 1900, p. 301.-Thomas.
Fenilles sout .• These. Paris, 19oo.-Pitard, l-'ericycle, These,. Bordeaux, 1901, p. 49.-Bouygnes,

I Concerning Sp~rgularia see also lasting, Inc. cit., pro 166 and 180.
 ADDENDA-CARYOPHYLLEAE

Petiole, These, Paris, 190:1, p. 18.-Clauditz, Blattanat. canar. Gew., Diss., Basel, 1902 (Si!cne).-
JOsting, Anat. deY Sperguleen etc., Beib. bot. Centralbl., xii, 190%, pp. 139-80 and Ta.b. iii-iv.-
Mardner, Phan .•Veg. d . .Kerguelen, Dw., Basel, 190:1 (Colobanthus).-Amar, Oxalate de calcinm,
etc., Compt. rend. Paris, cxxxvi, 1903. pp. 901, 903.-[Annari, Piante della reg. medit., Ann. di
bot., i, I9?3. p.: Ii et seq. (Dianlkus).l:-Theorin. ViUUriehom;, AJ:kiv for Bot., i, 1903, p. 17~; see
also loco Clt., Ill, n. 5, 1904, p. 9, and IV, n. 18, 1905, p. 6.-Freldenfeldt, Anat. Bau der Wurz.. ,
Bibl. bot., H. 61, '904. pp. 38-4s.-Solereder, Polycarpaea /iliform;s, BulL Herbier Boissier, 190..,
p. 435 -et seq.-Siissengutb, Behaarungsverh. der Wilrzb. Muschelkalkpfl., Diss., WUrzburg, 190-h
p. l2.-Sarton, Anat. d. pI. affines, Ann. sc. nat., ser. 9, t. ii, 1905, pp. 107-9 (Sajonan'a).-
Weberbauer, Veget. d. Rochanden Perns, in Engler, bot. Jahtb., xxxVIi, 1905. p. 6O_~t seq.-
Dauphine, Rhizomes, AnD. se. nat., ser. 9, t. iii, 1906, p. 355 et seq.-LUders, Syst. Untersuch,
liber die Caryophyll. mit einfachem Diagramm, Di.ss., Erlangen, 19°7, pp. 33-38.-[Maheu et
Combes) Format. su'bero.phelloa. anorm., Bull. Soc. bot. de Francel 190 7, p. 430 et seq.
( G)'psophila).]

PORTULACEAE (pp.111-113).
The genus Lenzia (with L. chamaepitys, Phil.) is placed. amongst the
Genera incertae sedis in Durand's Index, and is given as a doubtful member
of the Amarantaceae in Engler and Prantl, but is considered by Reiche to
belong to the Portulaceae. In this genus the structure of the axis is normal.
The transverse section shows a number of isolated vascular bundles arranged
in a ring, ~nd a continuous pericyclic ring of mechanical tissue. The leaves
are hard, and are provided with a membranous margin. On the lower sis}e
they have a hypoderm composed of thickened but nut lignified cells elongated
in the same direction as the leaf; on either side of the leaf this hypoderm projects
beyond the assimilatory tissue, which consists of rounded cells, and thus
constitutes the membranous margin of the leaf. The assimilatory tissue is
traversed by a median vascular bundle. Numerous stomata are found on the
upper side of the leaf.
With reference to the hairy covering we may mention Reiche's statements
regarding Calandrinia. The papillae on the multiseriate shaggy hairs are in
some cases rather strongly developed, so that • pili plumosi ' result, and occa-'
sionally a papillose branch terminates in a glandular head.
Literature: Reiche, Calandriltia, Ber. deutsch. hot. Gesellscb., 1897. p. 493 et seq.-Gasparis,
Tessuto assimil. del genere Portulaca, Rendiconti Accad. Sc, fis. e mat. Napoli, J901i' pp. 201, 20::1.
-[Holm, Clayton ia , Mem. Nat. Acado of Sc., Washington, x, 1905, pp. :17-37; abstr. in Bot.
Centralbl. 101, p. 5.]-Reiche, Syst. Stellnng von Lmzia, in Engler bot. Jahrb., xxxvi, 1905, pp. 8-4,
8S.-[Holtermann, Einflnss d. Klimas. 1907, p. 87,]

TAMARISCINEAE (pp.113-116).
2. STRUCTURE OF THE LEAF. In the first place we may add that in the
species of Tamarix and Myricaria which have been investigated, the stomata
are restricted to the upper side of the leaf (Vesque and Kohne).
Regarding Fouqu£era the following statements may be added. The
structure~of- the Jeaf varies from bifacial to centric. The stomata have no
subsidiary cells. Oxalate of lime is present, and is deposited in the form of
solitary crystals in the cortex, and of structures resembling sphaerites in the
midrib of the leaf. Unicellular clothing hairs have likewise been observed.
The conversion of the persistent part of the leaf into a thorn is due to the
presence of a zone of sclerenchymatous fibres; in the basal portion of the
petiole this zone occupies the whole of the space between the lower epidermis
and the fibrovascular system, whilst higher up in the petiole it gradually
narrows down so as merely to form a subepidermal group of cells separated
by parenchymatous ground. tissue from the vascular system, as seen in trans-
verse section; finally, the fibrous zone terminates in a point, which is either
located in the petiole or most frequently in the lower or middle part of the
SOU:RED~R 3H
 ADDENDA-TAMARISCiNEAE

midrib .f the leaf.. ~ At the end of the vegetative period the remaining parts
of the petiole and lamina become detached from this mass of sclerenchymatous
fibres, which then cCfstitutes the foliar thorn found in the species of Fouquiera.
"'he fibrous zone may be regarded as a continuation of the layer of scleren·
chymatous fibres found in \he primary cortex (see below).
3. STRUCTURE OF' THE BRANCH. In the first place we may notice the
characteristic structure of the Quter zone of the primary cortex, common
to all the species of' Fouquiera. This zone originates by secondary division
of ~he subepiderm,,:llaye, of cells ,and ~onsists ~ither ~iirely, or only at ce~tain
pOlOts correspondmg to· the leaves sItuated Immediately, above, of a tIssue
composed of elongated sclerosed elements with narrow lumina and varying in
the extent of its development according to the species. In older axes cork-
formation sets in on the inner side of this sclerenchymatous zone, the cork
consisting of cells with thin walls and wide lumina. The pericycle contains
isolated groups of bast-fibres only in certain species of Fouquiera (P" spinosa,
Torr_ and F. columnaris, Kell.). According to Van Tieghem, secondary hard
bast is not developed in Fouquiera. It still remams doubtful whether the
I hom-leaves' (HornbHi.tter) found in older stems of Fouquiera splendens.
Engelm, and investigated in detail by Schaer, are really part of the secondary
cortex, as this author assumes, or belong to the sderenchymatous tissue above
mentioned. These structures contain what is known as Ocotilla-wax, and
are composed of peculiar fibrous cells, glued to one another' by a substance
resembling wax; similar substances are also present in' the thick cellulose-
walls. For details as to the nature of the pericyc1e in Tamarix africana and
Myricaria germanica, see Pitard, loco cit.; in these two species the pericycle.
even in branches of slight thickness, includes a composite and continuous
sclerenchymatous ring, which subsequently gets split open.
Literature: Poisson, EpiQ.e5 de l'Idn'a (olulIlnaris, Bull. Mus. d'hist. nat., i, 1895, pp. 218• 279.
-Schaer, FoufJuim:z splcrukns, Archiv d. Pharm., 236, 1898, pp. I-B.-Kohne, Papillen, Mitteil.
deutsch. dendrolog. Gesellscb., 1899. p. SI.-Van Tieghem, fou'luieriacees, Jouro. de bot., 1899,
pp. ~93-301.-Petersen, Vedanatomi, 1901. pp. 4 2 , 43 (JJlyrt'caria).-Pitard, Pericycle, These,
Bordeaux, 1901) pp. 73. 74.-Jonsson. Anat. Ball d. Wiistenptl., Lunds Univers. Arsskrift, xxxviii.
1902, p. 39.-[Robinson, Spines of Foul}uitYa, Boll. Torrey Bot. C!ub, xxxi, 1904, pp. 45-~O.]­
Pic~ioli, Legnatni. Bull. Siena, I9O'J, p. I2.9.-[Holtermann, Einfluss d. Klimas, 1907, p. 9~
( Tamarix).]

HYPERICINEAE (pp. 1I7-120).

I. ANATOMtCAL FEATURES. In Endodesmia the cork develops in a sub~
epidermal position, so that superficial cork-formation also occurs in this Order.
The stellate hairs are accompanied by simple unicellular or uniseriate trichomes
with thin walls and wide lumina. Papillose differentiation of the epidermis
of the leaf-has recently been observed in species of Cratoxylon. Endodesmia,
Haronga, and Hypericum. Hypoderm has been recorded in the leaf in species
of Haronga, Hypericum and Psorospermum. Interxylary phloem has been
met w.ith in the wood of Endodesmia calophylloides, Benth. .
2. STRUCTURE OF THE LEAF. Nearly all the species of Hypericineae
investigated by Kexel and Weill 1 have bifacialleaf-structure, the palisade-tissue
in most cases consisting of a single layer of cells. Certain species of Hypericum
(H. Coris, L., H. ericoides, L., H. procumbens) have rolled leaves. In Hyperi-
cum ericoides the upper epidermal cells bear papillae which are solid. Papillose
differentiation of the lower epidermis is found in H'. procumb~ns (only in the
furrows), H. Roeperianum, Schimp., Haronga madagascariensis, Choisy, Cra·

! Weill'!i invC9tigations extend to all the genera.

 ADDENDA-HYPERICINEAE 835
toxylon polyanthum, Korth., and I have mys~lf found i~ also in E'ltdodesmia
calophylloides. In Hypericum Richerii, Vill., on the other hand, only isolated
epidermal cells on both sides of the leaf are produ<;ed mto papillae. I tin.d
the papillae present in Haronga madagascariensis to be particularly charac-
teristic; they are long, finger-shaped, and rather thick-walled structures,
which are frequently divided by transverse walls and are sometimes fused
with one another. Hypoderm is present on the upper side of the leaf in
Haronga madagascarienSlis, Hypericum nanum, Vismt'a dealbata, H~ B., V. ferru-
ginea, H. B., V. gu.ianen$is, DC., V. parvifiora, Cham. and V. viridiflora, Duch. ;
on the lower side of the leaf in Hypericum nanum and Psorospermum jebri-
fugum, Spach. Weill comes to the conclusion that the stomatal types found
in the Hypericeae on the one hand. and'in the Cratoxyleae and Vismieae on
the other, do not present so uniform a character as Vesque maintains. Weill's
investigations do not however appear to me to be quite .reliable from this
point of view, since in Endodesmia calophylloides, Haronga madagasca1iensis
and Vismia lerruginea I was able to demonstrate the occurrence of the
Rubiaceous type, which is contrary to Weill's statements. In exceptional
cases (Hypericum hyssopitolium, L.) stomata also occur in small numbers on
the upper side of the leaf.
In his description of the secretory cavities, which are filled with brown
contents, Weill again disputes their schizogenous origin, and states that they
arise from a group of secretory cells, iri -which the separating walls can ulti-
mately no longer be demonstrated.
The hairy covering invariably consists of clothing hairs with thin walls.
The hairs found'in the species of Hypericum are unicellular or uniseriate, and
in the latter case consist of from two' to many cells, which vary in length and
shape--sometimes even in the same trichome. I have examined the stellate
or tufted hairs found in Vismia (also V. ferruginea, H. B.) and Haronga;
they have numerous short ray-cells with thin walls and wide lumina, the rays
being inserted at different levels. Weill's statement as to the occurrence of
stellate hairs with a star-shaped terminal cell in V ismia lau1itorm~s, Choisy.
V. ferruginea, H. B. and Psorospermum febrifugum, Spach, as well as the
corresponding figures, are incorrect as far as V. ferruginea is concerned, and
the remaining cases require reinvestigation. In Psorospermum senegalense,
Spach 1, the same author records only simpler uniseriate hairs with cells of
unequal length.
3. STRUCTURE OF THE AXIS. The wood of Endodesmia calophylloides con-
tains exceptionally numerous islands of soft bast, the innermost being found
in the immediate neighbourhood of the pith, which has a four-rayed outline'
in transverse section; I am able to confirm this statement on the basis of an
investigation of material of this species collected by Zenker. The outer
(normal) soft bast is reduced in this case. The mode of origin of the interxylary
phloem yet remains to be determined. The wood of Endodesmia exhibits the
following structural features: (a) the meduUary rays are narrow ; (b) the vessels.
are isolated and some of them are of large size, the lumina invariably being
rounded; they have simple perforations, and the walls bear relatively large
simple pits as well as bordered pits in contact with parenchyma of the medullary
rays; (c) the wood-prosenchyma is covered with small, but distinct bordered
pits.
The pericycle in V ismia also contains a ring or isolated groups of bast-fibres.
According to my own investigation Endodesmia possesses a' composite and

I I did not observe any capitate hairs C' pail. capit~s a la face inferienre ') in E"dodes",ia.
a.lthough I met with papillose protrusion of the lower epidermal cells (see above).
3 H 2
 836 ADDENDA -H YPERICINEAE
continuous ring of sclerenchyma in the per~cycle. In Cratoxylon polyanthum,
according to Stepowski, a zone of bast-fibres is situated immediately beneath
the epidermis of the stem, and on the inner side of this zone there is a layer
of stone-cells, the inner tangential walls of which are thickened. The first
cork in Enjodesmia is developed in the subepidermal layer of cells and not in
the pericycle, as was previously stated; in Psorospermum febrifugum, on the
other hand, it arises in the pericyclic parenchyma. The cells of the cork in
Endodesmia exhibit ~ U-shaped thickening, involving the' outer tangential
and the radial walls. In Psorospermum tebritugum the cork consists of altere
nating layers of unsuberized cells and of uniformly sclerosed cells, which,
however, have fairly wide lumina; the former are elongated in the radial
direction, and are occasionally subdivided by a tangential wall. The periderm
in the root of Hypericum Przewalskii, Maxim. shows a similar structure.
Prior to this J. E. Weiss had already described the occurrence of·unsuberized
cells (phelloid cells) in species of Hypericum.
For the distribution of the secretory organs, whlch Weill classifies as' canaux '
(in the pericyc1e and bast, sometimes also in the pith)~ • poches fusifonnes ' (in the
primary cortex, rarely in the pith), and • poches spheriques on ovoYdes' (in the
mesophyll), and for the branching of the medullary secretory canals, see Weill's
paper cited below. '
Literature: Iodin. Org. secret., These, Mon tpe Hier, 1888, p. 65 et seq.-J. E. Weiss, KOlkbiJd.
Denkschr. bot. Gesellsch. Regensburg, vi, 1890, p. 21.-[Togntni, Stomi,Atti 1st. bot. Pavia, 1894.]
-KexeJ, Anat. d. LanbbI. u. Stengel d. Hyperic. u. Cratoxyleaeetc., Diss., ErlangeD, 1896. pp. 1-36
and Tab.-Meehan, Pellucid dots, Proceed. Acad. Philadelphia, 1897, ii, pp. 18I-3.-Petersen,
Vedanatomi, 1901, p. 4-$ (Hyjtricum).-Batgagli-Petrucci, Legnami, Malpighia, 1902, p. 335
(C,.atoxylon).-[Holm, T,.iadmum vi,.ginicum, Americ. Joum. Sc., 1903, p. 369 et seq.]-Weill.
Rech. hist. sur la fam. d. Ryperic., These, Paris, 1903, 189 pp.-Weill, Repart. d. app. secret. ciani
l'Hypericum calycinum, Jouro. de bot., 1903, pp. 56-6:1.-Siissenguth, Behaarungsverh. der Wiirzb.
Muschelkalkpfi., Diss., Wiirzburg. 1904, p. :l3.-SartOD, Anat. d. pl. affines, Ann. ie. nat., ser. 91
t. ii, J905. pp. 97, 98 (.h)opericum).-Stepowski, Veg. Org. d. Burseraceae etc., Diss., Bern, 1905,
pp. 106-9 {Cratoxylon).-[Moll and Janssonius, Mikrographie d. Robes, Heft I, Leiden, 1906.
pp. :139-49 (Crato,x),lon).]

GUTTIFERAE (pp. I2~I26).

3. STRUCTURE OF THE AXIS. In M esua !errea, L. the primary cortex

,contains a ting·of stone-cells, which is only separated from the epidermis by
a single layer of cells (Stepowski). The pericyc1e in j,fammea americana
and Rheedia laterifolia contains a composite and continuous ring of scleren-
chyma, which includes U~shaped stone-cells (Pitard). In Calophyllum Ino-
phyllum, L. the bordered pits on the vessels have a sieve-like structure
(Ursprung), whilst connecting bands of wood-parenchyma are found also in
Symphonia gabonensis, Pierre and Pentadesma butyracea, Don (Lecomte).
The fruiting axes of Tovomita guianensis show polystelic structure.
Literature: Leblois, Thylles d. can. secret., ·Bull. Soc. bot. de _France, 1887, p. 186:-Jadin,
Org. secret., These, Montpelli~r. 1888, p. 56 et seq.-Jonsson, Anat. Bau d. Bl., Acta Univ. Lund,
xxxii, " I896.-Ursprong, Anat. u. Jahresbild. trop. Holzarten, Diss., Basel, 1900, pp. 8-co
(CalophylJum Incphyllum).-Pitard, Pericycle, These, Bordeaux, 1901. p. 66.-Bargagli-Petrucci,
Legnamj, Malpighia, 1901, p. 336 et seq. (Ca/pphyllul'h, Ganima).-pjtard, PolysteHe, Act. Soc.
Linn. de Paris, set. 6, t. vii, 190:1, p. Ixviii.-[Lecomte, Quelques bois du Congo, Bull. Mus.
·d'hist. nat., 1903, p. 89; according to Bot. Centralbl., xcii, p. 0401.)-Are5choug, Trop. vaxt.
bladbyggn., Sv. Vet. Akad. Handl., 39, n. :z. 1905, pp. I8-:ll, Tab. i, ii, viii, ix (Carcinr"a).-
Stepowski, Veg. Org. d. Bu'rseraceae, Di:Pterocarpeae u. Guttiferae, Diss., Bern, 1905, pp. 97-122 '.
-[For additional literature, see p. II 70.J

1 The Cratoxylon polyanthtWl, mentioned by Stepowski in· this paper, diverges from the remain·
ing Guttiferae in its anatomical stnlcture, and is a member of the Hypericineae. The three-Ia)'ered
epidermis mentioned by the same alltbor :IS occurring in the stem of Clusia Criuva, Camb. is
probably of the natllre oC cork.
 ADDENDA 837

TERNSTROEMIACEAE (pp. 127-136).

I. The following ANATOMICAL FEATURES have been newly recorded:-
uniseriate, multicellular clothing hairs and glandular hairs in the genus Acti-
nidia; tufted hairs with unicellular rays in Gordonia Lasianthus and Lacathea
pubescens; and the occurrence of oxalate of lime in the epidermis of the leaf
in species of Ternstroemia. To the list of genera possessing sclerenchymatous
idioblasts in the mesophyll the following should be added: Camellia (inci.
Thea) and Eurya, which were omitted by mistake, and are considered under
the structure of the leaf; further, the genera Lacatkea, N abiasodendron. and
Ruyschia, and, if I understand Pitard rightly, also Adinandra, Pyrenaria and
Tremanthera.
2. STRUCTURE OF THE LEAF. The anatomy of the leaf in the species of
Thea has recently been investigated by Kochs I. According to him the
characteristic features of the genus are the bifacial structure of the leaf, the
absolute restriction of the stomata with their 3-4 narrow neighbouring cells
to the lower leaf~surface, and the sclerenchymatous idioblasts found in the
mesophyll. For the purposes of specific distinction the varying size, 'shape
and contents of the epidermal cells are more particularly made use of.
The occurrence of sclerenchymatous idioblasts in the mesophyll has
recently been recorded by A. Richter, Francken, Paoli, Pitard and myself
in the following additional species: Camellia Sasanqua, Thunb., Lacathea
pubescens (here extending from one epidermis to the other), M arcgfav£a recti-
/lora, Tr. et PJ.2, M. Sintenisii, Urb. and M. umbellata, L. (asterosclereids),
Nabiasodendron (,sclerites noduleux '). Ruyschia clusiae{olia, Jacq. (astero-
sclereids), Ternstroemia Toquian, ViII. (internal hairs); regarding the species
of Thea, see also Kochs, loco cit. -
The diverse anatomical structure presented by the dimorphic leaves of the
Marcgravieae has been reinvestigated by A. Richter and Paoli with reference to
Juers work. According to A. Richter, the most essential features of the leayes on
the orthotropic shoots of Marcgravia umbel/ata as compared with those on the
plagiotropic shoots are the presence of hypoderm on the upper side of the leaf
and of numerous branched idioblasts in the spongy tissue, the smaller number
of stomata on the upper and the larger numl]er on the lower side. the different
structure, of the guard cells. the somewhat stronger development of the palisade
tissue. and the smaller, though more numerous chloroplasts in the palisade
parenchyma. Paoli describes the following characters as distinctive of the leaves
on the fertile shoots as compared with those on the,sterile shoots: relati~ely small
epidermal cells on both sides of the leaf. a hypoderm on the upper side. a palisade
of elongated cells. and numerous sclerenchymatous idioblasts in the spongy tissue.
In the genus Stachyurus. which systematists in recent times have regarded
as constituting a separate Order (Stachyuraceae), oxalate of lime is excreted
in the form~of.clustered crystals (in the primary cortex and medullary rays
of the bast of the branch) (Van Tieghem). I have myself noticed small cells
filled with clustered crystals and placed singly or in pairs in the lower epidermis
of the leaf in Ternstroemia Toquian, ViII.; these cells often give the appearance

t Unfortunately Kochs' statements are not sufficiently explicit. To ensure proper comprehension
of his meaning, it is necessary to reinvestigate the material he examined. Thus he speaks of
, epidermal cells lying partly side by side and partly one above the other' (= a locally two-layered
epidennis), or of' isolated, relatively large cells, probably containing air (!) and frequently penetrat-
ing slightly into the tissue of the leaf' (perhaps mucilaginous epidermal cells), and so on.
Il The small septate tubes which are described and figured Lv A. Richter in the mesopbyll of
thi~ species, and which branch like the threads of a mycelium, are almost' doubtless mycelial fila-
ments: they are not mentioned by Paoli.
 ADDEND,A-TERNSTROEMIACEAE

of the clustered crystals being merely embedded in the common wall of contact
between the crystal~idioblasts and the adjoining epidermal cells. In -another
undetermined species of Ternstroemia, collected by Loher, I found solitary
crystals in eer,tain of the upper epidermal cells, which scarcely differed
from the others. According td Dunac, styloids occur also in Actinid~'a (A.
chinensis). ..
In amplification of the previous statements about the hairy covering we
may first notice that, according to Dunac, the genus Actinidia possesses the
following types of hairs. in addition to unicellular clothing hairs: uniseriate
trichomes; shaggy hairs (A. Kolomicta, A. polygama. A. tufa, A. strigosa,
A. tetramera); stellate hairs (A. Championi and A. chinensis); and also short
glandular hairs (A. Kolomicta). The tufted hairs mentioned by Pitard
(treatise II) as occurring in Gordonia Lasianthus and Lacathea pttbescens are,
according to my OWn investigation, of the first of these species, composed of
2-4 ray-cells, arranged like a fan.
According to Rochs, most species of Thea exhibit a tendency to form
cork-warts on the lower surface of the leaf. Extra-floral nectaries are found,
for example, in Marcgravia rectijlora, where they occur in scattered arrange-
ment on the lower side of the leaf, being IS in number and '5-1 mm. in
diameter; in M. umbellata, on the other hand, they are present to the number
of four or five at the base of the leaf (Paoli). In the latter species they consti-
tute small pit-like depressions, the epidermis of which is differentiated as
an epithelium.
Detailed statements as to the structure of the petiole are made by
Pitard (II), although his observations o1).ly extend to members of the Tribes
Gordonie;;te and Ternstroemieae, or in other words to Pitard's two groups,
Ternstroemiees and Thcees 1. In these groups the base of the petiole usually
contains a single vascular bundle, which is often very much reduced, and in
transverse section has the shape of a U or a widely open semilunar form.
InroHing of the margins of the furrow formed by the vascular bundle only
occurs in Freziera. In a few cases (F. reticulata) the vascular system is divided
into three bundles. According to Van Tieghem, three vascular bundles enter
the leaf in Stachyur'lts, these bundles being distinct from one another in their
course through the petiole.
3. STRUCTURE OF THE AXIS. In all the members of the Order investi-
gated by Pitard the wood contains isolated vessels with narrow lumina and
numerous very delicate medullary rays.
In Pitard's su.b-tribes Ternstroemieae, Adinandreae and Schimeae, the
development of the cork takes place in the subepidermal layer of cells, whilst
in the Haemocharideae and Camellieae the cork arises in the pericycle. In
Stachyurus I, according to Van Tieghem, the phellogen appears in the epidermis.
In the Camellieae the primary cortex is cast off at an early stage as a result
of cork-development, whilst in the -Haemocharideae it remains on the branch
for a long time. .
In the Ternstroemieae sens. str. and Schimeae, as well as in the Theeae,
the primary cortex contains abundantly branchedsclerenchymatous idioblasts
with long and pointed arms; in the Adinandreae, on the other hand, the

1 For the sake of brevity at this point and in my subsequent synopsis of Pitard's (ii) in-vestiga-
Hons on the structure of the axis, I here append the system of classification established by him, in which
Anatomica.l cbaract~r1I are taken into consideration: I, Ternstr~mieae: I, Temstroemieae scns.
strict.: Tcrnslroem£a, A"neslea i 2, Adinandreac; .Adi.nandro, VislUlJ, Cle,)'U'IJ, Eurya, Fnzura,
Trt"!anlhera; 3, Schimeae: Sclzinta, LacatJua, Gordonw. II. Theeae: J, Haemocharideae:
NtdJtl.fodendron, Haemocharis, Pyrenaria; 2, Camellieae: Camel/la, Thea. Siewariia.
~ In the ro()t of Slachyurus the development Qf the cork takes place in the pericycle.
 ADDENDA--TERNSTROEMIAtEAE
idioblasts in the primary cortex have a more rounded shape, and only exhibit
short and blunt branches. In all the species investigated by Pitard (II), the
pericyc!e of branches, which have attained a thickness of 5 mm.) is composed
of sderosed cells as well as of unlignified cells with thin walls. In the Adinan·
dreae a composite and continuous ring of'scleret1chyma is subsequently
developed; if I understand Pitard (in treatise J) rightly, a similar ring appears
in the species there enumerated by him and belonging to Caraipa, EUfya,
Haploclathra, Kielmeyera, Mahtt rea , Marcgravia, Marila, Norantea, Penta-
phylax, Ruyschia, Schima, and Saurauja pro patte, whilst isolated groups of
bast-fibres are stated to occur in species of A nthodiscus, Caryocar, Saurauia
(pro parte) and Ter-nstroemia. In Caraipa, Haploclathra, Kielmeyera and
Marila, according to Pitard (I), the sclerenchyma-ring includes stone-cells
.exhibiting U-shaped thickening.
In the Ternstroemieae sens. str. and Haemocharideae the secondary bast
contains short fibrous sclerites, provided with short branches (' sclerites fibri-
formes noduleux '), while in the Adinandreae and Schimeae there are numerous
long bast-fibres, which are only wanting in Eurya.
The pith is stated by Pitard to be in general homogeneous, while in the
Camellieae it is very heterogeneous.
It remains to mention the polystelic differentiation of the vascular system
in the fruit-stalks of Schima' Noronhae (Pitard), in which the normal ring of
vascular bundles is surrounded by rather numerous small steles.

Strasburgen·a.
This genus, which will best be discussed at this pOint, is included amongst
the Ternstroemiaceae in Durand's Index, being referred to the Tribe Gordonieae;
in the supplement to the Natiirl. Pflanzenfarnilien it is appended to the Ochnaceae
with a query; and recently it has been regarded as the type of a separate Order
(Strasburgeriaceae) by Van Tieghem. Anatomically its most noteworthy features
are the presence of'mucilage~cells in the ground·tissue and the occurrence of cortical
vascular bundles.
Regarding the structure of the branch, or rather of the cortex, the following
statements may be made. The mucilage-cells lie singly or in groups in the outer
portion of the primary cortex, and ar;: also distinguished from the surrounding
-cells by their larger lumina. The cork develops in the subepidermal layer of cells.
The pericy-de at first contains small isolated bundles of bast-fibres, but subsequently
a composlte and continuous ring of sclerenchyma is formed. The secondary bast
includes neither bast-fibres nor stone-cells. Crystals of oxalate of lime occur in
the pith, but there are no mucilage-cells.
The petiole contains three isolated vascular bundles and a number of smaller
bundles as well. Mucilage-cells are found in the petiolar parenchyma.
The lamina of the leaf presents the following structural features. The
.epidermis consists of large cells. Certain of the, upper epidermal cells contain
sphaerocrystalline masses of unknown chemical composition (not oxalate of lime).
Beneath the upper epidermis there is a one-Iarered hypoderm, the mucilaginous
>cells of wruch penetrate into the single layer 0 palisade-tissue: The stomata are
found only-on- the·loWer side of the leaf, and are not accompanied by any special
subsidiary cells. The structure of the leaf is bifacial. The spongy tissue contains
mucilage-cells.
Literature: Pierre, Flore forest. de- la Cochinchine. ix, 1888 (A'~{k)'laea).-Keller, Luftwurzeln.
Diss., Heidelberg, 1889, pp. 18-22.-Wijnaendts Francken, Sclereiden, Diss., Utrecht, 1890.
pp. 4o-s.-[Cav:ua, Idioblasti delle CameIlie, Atti R. Istit. bot. Pavia, serA ~. vol. iv, 1895, p. ~7;
according to Bot. Centralbl. Beih., v, p. 422.]-Tschirch·Oesterle, Atlas, 1895, p. 9 and Tab. iii.-
Ihmac, Adittidia, Com pt. rend., Paris, c:x:xviii, J 899, pp. 159B-looI.-A. Richter, Adatok a
Marcgraviaceae etc., Termeszetrajzi Fiizetek, 1899, pp. 27-87, and Tab. ii-v.-Van Tieghem,
At;tinidie et Sanravie, Ann. sc. nAt., ser. 8, t. x, 1899, pp. 137--4°; also in Journ. de bot., 1899,
pp. 170-3.-Kochs, Thea, in Engler. bot. Jabrb., xxvii, 1900, p. 606 et seq. (also Diss., Erlangeo,
63 pp.• especially p. 3.. et seq. ).-V nn Tieghem, Stachyuracees etc., Journ. de bot., 1900, pp. 1-6.-
Pitard (i), Pericycle, These, Bordeaux, 1901, pp. 60-2 and go.-Bargagli.Petrucci, Lt-gnami,
 ADDENDA __ TERNSTROEMJACEAE
Malpighia, 1902, p. 334 (Archyfaia).-BoIlYgues, Petiole. These, Paris, 1901, p. I 6.-Claudi t.r,
Blattanat. canar. Gew., Dils., Basel, 1903, pp. 37, 28 (Visnea).-Pitard (ii). Polystelie, Act. Soc.
Linn. Bordeaux, ser. 6, t. vii, 1903, C. R., p. lxviii; Rapp. et clas:aificat. des Temltroemiac., loco
cit., p. 1 et seq.; Nabia.s()(/endron, loco cit., p. Iv; Vt'sma etc., loco cit., p. Wx i and Caract. anat.
gen. des Temstroemiac., loc. cit., pp. lxxi-Ixxiv.-Poulsen, Bladkirtl. hos Marcgra1.lUJ umbellata,
Vidensk. Meddelels. Kj"benhavn, 190 a,. pp. 344-6.-Van Tieghem, Strasburgerie, Jount. de bot.,
'903, pp. 199, 2oo.-Paoli, Eterofillia, Nuov. Giorn. bot. Ital., xi, 1904, pp. 'alG-I6, and Tab. it.-
Arescboug, Trap. viixt. bladbyggn., Sv. Vet. Akad. Handl., ~9, n. 2, 1905, pp. 31, 33 (Tkta).-
Piccioli, Legnami, Bull. Siena, 1906, p. 149.-[}<'or additional literature, see p. Il7 3.]

DIPTEROCARPEAE (pp. I36-145).

I. ANATOMICAL FEATURES. Extrafloral nectaries occur in Shorea. Large
crystal-idioblasts, the inner walls of which are mucilaginous or merely thick-
ened, are found in the mesophyll in certain species of Doona and Hopea. Silica-
bodies are present in the parenchymatous tissues of the wood (according to
Bargagli-Petrucci) in species of Dryobalanops and' Cotylelobitlm.
2. STRUCTURE OF THE LEAF. Epidermal cells showing palisade-like
elongation, stomata with two subsidiary cells placed parallel to the pore, and
glandular hairs with a multicellular peItate head are found also in Vater,'a
Seychellarum (Fabricius). Guerin records gelatinization of cells of the upper
epidermis in further species of Dipteroca1'pus, and also in certain species of
Balanocarpus, Doona. Hopea and Shorea. The crystal-idioblasts observed by
the same author in the mesophyll of Doona nervosa, Thw., D. zeylanica, Thw.,
and-Hopea cernlla, Teijsm. et Binn., deserve special mention; these elements,
which are situated immediately below the u'pper epidermis, have mucilaginous
inner membranes and contain a solitary crystal. Similar crystal-cells are
found also in other species of Hapea (H. dryobalanoides, Miq., H. iuclmda,
Thw., H. M engarawan. l\Hq., H. nigra, Burck, H. Pier1'ci, Hance), but in these
case~ the thickened inner walls do not swell up in water (' mucilage fortement
condense,' according to Guerin) ; the cells, moreover, are commonly placed
several side by side and occur also next to the lower epidermis. Fmally,.
according to Guerin, in certain species of Balanocarpus, Dz'pterocal'pus, Doona,
H opea and Shorea, the ground-tissue of the petiole, and in some cases also
that of the midrib and lateral veins, contains cells with mucilaginous mem-
branes or whole rows of such cells (ct. under structure of the axis).
The extra floral nectaries situated on the lower side of the stipules and.
on the upper side of the foliage leaves in Shorea stenoptera, Burck, are disc-
shaped structures, consisting of two layers of cells derived from the epidermis.
viz. of an upper secretory layer composed of long prismatic cells and of a lower
Jayer of suberized ceUs, which are quadratic in section. The domatia found
in many members of the Order, such as species of Balanocarpus, Doona, Hopea~
[soptera, Pentacme and Shorea, must not be confused with the extrafloral
nectaries; these domatia appear on the under side of the leaves along the
midribs and in the axils of the lateral veins of the first order (see Guerin, 10c. cit.).
• 3. STRUCTURE OF THE AXIS. Guerin has recently investigated the distri~·
bution of the mucilaginous cells in the axis in certain species of Balanocarpus ..
Dipterocarpus, Doona and Shorea. They are found chiefly in the primary-
cortex, more rarely also in the pith. In Daona Guerin only met with a small
quantity of mucilage in the primary cortex; this is contrary to Brandis's
statements.
The course of development of the interxylary resin·canals has formed the
subject of investigations undertaken by Guerin ~m Dipterocarpus. According
to this authority the origin is schizogenous. the canals arising between four
cens of the cambium. In this respect, as well as in the occurrence of occasional
 ADDENDA-DIPTEROCARPEAE

anastomoses, these resin-canals resemble those of the genera Copaifera and

Daniella (Caesalpinieae). Interxylary resin-canals have been recorded by
Guerin and Bargagli-Petrucci also in species of Anisoptera, Balanocarpus,
Cotylelobium, Doona, Tsoptera, Monoporandra, Pachynocarpus, Pentacme and
Vateria.
Stepowski's statement (lac. cit., p. 92 and Fig. 47) as to the occurrence of
resin-canals with a small lumen in the secondary corte» of Vateria indica, L.
remains to be explained; for. to judge by the structure of the leaf, which is described
in the same paper. the material on which the investigation was undertaken would
appear to have been correctly determined.

ApPENDIX: I. Ancistrocladus (pp. 143, 144).

Van Tieghem's recent investigations, in which he splits up Ancisl,ocladus
into three genera, Bigamea. Ancistrocladus and Ancistrella, necessitate the addition
of the following statements. ' .
AIl parts of the primary cortex of the branch contain relatively large secretory
'.;ells, filled with hyaline contents and provided with lignified membranes; these
secretory elements may occur either isolated or in groups. In A ncistrella Barte1'i, V. T.
the primary cortex also includes a few isolated fibrous cells. The development of
the cork takes place on the inner side of·the endodermis, viz. in the outermost layer
of pericyc1ic cells. The pericycle is at first parenchymatous, but subsequently
comes to contain a number of sclerenchymatous elements, which in some cases
unite to form an almost continuous ring; occasionally these elements appear in
two forms, viz. as short cells with a tendency to stellate differentiation and as
fibrous cells.
In A ncistrocladus and Bigamea the leaf exhibits more or less distinct palisade
tissue on its upper side, and veins which are vertically transcurrent by means of
sclerenchyma. whilst in A ncistrella Barteri there is a one-layered hypoderm on
the upper side with a subjacent palisade layer. The peltate glands previously
described by me constitute a feature characteristic of all the Ancistrocladeae;
these glands are sunk in depressions on the surface of the leaf, and excrete wax.
The stomata are invariably situated on the lower side of the leaf, and are not pro-
vided with special subsidiary cells. According to Van Tieghem, a special character
of the fibrovascular system of the midrib is found in the occurrence of a/pericyclic
zone of fibres in the peripheral portion of which, in the lower part of Ithe curve,
5-19 small vascular bundles with inverse orientation of wood and bast are embedded.
These bundles can be traced to a varying distance along the midrib, and are only
absent in Bigamea hamata, V. T. ./
Three vascular bundles enter the Ie"af and at once unite to form a tube,
whiCh in Anc£strocladus can be traced along the whole length of the midrib. In
Bigamea the tube opens out and assumes the form of an inverted omega (u) ;
in addition to that, a vascular bundle is found at each of the two margins of the
fibrovascular arc, these bundles having their bast directed upwards and their
xyJem pointing downwards. In Andstrella the'tube remains closed, the vascular
system on the upper side of the curve exhibiting inverse orientation (bast below
and wood above, as in the lower portion of the curye).

Lophira (p. 144)

2.
The following additional details regarding the structure of the branch are
based on Van Tieghem's statements. The primary cortex includes branched
stone-cells containing crystals, as well as clustered and solitary crystals. The
number of cortical vascular bundles is 24 or more. The pericyclic groups of hard
bast subsequently unite to form a composite and continuous ring of sclerenchyma,
which at a still later stage again becomes split open. The development of cork
takes place in the subepidermal layer of cells. The bundles of medullary fibres
are accompanied by chambered crystal-fibres containing solitary crystals.
According to Van Tieghem's recent investigations, the earlier statement (by
Heim) regarding the occurrence of subsidiary cells to the stomata is incorrect.
The large number (as many as 16) of radial vascular bundles, participating III
 ADDENDA--DIPTEROCARPEAE
the formation of the fibrovascular system of the root. deserves mention; it is
exceptionally large for a Dicotyledonous plant.

3. Monotes (~ 144)-
As a result of renewed investigation Gilg again includes this genus amongst
the Dip\erocarpeae. though in the absence of resin-canals it certainly occupies an
anomalous position amongst the members of this Order. According to Gilg.
however, it is noteworthy that Welwitsch describes Monotes as • frut~x resinosus:
so that the question arises whether excretion of resin may not take place in older
parts of the steIll.
Literature: Leblois, Tbylles de can. secret., Bull. Soc. bot. France, 188,., p. 187.-Jadin, Org.
secret., These, Montpe1Iicr, 1888, P.54 et seq.-Pierre, Flore forest. de la Cochin<:bine, xv, 1890;
xvi, IS91; and ltvii. J~l.-Brandis, In Sitz.-Ber. Niederrhein. Gesellsch. Bonn, 1896, pp.6-8.-
Poulsen, Nektarier. Vidensk. Meddelelser, Kjllbenhavn. 1897, pp. 368-7o.-Gilg, MOHl)/ts,,.in Engler
Bot. Jahrb., xxviii, 1899, p. 117 et seq.-FigdOT, Anat. d, Stamwes de. Dammarpfl., Osterreich.
bot. Zeitschr., 1900, p. 74 et seq.-Pitard. Pcricycle, These. Bordeaux, 1901, pp. 40 and 104.-
Van Tieghem, I.qphira, Joum. de bot., 1901, pp. 169-94, especially pp. I7l-g.-Bargagli-Petrucd,
ConcreziQni silicee, Malpighia, 1902, p. :il3 et seq:; and Legnami, loco cit., p. 338 et seq. (Balano-
carpUJ, IJipterocarpuS, Dryo/Jalanops, Hcpea, Skona, Vattria).-Fabricius, Laubtlattanat., Beih.
bot. Centralbl., xii, £9°', pp. Z",S-Jo.-Van Tieghem, Ancistrocladecs, Jouro. de bot., 1903,
pp. 151-68, espt'cially pp. 155-6,s.-Areschoug, Trop. vax,t. bladbyggn., Sv. Vet. Akad. Hand!.,
39, n. :il, 1905. PP' 123, ui and -Tab. ltvii.-Guerin, Appar~i1 secret. d, DipteroCllrp., Compt.
lend. Paris, ex!, 19?5, pp. {'Zt>;2; and Ca,na12x secret. du ,boIS d. Dipterocatp.,. lac. cit., exlii,
1906, pp. ·I02-4.-Stepowsln. \ eg.~Org. d., Burseraceae, J)tpterocarpeae, etc., DIss., Bern, 1905,
pp. 53-9s--G12erin, Domaties den feuilles des Dipterocarp., Bull. Soc. bot. France, lq06, pp. 186-
9~; and Cellnles a mncilage des Diph!rOcarp., loco cit., 1906, pp. H3-5I.-IFor :tdditionallitera-
Hire, see p. ] J 70.] -

CHLAENACEAE (pp. 145, 146).

The pericycle of the axis contains isolated bundles of bast-fibres also in
SchizolQena elongata and Scleroolaena Ricnardi; in Sarcolaena etiopho1'a and
Rhodolaena lIumblotii sclerosed parenchyma occurs at some points between
the groups of primary bast-fibres (Pitard 1).

MALVACEAE (PP.146-152).
2. STRUCTURE OF THE LEAF. Nestler states that the glandular hairs of
the Malvaceae can also function as hydathodes, that short multicellular
glandular trichomes occur in species of Abutilon, Althaea, Hibiscus, Kitaibelia,
L«vatera, j_llalope, l}/alv«, Palava, Plagt:antkus and Sidaleea, while hairs with
a long stalk and a unicellular head are recorded in Abutilon and Kitaibelia
vitiloliCl \Villd.
t

According to Terraciano, extrafloral nectaries occur also in species of

Adansonia ChDrisia~ Ceiba and Pachira, being situated on the lower side of
t

the midrib of the leaf and on the dorsal surface of the petioles. The nectaries
on the midrib show a low type of differentiation and vary in position, number,
•and shape. The petiolar nectaries constitute one or more longitudinal furrows
or small eIIiptical pits; in the former case they are sometimes provided with
special excretory surfaces.
These nectaries, like those of Hibiscus, &c., which were mentioned in the
earlier part of this work, all possess a tissue in which the nectar is formed, and
numerous multicellular external glands serving the purpose of secretion t

.
IPitard, Pericycle, These. Boroeaux, 1901, pp. J02, 103.
; The tlectariei on the caly:lt, (In the other hand, have a secretory palisade-epidermis.
 ADDENDA-MAL VA CEA E

3. STRUCTURE OF THE AXIS. Bargagli-Petrucci has recently investigated

the distribution of the peculiar structural features exhibited by the medullary
rays of the xylem in the Bombaceae, which I was the first to observe. He
describes the occurrence of special tiers in the medullary rays in further species
of Boschia, Durio and Nees'':a, as well as in CQelostegia borneens'is, Bece., and
Cullenia exeelsa, Wight, both of which likewise belong to the sub-tribe Durio-
neae. On the other hand no tiers are developed in the AdansoDieae (species of
Adansonia, Bombax, Ceiba, Chorisia) and Matisieat: (species of Cava'1illesia,
Montezuma, Ochroma, Quararibea), or in Camptostemon aruense, Bece. and
Dialycarpa (Brownlowia) Beccarii, Mast.
According to Bargagli-Petrucci, silica-bodies are found in the wood-
parenchyma in Coelostegia borneensis, Becc.
We may also notice at this point that the vascular system in the fruit-
stalks of Adanso-nia d'igitata shows polystelic structure.
Literature; Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, p. 109 et seq.-
Barber, Corky excresc., Arm. oJ Bot., vi, 1891, p. 163.-Gerber, AdaltSonu" These, PariS, 1895.
pp. 33-51, pl. i-ii.-[Mirabelli1., Colleteri, Contribuz. 1st. bot. Palermo, ii, 1897, p. 15 et seq.;
... bstr. in Just, 1897, j, p. 513.1-Nestler, Wassertropfen an den Bl. d. Malvac., Sit:z:.-Ber. Wiener
Akad., cvi, I, 1897, pp. 387-96, with Tab.-[Zancla, Acutei, Con~ribuz. 1st. bot. Palermo, ii, 1897,
p. 1 et seq.]-Nestler, Schleimzellcn d. Laubbl. d. Malvac.; Osterreich. bot. Zeitschr., 1898,
PP.94-9 and Tab. vi.-Terraciano, Nett. estramlZ. neUe Bombacee. Contribuz. 1st. bot. Pa.lermo, ii,
189M, pp. 137-91, tab. xv-xviii.-Nestler, Sekrettropf., Sitz.-Ber. deutsch. bot. Gesel1s~h., 1899,
p. 331 et seq.-Kearny, in Contribut. U. St. Nat. Herb., v, '900, p. 303 (KoJ'ttle/.:.kya).-Pitard,
PolysttJie, Act. Soc. Linn. de Bordeaux, ser. 6, t. V, 1901, pp. lxi, Ixii; and Pericyc1e, These,
Bordeaux, 1901, p. -40.-Bargagli-Pehltcci, Concreto silicee, Malpighia, 190:11, p. l3 et seq.; and
Legnami, lac. cit., 1902, p. 337 et seq.-Knothe, Unbenetzb. Bt, Diss., Heidelberg, 1903, p. 13'-
The orin , Vaxttrichom., Arkiv for Bot., i. )903, p. 16o; and iv, n. 18, 1905, P. 2o.-Bargagli-
Petrucci, Oss. anat.-sist. suIle Bombacee, Nuovo Giorn. bot. Ital., xi, 1904. pp. 407-1S.-Chrysler,
Strand-plants, :BeL Gazette, xxxvii, 1904.0 p. 461 et_ seq. (HihisC1.ls).-Col, Faisceanx" Ann. ac.
nat., ser. 8, t. xx, 19°-4, pp. 108-Il.-Ursprung, Dickenwachst., Bot. Zeit., 19°4, p. 20l (Eru-
dtndron).-Areschong, Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 3, 1905, pp. 56-8
(.lJice//(istyks). pp. 64-6 (Tne.pt'sia).-Weberbaucr, Veget. del' Hochanden Perus, in Engler Bot.
Jahrb., xxxvii, 1905, p. 60 et seq.-[Fol' additional literature, see p. I1iI.]

TRIPLOCH ITON ACEAE.

Triplochito1J. scleroxylon, K. Schum. (Zenker, No. 298, Kamerun, Herb.
Berol.) 1 was examined. In the str:ucture of its bast and the possession of
intercellular secretory receptacles, some of which contain mucilage, Triplo-
chiton shows affinity to the remaining Malvales. The stomata are surrounded
by ordinary epidermal cells. Oxalate of lime is excreted in the form of clus-
tered and solitary crystals. No trichomes were observed.
Structure of the leaf. The epidermal cells are polygonal in surface-
view and none of them are mucilaginous.. Stomata are found only in the
lower epidermis. The mesophyll is composed of palisade tissue, the upper
layers of which are typically differentiated~ while- the lower ones consist of
conjugate 'elongated cells. The vertical transcurrence of the smaller veins is
especially characwristic; their vascular bundles are surrounded by a layer
of fibrous cells, which is prolonged on the upper side into a narrow plate of
fibres extending as far as the epidermis, whilst on the lower side, between the
bundle-sheath and the lower epidermis, there is mostly a secretory cavity
enveloped by special conjunctive tissue. Relatively large clustered crystals
are met with in the fibrous plates belonging to the veins, and smaller ones are
found in the bast of the larger veins; solitary crystals are rare. Secretory
cavities are situated also in the large lateral veins, where they occupy the

1 The second genus of the Order (1IIa1lSoma, Prain, indigenous in Bunna) was not available.
 ADDENDA-TRlPLOCHITONACEAE

same position as in the smaller veins; they are present in the bast as well.
Their contents, as far as they could be determined, are certainly in some cases
mucilaginous, whilst in others they are of a different nature.
The transverse section of the branch shows (empty) secretory spaces in
the inner portion of the primary cortex and in the pith. The phloem is strati-
fied into hard and soft bast. The cOJ.t~x contains clustered crystals, rarely
solitary crystals. The wood is soft and consists principally of: (a) vessels
with rather ,wide lumina, ~mple perforations and bordered pits in contact with
parenj:hyma of the medullary rays; (b) relatively broad medullary rays,
attaining a width of four cells; and (c) wood-fibres bearing simple pits.

STERCULIACEAE (pp. 152-155).

I. ANATOMICAL CHARACTERS. Extrafloral nectaries occur in Ptero-
spermum iavanicum~ Jungh. Silica-bodies are found in the wood-parenchyma
in species of H eritie'1a and Sterculia. The vascular system in the fruit-stalks
of Helicteres jamaicensis, Kleinhovia Hospita and Slercttlia ptatani/olia is
pol)lStelic. ,
2 and 3. STRUCTURE OF THE LEAF AND AXIS. The tufted and stellate
hairs, which are widely distributed in the Sterculiaceae. generally have uni·
cellular rays; in Dombeya (Astrapaea) IVallichii, Benth. and Hook., however,
1 have found the rays to be occasionally uniseriate with several thin trans-
verse walls.
The extraflQral nectaries. found in Pterospermum iavanicum. although not
present in other species of Pterosperfflum growing in Java, ale constituted by
one of the two stipules of the foliage-leaf. This stipule is hollowed out so as
to be goblet-shaped, and is clothed with abundant stellate hairs; the cavity
contains numerous pearl-glands in the form of relatively large glandular bodies
composed of numerous cells and provided with a short stalk (Raciborski).
Doussot publishes a few statements on the structure of the petiole.
According to him the fibrovascular system of the petiole in most cases (species
of Sterculia, Heritiera, Pterospermum, Klei-nhovia, Lasiopetalum, Theobroma,
Abroma, Hermannia~ Che';ranthodendron) consists of a dng of wood and bast,
Of of a ring of vascular bundles; in Cola aC1tminata there are two arcs of wood
and ba~ with the xylem portions turned towards one another, while in C. Bal-
layi there is a single arc of the same kind. In addition to these main systems,
the pith in some of the species of Sterculia contains one or more normally
orientated vascular bundles, in Heritiera a second ring of wood and bast, and
in Cola acuminata one or two smaller bundles.
In amplification of the previous discussion of the mucilage-receptacles
we may in the first place notice that, according to Doussot, the mucilageMcanals
of Sterculia aceri/olia, S. platanifolia, Cola acuminata, and' Pterospermum
saigonense' are partly lysigenous and partly schizogenous in origin, while the
mucilage-receptacles (canals and cavities) ()f 'Abroma orbicutaris ' are invariably
lysigenous. The development of the mucilage-receptacles in the individual
genera therefore requires further detailed investigation. According to the same
authority it is noteworthy that the number of mucilage~canals may vary at
different levels in one and the same plant, and that at some points the canals
may be replaced by mucilage~cavities.
To the review given on p. 153 of the detailed distribution of the mucilage-
canals in the axis, the- ~llowing additions may be made on the basis of Doussot's
and Ledig's Itatements. Mucilage-canals are found in the following additional
species :-( t) in the pith and primary cortex.: Sterculia aceri/olia. S. foetida.
 ADDCNDA-STERCULJACEAE

s. ' furcc:ta/ S. • monophylla,' and S. tomentosa 1; Cola Baflayi and C. gabonensis ;

Heli~te,es involuc"ata; Ptef'ospermum' saigonense '; Ruizia variabilis; Astrapaea.
WaJlichii. (2) in the pith, but..,not in the cortex: Heri"era littMalis; Klein}tovia
HosPita (~t with numerous muCilage.cavities in the primary cortex); Pte,o~
spennum ~berifolium (many mucilage-cavities in the primary cortex); Ab"oma
cwbicuJaris,
Before leaving this subject we may notice the occurrence of interxylary
mucilage-canals in the older portions of the axis WBrachychiton, populneum
and Theobroma Cacao (Mangin).
According to Van Tieghem, the (lysigenous) mucilage-cavities and mudl~e.
cells are absent il\. Herma,nnia, Mahernia, Rulingia, Battneria, LasiopetarUtnt
Thomasia and M elochia, and according to Doussot also in H eritiera littarah£
(in contradistinction to R·. macrophyllR). Doussot, on the other hanq, state~
that mucilage-cells occur in the pith of Hermannia candicans, and mucilage:
cells and cavities in the pith of Lasiopetalum ferrugineum. These facts show
that It is necessary to investigate the mucilage.receptac1es of the Sterculiaceae
in each individual species before they can be employed for generic diagnosis
and classification within the Order. Schizogenous mucilage~cavities~ situated
in the pith of the branches and petioles, are stated to be characteristic.()f ~ y.:nia
and Commersonia. Doussot records the occurrence of mucilaginous epid6lJl1a1
cells in species of Sterculia, Brachychiton, Cola, Theobroma, Abroma, Hermannia
and Dombeya.
A brief account of the mucilage-receptacles found in the root may be added
on the basis of Doussot's results. Amongst the" investigated Sterculieae the roots
of aU except Heritiet'a (Sterculia, Brachychiton, Cola) contain only mucilage-cavities
and mucilage~cells. The mucilage-cavities are confined to that part of the main
root which is situated immediately next to the • collet'; they may be present in
the primary cortex, in the medullary rays of the bast. and in the wood (here in the
wood~parenchyma or in the medullary rays). They have been' observed in the
wood in Sterculia Balanghas, S. loetida, S. 'monophylla,' S. ptataniloJia, Brach'Y.
chiton aceri/olium and B. populneum. In 'Alwoma orbicularis' mucilag~avities
only are found in the medullary rays of the bast. while in Hcrmqnia Icandicans
the· primary cortex and bast contain mucilage-cells alone. In Hlhtiera. Plero-
spe'w.um~ Theobroma Cacao, Dombeya spectabilis and Ches'yanthodendron platan;:"
folium there are no mucilage-receptacles whatsoever.
Literature: Ledig, Gummikaniile, Bot. Centralbl., "" 1881, ii, pp. 381-9.-Jadin, Org. secret,
These, Montpellier, 1888, p. 61 et seq.-Johannson, RiDden, Diss., Dorpat, 'J89I, p~ 8-1I.-
Mangin, Gemme chez les Stercul., Compt. rend. Paris, cxxv, 1891, pp. 13S-8.-BOrgesen og
Pau~ Veget. dansk.-vestind. <>er t Bot. Tidsskrift, xxii, 1898-9, 'po 93 (Mel«hitJ lomentostJ).-

~
artw.ich) Ersatz der QuUlajarinde, Schweizer. Wochenschr. f. Chemie, 1899, p. +9; abstr. in
ut.l 11599, ii, p. 28; cortex and wood of Sterculia cordi/olia, Cay. contain saponin I]-Raciborski,
Mt_rmek0l?hile Ptl.! Flora, 19oo,.]? 38 etseq ..-~itard, Pericyete, These, Bordeaux, I90 I ! pp. 106:-8.
~gag1i.PetruCCl, Concrez. Slhcee, Malplghla, 1902, p. 23 et seq.; and Legnaml, loco CIt.,
p.' 331 et seq. (Heritiera. Sterculia).-DoDssot, Appareil gommifere des Stercnl., These, Paris,
J90 2 , 75 pp.-Pitard, Polystelie, Act. Soc. Linn. Bordeaux, ser. 6, t. vii, I9Oa.-Barle1etti, Corteccia
dl Ptenspermum jJlalanifolium, Nuov. Giom. oot. It:Ll., x, 1903, p. S66.-Col, Faisceaux, Ann. sc.
naL, II!r. 8, t. xx, J904, p. U3.-Ursprung, Dickenwachstum, Bot Zeit., 190+, p. 205 (Melochia).-
~oug, Trop. viixt. bladbyggn., Sv. Vet. Akad. Ranell., 39, n. 2, 1905, p. 63 (Biit/neria), p. ~
(D0mh7a), pp. 79, 80 (Biittneria), pp. 118-20 (Pteroj'pn-mum).-Haberlandt, Uchtsinnesorg.,
I 90S, p. 96.-(Prain~ Mansoni.eae, Journ. Linn. Soc" xxxvii, 1905, p.. 250 et seq.]

1 The' septate latiferons tubes' recorded by Ledig in S/b'culi4 Balanglzas are mere1y:lysigenoUi
mucilage.canals.
 ADDENDA

TILIACEAE (PP.155-159).
3. S;rnUCTURE OF TIlE AXIs. A ccnnposite and continuous ring of sc1eren-
chyma. in the pericycIe is present! also in the genus Sloanea (Pitard.). Silica-
bodies have been obseIved in the wOQd-parenchytna in Brownlo'lJ)ia sp. (P. B.,
no. 3652)" (Bargagli-Petr'(ICci).
Literature: Bohnel, Gerberimlen, Berfm, 1880, p. n I et seq. {ElaeorQ.t1us).-Nanke, DikqO'le
UolZpB., D_, KOnigsberg, J886, P 17 et seq.- BOrgeseo og PllU)SfD, Veget. daJJsk.~v'e5tjJld. Oer,
Bot. TidliioSkrift, ;cxii; "I89S~9, p. 94 (C(It'ckorus In·7'sulUJ').-Pattl~, Struct. anat. de$ hybrides,
TheJe, Geneve, 1900, p. 69 (Til,a).-Pctersen, Veda.oatoml j .~I. p. 45 (Tilia).-Pitard, P.i-
cycJe, These, &rdeaRx, U)OI, pp. ~o and lo8.-&rgagli-Petrucd, C()ocrez. ailicee, M41p~hia, 1~2,
p. a3 et seq.; alld Legnaml, loco cit., p. 3:15 et seq. (Berrya, Brownlowia).-Bouyguts, Petiole,
These, Parl6;. 190:1, p. Io.-Fritsch, Plagiopteron jragralls, Ann. of Bot., xvi, 1902, pp. 1'9'1-80.-
Col, Faisoeau x, Ann. sc. Dat.,ser. 8, t. n, '904-, p. I 36.-Areschoug, TrQP. viix:t. bkdbr~.. Sv.
Vet. Akad }Iandl .. 39, n. 2, 1905, pp. 136-8 (Grt'lvia).-[Frornmel, Plantas text. ChiL, '90S,
p_3a.1-Haberlandt, Lichtsinnesorg., 1905, p. 97.-Piccioli, Legnami, Bull. Siena. 1,06. p. 141.-
[Holte~ann, Einfiuss d. Klimas, '907, p. uS (ElacQCarpus).]
- l

RHAPTOPETALACEAE 1.
The following anatomical features are common to the different members
of the Order: The bilateral structure of the branch, and in connexion with
this the ocCurrence of two cortical vascular bundles; the superficial de~elop­
ment of the cork; the presence of isolated groups of bast-fibres in the peri cycle ;
the stratification of \he phloem into hard and soft. bast; narrow medullary
rays and rather abundant development of wood-parenchyma; stomata with
three subsiwary cells of different sizes. The only kind of trichomes found are
unicellular clothing hairs. Oxalate of lime is deposited only in the form of
solitary crystals. There are no mucilage-cells. On the other hand, spicular
fibres have been observed in the mesophyll in species belonging to all four
genera.
Structure of the leaf. The mesophyll is in most cases bifacial, although-
the palisade;tissue is not always distinctly developed. In certain speci~s of
Scytopetalmn and Ottbanguia the epidermal cells are elongated and divtded
by tangential walls; Brazzeia has large epidermal cells, the inqer'" walls of
wbich ilTe convexly arched~ while some of the cells are filled with red contents.
Gelatiniz~~ion of the epidermis has not been observed. The stomata are found
either on .{)ot'li sides of the leaf or only on the lower side. In Oubang'ltia and
Rhaptopetalmn the three subsidiary cells of the stomata have violet or brown
contents, w4jle in Brazzeia such contents are found only in th~ smallest sub-
sidiary cell; in Scytopetalum they are altogether wanting. The spicular fibres
above mentioned as occurring in all four genera are connected with the scleren-
chyma of the veins; they traverse the mesophyll and ultimately spread out
beneath the epidermis. The fibres are not always present; for example, not
in Scytopetalum Pierreanum (De Wild.), V. T. In S~ Pierreanum the smaller
veins of the leaf are vertically transcurrent. whilst in Oubanguia they are
almost transcurrent.
An arc-shaped vascular bundle passes out into the leaf. The petiole
contains three bundles, viz, a 1arge median one and two smaller lateral strands.
Unicellular clothing hairs have been observed on the branches only in
species of Brazzeia and Rhaptopetalum..

1 This Order (Scytopetalaceae of Engler), founded by Pierre on the genus Rkapt()petalum

(which was formerly included among the Olacineae). and n few other genera, is related to the
Malvales. According to Van 'fieg-hem, it may be subdivided into the Oubanguieae with Oubanguia
and Scyto_petalum, and the Rhaptopetalea.e with Bra:~eia and Rlta/topetalllm. The above description
is ba!K:d on Van Tieghem's paper cited belo",.
 ADDENDA.--RHAPTOPETALACEAE
In dealing with the structure of the wood, Van Tieghem merely states
that the medullary rays are from one to three cells in breadth, and that wood-
parenchyma: is developed in some qualltity in all four genera. According to
Engler, the wood in Scytopetalum Klaineanum, Pierre contains the following
elements: (a) scalarilorm and reticulate vcssel$ with wide lumina and very
steep transverse walls exhibiting scalatiform perfoPaiions; (b) vessels with
narrower lumina· (c) trachcids; and (cl) Jibriform (i.e. wood-prosenchyma
with simple pits). The pith is lignified.
In the structure of the cortex theiollowing points may be mentioned. The
development of cork in O'IdJanguia takes place in the subepidermal layer of
cells, or locally in the second layer of the primary cortex, while in Scytopetalum,
Brazzeia and Rhaptopetalum it commences in the epidermis. III Rhaptope-
talum the cork includes cells with U-shaped thickening. The two genera of the
Oubanguieae are characterized by the fact that all the different P3f'ts of the
printary cortex (even the subepidermal layer and the endodermis) contain
cells with U-shaped thickening and with or without an enclosed crystal
of oxalate of lime (Van Tieghem's ' cristarque '-cells); in some cases (Scytope-
talum) the cells are uniformly thickened all round. In the Rhaptopetaleae,
on the other hand, cells of this type are wanting. For details as to th~ rovrse
of the two cortical vascular bundles, see Van Tieghem; regarding th~ pertCycle
and bast, d. above.
Literatnre: Engler. Scytopetalaceae, in Natlirl. Pflanzenfam., Nachtr. z. ii.-iv. Tetl, 18~)7,
pp. 242, ~43.-Van Tieghem, Rhaptopetalacees, Ann. sc. nat., ser. 9, t. i, 1905, pp. 331-88.

LINEAE (PP.159"-160).
1. The following characters, newly recorded in certain members of the
Order, may be added to the REVIEW OF ANATOMICAL FEATURES: deposition
of oxalate of lime in the form of clustered crystals; epidermal cells in the leaf
containing solitary crystals (Erythroxylon Coca); occurrence ot peg-shaped
cystolith-like bodies (E. obtusum); presence of hypoderm in the leaf..;' scleren-
chyroatous idioblasts in the mesophyll (species of Erythroxylon); occurrence
of cells exhibiting U~shaped thickening and containing solitary ~ystals,
i.e. 'cristarque' -cells (species of Erythroxylon and A ne'ulophus); cortical
vascular bundles (Erythroxylon). The development of the cork is iuperficial,
taking place in the subepidermal layer in Erythroxylon, and in the epidermis
in A neulophus.
2. STRUCTURE OF THE LEAF. Wilde has recently investigated species of
Linum, Radiola and Reinwardtia, whilst Hartwich has examined species of
Erythroxylon, and Van Tieghem the genera Erythroxylon and A neuloph~ts.
In certain species of Linum the leaf is centric in structure; it is also approxi-
mately centric in Erythroxylon tortuosum, in which the cells constituting the
lowest layer_:ot-the'mesophyll have the same form as the spool-shaped girder-
cells found in Papilionaceous seeds. According to Van Tieghem, gelatinized
epidermal cells are found on the upper side of the leaf in all the species of
Erythroxylon examined by him. as well as in A neulophus african us, Benth.,
Hugonia montana, Pierre, and Roucheria Contestiana, Pierre. A one-layered
hypoderm is situated on the upper side of the leaf in Ixonanthes cuneaia, Miq.
According to my own observation certain of the lower epidermal cells in the
leaf of Erythroxylon Coca contain solitary crystals of oxalate of lime, these cells
being sometimes arranged in pairs. Papillose differentiation of the lower
epidermis is of frequent occurrence in the genus Erythroxylon, e.g. in E. ovatum
and E. subrotundum (Hartwich). The stomata are accompanied by subsidiary
 ADDENDA-LINEAE

cells placed parallel to the pore, also in A neulophus, Radiola and Reinwardtia.
In Erythroxylon bolivianum, ~urck certain cells of the lacunar spongy tissue
are sclerosed (Hartwich). Aa::oqiing to ij:artwich, typical sclerenchymatous
idloblasts, usually assuming the. shape of a T, are found in Erythroxylon acuti-
lolium, E. C"itrifoliJ'm, E. mucronatum and E. squamatum; these elements
traverse the palisade-tissue, and then continue their course between the latter
and the. epidermis,. Van Tieghem also records • scIerites rameuses' in Ery-
throxylon amplum, E. campestre, E. lucidum, and E. suberosum, whilst Pierre
mentioI\s f rares spicules' in Hugonia montana, Pierre. According to Van
Tieghem, the vascular bundles of the veins are provided with bundles of scleren-
chymatous fibres in A neulophus and Erytkroxylon; in A neulophus and some
of the species~of Erythroxylon these fibres are accompanied by , cristarque '-
c~lls.
The clustered crystals above mentioned have been observed in the priI~.ary
cortex in..Reinwardtia indica, Dum., and in the secondary bast in Erythro~lon
pulchrum (Wilde and Johannson). The cystolith-like bodies appear as small
unlignified peg-shaped structures, arising from the middle of the outer wall
of the up·per epidermal celIs and occasionally showing stratification; th~y
have only 'been met witp in Erythroxylon obtusum (Hartwich).
In A.neulophus, Erythroxylon, Hugon£a, Ixonan.thes and RouclJeria, as well
as in the genus Ctenolophon, which 'has been transferred from the Olacineae
to the Lineae by Pierre, three vascular bundles branch out from the axis into
the leaf. Van Tieghem gives the following description of the petiole in Aneu-
tophus anti Erythroxylon. Of the three vascular bundles passing out into the
leaf in E rylhroxylon, the two lateral strands depart from the vascular ring of the
axis in the lower portion of the interndde; they consequently traverse the
upper part of the internode as cortical bundles. These two bundles each give
off a branch. to the stipules, and finally on entering tlle petiole fuse with
the median bundle, which leaves the axial vascular ring at the node, to form
ill arc of' wood and bast. In A neulophus the three vascular bundles unite
to form a similar arc of wood and bast, but here the bundles only emerge at
the node. In some of the species of Erythroxylon the ground-tlSSue of the
petiole contains irregularly scattered (cristarque' -cells, while in A neulophus
such ce]]s are found in the subepidermal and endodermallayers.
3. STRUCTURE OF THE AXIS. According to Wilde, the xylem contains
vessels with simple perforations and libriform also in Linum and Radiola ..
In Anettlophus and Erythroxylon the cens of the cork have thin walls (Van
Tieghem). Wilde records a distinct endodermis in Reinwardtia indica, and I

a pericycle including isolated groups of fibres in Linum and Radiola linoides,

Gmel. According to Van Tieghem, isolated groups of bast-fibres are present'
in the pericycle also in the species of Erythroxylon, while in Aneulophus a com-
posite and continuous ring of sc1erenchyma is formed at an early stage.
, Cristarque '-cells are only found in the primary cortex in part of the species
of Erythroxylon; in a few species (E. anguifugum, E. suberosum) they are
situated in the second layer of cortical cells, but in most cases (E. amplum,
E. campestre, E. citrifolium, E. deciduum, E. lucidum, E. nitidum, &c.) they are
irregularly distributed throughout the whole thickness of the primary cortex.
In the monotypic genus A neulophus the C cristarque ' -cells are confined to the
subepi~rmal layer and the endodermis (see above, under petiole). In Ery-
throxylon Coca cells with yellowish or yellowish-red contents are found in the
primary cortex, as well as in the pith. The secondary bast of. E. tortuosum
includes groups of stone-cells, while that of E. australe and other species con-
tains bundles of fibres; the latter are especially numerous ~ E. suberosum,
where they cause a stratification of t,pe bast (Van Tieghem). In E. pulchrum
the secondary bast includes fibrous cells and stone-celts (Jobannson); in
 ADDENDA-LINEAE

A neulaphus there is practically a second sderenchymatous ring exhibiting the

same structure as that found in the pericycle and having a layer of ( cristarque ' ..
cells on its outer side (Van Tieghem).
Literature: Johannson, Noch wenig bekannte Rinden, Diss., Dorpat, 1891, p. 3g.et seq. (Erylnr~.
X)'IDn).-[Tognini, Fasci libro-Iegn. prim. negli org. veg. del lino, Atti 1st. bot. Pavia, 1891,
:n pp.; according to Bot. Centralbl., 1, p. 337.}-Pietre, Flore forest. de la Cochinchine, xviii,
IB93.-(Tognini, Stomi, Atti 1st. bot. Pavia, I~ ..]-Schubert, Parenchymscheiden, Bot. Centralbl.,
1891, iv, p. 16.-Wilde, Beitr. z..·Anal d. Lineae, Diss., Heidelberg, 1903, ;;6 pp. and I Tab.-
Hartwich, Cocablatter, Archiv d. Pharm., 241, 1903, pf.. 617-30, and Tab. 1, h.-Van Tieghem,
Erythroxylactes, nouv. ex. de cristarque, Ball. Mus. d hist. nat., 1903, pp. 287-93.-[Greenish,
Structure of Coca leaves, Pharm. ]oum., 1904, pp. 493-6.]-S~gutb, Bebaarnnguerh. d.
Wmb. Mu.schelkalkptl., Din., Wiirzburg, 1904, p. 24.-Areschoug, Trap. viixt. bladbyggn., Sv.
Vet. Akad. HandL, 39, D. 3, 1905, pp. 38, 39 (Erythroxylon).-Tbeorin, Vaxttricbom' a Arkiv fOr
Bot., iy. n. 18, 1905, p. 7.-[0. E. Schulz, Erythroxylaceae, in Pflanzenreich, Heft 29, 1~7,
pp • .ot--6].-[Tammes, Flachsstengel, N atk. Verb. bolland. Mij. Wet., 1907, 285 pp.]

HUMIRIACEAE (pp. 160, 16r).

Colozza's 1 investigation of the structure of the leaf in the species of Humi·
riaceae contained in the Florence Herbarium has afforded the following ad·
ditional facts. The leaves are invariably bifacial in structure. In Jlumiria
and Vantanea the cells of ihe upper epidennis are larger than those of the lower,
and are differentiated as aqueous tissue; in Sacoglottis, on the other hand, the.
. epidermal cells on both sides of the leaf are small. In H umiria the cells of the
palisade.tissue are much elongated, whilst in Vanta1fea and Sacoglottis they
are not so long, or even scarcely elongated; the gehus last named has 2-3
layers of palisade. In Sacoglottis cuspidata, Urb. and S. guianensis, Benth ..
em. the mesophyll contains numerous sclerenchymatous idioblasts which run 9

from OJie epidermis to the other and have swollen or forked terminations.
The median vein in all the members of the Order includes a stele with an
enveloping ring of mechanical tissue. In certain species of SacoglotUs the leaves
bear short, conical unicellular clothing hairs.
For details as to the·. structure of the petiole, the transverse section of
which is circular in Vamanea, circular or semicircular in Sacoglottis, ,and trian·
gular in Humiria, see Colozza, loco cit.
According to Pitard t:, a composite and continuous ring of sclerenchyma in
the pericyc1e of the aDS is present also ill A1tb"ya gabonensis and in additional
-speCIes of Humi"ia and Vantanea. ./

MALPIGHIACEAE (pp. I61-167).

3. STRUCTURE OF THE AXIS. A composite and continuous ring of scleren-
chyma is developed in the pericycle in Heteropteris laurilolia (Pitard). Pierre's
statement (in Flore forest. de la Cochinchine, xvii, r892; see also xviii, 1893)
regarding tp.~ 9c_c~ence of 'poches a. contenu blanc special' in the cortex
of the branch of Aspidopterys costulata, Pierre, requires further investigation;
the contents 'of the sacs are stated to be coloured blue by iodine.
Note. In BibI. bot., Heft 56, 1902. p. 77 and Tab. vii-viii, Areschoug publishes
the results of examination of leaves belonging to a plant originally referred to
Derris uligt.'nosa. Benth., but subsequently stated by the same authority (in Flora,
1903. p. 302) to belcwg to Tristellate1'(J austt'alasiae, A. Rich. (Malpighiaceae); since
the leaves in question bear peculiar three-:celled trichomes of a simple type, the
ma.terial is certainly not referable to a member of the Malpighiaceae.

Colozza, Note anat. sulle roglie delle Humiriae., Nnovo Glom. bot. ttal., xi, 1904, pp. 235--45'
Pitard, Peri cycle, These, Bordeaux, 1901 , pp. 77, 78.
SOLEUDh 3. 1
 ADDENDA-MALPIGHIACEAE
Literature: HohneI, Gerberinden, .Berlin, 1880, p. Il3 et seq.-Pifard, Pericycle, These,
Bordeaux, 19()1, pp. 40 and 67.-Arescboug. Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39,
n. 2, 1905, P_E. 35, 26.-Viret, Liaisons d\l phloeme med. etc., Inst. de Bot. Geneve, 190-4, pp. 46 - 6 3
(Duella).-LFor additionalliteraturt see p. IIp.)

ZYGOPHYLLEAE (pp. 167-169).

STRUCTURE OF THE LEAF. In Nitraria Schoberi, L. and N. tridentata,
Desi., the mesophyll contains mucilage· cells like those found in N. retusa';
tanniniferous idioblasts have, however, only been obserVed in N. Schoberi
and N. retusa (Jonsson). We may add that according to the same author'
gelatinization occurs in the axis in N. Schoberi and leads to the formation of
large mucilage-lacunae, situated in the primary and secondary cortex. These
lacunae generally originate as intercellular spaces, and undergo subsequent
enlargement at the expense of the cells of the surrounding tissue; after this
process of resorption the contents of the lacunae in some cases include large
numbers of crystals of oxalate of lime. In the species of Fagonia, Guaiacum,
La"ea, POTlieTia, Tribulus and Zygophyllum investigated by Pantanelli t the
t

stomata are small and are either sunk or raised above the level of the epidermis;
they have no subsidiary cells. The cuticle of the t;;pidermis varies very much
in thickness. A specially noteworthy feature is the occurrence of spicular fibres
in the mesophyll in Nitraria Schoberi; these fibres branch off from the
sclerenchyma of the veins, and traverse the palisade-tissue on the upper
side of the leaf, penetrating as far as the epidermis (Jonsson).
According to Pantanelli, the fibrovascular system of the petiole is con·
stituted by: (a) a principal system composed of 3-4 (Fagont.·a) or 4-6 (Tribulus)
vascular bundles or of a ring of bundles (Zygophyllum, Porlieria, G1faiacum,
Bulnesia, Larrea); and (b) an auxiliary system consisting of two lateral vascu]ar
bundles, which are situated towards the upper side.
For the structure of the petiolar cushions of Porlieria hygrometra and
Guaiacum otficinale, L., see the papers by Paoletti, Pantanelli, and Rodrigue.
STRUCTURE OF THE AXIS. According to Pantanelli, the cork is in most
cases (including Zygophyllum album, L.) developed in a subepidermal posi6on,
but in Fagonia cretica, L. it arises on the inner side of the pericyc1ic groups of
bast-fibres; the latter feature has by the way been previously recorded by
Vesque (in Ann. sc. nat., ser. 6, t. ii, p. 194). In Bulnesia Retamo the primary
cortex contains stone-cells (Pantanelli), whilst in Larrea nitida there is a com-
posite :lnd continuous ring of sc1erenchyma in the pericyc1e (Pitard). In
Fagonia, Trib'ulus and Zygophyllum the pith is composed of a thin-walled
tissue serving the purpose of water-storage'; in P01'lieria it is made up of
coarsely punctate celis, while in Guaiacum officinale it includes stone-cells
(Pantanelli). According to Pantanelli, a tier-like structure is exhibited by the
xylem-mass also in Larrea cune~'lolz'a, Cav.
The vascular system of the root is diarch (Fagonia, Tribulus, and Zygophyllum)
~r triarch (Guaiacum" Pot#e,,~·a).
Literature: Paoletti, Por/uria nygroml!Jral Malpighia, iv, 1890, PP. 34-40, and Nuovo Giom.
bot. Ital., 1892, p. 65 et seq., especially pp. 68-71.-Wilion, Leaves and stipules of Larrea ml!n-
cana, Transact. and Proceed. Bot. Soc. Edin burgh, xix, J 893. pp. 18 5-90.-Gamper, Angosturadnden,
Disa., ZUrich, 1900, p. 60.-Pantanelli, Anat. fisiol delle Zygopbyll .• Atti della Societa dei natural.
e matemat. di Modena, ser. iv, vol. ii, 1900, pp. 93-181, tav. viii-xi; and Pulvini_ motori etc., loco
cit., p. 201 et ~.-PitardJ P~ricycle. These. Bordeaux, 1901, p. 76.-Jonsson, Anat. Bau d.
WustenpB., Lunds Univers. Arsskr., xxxviii, 1902, Afd. 2, n. 6, p. :u et seq, and Tab. ii-iii.-
Plitzold, Harz u. Hob von Guaiacum, Diss., Strassburg, 1902, 122 pp.-Rodrigue, Poriieria "J'~O­
metra [Arch. Sc. phys. et nat. Soc. helv., 1902, pp. 140-2; Actes Soc. helv. sc. nat., 190~, p. '2];
and .Bull. de l'Herbier Boissier, ii, 1902, p. 893.-[Holterm.ann, Einfluss d. Klimas, 1907, p. 83
(TnDulus).]
 .ADDENDA

GERANIACEAE (pp. 16g-174).

I. The REYlEW OF ANATOMICAL FEATURES requires the following additions.
The heads of the glandular hairs are for the most part unicellu1ar, but in ex·
ceptional cases (species of Oxalis) they may be multicellular. Glandular
shaggy hairs.occur in Biebersteinia. In Rhynchotheca oxalate of lime is deposited
in the form of styloids, which had hitherto not been recorded in this Order.
Secretory ~ arl1l found side by side with the secretory cavities in species of
Oxalis. The epidermis of the leaf in Tropaeolum maiust L. includes elongated
cells with mucilaginous contents. Septate laticiferous tubes (?) are present
in the tuberous roots of a few species of T1'opaeolum. Hard bast is occasionally
wanting in the pericycle.
2. The STRUCTURE OF THE LEAF in the Geraniaceae has recently been
studied in detail as 'follows: by Brunies in Biebersteinia, M onsonia, Sanftaulon,
Geranium, Erodium, Pelargonium, V iviania, Rhynchotheca, Wendtia, Balbisia.
and Dirachmia; by Magnus·in Tropaeolum; by Chauvel in the genera Hypseo.
charis, Oxalis, Biophytum, Eichleria, Ave"hoa, Connaropsis and Dapania,
which belong to the Oxalideae; and by Brunotte in Impatiens. In many
species of Oxalis, e. g. O. articulata, Sav. or O. montevidensis, Prog. (on the
upper side of the leaf) and O. brasiliensis, Lodd. (on both sides of the leaf),
the epidermis consists of remarkably large cells and is differentiated as aqueous
tissue. Hypoderm is found beneath the upper epidermis in Dapania scandtns,
Stapf. Papillose differentiation of the lower epidermis has recently bee'n
recorded also in Eichleria Blanchetiana, Frog., Rhymhotheca spinosa, R. et P.
(poorly developed), Oxalis articulata, Sav., O. corniculata, L. var. purpu¥ca,
O. halophylla, Arech., O. hirta, L. and O. Osteni, Arech. In Tropaeolum peTegri, ..
num, L. Chlorophyll is present in the epidermis. The stomata are placed at
different levels. In Averrhoa, Biophytum, and EichZeria, according to Chauv~l,
there are two neighbouring cells, or at least one such ce~ lying par.alIel to the
pore. A study of the course of development of the stomata in Tropaeolum
has shown that three neighbouring cells are formed before the moth'er-cell of
the guard-cells becomes differentiated (Magnus). Water-pores functioning as
hydathodes occur also in certain species of Oxalis, and are situated in the de-
pressions of the emarginate leaflets. The neighbouring cells of the water-pores
in Oxalis like those of TTopaeolum are occasionally (e. g. in O. Scn,ade7iana,
H. B. K.) papillose. The mesophyll is for the most part bifacial~ although
centric (homogeneous) structure, with uniform assimilatory tissue composed of
rounded cells, is also met with. Balbisia mic,ophyUa, Phil. has rolled leaves,
~e stomata being contained in two furrows, situated one on either side of the
principal vein on the lower side of the leaf. In Pelargonium co,onopifolium,
Jacq. the two halves of the leaf are revolute, so that a de€!p furrow is formed";
the stomatal in this species, ho~ever, occur in approximately equal numbers
on both sides~ of the~leaf. With reference to the veins we may mention that
enlarged terminal tracheids are characteristic of A ve"hoa, Biophytum and
Eichle,ia. According to Irgang, the drops, which appear on pricking or cutting
through the veins of the leaf, the petioles and relatively yolJD.g stems of Tropaeo.
lum majus, emanate from tubular cells situated in the xylem and provided with
rather wide lumina, a nucleus and protoplasmic contents; these cells are
merely the young segments of the vessels, which in this case persist in an
unaltered condition for an exceptionally long time.
We may now pass on to consider the clothing hairs. Unicellular and unj·
seriate tncho~es in some cases occur in the same species. According to
Brnnies, the Geraniaceae investigated by him have clothing hairs in which
the basal portion has a characteristic structure; it is inserted between the
3 12
 ADDENDA--GERANIACEAE
epidermal cells and is generally conical or shaped like a truncated cone" and only
in rare cases has a cylindrical form. As specially noteworthy,'we may mention
the unicellular trichomes, which Fredrikson met with on the bulb-scales in
certain species of OxaJis; the longitudinal wall of these hairs has an uneven
surface due to small (0. incarnata, L., lac. cit., Tab. I, fig. IS) or spinose (0. sp.
349, lac. cit., Tab. II, fig. 25) papillae. At this point we may also refer to the
short trichomes found in Impatiens J.l1ar~·annae, Reichb.; in their typical foml
these hairs consist of a large basal cell and a plano-convex lens-shaped cell, but
they occasionally exhibit transitions to longer uniseriate hairs or may be reduced
to a single large epidermal cell, the outer wall of 'which is arched outwards in
a spherical manner. These structures are regarded by Haberlandt as local
organs for light-perception (ocellae). Glandular hairs having a stalk of varying
length and a unicellular head are found also in the genera Averrhoa, Biophytum,
M onsonia, V iviania and U' endtia. The following special types of glandular hairs
have been newly recorded: unicellu1ar clavate hairs (Oxalis rhcmbifolia, Jacq.
·according to Chauvel, and O. sp. 346 according to Fredrikson); glandular hairs
with a very long stalk and a unicellular head (0. re/racta, St. Hil. var. tYPica
and O. suocorymoosa, Arech. according to Chauvel); hairs with a long stalk
and an ellipsoidal head, subdivided into three cells by means of horizontal walls
(0. amara, St. Hil. according to Chauvel); lastly, the glandular shaggy hairs
of Biebersteinia multifida, DC., described by Brunies as emergences, which
consist of a long multiseriate stalk and, a knob-shaped head composed of
numerous cells. Regarding the structure of the glandular appendages (extra-
floral'nectaries) occurring on the petiole and stem in the species of Impatiens,
see also Aufrecht and Brunotte, lac. cit.
Oxalate of lime is also deposited in' the form of styloids 1 (parenchyma of
the veins of Rhynchotheca spinosa). Large idioblasts containing clustered
crystals are found in the mesophyU in many species of Erod~tm, and in Pelar-
gonium anatymb£cum, Steud., while idioblasts with solitary crystals are met with
in certain species of Oxalis. Other features of special note are: the sphaero-
crystalline or clustered aggregates found in species of M onsonia and Erodium ;
the short, moniliform rows of cells filled with clustered crystals occurring in
the veins of the leaf in species of Wendtia; and the rows of crystal-cel1s (with
solitary crystals) accompanying the vascular bundles of the veins to which they
form a kind of sheath in Averrhoa, Biophytum and Eichleria.
Chauvel, Fenizia and Fredrikson have published new data on the dis-
tribution of the secretory cavities in Oxalis; Fredrikson, however, onlyex-
amined the bulb-scales. According to Chauvel, the secretory cavities are a
constant feature in the acauline species, although they also occur in certain
species which have a well-developed stem (e. g. O. cernua, Thunb. or O. Deppei,
Lodd.). The secretory cavities found in the bulb·scales are situated on the
dorsal side of the vascular bundles, and are often elongated so as to resemble
canals. Those occurring in the foliage-leaves are chiefly marginal in position;
more rarely they are distributed over the entire surface of the leaf (e. g. in O.
artz'culata, Sav. and O. hirta, L.), while in other cases again (e. g. in O. Deppei)
there is a single large secretory cavity at the end of the median vein. According
to Chauvel, the moqe of development of these glands is lysigenous and not
schizogenous, as was formerly stated. Chauvel failed to observe a lining layer

1 It may be pointed ont here that Hohnel met with' prismatic crystals- of oxalate o{lime, of
exceptionally large size and rhombic in section' (presumably also styloids) in Chureo bark, which is
imported from Chili and is rich in tannin; this bark is stated by Hohnel and Wiesner to be derived
from Oxa/iJ gigantea, Barn., while according to the (ormer authority its structure is quite identical
with tba.t of the cortex of O. Orlgt"esit". We may also note that according to Knothe the mesophyll
of Oxalis articulata, SaY. contains 'twin-crystals of calcium sulphate showing the well-known
swallow-tail form' (probll-bly also equivaIent to styloids of calcium oulate).
 ADDENDA--GERANIACEAE
of definite epithelial cells in the secretory cavities as described by Fredrikson.
The bulb-scales of certain species of Oxalis contain secretory €ells side by side
with secretory cavities, while in other £pecies of this genus secretory cells alone
are :present (Fredrikson). Magnus's statement as to the occurrence of anasto-
mosmg septat~ laticiferou5 tubes in the rooi-tubers of Tropaeolum brachyceras,
Hook. and T. Leichtlini, Herb. Kew, still awaits confirmation. In Tropaeolum
maius, L. the epidermis on both sides of the leaf contains tubular cells;\vhich
Irgang describes as mucilage-cells. Myrosin-cells are wanting in Oxalis
(Chauvel).
STRUCTURE OF THE PETIOLE. According to Brunies, medullary vascular
bundles (with centr:al phloem) are not present in all the species of Pelargonium.
In Tropaeolum the petiole contains isolated vascular bundles. In Averrhoa,
Biophytum, Co nnaropsis,. Eichleria and Tropaeolum the mechanical tissue is
developed in the form of a sclerenchymatous ring, while in the caulescent
species of Oxalis it appears as bundles of fibres developed in relati9n to the
individual vascular strands.
For the systematic anatomy of the bulb-scales of Oxalis see Fredrikson,
and also Chauvel.
3. STRUCTURE OF THE AXIS. Amongst the ~oody members of the Order,
Averrhoa, Connaropsis, Dapania and Eichleria possess pericyclic groups of bast-
fibres, forming a more or less closed mechanical ring. Dapania scandens shows
bands of secondary hard bast, while the pith of the same plant contains cells of
the nature of bast-fibres .
. Development of hard bast in the pericycIe is met with amongst the her-
baceous forms also in species of Biophytum, Hypseocharis, Oxalis and T,opae.o-
lum. In TrC!Pae;olttm peregrinum, L. the cork arises in the endodermis (Magnus).
Literature; Moller, Gerberinden, Berlin, 1880, pp. 124-7 (Churco-bark).-Costantin, Tiges
aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, 'p. 104 et seq.-Hildebrand, Schutzeinricht. bei den
Oxalis-Zwiebeln, Ber. deutsch. bot. Gesellsch.• 1884, p. Io8.-[Acqua, in Ann. R. Ist. bot. Roma,
1887.]-Aufrecht, Extraflorale Nekt., Diss., Ziirich, IS92, p. 20 et seq. (Impatiens glanduligera).-
[Drobnig, WUTzel-Knollen, Diss., Rostock, 1892, p. 49 et seq. (Oxa/is).]-Guignard, Principes.
act., Comptes rendus, Paris, cxvii, 1893, pp. 587 a.nd 751 et seq.-[Tognini, Stomi, Atti 1st.
bot. Pavia, I 894.]-Fredrikson, Anatom.-syst. Stnd.ofver Lokstammiga Oxalis·arter, A¥ad. Alb.
Upsala, 1894-5, 67 pp., Tab. i-ii.-Jonsson, Anat. Bau d. BI., Acta Univ. Lund, xx:di, 2,
J896.-Magnos Gust., Beitr. z. Anat. der Tropaeoiaceae, Diss., Heidelberg, 18g8 , 50 pp.-
Schwendener, Gelenkpolster von Phaseo/us u. Oxalis, S:tz.-Ber. Berliner Akad., 1898,- xii, p. '76
et seq.-Spanjer, Wasserapparate, Bot. Zeit., 1898, i, p. 54.-Brunics, Anal. d. Geraniaceen-Blatter,
Diss., Breslau, I~OO, 40 pp., 1 Tab:-Brunotte, Recb. ernbr. et anat. sur qu. esp; d'lmpatints et
Tropaeolmn, These, Paris, 1900, 178 pp., 10 pl.-Kliem, Regenerationsorg., Diss., Erlangeri, 1900,
p. I1.-Schleichert, Xerophyten bei Jena, Naturwiss. Wochenschr., 1900, p. 4 ....9 (G~ranium).­
Tunmann, Sekretdriisen, Diss., Bern, 1900, p. 23.-[ Arechavaleta, Fl. Vrug., Ann. ,Mus. rulC. de
Montevideo, iii, 19°1, p. 189 et seq.; cited from Chauvel.]-Haberlandt, Sinnesorgane, 1901, p. 88
et seq.-Molisch, Milchsaft u. Schleimsaft, 1901, fl. q ......:.Pitard, Pericycle, These, Bordeaux,
. 1901, pp. 48, 50, and 75.-Bargagli·Petrncci, Legnami, Malpighia, 19°2, p. 313 (Connaropsis).-
Buchenal1, Tropaeolaceae, in Pflanzenreich, Heft JO, 1902, pp. 4, :;.-[Fenizia, Corpnsculi resinosi
colorati nell' Oxalis esotiche, Riv. hal. Sc. nat., 1902, pp. 52 and 83; abstr. in Bot. Centralbl.,
xciii, p. 337.]-Hohlke, in Beih. bot. Centralbl., xi, 1902, p. 42.-Irgaug, Saftausscbeid. Elem. u.
Idiobluten bei Tr()pae()ltmi·'niaiiis~ Sitz.-Rer. Wiener Akad., exi, Abt. I, 1902, 9 pp., I Tab.-
Knothe, Unbeneub. -Blatter, Diss., Heidelberg, 1902, pp. 10, 1I.-Knoth, Geranium, in Engler,
Bot. Jahrb., xxxii; 1902, p. 190 et seq.-(Preston, in Bot. Gaz., xxxiii, 1902, pp. 150-.... j abstr. in
Bot. Centralbl, lxxxix, p. 516.]-[Ramaley, Trich. stntct. of Erodium cicutariuJn, Bot. Gaz., xxxiii,
1902, pp, 140-2.]-Chauvel, Rech. sur ia fam. des Oxalidacees, These, Paris, 1903, 205 pp.-[Van
Tieghem, in Bull., Mus. d'hist. nat., ix, 19°3, p. 287.]-Col, in Ann. sc. nat., ser. 8, t. xx, '904,
p. 128.-[Dore, Et. bot., chim. et phann. des Germdum at/antimm et C. matulattlm, These,
Toulouse, 1904-)-Haberlandt, Lichtsinnesorg., 1904, p. ]]2 et seq., and Tab. iii..;-Si.i.ssenguth,
Bebaarungsverh. d. Wiir.zb: Muschelkalkpfl., Diss., Wiirzburg, 19°4, p. 2+-Theorin, Vaxttricbom.,
Arkiv for Bot., iii, n. S, 19°4. p. ii; see also lac. cit., iv, n. 18, J905, p. 7.-Netolitzky, Dikotylenbl.
(Rbaphiden), 19°5, p. 33.-Porsch, Spaltoffnungstypus, lena, 1905, pp. II7-19 and Tab. iv..!-.
Sarton, Rech. expo sur I'anat. des pl. affines, Ann. sc. nat., ser. 9, t. ii, 19°5, Pp. 3.~ and 95 et seq.
(Ge:o-anium).-[Holtermann, Einfluss d. Klimas, '907, p. 217 (Impatiens fflacrophyila).]
 ADDENDA

RUTACEAE (pp. 174-182).

I. The following special characters may be added to the REVIEW OF THE
ANATOMICAL FEATURES ': glands, which do not project in the form of hairs, in
certain,species of BOTonia; sphaerocrystalline masses of varying chemical
composition (partly besperidin); branched sclerenchymatous cells (species of
Boronia) and enlarged terminal tra.cheids (Phebalium) in the mesophyll ;, groups
of fibrous cells situated in the pith of Evodia /I'Qzini/olia, Hook. f. The ordinary
clothing hairs are unicellular or uniseriate. In a few members of the Order
additional records of the differentiation of epidermal papillae or of the pre-
sence of hypoderm in the leaf have recently been published.
2. STRUCTURE OF THE LEAF. Schulze has recently investigated,the struc-
ture of the leaf iiI a large number of genera 1. The subsequent·description is
based on his work, as well as on the remaining papers .cited below. EpIdermal
cells with mucilaginous inner memQranes have been recorded in certain species,
of Acmadenia, Adenandra, Agathosma, Barosma, Coleonema, Diosma, EmpleuTum,
Flindersia, Macrostylis, Phellodendron, Skimmia, Toddalia and Zanthoxylon.
HeSperidin is not only found in the epidermis of the species of Barosma, for,
according to Schulze, It occurs in the same position also in Agathosma biophylla,
E. et Z., Calodendton capense, Thunb., Dictamnus Fraxinella, Per~i., Empleurum
ensatum, E. et Z., Ptelea trifoliata, L., Skimmia japonica, Thunb., Toddalia,
aculeata, Lam., Zanthoxylon fraxineum~ WiUd. and Z. Pterota, H. B. K., and,
according to Duval, in Pilocarpus Goudatianus, Tul., P. pennatifolius, Lem.,
P. trackylophus, Hemsl. and Zanthoxylon el.egans, Engl. According to Schulze,
as may be pointed out here, the sphaetocrystalline masses, observed by Schaal'S·
schmidt in alcohol-material of Ruta (Haplophyllum) Bt'ebersteinii, Neilr., and
regarded by him as inulin, consist neither of inulip nor of hesperidin; Geiger also
states that the crystalline aggregates, which are found in the epidermal cells'
of Pilocarpus trachyphyllus, Holmes and other species of this genus (also P.
pennatifolius, Lem.) and which have a'tufted, rayed or racemose structure,
cannot with certainty be regarded as consisting of hesperidin. In Eriostemon
saUcifolius, Sm. the upper epidermis consists locally of two layers. Hypoderm
is foun~ according to Duval, in P#ocarpus giganteus, EngI (composed of 2-3
layers), P. Goudotianus, Tul. (one layered) and P.latifolius, St. Hil. (1-2layers),
and, according to my own observation, in Pagetia medicinalis, F. v. M. (one-
layered) ; it occurs also in the neighbourhood of the larger veins in Evodia
obtusifoUa, DC.? and in the petiole of Ruta graveolens, L. Papillose differentia-
tion of the epidermal cells has been observed also in Boenni1Lgha,usenia albi/lora,
Rchb. (on both sides of the leaf) and Eriostemon salicifoliu5, Sm. (only on
. the upper side); according to Duval- (loc. cit., pp. 119 and 37 et seq.). and
other authorities, it is found i.n species of Pilocarpus as well.,.t::."..In Adenandra
the cUticle exhibits slight papillose irregularities. Schulze states that the
upper epidermis in Murraya exotica,.L. contains-peculiar groups of small cells,
and that below these groups the palisade-tissue is more strongly developed.
The stomata are more commonly found only on the lower side of the leaf
than on both sides. In most cases they are uniformly distributed over the
entire surface, but exceptions to this rule are furnished by Acmadenia and

1 vi", Cusparlcae: A liluitka , ErytlzrocMton, Ravia; Rllteae: Ruta, B(}tnningnausenia,

Did.mm41; Diosmeae: CaJ(J([endrffl, M(t(1'ostylis, Diosma, Coieanema, AC11lar/enia, Adena"ara.
BlnWma, Agrztwft1a. Empl~m; Boronieae: Zieria, Bortmia, Emstemon, Plubalium, C~;
ZaIlthoxyleae: Evodi., Cklisya, Zantltoxylm, Pi/()(arpus; Toddalieae: Toddalia. Plullode""""",
Plew. Skimmia; Aurantieae: MIINrZya, Parrznll~ Cilr«r; also the genus Flinderri'a, which
was fonnerly placed amongst the Meliaceae.
 ADDENDA--RUTACEAE Sss
Coleonemet, in which they are confined to a narrow median zone on both sides
of the leaf, and by Macrostylis, in which they occur only on two narrow strips,
situated to the nght and left of the midrib on the lower surface of the leaf.
Schulze describes the occurrence of subsidiary cells to the" stomata in the
following species: Citrus trifoliata, L. (4-5 subsidiary cells), E,iostemon buxi-
folius, Sm. (1-2 subsidiary. cells on either side of the pore and placed parallel
to it)·, Crowea saligna (two or more distinct subsidiary cells occasionally present),
Murraya exotica, L. (4-5 subsidiary cells), Pilocarpus pennatijolius, Lem. (s sub-
sidiary cells), Paramignya sp. (4-5 subsidiary cells), Ravia resinosa, Nees et
Mart. (z-4subsidiary cells), Skimmia faponica, Thunb. (subsidiary cells arranged
to form a rosette). The stomata of Erythrochiton brasiliense, Nees et Mart.
require special mention; in this species there is a single neighbouring cell
situated on either side of and parallel to the pore, while in surface-view a narrow
crescent~shaped area, which is faintly rose~coloured,.is superposed .on each
guard. cell ; the latter phenomenon is due to interference of light causing the
cell·membranes, which are thin at these points, to appear red. The meso-
phyU is bifacial or centric. In the leaves of Agathosma ledifoTmis, E. et Z.,
which are adpressed to the axes, the mesophyll exhibits an inversion of the
ordinary anatomical structure, since the palisade-tissue situated on the mor-
phologically,lower side is more strongly developed than on the upper side.
Schulze met with branched sc1erenchymatous cells, generally developed in
relation to the terminations of the veins, in Boronia crenulata, Sm., B. elatioT,
Bartl., B. ledifolia, Gay, and B. serruiata, Sm.; enlarged'terminal tracheids
have been observed in Phebalium, and stone-cells in the conjunctive paren-
chyma of the principal veins' in Almeidea rubra, St. Hi!.
Our knowledge as to the mode of deposition of oxalate of lime has been
extended by the following additional facts. Solitary crystals occur L."l the
epidermis of the leaf in Flinders-ia australis, R. Br.; the epidermal cells in this
case either contain·a single crystal inserted.in the thickened wall or, undergo
subdivision by irregularly orientated walls into a number of chambers, each
of which includes a crystal. On.the other hand, according to the investigations
of Pfitzer and Guttenberg the well-known crystal-cells of Citrus, (see also
Penzig, loco cit.), which are met with in surface sections of the leaves, are not,
as stated on .p. 176, epidermal structures, but belong to the gronnd tissue; it is
only in the course of subsequent growth that they push their way.between the
epidennal cells. Crystal-cells occupying the same position as-those of.Citrus are
found also in Atalantia buxifolia and Paramignya sp. According to Geiger
(see also Duval), features characteristic of (perhaps all) the species of the genus
Pilocarpus are the occurrence of transversely septate palisade-cells with
clustered crystals in the chambers, and the blocking up of the 'respiratory
cavitie() by small cells containing clustered c'rystals of oxalate of lime.
The . following features have been recently observed in connexion with the
trichomes. Erythrochiton brasiliensis, Nees et Mart. pas unicellular glandular
hairs, which are commonly situated above the secretory .cavities 'and have
a sphericarshape:--External glands with a clavate or spherical multicellular
head are found in Choisya ternata, Kth., Esenbeckia febrituga, Juss. (according
to Duval), Monniera trifolia, L. (according to Duval), Peganum Harmala, L.,
species of Pilocarpus, Zanthoxylon Pterota, H. B. K. and other species of this
genus, and Zieria lanceolata, R. Br. We may class with the glandular hairs
the glandular structures, commonly found on the lower side of the leaf in
Boronia crenulata, Sm., although only of isolated occurrence in B. datttJil, Bartl. ;
these glands are not however differentiated as hairs, ·being even slig'1ltly sunk
below the surface. In surface sections they appear as small round areas,
composed of. a small·celled tissue and surrounded by a .rosette of 4-5 narrow
epidermal cells. In transverse. sections of. the leaf these glands exhibit a thick
I
 ADDENDA--RUTACEAE
outer wall with a subjacent secretory tissue consisting of 2-3 layers of palisade M

like cells with thin walls,. and below that 2-3 further layers of rounded cells
with slightly thickened walls.
Schulze records stellate hairs also in Boronia ledi/olia, Gay and Crowea
saligna, Sm., -while those found in species of Correa and Zieria have been sub-
jected to detailed examination by the same investigator; in Crowea saligna
the hairs are of such small dimensions that the surface of the leaf appears smooth.
The ordinary clothing hairs are unicellular or uniseriate 1. .
In marlY of the genera Haberlandt has demonstrated a special mechanism
in the 'secretory' cavities II serving the. purpose of excretion. The glands, are
provided with two or more (mostly four) epidermal cells of peculiar shape,
which are differentiated as lid-cells, and the lateral walls of which are specially
modified. The turgescence of the cells forming the wall of the gland, and
-the' consequent pressure exerted on the secretion, together with movements
resulting in the bending of the leaf-surface, lead to the formation of clefts
between the lid-cells, through which emission of the secretion takes place.
According to Schulze, however, the lid-cells are not-present in all the members
-of the Order investigated by him. As regards the distribution of the secretory
cavitir.5 we may first notice that, according to Pierre, they occur also in Thorel-
d()'1a 'cochinchinensis, Pierre. Schulze's statement that Zanthoxylon (Fagara)
Pterota has no· secretory ca.vities- and only possesses secretory cells requires
some modification, the secretory cavities in this species being confined to the
notches between the leaf-teeth ~ the secretory cells are found both in the
bast and in the conjunctive parenchyma of the veins. Schulze again discusses
the question whether the secretory structures found in Pagetia are of the nature
of cavities or ceils, and in this relation, I may mention~ that I have recently
investigated an original specimen of P. medicinalis, F. v. M., and that it posses£es
secretory cavities and no secretory cells 3.
In most of the species examined by Schulze the petiOle contains a ring of
wood and bast.
3. STRUCTURE OF THE AXIS. -The description of the structure of the
cortex requires the following additions. The cork arises in the subepidermal
layer in P.ilocarpus~ and not in the epidermis, as was formerly stated (Geiger).;
cork.development also takes place subepidermally in Murraya exotica (Laborde).
The cells of the cork are thin in Galipea and Cusparia, while in Esenbeckia the
inner tangential walls are strongly thickened (Gamper); Toddalia has cork-
cells thickened in the form of a horseshoe (Bocquillon). The literature cited
below also furnishes additional data on the occurrence of s tone-cells and of
secondary hard bast. The presence of groups of fibrous cells in the pith of
Evodia jraxinijolia, Hook. f. requires special mention (Bocquillon).
Bocqtilll6n'g paper contains a few important facts about the secretory re·
ceptaclcs found i::i the ,axis. Secretcr"j caVIties are present in the pith also in
Evodia Iraxinifolia, Hook. f., Toddal£a aeuleata, Pers., T. paniculata. Lam. and
Zanthoxylon hyemale, St. Hi!. The' lacunes • recorded in the primary cortex in
certain species of . Zanthoxylon~ and the' lal'ges lacunes ' found in the wood in Zan-
thoxylon Budrunga, Wall. are no doubt in view of my earlier statements (p. 181) oi the
nature of mucilage-spaces. The' glandes oleiferes,' stated to occur in the bark in

, 1 Duval's ,statement (loe. cit., p. lf9, cf. p. :37) as to the occurrence of mniticellular clothing
hairs in PUocarpu! pemuzt'loli.ls, Lem., arid P ..Sel/oa1lus, Engler, is no, 'doubt only a misprint.
Uniseriate clothing hairs are, however, fOWld in this Order, c. g. in species of Monniera and
Zant!t0xy/oH. _ .
:ISecretory cavities bave I),ot been obse:-ved in the root of the Rutaceae (Van Tieghem).
. I The statements published by Tschirch'a pupils (recently Stepowski, and formerly Bechuaz and
Sieck) as to the occurrence of medullary resin-canals in Amyris balsamiflra probably depend on
in~rte~t determinati.on, of the materi.al used {OT the investigation.
 ADDENDA--RUTACEAE
many species, probably in all cases represent oil-cells; the statement as to the
presence of secretory cavities in the bast in Zanthoxylon Pentanome, DC. is certainly
Incorrect. What BocquilJon means by • nodules secreteurs,' described as occurring
in the pith in Z. Tingoassuiba, St. Hil. and Z. a/atum, Roxb., must be made the
subject of further investigation.
For the structure of the spines occurnng in the species of Zanthoxylon,
see Barber, Bot:quillon, Lothelier and Mitlacher; these spines subsequently
become elevated on a corky excrescence, which undergoes continual increase
in girth at its base (the same phenomenon moreover is met with also in Tod~
dalia aculeata). The structure of the leaf-spines of Citrus is dealt with by Mitt·
mann, that of the stem-spines of Maclura by Lothelier.
Regarding the occurrence of a • reseau de soutien' in the subepidermal
layer of the root in Choisya ternata, see Boudouresques, loco cit.
Literature: Penzig, Apparecchi ilIuminat., Atti Soc. dei Naturnlisti di Modena, Rendiconti,
1884, pp. I06-I,.-[Semenow, Pernambuco-Jaborandi, Zeitschr. d. Pharm., 1888, p. 67.1-Mittmann,
Anat. d. Pflanzenstach., Verh. bot. Ver. Brandenburg, 1889, p. 52.-C. de CandoIle, ·Intloresc.
epiphj'HC5, Mem. Soc. de phys. et d'hist. nat. Geneve, 18ge; vol. snppl., sctl. copy, p. 24 et seq.-
Barbel', Cot,:ky excresc. on stems of Za1lthoxylcm, Ann. of Bot., vi, 1892, pp. 154-66 and PI. vii, viii.-
Lothelier, Epines, These, Paris, 1893, pp. IS. 30 and 34.-[Togr.inl, Stomi, Atti I!t. bot. Pavia,
J89+]-Boudonre-sques, Choi~a temata, These, Montpellier, 1895, PV. ~6-37.-Weigt, RalJelaisia.-
Rinde, Diss., Er1angen, 1895, pp. i7-22 and :01 Tab.-Zenetti, Hesperidin in Folia Bncco, Arch. do
Pharm., J895, pp. 10+-10, 2 Tab. (incorrect interpretation of the gelatinized epidermis in the leaf).-
Knob13Llcb, Okolog. Anat. etc., Habilitat.-Schr., TUbingen, 1896, p. 15 et seq.-P!erre, Flore forest.
de 1a Cochinchine xxii, 1896 (T horeldcra).-Elfstrand, Heilpfl., Ber. deutsch. phann. Gescllsch ..
1891. p. 302 Uaborandi).-[Dohme, Hist. and pharm. of Buch~ leaves, DrUggist's Cire. and
Chem. Gazette, ) 897, n. 7; after Bot. Cen tI;.al bl., 1898, ii, p. 93; contains an incorrect interpretation
of the gela.tinized epidermis.]-Laborde, Et. bot. P.t chim. des .lI-lu1'Ta;'a exoli,'a et M. KOhIzg#,
These, Toulouse, .897, pp. 2o-3.-[Schneid(![, The offic. Jaborandis, Journ. otPbarma.col, x, 1897,
ii, n. (}; cfter Bot. Ceot!'a1bl., Beihefte, vii, p. 51B; and Just, 1897, p. 97.]-Schubert, Paren-
chymscheirlen, Bot.' CenI ralbl., 1897, iv, p. J 6.-[Zancla, Acu1ei, Contribuz. 1st. bot. Palermu, ii,
1897, p. I et seq.]-Geiger, }ab01'anJ.i-Bl., D1SS., Z~Iic.h, 1898, 74 pp., 3 Tab.-Habetlandt,
Entleernngsapp. der inneren Driiren einigcr Rutacee.'1, Sitz.-Ber. Wiener Akad., .Ed. evii, Abt. I,
1898, pp. 1221.-46 and 2 Tab.; <;ee also Bot. CentralbL, 1899, i, p. 263; and Ost. Bot. Zeitschr.,
1899, p. 117.-Kohne, Papi!1. u. oberseit. Spalt5ffn., Mitten. deutsch.· dendrolog. Gesellsch., 1899,
p. 58.-Gamper, Angostuntrinden, DlSS., ZUrich, 1900, p. 16 et seq. nnd Tab. i-ii; see also Hartwich
and Gamper, in Arch. d. Phann., 236, 1900, p. 568.-Keamy, in Contrib. U. S. Nat. Herb., v, 5,
1,900, p. 295.-PantaneIli, Anat. fi!;. delle Zygopbyllacee, 1900, pp. 165-74 (Peganum).-Bocquillon,
Et. bot. t!t phlmn. des Xanthoxylees, These, Paris, 1901, J~8 pp., -4 pl.--[Mitlacher, Vergl. Anat.
einiger Riltllce.en- Rinden, Zeit~hr. aUg. ost. Apot.hek.- V er., 190 I, p. 2'25 et seq. j after Bot. Centralbl.,
lxxxix, p. :P4.}-Pitard, Pericycle, These, Bordeaux, 1.901, p. 7I.-Bollygnes, Petiole, These, Paris,
1902, p. 12.-GuUenberg, Krystallz. im nI. von C#rw, Sit7..-Ber. 'Viener Akad., Bd. xci, AM. I;
1901, pp. 855-12 and Tab.-Knothe-, Unbenetzh. Bl., Diss" Htioelbe-tg, 19°2, p. g.-Poulsen,
Bladkirtl. hos E,.ythrodlilon brasiliense, Vidensk. Meddelels. Kj~benhavn, 1902, pp. 239-42.-
H. Schulle, Beitr. z. Blattanat. d. Rutaceen. Diss., .Heidelberg, 1902,50 pp., 2 Tab. (sep. copy from
Beib.illol. Centralbl., xii. 1902, p. 55).-"\Vidern, Pharmakogn.·chem. Stud. 'tiber die Verbreit. des
Berberins, msb. in der Gatt. Zaillhoxylon, Diss., Strassburg, 1902, p. 54 et seq.-Achner, Falsche
Chinarinden, Diss., Dei'll! 1904, p. Sr.-Duval, Jaboralldis, 190.5, !30 pp., 10 pl., ir! Perrot, Travaux,
iii, I906.-Stepowski, Veg. Org. d. Bnrserac. etc., Dis:>., Bern, 1905, p. 47 et seq.-Pk:cioli, Legnami,
:Hun. Siena, 19(>6, p. (35.--[For additional literature! see p. II72.]

SIMARUBACEAE (pp. 18z-188).

7.. In the REVIEW OF ANATOMICAL FEATURES a number of conections and
additions are made necessary, chiefly by ] adin's Lomprehensive paper \ The
following speclal types of stomatal apparatus have recently been recorded:

1 Jaclin's -inyestigations deal with the stru.::ture of the leaf and axis in the followiug genera:
Quasst'a. Simaba, HamJoa, ~llalznia, Simarttba, Ailanthus, Sallladert;, Hyptiandra, Castela, Hola-
cantha, Picrasma, Brucea, PicI"olelllma, .E'ulycoma. ·Cadd/ia, Stlriana, SOlllamen, Amargria,
Irvingia, Kir}u'a, IIa1'Tis{Jm·a, Picnlla, Picramnia, .JV"·odnuln",, as well as Alvaratioa, KlaineatJ.xa,
Old},mduz, and Picrocardti:; in Amerada the leaf onl) was eltamined.
 ADDENDA-SIMARUBACEAE

stomata with subsidiary cells, placed parallel to the pore (Castela, lroingia,
Klainedoxa, Picrodend,on); stomata with 3-5 neighbouring cells resembling
subsidiary cells (Suriana); and stomata arranged in groups (Castela, Soulamea).
Suriana constitutes an exception as regards the mode of development of the
cork, the phellogen in this genus arising in the inner part of the pnmary cortex.
A composite and continuous ring of sclerenchyma in the pericycle is present
also in Rigiostachys and Samadera Harmandii, Pierre; in the genera Irvingia,
I rvingella, Desbordesia and Klainedoxa, which belong to the Irvingieae, there
is a similar ring, which differs however in including sclerosed parenchymatous
cells exhibiting U-shaped thickening; Castela has a fairly continuous and com-
posite ring. Secondary hard bast is of frequent occurrence. In Guilfoylia
the walls of the vessels bear simple pits in contact with parenchyma of the
medullary rays; in Rigiostachys, as well as in Bruneltia and N eopringlea, some
of the wood~fibres are septate. Jadin records the occurrence of resin-c~anals
at the periphery of the pith also in Eurycoma, Hannoa and Oldj'tmdea.
Uniseriate clothing hairs are found side by side with the unicellular hairs.
The deposition of oxalate of lime in the special form of sma.ll crystals of varied
shape, which are situated in the mesophyll or in the epidermIs of the leaf, is
met with also in the genera of the Surianeae (Caaellia, Guilfoylia, Rigiostachys
and Suriana). Resin-cells have been demonstrated,also in species of Irvingia
and in Oldyendea, as well as in the genus Chamaelea (Cne()TU1n pulverulentum,
Vent.), which Van Tieghem rightly separates from Cneorum. Mucilage-cells
or spaces are of general distribution in the genera of the Irvingieae (Irvingia,
Irvingella, Desbordesia and Klainedoxa), a..'1.d in the" neW genus Perrie-rea, which
.is closely related to Picrasma. Other specially noteworthy features are as
follows: The vertical transcurrence of the smaller veins of the leaf in I rvingia,
Klainedoxa and Pierodendron; the extrafloral nectaries on the petiole in
Cadellia, and on the midrib and occasionally on both surfaces of tbe leaf in
Samadera; the heterogeneous pith found in H arrisonia; the subepidermal
groups of fibrous cells in the stem of the leafless genus Holaeantha·; and the
• cristarque ' -cells situated in the primary cortex in the genera of the
Irvingieae. .
2. STRUCTURE OF THE LEAF. In most cases the leaf is bifacial in structure,
rarely (A ilanthtes exeelsa, Roxb., Chamaelea -p'tdverulenta, V. T. and SU1iana
maritima, L.) centric. Papillose differentiation of the lower epidermis is
found in the following additiona.l species: Ailanthus Fauveliana, Pierre)
A. imberbifolia, F. v. M., A. malabarica, DC., Euryeoma longifolium, Jack,
lrvingia Oliveri, Pierre (=lrvingella Oliveri, V. T.), Kirkia Wiltiamsii, Engl.,
Oldyc1Jdea gahonensis, Engl., O. Klaineana, Pi€!n'e, and, according to Van
Tieghem~ quite generally in Desbordesia and Irvingella. Gelatinization of the
epidermis of the leaf no doubt occurs in a relatively large number of genera,
e. g. in Castrla erecta, Turp. (Borgesen and Paulsen) and in the four genera of
the Irvingieae (Van Tieghem). Unfortunately Jadin placed an incorrect
interpretation on these gelatinized epidermal cells, as is cleady shown by his
statements regarding Picroca,.dia resinosa, Radlk. and by his figures; he
regarded the gelatinized cells as divided epidermal cells and the mucilaginous
membranes as hypodermai cells,-an error which is very widespread in the
literature dealing with systematic anatomy (e. g. also in Vignoli'"s paper with
reference to Irvingia Oliveri) and is met with over and over again. Jadin's
statements as to the occurrence of tangential division~walls·.in the epidermis
or as to the presence of hypoderm in species of Ailanthus, Ama1'oria, Castela,
Hannoa, Irvingia~ Kirkia, Klainedoxa, Oldyendea, Picrocardia and SouJamea
must therefore be accepted with reserve and require critical revision. The
stomata are provided with subsidiary cells in only a very few of the genera. In
lrvingia and Klainedoxa (but not i~ Irvi1'!geUa and Desbordesia), as well as in
 ADDENDA-SIMARUBACEAE

PiCf'odendron, the stomata have subsidiary cells, which are placed parallel to
the pore; according to Borgesen, and Paulsen the same is true of Castela. In
Suriana there are 3-5 neighbouring cells differentiated like subsidiary cells.
Stomata are found on both sides of the leaf also in Chamaelea pulverulenta,
V. T. and Suriana maritima, L. In Casteta depressa, Turp. each group of
three stom.ata is surrounded by four or five neighbouring cells; in most of the
species of Soutamea, if I understand Jarlin rightly, the stomata are. arranged
in groups (stomates ... rennis en pl~ges avec 5 ou 6 ceUules -de bordure '),
whilst in S. Pancheri, Brongn. et Gris they are contained in pits, like those of
Nerium. Jadin has recently observed spicular cells in the mesopbyl1 in the
following additional genera: Hyptiandra, Irvingia and Oldyendea 1. In
lrvingia gabonensis, BailI. occasional cells of the spongy tissue are strongly
thickened and qifferentiated as sclereids, while in Castela longitolia, Gris and
C. ereeta, Turp. the same feature is shown by isolated palisade cells. In ITVin~
gella (according to Van Tieghem, but not in Ir'llf.·ngr:a, V. T. em.), Klainedoxa
and Picrodendron the smaller veins are vertically transcurrent by means of
sclerenchyma.
Regarding the mode of deposition of oxalate of lime we may add the follow~
ing information. Small crystalline bodies, sometimes resembling clustered
crystals, are found in the mesophyll and occasionally in the epidermis also in
the remaining Surianeae (Suriana, Cadellia, Guiltoylia). Relatively large
idioblasts occupied by solitary (e. g. in-Picramnia) or clustered crystals (e.g. in·
Brucea) are frequently present in the,mesophyll. In addition to the unicellular
clothing hairs uniseriate hairs also occur in this Order (e. g. in Hyptiandl'a,
Picrocardia and Soulamea), while in Cadellia the unicellular hairs are accom~
panied by bicellular trichomes with a short basal.cell. Unicellular, two-armed
clotrung hairs baving the shape of a Tor Yare found only jn CneoTum puJveTu~
lentum (=Chamaelea pu!verutenta, V. T.), C. tricoccum merely having ordinary
unicellular clothing hairs. The only additional records of the occurrence of
multicellular external glands are those of Brucea sumatrana, Roxb. (according
to Jadin) and of Cneorum tricoccum, L. (according to Van Tieghem). In
connexion with our account of the glandular hairs we may. notice the extra-
floral nectaries (?) found on the petiole in Cadellia pentastylis, F. ~. M., and
dppearmg to the naked eye as callosities. Their epidermis consists of narrow
paIisade·like cells, the lumina of which terminate in the thick outer wall in
the fomi of a cone; beneath the epidermis there are two, or in the middle
of the nectary three, layers of cells, exhibiting greater elongation and having
thick lateral walls which are yellow in colour. The nectaries found in Samadera
have not yet been subjected to a close examination; they. occur in large numbers
on both surfaces of the leaf in Locandi (Samadera) mekongensis; Pierre and
Samadera indica, Gaertn,~ while in other· species of the genus two of them are
situated on the midrib on_~he lower side of the leaf.
According to Jadin, the petiole for the most part has an annular :fibrovascular
system"'\\.:h,i~ll f!eg~~q tly (according. to Jadin, the only exceptions are Alvaradoa,
Ha"isonia, -Irvingia, Klainedoxa, Pic1'amnia and Picrodendron 1) encloses one

1 The species in which spicUlar cells were not known or at least were not expressly sta.ted to be
present in the mesophyll are as follows: E/annoa K/aineana, Pierre, H. uMulata, Planch.; Hypti-
andra Bitfwillii, Hook. f.; Mannia africana, Hook. f.; Old)'endea K/aineana, Pierre, O. gaoonensis,
Engl.; Quam'a tlfricana, Baill.; Sima/)a tlngustifolia, Spruce, S. Cedron, Planch" S. crustacea,
Engl., S.jlbribunda, St. Hil., S,foetida, Poepp., S. glandulifera, Gard., S. insipu's, St. Hil., S. t)/m..
vaJa, Engl., S. mnocmsis, H. B. K., S. sa/u6rz's, Engl., S. su!xymostl, St. Hil.,_S. U'armingJ'aIIIJ,
Eng!.; St'maruba jloribunda, St. Hil., S. officinalis, Macf., S. Tu/ae, Urb. According to Van
Tieghem, Jadin's statement as to the occurrence of spicular cells in Irving,'a Barleri, Hook. f. and
I. Oliveri, Pierre is incorrect. .
t The only point of disagreement with the abo\'e facts i~ th6 earlier statement (see p. 18...
860 ADDENDA-SIMARUBACEAE

or more medullary bundles. The species with medullary resin-canals in

the axis also have them in the same position in the petiole. The fibro-'
vascular system is commonly accompanied by pericyclic hard bast. In Sama-
dera (4 species) a characteristic feature is the occurrence of concentric vascular
bundles, with central phloem and peripheral xylem, at the margin of the pith.
According to Van Tieghem, medullary vascular bundles are wanting in all the
genera of the Irvingieae, but in the upper part of its course the closed vascular'
ring, fonned by the fusion of seven or more isolated vascular strands derived
from the axis, has two inversely orientated bundles situated in the peri-
cycle. Lastly, according to my own investigation, the petiole of Sur-iana con-
tains only a single arc-shaped vascular bundle, while in Rigiostachys there is
an annular vascular system, which may be either closed or open.
3. STRUCTlJRE OF THE AxIS. With regard to the structure of the
wood we may add that in lrvingia gabonensis, Baill. it shows alternating
zones of wood-fibres and wood-parenchyma (Lecomte, see also Van Tieghern),
that in lrvingella and Klainedoxa the wood-parenchytr..a is rather abundant
(Van Tieghem), and that septate wood-prosenchyma occurs also in Rigiostachys.
According to Jadin, the species investigated by him also for the most part
show isolated groups of bast-fibres in the pericyc1e. Jadin distinctly mentions
the occurrence of a more or less continuous and composite ring of sclerenchyma
in the following additional species: Castela depressa, Turp., Picramnia venicosa,
Tul., and Sa madera Harmandii, Pierre; according to Van Tieghem, a composite
and continuous ring of sclerenchyma, distinguished by the inclusion of U·shaped·
sclerosed cells, occurs in the four genera of the Irvingieae. In the leafless
species, Holacantha Emoryi, A. Gray, hard bast appears to be wanting in the
pericycle; instead there are subepidermal groups of fibres with intermediate
assimilatory tissue of the nature of palisade. The pericycle of Rigiostachys·
contains a composite and continuous· ring of sclerenchyma, which splits open
in the course of the subsequerit secondary growth. Regarding the pericycle
of Ailanthus glandulosa and Simaruba amara, see also Pitard, loco cit.
As a rule the cork arises subepidermally; this is the case in all the genera
investigated by Jadin with the exception of Suriana, Amaroria and Soulamui,
in the genera of the Irvingieae (according to Van Tieghem) and in Rigiostachys.
In Suriana the cork develops in the inner part of the primary cortex; in
A maroria and Soulamea its place of origin has not yet been determined. Cork·
cells with one~sided or U~shaped thickening (affecting the inner tangential walls)
occur in Irvingia and have also been observed in the Asiatic species of Irvingella
and in Klainedoxa Trillesii, Pierre (Van Tieghem). .
The primary cortex occasionally contains ordinary sfone~cells, which in
Castela and Samadera are thickened in the form of a !torse-shoe. In the genera
of the Irvingieae, according to Van Tieghem, there are two. layers of what he
calls cristarque' ~cells, i. e. cells which mostly exhibit U-shaped thickening,
f

and each of which includes a solitary crystal, or rarely ([rvingia) a clustered

crystal'; one of these layers is subepidermal in positiori, the other is endodermal.
These cristarque '-cells, it may be added, are also present in the petiolar tissue.
j

Development of secondary hard bast is very common, although rare amongst

,the Irvmgieae. Van Tieghem found stone-cells in the secondary bast in
species of Desbordesia and Klainedoxa.
The structure of the pith in Harrisonia requires special mention. The
medullary tis~ue is heterogeneous, small thick-walled cells being distributed
in a reticulate manner amongst larger cens with thin walls.

foot·note) as to the :lbsence of medullary bundles in Bruua; Jadin did not investigate the petioles
of BruneI/its, Castela, Dictyo/(Jma and Picnuna, in which medullary bundles have likewise been
preTiously recorded as absent.
 ADDENDA-SIMARUBACEAE 86r

In amplification of the earlier statements regarding the occurrence of

medullary resin-canals in the Simarubaceae the following details may be
quoted from J adin's work.
'4 _Resin-canals are found in the following additional species: Ailanthus caly-
cina, Pierre, A. exee/sa, Roxb., A. Fauveliana, Pierre, A. imbet'bi/olia, F. v. M.,
A. malabariea, DC.; Byueea paniculata, Lam., B. sumatrana, Roxb.; Euryeoma
longi/olium, Jack (in opposition to 5yst. Anat. p. I87); Hannoa Klaineana. Pierre.
H. undulata, Planch.; Oldyendea gabonensis. Pierre, O. Klaineana, Pierre; Pict'tlsma
ailanthoides, Planch., P. iavanica. Bl., P. nepalensis. Benn, P. quassioides, Berm.,
P. Tweed ii, Planch.; Simaba angustifolia, Spruce, S. fioribunda, St. Hil., S. glan-
dulifera, Gardn .• S. insignis, St. Hi!., S. obovata, Engl., S. ot'inocensis, H. B. K.,
S. subcymosa, St. Hil., S. 5uOruticosa, Engl. (contrary to Syst. Anat.• loc: cit.) ;
Soulamea amara, Lam., S. elegans, Vieill., S. Mulleri, Brongn. et Gris, S. Panchct'i,
Brongn. et Gris, S. tomentosa, Brongn. et Gris, S. trijoliata, Baill. On the other
hand, Jadin did not meet with resin-canals in the generaAlvaradoa, Cadellia, Castela,
Harrisonia, Holf!.eantha, Hyptiandra, Irvingia. Kirkia, Klainedoxa. Mannia, p~'cram­
nia, Pz'cl'clla, Picyodendron. Quassz'a, Samadera, Sur~'ana, and also not in R~'gioslachys
and Guilfoylia. As a general rule the presence of medullary resin-canals is a
generic character. The sole exception has proved to be Simaba, since some .0£ the
species of this genus have no medullary resin-canals (see Syst. Anat. p. 187;
according to Jadin, this is also the case in S. crustacea, Engl., S. joetida, Poepp.
and S. salubris, Engl.).

Resin.:.cells have been recorded by Jadin. Guerin and Van Tieghem in the
following additional cases :-in the mesophyll, in Ailanthus calycina, A. Fauve-
liana, A. imberbifolia and A. malabarica; in the primary cortex and in the
leaf, in Chamaelea,pulverulenta; in the primary cQrtex, in Harrisonia Brownei,
Juss., Irvingia Duparqueti, V. T., I. gabonensis, Baill. and I. tenuifolia, Hook.
f.; accompanying the 'pericycle in the axis and the vascular bundles of the
veins in the leaf, in Oldyendea Klaineana, Pierre. In the genera I rvingia,
Klainedoxa and Picrodendron (according to Jadin) mucilage-cells are found
in the primary cortex of the axis and in the ground-tissue of the petiole, while
mucilage-lacunae are present in the pith of the axis. Mucilage-spaces, similar
to those of I rvingia, occur also in Perriera (according to Guerin), being situated
in the axis, in the rachis of the leaf, in the petiole, and in the larger veins of
the leaf. The genera Desbordesia. IrvingeUa,. Irvingia and Klainedoxa (accord-
ing to Van Tieghem) have mucilage-cells which either are isolated or form
groups and are found in the primary cortex, and in some cases in the pith as
well.
For the structure of the spiny aerial roots of Klainedoxa spinosa, V. T., see
Van Tieghem, 1905. loco cit.

ApPENDIX: Koeberlinia.
A connected account of the anatomy of the genus _Koebe"linia may be given
at this point apart from the general description of the Simarubac~<_te. The affinities
of>the genus"are noC-yet quite Clear; but it.is regarded by Engler and Van Tieghem
as constituting an independent ,Order (Koeberliniaceae). The only species of the
genus, Koebellinia spinosa:. Zucco is a leafless and spiny shrub; from an anatomical
point of view, it is specially characterized by the possession of secreto.ry canals
situa.ted in the bast, the composite and continuous ring of sclerenchyma in the
pericycle, the peri cyclic cork-development, the simple perforations in the vessels
and the wood-fibres which have thick walls and slit-shaped pits with asrp,all,border.
The vascular bundles of the axis are separated by rather broad medul~ rays,
the outer ends of which are enlarged in·the form.of a wedge between the.bast-por-
tions. Opposite the hast-portions the pericycle contains massive bundles of hard
bast, which are joined to form a continuous strengthening ring by means of stone
cells. The bast-portions exhibit a peculiar stratification into darker zones, com-
posed of parenchyma and lig4tcr zgnes, consisting of Compressed sieve-tubes.
 ADDENDA--SIMARUBACEAE
The secretory canals originatein the parenchymatous.zones, and either lie singly
or several of them are placed side by side; in the former c~e they take up the
whole breadth of the band of parenchyma. In later stages sclerosed cells are found
in the bast and in the medullary rays of the bast; similar cells occur also in the
primary cortex. The cork develops in the pericyclic parenchyma on the inner side of
the strengthening ring and consists of cells, the outer tangential walls of which are
strongly thickened. Phelloderm. is present, its cells having lignified walls. The
epidermis consists of cells exhibiting palisade-like elongation and having all their
walls thickened, especialIy the outer ones. 1n the absence of leaves the outer
part of the primary cortex is differentiated as a palisade. tissue of several layers,
while the inner part consists of isodiametric cells. The only kind of trichomes
yet observed are unicellular conical clothing hairs with thick walls. Oxalate of
lime is found in the pith in the form of solitary crystals.
Literature: Yignoli, Cay·Cay (Irving-ia Oliveri), These, Montpellier, 1886, pp. :Z9-32 and
pl. i-ii.-Leblois, Tbylles d. can. secret., Bull. Soc. bot. de Frauce, 1887, p. 184.-]adin, Org.
secret., These, Montpellier, 1888, p. 52 et seq.-Barber, Corky excresc., Ann. of Bot., vi, 189:1,
p. 16s.-Pierre, Flore forest. de la Cochinchine, xvii, 1892, and xix, I 893.-Cla.udel, Quassia
africana etc., These, Montpellier, 1894, p. I I et seq.-Engler, Koeberliniaceae, in Naturl. Pflanzenfam.,
iii Teil, -Abt.. 6, 1895. pp. 320, 32I.-COrnu, Quassia afrieana, Bull. Soc. bot. de France, 1896,
p. 523 et seq.-[Hills, Holz von Picraena u. Qtlassia, Jouro. of Pharm., 1897.]-B~rgesen og
Paulsen, Veget. dansk.~vestind. aer, Bot. Tidsskrift, xxii, 1898-9, pp. 94,95 (Casfeta eree/a, Turp.).
-Macchiati, Uffic. dei peIi dell' antocianino e dei nettarii estranuz; dell' A~1antflus glanaulosa,
Bull. Soc. bot. Ital., 1899. pp. I03-12.-Van Tieghem, Cneoracees, Ann. sc. nat., ser. 8, t. ix, 1899,
pp. 363-9; see also Bull. Mus. d'bist. nat., 1898, p. 241 et seq.-Gamper, Angosturarinden, Diss.,
ZUrich, J9OO, p. 64-.-Van Tiegbem, Stacbyuracees et Koeberliniacees, Joum. de Bot., 1900, pp. 7-12.
a
-Jadin, Contribut. l'etude des Simarub., Ann. sc. nat., ser. 8, t. xiii, 1901, pp. '201-304.-Pitard,
Pericycle, These, Bordeaux, 1901, pp. 80, SI.-Van der Marck, Samadera indi&a, Archiv d. Pha.rm.,
239, 1901, pp. 96-10l.-Jadin, Classification des Simarub., basee sur les caract. anat., C. R. Assoc:
{ran.;. Ajaccio, 2" partie, 1902, pp. 477-8J.-(Annan, Piante (leIla reg. medit., Ann. di Bot., i, [903,
p. 17 et seq. (Cneortun).}!.·Lecomte, Qu. bois dui Congo, Bull. Mus. d'hist. nat., 1903, p. 890-
Tuzson, Spiral. Struktur d. Zellwande in den Markstr., Ber. deutsch: bot. Gesellsch., 1903, po 276.-
Achner, Falsche' Cbinarinden, Diss., 'Bern, 1904, p. _63.-[Bessey, Chimney-shaped stomata of
Ho/acantha, Bull. TotTey Bot. Club, ~xxi, 1904, pp. 523-7 and pI. 24.}-Col, Faisceanx, 'Ann. sc.
nat., 5er. 8, t: xx, 1904, p. log.--Courchet, Kirandro, Bull. Soc. bot. de France, 1905, p. 284.-
Solereder, Syst. Stellung der Gatt. Rigz'ostachys, Verb. bot. Ver. Mark Brandenburg, 190 5, p. 41
et ~.-Van Tieghem, Irvingiacees, Ann. sc. nat.; ser. 9, t. i, 1905, pp. 247-320 (Iroiflg-z'a,1ruin-
gella, Desbor'desia, .Klainedoxa).-Piccioli, Legnami, Bull. Siena, 1906, p. 173.-[Van Tieghem,
Agialidacees, Ann. sc. nat., ser. 9. t. iv, 1906, pp. 222-6o.1-[Van Ticgbem, Ailante et ,Pongele,
Ann. sc. ll3.t.) ser. 9, .t. iv! 1906, -pp: 272-80. ) - , '

OCHNA~EAE (pp. I88-190).

In the course of the last few years the Ochnaceae (sensu Bentham and Hooker'
have formed the subject of detailed systematic, morphological and anatomical
studies by Van Tieghem I. On the basis of his investigations Van Tieghem regards
the genera Ochna, Ouratea, Brackenridgea and Elvas~'a (belonging to the Ochneae)
as constituting an independent Order, the Ochnaceae (with no less than 57 genera) ;
Tetramerista (see 5yst. Anat., p. r89) is excluded, while the four genera above named
are s'plit up into a number of others. In the same way the genera of the Luxem-
burgteae with the exclusion of Wallacea and the addition of seven other genera
are established as a separate Order, the Luxemburgiitceae. In the fOllowing descrip-
tion, in which Van Tieghem's system of classification is adopted, we shall deal with
the anatomical characters presented by leaf and axis', first in the Ochnaceae sens.
str., then -in the Luxemburgiaceae, and lastly in Wallacea (Order: Wallaceaceae,
V. T.);· as well as in Euthemis (formei'ly Tribe Euthemideae, Order Euthehridaceae~
V. T.). . .

~'I merely adopt Van Tieghem's nomen'clature of the genera and species in the above.description
as a matter of convenience. Fo( a criticism of his systematic theories see Gilg, Beitl'. z: Kenntnis
d. Ochnaceen, Festschrift fLir Ascllerson, uipzig, 1904. p. 97 et seq. .
S As far as the strUcture of the wood is concerned there is nothing to add to the older statements
(Syst. Anat., p. 189), since ,Van Tieghem devoted very little attention to it.
• • r-
 ADDENDA--OCHNACEAE

I. OCHNACEAE, VAN TIEGHEM.

I. ANATOMICAL' FEATURES. Two features are primarily characteristic'
of the whole taxonomic group, viz. (a) the presence in the branches, petioles,
median and lateral veins of a layer of cells (' cristarque '), which is normally
situated in the second layer beneath the epidermis and is composed of cells
with U-shaped thickening, each of which encloses a clustered crystal; and
(b) the occurrence of cortical vascular bundles. The cork invariably develops
superficially, viz. in the epidermis or in the first layer of cells of the primary
cortex. A hairy covering is rarely present, and then consists exclusively of
uni- or multicellular clothing hairs. Oxalate of lime is mostly deposited
in the form of clustered crystals.
The anatomy of the genera Elvasia, Vaselia, Trichovaselia and Hostmannia,
which Van Tieghem groups together as the Elvasioideae, differs from that of
the remaining members of the Order (Ochnoideae) in the occurrence on the
upper side of the leaf of a hypoderm, composed of fibrous cells which show
a transverse arrangement, and in the presence in the pith of the petiole of an
arc of wood and bast, showing normal orientation (with the wood on the upper
side). In certain cases the leaf contains gelatinized or papillose epidermal cells I
or spicular fibres, while in some of the species of Trichouratea the stomata are
placed in pits. _
2. STRUCTURE OF THE AXIS. In view of the systematic importance of
the layer of cells, termed the cristarque,' the structure of the axis may in I
I

this case be considered before that of the leaf. The,' cristarque 1 constitutes J

the second layer of the primary cortex, -and is composed of lignified cells~
with U-sbaped thickening (on the inner tangential and radial walls) and
each enclosing a clustered crystal of oxalate of lime; the latter sometimes
shows a slight sphaerocrystalline structure. The cristarque J is not quite con-
I

tinuous, being interrupted by thin-walled passage-cells, which in general

correspond-in position with the stomata in the epidermis. Very considerable
diversity is shown in the mode of differentiation and in the position of the I
, cristarque in the individual" genera and within the limits of one and the
J

same genus, often varying, in fact, from species to species; these differences
are of systematic importance. The number of thin-walled cells may be small
or large; as a consequence, we get all transitions between an almost continuous
I cristarque' and a zone, composed only of a small number of ' cristarque '--cells
In certain species (e. g. of Ouratea, Campylospermum, Campylocerc'Zem, Cer- I

'canthemum, Cercinia, &c.) the' cristarque' is apparently not situated in the

second cell-layer of the primary cortex, since it is separated from the epidermis
by morethan one (2, 3 or 4) layer of cells; 'but an investigation of the course
of development in these cases shows that the subepidermal layer has under-
gone subsequent division into 2-4 layers of cells, so that the (cristarque'
nevertheless belongs to the second cortical layer. The thin-walled cells found
in the discontinuous cristarque undergo subsequent sclerosis in many species,
f J

the cells being affected equally on all sides. In certain species the' cristarque '-
sheath is further strengthened by uniform sclerosis of the cells of one or more'
layers of the primary cortex; these are either situated on the inner side of
the cristarque: or beneath the epidermis (including the layers of cells produced
I

by the division of the subepidermal layer), or in both these regions. Only

in very rare cases (Dip::-yllanthus Duparquetianus, V. T.) do the cells of the
(here well-developed) 'cristarque' contain prismatic crystals in place of the.

1 In the rhi%omes and roots which have been examined, the layer of cells known as ' cristarque '
is completely absent
 ADDENDA--OCHNACEAE
clustered crystals. As regards the remaining characters of the primary cortex.
we may notice that it frequently contains clustered crystals and sclerosed cells,
in some cases also prismatic crystals (species of Campylospermum, Spongo·
pyrena, Dipor£dium, ~1fonoporidium, Porochna) or cells in which the structure
of the wall is similar to that of the cristarque '-cells (species of Cercinia,
I

Ochnella, Vasel£a; in the last two genera occupied by prismatic crystals) or

unequally thickened cells containing prismatic crystals (Trichovaselia). The
endodermis is as a rule not distinctly differentiated; it is only in a few species
that it includes cristarque '-cells with clustered crystals; in these cases
f

there is a secondary endodermaI ' cristarque,' which is however only rarely well-
developed. The above-mentioned cortical vascular bundles owe their origin
to the fact that of the three bundles, passing out from the stem into the leaf
situated immediately above, the two laterals in most cases depart from the
vascular ring of the axis some considerable distance below the node (in Dipori-
dium alone do they arise only in the uppermost part of the internode); as
a consequence no cortical vascular bun.dles are, present in the lower part of
the internode. In most cases there are two of these cortical bundles, but
occasionally (Camptouratea) a larger number is found in the uppermost portion
of the internode owing to branching. The bundles are accompanied by groups
of bast-fibres. In most members of the Order the pericycle is formed by isolated
groups of bast-fibres, but occasionally an almost or completely continuous
and composite ring of sclerenchyma is developed (e .. g. in species of Trichou·
ratea, Dasouratea, Cercouratea, Microuratea, Gymnouratella, Campylospermum,
Diphyllanthu,s, Monelasmum, OchneUa) by sclerosis of the cells situated between
the groups of bast-fibres. The secondary bast sometimes contains clustered
crystals, but rarely (Diporidium purpureum, V. T., Porochna Autunesii, V. T.)
prismatic crysta1$; sclerosed cells 1 may also be present; fibres (Diporidium
purpureum, Hostmannia) or unequally sclerosed cells containing prismatic
crystals (Elvasia, Trichovaselia) are very rare. The mode of development ot
the cork, whether epidermal or subepidermal, is on the whole only a specific
character. The walls of the cork-cells are either thin, or the tangential walls
are sclerosed. Phelloderm mayor may not be present. It has either thin walls
or some or all of its cells exhibit U·shaped thickening; complete sclerosis
of the cells is rare; in a few cases (species of Porochna and Diporochna) the
cells of the phelloderm contain small prismatic crystals.
Epidermal cork.development has been recorded in: Camptouratea pro parte,
Stenoumtea, Notouratea, Plicouratc!J pro parte, Ancouratea pro parte, Diouratea,
Trichouratea (almost always), Pilouratea, Dasouratea, Ouratea pro parte, lsouratea,
Polyouralea, Tetrouratea, Cercouratea pro parte, M£crouralea pro parte, Setouratea,
OurateJla, Gymnouratelta, Bisetaria, Campylospermum pro parte, Campylocercum
pro parte, Cercanthemum pro parte, D£phyllopodium, S pongopyrena. Rhabdophyllum,
M oneJasmum pro parte, Exomicrum pro parte, Oehnella pro parte, Polyochnella
pro parte, Discladium, Diporidium pro parte, Monoporidz"um, Polythedum pro
parte, Heteropodium, Oehna, Diporoehita, Pleuroridgea, Campyloch n ella , Vaselia;
subepidermal cork-development is found in: Camptouratea pro parte, Plicouratea
pro parte, Ancouratea pro parte, Triehouratea /oliosa, V. T., Hemiouratea, Valken·
steinia, Ouratea pro parte, Cercouratea pro parte, M£crouratea pro parte, Campylo·
spermum pro parte, Campylocercum pro parte, Cereanthemum pro parte, Cet'Cinia,
No tocampylum , Diphyllanthus,1 J.,lonelasmum pro parte, Exomierum pro parte,
Oehne/la pro parte, Polyoehnella pro parte, Diporidiu"~ pro parte, Polythecium pro
parte, Porochna. Brackenridgea, Elvasia, Trichovaselia, Hostmanm"a.
The pith becomes lignified at an early stage. In addition to clustered
crystals the pith in many species includes 'cells which are sclerosed on all sides;
-----~--.

1 Van Tiegbem does not describe the detailed structure of the sclerosed cells in the bast;
regarding this point, see Syst. Anat., p. I89.
 ADDENDA __ OCHNA CEAE
in Notouratea undata, V. T. occasional cells exhibit thickening of the wall
similar to that found in the' cristarque' -cells.
3. STRUCTURE OF THE LEAF. Three vascular bundles pass out into the
leaf or petiole as the case may be ; the median bundle of the three branches
off from the vascular ring of the axis at the node, while the two lateral strands
traverse the upper portion of the internode as cortical bundles; each of the two
lateral strands gives off a small branch which supplies the stipules or the ligule.
The vascular bundles on entering the petiole soon unite to form a ring of wood
and bast, which is in most cases completely closed (except in Microuratea
cassini/olia, V. T.) and has a central pith; the lower surface of this annular
system is- convex, while the upper is concave or flat.; bundles of fibres, which
are mostly distinct from one another, accompany the bast on its outer side.
The Elvasoideae are specially distinguished from the Ochnoideae by the fact that
the pith of the petiole contains an arc of wood and bast, comprising several
vascular bundles, which exhibit normal orientation, the wood being placed
on the upper side. Medullary bundles, it is true, are exceptionally present also
in two members of the Ochnoideae (Campylospermum angulatum, V. T. and
NottJcampylum Mannii, V. T.), but the bundles in these cases show inverse
orientation, the wood being on the lower, the bast on the upper side. In this
connexion we may notice that the pith· contains a transverse band of fibres
in Trichouratea Gardneri, V. T., two bands of fibres in species of Spongopyrena,
and elements resembling' cristarque '-cells in species of Rhabdophyllum, Poly.
thecium and Diporochna. In Campylosperm'ltm nigrinerve, V. T., five cortical
vascular bundles, which are united to form an arc, are found on the outer and
lower side of tpe vascular ring. The cristarque' (and we may deal first with
I

the outer one, which is situated in the second cell-layer beneath the e~idermis)
is generally present also in the petiole, and then exhibits the same modIfications
as in the branch. It is more or less typically differentiated. Only in rare cases
is there merely a_single layer of cells between it and the epidermis; as a rule
there are 2,3, 4 or even 6-8 intervening layers of thin-walled cells, which have
been formed by subsequent division of the subepidermal layer. The outer
l cristarque' is rarely absent (e. g. in species of Camptouratea, Tetrouratea
and Campylospermum); still mQre rarely (Diphyllanthus) do the cristarque '. I

cells contain prismatic in place of clustered crystals. In some of the species

there is, .as in the branch, an inner (endodermal) 'cristarque' in addition to
the outer one, this inner cristarque' varying in the extent of its development.
I

Lastly, elements resembling 'cristarque -cells are occasionally found also in

J

the cortical tissue (species of Ouratea, Rhabdophyllum, Polythecium, Diporochna,

Brackenridgea, Trichovaselia); in other cases the cortex contains cells which
are sclerosed on all sides.
Regarding the structure of the lamina of the)eaf the following facts may
be mentioned. In most of the species the leaf is bifacial in structure, the pali.
sade-tissue consisting of a single layer of cells. Centric structure with palisade-
tissue on_ b-oth sides~of the ·leaf is found only in Pilouratea ovalis, V. T. and
Dasouratea Hassleriana, V. T.; in certain species of Cercouratea and Diphyllo-
podium the palisade-tissue is not typically differentiated. In many species I
a varying number of the epidermal cells of the leaf have mucilaginous inner
membranes, these cells sometimes penetrating deeply into the mesophyll.
Ouratea guianensis, Aubl., O. rubescens, V. T. and Cercouratea Magdalenae, V. T.

1 viz. species of Camptauratea, Stenouratca, lI'olouratea, Plicouralea, .£-I1uouratea, Tric/Jallratea,

Villauratea, Hemiotlralea, Oura/ea, Po/youYatca, Tetrouratea, CeYcouralea, ltJicrouratea, Setotlralea,
Ouratdla, Gymnourate//o, Campylosper11lum, Campylocercum, Cercanlhemum, Cercim'o, SponKO-
pyrena, AlolleiaslJltl, Ochl1ella, Palyoclmella, IJisdaJiulIl, IJijon"diunt, lUollojoridium, Poly/hecium,)
Odma, IJijorOt.-hna, Brad:mridgea, l'leut"oridgea, Campylochllella.
SOI.ERF.OU 3K
866 ADDENDA __ OCHNACEAE
a!'e distinguished by possessing sclerosed epidermal celIs, the inner ends of which
are DartOwedin fhe form of a cone and penetrate into the palisade-tissue. The
epidennal cells of NotouTatea inundata, V. T. have lignified thickening bands
QA their lateral walls. Formation of papillae has been recorded in Notou,atea
i~undata, V. T. (knob-shaped cuticular papillae on the lower side of the leaf),
Trichouratea j(jliosa, V. T. (on the upper side), Diphyllopodium Klainei, V. T.
and Exomicrum coriaceum, V. T. (on the lower side). Not uncommonly the
Walls of the epidermal cells are completely or partially lignifie~. In most of
the species the stomata are confined to the lower surface of the leaf. But in.
Pilouratea ovatis, V. T. and Isouratea humilis, V. T., they occur on both sidf".-S,
and in species of OUTatea, Cercinia, Polyochnella, Pleuroridgea, Campyloehnella,
Elvasia, Vaselia, Trichovaselia and Hostmannia, stomata are also present on
the- upper side, though only on and near the midrib (in CampylochneUa they
occupy the same position with reference to the lateral veins as well). Van
Tieghem's statement as to the absence of subsidiary cells is not quite in agree·
ment with my earlier observations. In Trichouratea subvelutina, V. T. the
stomata (also those on the axis) are found in special pits CSpaltoff-
nungskrypten '). The occurrence of a hypoderm composed (jf' trans-
versely placed fibrous cells, and situated beneath the epidermis of the leaf
in the Elvasioideae, has already been referred to abdve. Other special features,
noticed in the mesophyll, are as follows :-large solitary crystals (Ouratea
Leprieuri, V. T.); 'cristarque '-cells, occasionally forming groups (species of
Camptouratea, Ouratea, Campylospermum, Cercanthemum, Rhabdophyllum) ;
sclerosed isodiametric cells (Ou1'atea gigantophylla, V. T.); and lastly,
spicular fibres, which generally branch off Jrom the sclerenchyma of the veins,
run vertically through the mesophyll or traverse it in all directions, and in most
cases ultimately spread out beneath the epidermis (in a large number of
species 1). The vascular system of the lateral veins is provided both above
and below with a group of sclerenchymatous fibres, and is separated by two
layers of cells from the epidermis. The inner of these two layers, which
may be regarded as equivalent to an endodermis, is generally constituted
by a 'cristarque,' containing clustered crystals, and must be considered as ..
a continuation of the outer' cristargue ' of the branch, petiole and midrib.
The 'cristarque' of the lateral veins IS developed either on both the upper and
lower sides of the vascular system, or (very commonly) only on the upper
side; but in a few cases (viz. in the Elvasioideae, in which palisade-tissue'is
met with also in the lateral veins) it is confined to the lower side. The
• cristarque ' is very seldom imperfectly developed. Owing to sG:lerosis of the
two layers of cells situated between the bundles of fibres and the epidermis, the
lateral veins in many species 2 ultimately become vertically transcurrent. The

, viz.: Camptoura/ea agropkylla,.V. T., C. £licifoHa, V. T., C. spimtlosa. V. T.; Plic()urlllea

granulosa, V. T. i Ancouratea hemiodotz/a, V. T.; Diouralea eardiosperma, V. T.; Trickouralea
.JJJandu#ana, V. T., T. t"ou/ipila, V. T.,· T..Jl~ribJlnda, V. T., T._foliflsa, V. T., 1: rufiduJa, V. T.,
T. sa/iei/o!;a, V. T. j Dasuuratea Hass/eriana, V. T. j Vollumteinia Theoplzrasta, Reg.; Ouralea
anguJata, V. T., O. castantifolia, Engl.l O. (()cci1lea, Engl., O. crassijo/ia, Engl., O. (ubensls, Urb.,
O. distieha, V. T., O. Glaziovi£, V. T., O. neterotionta, V. T., O. macr()phylla, V. T., O. panamieIJ,
V. T., O. Purdieana, V. T., O. RudeNana, Engl., O. rupununiensis, Engl., O. Spruceana. Engl.;
b,uratea "umilis, V. T., I. sjectabilis, V. T.; Ctrcuuratea curvata, V. T., C. repens, V. T., C. wr-
rucu/osa, V. T.; ftlicrouralea }ygmaea, V. T.; Campylospermum angoula/um, V. T., C. BIJr6";,
V. T., C. Ckape/ieri, V. T., C. denudalum, V. T., C. Hi/tklJranatii, V. T., C. nigrinerw, V. T.,
C" ()'IJale~ V. T .• C. sculptum, V. T.; Cercantlumum lanet()lalum, V. T.; N()t«ampjIUIH M.".tdi,
V. T.; Rhalx/opltyl/um caloplzyllum, V. T.) R. panieulatum, V. T. ; Mtme/as"", ZnU6ri, V. T ••
A/. um/Jrico/a, V. T.; P()lJItludum tnadagascariense, V. T. j Brackenrid~a H",,~w;, A. Gmy,
B. nitidaJ A. Gray, B. }a/ustris, Hartel.
, viz. species of the genera: Camp/oura/ea, Tr;cllouralea, Duralea, Micruuralla (in almost all
the species), CamjJyksjJerlllum, Cercantlultlum, (ercin;a, Oelme/la, Po!J'«Imella, Disdadium,
LJiporidlum, PII/)Itnecium, Oc/ana, PorocAna.
 ADDENDA--OCHNACEAE
structure of the midrib of the leaf, at least in it:; lower pcN.i0ll. is iimilat
to that of the petiole (see above).
A hairy covering has only been observed in a few genera and species, and
is composed exclusively of clothing hairs. These are unicellular (TriQ,¥urateu,
Pilouratea, Villouratea?, Dasouratea?, Hemiouratea, Trichovaselia) orblcellulat,.
and a third cell often occurs in the latter case owing to the division of the lower
ceIl by means of a longitudinal or oblique wall (Diporochna). In ot4.- c~
(T,ichourateal) the hairs are uniseriate and consist of a larger nUI{lber of celfs.
The unicellular trichomes vary in length; in T,ichouratea they are sometimes
united in bUIldl~s of two or three.
2 .. LUXEMBURGIACEAE.
1. REVIEW OF THE ANATOMICAL FEATURES. The Luxemburgiaceae, like
the Ochnaceae, are characterized by the possession of cortical strands (leaf-
traces); cork~formation likewise takes place superficially in the epidermal or sub ..
epidermal layer of cells. The pericycle is formed by isolated groups of bast ..
fibres. The structure of the leaf is bifacial and the stomata are found ex-
clusivelyon the lower side of the leaf. There is no hairy covering (apart from
the glandular shaggy hairs). ,Oxalate of lime is deposited in the form of clustered
or solitary crystals.
The genera of the Godoyeae are distinguished by the presence of medullary
bundles in the branch; these are composed either of vessels and fibrous cells
(' fibrovasculaires,' e. g. in Godoya, Planchonella and Rutidanthera) or of a
strand of soft bast and fibrous cells Cfibrocribles,' e.g. in Cespedesia and Four·
nieria). Other characters distinctive of the Godoyeae are the stratification
of the phloem into hard and soft bast, and the occurrence of· characteristic
glandular shaggy. hairs (' franges secretrices') on the stipules and sepals.
On the other hand, medullary bundles and glandular hairs are wanting in the
two other subdivisions of the Luxemburgiaceae, viz. the Luxemburgieae
(with Luxemburgia, Periblepharis, Plectanthera, Epiblepharis and Hilai,ella)
and the Blastemantheae (with Blastemanthus and Poecilandra); the Blaste-
mantheae however, like the Godoyeae, have a stratified bast, while in the
Luxemburgieae there is no hard bast. J

Among special features 'of the structure of the leaf, we may name: the
gelatinization of the epidermis; the occurrence of spicular fibres in the meso-
phyll; the development of an endodermal ' cristarque ' in the lateral veins;
and the occurrence of vertically transcurrent lateral veins.
2. STRUCTURE OF THE LEAF. The leaf is bifacial in structure. Epidermal
cells wi~h mucilaginous inner membranes are found in Luxe1nburgia, Epible-
pharis and Hilairella. The stomata are confined to the lower side of the leaf;
in Godoya they are crowded together in groups"in the narrow meshes formed
by the network of veins. Sclerenchymatous fibres, running freely in the
mesophyll, have been observed in the genera Rutidanthera, Cespedesia,
Fournieria ...~·I!..Q_gljlslemanthus;- in· some <;:ases (Blastemanthus) they form a
continuous layer beneath the upper epidermis. The lateral veins are often
vertically transcurrent .by means of sclerenchyma (Luxemburgia pro parte,
Periblepharis, Godoya, Planchonella, Rutidanthera, Blaste~nthus, Poecilandra).
An endodermal cristarque t containing clustered crystals is stated to occur
j

in species of Luxemburgia, Plectanthera, Epiblepharu, HilaireUa, Cespeaesia,

Fournieria. and Blastemanthus, whilst Poecilandra alone has a 'c~tarque'
containing solitary crystals. In all the genera except Poecilandra this layer is
confined to tp,e upper side of the vascular system, while in Poecilahd,a it is
present on both sides. The vascular bundles, which pass out into the leaf,
unite low down in the petiole or in the midrib to form a ring, the pericycle of
which contains .fibrous cells. In Hilairella the pith of this ring of bundles
3K2
868 ADDENDA-LUXEMBURGIACEAE

contains a strand of bast With a band of wood on either side of it, while in
Godoya, Pia nchonella , Rutidanthera, Cespedesia and Fournicria it includes
from two to four arcs of wood and bast exhibiting diverse orientation and
situated one above 'the other; in Blastemanthus one or two vascular bundles,
and in Poecilandra an arc of normally orientated bundles, are found in the pith.
Theg]andu]ar shaggy hairs, occurring on the stipules and sepals in all the
Godoyeae, are .2-5 mm. ill length and are distinguished by having a secretory
palisade-llk:e epidermis. Beneath the latter lies a layer of cells containing
clustered crystals, and within this a vasculcir bundle, with an enveiopirtg sheath
of fibres.
3. STRUCTURE OF THE AXlS. The pericycle is invariably composed of
isolated groups of bast-fibres. The cork develops in the epidermis (Epible-
phari~, HilaireUa, Godoya, Fournieria, Blastemanthus) or in the subepidermal
layer of cells (Luxemburgia, Periblepkan's, Plectantkera, Planckonelta, Rut";-
danthera, Cespedesia, Poecilandra). The walls of the cork-cells are either· thin
or sclerosed; in the latter case they may be sclerosed equally on all sides or
in the shape of a U. Phelloderm mayor may no't be developed; when present
it is sometimes sclerosed. The primary cortex occasionally contains stone-
cells and oxalate of lime in the form of clustered or solitary crystals. 1n
Periblef!haris it includes isolated fibrous cells, while in Godoya the inner part
of the primary cortex is lacunar. The cortical bundles, which ar~ accom-
panied by groups of scier(>nchymatous fibres, vary in number; there .are
often 4-6; in Fournieria scandens, V. T. there are 16, in Blastemanthus and
Poetilandra only 2. The presence or absence of ,secondary hard bast has
already been dealt with in the review of the anatomical features. When hard
bast ·is present the appearance of a tran'sverse section through the phloem..
groups, and the intervening medullary rays 'with their expanded outer ends,
quite recalls the similar features seen in the branch of the lime.
The following details may be. added regarding the medullary bundles
found in the Godoyeae. In Godoya, Planchonella and Rutidanthe-ra the bundles
are made up of vessels and fibres. In the two genera first named there are 8-20
(mostly 10) of these bundles, arranged in a ring, while in Rutidantkera the
bundles are very numerous and irregularly distributed through the whole of
the pith, except m its central portion. Each bundle consists of a group of fibrous
cells and of a xylem-ray comprising a row of spirally thickened and pitted
vessels, which usually show radial arrangement; the vessels exhibit a pro-
gressive increase in size from without inwards, and are developed centripetally,
exactly as in the caSe of the xylem-rays in the radial bundle of a root. In
Godoya and Planchonella the xylem-ray is directed outwards, the group of
fibrous cells being placed at its inner end, while the outer end and the lateral
surfaces of the xylem-ray are surrounded by a layer of unlignified tissue
belonging to the pith. In Rutidanthero, on the other hand, the xylem-
ray is embedded in one side of the group of. fibres, the latter extending round
the lateral surfaces of the xylem-strand; the ray of wood in this genus
is moreover occasionally not situated on the outer side of the group of fibres,
but internal or lateral to it. The medullary bundles -found in Cespedesia and
Fournieria have an essentially different structure, being composed of a strand
of phloem and a group of fibres. In Cespedesia the phloem-group is for the
most part embedded in the inner margin of the well-developed bundle of fibres;
in some cases, however, it is situated at the outer margin or laterally and may
occasionally be absent altogether. The phloem-group develops in the centri·
fugal direction. In Fournieria the strand of phloem is generally surrounded
by a sheath of one or two rows of fibrous cells; it may, however, also be wanting.
The bundles in Cespedesia and Fournieria exhibit an irregular arrangement,
.
and are present in large numbers. It. is specially to be.. noted that the medullaty
 ADDENDA-LUXEMBliRGIACEAE

bundles of the Godoyeae are cauline, so that they are not connected with the
vascular strands of the stele.
We may add that in all the Godoyeae the meduJ\arybundles become converted
into typical vascular bundles in the axis of inilorescence. Strands of phloem
appear in relation to the vessels and fibres, constituting the medulla.ry bundIes of
Godoya, Planchonella and Rutidanthera, wllilst vessels are added to the groups of
phloem and. ttQ_res in the medullary bUildles of Cespedesia and Eou1lnieria. .But
the arrangeIrieut and pcevious orientation of the wood and bast is f6r the most
part retain~d. -
Un the genera Wallacea and Euthemt"s.
In its anatomical structure the genus Wallacea, which Van Tieghem excludes
from the Luxemburgiaceae, really shows quite a number of points of agreement
with the latter, as is evidenced by Van Tieghem's own statements; these points of
similarity are the cortical vascular bundles (six in number, each strengthened by
an arc of fibres), the nature of the pericycle (formed by small isolated bundles of
fibres), the superfi~ial (subepidermal) development of the cork, the cells exhibiting
U-shaped thickening, and also the presence of secondary hard bast (in small groups).
The OtIter ends of the primary medullary rays of the bast are not broadened in the
form of a wedge. The structure of the petiole is particularly striking; there is
~ rin~ of bundles, the pith of which contains two superposed vascular strands,
of which the IOVfer one exhibits normal, the upper one inverse orientation of wood
and bast. On either side of the vascular ring four cortical bundles are found;
these are situated one above the other, and consists of a central mass of :xylem.
enveioped by a ring 01 soft bast and fibres. The structure of the leaf of Wallaua
is bifacial, the stomata being confin·ed to the lower side. The lateral vein3 are not
vertically transcurrent. Oxalate of lime occurs in the form of clustered and solitary
erystals (the latter in the lateral veins).
According to Van Tii·ghem, the genus Euthem£s is specially distinguished by the
possession of large mucilage-celts situa.ted in the pith and cortex of the branches
and in the mesophyil.
Literature: Barteletti, Studio rnonogr. into aUa famiglia delle Ochnaceae, Malpighia" J9GJ,
rp. 105-74 and Tab. v-xi.-Pitard, Pericycle, Tllese, Bordeaux, I!)OI, p. 92.-Van Tieghem, Epible·
p~aride etc., Journ. de bot., 1901, pp. 389-94.:-Gerhard, B!::.!ta'l~t. v. Gew. des Knysnawald~t
1)1Is.) Basel, 1902, pp. 8-10 (Odtna).-Van Tlp.ghem, Ochnacees, Ann. se. nat., ser. 8, t. x't.J.,
190 2 , PP. I6I-4 I6. -Van Tit'ghem, Deux Ochnacees, Bull. Mus. d'hist. nat., 190 2 , PP./47-Sl.--.
Van Tieghem, Cristarque etc, Bull. Mus. d'hist. nat., 1902, pp. l66-73.-Va.'l Tieghem, UDe
()uratee etc., Bun. Mus. d·hist. nat., 1902, p. 61s.-Van Tieghern, Setoura.t~, Campyloopenne et
Bisetaire, Joum. de bot., 1902, pp. 33-47.-Lecomte, Bois eu Congo, Bull. Mus. d'hi!t. nat., 1903,
p. 89.:'_Van Tieghern, NOllv. obs. sur le~ Ochnacees, Ann. sc. nat., aer. 8, t. xviii, 1903, pp. J-60.~
Van Tieghelll, Esp. nouv. des Ochnacees, Bull. Mus. d·hizt. nat., 19°3, pp. 30, 70 and J~6 ct seq.-
Van ,Tieghem, Lnxembourgiacees, Ann. ~. nat., ser. 8, t. xix, 19°4, pp. I-¢.-Van Tieghem.
Wallacee, B!lIl. Mus. d'hist. nat., 1904, pp. 14S-so,-Van Tiegbem, }"aisc. med. des Godoyees.,
Joum.' de bot., 1904, PI>. .i:i3-64.-Van Tiegbem, Franges secretr., Joum. de bot., 19°4, pp. lC5-9.-
Areschoog, Trop. vaxt. bladbyggo., Sv. Vet. Akad.. H£lldl., 39, tI. 2, 19°5, pp. 29--90 (Ochna).-
[Pilger, in Natiirl. PfJam:en{am., Erg.. Heft, II, [907. pp. 219. 220.)

BURSERACEAE (PP.I9(}-I94).
2. STRUCTURE OF... THE LEAF. The following are additional details as to
the nature of ·the hairy covering (Syst. Anat., p. I9I}. In Boswellia
Carteri, Birdw. external glands with a short stalk and a bicellular head
divided by a vertiCal wall OCCUI" side by side with thick~wa1led clothing
hairs which are either unicellular or uniseriate (Ad. Peter). PfotiUHJ. set'1'atum,
Engl. has unicellular trichomes, which are sometimes united to form tufts,
while peculiar unicellular lanceolate hairs occur in Canarinm zeylanicutn
(Stepowski). .
3. STRUCTURE OF THE AXIS. Bargagli-Petrucci met with silica·bodies in
the wood-parenchyma of an undetermined species of Canarium from Borneo.
In Boswellia Carteri the bast-fibres composing the sclerenchymatcus ring are
septate like the wood-fibres. In Protium divaricatum there is a ring of stone ..
 ADDENDA __BURSERA CEAE
cells in the primary cortex (Stepowski). The resin-canals in Boswellia Carler;
are found also at the periphery of the pitli (protoxylem) .
. In BOS1lJcllia Carteri the cork arises subepidermally. According to Holmel
and Ad. Peter (see also Mohl), the peculiar exfoliation of membranous cork-
layers, already (Syst. AnaL, p. 193) described for species of Boswellia and
Commiphora, takes -its origin from single layers of phelloid-cells of peculiar
structure; the inner tangential walls and the adjoining portio~ of the radial
walls are strongly thickened, lignified and silicified, while the' remaining
parts of the wall are very thin; moreover the siJicified portions flre rarely
smooth, but have delicate ridges nmning in the vertical direction and occasion-
ally forking. The process of exfoliation does not, however, take place in every
!arer of phelloid cells. so that unruptured layers of these cells may be met
Wlth in the cork.
Literature: Mobi, in Bot. Zeit., J861, p. 229.~Hohnel, Kork, Sitz.-Eer. Wiener Akad., buc\·i,.
Abt. I, 1877, p. 605 etc.-Johannson, Noch wenig bek. Rinden, Diss., Dorpat, 1891, p. 31 et seq.-
Jadin, Terebinthacees, Journ. de bot., 1893, p. 382 t:t seq.-Boergesen og Paulsen, Vegetat. dansk.-
vestind. Oer, llot. Tidsskrift, xxii. 1898-9, pp. 97; 98 (Burseragummifera, L.).-Pitard, Pericycle,
These, Bordeaux, J901, p. 84.-HargagIi.Petrucci, Concrez. silicee, Malpighia, 1902. p. 23 et seq. ;
and Legnami, loc. cit., p. 3J4 et seq. (Cal1arill1Jl, Santida).-Poulsen. Luftrooder hos Canar'ium
C()fI""U1Je, Vidensk. Meddelels. Kjifbenhavn, 1902, pp. 331-5.-Ad. Peter, Anat. d. Veg. Org. von
Bo.rwe!Iia Cartr:ri, Sitz.·Ber. Wiener Akad., cxii, Abt. r, 1903., pp. 511-34 and Tab. i-iii; see also
Anzeiger d. Akad., 1903, p. I69--Areschoug, Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39,
n. 2, 1905. pp. I 34-6.-Stepowski, Anat. Untersnch. iiber die oberird. Veg. Org. der Burseraceen etc.,
Dlss., Bem, 1905, pp. IJ-,; l.-[Boorsma, Aloeholz, Bull. Depart. de l'Agric. aUK lodes neerland.,
vii, 1907, p. zR et seq. (Calla;ium~.J

MELIACEAE (l?P. 194- 198).

I. To the REVIEW OF TH~ ANATOMICAL FEATURES the following newly'
discovered facts may be added: Unicellular, two-armed hairs are found also
in Epichal'is. Hypoderm is present in the leaf also in certain species of Aglaia
and Sandoricum. Papillae occur on the lower epidermis of the leaf also in
Heynea.
2. STRUCTURE OF THE LEAF. Hypoderm is found also in Carapa obovata,
Bl. (according to Areschoug), Aglaia cambodiana, Pierre and Sandoricum
indicum. Cav. (according to Pierre). The latter author states that the lower
epidermis in H eynea trijuga, Roxb. bears rather long papillae. .
The secretory cells characteristic of the members of this Order are accord.
ing to Pierre in some cases surrounded by special cells resembling an epithelium.
Mucilage-cells are stated by Areschoug to occur in the palisade-tissue of Carapa
obovata.
To the section dealing with the hairy covering we may add the following
information. Simple unicellular clothing hairs are present also in species of
Chisocheton and DysDxylum, and unicellular two-armed hairs also in EPi-
charis Juglans, Hance, and E. hoaensis, Pierre. Glandular hairs, sunk, like
those of Cabralea, in deep pits, the apertures of which appear as dots on
the surface oi the leaf, occur also in Dysoxylum Lour.eiri, Pierre (Pierre).
According to Areschoug, peculiar hydathodes~are found on the lower surface
of the leaf in Carapa obtusa; they consist of a small-celled tissue situated
beneath the epidermis, which subsequently becomes resorbed at these points.
~ Yate. We may add that according to Pitard the fruit-stalks of Swietem:a
Ma7iagoni and Aglaia Raxburgh'ii exhibit polystelic structure. For the detailed
structure of the hom-shaped respiratory organs, found in Carapa moluccensis, Lam.
(and exhibiting a peculiar formation of intercellular spaces in the primary cortex),
see Karsten, loco cit.
Li.terature: ~arsteJ), Mangroveveget., I'ib!. bot., Heft :n, 1891, pp. 51, 52.-[]{usby, CobJentz
and Wilcox, Coccilb.na (Gua"ea), Bull. of l'hnrm., 1893, p. 350 et seq.l-·Pierre, Flore forest. de lao
 ADDENDA-MELIACEAE
Cochinchine, xxii, IS96, ano xxiii, 1897.-[Mitlachtt, Meliaceen-Rinden, Zeitschr. aUg. osterreich.
Apotheker-Ver., 19\)0, p. 573 et seq.; abatr. ~in Just, 1900. ii, p. 47.]-Pitard, Pericycle, These,
Bordeaux, 19<)', p. 93.-Areschoug, M~grovevegetat .• Bib]. bot., Heft 56, 190::' pp. 46, 47 and Tab.
iii-iv.-Dargagli-Petrucci, Legnami, Malpighia, J90OJ, p. 316 ct seq. (Carapa, Dysoxylum, Sant/()·
ricum).-Pitard, PoiysteIie, Actes Soc. Linn. de Bordeaux, ser.6, t. vii, 1901.-Areschoug, Trap.
vut. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2,190 5, pp. 98, 99 (ez:podessa).-Piccioli, Legnami,
Bull. Siena, I~, pp. 147, 149 and I63.-[For additional literature see p. IItl.]

CHAILLETIACEAE (pp. I98-zoo).

Literature: C. de CandolIe, Inllor. epiphylles, Mc;m. Soc. de phys. et d'hist. nat. Geneve, 1890,
vol. supp., sep. copy, p. '4 et seq.-Pitard, Pchicycle. These, Bordeaux, 190 1, p. 93.

OLACINEAE (pp. 200-209).

I. ANATOMICAL FEATURES. First, as regards the structure of the leaf,

we may add that stomata with subsidiary cells placed parallel to the pore,
previously recorded in Opilia, are pres en t also in Coula and other genera, hypo.
derm also in species of Scoopfia, spicular cells in Anacalosa puberula, Kurz,
and spiCUlar fibres running freely in_ the mesophyll in Minquartia and
Eganthus I. In the Olacineae recently investigated by Colozza, the wood-
prosenchyma also bears bordered pits in all cases. Laticiferous tubes. already
recorded in Endusa and Cardiopteris, are present also in Coula, Eganthus,
Minquartia and Ochanostachys,_ whilst SChizogenous secretory cavities, besides
occurring in Coula and Endusa, have been observed in Eganthus, Minquartia
and Ochanostachys. Amongst noteworthy types of: hairs we may mention
the branched multicellular trichomes of Ximenia calfra, Sond. and the tufted
hairs found on the branches of Coula, Ochanostachys and M inqt-tartia.
2. STRUCTURE OF 1'HE LEAF ~. Van Tieghem states that the leaves in Coula,
Ochanostachys and Minquartia have bifacial structure with the stomata on the
lower side. According to the- same authority the stomata are accompanied
by subsidiary cells placed parallel to the pore in Coula edulis, and according to
Pierre in A nacalos~ Clar~ii, Pierre, M elientha sttavis, Pierre, Ola_x (~b~cata,
RoXb. var. cambod'tana, PIerre, Schopfia jragrans, Wall. and S. MJeTSU, PIerre,
and according to Gerhard in Apodytes dimidiata. The two species of SchOpfia
just n.amed have a hypoderm composed of two layers. According to Colozza,
the mesophyU in Heisteria cauli/lora, Sm~ arid Anacalosa puberula, Kurz con-
tains • sclerenchymatous idioblasts: while in Minquartia and Eganthus, accord-
ing to Van Tieghem, it inCludes sclerenchymatous fibres, having --the same
structure as those of Endusa.
According to Van Tieghem, three vascular bundles pass out into the leaf
in Coula, Minquartia and Ochanostachys; the two lateral bundles branch
off from the vascular ring of the axis a little way below the node. Ac ..
cording to Pierre, there are likewise three bundles in A nacalosa, A podytes;
Erythropalum, Olax.and Strombosia. while in Melientha, ScMpfia and Ximenia
there is -only· a single bundle. The petiole, according to Colazza, contains
a stele in Coula, Heisteria, Ochanostachys, Scorodocarpus and Strombosia, while
in A nacalosa, Liriosma, Olax and X imenia the vascular bundles are arranged
to form an arc.
A few facts rnay be taken from the new records of the occurrence of

I Van Tieghem unites Cou/a, Egan/litis, ElIdttsa and Millqllal·tia to form a separate Order
(Coulaceae). .
• Cotozza's recent jnve.tigations all the structure of the leaf deal with the genera AnacallJsa.
Coula Htis/eda, Liri()sma, Ocha1Wstacnys, Olax, Sirombosia and Ximmia; Colozza moreover also
exami'ned the structure of the axis in these genera, as well as in SckOpfia.
872 ADDENDA--OLACINEA£
secretory organs mentioned above. A.ccording to Van Tieghem, the laticiferous
tubes found in Coula 7 Minquartia, Oeha-nostaehys and Eganthus are branched
and unseptate (' non cloisonnees') I; special emphasis is laid on the latter
pomt) at least as far as tile three first-named genera are concerned. The
latieiferous elements are present in the pith, primary cortex and. bast of the
branch; in the leaf they are met with especially in the veirut. According
to Van Tieghem, the secretory cavities occur in the primary cortex of the
branch II and in the mesophyll; in Coula, Minquartia and Ochanostachys their
contents, as in the case of Endusa, assume a blue colour after treatment with
Eau de Javelle.
Colazza has published a number of new statements on the mode of deposi-
tion of oxalate of lime in the axis, the lamina of the leaf and the petiole. Of
these we may mention that solitary crystals occur also in Coula, Lil'iosma,
Ochanostachys, Strombosia and Ximenia, and clustered crystals also in Anacalosa,
Coula, Heisteria and Ochanostachys.
We may lastly mention the speCial forms of I clothing hairs that have been
recently recorded. Colozza describes the trichomes of X imenia cagra as 4 peli
plut?c~~ari~, sempli.ci 0 ramificati,' Van Tieghem those of Coula, &c., as
polls unisenes, ramIfies ~ la base en forme de bouquet.'
3. STRUCTURE OF THE AXIS. The previous account of the structure of the
coI"!.eJC needs the following additions. I~ Coula, M inquartia and Ochanostachrs
~he development of the cork takes place m the subepidermal layer of cells; In
these genera cells with strongly thickened tangential walls are found amongst the
thin-walled cells of the cork (Van Tieghem). According to Van Tieghe~ the
pericycle in the three genera just named c:ontains a composite and continuous
ring of sclerenchyma; Colozza describes: bands of bast-fibres as present in
the pericycle in species of Liriosma, Olax, SchOpfia and Ximenia, and a ring
of sclerenchyma in species of Anacalosa, Heisteria, Scorodocarpus and Strom-
bosia, while Pitard records isolated groups of bast-fibres in the pericyc1e of
Heisteria coecinea and Olax imbricata (Fissilia psittacorum). Secondary hard
bast has been observed in Liriosma (Kleesattel).
According to Leisering, the interxylary phloem found in Sarcostigma
Kleinii is given off by the·cambium on its outer side, but subsequently pecQlIles
bridged over by a cambial arc; in other words, its mode of de~alopS)J.en\ ii
that characteristic. of the Stryehnos~type. .
Literature: Wijnaendts Francken, Sklereiden, Diss., Utrecht, 1890, p. 52.-Kleesatttl, Malta
Puama, Diss., ,Erlangen, 1892, 44 pp., 2 Tab.-Pierre, Flore forest. de la Cochmchine, xvii, 1893 .
.-Leisering, Interxylaeres LeptQm, Diss., Berlin, 1899, p. 17.-Van Tieghem, J;).eQJ: genres nouv.
pour Ia (am. d. Conlacees, BulL Mus. d'hist. nat., 1899, PP'.97-Joo.-Van 'rre~hem, Coulacees,
J~tlm. de ~t.) 1~99, pp ..69-79; a.nd Ann. sc. nat.,:et.8, :. ,., 1899, pp. Il5-36._:'H~kel,. ~ara.si­
tll.~me de Xzmema americana, Comptes rendus, Pans, CXXXI, 1900, pp. 764, 76s.-P'itar<l, ;fleCicycle,
These, Bordeaux, 1901, pp. 84 and 93.-Bargagli-Petrucci, Legtlami, Malplghia, 1902, p. 293
(S,orodocaryus).-Gerhard, BI:l.ttanat. v. Gew. d. Knysnawaldes, Diss., Basel, 1902, pp. 10, 11
(AJot/ytes).-Van Tieghem, Coulacees, Joum. de bot., 1902, p. 22S.-Colozza, Anat. delle Olacin.,
Nuovo Giom. bot. ltal., Xi7 1904, pp. 539-6,5.

OCTOCNEMACEAE.
We may follow Engler (Natiirl. Pflanzenfam., Erganz.-l;Ieft, i, I 900,
p. 19) and Van Tieghem in regarding the genus Octo enema as constituting
a special Order. Octocnema is characterized by the following anatomical

I Van Tieghem also describes the laticiferons tubes present in Etuiusa .s unseptate, but this is
not the case (see Syst. Anat., p. 202); he himself observed the occurrence of anastomoses in this
genus.
S Colozza's statement that In COllla and OchanostacR),s the secretory cavhies occur also in the
pith is no doubt incorrect.
 ADDENPA--OCTOCNEMACEAE
features: multicellular tufted or steUate clothing hairs; oxalate of lime in
the form of solitary crystals; stomata without subsidiary cells; absence of
cortical vascular bundles; a composite and continuous ring of sclerenchyma
in the pericycle; occurrence of secondary hard bast; and subepidermal
development of cork.
The two species, O. Klainea1ta, Pierre and O. affinis, Pierre, have been
examined by Van Tieghem. The hairy covering varies somewhat in the two
cases, being-floccose or tufted in O. Klaineana, and composed of stellate hairs
having their ray-cells spread out parallel to the surface of the organ in O. afftnis.
The mesophyll is compact on the upper side of the leaf and lacunar towards
the lower side; the outer layer of the compact portion contains thick crystals.
The stomata are confined to the lower surface of the leaf. The vascular bundles
in the lateral veins are accompanied by hard bast and provided with an
endodermis containing crystals. Five vascular bundles pass out into the leaf;
for the further course of these bundles, which varies slightly in the two species,
see Van Tieghem, loco cit.
As regards the structure of the axis, we may first note that the corl)., as
above mentioned, develops in the subepidermal layer of cells. The.' cells of
the cork are sclerosed on the outer tangential and radial walls. The primary
cortex contains stone-celis, which are either thickened uniformly or in the
form of a horseshoe; in the latter case they enclose a crystal. The endodeanis
is not distinctly differentiated, although many of its cells contain crystals.
In young branches the peri cycle comprises isolated groups of bast-fibres, but
in later stages a composite and continuous ring of sclerenchyma is developed.
The secondary bast of the thicker branches contains small groups of bast-fibres,
which are arranged in several layers and are enveloped by chambered crystal-
fibres with solitary crystals. The medullary rays of the wood are uniseriate.
Nothing is known as to the structure of the vessels and wood-prosenchyma.
The pith includes groups of stone-cells, whilst in its peripheral portion an arc
of lignified tissue is situated opposite each group of' primary xylem.
Literature: Van Tieghcm, Octocneme, Jouro. de bot., 1905, pp. 45-58, especially PP"/~7-50.

ILICINEAE (pp .. 20g-2II).

2. 'STRUCTURE OF THE LEAF. Supplementary observations on the structure
of the leaf areJ;ontained in Cador's work (loc. cit.) and in the literature cited
below. In sllrface-view the epidermal cells show straight or undulated lateral
margins; '.in· other cases they appear to be undulated at a high focus and
straight at a low focus, this feature being combined with the presence of marginal
pits. A two-layered epidermis on the upper side of the leaf has been recorded
also in !lex canariensis, Pair. andI. capensis, whilst:in 1. theezans, Mart. var.
tertiUs, Loes:rand yar. Riedelii, LoeS. the epidermis consists locally of two layers ;
in I. chamaediyfolia, Reiss. var. typica, Loes. occasional horizontal division-
walls are met with in the upper epidermis. A one-layered hypoderm is stated
to occur on the upper side of the leaf in I. platyphyUa, Webb et Berth. Gela-
tinization often affects almost all the cells of the upper epidermis, while in other
cases isolated cells of both upper and lower epidermis exhibit this feature.
Gelatinization has been recorded in the following species and varieties: !lex
afjinis, Gardn. var. genuina, Loes. and var. rivularis, Loes., I. amara, Loes. var.
longifolia, Loes. and var. lati/olia, Loes., I. Caroliniana, Loes., I. Cassine,
L. var. myrtifolia, Chapm., I. chamaedrylolia, Reiss. var. typica, Loes., I. cogllata,
Reiss., I. Congohinha, Reiss., I. conocarpa, Reiss., I. Cujabensis, Reiss., I. dumosa,
Reiss. var. Guaranina, Loes., I. glabUl, Gray, I. Glazioviana, Loes., I. Para-
 ADDEN04-ILICINEAE

;.ariensis, S1. Hil. vaT. genuina, Loes., I. Pseutiothea, 'Reiss., I. symPlociformis,

eiss. In some cases the epidermal cells ~xhibit a p~ade-like elongation in
ansverse sections of the leaf, e.g. in I. theezans var. tertitis. The cuticular
:lges occasionally form a kind of rampart around the stomata; in I. theezans
LT. typica the stomata are surmounted by a r\4ge-like elevation or a chimney-
aped space, as the case may he, due to the neishbouring cells projecting m
.e form of a ridge. The neighbouring cells of the stomata in some cases
aT a slight resemblance to subsidiary cells, e.g. in I. Paraguariensis. The
esophyll commonly contains fat-bodies. In I. Cassine var. myrtifolia the
'0 lowest layers of the spongy tissue consist of cells with thick pitted walls;
e palisade and spongy tissue in I. glabra contains round cells having wide
rnina and sclerosed on one side.
An additional feature presented by the hairy covering is the occasional
currence of long, unicellular trichomes. The clustered crystals of oxalate
lime are frequently enclosed in relatively large cells, which are ,differentiated
idioblasts. In IJex Cassine var. myrtifolia~ I. d'Umosa var .. Guaranina, and
paltario ides, Reiss., Cador met with sphaerocrystaIline masses (hesperidin 1)
the epide;rp.is of the leaf, while in I. Pseudothea he observed yellowish crystals
varyIng size and composed of an unknown _chemical substance.
Fer the distribution of the abundant cork-warts, which in certairl species
lSe a punctate appearance on the lower surface of the leaf, see especially
esener's monograph (in Nova Acta Leopold.-Carol. deutsch~ Acad., lxxviii,
)1)' regarding the occurrence of domatia and their structure, see Loesener
Biolog. Centralbl., 1893, p. 449 et seq.
3. STRUCTURE OF THE AXIS. According to Pitard, a composite and con-
, . ring of sc1erenchyma is present also in Ilex celastroides and Byronnr
"ous
,enSJs.
Literature: Pierre, Flore forest. de la Cochinchine, xviii, 1893.-Loesener, lIes paraguarim,m,
ubI. Berlin, D. la, 1897. p. 31..~ et seq.-C~dof, Anat. Untersuch. der ~atebl., piSS., Erla~"
)p.; sep. copy from Bot. Centralbt, 1900, IV, p. '41 et seq.-Kearny, In Contnbut. U. S. Nat.
'b. t V, 5, 19°1, p. 296.-Petersen, Vedanatomi, 1901, pp. 50. 5 1.-Pitard, Pericycle, These,
de~, 1901, p. 72.-Bargagli-Petrucci, Legnami, Malpighia, 190:1, p. 3~3.-Clauditz;Blattanat.
Lf. Gew., Diss., Basel, H)02, pp. :l3-6.-Gerhard, Blattanat. v. Gew. d. Knysnawaldes, Diss.,
~l, 1903, pp. I1-J41.-Neger and Vanimo. Paraguay thee, 19°3, pp. 449-53.-Piccioli, t;egnami,
I. Siena, J906, p. I36.-[For additional literature see p. u70.) :;

CELASTRINEAE (pp. 212-214).

Thanks to the recent investigations of Stenzel and Metz, the structure of

leaf and axis in this Order is now well known. Their work net:essitates the
owing alterations or additions in the GENERAL DIAGNOSIS:!. The development

1.The plant which Gerhard describes under the name of 'Il~x Cassini' (sphalm,. ex. ' 1. Ca.rsint .),
1 specimens growing in the Botanic Gardens at Basel, does not belong to the genus Ilu, u Is
;on by the anatomy of the leaf (spicular cells, peltate glands); except for the statsp_ertt as to the
nence of clustered crystals there is nothing against the pll1.nt ill question being a. member of the
~eae.

• Stenzel's and Metz's investigations extend to the following genera, which are enumerated in
serial ordtt of Loesener's system: Euonymus, Lopltopttalum, *GlyptojetaluJ", A/icrotroji':>,

.*
lJamia, CeiastyUS, 1I1aylemts, GYJ)ln()sjo,-ia, Plillerlz(_'kia, Catha, Pterocclaslrus, Po/ycardia,
rimia, Padr),stil1lo, .. Koi.'oona, Zillowiewia, Plenckia, Tri'pterygiulIl, IVimmeria, ElaetJdendron.
ine, lllys(rQxyloll, Mauroania, HartQgia, PltttroslylUz, ... Lallridia, ··GYlllind4, MJ'Cinda.,
111M/era, Iflorlolda, *Gloss()petalttm, Sc/ztUjfiria, ... PerrolleJia. Goupia, and Siplumotlim. The
ra provided with an * have been investigated .y Stenzel only. those marked with •• by Metz
; Stenzel's work de!).ls with the stnlcture of the leaf nnd axis, whilst that of Metz Is concerned
with the structure of the leaf.
 ADDENDA-CELASTRINEAE

of the cork occasionally takes place in. a deeply situated cell-layer of the primary
cortex. The hairy covering consists of unicellular or uniseriate clothing hairs,
the former often being short and differentiated as papillae; hairs are not of
common occurrence and are never' present in large numbers. Special forms
of clothing hairs (such as uniceJ.4¥ar one- or two-armed" hairs, and forked multi-
cellular trichomes) are very roce. ' A widely distributed feature is the occurrence
of small bodies, consisting of caoutchouc or fat, in the mesophyll. Special
features in the structure of the leaf may be briefly enumerated as follows:-
palisade-like differentiation of the epidennal cells; papillae on the epidermis
(very rare); mucilaginous epidermal cells (very rare); peculiar pit-canals in
the outer wall of the epidermal cells (Mortonia); hypoderm or an epidermis
of several layers; crystal-cells in the epidermis; spicular fibres in the meso-
phyll. The secretory receptacles are represented by: (a) long secretory sacs,
which are mostly filled with a substance resembling caoutchouc and capable of
being drawn out into threads Cspinning ') (Celastrus, Euonymus, Mystroxylon,
Wimmeria); (b) secretory canals (Mortonia, Pachystima)_; and (c) tannin-
idioblasts. The earlier statement as to the presence of 'resia-cells?' in
Kokoona has been traced to the occurrence of cork-warts on the lower side of
the leaf.
In the STRUCTURE OF THE LEAF the epidermis in the first place kftords
a large number of systematic characters. The size of the epidermal cells,
the nature of their lateral margins and the thickness of the outer'walls, are
features which are subject to variation. Palisade-like elongation qf the
epidermal cells is found in Cassine scandens, Eckl., Gymnosporia laurina, Szysz.,
'. KORoona zeylanica, Thw., M aurocem:a Frangula.ria, Mill., M ortonia Greggit',
Gray, Polycardia Hildebrandtii, Baill., species of Ptero celastr us, and Put'"-
lickia pyracantha, Endl. Striation of the cuticle is met with in species of
Microtropis, Polycardia and Zinowiewia, especially in the neighbourhood of
the stomata. In Catha edtdis, Forsk. and Goopia glabra, Aubl. the outer walls
bear linear pits; in Polycardia Hildebrandtii, Baill. and P.Ube1'a, O. Hoffm. they
are- provided with delicate pits. In Mortonia Greggii, Gray the outer walls
are tJ aversed by peculiar long pit-canals, which may be either branched or
unbranched, and run in all directions, though for the most part parallel to
the principal vein; the outer ends of these canals invariably come tp lie above
the lateral margms. Marginal pits have also been observed in forms having
epidermal cells with undulated lateral margins (species of Cassine, Elaeoden-
dron, Gymittda, Ku"imia; Lauridia). In some cases (species of Elaeodendron,
Gymnosporia, Maurocenia, May tenus, Microtropis, Myginda) the lumina of the
epidermal ,ells are considerably narrowed owing to the thickening of the walls.
Mucilaginous epidermal cells have only been recorded in Goupia glabra, Aubl.,
Perrottetia alpestris, Loes. and P. sandwicensis, Gray. According to Metz,
occasional epidermal cells are very commonly drawn out into papillae or short
papillose hairs: A typical papillose epidermis is, howeverl present only in
Wsmmeria 'confusa, HemS!. (on the upper side of the leaf) and in Siphonodon
celastrinus, Griff. (on the lower side of the leaf). In certain species of Gymno-
sporia, Ct'.ft}Ia, Elaeodendron and Plenck£a the epidermis consists locally of
two or three layers owing to the presence of division-walls parallel to the
surface of the leaf, while in Goupia glabra, Aubl., Gymnosporia linearis, Loes.,
G. senegalensis, Loes. and G. Wallichiana, Spreng. the epidermis is typically
two- or three-layered. In Goupt'a ~labra the horizontal division-walls are
accompanied by vertical ones; vertIcal walls are also present in species of
Euonymus, Gymnosporia, Kurrimia and Polycardia. Hypoderm is of frequent
occurrence. A continuous hypoderlp consisting of one or more layers and
situated either only on the upper or on both sides of the leaf has been observed
in certain species of Cassine, Celaslr'Us, Denhamia, Elaeodendron, Gyminda,
 ADDENDA--CELASTRINEAE
Gymnosporia, Maurocenia, May tenus, Myginda, Mystroxylon :and SchaeDeria 1.
In other species belonging to these genera, as well as in Ptc1Ickia populnea,
Reiss., the hypoderm is confined to certain parts of the leaf, situated especially
in the neighbourhood of the veins. A last feature requiring sp~cial mention is
the occurrence of crystal. cells in the epidernris of the leaf in species of Catha,
Denhamia, Elaeodendron, Euol1ymus, Gyminda, Kurrimia, Lophopetalum,
};[aytenus, M iCTotropis, M yginda, Pleurostylia, Siphonodon, and W immeria 2.
In most cases each of the crystal-cells contains a solitary crystal, rarely a
clustered crystal. These crystal-cells are found either in both upper and lower
epidermis, or only in the lower epidermis, and may be present in large or small
numbers. In certain species they differ from the remaining epidermaJ cells, this
being especially the case in the species of Elaeodendron. Here the crystal·cells
either form rows of as many as twelve cells or groups of two to six cells lying
side by side, or are isolated; in the latter case they are surrounded by a kind of
rosette formed by the neighbouring cells; the crystal-idiobl~ts are further
frequently· rounded and smaller than the other epidermal cells, and may even
have thickened inner walls in which the crystals are inserted. Gyminda
5hows similar features to those just described for the species of Elaeodendron.
In Sil;honodon the crystal-cells are distinguished from the other epidermal
cells By the absence of papillae. In Lophopetalum and Microtropis discolo1',
Wall. ordinary cells of the epidermis are divided up into two or three chambers
by means of delicate walls, each chamber enclosing a clustered crystal.
The stomata are as a rule confined to the lower side of the leaf. They are
developed in considerable numbers on the upper side of the leaf, for instance,
in species of Gymnosporia and May tenus, as well as in Mortonia. There is no
uniform type of stoma. In most cases the stomata are surrounded by 3, 4 or
more neighbouring cells, but in many species some cf the ,stomata are accom-
panied by subsidiary cells placed parallel to the pore. According to Met.z,
the Rubiaceous type is well marked in Kurrimia; in Lauridia there are mostly
two pairs of subsidiary cells arranged cross-wise, and in Mortonia the pairs of
guard-cells are surrounded by a rosette of smaller epidermal cells. .
The leaves for the most part show bifacial structure. Distinct centnc
structure with palisade-tissue situated on both sides of the leaf has been recorded

1 The species aTe: Cass;"ne btm'lara. L., C. "apens~'s, L., C. sjhatrvphylla, O. Kl:ze. ; Celastrus
australis, F. v. M., C. disjJerma, F. v.. M., C. papualla, '''arb.; lJenna11lia o/;scura, ~teissn:;
Elae,xU"dron australe, Vent., E. capens~s, Eckl. et Zeyh., E. croceum. DC., E. g/aucum, Pel'S.,
E. ilicifolium, Hochst., E. orientale, Jacq., E. pap£llusum) Hochst., E. ,]uadrangulatu"" Reisi.,
E. Schweinjurtnianzan, Loes'} E. xylocarpum, DC.; Cymin:Ja Grise/;acnU, Sarg.; Gym~spq;ia
allgu/aris, Loes., G. buxifllia, Szysz., G. Ctlnningnamii, Loes. G, lucida, L<>es., G. prtxumhens,
j

l.oes., G. venmaia, Szysz., aiso C. (1) octracilis, Pierre and C. llieko11gensit, l'ierre, according to
l'ierre_; Maunxenia .Frangularia, Mil1.; May tenus Db/usi/alia, Mart., M. pk)lUan(koifies, Benth.,
AI. vzscifolia, Griseb.,.AI. Vil£s Idaea, Grir.eb.; AIyginda Gaumen', Loes., AI. ilicifolia, LaDl.,
ill. /atijo/ia, Sw., A-I. pallms, Sm., M. Rhamma, Sw. ; lI:fystrcxylon jubescens, Eckl., M. sphatro-
p.ltyllmll, Eckl. et Zeyh. 13 /itorale, Harv. et Sond. ; Schae.lftrit:Zfrutescens, Jacq.
, The species in question are the following: Catna dulls, Forsk. (clust, cryst.); nenlzamia
ooscura, Melssn.; Elatodmtiron Qustrale, Vent., E. cajJI!1lte, Eckl. et Zeyh., E. C1'OC&lIm, DC.,
1:.'. g-lauClIm, Per:>. (dust. cryH. or solitary and dust. cryst.), E. iIicifoHum, Ten., E. or:'en/ale, Jacq.,
E. pajillosUfIl, Hochst., E. quadrangulatum, l{eiss.., E. Schweilt/uythiamml, Loes. (solitary and
dust. cl)'st.), E. xylo,arpum, DC.; Eflo'lzymus ame.-icaltus, L.; Gyminda GYisebochii, Sarg.;
.liurn'mia (according to Stenzel); Lclhop~talum jimbriatum, Wight, L. WightiauUf1I~ Am.;
fllaylenus acallfhophyIla, Reiss" /If. aquifolia, Mart., M. basiamtata, l{eiss .• /1[. CaiinKorurn, Reiss.,
lIf. Itorrida. Reiss., lJ. iliCl/olia, Mart., M. farois, Reiss., .At. llIacrophylla, Ma.rt., M. Miille"i,
Schwacke, ~'tI. 11lyrsincides, }{eiss., M.oUttsi/o/ia, Mart., If'f. rigMa, Mart., J!1. viscifdia, Grjseb.;
Alicrotropis eiva/vIs, Wall., Af. dms(!lora. Wight, M. disc%r, \'Ir"aJl., AI. latifo'ia, Wight. AE. ovali-
folia, Wight; l'.fyginda Cros!opetalum, L., AI, lali/olia, Sw. i Pleuroslylia W:;;/tIU, Wight et Am.
(acicular crystals); Sip/mIt-dolt celasln'nus, Griff. (dust. cryst.); 1-V~·m11le,ia serrolala, Radik.
(solitary and c1nU. cryst.). Except where there is a special statement to tr.e c()ntrary, only solitary
crystals are pcesent.
 ADDENDA-CELASTRINEAE

in species of Gymnosporia, M aytenus and M ortonia, while the mesophyll is

homogeneous in species of Gymnosporia, Myginda Gaumeri, Loes. and Zino~
wiewia integerrima, Turcz. Other noteworthy features are; palisade-cells
exhibiting transverse division (in species of M ortonia and Pachystima); layers
of rather strongly thickened cells (in the palisade-tissue in Elaeodendron Schwein-
furthianum, and in the spongy tissue in M aurocenia); and spicular fibres
(in species of Gymnosporia, Mauracenia, May tenus, Microtrapis, Pteroce!astrus,
and Schaejferia 1). The spicular fibres branch off from the sclerenchyma of
the veins and penetrate more or less deeply into the mesophyll; in some cases
they extend as far as the epidermis,or may even spread out between the epidermis
and the mesophyll. They are rarely present in any-considerable numbers.
According to Pierre, the petiole is supplied by three vascular bundles in
Kurrimia, but only by a single bundle in Celastrus, Elaeodendron, E~lOnymus,
Gymnosporia, M icrotrapis, Pleurostylia and Siphonodon. The vascular system
of the petiole and midrib is either of a horseshoe form or annular and closed.
In the latter Case medullary vascular bundles are occasionally (Goupia, Kur~
rimia, Lophopetalum) found within the 'vascular ring, and in Goupia glabra
there is also a cortical bundle situated in each of the two angles of the petiole.
The vascular bundles of the veins are for the most part accompanied by scleren-
chyma. Vertical trans currence of the smaller veins has been observed in species
of Cassine~ Plenckia, Tripterygium and Wimmeria.. Enlarged terminal tracheids
are present in species of Goupia, Myginda and Schaejferia. while similar re-
ticulately thickened cells accompany the vascular bundles of the veins in
Polycardia libera~ O. Hoffm.
The description of the structure of the leaf may be followed by a general
discussion of the hairy covering, the mode of deposition of oxalate of lime
and the secretory organs; these will be taken in the order in which they
have been named. The hairy covering does not include glandular hairs:l.
The clothing hairs-are confined to a relatively small number of genera and only
rarely form a dense covering. The most widely distributed type of clothing
hairs are short unicellular trichomes (species of Euonymus, Fraunhofera, (Joupia,
Mystroxylon), which are either papillose or of slightly greater length. Tri-
pterygium has papillose hairs composed of one or two cells, while in Fraunhofera
the papilla-like trichomes are accompanied by long hairs, which are either
unicellular or multicellular with thin ti"ans~erse walls. Myginda has unicellular
one-armed hairs, which in M. ilicifolia, Lam. are accompanied 'by unicellular
tri..::homes with two arms. In Wimmeria, lastly, the hairs are tubular multi-
cellular structures connected by numerous· transitional forms with unicellular
papillose hairs. In W. microphyUa, Radlk. they are commonly seated on two
basal cells,. the lower of which bears a spinose process- at its upper end; in
some cases this process is strongly developed; so that forked hairs result.
Oxalate of lime is deposited in the form of solitary or clustered crystals,
and as crystal-sand. The solitary crystals are generally rhombohedral, although
sometimes~~'~ery .long • (in the palisade-tissue of Kurrimia) or 'rod-shaped'
(in Frattnhofera) Of ' hastate or acicular) (in the tissue of the leaf of Glosso~
petalum spinescens, Gray and in' numerous species of May tenus). The crystal·
sand never occurs independently, but is invariably found together with a solitary

1 viz.: Gymnosporia tmarginata, Roth, G. laur£l1a, Szysz., G. tn'ala, Laws., C. Royleana,

Wall.; Afauroceuia Frangularia, Mill.; Jr,laytenus aca1lIIIOphyl/a, Reiss., Jr,f. amazonica. Mart"
11-1. laevis, Reiss.; Jr,Iict'o/ropis densijlora, Wight, AI. discolor, Wall., 11-£. lali/o/ia, Wig!lt, AI. Mla/j-
folia. Wight, Jr,f. ,amijlora, Wight; Pterocelas/NIS 'l'an'abilt's, Sond. var. li/oraNs and'var. ootusilolJus,
P. 1"Oslra/u!, \\o'alp. ; Sch{Uffiriafrut~sce1l!. Jacq .
• I Stenzel's statement regarding the occurrence of peltate glands in Rhacoma micropllJ11Ia, Loes.
is incorrect. The plant examined by Stenzel does not belong to the Celastrineae, but is probably
a member of the Verbenacene.
 ADDENDA-CELASTRINEAE

or clustered crystal, e.g. in species of Elaeodendron, Maurocenia, Microtropis,

Mortonia, Mystroxylon, SipJwnodon. Within the limits of the same genus
the tissues of the leaf and axis in some cases contain either solitary crystals
only or clustered crystals only, or the two forms occur side by side. In other
genera the forms of crystals found in the leaf and axis are different in the two
cases; most commonly the leaf contains clustered crystals only, while the
axis includes both solitary and clustered crystals. In Denhamia. Kurrimia.
LopJwpetalum and Pleurostylia solitary crystals alone are present 'in the leaf
and axis; with these genera we may class FraunJwfera and Plenckia, which
likewise have solitary crystals only, although in this case the crystals are con-
fined to the leaf. In Euonymus, Glyptopetalum, Pachystima, Perrottetia and
Tripterygium only clustered crystals are found in the leaf and axis; in Goupia
also only clustered crystals have been recorded, but they are only present in
the leaf. Finally, solitary and clustered crystals occur side by side in ·both
stem and leaf in Cassine, Elaeodendron and M aurocenia. The occurrence of
oxalate of lime in the epidermis has already been dealt with above. Other
features requiring mention are: the occurrence of relatively large crystal-idio-
blasts in the mesophyll (species of Cassine, Catha, Celastrus, Euonymus, &c.) ; sep-
tate palisade-cells, the chambers of which contain crystals (Elaeodendron capense,
Vent., Mortonia Greggii, Gray); cells with unilateral or uniform thickening.
which contain solitary crystals, and are found in contact with the epidermis
on both sides of the leaf (Kurrimia) ; the occurrence of complete layers of crystal-
cells in the mesophyll (Pleurostylia Heynei, W. et A., Ptcrocelastrus tricuspidatus,
Sond.).
Among the secretory organs we may first discuss the long secretory sacs
occurring in species of Celastrus, Euonymus- and Wimmeria, as well as in Mystro-
xylon eucleaeforme, Eckl. et Zeyh. In Celastrus (C. acuminatus, L. f.) they were
met with by Gerhard in'the veins of the leaf (in the pericycle). .In Euonymus
(E. alata, Thunb., E. americanus, L.\ E. atropurpureus. Jacq., E. europaeus, L.,
E. iaponicus, Thunb., E.latifolius, L., E. verrucosus, Scop.), according to Col.
they are confined to the bast of the stem and root and are not found in
the leaf; the secretory sacs of the stem are distinguished by their late appear-
ance, so that they are not met with in the young branches. In Wimmeria
they were first observed by Radlkofer in the bast of the axis and the veins of
the leaf. Metz publishes further details as to their distributioI}. in the leaf.
According to him. the secretory. sacs are for the most part restricted to the
veins and occur in .the position usually occupiedby the hard bast. They are
found exclusively at these points in W. con/usa, Hems!., W. microphylla, Radlk.,
W. persicifolia, Radlk., W. pubescen.s, Radlk. and W. serrulata, Radlk., while
in W. concolor, Cham. et Schlecht., W. cycloca,pa, Radlk. and W. discolor,
Cham. et Schlecht. they also run freely in the palisade and spongy tissues. The
secretory sacs of Mystroxylon eucleae/orme show a distribution similar to that
recorded in Wimmeria concolor, &c. In nearly all the species (excepting
Wimmeria -pubescens) the contents of these secretory sacs are of the nature of
caoutchouc; and are drawn out in the form of threads (' spinning '). when organs
containing large numbers of the sacs are broken in two. The secretory canals
of Mortonia Greggii, Gray run at the sides of the-veins and are found especially
within the induplicate margins of the leaves; the solid yellowish cop.tents are
soluble in alcohol and Eau de Javelle. Similar secretory canals,~which, however,
had no. contents, have been obser~ed within the duplicate margins of' the
leaves in Pachystima Canbyi, Gray and P. Myrsinifes, Raf. Metz-mentiOI1S the
------ ....,.-
--~ . .--::------~--

I The' metamorpbosed fibres' of Ellonynllls Dbovalus, Nutt. (= E. flllUricaHus), mentioned by

Moller, are identical with the caoutchouc-sacs above discussed. '
 A DDENDA-CELASTRINEAE

occurrence of tannin-idioblasts 'in the mesophyll in certain species of Cassine,

Eucnymus, May tenus, Microtropis, Myginda, Pachystima, Siphonodon, Wim-
meria and Zinowiewia; these elements are distinguished from the surrounding
cells by their size or shape, and in some cases also by the thickening of the wall.
STRUCTURE OF THE AXIS. The following facts may be added to the section
dealing with the structure of the wood (Syst. AnaL, p. 213) on the basis of
Stenzel's work. The diameter of the vessels varies from ·013 (Glossopetalum)
to .084 mm. (Goupia). Exclusively scalariform perforations, previously recorded
in Elaeodendron, Goupia and Kurrimia, are found also in Perrottetia (with 30-40
bars); as in the case of Elaeodendron glaucum, the scalariform perforations
found in Glossopetalum spinescens, Gray are confined to the neighbourhood
of the primary wood. The type of perforation in the vessels has recently
been employed by Loesener a.c; a distinguishing character between Elaeodendron
(with scalariform perforations, rarely accompanied by simple ones) and Cassine
(with exclusively simple perforations). Spiral thickening of the walls of the
vessels is found also in certain species of Euonymus, May tenus, Mortonia,
Pachystima and Tripterygium 1 • Apart from the'species of Microtropis, wood·
parenchyma is present in considerable amount also in species of Cas sine, Goupia,
Hariogia, K-urrimia and Perrottetia. .
Stenzel's investigations on the structure of the cortex have afforded the
following results. In most of the genera ~ the cork develops in the subepidermal
layer of cells, but it arises in the epidermis .in Euonymus~ in the second cell-layer
of the primary cortex in Elaeodendron, in the third in Lauridia and in a still t

deeper layer in lrfyginda and Tripterygium. The cells of the cork are generally
tabular, and for the most part have thin walls; cork-cells exhibiting one-sided
sclerosis (viz. on the inner tangential walls) are found also in species of Elaeo-
dendron, J{urrimia and Maytenus. The mechanical elements of the cortex
are represented by bast-fibres and stone-cells, but in some cases (Pachystima
Myrsinites, Raf., Tripterygium Wiltordi, Hook. f., Wimmeria discolor, Cham.
et Schlecht.) they are altogether absent. The pericycle very often contains
isolated groups of bast·fibres or an interrupted ring of fibres, while in Fraunho-
lera mulH/lora J Mart.~ M aurocenia Frangularia, Mill., &c., it includes a composite
and continuous ring of sderenchyma. Stenzel particularly mentions the occur-
rence of secondary hard bast in Denhamia obscura, Meissn.
According to Stenzel, the pith is homogeneous in most of the Celastrineae.
A heterogeneous pith is present only in a. few genera, such as Lophopetalum,
M icrotropis, Perrottetia, Polycardia, and Pterocelastrus, while in Gymnospo1ia,
for example, the pith may be homogeneous or heterogeneous; Tripterygium
Willordi has an empty pith. Groups of stone-cells areo found in the pith in
many species of M aytenus and in Kokoona zeylanica, Thw. Regarding the
formation of peculiar cavities (initiated amongst cells containing clustered
crystals) in the pith of Euonymus europaeus, see Kassner, lac. cit.
Literature: Kassner, Mark einig. Holzpfl., Diss.; Basel, 1884, pp, 19-21.-Nanke, Dikotyle
Holzpfl., Diss., Konigsberg, 1886. p. 6.~Gregory, Cork-wings, Bot. Gazette, 1888, pp. 31:1-16
(Euunymus).-C_. ·de-C:in-dol!e, InOor. epiphylles, Mem. Soc. de pbys. et d'hist nat. Geneve, 1890,
~upp1. vol., sep. copy, p. 18 et seq.-Pierre, Flore: forest. de la Cochinchine, xix, 1893.-Stenzel, Anat.
0. LanbbL u. Stamme der Celastr. u. Hippocrateacelle, Diss., Erlangen, without date, communicated

1 viz.: EUOtl)'TIlui alala, Koch, E. angustifolia. Vill. • .E. atropurjurea, ]acq., E. fimbria/a,
'Vall., E. garczOnioz·des. Hooxb., E. japo"ica, Thunbo, E. lati/olia, Seop., E, lIana, .Bieb., E. (h°d-
dentalis, Nutt., E. paudJlora. Maxim .• .E. pendula, \\'fil1., E. ,l'tlulill.1. )". et M., E. vtrt'ucosa,
Scop.: lL'aylenm ilill/olia, Mart.; }llor/ollt'a Crtggii. Grny, /If. seah,.dla, Gray: Packystima
Myrtiniks, Raf.; Tripterygt"um lI/iljiJ1'd£, Hook. f.
~ These genera are: Cass;'u. Calha. Celastrus, Dellhamr'a. Frallnhq/era, Goupia. GYlllttosporia.
Har/ogja, Kokoona, Kurrimia. L~ph()Petalu",. llfaJ1tmus, .JliC1'otropis, /ller/om'a. Pachystima,
Plend.:ia; Pleuroslylia, Poloymrdia, Pterordlls/1'Its. Putterlid:ia. Scliaeffiria. IVimtllf"ia.
880 ADDENDA __ CELASTRINEAE
to the unj.,oersity of Erlangen in 1892-3, 91 pp.-Loesener, in 'Engler and Prantl, Nacbtr. z. Teil
iii~iv, 1897, p. 223.-Boergesen og Paulsen, Veget. dansk.·vestind. Oer, Bot. Tidsskrift, xxii, 1898-9.
pp. 98 (~Iygin,{a pal/ms, Sm.) and 101 (Elaeoaendrol1 x)'/ocQrpum, DC.).-Beitter, Calha tclulis,
Dis!>., Strassburg, 1900, pp. 4O-3.-Col, Lacticiferes a cootenu special dans les Fusains, Comptes
rendu!>, Paris, cxxxii, 1901, pp. 1354-6.-Peterse1l,Vedanatomi, 1901, p. 49.-Pitard, Fericycle. These,
Bordeaux, 1901, p. 38.-Bouygues, I'etiofe, These, Paris, 19001, p. 16.-Gerhard, B[attanat.. v. Gew.
des Knysnawaldes, Diss., Basel, 1902, pp. q.-10 (CeiastrllS, Elaeodmdr01l, Pttroce!astrus).-Metz,
Anat. d. uubbl. d. Celastrineen etc., Diss., Erlangen, 19°3, 78 pp.; sep. copy from Beih. 1:. bot.
CentralbL. xv.-Siissengutb, Behaarungsv~h. d. Wlirzh. Muschelkalkpfl., Diss.,· W~rzbQrg, 1904,
p. 25.-Theorin, Vaxttrichom., Arkiv f. Bot., iii, 1904, p. 5.-Areschoug, Trop. va?'t. bladbyggn.,
Sv. Vet. Akad. Handl., 39, n. 2, 19, 5, pp. 131-3 (Celaslrus).-Piccioli, Legnami, Bull. Siena, 1()o6,
p. 166.-[Hollendonner, Kork einiger EV()11ymus-Arteu, Nuv!. Koz!. Beibl., vi, 1907; pp. [1]-[3])

HIPPOCRA TEACEAE (pp. 214-217).

The necessity for a detailed investigation of the anatomical features in
this Order induced F. E. Fritsch to undertake work in this direction. The
following statements are based on his paper as well as on Stenzel's dissertation,
with which I only became familiar after the publication of the main portion
of this book.
The CHARACTERS mentioned (on p: 2I4) as common to the Hippocrateaceae
as the result of a preliminary investigation of Hipppcratea and Salacia have
on the whole been proved to be of general application, and apply also -to the
genus Campylostemon. According' to Frits~h, however, a composite and con-
tinuous ring of sc1erenchyma occurs in the pericycle in certain species of Hippo-
cratea and Salacia, and the development of the cork may take place in deeper
cell-layers of the primary cortex. A new ordinal character mentioned by
Fritsch is the frequent occurrence of small caoutchouc-bodies in the cells of the
mesophyll, these bodies being either soluble or ms?luble in ether. According
to Fritsch, hypoderm occurs also in Salada and Campylostemon, while crystal-
cells are met with in the epidermis of the leaf in Salacia as well. The caout-
chouc-tubes are found in species of Hippocratea and Salada; tannin-sacs are
present in the bast in species of the same two genera. while special tannin-
idioblasts are met with in the mesophyU in species of Hippocratea only. Special
anatomical features occurring in certain species are as follows :~haracteristic
unicellular papillose hairs (Hippocratea aspera, Lam.); stellate hairs with
uniseriate rays (H. velutz'na, Afz.); unisenate hairs, some of which have a
peculiar basal portion with transverse, closely placed septa (ll,. iotricha, Loe~!) ;-
pitting of the outer walls of the upper epidermal cells of the]eaf (Salada oblonga.
Wight); branched or unbranched sclerosed cells belonging to the mesophyll,
or in other cases spicular fibres connected with the sclerenchyma of ·the veins
(species of Salacia); mucilage-cells in the mesophyll (HippoC1'atea t'elutina,
Afz.); cork-warts on the lower side of the leaf (besides occurring in Salacia
attenuata, Peyr., also in.S. obovata, Peyr.).
STRUCTURE OF THE LEAF. The epidermal cells of the leaf exhibit diverse
features as regards their shape when seen in surface-view, their height, the size
of their lumina, and the nature of the inner and outer walls: for information
on these points Fritsch's paper must be referred to .. The absence of gelatiniza-
tion in the epidermis and the scarcity of markings on the cuticle (striation
if.. a few species of Salacia) are noteworthy features. The presence of pits in
the outer walls in S. oblonga and the occurrence of cork·warts have already been
mentioned above. The crystal-cells found in the epidermis 1, and' likewise

, .. 1 Fritsch ~ves the followingsyn?psis of the occur~ence of crystals in the epide~is.:-(I).C.fystals

In on: v epidermal cell:;: (a) Solitary crystals: Ht}pocmtea as}era, Lam., H. bijtndmsu. Laes.,
If. 11n, "llha, Camb., H. obtusifolia, Roxb. var. oarbala, Henth., H. 7!e/utina, Afz.; Salaa'a
J..-ratlssii, Hochst., S. Slaudtialltl, Loes. (b) Clustered crystals: Hippocralea filSrtscens. Kurz.
 ADDENDA-HIPPOCRATEACEAE 88r

referred to in the preceding paragraph, contain solitary or clustered crystals.

They are either ordinary epidermal cells or are distinguished, not only by their
contents, but also betray their differentiation as idioblasts by their shape or
small size, or in .some cases by stronger thickening of the inner walls. In the
genus Hippocratea, in which by far the larger numbers of species:possess these
crystal-cells (exceptions': H. Boieri, Tul., H. ovata, Lam., H. pachnocarpa,
Loes., ,H. scandens, Jacq.), they are very commonly found in groups of two or
more, 'Yhile in Salacia they are mostly isola!_ed. Before leaving this subject
we may mention the occurrence of 'acicular crystals in the epidermal cells' in
Salacia cognata, Peyr. and S. Martiana, Peyr. Hypoderm 1 as a rule is present
only on the upper side of the leaf and is specially widely distributed in the
genus Hippocratea. It consists of one or. two layers; the walls of the hypo-
dermal cells vary in thickness, and the cells themselves may be larger or smaller
than the, epidermal cells. The stomata are generally confined to the lower side
of the leaf (exceptions: Hippocratea celastroides, H. B. K., H. tenuiflora, Mart.,
Campylostemon TVarneckeanum, ,Loes.). As a rule they have no special neigh-
bouring cells, but in exceptional cases, namely in certain species of Hippocratea,
and especially of Salacia, as well as in Campylostemon, subsidiary cells occur.
In most of these cases the pairs of guard 7 cells are surrounded by one or rarely
(Salacia dulcis. Benth.) two rings of four cells of. which two are placed'parallel
to the pore; the latter are occasionally very narrow and are divided by
walls, which are parallel to the pore, Qr even at right angles to' it. The meso-
phyll is generally bifacial in structure; it is subcentric only in a few species ot
Hippocratea and Salacia. Regarding the sclerenchymatous elements found in
the mesophyll the following facts may be mentioned. In their simplest form
they appear as enlarged and pitted_cells of the mesophyll. which do not show
any striking difference in' shape (Salada attenuata, Peyr., S. fluminensis, ~eyr.,
S. laevigata, DC., S. silvestris, Walp.). With these we may associate branched
spicular cells, which. are not connected with the sclerenchyma of the vascular
system (Salada dulcis, Benth., S. lacunosa, Peyr., S; laevigata, DC., S. obovata,
Peyr.), and spicular fibres (S. amygdalin,!, Peyr., S. crassi/olia, Peyr., S. elliptica,
Peyr., S. glomerata, Peyr., S. grandi/lora, Peyr., S. grandi/olia, Peyr., S. pachy-
phylla, Peyr.), which branch off from the sclerenchyma of the veins, traverse
the mesophyll in all directions, and frequently form such a dense plexus beneath
the epidermis that a kind of hypoderm results., The small veins of the leaf
are embedded in the mesophyll. There is usually no development of sc1eren-
chyma in the veins. A hairy covering is rare among the Hippocrateaceae, and
has only been demonstrated iIi a few species of Hippocratea. The extremely
small papillose hairs of H. aspera, Lam. (?) are conical in shape, and have only
a small lumen, which is compressed into'the lower portion of the trichome;

11. G1'alzami, Wight, H. i1ldica, Willd.; Salada dulds, Benth., S. gabunmsis, Loes., S. laevlKala,
DC., S. Roxburghii, Wall., S. lorluosa, Griff., S. vcrrucosa, Wight. ( :I) Crystals in special crystal-
cells: (a) Solitary crystals: Hippocratea (ampcstris, Peyr., 11. alastr(Jid(s, H. B. K., H. jlaaic/a,
Peyr., H .... Griubackii, Loes., H. inundata,. Mart.-, H. tenttiJlora, Mart., H. IVarmingii, Peyr.,
H. Wdwit!Chii, Engl. (according to Fritsch, p. 52, whilst on p. 66 he speaks of clustered crystals) ;
Salada by/uri, Spreng. _(b) Clustered crystals: Hippoeratea arborea, Roxb., H. excelsa, H. B. K.,
H. indi,a, WiUd., H. iotricha, Loes., H. ~btusijolia, Roxb., H. Scnimjel"iana, Hocbst.; Salada
jlavesten.t, Kurz, S. /lon'bunda, W. et A. var. dtnsijlt»'a, Wall., S. g-abunmsis, Loes. It may be
added that Stenzel afso records solitary crystals in the epidermis in Hippocratea jlQribunda, Benth.
and H. vtrrucosa, Griseb.
1 A hypoderm occurs in: Hippocratea arborea, Roxb., II. aspera, Lam., II. Bo/eri, Tul.,
H. celastroides, H. B. K., H. exee/sa, H. B. K., H. flaccida, Peyr., H. Grahami, Wight, H. Grise··
batkii, Loes., H. immdata, Mart., H. obtmifoHa, Roxb. var., 11. ovata, Lam., H. pach1UJcarpa, Loes.,
H. scandens, Jacq., H. velttlina, ACz., II. verrllcosa. Griseb., R.' Warmingii, Peyr., H. We/wi/scl:'-i,
Engl.; according to Pierre also in H. camlJodialla, Pierre, H. Ch~sJialla, Pierre, H. dinhensil, Pierre •
further, Salacia Rtgdiana, F'. Br. et K. Sch., S. Zt)IIud, Spreng.; Camp),losttmon Wanuckeanum,
Loes.
SQLERf.I>EK 3L
882 ADDENDA--HIPPOCRATEACEAE

they are silicified and are the cause of the rough surface of the leaf. 111 addition
to the stella~airs described above, H. velutina, Afz. possesses simple uniseriate
trichomes, corresponding in structure to the rays of the stellate hairs. The
uniseriate hairs found in H !"iotricha, Loes. are characterized by the fact that
their basal portion, which is sunk in the epidermis, is divided by numerous
transverse walls, following ,"'ery closely upon one another; the lower part of
the free body of the hair likewise consists of short cells.
Among the secretory organs the caoutchouc-tubes in particular require
discussion. Fritsch has investigated the details of their distribution and the
exact nature of their contents. They are found both in species of Hippocratea
and Salac.ia, a'ld in Campylostemon lVarneckeanum. In some of the species l
they have been observed only in the axis, while in others 2 they are present
both in the axis and in the leaf. The caoutchouc-tubes of the axis are generally
situated in the soft bast and pericycle; in the latter case they are sometimes
sUlToll..'lded by hard bast. In Salacia Regeliana thev are also found in the
int(Jfxylary phloem. They are rarely (S. obovata and S. serrata) present in the
nrimary cortex. In the leaf the caoutchauc·tubes are found mainly in the soft
bast of the veins or in cannexion with the bast, viz. either in the immediate
nei~hbourhood of the hard bast or between the celis of the latter, often taking
theIr place. In ffiOSt of the species (all those named except Hippocratea pachno-
carpa and Salacia micrantha) the caoutchouc·tubes accompanying the veins
send out branches into the mesophyll. On the grounds of an investigation
of the embryo of Hippocratea ovata Fritsch regards the caoutchouc-tubes of
the Hippocrateaceae as being of the nature of laticiferous cells. They have
_thin walls and a small diameter and are branched, but do not anastomose.
Their-bright, doubly refracting contents are .constituted by threads of caout-
chouc, which produce the phenomenon of I spinning' when the organs con-
taining these caoutchouc-tubes are broken in two; the two ruptured surfaces
are then seen to be connected by delicate elastic threads of caoutchouc, which
are capable of considerable extension. In concluding the discussion of the long
caoutchouc-tubes we may note that rows of shorter cells containing caoutchouc
are found in No. 347 b, as well as in Nos. 519 and 632 of H. Schenck's collection
of stems; according to Fritsch these stems certainly belong to members of the
Hippocrateaceae. 'fbi. occurrence of mucilage-cells in the mesophyll has
already been referred 'to above. According to Fritsch the tannin-sacs first
noticed by me in Salacia grandifolia, Peyr. are widely distributed in the Order;
they are situated in the neighbourhood of the pericyclic hard bast or in the
secondary soft bast; the longest sacs of this kind were observed by Fritsch
in Salacia micrantha, Peyr. Finally we may point out that in the leaves of
certain species of Hippocratea the tannin is localized in certain layers of the
mesophyll or is confined to the spongy tissue (H. Bojeri, Tul., H. indica, Willd.,
H. obtusifolia, Willd. var. barbata, Benth., H. Schimperiana, Hochst., H. War-
mingii. Peyr., H. Welwitschii, Engl.).
The vascular system of the petiole shows varying differentiation. It
consists either of a horseshoe~like group of vascular bundles (e.g. in HiPPo-
cralea ovata, Lam. or Salacia Calypso, DC), or of a ring of bundles enclosing
an inversely orientated medullary plate of wood and bast (e.g. in S. micrantha,
Peyr.), or of a flattened ring of bundles with 3-4 inversely orientated cortical
bundles on its upper side (e.g. in S. crassifolia, Peyr.).

J viz.: IJipjocralea aspera, Lam., II. ob/usi/olia, Roxb., H. TVarmilzgz'i, Peyr.; Salacia at-
letlua/a, Peyr., S.Jluminensz"s. Peyr., S. prineicUs, Jack, S. /orllUJsa, Griff.
~ viz.: fbpPocraiea ovaIa, Lam., H. pachnocarfa, Loes.; .Campy/oslenton IVarneckeanWlI ,
Laes.; Salacia anema/a, Peyr., S. Calypso, DC., S. Krauss;;, Hochst., S. macrocarpa, \Ve1w.,
S. micranlha, Peyr.) S. oOovala, Peyr., S. Regeliana, F. Br. et K. Sch., S. Jerrata, Camb.
 ADDENDA-HIPPOCRATEACEAE

STRUCTURE OF THE AXIS. With reference to the stru_cture of the wood

the folloWing information may be added. The vessels Jie isolattd or in pairs
in a transverse section of the branch; their diameter may be as much as
011-·r6 mIn. in thick stems. Wood·parenchyma is as a rule found only in the
neighbourhood of the vessels.
According to Fritsch and Stenzel the cork occasionally develops in the
second layer of cells of the primary cortex and according to Stenzel even in
y

the third or fourth layers. The cells of the cork have rather wide lumina;
their walls are either thin or exhibit uI)iform or ttllilateral thickening, the latter
affecting the inner or rarely the outer tangential walls. According to Fritsch,
the pericycle in Campylostemon, and in some of the species of Hippocratea and
Salacia, contains isolated groups of bast-fibres, which are more or less approxi-
mated to one another; in the remaining species of Hippocratea and Salacia
there is a composite and continuous ring of sclerenchyma. The primary cortex
frequently includes branched stone~cells, especially in its inner ·portion. In
many species the secondary cortex contains hard bast-fibres; and branched
sclerenchymatous cells also OCCUr9 e.g. in Hippocratea pachnccarpa, Loes~- or
Salacia obovata, Peyr. A specially noteworthy feature is foUnd in the :{>enetra-
tion of sclerosed tissue belonging to the medullary rays of the cortex mto the
medullary rays of the wood in Hippocratea Bojeri, Tul.; hand in hand with
this goes a sinuation of the edge of the _cambium, the convexities of which are
directed towards the pith.
The pith in some cases (Hippocratea pachnocarpa, Loes., Salada micrantha,
Peyr.) contains s,tone·cells.
Literature: Stenzel, Anat. d. Laubbl. u. SUimme d. Celastraceae u. Hippocratcaceae, Diss.
Erlan~nJ J8937~' especially pp. 8.+-8.-Pierre" FlQre (orest. de la Cochinchine, xix, 1893~­
LeiseflOg, Interxylares Leptom, Diss., Berlin, 1899. p. II.-F. E. Fritsch, Vork. v. Kautschuk bei
den Hippoaateaceen, verb. mit einer anat.~sysL Unters. etc., Diss., Munchen, 1901, 80 pp., I Tab.
(sep. copy from Beih. z. bot. Centralbl., xi).-Haberla~dt, Sinnesorga.ne, 1901, p. 13 •. -Pitard,
Pericycle, These, Bordeaux, -J901. p. 39.-F. E:Fritsch, Caoutchouc ill plants, New Phytologist,
ii, 1903, pp. 25-30.-ArtSch9'Jg, Trop. v3.xt. bladbyggo" Sv. Vr.t. Akad. Handl., 3~, ,no 2, 1905,
pp. 91,92 (Salada). / -

PENTAP_HYLACACEAE.
The monotypic-genus Pentaphylax (with P. euryoides, Gardn. et Champ.),
which is referred to the Temstroemiaceae by Bentham and Hooker,_has recently
been regarded as the type of an independen t Order by Engler and Van Tieghem ;
according to the former it has affinities with the Coriarieae, while,Yan Tieghem
places it near the Celastrineae. Pentaphylax exhibits the following anatomical
features :-simple, unicellular clothing hairs'; mucilage-cells in the primary
cortex; deposition of oxalate of lime in the '~orm of solitary and.:clustered
crystals; a composite and continuous ring of sclerenchyma in the: peri cycle ;
and subepidermal cork-development. ."
The follQwing'facts may be mentioned regarding the STRUCTUR~ OF TIlE
LEAF., Th-eleaves are bifacial and have a gelatinized epidermis. The mesopbyll
contains solitary crystals. Stomata are found only on the lower side of the leaf.
A single large arc-shaped vascular bundle passes ou~ into the leaf. The vascular
bundles of ,the veins are provided with a sheath of sc1erenchyma. - ,
STRUCTURE OF THE AXIS. The cork consists of cells with thin walls:""
The primary cortex contains' relatively large cells, which have thin walls and
hyaline mucilaginous contents, and are either isolated or' combined to form
groups. The endoderm is is not distinctly differentiated, but each of its cells
contains a solitary crystal. In young' branches the pericycle is formed by
a ring of fibres, which subsequently becomes replaced by a composite and con-
tinuous ring. of scIerenchyma. There is no secondary hard bast. Oxalate of
3 L2
 ADDENDA-PENTAPHYLACACEAE

lim,u)ccurs in the form of solitary crystals in the soft hast and in the 'form of
clustered crystals in the lignified pith and in the medullary rays of the bast. In
thicker branches sclerosed cells are also present in the primary cortex and in
the medullary rays of the bast.
Literature; Van Tiegbem, Pentaphylace et Corynocarpe, Jouro. de bot., 1900, pp. 188-93.

CORYNOCARPACEAE.
This Order, which is referred to the Sapinctales by Engler, 'and is placed
near the Geraniales by Van "Tieghem, consists only of the genus Corynocarpus
with C. laevigatus, Forst. and C.. similis, Hems!.; the anatomy of the first of
these species has been investigated. The special anatomical characters of
this species are: absence of secretory receptacles; stomata of the,Rubiaceous
type; vessels with simple perforations; wood~prosenchyma bearing simple
pits ;, subepidermal development of the cork; isolated groups of bast-fibres
in the pericycle; clustered 'and solitary crystals of oxalate of lime.
STRUCTURE OF THE LEAF. The leaf is bifacial in structure. The mesophyll
contains clustered crystals. A hypoderm composed of one or two layers is
found on the upper side _of the leaf, while the lowest layer at the spongy tissue
is,differentiated so as to resemble a hypoderm. The stomata are confined to
the ~ower epidermis and . are bordered by two subsidiary cells placed parallel
to the pore. Three vascular bundles pass out into the leaf ,; of these the median
one divides into·fi.ve bundles, whichjorm an arc open on its upper ,side.
STRUCTURE OF THE AxIS. The cork consists of cells with thin walls and
develops in the second cell~layer of the primary cortex. Phelloderm is present
and, like the primary cortex, contains clustered crystals and in later stages
-sclerosed cells 'as well. The endodermis is not ,distinctly, differentiated. At
the inner ends of the vascular 'bundles, 'which are separated by broad medullary
rays, there are strands of fibres analogous to the groups of pericyclic bast-fibre:i
and corresponding in position to them. The medullary rays of the bast 'Contain
clustered crystals, while those"of the wood have large solitary crystals. There
is no secondary hard bast. The main mass of the wood·is composed of fibres,
which may. either have thick, or thin walls. The pith, the peripheral portion of
which belA)mes lignified in.later stages, also contains clustered crystals ..
Literattu'e: Engler, Corynocarpaceae, in . Nachtr. z. ii-iv. Teil der NatUrl. Pflanzenfam., 1897,
p. ·u6.-V41l"Tieghem. Pentaphylace et Corynocarpe, Joum. de hot., 1900, pp. 193-7.-Hemlley,
CfryttQ(aYpus, Ann. of bot., xvii, 1903, pp. 748-5 T. and xviii, 190"'" pp. 179. 180.

STACKHOUSIEAE (pp. 217,218).

_ Ule anatomy of the two genera Stackhousia and Macgregoria has recently
been subjected to a careful examination by Pampanini and Bargagli-Petrucci 1
in the course of a monographic revision o( the Order. .
The following anatomical features constitute important DIAGNOSTIC
CHARACTERS OF .THE ORDER :-the absence of a special tyPe ,of stoma and of
external glands; the occurrence of unicellular clothing .hairs and of tannin-
cells, which in some cases are differentiated as idioblasts; the simple perfora-
tion.:) in the-'vessels; the fact that part at least of the wood-prosenchyma is
provided with -bordered pits; the absence of -medullary rays in the wood;
the almost universal occurrence of bundles of fibres in the outer part of the
primary cortex; and lastly. the absence of oxalate of lime.

1 Rull. de l'Herbier BoiS3ier, '90S, pp. 1I~6-60i and 1906, pp. 39-H, :load tn-b. xi,-'''.
 ADDENDA-STACKHOUSIEAE

Dealing first with the stem, we may note that the above·men tioned bundles
of fibres correspond with the ribs on the surface of the stem; the bundles
vary in the extent of their development, and are separated from the epidermis
by a hypodermal layer of cells; the fibres themselves have a polygonal outline
in transverse section, and are for the most part strongly thickened and abun-
dantly pitted. Such strands of fibres have been recorded in all the species of
Stackhousia except S, pulvinaris, as well as in Macgregoria. The primary
cortex shows varying differentiation, and contains palisade-tissue which is
most prominently developed in Stackhousia aspericocca var. incrassata and
S. Dielsi. The endodermis is occasionally composed ot large cells. In the
ll)ajority of the.species of Siackhousia the pericycle contains isolated bundles
bf fibres; in Macgregoria there are likewise small groups of fibres. Pericyclic
sclerenchyma is wanting only in Stackhousia Maideni and S. pulvinal'is; in
these species, however, there is a suberized endodermis instead. According to
Pampanini and BargagIi- Petrucci, the xylem in both genera consists of (a) vessels
with simple perforations, (b) a small amount of wood .parenchyma, and (c) wood·
prosenchyma bearing bordered pits, medullary rays being absent~ The pith
IS composed of large cells with thin walls.
The structure of the leaf is either bifacial or centric; the stomata are found
on both sides. The vascular bundles of the veins are not provided with scleyen-
chyma. In StackJeousia Brunonis and S, pulvillaris large tannin·cells occur in
the primary cortex, and in S. Bru,nonis they are present in the mesophyll as
well; in other species of Stackhousia. (e.g. S. Hztgelii) and in Macgregoria
1'ae-emigeYa the tannin-cells found in the primary cort.ex scarcely differ from
the cells of the surrounding tissue. The caoutchouc· bodies, ohserved by the
two authors aboye named in the ~ortical parenchyma and the tissue of the
leaf, are no doubt of the nature of fat·bodies.

RHAMNEAE '(pp. 218-22:1).

The structure of the leaf k now well known owmg to the recent r~earches
of Gemon" and Herzog 1. ·No new anatomical features common to all the
members of the Order have come to light, but for the purposes of special diagnosis
numerous characters have been recognized. The enumeration af spt::cial
features may be supplemented by the following statements. Secretory cavities,
provided with a papillose epithelium, besides occun°ing in Kaminpkia c;re
found also in Rhamnidium and Reynosia reticulata, Urb. Secretorv c.a~
with resinous contents and an epithelium re~embling that of the secretory
cavities in Karwinskia are present in the veins of the leaf in ReY1losia revoluta,
Urb. The palisade-tissue in some cases contains isolated enlarged ceIG filled,
with mucilage (species of COltdalia) or tannin (species of Condali:z, Phyllogeiton,
and Scutia j also in species of ..'Maesopsis, Pomade1"is and Rham·neUa, the cells
in question being more or less sclerosed in these three genera) or both IDlidlage
and tannin (species of Discaria and Talgutnea). In ZizyPhus ceJJidildlins, DC.
(incl. Z. timoriensis, DC.) a middle iayer of the leaf consists of large mncilage-

1 These inve!ltigations deal with the following genera, which are enumerated in the suid order
ofWeberbauer's sy.stem. (in Engler and Prantl, N atiirl. P lIanzen(am" ii i, 5) :-~ Ventilaginea~: Ventilago,
Smytltea; Zizypbene: Paliurus J ZizJ'}nus J CgrJalia J J.lfi,rorha,mlus, l{rugi4Jemiroll, Reynosia,
Sat tOfll/lta!us, Rlza11miJiu1Il, Aa,.-u,,'nskia, Berciumia, Ph,1I11ogdton (regarded as a section of Ber-
clumia in \Veberbauer's system), lIfaesopsis, La1Jltllisepa/ulIZ, RluZ1'lnella,-Da!lllChya ~ Rhamneae:
SagN"etia, S{,lltia, RJi(1lJlnu.r, Hot'tnia, Ceano!nus. Emmenosprmum. Noltea, Cl)lttorina. ConlUJntnlO,
PJeylica, L.:zsitJdist:U!, Alpkitam·a, Pomadcrris, Try'lIla/junt. 5..1t)II'idium. CrypaJUiya; Colleti~ae:
Talguenta, TrevM, Discaria, C(J/lttia; GOllllnieae: Ccu(lItia, Reisu1u'ol, Helinu!, Crummaria,
Ma"/othia. ,~
886 ADDENDA--RHAMNEAE
cells, while in Micr~hamnus the upper layer of the hypoderm is gelatinized.
Hypodenn and development of papillae are rare features in the ·Rhamneae.
The spicular cells which Blenk mentions correspond to the sclerosed tannini-
ferons idioblasts in the palisade· tissue (see above). True glandular hairs are
not met with on the surface of the leaf. The following forms of c1Qthin~ hairs
are worthy of special note :-trichomes, in which the body of the hair IS uni-
cellular and two.anned, and is seated on an eJ?idennal cell (Sagerelia); stellate
hairs with a specially differentiated basal portion and unicellular rays (in Poma-
derris, &c.); and tufted hairs, the rays of which are mostly uniseriate· and are
inserted directly in the epidermis (in a group of dosely allied species of Rhamnus).
Extra·ftoral nectaries (?) are found on the lower sIde of the leaf in Rhamn~
gla'ndulosa, Ait.
STRUCTURE OF THE LEAF. The stomata are for the most part confined to
the lower side of the leaf, but are found on both surfaces in certain species of
CoUeti~, Condalia., Cortnonema, Cr"menaN«, Discaria, Gouania, Marlothia,
Reissekia, Rhamnus, Scmia, Trevoa and Zizyph'us. Gemoll records stomata
of the Cruciferous type in Cryptandra obovata, Sieb. and stomata 'of the Rubia·
coous typ_e in Colletia spinosa: Her~og mentions .the occurrence of strikin~ly
small and crowded stomata ill speCIes of Reynosla and Sarcomphalus, whilst
he describes very large stomata in species of Condalia, Disca,ia, Emmeno-
spermum and Rhamnus. Taking recent investigations into consideration,
gelatinization of the epidermis'of the leaf has been observed in species of ' the
following genera: Alphitonia, Berchemia, Ceanothus, Colubrina, Condalia:Sect.
CondaJiqf>sis, Cormo~, Crume_naria, Cryptandra, Dalla~hy.a, Discana, E,,!-
menospermum, Gouanta, H01Jenla, La:mell1sepidum, Laswd'SoSCUS, M a~s"s.
Paliurus, Pomadmis, Reissekia, Reynosia, Rhamnella, Rhamnidium, Rhamnus
Sect. Frangula, Sageretia, Scutia Sect. I, Spyridium, Trevoa, Trymalium,
Vmtila~o, ZizYPhus ' .. The ~roces~r of gelatinization affects either all Of' only
a certam number of the epIdermal cells. The remaining characters of the
epidermal cells (viz. height, cubical dimensions, nature of the lateral margins,
thickenin~ of the outer walls and pitting of the lateral walls) are subject to
very considerable variation. The following features are specially noteworthy : -
papillose protrusion of the epidermal cells (in Illost cases only on the lower side
of the le~f) in species of Berchemia, Discariat HeUnus, Karwinskia and Mar-
lothia; development of typical papillae on the upper side of the, leaf in Cryp-
tandra obovaia, Sieb., and on the lower side in J(arwinskiaHumboldtiana, Zucc.;
a two-layered epidermis in Rhamnus Wightii, W. et A.; the occurrence of
paired crystal-cclls containing solitary crystals in the epidermis in Vemilago
leiocarpa~ Benth. Hypoderm (situated beneath the upper epidermis) is not
of frequent occurrence i it is present in MicTorhamnus e1'f,coides, A. Gray'(bere
the upper layer of the hvooderm consists of II1ucilage-cells). Reynosia revoluta.
Urb., R. septent,;onalis, l1rb., Rhamnus Alaternus, L. (in this species the hYJ>O':
derm is confined to the margin of the leaf and is' developed in the fonn of
sclerosed mechanical tissue), Sarcomphalus crenalus, Urb., S. domingensis,
Krug et Urh., S. laurinus, Griseb., S. rtticulatus, Urh., Ceanothus c,assifolius,
Torr. and other species of the subgenus C"astes (here tanniniferous). In
most of the species the mesophyll is bifacial, but centric or subcentric leaves
are also found. M iaorhamnus ericoides, Gray has rolled leaves which are
provided with a ftirr'ow on each side (to 'the right and left) of the median vein,
and show centric structure; regarding the occurrence of rolled leaves in Phylica,
see Knoblauch, loco cit. In the species of Ceatwlhus ~longing to the subgenus

1 In the previous (Syst. Anat., p. 318) enumeration of genera exhibiting this feature. the genas
Mi'rw-1l4ffC""f should be cancelled; the ~latintu.ti()n of the ~ptdenni' of the leaf in Hqt1fni~
(n. dulci's, Thunb.) bas only been lorrectly recorded by Blenk.
 ADDENDA-RHAMNEAE

Cerastes the leaf also presents a characteristic appearance in transverse section.

The lower surface of the leaf is provided with deep and narrow furrows, formed
by the projecting network of the veins; the epidermis which lines these furrows
includes the stomata.and bears numerous hairs. The mesophyll is composed
only of palisade tissue, which exhibits a larger number of layers at the bottom
of the furrows and at the top of the ridges (formed by the projecting portions
of the leaf-tissue) than along the lateral margins of the furrows. At the same
time the vascular bundles of the veins are accompanied by large parenchymatous
cells which contain tannin and merge into the hypoderm, situated on the upper
side of the leaf (see above). In certain species (see Herzog, Sep. Copy, p. 113)
-the palisade-tissue consists of exceptionally long or short cells. In Reynosia
and Sarcomphalus the spongy tissue is composed of elongated cells, which are
interwoven with one another in such a way as almost to resemble a hyphal
weft. Special features of the mesophyll are constituted by the above-mentioned
enlarged palisade-cells, which are filled with mucilage (Condalia lineata, Gray,
C. mexicana, Schlecht.) or tannin (Candalia lineata, C. tnexicana, Phyllogeiton
discolor, Herzog, Scutia Commersonii, Brongn., Maesopsis berchemioides, Engl.,
M. Eminii, Engl., Rhamnella fra ngulo ides, Weberbauer, PomadeN'is discolor,
Vent., P./erruginea, Sieb., P. lanigera, Sims and P. phylliraeoides Sieb.) or both
mucilage and tannin (Discaria discolor, Reiche, D. serrati/olia, Benth. et Hook.,
D. trinmns, Reiche. Talguenea costata, Miers). Cryptandra obovata, Sieb. has
a single layer of palisade-tissue, which is entirely composed of broad tannini·
ferous cells, while in Zizypkus celtidi/olius, L. there is a middle layer consisting
of mucilage-cells. In Herzog's summary the following structural characters
of the veins are stated to be of value for systematic purposes. Vertical trans-
currence of the lateral veins of the second order is found in Alphitonia, Ceano·
tAus, Colubrina, Cormonema, Crumenaria decumbens, Mart. (not in C. cJwrtroides,
Mart.), Helinus (with the exception of H. brevipes, Radlk.), Hovenia, Karwinskia,
Pomaderris, Reynosia, _Rhaff'.nidium, Sageretia, Sarcomphalus, species of Zizy-
phus; embedded veiz;ls occur in the Ventilagineae, Colletieae, Gouanieae (with
few exceptions), and also in Conda-lia, Emmenospermum, Krugiodendron, Lasio-
discus, Rhamnus, Scutia, Trymalium, species of Z.izyphus. There is a well-
developed sheath of sclerenchyma around the lateral veins of the first order in
the Ventilagineae t and also in species of Berchemia, Emmenospermum, speci~
of Gouania, Lamellisepalum, Lasiodiscus, species of Phylica, Reynosia, Sageretia,
Sarcomphalust species of Scutia and oCZizyphus; the lateral veins of the
second order-are provided with a sheath of large parench~atous cells, which
are rich in tannin, ,in Condalia, Krugiodendron, Microrhamnus, species"'of
Rhamnus (especially those of the subsection Cervispina) and species of Zizyphus.
The statements made in the earlier part of this work on the distribution of
oxalate of lime may be extended in the following directions. Large solitary
crystals occur in enlarged cells of the palisade· tissue in Karwinskia, Reynosia
N rwthropiana, Urb., Rhamnidium, Rhamnus Sect. Leptophyllius Subsect. Cervi-
spina, Scutia Sect. II, and Zi:yphus gla'bratus, Heyne 1. Large clustered crystals
are found in enlarged cells of the mesophyU or palisade-tissue, as the case m~
bet in Colubrina, Condalia, Cryptandra, Hovenia, Krugiodendron, Lasiodiscus:
Pomaderris, Rhamnus Sect. Alaternus and Sect. Leptophyllius Subsect. Espina;
Scutia Sect. I, T,.evoa~ T,ymalium, and species of Zizyphus. Large solitary
and clustered crystals occur in a small group of closely allied species of Rhamnus
(R. costata, Maxim., R. nipalensis, Laws., R. Wightii, \V. et A.). The genus

I The earlier statement (p. 219) as to the OCCllrrence of large solitary crystals in C,anol/zus must
be cancelled; it was based on Blenk', investigation of. plant described as • Cea1U)lnus tn4(rDJnylla.
WaU.,' which does not belong to the Rbamnea.e. According to GemoU, solitary crystals are
wanting in Ceanolhiu.
888 ADDENDA--RHAMNEAE
Relinus (a member of the Gouanieae) as well as M arlothia have elustered crystals
only and no styloids.
Among the internal secretory receptacles, the tannin- and mucilage-cells
have already been dealt with in detail in describing the mesophyll. Secretory
cavities are not only present in Karwinskia (three species), but also in Rham-
nidium (three species) and Reynosia reticulata, Urb. They have an epithelium
composed of several layers of flat cells, the innermost of which are drawn out
into long finger.shaped protrusions terminating freely in the secretory space;
the contents are brown and do not dissolve easily. The secretory cavities are
visible even to the naked eye as brown glandular dots. In Reynosia revoluta,
Urb. the cavities are replaced by secretory canals having a similar epithelium
and resinous contents, which are soluble in alcohol; these canals run beneath
"the, vascular system in the larger veins. Gemoll and Herzog have investigated
the detailed distribution of the mucilage-receptacles (described as canals)
found in the veins of the leaf. They have been recorded in the following
genera: Alphitonia, Berchemia, Ceanothus pro parte (mucilage-cells ?), Colu-
brina, Cormonema, Condalia pro par,te, Dallachya, Emmenospermum, Gouania,
Hovenia, Karwinskia, Lasiodiscus, Maesopsis, Paliurus, Phyllogeiton, Rham-
nella, Rhamnidium; Rhamnus Subgen. Frangula, Sageretia, Scutia Sect. I,
Ventilago pro parte, Zizyphus (in almost all cases). The mucilage-receptacles
are fo~ the most part restricted to the principal veins and to the lateral veins
of the ,first order. Their diameter varies, and an epithelium mayor may
not be present. They are situated in the collenchyrn~tous tissue of the veins,
and are either isolated or several of them lie side by side; in M aesopsis they
were met with in the phloem as well. Gres's paper also contains new statements
on the distribution of these ,mucilage-receptacles, especially in the species of
Rhamnus 1. The same authority figures large mucilage-receptacles, occasionally
containing clustered crystals, in different parts of the axis of certain species
of Rhamnus (here situated in the inner portion of the primary cortex, In the
pericycle, the pith and the secondary bast).
The following additional facts have become known regarding the hairy
covering 2. The glandular hairs mentioned by Gemoll as occurring on and near
the margin of the leaf in Ceanothuspapillosus, Torr .. et Gray, correspond to the
.glandular leaf-teeth found in other members of the Order. They are glandular
shaggy hairs with a multiseriate stalk containing the termination of a vascular
bundle, and a spherical head provided with a secretory palisade epidermis.
Special forms of clothing hairs are represented by the short papillose trichomes
occurring in many species of Cryptandra, and the short peg-like hairs of Zizyphus
/uniculosa, Ham.; the latter recall the 'characteristic trichomes found in the
Sapindaceous genera Pancovia arid Xerospermum, and have a swollen basal
portion, which appears striated owing to the presence of slit-shaped pits.
Unicellular two·armed trichomes, which are seated on epidermal cells, occur
in Sageretia and have already been noticed above. Mention has likewise been
rna ~e of the stellate hairs found in the Pomaderreae; these hairs. have a stalk
of varying length, and exhibit a variable number of ray-cells. Tufted hairs
with 2~8 rays are found in the fonowing species of Rhamnus: R. caii/ornica,
Eschsch., R. Palmeri, Wats., R. sedipetala, Mart. and R. sphaerosperma, Sw. ;
in R. cali/ornica the rays are unicellular, while in the other species they are

1 Gres's statements do not ~uite tally with those of Herzog. Thus, according to Gres, mucilage-
rece~tacles occur in the leaf of Rhanmus·cothartico, L., R. inftctoria, L. and R. ti'w:loria, W., as
well as in the axis of R. libalU>tica, Roiss. and R. Sibthorpiana. i. e. they are present in some of the
species of the subgenus EztrhammlS, whereas they are stated to be absent in R. lalifoHa. L'Herit.
(Snbgen. FraJlgula).
~ Swanlund's statement that' hAir1l terminating in a spherical end·cell' are present in Phylka
n£tida, Lam. requires verification.
 ADDENDA-RHAMNEAE

unisenate. The'~mall pits, observed by Clauditz in the axils of the lateral

veins on the lowe!" side of the leaves of Rhamnus glandulosa, Ait., are probably
extra-floral nectaries; they are provided with a palisade epidermis and contain
trichomes.
. For the structure of the stipular spines of Paliurus see Lothelier; regarding
the anatomy of the spiny branches of Colletia and Rhamnus, see Lothelier and Mitt-
mann. The localization of Emodin and Frangulin in the cortex of Rhamnus is
dealt with by Gres.
To the section dealing with the structure of the cortex we may add that the
pericyc1e contains only isolated groups of bast-fibres in the species of A/pMtom'a,
Berchemia, Ceanothus, Colletia, Colubrina, CondaJia, Emmenosperfflum, Hovenia,
Paliu1'us, Rhamnus, Scutia, Ventilago, examined by Petit, as well as in Zizyphus
sativa.
Literature; Borscow, Frangulin, Bot. Zeit., I87'h p. 33.-HohneJ, Gerberinden, Berlin, 1880,
p. 1I7.-Mittm:um, Ptlan,zenstacheln, Verh. bot. Ver. Brandenburg, 1889. pp. 48-So.-[Pirotta, in
Malplghia, iii, 1889, p. 61 et seq.; abstr. in Just, 1889, i, p. 606.]-Barber, Corky excresc., Ann. of
Bot., vi, 1892, p. 165.-Lothelier, Epines, These, raris, 1893, pp. 14 and 31.-Cabannts, Rhamnw,
These, Montpellier. 1896, 72 pp.-Knoblauch, Okolog. Anat., Habilitat.•Scbr., Tiibingen, 1896,
p. I I et seq.-[Planchon, Cascara Sagrada, Bull. d. Ph. d. S. E., i, 1896, n ... j abstr. in Just, 1897,
ii, p. 33. ]-[Sayre, Frangula and Cascara bark, Americ. Journ. of Pharm., 1897, n. 3; abstr. in
a
Just, J897, ii, p. 32.]-Gres, Contribut. retude anat. et microchim. des Rhamnees, These, Paris,
1901,10" pp., ~ pt-Petersen, Vedanatomi, 1901, p. 50.-Pitard, Pericycle, These, Bordeaux,- 1901,
pp. 39 and 90, 91.-.swanlnnd, Vegetat. Neuamsterdams u. St. Pauls, Diss., Basel, 19 01 , pp. 19-25
(Pllylica nilitk).-Clauditz, Blattanat. canar. Gew., Diss., Basel, 1903, pp. 31-3 (Rhamnus glandu·
losa).-Gemoll, Anat.-syst. Vnters. d. Bl. d. Rhamn. aus den Triben Rhamneen, Colletieen u.
Gouanieen, Diss., MuncheD, jn Beih. z. bot. CentralbJ., xii, 1902, pp. 351-424.-[Finlayson, Stem-
structure of some leafless plants, Transact. and Proceed. New Zealand Institute, 1903, p. 360 et seq.
(Discaria Toumatcu).]-Herzog, Anat.-syst. Vnters. d. Bl. der Rhamneen aus den Triben Ventila-
gineen, Zizypheen n. Rhamneen, Diss., Munchen, in Beih. z. bot. Centralbl., xv, 1903, pp. 9:;-::107.-,
Siissenguth, Behaarungsverh. d. Wiirzb., Muschelkalkptl., Diss., Wiirzbnrg, 1904, p. 25.-Theorlo,
Vmtrichom., Arkiv fOr Bot., iii, n. 5, 1904, p ....-Areschong, Trop. vaxt. bladbyggn., Sv. Vet.
Akad. Handl., 39, n. 2, 1_905, pp'. 58, 59 (Colubrina).-[Mitlacher, Verf. v. Cortex Frangulae,
Zeitscbr. osterreich. Apoth •. Ver., 1906, pp. 4-7 j abstr. in Bot. CeDtralbl., ci, p. 46l.]-Piccioli,
Legnami, Blll]' Sienat J906, pp~ J$1. 157-9, and 161.-[For additional Iiterature, see p. 1172.]

A~fPELIDACEAE (pp. 221-226).

1 ...~

1. !(EVIEW OF THE ANATOMICAL FEATURES. According to Gard, the

first cork'may in some cases develop in t~e bast and not in the pericycle. In
the pith of Tetrastigma Kalberlah met with secretory canals with mucilaginous
-contents side by side with groups of mucilage-cells.
2. STRUCTURE OF THE LEAF. According to Gard's recent investigations
the leaves in the species of Vitis are mostly bifacial in structure, the palisade-
tissue consisting of one or two layers; in rare cases the leaves exhibit a ten-
·dency towards centric structure. In V. Labrusca, L. the lower epidermis is
subpapillose. The petiole in the species of Vitis contains isolated vascular
bundles which are arranged to form a ring; in addition to the main system
there are ctwo cortical bundles on either side of the petiolar groove, and in rare
cases flirther smaller bundles occur in the same position.
The following statements may be added regarding the features presented
by the crystals. According to Gard, the shape of the individual raphides in
Cissus, A mpelopsis and V itis is characteristic of these genera, one end of each
needle being pointed, while the other is bidentate, as in the gypsum-crystals
found at Montmartre. According to the same authority, the raphides in the
leaves (especially in the veins) of the species of Euvitis, Planch. are accompanied
by clustered crystals, while in V. 1'otundifolia, Michx. and V. Munsonia1la,
SlDlS. (V. Muscad£nia, Planch.) they occur side by side with prisms of oxalate
()f lime.
&)0 ADDENDA-AMPELIDACEAE
i
3. STRUCTUlffi OF THE AXIS. According to Gard, the wood-fibres, and
the fibrous cells in the bast and pericycle of the t Vites verae,' are invariably
septate by means of tbin division-walls. ..
The cork develops subepidermally also in Leea (Hallier). Regarding the
formation of the cork in the bast and the detailed differentiation of the cork-
cells (with U-shaped or unilateral thickening) in Vitis, see Gard, n. ce.
According to Gard, the arrangement of the hard bast in the species of
Vitis belonging to the sections EuviJis and Muscadinia is different in the two
cases and characteristic of each section. In Euvitis the phloem is stratified
into hard and soft bast. In MUscadinia, on the other hand, the bast-fibres
form more or less regular radial rows alongsid~ of the medullary rays, and
exhibit a scattered arrangement in the inner portion of each bast-group.
literature; [Mitlardet, Histo des prine. var. et especes de "ignes d'origine aIJleric. etc., 1877-85 i
referred to by Gard, lac. cit.]-Ka!;sner, Mark, Diss., Brt'Slau, 1884, p. lI.-Keller, Luftwurzeln
einiger DikotyI., Diss., Heidelberg, 1889, pp. 23--6.-[Tognini, Stomi, Atti 1st. bot. Pavia, 1894.]-
llalUer, in Natuurkundig Tijdschr. voor Ned~-Indie, lvi, 1896, pp. 308, ,309·-Gnuchery, Notes
anat. sur qu. "ignes hybrides, Assoc. frans:. potlr l'avance:rn~nt d. $C., Congres' Nantes,. 1898, ii,
pp. 417-33.-Kalbedah, Ban von Tetrasfi'gma scariosufn, Zeitschr. f. NatuJ"W'., lxxi, 1898, pp. 161~
2I8.-Gard, in Actes Soc. Linn. de Bordeaux, lv, 1900, pp. c)(.vii, ccii and ccvii; and lvi, 1901. pp. x,
lxvii, lu, cnvii, and cxxx.-[Lopriore, Anat. iii alcune Ampelld., .Boll. Accad. GioeD. Sc. nat.
Catania, 1901,16 pp.; abstr. in Just, 1902) ii, p. 280.]-Molisc:b, PRa.nzen als Trin~quelleD, Deutsche
Arbeit, i, 1901, p. 78 et seq.-[Ravaz et Bonnet, Bois.; de la, 'figne, Ann. de l'Ecole' nllt. 'agr. de
Montpellier, DOllV. ser., i, 1901; abstr. in Bot. Centralbl., 1902, p. 4SI.}--Pollisen, Lnftllfdderne
bos Cissus sicyoJ(ks, Vidensk. Meddelels. KjlJoonhavn, 1!102, pp. ;138, 239.-Gard, ltt. fUlat. 'sur
Ies vignes, etc., These, Bordeaux, 1903. J34PP.; also in Actes Soc. Linn. de Bordeaux, lviii, 1903.-
Tondera, Innerer Ball d. Sprosses von Pitis vim/era, BUll. internat. Acad. s.c. Cracovie, 1904.
pp. 91-6, :I pI.; see also Abh. matb.-nllturw. Kl. Ahd. Krahn, ser. 3, Bd. iv, B, 1904, pp. 'U-54
(Polish).-Frommel, Plant. text. chil., 190,5, p. ,p.-Netolitzky, Dikotylenbl. (Rhapbiden), 1905,
pp. 3S-8.-Fries, Zwei siidamerikan. Lianeu, Botaniska Studier tillagn. Kjellman, 1906, p. 89 et seq.
-Picaoli, Legnami. Bull. Siena, 1906, p. 133.-[For additi onalliteratllre, see p. 1169,]

SAPINDACEAE (pp. 226-236).

ApPENDIX: On the Didiereae.
We may append to the Sapindaceae a discussion of the genus Didiefea (which
was formerly regarded as a member of this Order by Baillon), and of the allied genus
Alluaudia, both of which are distinguished by a Cactus-like habit. In the first
place. however, it may be pointed out that these genera must be removed from the
Sapindaceae; Radlkofer bolds that Didiefea is allied to the taxonomic group
~nstituted by the Amarantaceae and POlygonaceae. Anatomically the two
genera are characterized by the possession of mucilage-receptacles Or mucilage-cells.
the subepidermal development of the cork, and the occurrence of isolated bundles
of pericyclic fibres.
Our knowledge of the anatomy of Didierea is as follows. The 'primary cortex
.contains spaces which are' almost as big as a grain of millet and are :filled with
mucilage; near the surface there is a continuous layer of stone-cells, some of which
are of very considerable dimensions, while on the outer side of this layer are situated
cells containing large clustered crystals of oxalate of lime; finally, the great~ part
of the primary cortex consists of cells with brown tanniniferous contents, which
in the living plant are possibly of the nature of latex. Cells with similar brown
contents are also present in the pith. The bast-fibres form circular grou:ps of vary-
ing size. The wood possesses broad medullary rays, wood-fibres beanng simple
pits, and vessels with simple perforations. A traD:iverse section through one,of the
linear leaves shows a homogeneous mesophyll containing tannin- and mucilage-
cells~ and 7-9 isolated vascular bundles, which are arranged in the form of an almost
closed arc; 'the stomata are not present in large numbers and are only s1tghtly
depressed. ,
As regards the structure of the axis in A lJuaudt'a, we rnay add that in A. procera
the primary cortex contains mucilage- and tannin-cells, while in A. ascendens the
 APpENDA- SAPINDACEAE

mucilage-cells are -not is<?lated, but fuse to form mucilage-lacunae. The, epidermis
OD both sides of the leaf of A ltua~d~'a has a thick cuticle; beneath each epidermis
there is a one-layered hypoderm, which at certain points, and especially in the
neighbourhood of the stomata, shows an iilcrease in the number of -layers. On
the upper side of the leaf the mesophyU consists of indistinct palisade-tissue. The
stomata, which are ,found on the lower surface, are not numerous and are strongly
depressed. A feature worthy of special note is the occurrence of clustereq crystals
of oxalate of lime in the ,epidermis of the leaf as well as in that of the stem. ... ~
-
Literatnre; R. Schenck, Qumae&i cipO, Diss., Erlangen, t89..., 19 pp.-Radlkofer, in Natiirl.
Pftanzenfam., m.Tei}, Abt~ 5, 1896, p. "'6:l.~hwabacb,.in Bot. CentralbI., 1898, iv, PP·'357-9.-
[Radlkofer, Sap~daceae, in Martins, Flora brasil., xiii, 3, 1892-1900.]-Haberlandt, Sinnesor~e.
1901. p. Il6 et seq.-Pitard, Pericyc1e, These, Bordeaux, 1901, p. p.-Perrot et Guerin, Didurea,
100m. de bot., 1903, pp. 333-SI.-Areschoug, Trop. viixt. bladbyggn., Sv. Vet.-Akad: Handl., 39,
D. :It 1905, pp. 68,69 and Tab. ii (Nephelium).-Haberlandt. Lichtsinnesorgane, 190 5, P.97.-
Piccioli, Legnami, Bull. Siena, 1906, p. 176.-[Radlkofer, in Natiirl. Pflanzen£am., Erg.-Heft ii,
19o~. pp. ~02-9;
-
see
- ... zum ii-iv Ten, I897~ 228.]
also Nachtr. p: ,

HIPPOCASTANACEAE (pp. 236-237).

The stomata on the leaves of Aesculuscali/ornica, Nutt. are provided with
neighbouring cells which bear striate papillae, converging towards one another
in a radiate manner. -
Literature: Uhlworm, Entwicklungsgesch. d. Trichome, Bot. Zeit., Ifl73,- p. 8I8.-Hohne1,
Gerberinden, Berlin, 1880, p. lIS et seq.-Nanke, Blatt u. veget. Axen dikot. Holzpff., Diss.,
Konigsberg, 1886, p. 16 et seq.-Kobne, Papill. u. obers. SpaltoH'n., Mitteil. dendrolog. Gesellsch"
1899, p. s8.-Tunmann,. Sekretdritscn, Diss., Bern, 1900. pp. 35, 26.-Bouygues, Cert. merist. vase.
dans Ie petiole, Act. Soc. Linn. Bordeaux. lvi, 1901, pp. lvii, lviii.-Pitard, Pericycie, "These,
Bordeaux, 190J, pp. 87, 88!-Bottygues. petiole, These, Paris, 1902, pp. 8 and II:-Col, Faisceaux,
AnD. sc. nat., ser. 8, t.'xx, 19°4, p~ 140.-Piccioli, Legnami, Bull. Siena, 1906, p. '42.

ACEEA~EAE (pp. 238-240).

I. REVIEW OF THE ANATOMICAL FEATURES. According to Warsow,
secretory sacs are presel1t in the veins of the leaf in all the species of Acer,
although their contents are milky only in a small number of cases. Latex 4

cells situated in the mesophyll, besides occurring in A. campestre, L., are found
also in A. Pic.tum, Thunb. and A. neglectum-, Lang., which belong to the section
P1atanl?idea 1. Sausage-shaped or almos( spherical idioblasts containing
mucilage have been observ~d in the mesophyll in A. laevigatum, Wall. and
A_ oblongum, ·Wall. The clusters of calcium oxalate crystals occasionally
resemble sphaerites; another form of excretion of oxalate of lime is thatJ)f
small rod-shaped or a~icular crystals, which are found in the mesophyll. 'In
the genus 4cer the unicellular clothing hairs are accompanied by uniseriate
ones; two-armed hairs have been recorded also in A. parvijlor'ltm, Franch.
et Say. '
2. STRUCTURE OF' THE LEAF~ The following statements ar,e based on
·\Varsow's ,recenf,investigations, which' deal with eighty-five species and have
led, to certain improvements in Pax's system of classification.
The structure of the leaf is for the most part bifaCial and rarely centric
or subcentric. The palisade-tissue generally consists of a single layer of cells.
The stom~ta _are ~ost invariably c«?nfined to the lower side of the leaf; only
iD: a few,species (e.g. A. iaponicum, Thunb.) do they occur in small nUqlbers also
on -the upper side, where they are found in the neighbourhood of the veins.
The latera~ ~rgins of the epidermal cells are straight or undulated, but in
----'----_.--- ----_._
1 The nomenclature of the. sections adopted here and in the following description is based on Pax's
l'evi:ion of. the Acericeae in Engler's ( Plla~zenreich.' -
 ADDENDA-ACERACEAE

a few species (e.g. A. Trautvetteri, Medw. or A. nikoense, Maxim.) the cells

of the lower epidermis have jagged margins. The cuticle occasionally e$bits
striation, while in certain species it is covered by a layer of wax of varying
thickness; the latter is the cause of the white colour presented by the lower
surface of the leaf in A. Drummondii, Hook. et Am., A. niveum, BI. and A. sac-
charinum, L. 'Gelatinization of the epidermis of the leaf is remarkably common
in the genus Acer (for details, see Warsow's paper), either affecting both upper
and lower epidermis or only the upper epidermis. Papillose difierentiatlOn
of the lower epidermis has been recorded in the following additional species :
A. caesium, Wall., A. DureUi, Pax, A. glabrum, Torr., A: griseum, Pax, A. HeZ-
areichii, Orph., A. hybridum, Spach, A. insigne, Boiss. et Buhse, A. mand·
schuricum, Maxim., A. monspessulanum, L., A. nikoense, Maxim., A. niveum,
Bl., A. ramosum, Schwer., A. sericeum, Schwer., A. sinense, Pax, A. sutchuense,
Franch. and A. r.rautvetteri, Medw. Specially noteworthy features' are: the
occurrence of one or two small solitary crystals in cells of the lower epidermis
in A. erianthum, Schwer. and A. oblongum, 'Wall., these cells being generally
somewhat smaller than the remaining epidermal cells; and the presence of
tanniniferous idioblasts in the lower epidermis in A. villosuffl" 'Vall. Hypoderm
has not been recorded in any species of Acer. The larger veins almost invariably
contain a ring of sclerenchyma; but in most of the species belonging to the
section I ndivisa there are only arcs of sc1erenchyma or isolated sc1erenchy-
matous elements. In all the species of Acer the smaller veins are vertically
transcurrent by me~ns of tissue with thin or thick walls.
Oxalate of lime is developed in relation to the vascular bundles of the
veins; in a certain number of the species it is deposited mainiy in the form
of solita.ry crystals, while in the remainder it occurs principally in the form
of clustered crystals. Lar~e idioblasts which, in addition to smaller crystalline
masses, contain a large solItary crystal, often placed with its long a_xis at right
an.gles to the surface of the leaf, are found not only in A. N egundo, L., but also
in the remaining species of the section lVegundo (A. californicum, Dietr. and
A. mexicanunt, Pax), as well as in A. cissifolium, C. Koch; these element" cause
transparent dots in the leaf. Similar idioblasts containing clustered crystalS;
which occasionally resemble sphaerites, are present in the species of the section
I ndivisa (with the exception of A. carpinifolium, S. et Z.), in A. glab1um,
Torr. and in those of the section Macrantha. We have already pointed out
above that according to Warsow the secretory sacs no doubt occur in all the
species of Acer; they are situated in the bast in the vascular bm).(Ues of the veins
and probably in the axis as well. In transverse sections of the leaf, which have
,b,t.en bleached with Eau de Javelle, ,they show up distinctly owing to their
'large lumina, but only in a small number (12 out of 85) of the species do they
contain typicalla.tex. These species are the following: A. macfophyllum, Pursh
(which should perhaps be transferred from the Sect. Spicata to the Sect. Plata-
noidea); all the species of the section Platanoidea which have been investigated,
viz. A. laetum, C. A. Mey., A. Lobelii, Ten" A. Miyabei, Maxim., A. neglect'llm,
Lange, A. picturn, Thunb., A. platanoides, L., and A. truncatum, Bge.; A. cam-
pestre, L. (which should perhaps be removed from the Sect. Campestria and
pJaced in the Sect. Platanoidea); lastly, A. diabolicum, Bl., A. purpurascens,
Franch. et Sav. and A. Thomsonii, Miq., which belong to the s~ction LithocaTpa.
In the remaining species the contents are glassy and rather strongly refractive;
they are readily soluble in alcohol and water, and are best seen by mounting
dIS transverse --sections in olive-oil. The latex-cells and mucilage· idioblasts ,
or.rorring freely in the mesophyll in certain species, have already been referred
to above.
The most widely _distributed type in the hairy covering are unicellular
dothing hairs. In some cases they are developed in the 'form of papillae, but
 ADDENDA--ACERACEAE

this type of differentiation ~ only found in species which have a papillose epider-·
mis in the leaf, and in which the hairs constitute a transition to the papillae.
As·a tule, however, the clothing hairs are elongated structures, which are either
wavy or stiff; in the latter case they may be straight or bent after the manner
of a sabre. Uniseriate clothing hairs (composed of eight cells or less) are found
only in a small number of species (mainly in the section Platanoidea). The
ends of the clothing hairs are for the most part pointed, rarely (A. parvi/lo,um,
Franch. et Sav.) rounded. Two types of glandular hairs may be distinguished.
The first of these, which is the most widely distributed, comprises approximately
club-shaped glandular hairs with a uni- or biseriate stalk (mostly composed
of 5 or 10 cells) and a multicellular head. A modification of this type of hair
is found in the species of the section- Platanoidea, in which the cells of the stalk
are fiat and the head spherical. The second tyPe of glandular hair is constituted
by uniseriate external glands, which in certam species pass over into (multi-
seriate) shaggy hairs; a peculiar form of trichome, which may be noticed in
this connexion, is that of glandular shaggy hairs, which exhibit division into two
uniseriate glands at a point situated at a varying distance from the base of the
trichome. The second type of external gland occurs in the species of the section
Indivisa (with the exception of A. ca,pinifolium, S. et Z. and A. distylum,
S. et Z.)~ in A. Tubrum, L. and in the species of the section Macrantha (excepHllg
A. parvi/iorum, Franch. et Sav.).
For the vascular system of the petiole, see also Bouygues and Cot, u. cc.
Literature: Gregory, Cork wings, Bot. Gaz., 1888, pp. 28l, ::S:I.-[Tognini, Stomi, Atti Ist.
bot. Pavia, I 894.]-Kohnc, Papill. u. oberseit. Spaltoffn., Mitteil. deutsch. dendrolog. Gesellsch.,
1899, p. s8.-Hammerle, Acer Pmedoplatallus, Bibl. bot., Heft 50, 1900, 101 pp.-Petersen,
Vedanatomi, 11)01, p. 4i.-Bouygues, Petiole, These, Paris, J90l, p. n.-Pax, in Pftanzenreich,
Heft 8, 1902, p. ::.- Tuzson~ Spiralige Strukt. d. ZeU w. in den Markstr., Ber. deutsch. bot. Gesellseb.,
1903, p. :l76.-Warsow, Syst.-anat. Untersuch. d. HI. bei d. GatL Acer, etc., Diss., Erlangen. 1903.
109 pp.; scp. copy (rom Beih. z. Bot. Centralbl., xv, p. 493 et seq.-Col, Faisceaux, Ann. se. nat.,
ser. 8, t. xx, 1904, pp. 136-9.--Siissengnth, Behaarunglverb. der Wiirzb. Muschelkalkpfl., Disli.,
Wiirzburg, 1904, p. 24·-Haberlandt. Lichtsinnesorg., 1905. pp. 105, 106 and Tab. iii.-Theorin,
Vaxttrichom., Arkiv for Bot., iv, D. IS, 19°5, pp. 2, 3.-Piccioli, Legnami, Bull. Siena, 1906, p. 138.

STAPHYLEACEAE (pp. 242 , 243).

Literature: Piccioli, Legnami, Bull. Siena, 1906, p. 130-
./

SABIACEAE (pp. 243, 244).

The exact nature of the' Pooches secretrices.' mentioned by Pierre (in FIQ5d.....
forest. de la Cochinchine, xxiti. 1897) as occurring in Meliosma Cambodiana, ,
'J;D.ust be made the subject of renewed investigation, in view of the fact that in other
species of M eliosma Radlkofer has recorded the presence of cells having siliceous
contents (not resin.cells, see p. 243).
literature:~ Pierre,'loc. cit., ·.-897 .....:.Pitard, Pericycle, These, Bordeaux, 1901, pp. 76 and 9,...-
[Dihm, Blatt d. Gatt. Meli()sma in anat. Hinsicbt, Beib. z. bot. Centralbt., xxi, Abt. I, 1907, 31 pp.
and Tab. v, vi.]

ANACARDIACEAE (pp. 244-248).

1. REVIEW OF THE ANATOMICAL FEATURES. To the statements on the
hairy covering we may add that simple uniseriate clothing hairs, as well as
branched (glandular ?) multicellular trichomes also occur in this Order.
2. STRUCTURE OF THE LEAF. The mesophyll is bifacial in most of the
species of Rhus of the section Gerontogeae, Engl., which have been investigated
 ADDENDA~ANACARDIACEAE

by Diels; in R. incisa, L. f. it consists of palisade-tissue only. In the species of

Rhus, belonging to the section just named, the stomata are either confined to
the lower side or occur on both sides of the leaf. In R. Burkeana, Sand. they
are deeply sunk and provided with a vestibule, the inner margin of which 15
formed by the neighbouring cells of the stoma, which project considerably
beyond the level of the epidermis. Papillae are found on the lower side of the
leaf also in R. Cotinus, L. (according to Knothe) and R. Osbecki, DC. (according
to Kohne). Regarding the hairy covering we may add the following facts.
The uniseriate trichomes occurring in certain species of Rhus are hi- to multi-
cellular and have pointed of blunt ends. R. ho"ida, Eckl. et Zeyh. has stellate
trichomes with thin walls, the ray cells of which show a radial arrangement and
lie in a plane parallel to the surface of the leaf; similar trichomes, which,
however, exhibit irregular orientation of the fay-cells, are found in R. somalensis,
Engl., while R. incana, Eng!. has irregularly branched trichomes composed of
a large number of cells; all these different forms of hairs may possibly have
a glandular function. Among the types of glandular hairs observed by Diels
we may mention the external glands found in R. discolor, E. Mey; they
have a unicellular ·stalk and a unicellular glandular head of variable shape.
Small peltate glands with irregularly arranged glandular cells have been recorded
also in Campnosperma (Fabricius).
3. STRUCTURE OF THE AXIS. Silica·bodies are found in the wood·paren-
cbyma in M elano"hoea obtusifolia, Eng!. (Bargagli- Petrucci).
The contents of the secretory canals are sometimes (species of Rhus) of the
nature of latex; see also Molisch, loco cit.
Literature: [VogI, Gum of Quebracho colorado, Pharm. Joum. and Transact., 1880; abstr. in
Bot. Centralbl., 1880, p. I04:t.]-Jadin. Org. secret., These, Montpellier, 1888, p. ~~ et seq.- J. E.
Weiss, Korkbild., Denkschr. bot. Gesellsch. Regensburg. vi, 1890, p. 6..f..-Jadin, Q!il' Th'ebinth.,
Joum. de bot., 1893, p. 382 et seq.-Knoblauch, Oekolog. Anat. etc., Habilitat.-Sclff•• Tiibingen,
1896. p. II et seq.-Werner, Neuere Drogen, Diss., Erlangen, 1896, pp. 1-10 (Cortex CC1It«ladUu).-
Diels, Ephannose d. Vegetationsorg. bei Rhus, L. § GeYon/ogeae, Engl., in Engler, Bot Jahrb,. mv,
J898, pp. 568-647 and Tab. xiv.-Pierre, Flore forest. de la Cochinchine, x:xiv, 1898 (f)nuoH/lI-
me/um), and x'iii, 18941 (Mangijtra).-Kohne, Papill. u. oberseit. SpaltOfl"n., Mitteil • •eutlCh.
dendrolog. Gesellsch., 1899, p. S8.-Mobius, Der ja.pan. Lackbaum, Rhus vernidfera, A~.
Senckenberg. Gesellsch., xx, 1899. pp. 2IO-:J~.-Inuj, Gl1.mmibarzgang ll. Lackbaurnes, etc., Bot.
Centralbl., 1900, iii, p. 35:t.-Briquet, Anat. comp. de la fenille chez les Pis/ada LentisCUJ, etc.,
ser.
Bull. de 1'lIerbier Boissier, 2, t. i, IgOl, pp. 1301-S.-Molisch, Milchsa(t u. Schleimsaft, 1901,
p. '5. footnote I.-Pitl\rd, Pericycle, These. Bordeaux, 1901. pp. 40 and j4.-Bargagli-Petrncci,
Concrez. silicee, Malpighia, 19°2, p. 23 et seq.; and Legnami, loco cit., p. 319 et seq. (CamjmlJ-
spertt14. MelanQchyla, Jl.fdaniJrrkoea, Pmtaspadon).-Botlygues, Petiole, These, Paris, 1902, p. 12.-
Fabrl"ius. Lanbblatt-Anat.. Beih. z, bot. Centralbl., xii, 1902, pp. 311, 312.-Kn?the, Unbenetz·
bare m., Diss., Heidelberg, 190", p. lo.-Simon, in &t. deutsch. bot. Get;eIlsch., 19o1, p. l41.-
{"...-marl, Piante della reg. medit., Ann. di Rot., i, 1903, p. 17 et seq. (Pistacia).}-Qaanjer, Anat.
oonw, etc., Natllurk. Verhandel. Haarlem, iii. 5, 1903 (Gluta BenKkas and Bu{hanania sp.)._
Ursprnng, in Bot. Zeit., 1904, p. 19R.-Areschoug, Trap, vaxt. bladbyggn., Sy. Vet. Akad. llandl,
39. n. 2, 1905, pp. 76, 71 (Mangifira), and pp. 124. 125 (Anacardium).-Sarton, Anat. d. pI.
atllnes, Ann. se. nat., set. 9, t. ii) 1905, pp. 63-79 (Pislacia).-Piccioli, Legnami, Bull. Siena 1906,
pp. J.p, 150 and I6o.-[For further literature, see p. 1169']

CORIARIEAE (p. 249).

Oxalate of lime is found in the form of clustered crystals also in the rhizome
of Co ria ria myrtifoliiJ..
Lilerature: Villeneuve, Le Red-ouI, These, Montpellier, 1893,62 pp., especfaUy pp. 20-37.
 ADDENDA 895

MODINGEAE (pp. 249, 25 0 ).

Guignard's discovery of the occurrence of myrosin.cells in this Order
is of great importance in interpreting the taxonomic relations between the
Moringeae and the Capparideae.
Jadin has investigated the distribution of the myrosin in detail in Moring"
pte,ygosperma. According to him the myrosin.cells of the root are situated in the
bast and primary cortex, myrosin being distributed throughout the peripheral'
layers of the cortex. In the branch, myrosin~cells are likewise present in the bast
and the primary cortex (here especially in the subep~germallayer of cells). Similar
cells occur also in the mesophyll.
Literature: Jadin, Localisat. de la myrosine et de la gomme chez les Moringu, Comptes rendus
Paris, J900, I. sem" pp. 733-fi'

CONNARACEAE (pp. 250-253).

Pierre records small secretory cells and a hypoderm in the leaf in Cnestis
ramiflora, Griff., secretory cells and an almost sclerosed palisade.tissue com ..
posed of two layers in Aglaia Cambodiana.
A composIte and continuous ring of sclerenchyma is developed in the
pericycle also in Connarus paniculatus (Pitard).
Literature: Bartels, Cangoura, Diss., Erlangen, 1894, pp. 28, 29._':costeruli, ConHllrus, Ann.
Jardin Buitenzorg, Suppl. ii, 1898, pp. 109-12 and pl. iv.-Pierre,: Flore forest. de 14 Cochinchine,
xxiv, 1898.-Pitard, Peri cycle, These, Bordeaux, 1901, p. 8a,-Areschong, Trop. ya.:It. bladbyggn.,
Sv. Vet. Akad. Handl., 39, n. 2, 1905, pp. 1 I 1, 112 and Ta.b. xiv, xv (Connarus).-[Sperlicb, Opt.
Verh. in d. oberseit. Blattepiclermis, Sitz.-Ber. Wiener Akad., cxvi, Abt. I, J907, p, 718.]

LEGUMINOSAE (p. 253).

1. PAPIUONACEAE (pP. 253-28I).
I. REVIEW OF THE ANATOMICAL FEATURES. The following additional
facts have been recorded:-
Internal secretory organs (p. 254) :~Mucilage-cells are found in Cdragana,
and mucilage-lacunae in Alhagi and H aUmodendron. Secretory ca vi ties in the
form of ordinary intercellular spaces filled with secretion occur in A nthyllis
Genistae, Duf., while closed schizogenous secretory cavities are present in the
leaf in the monotypic genus Cordyla. Lastly, schizogenous secretory canals
are found in the primary cortex of the branc!i in Cordyla.
Oxalate of lime (p. 254) :-Small crystalline grains or needles are not
uncommon in the Podalyrieae and Genisteae, and oCC.U!, also,in members of the
Trifo1iea~,,.Loteae~llnd Vicieae.
Hairy ·-covering (pp. 254, 255) :-0£ special forms of clothing hairs three-
celled trichomes having a two-armed terminal cell are present also in certain
Podalyrieae and Genisteae. With reference to the distribution of the external
glands we may note that, taking the results of the earlier investigations into
consideration as well, they are of frequent occurrence only in the Tribes Tri-
folieae, Galegeae, Hedysareae, Vicieae, Phaseoleae and Dalbergieae, while
they are absent in the Podalyrieae, Sophoreae and Swart#e~e, and are only
rarely met with~in the Genisteae and Loteae.
Special anatomical features (p. 255) :-Pseudo-pitting or internal striation
of the epidermal cells of the leaf (certain Podalyrieae and Genisteae); epidermal
cells of the leaf having a prosenchymatous shape (species of [Athyrus) or elon-
 ADDENDA-LEGUMINOSAE

gated transversely to the midrib (species of Eutaxia and Trifolium) ; a peculiar

depression of the stomatal apparatus (species of Jacksonia); restriction of the
stomata to the upper side of the leaf (in certain Podalyrieae which have rolled
leaves with a furrow on the upper side); transverse arrangement of the stomata
on the leaves (species of Anarthrophyllum, Eutaxia, and Latrobea) or on the
assimilating axes (Daviesia divaricata, Benth.) ; spicular fibres in the mesophyll
(species of Pultenaea, DiUwynia, Bossiaea); special structural features pre-
sented by the veins of the leaf (' double vascular bundles' in Daviesia, annular
arrangement of the vascular bundles of the leaf, &c.) ; lastly, the occurrence of
sphaerocrystalline masses or of bodies resembling indigo or indican in the
mesophyll.
Anomalies in the structure of the stem (p. 255) ;-Successive rings of
growth occur also in Strongylodon.
2. STRUCTURE OF THE LEAF. In recent times a detailed investigation
of the structure of the leaf has also been undertaken in the Podalyrieae (by
Prenger I, Biirkle ~ and Huhner S), the Genisteae (by Schroeder 4, Cohn II,
Winkler 6, Levy', H. Schulze 8, W. Schulze 9 and Rauth 10), the Trifolieae (by
G, Fischer 11), the Loteae (by W. Schmidt 12) and the Vicieae (by Streicher 121).
The epidermis of the leaf in these groups sl}ows features similar to those
found in the Tribes previously investigated, the characters being for the most
part of value in specific diagnosis. In many Podalyrieae and also in certain
Gen~teae the cuticle exhibits verrucose thickenings; this feature is commonly
combined with the penetration of peg-shaped or lamella-like processes of the
cellulose-membrane into the outer cuticularized portion of the external wall ;
this -leads to a peculiar structure as seen in surface-view (' pseudo-pitting' or
'internal striation '), Lateral walls exhibiting angular folds have recently
been observed also in Anagyris loetida, Ten., species of Ononis, Trigonella,
M elilotus and Trifolium, and species of Securigera, Dorycnium, Lotus (see
Syst. Anat" p. 256, footnote r), Anthyllis, Hymenocarpus and Hosackia; epider-
mal cells of prosenchymatous shape have been recorded in species of Lathyrus;
epidermal cells elongated transversely to the midrib in species of Eutaxia and
Trifolium; special small (empty?) cells in the epidermis in certain species of
Lupinus. Formation of papillae is a common feature; the papillae vary in
s~ape, are sometimes solid or reduced to cuticular humps! and may be.I?~~~nt
eIther on both surfaces of the leaf or only on one, An Important poirit 'wIth
reference to the gelatinization of the epidermis lies in the fact that epidermal
cells.with mucilaginous inner membranes have not been: observed in any member
of the Loteae, Trifolieae or Vicieae.

I Species of the following genera were examined: Anagyris, PiptalltRus, Thermo/sis and
Baptisl'a (northern hemisphere); Cyclopia and Podalyria (Cape of Good Hope) j Brachysema,
Ox}'/olJium, Chon'::ema and lJ.firbtlia (Australia).
!l Species of lsotropis, GOIllPh%bi14m, Btlrt(mia.J(lrksoltia, Spkaerolobiunl, Vi1lliltaria, Dafliesia,
AotlfS, Phyllola.
S Species of GastroloDium, Pu/ttllata, La/robta, Eutaxia, DiIlwYllia.
I Species of Li'j)(wia, Pritstltya, A mplzitha lea , Lalh,,£o,vne, Coelid;um, Platy/obiu", , B(mUua,
Ttmpletonia, HMMa, Goodia. .
.i Species of BarDonia, Gafnia, Euch/IJl"fl, Lolommi.r, Rothia, LehcJda, V'-h"l"gia,
• Species of Crotalaria and Pn'{)Yitl"opis.
, Species of Aspalathus, BudunriJedera, lUd%bium, DieMllls, Hey/al1dia.
I Species of Lupinu! and Argyrolo/Jittlli.
• Species of Gmista with AnarlkropAyllum, Admocarjms, Ca/y(otollu. ~
10 Species of LalJuntum, Pet/tn'a, Sparliu111 , EriuQaa, Uter,. Cylims, IIj/,ocalyplm and
Loddl""g'csia.
'11 Species of Ollonis, Tri~dla, lrltdicago, ,ft/eli/Dtm, Tnfolium,
n Species of Anthyllis, Helmintn.()(ar;Um, Hymen()(arjJtu, Se(ungtra, Cylisopsis, DD1ymiul11,
Lotlfs, lIosacki"a. .•
13 Species of Cictr, "Pz'cia, Lms, Laln.yrlls, PlUWI, AII,.us.
 ADDENDA-LEGUMINOSAE

The pseudo-pitting or internal striation above referred to has been observed

in the following cases: (I) amon~ the PodaIyrieae, in species of Cyclopia, Podalyria.
Oxylobium, Gompholobium, Damesia, Aolus, Burtonia (here penetrating into the
papillae in the form of radiating tufts). Pultenaea; (2) among the Genisteae, in species
of Aspalathus, and in Lup'inus ,;on'aceus, Benth.
Papillose epidermal cells have recently been observed as follows: (I) among
the Podalyrieae, in species of Cyclopia, Brachysema, Oxylobium. Chorizema, Mil'belia.
Gompholobium, But'tonia, Sphaeyolobium, Dava'esia. A o/us, PkylJola, Gast,olob";um,
Pultenaea, Latrobea, Eutaxia, Dillwym'a; (2) among the Genisteae, in species of
Liparia, Priestley a, Coelidium, Bossiaea, Templetonia, Hovea, Goodia, Crotalaria,
Prioritropis (P. cytisoides, W. et A.), Lupinus (only isolated epidermal cells), A ~cy,o­
lobium (only isolated epidermal cells), Genista (also in G. pilosa, L. accordi~ to
Kohne, d. W. Schulze), Adenocarpus, Calycotome, Laburnum. Spartium, CytJSUS,
Hypocatyptus, Loddigesia; (3) among the Trifolieae, in species of Tf'ijoli14m; (4)
among ilie Loteae, in species of A nthyllis, Dorycnium, Lotus, H osacAia; (s) among
the Vicieae, in Abrus tenuifiorus, Spruce; (6) among the Galegeae, also in Indigo.
le,a Gerardiana. Wall. (according to Kohne I). Specially noteworthy forms are the
papillae of Burtonia scabra. R. Br. and B. villosa, Meissn., which are provided with
a coronulate apex and are connected by reticulate cuticUlar ridges, and the papillae
of Dillwynia, especially D. hispida, Lindl., which form longitudinal rows projecting
in the form of ridges.
Epidermal cells with mucilaginous inner membranes have been recorded in
the following cases: (I) among the Podalyrieae, in species of Cyclopia, Bl'achysema,
OxyloQ·ium, Ckori$ema, Mil"belia, Burtonia, Sphaerololyium, Vim:r:naria, Aotus. ,Phyl-
Iota, Gastrolobium, Pultenaea, Latl'obea, Eutax,'a, Dillwym'a; (2}among the Genisteae,
in species of Platylobium, Bossiaea, Templetonia, Hovea, Goodia, Borbonia. Ra/nia,
Euchlo,a. Lownonis, Rothia, Lebeckia. Viooygia, Aspalathus, Melolobium, Dic/u1us,
Heylandia. Cf'otalaria, Priol'it'Yop1·s, ArgYl'olobium, Genista, Adenocarpus, Calycolome,
Laburnum, Petteria, Spat'tium, Erinacea, Ulex, Cytisus.
Hypoderm has recently been recorded among the Podalyrieae in species
of the genera Daviesia (on both sides), Pultenaea, Eulaxia and Dillwynja
(here on the lower side, and filled with brown contents), as well as in Crotata,ia
lunulata, Heyne (on the upper side) and Cicer subaphyllum, Boiss. (here in the
leaf-spines). Thin vertical walls are found in the epidermal cells in species
of Pettel'ia (in the neighbouring cells of the stomata) and Cytisus,
There is again no uniformity in the type of stoma found ·'In the Tribes
Podalyrieae, Genisteae, Trifolieae, Loteae and Vicieae. In the Podalyrieae
the pairs of guard-cells are very often surrounded by a varying number of
ordinary epidermal cells, but the following additional types of stomata have
been noticed in this Tribe: stomata with four neighbouring cells, two of which
are parallel to and situated to the right and left of the pore (in species of
Brachysema, Oxylobium, Dil/wynia, and in Aotus cordi/olia, Benth.); stomata
with three neighbouring or subsidiary cel~s (in species of Jacksonia, Sphae,o-
lobium, Gastrolobium, Pultenaea and Latrobea); and stomata of the Rubia..
ceons type (in species of }acksonia t Euiaxia and Dillwynia). In the Genisteae
the following types are found: stomata approximating to the Cruciferous
type (in'species of- Rafnia, Borbonia, Lotononis, Lebeckia, Viborgia, Crotalaria,
Pl'ioritropis); stomata showing an approximation to the Rubiaceous type
(in Hovea, Hypocatyptus and Borbon-ia crenata, L.; in the last only some of the
stomata belong to this type); and lastly, stomata surrounded by a rosette
of epidermal cells of the nature of subsidiary cells (in Templetonia, Lebeckia
psiloloba, Walp. (on the axis), and species of Anarthrophyllum and Genista).
For most of the Trifolieae Fischer r~cords 3-4 (rarely more) neighbouring
cells arranged according to the RanUnculaceous type; Ononis alone follo\\s
1 In Kohnets paper (loc. cit.) a Dumber ot species of Colutea and Ro/Ji,,;a are also mentioned, aa
bring a papillose epidCJ1l)is ill the leaf; Weyland, who investigated the Galegeae, did not examIne
these species.
SOLIUt.bER 3 M
 ADDENDA--LEGUMINOSAE
the Rub:raceous type. According to Schmidt, th.e neighbouring cells (numbering
usually 3-4, rarely 2 or 6-7) of the stomata in the Loteae are not specialized.
Nor, according to Streicher, are those of the Vicieae; but in species of
Cice" Vicia, Lens and Lathyrus, a small number of stomata have been
observed with a parallel subsidiary cell situ~d to the right and left of ths.
pore; in Abrus there are four subsidiary cells, of which one is occasionally
placed on ,either side of and parallel to the pore. The stomata situated on
the asSimilating branches of certain species of ] acksonia are sunk and require
special mention. The pores of the stomata 3;re here placed parallel to the
vertical direction, and are accompanied either by two (in this case parallel
to the pore) or three subsidiary cells, which have remarkably thin walls.; the
guard-cells, together with their subsidiary cells, are sunk in the surface of the
branch in a deep pit, which generally has a lon~, narrow outline. On that
part of the margin of the pit, moreover, which IS situated nearest the base
of the stem, a "hair is inserted, the terminal ceU of which has thin ·walls
and wide hunina and covers in the pit. With reference to the distribution
of the stomata on one or both surfaceS of the' leaf, the following facts may
be mentioned: in the Trifolieae and Loteae which have been investigated,
the stomata are invariably present on both sides; in the rolled leaves of
certain genera of the Podalyrieae (viz. PuUenaea, Eutaxia and Dillwynia.
in which the leaves are furrowed on the upper surface), as well as in Coelidium
(Tribe Genisteae), the stomata are confined to the upper side 1. The stomata
are very commonly placed parallel to one another. their pores being at the
same time directed parallel to the midrib of the leaf. A more striking feature
is furnished by stomata which lie parallel to one another, but are arranged
transversely to the midrib of the leaf or to the vertical axis of the branch, as
the case may be; e.g. on the leaves in species of Eutaxia and in Lat,obea
tmella, Benth., and on the assimilating axes of Daviesia divaricata. Bentb.
(Tribe Podalyrieae), as well as on the leaves of Anarthrophyllum Cumingii,
Phil. f. and A. andicolum, Phil. f.
The following features of the mesophyll may be noted: the central pith-
like tissue, found in the terete ot narrow leaves of species of Daviesia and
Lebeckia; the parenchyma in the interior of the acicular leaves of species
of Aspalathus, which envelops the vascular system and resembles a pith;
the occurrence of weakly developed spicular fibres, which are connected with
the sclerenchyma of the veins, in species of Pultenaea, Dillwynia and Bossiaea,
and of sclerosed parenchymatous cells in species of Buchenroedera.
For details as to the structure of the veins, the reader is referred to the
papers cited below. The smaller veins of the leaf are vertically transcurrent
m certain Podalyrieae, Genisteae, Trifolieae (species of Trifolium) and Vicieae
(Abrus). Sc1erenchyma is occasionally not developed in the veins; this is
specially often the case among the herbaceous species. According to
Streicher, the sderenchyma in the veins of Vicia, Lathyrus, Lens and Pisum
is confined to the side on which the xylem is situated. In those species of
Daviesia (Tribe Podalyrieae), in which the leaves are approximately horizontal
and of some breadth, the median veins consist of two opposite systems of
vascular bundles with their xylem·groups abutting on one another (' double
vascular bundles '); the lateral veins either contain' double vascular bundles J

of the same type, or exhibit a row of simple bundles in transverse section,

the xylem.groups of these bundles being directed alternately towards the
upper and lower side of the leaf. An annular arrangement of the vascular

1 KOhne (loe. cit.) states that the stomata are confined to the upper side of the leaf also in
GtttiJla it.istJ(utica and G. Sritd, as well as. in Lespedeza seric~a (Hedysareae); according to
W. Schulze this is incorrect in the case of the first two species.
 ADDENDA--LEGUMINOSAE
bundles is found also in the leaves of the species of Lebeckia, which contaia
a tissue resembling 'a pith.~ their interi.r. Sheaths compOsed of large
parenchymatous cells occur in certain Podalyrieae, Genisteae and Loteae,.
and enlarged terminal tracheids or storage.. tracheids are present in certain
representatives of the same tm¥e Tribes.
In dealinfl with the secretory organs we will firft- consider the tannin-:
sacs. Their distribution in the Tribes which have been recently investigated'
is as follows.: They are present in the leaf in certain Podalyrieae and tOteae;
in the Tribe Genisteae they occur only in Goodia; in the Tribe Trifolieae'
distinctly differentiated sacs have been demonstrated only in Pa,ochetus;
and in the Tribe Vicieae they are altogether wanting.
To the synopsis on p. 260 we may add :-1 a. Podalyrieae: According to
Prenger, Burkle and Huhner, tannin-idiobla.sts are found in the leaves of certain
species of Cyclopia, BYacbysema, O~ylobium, ChoyizeJfJa, Miybeli~ Aotus. Ph"JJoJa.
Pultenaea, LatrobefJ, Eutaxia and Dillwynia. I b. Geliisteae: Schroeder records the
occurrence of sac-shaped tannin-cells in the palisade-tissue, and of a hypodermal layer
of cells :6.1Ied!with brown contents in the spongy tissue in Goodia, a~enus which
is anomalous also in the possession of trifoliate leaves, and should probably be
transferred to the Galegeae. 2. Trifolieae: Tannin-sacs have been observed by
G. Fischer only in the parenchyma of the veins of Parochetus communist HaIl'lilt- ;
they have wide lumina and are arranged in rows. 3. Loteae: According to
W.Schmidt, tannin-sacs occur in the leaves in Anthyllis pro parte. Helminthocal'pum,
Cyti5opsis~ DOl'ycnium~ Lotus pro parte. arid Hosackia (but not in Hymenocarpus
and Securigera). 6 a. Vicieae: According to Streicher, there are no tannin-sacs
in the leaf. 9. Swartzieae: According to Guignard, tannin-sacs are present also
in Cordyla (primary cortex and parenchymatous pericycle). ..
• •
Other types of secretory cells have recently been observed by :JOnsson.
viz. mucilage:-sacs in the palisade tissue of 'Ca,agana j,utescens, DC.' and
mucilage-cells in the secondary cortex of Alhagi camelorum, Fisch. With
these we may class' the mucilage-lacunae which the same author found in
older parts of the axis of Alhagi camelarum and. Halimodend,on a,cmkum,
DC.; these lacunae are situated in the secondary cortex in the'former species,
and in the pith and secondary. cortex in the latter. We may also/point out
here that Lebeckia? retamoides, in which G. Cohn found rounded secretory
cells with resinous contents, belongs to the genus Tephrosia (see Solereder,
loco cit.). /'
Secretory cavities have also been demonstrated in Anthyllis Genistae,
Duf. l They are confined to the leaf, and consist of ordinary intercellular spaces
filled with a bright yellow resinous secretion, which is soluble in alcohol;
they are connected with the remaining intercellular spaces and thus recall
t he secretory receptacles found in Derris, &c." According to Guignard, .closed
schizogenous secretory cavities occur in the leaf of Co,dyla africana, Lour.
(Tribe Swartzieae); in the branch of the same plant they are replaced by
schizogenQus. ?_ecr~tpry canals (situated in the primary cortex, but not in the
pith or wood).
New forms of crystals of oxalate of lime or typical clustered crystals

1 In view of the isolated occurrence of secretory cavities i.Jl this speci~ (the cavities are wanting
in the closely related A. c),Iisoides, L. and A. Htrmanniae, L.) it may be expressly pointed out that
a lcaRct of the original plant (Herb. DC.) has also been examined. ,
I Secretory cavities, provided with a loose papillose epitheliwn (similar to those fOUJld in certain

specie& of Lonclt~car'pus) have recently been recorded by Geiger (see also Dllval) also in Swc"/Jia
tftcipiem, Holmes, the leaves of which constitute a new form o{ adnlter.ation o( Jaborandi.leavea.
In all pr(}bability, however, S. detipims, Holmes no more belongs to the genua Swarlzia than does
the plant (Sptnce, No. 1197, Brasil) previously investigated by Kopff and described as S . .Iltrna,
Benth.: the latter exhibits • secretary m46S~ ~n,,~loptd by bracket-«lls,' at in cm..in species of
LOllcJrocarpUl and Dtf"I'is.
3 M2
 ADDENDA--LEGUMINOSAE
have·. not been recorded·t • All the Australian genera of the Podalyrieae
possess ordinary large rhombohedral crystals or hemitropic forms of such
crysta4;, which are often rod.shaped; the crystals'of this type are occasionally
accompanied by small prismatic or aciclllaf crystals. The latter are the only
forms of crystals pr~ent in the genera endemic in the region of. the Cape
(Cyclopia and Podalyria) and in those inhabiting the northern bemisphere
(Anagyris, Piptanthus, Thermopsis and Baptisia). Among the Genisteae
ordinary large crystals or their hemitropic forms occur only in Platylobs'um,
Bossiaea, Templetonia, Hovea and Goodia, while small, CUbICal, prismatic or
acicular crystals are found in L1:p_aria, Priestieya, A mp hithalea , LatA,iogyne,
Coelidium, BiJrbonia, Lebeckia, V,borgia, A sp ala thus, Melolobium, Crotalaria,
Prioritropis, Lupinus, Argyrolobium, Gens'sta, A narlhrophyllum, Adenocarpus,
Calycotome, Peteeria, Sparii'Um, Ulex, Cytisus, and Hypocalyptus. Among
the Trifolieae Parochetus is the only genus in which ordinary solitary crystals
are absent; to~ether with crystals of this type small crystals occur in the
meSopbyll or epIdermis. In the Tribe Loteae" ordinary solitary crystals have
likeWISe been demonstrated in all the genera with the exception of Helmin-
tlwcarpum; in certain species of Lotus and Dorycnium these crystals attain
a specially large size and have a distinct styloid-like shape j crystalline grains
have also been observed in certain species of A nthylUs, SecuTigeTa, Lotus and
Hosackia. Lastly, ordinary large solitary crystals have been recorded also
in the genera of the Vicieae· in species of C,:ce" Vicia and Lathy,.us small
crystalline bodies are found. '
Cells filled with rod-shaped crystals (cf. p. 266) have recently been observed
inof the palisade.tissue in the folloWlng species: among the Podalyrieae, in species
Isotrapis, Dal)1'es,,°tJ, Pultenaea, Latrobea and Eutaxia; among the Genisteae,
apparently (see Schroeder) in species of Platylobium, Bossiaea, Templet<mia and
H01I8(;1; among the Trifolieae, only in Ononis spe&iosQ, Lag. and a few species
of Trifolium; among the Loteae. in species,of Anthyllis, Cytisopsis, and Lotus
(L. trichoca,pus, Lag.); among the Vicieae, in species of Vi£:ia (V. Gera,di. Vill.).
Lathyf'US (L. a"reus, Stev.), and Abrus (A. Schimperi, Hochst.); among the Dal-
bergieae, also in Dems elliptica, Benth. (according to Quanjer).
To the enumeration (on p. 266) of species having large solitary crystals
in the epidermis of the leaf we may add Abrus precatorius, L. (Tribe Vicieae).
In this species both the upper and lower epidermis contain small groups 01
2-] (mostly 4) cells, each of which encloses a rod shaped crystal; the latter
4

is embedded m the thickened inner wall of the epidermal cell with its longi-
tudinal axis parallel to the surface of the leaf.
Of special features it remains to mention: (a) sphaerocr_ystalline masses
of unknown chemical composition which have been observed m the epidermis
of the leaves (for the most part in herbarium material) in species of the genera
Anagyris, Piptanthus, Thetmopsis, Cyclopia, Podalyria, Chorizema, PhyUota t

P1dtenaea, Latrobea, Eutaxia,· Dillwynia (Podalyrieae), and of the genera

H()1)etl, Aspczlathus, C,otalaria and Argyrolobium (Genisteae) j (b) small bodies
resembling indigo or indican have been found in the mesophyll of the dried
leaves in species of Melclobium, Crotalaria and P,ioritropis (Tribe Genisteae),
in species of H ymenocarpus, H elminthocarpum, Lotus and H osackia (Tribe
Loteae), and in species of Lens and Lathyrus (Tribe Vicieae); (c) lastly, sub-
stances resembling saponin occur as shapeless masses in the epidermal cells
of species of Aspalathus (Tribe Genisteae).

• For the occ:urrence of sphaerltes and clustered crystals i.o the COlbryo o{ the Papiliooaoeae, see
Caldarcra, in Atti Accad. Giocoia Sc. nat., set. 4, vol. xi, 1898 (abstr. in Just, 1898, ii, po, ~u) i
also Lind.inger~ Diss•• 1903, p. 10.
 ADDENDA--LEGUMLVOSAE

In the Tribes recently investigated the ordinary type of clothing hair

is again in the form of uni!er'iate trlchomes, consisting of three cells,. viz.
(a) a frequently enlarged epidermal cell, serving as a basal cell; (6) a short
neck· or stalk·cell, which is occasionally distinguished by having special
contents and by being suberized; and (c) an elongated terminal cell,. of
variable character. The terminal cell shows either uniform' or unilateral
thickening or is provided with a spiral thickening band, which mostly
however, makes only one or two turns; occasionally the terminal cell is
band-shaped and twisted, while in Abrus it bears '9. characteristic awn-like
process. The longitudinal wall of the end·eell frequently bears verrucose
or peg-shaped irregularities, sometimes resembling papillae, e.g. in certain
Podalyrieae (species of Thermopsis and Baptisia), Genisteae (species of Lupj'n,us)
and Trifolieae (species of Trigo nella, Medicago, Melilotus and T,ifolium).
The terminal cell of the trichomes found in Hosackia pubeTula, Benth. and
H. st,igosa, Nutt. is unevenly thickened, the thicker parts of the wall being
provided with two Of three peg-like protuberances, which project into the
lumen of the eeH. Socket~like prominences, formed by the subsidiary cells of
the hairs, are found in certain Trifolieae (species of Trifolium), Loteae (species
of Hymenocarj;us, Anthyllis, Dorycnium, Lotus (especially Tetragonolobus) and
Hosackia) and Vicieae (species of CieeT, Vicia and Lathyrus).
Deviations from the ordinary type of three-celled clothing hairs have
been recorded as follows: (a) among the Podalyrieae, in the genera GompJw.
lobium and Daviesia; the former has simple thick-walled trichomes, diVIded
either by a single thin transverse wall, which segments the hair into two equal
halv'es, or by several thin transverse walls; Daviesia has unicellular trichomes,
which, however, show' transitions to the papillae present on the epidelll1al
cells; (b) among the Genisteae, in Bosstaea scolopendria, Sm., which has
uniseriate triehomes, composed of 4-5 relatively short cells; (c) among the
Trifolieae, in Ononis, in which uniseriate trichomes are of general occurrence;
they are mostly formed by 4-8 cells of, approximately equal length, and are
generally narrow and flagelliform, more rarely conical; in all probability,
however, they may be derived from the glandular hairs which are J found In
this genus (regarding this point, see G. Fischer), these hairs having long uni·
seriate stalks; (d) lastly, among the Vicieae, the hairs which are intercalated
among the external glands in the extrafloral nectaries of the species of Vicia;
these hairs consist of a high basal cell and an elongated terminal cell, which
has relatively thin walls and is occasionally subdivided,by a transverse wall.
A characteristic type of three-celled clothing hairs has recently been
observed in the form of tricbomes with a distinctly two-armed terminal cell,
the arms being of equal or unequal length ; these occur in species of Oxylobium,
Clwrizema, MiTbelia, Jacksonia, Pultenaea and Dillwynia (Tribe Podalyrieae),
and in species of H DVea, Lotononis, Lebeckia, A spalathus, Buchenroedera,
Crotalaria. (rare), Genista (rare), Calycotome, Erinacea and Cytisus (Tribe
Genisteae), while the hairs in certain species of P,iestleya, Aspalathus and
Genista (Tribe Genisteae) only' show a tendency to develop this structure.
External glands are of general distribution among the Trifolieae and
Vicieae. Among the Genisteae they are found only in Adenocarpus and Melo-
lobium, and among the Loteae only in Securigera, while they are completely
wanting in the Podalyrieae. The glandular hairs occurring in the genera
Parochetus, Trigonella, Medicago, Melilotus pro parte and Trifolium (Trifolieae)
are small and have short stalks; as a rule they consist of (a) a small basal
cell, (b) a short, uni- or bicellular stalk which is more or less demarcated,. and
(c) an ellipsoidal or club-shaped head, which is elongated to a varying ex.tent
and is subdivided by horizontal walls and often by vertical walls as well. On
the other hand the glandular hairs found in Ononis, and in Medicago lupfdina, L..
 A DDENDA--LEGUMINOSAE
M. prostrata, Jacq., and M. scutetlata, All., mostly have long stalks and ace
visible to the naked eye; the stalk is uniseriate,·the component cells showing
a gradual decrease in bulk, breadth and thickness of wall towards the upper
end, while the multicellular ellipsoidal head is divided by longitudinal and
transverse walls; the hairs of this type are accompanied by other external
glands which have shorter stalks, but otherwise vary in structure. Among
the Vicieae, Cicer bas glandular hairs wruch are visible to the naked eye;
these trichomes have a long uniseriate stalk composed of 4-6 cells, and a
head, which in most cases comprises four multicellul,ar tiers. The remaining
Vicieae only have small external glands with a short stalk and a more or less
sharply demarcated head, which usually consists of two or more cells and is
club..shaped; only in rare cases is the head elongated and tubular (species
of Lathyrus and Pisum), or composed of four cells and peltate, the shield
having a lobed margin and showing excentric insertion on the stalk (Lathyrus
hi,sutus, L.). The external glands found in the genus Adenocarpu5 (Genisteae)
are confined to the ovary and the fruit; they are multicellular glandular
shaggy hairs, columnar in shape and broadened in a capitate manner at their
apex. The unicellular glands of Melolubium (Tribe Genisteae) consist of a short
stalk and a globular head; the base of these hairs is inserted either directly
in the epidermis or at the apex of a conical pedestru, composed of parenchyma.
The multicellular emergences occurring on the different parts of the axis in
the monotypic genus Securigera (Loteae) are recognizable with the naked eye;
their apex bears a cell containing secretion.
The extrafloral nectaries found in the species of V icia are formed by
shortly italked glands intermingled with ianomalous forms of clothing bairs ;
they occur in species of the section Euvicia and in V. Mthym'ca, L. The honey·
secreting organs discovered by Delpino (Malpighia, iii, 1889, p. 345) on the
lower side of the leaves in Glyc-me sinensis have not yet been closely examined.
For the structure of the spiny rachis of the leaf of Hal1.'modendron and Cara·
gana, see -Mittmann and Lothelier respectively; for the anatomy of the stipular
spines of Robinia, see Mittmann. We may add that the anatomy of the spiny
stems of Ononis is dealt with bv Mittmann. that of the stem-spines of Ulex and
Genista by Lothelier, n. cc. ~

3. STRUCTURE OF THE AXIS. New investigations on the structure of the

axis have been published especially by Prenger and Hubner (Podalyrieae 1),
by Schroeder Cohn, Winkler, Levy, H. Schulze, W. Schulze and Rauth
7

(Genisteae '), and by Streicher (Vicieae 3), 11, ce.

The general features presented by the structure of the 'wood in these
Tribes are identical with those which have been previously recorded. To
the statements made on pp. 272, 273 regarding the structure of the vessels
we may add that sieve-like structure of the bordered pits is found also in
species of Argyrolobium, Hypocalyptus and Laburnum, and spiral thickening
of the pitted vessels also in species of Amphithalea, Anagyris, Argyrolobium,
Coelidiu1tt:, Erinacea, Laburnum, Lathriogyne, Lebeckia, Liparia, Lotononis,
PeUe,ia, Platylobium and Priestleya. Another noteworthy feature is con·
stituted by the broad medullary rays of the wood in Lebeckia, microphylla,

1 Species of Anagyn's, Piptanthus, Tlur1Jlopsis, Baptisia, Cyclopia, Podalpn'a, h'nuk),sema,

Osy/qIJ;um, CluW';:;ema, Mirbdia, Gastrolobium, Pullmaea, Lalrobea, Eu/aria., -Dil/wynia.
2 Species of Liparla, Pries/ieya, AmpMllla/ea, Lalhrio8Jme, CoeHd~'um, Platylohium, Bossiaea,
Templetonia, HUflea, Goodia, Boro()nia, Rafnia, Euch/ora, LotolUmis, ./\ol"i4, Lekdeia, Viborgia,
CtWalarla, Priorilropis, Aspalalhus, BW:MlIroedera, lJlelo/obiulll, DieM/us, If17landia, Lt~iinus,
A~/obium, Genis/a, Admocarpus, Calycotome,' Labunlum, Pelteria, SpartiulIl, Erinatea, Ulex,
U,,}tKal7lhu, LDtidigrs~'a.
_
3 Speci~ of Cictr nng Almer.
 ADDENDA--LEGUMINOSAE
Eckl. et Zeyh.; these rays arise by the fusion of several medullary rays in
the outer part ot the secondary wood.
To the last paragraph but two on J? 274 we may add, that a tier·like structure
of the wood is Iound also in Pongamla glabra, Vent. (Quanjer); to p. 274, last
paragraph but one, that according to Saupe the wood quite generally has a very
loose texture in the Phaseoleae (also in Strongylodon, according to my own observa-
tion), the ground.work being formed by thin-walled wood-parenchyma in which
groups of wood-fibres are inserted; this structure is what we should expect in
view of the fact that the members of this Tribe are lianes.
Regarding the development of the cork the following new facts have been
published (d. p. 276).
Epidermal development of the cork is found in specie$ of Oxylobt:um (Tribe
Podalyrieae), El'inacea and Loddigesia (Tribe Genisteae), Abrus (Tribe Vicieae),
and according to Oberlaender also MYl'oxylon. The cork develops in the first
layer of primary cortical cells in species of Podalyria and Gastrolobium (Tribe
Podalyrieae), Lipa'liQ, Priestley a, Lath,iogyne, Codidium, Hovea, Goodia, Lotononis,
Cl'otafaria, PriOYitropis. Melolobium. Calycotome, Spal'tium and Hypocalyptus (Tribe
Genisteae); in the second layer of the primary cortex in species o:f ihermOPsis,
Dfllwyftia and Eutaxia (Tribe Podalyrieae)~ Amphithalea and Buchent'oedct't.J (Tribe
Genisteae); in the second to fourth layer in species of Puttenaea (Tribe Podal~eae),
Argyrolobium and Labul'num (Tribe Genisteae); in the middle of the primary
cortex in species of Anagyris and Piptanthus. (Tribe Podalyrieae); in a region of
the primary cortex situated near the pericycIe in Cieer (Tribe Vicieae); in the
innermost layer of the primary cortex in Halimodendron argenteum, DC. (according
to Jonsson). Pericyclic cork~development is found among the Genisteae, also in
the genera Boyboma, Lebeckia. Viborgia, Aspalathus~ Adenocarpus, Petleria and
Ulex.
For the development of the cork on the furrowed stems of the leafless Papi-
lionaceae, see the respective papers by Ross, &c., which have already been cited
in the earlier part of this work; also Levy and Roth,loc. cit. At this point we may
also refer to the development of a mucilaginous cork in Halz"modcndron argenteum,
DC•• on the basis of Jonsson's description'; the cells formed by the phellogen
become rounded off and filled with a mucilaginous substance, which results in
their ultimate detachment. Lastly, in a few species of OxyloMum (0. a,borescens,
R.Br. and O. l'etusum, R.Br.) Damm met with a '.cuticular epithelium' (see
under Loranthaceae), beneath which cork-formation takes place in later stages
For the structure of the cortex in the,Jeafiess Podalyrieae and Gemsteae
a.nd the anatomy of the winged stems, see also the papers by BUrkle, Schroeder.
Rauth and Van Tieghem (1905).
The following new facts (see also Pitard, loco cit.) regarding the nature
of the perlcycle are supplementary to those previously mentioned (on p. 278).
The following genera have isolated groups of bast-fibres: among the Poda-
lyrieae, Anagy1'is, Piptanthus, Thermopsis, Baptisia, Cyclopia. Podalyria, Oxylo-
bium, Chorizema. M£rbelia, Gastl'olobium. Latrobea. Eutaxia. Dillwynia; among
the Genisteae, A mphilhalea, _Lathn'ogyne, Coelidium, T empleton£a, Goodia, Ra/nia,
Euchlol'a, Lotononis. Rothia. Lebeckia, Viborgia. Cl'otalaria, Prio'Yitrop£s1 Aspalathus.
Buchenl'oedera, M elolobium, DieMlus, Lupinus, A rgYl'oJoMum, Gem'sta, A denocarpus,
Calycotome, Laburnum, Petteria, SPa rtium , Erinacea, Ulex, Hypocalyptus, Loddi-
glsia; among the Vicieae, Cicero At certain points between the groups of bast-
fibres groups of stone-cells are occasionally found; in Bossiaea microphylla, Sm.
the isolated groups of bast-fibres become secondarily united to form a continuous
strengthening ring owing to the outermost part of the secondary cortex undergoing
subsequent sclerosis. A composite and continuous ring of sclerenchyma is present
in the peri cycle also in species of Bl'aehysema and Puitenaea (Tribe Podalyr!eae),
Lip aria , P"itstJeya, PJatylob£um and Hovea (Tribe Genisteae), and Ab"us (Tribe
Vicieae); an almost continuous and composite ring in Borbonia (Tribe Genisteae).
It is interesting that the secondary thickening layer (which consists of cellulose).
in the fibrous cells constituting the T·shaped groups of bast-fibres found in Genist"
 ADDENDA--LEGUMINOSAE
po(ygalae/olia, DC. and G. ,adiata~ Scop., disappears in later stages, probably as
the result of the action of a ferment dissolving cellulose; this thicKening-layer
is therefore of the nature of reserve-cellulose (see W. Schulze and Rauth, H. ce.).
Astely occurs in the stem of Trifolium (Belli). Cortical vascular bundles
'are found also in Borbonia lanceolata, L. and Viborgia obcordata, Thunb.
(according to Cohn), as well as in Genista canariensis, L., G. pros/rata, I.am.,
and G. sagittalis, L. (according to Van Tieghem), but by no means in all
Papilionaceae with winged stems (see Van Tieghem, loco cit., 1905).
To the list of anomalous structural features found in the axis (p. 279)
we may add the occurrence of secondary strands of wood and bast in the
primary cortex of Derris uliginosa, Benth. (Tribe Dalbergieae), according to
Perredes, and of successive rings of growth in llfucuna gigantea, DC., and
Strongylodon sp. (Tribe Phaseoleae).
The secondary strands of wood and bast in Den'is utiginosa are found in parts
of the axis which are about Ii cm. or mpre in thickness. They are present to the
number of one or two, and occuPY a peculiar position; a band of sclerenchyma,
containing pericyclic bast-fibres, IS situated on tbeir inner side. while on their outer
side there is another sclerenchymatous band, which forms part of the composite
and continuous ring of sclerenchyma in the pericycle and contains no bast-fibres.
In view of these facts it seems probable that the secondary strands of wood and
bast are developed in the primary cortex.-Pieces of the stem of Mucuna gigantea,
collected by Loher in the Philippines and having a diameter of 4-5 cm., show one
or two more or less completely differentiated rings of growth; the first (nonnaI)
ring of wood in this species attains I a thickness of 7-9 mID. As in the species
examined by H. Schenck, the secondary zoneS of growth originate in the pericyc1ic
parenchyma. InterxyJary phJoem has not been met with in M. gigantea, although,
It occurs in M. pt'uriens (see H. Schenck, Anatomie der Lianen, p. 164), con-'
trary to the earlier statement on p. 280.-The anomalous structure observed in
Strongylodon was found in a piece of the stem, 21- cm. thick, likewise collected
by Loher (Montalban, Philippines); an analysis of the flowers present on the
corresponding herbarium-material has shown that this stem belongs to the genus
St1'ongy/odon. The normal ring of wood in this case attains a thickness of 4 mm. ;
it is followed by two concentric rings of growth, and occasionally the rudiments
of a third ring are present. It may be specially pointed out, that the secondary
rings of wood and bast here apparently arise in the bast of the zone of bundles
situated immediately internal to the one in process of development. Regarding
the artificial production (by means of wounds) of extrafascicular vascular bundles
in Phaseo/us, see Schilberszky, loco cit.

II. CAESALPINIEAE (pp. 281-291).

I. REVIEW OF THE ANATOMICAL· FEATURES. Schizogenous secretory
cavities occur also in Daniella, Detarium, Kingiodendron, Oxystigma and Hard-
wickia, schizogenous in terxylarysecretory canals also in Daniella, K ingiodendfon,
Oxystigma and Prioria. Spicular fibres are present in the leaf also in Oxy-
,stigma M annii, Harms. Uniseriate clothing hairs with an elongated terminal
'cell are found in Scorodophloeus Z enkeri, Harms.
2. STRUCTURE OF THE LEAF. In the species which I have recently investi-
gated the epidermal cells of the leaf exhibit the following characters. Undulated
lateral margins with marginal pits and the occurrence of thin vertical walls
are distinctive of the species of Copaifera pro parte, Kingiodendron, Oxystigma
andPrioria. Development of papillae on both sides of the leaf isf.eund in Btn'd·
wickia bi-nata, Roxb., while in Detarium senegalense, Gmel. and Scorodophloeus
Zenkeri, Harms, the lower side of the leaf bears papillae. The speCles last
named also has gelatinized epidermal cells, stomata with subsidiary cel1s placed
parallel to the pore, and uniseriate clothing hairs with a few short basal cells
and an elongated terminal cell.
 ADDENDA-LEGUMINOSAE 905--
The following details may be mentioned regarding the secretory organs of
Daniella, Deta,,-um, H a'fdwc-ckia, K ingiodendron, Oxystigma and P'l'ion·a. In Daniella
thurijeYQ, Benn. and D. oblonga, Oliv., according to Guignard, there are more or less
elongated secretory cavities (but not canals, as Heckel assumes) situated in the
primary cortex, and secretory canals, which are found at the periphery of the pitlJ..
(although belonging to the wood), and also in the secondary wood. Similarly.Pnori4
copaile,a, Griseb., which I have examined, possesses (a) secretory cavities situatedin
the primarjcortex and more or less elongated in the vertical direction, (b) numerous
secretory canals in the secondary wood, and (c) secretory organs of the nature of
canals lying at the periphery of the pith and in its interior; those situated at the
margin of the pith maybe regarded as belonging to the wood, while those occupying
the interior of the pith are at least in part merely seeretory cavities, which are very
much elongated. According to ,my own investigation, Hardwickia binata, Roxb.
has secretory cavities only (situated in the primary cortex and sometimes also in
the pith and the parenchymatous pericycle). interxylary secretory canals being
absent; Kingiodendron pinnat"m, Harms has secretory cavities, which are only
slightly elongated and are situated in the primary cortex, and interxylary secretory
canals with rather wide lumina; in Oxysttgma M ann#. Harms there are {a} secretory
cavities of varying length in the primary cortex, (b) interxylary secretory canals, and
(c) intercellular secretory receptacles of the nature of canals, which are found at the
margin of the pith and occasionally penetrate into the wood. In Detarium seneRalense,
Gmel. there are secretory cavities in the primary cortex and in the pith, but likewise
no interxy1a.rY canals. The interxylary secretory canals are developed from the
cambium, like those of Copaijera ; their origin is schizogenous. In all these genera
secretory cavities are found also in the mesophyll.-For the secretory canals, cavi-
ties and cells of Epu'ua jalcata, see Courchet, loco cit.
The occurrence of an ethereal oil containing sulphur in the cortex of Scorodo-
pAloeus Zenkeri, Harms, which smells of garlic, is interesting from a systematic-
chemical point of view, since an oil of this kind has not previously been demonstrated
in any member' of the Leguminosae. The oil is present in the cortical parenchyma
in the combined form as a glucoside, and is accompanied by a ferment, by the
action of which the oil is liberated in the presence of water. The cells containing
the ferment are of relatively large size, and their contents assume a deep coloration
with Iodine dissolved in Potassium Iodide (Hartwich).
Regarding the occurrence of concentric vascular bundles in the petiole
of Cercis Siliquastrum, see Bouygues, loco cit. /
3. STRUCTURE OF THE AXIS. For the structure of the stem-spines of
Gleditsckia, see Mittmann, loco cit. 1

III. MIMOSEAE (pp. 29I-299).

2. STRUCTURE OF THE LEAF. For the stomatal apparatus in the phyllo-

dineous Acacias, see Porsch, loco cit.; fOf. the structure of the stipular spines
of the species of Acacia, see Mittmann and Lothelier, n. cc.
3. STRUCTURE OF THE AXIS. Sie\'e~like pits are found on the walls of the
vessels also in Albe'zzia Lebbek, Benth. (Ursprung).
Literature ~Leguminoslle! see p. 300): Hiihnel. Gerberinden, Berlin, 1880, pp. 139 and Lp
et seq.-[Stowell, Jamaica Dogwood, Therapeutic Gaz., N. S., ii, 1881, p. 16; see Just, 1881, ii,
p. 6g:l.]-Costantin, Tiges ~.er. et sout., Ann. sc. nat., ser. 6, t. x..."i, 1883, p. 51 et seq.-Darwin,
Bloom and distribution of the stomata, Journ. Linn. Soc., xxii, 188" especially p. 115.-Mittmann,
An at. d. Pflanzenstacbeln, Verb. Ver. Mark Brandenburg. 1889, pp. 4 6 , 50, 56, 57, 59 and 60.-
Johannson, Wenig bek. Rinden. Diss., Dorpat, 1891, l)P' '7 and 34 (Ptenxarpus and Erylhrina).-
Aufrecht, Extraflorale Nektnrien, Diss., Zurich, 1892, p. :H et seq. (Acacia lo_phantha).-Rarber,
Corky excresc., Ann. of bot., vi, 1892, pp. 160 and J 6s.-Hiller.Bombien. Gt'~rcya-Rinden.
piss., Dorpat, 1892, p. J:; et seq.-Schilberszky, Kiinstlich hervorger. Bild. sek. Gefassb., Ber.
deutsch. bot. Gesellsch., 1892, PP.424-32 and Tab. -xxii,-Lothelier, Epines, These, Paris, 1893.
_.._._---- _._--- ~

I Ursprnng's statement as to the occurrence of b,mlere,l pits on the wood-proscnchyma in

AfuUa bijllga. Gray requires verification.

 ADDENDA--LEGUMINOSAE
pp. II, 13. :18 and 31.-Radlkofer. Struct. anom. de la tige d'une Ug. vois. des Baulzini"a, Comptes
Tendus Soc. Hel".. Bite, 1893,P' loo.-Brandt, Wenig bek. Rinden, Diss.,Dorpat, 18~)4. p. 4S et seq ..
-Latour, Sen~, These, Montpellier, 189,.. rp. 18 and 45 et seq."":""Oberlander, Tolubalsam, Archiv
d. Phann., 232, 1894, pp. S9b-9.-[TogniQl, Stomi, Atti 1st. bot. Pavia, 1894.]-[Geoffroy, Robinia
Ni((IU, Ann. Inst. col. Marseille, iii, 1895, p. I et seq.]-Lutz, Gommose dans les Acacias, Bull. Soc.
bot. de France, 1895. pp. .,.67-7J.-(Belli, Endodermale periciclo nel Gen. Trifolium, etc., Mem.
A.ccad. Sc. Torino, set. a, v. 46, 1896, p_ 253-]-Knoblauch, Oekolog. Anat. etc., Habilitat.~Schr.,
Ttibingen, I8!J(i, p. 1 5 et seq.-Werner, Neue Drogen, Diss., Erlangen, 1896, p. 36 et seq.
(Euehrtsla).-[Scbneider, Senna, Americ. Druggist, 1897, n. 7.]-Schubert, Parenchymscbeiden,
Bot. Centralbl., 1897, iv, pp. 13-16.-Schwendener, Gelenkpolster von ,Mimosa pudica, Sitz.-Ber.
Berliner Akad., 1~7, xiv.-[Zancla, Aculei, Contribuz. 1st. bot. Palermo, ii, 1897, p. I' et seq.]-
Biermann, Olzellen, Dns., Bern, 1898, pp. 61, 62.-Geiger, Jaborandiblatter, Diss., Zurich, 1898,
pp. <f7, 48 (Stoarlzza).-&bwendener, Gelenkpolster von Fhaseolus, etc~, Sitz.-Ber. Berliner Akad.,
1898, xii.-seUe, Fabae Impigen u. Wurzel von iJerris elliptica, Diss., Erlangen, without date,
teceivtd 1898--99, p. 19 et seq.-Spanjer, Wasserapparate, Bot. Zeit., 1898, i, p. 59.-Boergesen og
Paulsen, Veget. dansk.-vestind. 6er, Bot. Tidsskrift, xxii, 1898-9, pp. 9 (Canavalia 06lu#folia, DC.)
and l03 (Leucaena glauca, Benth. and Acacia tortuQsa, Willd.).-[Driessen-Mareuw, Cortex Lokri,
Nederl. Tijdscbr. voor Pharm., 1899; abstr. in Just, ]900, ii, p. 1,5. )-Fucbs, C)'tisus Ada»;;; Ost.
bOt. Zeitscllr., 18~~, p. 74; and Sitz.-Ber. Wiener Akad., cvii, Abt. I, 1899, pp. 1273-92 and
l Tab.-(Greenish, Senna-leaves, Phann. JO\1rn. 1899; abstl'. in Just, 1900, ii, p. 2~.]-Hirsch,
Entwicklung d. Haare, Diss., Berlin, I~9, p. 27.-[Hua-Henri. Fcuilles des Cesalpiniees-Cyno-
metrees, Bull. Soc. Linn. Paris, o. s., n. 7, 1899, pp. 55, 56.]-Kobne, Papillen u. obers. Spaltofl'n.,
Ber. dentscb. dendrolog. GeseUsch., 1899. p. 51 et seq.-Leisering, Interxylares Leptom, Diss.,
Berlin, 1899, p. 35.-Nestler, Wasserausscbeidung, etc. (Phaseofus), Sitt.-Ber. Wiener Akad., cviii,
Abt. 1, 1899, p. 69Q et seq. and Tab.-Nestler, SekretttQpfen an d~n Laubbl. van PIuut4lus, etc.,
Ber. deutscb. bot. Gesellsch., 1899, p. 332 et seq.-Perrot, Tissu crible, These, Paris, 1899, p. 1::11.-
Pierre, Flore forest. de la Cochinchine, xxv, 1899 (Coroya).- Joh. Schmidt, Latlzyrus maritimus,
Bot. Tidss1crift, xxil, 1898-9, pp. 14S-68-"':"_Terraciano, Note anat. biol. sol1' Aucnynomuu indica,
Contribuz. 1st. bot. Palermo, ii, 1899. pp. 195 and 247 et seq.-Hiepe, Senna, Diss., Bern, 1900,
pp. 18-55 and Tab. iv, v.-Pantanelli. Pulvini motori di Robt'nia Ps(udai:acia, Atti Soc. natural. e
mat. Modena, ser. iv, vol. ii, 1900, p. 181 et seq.-Thomas, Feuilles sout., These, Paris, ;:900.-
Ursprung, Anat. 11. Jabresringbild. trop. Holzarten, Diss.,,'Basel, 1900. pp. 13-18 (species of A/zelia
and Albi%!;ia).-Bi.irkle, Innere Struktur d. B1. ll. and. Assimilationsorg. hei einigen austral. foda-
Iyrieen, Diss., Erlangen, 1901. 91 pp. (also in Funfstiick, Beitr_).-Cohn. Vergl. Untersocb. von
Blatt u. Axe einiger Genisteen-Gatt., Diss' l Erlangen, 1901 ; also Beih. z. bot. Centralbl., x, 1901,
pp. 52S-6J.-[Driessen-Mareuw. Radix Lawno (MilleJtia). Ned. Tijdschr. voor Pharm. etc., J901 ;
abstr. in Just, 1901, ii, p. :u.]-[Gretni5b, CaJs;a montana, Pharm. Journ., 1901, p. ~4; abstr_ in
Just, 1901, ii, p. 39.)~Haberlandt, Sinnesorgane, 1901, pp. 79 and 136 et seq.-Hill, Anatomy of
the stem of Dalbergia panicu!ata. Annals of Bot., 19°1, pp. 183-6.-Hohlke, in Beib. z. hot.
Centralbl., xi, 14)01, p. 41.-H iihner, .Blatt - u. A xenstruktur einiger austral. Podal yrieen- Gatt., Diss.,
Erlangen, 1901 j also Belli. t.. bot. Centralbl., xi, 19°1, pp. 143-2[7, with Tab.-Laobert, Cytims
Adami. Beih. z. bot. Central!>l., x, 190~, p. IH et seq.-Ledoux, Annt. compo des org. fol. chez les
Acacias, Comptes rendus, Paris, cnxii, 1901, Sem. 1, pp. 7l:l-5.-Levy, Blatt- "'1. Axensttuktur der
Genisteen-Gatt. Aspa!allzul tl. einiger verwandter Genera, Diss., Erlangen, 1901 ; also Beih. z. bot.
Centralbl., x, IC)OI, PP' 3f2--66.-Perrcdes, Bark of Robinia Pseudacada, in WeUcome Research
LaWnt. London, 1901, II pp., 4 pl.-Petefllen, Vedanatomi. 1901, pp.65-8.-Pitard, Pericycle,
These, Bordeaux, 1901, pp. 57-60 and 100, IOL-Prenger, Syst.-anat. Untersuch. von Blatt u. Axe
bei den Podalyrieen-Gatt. der nardl. Hcmispb.are u. des Kapgebietes, sowie den vier attStral. Gatt.
Brachystma, etc., Diss.. Erlangen, 1901, II I pp.-Rautb, VergI. ~nat. einiger Genisteen-Gatt.,
Diss., Erlangen, 1901, 58 pp.-Hugo Schulze, Vergl. Anat. d. Gatt. Lup;nus and A rgyr%bium ,
Diss., E!langen, 1901, H pp.-W. Schulze, Verg1. Anat. d. Genisteeo-Gatt. GeniI/a, Admocarpus
and Ca/ycptD11le, Diss., Erlangen, 1901, 59 pp.-Winlder, VergL Anat. d. Gatt. Cro/alan'a und
Priorilropu, Diss., Etlangen, 190[, 81 pp.:-Ha.rgag1i~Petrucci, Legnami, Malpighia, 1902, p. 302
et seq. (AiJauria, A/selia, Dialium, KoomjJassia, Pi"lhu%bium, Pqngamia, Siudora).-Bouygues,
Form. vase. anorm. du petiole, These, Paris. 1902, p. 57.-Clanditz, Blattanat. canar. Gew., Diss.,
Basel, 1903, pp. 53-,5 (Adtnoca'fUS a.nd Spartocytisus)l.-Damm, Mehrj. Epid. b. d_ Dicoty1.,
Belli. z. bot. Centralbl., xi, 1902, especially pp. 239. 240.-G. Flscher, Verg1. Anat. d. BI. bei den
Trifolieen, Diss., Erlangen, 1902, go pp.-Guignard, .Daniellia et lenr appareil secreteur. Journ. de
bot., xvi, 190 2 , pp. 69-97; see also Comptes rendus, Paris. cxxxiv, Sem. I, 190 2 , pp. 885-7.-
HartWich, Bubimbirinde, Apotbekerzeitung, Berlin, 1902. D. 40, pp. 339. 340.-Heckel, Daniel/ia,
Compt~ ttndus, Paris, cn:xlv, 1902, pp. 784-6.- Jonsson, Wust.enpfl., Acta Lund., :xxxviii, 1901,
pp. 28-38 and Tab. iv and v.-Knothe. Unbenebb. BI., Diss., Heidelberg, 1902, p. Ig.-Perredes.
Anat. of tbe stem of Derris ulz~'nosa, in Wellcome Research Laborat., London , J902, sep. (OPYI

l The statement as to the occurrence of unicellular clothing hairs in Adm«l1f'jUs is inconect

(d. S)tt. Anat., p. l68, footnote 2); the same is probably true of the statement that glandular hairs
OttUr in Sjar/ocylisul nU!Jigenus. The pelt ate hairs_recorded in Spa,.tocylis#1 are no doubt really
two-armed hain.
 AQDENDA--LEGUMINDSAE go7
10 pp., 9 pl.-W. Schmidt, Blatt- u. Samenstr Jktur bei den Loteeo, Diss., ,Erlangen, 190'; allO
Beih. z. bot. Centralbl., xii, 1902, pp. 4::14-82..-Schroeder, Anal. Unt.enuch. d. BI. no '"d. Axe bei
den Liparieae u. Bossiaeae, Diss., Erlangen 7 190:1; also Beih. '" bot. Centralbl., xi, 19Oa, pp. 368-
417.-S01ereder, Anat. Charaktere d. BI. bel den Podalyrieen u. Genisteen. Beih. z. bot. Centralbl.,
xii, 190::1, pp. 27g-88.-S01ereder, Le6cckia 1 re/amoides, Boll. de l'Herbier Boissier, ser. a, '1903,
pp. II7-ao.-Strticher, Vergl. Anat. d. Vicieen,'Diss., Erlangen, 1902; also ~ib. z. bot. Centralbl.,
:xii, J90l, pp. 483-538.-Tobler, Ursplung des peripher. Stam.m~webes, Pringsheim Jahrb•• I~l,
p. 133 et seq.:""[Armari, Piante della reg. medit., Ann. di Bot., I, 1903, p. 17 et seq. (Sparlium,
Reta",a, Genis/a, AnI~J'/lis).J-[Fin1ayson, Stem-structure of some lealless plants, Transact. and
Proceed. New Zealand Instit., 1903, p. 360 et seq. (CarmitAaelia).]-QuanJer, Anat. Boaw, etc.,
Natnurk.Verhandel., Haarlem, v, 1903 {Ptmgrtmiagiabra, lJerris e/lipticQ., Soj'ahispitla).-[Ramaley,
Pubescence of..lpecies of Astraga/us, Torreya, iii, 1903, pp. 38-40; abstr. in Bot. Centralbl •• xciii,
p. 199.]-[Rippa, Nettarii estranuz. della Poind'nna, Gill., Ball. Otto bot. Napoli, i, (903, pp.
431-3.]-Schoute, SteHirtheorie, 1903. p. In.-Theorin, Viixttrichom., Arkiv for Bot.• i, 1903-
p. I63.-Tuzson, Spiral. Struktnr d. Zellw. in den Markstr., Ber. deutsch. bot. Gesellscb., Igo,3.
p. ~76.-Chrysler, Strand plants. Bot. Gu., xxxvii, 1904 (LatR)'yus).-Col, Faisceanx, Ann. sc.
nat., ser. 8, t. xx, '904. pp. na-15.-Heckel et Cordemoy, Double appareil secreteur, Comptes
rendl1s, Paris, cxxxviii, 1904, Sem. I, pp. 57-9; see also [Ann. lnst. colon. Marseille, ser.2, t. ii,
1904, p. 7I et seq.; abstr. in Bot. Centralb1., xcix, p. 287l-Sussenguth, Behaarungsverh. der
Wiirzb. Muschelkalkpfl., Diss., Wiirzburg, 1904, pp. as-7.-UrsPrung, in Bot. Zeit., 1904, pp. 303
and 208.-Areschoug, Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39. n. 2. 1905. pp. 44, 45
and Tab. xxii (Adena1l//zera), PP' 49, 50 (Cynometra), pp. 54-6 and Tab. niii (lJerr';s). p. 61 and
Tab. xxiii-xxiv (Brownea), pp. 69-71 (Copaifera) , p. 74 (lJuman/nus), pp. 101-3 (/no(arput).
pp. 125, 126 (Bauhinia).-[Coun:het, EpeTUa jakata, Ann. lost, col. Marseille, xiii, 1905, pp.
121-.. 7; abatr. in Bot. Centralbl., eii, p. 434.]-Dnval, Jabotandis, 19°5, in Perrot, Travaux. iii.
1906, pp. 64-7o.-Gnignard, CordJ'la ajricana, Joum. de bot., 1905, pp. log-'4.-[Pabisch,
Fischgiftwurz., Zeitschr. Osterreich. Apotbekerver., 1905, pp. 975,976 ; abstr.- in Bot. Centralbl., ci,
p. 239- ]-Porsch, SpalloffuuDgsapparat, Jena, 190 5, pp. #. 100-17 and Tab. jjj.-Sarton, Rech.
expo sur l'anat. des pI. affines. Ann. sc. nat., ser. 9, t. ii, 1905, pp. 44-53 (LaI~}'rus).-Theorin,
Viixttrichom., Arkiv fOr_Bot., iii, D. -5, 19°4. p. 28, and iv, n. 18, 1905, p. 12.-Vadas, Robinienhoh.
in Tubeuf, Zeitschr., 1905. PP' 303-8.-Van -Tieghem, Meristeles, corticales, Ann. &c. nat., ser. 9.
t. i, 1905, p. a8.-Van Tieghem, Stele ailee de la tige des quo Legumin., Joum. de bot., 1905.
pp. 18S-97.-Weberbauer, Veget. d. Hochanden Perus, in Engler, Bot. Jahrb., xxxvii, '905, p. 6o,
et seq.-Danpbine, Rhizomes, Ann. sc. nat., ser. 9, t. iii, 1906, p. 327 et seq.-Kanngiesser. Spartiu1Jl
r(()parium, in Tubenf, Zeitscbr., 1906, p. 176 et seq.-~Mentz, Genu/a·Typen, Bot. Tidsskrift, xxvii,
2, J906, pp. 153-201 ; abstr. in Bot. Centralbl., d, p. 428.}-Nevenny, Tn"gtmella coeruled, Ber.
naturw.-med. Vcr. Innsbnlck, xxix, 1903-5, Innsbruck, 1!)06, p. 130 et seq.-[Rosenthaler, Rinde
\'on Pitku()/obium lngeminum, Zeitschr. Osterreich. Apoth.-Ver., 1906, pp. 147. 148.]~Stockard,
Nectar glands of Vida FalJa, Science, xxiii, 1906. pp. 204, 305.]-PiccioU, Legnami, Bull;Slena, 1C)06.
pp. 130, 145, 157-9, 162, 17 1-3, lj'6, 179. 180·-LFor additional literature', see p. I1io'.]

ROSACEAE (pp. 30r-3IO).

2. STRUCTURE OF THE LEAF. Among numerous investigated species of
Rosa papjllose differentiation of the lower 'epidermis has been found only in
R. rugosa, Thunb. (Parmentier); gelatinization of the inner walls of the
epidermal, cells is recorded by the same authority in many species of Rosa
and by Knoblauch in species of CliOorlia. In Acaena atlscendens, Vahl papillae
are differentiated both on the upper and lower epidermis (Clauditz); accord-
ing to Kohne, the lQ~er epidermis is papillose also in A melanchier rotundifoZia,
Dum.·Cours., Cotoneaster thymi/olia, Wall., P,unus Padus, L., P'Y'us amygdali-
/ormis, ViII., Sorbus sambuci/oUa, Roem., Spiraea alpina, 'Pall., S. bella, Sims.,
S. bracteata,. Zab., S. canes~ens, Don, S. ~xPa_nsa, Wall., 3:n~ S. trist~s, Zab.
Hypoderm 15 present also m Pygeum Wtghtsanum; Photsnta Notomana, on
the other hand, only has an epidermis composed of very large cells (Holter-
mann). In Rosa the stomata are confined to the lower side of the leaf (except
in R. berberifolia, Pall.), and develop according to the RanuncuIa<*>us type.
In Acaena adscendens the stomata are likewise surrounded' by ordinary epi-
dermal cells.
Oxalate of lime. In Rosa Parmentier met with sphaerites side by side
-with clustered and solitary crystals, and Bargagli.Petrucci found silica-bodies
t in the wood-parenchyma of Parastemon urophyUus, DC.
 ADDENDA--ROSACEAE
ro the statement's regarding the hairy covering we may add :-Structures
resembling stellate bairs and external glands with a head composed of numerous
cells occur in Spiraea sorbifolia, L. (Theorin). Small external glands with
a uniseriate stalk and a head composed of one or two cells are found also in
Geum and Potentilla (Brautigam). ClitJortia arborea, Marloth has Malpigbian
hairs (Madoth), while C. talcata, L. only has ordinary unicellular hairs. Extra-
floral nectaries are present in the genus Grifjonia, a member of the Chryso-
balaneae (Mattei).
The petiole in many species· of Alchemilla contains three concentric vas ..
cular bundles, each provided with a pi th and surrounded by a typical endodermis
(' faisceaux concentriques vrais '). The petiole of Sanguisorba ~anadensis
.includes B-<) isolated bundles, One of which has the same structure as those
of Alchemitla, while the remaining bundles are hemiconcentric Cfaisceaux
hMliconcentriques '), i.e. each of them consists of an arc of wood arid ba.c;t,
which is likewise surrounded by a typical endodermis (Bouygues).
For the structure of the integumental prickles of Rosa and Rubus, see Mittmann
and Lothelier, n. ce.; these authorities also discuss the anatomy of the spiny
branches of Cf'ataegus, Cydonia, pYI'US, PYl'acantha and Prunus; regarding the
integumental prickles of Rosa, see also Duchartre, loco cit.
3. STRUCTURE OF THE AxIS. The cork develops in the pericyc1e also in
kI argyrocarpus, ClifjQrtia and Polylepis, which belong to the Poterieae
(Bouygues).
Burgerstein's recent investigations on the structure of the· wood in the
Pomeae (see the note on p. 308) and in numerous species of P,unus have
furnjshed the following facts. Among the Pomeae distinct spiral thickening
of the walls of the vessels is found in Amelanchier, Aronia, Chaenomeles,
Cotoneaster, Cydonia, Eriobotrya, Mespiltls, Micromeles, Photinia, Rhaphiolepis,
and Sorbus, while the spiral thickening is not well marked in Hesperomeles,
Osteomeles and Peraphyllum, and is wanting in Chamacmeles, Crataegus, Malus,
Pyrus, Pyracantha, and Stranvaesia. The wood in aU the Pomeae contains
scattered vessels, and in the ~()urse of each annual ring the latter showed
a gradual decrease in size. The medullary rays vary from one to three (mostly
one or two) cells in breadth j only in Mespilus are they from one to five cells
broad. The cells of the medullary rays are rather low in Chaenomeles, Cydonia,
Mespilus and Micromeles, and relatively high in Cotoneaster, E'Yiobotrya,
Hesperomeles, Photinia and Sorbus. \Vith reference to the structure of the
wood in PTunus, we may mention that in all the investigated species the vessels
show a spiral thickening band, and that they are either scattered or arranged in
annular zones; in the latter case the zone of vessels belonging to the spring-
wood appears as a rather sharply marked ring of relatively large pores.
Regarding the structure of the cortex we may add that Pitard records
a pericycle, comprising isolated groups of bast-fibres with intervening un-
lignified hands of parenchyma of varying width, in species of AmeZanchieT,
Cotoneastet, Crataegus, Eriobotrya, Kageneckia, LindZeya, Osteomeles, Photim'a,
Prinsepia, Prunus (sensu latiore), Pygeum, Pyr'lts (sensu latiore), Quillaja,
Rhaphiolepis, Rosa, Stranvaesia, Stylobasium and Vauquelinia. According to
the same authority pericyc1ic sc1erenchyrna is completely absent in Canotia
holacantha and Pterostemon mexicamls.
Literature y Costantin, Tiges aero et SOUL, Ann. sc. lIat., !>er. 6, t. xvi, 1883, p. E.9 et seq.~
Buchenau, Ausscheid. einer kryst. (lrg. Sanre im Holzkorper e. Eberescbc, Festschr. Vcr. f. N:lturk.,
Kassel, 1886, pp. 3i-9.-Macchiati, Nettarii estrarlor. delle Amygd., Nuovo Giom. bot. Ital., 1886
pp. 305-i.-Nanke, Dikotyle Holzpfl., Diss., Konigsberg, 1886, p. 40.-Keller, Luftwuf'''., Diss"
Heidelberg, 1889, pp. 36, 37.-Mittmann, Pflanzenstacbeln, Verb. Ver. Mark Brandenburg, 1889.
pp. 4 1 and 6j' et seq.-Barber, Corky eltcresc., .<\nn. of Bot., vi, }892, p. 16s.-Dnchartre, AiguiJ10ns
dll Rosa uriwl, Revue gen. de bot., 1893;Pp. 5-1 I .,.._Lothelier, Epines, These. Paris, 1893, pp. 17.36
 ADDENDA--ROSACEAE 909
and 38.-Orth, Al1atomie d. Gatt. Polen/ilIa, Diss., Kiel, 1893, 33 pp.-{Tognini, Stomi,. Atti 1st.
bot. Pavia, 189....}-[Bastin, Cherry barks, Americ. Joam. Pharm., 1895, pp. 435 -and 595 et seq.]-
[Burgentein, Gatt. d. Pomaceen, Wiener Illastr. Gartenzeit., 1896.]-Knob1aach, Okolog. Anat. etc.,
Habilitat.-Schr., Tiibingen, J 8~, p. I I et seq.-[Parmentier, Especes crit. ou lit., Mem. Soc. d'etnula-
tion Doubs. 1896, p. 337 et seq. (Crataegus).]-Brautigam, Anat. Cbaraktere der ROsaccen-Butarde,
Diss., Erlangen, 1897, 56 pp., 3 Tab.- Parmentier, Rech. anat. et taxon. sur lei Rosien, Ann. sc.
nat., ser. 8, t. vi; 1897, pp. 1-175, 8 pI.; [see also Comptes tendui, Congres des Soc. sav., 18gB,
p. 220.1-Schubert, Parenchymscheiden! Bot. Centralbl., 1897, iii, p •. 47~:-Banting, Cdtk tiline in
roots or some Rosaceous genera, ContrlbaL Eot. Lab. of PennsylvaDla, 11, 1898, pp. 54-65, pi. X.-
Hurgerstein, Holzstruktur d. Pomaceen, Sitz.·Ber. Wiener Akad., cvii, Abt. I, 1898, pp. 8-23 ; and
Xylotomisch-ayst. Studien tiber d. Gatt. d. Pomaceen, Jahresbericht Staatsgymnas. ii, Bezirk WieD,
1898. 35 pp.-Crepin, L'anatomie appliquee A la c1assificat., Bull. roy. de bot. de Belgique, xxxvii,
1898, pp. 7-15; nnd Idees d'un anatomiste, etc., loco cit., pp. 151-201.-Guffroy, Sorbus et hybrides,
Bull. Soc. bot. de France, 1898, p. 341.-[Kayeriyana, Discoid nectary of Jap. cherry leaves, Bot.
Magaz. Tokyo, :r.ii, 1898, pp. 281-4 ; and Disc-shaped glands in the leaves of Prunus PseudO€eraslls,
loco cit., xiii, 1899, pp. 316-18.)-Spanjer, Wasserapparate, Bot. Zeit., 1898, i, p. fi5.-Boergesen og
Paulsen, Veget. dansk.-vestind. Oer, Bot. Tidsskr., xxii, 1898-9, pp. 20,21 (Chrysobalanus ICQco,L.).
-Burgerstein, Xylotomie d. Pruneen, Verh. k. k. zoolog.-bot. Gesel1scb. Wien, 1899, sep. copy,
:; pp.-Kohne, PapiUen u. oben. Spaltotfn., Mitteil. deutsch. dendrolog. Gesellsch., 1899. pp. 56,57.
-Macchiati, Nettarii estranuz. del Pm"us LaurOctraS1ts, Bull. Soc. bot. Ital., 18 99, pp. 1#-7.-
&uygues, Anat. comp. de la tige et du petiole des Ruhees et Rosees, Act.- Soc. LlDn. de Bordeaux,
Iv, 1900, pp. xxxii-xl; Jd., Pol:ystelie d~ geme TAldum~'lIa, loco cit., pp.lvii!-lix; Id., !ige et pet.iole
da Neuratfa potumbellS, loco CIt., pp. ltx, Ix; ..d., Perldenne de quo Potenees, loe. Clt., pp. XCVI-C;
(d., Poly:;telie part. de Sanguisorba canadensis, loco cit., pp. cxlvii-cxlix.-Paulesco, Strnct. annt.
des hybridcs, These, Geneve, 1900, pp. 38 and 64 (~T".·osa, Sorbus).---Bouygue3, Polystelie, Act.' Soc.
Linn. &rdeaux, hi, 1901, pp. xxxv-xl.--Petersen, Vedanatomi, 1901, pp. 5S-64.-¥itard, Pmcyc\e,
These, Bordeaux, 1901 , pp. 65, 66, 68, 97-Ioo.-Barg1!.gli-Petrncd, Concrez. silicee, Malpighia,
190z, p. 26 et ieq.; and Legnami, loc, cit., p. 300 (Paraslemon).-Bouygues, Petiole, These, Paris,
1902, pp. 55 et seq.- and 99.-Clanditz, Blattanat. 'canar. Gew., Diss., Basel, ,190 2 , pp. 46, 47
Prunus ) .-Knothe, Unbelletzb. BI., Diss., Heidel berg, 19°2, pp. 17-19. -Mardner, Phanerog. Vegetat.
d. Kerguelen, Diss., Buel, 1902, pp. 22-5 ~Acama).-lMattei et Rippa, Nettarii estranuz. di aleune
Crl!obalanee, Bull. Orto bot. Univ. Napoli, i, 1902, pp. 286-91 i abstr. in Bot. Centralbl., .xcv,
p. 580.1-f Arman, Piacte della reg. medit., Annali di Bot., i, 1903, p. 17 et ieq. (P(1terium).)-
Freiden1'e!dt, ~t. Bau d. Wurzel, Bib1.bot., Heft 61,1904, pp. 5~, s6.-Siissengutb, Behaarungsverb.
der WUrtb. Muschelkalkpft., Diss., Wiirzburg. 1904, pp. 27-33.-Theorin, Vaxttrichom, Arkiv for
Bot., iii, no 5, 190+, p. 16.-Weberba~r, Vegetat. d. Hochanden Petus, in Engler, Bot. Jahrb.,
xurii, 1~5, p. 60 et_ seq.-Marlotb; Cli_fforlia, in Engler, Bot. Jahrb., xxxix~ 1906, p. 319.-Piccioli,
Legnanu, Bull. Siena, 1906, pp. 127, 130, 157, 16j.-5.-Holtermann, Eintluss des Klimas,etc., [907,
p. II!) (Pyteu1tl, PAo#"ia),-[For additional literature, see p. 1172.)

CROSSOSOMATACEAE.
Two features are important for the diagnosis of the Order and in support
of the view of a Rosaceous affini ty, viz: the presence of typical bordered pits
on the wood-prosenchyma, and the occurrence of scalariform perforations
(with few bars) or of malformed perforations of the-same type in the neigh-
bourhood of the primary wood, side by side with the simple elliptical perfora-
tions found in the secondary wood. The stomata are surrounded by several
ordinary epidermal cells. Trichomes, oxalate of lime, and internal glands are
~~. .
The structure of the leaf has been examined in Crossosoma Bigelovii,
Wats. (Parish, :n:-'10, Colorado) ~and C. calitornicum, Nutt. (Franceschi, n. 41,
Guadalupe) 1. The leaves in· both species show centric structure. ,Palisade
tissue is found on both sides of the leaf, and stomata are present in both upper
and lower epidermis. The outer waJls of the epidermal cells are stronQly
thickened. In C. calijornicum the epidermis contains sphaero-crystals, which
are insoluble in alcohol, but soluble in Caustic Potash and Rau de Javelle.
Peculiar small boclies of a. dark colour are frequently found in the .palisade
tissue in.hoth species; their nature is not known. The larger veins ar-e accom-
panied by a little sclerenchyma only in C. Bige/.ovii.
The structure of the axis could only be investigated in C. Bigelovu I.
, I am indebted to Geheimrat Engler for the materials for this inve'Jtigatioo.
 ADDENDA-CROSSOSOMATACE~E

Regarding the worrd.we may add that: (a) the vessels have very small lumina ;
(b) the wood.;fibres have thick walls ·and narrow lumfua; (c) the medullary
rays are narrow, con~tin~ of a single row of cells; and (d) the .walls of the
Vef;Se]s bear bordered pits m contact wiih parenchyma of the medullary rays.
The pericycle contains isolated bast·fibres. The cork is for the most part
composed of cells with thin walls, but also includes uniformly sclerosed cells.
Literature: Engler, CIOSiOSOmataceae, in Naturl. PfllUlzenfam., Nachtr. zu ii-iv. Teil, 1897,
P. I8S·

SAXIFRAGACEAE (pp. 310-320).

2 A. STRUCTURE OF THE LEAF. For the water-pores, see also Gardiner.
loc. cit. "
To the previous remarks on the hairy covering, we may add that fimbriate
appenda'kes, similar to those present on the stam4'lodes, are found on the foliage-
leaves, as well as on the sepals and petals in Parttassia; they occur in varying
numbers in the different species. Both the epidermis and the subjaCent layer
of cells participate in the formation of these structures, which do not show
any sign of swelling at their ends, although they secrete mucilage in early
stages (Dutailly) 1.
The petiole of Saxifraga sarmentosa resembles that of,AlchemiUa in con-
taining three steles, each of which is surrounded by a typical endodermis;
S. crassifolia shows similar features. Other species of Saxi/raga, such as
S. dentata, S. hirsuta and S. lasiophylla, have at least one hemiconcentric
vascular bundle, enveloped by an endodermis (Bouygues).
3 A. STRUCTURE OF THE AxIS. According to Schoute and Jeffrev
respectively, polystely is found in thenodes of the stem in Parnassia palust,is I .
3 B. STRUCTURE OF THE AxIS. The pith in Ribes rubrum, R. Grossularia.
&c., contains peculiar cavities; in early stages their place is occupied by
relatively large thin-walled cells containing clustered crystals, and the develop-
ment of the cavities is mainly due to the collapse of these crystal-cells (Kassner).
In Roussea simplex the primary cortex includes a ring of stone-cells bordering
directly on the endodennis. In Brcxia chrysophylla the pericycle comprises
a composite and continuous ring of sclerenchyma, while in other species of
Brexia the ring is interrupted (Thouvenin and Pitard).
For the structure of the spinous organs found in the species of Ribes, see
Lothelier and Mittmann.

D. THE ANOMALOUS GENUS PENTHORTJM (ApPENDIX TO p. 319).

The followin~ statements regarding the anatomy of Pentho,um are based
on Van Tieghem s and my own observations. The vegetative and specially
the reproductive branches bea{ glandular sh~ hairs with a multiseriate
talk, merging a:pically into a more or less distinct secretory head. The inner
portion of the pnmary cortex contains large intercellular spaces. Small groups
Df bast-fibres are found in the peri cycle, but there is no secondary hard bast.
The wood is traversed by narrow medullary rays, which are one or two rows
of cells in breadth. The vessels of the. secondary wood have small lumina,
.and are proVlded with small bordered pits (also in CO!ltact with parenchyma
of the medullary rays) and scalariform perforations (with numerous or very
nomerous bars). The wood-fibres, thou~h thick-walled, have rather wide

1 For details regarding the structure of the leaf, stem,-and:root in .Ptzrnass;a, see Van TieghcDl,
loe. cit.
 _
ADDENDA--SAXIFRAGACEAE 911

lumina and bear small but distinct bordered pits. The cells of the pith have
cellulose-walls, which subsequently become lignified, especially at the margin
of the pith. Oxalate of lime is present in the form of cl~tered crystals, which
occur in the pith and primary cortex, as well as in the mesophyll. The petiole
contains an arc-shaped vascular bundle. The leaf is bifacial, the stomata
being confined to the lower side. According to Van Tieghem. the vascular
system of the root is pentarch, the groups of soft bast being supported by
bundles of fibres 1.
Literatnre: Uhlworm, Entwicklungsgesch. der Trichome. Bot. Zeit., ) 873, p. 820.-{Gulliver,
Rh~hides ip Hydrangea, 100m. R. Microsc. Soc. London and Edinburgh, iii, 1880, p ....... ; abstr.
in Just, 1880, ii, p. 149.]-Hohnel, Gerberinden, Berlin, 1880, p. 108 et seq. (Weinmannia).-
Gardiner, Water.glands in the leaf of Sa..xifroga crus/ata, Quart. Journ. Microscop. Sci., .xxi, N. S.,
1881, pp. 4°7-14, pI. niii,-Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883. p. 77
et seq.-Kassner, Mark, Diss., Basel, 188..... p. 16.-Mittmann, Pflanzenstacheln, Verb. Va. 'Mark
Br:mdenburg, 1889, p. 65.-Thollvenin, Appareil de soutien dans les tiges des Sa.xifrages, Bull. Soc.
bot. de France, 1889, pp. U5-33.-C. de Candolle, mflor; epiphylles, Mem. Soc. de phys. et d'hist.
nat. Geneve, 1890, vol. It1lppL, sep. copy, p. 12 et seq.-J.,E. Weiss, Korkbild., DeWcschr. bot.
Gesellsch. Regensburg, vi, 1890, pp. 61, 6:il.-Lothelier, Epines, These, Paris, 189~, p. 37.-
Boergesen, Arkt. pI. bladbygn., Bot. TidM=kr., xix, 1895. p. 2I!). et seq.-Hallier, it! Nataurkundig
Tijdschr. VOOT Nederl. [ndie, 1896, p. ~IO et seq.-Van Tieghem, ~enthore, Joam: d~ bot., 1890,
'pp. 150--4; and Ann. sc. n.t., ser. 8, lllC, 1899, pp. 371-3.-Van Tleghem, Parnassracees, Journ. de
bot., 1899, pp. 326-32.-Bouygues, Merist. vase. dans Ie petiole, Act. Soc. Linn. BordeaU%, lvi,
]901, p. lvii-Dutailly, Parnassia. Assoc. fran~. Ajaccio, IC)OI: i, 1901, p. £26, and ii, 190', pp.
• S7-7 1.-Petersen, Vedanatomi. 1901, pp. 56, 57.-Pitard, Pericycle, These, Bordeaux, 19°1, p. 70.
-Bouygues, Petiole, These, Paris, 1901, pp. 13, 17,66, 97 and I07.-Ge-rharo, Blo.ttauat. v. Gew.
d. Knysnawaldes, Diss., Basel, I90~, pp. 20,21 (Platyl"phtts).--Schoute, Stelartheorie, 1903,_P' 120.
-Col, Faisceaux, Ann. SC. nat., ser. 8, t. xx, 1904, p. J 16.-Freidenfeldt, Anat. Baa der Wunel,
BibL bot., Heft 61, 1904, pp. S6--61.-Theorin, Vaxttrichom., Arkiv for Bot.. iii, n. 5, 1904, p. 3.-
Areschoug, Trop. Taxt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2, 1905. Pp. 15.-6 (Brexia).-
Haydo-., Saxifrara I, Denkschr. Wiener Akad., l.xxvii, 1905, pp. 616-.12 and Tab. i.-Maheu et
Gillot, Ascidles des Saxifrages, JOUIn. de bot., I~O:)J p. 33 et seq.-Piccioli, Legnami, Bun. Siena,
1906, pp. J30 and 156.-[For additional literature, see p. 1172.]

CRASSULACEAE (pp. 320-324).

,
I. REVIEW OF THE ANATOMICAL FEATURES. Both clothing arid glandular
hairs of the shaggy type occur in this Order.
2 •. STJiUCTURE OF THE LEAF. Brenner records a two-layered epidermis
in Crassula portulacea and a twica] papillose epidermis in Sedum Hillebrandii,
Fenzl, while according to L. Koch Sedum dasyphyUum only has isolated
papillae; Brenner.states that in Cl'assula pOl'tulacea distinct papillae are only
developed when the plant is placed in a moist chamber. According to Mardner,
the stomata in Tillaea moschata, DC. have three typical subsidiary cells (con-
trary to what is found in T. muscosa) and are situated principally on the upper
side of the leaf. Water-pores, moreover, also occur In tbis species.
With regard to the differentiation of the mesophyll, it may be pointe~
out that in certain species of Sedum and Sempervivum the assimilatory cells
exhibit~a'stratifted arrangement, while the intercellular spaces are lined by
a membrane, which we, must regard as the metamorphosed outermost layer
of the cell~wall.
The clothing and glandular shaggy hairs mentioned above are found in
SemperVivum al'achnoideum and have a biseriate structure. The glandular hairs
are short and slightly swollen at the apex. The clothing hairs constitute the
cobweb-like covering on the leaves and are much longer; the two rows of

1 Van Tieghem':; statement (11. Cc.) that such bundles of fibres have ail yet been observed only
in the aerial roots. of parasitic Loranthaceae and not in terrestrial roots is incorrect. The phloem.
groups in the tetrarch vascular system of the root of Vitia f~a are likewise supported by hard bast
{Stt Kn" Wand(l.Cc:l, lvi, 1&84).
91 2 ADDENDA--CRASSULACEAE
cells of which they are composed are not absolutely~parallel, but are slightly
twisted. The lower part of the. body of the trichome in both kinds of -hairs
consists of long cells, while the cells of the upper part are shorter. In the
course of their development the clothing hairs pass through all the stages
shown by the l?landular shaggy hairs; in a certain stage they even secrete
ethereal oil, whIch functions as a glue joining together the neighbouring hairs ;
subsequently, as a result of extremely rapid growth in length, the bodies of
the hairs become intertwined with one another and in tbis way the dense
hairy felt present on the. rosette of leaves is ultimately formed (Dintzel). .
Literature: Corda, in Sternberg, Flora der Vorwelt. 1838, Anhang, p. lxii and Tab. A.-
[Henri, Knospen, etc., Verb. Ver. f. Rheinlande 11. Westfalen, 1850, p. is; and Wurzelfascm -yon
Seaum Telephiu"" ete., loco cit., 1860.]-Irmisch, Sedum nuuimum, Bot. Zeit., \8S5, p. '.9 et ,seq.
and Tab. 2 A.-Schwendener, Meehan. Priaeip. 187i, p. I48.-[Gatdiner, in Quart. JOUln. Microsc.
Sci., 1881, p. 407 et seq.J-Costantin, Tiges aero et sout., Ann. sc. nal, ser. 6, t. xvi, 1883, p. 79
et seq.-]ost, ZerklUCt. einiger Rhiz. u. Wurz., Bot. Zeit., 1890, p. 503 et seq. (Sea'um).-Matteucci,
Placche sugbero.se, Nuovo Giom. bot. It at, 1897, p. 236 et seq.-Schubert, Parenchymscheiden, Bot.
Centrubl. 1897, iii, p. 475.-ROdler, Aasiro.ilatot. Gewebel~. Diu., Freiburg i. Schw., 18g~.
p. 38_Brenner, Fettp6anzeD, Flora, [900, pp. 389-98.-Schlelchert, Xerophyten bei Jena, Naturwiu.
Wochenschr., 1900, P'450 (Sedum).-Clauditz, Bl&ttanat. canar. Gew., Diss., Basel, 1902, pp.•' , i3
(Selll~rvivu"z).-Knothe, Unbenetzb. B1., Diss., Heidelberg, 1901, p.-:ro.-Mardner, Phan.-V~t.
d. Kerguelen, Diss" Basel. J902, p~. 26, 21 (Til/aea).-r Arman, Piante della reg. medit., Aonali di
~t., i, 1903, P. I; et seq. (Sea'um).j-Dintu:l, Hure a. d. Blattsp. von Semptrvi1lUm aracltndUk"""
Osterreich. bot. Zeitschr., 1905. p. 2S4 et seq. and Tab. 'II, vi.

DROSERACEAE (pp. 324-328).

The tentacular glands of Roridula Gorgonias, Planch., which have recently
been examined by Fenner, are of diff~ent lengths; they have a multiseriate
stalk, which exhibits corresponding variation in length, and an ellipsoidal
head, in which the inner part is formed by a group of approximately isodia ..
metric cells, constituting a continuation of the cells of the stalk, while the
secretory epidermis is differentiated like a palisade; tracheids are wanting
alike in the stalk and in the head. The middle layer in the digestive glands
of Aldro'Uanda consists of two rather high cells, which form a stalk; the number
of central and peripheral cells in the glandular disc is occasionally, greater
than 4 and 8 (viz. 5-8, and g-I4)· Pores are present in the cuticle on the
secretory heads of Drosophy/lum and Drosera rotunds/olia (Haberlandt ,and
Fenner). The genus Byblis. which up to the present has been included among
the Droseraceae, but which has been transferred; and no doubt rightly so, to
the Lentibularieae by Lang, has digestive glands, which differ. considerably
from those of other Droseraceae; according to Lang, their structure is. analo-
gous to thatof the glands occurring in the genus Pinguicula (Lentibularieae).
Both sessile and lo~g-stalked glands ar~ found in Byblis .. ~e simplest t~
of the former conSIsts ·of three cells, VlZ. a basal, a median, and a temunal
cell; in many cases, however, the terminal cell unde:l~s division into quad-
rants followed by the appearance of four anticlinal w , so that a disc-shaped
glandular body IS developed, which for the most part consists of eight 'cells ;
the basal cell at the same time divides into two cells. In the fonnation of the
long-stalked glands the basal cell divides into two cells by means of a transverse
wall, and the upper of these cells grows.,out into a long unicellular stalk, While
the (lower) sister-cell undergoes further division; the terminal cell becomes
segmented to form a glandular disc, composed of 16 or 32 cells. The cuticle
of the glandular disc, both in the short- and .long-stalked glands, bears a
pore on the upper and lower side of each cell. ,A feature deserving special
mention is that the ~lands on the leaf of Byblis are associated with thti ~tomata
to form parallel longitudinal roWS which are sunk below the surface and alternate
 ADDENDA--DROSERACEAE
with one to three rows of cells, containing neithet stomata nor glands. For
further details regarding the glands, see the papers by Lang and Fenner, 11. cc.
Regarding the tentacular glands of Drosera we may add that the epidermal
cells of the head are provided with marginal pi~ which are occupied by proto-
plasmic processes; these constitute the perceptive organs for the mechanical
stimuli, to which the movements of the tentacular glands are due (Habedandt).
The two- and four-armed glands of Aldrovanda, which were discussed
under the heading of ' other types of hairs,' have two basal cells and a stalk,
consisting of a single layer of two to four cells (Fenner). For details as to the
structure of the sensitive bristles of Aldrovanda and Dionaea, see especially
Haberlandt, loco cit.
Although the stomata of the Droseraceae were previously stated not to
have any subsidiary cells (p. 326), it has since been shown that in the genus
Byblis, which Bentham and Hooker place among the Droseraceae, the pair
of guard-cells has a single subsidiary cell on each side of the stoma and
parallel to the pore (Lang).
As regards the. detailed anatomical features of the leaf of Drosophyllum we
may add that the mesophyll is composed of spongy parenchyma with Jarge lacunae;
the cells of this tissue contain numerous acicular crystals, which are soluble in
a solution of chloral hydrate~ as well as a few large crystals of oxalate of lime
(A. Meyer and Dewevre).-According to Lang, the epidermal cells in the leaf of
Byblis are elongated in the direction of elongation of the leaf. The subsidiary
cells of the stomata have already been referred to above. The epidermis is followed
by a sheath of two or three layers of cells which show a slight palisade-like dif·
ferentiation; on the inner side of this zone lies a starch·sheath, and within this
again a medulla consisting of cells with wide lumina and enclosing 3-5 collateral
vascular bundles; the phloem-groups: in these bundles are surrounded on all sides
by a sclerenchymatous sheath. The swollen apex of the leaf of Byblis contains
only a single vascular bundle, whlch is apparently concentric (with central phloem),
and is enveloped-by a tissue serving for the storage of water; the walls of this
tissue bear PIts having a faint border or exhibit spiral to reticulate striation;
on its outer side lies a starCh-sheath, followed by two or three layers of rounded
assimilatory cells. The tip of the leaf bears a few elevated water-pores, but there
is no epithema on their Inner side (Lang).-The following detaifs regarding the
structure of the leaves of Rot'iduJa, the upper surface of which is furrowed, are
based on Fenner's statements. The mesophyU is composed of loose spongy tissue,
the lower portion coptaining large lacunae. The vascular bundles of the veins
are supported by groups of sclerenchyma. The stomata are confined to the lower
side of the leaf.
The vessels in Byblis have simple perforations (Lang)~ Astely is found
only in the peduncle of Byblis (Lang).
The structure of the root of Byblis is norplal, the vascular system being
triarch.
Literature: Groenland. Org. gland. du genre Dt'bstra, Ann. IC. nat., ser. 4. t. iii, 1855, pp. 291-
303, and pI. 9.-Trecul~Gl. poo.ic. du Dros~ra rolundi/o/ia, Ann. IC. nat., ser. 4, t. iii, 18S5, pp.
303-1I.-A. Meyer and Dewevre, I)fw~ltyllulllluJilaltitW"" Bot. CentraIbl., 1894, iv, pp. 33-4'.-
Macfarlane, Hybrids, &c., Contribut. Bot. Laborat. P~nnsylv.• ii, n. I, 1898, p. 87 et~. and pl. xii.
-[Rosen bug, D,."se,.a ,."JundijolitJ, Medd. Stockb. HOgskoL Bot. lnst.. ii, 1899, 126 pp., l Ta~;
abstr. in Just, 1899, ii, p. 207.)-Habedandt, SinnesorgaDe, 1901 , p. 94 tt seq.-Lang, PolypompluJlyx
u. Byhlir gt'fantta, Flora., 1901, pp. 179-92: also Diss., Munich.-Dutailly, Parnasna, Assoc.
franc;. Ajaccio, 1901 ; ii, 1902, pp. 471, 472.-[Hamilton, Byhlis grgan/ea, Proceed. Linn. Soc. Ntw
Sooth Wales, 1903, p. 680 et seq.; abstr. in Bot. Centralbl., xcvi. p. S79.]-Schoute, Stelirtheorie.
1903. p. II7.-Fenner. Laubbl. u. DrUsen tiniger Insc:ktivoreo, Diss., Zurich, 1904, pp. 33-91 and
Tab. xi-xxi; also in Flora, 1904.-Freidenfeldt, Anat. Ban d. Wurzel, Bibl. bot., Heft 61. 1904,
pp. 61 and 63.-Diels, Droseracellc, in Pflanzenreich, Heft 26, 1906, p. 4 d: seq.-[Haberbndt,
Sinnesorg'J 2nd edit., 1906.]

SOLEaEDF.. 3N
 ADDENDA

HAMAMELIDEAE (pp. 328-333).

The petiole of Liquidambar contains three vascular systems of the nature
of steles (Bouygues).
Literature! [Bohnel, Kork, Sitz.-Ber. Wiener Akad., lXlI'Vi, Abt. I, 1877, p. 603.]-Gregory,
Cork wings, Bot Gazette, 1888, pp. :l8~-7 and pI. xxii (Liqu~·tlambar styradflua).-Jadin, Org.
secret., These, MontpeUier, 1888, p .... 8 et seq.-Wijnaendts Francken, Sclereid~ Diss., Utrecht,
l890, pp. 51, 5~.-[Wassujewsky, P,",otia pel'sica, 1891 (Russian) ; abstr. in Just, 1893. i, p. 580.]
_[Mohr, Balsam von LiquidamlJar, Pbarm. Rundschan, N. Y., 1895. xiii, n. 3.1-Uensen, Rinde
von Hamamelis virginiana, Pharm. Archiv, '90r, n. 7; abstr. in Just, 1901, ii, p. 58.]-Pitard,
Pericycle, These, Bordeaux, 1901, p. IO:l.-Bouygues, Petiole, These, Bordeaux, 19°2, p. 6g.-
[Kramer, Mikr.-pharm. Beitr., Diss., Wtirzburg, 1907, p. 20 (Hamamelis).]-For Cercidiplty/lum
and o~er g~era belonging to the Hamamelideae, see under Trochoclendraceae.

BRUNIACEAE (pp. 333-335).

1. REVIEW OF THE ANATOMICAL FEATL'RES. According to Kirchner, the
apex of the leaf in all the Bruniaceae is suberized in a peculiar manner.
2. STRUCTURE OF THE LEAF. According to Colozza, the earlier statement
that the leaves of the Bruniaceae are traversed by three veins, is only partly
correct. In certain species of A udo uinia , Berzelia, Linconia, Pseudobaeckea
and Staavia five veins are found, while some of the species of Pseudobaeckea
have even more (9, 13, or 20).
The stomata, as already previously mentioned, are generally present on
_Doth sides of the leaf. On the basis of recent investigations the following facts
may be added regarding their distribution. In the pinoid, patent leaves of
Lin,conia cuspidata, Sw. and the thin flat leaves of Pseudobaeckea cordata, Niedenzu
the stomata are confined to the lower side. In the broad scaly leaves found
in certain species of Brunia, Lonchostoma), Pseudobaeckea and Raspalia t the
stomata occur only on the upper side, which is adpressed to the stem., In
certain species of Brunia and Diberara with subulate inflexed leaves the
stomata are practically restricted to the upper side, since stomata are found
on the lower side only at the base of the leaf, i.e. on that part of the leaf wpjch',
is covered by the tips of the leaves below (Kirchner). In Linconia cuspidata,
Pseudobaeckea cordata 3 and Thamnea gracilis, Olivo Colozza states that the
stomata are confined to the lower side, while in Lonchostoma acutiflorum,
Wickstr., Pseudobaeckea virgata, Niedenzu, and Raspalia Phylicoides, Niedenzu
they occur only on the upper side. With reference to the arrangement of the
stomata on the surfaces of the leaves, we may mention that according to
Kirchner they are irregularly scattered in Staavia nuda, Erkl. et Zeyh. and
Brunia sacculata, Bolus, while in Staavia capitella, Sond. and Brunia palust,is,
Schlecht., as well as in Staavia radiata. they show a marked transverse arrange-
ment.
J\.ccording to Colozza, the structure of the leaf.is either centric or bifacial,
or exhibits transitions between the two types. According to the same authority,-
the earlier statement that the palisade-tissue in all cases consists of a single
layer of cells is incorrect. Kirchner states that the palisade-cells are almost
'invariably placed obliquely to the surface of the leaf. In those leaves in

1 Regarding Londlostoma. which in recent times has been referred to the.llmniaceu by general
consent, see also the former appendix to the Solanaceae, pp. 581, 583.
I According to Kirchner (loC. cit., p. 15) these species are: • Brunia paluslris, Schlecht.,'
• B. saccu/ala, Bolus;' Londlo,sloma acuh]/orum, Wickstr., L. mOll(jslyle, Sond., L. 061us'f/4,rum,
Wickstr.; PseudlJlJaeew s'lualitia, Niedenzu, P. virga/a, Niedenzu; Raspalia ~~lalaJ E. Mey,-
R. nregeana, Niedenzu, R. m{crojltylla, Brongn., R. pltylicoidea, Niedenzu.
S See Colozza., loco cit. p. 20 (in contradiction, however, to the statement on p. '9)'
 ADDENDA-BRUNIACEAE
which the uppoa; surface is ad pressed to the stem, the palisade-tissue is naturally
confined to ~ lower side of the leaf.
Colozza records solitary crystals oi oxalate of lime in the neighbourhood
of the mechanical sheaths enveloping the vascular bundles of the veins in
Brunia nodiftora, L. and in the genera A udouinia, Berzelia, Diberara, Thamnea
and Tittmannia; the same authority mentions the occurrence of clustered
crystals in the mesophyll in Brunia globosa, Thunb. and B. lqevis, Tbunb.,
and in the genera Linconia, Lonchostoma, Pseudobaeckea and Staavia. In
Raspalia both solitary and cluster~d c:t:ystals are wanting. .
Long unicellular clothing hairs are recorded by Colozza in species' of
Brunia, Diberara, Pseudobaeckea and Raspalia.
Parenchymatous cells serving for water-storage are present at the apex
of the leaf in the neighbourhood of the terminations of the veins; in an un-
named species. of Brunia similar cells lie scattered in the spongy tissue.
The tip of the leaf in the Bruniaceae is formed by a cap of tissue; composed
of brown suberized cells; on the inner side of this cap lies a meristem, from
which new cork-cells are produced at the same rate as the outemlOst ones
become exfoliated.
Literature: Knoblauch, Okolog. Anatomie, etc., Hl;\biHtat.-Schr., Tiibingen, 18gti, p. 15 et &eq.
-Kirchner, Bdtr. z. Kenntnis de!" Brunisc. J Diss., Breslau, 1904. 29 pp.-Colo~ Bruniactac,
~nn. di Bot., ii, RODla, 1905, sep. copy, especially pp. 9-3'"

HALORAGEAE (pp. 335-339).

To the first paragraph on p. 335, in which the most important anatomical
features of the Order are sumrnarized, we may add the following details. The
Haloragoideae, Schindl.1 are distinguished by having uniseriate clothing hairs,
the Gunneroideae, Schindl. by having lmicellular hairs. The genera Callitriche
and IliPP'lf,is are characterized by the possession of peltate hairs; Schindler
advocates the exclusion of these genera from the Halorageae, and in this he
is probably right. Hippuris is also peculiar in the absence of clustered crystals
of oxalate of lime. The axis of Loudoni,l' is distinguished by having subepi ..
dermal groups of sclerenchyma consisting of a single layer of fibres.
Recent investigations have shown that the mesophyll rarely contaiJ'!..s
typical palisade-tissue with marked elongation of its cells j a distinct differentia-
tion into·palisade and spongy tissue is found in- Haloragis elata, Cunn. and in
Loudonia. Sclerenchymatous elements are completely absent in the leaf.
Schindlenalso states that the stomata in no case have characteIistic subsidiarv
cell~; we should mention, however, that in the Haloragoideae those walls
of the surrounding epidermal cells which converge on the guard-cells are
thin, while in the Gunneroideae the contoU!"~walls of the guard-cells the.m-
selves are thin; the remaining epidermal cells in both cases have thick lateral
walls. As a rule the stomata are present on both sides of the leaf. In the
extreme terrestrial species· of Haloragis and Loudonia the stomata situated.
on the upper side are not so large as tho~e on the lower, and occur ill smaller
numbers; in other cases they are only represented by rudiments.
Special internal secretory organ~ are absent, but tannin is widely dis-
tributed both in the land- and water·fonns. The clustered crystals of oxalate
of lime are mostly of small size, but very large ones are a]so found. (e.g. in
Gunnera). They are invariably composed of a large number of individual

1 Schindler, on whose work the (oUowmg additions are chiefly based, subdivides the Order as
follows: I. Haloragoideae: J, Halorageae! Lamionia, Haloragis (incl. Meio1uctn), MesielllJp
Laurembe"gia (Serpicula). Proserpinaca j 2, Myriopbylleae: Myn'ophyllullt. II. Gunneroideae:
Gunnera.
3 N2
 ADDENDA--HALORAGEAE
'Crystals with very narrow tips. Accordingto Schindler, hair-like cells con-
taining c1ustered crystals, which were previously' mentioned as occurring in
the cortex of the stem in lYyriophylium and.Serpicula repens, are found in all
the species of Laurembergia, Meziella, Myriophyllum and Proserpinaca, as
well as in certain species of H aloragis, having relatively large intercellular
spaces. It remains to ~ention the occurrence of aggregates of acicular crys~
tals, composed of an unknown chemical. substance, ,in Gunnera H amiltonii
(according to Schnegg).
We have next to discuss the trlchomes, 'and will deal first with' the clothing
hairs. We have already stated above that according to Schindler uniseriate
clothing hairs are characteristic of the Haloragoideae, and unicellular hairs
of the Gunneroideae. It should be noted, however, that the,.aquatic forms
(Myriophyllum and Meziella) have no clothing hairs whatsoever, and that in
some species of Haloragis unicellular hairs are also found. The.latter are,
however, connected by transitional forms with the epidermal papillae, which
are of frequent occurrence in Haloragis, and differ very essentially from tho
haits of 'Gunnera (which have thin walls and wide lumina) in being of small
size and having thick walls and narrow lumina. The unicellular trichomes
of the New Zealand species of GUnnera examined by Schnegg are for the.most
part club-shaped and cro'wned by a short awn. ,According to Schindler, the
distinction between Hippuris maritima and H. vulgaris (mentioned in the
earlier part of this work) based on the structure of the rays of the peltate hairs
cannot be maintained.
According to Schindler. glandular shaggy hairs, having the same structure
as those of Myriophyllum and Gunner«, occur in all the Haloragoideae with
the exception of Loudania; they are situated in the excavations on the leaf-
teeth, &c., and a.re visible even with a lens as small lappets or dots. Other
forms of hairs to be mentioned here are: (1) the soft cylindrical hairs of
Gunnera, which had already previously been observed by Uhlworm, and
consist of four regular rows of cells; (2) the emergences recorded by Schindler
in Haloragis exaiata, F. v. M., and occurring on the axes in the form of coarse
knobs having the shape of an inverted flask; and (3) the multicellular trichomes
described by the same author in certain terrestrial species of Halo;agis (H. con~
tertifolia, F. v. M., H. elata, Cunn .• and H. exalata, F. v. M.); these trichomes
resemble a blackberry in form and are purely epidermal. structures.
In turning our attention to the structure of the axis we may once more
emphasize the absence of intraxylary phloem in view of. the Clcse affinity of
this Order with the Onagrarieae. According to Schindler, ParmentIer's
statements as to the occurrence of internal soft bast in H aloragis and Loudonia
(d. footnote I on p. 335) are incorrect. Schindler states that the vessels have
simple perforations in all the members of the Order. Among the features
presented by the cortex. (da. p. 339) the occurrence of subepidernlal groups of
sclerenchymatous fibres in Loudonia has already be~n refen-ed to above: Most
of the Haloragoideae are characterized by having very large air-cavities in
the primary cortex, the cavities being arranged to form a single ring (except
in Haloragis salsaloides, Benth.). These interce1111lar spaces are most prominent
in the aquatic forms, but are also found in the large majority of terrestrial
species of Haloragis, although not so strongly developed. They are absent,
-however, in 'Loudonia, Haloragis loli-osa, Benth., H. Gossei, F. Y. M. and
H. pycnosiachya, F. v. M. In many Halorageae (sensu Schindler)-intercellular
spaCes are present in the pith as well, but they are w31l.ting in Meziella, as
well as in the Myriophylleae (Myriophyllum). In contrast to the typical
Halorageae, Hippuris has several layers of intercellular spaces in the cor~tex ;
the same applies to Haloragis salsoloides, Benth., a species already referred
to a b o v e . ' .
 ADDENDA--HAL ORA GEAE
A brief description of the vascular system of the petiole and stolons in the
species of Gunnera, examined by Schnegg, may be given, the reader being referred
for details to Schnegg's paper. The larger petioJar strands are distinct steles;
in the simplest case {e.g. G. dentata} they consist of a concentric vascular bundle,
surrounded by an endodennis and provided with central xylem, the innermost
vessels of which are situated in an apparent pith and exhibit compression; in
G. lobata the steles include a pith composed of sclerenchymatous fibres and have
a collenchymatous sheath; a still higher degree of differentiation is attained in
G. magellanica, in which the place of the pith is taken by a vascular system, sup-
ported by two groups of sclerenchymatous fibres and composed of central strands
of phloem with peripheral xylem-vessels, while the collenchyma-sheath is accom-
panied by a second sheath, provided with Caspary's dots on its radial walk The
stolons, with some few modifications, as a rule show two rings of wood and bast,
the inner one of which is inversely orientated as regards the position of xylem
and phloem; in G. Hamiltonii there is only a single stele, while in G. cMlensis
there are several (3-7).
Literature: Ublwonn, Entwicldungsgesch. d. Trichome, Bot. Zeit., 1873, pp. 769-73 and Tab.
be; also Diss., Leipdg.-Costantin, Tiges de: pI. aquat" Ann. sc. nat., ser.6, t. xix, 1884, p. 287
and pI. xiv, xv.-fDanie1li, Certi org. della Gunnera !Cabra, Atti Soc. Tosc. Sc. Nat., vii, 1885,
J7 pp.; abstr. in Bot. Centralbl., 1885, iii, p. 303.]--Scott, Polystely, Ann. of Bot., v, 189~I,
p. 5J4 et seq.-[Parmentier, in Le Monde des Pl., 1897. p. 178 et seq.]-Wcinrowsky, Scheiteloffn.
hei Wasserpfl., Diss., Berlin, 1898, p. 24.-Roedler, A$Similator. Gewebes}1it., Disa., Freiburg i. d.
Scbw., 1898-9, p. 34 et seq.-Minden, Wassersec. Organe, Bib1. bot., Heft 46, 1899, p. 10
(Callitriche).-Perrot, Org. app. des feuilles de cert. Jl.lyriophyUunt, Joum. de Bot., 1900, pp. 198-
20l.-W. B. Me. Callum, Proserpinaca palustris; Bot. Gaz., 1902, pp. 93-108.-Knothe, Unbenetzb.
Bl., Diss., Heidelberg, 1903, p. 20•.:.....[Porsch, Spaltoffnungsapp. submeJ1l. Pfianzenteiie, Sit:t.-Ber.
Wiener Akad., exii, Abt. 1,1903, pp. 103-7 (Callitricht, Hippuris)).-&hnegg, Beitr. lo. Kenntnis
d. Gatt. Gunnrera, Flora, '"c, 1902, p. 161 et seq.; also Diss., Munich.-Scboute, Stcliirtheorie, 1903,
pp. 131-2.-Schindler, Abtr. d. Hippuridaceen von den Halorag., in Engler, Bot. Jahrb., xx\iv,
1904, Beibl. 77, 77 pp., especially pp. 53-69 and 74-5; also Diss., Erlangen.-Giinther, Anat. d.
Myrtifloren, Diss., Breslau. 1905. p. 27 et seq.-Sehindler, Halorrhagaceae. in PB~nreich, Heft 2~"
1905, pp. 5"'9·-Geneau de la LaJllarliere, Membr. Cllt d. pI. aquat., Revue gen. de Bot., 1906,
p. 289 et seq.

RHIZOPHORACEAE- (pp. 339-343).

2. STRUCTURE OF THE LEAF. The following additional facts have become
knO\\lll. The outer appendicular ridges on the stomata of Ceriops Candolleana,
Arn.~ Bruguiera gymnorhiza, Lam., and 'Rhizophora mucro nata, Lam. are split
into two, so that the front cavity is divided into two parts. An aqueous tissue
belonging to the spongy parenchyma is found also in the interior of the meso-
phyll of Kandelia and Bruguiera. The palisade-tissue of Rhizophora consists of
a layer of very much elongated cells, which exhibit more or less abundant
transverse divisions. Lastly, according to Areschoug's statements, hypoderm
occurs on the lower side of the leaf more frequently than was previously stated.
3. STRUCTURE OF THE AXIS. According to Pitard, the peri cycle in Rhizo-
theTa Mangle ·and-·Bruguiera parviflora contains only isolated groups of bast-
fibres; sclerosed stone-cel1s, which are occasionally branched, are found in
their neighbourhood.
Literature: Bohnel, Gerberinden, Berlin, 1880, p. 129 et seq.-[Karsten, M;..ugroveveGetat.,
Bibt. bot., Heft 22, 1891.]-Pitard, Pericycle, These, Bordeaux, 1901, p. 7J.-Areschoug, Blaltbau
d. Man~ovepfl., Bibl. bot., Heft 56, 1902, pp. 42, 57 and 66 et seq., Tab. i-iii.·1:I: ~d xiii.-
Bargagh-Petrucci, Legnami, Malpighia, 1902, p. 3.5+ (Brugu iera).-H 01 termann , Anat.-phyaiol.
U ntersuch., Sib.· Ber. Berliner Abo., 1902, i, p. 671.-[Blatter, Mangrove of tne'Domba,. Presidency,
Joiun. Bombay Nat. Hist. Soc., xvi, 1905, pp. 644-56 a!ld pl.~-Giint~er, A~t. d. Myrtifl.oren,
Diss. Breslau, J905, p. 2s.-Giirtler, Interzellulare Haarblld., DIS!., Berlin, 1905, p ••o.-Poulsen,
Stlrtt~tJdern," hOI Rhi%IJplzq"(l. Vidensk. Meddelels. Kj~benhavn, 1905, pp. 153-64 and Tab. v.-
Holtermann, Einfl.uss d. Klimas. etc., 190" pp. lO, ~'1, 60 and Tab. lx. tBrnguitrG, Ceriops,
IfAizo/"ora).
 9 18 ADDENDA

COMBRETACEAE (pp. 343--350).

2. STRUCTURE OF THE LEAF. The absence of a typical hypoderm is
remarkable. Aqueous tissue is present in the middle of the mesophyll in
Laguncularia l'acemosa, as well as in L-umn#zera coccinea and L. racemosa
(Holtermann). We may also mention, likewise on Holtermalln's authority,
the occurrence of large terminal tracheids in Laguncularia racemosa, and of
hyd.thodes provided with an epithema in species of Laguncularia and
Lumnitzera. To the previous account of the hairy covering we may add t~at
cap·shaped glandular hairs, similar to those found in Laguncularia rar.emosa,
occur also in Conocarpus erectus, though the hairs of this species are far less
sunk, and consist of a smaller number of cells (Boergesen and Paulsen). Details
as to the differentiation of the shield in the peltate glands of certain African
species of Combretum are given by Engler and Diels, loco Cit. According to
Boergesen and Paulsen, special epidermal idioblasts occur on both surfaces of
the leaf, but mainly on the upper side, in Con1ocarpus erectus (hydathodes ??) ;
they are distinguished from the remaining epidermal cells by their rounded
outline and by the presence of a conical elevation arising from the middle of
the cell; this protuberance has a striated cuticle, and. often includes a small
space which is separated from the rest of the lumen of the cell. The petiolar
glands of Conocarpus erectus constitute two depressions on the _upper side of
the petiole, and are provided with an epithema and the termination of a vascu·
1ar bundle (Areschoug); regarding the glandular stipules found in species
of Comoretum, see Mirabella, Ioc. cit.
For the structure of the petiolar spines of Combretu,m, see Heiden, Holmes,
and Lefevre, 11. cc.
3. STRUCTURE OF THE AXIS. The interxylary;>hloem in the specimen
of Combretum salicifolium, E. Mey (d. Holtermann s statement on p. 350);
examined by Leisering, develops in the same ,vay as in Guiera. According to
Lefevre, the same mode of development also obtains in Combretum glu#nosum,
Guill. et. Perr.
Phelloid·cells have been demonstrated in the cork of Quisqttalis (J. E.
Weiss). Lefevre states that the cork develops sub-epidermally in Cona-
carpus leiocarpus~ DC. (= Anogeissus leiocarpus, Guill. et Perr.) and Anogeissus
pendula, Edge-w.
According to Lefevre, intercellular mucilage--spaces of schizogenous origin
similar to those present in the veins of the leaf occur also in the intraxylary
phloem of Terminalia Bellerica, Roxb., T. procera, Roxb. and T. tomentosa,
W. et A., as wen as in the interxylary phloem of Combretum glutinosum~
Guill. et Perr. ,
According to Karsten, the negatively geotropic respiratory roots of Lumnitzera
racemosa, Willd. exhibit a well differentiated system. of intercellular spaces in the
secondary cortex in later stages. This intercellular system is specially distinguished
by its mode of development. In the first place the cells commence to separate
from one another at certain points and especially at the corners, so that they ocly
remain connected by narrow processes; in this way the tissue acquires the appear-
ance of a parenchyma the cells of which are provided with numerous processes
emanating in all directions (' conjugate parenchyma '). The intercellular spaces
gradually increase in size, and the connecting processes become longer; diVlsion-
walls appear in the latter and thus a strongly developed system of intercellular
spaces 1S ultimately formed. According to Boergesen and Paulsen, the intercellular
spaces in the primary cortex of the' asparagus-like roots' of I.aguncularia t'acemosa,
Gaertn. f. develop in exactly the same way.
Literature: Hobnel, Gerberinden, Berlin, 1880, p. 127 et seq.-liohnel. Neue Gerbebl., in
Dingler, PolytechD. ]ollrna], ccxl, J881, pp. 388-91.-J. E. Weits, Korkbild., Denkschr. bot.
 ADDENDA-COJIBRETACEAE 919
GaelllCh. Regea.bari, vi, J890, sep. copy, po 16.-Kanten, MangroftVegetat., Bib!. bot., Heft 22,
1801. pp. 51-1 and Tab. ix.-Brandt. Wenig belc. Rioden, Diss., Dorpat, 18~, p. 30 et &eq.-
[llirabel1a, Colleteri, Contribu.z. 1st. Bot. Palermo, ii, 1897, p. 15 et seq. ; abstr. in Just, l897,

~Int"""M~fl.,
. "I';']-Boergeseo og Paulsen, Vegetat. dansk. vestind. Oei, Bot. Tidaskrift, xxii, 1898-9, pp. 43-8
!Al~ racemMa, Gaertn. C.) and pp. 53--4 (Cmoc
er Bibl. bot., Heft S6, 1899, pp. 71-7 anm. ",dw, L.).-Areschoug, Blattbau
ix':xiii.-Engler and DieIs, CqM-
in EDgIer, MODogr. afrikan. PAan.tenfamilien u. Gatt., iii, 1899.-Leisering, Interxylares.
Leptem, Diu., ~, 1899, pp. 1.-17 ....... Baranetzky, Faisc. bicoll., Ann. sc. nat., ser. 8, t. xii,
1900, p. 300.-Bargagti·Petrncci, Legnami, Malpighia, J90~, p. 356 (Lumnitura).-Penzig, Piante
acuoftle, Malp~bia, 1902, p. H6 et seq. (Terminalia).-Areschol1g, Trap. van. bladbyg~., Sv..
Vet. Akad. H udl., 39. n. " 1905, pp. '5, J 6 and 75 (ComDnlum), pp. 9.., 95 (Tffmi1aHa) ,
pp. JJ5-}6 (Qllt·J~iJ).-Gijnther. Anat. d. Myrtiiloren, Diss., Breilau, 1905, pp. '5, a6.-
~te, Et. anat. d pharmacol. des Combret., 1905. in Perrot, Travau~, iii, 1906, 126 'pp.-[Drabble,
Anat. of the Kinkeliba, C"",!Jrelum Raimbaulli, Quart. Journ. Carom. Research In the Tropics,
LtT~l, ii, 1907, pp. 66-7 0 , I pl.J-Holtermann, Einfluss des Klim:tS, etc., 1907, pp. 3 2, 55, 58,
59 (Lagwuularia, Lumnitura).

MYRTACEAE (pp. 350-358).

I. REVIEW OF THE ANATOMICAL FEATURES. The unicellular clothing
hairs found in certain members of the Myrtaceae sens. str. are two-chambered
in the same way as in the Combretaceae. Unicellular Malpighian hairs, showing
transitions to ordinary unicellular trichomes, are found also in Eugenia co"eae-
folia, Hook. et Arn. The hairs of Bertholettia excelsa, Humb. et Bonpl., exhibit
a tendency towards a tufted arrangement, while those of Lecythopsis rutescens
show indications of branching. Shaggy hairs are present in Lhotskya gene~
thylloides, F. v. M.
Among the anomalous genera (see p. 351) Gaslondia and Psiloxylum have
recently been investigated 'by Van Tieghem. Both of them have proved to be
members of this Order, as evidenced by the presence of secretory cavities in the
primary cortex and rnesophyll, and by the occurrence of intraxylary soft bast.
Of the remaining anatomical features of the two genera we may mention th.e fol-
lowing. The cork in Gaslondia develops in the second cortici!l.layer and is composed
of alternating strata of quadrangular thin-walled cells and fiat lignified cells, whilst
in Psiloxylum it arises in the pericycle and consists of layers of fiat thin-walled
cells and thickened cells with wider lumina. The pericycle in Gaslondia contains
isolated groups of hard bast, and the phloem is stratified into hard and soft bast,
while in Psiloxylum bast-fibres are completely absent, and the stratification of
the phloem, as in 'Cortex Granati,' is due to cells containing clustered crystals. In
both genera, lastly. the stomata are found only on the lower side of the leaf; the
palisade.tissue in Gaslondia includes relatively large cells with clustered crystals.

A. MYRTACEAE SENS. STR. (pp. 352-355).

Holtermann records remarkably large epidermal cells in Eugenia subavenis.
and stone-cells in the mesophyll in E. rotundifolia and E. sderophylla.
,1 have myself investigated the hairy covering, and may mention in the
first place that some of. th~ simple unicellular clothing hairs are two-chambered,
like those found ih the Combretaceae. As in that Order the basal portion of
the hair contains a body which varies in length and has the appearance of
a cell, and in this way the duplication of the hair is brought about. This
feature is found: (a) in the narrow, elongated clothing hairs of Leptospermum
grandifolium, Hart., which have thick walls and narrow lumina and give rise
to the silky covering on the leaves; (b) in the short curly clothing hairs on
the leaves of Metrosideros tomentostl, A. Rich.; (c) in the sim.i1ar hairs of Kunzea
ericifolia, Reichb.; (dl in the mostly one-armed hairs on the leaves of Psidium
Guaiava, L.; (e) lastly ralso in the hairs on the fruits o(the Pimento (belonging
to Pimenta officinalis, Lind!.; in tp..is case they are figured by Rosen on his
I Wandtafel: xxvi. Fi~. B and C. but are not considered in the corresoondinlZ
 ADDENDA-MYRTACEAE

text, p. 188); these hairs are one-armed 'or indistinctly two· armed with arms
of unequal length. On the other ha.nd~ the unicellular clothing hairs found
on the ovary of Pileanthus filifolius~ Meissn. do not show the two chambers.
Unicellular clothing hairs having equal or unequal arms with wide lumina,
and showing transitions to crop-hairs, are found in Eugenia coneae/olia, Hook.
et Am. These hairs are specially remarkable in .that the membrane which
causes the duplication in the cases aboveJrnentioned can still be demonstrated
in t:~em with certainty, but since it is apposed. to the longitudinal walls of the
body of the hair along its whole length, it fails to produce duplication. Another
new form of trichome is that of the shaggy hairs which have a clothing
function, and are the cause of the hairy covering on the branches and the
cillation of the margins of the leaves in Llwtskya genethylloides, F. v. M. They
are multiseriate structures, only one cell in breadth at the apex, and are
characterized by the fact that the ends of the component cells starid off
from the body of the hair, in a papillose manner.
According. to Porsch, the secretory cavities found in the species of Euea·
lyptus have an excretory mechanism similar to that found in the Rutaceae;
it is composed of two parts, viz. (a) a passive portion, which in this case com·
prises the lid of the gland (·with 2-4 lid-cells) as well as those cells of the wall
which are situated immediately below the lid, and (b) an active, portion con-
stituted by the wall of the gland. Another noteworthy point is that in this
case both the inner and outer walls of one or both lid· cells become split open,
the outer walls tearing at certain points, pre-determined by their histo-
logical structure; the emission of secretion takes place through the slits thus
formed, and not, as in the case of the Rut.aceae, through' preformed clefts
between the walls.' Secretory cavities are present also in the genus Aphano-
myrtus, which was formerly regarded as a doubtfullTl':mber of the Myrtaceae
(Koorders and Valeton).
The cork commonly contains unsuberized cells"(phelloid-cells). In certain
species of Eucalyptus (e. g. E. globulus, Labill.) the secondary bast includes
numerous slightly sclerosed spicular cells, of an irregularly lobed, parenchy-
matous shape. .

B. LECYTHIDACEAE (pp. 355-357).

The following additional details regrading the clothing hairs are based
on incidental investigations of my own. The hairy covering on the stems of
Lecythopsis rufescens, Berg is constituted by 1-3·celled clothing hairs which
vary in length and are provided with transverse walls of varying thickness.
In some cases these hairs are united to form tufts (the ra'y-cells then occasion·
any consisting of two cells). The bicellular hairs further show a tendency to
branch, since the longitudinal wall of the lower cell is sometimes drawn out on
one side,into a short point,ed papilla. The ~lothing hairs on the fioralbuds of
Bertholettia execlsa; Humb. et HonpI. are for the most part unicellular, more
rarely bi~ellular, with a thin transverse wall, and are peculiar in the fact that
their longitudinal walls exhibit transverse foJds at certain points; these folds
penetrate into the lumen of the cell to a varying extent,and are placed at right
angles to the longitudinal axis of the body of the hair.
According to Areschougt secretory cells are fonnd in the mesophyll in
N apoleonea H eudelotii, Juss.
Literature: Hahnel, Gerberinden, Berlin, 1880, pp. '32 and 13-4 et seq.-Lamounette, Liber
interne, Ann. sc. nat., ser. 7, t. xi, 1890, pp. 260-I.-J. E. 'Weiss, Korkbild., Denkschr. bot.
Gesellsch. Regensburg, vi, 1~90, p. 16 et seq.-Johannson, Wenig bek. RiDden, Diu., Dorpat, 1891,
pp. II and +2 et seq.-[Korpow, M,laleu(a Cajeput'~ Dis!., Moscow, 1894.1-Wartenberg, Psidiu1I'
A"araJ Diss., Erlangen, 1895, pp. 17-3 ...-Matteucci, Placche lugherose~ Nuovo Gicm. bot. Ital.,
 ADDENDA-MYRTACEAE
%897. p. 23.... -[A. Schneider, Leaves of Et«aI)'l'm gltJl.uhu, Jouro•. of Pharmacol., New York, 1897,
n. 7; abstr. in Just, 1897, ii, p. l07.]-Koordersand Valeton, Apkanomyrlus, Ann. Jardin Buitenzorg.
Supp). m. 1~8, p. l~5.-Le.nz, Folja Djamhu, Bet. deutsch. pbllrm. Gesellscb~ ]899, pp. 125-36,
Tab.-Baranctzky, }~aisc. bicoll., Ann. sc. nat., ser. 8, t. xii, 1900, pp. 299-3°o.-[Palmieri, Anat.
camp. del genere Etaalyplus, NapoU, 190", ]9 pp.l.-Unprung, ADst. u. Jahresringbild., dc.,
Diss., Basel, 1900, pp. 18-20 (Psidium /t)11liftrum, L).-(Baker and S~ith, Eucalyp.ts, Technic.
Educat., SeT. 13. Techn. Mus.., New South Wales, Sydney, 190,).]-Hargagli-Petmccl, Legnami,
Malpighia, Iyo:a, p. 356 et seq. (Eugenia, Tristania).-Fabricills, Laubbl.-Anat., Beih. z. bot.
Central bl. , xii, ]902, P. 330 (B4rringiPnia).-Knothe, Unbenetzb. Bt, Diss.; Heidelberg, 19(1:1,
p. ~o.-Porsch, Entleerungsapp. innerer Drusen, a.t. bot. Zeitschr., 1903, pp. :156 and 318 .e.t seq.,
Tab. jx.-Col, Faisceau%, Ann. sc. nat., set. 8, t. U, J904, pp. 195-:-.:105 and ~.1-5.-Van Tle}rhem,
Gaslondie et Psiloxyle, Ann. sc. nat., ser. 8, t. xix, 1904, pp. 349-60.----:Areschoug, Trop. vaxt, blad-
byggn., Sv. Vet. Akacl. Bandt, 39, D. ~, ]905, pp...6, 47, Tab. v. (Napoleonea), pp. IIo-Il and
Tab. xvi (Psidium), pp. 130, 131 (Eugmia).-[BourdilloD, Timber of !Jarring/Mia, Indian Forester,
xxx;, 1905, pp. 89, 9o.]-Giinther, Anat. d. Myrtifloren, Diss., Breslau, 1905. pp. "4, 35, and :18-3 1 .
-Porsch, Spaltoffnungsapparat, lena, 1905, p. +4.-[H. G. Smith, Calcium oxalate in the barks of
tbe Eucalypts, Abstracts Roy. Soc. N. S. Wales, May, 1905, pp. ~, ..J-[Pampanini e Pampaloni,
Xanl!uJstemon, Nuovo Giorn. bot. Ital., n. s., xiii, 1906, pp. 121-37.j-Picdoli, Legnami, Bull. Siena,
l~, pp. ]SJ, 15:J.-Holtermann, EinfJoss cles Klimas, etc., 1907, p. 120 et seq. (Eugenia).-[For
additionalltteraturc, see p. 1171.]

}IELASTOMACEAE (pp. 358-368).

1. REVIEW OF THE ANATOMICAL FEATURES. The occurrence of a
characteristic stomatal apparatus, in which the pairs of guard· cells are almost
completely surrounded by a single epidermal cell, is worthy of special '-note.
A very striking feature of the hairy covering is the great scarcity of unicellular
clothing hairs'; uniseriate hairs are also of infrequent occurrence. Shaggy
hairs showing manifold types of structure, as well as stellate, tufted, candee
labra, and peltate hairs are far commoner. The hollow shaggy hairs of
Heterotrichum strigosum, Tr., and the combinations of clothing and glandular
hairs~ found more especially in many of the Miconieae, also require special
mention. External glands are widely distributed; they are clavate or their
head is spherical, discoid, or otherwise shaped or (rarely) differentiated as
a vesiCular integumental gland; in other cases the glands are represented
by glandular shaggy hairs. Peculiar retort~shaped glands' are fQund in
j

Bellucia~ papillose or hair-like appendages on the cells of the stalks of ex-

ternal glands-in species of Mico1U'a, &c., composite glandular hairs with two
or more heads on a common stalk in spe~es of Calycogonium, Charianthus,
Henriettella, ~"1iconia. Of internal secretory organs, tallnin-idioblasts have
recently been observed in species of Clidemia and }fliconia. The only addi·
tional fecord of gelatinization of the epidermis of the leaf is afforded by
Pternandra. Typical soljtary crystals of the ordinary form appear, to he
wanting in the Melastomaceae, at least in the-leaf; but in some cases the
clustered crystals exhibit a reduction to short thick solitary crystals. To the
enumeration of special features, which are of value in detailtd diagnosis, we
may add: the reticulate thickening of cells of the palisade tissue (species of
Grafjenriedliwaiid Merian,a); the sclerosis of cells of the mesophyll; the
presence in certain members of the Order of spongy tissue, provided with
peculiar collenchymatous thickenings; the occurrence of spiral tracheids in
the roesophyU (species of Bellucia, H enriettea, and Sonerila).
2. STRUCTURE OF THE LEAF. The structure of the leaf has recently been
investigated by PalCzieux in numerous genera belonging to the Tribes
Osbeckieae, Rhexieae, Merianieae, Oxysporeae, Sonerileae, Bert.olonieat. Disso..
chaeteae, and Blakeae, as well as in the Astronieae and Memecyleae 1, and by

1 The (ollowing genera were examined: Osbeckia, RJu)(/osepala t OtantlUF"fI, Muasl.",..

Tn'slert1fluz, IJusotis, IJicfatUtantlura, IHnopho,.a (Tribe Osbeclcieae); Rluxia, Pac"yltmUJ. M,..
(katlum (Tribe Rhexieae); HulJtria, Belmria, Opisthocentra, Adt/()/Jotrys, Men'ania, G,.ajfiterkt/jJ'
 ADDENDA-MEL ASTOMACEAE

Gottschall in numerous genera of the Miconieae.' These ,new investigations

help to confirm and extend the statements made in the earlier portion of:
this work. '
According to Palezieux and Gottschall, the stomata. vary very much in
Slze. As regards the neighbouring cells both authQrs in the first place state
that the pairs of guard.cells are frequently surrounded by three or mo~e ordinary
epidermal cells; side by side with stomata of this type, others'with subsidiary
cells placed transversely to the pore occasionally occur in one alld the same
species. The type of stoma just mentio'ned,is found in most of the species of
the Tribes Oxysporeae, Sonerileat', and Bertolonieae, as well as in Medinil/a.
and many of the Miconieae. In stomata of this kind one bf the two trans ..
versely placed cells often surrounds the greater part of the guard-cells, so that
the other neighbouring cell merely touches. them; in some cases the latter
\mdergoes division by a wall approximately coincident with the direction' ()f
the pore, so that three neighbouring cells are found, one of which surrounds
the pair of guard-cells in the way previously' described, ~tomata with neigh-
bouring cells placed parallel or ohliq.uely tb the pore
are rare (Micomeae).~
In certain species of Osbeckia and Dsssotis the stom.lta are arranged parallel
to the midrib. In Clidemia hetCfoneura, Cogn. the pairs of guard-cel1S occur
singly in crateriform depressions in the ~urface of the leaf, while in 'species of
Clwemia and Miconia they are surrounded by a cuticular ridge. Crowding
of the stomata in groups is met with on the mner and (rarely) outer side of
the domatia (inhabited by ants) found in species of Tococa and Mymiidonf., .
as well as on the lower side of the leaves in Leand,a acuti/lQrfJ, Cogn., ~4ly­
cogoniutn Krugii, Cogn" and OsStU4 Krugiana, Cogn.; in the two species last
named this phenomenon is due to the partial sclerosis of the lower epiderntis.
Palezieux and Gottschall also mention the occurrence of epidermal cells
with wid~ l~na and the gelatinous. appearance. of the. cellulose·membranes
of the epIdernus as features charactenstic of certam Specles; other characters
are the cuticularization of the outer wall, the pehetration of the cuticle into
the lateral walls in the form of ridges or pegs, and the varied types of striation
presented by the cuticle. The epidermal cells, found in the neighbourhood of
the .bairs and those situated above the crysta1~idioblasts of the mesophyll,
occasionally ha.ve a special sbape. The case of Huberia laurina, DC. may be
particularly noted in this connexion; here rows of elongated epidermal cells
extend from gland to gland on the upper side of the leaf, these rows iri their
entirety forming a network. The only additional record of gelatinization
of the epidermis of ~he leaf is furnished by Pternandra, so that mucilaginous
epidermal cells are only of very rare occurrence in this Order. Upper epidermal
cells showing palisade-like elongation are found also in species of CaJycogonium,
Clidemia, Leandra, lrliconia, Ossaea, and Tet1azy~ia; such cells either 'form
the whole of the epidermis or only occur in certam parts, which are situated
above the idiobiasts (containing clustered crystals) in the palisade~tissue and
above the veins; the cells in this case are occasionally divided transversely
or the lateral walls exhibit a concertina·like folding. Palisade-like elongation
of the lower epidermal cells has been demonstrated only in certain species
of Miconia. Papillose differentiation of the upper and lower epidermis and

(Tribe' Merinnieae); Allo1tJo1'j/u'(J, OXjs}>tmJ., Bm-dea, Blastus, Od,lh.ocllaris, VtprlCtlla. (Tribe

Oxyspoteae); Saneri/a, Sarco/'J'ramis (Tribe Sonerileae); .B~"lo!()niat Salpinga, T,.;olttta (Tribe
Dertolonieae) j Martlmia. DiJ1iUluula. AmJledf'llm. Ompha/()jJu!, Medinilla (Tribe Dissochaeteae):
BJaJua (Tribe BWceae) j Astro1f;a, Pternandra, Xi/)lssia (Tribe Astronieae) ; M~riria, Nmmykn
(Tribe Memecyleae). .
1 The falIowin" genera: l~andra, C(;n()!/~sia, Cllar;anllzuJ, Ttlrazyg;a, AlUott;a, C4lycogoniulIl,
Hetel'oIric"UIII, Tococa, A/ajl/a, Mpmidone, A/Uranium, Clidt/Jl;d, Bellufia, LortYIJ, Htnritllla,
HmrielltJ/IJ, OmuD, ltfyriasjcra.
 ADDENDA--MELASTOMACEAE
the occurrence of a hypoderm composed of one or more layers have been
observed in many species.
The upper epidermis is papillose in the following species: Allomorphia umbel-
lulata, Hook. f., Bt1'toioriia maf'mM'ata, Naud., Clidemia cymiferu., Donn.-Smith,
Conostefia subcrust,,'a, Tr., C. viridis, Cogn. and C. xalapensis, Don, Heterotrichum
Egge,s;s, C9gn. and H. pallens, DC., Leandra 50lenifera, Cogn., M"i1lill" ast,o-
nioides, Tr., Opistlwcentra cJidemioides, Hook.. f., Sonerila obliqutJ, Korth. and S.
secunda, R. Br.; a papillose lower epidermis occurs i~: Bellucia lwasiJi~"sis, Naud.,
B. dichotoma, Cogn., II. grossularioiaes, Tr. and B. imperialis, SaId. et Cogn., Dichae-
tantltera altissi'ma, Cogn., Dissocluuta pall'ida, BL, Henriett4a panulata, Berg.
H. Ma,.tii, Naud., H. multiflora, Naud., H. Spruceana, Berg and H. $",cosa, DC.,
Kibessia 1Ii"lIlla, Cogn., Mecranium haemantnum, Tr., Miconia guiarunsis, Cogn.,
M. macrophyUa, Tr. and M. Regelii, Cogn., Ptern"ndra paniculata, Benth. (sub-
papillose}, T#ITa.r"pa bic()/ot', Cogn. and T. paJJens, Cogn. The papillae vary in
shape, but are ma.lnly conical or finger-shaped.
HrPOderm has been observed on the upper side of the leaf in: Anplutrum
tUsamscum, Clarke, Astronia Candelleana, Cogn., Blakea pulve,ulenta, Vahl and
B. t,inervia, L. (both species with a two-layered epidermis), Calycogonium squamu-
losum, Cogn., Charianthus coccimus, Don, C. corymbosus, Cogn., C. lonti/olius.
Cogn. and C. nodesus, Tr., Clidemia blephat'odes, DC. (cells remarkably large and
showing concertina·like folding), Ccmostegia Donnell-Smithi;, Cogn." C. procera,
Don, C. subhi,suta, DC. and C. kalapensis, Don, Dichaetanthera altissima, Cogn.,
D. lati/ol;a, Cogn. and D. ,osea, Cogn., Dissotis incana, Tr. (lateral walls exhibiting
concertina-like folding), Gralf_enrieda boliviensis, Cogn. and G. ema,ginata, Tr.,
HmrUttea angusti/OJia, Berg, H. gronulata. Berg, H. Martii, Naud .• H. muitiflOt'a.
Naud., H. Spru&eana, ~ and H. $uccosa, DC., Henriettella fascicularis", Tr.,
H. Macfadyeni;, Tr. and H. mefn/)ranilols'a, Cogn., Loreya n'ig,icans, Tr., MIIf'Umia
paehYK'Yna, Korth., Me&f'anium amygdalinum, Tr. and M. haemanthum, Tr., Medi-
nilla ast,07Iioides, Tr., M. magnifica, Lind!., M. mYl'tijonnis, Tr., M. pap;llosa,
Bak., M. parvi/oJia, Bak.., M. pauciflo,4, Hook. f. and M. rubicunda, BI., M~/astoma
imm&alum, Wall., M. MaJabal}'l'icum, L., M. normaJe, L. and .1.11. olJvolutulT), Jack,
Meriania bo/iviensis, Cogn. and M. calophylla, Tr., Miconia "{finis, DC., M. cubensis,
Sauv., M. difficilis, Tr., M. joveolata, Cogn., M. glandulifeya, Co~., M. hi,tella.
Cogn., M. lifacina. Tr., M. quadrangularis, Naud., M. Sintenisis, Cogn. (in the
species last named the cells of the one-layered hypoderm have very wide and deep
lumina, are sometimes transversely divided, and are further provided with exception-
ally thick and gelatinous inne:- walls which have a milky white colour and extend
upwards in the form of a wedge between the lateral wallS) and M. theaeztJfls. Cogn.•
It?ouriria dotningensis, Spach., M. grandifiora, DC., M. guianensis, Aubl. and
M. mYt'tilloides~ Poir., Ochthocharis borneensis,. '81., Paehyloma coriaceum. DC.
and P. huberioides. Tr., Sonerila Irandiflora. "R. Br. and S. moluccana, Roxb.,
Tetrazygia angu.stijolia. nc.. T. bicolor, Cogn.. T. ,rotani/alia, DC., T. discolor.
DC., T. elaeagnoides, DC., T. hispida, Mad. and T. pallens, Cogn. According to
Holtermann, Kendn:cMa WalluN also has a hypoderm, which occupies half the
thickness of the leaf.
In Miconia tetrandra, Naud. the upper e1?idermis in transverse section consists
of a ~g number of layers. the cells of which are of unequal size ; in this species
cells of varying size are cut olI irregularly from the outer· side of the larger epidermal
cells.
The following structural features of the mesophyU require special mention.
The species of Bellucia have a palisade-tissue, composed of two or three
layers of cells, which are fitted in among one another in such a way that they
are finnly joined together. Reticulate thickening of isolated palisade-cells,
similar to that occurring in Clusia ,osea, L. (Guttiferae), is found in M eriania
paniculala, Tr. var. parvi/olia, Cogn., and GratJenrieda emarginala, Tr. In
Medinilla papillosa, Bak. isolated cells of the palisade~tissu~ are !c1erosed.
Special tanninaidioblasts, which are filled with brown contents in h~rbarium·
material, have been observed in the palisade-tissue in Clidemia KapPleri,
Cogn., and Miconia compressa, Nand., whilst in other members of the Order
all the palisade.cells have brown contents, and at the same time show -folded
 ADDENDA--MELAS TOMA CEAE
lateral walls; large lenticu¥tr, starch-grains occur in the mesophyll in spedes
of Meconium and Miconia. :ollenchymatous thickening of tpe spongy tissue,
similar to that found in Pachyloma coriaceum (see p. 360) is recorded by
Palezieux also in species of A nplectrum, GraUenrieda, M edinilla, M ela!toma,
Meriania and Ochthocharis. Spiral tracheids have been observed at the limit
of palisade and spongy tissues in Sonerila molu.ccana, Roxb., and iIi the spongy
tissue in BeUucia imperialis, Cogn. et Sald. and Henriettea angustifolia, Berg.
In Medinilla papillosa, Bak. and M. parvifolia, Bak. the middle. layer of the
spongy tissue is sclerosed, while in species of Leandra, lI-ficonia, and Ossaea
the walls of all the cells of the spongy tissue are remarkably thickened; in
lIuberia semiserrata, DC. the spongy _tissue only. contains isolated sclerosed
cells. Gottschall lastly mentions the occurrence of strongly sclerosed and
pitted cells, having an undulated outline, in the spongy tissue of Henriettella
glabra, Cogn., and of strongly sclerosed cells of a serpentine form in, that of
Miconia tetrandra, Naud.
A consideration of the data recentJy publi~hed seems to show that scleren-
chyma is more frequently found accompanying the veins than was formerly
supposed, although the accompanying tissue for the most part exhibits
collenchymatous differentiation. Gottschall states that the veins are vertically
transcurrent in species of Charianthus, Clidemia, Henriettea, Leandra, ~[iconia,
Tetrazygia, and Tococa ; according to the same author, the more delicate veins
are frequently provided with a distinct parenchyma-sheath, which is occa-
sionally sclerosed. Gottschall and Palezieux also mention the occurrence of
more or less elongated rod-cells in the parenchymatous ground tissue of t~e
veins in species of numerous genera 1, while Gottschall records spindle-shaped
sclerenchymatous cells in the same position in Miconia rhytidophylla, Naud.
In the Tribes recently investigated oxalate of lime is also for the most
part deposited in the form of clustered crystals, which vary in size and shape.
They are found in the mesophyll, the veins, the hypoderm (e. g. in species of
Blakea, Dichaetanthera and Melastoma), and more rarely also in the epidermis
(embedded in the stron§ly-thickened inner wall in Micon£a campestris, Tr.,
according to Gottschall ; and in the epidermis of the veins in species of
Dichaetanthera, according to Palezieux). The large cells of the palisade-tissue,
which are occupied by clustered crystals, commonly penetrate between the
epidermal cells after the manner of a wedge, and occa.c;ionally give rise to
transparent dots in the leaf. A peculiar arrangement of the clustered crystals
has been observed in the leaf in species of Blakea; the crystals are contained
in three horizontal layers (viz. (a) in the hypoderm or in the hyer of palisade
situated beneath the upper epidermis, (b) in the layer of cells adjoining the
palisade-tissue, and (c) in the cell-layer lying above the ]ower epidermis).
Sphaerites are found in the bases of the shaggy hairs (I Zottenfiisse') also
in the Osbeckieae and Rhexieae. Other noteworthy features are as follows:
the occasional reduction of the clustered crystals to a short and thick solitary
crystal (especially in lJ.!icon£a atrata, Wawra and Ossaea micrantha, Mad.);
the presence of small monoclinic crystals in the cells containing clustered
crystals in Miconia buddleioides, Tr. and M. tricnotoma, DC., and the styloids,
recorded by Gottschall, in Calycogonium squamulosum, Cogn.
The hairy covering is again very manifold in the Tribes investigated by
Palezieux and Gottschall. The clothing hairs are represented by the same
1 1'hese genera are: Anplectrum, Behuria, Clidemia, Conostegia, Dic"adanthera, DimuMeta,
Hmritttella, Hu/Jeria, Leandra, Lortya, Marulllia, A-liconia, Om/Ita/opus, Pachyloma, Tttra~J'lia,
TDCDetJ, Vtpre(dkz.
t Gottschall mentions the occurrence of cells containing clustered crystals a.nd causing large hemi..
spberical protuberances on the upper side of the leaf in Helerotn"clzum Eggers;;, Cogn. and H. patens,
Cogn., but he does not state whether they belong to the epidermis.
 ADDENDA-MELASTOMACE4E

tyPeS as those previously enumerated on the ground of the older investiga·

hons (see p. 36r). Unicellular or uniseriate tri~"omes are also rare in the
Tribes recently investigated (and consequently in the Melastomaceae generally).
Palezieux records unicellular and uniseriate clothing hairs in Kibessia echinata,
Cogn. and uniseriate trichomes, the ,cells of which are in part ventricose and
often striulate, in species of Sonerila and VepreceUa, while Gottschall describes
unicellular hairs in Miconia plumi/era, Tr. var. Bangii, Cogn. The following
modifications of the uniseriate hairs are found ~trichomes, the ends of which
develop a few ray·cells. among which a club-shaped multicellular gland is
occasionally (VepreceUa microphyUa, f.,ogn.) included (species of VepreceUa);
and trlchomes consisting of a row of rounded cells, the terminal cell being
glandular (Sonerila obliqua, Korth. and S. secundtt, R. Br.). The shaggy
hairs, like those formerly observed, are either parenchymatous or prosenchy·
mato.us; their mode of insertion, as well as their shape and length, varies;
in some cases they are bent in the form of a hook and they may have the shape
of a short awn or wart, &c. In the larger types of shaggy hairs there is a
vascular bundle in the lower portion (species of Majeta, Myrmidone and
Tococa). PaIezieux records shaggy hairs with a papillose epidermis in species
of Dichaetanthera, Dissotis, Osbeckia, Rhodosepala and Tristemm~, brush-like
shaggy hairs in species of Dichaetanthera, Dissotis, Melastoma, Monochaetum
and Dtanthera, shaggy hairs of the abietiform or candelabra·type in species of
Dichaetanthera, Disso cha eta, Dissotis, K ibessia, M arumia, }r[eria1'tia and
Omphalopus, and shaggy hairs resembling the leaf of a moss in species of
Dichal'~anthera and M elastoma. Gottschall describes the following types:-
smooth shaggy hairs composed of two or three rows of prosenchymatous cells,
which are subdivided by two or three thin transverse walls, such hairs
commonly bearing a glandular head at their apex (species of Clidemia and
Tococa); prosenchymatous shaggy hrors with a papillose epidermis (species
of Clidemia, Conostegia, Henriettea, Leandra, Miconia and Ossaea); prosen·
chymatous shaggy hairs terminating in a stellate trichome (species of
Concstegia and Leandra); prosenchymatous shaggy hairs of candelabra· or
antler~like form (species of Clidemia, Leandra and Ossaea); parenchy-
matous brush~like shaggy hairs (species of Miconia and Tococa); paienchy-
matous shaggy hairs with mOle or less pronounced dendroid branching
(species of Conostegia, Leandra and Miconia). The different forms of hairs
just referred to are moreover occasionally (species of CUdemia, Leandra,
Miconia, Ossaea) seated on conical prominences of varying height on the surface
of the leaf, while small shallow pits correspond to these ,prominences on the
lower side of the leaf. The hollow, bristle·like shaggy hairs of Heterotrichum
strigosum, Tr. still require special mention; they are placed with their tips
pointing towards the apex 1)£ the leaf and are adpressed to its surface;,
the epidermis of these hairs is formed by elongated prosenchymatous cells, .
beneath whic.h a layer of short palisade·cells is situated, while the interior
of the hair is ',occupied by a _large intercellular space. Regarding the stellate,
tufted, candelabra· ana peItate hairs observed by PaIezieux and Gottschall
we may mention the following details. PaJezieux records stellate or tufted
hairs with a short, mostly biseriate stalk and a varying number of ray-cells
in species of Anplectrum, Astronia (probably on Cogniaux's authority and
not on the basis of personal observation), Dissochaeta, M arumia {trichomes
occasionally of the candelabra-type) and Omphalopus (trichomes likewise
occasionally resembling candelabra·hairs), while pelt ate hairs occur not only
in the plant investigated by Bachmann, which belongs to Astronia CandoUeana,
Cogn. (not A. papetaria), but also according to Cogniaux in many species of
Astronia. The types of hairs last discussed show much greater diversity of
form among the Mkonieae, where they are often connected by transitional
 ADDENDA--MELASTOMAcEAE
forms, which occur even in the sam~ species or on one .and the same leaf.
Gottschall in the first plal:e mentions stellate hairs wit1l a sMt, mostly
biseriate stalk: and a few ray-cells, which occasionally ihclude a gland, in
species of Calycogonium, Clidemia, Conostegia, HeterotTichum, Ll'4ndTa, Maida,
Miconia, Myrmidone, Ossaea (described as tufted hairs in the special part of
Gottschall's paper), Tetrazygia, and Tococa. The followi1'l.g are special forms
of the stellate hairs :~tellate hairs, in which the ray-cells are swollen in
a bulbous manner at their base (species of Clidemia, Miconia campestris, Tr.) ;
stellate hairs with a very long parenchymatous or· prosenchymatous stalk
(species of Calycogonium, Clidemia, MiC()nia); stellate hairs in which the
basal cells of the comparatively short stalk are divided into compartments by
numerOllS transverse walls (species of Miconia); stellate hairs, the ray-cells
of which have narrow lumina and a serpentine outline (species ·of Clidemia
and Miconia) ; lastly, the pulvinate stellate hairs of Miconia rugosa~ ·Tr. and
M. tomentosa, Don, in which the basal portions of the cells, which radiate
out in all directions, are fused to form a hemispherical cushion. Trichomes
described by Gottschall as tufted hairs', but n6t very sharply distinguished
from the stellate hairs (apparently only by the. larger number of ray-cells),
are found among the Miconieae in species of Calycogonium, Charianthus,
Clidemia, Conostegia, Mironia. and Tetrazygia. Candelabra-hairs with ray-
cells, which are horizontal or occasionally even point upwards, are present
in species of Clidemia, Conostegia, Miconia and Tetrazygia. According to
Gottschall, distinct peltate hairs occur in Miconia tulva, DC., M.lepidota, DC.
and M. tiliaetolia, Naud. (in the :first two species with long stalks; in the
third species with stalks of varying length),' and in Tetrazygia bieolor, Cogn.
(in small pits in the surface of the leaf). They are derived from tufted hairs,
tn which the upper ray-cells have undergone reduction; these abbreviated
ray· cells are seated on the top of the shield, and in some cases form a more
or less distinct small ' upper scale, which in ~f. tulva even includes an external
I

gland; in the same way the uppermost tier of cells in the stalk occasionally
(M. lepidota) grows out to form a small' lower scale.' Tufted hairs resembling
small scales occur also in other species of Miconia, as well as in Calycogonium
squamulosmn, Cogn. and HenrietteUa Mactadyenii, Tr. It remains to mention
that all the diverse forms of clothing hairs found in the Miconiea~ (with the
exception of the hollow shaggy hairs of H eterotrichttm and the bristle-hairs of Caly-
cogonium Krugii, Cogn., and a few species of Ossaea) occur also in combination
with glands; in the Tribes investigated by Palezieux such combined forms
of clothing and glandular hairs have been demonstrated only in Veprecella
(see above). We shall return to these intermediate forms once more below.
, The following details may be mentioned regarding the external glands.
The species belonging to the Tribes examined by PaIezieux for the most part
have small, multicellular external glands, which are either davate or exhibit
a distinctly demarcated stalk and a multicellular head. Uniseriate glandular
hairs in which the cells of the stalk are ventricose and frequently striulate,
are found in Sonerila; glands with a long stalk and a bicellular head, divided
by a vertical wall, occur in species of Allomorphia, Medinilla and Oxyspora;
glandular hairs with a disc·shaped head divided by vertical walls in Barthea
chine1tsis, Hook. f. (glandular disc composed of four cells), species of lIuberia
(glandular disc composed of a larger number of cells, occa.<;ionally with a lobed
margin) and species of GraUenrieda (glandular disc placed excentrically on the
stalk); vesicular integument'll glands, similar to those found in the Labiatae, in
Blastus cochinchinensis, Lour. Palezieux mentions the occurrence of glandular
shaggy hairs only in species of Rhexia and Sonerila. According to Gottschall, the
external glands found in the Miconieae exhibit a much greater diversity of shape.
Uni- or biseriate, filiform external glands have been observed in species of
 ADDENDA--MELASTOMACEAE 'P7
Henriettea~ MscatIUI ud TetrazYgsa. 'Quite a special form of hair is constituted
by the u~e'riate' tdQrt-shapecJ..glands (hydathodes?) of the species of Bellucia;
the ventricose porH~tn of these glands ~iDserted among the epidermal cells and is
divided by a lar~e ftUmber of thin transverse walls into low and broad cells,
which pass avel row cylindrical cells in the region of the neck, while the apex
of the trichome is occupied by a slightly spherical terminal cell; the cells of
the neck and of the ventricose portion have thick walls except for the trans-
verse walls in the latter and for small rounded areas which remain un-
thickent'd in the middle of the transverse walls of the lower cells of the neck;
the uppermost cells of the neck and the terminal cell have thin walls. Bent
uniseriate glandular hairs with a cylindrical head and bearing a few backwardly
directed papillae at the point of bending are found in species of HenrietieUa,
Lo,eya, and 1I1yriaspora; biseriate, club-shaped glandular hairs occur in species
of Calycogoni1tm, Clidemia, Hetcrotrichum, Maieta, Miconia, Myrmidone and
Tococa. Here we may also include glandular hairs, the head of which is bent in
relation to the biseriate stalk in a more or less geniculate manner, while the stalk
often bears one or more ray-cells. The head in these hairs exhibits the following
types of structure; it is :-small, oval or spherical, either unicellular or bicellular
owing to the presence of a division-wall in the plane of bending of the stalk, in
species of Lea'mira, Meconium (?, d. Gottschall, p_ 137), ~iico1Zia, Ossaea and
Tetrazygia; ellipsoidal, clavate or cylindrical, and multicellular in species of
Calycogonium, CUdemia, Conostegia, Heterotrichum, Leandra a1_!4 Tetrazygia;
ligulate and hi- to multicellular in species of Clilemia, Leandra, Miconia
and Ossaea; disc-shaped and multicellular in species of Conostegia; reniform
to spathulate and multicellular in species of Micon·ia (in certain specIes,
especi3.l1y M. quadrang1.£larts, Naud., the stalk bears hairy appendages exhibit-
ing dendroid branching); lastly, cordate and multicellular in species of Leandra,
Other noteworthy forms are constituted by external glands with a multi~
cellular head resembling a blackberry in' shape (species of Miconia and Tetra-
zygia) and glands in which four or more multicellular heads are borne on
a common stalk (Calycogonium squamulosum, Cogn. with four heads, which
are fused to form a rosette; Henriettella Macjadyenii, Tr. with 2-5 heads~
which are disc-shaped and include gelatinized cells in the glands situated
above the veins; species of Charianthus with a varied number of heads corre-
sponding to the number of rows of cells in the stalk; jt!';conia annulata, Tr.
with several reniform or spathulate heads). /It remains to mention that the
two basal cells of the stalk in the glandular hairs of certain species of lI-ficonia
a.re transversely septate by means of a large number of walls. To return
to the combined clothing and glandular hairs found iIi the Miconieae we
may in the first place ·point out that the external glands are very commonly
combined with stellate, tufted, candelabra, and peltate hairs. Smooth
glandular shaggy hairs with a terminal external gland occur in species of
Clidemia, H enriettea, H eterotrichum, M ajeta~ ~1iconia, ~1ynnidone, Ossaea and
Tococa; the gland is either uniseriate and filiform, or biseriate and club-
shaped, or large_ and"'spherical -to-ellipsoidal and provided with a palisade
epidermis. In species of lt1ajeta, Myrmidone and Tococa the stalk of these
glands contains a vascular bundle, while in Heterotrichum Eg~ersi£, Cogn.
and l.1iconia rhytidopkylla, Naud. it includes pitted fibrous cells. Finally.
shaggy hairs of the candelabra type provided with a terminal gland are found
in species of H enr£ettea and H enriettella.
For the structure of the ant-domatia of Majeta. l'IJyrmidone and Tococa, see
Gottschall. loco cit., pp. 27. 28.
3. STRUCTURE OF THE AXIS. Formation of phelloid·cork take? place also
in the Melastomaceae (F. E. Weiss and Gunther, ll. cc.). According to Van
 ADDEN[JA-MELASTOMACEAE

Tieghem, interxylary phloem i$ found also 10 the wood of the root in 'flrI emerylon
ramiflartlm.
Literature: Went, Haft· u. Nahrwurzeln, Ann. Jardin Buitenzorg, xii, 1895, p. 57.-Jo05son,
Anat. Bau d. Bl., Acta Univ••Lund., xxxii, 'I,' J896.-Pale?ieux, Anat.-syst. Untersnch. d. BI. del'
Melastom. mit Ausstbluss der 'rriben det Microlicieen, Tibouchineen und Miconieen, Bull. lterbier
Boissier, viifl899, App. v; Diss., Munich, 85 pp., 3 Tab.-Gottschall,Anat.-syst. Untenouch.d. m.
d. Melastom. :iUS det Tribus der Miconieae, Mem. Herbier Boissier, 1900, n. 19; Diss., Munich,
175 pp., 3 Tab.-Pitard, Pencycle, These, B()r~aux, I90I, p. 68.-Fabricius, Lau.bbIatt-Anaf:.,
Beih. z. bot. Centralbl.. xii, 190"', pp. 328-:J9.-GUnther, Anat. d. Myrtifloren, Diss.• Breslau, 190.5,
pp. 26-:;I7.-Holtermann, Einflnss des Klimas etc., 1907, pp. 13+-135 (Kendrickia, lJ-Iedinilla).-
[For further literatnre, see p. II7I.]

LYTHRARIEAE (pp. 36g-373).

I. THE REVIEW OF THE ANATO!dICAL FEATURES requires the following
addi tions. The cork is for the most part differentiated as phell aidecork. There
is nQ uniform and characteristic type of stoma in tHe Lythrarieae. Gelatinized
cells are occasionally found also in the tissue accompanying the veins. Oxalate
of lime is excreted also in the form of sphaerites or of small acicular or other-
wise shaped crystalline bodies. As regards the hairy. covering we may point
out that small external glands do not occur in this Order, that short, uni- or
bicellular papillose hairs are common, and that slightly branched multicellular
clothing hairs are present in Decqdon; the tufted hairs found in Lagerstroemia
are accompanied by trichomes exhibiting sympodial 'branching or resembling
a fir-tree. To the previous enumeration of special features given at the end
of the general review the following may be added :-the occurrence of division-
walls in the epidermis of the leaf; presence of hypoderm in the leaf (also in
species of Ginoria and Lagerstroemia); formation of papillae on the epidermis
of the leaf (also in species of Diptusodon and Lagerstroemia) ; crystal;idioblasts
with clustered or solitary crystals in the mesophyll; groups of cells containing
clustered crystals in the mesophyll (Pemphis); occurrence of ordinary solitary
crystals (Lagerstroemia lanceo/ata, Wall.), and of sphaerocrystalline or tufted
masses of an unknown substance (species of Decodon, Diplusodon, Heimia~
Lagerstroemia, Lythrum) in the epidermis of the leaf.
2. STRUCTURE OF THE LEAF. Eberlein's recent investigations, on wruch
the following ilescription is ba5ed, deal with the gertera Lythnem, W ood/ol'dz'a,
Pleurophora, Galpinia, Pemphis, DiPlusodon, Physocalymna, Lajoensia, Crenea,
Nesaca, Heimia, Decodon, Grislea, Adenaria, Ginoria, Lagerstroemia, and
Lawsonia.
The statement made in the earlier portion of this work that there is no
special type of stoma requires modification in the sense that stomata of a special
type are not of general occurrence. In Lythrum nummulariijoliwn, Lois.,
however, the pairs of guard-cells are accompanied by three neighbouring
cells, which are all of different sizes; in Pleurophora and Heimia myrtijolia,
Cham. et Schlecht., they are surrounded by three ordinary neighbouring cells ;
in Crenea there are commonly four neighbouring cells, and 50 on; in Lajoensia
punict'lol£a, DC. and L,lgerstroemia spec~·osa, Pers., the neighbouring cells are
.narrow, while in Sonneralia, according to Areschoug, they appear arranged
in a rosette. The stomata are either ah~ent on the upper side of the leaf or
are present in varying numbers; they either lie on a level ~i1h the 'epi9.erruis
or are slightly raised above it, or are depressed; occasionally pairs1 of guard~
cells of two sizes are found on one and the same surface of the leaf; i~ some
cases (species of Diplusodon, H eimia, Lythrum, N esaea~ Pleurophora) the
stomata are, placed with their pores directed approximately parallel to the
principal vein. The degree of thickening presented.by the outer wall of the
 ADDEND~-L YTHRARIEAE

epidermis and the varied nature of the cuticle. which is occasionally granular
o.r s~riated, furnish characters !or spec~c di~gnosi3; a note~orthy feat~re
lies m the occurrence of what IS called an mternal granulatIOn' m speCles
of DiPlusodon. Eberlein demonstrated gelatini$ation of the epidermis of the
leaf in certain species of all the genera investigated hy him with the exception
of Grislea. and' Woodfordia. In certain species of Ginoria, Lagerstroemia and
Pemphis the upper epidermal cells undergo local divisions by means of hori~
zontal 'w~ls. -Hypoderm is developed on the upper side of the leaf also in
Ginoria Rohrii, Kohne (one-layered) and Lagerstroemia- lanceolata, Wall.
(I-2-layered). Typical papillae are found on the lower surface of the leaf also
in Diplusodon epilobioides. DC. (papillae here short and pectinate, and serving
as centres for the cuticular striation), as well as in Lagerstroemia hypoleuca,
Kurz, L. lanceolata, Wall. and L. parvifiora, Roxb. The mesophyll varies
from bifacial to centric in structure., Sclerenchyma mayor may not be present
in the veins. In species of Decodon, Grislea, Lagerstroemia. and Physocalymna,
ev~n the smaller veins· are vertically transcurrent. In the larger veins the
lower group of soft bast in the bicollateral vascular system occasionally exhibits
redu~tion, so that only that situated on the upper side (the inner bast) is
strongly developed. Another striking feature observed in certain species is
the occurrence of numerous mechanical elements or of spiral tracheae with
wide lumina in the wood of the .larger veins. Storage and terminal tracheids
in the form of enlarged pitted cells are found in species of Crenea, Diplusodon,
and Lawsonia. Gelatinized cells occur als0l in the tissues accompanying the
veins in species of Diplusodon, Lagerstroemia (here also in the hypoderm)
and Nesaea.
Oxalate of lime, as already previously stated, is generally deposited in
the form of clustered crystals. The latter vary both in size and structure;
they are either star-shaped or built up of small and delicate needles; in other
cases again they are more of the nature of sphaerites or consist of granular
con~lomerates of individual crystals. In addition to these forms we have
tYPICal sphaerites (especially.in Pleurophora, also i~ Grislea and WoodI01dia),
ordinary solitary cry~tals of the rhombohedral type, or presenting some' other
phape, and small crystalline bodies assuming the shape of rods, needles or
granules; the latter are found both in the mesophyll and in the integumental
tissue (in species of Adenaria, Decodon, DiP!,usodon, Heimia, Lafoensia, La'W~
sonia, Lythrum, Nesaea, Physocalymna, Pleu1'ophora, WoodI01'dia). Other
features requiring special mention are: the occurrence of small groups of
cells filled with sma}l clustered crystals in the m~ophyll, which is specially
characteristic of Pempkt"s (P. addu/a, Forst.); the occurrence in the mesophyU
of largf' idioblasts, containing clustered (species of DiPlusodon, Galpinia and
N esaea) or large solitary crystals (Lafoensia nummulariifolia, St. Hil., and
certain species of Lagerstroemia), these idioblasts in some cases giving nse to
transparent. _9.0~s in the leaf (in Lagerstroemia, -Kohne's (folia glanduloso-
punctata '); -lastly, the presence of relatively large solitary crystals in the
epidermis in Lagerstroemia lanceolata, Wall., these crystals being enveloped
by a cell~membrane in the same way as in Rosanoff's crystals. Before leaving
the discussion of the modes of excretion of oxalate of lime, we may refer to the
sphaerocrystaUine, tufted or variously shaped masses of an unknown substance,
which Eberlein observed in the integumental tissue in certain species of
Decodon, Diplusodon, Heimia, Lagerstroemia and Lythrum (in Lythrttnt both in
the dried and in the living 1eaf).
Am{'og the types of clothing hrurs found in the genera recently investi-
gated, simple unicellular or uniseriate forms are again the most important.
Short unicellular papillose hairs, which are occasionally divided into two
cells by a transverse wall, are very common (species of Adenaria, Decodon,
so(..£aEo£a 3 0
930 ADDENDA-LYTHRARIEAE
DiPl~ don, Grislea, Ginoria, Lagerstroe-mia, Lythrum, N esaea, Pleurophora, Physo·
calymnal Wood/ordia). The longer hdlrs occurring in this Order are either
unicellular or uniseriate; the latter may have thin or thick division-walls, and
their com.ponent,cells ife occasionall} articulated with reference to one another.
The surtace of t.e clotlung hairs is not uncommonly granular or verrucose.
The appearance of relatively thin transverse and longitudinal walls in the
lower portion of the 'body of the hair jt, Lagerstroemia indica, L., leads to the
production of shaggy hairs. The species of Lagerstroemia belonging to the
section Trichopodium have branched hairs. In L. Engleriana, Kohne, these
hairs are of sympodial structure; their main stem is uniseriate and consists
of a t'arying number of cells, which, with the exception of those at the base,
are 4lrawn out into rays emanating in various directions and more or less bent;
in this way forked o~ antler-shaped hairs are produced. Among these .trichomes
we may mclude certain hairs found in other species of Lagerstroemia·;. the
latter are either tufted or abietiform and consist of a miseriate stalk and
a varying number of tiers of cells, which are drawn (Jut into rays. Little
branched multicellular clothing hairs, which are either forked or antler-shaped,
are found side by side with unbranched trichomes in Decodon tJerticillatus, Ell.
In some cases (DiPlusodon, Physocalymna) the surrounding cells are prolonged
on to the clothing hairs after the manner of subsidiary cells. The leaves of the
species of Grislea bear black glandular dots which had not previously been
subjected to a close examination; according to Eberlein, they are due to
shortly stalked external glands which have a oneelayered wall and a large
intercellular secretory space in their interior, and are thus quite similar to
the glands found in the genera Adenaria and W ood/ordia 1. Small external
glands- are completely wanting in this Order.: Areschoug mentions the occur-
rence of cork-warts on the leaves in Sonneratia caseolaris, Eng!. and S. [anceo-
lata, Miq.
3. STRUCTURE OF THE AXIS. Gunther has recent1y examined the structure
of the axis, more especially with reference to the development of the cork, in
the following genera: Rotala, Ammannia, PePlis, Adenaria, Woodfordia,
Cuphea, Lythrum, Pleurophora, Nesaea, Heimia, Decodon, Crenea, Ginoria,
Pemphis, Diplusodon, La/oensia, Physocalymna, Lawsonia and Lagerstroemia
The vascular bundles are invariably bicollateral, and the perforations of t~
vessels are always simple. The groups of pericyclic fibres are occasionally
united to form a ring (species of Ammannia, DiPl~tsodon, La/oensia). In
Laloensia Vandelliana, Cham. et Schlecht. the transverse section shows several
mechanical rings composed of bast-fibres intermingled with stone-cells. In
Lagerstroemia and Diplusodon the cells of the pith are strongly sclerosed, while
in Adenaria floribunda, H. B. K. there are isolated stone-cells ttl the pith.
According to GUnther, the concentric arrangement of the crystal-cells (contain-
ing solitary or clustered crystals) in the bast, as seen in a transverse section of
the branch, is a feature deserving special notice.
To the previous account (on p, 372) of the cork we may add the following
details. The cork develops in deep layers' (probably in most cases in the
4

pericycle), also in Cren~a, Cuphea, Diplusodon, Ginoria and Heimia, The

cells of the cork are for the most part tabular, but in some cases they are
cp.bical; they are very small in species of Ammannia, Crenea, Nesaea and
Peptis. GUnther distinguishes the following four types of differentiation of
the cork. Crenea surinamensis, Kohne and P~plis Port'ula, L. have a single
layer of cork, which may be regarded as an endodermis, whue on its outer.l
side there is a massive primary aerenchyma. Decodon verticillatus, Ell. has

1 The cause of the 4 folia utraque pagina nigrOepunctulata • {oued in Pem/lris tnadagascariem;s,
Kohne still remains to be determiued,
 ADDENDA-LYTHRARIEAE

a normal many·layered periderm, on the outer side of which primal.!.' aeren

chyma :is again strongly developed. Typical cork consisting of a ~tdl large-t
number of layers is found in Pemphis aciiluZa, Forst. and PleuTophora pungens,
Don. In the remaining members of the {)rder, inv6Stigated by Gfutther, the
cork contains phelloid·cells (see also J. E. \Veiss, loco cit.). The cells given off
hy the phellogen on its outer side undergo subsequent- division by means of
tangential walls so that each gives rise fo three cells; of these only the middle
one develops into a typical cork-cell, while th~ outer and inner cells retaift
their cellulose-walls (Lythrum) or become lignified (Cuphea, Heimia) or sclerosed
(in the remaining genera). In the -fourth type sman intercellular spa~ are
generally present between the cells of the periderm.
Typical (secondary) aerenchyma is developed also in the floating stems of
Nesaea verticillata (Schrenk).
4 .. STRUCTURE OF THE ROOT. The interxylary phloem in the wood of the root
of L-ythyum Sa/£caria is not given off by the cambium on its inner side, but arises
by a process of secondary differentiation in the wood-parenchyma.-For the
structure of the. respiratory roots of Sonneratia, see also Westermaier, loco cit.
Among the results of his investigations we need only mention that there are no
-spicular cells in those parts of the roots which are embedded in the mud at the
bottom of the water, their place being taken by special elongated cells having
thick walls and acting like springs.
Literature: Hahnel, Gerberinden, Berlin, 1880, p. 137 et seq.-Costantin, Tiges d. pI. aquat., Ann.
sc. nat., ser. 6, t. xix, 1884, p. 287 et seq. and pl. xiv.-[Schrenk, Float. tissue of Nmua, Bull.
Torrey Bot. Club, xvi, 1889, pp. 315-23 and pI. xcv-vii; abstr. in Bot. Centralbl., 1890, iii, p. 120.]-
J. E. Weiss, Korkbild., Denkscbr. bot. Gesellsch. Regensbnrg, vi, 1890, p. 6 et seq.-Leisering,
Interxylares Leptom, Diss., Berlin, 1899, p. 30.-KearnYi in Contrib. U. S. Nat. Herb., v, D. 5,
1900, p. 303 (Ammanma).-Westennaier, Pneurnatophoren, Freiburg i. d. Schw., I900, 53 pp.,
3 Ta.b.-Areschoug, Mangrovepfl., Bibl. bot., Heft 56, 1902, pp. 67---9 and Tab. xi-xiii (&nm"atia,
Pemphis).-Bargagli-Petmcci, Legnami, Malpighia, ,1902, p. 355 (Sonneratia).-Kohne, Lytbraceae,
in P.RlUlzenreich, Heft I7, 1903, pp. 4. s.-Eberlein, Bdtr. z. ana.t. Cha.rakt. d. Lythmr., Diss.,
Erlangen, 1904, 78 pp.-Areschoug, Trop. 'Vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2. 1905,
pp. IIg. 120 and Tab. iii and xxii (Lawsonia), pp. 122-3 and Tab. iii, iv (LafomsM).-Giinther,
Anat. d. MyrtitIoren mit bes. .Beriicks. d. Lytbrar., Diss., Breslau, 1905, 39 pp., esp. pp. 5-20.-
Piccioli, Legnami, Bllll. Siena, 1906, p. 147.-Holtermann, Einiluss des Klimas, etc., 1907, p. 92
(P~mpkis).-[For further literature, see p. I1io].

ONAGRARIEAE (pp. 373-376).

",..

In the summary of the anatomical features of the Onagrarieae given at

the beginning (p. 373) of the former description the statement as to the absence
of external ~ands must be cancelled. The simple perforations of the vessels
are accompaaied by a small number of scalarifonn perforations in certain
members of the Order. The cork of Trapa, unlike that of the other Onagra-
rieae, develops beneath the epidermis.
An account of the stnlcture of the leaf, stem, and root in this Order has
been pub1ished also by Grosse. whose work was not taken into consideration
in the previg-us--destfiption. Tile following details are taken from his paper
as well as from the other new literature. The cJosure of the stomata in
Trapa, as in other floating plants, takes place solely by means of the cuticular
ridges on the guard-cells (Haberlandt). Raphides are found also in Euchari-
dium. It remains doubtful whether the small prismatic solitary crystals,
observed by Netolitzky in the apex of the leaf of Isnardia (Ludwigia) palustris,
L, are of constant occurrence. In glycerine·preparations of the leaves of
Oenothera biennis, L., Netolitz\;v met with yellowish crystalline deposits of an
unknown substance. Side 'by side with the raphide·sacs one occasionally
(e.g. in EPilobittm hypericifolium, Tausch) finds mucilage-sacs containing no
raphides or occupied only by a small number of short needles. The unicellular
302
932 ADDENDA-ONAGRARIEAE
clothing balrs comruonly have a granular or verrucose surface. The short
club-shaped ·unicellular hairs of Oenothera Ramanzowii, Ledeb. and Clarkia
pielchella." Pursh. &c., as well as the rather longer unicellular trichomes of
Qencther4 bientlis" L., ~n :Which the apex is slightly swollen, have thin walls.
are fillet! with· granular contents in the living plant and probably have a glandu-
lar function.; Short dub.shaped trichomes are recorded by Netolitzky also in
Circaea, while 10ng ones are stated to occur in Epilobium (see also Theorin,
Ioc. cit.). Structuu~s resembling stipules are found at the base of the petiole
in Lud'llll(~a palustris, Ell., and in the species of ] ussiaea (Meeban), but they
still requre caIeft~1 investigation. In Jussiaea tl~se structures are somewhat
flattened.and .ovate and have a green colour; they do not contain any vascular
bundle9'; although there are numerous raphide-sacs, and they dry up at an
early stage. The' secretOl y receptacles' described by Grosse as occurring
in the axis of Trapa are probably in reality a local pathological formation of
cork. According to Grosse, carotin-crystals are present in the tissue of the
leaf in species of Jussiaea, Godetia and Fuchsia. Sclerenchyma is developed in
the veins of the leaf in certain species of Fuchsia, but is otherwise absent.
Grosse also examined the structure of the wood in the herbaceous
geitera, and found that as a general rule they show· features identical
with those which I demonstrated in the woody m~mbers of the Order, viz.
narrow medullary rays, vessels with simple perforations, and wood-fibres
bearing simple pits. Grosse, however, mentions the occurrence of a relatively
small number of scalariform perforations in the vessels, in species of Oenothera
and Gogetia. '
In many members ot the Order the ti')sue of the cork includes both
suberized and unsuberized (phelloid) cells, and in some cases (species of -Puchsia)
there is a regular alternation of layers of cork-cells and pheUoid-celL~.
These phenomena already give indication of the tendency to develop aeren-
chJrma, a tissue which H. Schent-k has observed also in EPilobium hirsutum, L.
According to H. Schenck the aerenchyma in the aerotropic roots of Jussiaea
is developed from the primary cortex, while according to Grosse it is subse-
quently increased by the development of an aerenchyma from the phellogen.
Interxylary phloem has recently been observed by J. E. Weiss in the
stolons of EPilobium angf/'sti/olium, L., and in the root of E. hirsutum, L .•
and by Grosse L'l the root, in Epilobium palustre, L., further species of Oenothera
and species of Gaura and Lopezia. According to Leisering, Chodat, and Fre·
mont, the interxylary phloem no doubt invariably arises from the wood-
paren~hyma by a process of secondary differentiation (though in some cases it,
appears very soon after the development of the parenchyma from the cambium),
and is not given off directly by the cambium on its inner side, as was formerly
supposed.
~ The vascular bundles in the delicate lateral :roots of T'fapa show a very
remarkable structure (Queva). The radial vascular system exhibits unipolar dif-
ferentiation, i. e. it comprises only a single group of xylem and a single group of
phloem-a type of structure which is unique among the Phanerogams. Both the
wood- and bast-portions are very much reduced, the former consisting of a single
trachea. the latter of 3-10 soft-bast elements. .
Literature: Costantino Tiges aero et Bout., Ann. se. nat., ser. 6, t. xvi, 1883, p. 68 et seq.-
(Meehan, Pet. glaqds in some Onagrar.~ Proceed. Acad. Nat. Sc. Philadelphia, 1886, pp. 349-50;
abstr. in Just, 1~86, ii, p. 705.]-Hahetlandt, Spaltoffn. d. Wasserpfl., Flora, 1887, p. 103.-
Lamounette, Liber interne, Ann. sc. nat., sere 7, t. xi, 1890, pp. 261, 262.-J. E. Weiss, Korkbild. t
Dcnlarchr. bot. GeseUSch. Regensburg, vi, 1890, p. ii et seq.-Grosse, Beitr. z. vergl. Anat. d.
Onagrar., Dis!•• Erlangen, 1895, 61 pp.-[Pannentier, in Mem. Soc. d'emulation Doubs. 1896,
p. ,327; and in Le Monde, 1896, p. 29, and 1897, p. 178.J-Ramaley, Stem·atr~tomy of certain
Onagrar., Bot. Gazette, xxii, 1896, p. 229.-Schubert, Parenchymscheiden, Bot. Centralbl., 1897, iT,
p. 17.-Wollenweber, Anat. d. Schwimmbl., Diss., Freibtttg i. Br., 1897, pp. JO-I.-Spanjer,
 ADDENDA-ONAGRARIEAE 933
Wasserapparate, Bot. Zeit., 1898, i, p. 45.-Leisering, Interxylares Leptom, DiK., Berij,n, 1899.
pp. 29-30.-Minden, Wassenez. Org., Bibl. bo•. , Heft 46, 1899, p. #:-Kearny. in Contril>'I1t.
U. S • Nat. Herb., v, n. 5, 1900, p. 297.-[Guffroy, Papilles chez les Epilobes, Bull, laterrlat. Geogr.
Dot., J901, p. 9.J-[Uveillt', Monogr. du genre Oenothera avec la coll:f.b. po~r la partie anat. de
Geoffroy, Le Mans, Fasc. I, 190:1, 138 pp., and Fasc. :I, 19°5.]-Qu(',va. Ra\1ilitlles <k: h. M4cre,
Comptes rendtls, Paris, cxxxvi, 1903, I. Sem., pp. 8 .. 6, 827; see also Bull. Sqc. d'hist. nat. Autoo,
xvi, 19°3.-Theorin, Vaxttrichom., Arkiv foc Bot., i, 1903. p. J 60; iii, n. 5, 190+, p. 11; ..and iv,
D. 18, 1905, pp. 19-20.-Col, Faisceaux, Ann. sc. nat., ser. 8, t. xx, 190...., pp. fS ....-1 and 209-10.-
Fteidenfeldt, Anat. Baud. Wurzel, Bibl. bot., Heft 61,1904, pp. 64-6.-[Getschon-5eUber, Justiaea
repens, Diss., Halle, 19°5.54 pp. j also in' Nova Acta Acad. Leop., Ixxxiv; abstr. in Bot. Ceptralbl.
cH, p. 2 .........)-Giinther, Anat. der MyrtifioreD, Diss., Breslan, 1905. p. 21-Netolitzk,. Dikotyle'·
donenbl. (Rhaphiden), 1905, pp. 39;-52.-[For additional literature, see p. lI71].

SAMYDACEAE (pp. 376-378).

I. REVIEW OF THE ANATOMICAL FEATURES. The stomata either have
subsidiary cells, arranged according to the Rubiaceous or Cruciferous type.
or are merely surrounded by ordinary neighbouring cells. Schizogenous
secretory cavities are found also in Zuelania; on the other hand, the earlier
record of their occurrence in Lunania pro parte must be cancelled. As regards
the hairy covering we may add that simple uniseriate clothing hairs with
t.hin division· walls are of occasional occurrence, while unicellular Malpighian
hairs are found also in Banara guiane'nsis, Aubl., and stellate hairs also in
B. glauca, Benth. Additional features which can be employed in _specific
diagnosis are as follow: the occurrence of a gelatinized epidermis in the leaf
(only in the monotypic genus Gerrardina); the presence of hypoderm on
the upper side of the leaf (species of Abatia, Banara, Byrsanthus, Casearia,
Euceraea, .Homalittm, and Zuelania); idioblasts occupied by small clustered
or solitary crystals in the epidermis of the leaf (besides occurring in species of
Homalium and Samyda, also in species of Bembicia, Casearia, Lunania, Ophio-
botrys, Osmelia, Pyramidocarpus and Zttelania); the occurrence of sclerenchy-
matous fibres in the mesophyll (species of Calantica, Casearia, Homalium, and
Zuelania); and lastly, the peculiar silicified structures resembling cystoliths
found in the mesophyU of H omalium donquaiense, Pierre. I
2. STRUCTURE OF THE LEAF. Brandlein's recent investigations, which
deal with the genera enumerated belowl, have furnished the following facts.
The epidermal cells have straight or undulated lateral margins. The structure
of the cuticle may frequently be compared with the markings on an etched glass
plate, so that in these cases we may describe it as fetched'; but striation
and ,granulation also occur. The outer and inner walls vary in thickness,
the latter occasionally presenting a swollen appearance, although they are
gelatinized only in Gerrardina loliosa, Olivo Marginal pits are not uncom~
mon, and the same statement applies to the occurrence of relatively thin
secondary division-walls, both horizontal and vertical. The surface-view
of t_he epidermis is particularly striking in (a) the epidermis of the leaf -of
Banara -brasiliensis; :Benth. "(owing to the penetration of the thick cuticle into
the lateral walls in the form of strongly developed wedge-shaped ridges),
(b) the lower epidermis of Banara portoricensis KIUg et Urb. (here the stomata
t

with their thin-walled subsidiary cells appear as islands in the remaining

tissue, which has thick and pitted walls), and (c) the epidermis of Casearia
densiff,ora, Benth. (the lateral walls of which appear thin at a high ioc.tis, being
provided with peculiar lobes, which are cuneate and have a narrow base and an

1 These genera are: Tribe I : Casearieae: Casearia, Zuelallia. Osmdia, Euceraea, Lunania,
Tetrathylacium; fJjJhic/Jofrys, Samyda j Tribe:l: Banareae; Banara, P)'ramidocarpus; Tribe 3:
Abatieae: Abat£a, Aphaerema; Tribe 4: Homalieae: Calanlica (incl. Bivinia)) Gerrardina,
Dissomeria, Homalt'u1JI, Byrsanthus, BemOida.
 934 ADDENDA-SAJIYDACEAE

emarginate apex; at· a low focus, On the other hand, they appear straight,
and are rather thick and pitted). The occurrence of hypoderm has already
been mentioned above; an enumeration of the species jn which it is found
is given below l . The hypoderm for the most part consists of a single layer of
celis, although it jg composed of three Of four layers in Homaliu1n tomentosunz.;
it is confined to the upper side of the leaf. In Euceraea nitida the cells of the
hypoderm, as well as the majority of the cells of the spongy tissue, are provided
with hippocrepiforrn thickenings on the side facing the middle of the trans-
verSe secti~m of Hie leaf. In some species of Casearia and Lunania small
numbers of stomata occur on the upper side of the leaf. The Cruciferous
type is distinctly differentiated in Osmelia, and the Rubiaceous type in Abatia
(here occasionally ill-defined), Banara, Bembicia, BY1'santhus, Calantica,
Casearia, Dissomeria, Euceraea, HomaUum, Samyda and Zuelania. The
structure of the leaf varies from bifacial to centric. The palisade-tissue cQn:
sists of one or more layers. In Abatia there is invariably a single layer of
palisade· tissue, which exceeds the spongy tissue in thickness. An indication
of arm·palisade-tissue has been observed in the third p'alisade layer of Casearia
ar~uta, H. B. K., while the cell~walls in the lowest palisade layers of the centric
mesophyll of C. corymbosa, H. B. K. show peculiar sinuations appearing as
ridges and giving rise to '·pseudo-pits.' In Homalium 1QCemosum, Jacq. the
lowest layer of the mesophyll presents a palisade-like differentiation and is
developed in the form of conjugate parenchyma composed of short cells. The
intercellular spaces of the spongy tissue vary in size, and the walls of the cells
are thickened to a varied extent. In'many species of Lunania the upper layers
of the spongy tissue are rather strongly thickened and filled with brown con~
tents, so that they form a kind of middle layer lin the leaf. Peculiar silicified
protuberance~ resembling cystoliths occur in Homalium donquaiense, where
they are found especially in the palisade-tissue and conespond with one another
in neighbouring cells. The vascular system of the veins is in most cases provided
with a thick ring of sclerenchyma (the chief exceptions being Abatia and
Apharema, which have little or no sclerenchyma). The sclerenchyma of the veins
occasionally exhibits well-marked stratification of the wall (species of Casearia
and Homalium) or consists of an outer envelope and a detached thickening layer.
In the species enumerated below 2 the sclerenchyma of the veins branches off
into the mesophyU to a varying extent. These spicular fibres also occasionally
show well-marked stratification; in some cases they penetrate as far as the
epidermis and may even continue their course beneath the latter. Bdindlein
observed vertical transcurrence of the smaller veins (lateral veins of the third
order), which in some cases are provided with perfect plates of sclerenchyma,
in species of Abat£a, Banara, Byrsanthus, Calantica, Casearia, Homalium and
Samyda.
The following details may be added regarding the hairy covering. Uni-
cellular clothing hairs, which for the most part have thick walls and narrow
lumina, have been recorded in the genera A bat£a, A phaerema, Banara, Bembicia,

1 The species are: Abatia lomen/osa, Mart. (locally); Batulra gta'anensis, Aubl. (locally);
B7na1tthus sp. (Demease, Congo); Casearia Bule, Gilg, C. comocladi/olia. Vent., C. corymoosa,
H. B. K., C. glomerata, Roxb., C. graveolens, Dab., C. ilicifolia, Vent., C. lucida, Tu}., C. Mcli~
slaut'U11Z, Spreng., C. fIlollis, K. Sch. var. glabra, K. Sch., C. 1tilt"da, Jacq., C. rubescens, Dalz.
(locally), C. lommtosa, Roxb., C. Zenkeri, Gilg; Euceraea nit£da, Mart; Homaliu11l Barandae,
Vid., H. brevifrdu1tculatulll, Scott~EIl., H. .Dejlanchei, "Tarb., H. '"tU'1Yplerum, GiIg, H. /(/1/(m-
tOSU'll, Benth., H. vilienu, Benth., H. Zmken', Gilg; Zuelallia crtnata, Griseb., lattz"oides,
A. Rich.
I Calantii:a cerasifolia, Tul.; Casear£a attenuata, Rusby, C. longicuspidata, Gili!', C. Selloana.
Eichl. (short fibres), C. sylvestn's, Sw.; Homah"um brevlpedunw/alum, Scott-Ell., D. micran/hunt,
O. Hoffm., H. mic1'ophyllmll, O. Hoffm., H. panimlatum, Benth., H. Parkeri, Bak. (short fibres),
H. urceolatum, Scott.Ell. ; Zttdania laetioides, Rich. (short fibres).
 ADDENDA-SAM YDA CEAE 935
Calantica, Casearia, Euceraea, Homalium, Osmelia, Sa tnyda , Tetrathytacium
and Zuelania, while uniseriate hairs which are septat~ by means of relatively
thin division-walls occur in species of Casearia, Samyda and Zuelania. The
trichomes of Banara guianens'is, Aubi. show all transitions betweeR .ordinary
unicellular hairs and Malpighian hairs with equal arms. f ' Diverse types of
tufted hairs having two or more ray-cells with narrow lumfna are found also·
in Abatia boliviana, Britt., A. parvifiora, Ruiz et Pav., Banara glauca, Benth.,
and certain Central and South American species of Casearia belongill.£ tQ the
section Pitumba. In the two species of Abatia just named th8'\ wficellular
and tufted hairs are accompanied by other types of tufted hairs, which have
a longer, multiseriate shaggy stalk; the protrusion of a_ certain number of the
superficial cells of the stalk of these trichomes into rays leads·to the piOduction
of candelabra-hairs.
There is no oxalate of lime in the leaf in Abatia, Aphaerema, Banara.
pyramidata, Rusby and Casearia attenuata, Rusby. Small crystal-idioblasts are
found in species of many genera 1; they occur either in both upper and lower
epidermis or only in the one or the other and are either isolated or arranged
in rows or groups; in most cases each of them contains a clustered crystal,
rarely a solitary crystal (the presence of solitary crystals is indicated in the
list below by the abbreviation cr.). The crystals are .occasionally enclosed in
an envelope of cellulose after the manner of Rosanoff's crystals, or they may
be suspended from beams of cellulose. Other noteworthy features are: the
occurrence of complete layers of cells containing clustered crystals in the
mesophyll (species of Casearia and Homalium); the occurrence of relatively
l~ge idioblasts with clustered crystals (species of Casearia), and the presence
of ceUs exhibiting one-sided thickening and enclosing solitary crystals (' cris-
tarque '-cent), in association with the sclerenchyma in the veins of the leaf
(species of Casearia and Homalium).
Secretory cavities have been demonstrated in Casearia pro parte, Euce1.,J}ea,
Samyda and Zuelania. Although Bokorny and \Varburg record secrefory
cavities in Lunania, Brandlein did not meet with these or any other types of
secretory organs in any of the seven species which he examined. The earlier;
statement as to the occurrence of secretory cavities in Osmelia (according to
Warburg) also requires confirmation, since they are absent in Osmelia Ma~·ngayi,
King, accordinito Brandlein. According to Brandlein (d. the earlier statements),
all the species _of the genus Casearia belonging to .the section Piparea (with the
exception of C. eriophora, Wr.), as well as C. cuspidata, Gilg and C. rubescens,
Dalz. (which belong to the section Pitumba), are characterized by the absence
of secretorYI cavities. Brandlein states that the secretory cavities invariably
have a distinct epithelium. Their yellowish and strongly refractive contents
are soluble in alcohol, so that they are of the nature of resin. In Casearia
sylvestris, Sw. secretory cells occur in the mesophyll side by side with the
secretory cavities.
-----------------------------------------------------------------
1 viz.: Bembida axt.ll~ris~ Oliv-:";- Casearia bic%r, Urb., C. Bule, Gilg, C. comodadiflJ/ia, Vent.,
C. corymbosa, H. B. K., C. tientafa, EichI. (cr.), C. (scu/mfa, Roxb., C. glomerata, Ro,c:b., C.
graveo/ens. Dalz., C. guiancnst's, Rusby, C. MYla, Sw., C. ilicijol£a, Vent., C. Lobbiana, Turez.• C.
maCYbpkylla, Vahl, c. ~felistaurum, Spreng., C. moms, K. Sch. var. glabra, K. Sch., C.oblongifolia,
Camb., C. bOovata, Poepp., C. ramijlora, Vahl, C. rubescens, Dalz., C. fomenlosa, Roxb., C. Vareca,
Roxb., C. ZenNeri, Gilg; Homalium dmsijlorum, Benth. (cr., side by side with clustered crystals),
H.jodidum, Benth., H. pedicel/alum, Benth., H. raU1IlOSU1Jl, Jacq., H. Racoubea, Sw.; Lunania
(usjidala, Warb., L. dit!rJ'ricata, Benth., L. dodecandra, \Vright, L parvijlora, Spruce, L. racemosa,
Hook .• L. Sauvalii, Griseb.; Ophz'obotrys Zmkeri, Gilg j OSlllelia fi/aingayi, King; PY'Yamiaocarpus
Blacki;, Oliv.; Samyda glabrala, Sw., S. gralldijlora, Griseb., S. rose a , Sims., S. serru lata , L.;
Z.ulania cre1flJ/a, Griseb., Z. ltulioidts, Rich.
I Tbe species are: Cauan'a Bri'chami', Wats., C. Commersoniana, Camb., C. densijl()ra, Benth.,
C. dentala, Eiehl., C. ja'lJilensis, H. B. K., C. lae/£oides, Warb., C. /aun/olia, Benth. (e:u.mined by
Harms) C. MtU~'miHani, Eichl. and C•.S'prucea1Ul, Benth.
 ADDENDA-SAMYDACEAE

3. STRUCTURE OF TilE AxIS. Regarding the occurrence of a composite

and continuous rini of sc1erenchyma, which also includes cells with U-sbaped
thickening, in the pericycJe in species of Casear-ia, Homalium and Samyda,
I as well as of an interrupted ring in the pericycle in species of Bana,a and
Casearia, see also Pitard, loco cit.
Literature: Elfstrand, Heilpfl., Ber. deutsch. pharm. Gesellsch., 1897, p. 3Ii.-Pitard, PericycIe,
These, BordeaU:lt, 1901, pp. 67-92.-[Pitard, Astetopeiees, Act. Soc. Linn. Bordeaux, lviii. 1903,
p. Hi et seq.]--Briindlein, Syst.-anat. Unterstlch. d. Bt der Samydaceen, Benth. et H"ok., DiSl.,
Erlangen, 1907, 69 pp.

TURNERACEAE (pp. 381-383).

Literature: Pitard, Pericycle, These, Bordeaux, 1901, p. 93.

PASSIFLORACEAE (pp. 383-388).

2. STRUCTURE OF TlIE LEAF. Papillae on the lower side of the leaf" are
found also i!l Passiflora trilasciata, Lem. (Knothe). Jumelle has recently
also demonstrated internal secretory receptacles (see p. 38S) in the axis and
leaf of the newly established species, Ophiocaulon Firingavelense, Dr. de Cast.
He describes them throughout as secretory cells, and states that they give
rise to the wax-like or more properly (as he points out) resinous masses, which
are repeatedly mentioned by systematists in their descriptions as covering the
different parts of the axis in the species of Ophiocaulon.
Extrafioral nectaries are, according to;Harrns, present on the leaves also
in Paschanthus, HoUrungia and Tetrastylis.
LiteratUre: Aufrecht, Extrafior. Nekt., Disl., Zurich, 1892, p. 29 et seq. (Passifora c()erulea).-
Pitard, Pericyde, These, Bordeaux, 1901, pp. J9aJld 77.-Kllothe, Unbcnetzb. .B1., Diss.,Heidelberg,
J9()l, p. 14.-Jumelle, Passifiote a resine. Comptes rendus, Paris, cxxxvii, 1903,:3 Sem., P,P' ~06-8.
-Col, Faisceaux, Ann. sc. nat., ser.8, t. JOt, 1904, p. I09.-Paoli. Eterofillia, Nuovo GlOm. bot.
Ita!., xi, 1904, p. :n6.-[Hann s, in Natiirl. Pflanzenfam" Erg.-Heft ii, 1907, p. ~34.]

PAPAYACEAE (pp. 388, 389).

Literature: A translation of Schacht's paper will be found in Ann. sc. nat., ser..... , t. viii, 1857.
p. 16.f. et seq. and pI. vii, viii.-Moliscb. Milchsaft u. Schleimsaft, 1901, p. 60.-Areschollg, Trop.
vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2. 1905, pp. 30-1 (Carica).

CUCURBITACEAE (pp. 389-397).

I. R:EVl£W OF THE ANATOMICAL FEATURES. Glandular hairs with rela-
tively long 5 talks are also found in this Order. Extrafloral nectaries are present
on the leaves in certain species. The bundles of intraxylary phloem become
secondarily changed into inversely orientated vascular 'bundles also in certain
'species of Coccinia, Cucurbita, Kedrostis and M elothria. In the older parts
of the axis of Actinostemma biglandulosum, A. racemosum and Momordica
Charantia secondary vascular bundles are developed from a secondary meristeni
arising in the endodermis.
2. The STRUCTURE OF THE. LEAF in the Japanese species 1 has recently.
been examined in detail by Yasuda. The epidermal cells in these species are
often of large size (the largest being .07 mm. in diameter in Gymnostemma

1 viz. species of the genera: Adi1lostemma, Benz'ncasa, Citl'U .....!.l Cucumis, CUlu,oitd, G7mno-
sttmma. Lagenaria, Luffa, ~/elothria, Momordica, SCh£ZDjJejtm, -.rrid,osanthes. The species in
question are mentioned in the text above.
 ADDENDA-CUCURBITACEAE 937
cissoides), and have straight or undulated lateral margins. In Ttichosanlhes
cucumeroides the epidermal cells on the upper side of the ]~af are drawn out
into blunt conical papillae. The stomata are either pre~ent on both sides
(varying in number on the upper surface) or are confined to the lower side of
the leaf. We may add that the stomata on the stem are commonly situated
at the apex of raised pedestals, formed by the epidermal cells (Benincasa
ceri/era, Cucurbita Pepo, &c.). For the occurrence of water-pores in the
Cucurbitaceae, see Spanjer, loco cit. According to Yasuda, the nature of
the vascular system of the midrib may be advantageollSly employed for
systematic purposes. He mentions the following types: A single vascular
bundle in Actinostemma racemosum and Schizopepon bryoniae/olius; two
vascular bundles, viz. a large one with a small strand above it, in M elothria
iaponica; one large bundle and two small lateral strands in Gymnostemma c~'s­
soides; three vascular bundles, viz. a large one and two small bundles situated,
vertically above the former, in Benincasa ceri/eta, Cucumis salivus and Lage-
naria vulgaris; four bundles, arranged to form an almost right-angled cross,
the lowest bundle being the largest, in M omordica Charantia, species of LuUa
and T,icMsanthes; seven vascular bWldles forming a ring, the largest bundle
being situated below while the remaining strands above become successively
smaller and show a symmetrical arrangement, in Citrullus vulgaris and Cucurbita
Pepo. Yasuda met with oxalate 0'( lime only in the older stems; it is specially
abundant in Momordica Charantia, w:9.ere it occurs in the form of solitary
crystals of diverse shape. In the hairy covering Yasuda distinguishes uniseriate
clothing hairs, which are pointed or blWlt. and external glands with stalks
of varying length. He found the following types of glands: shortly stalked
glandular hairs with a multicellular ovate head, in all the species; glands
with a long stalk: and a multicellular ovate head, in species of Benincasa,
Citrullus, Cucumis, Cucurbita, Lagenaria, LufJa, Momordica, and Trichosanthes ;
glands with a long stalk and a head showing a special type of structure,
the head being unicellular in Trichosanthes cucumeroides bicellular by means
t

of a transverse wall in Cucurhita Pepo, and lastly, multicellular with' the two
uppermost cells drawn out into blunt processes, in Benincasa cerijerao/
In connexion with the subject of glandular hairs we may notice that
extrafioral nectaries occur on the leaves ;n species of Abobra, Adenopus,
Alsomitra, Bryonia, Cephalandra, Cucurbita,· Feuillea, Lagenaria, LuUa,
M omordica, Sphaerosicyos, Trianosperma;and Trichosanthes; they are found
either on the entire lower surface or are restricted to the bases of the leaves ;
in some cases they are also present on the sepals Cl1ld bracts, the latter being
sometimes completely transformed into nectaries (see Dutailly and Delpino,
11. cc.). These nectaries have long been known to systematists and biologists,
but have not yet been closely examined. In the same way the folia subtusI

glandulosa' mentioned by Bentham and Hooker in species of Cephalandra

and Trianosperma require detailed investigation; possibly they are merely
due tt? _th~._ex!l?rnal..glands. .
Internal secretory receptacles have not as yet been recorded in this Order.
Hallier, however, informs me, that he has seen an exudation of latex from
a fruit of Trichosanthes, which had been cut open; so that Trichosanthes may
in the first place be further investigated for the occurrence of secretory organs.
3. STRUCTURE OF THE AXIS. We may add the following details regarding
the occurrence of bicollateral vascular bundles in the Cucurbitaceae. Accord-
ing to Baranetzky a cambium, which produces woody tissue externally, appears
at the outer margin of the inner soft bast in Cuc'Urbita perennis (=C. toetidissima,
Kth. ex Syn.).. Bryonia abyssinica (=Coccinia abyssinica, Cogn.), Rhynchocarpa
dissecta (= Ked1ostt's atr1:cana, Cogn.) and Zennerz'a suavi~ (=Jt.Ielothrz'a punctata,
Cogn.). The vascular bundles thus produced are situated at the periphery
 ADDENDA-CUCURBITACEAE

of the pith, and exhibit inverse orientation of the wood and bast; even in
the thicker stems, however, they are not found in all parts of the transverse
section. We may further note that the intraxylary phloem in some cases
(e. g. in Actinostemma biglandulosum, see below) only appears at a late stage.
This leads us to consider the anf'maly recorded by Fries in the older stems
of Siolmatra brasiliensis, Baill., which ultimately results in the appearance
of inversely orientated medullary bundles of wood and bast, devoid of vessels.
Siolmatra brasiliensis, like Alsomitra, &c., has ordinary collateral vascular
bundles. Secondarily, however, a zone of cambium, which is at first interrupted,
but subsequently becomes continuous, is developed at the. margin of the pith;
this cambial zone first produces groups of phloem internally and later groups
of libriform externally.
Siolmatra, like the other members of the Order, shows two rings of vas·
cular bundles in the transverse section of the stem. Of these the bundles. of
the inner ring are approximated in pairs, while those of the Quter ring are
inserted in the broad primary medullary rays, separating the paired bWldles
of the inner ring from one another; the outer burtdles are divided into two
lamellae, the inner ends of which are joined to~ether so that they appear like
the arms of a V, which is open towards the outslde.
According to Yasuda, the wood contains vessels with a diameter of
'14-'5 mm., and formation of tyloses is of common occurrence.
In young parts of the axis the pericyc1e contains a closed ring of scIeren-
chyma composed at fibrous cells, but, according to Yasuda and Pitard, this
ring becomes split open at later stages. In older stems, according to Yasuda,
fibrous sclerenchyma is occasionally found also on the inner side of the vascular
bundles. Secondary scIerenchyma, composed of short cells, is moreover
commonly developed on the inner side of the primary sclerenchyma, e. g. in
species of Actinostemma, Citrullus, Gymnostemma, LuUa, Melothria, Momordica,
and Trichosanthes. In the species of Trichosanthes the secondary sderenchyma
in the older stem forms a ring, which is, however, not complete.
The outer portion of the primary cortex frequently contains well·dif·
ferentiated collenchyma.
Yasuda observed formation of cork on the outer side of the sclerenchy.
matous ring in the species of Trichosanthes.
A central cavityuften appears in the body of the pith, sometimes (Benin-
casa cerifera, Cucurbita Pepo, Lagenaria vutgart."s) even in young stems.
To A. Fischer'S synopsis of the distribution of sieve-tubes in the Cucurbi·
taceae we may add that according to Yasuda entocyclic and commissural
sieve tubes occur also in species of Actinostemma, Gymnostemma, Momo1dica,
Schizopepon and Trichosanthes.
The following details may be added regarding the anomalous structure of
the axis mentioned above as occurring in species of Act£nostemma and MomoYdica.
The structure of the stem of Actinostemma b-iglandulosurn may first be considered
(according to Wallace). In the young axes there are two rings of five bundles each,
the bundles altemating with one another. The inner ring consists of three col·
lateral bundles with wood and bast and two phloem-bundles, the outer ring of five
collateral bundles of wood and bast. At a later stage the bundles of both rings
become bicollateral, the phloem-strands having in the meantime developed into
collateral vascular bundles. In this case therefore the intraxylary soft bast appears
at a relatively late stage. At the base of older stems accessory vascular. bundles
are developed from a meristem arising in the endodermis, i.e. immediately external
to the ring of pericyclic sclereochyma. These vascular bundles show a radial
arrangement and are situated on the outside of the primary bundles; they ulti-
mately become bicollateral, and then undergo increase in the normal way by means
of a cambium j to the naked eye they appear as ribs.-Yasuda records the develop-
ment of such secondary vascular bundles, which are visible externally as ribs, and
 ADDENDA-CUCURBITACEAE 939
are situated on the outer side of the sclerenchymatous ring also in old stems of
Actinostemma racemosum and Momordica Charantia.
Tondera has published at length on the structure of the tendrils; he
utilizes the fact that the stalk of the tendril resembles the petiole in containing
only a single vascular ring, formed by the leaf-trace bundles situated in the
angles of the stem, to support the view that the tendrils are metamorphosed
leaves. 'For the sensitive pits on the tendrils, see Haberlandt, loco cit. .
Literature: Dutailly, Ecailles glandulif., Bull. Soc. bot. Linn, de Paris, 0:"6. 1875.-[Arthur.
Trichomes of EchiltOcys/is loba/a, Bot. Gazette, vi, 1881, pp. 180-3, 1 pl.]-Habcrlo.ndt, Einkap-
se~tmg~. P!otopl., Sitr..-Bet. Wi~net Akad:, xcviii, Aht. 1, 1889. P2. 1~8 and Tab.-Lamoun~tt~,·
Liber mterne, Ann. sc. nat., ser. 7, t. XI, 1890, pp. lI49-52'-L.DrobDlg, Wurzelknolleo, Diss.,
Rostock, 1892, p. 29 et seq. (Tltladia1ZIha, Eciiallium).J-Tognmi, Stomi, Atti 1st. bot. Pavia,
1894.]-Elfstrand, Heilpfi., Ber. deutsch. pharm. Gesellsch., 1897, p. 293 (TrUmosftrma).-
Spanjer, Wasserapparate, Bot. Zeit., 1898, i, p. 53.-Baranetzky, Faisc. bicollat., Ann. se. nat.,
ser. 8, t. xii, 1900, pp. 274-8 and 301-4.-Wallace,Stem.strueture of Actinostemma oiglandulosum.
Ann. of Bot., xiv, 1900, pp. 639-45 and pl. xxxiv.-Borzi, Apparato senso:'mot. dei cirri delle
Cucurbit., Atti R. Accad. Lincei, Rendiconti, x, I. Sem., 1901, pp. 395-400.-De1pino, Org.
caratterist. di alc. Cucurbit., Mem. Accad. Sc. Bologna, Ser. 5, iX,l901-190l, p. 383 et seq. and Tab.
i-iii.-Haberiandt, Sinnesorg., 1901, pp. 126-32 and Tab. v.-Pitard, Faise. liber. tert. d. riges des
Cucurbit., Act. Soc. Linn. Bordeaux, lvi, 1901, pp. civ-eviii; and Pchicyc1e, These, Bordeaux, 1901,
pp. 46, 47.-[Pollock, Fiprovasc. bundles in the root and hypocotyl in Ec/zinocystis Io/Jala, Report

J
Michigan Acad. Sc., iii, 19°1, pp. 40-2.]-[Yasuda, Compo anat. of the Cucurbit. etc. (Japanese),
Bot. Magaz. Tokyo, xv, 19°1, P' 88-9I.]-[Borzi, Apparato senso-mot. etc., Contribuz. 1st. bot.
Palermo, iii, I, 19°2, pp. I19-76. - Tondera, Gefassbiinde)syst. d. Cueurbit., Sitz.-Ber. Wiener Akad.,
cxii, Abt. 1, 1903, pp. 23-59 an Tab. i-v.-Yasuda, Compo anat. of the Cucurbit. etc., Jouro. CoIl.
of Sc. Imp. Univ. Tokyo, xviii, 1903,56 pp. and 5 pl.-Col, Faisceaux, Ann. sc. nat., ser. 8, t. XX,
1904, pp. 20s-6.-Faber, Bikoll. Gefassb. von CZlclIt'bita Pepc, Ber. deutsch. bot. GesellMh., 1904,
pp. 296-3°3, and Tab, XVI, xvii.-[Nemec, Starkescheide d. Cucnrbit., Bu.l1. into Acad. ie. Boheme,
jx, 1904, I2 pp.]-[Bernet, Obs. anat. nouv. sur la tige des Cucurbit., Ball. Herb. Boiss., 3 1 ser., V,
1905, p. 312; and Comptes rendus Soc. bot. Geneve.J-Fries, Zwei aiidamerikan. Lianen, Botaniska
Studier tillagn. Kjellman, Upsala, 1906, pp. 89 and e~pecially 96 et seq.

BEGONIACEAE (pp. 398-406).

Lit~rature: Keller, Luftwurzeln, Diss., Heidelberg, 1889, pp. 17, IS.-C .. de CandoUe,
Infloresc. epiphylles, Mem. Soc. de phys. et d'hist. nat. Geneve. 1890. vol. snppl.. sep. copy.
p. 20 et seq.-Jonsson, Anat. Bau d. Bl., Acta Univ. Lund., xxxii, 3, I 8!}6.-Minden , Wasaersez.
Org., Bibl. bot., Heft 46, 1899, p. 45.-Rechinger, Trichome d. Gesneriaceen, Osterr. bot. Zeitschr.,
1899, sep. copy, p. 2, foot-note 2.-Haberbndt, Lichtsinnesorg., 1905, p. 75.-Theorin, Vaxttrichom.,
Arkiv for Bot., iv, n. 18, 1905, p. 8. ./

DATISCEAE (p. 406).

Regarding the water~pores. see Spanjer, loco cit.
Literature: Spanjer, Wasserapp:lrate, Bot. Zeit:, 1898, i, p. 54.-Montemartini, Studio
anat. sulla /)atisc~ cannabina, Annali di Bot., iii, 1905, pp. 101-12 and Tab. xi, xii.

- CACTEAE (pp. 4°6-415).

Regarding the differentiation of the hypoderm, see also Preston, loc. cit.
He records a hypoderm of 6-7 layers in Echinocactus Wislizeni, Engelm., and
one of 12-14 layers in Cereus giganteus, Engelm.; he also mentions the occur-
rence of oxalate of lime in the hypoderm in Mammillaria Grahami, Engelm.,
Opuntia arborea, Engelm., O. julgida, Engelm. and O. phaeacantha,: Engelm.
In Opuntia fulgida and O. leptocaulis, DC. the epidermis. contaiqs crystals
which in the case of O. leptocaulis are found in special idioblasts.
The peripheral ground tissue is frequently differentiated as palisade-tissue,
e. g. in Cereus giganteus.
According to Preston's statements on the structure of the wood the
 ADDENDA-CACTEAB

characteristic elementary organs, described under the name of tracheids,

occasionally occur also in the. gen'us Cereus (C. Fendleri, Engelm.). In the
species just named narrow zones composed of spiral vesiels alternate in trans-
verse section with broader zones consisting of tracheids with a flat spiral band.
It remains doubtful whether these tracheids possess a lining layer of proto-
plasm and a nucleus (d. p. 410) during the whole of their existence or whether
such contents are present only in the young condition (see tDarbishire, loco cit.).
To the remarks made on p. 411 on the morphological nature of the thorns
we may add that according to Rudolph the thorns of Opuntia missouriensis
are neither leaves, nor emergences, but merely simple trichomes.
From the systematic-chemical point of view the demonstration of saponin
in Cerct's gummosus, Engelm. is interesting (Heyl).
Literature: Keller, Luftwurzeln, 1889, pp. 33-5.-Mittmann, Pflamen&tacheln, Verb. bot. Ver.
Mark Brandenburg, J 889. p. 6o.-Barber, Corky excresc., Ann. of Bot., vi, 1892, p. 166.-Lothelier,
Epines, These, Paris, 1893, p .•p.-Ganong, in Bot. Gazette, 1895. pp. 3I3-:n.-Matteucci, Placche
logher05e, Nuovo Giorn. bot. Ital., 1891, p. 224 et seq.-K. Schumann, Gesamtbescbr. d. Cacteen, .
1899, pp. I3-15.-Preston, Root·system of cenain Cacteae, Bot.IGazette, 1900, pp. 348-51.-
HeyJ, Alkaloide u. Saponine in Cact., Arcbiv d. Pharm., 239, 1901, p. 471.-Preston, Suuct.
studies 00 Southwestern Caet., Bot. Gazette, 1901, pp. 35-55' (species of Cereus, EcM"ocadus,
Alammi/laria, OPUnl,a).-[Hray, Plants of the Sotol region, Bull. Torrey Bot. Club, xxx, ]903,
po 621 et seq.]-Rudolph, Stacbelbild. bef Cact., Oesterreich. bot. Zeitschr' J 190.~, pp. 105-9 and Tab. i.
-Darbisbire. Mammillaria elongata, Ann. of Bot., 19°4. pp. 37!i-.p6 and pI. xxv, xxvi.-Porsch,
Spaltoffnnngstypu5, Jena, 1905, pp. 119-23 and Tab. iv.-[Arcangeli, Cereus jeruvianus, Atti del
Congresso dei Naturalisti ital., Milano, 1906, p. 403 et seq.]

FICOIDEAE (pp. 415-419).

According to Gulliver, styloids occur also in species of M esembryanthemum.
Literature: Schubert, Parenchymscheiden, Bot. Centralbl.. 1891, iii, p. ..72.-Baccarini e
ScUlama, Glimu I~toidtl, Contribnt. 1st. bot. Palermo. ii, 1898, pp. 83-u9 and Tab. iJ;-xiv.-
W. Meyer, Vergl. Anat. d. Cary ophyUac. , etc. Diss., GOttingen, 1899 (TeltpM..m).-BreIl.Iler,
FettpB., Flora, 1900, pp. 398--4-02 (A-fut'mbtyantMmul1l).- jOsting, in Beih. z. Hot. C~ntralbl., xii,
J902, p. 1-41 (Tdt'plzium).-Kearny, in Contrib. U. S. Nat. Herb., v, 5, 1900, p. 302 (Sesuvium).-
[Bergamasco, BioI. delle Mesembr., Bull. Orto bot. Napoli. 1904, p. 165 et seq.; abstr. in Just,
1904, I, p. 71!).]-[Pilger, Neue Gatt. der Aizoaceae, in Engler, Bot. Jabrb., xl, 1908, pp. 396,
397 (GliscnrolnamKus).)

UMBELLIFERAE (pp. 419-426).

I. To the list of anomalous structures enumerated at the end of the
REV1EW OF THE ANATOMICAL FEATURES we may add the fission of the xylem-
mass in the axis and root of Azorella Selago, Hook. fi1. The very rare occur-
rence of glandular hairs may also be mentioned on Siissenguth's authority.
2. STRUCTURE OF THE LEAF. A one-layered hypoderm is developed
above the lower epidermis in Bttpleurum petraeum (according to Briquet)
and B. gramini/oiium, Vahl (according to David). The upper epidenrus is
papillose in Bupleurum lalcatum, L. and B. montanum (David); it consists
of larae cells in Hermas villosa (Duboule). The mechanical elements in the
veins of the leaf are collenchymatous or scIerenchymatous.
For the structure of the septate phyllode-like leaves of Oxypolis ftlijo1'mis,
Britt., see Rennert'; for the structure of the leaves of Azorella, see Ternetz,
and for those of Hermas, see Duboule, n. ce.
In connexion with the subject of oxalate of lime (p. 421) we may add
that according to Ad. Meyer and Nestel the leaves of Aethusa Cynapium, L.,
Conium maculatum, L., Seseli Libanotis, Koch and Trinia glauca, Reichb.
contain small, sometimes plumose excretions, which occur both in herbarium·
.and alcohol~material. AdJIeyer considers these masses to be hes~ridin, while
Tschirch
I
looks upon • tltem as consisting of a new substance of unknown nature.
 ADDENDA~UMBELLJFERAE 941
As regards the hairy covering we may note that Bourdin figures simple
uniseriate clothing hairs in O~nanthe.globosa, L...,. while Duboule records three-
celled clothing hairs with'a lJasal cell, a' short stalk-cell and a long ·terminal
cell in H ermas gigantea. SiissenguUi .states that he has observed small glandu-
lar hairs with a head composed."Jf 2~4 cells o!J: the lower side of the leaf of
Pimpinella Saxifraga. .
3. STRUCTURE OF THE Mns; Among the species :having medullary vas-
cular bundles (d:.:p. 424) we may include 1\1agydaris panacifolia, Lge. (according
to Kiimmerle; this species has' numerous bund1e~, which are ei,fher cbllateral
or concentric with central phloem), as well -as Apium graveolens, L. and
Hacquetia Epipactis, DC. (according to Neste!). ,
According to Ternetz, Azorella Selago, as already mentioned above, exhibits
fission of the xylem-mass in 'the older parts of"the stem and in the roots. This
anomaly is closely rela~ed to the anomalous structure of the root previously
known to occur in certain species of Oenanthe, &c., and already described in
section 4, p. 425 Csecondary transformation of the fibrovascular system into
concentric vascular bundles '), although it has 'quite a' distihct· aspect in trans-
verse section, especially in. the older parts o.f the plant. The common features
of both types of anomaly are 'the ~leav?-ge of theorigfnal fibrovascular.system
and the appearance of. secondary meristems, producing wood and bast.
~ , , iIII

According to Ternetz, the fission. of ·the xylem-mass in the .axis of. Azorella
Selago takQS place as follo\V!;.In the y~ung aXlS the vascular bundl~s are loosely
arranged in the normal way to. form a ring. In .the cou~se of gro\\-'th in thickness
and the simultaneous development of .cork. a process of disorganization is started
in the parenchymatous ground-tissue. and in the outer portions of the secondary
cortex, both of which become partly transformed into a kind of complementary
tissue; at the same time clefts are formed, which have an a,{Jproximately radial
course and also penetrate between the groups of·wood in the dlrection of. the pith.
When the axis has attained a thickness of 5~ mm .• meristematic tissue cQ01.-
mences to 'develop in. the region 'of, the xylem-ma~. viz. alon~. the radial clefts
and on the inner side of the.xylem-segments; this meristematic tissue gives rise
to a parenchyma. which undergoes dilatation. The appearance of secondary meri-
stems forming wood and bast in this parenchyma and their junction 'with ,the
normal cambium of the vascular bundles then leads to the development of a kind
of polystely; in consequence of· further' cleavage, however, this structure soon
becomes dlSturbed and, gives ,way to a maze of separate segments of wood and
bast irregularly interwoven with one another.-~e may also note that the xylem-
mass of AZ01'ella contains only' ''annular and spiral vessels' and no reticulate or
pitted vessels; apart from these vessels it consists solely of unlignified parenchyma.
4. Sl'RUCTURE OF THE ·ROOT. In Magydal'is panaci/olia the older roots have
a structure similar to that found in Oenanthe. croca(a, the transverse section show-
ing two rings of concentric vascular bundles with central xylem. The mode of
develo~ment of this structure is ~the same as in Oenanthe.

Jjterature: Ad. ,Meyer; Anat. Charaktcre offiz. Bl. u. Kratltei', Abb. naturf. Gesell.Jsch.. Halle,
xv, 1882, sep. copy, pp. 7-10.-Coltantin, Tiges aero et sout., Ann. Se. nat., s~r. 6, t. xvi, 1883,
p. 73 et seq.-Costantin, Ti~es d. pI. aquat.,.Ann. sc. nat., ser. 6, t. xix, 1884, p. 287 et seq. and pi.
xYi.-Jadin, Org. secret:," These,"Molltpellier, 1888, po 36 et seq.-K.. Miil}er, Freie Gefassb. in den
Blattstielen, Sitz.-ller. naturf. Freunde Berlin, 18901 p. 13I.-Lotheller, Epmts, These, Paris, 1893,
p. 25 (Eryngium).-[Pohl,Pharmakognosied. Umbellif.-Wnneln,Lotos, xiv, 189i, pp.89-98, 2 Tab.;
abstr. in Just, 1894, i, p" 48i.]-:&urdin, o mbelliferes , etc., Anat. compo de la feuille, These,
MOD~pellier. 1897, pe. sr-81.-{Briquet, Monogr. des Bupleures des Alpes in aritimes, &.Ie. et
Geneve, 1897. 131 pp.]-Briqnet, Theorie phyllodique, et-:., Bull. Herbier Boissier, v, 1897, p. 235
et scq.-Schubert, Parenchyrnscheiden, Bot. Centralbl., 1897, iv, p. 17.-Duboule, Anat. compo de la
fcuille dans Ie genre Htrmas, Arch: $C. phys. et nat. Geneve, ser. 4, t. vii, 1899, pp. #6-80 and pI. iv;
al~ in Bull. ~abor. de bot. gen. de Geneve, ii, 1899, pp. 37-7l.-:-Minden, \\assersez. Orgue, Hibl.
bot., Heft ,46, 1899. p. I I (Be~la).-SchleichertJ Xerophyten bei lena, Naturwiss. ''Yochenscbr.•
1900, p. 149 (Bupleurum).-Thomu, Feuilles soul, These, Pans; 19oo.-David, Etude anat.
du genre Bujkurum, These, Paris, 1901,95 pp.-[Goris, \\'unel von Asa/oetida, Joum. de Pbarm.
et Chimie, 1901, n. 12; abstr. in Just, 1901, ii, p. 37.]-[Holm, Ilrigenza bulcosa, Ameri.c.loum.
of Sc., xi, 1901~ p. 63.]-Pitard, Peri cycle, 1'~ese, Bordeaux, 01901, p. 39.-KnQthe, Unbenetzb.
 ADDENDA-UMBELLIFERAE
B1., Diu., Heidelberg, 1903, p. 21.-Kiim,Plerle, Beitr.~. Kenntnis d. Anat. d. Umbellif. (Magyaaris),
Novenytani Kozlemeoyek, 1902, Hungarian, sep. copy, 18 pp.-[Petersen, BIadnerv. hos arter af
slagten .Eulle~rum, Bot. Tidsskrift, xxvi, 1902, pp. M3-76.]-Ternetz, Morphol. u. Anat. d.
AZ()"ella Selago, Hook. i., Bot. Zeit., 1903, pp. 1-20 and Tab. i.-Modrakowski, Vergl. Untersuch.
d. dem Cmium macula/um ahnlichen Umbellif., Zeitschr. aUg. osterreich. Apothekerver., 1903,
n. ",5-50, aep. C()py, 2i) pp.-[Rennert, Phyllodes of Oxypolisjili/ormis, Bull. Torrey Bot. Club, xxx,
1903, pp. 40 3- 11 ; abstr. in Bot. Centralbl., xcv, p. 247.J-1'heorin, Vaxttrichom., Arkiv for Bot.,
i, 1903, p. 160.-Col, Faisceaux, Ann. sc. nat., ser. 8, t. XX, 1904, pp. 173-9.-Freidenfeldt, Anat.
Ban d. Wurzel, Bibl. bot., Heft 61, 1904, p. 66.-Siissenguth, Behaarungsverh. d. Wiirzburger
MIDlc:helkalkp1t, Diss., Wurz:burg, 1904, pp. 3l-5.-Nestel, Stengel- n. Blattanat. d. Umbellif., Diss.,
Zurich, 1905, u6 pp., I Tab.-Weberbauer, Vegetat. d. Hochanden Perus, in Engler, Bot. Jahrb.,
xxxvii, 1905, p. 60 et seq.-[For further literature, see p. 1172.]

ARALIACEAE (pp. 426-432).

I. REVIEW OF TaE ANATOMICAL FEATURES. The recent investigations
00. the structure of the leaf and axis undertaken by Giissow 1 and Viguier 2
render a number of additions necessary. The superficial development of the
rork in the stem and the frequent presence of co])enchyrnatous tjssue ill the
priJnary cortex ate features characteristic of the Order. Oxalate of lime is
excreted not only in the form of ordinary solitary and clustered crystals, but
occasionally also in the form of prismatic crystals of varying size (epidermis
of the leaf of Astrotricha) and of typical crystal-sand (Apiopetalum., Boerlagio-
tfendron, Mothemellia, Tetraplasandra). According to Giissow, secretory
<:avities are found in the mesophyll also in species of Cussonia, Eremopanax,
Heteropanax, Kissodendron and Pseudosdadium. As regards the hairy covering
we may add that unicellular clothing hairs and typical external glands are
wanting, that shaggy hairs occur also in species of Kalopanax and Stilboca,pa,
and fhat small scale~like trichomes are found in Oreopanax xalapensis, Decne.
et Planch. According to Giissow (although not in agreement with Viguier)s
medullary vascular bundles, showing varied orientation of wood and bast,
are present not only in species of Aralia and Arlhrophyllum, but also in species
of numerous other genera. Certain species (especially those of Oreopanax),
moreover, are distinguished by the possession of cortical vascular bundles;
for details on this point, see under the structure of the axis.
According to Viguier, the genus Aralidium, in which both Giissow and Viguier
found no resin-canals, must be e~cluded from this Order. Special anatomical
features recorded in this genus are: sman cortical vascular bundles exhibiting
normal orientation; isolated groups of bast. fibres in the pericycle; broad medullary
rays in the wood, and bundles of medullary fibres.
The STRUCTURE OF THE LEAF is again bifacial in most of the species
2.
.investigated by Giissow, but in some cases it merges into homogeneous-centric
structure; isolateral structure with palisade-tissue situated on both sides of
the leaf has not been recorded in. any member of the Order. Arm-palisade
tissue occurs also in Acanthopanax sessili/lortts, Se~m. and Pseudopanax
laetevt'rens, Benth. et Hook. The intercellular spaces of the spongy tissue

1Giissow's investigations extend to the following genera, which are enumerated in the serial order
of Harms' system: t. Scheffiereae: Tupidantnus, Plera1uira, Tetraplasandl'a, ReynoltUia, Ptero-
tropia, Boedagiodendron, Tnvesia. Fatsia, Tetrapanax, Meryta, Scluffiera, Oreopanax, Gilihertia,
Hedera, Brassaiopsis, Gastonia, Polyscias, Kissotiendro1t, Pseudopanax, lJ-facropanax, ll!otlzopanax,
CheirodmdrolZ, Ast1'()tricha, Acanlnopanax Kalopanl7x, Didymopanax, Heteropanax, Cussonia,
Arthrophyllztm, Eremopaltax. II. Aralieae : Penlaptmax, JI,/otnerwelHa, Aralia, Slilboca"pa, PantU,
Aralidium. Horsfieldia (-Har11lSiopanax), Myodocarpus, Delarbrea, Porospermum. III. Mackin-
layeae: Mackinlaya, Apiopetalu1lts Rsmdosciadium. .
S Viguier's work is essentially concerned only with the djfferentiation of the vascular system and
the distribution of the resin·cantls in the axis and petiole i his investigations were, however, also
undertaken on material repr~etlting very numerous genera.
 ADDENDA-ARALIACEAE 943
are usually small, although in some· cases of large size. Stomata are rarely
found on the upper side of the leaf and then only occur in small numbers
(Aralia humilis, Cav., Gastonia cutispongia, Lam., Tetrapanax papyrifer,
K. Koch). In certain species of M acropanax, SchetJlera and Tupidanthuf,
the stomata are exceptionally numerous, so that the lower epidermis is formed
entirely by the stomata with their subsidiary cells. The pairs of guard-cells
either lie on a level with the epidermis or maybe somewhat raised or a little sunk.
Stomata. having subsidiary cells arranged according to the Rubiaceous type
were observed by Giissow also in species of Aralia, Astrotricha, Cheirodendron,
Echinopanax, Eremopanax, Fatsia, Gastom'a, M ackinlaya, Nothopanax, Oli-
goscias and Psorospermum. The old and new investigations (see below) have
shown that the occurrence of hypoderm on the upper side of the leaf is a very
widely distributed feature in the Araliaceae. On the other hand, papillose
differentiation of the epidermis is not frequently found ; to the previous records
we may add the slightly papillose protrusions found on the upper side i.q
Reynoldsia sandwicensis, A. Gray, on the lower side in Polyscias sambuci/olia,
Harms, and on both sides in P. tarino!a, Harms, as well as the papillae
present on the lower side of the leaf of Pentapanax parasiticus, Seem.; the
laUer resemble those 6£ Aralia canescens. Giissow also failed to meet with
gelatinization of the epidermis of the leaf in any member of the Order.

Giissow mentions the occurrence of tangential division-walls in the epidermis

in Nothopanax dive'J'sifolius, Harms, and of a one-layered hypoderm, which is only
developed locally, in certain species of Artkrophyllum, Oreopanax and Polyscias.
He further records: a one-layered.hypodenn, m Arthrophyllum Blumeanum, Zollo
et Morie., A. dive'Ysijolium, Bl. (sphalm. • quinquejolium '), Cheirodend-ron Gaudi-
chaudii, Seem., Cussonia spicata, Thunb., C. umbelU!era, Sond., Delarbrea cotUna,
Vieill., Didymopanax vinosus, March., Gastonia dupUcata, Pet.-Thouars, G£libertia
lau,ifolia, March., Meryta. micyocaypa. Baill., Myodoc,aYpU5 pinnatus, Brongn.
et Gris., Nothopanax arboreus, Seem., Reynoldsia sandwicensis, A. Gray, Schefflera
tomentosa, Harms, Tetrapanax papyri/er, Koch; a 1-2-layered hypoderm, in
Chei'Yodendron. platyphyllum, Seem" Eremopanax otopyrenus, Baill., Oreopana:f
Rusbyi, Britt., Schefflera Volkensii, Harms; a 2-layered hypoderm, in 'Aralia
t'Yifoliata/ Didymopanax Morototoni, Decne. et Planch.• Ptcrotropia Ikavaiens£s,
Hillebr.• Tetraptasaniira mdandra, Hillebr.; a 2-3-1ayered hypoderm, in Tupidan-
thus calyptratus, Hook. f. et Th.; a 3-1ayered hypoderm, in Apiopetalum velutinum,
Bailt (cells 7-8 times the size of the epidermal cells) ; a 4-5-1ayered hypoderm in
Gastonia cutispongia, Lam. ./
Viguier mentions the occurrence of hypoderm also in the following species.
which have not been named in the preceding paragraph: Apiopetalum glabratum
(3-4-layered), Cussonia vantsilana, Eremopanax Balansae (2-layered), Fatsia
iaponica (locally), Mackinlaya mac'Yosciadea (3-layered). It-!esopanax cap£tatum,
Myodocarpus crassifotius, M. floribundus. Nothopanax s£mplex (locally), Oreopanax
Sanderi (I-layered), O. xalapense (2-3-1ayered), Pseudopanax Lesson#, PtcYotrop£a
dipyrena, Schefflera Humblotiana (situated on both sides, and composed of one
or two layers of lignified cells). Sciadopanax Grevei, Tieghemopanax lVeinmann£ae;
Holte~ann also records hypoderm in Heptapleurum emarginatum (2-3-layered,
as in H. stellatum).--

Oxalate of lime is found not only in the form of clustered and ordinary
solitary crystals, but also as cryst.al-sand and prismatic crystals. Cells con-
taining crystal-sand have been observed in the mesopliyll in Apiopetalum
velutinum, Baill. t Boerlagiodendron Warburgii, Harms, lvlotherwellia haplo-
sciadea, F. v. Mull. and Tetraplasandra meiandra, Hillebr.; the crystal-sand
is composed of fine grains, and includes clustered crys'tals in the species of
Boerlagiodendron and TetrapIasandra just named. The clustered crystals
found in the pith are occasionally grumose, while those in the mesophyll in
some cases attain a considerable size. Ordinary solitary crystals are of much
944 ADDENDA-ARALIACEAE

more frequent occurrence th~ri was formerly supposed~ The epIdermis of the
leaf contains clustered cr~tals .also in Acanthopanax sessilijlorus, Seem. and
Pseudosciadium Balansae, Baill., the above-mentioned prismatic crystals in
speci~ of Astrotricha, and ordinary solitary crystals in Eremopanax otopyrenus,
BailL, as well as in A,canthop.a_nax ricinitolius, S. et Z., Arthrophyllum diversi:'
folium, :Bl. and Kalopanax ricinilolius,~ S. et Z. (in the last-named species in the
epiderm is covering theNeins, of the leaf). Clustered crystals, lastly, are found
also in the hypoderm in Apiopetalu.m velutinum, Cheirodendron platyphyllum,
Potyscias /arinosa ari& Tupidantkus calyptratus. _
We have next, to consider the secretory canals -found in the veins of the
leaf (see p~ 428). According to Gtissow, they ,are con~ned. to the midrib also
in Acanthopanax"pentaphy/lus, March., -Aralia -dasyphylla, Miq., A. humitis,
Ca~., Cu~sonia spicata, Thunb:, and Pentapanax angelicifolius, Griseb., while
they, appe-ar to be absent in the leaves of Polyscias larinos'a" Harms, P. sanibuci-
folia H~rms, Pseudopanax laetevirens, 'Benth. et Hook:" and P. valdiviensis,
J

Benth. et Hook. According to Giissow~ secretory cavities~ resembling those

of.Gilibertia,. but of small-size, are found in the mesophyll in Cusso1zia umbelli-
fera, Sond.,. Eremopanax otopyrenus, Bam., Heteropanax fragrans, See~.,
Kissodendron australianum, Seem. and Pseudosciadiu,m- Balansae,_ Baill.; they
do not give rise to transparent dots 1. Gilibertia protea, Harms (see p. 429),
which differs from the other species of the genus in not having secretory
cavities, is placed by Viguier in the new genus Mesopanax,·which also includes
certain ~pecies of Oreopanax. .' .
The mostimpo~tant feature of the hairy covering is the. absence.of u.ni-
cellular hairs and of typical external glan,ds. GtissoW' records shaggy hairs
(the terminal cells of which are, however, not glandular) in Acanthopanax
sessilifolius, Seem., Aralia nudicaulis,. L., Kalopanax ricinitolius, S. et ~Z.
and Stilbocarpa polaris, Decne. et Planch. Two-armed hairs, like those of
Didymopanax longepetiolatus, are found also in D. Morototoni, Decne. et Planch.
and D. vinostls, March. According to Glissow, tufted and stellate hairs are
widely distributed. Scale~like; trichomes, having the same .structure as the
pelt ate hairs of H edera Helix, but small and orange-coloured, occur_ in Oreo-
panax xalapense's, Decne.' et ,Planch.,- while uniseriate clothing hairs, which
have'thin walls and are occasionally branched, are found in Kalopanax- ricini·
folius, S. et Z. Viguier's statements about the trichomes are too vague- to
demand further consideration.
. The petiole (d. P.,430) has recently been subjected to a, v~ry thorough
investigation by Viguier. The, normal vascular bun,"es of -the petiole are
generally arranged in a 'ring,. rarely~ in the form of, a .horseshoe (as in Pseudo-
panax pro parte, or Apiopetalum). They are either quite distinct. ·from one
another or are. united by an. inner or Quter ring of sc1erenchymatous fibres ;
in some cases (as in Tupidanthtts) .they are embedded in a special :ring of
lignified parenchymatous tissue. The number of bundles is for the most part
relatively large, but in Astt'otricha ledifolia there are only three vascular
bundles, which almost touch one another. In Oreopanax § Digitatae, and
M acropanax the va~cu!ar bundles of the normal ring are alternately large and
small. Other anatomical features ()f the petiole. which•. besides those already
named, are employed by Viguier for purposes of classification, are as follows :
the occurrence of subepi4ermal collenchymatous tissue; cambi.il activity in
the vascular bundles of the normal ring, (particularly marked in Delarbrea
spectabilis); the distribution and size of the reSiri·canals; the occurrence
of a central lacuna; and lastly,! the ~evelopment •. structure, number, and

1 Viguier (loc. cit.) does not mention these features.

 A DDENDA-ARALIACEAE 945
arrangemenf of the medullary' vascular bundles. As regards the latter we may
note that in some cases there is only a small 'central bundle (e. g. in Anoma-
panax and Pseudosciadium);' in ,other cases there is a ring of medullary
bundles showing mverse orientation of the wood and bast (e. g. in Aralia,
ArlhrophyUum, Kissodendron, and Pentapanax), or a normally orientated
ring of medullary bundles with the 'xylem-groups pointing'inwards (e. g. in
Cussonia and Gamblea) or medullary bundles showing an irregular arrangement
of the wood and bast, arid so on.
Viguier records medullary vascular bundles in .the petiole in the following
genera: Anomopanax (I v;l.scular bundle), Apiopetalum (some of the bundles
inversely orientated, others irregularly arranged), Aralia excl. A. ferox (rachis
containing a single ring of inversely orientated bundles), Al'throphyllum (single
ring of inversely orientated bundles), Boerlagiodendron, Brassaiopsis (single ring
of irregularly arranged bundles), Cuphocarpus (numerous irregularly arranged
bundles), Cussonia (single ring of normally' orientated bundles), Didymopanax
(a few, more or less coalescent bundles), Dizygotheca (outer bundles inversely, inner
bundles normally orientated, or "two rings of inversely orientated bundles), Eremo-
panax (numerous scattered bundles in the petiole), Gamblea (single ring of normally
orientated 'bundles), Gastonia (numerous bundles exhibiting varied orientation;
some of th'eni concentric with central phloem), Gilibertia pro parte (a few 'small
inversely orientated cundles), Heteropanax (rachis with a single ring of inversely
orientated.bundles, which are intercalated between "the bundles of the normal ring).
Kissodendron (single ring of inversely orientated bundles), Mackinlaya (irregularly
and inversely orientated bundles), Meryta (numerous irregularly arranged bundles),
Mesopanax (a few irregularly arranged bundleS or a single ring of inversely orientated
bundles), Myodocarpus (numerous irre~larly arranged bundles, occasionally not
prominently aeveloped), Octotheca (like Dizygotheca), Oreopanax § Digitatae
(single ring of inversely orientated bundles), Pentapanax (single ring of inversely
orientated bundles), Plerandra (normally orientated -bundles), Polyscias (few or
many irregularly arranged bundles), Pseudopanax pro parte (three small normally
orientated bundles), Pseudosciadium' (one bundle), Pterotropia (one or two large
bundles). Schetflera (one or two rings of inversely orientated bundles), Sciadopanax
(a few small bundles). Strobilopanax (like Mer'yta). Tetrapanax (somewhat irregu.,
larly arranged bundles), Tetraplasandra (many irregularly arranged ,bundles),
Tieghemopanax (numerous irregularlx' arranged bundles), Tl'evesJoa (some of the
bundles inversely orientated). Tup~danthus (some of the bundles with varied
orientation, others inversely orientated). On the other hand, Viguier states that
medullary - vascular bundles are absent in: A canthopanax. A raUa /erox, A stro-
t1'icha, Bonnierella. Cephalaralia. Cheiyodendron, Delarbrea, Echin<?panax, . Fatsia,
Gilibel'tia pro parte, Hedera. Macropanax, Nothopanax, Oreopanax § Lobatae,
Panax. Pseudopanax pro parte, Stilbocarpa.
According to Viguier, medullary resin-canals are found ,in the petiole in:
A canthopanax, A nomopanax, A I'thl'ophyllum, Cheil'odendl'on, Cuphocarpus, Cussonia,
Delarbrea, Didymopanax pro parte. Dizygotheca, Eyemopanax, Gastonia, Hetero-
panax (rachis), Kissodendron, Mackinlaya, Macropanax, Meryta, Myodocarpus.
Orcopanax ,pro parte, Pentapanax, Plerandra, Polyscias, Pseudosciadium. Ptero-
tropia, Schefflera pro parte, Sciadopanax, Tetrapanax, Tet1'aplasandYa. ,Tieghemo-
panax,. Tl'eve.sia pro parte, T_upi4_411.thus. Regarding the arrangement and size of
the resm~canals, see-Ioc.,cit. ,
The" green spots occurring on the petioles and on the verns of the leaf in many
species of Meryta are due to a,local interrupti'on of the subepidermal layer of
collenchyma. which at these points is replaced by cells with green contents. The
swellings found on the principal veins and on the larger secondary veins in Meryta
and in Strobilopanax are mainly caused" by a strong development of the inner
ground tissue, which is differentiated as aqueous tissue at these points;

3. STRUCTURE OF THE AXIS. Numerous details regarding the distribution

of the resin-canals ·in the various tissues of _the axis are to be found especially
in Viguier's work. According to him the most important systematic point
lies in the presence or absence of resin-canals in the outer; collenchymatous
SOLEI£D£It 3 P
 ADDENDA-A RALIA CEAE

portion of the primary cortex and in the pith 1. Viguier observed radially
elongated secretory cavities C poches '§ecretrices ') in the broad medullary
rays of the wood only in Arthrophyttum (A. diversilolium); this observation
requires confirmation, as it is not impossible that the structures in question are
merely resin-canals, running from the cortex towards the pith.
According to Giissow, medullary vascular bundles are more wideJy dis-
tributed in the axis of the Araliaceae than was previously supposed. This
author was also the first to demonstrate cortical vascular bundles in certain
members of the Order. The following synopsis summarizes the results of his
investigations on tht; distribution of the vascular bundles as seen in the trans-
verse section of the stem.
1# Only a single ring of normally orientated vascular bundles (i.e. with the
bast directed outwards and the wood pointing inwards) is found in certain species
(which are named on pp. 29. 30 of the paper cited below) of: Acanthopanax, Aralia,
Astrotricha, Cheiro dendron , Didymopanax, Fatsia, Gilibertia, Hedera, Hot'sfieldia,
Kalopana,;, Afacropa'nax, ftfackinlaya, Motherwdlia, Nothopanax, ONgoscias, Panax,
Polyscias, Porospermum, Pseudopanax. Pseudo sciadium , Schefflera and Tetra-
pJasandra.
II. In addition to the normally orientated ring of vascular bundles there is a
medullary ring showing inverse orientation (with bast on the inner and wood on the
outer side) in: Aralia chinensis, L. (in this species there are also numerous small
vascular bundles showing irregular arrangement of the wood and bast between the
two rings), A. dasyphytla, Mig., Arthrophyllum pinnatum, Clarke and Eremopanax
otopyrenus, Baili.
III. In addition to the normal ring of bundles there is a medullary ring, which
is likewise normally orientated, in: Arthrophyllum diversifolium, Bl. and Cussonia
spicata, Thunb. . -
IV. Besides the normal ring and a ring of medullary bundles at the periphery
of the pith (which is normally orientated?, see Giissow, loco cit., p. 30), scattered
vascular bundles with irregular orientation of the wood and bast are found iil the
pith in: Aralia cordata, Thunb., A. kumilis. Cav., Didymopanax Morototom:. Decne.
et Planch., D. vinosus. March., Kissodendron austrah"anum, Seem., pterotropia
kavaiensis, Hillebr.
V. In addition to the normal ring of vascular bundles the pith contains scattered
bundles with irregular orientation of the wood and bast in: Gastonia cutispongia,
Lam., Meryta microcarpa, BailI., Plerandra Stahl-,:ana, "Varb., PolysC£as farinosa,
Harms, Trevesia palmata, Vis.
VI. Besides the normal ring of bundles there are cortical vascular bundles
in: A p":opetalum t'elutinum, Bail!.. Brassaiops":s H ainla, Seem., Cussonia umbelli-
fera, Sand., Delarbrea callina. Vieill., Heteropanax jragrans, Seem., Ml,'odocarpus
pinnatus, Brongn. et Gris.. Oreopanax xalapensis, Decne. et Planch., O. Rushyi,
BritL, Pentapanax angelici/oJius. Griseb., Stilbocarpa polaris, Decne. et Planch. (?).
VII. In addition to the normal ring of bundles both medullary and cortical
vascular bundles are found in: Aralia racemosa, L., Boerlagiodendron Warburgii,
Harms, Brassaiopsis speciosa, Decne. et Planch., Tupidanthus calyptratus, Hook.
f. et Th.
Viguier, on the other hand, who likewise examined the structure of
the axis in abundant material comprising numerous genera and species
and to whom Giissow's work was unfortunately unkno'WIl, demonstrated
medullary vascular bundles (constituting a single ring with inverse orienta-
tion) only in the vegetative axes of certain species of Aralia, which he
groups together as Euaralia (A. cachemirica, A. chinensis, A. cordata,
1 Viguier demonstrated medullary resin-canals, which e .. hibit diverse arrang_~ment (viz. scattered
in the pith or occupying a periphl"ral position and in the laU. r Clse sotnetimf's in contact
with the protoxylern of the ring of bundles) and vary in numher and size, in species of the following
genera: Acanlhopanax, AtuJmopanax. Apio/,etalum, AraH{~t Arlhrophyllum. Bonmerella,
Cluirodendron, Cuphocarpus. Cussonia, Didymopanax, Iiremoptmax, Gild'erlia. Har1llsiopsis,
Hulera, Kissodend)'(nt, Macrojanax, MackilZlaya, Meryta ••Myodocorpus. Oreojallax, Pentapanax,
P"/yscias, Puucibs(iadium, Sc/zej//era, l'itghe11lopanax, Tttjidantlms.
 ADDENDA-ARALIACEAE 947
A. dasYPhytttJ, A. hispida, A. h'Ypoleuca~ A. montana, A. nutiicauiis, A. sp",'nosa
and A. urticaefolia, but not in A. ferOlC}, as well as in the axis of inflorescence
of Didymopanax M orototoni, K issodendron australianum and Polyscias nodosa ;
it remains an open question whether medullary bundles occur also in the
vegetative axes of the three species last named. Viguier records cortical
vascular bundles (leaf-trace bundles belonging to leaves situated higher on the
axis) only in Oreopanax. Giissow's statements therefore still reqUire con-
firmation 1.
The bundles of the normal vascular ring are no doubt alwaY5 simple and
collateral 2. According to Viguier, more or less strongly peveloped groups of
pericyclic bast~fibr-es are found on the outer side of the bundles in most of the
species, but in some cases (e. g. in Fatsia iaponica and species of Echinopanax,
Gilibertia, Nothopanax and Panax) there is no sc1erenchyma in the pericycle.
The perimedullary tissue on the inner side of the vascular bundles is occa-
sionally lignified or includes arcs of fibres similar to those found in the pericycle
(e. g. in Acanthopanax divaricatus).
According to Gtissow 3, the first cork invariably develops superficially, for the
most p:lTt in the first layer of primary cortical cells, more rarely in the epidennis
(Apiopetalum velutinum, Baill., Delarorea collina, Vieill., },facropanax undu~
latus, Seem., Polyscias xanthoxyloides: Harms). The cells of the cork often
have thin walls; in Aralia /eumilis, Cav. some of them exhibit a palisade-like
elongation; in other cases the cork~cells are thickened on the tangential walls
(species of Oztssonia, Gilibertia, Oreopanax, Sche!ftera) or on all sides (Pseudo-
panax laetevirens, Benth. et Hook.); thick-walled sc1ereids are found in the
cork in Oreopanax xalapensis, Decne. et Planch. and Polyscias xanthoxy-
loides, Harms. Certain species have a more or less closed ring of stone-cells
belonging to the phelloderm. Stone-cells likewise occasionally occur in the
primary cortex, but the characteristic mechanical tissue of the latter is
collenchyma, which is rarely apsent and for the most part forms a com~
pletely closed ring, 5-6 layers of cells in thickness, in the outer portion of the
primary cortex.
The following additional details regarding the structure of tho/ wood 4
are based on Giissow's statements 3. The medullary rays vary from one to seven
cells in breadth. Spiral thickening of th,e walls of the vessels has been demon~
strated in Aralia humilis, Cav., Astrotricha jloccosa, DC., A-. ledifolia, DC. and
Nothopanax diversifoUtlS, Harms; the perforations in the vessels show a transi~
tion to the scalariform type also in species of Horsfieldia, Trevesia and Tupi~
danthus. The wood-parenchyma is mostly restricted to the neighbourhood
of the vessels, although it is occasionally developed in greater abundance,
e. g. especially in Nothopanax diversilolius. Tracheids are absent. Giissow
invariably found simple pits on the wood-prosf.D.chyma. The same authOl ity
observed septation of the wood~prosenchyma together with storage of starch,
' __ _..
. ---~
I Gussow gives no details as t() the exact nature of the material employed in his investigations
.of the stem. :From what has been lDentioned above it is in no way improbable, that Giissow's state-
ments in part at least refer to the 4xis of inflorescence and not to the vegetative a.xis; nor is it quite
impossible, that Giissow occasionally investigated petioles in place of axes.
, According to Gi.issow, bicollateral Inmdles are not uncommon in the Araliaceae, even in the
normal vascular ring. Among the species enumerated by GiilSOw (loc. cit., PP' 3 ...., 35) in this rela~
tion Panax qut·nquefllius, L. (Arafla tjut"nquefo1ius) was the only one at my disposal. In this
species the outer soft bast extends round the body of the wood in the shape of a horseshoe, but the
bundles are not typically bicollateral.
3 Regarding the material on which Gi.issow's investigation of the axis was undertaken, see foot-
note t, above. . . ., . .
• Viguier records "ai, bois homoxyle' devoid of'secondary vessels in Not1t.opanax anomalus and
N. microp"ylltts; thick axes were, however, not examined I
3P2
 ADDENDA-.,ARALIACEAE.
also in species of Arth,ophyllum, Didymopanax, E,enwpanax, Gastonia, Gilibertia,
Kalopanax, ,Macropanax, N o~nax, Oligoscias, Oreopanax,. Pseudapanax,
Schefflera., Tupidanthus, &c. ..
The pith coDsists of. cells with thin or thickened walls; starch is occa-
sionally stored in .the peripheral cells.
Literature; 'Mobius,'Mechao. Scbeiden der Sekretbeb., Der. deutsch. bot. Gesellscb., 1884.
Geoeralversammlungsbeft, p. ;uvi.-C. de CandoHe, Infloresc. epiphylles; ~Iem. Soc. de phys. et
d'hist. pat. Geneve, 1890. vol. luppt., sep. copy, p. 5 et seq.-Barber, Corky excresc., Ann. of Bot.,
'Vi, 1893, p. 166~-Went, .Han- u. Nlibrwurzeln, ADD. Jardin. Buitenzorg, xii, ,18 95, pp. 55-6.-
[Zancb, Aculei, Conlribllz. lstit. bot. Palermo, ii. 1897, p~ I et seq.]~Baranetzky, Faisc. bicoll.,
Ann •.co nat., aer. 8, t. xii, 1900. pp. 304-i.-Giissow, Beitr. z. vergl. Anat. d. Araliac., Diis., Breslau,
1900,68 pp., I Tab.-Petersen,· VedaDatomi, 1901, p. io.-Clauditz, Blattanat: canar. Gew., Diss.,
Balel; 1901, pp. 35~ (Hed~).-(Chrysleri Central cylinder of. AraI,- and Lili~ceae, Bot. Gazette,
1904,.PP. _161.-8S.)-Col, FaJlceaux, Aoo. Be. nat.; ser. 8, t. xx, 1904, especially pp. '79-81.-
Areschoug, Trap. vaxt. bladbyggn., Sv. Vet. Akad. HandL, 39, o. 2, 19°5, pp. 150; lSI (Arlnro-
pk,.Oum).-Picdoli, Legnami, Bull. Siena, 1906, p. 13:1.-Viguicr, Rech. anat. sur In classification des
Araliac., Ann. Ie: oat., ser. 9, t. iv, 1906, pp. 1-207.-Holtermano, Einfluss des Klimas, etc., 1907,
p. 135 (HtJlopleurum).-[For further 1i~erature, see p. 1169.]

CORNACEAE (pP'. 4~2-439)·

I. REVIEW OF THE ANATOMICAL FEATURES. On the basis of Wangerin's
:work we may add that simple perforations occur in the vessels in Kaliphora also,
crystal-sand also in Kaliphora and M elanop,hyUa, and stellate haiI:s 'with a
varying number of r~ys (a, type of hair new to the Cornaceae) in Alangium
costatum, Wangeriq. ,
2. STRUCTURE OF THE LEAF. The earlier inexact statements on the
occurren~e of papillose, epidermal cells, (o:u p. 433) may be replaced by the
following details quoted from Kohne's and Sertorius' observations.
. Papill?Se developmen~ of the uppe_r epidermis is f?und. in: . Garrya ellipt£ca,
Dougl., G. {lavescens, Wats., G. laur'Jjoha, Benth., G. L~ndhe'Jmerl, Torr., G. ovata,
Benth., G. Wrightii, Torr. (the species last named has solid peg-shaped papill~e),
Manea bego-niae/olia. Roxb., M. macrophylla, S. et Z., M. platan#olia. S. et Z. and
Cornus canadensis, L. In Garrya the ,?uter walls of the lower epidermal cells ••
like those of the upper epidermis, are more or less distinctly par.i11ose,' and this is
even the case in species which show no tendency to form .paplllae on the upper
side. The lower eJ>idermal cells are slightly papillose also in Aucuba iaponic(J,
Thunb., A. himalal,ca, Hook. f., Nyssa capitata, Walt., N. unifiora, Wang., Torri-
cellia til1'aeJolia, DC., Comus alterni/olia, L. f., C. jlor,,'da; L .. and C. stolonifera,
Michx. In other species of Cornus the lower side of the leaf bears distinct
papillae, which- are generally provided with a corona and stand in connexion with
one another, by means of reticulately' arranged cuticular ridges;, these species
are: C. alba, L., C. asperifolia, Mi~., C. brachypoda, C. A. Mey., C. 'dtlli/orm'ca,
C. A. Mey., C. candidissima, Mill., C. capitata, \Vall., C. drcinata. L'Herit., C.
disciflora, DC., C. Hessei, Kohne, C. ~'gnorata, Koch, C. macrophylla, Wall., C.
oblonga, Wall., C. paniculata, L'Herit., C. pubescens, Nutt., C. Purpusi, Kohne,
C. siln',-ica, Loddig., and C. tatarica, Mill.
In_Mastixia Cambodiana, Pierre the petiole is supplied by three vascular
bundles which ultimately form a.central cylinder (Pierre). .
3. STRUCTURE OF'THE AxIS. According to Wangerin, 'the cork'also arises
in the subepidermal layer of' cellS in Kalip~ora ~d M elanophylld. , The
same author states that the yesse1s of the second?IY wood have' SImple
perforations in Kaliphora, and scalariform perforations iIi M elanophylta, as
well as in Alangium (Mayle-a) Mezianum, Wangerin' and A. costatum, Wangerin.
According to Pierre, M astixia Cambodian a, Pierre, like the other species
of Mastixia, is distinguished by ~ving medullary secretory canals a,nd' by the
stratification of the phloem into hard and soft bast~ .,
 ADDENDA-CORNACEAE 949
Literature: Pierre, Flore fol'eit. de la Cochinchine. xvii, 1 89:1.-[Tognini, Stomi, Atti 1st. bot.
Pavia, 18g4.J-Kohne, Papill. u. obers. SpaltoffD., Mitteil. deutsch. dendrolog. Gesellsc:h., I8~X,.
p. 59.-Petersen. Vedanatomi, 1901 , p. 68.-Bouygnes, P ~tlole, These, pa__..js, 1901, p. 13.-Gaueher)t
Hybridite, Assoc. franer. Congres- Ajaccio, 1901, ii, ed. 1903, pp. 403-8 (Garrya elfipticQ x Patlyma).
-Gerhard, Blattanat. von Gew. d. Knymawaldes, DIss., Basel.. J902) pp. 2I-l (Curliti4)..-
Tbeorin, VaxUrichom., Arkiv for Bot., i; 1903, p. 1.59.-Stissenguth, Bebaarungsverb. d WUrzba~
Mttschd.ka1kp~., Diss., Wu.rzburg, '90.... p. 35.-Pieciol!, ~ami. ~all.' Siena, 19Q6, p. 16.z.-
WangerIn, Umgrenz. u. Gliederung d. Faro. d. Cornac., DI!IS., Hal.e, 1906, 9l pp., especIally
pp. 5Q, 65, 7'l, 79 and 82; sep. copy from Engler, But. Jahrb., xxxviii.

~APRIFOLIACEAE (pp. 43~444).

t. REVIEW OF THE ANATOMICAL FEATURES. The mention of Aioxa
among the exceptions to the' occurrence of glandular hairs' must be cancelled.
2. STRUCTURE OF THE LEAF. The mesophyll also inc1.udes ann~pali·
sade parenchyma in Adoxa Moschatellina (here composed of low cel1s, Lager-
berg) and Viburnum pauci{lorum (Schwartz-Clements). Papillose diff~rentia·
tiOD of the epidermis is found on both sides of the leaf in Lonice't'a: syringantha,
lfaxim. and L. tomentula, Hook. f. et Th., on the upper side only in L. arborea
Boiss. and L. Standishi, Carr.~ and on the lower side only in l. fioribunda,
Boiss. et Buhse, L. 1upicola, Hook. f. et Th. and 1.. Zabeli, Rheder; in S'1'I1,-
plwricarpus Heyen, Dipp. and S. occidentaiis, Hook., thick conical papillae
occur in the neighbourhood of the stomata, while in S. orbiculatus, Moench,
there are weakly deVeloped papillae on the lower side of the leaf (Kohne).
The ,stomata,-and more emphasis can be laid on thjs point than formerly,-are
in no case exclusively provided.,_with subsidiary cells of the Rubiaceous type.
The club-shaped external glands found on the leaves of Adoxa are pro-
vided with a short stalk consisting of 2 or 3 cells and a bicellular head diVided
by a vertical wall (Lager berg). No~ak had previously met with external
glands, with more numerous cells, at the base of the petals.
3. STRUCTURE OF THE AxIS. The vascular system in the rhizome of Ado.:ra
M oschatellina consists of two collateral arc-shaped vascular b~ndles, which are
enveloped by a. narrow parenchymatous pericycle, a.n endodennis 'pro.Jided with
Caspary's dots and a cortex rich in starch. It is noteworthy that even the pitted
tracheae are devoid of perforations, so that they are merely elongated tracheids.
y
Likratine: Costantin J 'fjges !ler. et sout., Aim. sc, nat., sir. 6, t. xvi, J88:~, p. il et $eq.
-Kassntr, Mark, Dis!>., Breslau, 1884, pp. 15-16.-NanC'ke, Dikotyle Holzpfl .. Diss., Konigsberg,
l886, p. 3r.-Anfrec.b.t, Extraflorale Nektar., Diss., Zurich, 1892, p. ~5 et seq. (Viburnum Opu/IIS);
-Hartwich, Falsche Sen.egawurzel, Archiv d. Pharm., l33, 1895, pp. J 21-3 and Tab.-[Sayre,
Viburnum, Americ. Joum. Pharro.,.1895, p .....65.)-[Moller, Attichwurzel. Phs.nn. POllt, 1895.
p. 113 et seq.J-[Dermiston, Compo sUuet. of the barks of cert. AICeric. Viburnums, l'hann.
Archives, t, 1898.]-Kohne, Papin. u. obets. Spaitbffn., Mitteil. deutsch. dendrolog. Gesellsclt.,
1899. p. 6c.~TU!lma.lln, Selcretdriisen, Diss., Bern, 1900, pp. 43, .f4.-Petersen, VedaD~it 1901,
pp. 87-93.-Clauditz, BJattanat. kar.ar. Gew., Diss., Basel, 1902, pp. 36, 37 (Viburn,p,,).-Kuotbe,
UDbenetzb.~ Hi., Diss., Heidelberg, 19t?2, p. 9.-[Novak, Ado:ra Mtlfclzatellind, in Theodora
Novaka stati nylerana, Frag, 190:J, pp. 148-60 (Bohemian'; abstr. in Bot. CentralbL, xc, p. 546).1
-Th ou venin, Glar..des pet. du Vi.burnum Opulus, Revne gen. de bot., xv, 1903. pp. 97-I03.-Col,
FailCe1l11x, Ann. sc. cat., ser. 8, t. xx, 19°4-. pp. 139 and 13.3.--Lagerberg, Adoxa M()sckaitllina,
Arkiv f~r Dot., iii, 1904. sep. copy, 28 pp. (Swedish).-Paoli, Eterofillia. Nuovo Giom. bot. Ital.,
xi, 1904, p • .u6.-Siissellguth. BebaarnnlZ'Sverh. d. Wiirzh. MuscheikalkplJ., Dis!!., Wurzburg, 1904,
p. 3s.-Theorin, Vaxttrichom., Arkiv tOr Bot., iii, n. 5. 1904, p. 6.-Guttenberg, Lichuinnesorg.,
Her. deutsch. bot. Gesellsch., 1901\, p. ~65 et seq. and Tab. x, xi.-Haberlandt, Lichmnnesorg., 1905.
p. 69 and ·Tab. i.-Schwartz.Clement;. Relation of leaf-str>.lcture to phys. fact., Transact. Amenc.
Microscop. Soc., 1905, p. 63, pI. vii, 5 a,-Dauphine, Rhizomes, Ann. sc. nat., ser. 9, t. iii, 1906,
p. 359 {Adoxa).-Piccioli, Legnami, Bull. Siena, IS106, pp. 153 and IS6.-[Foc further litt!8.ittre..
see p. Il69.]
 ADDENDA

RUBIACi.AE (pp. 444-454).

2. STRUCTURE OF THE LEAFJ ..... Centflc structure with two layers of palisade-
cells on both sides of the leaJ occurs for instance in Asperula cynanchica
(Netolitzky), while in Bo"eria verticil/uta the mesophyll is homogeneous
(Saint-Just). Numerous papillae are found on the upper epidermis above
the larger veins in Galium cruciatum, Scop. and G; pedemontan'ltm, All. (Neto-
litzky); hypoderm is developed on the upper side of the leaf in Crateri-
spermum microdon (Fabricius), as well as in Cinchona caloptera, Miq., Hydno-
phytum tormicarum, Jack, Myrmecodia echinata, Gaud., and Scyphiphora
caryophyUacea, Gaertn. (Areschoug) and in Leucocodon reticulatum (Holter-
mann). Sclerenchymatous fibres running freely in the mesophyll are mentioned
by Fabricius as occurring in a plant described as ' Pyrostria sp. n.' and ()ne
incorrectly determined as Gardenia A nnae 2, while Frohner records them in
CoUea brackyphytla, Radlk. and C. Zanguebariae, Lour. For the structure of
the rolled leaves of Anthospermum and Nenax, see Knoblauch, loco cit ..
To the section on the hairy covering (p. 447) we may first add that the
bent unicellular hairs, found in the Steliatae, are not always typical hooked
hairs, i. e. curved like the horn of a chamois, but are sometimes merely hamose
or sickle-shaped (Netolitzky). Glandular shaggy hailS of somewhat simpler
structure than those depicted in Fig. 101 A ha~e been observed by Mardner
in Galium antarcticum, Hook. f., where they lie'at the base of the four leaves
of the whorl, and by Holtermann in H edyotis verticillaris, where they ~. found
at the bases of the leaves, which are here united to form a small ochrea; these
-glands therefore occur also on the lamina of the leaf. Regarding the glandular
shaggy hairs, see also Mastrostefano and Mirabella, ll. ce.
Elongated secretory sacs having wide lumina, filled with bright brown
contents, are found also in the veins of the leaf in Pyrostria (Fabricius), and
secretory sacs, like those present in the cortex of Cinchona, occur also in the
cortex of Corynanthe ]ohimbe, K. Sch. (Gilg); epidermal secretory sacs are found
.at the apex of the leaf in Galium hirsutum, Ruiz et Pav. (Weberbauer); the
well-known elongated secretory sacs of_ the Cinchonas are present also in the
pith (Meyer). Saint-J ust's statement as to the occurrence ~f t canaux secre-
teurs' in Manettia luteo-rubra is certainly incorrect; the same author records
secretory cells in species of Borreria, Coprosma, Ernodea, HoDmannia, Morinda,
Palicourea, SPermacoce and Vangueria, but his statements on this subject are
inexact and hence scarcely" reliable, so that we can do no more than briefly
mention them (see also under' secretory cells filled with brown contents' on
p. 448). Netolitzky associates with the secretory organs peculiar groups of
strongly enlarged epidermal cells, which have greenish, strongly refractive
contents in the form of small drops, and occur on the lower side of the leaf-tip
in certain species of Galium (G. boreale. L., G. cruciatum, Scop., G. pedemon-
tanum, All., G. rot undifoliu m; L., G. rttbioides, L.). We may likewise include
here the peculiar crypts found by Saint-Just 'on the lower side of the leaf- in
Coprosma, lucida, but not very accurately described by him; these structures.
which occur at the base of the secondary veins, are lined with a two-layered

lOur knowledge of the structure of the "leaf in the Ruhiaceae is still very insufficient. Netolitz:ky
has recently investigated the an:ttomy of the leaves of the endemic Stellatae. The statements in
Saint-Julit's paper, which also deals witk tbis Order, are unfortunately very inexact and in part
incorrect; moreover some of his plants are certainly incorrectly determined, as may be concluded
from the mode of deposition of oxalate of lime.
, In view of the presence of the characteristic crystal-hairs this plant must be one of the
Guettardeae. the' secretory cells with dark granular contents' observed by Fabricius are doubt1es~
sacs containing crystal·sand.
 ADDENDA-RUBIACEAE 95 1
epithelium of elongated palisade-like cells, and open towards the outside by
means of a narrow aperture, covered in by trithomes.
To the synopsis of the diverse crysta1lino forms in which oxalate of lime is
deposited in the genera of the Rubiaceae (pp. 450, 45 I) we may add: Under
I. Crystal-sand: after Condaminea: Cor'Ynanth~; further, under 'together with
the crystal-sand, clustered crystals': Cinchona. Under II. Raphides : after Cruck-
shanksia in -brackets: C. Monttiana, Gay; further at the points determined by
the alphabetical se~uence: Coprosma, Hydnophytum. Myrmecodia and Oreopolus
(Cruckshanksia glac1,alis, Poepp.). Under III. Clustered crystals: Pyrostn·a. We
roay also note that in some cases there are no raphides in the raphide-sacs, so that
the lattc;r are accompanied by mucilage-sacs (e. g. commonly in the leaf of Asperula
odorata, L.).
Netolitzky met with sphaerocrystaUine masses having the appearance of
hesperidin.. crystals in alcohol-material ·and glycerine-preparations of many
of the Stellatae (e. g. Galium lucidum, All. and G. palustre, L.). In the bast
of some kinds of Cinchona-bark, which are rich in alkaloid, the latter occa-
sionally crystallizes out.
3. STRUCTURE OF THE AXIS. According to Saint- JUS4 the cork develops
superficially (viz. in the subepidermal layer or in the second layer of primary
cortical cells) in the species of Chione, Faramea, Gardenia, Genipa, Guettarda,
/xora, Laugtt.eria, Pentas, Randia, Rondeletia, Rudgea, Strumpfia examined
by him, while it arises in the pericyc1e in the species of Coprosma, Leptodermis
and Paederia. In the species, of Chimarrhis, Faramea, Guettarda, Hamelia,
Hyptiafllhera, [:;cora, Morinda, Mussaenda, Paederia, Pavetta, Plectronia,
Posoqueria, Psathura, Psychotria, UrophyUum and Vangueria investigated by
Pitard the peri cycle contains isolated bundles of hast-fibres j the laUer are
wanting, however, in the officinal Cinchona-bark. As regards the structure of
the wood we may mention that Craterisperm1.tm microdon, Bak., like the other
Vangueriea~, has simple perforations in the vesselS and wood-prosenchyma
bearing bordered pits (Ursprung).
According to Goldstein, cortical vascular bundles showing concentric
structure are found in Sickingia. Baranetzky's statement as to the occurrence
of bicollateral vascUlar bundles in a member of this Order (Plt3ctronia ventosa)
is incorrect; the plant examined by him is not Plectronia ventosa, nor even one
of the Rubiaceae (Solereder). /'
Literature: Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, p. I.p et set{.-
Goldstein, _Rinde von Arariba, etc., Diss., Erlangen, 1892, 30 pp., 2 Tab.-Brandt, Wenig bek.
Rinden, Diss., Dorpat, 1894, p. 52 et seq.-[Tognini, Stomi, Atti I.~t. bot. Pavia, 1894.]-Jonsson,
Anat. Ban d. H1., Acta Univ. Lund., xxxii, 2, I 896.-Knoblauch, Okolog. Anat., Habilitat.~Schr.,
Tubingen, 1896, p. 9 et seq.-Elfstrand, Heilptl., Ber. deutsch. pbarm. GeseUsch., [897, p. 29 1
(PaiicoU,.tll).-[Mastrostefano, Stellate, BalleU. Soc. Natural. Napoli, 1897, p. 75; abstr. in Just,
1898, ii, p: 27o.)-[Mirabella, Colleteri, Contrib. 1st. bot. Palermo, ii, 1897, p. IS et seq.; abstr. in
Just, 1897, i, p. 5I3.]-Scbubert,J'arenchymscheiden, Bot. Centralbl., 1897, iv, p. 63.-Boergesen og
-J1auisen, Veget. dansk~vtstind. Oer, Bot. Tidsskrift, xxii, 1898-9, pp. 96, 97 (Rant/ia acultala,
L.).-Frohner, Cojfia, in Engler, Bot. Jahrb., xxv, 1898, p. 236 et seq.-Tichomuow, Meehan.
Elemente- bei Cinchona, Bot. Centralbl., 1899,,1, p. 60.-Baranetzky, Faisc. bicoll., Ann. sc. nat.,
ser. 8, t. xii, 1900, pp. 292-4.-Charpentier, Et. anat. et microchim. d. quinquinas de cult., These,
Paris, 1900, 60 pp., 2 pl.-Gamper, Angostorarinden, Diss., Zurich, 1900, pp. 6:1; 63.-G. Meyer,
Anat. d. auf Java knIt. Cinebanen, Zeitschr. f. Naturw., lxxii, 1900, pp. 409-+1; see also Hartwich
and G. Meyer, in Arehiv d. Pbarm., 238, 1900, p. 253. - Ursprung, Anat. u. Jahresringbild. trop.
Holzarten, Diss., Hasel, 1900, pp. 23-5 (Craten'sptrmum micriJdon, Bak.).-Gilg and Schumann,
Johimberinde, Berliner Notizbl., iii, no. 25, 1901, pp. 92-4; see also Gilg, ill Ber. deutsch. Pbarm.
Gesellscb., 1901, p. 2I 2.-Pitard, Pericycle, These, Bordeaux, 1901, p. 9J.-Areschoog, Mangrovepfl.,
Bibl. bot., Heit 56, 11,102, pp. 52-5 and ~b. v, vi.-Bargagli~Petrucci, Legna.mi, Malpighia, 190 1 ,
p. 565 et seq. (.ll.Juss~ndtlpsis, Sarcocephaiw).-Clauditz, Blattanat. kallar. Gew., Diss., Basel, 1902,
pp. 36, 37 (PhyIHs).-Fauricius, Laubbl..Anat., Beih. z. Bot. Centralbl., xii, 190 2 , PP. 3[4- 1 7.-
Gerhard, Blattanat. v. Gew. d. Knysnawaldes, Diss., Basel, 1902, pp. 22, 23 (Plectronia).-Kno~he,
l]Dbenetzb. Bl., Diss., Heidelberg, 19°2, p. 9.-Mardner, Phan.-Vegetat. d. Kerguelen. DlSS.,
Hasel, 1902, pp. 27.28 (Gczlium).-[Armari, Piante della reg. medit., Ann. di bot., 1,1903, p. 17 et
 ADDENDA-RUBIAcEAE
seq. (Putof';a).J-GorrisetReimen, Cinc/t(Jna rtJ/Jus/a, Bull. Soc. pharoulcol., 1903. pp. 38....-6; 'also
in Perrot, Travaux, II.-[Greensill, Struct. of leaf of cert. spec. of CO/Jrls11Ia, Transact. and ProCeed.
New Zealand lost., xxxv, 1903, PP. 34'-.55 and pi. xli-xliv.]-Soleteder, Berichtig., Bull. Herbier
Boissier, ae str., 1903, pp. 341 ...., 3:Js.-AcliD~r, Falsche Cbinarinden, Dis.o., Bern, 1904, 109 pp.-Col,
Fais()eaux, Ann. sc. 03.t., Set. 8, t. xx, 19Oi. flP. I19-:n.-Saint-Just, Rech. anat. sur l'appareil veg.
aerien des Rubiac., These, Paris, I90"h 70 pp., ... pl.-Sussenguth, Behnarungsverh. d. Wiirzb.
Mnsc;:helkaL~pfI., Diss., Wiirz,burg, 190., p. 36.-Areschoug, Trop. vaxt. bladbyggn., Sv. Vet. Alcad.
Handl., 39, D. ~, 1905. pp. 50, 51 (Spermac~), pp. 73, 74 (Den/ella), pp. 86, 87 and Tab. xiii
(Cinckona) , pp. 90,91 (CI1/fia), p. 108 (ilfyrmecodia), and pp. 144-6 and Tab. vi-viii(HJ!dnophylkm).
-Nttolitzky, DikotylenbT., Vienna, 1905. pp. 9-33.-Russell, Principes actifs de In Garance, R~De
gen. de bot., 1905, p. :i54 et seq.-Theotin, Vaxttrichom., Arkiv for Bot., iv, n~ 18, 1905, p. li.-
Webetbauer, Vegetat. d. Hochanden P~rns, in Engler, Bot. Jahrb., xxxvii, lQOS, p. 60 et seq.-
Holterman~, EinfJnss des Klimas etc., 1907, pp. 59, 83, 130 and 135 (SCJlj>kipk0trt. Sprmacoce-,
Hedyotis, Leucocodon).-[For fllrther literature, see p. II71.]

VALERIANEAE (pp. 454, 455).

, The following details 'may be added to the account given in the earlier part
of this book; they are mainly extracted from Vidal's work. In the species of
Patrinia, 'Centranthus, Valerian~, Pedia and ValerianeUa examined by Vidal the
structure of the leaf is again mostly bifacial, although it is centric in Valericma
saliunca and almost homogeneous in V.: dioica. 'Twin stomata are frequently
met with on both surfaces of the leaf in Valerianella; water-pores ~e fOlUld
also in Valeriana Phu. The lateral margins of the epidermal cells are either
straight or undulated. Vidal observed glandular hairs in all the species; the
multicellular ~ea9s of these hairs are either elongated or spherical and are for
the most part divided both'by horizontal and vertical wails, rarely (Valeriana
tripteris) by vertical walls only. As regards the fibrovascUlar system of the
leaf. we may note that there is no mechanical tissue in the veins apart from
collenchyma. The leaf is supplied by three vascular bundles, which in the
upper portion of the petiole pass (wer, into a widely open arc com~ed of. five
bundles. In Valeriana officinalis, according to Bouygues, the petIole contains
several concentric vascular bundles (steles). some of which are not quite com-
pletely differentiated j each of these bundles is provided with ,a pith and an
enveloping endodennis. Oxalate of 1ime has been found in the vegetative
organs 1 only in Patrinia, where 'it was first observed by Chatin; it occurs
in the form of clustered crystals, which are present in the root, stem, and
leai. On the other hand, structures of unknown chemical composition,- but
resembling sphaerites, appear in the leaves and other organs of many Valerianeae
(e. g. Valeriana saliunca) after treatment with alcohol;' these bodies recall the
sphaerocrystals of inulin. Oi1~cells, first observed by Zacharias in the root of
VaZeriana and distinguished from the neighbouring cells by their contents
and their suberized wall, are, to judge by Vidal's statements, present in the
root also in other species 'of Valeriana, as well as in Pat,-inia~ Centranthus,
Fed,'a and Valer,'anella; they are situated eithe~ in the second cell·layer of
the primary cortex beneath the suberized outermost layer, or are found in
a slightly deeper layer. According to Vidal, however1 the chief production
of oil in the roots takes place in the cork, which explains the large amount of
oil found in the older roots.
For details as to the structure of the stem, rhizome, and root, see Vidal,
loe. cit.; it will suffice to mention the following facts here. Apart from the
superficial development of the cork in the roots and rhizomes of Valeriana
otficinalis and V. Phu, the cork invariably arises in the _pericycle in both'organs.
There are no mechanical elements in the pericyc]e. The lnner 'portion of the

1 According to Vidal, the pericarp of Cm/rantnus, Va/eriana, Fedia and Valtritllulla contains
oxalate of lime in the fonn of solitary crystals, although there is no oxalate of lime in th~ vegetative
organs. ' C '
 ADDENDA-VALERIANEAE 953
pith of the stem consists of unlignified cells and subsequently becomes fistular.
In the- roots of Vale,iana otficinalis and V. Phu the cells of the endodennis,
which are provided with Caspary's dots, are persistent in. consequence of the
superficial development of the cork and etfilbit secondary diVlSionewalls in
the same Ylay as in: certain Gentianeae.
Literature: Castanon, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, p. I..j.6.-[Tognini,
Stami, Atti _1st. bot. Pavia, 1894.]-Biermann, C:>lzellen, Diss., Bern, 1898, pp. 30-5.-Tho1Das,
Feuilles sont., These, Paris, 1900.-Dye, Unterird. Org. yon V41eriana etc., Diss., Bern, 1901,
pp. 8-35 and Tab. I.-Bouygul!S, Petiole, Tbese, Paris, 190:3', p. 1'.-L. Vidal, Anat. des Valerian.,
Ann. de l'Univ. de Grenoble, xv, 1903, sep. copy, especially pp. ~31; see also the authors abstract
in Bot. CentT1}lbl., xcv, p. I 46.-Co1, Faisceaux, Ann. sc. nat., ser. 8, t. xii, 1904, p. 136.-
Webert~a:ner, Vegetat. d. Hochanden Perus, In Engler, Bot. Jahrb., xxxvii, ]905, p. 60 et seq.

DIPSACEAE (pp. 455, 456)~

Szabo's recent investigation of the anatomy of the genus Knautia has
afforded the following_results. External glands with a short stalk and a head
tomposed of four or more cells occur also in this genus. K. orientalis, unlike
other species of the genus, has scalariform perforations iI! the vessels. Tb:e
cork in K. longifolia (rhizome) develops jn the pericycle.
Literatnre: Costantin, Tiges aero et sout.) Ann. sc. nat., ser. 6, t. xvi, 1883, p. 1-45 et seq.-
Mittmann, Pilan:renstacbeln, Verh. bot. Ver. Brandenburg, 1889, p. 67 and Tab. I.-Lothelier, Epines,
iii.-~~acace, Contri~. stud. d:. Dips.acee, Ric. ~tol.. R. Orto bot.
These,. Paris, 1893, p. 3.9 and pI.
Napoli, 1898, I:I pp.J-Rostock, Drusenh. -Yon DipstICW .II/vesJru, Bot. ZeIt., 1904, Abt. I,
pp. I I-~O.-SijsseDguth, Behaarungsverh. d. Wiirzb. Muschelkalkpfl., Diss., Wiirzburg, 1904, p. 36.-
~zabo, Monogr. d. Gatt.l!nautia, in Engler, Bot. Jihrb.,/xxvi, 1905, pp. 401-8.

CALYCEREAE (p. 456).

According to Reiche, • the absence of special bast-strands in
the cortical paren·
chyma' is a feature common to all the Calycereae. Boopi$ australis, Decne.,
B. g"acilis, Phil., B. m'ulticaulis, Phil. and B. g"aminea, Phil. possess' a lignified
strengthening ring. in which the vascular bundles are embedded. I The fleshy
stems found in various species of Nastanthus are due to abundant development
of the medullary and cortical parenchyma.
Literature: Reiche, Calycereen, in Engler, Bot. Jahrb., xxix, 1900, p. lto.

COMPOSITAE (pp. 4-56-4-69).

I. REVIEW OF THE ANATOMICAL FEATURES. Secretory cavities also occur
in the leaves of Trilisa and species of Athanasla; in certain members of
the Order. moreover, they take the place of ,the secretory canals in the sub-
terranean organs. Secretory sacs, which are often much elongated and' have
milky or resinous~coIitents', are-:-found in species of Gazania (Tribe Arctotideae,
in this case together with transitions to laticiferous vessels), and Vernonia (Tribe
Vemonieae), as well as in members of the Tribe Cynaroideae (Alft'edia, Arctium,
Atractylis, Bel'ardia, Carduus; Carlina. -Chal'dinia, Cil'sium. Cousinia, Galactites.
]u,inea, Lappa, Onopordon., Silybum, Staehelina, lYarionia; in Atracfylis and
Carlina, together with transitions to laticiferous vessels). Beyond the limits
of the Cichoriaceae and apart from the genus Gundelia (Tribe Arctotideae),
which was already referred to in the. earlier part of this work, laticiferous
vessels occur in Gazania, which belongs to the same Tribe as Gundelia, and
in Carlina and Atractylis (Tribe Cynaroideae, Sub tribe Carlineae). Unicellular
trichomes appear'to be wanting in the hairy covering of the Compositae.
Recent investigations have shown that multicellular clothing hairs with a
954 ADDENDA-COMPOSITAE

two-armed or stellate terminal cell occur in many additional genera. Vesicular

integumental glands, showing the same type of structure as those found in
the Labiatae, are present in Centauropsis, while extrafloral nectaries occur in
Helianthus. To the list of special or anomalous structural features presented
by the axis we may add: the arrangement of the vascular bundles in indistinct
rings, as in the stem of a Monocotyledon, in species of Scorzonera. and Cen-
taurea, as well as in speci€} of Cynara; and the occurrence of medullary or
cortical vascular bundles in species in which .these anomalies were not pre-
viously known. Anomalous structure has recently been demonstrated in
the fleshy adventitious roots of Thrincia tuberosa, DC. (transformation of the
primary xylem-groups of the vascular system into stele~) and in th~ tap-root
of Atractylis gummilera, L. (appearance of concentric secondary meristems
in the wood of the root, and development of rings of vascular bundles, which
alternately show normal and inverse. orientation).
2. SPECIAL ANATOMICAL DIAGNOSIS. A very important feature of the
hairy covering (see p. 457) lies in the fact, that unicellular hairs really
appear to be absent in the Compositae, just as in the Papilionaceae. G. Fischer's
statements as to the presence of unicellular hairs in certain species of
genera belonging to the Vemonieae and Eupatorieae (Agrianthus, Bolanosa,
HoPlophyllum, Piptolepis, Ver-nonia) have proved to be incorrect upon rein-
vestigation, the basal cells of the hairs having been _overlooked. Recent
observations have shown that simple uniseriate clothing .hairs with a long
terminal cell (flagellum-trichomes) are very widely distributed. They occur
in many genera of the Vernonieae and Eupatorieae, e.g. in species of Adeno-
styles, Bothriocline, Brickellia, Centauropsis, Elephantopus, Liatris, Rolandra,
Soaresia, Telmatophila, Vernonia, &c. (G. Fischer), as well as in species of
Anacyclus, Athanasia, Culcitium, Hieracium, Lucilia and Senecio. 'In Balanosa
Coul/eri, Gray and Stilpnopappus specioslts, Bak., the terminal cells of these
flagellum-hairs exhibit a nodose thickening at their base (G. Fischer), while
in Cirsium horridum, M. Bieb., the basal portion of the·terminal cell is enlarged
in a campanulate manner; in certain species of Senecio (Wagner) and Anacy-
clus (Feuilloux) the terminal cells are placed at an angle to the stalk, in the
same way as in the hairs of Brocchia cinerea (see p. 458). Uniseriate hairs
with a two-armed terminal cell, which in other respects shows varied structure,
have been demonstrated also in certain species of Albertinia, Centauropsis,
Eremanthus, Lychnophora. Oliganthes, Piptolepis, Pithecoseris, Vanillosmopsis,
and Vernonia by G. Fischer, and in certain species of Anacyclus and Athanasia
by Feuilloux; I have found them also in Senecio incanus, L. and S. uni/lorus,
All.) which is not in agreement with P. Wagner, who speaks of pelt ate hairs
as occurring in these species. Uniseriate hairs with a stellate terminal cell,
which varies in the thickness of its walls and in the number of rays, occur
also in species of Blanchetia, Haplostephium, Lycnophora, Piprocarpha and
Piptolepis (according to G. Fischer), in species of Artemisia and Athanasia
(according to Fe uilloux) , in species of Vernonia (according to Grimm), and
apparently also in the genera Baccharis and Santolinum (according to A. Weiss) ;
uniseriate hairs with a candelabra-like terminal cell are found in Scorzonera
hispanica, L. (Theorin). These different· forms of hair in which the terminal
cells aTe simple, two-armed or stellate are as a' rule only characteristic of the
species and not of the genus. Thus, according to Feuilloux, all three types of
hair are found within the limits of the genera Artemisia and Athanasia (hairs
with a simple terminal cell, for example, in Artemisia ramosa, Sm. and ' Atka-
nasia parvi/lora, L.'; hairs with a two-armed terminal cell in Artemisia
Absinthit,m, L. and Atlzanasia pin nata, L.; and hairs with a stellate end·cell
in Artemisia glauca, Pan. and Athanasia pubescens, L.), while trichomes with
a simple and with a two-armed terminal cell.both occur in the genus Anacyclus
 ADDENDA-COfifPOSITAE 955
(hairs with a simple end·cell, for example, in A. clavatus, Pers., and two-armed
hairs in A. pedunculatus, Pers.); further, according to G. Fischer, trichomes
with a two-armed and with a stellate terminal cell are found side by side in
Lychnophora salic~jolia, Mart. and Piptolep~'s er£coides, Schultz Bip., and hairs
with simple and two-armed. end-cells side by side in Centauropsis lanuginosa,
Boj. Clothing hairs of the shaggy type have recently been observed by
G. Fischer also in Hofmeisteria pluriseta, Gray (here provided with a papillose
surface), Piptocarpha lucida, Bak. (likewise papillose) and Vernonia inulaefolia,
Steud., as well as by Noe von Archenegg in C~'rs.ium horn'dum, M. Bieb., and
other species of. Cirsium belonging to the section Epitrachys. The shaggy
hairs found in the species of Cirsium occur on the upper side of the leaf and
are composed of pitted sclerenchymatous fibres, which penetrate into the
mesophyll in the form of a bundle and thus constitute a base to the' shaggy
hair, similar to that found in many Melastomaceae; ultimately the bundle
of fibres becomes apposed to the vascular bundle of a vein. Both the epidermis
and the subepidermal layer of cells are concerned in the formation of these
shaggy hairs.· A transition to the shaggy hairs is represented. among other
fonDS. of trichomes, by the clothing hairs of Artemisia argentea, L'Herit.,
which are provided with a narrow elongated t~rminaI cell and are seated on
a high pedestal, the lower tiers of which are formed. of. several cells. Charac-
teristic shaggy hairs, showing a reduced type of structure and similar to those
found in Spilanthes oleracea (Fig. 103, Nand p. 460), appear to be rather widely
distributed on the floral parts (ovary) in the Compositae; they consist of two
basal cells, which are prolonged upwards to an unequal extent, and two long
hair-cells, which are joined together lengthways along one side, but diverge
apkally so· as to resemble a swallow's tail; hair£. of this kind are found, for
example, a}!O in Atractylis gummifera, L. and Carlina acaulis, L. (Kerckp.off).
With these hairs we may class the peculiar trichomes, observed by Schinz on
the same organs of the plant in Sphaeranthus epigaeus, Schinz; their dis-
tinctive feature lies in the fact that the two terminal cells, which have rounded
ends, are spirally wound around one another (with two turns to the spiral).
We will next, consider the glandular hairs. As far as can be. gathered
from G. Fischer's statements, which are unfortunately in part incompletE}
and not clearly presented, the characteristic biseri'J,te vesicular integumental
glands were observed by him in certain Vernonieae {nd Eupatoneae; Tunmann
also met with them in A chillea. If we imagine the appearance of vertical
and oblique ,division-walls in these biseriate glands we obtain the shortly
stalked external glands found in Haplopappus and Gochnatia; in these glands,
which were not noticed in the previous review, the cells are arranged in
the three directions of space (Volkens). Multicellular glandular hairs with
a unicellular head are found in species of Brickellia, Kuhnia, and Trichocoronis
(according to G. Fischer), in Senecio Boissieri, DC. (according to P. Wagner).
and in Centaurea (according to Briquet); almost sessile glands with two
short basal cell? and a secretory head-cell are :figured by Feuilloux in Athanas1:a
leucoclada arid glandular hairs with a- uni- or biseriate stalk and a spherical
head, divided by horizontal and vertical walls into 4-8, cells, by G. Fischer
in Hofmeisteria pluriseta, Gray; vesicular integumental, glands having a short
stalk and resembling the glands of the Labiatae, occur in Centauropsis la-wu·
ginosa, Boj. The extrafioral nectaries, observed by Delpino in Hel£anthus
giganteus and H. tuberosus, have not yet been closely examined; they are
situated on the uppermost leaves (on the lower side near the base oi,the leaf)
in the neighbourhood of the floral region.
Our previous knowledge regarding the internal secretory organs of the
Compositae (see p. 460 et seq.) has been very considerably extended, especially
by Col's recent investigations.
 ADDENDA-COMPOSlTAE

Schiwgenous secretory canals are found in certain members of all the

thirteen Tribes include. in Bentham and Hooker's system, although they
are of rare occurrence m the Cichoriaceae~ In this Tribe they are to some
extent replaced by laticiferous vessels, just as in certain Cynaroideae their
place is taken by secretory sacs with milky or resinous contents. The charac~
teristicposition in which they occur is in the root; this statement must not,
however, be taken to imply that they are not very frequently found traversing
the lubaerial portions of the plant as well (i. e. axis and leaf). No case is known
among the Compositae of the occurrence of secretory canals in the shoot when
they are absent in the root; it has also been shown that in those forms which
have still other types of secretory organs, such as laticiferous vessels or secretory
sacs, the secretory canals, so to say, withdraw mto the lower organs of the
plant and ultimately become confined to the root. These statements already
serve to show that, when undertaking a systematic-anatomical inves~ation
of the Compositae, an examination of the root is of .primary importance,
while, if possible, the entire vegetative organs should be examined from
below upwards, in order to determine the distribution of the secretory canals.
and of the internal secretory receptacles in general. As far as can be gathered
from the literature, moreover, the secretory canals of the root are always
distinctly endodermaLin origin, while this is .not always so in the case of the
secretory (..'atlals of the axis. In many cases,' it is true, the secretory canals
are observed to originate in the endodennis ill the axis also (see the examples
mentioned in Col's Thesis, p. 99), the canals thus formed being for the most
part narrow. In other cases, however, the secretory canals of the axis develop
at a very early stage and have no connexion with. the endodeimis, which only
appears subsequently; they seem to. belong to the primazy cortical tissue
and are of wide diameter. This occurs very commonly in th~ Heliantheae,
which sometimes have medullary canals as well, so that the secretory system
of the axis, as in the case of SiZphium, recalls that of the Umbelliferae.
Th.e previous synopsis of the arrangetnent of the cauline resin-callals in relation
to the fibrovascular system (see the second paragraph of small print on p. 461)
has been extended and improvtd by Col, who likewise attributes systematic impor4
tance to these features; we Can do no more than briefly refer the reader to pp. 90-3
of his thesis.
We may also point out here that in some cases in conjunction with repeated
divisions in the endodennal cells th.e resin-canals of the root may occur in two or
more rows, e. g. in Platycarpha. glomerata, Less., Aretatis stoechadi/olia, Berg and
Venidium calendulaceum, Less.
It has aJieady been mentioned in the earlier portion of this work (pp. 46r,
462) that, side by side with the endodennal secretory canals,.resin-canais have
been demonstrated in other positions in L'le stem, rhizome, or root of certain
members of the Order; these canals are situated in the primary cortex, in
the periphery of the pith, in· the secondary tissues (wood and bast) of the
vascular system, or between the pericycle and the phloem. The earlier state-
ments on this subject require a few additions and ·alterations. Col mentions
the occurrence of medullary secretory canals in species of a few further genera,
vi.z. Baltimora, Carlina, Centaurea, Chrysanthemum, Cirsium, Cynara, Gail~
lardia (rhizome), .Ha.rpalium (= Helianthus), Podachaenium, Polymm:a, Tridax
and Zaluzania, while the secretory organs present in the pith of the rhizome
in l1t.ula Hdenium, L. and Helenium autum-nale, L. are of the nature of cavities.
Secretory canals'are ~tributed in the secondary tissues. of the vascular system
as follows: (a) in the secondary bast: in the stem and 'root of H dianthus
tuberosus and· Centaurea airopurp,urea, in the rhizome of Carduncellus mo~
spelie1'.,sium, All., andin the rl}Ot of Kentrophyllum lanatum, DC.; (b) in the
medullary rays of the wood and bast: in the fleshy roots of Dahlia; and
 ADDENDA-COMPOSITAE 957
(c) in the wood and bast: in' the tubers of Helianthus tuberosus. Secreto.ry
cavities are found in place of the secretory canals: (,) in the medullary rays,
of the bast: in the root and rhizome of Artemisia. Dracunculus, and in the
subterranean organs of Atractylis gummifera, L.; (b) in the medullary rays
of the bast and wood: in the root of A nacyclus Pyrethrum and CarUna acaulis ;
and (c) in the bast and wood: in the root and rhizome of Inula Helenium.
We may replace the former synopsis (d. p. 462) of the genera of Composi-
tae in which the subaerial parts of the plant contain no resin-canals, by the
following enumeration taken from Col's thesis. The resin-canals do not pene-
trate into the subaerial axis in species of: Acroclinium (= Helipterum),
A ctinclepis, A mblyolepis (= H elenium). A mmobium, A naphalis, A ntennaria,
Anthemis (pro parte?) 1, Arctotis, Asteriscus (= Odomospermum), Baeria,
Calendula, Calocephalus, Cardopatium, Cephalophora, Cryptostemma, Cupularia
(= I~). Dimorphotheca, Echinops, Elephantopus, Ethulia, Eurybia (=Olearia),
Evax, Filago, Gaillardia, Gnaphalium, Helenium, Helichrysum, Hel~m,
Hum-ea, Jasonia.lnula (pro parte). Layia (?)2, Leontopodium, Leyssera, Mtcropus,
PaUenis, Phagnaion, Pinardia (= Aster 7), Podolepis, PotUJtheca, Rlwdanthe
(= H elipterum). Rhynchopsidium (= Relhania); Sphencgyne (= Ursinia), TaT-
clwnanthus, Tripteris, Venidium, Vernonia pro parte {V. anthelmintica, Willd.).,
Xeranthemum. To this list we may add the species of Achyropappus, Madaria,
Madia and Schkuhria, which have been examined, although m these cases the
secretory canals are absent only from the upper parts of the axis. In the new
enumeration, just given, the Cynaroideae which are provided with secretory
sacs have not been taken into consideration (see under secretory sacs, below) ;
moreover, the genera Centaurea, Eupatorium and Mada,ia which were included
in the earlier list have been omitted, since they possess secretory canals in
the different parts of the axis; neither have I included the genera of the
Labiatifiorae (Tribe Mutisiaceae), mentioned On p. 462 as having no resin·
canals in the stem and leaf, since no new investigations dealing with this group
have been published and the old statements retain their value. The number
of species and genera, as yet examined in the various Tribes, is far too small
to admit of a correct appreciation of the systematic value to be attached to
the absence of the secretory canals in the subaerial organs. Yet it is'a striking
fact that resin-canals are wanting in the subaerial organs in all the Calendulaceae
which have been investigated, as well as in those Inuloideae, which are included
in the .. first seven subtribes (Tarchonantheae to Athrixieae) of Bentham and
Hooker's system. But, on the other h:and, we may also point out that,
according to Col, within the limits of the genus Inula, I. crithmoides, L., for
instance, has secretory canals in all the vegetative organs, while in I. hirla, L.
they are confined to the rhizome and are not found in the subaerial shoots.
As regards the occurrence of secretory canals (side by side with latici·
ferous vessels) in certain Cichoriaceae (d. pp. 462, 463), we may add that they
are present not only in Scolymus grandi/lorus, but also in S. hispanicus and
S. maculatus,- that-is to say in-all the species of the genus (Col and Kniep).
Doubling of the endodermis in the absence of resin·canals has also been observed
in the root of PodospeTmum laciniatum (Col).
According to Feuilloux, secretory cavities (see p. 463) also occur in the leaf
in numerous species of Athanasia (but not in A. leucoclada and A. pinnata, L.),
and, if 1 understand Paschkis rightly, they are present also in the mesophyll
of Liatris odoratissima, WiUd. (= Trilisa odoratissima, Cass.). It'remains to
investigate whether the cavities in these plants and in those forme~ly named
take the place of the secretory canals in the leaf or whether they exist side by

1 According to Co), Alllnemis uooilis has resin· canals in the base of the stem Ol1Ir.
• The ba5.e of the stem ha5. not yet heen investigated in this case.
 ADDENDA-COMPOS/TAE

side "ith them. For it' has been shown that in other organs the secretory
cavities are occasionally vicarious in their occurrence.
Secretory cavities have been observed in organs oilier than the leaf in. the
following Compositae: A nacyclus Pyrethrum (medullary ra.ys of the wood and
bast of the root); A rtemisia Dracunculus (primary CQrtex and medullary rays of
the bast of the root, medullary rays of the bast of the rhizome); Atracty/is gum-
mite1'tJ, L. (medullary rays of. the bast of the subterranean organs); Carlina acaulis
(medullary rays of the wood and bast of the root); Cirsium -lanceolatum, Scop.
(at certain levels in the stem in place of the secretory canals), C. rivulare, Link
(in the rhizome, replacing the secretory canals); Echinops Ritro, L. and E. sphae1'o-
cephalus, L. (endodermis of the rhizome); Ga#lardia pt'lche/la (in the axis of the
shoot to the right and left of the points of insertion of the leaves, somewhat of the
nature of canals); Helenium autumnale, L. (pith, primary cortex, and endodermis
of the rhizome); Inuta Hetem;um, L. (bast and wood of·the root and rhizome; in
the latter also in the pith); Inula britanm·ca, I. Conyza. DC., also I. bifrons, L. (endo-
dermis of the rhizome); Onopordon AcantMum, L. (cndodermis of the base of·the
shoot).
, . ,.
There is nothing to add regarding the laticiferous vessels of the Cichoria-
ceae (p, 463), but a few new facts as to the occurrence of these secretory organs
outside this Tribe (d. p. 463) have become known. Laticiferous vessels had
previously been demonstrated in the pericycle and' bast of the axis, as well
as in the veins and ground tissue of the leaf in Gttndelia Tournejortii, L., a
member of the Arctotideae. Col has since found them in a!}other genus of the
Arctotideae (viz. in Gazania splendens x Hort. AngJ.1) -as well as in Carlin a
(Tribe Cynaroideae, Subtribe Carlineae 2), while Kerckhoff records them in
Atractylis, a genus very closely related to Carlina.
The laticiferous vessels of Ga.:ania splendtm.s show a primitive type of struc.;
ture. Where the cells abut upon one another their longitudinal walls are locally
resorbed, but true transverse bridges are not formed; some of the .transverse
walls, moreover, are persistent. Further, the laticiferous vessels of Gazania
are confined to the pericycle of the axis and the veins of the leaf; in the
root they are replaced by latex-cells, which are either isolated or arranged in
longitudinal rows and are situated in the secondary bast. According to Kerckhoff,
the laticiferous vessels of Atractylis gummifera are identical in structure with those
of Gazania; they OCC1J.r in the secondary bast. of the subterranean parts of the
plant, especially in the tap-root and in the axial parts of the root-stock.
As regards Car/ina, lastly. Col publishes the following details. In C. caule-
scens, Lam. the pericycle of the subaerial axis includes elements of the nature of
laticiferous vessels, although their origin by fusion is very difficult to determine.
In C. acanthifolia. All. the root and shoot contain laticiferous vessels composed
of relatively short cells in which some of ·the transverse walls are Dot absorbed;
these elements occur in the bast and pericycle respectively. The fact that in
Gazam'a the laticiferous vessels are replaced by latcx-cells and that CarUna vulgaris,
L. has latex-cells only (in the pericycle of the axis) and no laticiferous vessels,
indicates the close relationship of the two types of secretory elements. In order
to complete the account of the secretory organs found in those genera which are
provided with laticiferous vessels, we may mention that in Gazania endodermal
secretory canals occur only in the root (in addition to the latex-cells in the bast
already referred to above), that in Atractylis secretory cavities are found in the
subterranean organs, and that Ca1'l£na has both secretory cavities and endodermal
secretory canals,-the latter, for example, in the Toot of C. caulescens and in the
root arid a.."'{is of C. vulgaris.

1 Other members of the Arctotideae examined by Col do not possess "these· secretory organs,
e, g. Platycarpka, in which endodennal secretory canals have been. demonstrated in the root, and
A,·c/olt's, CrYPlosltlllma and Veuidium, which have endodennal,secretory canals in the root only and
not in Ihe stem.
, ~ In other members· of the- Subtribe Carlineae, which he investigated, Col did not observe
hticiferons vessels, but mNely rows of latex·cells, nor ,,·ere these found in every case.
 ADDENDA-COlffPOSITAE 959
In place of the earlier statements regarding secretory cells with resinous
or milky contents (see the first paragraph on p. 464) the following details
may be inserted. The secretory sacs in question, which are often the cause
of a very considerable exudation of milky juice from the organs in which they
occur, are closely related to the laticiferous vessels by 'means of the transitional
forms above discussed; they are 'for the most part much elongated, and,
as a rule, occupy the same position as the laticiferous vessels, i. e. they -are
found on the inner side of the endodermis, external to ihe bast-groups of the
vascular ;system. \Vhenever there is a development of primary hard bast,
the secretory sacs are apposed to its outer side; in some cases they even
penetrate into the group of primary hard bast itself or are embedded in it.
In certain species the secretory sacs are found also at the periphery of the
pith, very rarely in the bast as well. As regards their distribution we may
first mention that they occur only in a few representatives of the Arctotideae
and Vemonieae, but are more commonly found in the Tribe Cynaroideae;
in the latter, ·however, they are confined to the Subtribes Carlineae and Car-
duineae (being absent in the Subtribes Echinopsideae and Centaureeae) 1.
Secretory sacs have been observed in the following species: I. In the tribe
Arctotideae: in Gazania sptendens (in the secondary bast of the root; for the
laticiferous vessels present in the pericycle of the axis, see above). II. In the Tribe
Vernocieae: in Vernonia praealta (outer portion of the primary bast of the root;
pericycle of the rhizome; pith, primary cortex, and bast of the axis of the shoot;
parenchyma of the veins and mesophyll of the leaf), V. arkansana, DC., V. eminens,
Bisch .• V. j1e:tuosa, Sims., V. noveboracen~is. Willd, , but not iri V. anthelmintica.
Wilid. III. In the Tribe Cynaroideae: I. Subtribe Carlineae: in Atractylis cancel-
lata; L. (pericycle of the axis). Chardinia xeranthemoides, DesL (as in the preceding
species), Carlina graeca, C. racemosa, L. and C. 1Julgaris, L. (as in the preceding
species; for the latlciferous vessels of C. acantkaefolia and C. caulescens and the
transitional forms related to them, see above). 2. Subttibe Carduineae: in AI/redia
. solenopis! (? sphalm. ex ' stenolepis '; pith imd peticycle of the axis); Arctium
'lanuginosum, 'DC.' (pericycle of the axis); Berardia (axis); Carduus nutans. L.
and C. tenuz"fiorus, Curt. (pericycIe of the axis); Cirst'um arvense, Scop. (peri cycle
and pith of the upper parts of the axis and the leaves), C. eriophorum, Scop., C.
lanceolatum, Scop., C. monspessulanum, All., C. oleraceum, C. palustre, Scop. and
C. rj'vulare. Link; Cous1.·nia Hystrl,x, C. A. Mey. (axis) ;, Galactites totnentosa (peri-
cycle and pith of the axis); furinea alata (axis); Lappa minor, DC. (pericycle
and pith of the axis), L. major; 'Onopordon Acanthium, L. (pericycle and pith of
the axis and leaf); Sitybum 1lfarianum,- Gaertn. (pericyc1e of the upper parts of
the axis, not in the leaf); Staehelina dubia, L. (pericycle of the veins of the leaf,
not in the stem); TVariont'a (axis).
Within the limits of the Tribe Cynaroideae secretory sacs have been
shown to be absent, in species of Cardopathium and Xeranthemum (Subtribe
Carli.-'1eae) and in species of Cm:cus (Chamaepeuce), Cynara and Saussurea
(Subtribe Carduineae), as well as in the Subtribes Echinopsideae (species of
Echinops) and Centaureeae (species of CardunceUus, Carthamus incl. Kentro-
phyllum;_Centaurea- and- Serratula).
Secretory sacs have not as yet been demonstrated in the root in any
member of the Cynaroideae, as will be seen by reference to the synopsis given
above; in certain species, moreover, they are confined to the upper regions
of the shoot, viz. to the upper part of the axis and the upper leaves. The
feature last mentioned of course very considerably restricts the value of the
secretory sacs_for practical systematic purposes; its explanation is to be found
in the fact that in the more closely investigated Cynaroideae (belonging to the
genera Carduus, Carlina, Cirsium, Lappa, Onopordon and Silybum), in which

1 It is very much to be desired that a more detailed invesiigation of the secretory organs in these
Tribes and Subtribes ShOllld be undertaken on material from numerous genera.
 ADDENDA-COMPOSITAE

the secretory sacs are accompanied by secretory canals, the 'two types of
secretory organs are vicarious in their occurrence, inasmuch as the secretory
sacs more or less replace the canals as one advances from the base towards --the
apex of the plant.
Thus in Carlina vulgaris, for example, the root contains endodermal resin-canals.
which even penetrate into the lowest portions of the axis; higher up in the stem
they are replaced by secretory cavities and ultimately by latex-cells aituated in
the pericycle. The rhizome of Cirsium arvense has endodermal and medullary
resin-canals. which extend into the lower portion of the subaerial axis, but soon
exhibit a decrease in number and at a higher level become crowded out by
medullary and pericyclic secretory sacs. A similar relation, although it is not
so marked, obtains between the secretory canals and the laticiferous elements
in Gazania and Vernonia; this finds its expression in the fact that .endodermal
resin-canals occur only in the root (here side by side with laticiferous elements) and
not in the stem, which is rich in latex.
To complete our synopsis of the secretory organs found in the ,Cynaroideae
it remains to mention that those members of the Tribe 'Which have no secretory
sacs either possess secretory canals or have no secretory organs whatever. Secreto;ry
canals alone, not a.ccompanied by laticiferous sacs, have been observed in: Ecln-
nops lSubtribe Echinopsideae, here confined to the root), Cnicus, Cynaya and
Saussuyea (Subtribe Carduineae), and the investigated Centaureeae (see above);
neither secretory canals nor secretory sacs are present in species of Cafdopat£um
and Xef'anthemum (Subtribe Carlineae).
In concluding the discussion of the secretory organs we have still to refer
to the occurrence of ordinary cells of which the contents are resinous or
resemble latex, although the cells are not differentiated as idioblasts. In
some cases such cells appear to occur in ,place of secretory canals or latex-sacs.
To this category belong the elements found in Eckinops exaltatus and Tagetes
patula and described as resin-cells on p. 462. as well as the oil-cC!tfaining
endodermis of Chaptalia tomentosa referred to on the same rage. Such cells are
also found in the following species: Bal'nadesia 'Yosea (bast 0 the root); Cous2'nia
Hyst1'ix (medullary rays of the wood and bast of the rhizome); Eurybia afgophylla,
Cass. (medullary and cortical parenchyma of the axis and petiole); StaeheUna
dubia. L. (in the bast (here tubular). and in the pericyc1e of the axis); Tarcho-
nanthus camphoyatus, L. (in the bast of the axis and leaf).
Among other forms of excretion of oxalate of lime (see p. 464) clustered
crystals, some of which were of large size, were found by G. Fischer in numerous
members of the Vemonieae and Eupatorieae (species of Bolanosa, Eremanthus,
L)lchnophora, Pacourina, Vernania, etc.), and by Heering in species of Baccharis.
Bundles of small acicular crystals occasionally (Cirsium a1vense) occur also
in the epidermis of the leaf. The small crystals covering the leaves in Liatris
odoratissima consist of cumarin.
To the statements on the structure of the leaf (p. 464) we may add th.
fonowing details. According to Greenman, the upper epidermis in Senecio
chalapensis, Wats. va-;-. areolatus, Greenm. is papillose. According to Grimm,
Vernonia Luschnathiana has a two-layered hypoderm on the upper side of the
leaf. The vascular system in the veins of the leaf frequently possesses a distinct
parenchyma~sheath. For the occurrence of water-pores in the Compositae,
see Spanjer, loco cit.
New details as to the structure of the ground tissue and fibrovascular
system in the stern of the herbaceous members of the Order are to be found
in the papers of Keseling on Achillea Sect. Ptarmica, of Peter on SCOfZ01tera.,
of Kruger on the Ckhoriaceae, and of P. Wagner on Senecio. The structural
variations, which can be employed for specific diagnosis, refer especially to
the development of collenchyma and hard bast~ the nature of the endodermis
and of the tissue composing the medullary rays, and so on. According to
 ADDENDA-COMPOSITAE
•
Mobius a feature deserving special mention is the peculiar occurrence of
assimilatory tissue in the primary cortex of X anthium strumarium; this tissue
is confined to short strips on the stem. the epidermis above these strips alone
containing stomata.
Septation of the pIth has been observed by Harshberger and Greenman
in the xerophilous species, Senecio praecox, DC., while in other species of
Senecio the central portion of the pith commonly undergoes resorption. The
dividing septa in S. praecox are formed by cells which store up water.
We will now tum our attention to the anomalous structure presented
by the axis in many Cichoriaceae and certain representatives of other tribes,
the previous· description of w!llch was comprised under the heading of
'medullary bundles' (phloem- and vascular bundles, see p. 467). We may
first mention that recent investigations dealing 'with these anomalies have
been published by A. Peter (on Scorzonera) , Kriiger (on the Cichoriaceae),
and Col (on the Tubuliflorae). In addition to the modifications previously
described, a further one has been observed in certain species of the genus
Scorzonera; in this case medullary vascular bundles do not stand out plainly
in the transverse section of the stem, all the bundles being arranged in two
or m@re not very distinct rings; a similar structure is met with also in certain
species of Centaurea, e. g. C. acaulis and C. alata (see under' cortical bundl~:
p. 468). According to Peter and Kruger some of the manifold variations,
found even within the limits of one and the same genus in the Cichoriaceae,
are of value in specific diagnosis; in illustration of this statement the synops~
of these characters in the genus Scorzonera may be quoted from Peter's
work.

The species of Scorzonera may be classified in the following four groups:

I. Vascular bundles collateral with normal orientation (i. e. wood on the inner and
bast on the outer side) and arranged in several indistinct rings, as seen in the trans-
verse section of the stem: S. eriosperma, M. Bieb., S. hirsuta, L., S. ensilolia, M. Bieb.,
S. nervosa, Trev., S. cretica, Willd., S. tomentosa, L. II. Vascular bundles of unequal
~ize, co~ateral, an~ ,normally orientated, sho~ng a rather distinct ,arrjtngement
10 two nngs: S. Y'lgtda, Auch., S. pygmaea, Slbth., S. subaphylla, BOlSS., S. ramO'-
sissima, DC., S. cinere.a, Boiss. III. Vascular bundles in a simple ring with normal
orientation, the pith containing scattered strands of soft bast with or without
a rudimentary xylem-mass, the medullary bundles in the former case being in-
·versely orientated; in this group the peripheral vascular bundles are either (a)
bicollateral: S. latitolia, Vis., S. mollis, M~ Bieb., S. elata, Boiss., S. hispanica,
L.,. S. papposa, DC., S. incisa, DC., S. limnopMla, Boiss.; or (6) collateral: S.
aristata, Ram. IV. Vascular bnndles in a simple ring 'with normal orientation,
medullary strands absent; the vascular bundles being either (a) bicolla.teral':
S. st'Yicta, Homem.• S. inaequiscapa, Boiss:, S. croct/oh'a, Sibth., S. macfo~ePhala,
DC.; or (b) collateral: S. lanata, M. Bleb., S. tuberosa, Pall., S. hum'l12s, L.,
S. parvilotia, Jacq., S. se'Yicea, Auch., S. viltosa, Scop., S. austriaca, Willd., S. pur-
purea, L., S .. ciliC£ca, Boiss., S. criophora, DC.
-To the Jist_ (p. "467) ,of Cichoriaceous genera, in which some of the species at
least have medullary bundles (in the broader sense) in the stem, we may add
Hymenonema; among the genera of Cichoriaceae in which medullary bundles
are wanting we may include: Aposen's, Haenseleria, Hispidella, Lagoseris, Lygo-
desmt'a, Metabasis, Notobasis, Podospermum, Richardia, Sonchus, Tragopogon and
Urospermum.
Outside the Cichoriaceae (see p. 468) medullary vascular bundles have been
recorded by Col in Cardopatium corymbosum and Kent-rophyllum lanatum. Accord-
ing to the same author the leaf-trace bundles in many Tubuliflorae exhibit strands
of soft bast on their inner side prior to entering the ring of bundles, these strands
occasionally joining up with the outer phloem in an annular manner; in some
cases these intraxylary strands of soft bast continue their course on the inner side
of the ring of bundles for some little distance (Balduina multiflora) or they may
persist a1 together (A ctt'nome"is altern i folia).
SOLEREDER 3Q
 ADDENDA-COJrfPOSITAE

To the category of cortical vascular bundles we may refer the bundles

situated in the peri cycle in Achillea filipendulina (according to Vuillemin),
l\ladia sativa, Atractylis cancellata and species of A nthemis (according to Col),
and the vascular bundles occasionally occurring in the cortex of llelenium
autumnale (consisting of one. large bundle with smaller ones arranged in a
ring; likewise according to Col). We may also mention here the anomalous
structure presented by the stem of Cynara Cardunculus, L., and C. Scoly-
mus, L. The transverse section shows irregularly arranged vascular bundles
which are of unequal size and occasionally lie together in groups of two or
three; the bundles in these groups may have either the xylem- or phloem-
portions directed towards one another. Cortica.1 vascular bundles, lastly, are
found not only in Senecio Doria (see the earlier part of this work), but also in
S. coriaceus, Ait., and S. macrophyllus, M. Bieb., both belonging to the same
subsection Sarracenici (P. Wagner).
Note (d. p. 468). New examples of anomalous structure in the root have
been observed in Thrincia tuberosa. DC. by Maige. Gatin, and Carano, and in
At"a;tylis gummifera, L. by Kerckhoff. In the swollen parts of the adventitious
roo'b;, arising' from the base of the stem in .Thrincia tuberosa, the primary xylem-
groups of the vascular system become surrounded by a meristematic ring, which
is derived from the normal cambium and produces weod internally and bast exter-
nally. In this way the primarv xylem-groups become transformed into steles.-
The anomaly shown by the tap~root of A tracty/is guml?Ji/era consists in the occur-
rence of successive, concentric rings of. vascula.r bundles in the wood. The bundles
of the first ring show inverse orientation (the xylem being on the outside), those
of the second ring normal orientation, those of the third again inverse orientation,
and 50 on. In this way Kerckhofi was able to distinguish five rings of bundles in
a root of some thickness. At certain points groups of vascular bundles belonging
to the two inner rings unite to form actual .ste1es. The anomalous structure just
described can also be deteCted in an early stage of development. in axial parts of
the root-stock.
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Tr~pogon pom".folius) Comptes rend us Paris, lxxxviii, 1879, pp. 269-72 j . [also M em. Acad. d. sc.,
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 ADDENDA-COMPOS1TAE
..
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I"
(Kleinia):
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1904, pp. 566-70 and)'fab. xxv.-Siissenguth, Behaarungsverh. d. Wiirzb. Mtlschelkalkpft., Diss.,
Wiirzburg, 1904. pp. 37-43.-Theorin, Vaxttrichom., Arkiv for Bot., iii, n. 5, 1904, pp. 8, 12 and
15.-Areschong, Trop. vaxt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. :I, 1905, pp. 139, 140
(Vernon;a).-Kniep; Milchsaft, Flora, 1905, pp. 171-6.-Mayus, Milchrohr. in den Bl., Beih. z. bot.
Centralbl., xviii, 1. Abt., 1905, pp. 28:il- 5.-Sarton, Anat. d. pI. affines, Anc; sc. na.t., ser. 9, t. ii, J 90 5,
pp. 94. 95 and 105, 106 (Adu'//ea, Toraxacum).-Theotin, Vaxttricb.om., Arkiv for Bot., iv, n. 18,
J905. pp. 3, 5, 9. 10 and II.-Weberbauer, Vegetat. d. Hochanden Perns, in Engler, HoC. Jahrb.,
xxxvii, 1905, p. 60 ct seq.-[Altan, Sweda jacobaea, Phllrm. Post, 1906, p. 48S.]-Boodle, Lignifica-
tion of phloem in lIe/ialttkus, Ann. of bot., xx, l!)OO, pp. ;:I19-21.-[Brezina., Beitr. z. Anat. d. Holzes
der Composit., Sitz. Ber. Wiener Akad., 1906, 19 pp., 3 Tab.]-Dauphine, Rhizomes, Ann. &C. nat.,
ser. ?, t. iii, J906, pp. 327,349 aad 352 et seq.-[For additionalliteratnre, see p. 1170 .)

CANDOLLEACEAE (STYLIDIACEAE) (pp. 469-471).

Burns's recent investigations on the structure of the axis and leaf in the
genus Stylidium form a valuable extension of our previous knowledge.
The anomalous structure -of the stem in Stylidium adnatum, R. Br., &c.
does not, however, appear to be quite fully explained even by Burns's. work.
According to }ljm the broad ring of sclerenchymatous fibres -(some of which
bear bordered pits in certain species) does not arise from an actual secondary
meristem, but is merely a pericyclic strengthening ring, which ·adjoi~ the,
fibrovascular system on its outer side. The groups of soft bast ·and veJ!:iels,
which. are found enclosed in the sclerenchymatous ring in old vegetative stems
of S. adnatum and S. ja scicula tum, R. Br., are stated by Burns to belong to
the leaf-trace bundles. According to the same authority the vascular bundlas
in the vegetative axes and peduncles are occas~onally arranged in two rings;
in other cases they. are more-irregularly distributed, much as in a MonoQ)tyle-
donous stem;- Their structure shows no uniformity in the relative positions of
wood and bast; one meets with collateral vascular bundles accompanied by
transitions to concentric bundles with central phloem; vessels may occasionally
be observed in the phlOem and now and then (e.g. in S. calcaratum, R. BL)
isolated vessels occur also in the pith. The strengthening ring above men-
tioned is 5 to 20 cells in· thickness. In those species which have reduc.ed leaves
palisade.parenchyma composed of long cells is found in the stem.
Regarding the structure of the .leaf in the species of Stylidium we may
mention the following details. The epidermis always consists of a single layer
of cells, although in many cases it appears to be composed of two or more layers.
The apparent presence of several layers is due to the fact -that the epidermal
3Q2
 ADDENDA--CANDOLLEACEAE
cells (as is evident in longitudinal sections~ are tqil ,and are placed
obliquely to the surface of the leaf; two or more epidermal cells are conse':'
quently cut through one afte!' the other by a tra.nsverse ~ction and -see'm;
to lie above one another. This phenomenon is copfleCbtd with the prolonged
apical growth of the young leaves. The shape of t:'e imbricating cells varies
fr-om rhombic (e. g. S. iunceum, R. Br.) ,to fibrous (5. eriopodum, DC., S. pilosum,
Lapill., etc.); their lumina may be rather wide (e. g. S. saxi/ragoides, Lind!.) or
narrow, the cellulose-wall in the latter case exhibiting a correspondingly
strong thickening (e.g. S. eriopodum, DC.). Owing to uneqmil growth in
length the ends of the epidermal cells in some cases appear as uneven pro-
minences on the surface (5. scariosum, DC.) or margin (5. gramini/olium, Sw.)
of the leaf. A typically one-layered and an apparently many-layered epidermis
are found in species in which petiole and lamina of the leaf are differen-
tiated (5. calcaratum, R. Br. and S. petiolare, Sond. with a one-layered
epidermis; S. pilosum, Labill., S. reduplicatum, R. Br. and S. saxijragoides,
Lindl. with an apparently many-layered epidermis), as well as in species,
which do not show such a differentiation (5.1 scandens, R. Br., with a one-
layered epidermis; S. eglandulosum, F. v. M., with an apparently many-
layered episiermis). In certain species (e. g. 5. streptocarpum, Sond.) the
epidermal cells have a jagged outline in surface yiew. the apices of the_angles
in the notches being thickened. In some cases (specially distinct in 5. strepto-
carpum) the lateral walls of the epidermal cells bear true bordered pits side
by side with simple pits. The stomata show differences of structure
within the gen~ 5tylidium. In certain cases (as in S. eriopod'um, DC.) the
mother-cell of the guard-cells is formed by the first division in the dermatogen-
cell. In most of the species (e.g. in 5. saxijragoides, Lind!.) the development
of the stomata takes place according to'the Rubiaceous type, which is, how-
ever, occasionally somewhat obscured in the mature leaf. The distribution
of the sto~ata on the surface of the leaf varies; for the most part they are found
on both sides of the leaves, but in certain species they are confined to one side,
which may be either the morphologically lower or the morphologically upper
side, the latter under these circumstances being physiologically the lower side
owing to twisting of the leaf; .the latter case is found in the reduplicate leaves
of S. pilesu"" Lahill. and S. reduplicatum, R. Br. In those species in which
the apparently many-layered epidermis has a mechanical function owing to
the sclerosis of its cells, the stomata are not equally distributed over _the
surface of the leaf, but occupy longitudinal band-shaped zones consi~ting of
a single row _of cells; thus, two such zones are found on either side of the
leaf in 5. eglandulosum, F. v. M. and 5. saxijragoides, Lindl.. while in S. pilos'Um
and S. reduplicatum there are only two zones, which are confined to the
morphologically upper side of the leaf. As a general rule the stomata
ar) arranged with their pores directed parallel to one another and to the
long axis of the leaf. The mesophyll is either centric with palisade tissue on
all sides (e.g. in the acicular leaves) or more or less distinctly bifacial or almost
homogeneous. Arm-palisade parenchyma, occupying either a vertical (5.
calcaratum) or a horizontal (5. eriopodum) position, is occasionally distinctly
differentiated. In S. pilosum the walls of the assimilatory cells are thickened
and pitted, much as in the Cycads, and the same sp~cies has spicular cells,
which function as a support to the epidermis. In the long-stalked leaves the
veins show an abundant development of mechanical tissue around the vascular
bundles, while in those leaves, in which the integumental tissue is sclerenchy-
matous~ there is a reduction of the mechanical tissue in the veins.
Bums observed oxalate of lime in the form of clustered crystals In the
ground tissue of the stem in certain species of 5tylidium (e. g. S. eriopodum).
The hairy covering in the genus Stylidium comprises_ glandular hairs (on the
 ADDENDA--CANDOLLEACEAE
axis oi infiorcscenctJ and on the leaves) and mucilage-hairs {at the growing-
points and in t~ axils of the Jeaves}. The gJandular hairs vary very much in
shape. The sman glands of S. calcaratum, R. Br., have a biseriate stalk and
a bicellular head divided :by a vertical wall; with them we may class the
external glands found in S. amoenum, R. Br.~ S. diversifolium, R. Br. and
S. redupticatum, R. Br., in which the stalks are longer, though likewise hi-
seriate, and the heads are divided by numerous vertical walls into a large
number of,approximately prismatic cells, which show a vertical arrangement.
The glandular hairs of S. pilosum, Labill. and S. lineatum, Sw. exhibit a dif-
ferent type of structure; the first of these species has long club~shaped external
glands provided with a biseriate staJk, which gradually merges into the head,
the latter being divided by walls showing varied O1'ientation; S. lineiztum
has glandular hairs with a long biseriate stalk and a spherical head, which_is
likewise divided by walls running in va!'ious directions. In the mucilage-hairs
the structure of the head resembles that of the external glands found in
). amoenum, etc., but the staJk is uniseriate. In the simplest case (S. a:dnatum,
R. Br.) the head of these mucilage-hairs consists of a varying number of rod-
shaped cells, which are developed by a series of vertical divisions in the terminal
cell of the young hair. These cells become detached from one another, while
the cuticle is raised in a vesicular manner. With the bursting of the cuticle
and the emission of the mucilaginous secretion the function of the- mucilage-
hairs in S. adnatum comes to an end. Other species, howeyer, show certain
points of difference from the case just described. The rod-shaped cells
in the glandular heads of S. pilosum, Labill. first undergo transverse division
into two cells; this leads to the bursting of the cuticle, remains of which
persist at tl;le base of the head. Thereupon the :rays of the head (which are now
bicellular) become enveloped by a new cuticle. In other species (e.g. S. ~axi-
fragoides) the transverse division of the cells and the formation of a new cuticle
is repeated several times, so that a third or even fourth cuticle may be excreted,
the remains of the old cuticles being found at the base of the individual cells
in the rays of the hair.
Literature: Raunkjrer, Cellekjxrne-Krystalloider, Bot. Tidsskrift, xvi, 1887, pp. 41-5.-
Leisering, ImerxyHires Leptom, Diss., Berlin, 1899, p. 46.-Burns, Beitr. z. Kenntnis d. Stylidiaceen,
Flora, 1900, pp. 313-54 and Tab. xiii. xiv; also Diss., Munich.

GOODENIACEAE '(pp. 471-473).

Literature: Leisering, Interxyl. Leptom, Diss., Berlin, 1899, pp. 26, ~ 7.-[Colozza, Studio anat.
delle Gooden., Nuovo Giorn. bot. Ital., N.S., xv, 3, '907, pp. 304-26; see also the same author,
Bnmonia, loco cit., pp. 296-303.]

CAMPANULACEAE (INCL. LOBELL.~CEAE) (pp. 473-476).

The distribution of the laticiferous vessels in the leaves of the Cam-
panu1oideae,~ Engler and Prantl has recently been determined from the
systematic point of view, especially by H. Schmidt in connexion with his in-
vestigations 1 on the structure of the leaf in the group referred to. He met with
laticiferous vessels in all the genera available for investigation, with the
single exception of Sphenoclea-a genus which also differs from the rest of
the Campanuloideae in the possession of large clustered crystals Of o_xalate of
lime, and constitutes a special group (the Campanuloideae-Sphenocleae) in
Schonland's system; Ostrowskia likewise has laticiferous vessels, as I have
found by personal investigation. The correctness of Poulsen's statel1Jf"nt as

I They deal with all the genera excepting Ostrowskia.

 ADDENDA--CAMPANULACEAE
to the absence of latex in Pentaphragma ellipticum, Poulsen, is open to doubt,
since H. Schmidt demonstrated laticiferous vessels in the l~ger veins of the
leaf in P. begoniaetolium, Wall. ' The laticiferous tubes of the Campanuloideae
are found in the bast both in the large and small.,.eins, and frequently show
fine examples of anastomosis. H. Schntidt observed penetration of the latici-
ferous tubes into the mesopbyll, only in CanaTina, where Trecul had previously
demonstrated the same ehenomenon; so that the occurrence of these elements
in the mesorhyll is certainly very rare in the Campanuloideae.
Y drac has recently published observations on the latic:iferous system of
the LobeUaceae. He records laticiferous tubes in species of Centropogon,
Clermontia, Cyanea, Delissea, Downingia, Haynaldia, Heterotoma, Isotoma,
LaUrentia, Lobelia,2 Pratia, Sclerotheca, and Siphocampylus. The principal
system of anastomosing laticiferous vessels in these genera traverses the bast
of the vascular bundles both in the stem and leaf. This principal system
undergoes ramification, and from the branches of the first order Crameaux
laticifu'es ') thus formed further branches (' branches laticiferes ') arise, which do
not anastomose and are not developed by a process of fusion; these branches
are found in the ground tissue (pith and primary cortex), pericycle and wood
of the stem, as well as in the mesophyll.
The following details regarding the structure of the leaf in the Campanu-
loideae are based on H. Schmidt's and Feitel's investigations. The epidermal
cells as a rule are of medium size and relatively low; their lateral walls
may be straight or strongly undulate~ while in some cases (e. g. in Light/ootia
juncea, Sond., Wahlenbergia Tabusta, Sond.) the outer walls are considerably
thickened. The ..cuticle is for the most part thin, its surface being smooth or,
striate~ or more or less coarsely granular. In certain xerophilous species,
especially those belonging to genera endemic in the region of the Cape
(e. g. M erciera, M icrocodrm, RoiUa, Wahlenbergia), the upper epidermis in the
middle of the surface of the leaf consists of cells with wide lumina and straight
lateral walls, while towards the margin of the leaf the cells show a progressive
decrease in size and have more or less undulated lateral walls. H. S~hmidt
records nodose thickenings at the points of intersection of the. lateral walls in
Pentaphragma begoniaejolium, Wall. and Trachelium rumelianum, Hampe.
Papillose differentiation of the epidermal cells is of frequent occurrence, but
according to H. Schmidt it always only affects isolated cells OT (near the
margin of the leaf) groups of cells; a typical papillose epidermis 'was not
observed in any species. The pa:r.i11ae are conical or hemispherical and in
some cases are reduced to solid silicified knobs. A point deserving special
mention is that gelatinized epidermal cells were not observed in any member
of the Campanuloideae. The stomata as a general rule are not provided with
special subsidiary cells.. The only exception is. furnished by Pentaphragma
(P. begoniaefolium)~ which is distinguished also by the nature of its hairy
covering (see below) and constitutes the group Campanuloideae·Penta~_
phragmeae in Schijnland~s system; in this genus there are three narrow
subsidiary cells. In the Cainpanulaceae investigated by Vesque, moreover,
the mother-cell. of the guard-cells is likewise not formed by the first division,
but only appears after a few epidermal cells have been cut off from the der-
matogen-cell. The stomata are found either on both sides of the leaf or only

1 Vdrac also enmined. the s.tructure or the root, uem aDd leaf. Hi.s inves.tigations. deal with the
. genera above enum~rated, in which he stadied the latici{eroa. system, as' well as witb the geOett
AP4/4JU'a. Bn"ghami'a, and R/u'z«ephalum (all the genera being taken in the sense of Darand's
Index).
y.
• Ydrac's statement (Ioc. cit., 106) that laticiferou5 tubes are absent in LDlJelia iJorlmonna,
L. is incorrect, since the Dle9l)phyl in this species is traversed by a very well developed net\\'ork of
latic:iferous tubes.
 ADDENDA-CAMPANULACEAE 961
on the lower side; according to Wettstein H edraeamhus Pumilio, Porta is
exceptional in having the stomata confined to the upper side. The stomata
generally lie on a level with the epidermis or are slightly raised or (rarely.
e. g. in Sphenoclea) depressed. In the species provided with narrow leaves
the pores of the stomata are placed parallel to the midrib. Water~pores are
present in the majority of the species, but are mostly of no great size; varying
numbers (for the most part 2-6) of these pores are associated with undulated
epidermal c~ to form groups, which are invariably found on the upper
surface near the margin of the leaf and are occasionally situated on special
prominences of the leaf~surface or on the.leaf·teeth. The mesophyU may be
centric, bifacial or hom~geneous. A specially noteworthy feature, not previously
recorded in this Order, is the occurrence of arm·palisade parenchyma, in
species of Adenop~ra, Campanula, Campanumaea, Canarina, Cephalostigtna,
Codonopsis, Heterocodon, jasione, Leptocodon, Musschia, Ostrowskia (according
to my own investigation), Pentaphragma (P. ellipticum, according to Poulsen),
Peracarpa, Platycadan, Prismatoc,arpus, Symphyandra and Wahlenbergia.
Peculiar spongy tissue formed by rows of hypha~like cells and traversed by
large intercellular spaces is found in Light/oatia ciliata, Sond., L. lasciculata,
DC., L. rubioides, DC., &c. In WaJde{'bergia pro$tra~a, DC. the transverse
section of the leaf, which is roughly four·sided, shows an extensive semilunar
complex of aqueous cells with wide lumIna, beneath the upper epIdermis;
below this hypoderm one first meets 'Yith spongy tissue and then with.·a single
layer of palisade tissue. A similar hypodermal layer of aqueous tissue, which,
however, in this case still contains chlorophyll, is found on the upper side of
the leaf also in W. capiUacea, DC. Other xerophilous members of the Order
ha~e a more or less strongly developed hypoderm composed of sclerenchy·
matous fibres, e. g.. in certain species of Cephalostigma, Light/ootia, M erciera,
Microcodon, Prismatocarpus, Roella. and Wahlenbergia. This hypodermal
sclerenchyma either merely forms crescent~shaped masses at the margin ,of
the leaf, or extends somewhat onto the upper surface, where it ultimately gives
rise to a continuous band of sclerenchyma. In some cases it is also found-
on the 10~er•. side pf.,the leaf_ p~.!l~ath the vascular system of the midrib, but
separated fr9m the-latter by assimilatory tissue. Hard bast is not found
accompanying the vascular bundles in the veins of the leaf in any member of
the Campanuloideae. In its .place the xylem~groups themselves occasionally
include sclerenchymatous fibres, e. g. in species of Lightfootia, MiC1'ocodon,
Rhigiophyllum ~Tld Wahlenbergia. In some cases the median vein is enveloped
by a distinct parenchyma-sheath (e. g. in Rhigiophyllum squarrosum, Rochst.).
Oxalate of lime, which prior to this had been observed only in Hed1'aean~
thus 1 by Wettstein, is found especially in the form of fine Clustered crystals
in Sphenocl~. H. Schmidt also met with small prismatic or acicular crystals
of the same salt in the mesophyll of certain species of Canarina, Heterochaenia,
Leptocodon and Phyteuma; crystals of this type are ,no do~ more widely
distributed, but are easily overlooked. Sphaerocrystals of unknown chemical
composition have been demonstrated also in species of Adenophora, Canarina,
M usschia, Phyteuma, Specularia, Symphyandra an~ Trachelium.
Schmidt also failed to find any glandular hairs in the hairy covering.
On the other hand, in Campanumaea celebica, Bl., he met with long coni~
clothing hairs composed of a single row of 4-7 cells with thin walls and
wide lumina, while fu Pentaphragma begoniaefolium, Wall., he observed multi~
cellular clothing hairs with two_or more arms; the latter consist. of a short stalk,

1 Wettstein Ooc. ciL, p. 187) refers to the rare occnrrence of crystals j he records clustered
cryst:lll in the stem of H. str6icur, Kern., MU ' raphides' (probably not true rapbides, see above)
in tbe ant ben of H~ ru",Uio. Porta.
 A DDENDA-CA MPANULA CEAE

one or more cells in"""_breadth, and uniseriate arms, which are placed more
or less parallel to the, surface of the leaf and are composed of rather short
cells, with moderately"thick walls and wide lumina. For the rest, how-
ever, only unicellular dothing hairs have been found; these vary in length,
in the thickness of their walls and in the width of their lumina, and
have a smooth, striate, granular or verrucose surface. In most cases these
hairs are pointed, rarely blunt (elongated and finger-shaped in Codonopsis;
provided with a slightly swollen capitate end in P.hyteuma pinnatum, L.).
In the species of C(Jmpanula the walls of the hairs are commonly calcified
and in some cases the neighbouring cells are raised to form a pedestal bearing
the. hair. The bases of the hairs, moreover, occasionally contain silicified
protuberances resembling cystoliths; in certain species the latter occur also
in the papillose and non-papillose epidermal cells' of the margin of the leaf
(being situated either in the corners or in the middle of the outer wall). or in
the subsidiary cells of the hairs (in the same way as in Lithospermum).
For the occurr~nce of steles in the petiole 10£ Campanula rapuncvloides,
see Bouygues. loco CIt.
With reference to the structure of the stem in the Campanuloideae we
may add that according to Pitard the pericycle contains a ring of fibres only
in Platycodon; at later stages stone-cells become intercalated in this ring.
The following account of the structure of the leaf in the Lobeliaceae is
based on the results of Ydrac's work. The stomata are either confined to the
lower side Of occur on both sides; they have no subsidiary cells. Water-pores
are present on the leaf-teeth. The mesophyll is bifacial Of homogeneous.
Oxalate of lime has not been observed. The hairy covering consists only of
clothing hairs. which are generally unicellular (1 soioma, Lobelia, Siphocampylus)
or uniseriate (Pratia, Siphocampylus) and have scarcely thickened walls, which
frequently show a verrucose surface. A special form of hair is distinctive
of Siphocampylus Columnae, i.e. tufted hairs having eight or more rays and
seated on a multicellular stalk (see also Bentham and Hooker, Gen. Plant., ii,
P··548). -
According to Ydrac the structure-of the stem in the Lobeliaceae likewise
shows very few striking features. The cork, which is rarely formed, arises in
the outer parenchyma of the primary cortex (Clernumtia grand{jlora, Gaud.
and C. macrocarpa, Gaud.), or in some cases in the subepidermal layer of cells
(Apetahia raiateensis, Baill., Lobelia excelsa, Lesch., and Siphocampylus biser-
Tatus, A. DC.). The primary cortex is not uncommonly ccllenchymatous,
while in Downingia elegans, TOTr. and Lobelia Dortmanna, L. it contains'
large lacunae. In most cases the endodermis is distinctly developed and
provided with Caspary's dots. The pericycle includes sclerenchyma (with
wide lumina) only in Lobelia Dortmanna. There is no sclerenchyma in the soft
bast. In Pratia montana, Hassk. and Isolobus radicans, A. DC. the fibro-
vascular system consists of two arcs of wood and bast. The cel1s of the pith
mayor may not be lignified; in some cases the central portion of the pith
becomes resorbed and gives way to a lacuna (Lobelia guadalupensis, Urb., etc.).
According, to Col medullary phloem- and vascular bundles occur also at
some points in the stem of Campanula rapunculoides, and are present in the
petioles and veins of the leaf not only in those members of the Order. which
have axes with anomalous structure, but also in many which are normal.
In the members of the Lobeliaceae, investigated by Ydrac, these bundles are
absent. In the wood of the root of Campanula pyramidalis J. E. Weiss
observed concentric vascular bundles, which form a continuation of the
medullary bundles of the stem.
Literature: Costantin, Tiges aero et sout., Ann. $C. nat., St:f. t$, t. xvi, 1883, p. 161 et ~.­
Boergesen, Arkt. pI. bladbygn., Bot. Tidsskrift, xix, 1895, p. 219 et seq.-Spanjer, Wasserapparate,
 ADDENDA--CAMPANULACEAE
Bot. Zeit., 1898, i, p_ 55.-Minden, Wassersez. O~ane, Bibl. bot., Heft 46, 1899. pp. 23 a.nd 34
(Lobelia, Campanula).-Baranetzky, Faisc. bicoll., Ann. sc. nat., set. 8, t. xii, 1900, pp. "95-9.-
Feitel, Vergl. Anat. d. Laubbl. bei den Campanul. der Capflora, Bot. Centralbl., 1900, pp..., 41, 91,
129, and ~ 61 et $eq.; also Diss., Kie] ( WaJ,.lenbffgia, Lig-hifoolin, MicrocodPn, .Roj/la, PrismaltKarjus).
~Pitard, Pericycle, These, :Bordeaux, 1901, p. 49.-[&ar, Milchrohr., Sitz. Ber. Lotos f. BOhmen,
xxii, 1~2. n. 4, 5.]-Bouygues, Petiole, T!lese, Paris, 190", pr. 13 and 73.-Poul~n, Pentaphragma
eI#jJti~um, Vi~ensk. Med~ele~s. NatUJ:'~. Forening, Kj~benhavn, 1903, ~p. 319-30, pI. iv, v'41ven-,
Lol,elta Dqrtmann(J, ArklV for Bot., 1, 1903. p. 377 et seq.-Col, Fmscea\\x, Ann. se. m.t., ser. 8,
t. xx, 1904, pp. 19-97, :US and 2:;9-61.-Freidenieldt, Acat. Bau d. Wurzel, Blbl. bot., Heft 61,
19Q4, pp. 73, 74.-Paoli, Eterofillia, Nl.1ovo Giom. bot. ltal.,N. S., xi, 1904, p. l27.-H. Schmidt,
Syst.·anat. Untersuch. d. BI. der Campaooloideen, Diss., Erlangen, 190.40 103 pp.-Siissenguth.
Behucullgsverh. d. Wiirzb. MuS<.:he1kalkpfl., Diss., Wiirzbllcg, 1904, p. 43 ......:.::.HaberIandt, Lichtsinaes·
org., 1905, p. 70, Tab. I.-Kniep, MHcbrohr., Flora, 19°5, pp. 169-71.-MttyuS, MilchrOhr., Beih. z.
bot. Centralbl., xviii, Abt. I, 1905, pp. 281, 28z.-Theorin, Vaxttrichom., Arkiv.fOr Bot., iv, n.18,
1905, p. lI.-Ydrac, Appareillaticifere des Lob€liacees, Jouro. de bot., '905, pp. I :2-l0; also in
Penot, Travaux, ii.-Ydrac, Hech. anat. sur les Lobcliacees, 1905, in Perrot, Travaux, iii, 19()6,
165 pp. j also. These, P.aris,--HQ]termann, Ein/hlS$ des Klimas, etc., 19°7, Tab. ix, Fig. 45.-(For
further literature, see p. 1I6_?]

VACCINIACEAE (pp. 476-479).

According to Poulsen extrafloral nectaries, appearing as disc-shap~d
structures with a raised rim, are found at the base of the lamina o{ the leaf in
Vaccinium Teysmanni, Miq. They are provided With an irregular epidermis,
below which lies a spherical complex of small-celled tissue jn which the nectar
is secreted, while the inner part of this tissue is enclasped by.the ramifications
of a vascular bundle which branches off from the petiolar system.
Literature: Latour, Sene, These) Mcntpellier, 1894, p. 40 et seq.-Boer~esen, Arkt. pI. bladbygn.,
Bot. Tidsskrift, xix, 1895, p. 219 et seq.-Po1l1sen, Extraflorale Nektaner, Vidensk. Meddelel&.,
Naturh. For. Kjllbenhavn, 1897, pp. J6S-S.-Roedler, Assimilator. GewebesyGt., Diss' t Freiburg i.
d. Schw., !898-9, p. 37.-Petersen, Vedanatomi, 1901, pp. 81-3 (Vaccinium, Oxy,occol).-[For
further literature, see p. 1172.]

ERICACEAE (pp. 479-488).

2. STRUCTURE OF THE LEAF. The most important recent work is Rom-
mel's investigation of the structnre of the leaf and stem in the Pyrol~Y'Qla,
Moneses and CJ»maphila) and in Clam-a, and Linsbauer's paper on ehilled
structure -and course of development of the pocket-shaped leaves of Cassiof>e
tetrago-na (d. pp. 48 2, 483). ./
The structure of the leaves in the Pyroleae is either bifacial (Pyrola pro
parte, ChimaPhila) or homogeneous (Pyrola pro parte, Moneses) , while in
Clethra it·is bifacial. Cletkra brasiliensis, Cham. et Schlecht. has a single layer
of hypoderm on the upper side of the leaf. Rommel's statement that the
,stomata in Ctetkra are invariably provided with two subsidiary cells, which
are placed parallel to the pore, is incorrect, as I have found by an investigation
of C. arborea, Ait. There is no sc1erenchyma in the veins -in the Pyroleae,
whereas _~ ....Cl!thra, tt
is fo_und ll_('.companying the vascular bundles. For the
occurrence of rolled leaves in die Ericaceae and their structure, see also Knob-
lauch, loco cit.
Oxalate of 1ime is present in the form of clustered crystals also in the
Pyroleae and in Clethra. For the oc('"llrrence of nuclear ("'rjstalloids in the
vegetative organs of Pyrola and ChinUlphila, see Raunkjrer, loco cit.
In connexion with the section on the hairy covering we may 15pecially
draw attention to Boergesen's paper, which also deals with the mode of develop-
ment of the hairs. Unicellular bristle-hairs are found inspedes of Cassiopc,
Erica, Kalmia, Ledurrt, Loiseleuria, M e_nziesia, Phyllodoce and Rr"ododendron,
while multicellular woolly hairs occur in· species of Azalea, Clethra, Eric;a,
Ledum, I.eucotlwif, Lyonia and Rhododendron. Long club-shaped glandular
970 ADDENDA-ERICACEAE-

hairs composed of two rows of cells are present also in Loiseleuria procumbens,
Desv. and PhyUodoce coerulea, I Gr. et Godr.'; sessile external glands varying
in shape from ellipsoidal to ovate and formed by a sjngle row of cells, in Cas·
stope tetragona, Don; glandular hairs with a multiseriate stalk and a spherical
head composed of numerous cells, in Epigaea repens, L.
3. STRUCTURE OF THE AXIS. Rommel's paper furnishes new data on the
structure of the axis in the Pyroleae and in Clethra, while Petersen has recently
examined the wood in a relatively large number of species.1
In the species recently investigated the wood likewise includes vessels
which have relatively small lumina and are for the most part provided with
scalariform perforations, while the ground-mass consists of wood-prosenchyma
with bordered pits (tracheids). Spiral thickening of the walls of the vessels
(at least of some of them) and tracheids is found in the species of Arctostaphylos,
Arbutus and Daboecia cited below, as well as in Chitnaphila umbellata, Nutt.2 ;
wood-fibres bearing simple pits have been observed in Moneses, Pyrola, and
Arbutus Unedo, L.
I

The perforations of the vessels are stated to be scalariform only in: Andro-
meda polifolia (with IQ-ZO bars, which are specially closely placed), Cassiope
tewagona, Cnamaedaphne calyculata (with delicate, closely placed bars), Chimaplu'la
umbellata (bars not numerous), Clethra (with numerous, bars), Ledum palust1'e (as
in Andromeda polilolia), Loiseleufi'ia procumbens (with numerous delicate bars),
Moneses grandiflora, Salish. (with numerous bars), Phytlodocc &oerulea (with lDany
bars). Pyrola (occasionally with 12-15 bars, which may anastomose in a reticulate
manner), Rhododendron lappon£cum (with as many as IO bars, but mostly less;
bars occasionally reticulate); simple perforations occur side by side with scala.ri.
form perforations having few bars in: Arctostaphylos uva ul'si (scalarifonn per-
forations with one or two bars), Calluna vulgaris (small scalariform perforations,
which may also have a reticulate structure); simple perforations only are' present
in: Ayb'iltus Unedo (elongated elliptical), Daboecia cantabrica, Erica Tetralix.
In some of the species of Pyrola the cortex includes a ring of pericyclic
scierenchyma, which occasionally gives rise to radial processes penetrating
between the vascular. bundles; in M oneses grandijlora and in Chimapkila, on
the other hand, there is no such sclerenchymatous ring.
Literature: [Pasc11kis, Pharmakogn. Beitr., Zeitschr. osterreich. Apothek.-Ver., I88tJ, n. 27, 28;
abstr. jn Bot. Centralbl., J881, i, p. 54.]-[Raunkjrer, Krystalloider, etc., Vidensk. Meddelels.
Naturh. For. Kjebenbavn, J882, p. 70; Ilbstr. in Bot. Centralbl., 1883, ii, p. 267.]-Boergellen, Nogle
Eric.-Haars Udvikl., Bot. Tidsskrift, xvii, 1890, pp. 307-1+.-Wijnaendts ,Francken, Sclereiden,
Disl., Utrecht, 1890, pp. 58, 59.-Boergesen, Arkt. pI. bladbygn., Bot. Tidsskrift, xix, 1895, p. :.319 et
leq.-[Andersen and Kallstroem., Folia nVIl ursi, etc., Nord. Farm. Tidsskr., 1896, p. 33.]-Knoblauch,
Okotog. Anat. etc., Habilitat.-Schr., Tiibingen, 1896, p. 5 et seq.-Schnbert, Pa.renchymscheiden,
Bot. Centralbl., 1897, iv, p. 19.-Ho1m, Pyro/a aphylla, Bot. Gazette, 1898, p. 2+9.-Rommel,
Anat. Untersllch. tiber d. Gr. d. Piroleae u. Clethra~ae, DiS!>., Heidelberg, 1~98, 53 pp. and I Tab.
-Kohne, Papill. u. oberseit. Spaltoffn., Mitteil. deutsch. dendrolog. Gesellsch., J899, p. 59 (P,apiltae
in species of GMdQt/endrou).-Linsbauer, VegetatioDsorg. von CQssjope le/ragDIta, Von, Sitz. Ber.
Wiener Akad., cix, Aht. i, 1900, IS pp., 2 Tab.-Paulesco, Struct. anat; des hybrides, These,
Geneve. 1900, p. i 3 (If Jwdodmdrtm).- Tunmann, Sekretdriisen, Diss., Bern, 1 goo, pp. 36-8.-P~ter~
sen, Vedanatomi, 1901, pp. 72-80.-Clauditz, Blattanat. C~Dar. Gew., Diss., Basel, 19°::1, PP.33 and
47 (Erica, .Arlutus).-Simon, Sommer~ u. wintergriine Gew., Her. deutsch. hot. Gesellsch., 1902,
pp. 239-40--Theorin, Vlxttrichom., Arkiv for Bot., i, I903, p. I 59.-[Gyorffi , Phys.-anat. Verh.
von RhtJt/odenlirull' myrttfo/itlm, etc., Diss., Koloszvar, J904 (HlUlgarian); abstr. in Just. 1~4, i,
P. 77 I .]-Andrews, Epiga.:a ,.ejms, Beih. z. bot. Centralbl., xix, i. Abt., 1905, pp. 31.4,-20.-

t viz.: Andromeda polifolla, L., Arbulus UnedD, L., Arc/()staphy/os lIva u,;s£, Spr., Cal/una
vulgaris, Salish., Cassiope Ie/ragona, Don, Chamatdaphlle calJ'culala. Moench, Clz.imaphila um6ellala,
Nutt., Dtzhoecia cantaf»>ica, C. Koch, Edca TeN-a/ix, L., Ledum paluslrl':, L., Loiseleuria frtJCUtll-
Oms, Des.v., PAyllodoce ctKrt4lea, Bab., RWociendro'tt lapponicum, Wahlenb. .
~ Romtnel's statement as to the absence of vessels in the seconClary wood of this species h in-
correct, and the same ilJlplie!\ t(l the record of the occurrence of n. ring of sclerenchyma, etc. See also
Peter~en, loc. cit. '
 A DDENDA-ERICACEAE 971
Kanngiesser, Calluna 'lIulffaris, in Tubeof, Natorwiss. Zeitschr., 1906, pp. S!i-60.-PicooJi,
Legnami, Bull. Siena, I9O(J, pp. 150 and 166.-Holtennann, Einfloss des Klimas, etc., 1907,
pp. 76 and 115 (Rhododtndron).-[Guttenberg, Immergr. Lanbb1. d. MediterranfJora, in Engier,
Bot. Jamb., xxxviii, 1907, p. 434 (Arbutus Unedo).]

MONOTROPEAE (pp. 489, 490).

Literature: MacDougal, Symbiotic saprophytism (Pterospora andronledea, Nutt.), Anb. of hot.,
xiii, 1899, pp; 31-8.-Thomas, Feuilles sout., These, Paris, ,190o.-Porsch, Spaltoffnungsapparat,
Jena, 1905, PP. 78- 80•

EPACRIDEAE (pp. 490-494).

1. REVIEW OF THE ANATOMICAL FEATURES. Bordered pits have also
been obser.red on the walls of the fibrous cells in the pericycle and secondary
hard bast (in the axis of many Epacrideae). The infernal development of the
cork (in the pericycle) constitutes an ordinal character. Oxalate of lime occurs
in the form of solitary as well as of clustered crystals.
2. STRUCTURE OF THE LEAF. For the structure of the epidermis, see also
Baccarini, loco cit., p. 81 et seq. and Tab. IV; this author deals especially
with the peculiar secondary thickenings of the cell-walls, which occur either
on all the walls, or only on the outer, or on the outer and lateral walls or on the
outer and inner walls; these thickenings are occasionally ~aversed by pits.
In the structure of the mesophyll we may specially note· that according to
Baccarini the palisade-cells show bellows-like foldings in very many species,
and that spicular fibres (' tracheidi ') extending up to the epidermis occur in
A ndersonia aristata and A. pamfolia. According to Baccarini the presence
of bordered pits on the fibrous cells ,forming the sclerenchymatous sheaths of
the veins is not a feature of such general distribution as was previously
maintained by Simon.
Baccarini's paper contains data for the occurrence of oxalate of lime in
the leaf, which very considerably add to our previous knowledge. Only solitary
crystals are found accompanying the vascular bundles of the veins; in those
species in· which the veins have strongly developed sclerenchymatous' sheaths
(Monotoca lineata, etc.) the crystals are found in the epidermis. The mesophyll
contains both solitary and clustered crystals. 'Macle scheggiate' (clustered
crystals ?) are recorded in Epacris heteronema and E. paucifl,ora, as well as in
Leucopogon /lavescens and L. gracillimus; short prisms are stated to occur
in Decaspora thymi/olia and Styphelia trijlora, octohedral crystals in Cyathodes
dealbata and Dracophyllum verticillatu,m, crossed twin-crystals in Andersonia
aristata, A. micrantha, Aerotriche cordata, A. serrulata, etc. In Prionotes there
is no oxalate of lime. - ,
, 3. STRUCTURE OF THE AXIS. New statements on the structure of the
axis have been published by Liiders and Baccarini. The former, on whose
investigatiofl:s the _follqwi!lg description is mainly based, examined species
of-all the tweiity-sl.X'genera enumerated in Durand's Index, as well as WooUsia
and Sphenotoma, which Drude regards as independent genera. No essentially
new features have, however, been discovered.
We will first consider the structure of the wood. The vessels have rela·
tively small lumina and are mostly arranged in distinct radial rows; a scattered
arrangement is rarer. In the different species the perforations of the vessels
are described as being exclusively simple or exclusively scalariform or both
simple and scalariform; it still remains to be investigated whether those
species which are stated to have simple perforations only, do not also possess
scalariform perforations in the primary wood or in the neighbourhood of
the latter. Anastomosis of the bars in the scalariform perforations frequently
 ADDENDA-EPACRIDEAE

leads to the production of reticulate or reticulate~scalariform types of perfora-

tion. The only additional species in which Liiders demonstrated the modified
scalariform perforations. figured for Epacris heteronema, Labill. in Fig. 112,
is E. lanugi,wsa, Labill., although Baccarini does not mention them (see also
Rodham, in Ber. deutsch. bot. Gesellsch. 1890, p. I90); in these perforations
the slits; which show a scalariform arrangement, are broken up into very small
pores by numerous delicate and filiiorm bars. Liiders observed spiral thicken-
ing of the walls of the vessels in Acrotriche aggregata, R. Br., Epacris paludosa,
R. Br., E. purpurascens. R. Br., Lysinerfl,a ciliatum, R. Br. and L, elegans, Sond.
Lfrders found only simple perforations in the vessels in species of A nderscmia,
A rcheria, A steroloma, Brachyloma, Coleanthera. Conoste!,hium, LeucopogQn, Lis-
sanelu, MeJichrus, Needhamia, Styphelia; for the most part simple perforations,
but accompanied by scalariform perforations with 2-5 bars or in some cases only
a single bar, in species of Acrot"iche, Andersonia, Conostephium, Cyathodes, Leuco-
pogon, Lissanthe, Lysinema, MelichYus (malformed perforations). Oligarrhena,
Seyphelia; mostly scalariform perforations in Monotoca (with 2-6 bars); only
scalariform perforations in species of Cosmelia (numerous bars), Cyathopsis (4-10
bars), Dracophyllum (15-20 or more bars), Epacris (mostly 4-15 bars, occaSionally
1-3), Lebeta,nthus (numerous bars)~ Pentachondra (IO-20 or more bars), Pn"onotes
(numerous bars), Richea (10-20 bars), Sphenotoma (10-15 or more bars), Sp)'engelia
(4-20 or more bars), Trochocarpa (2-12 bars), Woollsia (8-12 or more bars). See
also Baccarini's statements on this point, loco cit., p. 101.
The medullary rays are mostly ~me or two cells broad. Baccarini distinguishes
two types of medullary rays in the wood of the Epacrideae. In the first type
the medullary rays, which,are one or two cells in breadth and vary from one to
four cells in height, are composed of prosenchymatous cells (A ndeYsonia aristata,
A. prostrata, Brachyloma ericoides, Coleanthera myrtoides, Epacris impressa,
E. mucronulata, E. obtusifolia J E. pulchella, E. I splendens: Woollsia pungens);
in the second type which is found in the majority of the Epacrideae, the cells
of the medullary rays, as seen.in longitudinal section, are rectangular and elon~
gated in the vertical direction. The medullary rays of the second type occa-
sionally attain a breadth of 4-5 or even (Trochocarpa laurina) eight cells. In
some cases, however, both types of medullary rays are found in the same species
(e.g. in Epacris impressa). The primary medullary rays of the bast frequently
broaden outwards in the form of a wedge; this is the case in Lissanthe montana,
R. Bt'., and Monotoca. scoparia, R. Br. according to Liiders, and in Leucopogon
Richei according to Baccarini. The wood~parenchyma 1 is, for the most part,
not prominently developed,. but is present in some quantity in Prionotes
cerinthoides R. Br. according to Liiders, and in Dracophyllum U11Jilleanum,
Epacris longiflora, Richea Gunnii, and Trochocarpa laurina according to
Baccarini. In certain species of Acrotriche, Leucopogon, and Monotoca cham-
bered parenchyma containing crystals is found in the wood (Liiders). According
to Liiders the entire wood-prosenchyma, constituting the ground-mass of the
wood, bears bordered pits; according to the same authority, it is prov..ded with
a spiral band also in Epacris paludosa, R. Br. On the other hand, Baccarini
also observed mechanical fibres bearing simple pits, although such cases ar~ of
rare occurrence (Monotoca scoparia, Sprengel1,'a incarnata, W oollsia pungens, and
species of Epacris).
Regarding the structure of the cortex the following details may be men-
tioned, In all cases in which the primary cortex had remained intact in the
material employed for investigation (viz. in species of all the genera except
A crotriche, Brachyloma, N eedhamia, and Sprengelia) , Liiders was ~ble to demon-
1 Lijdels' statement that the wood-parenchymn occ:lsionally· bears bordt'red pits, is probably
incorrect and may perhaps be attributed to the presence of one-sided bordered pits on the walls of the
vessels where they are in contact with parenchyma.
 ADDENDA--EPACRIDEAE 973
strate the pr~sence of a continuous and composite ring of sclerenchyma or of
.isolated groups of sclerenchymatous fibres 1 in the pericycle, and of cork situated
on the inner side of the latter. The fibres of the pericyclic sclerehchyma, like
the ~c1erenchymatous fibres accompanying the vascular bundles of the veins (see
p. 493 and Fig. III, e), bear either bordered pits only (species of Andersonia,
Epacris, Lysinema, Woollsia) or bordered pits side by side with s~ple pits
(e.g. in species of Conostephium, Dracophyllum, Leucopogon, Richea, Sphenotoma).
The cells of the cork have uniformly thickened walls, and in most of the genera
do not show a distinct radial arrangement; the only exceptions in t~is respect
are afforded by Archerja, Lebetanthus, and Prionotes. Liiders' investigations
on the cqurse of development of the cork in those cases in which the cork~cells
exhibit an irregular arrangement, although not yet sufficiently extensive, have
shown tha\ a typical cork~cambium is really (d. also p. 493) not pJ;'esent iI) these
forms; regarding this point, see also Baccarini, loco cit., pp. 96, 97. .In many
species repeated formation of cork takes place in the bast. In numerous
members of the Order the secondary bast contains bast-fibres, which frequently
constitute complete annular zones and give rise to a stratification of the phloem,
while in other species they merely form groups of varying size. Bast-fibres
were not observed in a number of species, but this may possibly be due to the
slight thickness of the branch examined. According to Liiders the se~ondary
bast-fibres in most members of the Order likewise bear bordered pits side by
side with simple pits. In certain species the'secondary bast contains chambered
parenchyma with solitary crystals.
According to Liiders the pith in Cosmeria, DracophyUum, and ·,Richea
contains relatively small celis, which are distributed, in a reticulate manner
between larger elements, while in Dracophyllum it includes peculiar crystalline
conglomerates, which have a corroded appearance, and in some cases resemble
clustered crystals; in other cases ordinary crystals of oxalate of Ii,me occur in
the pith. According to Baccarini the cells of the pith are occasionally collen~
chymatous (Monotoca serrulata) or provided with thin walls (Leucopogon lanceo-
tatus and L. amplexicaulis); in Coleanthera myrtoides, moreover"they contain
crystalline masses, which Baccarini regards as consisting of carbonate 9£ lime.
owing to their behaviour when treated- with acetic acid (solution accompanied
by an evolution of gas).
Literature: Liiders, Stammanatomie der Epacrid.,_.. Diss., Heidelberg, without date Cll)~-l).
pp. 27-82 and Tab. ii.-Baccarini, Anatomia delle Epacrid., Ntlovo Giorn. bot.lta1., N. S., ix, 1902,
pp. 81-114 and Tab. iii-yo

DIAPENSIACEAE (p. 494).

Literature: Boergesen, Arkt. fll. bladbygn., Bot. Tidss~rift, ",ix, 189:;, p. 219 et seq.

PLUMBAGINEAE (pp. 495-5 01).

To the previous account of the structure of the. leaf (p. 499) we may add
that in St'!'tice sinJta!a. L. the cauline leaves develop palisade tissue and stomata
on both SIdes, while the radical1eaves have paJisade tissue (of a reduced, t~)
only on the upper side and stomata only on the lower side (Paoli). Redlich
has demonstrated. cortical vascular bundles (see p. 500) in additi0n¥ species
of Armeria and Statice (incl. Goniolimon). as well as in species of .Li11Jonit~str!4m
(here the bundles only traverse the cortex for a very short distance).

1 According to Baccarini there is no mechanical sheath in the pericycle in Leu(opogrm revo/ulus.

Oligarrl&ena micra'ltlza, Afl)110tOia sert'1tiata (= Acrolricke !erruiata), and_c AntierS(Jm'4 mtdr·a.'
914 ADDENDA-PLUMBAGINEAE
Literature: Redlich, Gefassbundelverlauf bei den Plumbag., Diss., Erlangen, without date
(commnn. to the universitrlibrary of ErJangen in 1896),30 pp.-Schubert, Parenchymscheiden, Bot.
Centralbl., 1897, iv, p. 19.-Wagner! Neuere progen, Diss., Erlang~n, ] 897, Pp. 13-21 (P/~1Jlba!JO).
-Leisering, Inteuylares Leptom, DlSS., Berhn, 1899, p. H.-Paoh, Eterofiil:a, Nuovo Glom. bot.
]tal., N. S., xi, 1904, pp. 217-19.

PRIMULACEAE (pp. 50I-509).

I. REVIEW OF THE ANATOMICAL FEATURES. The following additions are
necessary. Branched multicellular clothing hairs are found also in species of
A retia, Coris, Dionysia, and Douglasia. Oxalate of lime is absent only in the
vegetative organs, since in certain members of the Order (e.g. Cons, Glaux,
Hottonia, and Primula) the subepidermal layer of the testa contains .crystals of
this salt. Development of cork is rare, and takes place in differ~nt po~itions,
viz. in the pericycle or in the superficial cell-layers of the primary cortex.
Anomalous structure of the stem (more or le~s distinct polystely, combined
with the occurrence of a 'reseau radicifere ') is found also in Bryocarpum and
Dodecatheon. Arm-palisade parenchyma has been observed in the mesophyU
in species of Lysimachia and Trientalis.
2. STRUCTURE OF THE LEAF. The follo'\\ing supplementary details are
taken from Decrock's work. In the xerophilous species the epidermal cells
are provided with straight lateral walls. The cuticle, in some cases, shows
striation and punctation. The stomata are occasionally raised (Cortusa Mat-
thioli, L., Primula sinensis, Lind!.), while in the xerophilous species, Dionysia
revoluta, Boiss., they are confined to two furrows on the lower side of the leaf
Water-pores are found 8.t the end of the_J median vein in all the Primulaceae,
and in many cases may be met with at the ends of the larger lateral veins as
well. The leaf is for the most part bifacial in structure, although centric leaves
occur also among the xerophilous species. The palisade tissue consists of one or
more layers of cells, which vary (onsiderably in length. Decrock figures arm-
palisade parenchyma composed of short cells in Lysimachia punctata and
I have myself observed typical arm-palisade tissue consisting of short cells in
TrientaJis europaea, L. and T. americana, Pursh, and indications of arm-palisade
cells in the first and especially in the second layer of the mesophyll in Lysimachia
nemorum, L. The vascular bundles of the veins are occasionally provided with
sclerE'Ilchyma. .
According to Decrock those species which have a distinctly differentiated
petiole exhibit only one vascular bundle at the point of insertion of the leaf.
The petiole of Prim'ula rubra, according to Bouygues, contains a median hemi-
concentric vascular bundle, provided with a pith, and two lateral concentric
bundles, devoid of a pith; each of these bundles is surrounded by a typical
endodermis.
As regards the hairy covering the following facts deserve mention. Decrock
also records branched hairs in A retia pubescens, L. (side by side with unbranched
hairs; for the trichomes of Androsace § Aretia, see also Jeanpert, loco cit.),
Coris monspeliensis, L. (on the calyx), Dionysia and DougZasia Vitaliana, Hook.,
whilst glandular hairs were demonstrated by him in. species of all the 28 genera
included in Pax's monograph. His statements as to the structure of the head
of the external glands in the individual genera are unfortunately not sufficiently
comprehensive; we may, however, note that in Androsace villosa, L., both
unicellular and multicellula.r heads occur side b)7 side. In Hottonia palustris, L.
the heads of the glands are unicellular only, and not bicellular, as was stated
on p. 503. From a systematic-chemical point of view it is interesting to note
that the four species of Primula, in which Nestler demonstrated the presence of
a secretion, having an irritating effect upon the skin (viz. P. cortusoides, L.,
 ADDENDA-PRIMULACEAE 975
P. obconica, Hance, P. Sieboldii, Morren, and p, sinensis, Lindl.), all belong to
the section Sinenses.
3. STRUCTURE OF THE STEM. The structure of the normal stem in the
Primulaceae has recently been investigated from two distinct sides, by W. Meyer
and by Decrock, the two authors choosing different anatomical features as a
basis for classification. Their results are briefly summarized in the following
paragraphs.
In W. Meyer's system the presence or absence of a pericyclic strengthening
ring is the cJriei character employed in classification. Other points taken into
consideration are i the position of the strengthening ring (whether shifted towards
the outside or towards the inside); and the relation of the vascular bundles to
one another (whether the xylem-groups are united to form a ring by means of
interfascicular wood, or the bundles are separated by unlignified or sclerosed m.edul-
lary rays). W. Meyer records the presence of the strengthening ring in species of
AndYosace, At'etia, Centunculus, Coris, Cortusa, Hottonia, Lysimachia, Primula,
Samolus, Soldanella and Trientalis, and its absence in species of Anagallis, Andro~ace,
Aretia, Cyclamen, Glaux, Gregoria (= Dionysia)~ Lysimachia, Primula and Soldanella.
Decrock distinguishes three physiological types, corresponding to the exomor-
pbic features presented by the shoot. They are as follows:· I. Type Pl'imula ~
Short shoot with radical leaves; endodermis normal, peri cycle mostly paren-
chymatous. wood and bast forming a ring: Androsace pro parte, Ardisiandra,
Bryocarpum, Cortusa, Dodecatheon, Hottonia, Kaufmannia, Pomatosace, Primula
pro parte, Soldanelta, Stimpsom'a. II. Type Lys'imacMa: Rhizome with leafy
shoots; primary cortex narrower than in I, soft bast less developed, peri cycle in
the subaerial shoots invariably sclerosed: A nagallis. A pochoris, A sterolinum,
Centunculus, Glau:r, Lubz·nia, Lysimachia, Naumburgz·a, Pelie#era, Stelronema,
T"ientalis. III. Type Aretia: cushion-plants with acicl.llar leaves; cortex exfoli-
ating as far as the endodermis, which divides by radial and tangential walls, while
its inner walls become thickened; bast present in very considerabJe quantity and
collenchymatous; wood strongly developed with non-lignified wood-parenchyma:
Andyosace pro parte, A1'etia, Caris, Dionysia, Douglasia.
For the structure of the tubers of Cyclamen~ see Hildebrand, 'po 97, and Decrock,
p. 179 et seq. Regarding the aerating tissue in the primary cortex of Lysimachia
vulgaris, L., see Witte, loco cit.i
Formation of cork (see p. 504) is rare, although a number of cases are
mentioned by Decrock. The place of development of the cork varies'; it may
be the epidermal or subepidermal layer of cells (Lysimachia Ephemerum, L. and
L. vulgaris, L" or Dodecatheon), the outer ~one of the primary cortex (Pricnu1a
acaulis, Jacq.), or the pericycle (Douglasia Vitaliana, Hook., Primula bullata,
Franch.).
In dealing with the anomalous structure of the axis, I must, in the first
place, refer once more to the anomalies which are shown by the stem in
certain species of PI'£mula (polystely and 'reseau radicifere '), and have been
elucidated especially by Van Tieghem and DouIiot (d. pp. 504, 50S). The
groups which Van Tieghem establishes on the basis of these anatomical features
do not coincide with the natural groups in Pax's system of classification. I
append here:_~an enutfieration of the species of Primula which have a stem with
anomalous structure (alter'Van Tieghem).
The members of the Officinales, Van Tieghem which have a · reseau radicifere
are: Primuta acaulis, P. amoena, P. auriculata, P. elatioY, P. eltiptica, P. macrocalyx,
P. maJvacea, P. officinait's, P. pe#olart's, P. st'kkimenst"s.
Among the members of Auricula, Van Tieghem, which likewise have a 'reseau
radicifere,' Primula f'eptans still shows monostelic structure with a reduced pith.
Polystelic structure occurs in the following modifications: (I) More Qt Jess numerous
steles. which are either irregularly scattered or exhibit an annular arrangement and
are fused at some points to form small arcs, in: Primula algida, P. Allioni, P. angusti-
.folia, P. Balbisii, P. calycina, P. carniolica, P. Clusiana, P. commutata, P. cunei/alia,
P. daonensis. P. Delavayt'. P. erosa, P. Floerkeana, P. glabra, P. glutinosa~ P. hirsuta.
 ADDENDA-PRIMULACEt4E
P. integl'ifolia, P. Kitaibeliana, P. tati/olia, P. marginata, P. minima, P. minutissima.
P. Muretiana, P. nivalis, P. Palinuri, P. Parryi, P. pedemontana, P.\t*be~cens.
P. spectafn"Jis, P. tYl'olensis, P. unifiora, P. ' ursi,' P. venus/a, P. viscosa, P. yunnanen·
sis. (2) Steles broadened in the shape of an arc, not numerous, and arranged to
form a ring, in some cases fusing to form still wider arcs, in: Primula amethystina,
P. bella, P. calliantha, P. capitellata, P. Davidi, P. denticuiata, P. D,·cMeana, P. fari-
nosa, P. glacialis, P. Heydei, P. incisa, P. involucrata, P. longiflol'a, P. macrocarpa,
P. Maximoun'czii, P. memb,anilolia, P. Moorkraltiana, P. moupinensis, P. ovali·
/olia, P. pinnatifida, P. Poissoni, P. secundi(lof'a, P. sibirica, P. sonc_hifolia. P. spicata,
P. stricta, P. Stuartii, P. viscosa. (3) Steles fused to form a mO~:?a~r less complete
ring, the ' reseau radicifere' being almost absolutely ann ,in: Primula
japonica, P. nutans, P. obtusilolia, P. prolilera, P. purpurea, P. setralifoUa.
Decrock observed polystelic structure also in P. capitata, Hook., P. Faunae, Franch.
and P. petiolaris, Wall.
Anomalous structure of the stem, similar to that found in the Auriculag
(viz. polystely, combined with the presence of a reseau radicifere') -was
I

observed by Decrock also in Bryocarpum hymalaicum, Hook. f. et Th. and

Dodecatheon Meadia, L.; for details, see loco ci~. I
Literature: Costantin, Tiges :u!r. et sout., Ann. sc. nat., sere 6, t. xvi, 1883, p. 110 et seq.-
Costantin, Tiges d. pI. aquat., Ann. sc. nat., ser. 6, t. xix, I8H4-, p. 287 et seq. and pl. 15 and 17.-
Darwin, Bloom and distribution of the stomata, Journ. Linn. Soc., xxii, 1887, p. 1I4.-Pax,
Primula, in Engler, Bot. Jahrb., x, 1889, p. 75 et seq.-Seott, Polystely, Ann. of Bot., v, 1890-1,
p. 516 et seq.-[Blasdale, Leaf·hair Btruet., Erythrea, i, 1893. p. 25:1 et seq.; ahstr. in Bot. Centralbl.,
1894, ii, p. 402.]-Boergesen, Arkt. pl. bladbygn., Bot. Tidsskrift, xix, 1895, p. 219 et seq.-G'llffroy,
Pri11lula et Lysimachia de III flore paris., Hull. Soc. bot. de France, r898, pp. 341, 3.p.-Hildebrand,
Cydamm'jena, 1898, pp. 97, 114, ete.-Spanjer, Wasserapparate, Bot. Zeit., 1898, p. SI.-Hirsch,
Entwickl. • Haare, Diss., Berlin, 1899. p. 34.- W. Meyer, Beitr. z. vergl. Anat. d. Caryophyllac. .
u. Primnlac., Diss., GottingeIJ, 1899, pp. 43-59 and 68-7o.-Minden, Wassersez. Org., Bibl. bot., t
Heft 46. 1899, pp. 19 (Hpllonia) and 61 (Glau.r).-Paulesco, Strnct. an at. des hybrides, These,
Geneve, 1900, p. 71 (Pr;mula).-Thomas, Feuilles sont., These, Pari~. IQoo.-Decrock, Anat. des
Primulac., Ann. se. nat., ser. 8, t. xiii, 1901, pp. 1-199.-Pitard, Pchieyclc, These, Bordeaux,
190 1 , p. 50.-Bouygues, Petiole, These, Paris, 19°2, PP.14 and 99.-Nestler, Sekrrt d. Drilsenh. d.
Gatt. Primula, Sitl.-Ber. Wiener Akad., cxi, Abt. I, 190:1, pp. 29-51 j see also Ber. deutsch. bot.
Gesellscb., 1900, p. 189 et seq. and Tab. vii, viii, and p. 327 et seq.-Schoute, Stelartheorie, 1903,
p. I:33·-Brockschmidt, nO/lonia, Dill!., Erlangen, 1904. pp. 9-17.-Freidenfeldt, Anat. Baa d.
Wurzel, Bibl. bot.) Heft 61, 1904, pp. 66-8.-Nestler, Hautreizende Primeln, Berlin, I904.-PU
and Knuth, Prim ulaceae, in Pfianzenreicb, Heft "2, 1 905, pp. 3-5.-Theenn, Vaxttrichom.. Arkiv
for Bot., iv, n. 18, 1905, pp. 9, 10 and :n.-Dauphine, Rhizomes, Ann. &c. nat., str. 9, t. iii. 1906,
p. 342 et seq.-Geneau de Lam arliere , Membr. Cllt. des pl. aqa., Revue gen. de bot., 1906, p. 28y
et seq.-Jeanpert, Androsact du groupe AreNa, Bull. Soc. bot. de France, 1906, 'Pp. I~:I-"'.­
Witte, Aerenchymat. Gewebe bei L~simacltia vulgaris, Botaniska Studier, tillagn. Kjellman, Upsala,-
1906, p. 265 et seq.-[For further literature, see p. 1171.]

MYRSINEAE (pp. 507-5I2).

1. REVIEW OF THE ANATOMICAL FEATURES. The following details may
be added. The hypodermal sclerenchyma, which is a characteristic feature of
the leaf in the Theophrasteae, is absent only in a small group of species of
Claviia, and occurs also in the Tribe Eumyrsineae (viz. in Weigeltia Schlimii,
Mez). In addition to the glandular hairs with a multicellular head, others with
a unicellular head are exceptionally (Jacquinia pungens, Gray) found. New
types of clothing hairs have been observed in certain species of ] acquinia in the I

form of multicellular trichomes, which are either forked or branched like

antlers. The peculiar branched hairs previously described as occurring on the
stem in Jacquinia barbasco, Mez (Syn. J. armilla,is, Jacq.) are characteristic
of t",:o .gro~ps of closel:y allie~ species of l.acquinia. We may aJ:;o note that
gelatImzatIOn of the epldermIs of the leaf 15 completely wantmg m the Theo-
phrasteae, and that the occurrence of crystals of oxalate of lime in the epidermis
of the leaf in Jacquinia and Deherainia constitutes a generic character.
2. STRUCTURE OF THE LEAF. Votsch has recently published a careful
 ADDENDA-MYRSINEAE 977
investigation of the structure of the leaf in the Theophrasteae in connexion with
Mez~s Inanograph in the' Pftanzenreich.' The following details are abstracted
frum his paper and from the remaining literature.
In most of the Theophrasteae the mesophyll (d. p. 507) is not differentiated
into palisade and spongy tissue, although in certain species one or more layers
of palisade tissue are found. The epidermal cells in the Theophrasteae, for
the most part, have straight lateral walls. The cuticle is frequently strongly
striated, rarely ,(species of Jacquinia) granular. Peculiar local thickenings of
the walls of thaepidermal cells showing distinct str3:tification and recalling the
cystotyles of the Begonieae occur in C1aviia boliviensis, Mez. Theop"hrasta
/ussieui, . LindI. exhibits a peculiar calcification of the membranes of the
epidermal cells; surface-sectIons of the epidermiS, after being exposed to a red
heat, show the cellular framework, which on treatment with sulphuric acid
becomes transformed into needles of gypsum. A parenchymatous hypodenQ,
previously recorded in certain species of Claviia and J acquinia, is rather widely
distributed in these two genera, and occurs also in Theophrasta /ussieui, Lindl.
As a general rule, it is confined to the upper side of the leaf, but in Clavija
Kalbreyeri, Mez and C. nobilis, Mez it is found also on the lower side. The
cells of the hypoderm are larger than those of the epidermis, only in Claviia
spathuJata, Ruiz et Pav.
Accordi~ to Votsch a one-layered hypoderm is found in: Clavija boliviensis,
Mez, C. ,auli (Wa, Re~el, C. Hassleri. Mez, C. integrilalia, Mart. et Miq., C. Jelskii,
Szyszyl., C. ehmanmi. Mez, C. longi/oli<I:. Mez, C. nobilis, Mez, C. parviflora, Mez,
C. Poeppigii, Mez, C. Radlkoleri, Mez, C. Rodekiana, Lind. et Andre, C_ se"atijolia,
Mez, C. spathulata, Ruiz et Pav.; Jacquinia acuieata, Mez, J. barbasca, Mez, J. iwa-
siliensis, M~z, J. Eggersii, Urb., J. flammea, Millsp., }. linean's, Jacq., 1. ovalifolia,
Mez, J. pubescens, H. B. K •• J. revoluta, Jacq., J. Schiedeana, Mez, J. Seleriana, Urb.
et Loes.; Theophrasta Jussieui. Lindl; a I-2-1ayered hypoderm in; Clavija
Schwackeana, Mez; llUquinia keyensis, Mez, 1. Sprucei, Mez; and a typical 2-]ayered
hypodenn in; ClafJ'Jja Kalbreyeri, Mez and C. Ruiziana, Mez. Alocal development
of bypoderm is found in the neighbourhood of the veins in Clavija macrophylla,
Radlk. and Jacquinia incrustata, Urb., and near the margin of the leaf/in Claviia
parvuJa, Mez (see Votsch, pp. 14 and 34) and C. tarapotana, Mez. f
If we adopt the synonymy given in Mez's monograph, the only species pre-
viously recorded (see p. 507) as having hypoderm, which are not included in the
preceding lis4 are Clavija spinosa, Mez (formerly cited as C. Riedeliana and C. cala-
neural and C. umlwosa. Reg., and, outside;of the Theophrasteae, Aegice,as majus,
Gaertn.
In Myrsine heberdenia some of the epidermal cells are subdivided by a horizontal
wall, the lower cell. which is the larger of the two, occasionally including a crystal
of oxalate of lime (Clauditz).

The stomata in the Theophrasteae arc confined to the lower side of the
leaf. In the species of J acquinia they are deeply sunk, while in the other genera
they project above the level of. the epidermis by means of strongly developed
horn·likecprocesses:- In the Theophrasteae the vascular bundles, both of the
larger and smaller veins, are quite generally accompanied by sclerenchyma.
In the genera of the Clavijeae, A. DC. (Clavila, Theophrasta, Neomezia) the
median vein is traversed by two or more vascular bundles, while in the genera
of the Ja cquinieae, A. DC. (Deherainia, J acquinia) it comprises only a single
bundle. Among the Theophrasteae oxalate of lime has been observed only in
Claviia, Deherainia, and / acquinia (but not in N eomezia and Theophrasta); It is
deposited as clustered crystals in ] acquinia only, while in all other cases it is
found solely in the form of very small rhombohedral, prismatic, or acicular
crystals. In Jacquinia and Deherainia, as well as in Clavija serrati/olia, Mzz,
crystals of oxalate of lime (clustered in the case of Jacquinia) occur quite'
generally in the epidermis of the leaf, but are found also in the mesophyll.
SOl.U~OE. 3R
 ADDENDA-MYRSINEAE

The sclerenchymatous hypoderm, found in the Theophrasteae (see

p. SO<}), has been subjected to a thorough re-investigation by Votsch. The most
Important result, from the systematic point of view, lies in the fact that this
feature occurs in all the Theophrasteae with the exception of two small groups
of very closely allied species of Clavija, namely: C. Hassleri, Mez and C.
Jelskii, Szyszyl.; and C. lulgens, Hook. f. and C. boliviensis, Mez. In C.
;nlegrifolia, Mart. et Miq. the absence of tl)e hypodermal sclerenchyma is only
apparent, since in tbis species it is reduced to isolated fibres, lying near the
vascular bundles of the veins. Votsch describes the hypodennal sc1erenchyma
in the following words: 'It consists of elongated fibres, which are strongly
thickened (more rarely slightly thickened, as in Clavifa), provided with
oblique pits, and run mdependently of the sclerenchyrnatous sheaths of the
vascular bundles. They are united to form larger (Theophrasta, N eomezia,
]acquinia) or smaller (Claviia, Deherainia, Jacquinia) bundles, or (more rarely,
in Claviia Rodekiana, C. se"ata, C. grandis, Jacquir.ia aculeata, J.linearis) are
arranged in one-layered strata, in the latter case co~tituting sclerench::,...matous
plates.' In most of the Theophrasteae (especially in Theophrasla) the hypo-
dermal sclerenchyma on the two sides of the leaf is joined by means of spicular
fibres, which enter into connexion also with the sclerenchyma of the veins. In
concluding the description of the hypodermal sclerenchyma of the Theophrasteae
we may add that in Weigeltia Sck/imii, l'rfez-the only species of the subgenus
Triaclophora (Tribe Eumyrsineae)-Me z observed sclerenchymatous fibres
beneath the upper and lower epidermis of the leaf, just as in the Theophrasteae,
and that Areschoug figures moderately elongated spicular cells of a parenchy~
matous shape in the mesophyll of Aegiceras majus, Gaertn.
In most of the Theophrasteae the margin of the leaf is strengthened by
a single thick strand of sclerenchyma or by several sclerenchymatous strands,
but in some species of Claviia (e.g. C. biborrana, OeTst., C. grandis, Decne., and
C. Lehmannii, Mez) it is supported by a vascular bundle, from the sclerenchyma
of which fibres are given off to the margin of the leaf; in a transverse section
these fibres are cut through both longitudinally and transversely.
The section dealing with the hairy covering (see p. 509 et seq.) likewise
requires a few important additions. Unicellular clothing haIrS are wanting in
the Theophrasteae; nor are uniseriate trichomes very abundantly developed
on the leaves, although they are more commonly found on the vegetative axis
and in the floral region. A transition to the characteristic branched trichomes
of Jacquf,'nia barbasco, Mez (d. Fig. 116 c on p. 510) is afforded by the un i-
seriate hairs of J. Schiedeana, Mez, in which the terminal cell is divided by a
longitudinal wall. Peculiar branched trichomes, having thick walls and resem-
bling those found on the axis in Jacquinia barbasco, have been demonstrated by
Votsch in the species included under Nos. 1~ in Mez's monograph (J. aculeata,
Mez, ,. barbasco, Mez, J. Berterii, Spreng., J. brasiliensis, Mez, J. Eggersii, Urb.,
]. incTustata, Urb., J. keyensis, Mez, J. linearis, ] acq., J. revoluta, Jacq.) 1.
With these trichomes we may class the multicellular clothing hairs occurring in
the species of Jacquinia, described under Nos. 31-33 of Mez's monograph (viz.
J. Seleriana, Urb. et Loes., J. Sprucei, Mez, and J. pubescens, H. B. K.); these
hairs are either dichotomously branched or resemble an antler in shape.
Glandular hairs have been observed only on the axis of ] q,cquinia pungens,
Gray; they are not sunk, and are provided with a unicellular stalk and a uni·
cellular head.
3. STRUCTURE OF THE AxlS. According to Pitard a composite and con-

I According to Votsch (p. 532) trichomes of the same type as those found in /atguinia 6a,,6as(1J
occur also in J. flammia, Millsp. and J. Sit ItOpltylla , Urb., while on pp. 521 and 524 they are
stated to be absent in these species. Tht'se contradictory statements remain t(l be explained.
 ADDENDA-MYRSINEAE
tinuous ring of sclerenchyma is also developed in the pericycle in species of
A,disia, Cononw,pha, and Embelia. The statement as to the excretion of car.
bonate of lime in the lumina of the vessels in Myrsine Grisebachii, Hieton. must
be canceiled, since, according to Mez, M. Grisebachii-does not belong to the
Myrsineae, but is a member of the Sapotaceae (ChrysophyUum G,isehachii, Mez).
It remains to mention that, according to d' Arbaumont, the cells of the cortex
and pith in Myrsine a/ricana show a passing deep indigo-blue coloration on
treatment with caustic potash.
For the structure of the terrestrial roots of A egiceras maius (annular thickenings
in the cells of the cOrtical parenchyma), see Karsten, 'loc. cit.
,
Literature: Wijnaendu Francken, Sclereiden, Diss.. Utre(b.t, 1890, pp. 56, 57.-Karsten,
~:uJgrovevegetation, Bib1. bot., Heft n, 1891, p. 5o.-Boergesen og P&ulseu, VegetAt. dansk-veatind.
Oer, Bot. Tidsskrift, xxii, 1898-9, pp. 33, 33 UO<fJuinia tJrmil/aris, Jacq.)-D'Arbaumoo.t, Mynine
a.fricflna, Joam. de bot., 1900, pp. 361-8.-Pitard, Perieycle, These, Bordeaux, 1901, D. 6+-
Areschong, Mangrovepll., Bib). bot, Heft 56, 190~, pp. 55-1 and Tab. iv (.AegUenul}.~lauditz,
Blattanat. canar. Gew.• Disl., Basel, 1901, pp. :18-31 (HelJerdmia, Ple~risl).-Mez... Myninaceae,
in PfianzeJlreich, Heft 9, 1901, pp. 3, ....-Meonecb~t, Poils ~pid. des Myrsln. etc., Jaura. de bolo.
190 2 , pp. 355-7.-Mez, Theophrastaceae, in Pfianzenreich, Heft 15. 1903, PP' 3, +-Votsch, Syat.-
an at. Unters. von Blatt u. Axe del Theophrastaceae, in Engler, Bot. J&hrb" nxlll, 1904, pp. 503-..6 •
• lso Diss., Erlangen, 1903.-Joh. Schmidt, Verdens Mangrove trreer, Bot. Tidsskrift, 1)10..,.
pp. 106-13 (salt-glands of Aegiceras 11lajus).-Areschoug, Trap. vaxt. bladoyggn., Sv. Vet. Akad.
J-Iandl., 39. n. 2, 1905, pp. i33, 13. (ThUJpkrasta), pp. 1 ...8,1 ...9 (Arrtisia), pp. 15l-~ (JtM(juini4).
-[H. Weiss, Aegiceras majus, Dil3., Strassburg, 1906; extract in Archiv d. Pharm. 1906, p~ ut.]-
Holtermaon, Einflasa d. Klimas etc., 1907, p. 58 (Aegiceras).

SAPOTACEAE (pp. 512-5I5).

Engler records hypoderm in the leaf also in Buty,ospe,mum Pa,kii (one-
laye~ed), Palaqnittm oblong'':/olium (one-layered) and M,musaps Commersonii
(3-4 layered); and sclerenchymatous fibres in the mesophyll in Synsepalum
dulciforme. Daniell.
Engler observed lone-armed' hairs with transitions to ordinary simple
trichomes on the ovary of 1\1alacantha lVa,.neckean~, Eng!. and otlier species of
this genus. According to the same authority simple unicellular hairs almost
completely cover the different parts of the plant in Delpydo,a, Pierre.
Recent investigations on the laticiferous elements have been published
by Charlier, who showed that the sacs are present in the root also (primary
cortex and bast). According to Charlier, the occurrence of Iaticiferous sacs
running freely in the mesophyll is by no means so rare as has hitherto been
supposed. In the root, the Iaticiferous sacs of the bast are arranged in longi-
tudinal rows, which are united to form a network, or lie with their longitudinal
walls apposed to one another, the common walls of contact in the latter case
exhibiting tbin areas (' laticiferes anastomosees '); the laticiferous elements of
the primary cortex, en the other han<L consist of rows of cells, which as a rule
do not enter i.~to connexion with one another. In the axis, the bast likewise
contains • laticiferes anastomosees/ the longitudinal and transverse walls of
which show thin areas, which are occasionally even perforated; Charlier thinks
it not improbable that in the older axes these cells ultimately fuse to form a
system of laticiferous vessels. The primary cortex and the pith for the most part
include simple rows of laticiferous sacs only. The axis of certain species of
Bumelia is particularly remarkable in containing groups of irr~guIarItarranged

1 In Aegiuras Areschoug records secretory c~lls instead of s~cretory cavities, which is no doubt
a mistake.
, In this paper the !Oecretory ca.vities 3re likewise incorrectly described by C!auditz as secretoryceU ..
3R2
 ADDENDA- ··SAPOTACEAE

laticiferous sacs, which undergo fusion at an early stage. In the case of the
rows of laticiferous sacs· occurring in the leaf, it. is not certainly established
whether absorption of the transverse walls really takes place 'occasionally.· In
some members of the Order isolated laticiferous sacs are found in the meso,,:,
phyll, e.g. in the meshes formed by the network of the veins in Sideroxylon
brevipes, Bale (here side by side with groups of two or three laticiferous sacs).
According to Charlier crystal-sand occurs especially in the laticiferous sa~
of the leaf; it is rare in those of the axis, but is found in the sacs situated
near the epidermis. It remains to mention that the genus Tridesmostemon
likewise possesses laticiferous sacs.
For the excretion of carbonate of lime in the lumina of the vessels
in Chrysophyllum Grisebachii, Mez, see under Myrsineae. Bargagli-Petrucci
met with silica-bodies in the wood of Palaquium sp. and Bass£a crassipes, Pierre.
Literature: Bohnel, Gerberinden, Berlin, 1880, p. 106 et seq.-Brandt, Wenig bek. Rinden,
Diss., Dorpat, 1894, p. 9 et seq.-[Rempel, Payena ueri;, etc., These de pharm., MosCow, 1898
(Russian); cited by Grelot and Charlier.]-Grelot, Caontchollcs et Gnttapereha, These, Paris, 1899,
p. 240 et seq.-[Obach, Guttapereha, Dresden, '1899.]-Utsprnng, Anat. n. Jahresringbild., Diss.,
Basel, 1900, pp. 20-3 (Fm/JYi£aria maxima, Poir.).-Bargagli-Petrucei, Concrez. silicee, Malpighia,
]902, p. a3 et seq.; and Legnami, lac. cit., p. 360 et-seq.-Fabricius, Laubblattanat., Beih. z. Bot.
Centralbl., xii, ]902, pp. 306-8.-Col, Faisceanx, Ann. se. nat., ser. 8, t. xx, 1904, p. IH.-Engler,
Sapotaceae, in Monogr. rurik. Pflanz.-Fam. u. Gatt., viii, 1994, pp. :.I-5.-Areschoug, Trop. va.xt.
bLidbyggn., Sv. Vet. Akad. Handl., 39, n. 2, 1}}05, pp. 37. 38 and Tab. v, vi (Bassia) , pp. 106,
107 and Tab. xii (ChrysuphJ'llum).-Charlier, Et. anat. de_s pI. aGuttapercha, etc., These, Paris,
1905, 160 pp.; also Jaum. de bot., 1905 (Achra.r, Argan;a, Bassia, Bumelia, Cltrysoplzyllum,
Hormogyne, Lucuma., Mim"sops, Palaquium, Payena, Sz(leroxy!on) •...;..EngIer, Tnaesmostemtm,
in Engler, Hot. Jahrb., xxxviii, 1905, p. 99.-Ho1terma[ln. Einfluss d. Klimas, etc., '907, p. 179
(.8as.r;a).-[For further literature, see p. 1I 72.]

EBENACEAE (pp. 516-519).

3. STRUCTURE OF THE AXIS. Pitard records a composite and continuous
ring of sclerenchyma in the pericycle in several species of Diospyros and in M aba
buxilotia.
Literature: ,Knoblauch. Okolog. Anat., Habilitat.-Schr., Tiibingen, 1896, p. 2J et scq.-Pitard,
Pericycle, These, Bordeaux, 19°[111 p. 82.-Bargagli-Petrucci, Legnami, Malpighia. 1902, p. 361
(DwspyrQs).-[Wright, Diospyros, its morphology, anatomy, and taxonomy, Ann. Bot. Garden,
Peradeniya, ii, 1904, pp. I and 13.\ et seq.; abstr. in Just, 1904, i. p. 769.J-Piccioli, Legnami, Bull.
Siena, 1906, p. LJ.I.-[Elsler, Extraflor. Nektar. u. Papill. d. Blattunterselte vou Diospyros disc(}/(w,
Willd., Anzeiger Wiener Akad., 1907. p. 419; ,and Sitz.-Ber. Wiener Akad., cxvi, Abt. I, 1907,
pp. 1563-90, :.I Tab.]

STYRACEAE (pp. 519-52I).

The following new features have been observed: alumina-bodies in the
mesophyll and cortex in the species of Symptocos; papillose differentiation of
the lower epidermis of the leaf; development of hypoderm on the upper side of
the leaf; differentiation of arm-palisade parenchyma; spicular fibres branching
off from the sc1erenchyma of the veins; secretory organs resembling intramural
glands at the mar~ of the leaf or in the leaf-teeth; occurrence of solitary and
clustered crystals ill the epidermis of the leaf; hydathodes of peculiar structure.
All these features have been recorded in certain species of Symplocos.
The STRUCTUR.E OF THE LEAF in SymPlocos has recently been investigated by
Cador and especially by Wehnert. The leaves in this genus are invariably
bifacial. The palisade tissue consists of one or more layer~, and in many
species is differentiated as typical arm-palisade parenchyma, especially in the
lower layers. The spongy tissue likewise shows diverse structure; in certain
species it is composed of well·developed stellate cells, which (especially in the
lower layers) are occasionally somewhat sclerosed. The epiderma1.cells.have
straight or undulated lateral walls. Gelatinization of the epidermis has not
 ADDENDA-STYRACEAE gSl
been observed. Features deserving special mention are: the large epidennal
cells found in Symplocos con/usa, Brand; the epidermal cells of S. po!yanara,
Brand, with lateral walls bent in a zigzag manner and provided with ridge-
like processes in the apices of the angles; the papillae on the lower side of the
Jeaf of S. neriilolia, S. et Z., which are connected by means of cuticular ridges;
lastly, the nature of the cuticle, which is either provided with delicate striae or
with thick ridges, and often presents a very characteristic appearance (e.g.
especially in S. Itatiaiae, Wawra). The stomata are confined to the lower side
of the leaf, and are provided with two subsidiary cells placed parallel to the
pore, as in the Rubiaceous type, the--subsidiary cells in some cases undergoing
transverse division. According to Wehnert a typical one-layered hypoderm is
found beneath the upper epidermis in Symplocos Hohenackeri, Clarke, S. micro-
phylla, Wight, S. rubiginosa, \Vall., and S. Stawelii, F. v. M.; Holtermann
records a hypoderm also in S. obtusa (continuous and often even consisting of
two layers) and S. spicata (here locally developed). The vascular bundles of
the lateral veins are sometimes vertically transcurren~ on both sides of the
leaf, or at least on the one side or the other. \Vehnert demonstrated branching
of the sclerenchyma of the veins, the branches penetrating into the mesophyll,
in S. neriilolia, S. et Z. and S. rubiginosa, Wall., as well as in S. colorata, Brand.
Oxalate of lime is excreted in the form of large or small solitary crystals, which
are, for the most part, rhombohedral, although occasionally almost cubical,
while in other cases it is dep9sited as clustered crystals; in certain species of
Symplocos it occurs also in the epidermis of the leaf in the form of solitary and
small clustered crystals. The only type of trichome found in Symplocos is that
of simple sclerenchymatous clothing hairs provided with thin transverse walls.
With these we may associate the peculiar hydathodes, observed by Wehnert on
both surfaces of the leaf or only on the upper surface in S. adenophylla, Wall.,
S. nerii/olia, S. et Z., ai'ldS. tenuilolia, Brand. These hydathodes are unicellular
structures, having the shape of an inverted pistil; their outer end shows a
bladder-like enlargement, while internal to this the hydathode is provided with
a narrow neck, which is wedged into the centre of a rosette of 7-8 very small
epidermal cells, and a slightly enlarged basal portion. The secretory organs
above mentioned have been demonstrated at the margin of the leaf in S. adena-
phylla, Wall., and in the leaf-teeth in S. glomerata, King. They consist of
rounded spaces, traversed by large numbers of elongated tubular cells, which
form a secretion resembling gum, and are irregularly interwoven with one
another; the structures thus recall the intramural glands found in Psoralea,
and the glands of Milletia, &c. (see under Leguminosae).
The alumina.bodies, referred to above, were discovered by Radlkofer in
the mesophyll of Symplocos; according to Wehnert they probably occur in all
the species and are found in the cortex as well; In transverse sections of the
leaf, cleared with Eau de Javelle (which should contain no carbonate of soda)
they appear as grumose, pateriform, or placentiform bodies which are colour-
less, have~ an _irregular an gular- or rounded outline, and are insoluble in
water j they:fill the greate~ part of the palisade-cells, in which several of
these bodles lie one above the other, while they often take up almost the entire
breadth of the cell; in appearance they are not quite unlike the deposits
of solid fat, which occasionally occur in the cells of the leaf (Radlkofer,
loc. cit., p. 216). According to Wehnert they assume a bright red colour after
treatment with a weak alkaline solution of Brasilin, and are then easily recog-
nized. We may add that in many species of Symplocos fat is_met with in the
epidermis and mesophyll.
The scalariform perforations, previously recorded as occurring in the vessels
in SymfJZocos, were found by Wehnert to be of general distribution; the number
of bars varies from 15 to 30.
 ADDENDA--STYRACEAE
Uteratare: Pierre, Flore (mat. de la Cochinchioe, xvii, 189,.-LUdy, Sumatra.·Ben~,"{)i.Q •. ,
Bcru 1893, pp. 5:1--4--Barthelemy, S17r'U officinalis, These, Montpellier, 1895, pp. :J.3-33.-C~4or,
Mat~blatter, Bot. Centralbl., 1900, iv, pp. ,~, 345 and 369 et seq.; ,also Diss., ErllUlgen.-B~d,
Symplocacea.e, in Pflanzeoreich, Heft 6, 1901, p. 3.-Pitard, Pericycle, These, .Bc>rdeaux, 1901,
p. 9+-Radlkofer, TODerdek~, Her. deutsch. bot. Gesellsch., 1904. pp. :n6-:l4.-TlChiIch.
Han1I11I&, Flora, 1904, pp. 18~1.-WehDert, Anat...syst. Untersuch. d. BI. d. Gatt. S),1Il.plocl1s,
Dis.., MUnoban, 1900, 57 pp.-Holtermann, Einfluss d. Klimas, etc., 1907, p. lI8 (S)'1Itplocos).-
[Perkins, Styri.caceae~ in Pftanzcoreicb, Heft 30, 1907, pp. 5, 6.]

OLEACEAE (pp. 521-526).

Papillose differentiation of the lower epidermis of the' leaf (coronulate
papillae united by a network of ridges) occurs only in three cases among the
different species of ash, viz. in Fraxinus americana, L., F. juglandilolia, Lam.
(non Willd.) and F. Texensis, Sarg. (Kohne). Spicular cells are also found in
the mesophyll in Olea Gardneri and O. laurijolia (Holtermann, Gerhard). To
the previous description of the external glands (see p. 523) we may add that in
some cases (e.g. the glands on the buds of Fraxinus Ornus) they have a some~
what spherical h~d, divided by one or more vertical walll) in to the corre-
sponding number 'of cells. Regarding the occurrence of extrafloral nectaries
on the leaves of a species of Fraxinus, see Delpino, loco cit.
According to Baldacci the development of the cork takes place in the
epidermis in FOTsythiaeuropaea (cf. p. 525); According to Pitard the pericycle
also 'contains isolated groups of bast-fibres in Fontanesia phillyraeoides and
F01'sythia suspensa, while in Chionanthus~ Forestiera porulosa, Fraxinus, Jasmi-
num, Linociera compacta, Olea americana, O. europaea, and Phillyrea there is
a composite and continuous ring of sclerenchyma; in ] asminum the latter does
not persist for any great length of time, while in Olea europaea and Phillyrea
angustifolia the scierenchymatous ring is still found in axes of 4 cm. diameter,
and in Fraxinus excelsior even in those of 3S cm. diameter. MoUer's earlier
statements on this point (see p. 525) refer for the most part to pieces of bark
from old stems.
Literature: Wijnaendts Francken. Sclereiden, Diss., Utrecht, 1890, pp. 53-:;.-Stock, Protein-
krystalle, in Cohn, Beitr., vi. I 893.-[Tognini, Stomi, Atti 1st. bot. Pavia, J89-4.]-[Denniston,
Ff'tZXinus americana, Phann. Archives, 1898, n. J : abstr. in Just, 1898, ii, p. 13.)- Kohne, Fraxinus-
Arten, in Wittmack, Gartenfiora, 1899, pp. 383-8.-Baldacci, Fwsythia europaea, Degen et Bald.,
Mem. Accad. Sc. Bologna, ser. v, t. viii, 1900, pp. 481-90 and tab. i, ii.-Tunmann, ,sekretdriisen,
Dia, Bern, 1900, pp. 29-33.-Petersen, Vedanatomi, 1901, pp. 85-7.-Pitard, Pericycle, These,
Bordeaux, 1901, pp. 75, 76 and 86, 87.-Clauditz, Blattanat. canar. Gew., Diss•• Basel, 1902, pp. 26,
:17 (.P.cconia).-Gerhard, Blattanat. d. Gew. d. Knysnawaldes, Diss., Basel, 1902, ep. ~4-6 (O/ea).-
[Armari, Piante della reg. medit:. ~nnali di Bot., i, 1903, p. 17 et se!!. (Pkillyrea).J:-(Uelpino, Not.
fitobiol .• Bnll. Orto bot. NapolI, I, 1903, p. oOpS et seq.; abstr. In Jost, 1903, I, p. 38S.]-Col,
Faisceanx, Ann. sc. nat., ser. 8, t. xx, 1904, p. 1I7.-Sussenguth, Behaarungsverh. d. Wiirzb.
Mnschelkalkpft., Diss., Wiirzbarg, 1904, p. 44--Kohne, Forsythia, Gartenfiora, 1906, p. 199 et seq.
-Picc:ioli. Legnami, Bull. Siena, 1906, pp. 146, 154, 177, 178, 180, and I81.-Holtetmann, Einfluss
d. Klimu, etc., 1907, p. 119 (Olea).-[For further literature, see p. 1171.]

SALVADORACEAE (pp. 526-528).

For the strllcture of the leaf-spines of Azima tetracantha, see Lothelier.
Interxylary phloem is found also in the root of Salt/adora.
Literature: Moller, Rin~enanatomie. 188:3, p. 126.-Scott and Brebner, Str),ch1tOS, Ann. of Bot.,
ill. J88g, p. :a¢.-Lothelier• .c.piDCI1These, Paris, 1893, p. 27.-Brandt, Wenig bek. Rinden, Diss.,
Dorpat, 1894, p. ,S et aeq.-Leisering, Interxylares Leptom, Diss., Berlin, 1899, pp. '3-6.-Holter-
JIWID, Einftua d. Klimu, etc., '907, p. II 1 (Salvadora jJersica).
 ADDENDA

APOCYN ACEAE (pp. 528-534).

I. REVIEW OF THE ANATOMICAL FEATURES. The following additions are
necessary. Gelatinization of the spongy tissue has been observed also in
species of the genera Carpodinus, Chilocarpus, Cylindropsis, LandD/lphia, and
JViUoughbeia; hypoderm occurs also in the leaf of species of Bou~gonia, Carpo-
dinus, Chavannesia, Chilocarp us, I chnocarpus, Lepiniopsis, Leuconotis, M icrechites,
Nouettea, Para barium, Sclerodictyon, WiUoughbeia; papillae are developed on
the lower side or on both sides of the leaf, as the case may be, in species of
A pocyn,("m, Landolphia, W illoughbeia; spicular fibres are found in the meso-
phyll in species of Bousigonia, Micrechites, Neocouma, Sclerodictyon, Trachelo-
spermum. Bast·fibres, which are completely enveloped by crystal-cells, occur
in Quebracho-bark. Secretory cells, forming a layer beneath the palisade
tissue, are found in speciE:;s of Baissea, Echites, Kopsia, and T abernaemontana,
. mucilage-cells in species of Apocynum, Carissa, and Ichnocarpus, and lysigenous
mucilage-cavities in species of Bousigonia, Chonemorpha, M icrechites, Pottsia,
and Rhynchodia. Solitary crystals occur in the epidermis of the leaf in species
of Alstonia, Cerbera, and Hunteria. Cork-warts are pres~t on. the leaves in'
species of Carpodinu.s, Chilocarpus, Clitandra, Landolphia, Leuconotis, and
Pyc1wbotrya. Oxalate of lime is occasionally excreted also in the form of small
solitary crystals.
2. STRUCTURE OF THE LEAF. The followi'lg additions to the earlier state-
ments are principally based on the papers by Garcin, Hallier" Payrau, and Spire 1.
The epidermal cells of the leaf have lateral walls showing diverse struc-
ture and lumina of varying size. In the xerophilous species the cuticle is often
strongly developed and 'frequently striated as well. Curved lateral walls pro-
vided with marginal pits are recorded by Hallier, for example, in all the species of
Landolphia, with the exception of L. gummifera, K. Sch., while st!rrated cuticular
prominences of quite a special type are stated to occur on the lower side of the
leaf in M elodinus orientalis. Bl. Gelatinization of the inner walls of the epidermal
cells has, curiously enough, not hitherto been observed in any memb~r of the
Order. In Chone1twrpha Griffithii, Hook. (Spire) and Chilocarpus costatus, Miq.
some of the upper epidermal cells of the leaf are divided by horizontal walls.
Hypodermisverywidelydistributed; it has recently been observed in the follow-
ing additional species: Ichl1.ocarpus trutescens, R. Br. (one-layered) and Nerium
odcrum, Soland. (3-layered on the upper side, 2-3-1ayered on the lower side) by
Garcin; Carpodinus lanceolatus, K. Sch. (one-layered), Chilo carpus suaveolens,
Bl., and C. verlticosus, HI. (one layer of ceJls with wide lumina), Chilocarpus sp.,
Hose n. 3049, Lepiniopsis ternatensis,Valet. (one layer of cells with wide lumina),
Leuconotis anceps, Jacq. (one or two layers of cells with wide lumina), Micrechites
micrant1r.a, Hallier f. (2-3-layered on the upper side, one-layered on the lower
side), M. polyantha, Miq. (2-3-layered), Sclerodictyon Grifjoniam/,m (according
to Pierre) and Willo_ug~~~i!1 gfandiflora, Dyer (I-2·1ayered) by Hallier; Bousigonia
angusti/olia, Pieiie,~ B. mekongensis, Pierre, Chavannesia esculenta, DC., Micre-

1 Garcia investigated species of Acokan/hera (TtJ,xicophltMa), AlJtxia, Amsmia, AfJD7nur,r,

Can'ssa (with Ardui'IUZ), Cerbera (Tangkinia), Ecnites, Fon/crania, .Ichnot:arptls, Mandevillo.,
Melodinus, Neril4m, Plumiera, Rauwoljia (Oph~x7/on), Tahernamumtana, Thevelia, Tracl#lo-
spermum (Rhynchosperml4m), Vinca j Hallier examined species of Carissa, CarpMima, CAiIoca,P"r,
Cluandra, Craspidospermum, C),lindrops,'s, Huntma, Lanao/(Jhia, upiniolsis. LeuctJ1Ultis,
Me/Dtiinus, Otope/alum (=ltlicrecniJes), WiJ/oughbe£a, Winchia; P"yrau examined species of
S/TlJpIJant}.us; and Spire species of the Indo·chinese genera Aganlmerion, Amaioca/yx, B(lUsigpni(J,
CItDnerJlbTjJlza, Ectiysa1'llMrtJ, Mel«/a'nus, MicrtcAites, Ntnlelt~} Paraha,.ium, Par4mena, R "ytteWWj
X;h"no/Jaria. The inve.rigations of Garcin, Pay!"ll.u, and Spire deal_also with the structure 0(
the axis.
 ADDENDA--APOCYNACEAE
ckites Jacqueti, Pierre(one-layered)~ Nouettea cochinchinensis, Pierre, PaTabarium
Spireanum, Pierre, P. Tournieri, Pierre and P. Verneti. Pierre (composed of
large cells) by Spire;" Chilocarpus atroviTidis, BI. (2-1ayered) by Areschoug.
Papillae are distinctly differentiated on the lower side of the leaf in Apocynum
cannabinum, L., Landolphia ochracea, K. Sch., and Willoughbeia grandifiora,
Dyer, ar..d on both sides of the leaf L~ Apocynum androsaemijotium, L. (Garcin
and Hallier). The papillae found in Willoughbeia grandi/lora show a peculiar
type of structure; they are narrow and columnar in shape, have narrow lumina,
and bear two or more spherical heads, thus resembling a femur in appearance.
Regarding the papillose prominences arising from the middle of the outer wall of
the lower epidermal cells in Vinca maioT and V. herbacea (not in V. minor), see
Haberlandt, lac. cit.; in these structurts, which can scarcely be described as
papillae, and which function as organs for the perception of light, the cellulose-
Jayer is produced into a stopper-like process which penetrates into the cuticle.
Stomata of the Rubiaceous type with a subsidiary cell situated to the right and
left of and parallel to the pore have, recently been demonstrated in species of
Carpodinus, Chilo carp us, Chonemorpha, Clitandra, Cy'lindTopsis, Landolphill., Lepi-
niopsis, Micrechites (with Otopetalum), Para barium, Strophanthus, Willoughbeia,
and WineMa; on the other hand, stomata with three or four neighbouring cells
are recorded in species of Carissa, Craspidospermum, Ecdysanthera, Hunte,ia,
Leuconotis) Melodinus, and Rhynchodf,'a (Hallier, Payrau, and Spire). 'The
stomatal pits, characteristic of the common oleander, occur not only in this
species, but also in IIerium odorum, Soland. Hallier describe.; stomata which
are deeply sunk and have a vestibule abov~ the guard-cells in Leuconopsis
anceps, Jack and longitudinal ridges, covering in the pairs of guard-cells, ia
Landolphia eapensis, Oliv. According to ,Hallier the gelatinization of the
spongy tissue, :first ob3erved by Radlkofer in certain members of the Order, as
a rule constitutes a specific character only; it is widely distributed in the genera
Carpodinus, Chilocarpus, Clitandra, Cylindropsis, Landolphia, and W illoughbeia 1•
In certain species of Carpodinus, Chilo carpus, Melodinu.s, and Willo1.ighbeia
fissures due to dryi.'1g have been recorded in the palisade tissue. Spicular fibres,
which have a varied course and branch off from the sclerenchyma of the veins,
are found in Bousigonia rnekongensis, Pierre, Micrechites micfantha, Hallier f.,
M. polyantha, Miq., N eocouma ternstroemiacea, Pierre, SckrodictyonGriffonianum,
Pierre, and Trachelospermum iasminoidcs, Lem. (Garcin, Hallier, Pierre, Spire).
According to Hallier cork-warts, like those found in PycnobotTya (see
p. 529), occur on the lower side of the leaf also in all true species of Cllrpodinus
and Chilocarpus (in Carpodinus ligustTi/ol-ius, Stapf on the upper side as well),
, as well as iu Clitandra myriantha, Pierre, Ldndolphia owariensis, Hallier f., L. Teti-
cutata, Hallier f., Leuconotis anceps, Jack, and L. eugenifolius, DC. The varia-
tions recorded in Landolphia owariensis are related to the more or less advanced
stage of development of these structures.
The structure of the petiole has been examined in some detail by GarcL"l,
Hallier, Pierre, and Spire. In Nerium the petiole is traversed by 5-7 vascular
bundles, but in many ApocY!lz.ceae there are only three bundles1 of which the two

1 The !ipeci~s in wbich thjs feature has been observed are as follows: Carftldinus Buleri, Stapf,
C. /uJvus, Pierre, C. /ameoJatu.r, K. Sch., C. /4j1anlhu.1, Stapf, C. lz'gwtn:tl)/iu.1, Stapf,
e. mtuimus, K. Sch. (vix I), C. sl4brep4ndus, K. Sch. (vix I), C. lurbt'natus, Stapf, C. vlDkceus,
K. Sch.; Chilocarpus an-OfJ£ridis, Bt, C dmuda/us, BI" C. s'.laveo/ens, Bl., C. 'lIerni~osus, Bl.;
Llitandra Buchanani, H:l.Uier f., C, cir,hos-.z. Radlk., C. jlavidiJIora, Hallkr f., C. gracilis,
Hallier f •• C. landolplticides, Hallier f., C. te.ptant,~a~ Hal1ier f. (vix I), C:7nyrianiM, Pierre,
C. SdJ'l!JdnjuYlltii, St.apf, C. 'lIisdjlua, K. Sch.; Cylindropsis parvifolia, Pierre, C. tOC'JIana,
Hamer r., C. fVa/soniana, Hallier f.; LandoljJlzia ir.-acleala, Dew., L. capensis, Oliv., L. C1'a.rsipes,
K. Sell" L. EminiantJ, Hallier r., L. gumm~lerat K. Sch., L. Henri'lu(s;ana, H allier f., L. ludda,
K:. Stb., L. odmuca, K. Sch., l.. reticulata. Hallier roo L. ·scamkn.r, Didr.; Willotlglabeia apiculata,
Miq., W. grandijlQra, Dyer, W.ja1.Janica, lH.,}V.lmuijlora, Dyer.
 ADDENDA-APOCYNACEAE 985
lateral strands are small and often show concentric structure; in other memberS
of the Order again there is only a single vascular bundle in the petiole. Pierre
attributes great systematic value to the nature of the principal vascular bundle,
according as it forms (a) a tube which is either closed or only provided with a
very narrow groove on its upper side, or (b) a mass of wood and bast which has
a wide groove on its upper surface. Although Pierre goes too far in this respect,
Hallier likewise describes a closed or b'!lrely open vascular tube as characteristic
of the genus.Clitandra, and a grooved mass of wood and bast as characteristic
,.of Carpodimes.
We may next deal with the hairy covering. Uniseriate clothing hairs,
similar to those figured for Ecki~ peltata, VeIl. in the earlier part of this work
(p. 530), have been observed by Hallier o.lso in Micrechites polyantha, Miq. The
uniseriate hairs fotmd in Chonemorpha megacalyx, Pierre are remarkable, since
the upper epidermis of the leaf consists of several layers near the point of inser-
tion of the hair, while the lower epidermal cells and those situated above the
veins are prolonged onto the basal portion of the hair, occasionally even forming
true cushions (Spire). According to Valeton (see also Mirabella) the glandular
shaggy hairs, already mentioned. are present in very many members of the
Order, being situated on the illterpetiolar line between the opposite leaves and
. on the inner side of the petiole.
'- Inas_ldition to the forms of excretion of oxalate of lime, previously observed
(p. 5jo), small prismatic crystals occur in eertliin Apocynaceae (e.g. species of
Aganonerion, Parabarium, Xylinabaria, &c.). An investigation'based on more
abundant specific material can alone settle how far the occurrence of solitary
crystals only, or of clustered crystals only, or of both forms side by side are
features which can be employed in generic diagnosis. 'Ve may, however,
mention that Hallier, for, instance, records only solitary crystals in the meso-
phyll in Clitandra and Landolphia, and clustered crystals in the spongy tissue
in most of the species of Melodinus and in all the species of Hunteria, and that,
according to Garcirl, oxalate of lime appears to be wanting in Apocynum. The
solitary crystals found in the mesophyll are occasionally of large size, and
in Landolphia gummitera, K. Sch. and L. crassipes, K. Sch., for example, they
occupy the whole thickness of the leaf; large idioblasts, containing ci,istered
crystals, and giving rise to transparent dots in the leaf, are found in Hunteria
ajricar.a, K. Sch. P-.nother important systematic feature is the occurrence of
solitary crystals in the epidennis of the leaf_in eertain members of the Order;
apart from the species previously mentioned in this connexion (Alstonia scholaris
and Cerbera Manghas), this character is found in all the species of Hunt~ia
(according to HaUier)~ in Cerbera Tanghin, Hook. (according to Garcin), and in
Gonioma [(amassi (according to Gerhard).
To the section dealing with the non-articulated latidferous tubes we may
add the following details. In the first place we may note that Spire mentions
the occurrence of anastomoses in the laticiferous system of the axis, the leaf,
the floral parts and the fruits of certain Apocynaceae. He observed ollly two
examples of-anastomosis in·the axis, viz. in the laticiferous tubes situated lfl the
pericycle of t· Alstonia H oedtii, F. et B.,' and in the case figured by him in
Fig. I, pI. XXXVI (C'ercocoma macrantha~ Teijsm_ et Binn.). The firSt case
did not admit of verification; the second, judging by the figure, is merely an
instance of apparent anastomosis, which can be brought into relation with the
branching of one of the non-articulated laticiferous tubes. On the other hand,
Figs. I, 4, and 5 on pI. XXXIV, which refer to the laticiferous tubes in the
petrus of RhyncJwdia Capusii, Pierre, Aganosw..a marginata, Don and Parabari'um
Verneti, Pierre, appear to show true anastomoses. According to Spire, however,
anastomOSES in the floral organs and fruits are likewise of rare occurrence.
Leuconotis eugeni/olius, DC. and t Ochrosia glomerata' are.given as examples
 ADDENDA--APOCYNACEAE
of the occurrence of anastomoses in the leaf. A careful reinvestigation of these
cases of anastomosis is very much to be desired. With reference to the- distri..
bution of the laticiferous tubes in the axis we may add that they occur abun,-
dantly also in the pericyc1e. Laticiferous tubes running freely in the mesophyll
have been observed in the following additional species: in Acokanthera sputa ..
biUs, Hook. bv Garcin; in Chilocarpus denudatus, Bl., Dipladenia atropUrp"'t;a,
• Miill. Arg.,' Parabarium latitolium, Pierre, and P. Tournieri, Pierre by Spire 1;
in Hunteria ambiens, Hallier f., H. pleiocarpa, Hallier f., H. pycnantha, K. Sch.,
Lepiniopsis ternatensis, Val., Lemonotis anceps, Jacq., and L. eugenitolius, DC.
by Hallier. For the distribution of the laticiferous tubes in the petiole and in
the veins of the leaf, see especially Spire, loco cit., p. 155. The diameter of the
laticiferous elements varies with the species and with the tissue in which they
occur. The largest diameter is 50 p. (laticiferous tubes in the pith of Heligme
buruensis, Teijsm. et Binn.), the smallest 5-1 p. (cortex of Landolphia- Ki,kii,
Dyer). The secretion is not always white, but may be bright yellow (Leuco1Jotis
eugenifotius, DC.) or pale red (Parabarium, Parameria) or greenish (T1'achelo-
spermum jasminoides, Lem.). Starch-grains were I observed by Molisch in the
latex only in Nerium Oleander, L. and Allamanda Schottii, Pohl j the same
authority found small crystalloids in the latex of the oleander, and indican in
the latex (as well as in the mesophyll) of Echites religiosa, Teijsm. et Binn. The
walls of the laticiferous tubes for the most part consist of cellulose, but are
occasionally suberized.
The previous list of Apocynaceous genera, in'which non-articulated laticiferou!)
tubes and intraxylary phloem have beo;:n sho'wn to occur (see footnote on p. 531)
may be amplified by the addition of the following genera on the basis of Carcin's,
HaJlier's, and Spire's investigations: A cok_an thera , A gan ontlrion , Amalocalyx,
Bousigonia, Carpodinus, Chilocarpus, Chonemorpha, Clitandl'a, C,asp-idospermum,
Oylindropsis. Ecdysanthera, Hunter'ia, Ltlpiniopsis, Leuconotis, Micrechites, NoueUea,
Pa,abanum, Rhazya, Winchia and Xytinabaria. Spire records latidferous tubes
also in species of Aganosma, Camera,ia, Holan'kena. l(ickxia, Roupell'ia, Vallaris
and Voact:nga (under Orchipeda). We m,ay add here that according to Spire Blonders
statemE':nt (Connessie, in Les nouv. remcdes, 1887, p. 427) as to the occurrence of
transverse walls in the laticiferous tubes of Holarrhena antidysenterica. Wa.ll. is
incorrect.
In addition to the non-articulated laticiferous tubes other types of internal
secretory organs (viz. secretory cells with contents resembling latex, mucilage-
cells and mucilage-cavities) are found in certain members of the Order. Secre-
tory cells were already previously (p. 531) mentioned as occurring in ASPido-
sperma Quebracho, Schlecht., and Geissospermum Vellosii, Peck. According to
Spire the following elements belong to the same category: (a) cells filled with
granular contents and forming a complete layer below the palisade tissue in
Cleghornia cymosa, Wight (= Baissea acuminata, Benth.), Echites ,cligiosaf
Teijsm. et Binn., Kopsia fruticosa, DC., and Tabernaemontana sphaerocarpa, Bl. ;
(b) the isodiametric secretory cells found in the floral organs in species ofAganosma
and Melodinus; and (c) the elongated secretory cells present in the fruits of
Melodin'us and Tabernaemontana. Garcin observed mucilage-cells in Apocynum
venetum, L. (pith, bast, and primary cortex), Carissa Ard!iina, Lam. (primary
cortex and spongy tissue), C. Carandas, L. (subepiaelmal ceil-layer of the
cortex), C. ovata, R. Br. (primary cortex, peri cycle and tissue of the leaf), and
Ichnocarpus trutescens, R. Br. (pericycle). According to Spire -the mucilage-
cavities are lysigenous in origin, although surrounded by an epithelium. and
are occasionally of very considerable width. They are found at the margin of
the pith or in the intraxylary phloem in Bousigonia angustijolia, Pierre, Chone-
---~----------,.-- ._-
1 Spire's remaining statements (loc. cit., p. 156) concerning this point are not clearly presentee.
 ADDE/jDA-APOCYNACEAE
morpha Grandieriana, Pierre, C. megacalyx, Pierre, and M icrechitls Jacqueti,
Pierre, and in the pith, outer portion of the bast, and wood, as well as in the
parenc:hyma of the leaf in Rhynchodia Capusii, Pierre and PoUsia cantonen#s,
Hook. et Am.
Greshoff has shown that th~'demonstration of the presence or absence of
alkaloids in a member of the Apocynaceae by means of microchemical tests
may also be of importance in the solution of systematic problems "(see Hallier,
loco cit ).
3. STRUCTURE OF THE AXIS. To the statements on the structure of the
wood we may add that the wood-prosenchyma bears bordered pits also in
Aspidosperma (Hansen).
As regards the structure of the cortex we may first note that the cork
develops in the epidermis also in species of A cokanthera, A msonia, M elodinus,
and Trachelospermum, in the subepidennallayer of cells in species of Carissa,
Ecdysanthe t i, Melodinus, Micrechites, Parabarium, and XyUnaba,ia, and in
a slightly deeper layer of cells in certain species of Para barium and X ylinabaria
(Garcin, Spire). The phelloderm is occasionally sclerosed (Alstonia cons#ricla,
F. v. M.) or its cells are filled with solitary crystals (Alstonia sclwlaris, R. Br.).
The primary cortex frequently includes stone-cells or groups of these elements ;
in some cases there is even a ring of stone-cells (e.g. in Micrechites Jacqueti,
Pierre, according to Spire). In Forsteronia corymbosa, Mey. an annular zone
containing clustered crystals is found in the cortex. The groups of pericyclic
bast-fibres are frequently arranged in one or more annular zones; in other cases
>.they lie singly, or in small scattered groups in a broad pericycle, or unite to form
l.l ring. Side by side with the bast-fibres, the walls of which are often not
lignified, the pericycle contains crystal-cells, stone-cells and laticiferous tubes.
Spire's statement as to the absence of pericyclic fibrous cells in Amalocalyx
microlobus, Pierre is probably based on the investigation of a branch of slight
thickness, since the differentiation of bast-fibres in the pericycle of the Apocy-
naceae often takes place only at a late stage; a reinvestigation is required to
determine whether the ring of stone-cells, recorded by the same author in the
pericyc1e of Bousigonia mekongensis, Pierre, really belongs to the Ipericycle.
Hard bast seems to occur more frequently in the secondary phloem than was
formerly supposed, judging by the statements of Garcin with reference to
Alstonia scholaris and A. constricta, and of Spire with reference to Bousigo-tUt
mekongensis. In Aganonerium polymorphum, Pierre the secondary bast con-
tains a ring of stone-cells (Spire). The bast-fibres of the Quebracho-bark,
which either occur embedded in groups of stone-cells or lie isolated, are quite
specially distinguished by the fact that each fibre is completely enveloped by
,.a one-layered sheath of crystal-cells, each of which contains a solitary crystal
tHansen, Moller, Garcin, &c.); according to Moller the only other known
example of this type of structure is found in 'Cortex Chinae albae de Payta.'
The mode of development of the anomalous structure of the axis found in
Condy1ocarpon sp._.(see_Fig. 123_on p. 533) has been investigated by Leisering ;
according to him the enclosure of the phloem·groups is not due to the activity
of a new cambial arc (as in Strychnos), but to proliferation of the tissue of the
wood. With referenQe,tQ the interxylary phloem of Lyonsia straminea, the same
author states that the cambium at first merely produces-groups of thin-walled
parenchyma on its i,'mer side, and that the transformation of th~se groups into
leptome takes place at a later stage l ,

] The statement (in Perro~. Tissu crible. These, P.ri3, 18Q9. p. 186) to the effect that illterxylar,
phloem occurs in Apocynum (~binttm and Willoughbeiafirma is incorrect, cf.-Scott and BrebDet,
10 Ann. of Bot., 189(, p. 283 ef&eq.; these authors only mention the secondary transformation of the
intraxylary phloem-bundles into inversely orientated medullary vascalar bundles.
 ADDENDA--APOCY~CEAE

For the structure of the spiny branches of Carissa, see Garcin and Lothelier,
n. ce.
Literature: Hansen, QuebrachQ·Rinde, Berlin, 188q,. 204 pp., 3 Tab.-Hohnel, Gerberinden.
Berlin, 1880, p. 103 et seq.-MoUer, Westind. Btlcbsho17..t.AspUiC1s1enlla Vargasii, DC.), in Dingler,
Polytechn. Joum., ::138, 188o, pp. 59-63.-Costantin, TtPs aero et sout., Ann. sc. nat., ser. 6. t. ni,
1883. po J,fI.-Gal'ciD, Apocynacees, These, Lyon, J889, ::156 pp., ~ pl.-LamounflUe, Liber interne,
Ann. ae. nat.tier. '1, t. xi, 1890, pp. 257, ::I58.-Lotbelier, Epines, These, Paris, 1893, p. 18.-[;1'0-
guini, StomJ, Atti 1st. bot. Pavia, 1894.]-Bormann, Cerbera ()Va/a, Diss., Erlangen, t895. 30 pp.-
ValetO!l, Oc/mula, Ann. Jardin Baitenzorg. xii, 1895, pp. z:a6-g.-[Mirabella., Colleteri, Contribtrz.
bt. bot. Palermo, ii, 1897, p. IS et seq.; abatr. in Just, 1897, i. p. 513.]-Pierre, Landolphiees, &11.
Soc. UnD. de Paris, 1898, n. 5, p. 33 et seq. and n. 1 r, p. 89 et seq.-Grelot, Caoutchoucs et Gutta·
Percha, These, Paris, 1899. p. 159 et seq.-HaIlier, Kautschuklianen, Jahrb. Hamburg. wiss. Anst.,
xvii, 3. Beiheft, 1899. Hamburg, 1900, l16 pp.-Leisering, Interxyl. Leptom, Dis!!., Berlin, 1899,
pp. 1~ and 37.-Bannetzky, Faisc. bicoll., Ann. sc. nat., ser. 8, t. xii, 1900, pp. 289--92.-Gamper,
Angolturarioden, Diss., ZUrich, 1900, p. 6s.-Payrau, Strophanthus, These, Paris, 1900, pp. 37 and
47 et seq.-Tbomaa, Feuilles sout., These, Paris, 1900.-Molisch, Milchsaft u. Schleimsaft, 1901,
pp. 30, 17 and 71.-Petersen, Vedanatomi, 1901, p. 84.-Bargagli-Petrucci. Legnami, Malpighia,
190::1, p. 363 et seq. (Dyera, Ce,,6era).-Pentig, ~iante ac~~le, Malpighia, 1902, p. 450 (Ta"~rnae­
mcmlana}.-Gerhard, Hlattanat. v. Gew. d. Knysnawaldes, D1SS., Basel, 1901, pp. 26,27 (Gcnwma).
-QwuljeT, Anal. Bouw, etc., Natuurkund. Verbandel. liaarlern, iii,S, 1903 (Cerbe,.a Odolkm).-
Achner, Falsche Chinarinden, Diss., Bern, 190-j., p. 64 et seq.-Col, Faisceallx, Ann. !Ie. nat., ser. 8,
t. xx, 1904, pp. r9l, 193.-Areschoug, Trap. viixt. bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2,1905,
pp. ~6, a., (Al.rtMia), pp. a9,30 (Ckz1<?Car_tNs).-H a beTlandt, Lichtsilmesorg&lle, '905, p. 6S and
Ta~ i.-Kniep, Milchsaft. "-lora, 1905. pp. 166, 167.-Mayus, Milcbrohr. in den Bl., Beih. L. bot.
Centrelbl, xviii. Abt. I, 1905, pp. 279, l80.-Spire, Apocynacees, These, Paris. 1905, 186 pp. (also
in Perrot, Tranux, ii).-Piccioli, Legnami. Bull. Siena, 1906, p. 146.-[Fodnrther litera.tnre, see
po u6g.]

ASCLEPIADEAE (pp. 534-537).

I. REVIEW OF THE ANATOMICAL FEATURES. To the enumeration of special
anatomical features (po 534) we may add the occurrence of intercellular spaces
containing mucilage (lforrenia odorata, Lind!.), of secretory cells in the meso-
phyll (Solenostemma Arghel, Heyne), and of long prismatic crystal3 of oxalate
of lime (At! enabea). Among the anomalies in the structure of the wood we may
include the occurrence of islands of soft bast in the wood of the root of Asclepias
syriaca and M orrenia brachystephana, Griseb.
2. STRUCTURE OF THE LEAF. Stomata having 4-5 neighbouring cells are
{bQnd in Vincetoxicum palustre, Gray (Kearny), and stomata with 2-5 neigh-
bouring cells in Solenostemma A_rghel (Tschirch). The mesophyll in M o"enia
bt-achystephana contains fat·bodies (Keller).
The prismatic crystals mentioned above appear to be of the nature of
styloids. and are found in the bast of the root of M enabea venenata, Baill.~
while the tissue of the leaf in this plant contains clustered crystals. The
clustered crystals found in the leaf of Solenostemma Arghel have the shape of
a sphaerite. The non-articulated laticiferous tubes, which side by side with
intraxylary phloem 1 have recently also been demonstrated in Conchophyllum,
Menabea, and Morrenia, penetrate into the mesophyll also in Periptoca graeca
and Hoya carnosa (Kniep, Lehmann). Intercellular spaces containing mucilage
were observed by Keller in the primary cortex of the terresbial roots of M orreni-a
odorata; they are situated on the outer side of the endodermis and attain a con-
:riderable size. The secretory cells found in the mesophyll in Solenostemma
Arghel are either rounded or elongated; they have suberized w_alls and contain
a yellow secretipn.
3. STRUCTURE OF THE AXIS. The wood contains vessels with simple
perforations and wood-prosenchyma bearing bordered pits in Gymnema also.
1 The petiole of S()/mI)slemma Ar~he/ likewise contains laticiferous tubes and an a.rc-shaped vas-
cular bundle showing bicollateralsttucture.
 ADDENfJA-ASCLEPIADEAE

The cork arises in a subepidermal position in Gymnema. Chambered crystal.

fibres containing clustered crystals are found in the secondary bast in Gymnema,
and similar elements with solitary crystals in the secondary bast in Periploca
graeca.
No details are known as to· the mode of development of the interxylary
phloem in the wood of the root of M orrenia brachystephana (see 'Hantzschel,
loco cit.). The differentiation of the isiands of soft bast, observed by Kny in the
wood of the root of Asclepias syriaca, only takes place at a late stage.
Literature: Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, J883, p. 14o.-Koy,
Bot. Wandtafeln, vi. Abt., Text, 18>1<1, pp. ;;137, 238.-Leitge~ Spharite, Mitteil. bot. lost. Graz;
Heft. l, 1888, p. 314- et seq.-Keller,Luftwurzeln einiger Dikotylen, Diss., Heidelberg, 1889, pp. 8-u.
-Lamouoette, Liber interne, Ann. sc. fiat., ser. 7. t. xi, 1890, l'P. 255-7.-Wijnaeodts Francken,
Sclereideu, Diss., Utrecht, 1890, pp. 57, s8.-Latour, Sene, These, Montpellier, 1894, p. :15 et seq.
(Argbel); lee also Tschirch and Oesterle, Anat. Atlas, i, 1895, p. 26 and Tab. 7.-[Tognini, Stomi,
Aui 1st. bot. Pavia, 1894.]-Busch, Gyml1ema sylvest,.e, etc., DiS!l., Erlangeu, 1895. pp. '4 and SI
et seq.-Kal'iten, Epiphytenformen, Ann. Jardin Buitenzorg, xii, 1895. p. 15 .. et seq.-Hlintzachel,
MtwTenialmuhystephana, Diss., Erlangen, 1895, pp. 12 and· 35 et seq.-Lebrnann, PeripkJta ~a,
Archiv d. Pharm., :335. 1897,'p. 159 et seq.-[Mirabella, Colleteri, Contrib\l%. lst. bot. Pale:mo, ii,
1897, p. 15 et seq.; abstr. in Just, 1897, i, p. 513.]-Schubert, Parenchymscheiden, Bot. Cent&albl.,
1897, iv, p. 2o.-[Tassi, Hqya carnosa, liull. Lab. Otto bot. Siena, 1898, pp. 151-7, :I tab.; abstr.
in Just, 1899, ii, p. 245.]-Grelot, Caoutchoucs et Gutta Percha, These, Paris, 1899, p. 163 et seq
-Leisering, Interxyllires Leptom, Diss., Berlin, 1899, p. 38.-Kearny, in CContrib. U. S. Nat. Herb.,
v, n. !;, 1900, p. 503 (Vincelqx;cum).-Perrot, Menabea, Comptes rendus, Paris, cxxxiv, 190 3 , p. 305;
llee also 100m. de bot., 1903, pp. 114, Il5 .....:..Col. Falsc. bieoll., Ann. lie. nat., ser. 8, t. xx, 1904,
pp. 19:J-4.-Paoli, Eterofillia, Nuovo Giom. bot. ltal., xi, 1904-, p. 319 (Hemidesmus).-SUssengutb,
Bebaanmgsverh. der Wlirzb. Musehelkalkptl., Diss., \Viirzburg, 1904, p. 44.-Kniep, MilchrObren,
Flora, 1905, pp. 166. 167.-Mayus, Milchrohr. in den BI., Beih. Z. bot. Centralbl., xviii, Abt. J, 1905.
pp. 380. 381.-Sarton, Anat. d. pl. affine" Ann. sc. nat., ser. 9, t. ii, 1905, pp. 99, 100 (V;nce/oxicu",).
-[Hurrier et Perrot, Ginseng, Buli. se. pharmacol., 1906, p. 665 (TJ'lqphcra).]-Holtennann, EiD-
flus des Klimas, etc., 1907. p. 136 (Hoya).

LOGANIACEAE (pp. 538-547).

I. REVIEW OF THE ANATOMICAL FEATURES. Oxalate of lime is.'deposited
in the form of small needles in Spigelia also, while in the newly established genus
Crateriphytum, Scheff. MS. ed. Koorders, as in its nearest ally, the genus
Couthovia, it occurs in the form of crystal·sand. In the stem of Gelsemium
semperoirens and ' Spigelia dichotoma ' tlie in traxylary phloem becomes trau-
fonned into medullary vascular bundles, showing inverse orientation. Peculiar
extrafloral nectaries taking the form of small pits and provided with a secretory
palisade layer are fOlUld in F agraea.
2. STRUCTURE otTHE LEAF. A mucilage-layer situated beneath the upper
epidermis is stated by Morelle to occur also in M ostuea ( gabonica: M. Pervilleana,
M-itrasacme pilosa and M. prolifera; he likewise mentions the occurrence of
cells having partially gelatinized walls in the mesophyU of the two species of
Mitrasacme just named. In Mitrasacme polymorpha (M. cinerascens) Morelle
observed a-layer of mucilage, though not so strongly developed, also above the
lower epidermis.
The short unicellular hairs found in certain species of ~l1ostuea and Mitra-
sacme are true papillose·hairs.
The extrafloral nectaries,which occur in Fagraea crassifolia,Bl., F. elliptica,
Roxb.~ F. iml>erialis, Miq., F. lanceolata, F. lito1alis, Bl., F. monant}ja, Miq.,
F. obovata, Wall., and F. peregrina, Bl., have been examined espi:dally by
Zimmennann (see also Poulsen and Areschoug); they are found on both surfaces
of the leaf, at the base of the petiole (here two or more in mtmber) and on the
stem below the stipules. They constitute lobed pit-like depressions which
vary in shape and open to the exterior by means of a small aperture. The
 A DDENDA-LOGANIA CEA £

nectaries situated on the surface of the leaf show a short canal leading into
a cavity, which is placed paranel to the surface of the lamil13" and from which
lateral ~anals arise; in the case of the nectaries occurring on the petiole and stem
the vertical canal is longer, but its branches ate less numerous. The wall of these
pits is formed by a layer of secretory palisade-cells which have no cuticle, while
the layers of cells adjoining the palisade·Jayer are rich in clustered crystals of
oxalate of lime. The starting·poi"rlt in the development of the nectary is
a single epidermal cell, which increases in size, and becomes elongated in a
direction at right angles to the surface of the organ. The cells of the epidermis
and of the inner tissues bordering on this inItial cell likewise undergo elongation
and division, and thus give rise to the epithelium. The main cavity of the gland
is formed by the separation of the epithelial cells from the epidermal cell, which
initiates the formation of the entire glandular mechanism, and may still occa·
sionally be found in the cavity of the mature gland; the lobes are,formed
solely by the separation of the epithelial cells from one another.
The occurrence of oxalate of lime in the fo,"m of small needles in Spigelia
(according to Morelle) and of crystal·sand in Craterispermum moluccanum,
Scheff. (Koorders) has already been referred to above.
3. STRUCTURE OF THE AXIS. On the outer side of the intraxylary soft bast
in the older parts of the stem and in the rhizomes- of Gelsemium sempervirens
a cambium is formed, which not only produces phloem internally, but at some
points also gives rise to wood externally (Bolling). In this way the groups of
intraxylary soft bast become transformed into inversely orientated vascular
bundles. Such inversely orientated bundles are stated by Morelle to occur
also at the margin of the pith in ' Spigelia tf,ichotoma.' ,
As regards the structure of the cortex we may add that the cork in Spigelia
Sellowiana, Cham. et Schlecht. arises in the subepidermal layer, and that in cer-
tain xerophilous species of Spigelia which have reduced leaves (e.g. S.linarioides
and S. pulchella), subepidermal groups of sclerenchyma (visible to the naked
eye as ribs) and palisade tissue are developed in the primary cortex (Morelle).
Note. According to Morelle noteworthy features in the structure of the root
are the large lacunae in the primary cortex of Mitrasacme montana and species of
Mitre o/a , and the peri cyclic development of the cork in Mitrasacme cineyascens
(= M. polymot'pha).
Literature: Rothrock, Internal cambium-ring in Gelsemium, Proceed. Acad. sc. Philadelphia,188S,
pp. :12, 23.-Garcin, Apocynacees, These, Lyon, 1889, p. 206 (the plant described in this paper as·
Gelsemiu,." semjet'Virens contains laticiferous tubes and is therefore incorrectly determined).-Wij-
naendts Francken, Sclereiden, Diss., Utrecht) 1890, pp. 55, S6.-Ilschert, SIrycJmos oTieute, Diss.,
Erlangen, 1894, ' . pp.-Elfstrand, Studier Mv. Alkaloid. 10kal. etc., Upsala Univers. Arsskr., 1895,
126 pp., 2 Tab.-Went, Haft- u. Nahrwurzeln, Ann. Jardin Buitenzorg, xii, 1895, p. 6:1 (FagrtUa).-
[Dohme, in Druggists' Cire. and Chem. Gazette, 1897. n. 7 (Gelsemium); abstr. in Just, 1897. ii,
p. Il.)-Matteucci, Placcbe sugherolioe, Nuovo Giorn. bot. Ital., 1897, p. 235.-Poulsen, ExtrntIor.
Nektar.-Stud., Vidensk. Medde1els. Kjllbenhavn, 1897, pp. 360-4 and Tab. iH.-[Sayre, Celsemium,
Americ. loum. of Pharm., 1897, n. I; abstr. in Just, 1897, ii, p. i-6.]-[Serberling, Gelsemium,
Americ. Joum. of Pharm., 1898, n. 8; abstr. in Just, 1898, ii, p. 5I.]-Thompson, Internal phloem
in Gelsemium, Contrib. bot. Labor. Univ. of Pennsylvania, ii, 1898, pp. 41-53.-Leisering, Inter·
xyliires Leptom, Diss., Berlin. 1899, pp. 18-23.-BOlling, Alkaloidhalt. PH., Diss., Erlangen, 1900,
pp. 33-5 (Gelsemium).-Gamper, Angostnrarinden, Diss., Ziirich, 1900, p. 51 et seq. (Strydz1WS).-
Gerhard, Blattanat. v. Gew. d. Knysnawaldes, Diss., Ba.sel, 1902, pp. 27, 28 (Nuxia).-Koorders,
Craleriphylum, Bull. Instit. bot. de ~uitenzorg, n. xvi, 1902, sep. copy. p. 3.-Poulsen, Bladkirtl.
hos FagrtUa obovata, Vidensk. Meddelels. Kjgbenhavn, 190:.1, pp. 242-4.-Zimmermann, Exttafiorale
Neklar. einiger Fagraea-Arten, Ann. Jardin bot. Buitenzorg, xviii, 1902, pp. t-7.-Achner, Falsche
Chinarinden, Diss., Bern, 1904, p. 76 et seq.-Morelle, Histol. compo des. Gelsemiees et Spigcm~,
These, Paris, 19Of, 16:.1 pp.; also in Perrot, Travaux, ii.-Areschoug, Trop. viixt. bladbyggn., Sv.
Vet. Akad. Handl., 39, n. 2, 1905, pp. 140-2 and Tab. xix-xxi.-Holm,in Americ. Joum. oCPharm.,
J~, p. 553 et seq. and 1907, p. 51 et seq. (Spt"ge/ia marilandica).-Holtermann, EinHuss d. Klimas,
etc., 190 7, p. 13f (Jt'agraea). -
 ADDENDA 991

GENTIANEAE (pp. 548-550).

Our knowledge of the anatomy of the Gentianeae has recently been quite
considerably enlarged, especially by Perrot's investigations l , which deal with
the structure of the leaf, stem, and root. The two suborders Gentianoideae
and Menyanthoideae differ anatomically not only in the presence (Gentianoideae)
or absence'(Menyanthoideae) ofintraxylary phloem, but aIsoin other anatomical
characters. In the Gentianoideae the sieve-tubes both of the outer and intra-
·xylary soft bast (when present, of the interxylary bast as well) almost invariably
form small groups of cells wi th narrow lumina, these gr_oups in a transverse section
being approximately of the same size as one of the neighbouring cells of the phloem-
parenchyma; in the Menyanthoideae, on the other hand, the sieve-tubes have
wide lumina, and are irregularly distributed in the tissue of the soft bast. In the
Gentianoideae the formation of the pair of guard-cells is quite generally preceded
by two or three divisions in the dermatogen-cell, ~hile in the Menyanthoideae
the mother-cell of the guard-cells originates with the appearance of the first
division-wall in the dermatogen-cell. Hydathodes, provided with an epithema
and wat~r-pores~ occur only in the Menyanthoideac, and are wanting in the Gen-
tianoideae. The root in the Gentianoideae has a diarch vascular system, while
in the Menyanthoideae the vascular system of the root is polyarch (S-9-arch);
secondary division-walls, which are radial or more rarely tangential, occur in
the cells ~f the endodermis only in the Gentianoideae. To the earlier statements
on the structure of the wood (see the general diagnosis of the Order) there is
nothing to add. The sen tence ' a special type of stoma does not occur' must be
modified in the sense that distinct subsidiary cells are rarely found. In opposi..
tion to the earlier statements it has been shown that oxalate of lime is not
absent in this Order. In most of the genera. of the Gentianoideae it has
been observed as small crystals, taking the shape of prisms, octahedra, needles
of varying thickness, or gr~mules, as well as in the form of small clustered crystals.
Large numbers of the acicular crystals occasionally·lie irregularly scattered in
cells with mucilaginous contents, while large quantities of the granul~r crystals
in some cases form a crystal-sand similar to that of the Solanaceae. In the
Menyanthoideae oxalate of lime appears to be absent. Typical internal secre-
tory elements do not occur in the Gentianeae, but resinous substances are
found in the tissues of the Toot in species of Gentiana and Sweer#a, isolated
tannin-cells or rows of these elements in the Menyanthoideae, and mucilage-
cells, as well as gelatinized epidermal cells, in certain species of Gentiana. In
certain genera. of the Gentianoideae, moreover, the epidermis of the ovary
contains latex. Recent observations have also demonstrated the rare occur-
rence of clothing hairs (unicellular or uniseriate hairs, the latter composed of
a small number of cells); there are no typical glandular hairs, apart from
club-shaped bodies, which consist of a large number of cells and have hitherto
been obse!yed_onlyJn~th~_axi]§_ of the leaves in Obolaria and on the leaves in
Bartonia. -To the previous list of special features we may add: the occUlTence
of islands of soft bast and of vascular bundles in the pith (the latter in species
of Gentiana); the occurrente of cortical va5cular bundles (in certain Meny-
anthoideae); the anomalous structure of the axis found in the species of

1 These investigations extend to the following genera (in the serial order of Gilg's system) :
Exacum, Stbaea, Be/montia, Ent"co.rttmma, .Faroa, Aficrocala, Curtia, lVeurotlteca, GenieS/CillO",
Ciundia, Sallbatia, Lapithea, Erythnua, Chlora, Schi,zziella, Camcora, Bartom'a, Obdlan'a, Chiren:'a,
Orpkium. Cra'lo/urdia, Genlz'ana, IXllnthus. Pleurogy1t£, Sweertia, llalenia, Hockim'a, Lisianthus,
Eustoflla, Zyg-ostig-ma, Zonant/ws, Ru.rbyar..tlws, Pnpusa, Se1Jtua, ScJm/lflia. C(lutl)u/tea, Purdiealt-
11m!, Lagenanlhus, Deianira, Lehmanniella, Voyriella, Ldphaimosj Nephrophyllidi14m, Mmyanl}us,
Villarsia, Lil1lna1zthemum, Lipa1·Ophyl!um.
 ADDENDA-GENTIANEAE

Limnanthemum belonging to the section N ympkaeanthe and recalling the structure

of a Monocotyledonous stem; the peculiar discoid groups of small cells occurring
on the lower side of the leaf in species of Limnanthemum and V illarsia; and
the development of interxylary phloem in the axis in Chironia, Crawturdia,
] xattthus and Orphium, and in the root in many members of the Order.
The following details may be added regardmg the STRUCTURE OF THE LEAF
in the two suborders. In the Gentianoideae the upper epidermal cells are
provided with straight or slightly undulated lateral Walls, while the lower
epidermal cells have strongly curved walls. Papillae are developed in some
cases, especially at the margin of the leaf; the cuticle is occasionally striated.
The development of the stomata was described above; they are for the most
part confined to the lower side of the leaf, although in some cases present on
both sides, and are surrounded by 3-4 neighbouring celis. The mesophyll is
lacunar and either bifacial or homogeneous in structure. The epidermal cells
of the Menyanthoideae have rather straight lateral walls. The stomata; the
mode -of development of which has likewise been considered above, occur on
both sides of the leaf, but are developed only on the upper side in the floating
leaves; the guard-cells are surrounded by 4-6 neighbouring cells. The
structure of the leaf in this suborder is markedly bifadal, and the spongy tissue
is very lacunar. In none of the Gentianeae are, the stomata ever strongly
depressed or elevated. Large guard-cells are found, for example, in Gentiana
pyrenaica, L., small ones in Crawfurdia iaponica, S. et Z. Stomata have also
been observed in the typical saprophytes (in Cotylanthera by Figdor, irt Obolaria
by Holm and prior to that by Chatin, in Leiphaimos and V oyria by Svedelius
and Porsch). The veins of the leaf rarely contain fibrous cells (Deianira,
Senaea coerulea, Taub.), the latter being found in species in which mechanical
elements are also developed in the pericycle of the axis. In the Menyanthoideae
enlarged terminal tracheids occur at the ends of the veins.
The occurrence and mode of deposition of oxalate of lime have already
been referred to above!. The quantity of the salt excreted varies very con-
siderably. It is met with in the ground tissue and occasionally,in the bast
and epidermis as well. Mucilage-cells containing acicular crystals have been
observed, for example, in Gentiana cruciata, L., G.lutea, L. and G. Saponaria, L.,
while crystal-sand has been found in species of Gentiana, belonging to the
section Chondrophylla, and, as it seems, also in Sabbatia; small clustered
crystals occur in species of Gentiana, Lehmanniella and Purdieanthus. There
can be little doubt that the crystalline deposits do not possess that degree of
systematic importance which Kusnezow attributes to the occurrence of oxalate
of lime in the mesophyll in the genus Gentiana; according to him oxalate of
lime is present in all the sections of the subgenus Eugentiana with the exc~ption
of the section Cyclostigma, while emphasis is laid on its absence In the section
just named and in the subgenus Gentianella.
Perrot's statements as to the exact nature of the mucilage-cells are not
always quite clearly presented; according to him, moreover, the demonstration
of these elements in herbarium-material was attended with difficulty. Mucilage
is found mainly in the leaf and in the root. As regards the former, Perrot, in
the first place, records slightly gelatinized cells in the mesophyll in Belmontia
cordata, E. Mey., and leaves containing mucilage in Canscora, Gentiana Parryi,
Engelm. and G. tri{iora, Pallas (sect. Pneumonanthe), and species of Gentiana

1 Perrot and other authorities have demonstrated oxalate of lime in species of the fcllowing
genera: Exacum, Stbaea, Belmolzlia, Enicoslemma, Faroa, Microca/a, Neuyolhec4, Sa/Jbatia,
Erylnraea, Chiara, Sdtinzitlla, Crawfurdia, Genlia11a, IzaTt/nus, Sweerlia, Halenia, l?usbyanl;"us,
Sc/Jultesia, Purdieanlhus, Lagena~llUs, Deianira, Lehmanniella. Perrot dislinctly mentions the
absence of oxalate of lime in CCtitoltoea, Prepusa, Smaea and in the Menyanthoideae.
 ADDENDA-GENTIANEAE 993
belonging to the section Stenogyne. Epidermal cells of the leaf having mud·
laginous inner walls occur in Gentiana bavarica, L., G. excisa, Presl and G.
verna, L.~ while in G. pyrenaica, L., a complete layer of mucilage, formed by
the gelatinization of the inner walls of the epidermal cells and the walls of the
adjoining mesophyll-cells, is found beneath the upper epidermis. A similar
layer of mucilage is present also beneath the epidermis of the stem in Gentiana
saXQsa, Forst. In the species of Gentiana belonging to the section Chondro-
phylla, whicp are endemic in the region of the Himalayas an~ on the high
mountains of Western China, the tissue of the leaf is gelatinized to quite an
exceptional extent. The process of gelatinization in this case only affects the
middle portion of the lamina and does not extend to the margin of the leaf,
which has a horny appearance characterbtic of these species; the extent of
gelatinization is such, that transverse sections mounted in water as a rule
show nothing but mucilage, the cuticle of the upper and lower epidermis, and
the vessels. The thick roots of the species of Swecrtia belonging to the section
ophelia also contain a little mucilage, while the cOIk·Hke tissue found on the
surface of the roots in N ephrophyUidium crista-gatti, Gilg, includes a mucila-
ginous substance.
The cartilaginous margins above mentioned as characteristic of the leaves
in the species of Gentiana, sect. Chondropkylla, vary in breadth, and either
contain a tissue with cartilaginous walls (G. quadrijaria, Bl.) or consist merely
of the thick-walled upper and lower epidermis (G. albescens, Franch.). Thick
cell·walls having the same cartilaginous texture are met with also in the cortical
tissue of the stem (primary cortex and bast) in G. pap~llosa, Franch., and other
species belonging to the same group.
Perrot mentions the occurrence of (a) resinous substances in the cortical
tissue of the root in Gentiana lutea, L. and G. purpurea, L., as WE:ll as in Sweertia
Chirata, Ham., S. Hookeri, Clarke, S. Kingii, Hook. and S. multicaulis, Don;
(b) tannin-cells in the rhizome of Nephrophyllidium and Villarsia; and (c) latex
in the epidermal cells (which are here elongated at right angles to the surface)
of the ovary of Canscora, Chironia, Chlora, Erythraea, Lisianthus, Schultesia
and Sweerlia, and to these we may, on Baillon's authority, add' Sabbatia 1.
The branched .spicular cells, which are differentiated as intern'al hairs
(p. 548), do not occur in the root. The tissue of the latter contains only un·
branched sclerotic cells, and these are not abundant. Perrot observed rather
long uni" or bicellular trichomes also in a few species of Sweertia, whose names
are not mentioned. The unicellular hairs in this case are either elongated
trichomes with a bulbous swelling at their base, or are connected with papillae
by means of transitional forms. Papillae" varying in shape from conical to
elongated-cylindrical and commonly striated, are present on the epidermis
of the stem Of leaf (here especially on the margin), or on both in many Gen-
tianoideae, e.g. in species of Curtia, Exacum, Gentiana (species belonging to
various sections, such as Pneumonanthc, Stenogync, Frigida J ]somert'a, Chon ...
d10phylla, _..Andicola .. and Amarella), Halenia, Hockinia, Ixanthus, Orphium,
PU1dieanthus, and Sweertia. Another noteworthy feature lies in the occurrence
of glandular shaggy hairs in the axils of the sepals (bracts according to Baillon's
interpretation) and foliage-leaves in Obolaria virginica, L. (Ballion, Knoblauch),
and on the leaves in Bartonia verna, Mi.ihl. and B. lanceolata, Small (Holm).
Their structure is similar to that of the glands which are situated in the depres-
sions between the corolla~lobes, and are figured by Holm. In shape they recall

1 \Ye may notice here that according to Figdor the st3.rch-grains in the stem and root of
Cotyianthera become coloured red by Iodine solution, and consequently consist of amylodextrin.
Starch-grains showing the same re3A.i:ion had previously been observed by Russow and A. Meyer in
Genliana lutea and Swurtia permnis.
SOLEREOER 3S
 ADDENDA--GE~TIANEAE

the well-known glandular shaggy hairs found in the Rubiaceae, &c., but differ
from them in the absence of a vascular bundle and in the fact that the epidermis
of the body of the gland, which is composed of numerous cells, is not developed
as a layer of palisade-cells. Glandular bodies of this type are probably more
widely distributed in the Order. Hydathodes, as already stated above,
while completely absent in the Gentianoideae, are characteristic of the Meny-
anthoideae. They consist of the termination of a vein, associated with an
epithema including a varying number of spirally thickened cells, and of a
'small group of water-pores; they ant found either on the leaf-teeth or on
the margin of edentate leaves, and occur either on the upper or lower surface.
The peculiar groups of cells previously mentioned as present in the lower
epidermis of the Jeaf occur in Limnanthemum tacunosum·, Griseb., L. Hum-
botdtianum, Griseb. and L. nymphae<r.,des, Link, as well as in V illarsia parnassi-
folia, R. Br.; they constitute disc-shaped areas, composed of cells which
differ from the ordinary epidermal cells (which contain a violet cell-sap) in
having a smaller polygonal outline in surface-view and tanniniferous contents.
They are the cause of the shagreen-like appeatance of the. lower surface of the
leaf, which is noticeable even to the naked eye.
In amplification of the earlier statements on the structure of the petiole
we may firstly mention,- that in the Gentianoid~e it contains an arc of wood
and bast showing bicolla.teral stracture, two small lateral bundles being occa-
sionally cut off from the ends of this arc. Among the Menyanthoideae ·the
vascular system of the petiole consists of a more' or less open arc of isolated
vascular bundles supported by fibrous cells in M enyanthes and N eph1ophyl-
lidium, while in the species of Limnanthemum belonging to the section Nym-
phaeanthe the bundles show a scattered ;arrangement like that in the stem of
a Monocotyledon, and in Villarsia lasiosperma, F. v. M. the bundles are i~olated
and arranged in two concentric arcs with their xylem-groups directed upwards.
With reference to the distribution of bicollateral vascular bundles in the
Gentianoideae we may note that Figdor and Perrot have recently demon-
strated the occurrence of intraxylary phloem also in those ge:nera, which were
not investigated by Gilg (yiz. Cotylanthera, Voyriella and Lagenanthus); more-
over the previous mention of ' Voyria' as an exception (see footnote I on p. 548)
may be cancelled. since according to Gilg the species of V oyria having con-
centric vascular bundles in their axes (referred to on p. 550) belong to the
geuus Leiphaimos. But even these sp~cies of Leiphaimos, in which the vas-
cular bundles are concentric with central xylem, as well as Leiphaimos para·
sitica, Cham. et Schlecht., in which the bundles...have strands of phloem only
on their inner side, do not constitute real exceptions to the rule; for we must
bear in mind that the concentric vascular bundles can easily be derived from
bicollaieral ones, such a derivation being justified from the biological point of
view, while the case of Leiphaimos parasitic a fInds a certain amount of analogy
in the reduction of the outer soft bast, occurring also in other Gentianoideae
with a blcollateral vascular system, although the reduction in these cases is
not so far-going. Groups of soft bast situated at a greater depth in the body
,of the pith are found in Cotylanthera ten1~is, Bl., Erythraea Centaurium, Pers.,
Ge·ntiana cruciata, L. and other species of the section Aptera, G. £xcisa, Presl,
Obolaria virginica, L. and in species of Bartonia. In certain species of Gentiana,
moreover, some of the phloem-strands situated at the margin or in the interior
Qf the pjth also include' vessels, and thus become changed into medullary
vascular bundles, as is so frequently the case in the Melastomaceile; this
feature has been observed in Gentiana Andre'Wsii, Griseb." G. bavarica, L.,
G. BU1seri, Lapeyr., G. Delavayi, Franch., G. germanica, Willd., G. lut~a, L.,
G. Pneumonanthe, L., G. p·unctata, L., G. purpurea, L., G. stylophora, Clarke.
Fibrous cells are rarely (Deianira nervosa, Cham. et Schlecht., Lisianthus
 ADDENDA-GENTIANE.4E 99S
nig1escens, Cham. et Schlecht.} found accompanying the Pmer soft bast; in
some cases the groups of intraxylary soft bast appear to be embedded in the
ring of wood owing to sclerosis of the neighbounng cells (Exacum tet,agonum,
Roxb., ETythraea CentauTium, Pers.), or the internal phloem may be supported
by a ring of sclerosed medullary celis, which extends up to the central lacuna
of the pith (Eustoma).
We may next tum our attention to the structure of the wood. Most of
the Gentianeae (i. e. apart from a few semi..saprophytic or holo.saprophytic.
Gentianoideae and the Menyanthoide.re) have a continuous ring of wood. the
inner part of which is generally (with the exception of those species, in which
the cambium produces a considerable increment of secondary wood) rich ill
vessels, while the outer part consists of mechanical fibrous tissue. Perrot
records uniseriate medullary rays also in species of Rusbyanthus, Senaea and
Zonanthus, and Figdor mentions the occurrence of reticulate perforations in
the vessels in Cotylanth.e,a tenuis, BI. For the structure of the wood in the
saprophytic species and in the Menyanthoideae, see under the discussion of the
v~cular system in the stem of these forms.
In those members of the Order which are provided with a rin~ of wood
the outer bast is reduced (especially in the parts of the axis belongIng to the
Boral region) to a few layers of parenchymatous cells, surrounding the sfnall
groups of sieve-tubes. The latter occasionally constitute the entire outer bast,
and in this case appear to be embedded in niches in the xylem ..t~, which
extends right up to the endodermis (species of Chi,onia, Cntora, G~n,_ostemon,
Gentiana, NeUTo theca , Schultesia, Sweertia). Among the Gentianoideae such
small groups of sieve-tubes with narrow lumina are absent only in Rusbyanthus,
which has isolated sieve-tubes (Perrot).
The primary cortex in the Gentianoideae is more or less lacunar, and
occasionally contains mucilage; in other cases it is compressed, and consists
of a tissue resembling horn-bast. In the Menyanthoideae large air-canals
interrupted by transverse diaphragms, are present both in the stem and rhizome.
The stems of the Gentianoideae are very commonly provided with four ribs or
wings, which are either formed solely by two epidermal layers (Sebaea oVata, R.
Br.) or contain collenchymatous tissue (5. albens, R.Br., S. aureil,R.Br.). There
is often a distinct endodermis, provided with Caspary's dots j the endodermal
cells rarely (Crawlurdia, Gentiana pyrenaica, _L.)show secondary radial walls, and
when present they occur only in small numbers (lor 2). The pericyc1e generally
consists of thin-walled tissue, and often comprises only a single layer of cells.
According to Perrot a small number of fibrous or sclerertchymatous cells are
found in the pericycle in species of Chi,onia, Deiani,a, Lagenanthus, Lehmanni-
ella, Lisianthus, P,epusa, Purdieanthtts, Rusbyanthus, Senaea and Zonanthus,
while in some of the saprophytic Gentianoideae and certain Menyanthoideae
(see below) theopresence of strongly developed bundles of fibres or of a scleren-
chymatous ring in the pericycle compensates for the absence of mechanical
tissue in -the ~wood. ----
Interxylary phloem is developed in the axis in numerous species of
Chironia, in Craw!u,dia 'Oolttbilis, Gilg and C. japonica, S. et Z. (here according
to Perrot), Ixanth-us viscosus, Griseb. and O,phium /1utescens, E. Mey. Islands
of soft bast are much more widely distributed in the wood of the root. Accord-
ing to Perrot's investigations they are found in species of Canscora, Chironia,
Chlo,a (C. per/oliata, L. and C. se,otina, Koch), Cicendia (C. pusill-a,. Griseb.),
C,aw/u1dia, Erythrflea (species of the sections EuerythraetJ, Trichostylus and
Spica,ia), Eustoma, ~Exacum, Gentiana (species of the sections Coelanthe and
Pneumonanthe, Gentiana crinita, Frol., belonging to the section Crossopetalum),
H alenia, Sabbatia, Sweertia (species of the sections Ophelia and Eus'UJeertia,
which have small tuberous roots). The recent observations have shown that
~~2
 ADDENDA-GENTIANEAE
the islands of interxylary soft bast no doubt in all cases arise from groups of
unlignified tissue, which are produced internally by the cambium and in which
sieve~tubes' are secondarily differentiated. The islands of soft bast found
in the wood of the axis are invariably of small size and are more abundant in
the lower part of the stem as a result oi its growth in thickness than in the
upper part. In those roots in which the wood contains lignified mechanical
tissue in addition to the vessels, the islands of interxylary soft bast for the
most part form slightly larger groups, consisting of thin-walled cells and sieve-
tubes, near the centre of the root (e. g. in the soecies of Chlora, Cicendia,
Erythraea, EUstoma, Exacum, H alenia, Sabbatia); more rarely the phloem~islan~
are distributed throughout the entire mass of the wood, being in this case
arranged in lllore or less irregular rings (Chironia). In those roots in which the
body of the wood is unlignified (e. g. in Genti(ma, lutea, &c.) numerous small
islands of soft bast are found irregularly distributed throughout its entire mass.
It may still be noted that in the roots of certa.in Gentianoideae (Deianifa,
Gentiana campestris, L.) in which the xylem-mass is strongly lignified, groups
of unlignifted tissue, which do not however include any sieve-tubes, occasionally
occur near the centre of the wood; these must not be confounded with inter-
xylary phloem, _
In the herbaceous Gentianoideae the pith as'a rule becomes fistular when
the period of flowering is over. Under these circumstances the medullary
strands of rnestome (see above) likewise disappear. In Sabbatia a complete
border of corky tissue develops at the periphery of the central air·canal.
Isolated sclerenchymatous cells are of rare occurrence (Gentiana Pneumonanthe,
L.) in the pith, but the periphery of the pith is occasionally sclerosed. In
Lehmanniella ac-uminata, Gilg and Sena'ea coerulea, Taub. the entire pith is
sclerosed. The pith of the Menyanthoideae is traversed by large air-canals
like those found in the primary cortex in this group.
The following details may be mentioned regarding the fibrovascular system
of the axis in the Menyanthoideae; the latter shows anomalous structure only
in the species of Limnanthemum belonging to the section Nymphaeanthe. As
a general rule the vascular bundles are isolated. In N ephrophyllidium crista-
galli, Gilg the isolated bundles found in the axis of the inflorescence are
inserted in a ring of peri cyclic sc1erenchyma. A transverse section through
the rhizome or the axis of the inflorescence in M enyanthes tri/oliata, L. shows
ifiolated vascular bundles, which are supported both on their inner and outer
~des by strongly developed strands of fibres. The isolated bundles in the
rhizome and axis of the inflorescence of Villarsia have arcs of bast-fibres or
a ring of sclerenchyma on their outer side. In Limttanthemum nymphaeoides,
Link, the only species of the section Waldschmidtia, the rhizome contains an
almost complete ring of wood and bast without sclerenchyma, while the peduncle
and the axis of the shoot exhibit a ring of isolated vascular bundles surrounded
by a starch-sheath. On the· other hand, in the species of Limnanthemum
belonging to the section Nymphaeanthe the isolated vascular bundles, as already
stated in the earlier part of this work (p. 550), show an irregular scattered
(arrangement in a transverse section through the axis, resembling that of a
Monocotyledon, although the bundles are open. The centre of the stern in
these species is almost invariably occupied by a large' double vascular bundle,'
formed by two bundles with their'xylem-groups directed towards one another;
this large bundle may be said to constitute the stele, the remaining bundles
being regarded as cortical. The mode of arrangement of t~e vascular bundles
in the axis of the species of Lz'mnanthemum belonging to the section Nym.
,phaeanthe is doubtless an advanced adaptation to life in water. In the vascular
bundles of the Menyanthoideae a relatively large group of unlignified paren·
chyma is generally situated on the inner side of the xylem; this parenchyma
 ADDENDA--GENTIANEAE 997
does not include any primary vessels, and may be looked upon as being homo-
logous with the internal soit bast. Schizogenous intercellular canals occa-
sionally (Menyanthes trifoliata) arise in this tissue, and in some cases this is
followed by lignification of the walls of the adjoining cells_
Cortical vascular bundles (see p. 550) are found also in the stem of
Limnanthemum nymphaeoides and in the rhizome of Villarsia exaltata, F. v. M.
In the latter species the cortical bundles are inversely orientated and inserted
in the ring of pericyclic scierenchyma, as it were on the back of the normal
vascular brindles.
The previous account (p. 550) of the structure of the stem in the sapro-
phytic Gentianoideae, apart from the statement concerning the bicollateral
vascular system of Voyria rosea, Aubi. (on Gilg's authority), dealt exclusively
with the genus Leiphaimos, since, according to Gilg, the species investigated
by Johow and formerly referred to Voyria belong to Leiphaimos. We will
now once more summarize the features presented by the structure of the axis
in the saprophytic genera, taking the results of recent work into consideration.
Those cases in which there is no strengthening ring in the stem may first be
considered. The vascular system then consists either solely of four concentric
vascular bundles with a few vessels in the middle and a few peripheral phloem-
bundles (Leiphaimos azurea, Gilg, L. fiavescens, Gilg and L. tenella, Miq.), or
of an almost annular and closed ring of vascular bundles showing bicoll3.teral
structure and not containing any mechanical tissue (Cotylanthera tenuis, Bl.).
In other cases a strengthening ring is present, and here we may likewise dis·
tinguish two modifications, according as the strengthening ring is developed
as a ring of pericyclic sclerenchyma or as the xylem-ring of the fibrovascular
sytem. A pericyclic strengthening ring is found in Leiphaimos aphylla, Gilg,
L. parasitic a, Cham. et Schlecht. and L. trinitensis, Gilg; in two of these species
(L. aphylla, L. trinitensis) six concentric vascular bundles with central xylem are
apposed to the inner side of this ring, while in the third species (L. parasitica)
there are six vascular bundles, in which phloem-groups are developed only on the
side facing the pith, while the xylem-groups are embedded in the strengthening
ring. On the other hand, Voyria coerulea, AubI., Voyriclla parvifiora, Miq.
and the semi-saprophytic species Obolaria virgi'tica, L., have a bic~llateral
vascular system, the xylem of which constitutes the strengthening ring. Voy,ia
1'osca, Aubl., which according to Gilg has four bicollateral vascular bundles
• not actually in connexion with one another, although very close together:
should probably be classed with Cotylanthera. Among the species of the semi-
saprophytic genus Bartonia, B. lanceolata, Small more or less conforms to the
type of stem-structure found in Obolaria 1}irginica; in B. ver1ta, Miihl. neither
the wood nor the pericycle contain mechanical tissue, while the bundles in the
higher internodes of the stem show an irregular differentiation, since one finds
both collateral and bicollateral vascular bundles as well as mere phloem- or
xylem-groups on the inner side of the endodermis (for details, see Holm, loco cit.).
In dealirig with the structure of the root in. the Gentianeae an important
point to noti~.~ is thatcthe radial vascular system in the Gentianoideae consists
of two xylem- and two phloem-groups, while in the Menyanthoideae the
number of groups is larger (5-9)' In many of the Gentianoideae the structure
of the endodermis is very characteristic. Its cells, which are provided with
Caspary's dots on their radial walls, in the first place grow stt:,ongly in the
tangential direction; this is followed by the appearance of secondary radial
division-walls in each cell in numbers, which vary from 2 to 20 according to the
species, and even show variation within certain limits in one and the same
species; tangential division-walls are more rarely found side by side with tho
radial ones. Perrot mentions the occurrence of radial divisions in the endo-
dermal ceDs in species of Crawjurdi«, Erythraea, Exacum, Gentiana (species
 ADDENDA-GENTIANEAE

of the secttons Coelanthe, Pneumonanthe, Stenogyne, Frigida, Aptera, Isomeriil,

Chondroph'Ytld, Thylacites, Cyclostigma, Andicola and Amarella), Pleurogyne
and Sebaea, while' endodennal cells with radial and tangential division-walls
are recorded by the same authority in Ge1Jtiana terni/olia, Franch., and by
Holm in Obolaria viTginica, L. In SWtertiCl ronnatCl, F. v. M., S. ptTennis,
L. and other species of Sweertia belonging to the section E uS'UJeeftia there are
two endodermal zones, in which radial division-walls are found, the two zones
being separated from one another by a few parenchymatous layers. Perrot
expressly mentions the occurrence of undivided endodermal c~ in Belmontia,
Cvrlia, Microcala and Sabbatia; in Chi,onia some of the endodermal cells
have no secondary radial walls, while others have a feW'; Cell-divisions, similar
to those observed in the endodermis, occasioriiilly (Gentian~' ciliata a,ccording'
to Vuillemin) occur also in the outermost~ cell;layer' of ..the :-primary Cortex.,
In the Menyanthoideae only simple endpdebnil 'cells are fOll\ld; Development
of cork in the root has been obseryed onlY~ii1:fGtJtUiat&a lute;', G. purii'UTea, &c.~
where it takes place in the pericycle. In nre Gentianoideae the primary
cortex of the root mostly decays and tbe 'part external to the endodermis
becomes detached. In those cases in which, the endodermis is thrown off as .
well the cell-layers of the peri cycle become successively suberiud. ·~n'Halenia
elliptica, Don the primary cortex of the tap-root contains sclerosed, cells.
Bast-fibres or other types of sclerenchymatoUs,elements are sometimes found
also in the peri cycle (species of Chironia, Enicostemma, Exacum). The struCture
of the wood and the occurrence of islands of interxylary soft bast have already
been discussed above. A pith is present in the root in the Menyanthoideae
and occasionally also in the Gentianoideae. ~ regards the saprophytic forms
we may add that distinct root-hairs "arel absent in Obolaria and Bartonia
according to Holm, anq. in Leiphaititos according to Johow and Svedelius,
whilst in Cot yla nthet'a , according to Figdor, root-hairs of a reduced type are
found. According to Stahl, moreover, root-hairs do not appear to be developed
either in the smaller species of Gentiana (e. g. G. asclePiadea, G. ciliata, C. C7U-
ciata, &c.), whilst a mycorrhiza is present not only in the saprophytic species,
but is widely distributed also among the green Gentianoideae (species of Chlora,
Erythraea, Gentiana).
Literature: C:?Stantin, Ti~ ab. ct lout., Ann. Ie. nat., ~r. 6.}. xvi, 1883, p. 139 et se:t.-
Haberlandt, Spaltoffn. d. Scbwtmmpft., Flora, 1887,P. I03....:.;,Baillon, In Bull. Soc. LIOn. de Pans, 1~
1888. p. 756; see also Hist. d. pl., X, ~891. ~. t:24'7Johow, Humnsp~., in ~ringsheim Jahrb., n,
1889. p. 415 et seq.-Solereder. Logamaceae. 1Il Naturl. Pflanzenfam .• IV. Tell, Abt. 2, 189 2 , p. 27.
"otnote.-Boergesen, Arid. p1. bladbygn., Bot. Tid Iskri ft , xix, 1895, po 219 et seq.-A. Meyer.
Starkekorntt, 189S, p. 80.-Kusnetow, Gentz'allo, in NahirL Pflam:enfam., IV. Teil, Abt. 2, 1895, p. 81
et seq.-Figdor. Co~ylanl"era. Ann. J.ardin bot. Buiten:tO~. xiv, 1897, pp. 213-.0, esJX7ially p. 2U
et seq. and Tab. ltVl.-Holm, Ok/ana, Ann. oC Bot•• Xl. 1897. pp.•169-83 and pi. XiX; see also
Proceed. Americ. Assoc., xliv, Salem, 1896, p. ,8g.-Perrot, Tissn e:draliberierl, etc., Compt~
rendus. Paris, 1897. P. I 115 et seq.; and Tissu condnctenr sumum., Joum. de bot.• 1897, p. 37 .. et seq.
-Schubert. Parenchymscheidell, Bot. Centralbl., 1897. iv, pp. 19, :lo.-Wollenweber, Anat. del
Schwimmbk. Diu., Freiburg i. Br., 1891, pp. I9-2o.-Perrot, Anat. compo des Gentian., Ann. sc.
nat.', ser. 8, t. vii, 1898, pp. 105-293 and pl. I-is.; also These, Paris; see also Assoc. fran~. Nantet,
I898.-Leisering, Interxylares Le~tom, Diss., Berlin, 1899. pp~ 33-5.-Stahl, Sinn der Mykorrbizen-
bild., in 'Pringsheim Jahrb., XXXIV, 19(>0. P. 5~ ct seq.-Clndftz, Blattanat. canar. Gew., Diu.!>
Basel, 190', pp . • ', 43 (l.ranthus).-Svedeli.', Sapropbyt. Gentian., Bib. Svcnekn Vet.-Akad.
Handlingar, xl.{\·iii, Afd. iii.. D. 4, 1~2, I~ W-:...... Vllillemill, Bois mtermed., Cowptes rendm, Paris,
cxxxv, 190', pp. 136 7""9; .d., GentUJlltJ CIliata, Bull. Soc. bot. de France, 1902, pp. 274-80, and
Boll. Soc. Se. de Nancy, iii, 1902, pp. 158-H and pl.-[Ponch, SpaltOffnungupp. submers. Ptlan-
zenteile, Sitz. Ber. Wiener Akad., ex.h, Abt. J, '903, p. 124 (IImy4lft/Aes).]-Scboch, Chinm;a, Beib.
z. bot. Centralbl., xiv, 1903, pp. 183, 184.-Schoute, Stelartheorie, 1903, p. uS.-[Aug. \-ogl, Kalk-
oxlllat bei Gentian., Zcitschr. Osterreich. Apothekerver. Wim, 1903, n. 6 and 7; abstr. in Bot. Cen-
tralbl., xcii, p. 43,.]-Col, Faiseeanx, Ann. 1<:. nat., str. 8, t. xx, 1904, pp. 188~2, etc.-Siisaenguth,
llehaaruo,-O'Sverh., etc., Diss., Wurzburg, 19O~, p. H.-Theorin, '"al'.ttricbom., Arki... tor Bot., iii, n. 5.
1904, p. :lIt.-Giirtler. lntenellulare Haarbild., Diss., Berlin, 1905, pp. 3o-.0.-Porsch, Spaltoffnun~.
typus, Jena. 19°5, pp. 8O-3.-Ho1m, Darto1lia, Ann. of Hot., xx. 1906, pp. 441-8 :lnd pt. XXXiii,
xuiv.-(Pilgtr, in Natiir1. P8anzenfam., Erg.-Heft ii, 1908, p. 292.1
 ADDENDA 999

POLEMONIACEAE (pp. 55°-552).

The paragraph dealing with the essential ANATOMICAL FEATURES of the
Polenioniaceae requires the following addltions or alterations. Oxalate of
lime, though not abundant, does occur in the Polemoniaceae, being found for
the most part in the form of small acicular, prismatic, or otherwi~e shaped
crystals. (species of Bonplandia, CoUomia, Gilia); typical small clustered
crystals have also been observed in Bonplandia. True cystoliths are absent,
the statements in the literature referring to tD.eir occurrence being incorrect.
On the other hand, cystolith·like protuberances are found in the cells ()f the
epidermis and of the hairs. The complete absence of gelatinized epidermal
cells in the leaf is remarkable in view of the xerophilous character of many
Polemoniaceae. The hairy covering consists of unicellular (these, however,
never constituting the only forms of hairs) and uniseriate clothing ~airs, as well
as of glandular hairs. Specially noteworthy forms of clothing hairs are as
follows: the uniseriate trichomes of Gilia congesta, Hook., which are fused
in pairs at the base; the uniseriate hairs of certain species of Phlox and Cilia,
which-showa tendency to branch; the bicellular whip-hairs of Cilia densifolia,
Benth., and other species, which are provided with a short basal cell and a long
tenninal cell; and the urn-shaped trichomes of Cilia p'innatifida, Nutt.
The STRUCTURE OF THE LEAF has repeatedly been examined by Wolfel
and especially by HUller; the subsequent description is based on Hiiller's
statements. In those species which have narrow leaves the epidermal cells
are generally elongated in the direction of the median vein and occasionally
appear almost prosenchymatous in surface-view; in the species with bro-ader
lea.ves the lateral walls of the epidermal cells are often undulated (occasionally
showing angular bends and jagged), more rarely straight. The cuticle is
smooth or finely striated, or it may be granular or verrucose; in many species
of Collomia, Gilia and Phlox the part of the ()\iter wall lying .on the inner
side of the cuticle is strongly thickened and capable of swelling, and has a
gelatinized appearance. But gelatinization of the epidermis does not
occur. Isolated epidermal cells showing papillose differentiation are not un·
commonly' found on the margin and ,near the apex of the leaf, and in some
cases a sdicified protuberance resembling a cystolith is associated with these
papillae (species of Bonplandia, Collomia, Gilia, Loeselia and Phlox). Papillae
are never differentiated on all or almost all the epidermal cells on the surface
of the leaf. The stomata are found on both sides of the leaf, especially in the
species with narrow leaves; only in a few cases are they confined to the lower
surface. Special mention may be made ·of the arrangement of the stomata in
Phlox Hoodii, Richards.; this species has narrow leaves which are somewhat
adpressed to the stem and cover one another in imbricate fashion, the stomata
which, occur .on- both surfaces being almost confined to the base of the leaf.
which is·protected by means of hairs. In those species, which have narrow
leaves, the pairs of guard-cells are arranged approximately parallel to the
median vein; a really irregular arrangement of. the stomata is altogether rare
in this Order. There are no subsidiary cells, the stomata developing according
to the Ranunculaceous type (Cobaea, Gilia, Phlox, Polemonium). The pairs
of guard-cells for the most part lie on a level with the epidermis, and deeply
sunk stomata do not occur. The mesophyll is bifacial or centric. The spongy
tissue only rarely (Phlox amoena, Sims.) shows very large intercellular spaces.
In Phlox longifolia, Nutt. and a few species of Gilia (e. g. G. Parishii, Pet.) an
aqueous tissue composed of large cells devoid of chlorophyll is found in the
middle of the mesophyU; this tissue envelops the veins of the leaf and some·
times even surrounds the principal vein. Hypoderm occurs ()nly at the
 1000 ADDENDA--POLEMONIACEAE
margin of the leaf and above the median vein; as a rule it consists of fibrous
c~lls~ provided -with strongly thickened ~ulose ..walls, which are cap::tble of
swelling-. The vascular bundles of. the veins are commonly accompanied by
sc1erenchyma, which is apposed to the xylem· and phloem·groups. In some
species a group of sc1erenchymatous fibres is found also.in the xylem of the
larger veins, these fibres adjoining the tracheae and being in contact with
the soft bast. In this connexion we may mention a few anomalous cases
conwnmg the appearance of the vascular bundle of the median vein, as seen
in transverse sectiqD; these anomalies have been demonstrated in the upper
two-thirds of the leaf in Phlox Drummondii, Hook. and a few species of Cilia.
In some cases the group of sclerenchyma above mentioned enters into con-
nexion with the sclerenchymatous layer below the soft bast by means of a
bridge of sclerenchyma. and in this way the strip of soft bast becomes divided
into two groups, as in the vascular bundles of certain Palms. In other cases
vascular bundles have been observed, consisting for the most part of scleren-
chyma, a group of annular and spiral tracheae being apposed to the upper
side of the latter, while two to four small phloem~bundles accompanied by
spiral and annular tracheae are situated at its sides. Parenchymatous sheaths
consisting of large cells have been observed in certain species of all the genera
except Bonplandia. In some cases they are differentiated like an endodermis
(cells suberized in Phlox amoena, Sims. and P. Drummondi~'t Hook.; cells
thickened in the shape of a horseshoe in G#ia pungens, Benth.).
The following facts may bel mentioned regarding the hairy covering.
The clothing hairs, apart from the more or less hair-like papillae above dis ..
cuSsed, include only uniseriate trichomes. Unicellular clothing hairs have
not been recorded as the only form of trichome in any member of the Order.
The ·uniseriate hairs vary in length, being composed of long or short cells ;
the following types are found :-short conical hairs, consistingoi a small number
of cells and mostly placed obliquely to the surface with their apex pointing
towards the tip of the leaf; stiff hairs, which are longer and are either straight
or curvec like a sabre; and woolly hairs, bent in a vermicular manner. The
walls of the hairs vary in thickness; they are never calcified, but occasionally
slightly silicified. Regarding the special forms of clothing hairs above referred-
to we may add the following details. In Phlox macttlata, L. only one or two
of the 3-5 cells composing the trichomes bear short protrusions, while in Gilia
dianthoides, Endl. each cell of the multicellular trichome has such a protrusion,
so that hairs are formed which ·have a kind of sympodial structure. The
urn~shaped trichomes of GiNa pinnatifida, Nutt. in the simplest case consist
of two bluntly conical cells having a common ,basal surface, but a cylindrical
cell which is slightly narrowed in the middle of its length is sometimes in-
serted between the former; in the three-celled hairs, moreover, the lowest cell
is occasionally also cylindrical. It remains to mention the uniseriate trichomes
of Cilia Larseni, Gray, which are composed of barrel-shaped cells,. and the
uniseriate hairs of G. floccosa. Gray, in which the longitudinal walls are provided
, with numerous small papillose protrusions. The glandular hairs show the
same type of structure as the clothing hairs; in some cases (species of Cilia)
transitIOnal forms between the two kinds of hairs are found at the base of the
leaf. The variations in structure exhibited by the glandular hairs concern
the length of the stalk and the number of cells composing it, and especially the
nature of the head. In some of the hairs the head is unicellular' and spherical,
ellipsoidal or clavate in shape; glandular hairs with heads of this type are
the only forms of external glands found in Bonplandia, Cantua and Cobaea
(in the last of thes~ genera they constitute the only type of hair), but they
also occur side by side with glandular hairs having multicellular heads in the
remaining five genera of the Polemonlaceae. Glandular hairs \vith spherical
 ADDENDA-POLEJI;/ONIACEAE 1001

heads, divided into 2-8 cells by one or more radially arranged vertical walls,
are foUnd in many species ot Collomi.a, Cilia, Loeselia, Pltlox and Pole.nonium.
Lastly, in certain species of Collomia, Giti4 and Phlox glandular hail;s..are ff)und
in which the heads are obconical, or m~re rarely discoid or almost spherical,
and are divided in the first place by horizontal walls into 2-4 tiers, which
undergo further division by vertical walls.
Oxalate of lime is found in certain species of CoUomia and Gili4 in the
form of the small crystalline bodies above mentioned, the latter OCCUlTing in
the mesophyll and occasionally in the epidermis and tril;homes as well. In
Cilia atractyloides, Steud. these small crystals are united to form structures
resembling clustered crystals. Distinct clustered crystals of small size occur
in Bonplandia gemiHi/lora, Cav., lying singly in each cell of the palisade tissue;
in addition to them the small crystals are likewise present in the mesophyU of
Bonplandia. Greenish and Morelle's statement to the effect that typical
calcified cysloliths OCCUf in ' Phlox carolina J is incorrect according to Holm,
Hiiller and Stockberger; the material, which Greenish and Morelle. used in
their investigations under the name of 'Phlox carolina,' does not belong to
a member of the Polemoniaceae, but to one of the Acanthaceae (according to
Holm, Ruellia ciliosa, Pursh).
For t!1e STRUCTURE OF THE STEM, see also the detailed statements by
Wolfel, loco cit. According to \-Volfel the vessels have a small number of
scalariform perforations (with 1-10 bars) side by side with the simple per-
forations also in Polemoni-um and Cilia.
Literature: Wolfel, Vergl.Anat. d. PolemoD.,Diss., Heidelberg, 1901,62 pp., 2 Tab.-Theorin,
Vaxttrichom., Arkiv for Bot., I, 1903, p. 159.-Co1, Faisceaux, Ann. sc. nat., ser. 8, t. XXI 1"904,
p. Jl3.-Morelle, Histol. camp. des Gelsemiees et Spigeliees, These. Paris, 190 4, ,Pp. 144-7; alsotn
Perrot, Travaux, ii, I905.-[Stockberger, in Proceed. Americ. pharm. Assoc., Hil, I90~, p. 3:J".}-
Holm, Root~stmctDre of Sprge/t'a marilandica, Phlox ovata, etc., Amenc. Journ. of pharm., J9od,
p. 553 et seq.; and Internal strncture of the stem and leaf of Ruellia ciliosa, Pklox ova/a, etc., loc. cit.,
1907, p. 51 et seq.-Hiiller. Beitr. z. verg1. Anat. d. Polemon., Diss., Erlangen, 1907, 75 pp., 1 Tab.,
sep. copy from Beih. z. bot. Centralbl.-Brand, Polemoniaceae, in Pllanzenreich, Heft 27, I90 7, pp. 3, 4.

I
HYDROPHYLLACEAE (pp. 552-554).
Literatnre: [Ritter, Emdictyoll glutinosulII, Americ. Joum. Pharm., 1895, p. .565 et seq.]-
Minden, Wassersez. Org., Bibl. bot., Heft 46, 1899, pp..-26 and 38.-Theorin, Vaxttrichom., Arkiv f<ir
Bot., iii, n. S. 19(14, p. 26, and iv, n. 18, 1905, pp. 16, 17.

BORAGINEAE (pp. 554-561).

2. STRUCTURE OF THE LEAF. Epidermal cells which have jagged lateral
walls with .folds compressed like ridges in the angles, are found on the lower
surface of the leaf in Lithospermum purpureo-coeruleum and Pulmonaria (J odin).
For theohairy covering see "also Revedin's detailed statements (Ioe. cit.),
which refer~to species of Alkanna, Amsinckia, Anchusa, Asperugo, Caccinia,
Echinospermum, Echium, Heliotropium, Lithospermum, Lycopsis, Afy()sotis,
Nonnea, Omphalodes, Onosma, Onosmodium, P'ulmonaria, SOlena1!thus, Sym-
phytttm, To ltrnefortia, Trachystemon and Trichodesma.
To the special description cf the trichomes we may add that bracket~4airs
occur side by side with other forms of trichorries also in Anchusa officina/is, L., and
that almost spherical cystolith-hairs, provided with a short pointed" filiform process,
accompany other types in Symphytum officinale, L. In the glandular hairs ot
Nonnea pulla, DC. which have uniseriate stalks and unicellular heads. the lowest
of the four or five cells composing the stalk is very strongly developed and forms
a kind of pedestal. The stalk of the glandular hairs, found on the calyx in Tourne·
1002 ADDElVDA-BORAGINEAE
fortia /ruticosa, R. Br., likewise shows a special type of differentiation, two of the
four component cells (viz. the lowest and the next but one above it) being distended
in a barrel-shaped manner near theit: upper end, while the other two are developed
as short neck-cells.
In the species investigated by J odin the petiole may contain as many
as twelve isolated vascular bundles (generally 5-6 or only 3), the number being
connected with the differentiation of a furrow of varying breadth; the median
vascular bundle alone is strongly developed as an arc of wood and past.
3. STRUCTURE OF THE AXIS. For the detailed structure of the stem in
the herbaceous members of the Order, see ] odin. The primary cortex in
these forms is loose and occasionally contains palisade tissue; in Echium
vulgare it includes strongly sclerosed cells. The pericycle is generally paren-
chymatous, and rarely (Heliotropium curassavicum) contains groups of, fibres.
The vascular bundles are either isolated or fused to form a ring of woon and
bast. A composite and continuous ring of sclerenchyma has recently been
observed,by Pitard in the pericycle of Tournefortia hirsutissima.
Note. According to Jodin the root of the herbaceous Boragineae generally
contains a diarch vascular system, althollgh this does not apply to the adventitiQus
roots of the rhizome, which show 5-6, sometimes even as many as twelve, radial
vascular bundles, two bundles only being found as an exception (Symphytum
tuberosum). The development of the cork in the root takes place in the pericyc1e
(Alkanna, BOJ'ago, Echium, Pulmonaria). Regarding the red coloration shown by
the roots of many Boragineae, see Norton (in l\Essouri'Bot. Garden Report, 1898,
pp. 149-5 I ) and other au thori ties.
I
Literature: Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, p. 134 et seq.-
[Tognini, Stomi. Atti 1st. bot. Pavia, t894.]-Schubert, Parenchymscheiden, Bot. Centralbl., 1897,
iv, p. lo.-Minden, Wassersez. Org., Bibl. bot., Heft 46, 1899, p. 2,3.-Pitard, Pericycle, These,
~rdeaux, 1901, pp. 39, 65 and I04.-[Baldacci, Symphytum adm/ale, Rendiconti Accad. Bologna,
iv, pp. 74-8; nbstr. in Just, 1902, ii, p. :ISo. ]-Clauditz, BlattanaL cana.r. Gew., Diss., Basel, 1901 ,
pp. 49, 50 (Echium).-Preston, in Bot. Gazette, 1902, pp. Ifjo-4 (Amsinck,·a).-Revedin, Peli delle
Boragin., Nuovo Giom. bol Ital., N. S., ix, 190:1, pp. 301-IS.-Jodin, Rech. an at. sur les Boragin.,
Ann. sc. nat., ser. S, t. xvii, 1903. pp. 263-346, pL 5-9 (deals only with the European genera, and with
A 11lsinckia).-Theorin, Viixttri ch om. , Arkiv for Bot., I, '9°3, P. ],58; ii i, o. 5. 19°4-, pp. :16, 'J 7; and
iv, n. 18, 190_5, p. 22.-Freidenfeldt, Anat. Ban d. Wurzel, Bibl. bot., Heft 61, 1904, PP' 68, 69.-
Sussenguth, Behaarungsverh. d. Wiirzh. Mnschelkalkp6., Diss., Wiirzburg, 1904. p. 4S.-[Vidal,
EritrichiulIl na1zum, Assoc. fran y• Cherbourg, 19°5. pp. 47:1-Jii.]-[Kimptlin. Affinites des Boragin.
et des Lamacees, Assoc. Avane., 35 Sess., Lyon, 1906, pp. ,pS-3I.]-[Kramer, Mikr.• pbarm. Beitr••
Diss., Wiirzbnrg, 1907, p. 39 (Pulmonaria).]

CONVOLVULACEAE (PP.562-573).
To the discussion of. the internal secretory system (see pp. 565, 566) we
may in the first place add that according to a written communication from
Halliet the axes in all the higher Convolvulaceae (Ipomoeeae, Argyreieae,
and M erremia) contain a white or whitish sap. The secretory organs found
in the flower have been examined in detail by Grelot. According to him the
floral organs likewise contain secretory cells (isolated and in rows), as well as
cell·fusions~ the latter occurring in Falkt"a and Dichondra. Grelot, like the
earlier observers (d. footnote 3 on p. 565), maintains that he has encountered
resorption of the transverse walls in the rows of laticiferous sacs.
As regards the glandular hairs (see pp. 568, 569) reference may be made
to the large external glands, which are situated on the lower side of the leaf
in Stictocardia tiliaelolia, Hallier, and are provided with a head showing vertical
division, and to the pore-like perforation of the cuticle observed in the ordinary
peltate glands of Operculina Turpethum, Peter (Svedelius).
 ADDENDA--CONVOLVVLACEAE 1003
ApPENDIX: I. CUSCUTE:AE (p. 573).
-,
'Our knowledge of the anatomy of the Cusauteae has been quite materially
improved by recent investigations undertaken by Cornu and especially by
Mirande 1.
_ The ,secretory organs of the Cuscuteae consist of secretory cells, which
may be distinguished as cortical (situated in the primary cortex), pericyclic
and medullary according to their point of occurrence in the stem. Cortical
and pericy~c secretory cells are found in all the spe{:ies of Cuscuta, and con-
stitute two characteristic zones in the thin stems of these forms, while medullary
secretory cells have been observed only in C1t.Scuta americana; the latter
show the same structure as the elements sltuated in the cortex. In the young
parts"of the axis the zone of cortical secretory cells lies in the peripheral portion
of the primary cortex. In the older parts of the axis these elements are more
or less separated both from one another and from the epidermis as a resUlt
of the cell-divisions which have taken place in the primary cortex. In the
young parts of the axis the cortical secretory cells are moderately elongated,
and form. vertical rows of ceils, whic.h can be distinguished from the neighbour-
ing~ cells even by their size; in the l\Ionostyleae (Subgenus Monogynella,
Engelm.) their nucleus occupies a peripheral position, while in the Distyleae
(Subgenus Grammica and Cuscuta) it lies in the centre of the cell. In later
stages the cells .often elongate very considerably. The neighbouring ShOTt
cells of the cortex then frequently exert a pressure on the longitudinal walls
of the secretory sacs, so that dents are produced, which in the case of secretory
sacs isolated by maceratio.n .a~pear a~ a system of facets, corr~sponding to the
cellular network of the adjOInIng cortIcal cells. Except for thls the wall of the
secretory sacs is either smooth or finely punctate. The pressure exerted by
the turgescent cortical cells also commonly leads to the almost complete com-
pression of the cortical secretory cells in the older parts of the axis, so that
. they easily escape observation. The zone of pericyclic secretory cells is like-
wise typically developed only in the young stems, but for them it is highly
characteristic, the cells being readily recognized owing to their wide ,lumina.
The peri cyclic secretory cells also become separated froID.one another in later
stages; their lumin, then occasionally become narrowed as a result of inter-
calary growth, or the cells may acquire very thick walls and come to resemble
fibres; in other cases they retain their thin walls, but lose their rounded
outline in transverse section and look like star+shaped intercellular spaces. This
accounts for the fact that the earlier observers failed to,.place a correct inter-
pretation on the pericyclic secretory cells. In the Monostyleae the membranes
of the pericycli~ secretory cells become more, or less lignified, while in the
Homostyleae (Subgenus Cuscuta) they generally continue to consist of cellulose;
in'the Heterostyleae (Subgenus Grammica) the membranes ultimately become
thick, cartilaginous and refractive, and acquire the property of swelling up
in the preseI].ce_ pi water~ the cells at the same time assuming a fibre-like
appearance. -In the last stage the peri cyclic secretory cells constitute long
tubes, the length of which in the Monostyleae and Homostyleae occasionally
equals that of five internodes, althcugh shorter in the Heterostyleae. Neither
1 Mirande's investigations denl with the following species: T. Monostyleae: Cuscuta cass),thoides,
C. exaltata, C. faponica, C. Lenmannia1J(Z, C. lttfuliformis, C. 1ItOIU)GYIZQ, C. rejlexa, C. ti11Ioren.sis.
II. Distyleae: a. Homostyleae: C. ahyssinica, C. ajricana, C. arabica, C. babylonica, C. /Jrtvistyia,
C. capitata, C. Epilinum~ C. Epithymum~ C. eztt"opaea, C. m·tida, C. palaestt'na, C. ptl'roifora,
belonging to the type of C. eU1"opaea; b. Heterostyleae: C. decora, C. Gronovit", C. injlexa~ belonging
to the type of C. GrtmbV£i; C. califlrnica~ C. dzilmtt"s, C. cMnmsis, C. corymbosa, -c. ctlspidata. C.
jioribu1uia, C. glomerata~ C. lzyalina J C. ia/ajJeusis, (.:. adora/a, C. rostrata, C. sQ1tdwir:hiana, C.
tenftijiora, C. umf>elfata~ belonging to the type of C. {kimnsis; C. amel"icana, forming a special type
of its own.
 ADDENDA--CONVOLVULACEAE
branching nor anastomosis has been observed in the secretory cells. A very
noteworthy feature lies in the fact that the pericyc1ic secretory cells contain
several ellipsoidal or fusiform nuclei in place ·of a single one. The chemical
nature and surface-markings (punctation or striation) of the wall are subject
to variation, occasionally even in one and the same cell. Of the secretory
cells occurring in the stem as a rule only the cortical elements can be traced
into the scaly leaves. In the latter, and especially ~ the lower part of the
scale-leaf, they frequ.ently form a complete hypodermal layer of. rather short
cells with wide lumina above the lower epidermis; in the upper part of the
leaf the secretory cells decrease in number and ultimately occur only as isolated
elements. The protQplasmic contents of the secretory cells soon disappear,
the mature elements chiefly containing oily substances, tannins and resins.
An, important systematic feature concerning the differentiation of the
fibrovascular system of the stem is that in the Monostyleae (the stem being in
that stage of development in which the vascular bundles are fully differen.tiated,
but have not yet begun to grow in thickness) the xylem-groups qf the '\\ascular
bundles, which show an annular arrangement,1 are united to form a strengthen-
ing ring by means of groups of elongated sclerenchymatous cells resembling
fibres, while in the Distyleae this is not the case. In the Distyleae, moreover,
the entire fibrovascular system, and especially the xylem-groups of the bundles,
show a greater degree of reduction than in the Monostyleae.
Among the Monostyleae the vascular bundles are most strongly developed in
Cuscuta japonica, in which they $0 show growth in thickness in later stages.
Even in this species, however. strands of mestome, which are essentially of the nature
of phloem-bundles, occur side by side with the fully differentiated bundles (i. e. those
containing both wood and bast) of the vascular ring; these phloem-bundles are either
apposed to the outer side of the sClerenchymatous strengthening ring or are embedded
rin It. The xylem-groups of some of the vascular bundles. moreover, do not extend
through the entire thickness of the strengthening ring, being separated by a bridge
of sclerenchyma from the corresponding bast-group. The process of secondary
growth does not take place uniformly in the vascular bundles of C. iaponica. This
results in the bursting of the strengthening ring, while the vascular bundles them-
selves present a varied appearance in transverse section. Side by side with the
normal bundles of wood and bast, in which the secondary wood adjoins the primary
wood in the ordinary way, one meets with the fonowing types: (a) bundles in
which groups of sclerenchyma belonging to the original strengthening ring are
inserted between the primary and secondary wood (these being the bundles, in which
the xylem-groups did not extend through the entire strengthening ring prior to
the commencement of secondary growth); (b) bundles of wood and bast, which
cOl;ltain no primary wood and are derived from the original phloem-bundles; 'and
lastly, (c) groups of interxylary phloem, which owe their development to the cessation
of cambial activity in the median of three vascular bundles, lying next to one
another, while the two lateral bundles undergo considerable growth in thickness,
the new wood thus formed covering in the phloem-group belonging to the median
bundle.
. In the remaining Monostyleae the vascular system is more or less reduced,
as evidenced not only in the absence of growth in thickness, but also in the smaller
number of bundles of wood and bast or of phloem and the smaller number of tracheae
in the xylem-groups. In the Distyleae reduction has gone even further than in the
Monostyleae.

The pericycle in the l\lonostyleae besides containing secretory sacs also

includes groups of bast-fibres. The sieve-tubes, which are often developed in
large numbers and frequently have wide lumina, have horiiontal or oblique
sieve-plates with one or more sieve-areas; their longitudinal walls show
manifold types of structure, but bear sieve-areas only in the Monostyleae.
The structure of the pith varies in the monostylic species. In Cusc'llta
cassythoides, C. Lehmanniana, C. monogyna and C. timOTe1f,Sis it is sclerosed
 ADDENDA--CONVOLVULACEAE r005

except for small islands of thin-walled 'tissue in 'the neighbourhood of the

primary xylem-groups, while in the remaining species the walls of the medullary
cells consist of cellulose.
, According to Mirande the nature of the intercellular system of the stem
is likewise a feature of systematic importance. "The Monostyleae and Homo. .
styleae have small lacunae, occurring especially in the primary cortex; in
hath cases these lacunae are enveloped by parts of the walls of the adjoining
cells, th~se portions of the wall being suberized in the Monostyleae and cuti..
nized in the Homostyleae. In the Heterostyleae of the type of C. chinensis
and C. americana the primary cortex and bast contain canals and cavities,
having cutinized walls. Very commonly, moreover, lacunae may also be
observed on the inner side of the vascular bundles, these cavities arising by
abortion of the primary vessels, in the same way as in many Monocotyledons
and Dicotyledonous aquatics.
Finally, according to Mirande, the axes of the Cuscuteae are characterized
e
also oy' the lIl"esence of. a peculiar sheath of cells gatne nourriciere') having
abundant ptbtoplasmic contents and developed as a complete envelope to the
individual vascular bundles; this sheath is most typically differentiated in
the Monostyleae.
The structure ot the scale-leaves, which are devoid of Chlorophyll, may
ti~t be described in Cuscuta japonica, where they are most strongly developed.
A transverse section through the base of. the leaf in this species as a rule shows
a lower epidermis composed of large cells and an upper epidermis in which
the cells are smaller; aboV'e the lower epidermis an almost continuous layer of
secretory cells is situated~ while the vascular bundles are embedded in a homo~
geneous mesophyll consisting of rounded cells. The cells of the lower epidermis
are divided by numerous vertical walls, only a small number of which occur
in the .upper epidermis. Only the lower epidennis contains stomata, which
are'rather numerous; stomata are altogether not as rare in the Cuscuteae-
even on the different parts of the stem-as has hitherto been supposed. As
we approach the cucullate apex of the scale ..leaf the subepidermal secretory
sacs show a considerable decrease in number; the main mass of the apex
is constituted by a group of small-celled tissue, which occupies a central position
and is formed by the division of the tissue accompanying the fibrovascular
system. Occasional leaves, which arc particularly strongly developed, even
have typical palisade tissue containing.scIereids on their upper side. Of, the
vascular bundles entering the leaf, the median' one at nrst retains its complete
structure; in its further course, however, its xylem soon disappears, and
a phloem-bundle remains, from which branches are given off to the group of
tissue situated in the centre of the cucullate apex of the leaf. The two lateral
vascular bundles lose their xylem-groups on entering the leaf, while the
remaining lateral bundles passing into the leaf already leave the axis in the
form of phloem-bundles. In C. exaltata the structure of the leaf is simiku
to .that ~C?f ,C. jaPo.'!'t-ica.. In the remaining Monostyleae and in the Distyleae
the structure of the leaf is reduced to a varying extent.

2. NOLANEAE (p. 573).

According to Mirande alcohol-material of Nolana paradoxa contains
sphaerocrystalline masses, consisting of calcium malophosphate and neutral
calcium malate, of which the former is more abundant.
Literature: Decaisne, Struet. Anat. de Ia Cascote, etc., Ann. &c. nat, .er. 3, t. v, 18,46, pp. :147-9-
-Costantin, Tiges aero et sout., Ann. SC. na.t., ser. 6, t. xvi, 1883, p. 1~6 eot seq.-Gudtner, Petiolar
glands of the 1pornoeas, Proceed. Cambridge Phil. Soc., v, 1887, p. 8a.-:-Lamoanette, Liber interne,
Ann. sc_ nat., Set. 7, t. xi, 1890. pp. 2S8-60.-[Borzi, Ctistalloidi nucleari ell CtJn1Mlvulus, Contribuz.
1st. bot. Palermo, Fas·c. i, 1894. p. 6S.]-[Tognini, Stomi, Atti 1st. bot. Pavia, 1894.]-Comu, Une
 ADDENDA--CONVOLVULACEAE
CUSCllte du Turkestan, ii, Bull. Soc. bot. de France, 1896, pp. 70i-20 and pl. xv, xvi.-Biennann,
Olzenen, Diss., Bern, 1898, pp. 60, 61.-Miranoe, Melate neutre de Calciu~, etc., Joum. de bot.,
1898, p. 6' et seq.-Mirande, Laticiferes et tubes cribIes de. CtUCutes monogynees, Joum. de bot.,
'1898, pp. 70-80 and 8o-90.-Hallier, Bombycospermum, Jahtb. Hambutg. wiss. Anst., xvi, tS9~, 3.
Beih., 1899. pp. 59-62 (demonstration C?f B()mbycospermum as a member of the COllvolvulaceae an~
as a speeies of Ipomoea belonging to the section Ernsptrmum, on the basis of the a.natomical
features).-Boergesen og Paulsen, Veget. danalc.-vestind. Oer, Bot. Tidslikrift, xxii, 1898-9, p.p. .1:06,
107 (Evolvulus nummularius, L.).-&ranetzky, Faisceaux bicoU., ADD. sc. nat., ~r. 8, t. Xli. 1900,
p. 294.-Mirande, Rech. pbys. et anat. sur les Cu~clltacees, These, Paris, 1900, 284 pp., 16 pl.,
especially p. uo et seq.; also Bull. sc. de Ia France et de Ia Belgique. xxxv.-Beuhlygue, Calyste.fia
S#/daneIl4, These, Montpellier, 1901, pp. 22-30.-[Grelot, Laticiferd de la fleur des Convolvulac.,
Nancy, 1903, 23 pp.; abstr. in Bot. Centralbl., xcii, p. 83.]-Knothe, Unbenetlb. Blatter, Diss.,
Heidelberg. 19°2, p. 8.-[Miiller, Rhodiserholz, Pharm. Post, 1903. p ...pi et seq.; abstr. in Bot.
Ceutr.lbl., xcv, p. 107.]-Chrysler, Strand plants, Bot. Gazette, uxvii, 1904, p. 461 et seq. (Con-
vDlvulus).-Col, Faisceaux, Ann. sc. nat., ser. 8, t. xx, 1904, p. 18i.-Frommel, Plantas text. chiLo
J905, p. i 3 .-[Karoly RessO, Cuscuta IlulVtolms, Kiserletiigyi kozlemenyek, Budapest, viii. 1905,
pp. 604-~3, 3 tab. (Hungarian); abstr. in Just, 1905, ii, p. -4S.]-Porscb, Spaltoffnungstypas, Jena,
190!t, pp. j2, 73.-Svedelius, Postfior. Wachstum der Kelchbl. einiger Convolvulac., FIku;a. xcvi,
19OO, pp. ~3~59, especially p. 24 1 et seq.
1

SOLANACEAE (pp. 575-583).

I. REVIEW OF THE ANATOMICAL FEATURES. To the list of special features
found in certain members of the Order we may add the occurrence of: arm·
palisade parenchyma (Protoschwenkia and species of Schwenkia); peculiar
thickened palisade~cells (species of Dyssochroma, Juanulloa and Ma,kea);
cells of the spongy tissue with swollen areas on the walls (Melananthus guate-
malensis, Solered.); and tannin·sacs (Solandra grandijlora, Sw.).
2. STRUCTURE OF THE LEAF. ArI1:l-palisade tissue is developed in Proto·
schwenkia Mandoni, Solered., as well :as in Schwenkia brasiliensis, Poir., S.
divaricata, Benth., S. grandijlora, Benth., S. Karstenii, Vatke, S. mollissima,
Nees et Mart. and S. patens, H. B. K. With reference to its differentiation
we may note that the folds, which are united to form lamellae, as a rule project
singly from above and below into the lumen of the cell, and that the two
compartments thus formed in the palisade-cell communicate with one another.
by means of an approximately circular aperture. In Dyssochroma viridiflol'a~
Miers, Juannulloa aurantiaca, Otto et Dietr., J. membranacea, Rusby and
M arkea coccinea, Rich. the cells of the palisade tissue are distinguished by the
fact that their walls, and especially the longitudinal ones, are strengthened
by means of thickening ridges, which run in a direction parallel to the length
of the cell and are occasionally connected by means of transverse ridges. In
T,ianaea nobilis, Pl. et Lind. and T. speciosa, Solered. the mesophyll, and
especially the spongy tissue, includes sclerosed cells. In. M elananthus guate·
malensis~ Solered. the spongy tissue likewise exhibits a peculiar type of struc·
ture; at certain points- the walls of the cells show swellings, recalling the'
thickened parts of the wall in collenchymatous tissue (Solereder).
In the species of Datura (sect. Brugmansia) examined by Lagerheim the
lower epidermis of the leaf contains chlorophyll. The stomata in the species
of Trianaea are distinguished by having peculiar coma-shaped depressions, which
are situated at both ends of the two guard~cells and run obliquely to the direc-
tion of the pore, the thickened part of the coma being directed outwards
(Solereder).
No new types of hairs (see p. 576) have been recorded. I have observed
branched multicellular hairs also in A nthocercis albicans, A. Cunn., A. littorea,
Labill., A. tasmannica, Hook. fil., Anthotroche pannosa, Endl., Juanulloa
membranacea, Rusby, J. Sargii, J. D. Smith, Solandra grandiflora, Sw. (here
side by side with unbranched hairs); according to Kearny they occur also in
Physalis viscosa, L. (here showing dichotomous branching). -Combined clothing
 ADDENDA-SOLANACEAE 100']

and glandular hairs occur in Solandra grandijlora, the branched trichomes of

this species commonly bearing spherical glandular cells at the ends of their
branches. Hairs of this type are found also in Nicotiana T abacum, but in this
species the glands, borne on the branched trichomes, are bee1 unicellular and
multicellular. External glands having a unicellular head have recently been
observed also in species of Datura belonging to the section Br'ltgmansia (here
side by side with glands having multicellular heads), in Isandra Bankrottii,
F. v. M. and Protos.chwenkia Mandoni (here together with glands having multi-
cellular heads); glands having a multicellular ellipsoidal head, divided by
horizontal walls only or both by horizontal and vertical walls, have been found
also in species of Anthotroche (with rather long stalks)',. Protoschwenkia (with
stalks of varying length), and Dyssochroma, Juanulloa, Markea, Soland7a,
T,ianaea (stalks in these cases short). The cells of the external glands occa-
sionally (Nicotiana Tabacum) contain oxalate of lime in the form of small
clustered crystals.
Por the spines of Datura and Solanum, in the formation of which ground
tissue participates, see a1so Lothelier and Mittmann.
The tannin-sacs above mentioned are stated to resemble those of the
Leguminosae and have been recorded by Pistone in the stem and root of
Solandra grandiflora.
To the earlier statements regarding the occurrence of oxalate of lime in
this Order we may add that typical crystal-sand has noW been demonstrated
also in Dyssochroma, Iuanulloa, M arkea and Trianaea, and that in Proto·
schwenkia ox ala te of lime occurs only in the axis, being presen t in the fonn
of a crystal-sand composed of fusiform, prismatic or acicular crystals of varying
size. In Anthot1oche pannosa, as well as in other members of the Order, the
typical crystal-sand includes relatively large tetrahedral crystals (Solereder).
3. STRUCTURE OF THE AXIS. Recent observations have shown that the
vascular system exhibits bicollateral differentiation also in Protoschwenkia
and Trianaea (d. footnote 3 on p. 579).
Leisering's recent investigations have likewise failed to completely elucidate
the mode of development of the interxylary phloem (see P.580). The latter
occurs also in the wood of the root of Browallia viscosa, H. B. K. (Scott and
Brebner), and in the rhizome of Atropa Bellado'n-na (Beauvisage).
The wood-prosenchyma bears bordered pits in Protoschwenkia, and simple
pits in Trianaea. In both genera the perforations of the vessels are simple, and
the medullary rays of the wood narrow.
To the section dealing !Vith the structure of the cortex (p. 58!) we may
add that the development of the cork commences in the epidermis also in
Protoschwenkia.
Literature: Uhlworm, Entwicklun gsgesch, d. Trichome, Bot. Zeit., 1873, p. 801 et seq.-
Costantin, Tiges aero et soul, Ann. sc. nat., ser. 6, t. xvi, 1883, p. 132 et seq.-Kassner, Mark,
Diss., Basel, 1884, £p. l3~5.-Radlkofer'oBeitr. Z. afrik. Flora (J883), Abh. naturwiss. Ver. Bremen,
viii, 1884, p. 'Pi; footnote (refers to the interxylary phloem of Atropa Belladomza).-MiUmann,
Pfianzenstaeheln. Verh. Ver. Mark Brandenburg, 18S8, p. 66.-Beau'i'lsage, Faise. eribles dans Ie bois
sec. de 1a Bell ad anne, Joum. de bot., 1891, pp. 16I-3.-Lamounette, Liber interne, Ann. sc. nat.,
ser. 7, t. xi~ 1891, pp. 252-S.-Scott and Brebner, Internal phloem, Ann. of Bot.• 1891, pp. 265-9.-
Lothelier, Epines, These, Paris, 1893, p. 3s.-11ehrens, Tabakblalt, Landwirtschaftl. Versuchsstat.,
xliii, 1894, p. 271 et seq.-[Nevinny, Scopolia, Pharm. Post, 18941 p. 333 et seq.; abstr. in Just, 1894-,
i, p. 483.]-[Pistone, Solalldra, _Contribuz._ 1st. bot. Palermo, i, 1894, pp. 119-22; abstr. in Just,
1894, i, p. 27 I et seq.)-[Tognini, Stomi, Atti 1st. bot. Pavia, 1894.]-Tswett, Anat. de LyciulII,
Bull. Herbier Boissie r, 1894, pp. I 75-9.-Lagerheim, Bru.rmansia, Pers., in Engler"Bat. Jahrb.,
Xlr, 1895, pp. 656, 6S7.-Sabria, Uelladanne, These, Montpellier, 1895, pp. 17-2I.-Went, Haft- u.
Niihrwurzem, Ano. Jardin Buitenzorg, xii, 1895, p. 59.-[Johnson, Solaltum carolilleme, Americ.
Journ. of Phann" 1897, n. 2; abstr. in JUiit, 1897, ii, p. 52; root with anomalous structure and
other curious statements !1-Schubert, Parenchymscheiden, Bot. Centralbl., 1897, iv, P. 61.-
Solereder, Zwei Beitr. z. Syst. d. Solanaceen, Ber. deutsch. bot. Gesellsch .• 1898, pp. 242-60.-
Boergesen og Paulsen, Veget. dansk-vestind. Oer, Rot. Tidsskrift, xxii, 1898-9, pp. go nnd <)2
1008 ADDENDA-SOLANACEAE
(So/anum).-Hirsch, Entw.lcJclung d, Haare, Diss., BerJin, 1899, p. 3:1.-Leisel'lng, Inter.l(yliires
Leptom, Diss., Berlin, 1899, pp, 3Q-3z.-Minden, Wassersez. Org., Bibl. bot., Heft 46, 1899, p. 38
et seq.-Tswett, Leptom der Solanac., Ber. deutsch. bot. Gesellscb., 1899, Genena.lversammlungs-
heft, pp. (231)-(235).-Baranetzky, Faisc. bicoll., Ann. sc. nat., ser. 8, t. xii, 1900, pp. 283-9.-
Kea.my, in Contribut. U. S. Nat. Herb., v, n. 5, 1900, p. 298.-[Kraemer, Crysta.ls of /)atur4
Stra11lq-nium, Bull. Torrey Bot. Club, 1900. p. 37; abstr. in Just, 1900, ii, p. 89.]-Tuumann,
Sekretdriisen, Diss., Bern, 1900, pp. 33-6.-Thomas, Feuilles sout., These, Paris, 1900,-[Hartwich,
Verf. d. Folia Belladonnae, Schweizer. Wochenschr. f. Chemie u. Pharm., 1901, p. 430; abstr. in
Just, 1901, ii, p. 47.]-Peterse~ Vedanatomi, 1901, p. 83.-Siim-Jensen, Hyoscyamus niger, Bibl.
bot., Heft 51, 1901, 90 pp., 6 Tab.-[Bames, Potato, Ann. Report and Trans. North Staffordshire
Field Club, xxxvii, 1902-3, pp. 96-106. )-(Geremicca, Juanulloa Qurantiaca, Boll. Soc. Nat.
Napoli, set. i, '/.v. 1902, pp. 61-76 and tab. iii-v; abstr. in Bot. CentralbJ., xcii, p. I.]-[Marcello,
1st. di ale. Solanum, Cava dei Tirreni, 1902, p. 10, tab. ; abstr. in Bot. Centralbl., xciii, p. 161.]-
Martel, Notes sur l'ana~ des Solanac., Joum, de bot., 1903. pp. 211-)4'-Theorin, Vaxttrichom.,
Arkiv- tOt Bot., i, 1903, p. 155.--Col, Faisceaux Ann. lOC. nat., ~r. 8, t. xx, 1904, p. 1,94< etc.-
Paoli, Eterofillia, Nuovo Giom. bot. ltal., N. S., xi, 1904. p. :220.-Viret, Liaisons du phloeme med..
etc., lnstitut de bot. Geneve, 1904, pp. 64-7o.-Theorin, Vaxttrichom., Arkiv fOr Bot., iii, n.. 5. 1904,
p. 13; and loc. cit., iv, n. 18, 1905, p. :n.-[Marcello, Ric. anat. pre!. sulla Cyph"mandrctflelacea,
Sendtn., Boll. Soc. Nat. Napoli,, xix, 190<), pp. I4:2-9.]--[For further
I
literature, see p. 117:2.]

SCROPHULARINEAE (pp. 583-589).

2. STRUCTURE OF THE LEAF. The peculiar hydathodes of Herpestis
Monnieria, H. B. K., which are described by Boergesen and Areschoug and
consist of a small conical cell situated above an ordinary epidermal cell, require
further investigation. The same applies to the stalked club-shaped bodies
resembling cystoliths, observed ,by Ward and Dale in Crate1'ostig~ pumiJum,
Hochst., where they are found attached to t~e outer wall of the epIdermal cells.
Isolated stomata are fonnd on the scale-leaves in the parasitic form, Hyobanche
sanguinea (Chatin).
According to Heinricher protein~crystals are found embedded in the nuclei
in all the organs of Lathraea and Tozzia; in Lathraea they occur also in the
protoplasm and in the leucoplasts. Red granules of a substance not quite
identical with carotin have been demonstrated in Craterostigma pumilum;
they are chiefly found as a lining to the intercellular spaces in the cortex of the
root, "being the cause of the coral-red coloration of the latter, but they also
occur in the axis in the same position as in the root and in the lower half of
the mesophyll of the leaf. Rothert's and Zalenski's (Bot. Centralbl., 1899,
iv, p. 246) statement as to the presence of styloids in the pith and cortex or
Phyllopodium (' P. rigidum ') still awaits confirmation.
Glandular hairs of the ordinary type are present also in Peliostomum
(Weber).
3. STRUCTURE OF THE AXIS. In Capraria bi/lora, L. Ph. Wagner records
subepidermal development of the cork, isolated groups of bast-fibres in the
pericycle, and groups of stone-cells in the pith. In Herpestis ItJo1mieria Duval
mentions the occurrence of peculiar mechanical cells situated in the lacunar
tissue of the primary cortex; these elements are parenchymatous in shape
and are provided with ridge-like thickenings. Accordj.ng to Weber the peri.
cycle contains isolated bundles of fibres or a more or less interrupted ring of
fibres in A ptosimum and Peliostomum, while in some of the species of Peliosto-
mum (P. origanoides, E. Mey., P. virgatum, E. Mey., P. viscos'ltm, E. Mey.) sub·
epidermal groups of fibres are found in the tips of the branches, although not
of constant occurrence even in one and the same species.
,!.
Literature: Costantin, Tiges a.er. et sout., Ann. sc. nat., ser. 6, t. xl'i,-,r8S 3 lZS et s~.­
Costantin, Tiges d. pl. aquat., Ann. sc. nat, ser. 6, t. xix, 1884, p. 287 et seq. an pI. xvi, xvh.-
Scherffel, in liot. Zeit., 1890, p. 417 et seq.-Stock, Proteinkrystalle, in Cohn, Beitr." vi, 1893.
p. :2 13 et seq.-[Breithaupt, Struct. of Lejlandra, Phann. Jonm., lxix, 1897, n. 5 (Veronica
1Ji,.gin;ca, L.); abstr. in Just, 1897, ii, p. 54.]-Schobert, Parenchymscheiden, Bot. Centralbl., 1897,
 ADDENDA-SCROPHULARINEAE
iv, p. 63.-Ph. Wagner, Neuere Drogen~ Diss., Erlangen, 1897, pp. 35-4-7 (Capran'a bijlora).-
WcUU'owsky, Scheiteloffn. bei Wasserpfi. t Diss., Berlin, 1898, p. 33 (Veronica Anag-allir).-Hirsch,
Entwickl. d. Haare~ Diss., Berlin, 1899, p. ~8.-Lavadoux, Appareil pilifere des Verbascees, Joum.
de Bot. 1899, pp..:U6-18.-Minden, Wassersez. Org., Bibl. bot., Heft 4 6 , 1899, p. 36.-Ward and
Dale, Craterostlgma pumilum, Transact. Linn. Soc., v, 1899, pp. 343-55 and pI. xxxiv, xxxV.-
Heinricher, Eiweisskrrstalle bei Lathraea, in Pringsheim Jabrb., xxxv, 1900, pp. 28-46.-Keamy, in
Contribut. U. S. Nat. Herb., v, n. 5, 1900, p. 305 (Herpesti$ .II/I11IIli'eria).-Thorn&S, FeuiHes sont.,
These, Paris, I9oo.-Heinricher, Anat. d. To:;zia, in Pringsheim Jahrb., xxxvi, 1901 , pp. 713-17.-
Pitard, Pericycle, These, Bordeaux, 19°1, p. 8g.-Areschoug, Mangrovepfl., Bib!. bot., Heft 56,
1902, pp. 78, 79, Tab. viii.-SperIich, Inhaltsst. in den Saug-org. d. gr. Rhinanthac., Beih. z. bot.
Cenltalbl., xi, 190~, p. 437 et seq.--[Armari, Piante della reg. medit., Ann. di Bot., i, rgo3, p. 17 et
seq. (Linaria).]- Theorin, Vaxttrichom., Arkiv for Bot.. i, 1903, p. 15.5.-Col, Faisceaux, Ann. sc.
nat., ser. 8, t. itx; 1904, p. lIl.-Freidenfeldt, Anat. Bau d. Wurzel, Bibl. bot., Heft 61, 1904,
pp. 71-3.-SUssenguth, Bebaarungsverh. d. Wunb. MUllcbelkalkpfi., Diss., Wiirzbnrg, 1904, pp. 45,
46.-TheDrin, Vaxttrichom., Arkiv for Bolo, iii, n. 5, 1904, pp. 16 and 28.-Daguillon, Ltnaria
striato·f)U19aris, Revue gen. de bot., 1905, pp. 508-18.-Porsch, Sp;lltotlnnngsapparat, Jena. 1905,
pp. 65-B.-Weberbauer, Veget. d. Hochanden Perus, in Engler, Bot. Jahtb., xxxvii, 1905, p. 60 et
Ik.~.-Dilval, Jaborandis, in Perrot, TravlllIx, iii, 1906, pp. 90-4 (.flcrjertis).-[Bohny. IJigitalis-
Blatt Diss., Zurich, 1906.]-Piccioli, Legnami, Bull. Siena, 1906, p. I76.-E. Weber, Aptosimu·m
und Peliostomum, Beih. z. bot. Centralbl., xxi, Abt. 2 ; also Diss" Zurich, 1906, 101 pp., 3 Tab.
-Sperlich, ZelIkernkrystaUoide VQn Alecfor%p!zus, lkih. ~. bot. Centralbl., ui, Abt. r; [907,
p. I et seq.-Kranzlin, in Pflanzenreicb, Heft 28, 190~, p. 2.-[H\lchede, Veroniques et Gratiole.
These,l)aris, 1907, 137 pp. ; also in Perrot, Travaux, v.]

OROBANCHACEAE (pp. 589-591).

Among the Orobanchaceae, which havt a single ring of vascular bundles
In their axis (d. p. 590), we may include Aphyllon uniflo7um, Gray, in which
the bundles form a more or less continuous ring (A. C. Smith). Cooke and
Schiveley's statement as to the occurrence of bicollateral vascular bundles
in the stem of Epiphegus virginianus requires confirmation, the more as the
authors named did not observe distinct sieve-plates even in the outer phloem.
Contrary to the earlier statement, stomata are found on the floriferous
axis (although not on the leaves) in Conopholis americana; they are also
present in A phyllon 'ttnitlorum, where they occur on the axes, bracts and
floral organs. I
Literature: Ducharte, Anat. de l'Orobancne Eryngii, Ann. sc. na.t., ~r. 3, t. iv, 18....5. PP.74-9.
-Lory, Resp. et street. d. Orobanch., Ann. sc. nat., ser.3, t. viii, 1847, pp. IS8-72.-Wilson.
C(lftopho!is af!l~ricana, Contribut. Bot. Laborat. PennsyJvania, ii, n. I, 1898, pp. 3-I9, pI. i-vi.-
A. C. Smith, Apkyl/on unijlorum l Contribut. Bot. Laborat. Pennsylvania, ii, n. 2, 1901, pp. 1 I 1-2 I,
pl. xiii-ltv.-Cooke and Schiveley. Epipkegus virginiana, Contribut. Bot. Laborat. Pennsylvania,
ii, n. 3, 19°4, PP. 352-98. pl. xxix-xxxii.-Porscn, SpaltOffnungsapparat, Jena, 1905, PP. 60-5.

LENTIBULARIEAE (pp. 591-598).

To the review of the anatomical features we may make the following
additions. In connexion with the remarks on the uniform type of structure
shown by the. glan dular _hairs1 we-may note that in place of a single stalk-cell
there may be-several. Dissociation of the vascular bundles of the stem into
xylem .. and phloem-groups, which are independent of one another, occurs also
in Polypompholyx 1.
PINGUICULA (pp. 59I, 59z)
1.
The structure of the leaf in P. vulgaris has recently been examined by Fenner.
In rectification of the earlier statements we may nrst mention that chlorophyll
is found in the epidermis of the leaf in the neighbourhood of the midrib, and that
stomata are present on both sides and are even more abundant on the upper side;

1 For Byhlir, which has been tmnsferred to the Lentibularieae by Lang, see under Droseraceae,
Po 9 11 •
SOLERI!:DltR
{QIO ADDENDA-LENTIBULARIEAE
they are absent only from the marginal rones on ooth surfaces of the leaf. The
glandular hairs occurring on the upper side of the leaf are either stalked or sessile.
The stalked glands, which function as organs for the capture of the insects, consist
of (r) a basal cell, (2) a stalk composed of one to three cells, (3) a columella~ceU,
and (4) a head comprising 16 radially arranged cells. The shortly stalked glands
have a basal cell, a stalk-ceIl, and a glandular disc of 4 or 8 cells; these are the glands
which perform the actual Erocess of absorption. The same function is carried on
also by one to four rows of upper epidermal cells lying next to the ma.rgin of the
leaf and styled glandular celIs by Fenner. The lower side of the leaf only- bears
small external glands, which are provided with a head composed of four. cells and
function as hydathodes. Features deserving special mention are the sieve~like
perforation of the cuticle of the glandular head, and the fact that tracheae of the
vascular system extend up to all the glands (both sessile and stalked) on the upper
side of the leaf. Multicellular trichomes, which do not have a secretory function,
occur on the petiole and on the base of the lamina; these hairs either consist
of a. row of 3-6 cells or bea.r a rudimentary head, which is unicellular or composed
of a smaIl number of cells; the nuclei in these hairs contain crystalloids. According
to Russow and Klein, moreover, nuclear crystalloids are found also in the epidennis
of the leaf, &c. i

2.GENLISEA (pp. 592-594).

For the structure of the ascidiform leaves of G. vialacea, St. Hit.. which is
essentially the same as in G. ornata, see Goebel, Ioc. cit.

3· UTRICULARIA (pp. 594-597)'

_ The most important point in Meierhofer's recent investigations on the anatomy
of the bladders is the statement that there is no stalk~cell in the two~anned glandular
hairs (with upwardly directed arms) occurring on the inner wall of the lower lip
(which faces the interior of the bladder) and in the four:rayed absorptive glands on
the • inner wall of the bladder.' AccordingtoMeierhofer the apparent stalk in the
two types of hairs is-in Utricularia vulgar"s at least-formed by the lower portions
of the 2 or 4 ray~ceIls, which are narrowed so as to resemble a stalk. According
to the same authority. moreover, thelid in U. vulgaris, which, like that of U. {lexuosa.
is composed of two layers of cells provided with peculiar sti:ffenin~-mechanisms.
bears glands which for the most part consist of three cell!5, the head bemg uIDcellular
and of varied shape, viz. spberical, ellipsoidal or like the terminal portion of a two~
armed hair. In some cases the head of these glands is bicellular owing to the
presence of a vertical wall. or the stalk~cell (joint-cell) undergoes secondary division
lnto four cells by means of vertical walls. The four~armed absorptive glands,
which occur on the inner wall of the bladder. may baused as a means of distinguishing
the species; in Utricularj'a BremiJ', Heer they show the same arrangement of the
ray~cells as in U. minor (see Fig. 139. D, on p. 596), while in U. intermedia, Hayne
the ray-cells are arranged in pairs in two parallel rows in the same way as in U. mon-
tana (Fig. 139, E). On the surface of the bladder of U. £ntermed£a Meierhofer
observed glandular hairs with 4-celled heads in addition to those having z ..celled
heads.

4. POLYPOMPHOLYX (p. 597)

The anatomy of this genus ci the Lentibularieae was hitherto little 'known j
two Western Australian species, P. multifida F. v. MUll. and P. lenella, Lehm.,
f
have. however. recently been examined by Lang. The two species agree with one
another as regards thelI' anatomy.
In the structure of the axis the vascular system is specially distinguished
by the absence of proper vascular bundles, the xylem~ and phloem~groups running
independently of one another as separate strands. The xylem~ and phloetn~bundles
are present m equal numbers, the former consisting only of one vessel apiece.
while the phloem~bundles comprise a group of small cells including sieve~tubes.
Both kinds of bundles are inserted in a ring of sclerenchyma composed of cells
resembling fibres; on their inner side these cells merge into a pith rna-de up of
large cells. The bundles alternate with one another like the wood- and bast~groups
 ADDENDA-LENT/BULARIEAE 1011
~

in the radial vascular system of a root, the phloem-bundles lying at the periphery'
of the sclerenchrmatous ring and bordering on the sheath of large parencliymatous
cells which adjOInS the ring of sclerenchyma on its outer side. The primary cortex
contains intercellular spaces which, near the points of insertion of the leaves and
stolons, are of large size and are arranged in a ring; they are separated from one
another by lamellae composed of a single layer of cells, and border on the su~
epidermal layer externally and on the sheath of large parenchymatous cells interna.llv.
The vascular system of the stolons consists of a single bundle showing concentric
structure. the middle being occupied by a. solitary vessel, the place of which is
subsequently taken by an intercellular space.
The foliage-leaves have epidermal cells (occasionally containing chlorophyll)
with more or less undulated lateral walls and bear stomata. on both sides. The
meso:phyll is homogeneous and is provided with cavities which are particularly
proaunent on the lower side and attain considerable width in the neighbourhood
of the petiole. The solitary median vascular bundle of the leaf contains only
one vessel and is-at least at certain points of its course--collateral. The ~
covering of the leaf is constituted by external glands consisting of three cells. viz.
a:sunken basal cell, a stalk-cell. and a capitate cell secreting mucilage.
For the shape of the ascidiform leaves or bladders, see p. 597. The wall of the
bladder cop.sists of four layers of cells and is traversed both on tru; dorsal an4
ventral side by a well-differentiated vascular bundle, running in the median plane.
The outer wall of the bladder bears the ordinary 3-celled glands, while at points
situated opposite the ciliate margin of the cap one finds bristle-hairs like those
occurring on the latter and consisting of a basal cell, a short stalk-cell and a long
terminal cell. The inner wall of the bladder is occupied by the same characteristic
four-armed absorptive hairs, as are found in Utrt'culayia; in the species examined
by Lang the arms of these hairs are arranged in the form of an x. The Wing-like
extensions of the cap have the ordinary external glands on their outer sunace.
while the inner surface bears numerous glands secreting mucilage and exhibiting
all transitions between the ordinary capita.te glands and glandular hairs with
fiagelliiorm terminal cells; the terminal cell may be spherical. ellipsoidal or clavate
or in extreme cases flagelliform. The inner wall of the stalk, which here likewise
participates in the formation cf the wall at the entrance to the bladder. also bears
glands with terminal cells, which are flagelliform and are often bent in the form
of a crook; in addition to these one finds glands with two-armed .terminal cells, the
arms being placed parallel to the surface. The inner wall of the 'lid: which
is composed of two layers of cells, strengthened by means of annular, or s~al
thickening bands, and the inner wall of the 'lower lip' (' Widerlager ') are likewise
occupied by numerous fJ.agelliform mucilage-hairs. On those parts of the lip. whicb
are sItuated nearer the interior of the bladder, the mucilage-hairs are replaced by
glands with two-armed panduriform heads divided by a. transverse wall into two
<:ells, while the place of the mucilage.hairs on the lower lip is taken by a tessellated
epithelium composed of closely -crowded capitate glands of the ordinary type.
Finally, on that part of the lower lip, which faces the interior of the bladder, we
-find the same two-armed glands provided with upwardly directed arms, as occur in
UtriculaYia {lexuosa, &c.
Literature: J. Klein, ZeUkeme voD Pinguicula u. Vlr-icularia, Bot. CeJ].tralbl., 1880, iii,
pp. 1401-4.-Russow, Krystalloide bei. Pinguicula, SiU.-Ber. Dorpat. natnrf. GesellJch., 1880. PP.417•
... IS.-Goebel, ,Genlisca, Flora, 1893. pp. 208-Il.-[Endo, Ulricularia IJifoJa, Bot. Magu. Tokyo.
xii, J898, pp. 1-4. I p1.)-Weinrowsky, Scheiteloffn. bei Wasserptl., Diss., Berlin, 1898, p. ~6.­
Meister, Ulricularia, Bull:Herbier BoisSier. 1900, n. 12,40 pp., 'ftlb. i-iv.-Lang, .Po1y~",pMlyx
u. B,oli.s, Flora~ 1901, 'Pp. 15,,-66, Tab. xii.-Meie-rhofet, Utricularia.Bla.im, Flora, 1902,
pp. 84- 11 3. Tab. ii-v.-Dutailly. in Assoc. fran~. Ajaccio, 1902) p. 4-S7.-Fenner. Laabbl. u. Drt1Jen
~iniger Insektivoren, Flora, J904, p. 335 et seq.; also as Diss" Ziirich, especially pp. 6-20,
'Tab. vi-viii.

COLUMELLIACEAE (p. 598).

Van Tieghem has recently examined the remaining knowri species, viz.
Columellia oblonga, Ruiz et Pav., C. obovata, Ruiz et Pav. and C. seficea, H. B. K.
In these species he records the following features which agree with those observed
by me in C. serrata, Rusby:-uniseriate medullary rays in the wood; peri-
3T2
 ADDENDA-COL UMELLIA CEAE
cYClic development of tne cork, WnIcn has the same composition as in C. serrata;
absence of secondary hard bast; clustered crystals of oxalate of lime; bifacial
leaf-structure; stomata confined to the lower side of the leaf and without
subsidiary cells; simple unicellular clothing hairs adpressed fu the surface
of the plant; noglandular hairs; and lastly, in all the species, a single layer
of hypoderm on the upper side of thp. leaf.
The following details may still be added. The mode of development of
the cork in Columellia obovata recalls that in Dodonaea (Order: Sapindaceael.
The pericycle js parenchymatous. One of the outer layers of pericyc1ic cells
gives rise to the first phellogen and forms a periderm on its outer side in which
there is a regular alternation of layers of suberized cells, showiag marked
radial elongation, and small fiat cells with cellulose walls, which subsequently
become sclerosed. This is followed by the development of a first continuous
ring of fibres from pericyclic tissue adjoining the cork-cambium; the first
phellogen then ceases to be functional and a second one arises on the inner
side of ~he ring of fibres from tissue belonging .either to the pericycle or to
the bast; this second phellogen produces a new periderm, ,tl!e structure of
....,hich is idl(ntical with that of the first-formed one. While the second peri-
derm is in courSe of formation a second ring of sc1erenchyma is differentiated
on the inner side of the second cork-cambium; 'internal to this sclerenchy-
matous ring a third cork-cambium is produced after the activity of the second
has been brough.t to a close, and so on. In the other two species Van Tieghem
observed noteworthy modifications; thus in C. sericea the (first) ring of pericyc1ic
lJ.uieS is present before the development of cork commences, while in C. oblonga
it continues to thicken for some considerable time before the second phellogen
appears. In the three species, which he examined, Van Tieghem also records
an endodermis composed of large cells with lignified radial walls and situated
on the outer side of the peri cycle. The leaf is supplied by a vascular bundle,
taking the form of an arc of wood and bast. The vascular bundle found in the
median ·vein of the leaf is provided with an arc of fibres.
Literature: Van Tfegbem, Cohunelliac«s, Bun. Mr;s. d'hist. nat., (903, pp. '33-9, and Ann.
St. nat.,ser. 8, t. xviiit 1903. pp. 155-64.

GESNERACEAE (pp. 599-601).

To the SUMMARY OF THE ANATOMICAL FEATURES of the Gesneraceae~we
may add the following details. Typical bundles of raphides occur in N apean.
ihus repens. J. D. Smith (genus novum Gesneracearum); true cystoliths have
been observed in the two very closely related genera Klugia and Rhynchoglossum;
secretory canals, developed in relation to the vascular bundles of the leaf and
stem, as well as medullary vascular bundles in the stem, are found likewise in
Klugia and Rhynchoglossum, the medullary bundles being present also in Mono-
phyllaea and Coleus; a single cortical vascular bundle occurs in the stem in
RhynchoglossUtn oblit{uum, Bl.; stomatal groups are met with on the surface of
the leaf in the specles of the genus Napeanthus; in addition to that certain
species have hypoderm in the leaf and spicular fibres in the mesophyll (the laUer
in Stau7antheTa Rrgyrescenst Hallier f.). As regards the clothing hairs we may note
that in the uniseriate trichomes the thick-walled terminal cell, and in some cases
also the cell adjoining it, have calcified and silicified walls. Uniseriate trichomes,
in which the terminal cell is bent like a hook, occur in Epithema and Loxonia ;
simple shaggy hairs are found in certain species of Klugia and branched multi-
cellular trichomes in Klugs'Q zeylam:ca t Thw. The heads of the glandular hairs
are occasionally unicellular. Special mention may be made of the forked
external gla.nds of P,imulina Tabacum t Hance, and of the • peltate glands'
of Monophyllaea, which excrete chalk.
 ADDENDA-GESNERACEAE

The STRUCTURE OF THE LEAF is again bifacial in the species which I have
recently examined. In the species of Jeydonia and Napeanthus the cells of
the spongy tissue are rounded and are peculiar in being connected with one
another by means of short peg-shaped processes. According to K. Fritsch (1oc.
dt.), m.oreover, similar .features are found also in species of Monophyllaea,
Streptocarpus and Saintpaulia. There is no sclerencbyma in the veins in the
species which I have examined; the larger veins commonly (species of Jer-
donia, Klugia, .Loxonia, Napeanthus, Rhynchoglossum) include two collateral
vascular bundles, which either both show the same orientation (with the
xylem on the upper side) or have their xylem-groups directed towards one
another. The spicular fibres mentioned above as occurring in the mesophyll
in Stauranthera argyrescens were observed by Hallier. In Cyrtandronwea
decurrens, Zollo the epidermal cells on the upper side of the leaf are provided
with a small papillose protrusion, arising from the middle of the outer wall.
In Saintpaulia ionait~ha the upper epidermis likewise serves for water-storage
and consequently consists of large cells (Fritsch); in the species of Napeanfhus
both the upper ~!ld lower epidermis show the same feature, the two epidermal
layers here occupying the greater part of the transverse section of the leal.
Hypoderm is found also in Codonanthe Devosiana (according it> Schim~er),
Aeschynanthus purpurascens, Hassk. (according to my own observahon;
here comp,?sed of several layers ap.d taking up half the thickness of the leaf),
Monophyllaea Horsjietdii, R. Br. (according to Fritsch, two-layered), and
Streptocarpus Kir.kii, Hook. f. (according to Fritsch). In Saintpaulia ionantha
and Aeschynanthus purpurascens the spongy tissue likewise stores up water.
The stomata are often of large size. In N apeanthus an excellent generic character
is afforded by the stomatal groups, which are often visible even to the naked
eye as small spots on the lower side of the le-af and .recall the well-known
stomatal groups found in certain Begonias. The stomatal groups of N apeanthus
are constituted mainly by 2-18 pairs of guard-cells together with their sub-
sidiary cells, and are absent only in N. repens, J. Donn. Smith, which does not
belong to the genus Napeanthus (see above).
Regarding the wide distribution of anthocyanin in the Gesneraceae, see
Hassack and K. Fritsch, 11. cc. I
I have observed rather long acicular cry:;tals of oxalate of lime also in
Phylloboea amplexicaulis, Clarke, where they completely fill the cells of the
spongy tissue; small but rather distinctly"'differentiated clustered crystals
occur side by side with other small crystals also in the palisade tissue of
Cyrtandromoea decurrens; relatively large solitary crystals are found in the
epidermis of the leaf of Primulina Tabacum, and in the palisade tissue of
Acanthonema strigosum, Hook. f.; large crystalline bodies, which refract the
lighCdoubly and give the reactions of oxalate ot lime (though still requiring
more detailed .investigation), are present in the subsidiary cells of the hairs
and in enlarged cells of the mesophyll in Acanthonema strigosum; lastly,
typical bundIes..._of raphides. have "b~en observed" in spindle-shaped cells in
Napeanthus repens (genus novum).
Cystoliths have been found among the Gesneraceae in the two genera
Klugia and Rhynchoglossum 1. The cystoliths vary in shape, being ellipsoidal
or more elongated or slightly branched, while in Klugia ampliata, Clarke
they have a pec~iar' racemose form; they mayor may not be calcified, and
are frequently provided with distinct stalks. In Klugia zeylanica, Gardn.
(' Klugia Notoniana,' Hortorum) they are found principally in the palisade
tissue, but occur also in the spongy tissue and in the lower epidermis, while

1 K. Fritsch's paper (' 'Uber das Vorkommen 'Von Cystolithen bei Klugia zeylanica' in Wiesner·
Festschrift, Vienna, 1908, p. 412~ \vas nnh1io;hed while these sheets were in the press.
101 4 ADDENDA-GESNERACEAE

in Klugia azurea, Schlecht." K. major, SoIered. and K. Notoniana, Gardn. p

as well as in Rhynchoglossum obliquum, Bl. (with var. p(l.rvif/,ora, Clarke) and

R. klug·ioides, Clarke they are confined to the lower epidermis, and in Klugia
ampliata, Clarke to the palisade tissue. I have examined the genera N ape-
anthus, Loxonia, Phylloboea, Cyrtandromoea, Jerdonia, PIa tystemma , Acantho-
~ma, and Primulina, which are placed immediately next to Klugia and
Rhynchoglossum by Bentham and Hooker and other authorities, and find that
they do not possess any cystoliths.
Another new anatomical feature, not hitherto recorded in this Order,
is constituted by the resin-canals found in Klugia and Rhynchoglossum.
I was able to demonstrate their presence in all the species with the exception
of Rhynchoglossum klugioides" Clarke, of which only a badly preserved fragment
of a leaf traversed by Fungi was available. The secretory canals have
lumina of varying width and are occasionally surrounded by cells resembling
an epithelium; their contents are of a resinous nature. They are found in
-the first place in the veins of the leaves;i in the larger veins they are
situated between the two vascular bundles (see above), while in the smaller
ones they lie above the xylem of the solitary bundle. In the stem the
secretory canals in part accompany the medullary vascular bundles and the
bundles of the vascular ring, being placeQ next to the xylem; but they
frequently have an apparently independent course, and run sdme considerable
distance internal to the xylem-groups of the vascular ring.
The hairy covering of the Gesneraceae has recently been examined by
Rechinger, more especially with reference to the uniseriate clothing hairs,
which are provided with strongly thickened terminal cells. According to him
the strong thickEminglof. the terminal cell is due to an obliterating membrane
r Ausfiillung '), which is both silicified and calcified. Trichomes of this type,
showing obliteration of the lumen in the terminal cell, do not, however, occur
in all the members of the Order. In certain species uniseriate hairs are
altogether wanting; in other cases one finds uniseriate hairs in which the
terminal cell shows no -such obliteration of the lumen, while the remaining
cells of the hair have walls of varying thickness. Trichomes, in which both
the terminal cell and the cell adjoining it, and in rare cases even the third
cell from the apex of the hair, show obliteration of the lumen, while the
remaining cells have thin walls, were observed by Rechinger in certain
species of Alloplectus, Corytholoma ( = Gesnera I), Gesnera, N aegelia, Pearcea
(= Isoloma), Roettlera" (= Chirita), Sciadoc_alyx (= Isoloma). Sinningia,
Streptocarpus, and Tape~'notes (= Ss'nningia); trichomes, in which the mem-
branes of all the cells are thickened and the lumen of the terminal cell is
obliterated, were met with in certain species of A chimenes, Alloplectus, Centro ..
solenia (= Episcia), Drymonia. Jancaea (= Ramondia), Isoloma, Penta-
rhaphia, Ramondia, Rottllera (= Didymocarpus). Streptocarpus and Tydaea
(= Isoloma). The spec~al structure of the ordinary uniseriate clothing hairs,
which has just been discussed, is in general only of v-alue for specific diagnosis.
_According to Rechinger the same is true also of other structural features shown
by these hairs, such as nodular thickenings on the walls (A chimenes lanata,
lianst.) or nodose swelling of the basal portions of the individual cells (Boea
hygrometrica, R. Br., Jancaea Heldreichii, Boiss.). Among the members of
the' Order, which I have recently examined, Napeanthus repetfs alone has long
clothing hairs with a terminal cell showing obliteration of the lumen, but in
species of Acanthonema and Platystetnma a slight incrustation may be detected
at the apex of the terminal cells. Other peculiar types of hairs are: (a) the
very short pointed hairs of Napeanthus apodemus, J. D. Smith, N. subacaulis,

1 According to the Index Kewensis.

 ADDENDA.-GESNERACEAE 1015

Benth. et Hook., &c., which are composed of from one to three cells and are
seated on the middle of large epidermal cells; (b) the short clothing hairs
found in species of Klugia and Rhynchoglossum, which consist of a small number
of cells, the terminal one of which is somewhat bent, and are inserted obliquely
on the surface of the leaf; (c) the very long uniseriate clothing hairs present on the
margin of the leaf in Jerdonia indica, Wight, some of the long segments com-
posing these hairs being septate by means of thin transverse walls (to the number
of 6 or 7). According to H. Schmidt the uniseriate bracket-hairs already referred
to above, in which the terminal cell is bent in the form of a hook, are found
in Epithema Brunonis, Decne. and E. carnosum, Benth.; they constitute
a good distinguishing feature between Epithema and the genus Pentaphragma
(Campanuloideae), which is provided with branched clothing hairs; and is
often confused with Epithema in herbaria. I have observed bracket-hairs
also in Loxonia acuminata, R. Br. Uniseriate trichomes, in which each of
the lower cellS is divided longitudinally into two, occur in Klugia Notoniana,
and constitute a transition to the shaggy hairs, which have been demonstrated
in K. azurea. Schlecht. and K. major, Solered. Klugia zeylanica, Gardn. (' K.
Notoniana • Hortorum. see Rechinger, loco cit.) is distinguished from the true
K. Notoniana by the possession of more or less richly branched clothing hairs;
th,e lower part of the stalk in these hairs is frequently biseriate and, like
the uniseriate branches, consists of cells with thin walls; I did not see the
transparent (' vitreous ') unicellular cilIa observed and figured by Rechinger,
according to whom they are attached to the walls of the hairs with their ends
pointing downwards (strange bodies ?)1. Glandular hairs have recently been
demonstrated also in species of Acanthonema, Achimcnes, Boea, Cole·lts, Col-
landra (= Columnea), Cyrtandromoea, Ep£thema, Gloxinia, jerdonia, Kltlgia,
Loxonia, M onophyllaea, N apea1zthus, Phyllol;oea, PIa tys temma , Plimulina,
Rhynchoglossum, Roettlera (= Chirita), Saintpaulia and Tydaea (= Isoloma}
(Rechinger, H. Schmidt~ Chevalier et Perrot, Solereder). V.fe- may add the
following details regarding the differentiation of the head. Unicellular spherical
glandular heads are found in species of Cole1tS, where they are borne on very
short s ta!ks t ~.nd in N apeanthus repens, where they are placed on very long
stalks; long glandular hairs having cylindrical terminal cells, the apex of
which is swollen or lageniform, occur in Prim1elina T abacum, Hance. This
::;pecies also possesses the forked glands mentioned above; they are long
trichomes with a uni·, Of rarely bicellular,""'stalk and two cylindrical terminal
cells~ which show a pistillate swelling at their apex, are fused for about half
their length and diverge beyond that point in a dichotomous manner. In the
members of the Order which I have examined the heads of the shortly stalked
gla1'l.cs consist of from two to four cells. In the simplest case the bicellular
head, as seen from above, has an elliptical outline, while the four-celled head
has a rounded or rosette-like outline; as a result of elongation of the glandular
cells we obtain glandular heads, which are either bicellular and panduriform
Of hammer-shaped~,or- two-armed (Epithema, Klugia, Loxonia, Napeanthus,
Rhynchoglossum) or on the other hand are provided with three or four ra}'"S
(Klugia, Napeanth-us, Rhyncnoglossum). The peltate glands of Monophyllaea _
Hot'sjieldii, R. Br., which were referred to above, belong to a special category;
as Fritsch was the first to show, they excrete carbonate of liILe. I have
examined them and find that their strncture corresponds completely with that
of the. well-known pelt ate glands of the Rhinanthaceae; the epidermal celi,
1 The I sc91cs' described by Rechinger as occurring on the leaf oi Mbltoplzyllaea Horsfieldii, R. Br.
have been shown by Fritsch (loc. cit.) to be sr.'laH chalk-scales, which nre excreted by peculiar
glandular hair5 functioning as bydathodes (for details regarding the -hairs, see the text below).
Rechinger'li fnrther statement as to the occurrence of small scales in 'Boea speciosa' still requires
reinvestigation.
 r016 ADDENDA-GESNERACEAE

which constitutes the base ot the trichome, is followed by a low middle-cell which
has a rounded out1ine and bears the shie1d; the latter is of the same size as
the middle-cell and is divided into two' lid-cells' by means of a vertical wall.
For the occurrence of steles in the fibrovascular system of the petiole
of ' Ges1'teria alba" see Bouygues, loco cit.
According to my own investigation medullary vascular bundles are
fOllnd both in K.lugia (K. ampliata, Clarke, K, azurea, Schlecht., K. malor,
Solered., K. Notoniana, Gardn., K. uylanica, Gardn.) and in Rhynchoglossu~
(R. obliquum, Bl. with var. parvi/lorum, Clarke), while according to K. Fritsch
they occur! a150 in M onophyllaea H ors/ieldii, R. Br., a species which has
only one large foliage-leaf. In Rhynchoglossum obliquum (also in the var.
parvi/lorum), moreover, a projecting ridge on the stem contains a cortIcal
vascular bundle showing concentric structure.
For the structure of the characteristic runners of Aclu'menes coccinea,
which are occupied by scaly leaves, see Kliem, loco cit.; for the structure of
the tubers found in species of Coleus, see Chevalier and Perrot, loco cit.; for
the structure of the aerial and terrestrial roots, occurring in species of Aeschy-
nanthus and Columnea, see Keller, lac. cit. The only feature requiring special
mention is the occurrence of meduUary vascular bundles showing collateral
structure in the tubers of Coleus langouassiensis, thevalier.
Literature: [Pedicino, Sc1erenchymi nelle Gesnerac., Rendic. ACCIld. sc. fis. e mat. Napoli, 1879.
fase. :I; abstr. in Just, 1879, i, p. 24.]-Hassack, Anat. Bau bunter Laubbl., Bot. Centralbl., 1886,
iv, p. 84 et seq.-L. Keller, Luft.wurzeln, Diss., Heidelberg, 1889, pp. I::J-15.-Hallier, in Ann.
Jardin Buitenzorg, xiii, 2, 1896, p. 288.-Jonsson, Anat. .. Bau des Blattes, Acta. Univ. Lund .•
xxxii, 2, 1896, 20 pp.-Rechinger, Trichome der Gesnerac., Osterreich. bot. Zeitschr., 1899, p. 89 £t
seq., sep. copy, 18 pp. and Tab.-Kliem, Vegetat. Regenerationsorg., Diss., Erlangen, 1900, pp. 49,
50.-Bouygues, Petiole, These, Paris, 1902, p. 78.-Pischinger, Stnptocarpus and JllonophylltUa,
Sitz.-Ber. Wiener Akad., cxi, Abt. I, 1902, pp. 287,291, etc. and Tab. i, ii.-K. Fritsch, Keim-
pflanzen der Gesnerae., Jena, 1904, especially pp. 132-47.-H. Schmidt, Campanuloideen, Diss.,
Erlangen, 1904, pp. 101, I02.-Chevalier et Perrot, Coleus, in Perrot, Travaux, iii, J 906; also in
Vegetaux utiles de l'Afrique trap. fl'an~., t. i, 190:;, pp. 100-5:1, pI. 1-8, especially p. -130 et seq.-
Solereder, Zur Anatomie u, Systematik einiger Gesneraceen-Gatt., Manuscript, 1907.-[K. Fritsch, ill
Natiirl. Pflanzenfam .. Erg.-Heft II, 1908, p. 31 i.]-K. Fritsch, Cystolithen bei KINgia Eey/anica, in
Wiesner-Festschrift, Vienna, 1908, p. 412.
\'
BIGNONIACEAE (pp. 601-611).
I:To the REVIEW OF THE ANATOMICAL FEATURES we may add that stomata
having two subsidiary cells placed transversely to the pore also occur in this
Order (Kigelia). Among the list of special ~eatures we may include the occur-
rence of groups of silicified cells in the. mesophyll and of solid papillae on the
epidermis of the leaf (both features likewise observed in Kigelia).
2. STRUCTURE OF THE LEAF. According to my own investigation the
mesophyll is centric in structure in Kigelz'a p,'nnata, DC. and K. a/ricana,
Benth.; it consists of 5 or 6 layers of moderately elongated palisade-cells,
the longitudinal walls of which show a bellows-like f9lding, and at some points
include small intercellular spaces between one another after the manner of
conjugate parenchyma. Spicular fibres running freely in the mesophyU,
besides occurring in Colea Commersonil (where they may even spread out
beneath both upper and lower epidermis), are ·found in Colea pedunculata
(according to Fabricius) and in Crescentia regalt's, C. macrophylla (= Amphi-
tecna macrophylla, Miers) and Phyllarthron comorense, DC. (according to Cornu).
In Colea pedunculata there~ are also scattered stone-cells in the mesophyll.
Another noteworthy feature is constituted by spherical or otherwise shaped
groups of silicified cells, which were first observed by Corriu in the mesop[lyU
of KigeUa pinna/a, although his interpretation of them was not quite correct,
and which are present also in K. africana. 'Their structure recalls that of the
groups of silicified cells, figured-for Aristolochia acutilolia, Duch. in Fig. 166, G
 ADDENDA-BIGNONIACEAE rOl7
(on p. 684) in all respects. The constituent cells show an approximately radial
arrangement, while those parts of their walls, which are directed towards
the centre of the group of cells, and the adjoining parts of the radial walls
are thickened in the form of a horseshoe and are stratified and silicified. The
groups of cells in question either lie freely in the mesophyU or more commonly
OCcur in connexion with the vascular bundles of the veins, being situated both
on their upper and lower side; the number of cells composing the groups
varies. In Kigelia the upper and lower epidermis are likewise silicified; in
surface·view the cells of the upper epidermis have undulated lateral walls and,
like the epidermal cells situated above the larger veins on the lower side of
the leaf, are provided with blunt conical and solid papillae, while the lower
€pidermis consists almost exclusively of the exceptionally numerous stomata
with their ,subsidiary cells. The pairs of guard-cells in Kigelia are raised above
the surface of the epidermis, and are supported for the most part by two (more
rarely by three) neighbouring cells, which are placed transversely to the pore.
The hairy covering in /{igelia comprises unicellular clothing hairs with
rather thick walls and narrow lumina, and relatively large disc-shaped glands
showing radial arrangement of the cells and a slight elevation of the cuticle.
3. STRUCTURE OF THE AXIS. As regards the occurrence of anomalous
structure in the wood of the root in the Bignoniaceae (p.60S) we may note
that according to Hill bast·wedges are iound in this position.also in Bignonia
fmguis and B. vem~sta. -
The cortex of Oroxylmn indicum, Vent. contains deposits of green sub·
stances which resemble wax, and are the seat of the active principle (oroxylin)
in this bark (0. Werner). It still 'remains to be determined whether the rubber,
which exudes on wounding the cortex in Stereospermum euphorioides, is deposited
in special receptacles (see Jumelle, loco cit.).
Litemture: [Stowell, Folia Carobae (Jaccaymtda CaYObM), Therap. Gatette, N. S., 1881, h. ~,
p. 42; abstr. in Just, 188T. ii) p. G91.]-KeHer. Luftwurzeln, Diss., Heidelberg, 1889. pp. 15-17.-
[Tognini, Stomi, Aui 1st. bot. Pavia, 1894.J-Went, Haft- u. Nahrwur.zeln, Ann. Jardin Buitenzorg,
xii, 1895, pp. 59, 60.-Cornu, Co!ea jloribunda, etc., iii and iv, Bull. Soc. bot. de France, 1896,
pp• .4.06-14.-Jonsson, Anat. Ban des BI., Acta Vniv. Lund., xxxii, 2, IS96.-Werner, Neuere
Dfogen (Cortex Oroxyli), Diss., Erlangen, 1896, po. :27-33 and T:lb. iii, iv.-HiU, Roots of
Bi8nonia, Ann. of Bot., 1898, pp. 32'3-8 and pI. xxii.":"Bcergesen og Paulsen, Veget. dansk-vestind.
Oer, Bot. Tidsskrift, xxii, J898-9, p. 101 (Tecoma /eucoxy/on, Mart.).-Lcisering, Interxylares
Leptom, Diss,) Bertin. 1899. pp. 27, 28.-Pitard, Pericycle, .These, Bordeaux, 19~1, p. 98.-
[Anonymous, Glands in Tec()11la, Gardeners' Chronicle, xxxii, 1902, p. 44.1-Fabricius, Laub~
blattanatomie, Bein. z. bot. Centmlbl., xii, 190~, p. 313 (C()leatedftnculata),-Col, Faisceaux, Ann.
sC. nat., ser. 8, t. xx, 1904, pp. lIl, 112 and 152-4.-Paoli, Etcrofillia, Nnovo Giorn. bot. Ita!.,
N. S., xi, 1904. p. 220 (species of Bi_gJtonia and lUacrodis~-us).-:FtommE:I. Plantas text. chil., 1905.
p. 4o~-Jumelle, Une Bignon. a gomme, Comptes rend us Paris, 1905, l~r sem., pp. 17O-:.r.-
lPenhallow, Hybridization of Ca/.a/pa, Arneric. Nat., xxxix, 190~, pp. 1Ii\-.~6, abstr. in Bot.
Ce ntrn.lbt.! c-,:.ix, p. 32I.l-Pkcioli, Legnami, Bull. Sitll?, 1906, p. l~l.-Haberlandt, Sinnesorg.,
second edition, 1906.-[Schwendt, Extraflor. Nektarien, Beih. z. bot. Centralbl., nii, Abt. I, 1907,
pp. 25 6-9.1

PEDALINEAE (pp. 611-613).

GrUnewald's paper cited below deals principally with the anatomy of
the leaf and axis in the genera AJartynia, Cra1lio/aria, Pedalium, Pterodiscus,
Harpagophytum, Rogeria, Sesamum, Ceratotheca, Putrea, and Josephinia; it
does not contain any essentially new facts.
The characteristic mucilage· hairs appear to occur also in Pterodiscus
(' P. angustifolius '). whilst in Harpagophytum the external glands are of the
same type as those of Craniolaria annua. The structure of the leaf is for the most
part bifacial, the stomata generally occurring on both surfaces. In Crant:olaria
lln1ttta the epidermis of the leaf contains chlorophyll, and in Pre/rea eriocarpa
and' PterodisctlS angusti/olius' crystals of oxalate of lime. Oxalate of lime
1018 ADDENDA-PEDALINEAE

is occasionally found in the form of crystals resembling raphides (pith of

Rogeria adenophylla).
As regards the structure of the axis it appears that ba~t-fibres are invariably
found in the pericycle, and that the wood-prosenchyma always bears simple
pits.
Literatnre: Grunewald, Vergl. Anat. d. Martyniaceae u. Pedaliaceae, Diss., Erlangen. 1897.
43 pp.

ACANTHACEAE (pp. 6I3-624).

2. STRUCTURE OF THE LEAF. In the species which have been examined
the leaves are for the most part bifacial in structure, rarely (Acanthopsis horrida,
Nees) centric with palisade tissue' on both sides. Friedrich records a papillose
epidermis in 'the leaf in Aphelandra Fascinator, Lind. et Andre and Ruellia
Devosiana, Lindl. According to Holtermann a many-layered hypoderm is
found on the upper side of the leaf also in Strobilanthes sexennis var. argutus.
According to Dethan the parenchyma of the veins in Acanthus mol/is, L.
contains membrane~mucilage. At this point we may also mention the occur~
renee of groups of mucilaginous celis on both sides of the leaf in ilcanthus
ilicifolius~ L., a feature which still requires further investigation; on th,e
upp~r c;ide of the leaf these groups are inserted among the cells of the
bypoderm, while on the lower side they are formed mainly by epidermal cells
showing palisade-like elongation; in later stages they give rise to organs
resembling lenticels (Arescboug). ,
Oxalate of lime (d. p. 614) is found in the form of large clustered crystals
in Thunbergia lauritolia, Lindl. and Eranth'emum 'igneum, Lind. (Friedrich).
Small crystalline· bodies occur very abundantly in the epidermis and in the
clothing hairs. The acicular crystals are often differentiated like raphides
(see especially A. Weiss, loco cit.).
To the list of special forms of clothing hairs (5ee p. 6I4) we may add:
(a) the unicellular trichomes of Lasiocladus, which have thick walls and narrow
lumina; , (b) the llni- or multkel1ular la!lceolate hairs of ACdnthopsis hOTTida.
Neest each of which is seated obliquely on an epidermal cell, with the apex of the
hair directed towards the tip of the leaf; and (c) the trichomes of Fittonia
Verschaffeltii, which according to Haberlandt function as ocellae; these
hairs ronsjst of (1) a jarge epidermal celli which has a rounded outJine, as seen-
in surface-view. and projects strongly after the manner of a papilla above the
small-celled epidermis, and (2) a very small cell, which is inserted in the apex
of the epidermal cell and is shaped like a biconvex lens. The external glands
of the second type in some cases have a bicellular head which is divided by
a vertical wall, and, are sometimes (e.g. in Ruellia p,ltula, Jaeq. or, Strobilanthes
glomeratus, Anders.) provided with long stalks.
In connexion with the section on the cYRtoUths (see also the table on
p. 616) we may :first describe the forms of cystoliths recorded hy Friedrich
in a number of genera not examined by Hobein (Glossochilus, Lasiocladus,
Petalidium, Sautiera. Stenostephmms and Tetramerium). Gloss(lchilus Bur..
chellii~ Nees. which belongs to the Barlerieae, has the double cystoliths charac·
teristic of this group. The genu;5 Lasiocladus (L. acanillospe-rmi/ol£-us, Boj.
and L. chlorotrichos, Boj.), which Bentham and Hooker place at the end of
the Order with the addition of a query, shows affinity with the Barlerieae in
having double cystoliths. Petalidium barlerioides, Necs and Sautiera Decaisnci,
Nees (Tribe Ruellieae) have fusiform cystoliths; in Stenostephanus lobe~
liaeformis, N ees (Tribe Justicieae) the cystoliths show a longitudiHal arrange-
ment, while the cystoliths in Tetramerium nen,losum. Nees (Tribe Justicieae) are
elongated and either pointed at one end or rounded.
 ADDENDA-ACANTHACEAE 101 9

Molisch's recent investigations on the blue-green colouration of the

cystoliths, previously observed by K. Richter and A. Weiss in species of
Goldlussia and Sanchezia, necessitate the addition of the following details.
The lithocysts of Santhezia nobilis, Hook., Strobilanthes Dyerianus, Hort.
and Goldtussia flnisophylla, Nees contain. a colourless chromogen (pseudo-
indican) which, on coming in contact with the air as the result of injury to the
cells containing it, gives rise to a deep blue-green colouring-matter which
appears on the surface of the body oi the cystoliths or in their immediate
neIghbourhood. On the other hand, the greenis~ coloration shown by the
cvstoliths of Ruellia ochroleuca, when seen in sections~ is due to the antho-
cyanin contained in the surrounding cells. the violet colour of the anthocyanin
changing to green in contact with the cystolit:r..s owing to their alkaline reaction.
Dethan and Friedrich have recently demonstrated acicular fibres (raphi-
dines, see p. 618) in a large number of additional genera and species.
These species are as follows: A canthopsis horrida, N ees, A ndrographis pani-
cutaia, Nees, Anisacanthu$ virgutaris, Nees, Aphelandra auran#aca, Nees and
A. Fastinator, Lind. et Andre, Barleria cristala, L. and B. Prionotis, L., Cross(md,a
in/undibuliformis. Nees and C. undulaefOlia,_ Salisb., Cystacanthu5 turgida, Nichols.,
Dian/hera pectoralis, Gmel., Dicli-ptera acufflina/a, Juss., Ec'boliuffl L-inneanum, Kurz.
E"anthemum sanguinolentum, VeItch, patonia argyroneura, Coem., Justitia ca,nea,
Limli. and J. Gendarussa, Burnt., Pet'istrophe b",'calyculata, Nees, Rhinacanthus
ccm-munis, Nees, Rueltia formosa, Andr. (Arrhosto%ylum formosum, Nees) , Santhezia
nobilis, Hook. f., Sautiera Decaisnei. Nees.
For the occurrence of steles in the petiole of Acanthus mollis, see Bouygues.
loco cit.
3. STRUCTURE OF THE AXIS. Dethan has recently examined the structure
of the cortex in certain officinal species. The cork develops in the epidermis
in Strobilanthes callos us, Nees, and in the subepidermal layer of cells in Cross-
andra undulaefolia, Salisb. and ]acobint"a Mohintli, Benth. et Hook. The
outer portion of the primary cortex is frequently differentiated as collenchyma.
I~ Strobilanthes callosus, Nees there is a complete subepidermal layer of litho-
-i'cysts. The occurrence of fibrous cells in the collenchyma of the primary
~ortex in Nelsonia campestris, R. Br. and Thyrsacanthus nitidus, Nee's recalls
the hypoderm of fibrous cells characteristic of Thunbergia, Stone-cells or
slightly sclerosed parenchymatous cells have been observed in the primary
cortex in Dianthera pectoralis, Gmel., Jacobinia M ohintli, Benth. et Hook.,
Justicia Gendarussa, Burm. and Strobilanthes callosus, Nees. The species of
Hygrophila are distinguished by having a lacunar cortex with wide inter-
cellular spaces. In H. spinosa, Anders. the walls of the meshes are formed
by cells having a particularly characteristic shape, which in a transverse
section of the branch resembles that of a thigh-bone, while in the corners of
the meshes the cells are generally three-armed and likewise have swollen
ends; these cells, moreover, are dovetailed with one another by means of
the -undUlated surfaces of their walls. In most of the species the pericyc1e
contains isolated bast-fibres or groups of bast-fibres, but occasionally primary
hard bast is absent. In the latter case a distinct endodermis is sometimes
(Barleria cristata, L.) present. Bast-fibres and acicular fibres are of frequent
occurrence in the secondary bast.
As regards the occurrence of anomalous structure in the axis (p. 620)
we may note that Dethan's statement as to the presence of intraxylary phloem
in Jus#cia Gendar-ussa, Burm. is incorrect, and that, according to Leisering, in the
formation of the interxylary phloem in Thunbe1'gia coccinea, Nees, at least,
the cambium at first develops nothing but thin-walled parenchyma on its
inner side, sieve·tubes only becoming differentiated in this parenchyma in
later stages.
·1020 ADDENDA--ACANTHACEAE

For the structure of the terrestrial roots of Acanth~ t.licifolius (occurrence

of peculiar thickening-plates, traversed by slit-shaped pits, in the cortical
parenchyma), see Karsten, loco cit.
Literature: A. Weiss, Kalkoxalatm. in Acanthac., srtz.-Ber. Wiener Akad., xc, Abt. J,
ISS....,
pp. 19-90 and Tab.-Gardine., Secret. hairs of Thunbel'gia launfo!i4, I'roce.ed. Cambridge Phil. Soc.,
v, 18~4, pp. 184, and 1887, p. 8l.-Karsten-, Mangrovevegetat., Bil1l. Dot" Heft :U. 1891, p. !'!o.-
Dethan, Acanthllcees med., These, Paris, 1896 (28 ed., 1897), I~ pp.l-Leisering; k\erxyllires
IJeptom, DiIS., Berlin, 1899, pp. 39-4~.-Molisch, Pseudoicdican, Sitz.-Ber. Wiener Akad., cviii,
Abt. 1,1899. pp. 479-90 and Tab.-Friedrich, Blattnnatomieder Acanthac., Diss., Heidelberg, 1901,
62 pp., 1 Tab. 3_ Pitard, Pbicycle, These, Bordeaux, 1901, p. 73.-Areschoug, Mangroveptl., B~l.
bot., Heft 56, 1902, pp. 48-50 and Tab. v (Acanthus ilieifolim, l ....).-Bouygues, Petiole, These,
Paris, 19Cz, p. 79.-Clauditz, Blattanat. canar. Gew., DISS., Basel, 190z, pp. 10, II (Adhal(I(/Q).-
Col, Faisceaux, Ann. sc. nat., ser. 8, t. xx, 1904, p. 159, etc.-Vi ret, Liaisons du phloeme med., etc.,
Jnstitut de Bot. Geneve, 1904. pp, 71-96 (Acanthus and Thunbergia).-Haberlandt, Lichtsinnes·
organe, 1905, pp. 107-J1 and Tab. iii.-Holm, in Americ. Journ. of Pharm., 1906, p. 553 et seq.,
and 1907. p. PiJ et seq. (Ruellia ciliosa).-Holterman~ Einfluss d. Klimas; 1907. p. :u6.-[Holm,
Rtleliia and Dianllzera, Bot. Gazette, xliii, 1907, pp. 308-29 and pI. :lti-xiii.]

MYOPORINEAE (pp. 624-628).

Literature: Boergesen og _Paulsen, Veget. dansk-vestind. Oer, Dot. Tidsskrift, xxii, 1898-9.
pp. 31-3 (Bonlia dapknoidts, L.) ..

SELAGINEAE (pp.628-630).
The epidermis of the leaf of Glob1.tlaria sa{icina, Lam. consists locally of
two layers. I

Literature: Latour, Sene, These, MontpeUier, 1894. pp. 36-40 (GloDu/aria, .Alypum).-
Clauditz, Blattanat. canar. Gew., Diss., Basel, 1903, pp. 9, 10 (Globularia salicilta, Lam., IIpbalm.
salicij()lia).

VERBENACEAE (pp. 630-636).

I. REVIEW OF THE ANATOMICAL FEATURES. In the genera SymphoTema
and Congea (Symphoremeae) large secretory cells occur in the bast and primary
cortex of the branches. Teijsmanniodendron has large solitary and clustered
crystals, as well as medullary' bundles of mestome.' Tufted hairs are also
found in Symphorema. _
2. STRUCTURE OF THE LEAF. Van Tiegbem has recently undertaken 'a
detailed examination of the structure of both leaf and axis (see below) in the
Symphoremeae and Avicennieae. The following details are abstracted from
hIS investigations and the remaining new literature. According to Areschoug
stomata with two subsidiary cells placed transversely to the pore are·
found in Avicennia nitidu, Jacq.; stomata having three neighbouring cells
occur in Teijsmanniodendron bogoTiense, Koord., and stomata with several
ordinary neighbouring cells in Lippia Geisseana, Solered. According to Van
Tieghem the many-layered hypoderm found in Avicennia occupies half the
thickness of the leaf in A. ofJicinalis, while it is not so thick in the two species
A_ nitidus and A. tomentosus, which are grouped in the subgenus Hilairanthus.
1 This paper deals with species of the following genera: Acanlhus, Adhaloaa., Andrographis,
Bar/eria, Blepkaris, C),stacanthus, Dia,,/Jlet-a, Didip/era, Ecbolium, Graplopk),llum, HY.{r{Jplzila,
Jarobinia, Jus/ida, .file/sonia, Rhinacanthus, Ruellia, Rungia, Strobilanllzes, Thunbergia, TIz)'rs-
acanthus. '
:t This investigation deals with species of AeantluJpsis, Acan/hus, Adhalotia, AnisaeaniAus,
Aplulatuira, Barleria, Bkpharis, Crossandra, nialz/Izera, DieHptera, Era1ttlumum, . Fitttm;a,
Geissomtria, G/ossoclzilm, G),11Illostach),um, Hemigraphis, byjof"sttJ, Justicia, LasiodadtlS, Libo"ia,
Ptr;slrophe, Petalidium, Rlzinacanthus, Rudlia, Rungia, Sauckezia, Sau/iera, Slenandrium,
Slenos/~p"a,.us, SI,obilanllt.es, Tetranu,,'unl, Tlzunl)/~·gia.
 ADDENDA-VERBENACEAE 1021

Three· vascular bupdles enter the leaf in A vicennia, as well as in Symphofema

a,nd Congea. Ip the petiole of A vicennia they form a median bundle, which is
closed in an annular manner, a-nd a few small cortical bundles. while in the petiole
of Symphorcma aad eo",ea ~h~y ~fye rise to a large arc of wood and bast.
Ii. view of t)le earlier statement as to the absence of internal glands the
eiemOJlstration of Jarg~ lecretory cells in the bast and primary cortex of the
branches in $ymphoremc.rand Congea (but not in Sphenodesma) by Van Tieghem
is very noteworthy. In Lippia Geisseana afld (according to Quanjer) also
in Vitex tritolia, L. oxalate of lime is found in the mesophyll in the form of
small clustered crystals, whilst Teiismanniodendron bogoriense has large
solitary crystals in the cortical parenchyma and clustered crystals in the
tissue of the pith. Regarding the occurrence of sphaerocrystalline masses
in the epidermis of the leaf in species of Campylostachys, see Knoblauch,
loco cit.
With reference to the hairy covering of the Verbenaceae (p. 631 et seq.)
we may add the following details. Uniseriate clothing hairs are found also
in Congea tomentosa, Roxb., while tufted hairs with unicellular rays occur
in Symphorema involucra tum, Roxb. According to Haberlandt the small
silicified trichomes inserted in the outer walls of the epidermal cells in Petraea
volubilis (Fig. 151, A-B) are local organs for the perception of light; to judge
from Koorders' statements similar trichomes appear to occur also in Teijs-
manniodendron. Unicellular cystolith-hairs which have calcified tips and in
part contain cystolith-like protuberances (the latter occasionally occurring
In the subsidiary cells of the hairs as well) are present also in LipPia Geisseana.
Glandular hairs with unicellular heads are found also in Lippa (L. Geisseana,
L. saturejaelolia, Mart. et Schauer, L. thymoides, Mart. et Schauer) and in
Lantana macrophylla, Schauer and L. involucrata, L.; peltate glandular hairs,
like those of Avic.ennia, &c., occur also in Citharexylum citlcreum, L., and
vesicular integumental glands resembling those of the Labiatae also in V£tex
trifolia, L. The glandular hairs found on the parts of the flower occasionally
differ from those present on the vegetative organs. Thus on the bracts of
Lippia Geisseana_ the heads of the glands are both unicellular and' composed
of a small number of cells owing to vertical division-walls,. whilst on the clavate
processes of the anthers they show a still larger number of vertical division-
walls. An even more striking feature lies in the occurrence of external glands
with elongated ellipsoidal or tubular heads divided both by horizontal and
vertical walls on the processes of the anthers in Tamonea spicata, Aubl. and
on the connectives of the anthers in Casselia integrifolia, Nees et Mart. We
must regard these external glands as forming exceptions to the general rule,
since the glands of the Verbenaceae in most cases have heads, which are either
unicellular or divided exclusively by vertical walls (see Solereder, loco cit.,
p. 627, footnote I). .
3. STRUCTURE OF THE AXIS. For the structure of the stem in Lippia
Geisseana;"a" Chiliarr desert-plant (having radial plates of sc1erenchymatous
fibres corresponding in position with the ribs on the stem and palisade tissue
in the outer part of the cortex, &c.), see Solereder, loco cit.
The cork develops in the subepidermal layer of cells also in Avicennia,
~_phorema and Congea. In the species of Symphorema and Congea men-
tidded above the perlcyc1e contains an annular zone of lignii\ed fibres, while
in Sphenodesma unguiculata, Schauer it includes a composite and.: continuous
ring of sclerenchyma. In Symphorema involucratum the 'secondary bast con-
tains stone-cells, and in Sphenodesma unguiculata, Schauer bands of scleros~d
cells. Pitard records isolated bundles of bast-fibres in the pericyde in species
of Aegiphila,', Clerodendron, Cornutja, Oxera, and Verbena, and a composite
and interrupted ring of sclerenchyma in the pericycle in Clerodendron aculea-
 1022 ADDENDA-VERBENACEAE

tum, C;. Bungei, and Vitex agnus-castus. The vessels have scalariform per-
forations also in Teiismanniodendron (Koorders).
The medullary mestome·bundles mentioned above as occurring in
Teiismanniodendron are present in large numbers (about 25) and consist of
libriform and wood-parenchyma.
Note. For the anatomy of the aerial and terrestrial rODts of A vicennia tomentosa,
see also W. Brenner, loco CIt.; regarding A vz'cennia m'Nda, Jacq., see B<n!rgesen and
Paulsen, loe. cit.
Literature: Costantin, Tiges aero et sont., Ann. sc. nat., ser. 6, t. xvi, 1883, p. I23.-Karsten,
Mangrove-Vegetat., Bibl. bot., Heft :;12, 1891, p. 50.-Knoblauch, Qkolog. Anat., etc., Habilitat.-
Scbr., Tlibingen, 1896, p. 8 et seq.--Schtlbert, Parenchymscheiden, Bot. Centralbl, 1897, iv, p. :10.-
Solereder, Buddleia Geissf!ana, Bull. Herbier Boissier I 1898, pp. 633-9.-Van Tieghem. A vicen-
niacees et Symphoremacees, Joum. de Bot., 1898, pp. 34S-6s.-Boergesen og Paulsen, Veget.
dansk-vestind. Oer, Bot. Tidsskr., xxii, 1898-9, pp. 37--9 (Avicennia nilida, Jacq.), p. 100
(Cillzal'exylum cimreum, L.) and p. 90 et seq. (Lantana).-Keamy, in Contribut. U. S. Nat. Her-b.. v,
D.5, 1900, p. 304 (Lippia nodiftora).-Thotrul$, Feuilles sout., TMse, Paris, I900.-Pitard"pericycle.
These, Bordeaux, 19°1, p. 88.-Areschoug, Mangrovepfl., Bibl. bot., Heft 56,'1903, pp. ,!i0-3 and
Tab. iv and vii (Avicennia nilida).-Bargagli-Petrncci, Legnam'i, Malpighia, 1902, p. S65 (Viteae).
-Penzig, Piante acarofile, Malpighia, J902, p. 451 (Ci'tltarexy!um).-W. Brenner, Luftwurzeln von
Avicennia lomen/osa, Ber. deutsch. hot. Gesellsch., 1902, pp. 175-88 and Tab. vi-viii.-Quanjer,
Anatomiscb Bouw, etc., Natuurkund. Verhandel. Haarlem, iii, 5, 1903 (Vilex trifoHa).-Theorin,
Vaxttrichom., Arkiv for Bot., i, 19°3, p. IS6.-Urtprung, Dickenwachsttlm. Bot. Zeit., 1904, p. 19:1
et seq. (Tee/(Jna grandiS', L.).-Col, Faisceaux, Ann. sc. nat., sec. 8, t. xx, 1904. pp. u8, og.-
Koorders, Tei/smanniodentiron, Anu. Jardin Buitenzorg, xix, 1904. pr.' :13, 23·- Haberlandt,
Lichuinnesorgane, 1905, pp. 73-5 and Tab. ii.-Piccioli, Legnami, Bul. Siena, 1906, p. 150'-
Holtermann, Einflnss des Klimas, J907, pp. 31, 59 and :2 1:2.

LABIATAE (pp. 636-642).

I. REVIEW OF THE ANA'tOMICAL FEATURES. To the list of special features
we may add the occurrence of characteristic secretory cells in the mesophyll
of Pogostemon; these cells are differentiated as internal (glandular) hairs,
being 'provided with a stalk composed of a small number of cells and a uni-
cellular vesicular and glandular head of varying shape. .
2. STRUCTURE OF THE LEAF: To p. 637 et seq. the following details may
be added. Small clustered crystals of oxalate of lime have recently been
recorded also by Clauditz in Salvia canariensis, L. A noteworthy feature
from the systematic point of view is the occurrence of a special body in the
form of Molisch's scutellarin in all the species of Scutellaria which have been
examined, as well as in Galeopsis Tetrahit, L. and Teucrium Chamaedrys, L.,
while the investigation of certain species of Ballota, Brunella, Glechoma, Lamium,
Leonurus, Mentha, Nepeta, Teucrium, and Thymus gave a negative result;
the chemical composition of this body has been examined in detail by Gold-
schmiedt. The chief seat of the scutellarin is in the epidermis of the leaf and
especially in the lower epidermis. On treating sections with a IO % solution
of hydrochloric acid the scutellarin crystallizes out in the form of sphaerites
or of needles, which are grouped in a tufted or stellate manner. Regarding
the occurrence of masses, which resemble sphaerites and show similarity with
hesperidin, in the mesophyll of Hyssopus officinalis, see Tunmann, loco cit.
I observed the above-mentioned secretory ceJIs, which are differentiated as
internal glandular hairs, in the Patchouli-drug, which is derived from the
leaves of Pogostemon Patchouli, Pellet, as well as in the cortical tissues of the
same plan t. In the leaf these secretory cells belong to the mesophyll;
they are intercalated between the cells of the palisade and spongy tissues,
frequently projecting into the intercellular spaces. They have a short
stal~ which is composed for the most part of two or three low cells
witll suberized walls and forms the means of attachment to the cells of the
 ADDENDA-LABIATAE 1023

mesophyll, and a variously shaped unicellular head; the latter is either sac-
shaped or lobed, and shows a cuticularized membrane, which is raised above
the cellulose-membrane like a bladder, in the same way as in the rather large
vesicular integumental glands with a unicellular head found on the surface
of the leaf; the resinous or oily secretion is soluble in alcohol and fills both
the space between the cellulose- and suberized membranes and the lumen
of the heap-cell.
Clauditz has demonstrated branched hairs also in Sideritis (Leucophac,
Webb). The vesicular integumental glands in some cases (Pogostemon) merely
have a relatively large unicellular head.
For the occurrence of steles in the petiole of Phlomis Russeliana, see
Bouygues, loco cit. .
Literature: Paschkis. Minder bek. Blatter, Zeitschr. osterreich. Apotheker- Ver., IS 79, p. 415
et seq. (Pogostemon).-Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi, 1883, p. Il3 et seq.
-Costantin, Tige!! d. pI. aquat., Ann. sc. nat., ser. 6, t. xix, 1884. p. 281 et seq. and pI. xv.-Jost.
Zerkliiftung einig. Rhi:r.. u. Wur:r.., Bot. Zeit., 18<}O, p. 501 (Sal'Via).-[TQgnini, StQmi, Att.~ 1st. oot.
Pavia, I894.]-Spanjer, Wassera.pparate, Bot. Zeit., 1898, i, p. so.-Hirsch, Entwickl. d. Haare,
Dis!!., Berlin. 1899. p. 19 et seq.-Keamy, in Contrib. U. S. Nat. Herb., v, n. 5. 1900, p. 291
(TtNCrium Nashii, Kearny).-Tunmann, SekretdTiisen, Diss., Bern, 1900, pp. 38-43.-Molisch u.
Goldschmiedt, Scntellarin, Sit:z:.-Ber. Wiener Akad., ex, Abt. J, 190 1 , espeCially pp. 185-9,-
Petersen, Vedanatomi, 1901, p. 84.-Pitard, Pericycle, These, Bordeaux, 1901, p. 38.-Bonygues.
Petiole, These, Paris, 190:', p. 7s.-Clauditz, lUattanat. canal. Gew.• Diss., Basel, 1902, pp. 16, 1'1
and 48, 49 (Leu(ophai, Micromtn'a, SalVJ:·a).-[Armari, Piante della reg. medit., Ann. di. Bot., i,
19°3, p. 17 et seq. (Pklomis., TA:ymus).]-Theorin, Viixttrichom., Arkiv for Bot., i, 1903, p. 156.-
Col, Faisceaux, Ann. sc. nat., ser. 8, t. xx, '904. p. 1I8.-Freidenfeldt, Anat. Bau d. Wurzel, Bib1.
bot., He~t 61, ]9°4, pp. 69-7I.-Silssenguth, Behaarungsverh. ,d. Wtirzb. Muschelkalkptl., Diss.,
Wiirzburg, 1904, pp. 47-5o.-Sarton, Anat d. pl. affines, Ann. sc'. nat., ser. 9, t. ii, 19°.5, pp. 79-85,
86-93, IOI-J04 (Ajuga, Calamintlt.a, Galeopsis, Lavandula, Teucn·um).-Daupbine, Rhizomes. Ann.
sc. nat., s~r. 9, t. iii, 1906, pp. 360 Md 362 et seq.-Piccioli, ugnami, Bull. Siena, 1906, p. ril.-
[Tunrnann, H;lSSopus o.fficina!is,Zeitschr. aUg. osterreich. Apoth.-Ver., 1906, n. 30-3li abstr. in Bot.
Centralbl., civ, p. 63.)-[Borde, Rech. histochim. sur les Labiees astringentes, Toulouse, 1907.]-
Solereder, Die inneren baarart. Sekretdriisen des Patschuliblattes, Arcbiv d. Pharm., 245, 1907,
pp. 406-9.-[For further literature, see p. 1170.]

PLANTAGINEAE (pp.642-644).
In the first place we may add the following details to the section dealing-
with THE STRUCTURE OF THE LEAF on the basis of Pilger's recent investiga-
tions on the anatomy of the species of Plantago. The leaves are for the most
part ceritric in structure; they either have palisade tissue on both sides and
spongy tissue in the middle, or the mesophyll consists of palisade tissue only;
in many cases t moreover, it is homogeneous, being composed of 'uniform
assimilatory tissue with rounded cells. In all the species the stomata are
situated on both surfaces of the leaf. The vascular bundles of the veins are
invariably provided with sclerenchymatous layers on their upper and lower
side or are surrounded by a ring of sclerenchyma.; in their course through the
petiole they~at.:e _distinct from one another. A type of clothing hair, which is
specially noteworthy from the systematic point of view, is constituted by
trichomes having a short stalk·cell, which is seated on an epidermal cell as base,
and a long flagelliform terminal cell with thick walls. These clothing hairs,
which have already previously been recorded in Plantago arctica, are charac-
teristic of the perennial species of the section 01eades, Decne., and of the species
of the sections Arnoglossum, Decne. and Le·ucopsyUium, Decne. The flagelli-
form trichomes of Plantago Bismarckii, Niederl. (section Bismarckiophyton,
Harms) differ to a slight extent in the fact that the basal cell, which projects
above the level of the leaf, is divided by a wall running parallel to the surface.
Another type of trichome is found in the species of the subgenus Psyllium t

Harms (e.g. Plantago mauritanica) in the form of papillose hairs; these consist
of a large thick-walled cell, which is seated on several epidermal cells and is
 .It DDENDA-PLA NTAGINEAE

arched outwards. Glandular hairs occur only in species of the subgenus

Psyllium, Harms. Pilger remarks upon them as follows: 'One finds either
multicellular sessHe glanns, or a multicellular head inserted on a delicate
unicellular stalk, or a unicellular head on a multicellular stalk; in the latter
case one or two of the cells below the head have thin walls, while the epidermal
cell and the first few cells of the hair are broadened and have thickened walls.·
As regards .THE AXIS of the genus Plantago (p. 644) we may note in the
first place that certain differences in structure go hand in hand with the varied
morphological nature of the stem, which is differentiated either as a rhizome
or as a subaerial axis. The earlier statement as to the absence of medullary
rays must be limited in the sense that there are no secondary medullary rays.,
The xylem.rjng shows interruptions, which are of primary origin (primary
medullary rays), although they are not to be found in every transverse section;
the parenchyma composing these rays, or at least some of it, often becomes
lignified. In the annual species belonging to the subgenus Psyllium a typical
multiseriate ring of bast-fibres is generally developed in the pericycle on the
inner side of the endodermis, which is devQid of chlorophyll; to judge by
Reiche's statements the same is tpe case in Plantago Fer na ndezia , Bert. In
other cases typical hard bast appears to be wanting, and it is only in certain
species of the section Leucopsyllium (e.g. Plantago albicans), In Plantago
Bismarckii, Nieder!., P. princeps, Cham. et Schlecht. and in certain shrubby
species of the subgenus Psyllittm (e.g. Plantago sinaica, Barp.) that one meets
with sclerenchymatous elements at the outer limit of the bast-; these elements
are pr9bably for the most part parenchymatous, being differentiated from the
outermost portion of the soft bast. In those species, which have a thick mas$
of wood, the phloem is feebly developed .. In the fleshy rhizomes there.are long
rows of cambiform cells, the outer layers of which show collenchymatous
thickening. The tissue of the pith either persists as thin·walled .parenchyma
or (in the shrubby species) becomes lignified; the cells occasionally bear large
pits. Medullary cambiform.bundles occur chiefly in the fleshy rhizomes of
the species belonging to the section Polyneuron, Decne. (exceptions: Plantago
palmata, P. cordata, Lam., &c.), although they are sometimes found also in
,species of the section Heptanettron, Decne" and in Plantago princeps, Cham.
et Schlecht., which together with P. Fernandez~·a, Bert. (which is distinguished
by having medullary and cortical bundles, see p. 644) constitutes the section
Dendriopsyllium, Decne. In 'Plantago CornUti, Gouan there are all transitions
:between these medullary bundles and true vascular bundles; the latter are
Irecorded by Pilger also in P. ·princeps. Groups of stone·cells are' found in
the pith of the fleshy rhizomes of species belonging to the section Oreajes,
.Decne. "<U).d in the pith and primary cortex of the fleshy rhizomes, of species of
the section Arnoglossum, Decne. (also in the annual species Plantago Lagopus,
P. lusitanica, Willd., &c.).
We may finally mention a phenomenon, which has been observed in a few
!Species of the section Oreades (e.g. Plantago saxatJlis. Bieberst.) and in Plantag/}
alpina, L. (section Coronopus, Decne.); in these forms the rhizome splits up into
,several large bundles owing to the formation of medullary cork, which enters
,into connexion with the outer periderm by way of the primary medullary
Tays (for details, see Pilger, loc. cit.). We may also add a statement of Reiche,
which was overlooked in the earlier part of this work, and according to which
abundant' crystals' are present in the pith of Plantago Fernandezs·a.
The flowering scapes of Plantago show a more or less developed ring of bast·
bres adjoining the ring of vascular bundles, while on the outer Slde of the ring of
bres there is an endodermis which does not contain any chlorophyll. Numerous
f:
trands of sieve·tube~ are found between the vascular bundles being situated at
he limit of the pith and the ring of bast-fibres.
 ADDENDA-PLANTAGINEAE 1025

Literature: COltantin, Tiges aero et lOut., ADn. sc. nat., ser.

6, t. xvi, 1883, p. 124- et seq.-
Schobert, Parenchymscbeiden, Bot. CentraJbJ., 1897. iv, p. 63.-PiJger, Vergl. Anat. d. Gatt.
Plan/ago, in Engler, Bot. Jahrb., xxv, 1898, pp. 396-35I.-Minden, Wassersez. Org., Bib\. bot.,
Heft ...6, 1899, p. IS (Lillorella).-Swaolund. V~etat. Neu-Amsterdams u. St. Pauls, Diss." Basel,
19oI, pp. 36-9 (Plantago Stauntoni, Reichenb. I ).

NYCTAG INEAE (pp. 645-649).

2. STRUCTURE OF THE LEAF. The following details are taken from
Heimerl's and Gidon's recent investigations. The leaves show centric structure,
for example, in Bougainvillea patagonica, Decne., Phaeoptilon spinosum,. Radlk.
and A bromia umbellata, Lam. In Boerhaavia viscosa, Lag. et Rodr. the cells
of the palisade tissue are provided with ,short processes, which are sharply
demarcated from the cell proper and form the means of connexion between
the cells; similar short processes are found also on the cells of the spongy
tissue in this species, and altogether appear to be of rather frequent occurrence
on the cells of the leaf-tissue in the Nyctagineae (Bougainvillea spectabilis,
Willd., Oxybaphus visco sus, L'Herit., &c.). In Boerhaavia viscosa the vascular
bundles of the smaller veins are surrounded. by a.sheath of large parenchyma-
tous cells, which is only interrupted opposite the bast, while the sheaths of
the larger veins are less complete; a weakly developed sheath of large paren-
chymatous cells has' been' recorded also in Bougainvillea spectabilis, and a
distinctly differentiated sheath in Phaeoptilum spinosum. In Bougainvillea
patagonica, Boerhaavia viscosa and Phaeoptilum. spinosum the stomata are
found on both surfaces of the leaf. The last of these species has a hypoderm
of large cells with rather thick walls 'beneath the epidermis. According to
Heimerl uniseriate trichomes with a swollen, vesicular terminal cell, which
has a rather thick stratified wall, so that the amount of secretion is insignificant,
are very widely.·distributed in the species of Bougainvillea. -The same author
also records small uniseriate trichomes with a few short stalk-cells and a clavate
or vesicular terminal cell, wh.ich sometimes has thick stratified walls, in I

Bougainvillea patagonica; these hairs occasionally bear lateral vesicular cells

in addition to the terminal one, and thus approximate to the branched hairs
with a glandular function, previously described in Pisonia tonuntosa (see
Fig. 154, B, p. 646). The deposits of oxalate of lime in this Order appear not
uncommonly to assume diverse _crystalline forms in one and the same organ
(see the earlier statements regarding Pisonia nitida). In species of Bougain-
villea Heimerl observed the following types occurring side by side: raphide-
saC6 of varied size and differentiation, short sacs containing a large number
of delicate needles, sacs with distinct prismatic crystals, &c. In the veins
of the leaf of Phaeoptilum, spinosum short prismatic crystals are found side by
side with bundles of raphides. Elongated styloids, traversing the mesophyll
at right angles to _t!le surfac~ of the leaf, are characteristic of Bougainvillea
modesta, Heimerl, and B. stipitata, Griseb.
For the arrangement of the vascular bundles in. the median vein and their
course, see Gidon, loco cit.
Literatnre: [Ormandy, Schlaachgef. v. Afinwilis, Koloszvar, 1881, 31 pp. (HuDgarian).]-
Gidon, Appareil conductenr dans la tige et dans la {euil1e des Nyctagin., These, Caen, 1900, 120 pp.,
pL i-vi; see also Me'li. Soc. Linn. de Normandie, xx. J90o.-Heimerl, Menoge. d. Nyctagin., I
(BIJUC(Jinvill~a, Phaeoptilum, Colirnonia), Denkschr. Wienet Aka.d .. lxx. 1901, p. 91 et seq.-
[Riasner, Beitr. z. Anat. d. Bl. mancher Nyctagin., Societas historico·naturalis croatica, xii, 1901,
pp. 1-24, 3 Tab. (Croatian); abstr. in Bot. Centralbl., lxxxix, pp. q6 and 357.]

J The statements in this paper as to the occnrreoce of crystals of 'carbonate of lime in the axis of
the inflorescence and in the mesophyll are no doubt incorrect.
50lER£DEIt . 3u
 ADDENDA

ILLECEBRACEAE (pp.649-651).
To the first paragraph which summarizes the ANATC~ICAL FEATURES
. OF THE ORDER we may add the following details. Pericyclic development of
the cork has been observed in the axis of Gymnocarpos, Habrosia, and Scleran-
thus. Stomata conforming to the Caryophylleous type occur also in Habrosia.
Branched hairs are found also in A chyronychia, and uniseriate glandular hairs
with a unicellular head also in H abrosia. Masses of crystal-sand, similar to
those found in Gymnocarpos, are present also in Habrosia and Dysphania.
Lastly, anomalous structure of the vascular system (successive rings of growth)
has been observed also in the axis of Cometes, and in the root in species of
Acanthonychia (Pentacaena), Achyronychia, Cometes, Corrigiola, Dysphania,
·Haya, and Pollichia.
The STRUCTURE OF THE LEAF (as well as that of the axis) has recently
been examined especially by Josting and Liiders, their investigations dealing
with species of Illecebrum, Acanthonychia (Pentacaena), Dysphania, Pollichia,
H aya, A chyronychia, Paronychia, H e_rniaria, Siphonychia, A nychia, C orrigiola,
Gymnocarpos, Sclerocephalus, Pteranthus, Cometes, Dicheranth-us, Scleranthus
and Habrosia. The structure of the leaf is centric, more rarely (Siphonychia)
bifacial; according to Josting distinct palisade tissue is more commonly
developed than Liiders' recent statements lead one to suppose. Stomata
of the Caryophylleous type are found also in Habrosia spinuli/lora, Fenzl, while
in Siphonychia americana, Torr. only some of the stomata are surrounded by
distinct subsidiary cells of relatively small size. In the remaining members
of the Order subsidiary cells are wanting. In 11abrosia spinuli/lora the stomata
on the leaf show the same arrangement as in Scleranthus; the same feature
has been observed also in A nychia dichotoma, Michx. and Gymnocarpos truti-
cosum, Pers. (= G. decandrum. Fotsk.). Deeply sunk stomata are found also
in Pteranthus echinatus. In the species examined by Liiders the stomata are
always present on both surfaces of the leaf. The epidermal cells in the specJ,es
investigated by lasting are of large size in all the members of the Tribes
Pollichieae and Paronychieae (with the exception of Paronychia serpyllifolia, DC.
and Herniaria glabra. L.), as well as in Cometes abyssinica, R. Br.; Pter.,anthus
echinatus has high epidermal cells, whilst the cells of the epidermis a.re l,1J1usuaUy
large in Habrosia spinuliflora and Scleranthus perennis, L. Strong punotation
of the cu tide is characteristic of Illecebrum (Liiders), while in other cases.a ~gh t
punctation or striation may be observed. Papillae are found at the m,argin
of the leaf and beneath the veins in species of the genera Achyronychia, Anychia,
Cometes, Corrigiola, Habrosia, Pollichia and Scleranthtts. In Cometes aby~s~'nica,
Dicherantkus plocamoides, Webb, and Pteranthus echinatus the epidermis is
covered by a thick deposit of wax. The vascular bundles of the veins are
occasionally accompanied by sclerenchyma (species of Acanthonychia, Habrosia,
Paronychia, Pollichia, Scleranthus, &c.).
Unicellular clothing hairs have been observed also in Siphonychia, and
uniseriate hairs also in H abrosia; transitional forms between unicellular
trichomes and richly branched multicellular hairs have been found in Achyro-
nychia Parryi, Hems!. The different parts of the stem in Cometes abyssinica
bear peculiar uniseriate trichomes, consisting of a large and broad barrel-
shaped basal cell, followed by four further. cells, which show -a successive
decrease in size, and ending in a pyriform terminal cell; the trichomes found
iOn the surface of the leaf, on the other hand, are reduced to the pyriform cells,
which bear scattered grains of wax. Glandular hairs with a uniseriate stalk and
a unicellular spherical secretory head are present also in H abrosia spinuli/lora.
Clustered crystals of oxalate of lime are widely distributed both in the leaf
 ADDENDA-ILLECEBRACEAE
and axis. A feature deserving mention is the occurrence of rows of clustered
crystals a t the margin of the leaf in A nychia dichotoma, Michx. and H abTosia
sp in uli/lrr,a , The characteristic masses of crystal·sand have also been found'
by JOsting in Htibrosia spinulifiora (in !he axis and leaf) and by Lliders in
Dysphania.
We may now consider the STRUCTURE OF THE AXIS. In most of the
species the. vascular bundles unite to form a ring, but in Corrigioia capensis,
Willd. the transverse section of the stem shows six isolated bundles, which
are alternately large and small and are arranged in a single ring. As
regards the structure of the wood we may first note that jasting likewise
failed to observe typical medullary rays in any of the species. T~le vessels
invariably have simple perforations. In almost all the members of the Polli-
chieae, Paronychieae and Sc1erantheae, which have been examined, the
wood·prosenchyma bears bordered pits; the only exceptions are Hernia.,ia
glabTa, L., in which Josting did -not' find any lignified wood·fibres whatsoever,
and Achyronychia Parryi, Hemsl., in which the wood~fibres are not pitted.
The relative numbers of vessels and wood·ftbres participating in the formation
of the wood are subject to variation. An endodermis composed of large cells
i$ developed in very many members of the Order. The sclerenchyma in the
peric.ycle for the most part forms a closed strengthening ring consisting of
from one to five layers of fibrous cells; in the thicker parts of the axis this
ring may become burst open. In Corrigiola capensis the pericycle contains
isolated groups of fibres only, while in Pteranthus echinalus the gaps in the
sclerenchymatous pericycle are closed by sclerosed parenchyma. On the
inner side of the pericyclic sclerenchyma one occasionally meets with paren-
chymatous tissue belonging to the pericycle, and in Illecebrum verticillalum, L.
this tissue consists of remarkably large cells; within the parenchymatous
peri cycle lies the soft bast, which shows collenchymatous differentiation in
many of the species. Subepidermal development of the cork has been observed
also in ~nychia (according to Regnault), Herniaria glabra and Paronychia
supylli/.olia, while the origin of the cork is pericyclic in Gymnocarpos fruti-
c:osum, .Uabrosia spinuliflora and Scleranthus perennis. In many members of
the Order there is a cavity in the pith.
Anomalous structure of the vascular system, consisting in the formation
of 'CoI1cf.iltric rings of vascular bundles, is"found also in the axis of Gometes
(according to Liiders), in the root of Achyronychia Parryi, A. Gray, Corrigiola
capMsii, Willd. and PoUichia campestris, Sol. (according to Josting), and in
the mOI"Q, strongly developed roots of certain unnamed species of Acanthonychia
(Penlacaena), Cometes, Dysphania and Haya (according to Liiders).
Lil&ature: Christ, Laubblattstengel der Caryophyllinen u. Saxifrageen, Diss., Matburg, 1887,
pp. 6o-S.-W. Meyer, Beitr. z. vergl. Anat. d. Caryophyllaceen u. Primulaceen, Diss., Gottingen,
1899. pp. 39-4J.-[ParmentierJ Gnavelles de France, Ann. Soc. bot. Lyqn. xxiv, 1899, p. 83
(Sduanthus).)- Josting, Anal. d. Sperguleen, Polycarpeen, Paronychieen, Sderantheen u. Pteran.
thee-n, Beih. z.-Bot. Centralbt, xii, 1902, pp.-i49-S6, 159-62, 163 and 171 et seq., also Tab. ii [or 4].-
H. Luders, Untersuch. liber d. Caryophyllaceen mit einfachem Diagramm, Diss., ErllLDgen, 1906.
pp. 33-8; scpo copy from Engler, Bot. Jahrb., xl.

AMARANTACEAE (pp. 651-655).

2. STRUCTURE OF THE LEAF. Capitate hairs with a uniseriate stalk com-
posed of a small number of cells and a tubular unicellular head ar(...tound also
in lrcsine (1. spiculigera, "Seub.). Uniseriate clothing hairs having pointed
ends and consisting of several short basal cells and one or more longer terminal
cells, which are either smooth or papillose, occur also in species of GomphTena.
FToelichia tomentosa, Moq. has uniseriate clothing hairs composed of a basal
.3U 2
 ADDENDA-AMARANTACEAE

cell which is differentiated as a pedestal an<t is followed by two or three short"

and narrow cells, and a longer pointed terminal cell (Seubert).
Literature: Seubert, in Martias, Flora brasil., Y, i, 1875, Tab. So et seq.-ViretJ Liaisons du
phtoeme med. etc., lnstitut bot. de Genhe, 1904, pp. 18-35 (Ackyrant/l.es).

CHENOPODIACEAE (pp. 655-663).

I. To the REVIEW OF THE ANATOMICAL FEATURES we may add that
unicellular hairs occur in this Order, but are only very rarely found (Salsola).
2. The STRUCTURE OF THE LEAF has recently been examined especially
by Montelli, and further statements will also be found in Solms-Laubach's
work (lac. cit.). The structure of the mesophyll in Kirilowia eriantha, Bunge
(Tribe Camphorosmeae) conforms to the second of the special types mentioned
in the earlier part of this work, while Camphorosma monspeliacum, L. belongs
to the third type (Mantell). According to Mo,ntell, however, ordinary types
of leaf· structure are also found in the Order, e.g. : bifacial structure with one
or more layers of palisade tissue on the upper and rather dense spongy tissut'
on the lower side (species of A 1~thochlamys, Beta, Chenopodium, Spinacia, &c.).;
a homogeneous mesophyll composed of rounded cells (Axyris hybrid a, L.);
centric structure with one or more vascular bundles occupying a central
position and a sheath of one or more layers of palisade tissue beneath the
epidermis (Lophiocarpus pOlystachyus, Turcz., Kochia hil'suta, Nolte) or centric
structure with a mesophyll formed entirely by short palisade-cells (Beta
maritima, L.). j

The following details are taken from Solms-Laubach's investigations. In some

of the species "of Suaeda (5. Foyskalii, Solms, S. pruinosa, Lge. and S. vermiculata,
Forsk.) as well as in Schan~inia baccata, Moq. and S. hortensis, Moq.. the arc·
s~a~d vascular system, whi,:-h occupies a ce?tral position, is enveloped. from
WIthin outwards by the folloWlng sequence of tissues: (a) first by aqueous tissue;
t~en (0), by a. sheath of collecti~g cell?; and (c) finally ~y a single layer of pali~ade
tissue sItuated beneath the epldernus. In other species of Suaeda (S. a,lhsslma,
Pall., S.!ruticosa, L'l S. mafitima, L., S. Physophora, Pall., S. salsa, Pall., SFsetigefa,
DC.), on the other hand. there is nothing but palisade tissue (comprising several
layers of cells in the radial direction) between the epidermis and the vascular
network. The same structure of the leaf, as is figured m Fig. 158. B (p. 656) aRer
Volkens for a plant described as 5alsola·longifolia,. Forsk., but considered by Solms
to belong to S. Siebey;, PresI, is found according to the latter authori~ also in
Salso/a opfJas#ilolia, Desf., S. Sckweinlurtkt'i, Solms, S. letragona. Delile, S. VeI'-
miculata, -L., Halogeton alopecut'oides, Moq., Tyaganum nudatum, Delile and Sevada
Schsmperi, Moq.; these species likewise show vascular strands which branch off
from the median vascular bundle and run in the aqueous tissue.
Special sheaths like those previously described are developed in the veins
of the leaf also in species of COTispermum (Corispermeae). Sclerenchyma is
found accompanying the median vein in CeratocaTpus aTenarius, L. and Grayia
polygaloides, Hook. et Arn. (Montell).
The transverse ~angeme~t of the stomata mentioned on p. 658 has
been observed also on'1~~ stem of Camphorosma mon~peliacum (Cassan).
To the' section d~ng with the hairy covering we may in the first place
add that short unicellular clothing hairs, appearing to the naked eye as small
bristles, actually occur in Salsola (5. Tragus, L.), so that contrary to the earlier

1 Montell's investigations deal with the following genera: /I hacodia, LOf>hiocarpus, Clunopo-
dium incl. Bli/u"'" !f()u~inJa, B.da, Oreobli/"N,. Spiltada, Alrip!e:x, (/rayia, Cetutacarplls; AXJln's,
Ca1np~DrDSma, K,nlawla', Corupermum, An/leoch/amys j Polycnemuf1,l; 'CIImo/ta incl. Echinopu·'on,
Kuch1a, Ellchylaenq. j Suaedaj Sa/sC'/a, Anaoast·s, Nanoplzyfum, Sympegma.
 ADDENDA-CHENOPODIACEAE 1029
statement unicellular trichome$l are not wanting in all the Chenopodiaceae.
Among the members of the Camphorosmeae, in which Montell records clothing
hairs with a long terminal cell bearing papillae, Enchylaena tomentosa~ R. Br.
deserves special mention, owing to the great length of the papillae in this
specIes.
Vesicular hairs in which the terminal cell shows varied differentiation
(sometimes even in one and the same species) were observed by Montell also
in species of Roubieva, Oreobliton, Spinacia and Chenolea (Echinopsilon) , In
some of the vesicular hairs fo~nd in Chenopodium anthelminticum, L. and
Roubieva multijida, Moq. the terminal cell is two-armed.
True glandular hairs, which secrete oil and have a structure similar to that
figured in Fig. 159, 0, are found in Roubieva multifida, which I have myself
examined.
In certain members of the Order the clustered crystals of oxalate of lime
fill large idioblasts in the mesophyll. Cells containing crystal-sand were
observed by Montell in the mesophyll in the following additional cases: Beta
Ero parte, Corispermum (pro parte, side by side with clustered crystals),
Enchylaena tomentosa, R. Br., Nanophytum iuniperinum, C. A. Mey., Orco-
bliton thesioides, Dur. et Moq. and Suaeda altissima, Pall. (whilst other species
of Suaeda have no oxalate of lime in the leaf). A subepidermal layer containing
clustered crystals occurs also in the leaf of Salsola Soda, L. (but not in that
of S. Kali, L.).
3. STRUCTURE OF THE AXIS. The development of the cork, which is
described as pericyclic on p. 662, has been carefully studied by Leisering.
It is a familiar fact that the cork in this case occasionally arises so near to the
anomalous vascular system that it appears to take its origin in the meristem,
which produces the anomalous secondary growth of the latter. It has now been
shown that the cells of the cork-cambium either belong to the parenchymatous
pericycle or are given off on its outer side by the meristem just referred to.
In At"tplex hastata, L. the place of origin of the cork varies in one and the
same transverse section; at certain points the cork is found immediately
extern~ to the groups of pericyclic sclerenchyma, while at other points it is
. situated immediately on their inner side. According to Leisering Jthe cork in
Eurotia ceratoides, C. A. Mey. includes lamellae composed of one or two layers
of uniformly sclerosed cells. In the species just named, as well as in Haloxylon
A mfnOdendron Bge. and H. articula tum, Bge., Jonsson records a peculiar
t

process of gelatinization among the cells of the cork. In these species the
cork besides containing celIs of the normal type includes 'phelloid-celIs'
(occasionally containing small crystals of oxalate of lime); which ultimately
come to have a spherical shape and become separated from one another as
a result of the gelatinization of their middle lamellae _and of the inner layers
of their membranes.
As regards the occurrence of tracheids with a spiral strengthening band
(see p. 663), ~we may add that- in Salicornia herbacea Montell records a small
number of imperfectly differentiated tracheids, while in S. macrostachya he
figures spiral tracheids side by side with spiCUlar cells.
The following additional details regarding the anomalous structure of
the fibrovascular system of the root (see p. 663) are taken frdID Fron's
investigations. The arrangement of the bundles of wood and bast is the same

1 Montell (loc. cit.• p. lO) correctly records unicellular trichomes in Salsala; on the other hand
his statement to the- effect that such hairs occnr in Koclu"a scapa,..ia, Schrad. is incorrect, as I have
found by an investigation of this species. The capitate hairs, which Montell (loc. cit., pp. 39,; 49
and 70, 71) figures as nnicellnlar in species of Clunopodium. Oreohlilon and Atriplex, no doubt
in all cases have a distinct uni- or multicellular stalk; reinvestigation of Alnp/ex ~rlula(aidt!, L., at
all events, showed Ihis to be the case in ,this species.
1030 ADDENDA-CHENOPODIACEAE
as in .. the stem. Concentrjc rings of bundles (Type I) are found in species of
Beta, Camphorosma, Chenopodium, Corispermum, Kochia, and Spinacia, being
incomparably more abundant than in the stem; but the second type, in
which the vascular bundles are embedded in prosenchymatous conjunctive
tissue, has also been observed, the bundles not uncommonly exhibiting a spiral
arrangement. Fron has shown that this spiral arrangement takes its origin
in an irregular differentiation of the primary (diarch) vascular system, and
becomes intensified in the first place during the course of the normal growth
in thickness and subsequently during the anomalous growth; the spiral
arrangement may, moreover, be explained as a result of the position of'the
radicle in the seed and of the mechanical pressure, which is exerted upon it
by thb cotyledons. This spiral arrangement has been found both in the main
root, developed from the radicle, and in the lateral roots; it is only a specific
character, being recorded by Fron in species of Anabasis, A triplex, Chenopodium,
Haloxylon, Obione, Salicornia, Salsola and Suaeda. It remains to mention that
in 'One and the same species the vascular bundles of the root are occasionally
arranged according to the' first type, while those of the stem are arranged
acmrding to the second type; this is the case in Beta, Blitum, Spinacia, and a
few species of Chenopodium (e.g. C. Bonus Henricus, C. murale and C. rubrum).
~

Literature: [Paschkis, Pharmakogn; Beitr., Zeitschr. Osterreich. Apotheker-Ver., 1880, n. 27,

28; abstr. in Bot. Centralbl., 1881; i, p. 54 (Chtnopodt·um).]-Mangin, Celloles spiralees, Bull. Soc.
bot. de France, 1883, p. 16 ; and Ann. sc. nat., ser. 6, t. xiii, 1883, pp. 'Jl4, 'l15.-Schnlz, Epidermis-
ullen bei Salicornia herbacta, Ber. deutsch. bot. Gesellsch .. i 886, p. 53 .-(Vi bouchevitcll, Plan tea
des terrains ,;a Iants, Rev. se. nat. appliquees, 1893; cited from Montell.]-(Pammel, Russian Thistle
(Sa/so/a Ka/i), Bull.. Iowa Agricul.t. College Exp. Stat., n. 36,.1~94, 33.Pp.]-:-[Pontebnia, "!-t.
surles
halophytes de la Cnmee. 1894; CIted from MonteU.]-(Tognlnl, Stoml, Atti fst. bot. Pavia, 1894.]
-Fron, Racine des Suaeda et des Salsola, Comptes rendus Paris, CXX1', 1897, pp. 366-8.-FroD,
Struct. spirnlee d~ rae. d. cert. Chenopodiac., Comptes rendas Paris, cxxvii, 1898, pp. 563-5; and
BetteJave, loc. cit., pp. 397-400.-Fron, Rech. anat. sur la racine et les tiges des Cbenopodiac.,
Ann. sc. nat., ser. 8, t. ix, 1899, pp. 157-240 and PI. v-x.-Herzog, Monogr. d. Zuckerriibe,
Hamburg, 1899, p. 4 et seq.-Hirsch, Entwickl. d. Haare, Diss., Berlin, 1899, p. 29.-Leisering,
Korkbild. bei den Chenopodiac., Ber. deutsch. bot. Gesellsch., 1899, pp. l~3-55 and Tab. xix.-
Leisering, lnterxylare, Leptom, Diss., Berlin, 1899, pp. 9- 1 I .-Cassan, Camp},qrosma monspelitUtJ,
Th~, Montpellier, 1901, pp. .31-46.-Solms~Lanbacb, SpilOlobe Chenopodeen, Bot. Zeit., 1901, i,
pp. 168-7o.-Jonsson, Wiistenpfl., in Lunds Univers. AT!skrift, xxxviii, Afd. 2, n. 6, 1903, pp. 6-18
and Tab. i, ii (Haloxylon, Eurotia).-Pons, A/riplex, Nuovo Giorn. bot. ]ta1.. N. S., ix, 1903,
pp. 35-48 and Tab. i.-Theorin, Vii.xttrichom., Arkiv for Bot., i, 19°3, p. I73.-Chrysler, Strand-
plants, Bot. Gazette, JtXxvii, 1904. p. 461 et seq. (A/rip/ex).-Sarton, Anatomie d. pI. affincs, Ann.
$C. n~t., $r";". 9, t. ii. 1905, pp. 53-9 (A/riple:c).-Holtennann, Einflass d. Klimas, 190 7, pp. 81,85
and 86 (Aln'pkx, Arl/lrlunemum, Salicornia, Suatda).-Montell, Anat. compo de la feuille des
Chenopodiac., These, Paris, 1906 j also in Perrot, Travaux, iv, 1907. 156 pp.

BASELLACEAE (pp. 663,664).

Large spherical mucilage-cells are found in the mesophyll also in Basella
paniculata, Volkens.
Literature: VollceM, Baullaptllliculala. in Englec, Bat. Jahrb., xxxviii, 1905. p. 81.

PHYTOLACCACEAE (pp. 664-668).

I. REVIEW OF THE ANATOMICAL FEATURES. We may first add that
anomalous structure of the ,-oot has been recorded also in Agdestis, Petive,ta
and Rivina, and anomalous structure of the stem also in Agdestis and Barbeuia.
The earlier statement to the effect that clustered crystals of oxalate of lime
art; absent requires modification since they have been observed in Stegnospetma
by Walter. According to the same authority sphaerites occur in Barbeuia.
2. STRUCTURE OF THE LEAF. Walter has recently undertaken a careful
 ADDENDA-PHYTOLACCACEAE 1031
•
investigation of the stomatal apparatus In this Order and finds that . .the
Rubiaceous type is not sufficiently widely distributed in any genus to admit
of its bemg regarded as a true generic character. He observed that the stomata
are in all cases provided only with a number of ordinary neighbouring cells
in the genera A chatocarp.us, A,gdestis, A nisomeria, Barbeuia, Didymotheca,
ErciUa, Gyrostemon) MicrQtea, Plw.ulothamnus, Se~uieria, Stegnosperma and
Tersonia.
The same author makes the foHowing statements (some of them new)
regarding the mode of excretion of oxalate of lime. The genera Phytolacca,
Anisomeria and Ercilla (Phytolacceae) possess raphides, while Barbeuia belong·
iug to the same taxonomic group has sphaerites. The typical members of
the Rivineae (Gallesia, Ledenbergia. Mohlana, Monococcus, Petiveria, liivina,
Schindleria, Seguieria, Villamilla) are distinguished by having styloids, whilst
the genus Stegnosperma, which forms the group Stegnospermoideae in \;Valter's
system of classification, is characterized by the possession of clustered crystals
(sphaerites according to Schulze, see Syst. Anat., p. 665). The typical members
of the Gyrostemoneae (Codonoca"pus, Didymot1ieca, G)'1'ostemon, Tersonia)
have no oxalate of lime at all. Regarding the remaining genera, whkll ~are
excluded from the Phytolaccaceae by Waiter, we may mention the following
details on his authority. Agdestis (' Genus anomalum Phytolaccac.' in Bentham
and Hooker, Gen. Plant.), and Gisekia (a member of the Ficoideae in Bantham
and Hooker, Gen. Plant.) are provided with raphides; Limeum and Semonvillea
(members of the Ficoideae in ,Bentham and Hooker, Gen. Plant.) have clustered
crystals, while Microtea has sphaerites. In Phaulothamnus Walter observed
only small crystals of oxalate of lime (in the ovary), while in Achatocarpus,
Adenogramma and Polpoda he failed to find any oxalate of lime at all.
3. STRUCTURE OF THE AXIS. Regarding the recently discovered cases of
anomalous structure in the fibrovascular system of the stem and root, see
above under 1, and_ Walter, loc. cit. ; for Agdestis, see also Cobau, lac. cit. In
the stem and root of Agdestis clematidea, Mo~. et Sesse the secondary zones of
vascular bundles develop in the pericycle (Cobau). Walter failed to observe
anomalous growth in thickness in the axis of the following genera: Achato-
carpus, Adenogramma, Codonocarpus, Didymotheca, Gisekia, Gyrostemon,
Ledenbergia, Limeum, Microtea (here also wanting in the root), ..Uonococcus,
Personia, Phaulothamnus, Polpoda, Psamtnotropha, Rivina, Senzonvillea,
Stegnospelma, Tersonia and Villamilta.
The perforations of .the vessels are invariably simple also in the genera
A chatocarpus, A denogramma, Agdestis, Barbeuia, Didymotheca, Gisek1.·a, Leden-
bergia, L'Imeum~ M oklana, M onococcus, Petiveria, Phau lothamn us, Polpoda,
Psammotropha, Schindleria, Semonvillea, Stegnosperma, Tersonia and Villamilla.
In Adenogramma. Agdestis, Anisomeria, Barbeuia, Didymotheca, Ercilla, Gisekia,
L1meum, M icrotea, M onococcus, Petiveria, Phaulothamnus, Polpoda, Psammo-
tropha, Semonvillea, Stegnosperma and T ersonia the walls of the vessels, where
in contact withllarencbYma, are occupied by bordered pits; in Achatocarpus
and Villamilla there are both bordered and simple pits at these points, and
in Ledenbergia, Mbhlana and Rivina simple pits only.
Literature: Cobau, Anatomia della Agdelli.r dl!malidta, Boll. R. Orto bot. 4.i Palermo, ii,
1898) pp. 111-,.,.-[ Kraemer, The pith-cells of Plzyto/tuCIJ dtcandra, Torreya, ii, I902i l'p. 1",1 -3.]
-Col, Fai~a.ux, Ann. sc.. nat., ser. 8, t. xx, 1904, p. J.67.-Netolitzky, Dikotylenbl. (Rhaphiden),
1905, p. 38.-Walter, Diagramme der Phytolacc., Diss., Erlangen, 1906, PP. 58-61; also in Engler,
Bot. Jahrb., xxxvii.-[Sen{t. Radix Phytolaccae decandrne, 'harm. Post, 1906, p. 281.]-[Holm,
PAytolar,a tkc4nara. in Merck's Report. xvi., 1907. pp. 3U-14.]
 ADDENDA

BATIDEAE (pp, 668,669).

According to Van Tiegbem a particularly noteworthy feature lies in the
arrangement and course of the vascular bundles in the quadrangular stems,
which bear opposite leaves. Corresponding to each of the angles of the stem
there are 3-5 bundles i, the vascular ring, these bundles being separated from
one another by multiseriate medullary rays, while opposite each lateral surface
of the stem there is a single smaller vascular bundle, The two smaller bundles
belonging to two opposite sides of the stem show a decrease in size as they
are traced up towards the next node above; they first loose their xylem,
, and after that their phloem, while Ultimately-in the neighbourhood of the
node-even the strand of pericyclic bast-fibres belonging to the bundle dis-
appears. These smaller vascular bundles therefore do not pass out into the
petiole, which is supplied by the two nearest of the bundles corresponding to
the angles of the stem. These bundles fork and the two middle bundles thus
produced fuse to form a single median bundle, the lateral branches thereupon
forking once again. In this way the base of the leaf is supplied by five vascular
bundles, which are arranged in an arc.
Van Tieghem records clustered crystals of oxalate of lime in the pith
and primary cortex, and solitary and clustered crystals in the medullary rays
of the bast. The peculiar transverse arrangement of the pairs of guard-cells
is found also on the axis.
Literature: Van Tieghern, Batidacees, Journal d~ Bot., 1903, pp. 363-76, especially pp. 365-7.
,

POLYGONACEAE (pp. 669-674).

1. REVIEW OF THE ANATOMICAL FEATURES. We may add that secretory
cavities containing a resinous secretion occur in species of Polygonum, a..'1d that
simple uniseriate clothing hairs and shaggy hairs have been observed in this
Order. Cortical vascular bundles, situated in the pericycle, are found also in
Rumex bilormis.
2. STRUCTURE OF THE LEAF. The upper epidermis of the leaf contains
numerous cells with gelatinized inner membranes also in Polygonum acre,
H. B. K.; these cells are almost spherical and penetrate rather deeply into the
mesophyll, While their lumina in most cases include a body, which resembles
a clustered crystal and refracts the light doubly; this body is soluble in
hydrochloric and sulphuric acids, as well as in caustic potash (in the latter case
a simply refracting skeleton remains undissolved). The stomata in Polygonum
acre occur in small numbers on the upper side of the leaf as well, and are pro-
I vided with subsidiary cells arranged according to the Rubiaceous type. Per-
drigeat describes three neighbouring cells as of qui~e general occurrence.
'I Unicellular clothing hairs are found in Antigonon (Perdrigeat). The long
conical shaggy hairs present on the margin and median vein of the leaf in
Polygonum acre are composed of lignified fibrous cells with thick walls and narrow
lumina. The same species also has discoid glands with a low bicellular stalk
a.nd a multicellular head, divided by vertical walls.
To the section dealing with the internal secretory receptacles we may
add that Peltrisot has observed secretory cavities, which are of-quite a peculiar
type and have oily-resinous contents, in the leaves and branches of Pol'Y~onum
\ Hydropiper, L. These cavities are schizogenous in origin and are surrounded
by four epidermal cells, which penetrate into the primary cortex or mesopbyll
as the case may be. To judge- by' analogy the' secretory cells (or secretory
cavities),' previously recorded in two other species of Polygonum (P. acre and
 ADDENDA--POLYGONACEAE 1033

P. punctulatum) are probably also of the nature of cavities in all cases. This
is at least true of P. acre, as I have found by an investigation of this speci~.
In this plant the secretory cavities are likewise situated in direct contact with
the two surfaces of the leaf, but they are surrounded by a relatively large
number of low epithelial cells; it remains to be investigated whether all these
epithelial cells and consequently the secretory cavities themselves are of
epidermal origin in this case. Tannin appears to be very widely distributed in
the Polygonaceae, especially among the desert-plants ..
The clustered crystals of oxalate of lime present in the primary cortex
and mesophyll occasionally attain quite a considerable size (e.g. in Polygonum
acre).
For the stmctureof the petiole (the vascular bundles of which are invariably
isolated) and of the stipular ochrea, see also Perdrigeat, lac. cit.
3. STRUCTURE OF THE AXIS. Numerous new statements on the structure
of the stem Will be found in Perdrigeat's paper, which deals with species of all
the genera enumerated in Dur~nd's Index with the exception of Hollisteria.
'1]Us piece of work is mainly concerned with the vascular system of the axis
and leaf and with the structure of the cortex, while the treatment of the hairy
covering and the secretory organs is inadequate.
The following details regarding the structure of the cortex are taken from
Perdrigeat's paper and the remaining literature. Assimila tory tissue in the
form of palisade-parenchyma is found in the primary cortex not only in Cal-
ligonum comosum (see Fig. 162 on p. 673), but also in species of Chorizanthe
and Oxytheca. In an undetermined species of Calligonum examined by Jonsson
the outer parts of the walls of the epidermal and, hypodermal cells become
swollen, so that the cell-cavities, which are surrounded by the inner parts of
the walls, appear as though they were embedded in a mass of mucilage. The
endodermis may be provided with Caspary's dots on its radial walls (e.g.
in Koenigia islandica, L.); in Rumex Patientia, L. it is sclerosed. According
to Perdrigeat the pericycle is invariably sclerenchymatous in the subaerial
parts of the stem; it contains either isolated bundles of fibres or a closed
or slightly interrupted ring of fibres or a composite and continuous ring of
sc1erenchyma; the latter has been recorded in Campderia floribunda, Coccoloba,
E,iogonum sphaerocephalum, Doug!., Leptogonum domingense, Benth., and
Podopterus mexicanus, Humb. et Bonp!. ~ The bundles of pericyclic fibres
show a radial development similar to 'that found in Calligonum comosum
(see Fig. 162) also in species of Coccoloba and Pterococcus. The cork arises
subepidermally also in Coccoloba, Rumex tingitanus, and Triplaris americana,
I superficially' (probably in part in a subepidermal posHion) in Calligonum

comasum, L'Herit., Eriogonum 'panitolium,' Leptogonum domingens~, Benth.,

Podopterus mexicanus, Humb. et Bonpl., Pterococcus, Ruprechtia apetala and
Symmeria paniculata, while it develops in the peri cycle also in Harlordia
macroptera, Greene et Parry and Pteropyrum scoparium, J aub. et Spach. The
cells of the~cork' for-the most part have thin walls. In Ruprechtia apetala
there is an alternation of layers of cork-cells with thick and thin walls; in
Harlordia macroptera the cells of the cork have strongly thickened tangential
walls, while in Coccoloba Schomburgkii they are thickened in the shape of a
horseshoe. In the species examined by Jonsson (Calligonum sp., Atraphaxis
canescens, Bge. and A. spinosa, • Eichw.') the cork contains mucilaginous
j

cork-cells J (see under Chenopodiaceae, p. 1029). The secondary bast not

uncommonly includes chambered parenchyma with clustered crystals, but
bast-fibres are rare (Rumex scutatus, L.).
In many members of the Order and especially in the herbaceous forms
there is ~ lysigenous cavity in the pith, whilst in Nemacaulis NuttaUii, Benth.,
for example, the entire pith becomes lignified at an early stage and is therefore
1034 ADDENDA-POL YGONACEAE

persistent. The occurrence bf bundles of fi.1>res, which are often very strongly
developed, or of a zone of fibres at the margin of the pith is a very common
feature (e.g. ¥t species of Campderia, Coccoloba. Emex, Oxyria, Podopterus,
Pteropyrum, Rumex, Triplar-is, &c.).
Anomalous structure of the stem has recently been demonstrated also in
Rumex bi/ormis (in the form of variously orientated vascular bundles, which are
enclosed in the pericyclic strengthening ring), as well as in Rumex conglomeratus,
MUlT., R. intermedius, R. obtusifolius, R. purpureus, Poir., Rheum hybridum,
MUlT., R.leucorrhizum, Pall. and R. undulatum, L. (medullary vascular bundles,
the records in the species of Rheum referring to the a.."{is of the inflorescence)
(Baranetzky, Perdrigeat, Saget).
Literature: [Theorin, Vaxtslem uti knoppame hos fam. Polygon., Stockholm, I87:.J.l-Schubert,
Parenchymscheiden, Bot. Centralbl., 1897, iv, pp. 469-7 I.-Wollenweber, Anat. d. Schwimmbl.,
Diss., Freiburg in Br., 1897, pp. 33, 34.-Hammerle. Polygonum cuspidalum, S. et Z., 'Diss.,
Gottingen, 1898, 70 pp.-Montemartini, Fnsto del Folygonum SieofJidii, Reinw., MaJpighia, 1898,
pp. 78-80 and Tab. iii.-Boergesen og Paulsen, in Bot. ,Tidsskrift, xxii, 1898-9. pp. 16-18 (CoccolclJa
"'Dirt,,,, Jacq.).-B~ranetzky, Faisceaux bicollatl~raux, Ann. sc. nat., sere 8, t. xii, 1900. pp. 3°7-14.
-fBernalsky, Anal. Bestimm. eil1heim. Pdy(Onum-Arten, Termesz. Fiizetek, xxiii, 19oo, pp. i, 11
nna pp. 66-74']-Thomas, Fenilles sout., These, Paris, 1900.-Dye. Unterird. Org. von ValeriaM;
Rheum u. I nula, Diss., Bern, 190 I, pp. 36-64 and Tab. i. ii.-Perdrigeat, Anat. compo des Po)ygonac.,
Act. Soc. Lion. de Bordeaux, lv, I9ot, 94 pp., 3 pl.; also These', Bordeau:x.-Pitard, Pe ricyc Ie,
These, Bordeaux, 1901, p. 48.-Jonsson, Wiistenpfl., in Lunds Univ. !rsskr., xxxviii, Afd. 2, n. 6,
1901, pp. J8-l:ll and Tab. ii (Calligonum, Atraphaxt's).-[Mitlacher, Herba Polygoni avicularis,
Pharm. Post, J902, D. 56; abstr. in JUSt, 1902, ii, p. 4s.]-Peltrisot, Org. secret. du Po!ygtmU11I
H)'dropiper, L., Joum. de Bot., 1903, pp. :J23-8.-Saget, lit. anat. des l1umex crispus et R.
oNusifo/ius, etc., These, Montpellier, 1903, pp. :.J6-3I.-Chrysler__. Strand-plants, Bot. Gazette,
xxxvii, J904, p. 461 et 5eq. (Poo-gonum).-Eijken, Rhabarberrhizome, Diss., Bern, 1904, p. 45 ct
seq.-Freidenfeldt, Anat. Bau d. Wurzel, Bibl. bot., Heft_:6I, J 904, pp. 37 J 38.-Cristofoletti, RAeum
mponticum, Diss., :Bern, I905.-POlSCh, SpaltDffnungsapparnt, !e-'!.a., J905, p. 87 and Tab. i i -
. [Litscbauer, Eingesenkte epid. Drusen bei POlJ'gonum Hydropiper, Osterreich. hot. Zeitschr., 1907.
pp. 201-4.]-[For further literature, see p. It71.]

PODOSTEMACEAE (pp. 674-676).

The following details are taken from recent papers on the anatomy of the
Podostemaceae. According to Mildbraed the drifting stems are constructed
so as to have tensile strength. In the axis of M ourera fluviatilis, Aubl. this
author observed a central cylinder, which consists of long pitted mechanical
cells and is traversed by air~cana1s, while the peripheral tissue contains a large
number of isolated vascular bundles which are arranged in three indistinct
rings and show the ordinary type of structure; in the neighbourhood
of the air~canals of the central cylinder, moreover, spiral and annular vessels
were found. The stems of the Apinagieae examined by Mi1dbraed (especially
Apinagia Riedelii, Tul.) have a central vascular bundle, which includes an air~
eanal and is enveloped by a rather thick stereom~cy1inder composed of lignified
cells bearing slit-shaped pits. These statements serve to show that mechanical
cells are not as rare in this Order as has hitherto been supposed.
. H. Moller observed cubical and prismatic crystals of oxalate of lime in
the epidermis of the root of Cladopus Nymani, Moll.; previous to that only
clustered crystals had heen recorded in the Podostemaceae. In the event
of fresh investigations being undertaken on the members of this Order it will
be well to devote special attention to the occurrence of interce11ular secretory
receptacles. Secretory spaces resembling. resin~canals had previously been
observed by Wachter in the root of WeddelUna squamulosa, and on Mildbraed's
authority we may class with them the following two types of elemen~: (a\Secre-
tory cavities, which have yellow resinous contenfs so}uble in alcohol and are
apparently lysigenous in origin, in the outer tissues and central cylindetof the
scape of MOUfera fluviatilis; (b) the secretory cavities, which occur in the
 ADDENDA-PODOSTEMACEAE I035
thallus of Castelnavia Lindmaniana. Warm., and appear in the form of brownish
streaks in alcohol-material; they arise ly~igenously from a row of about 3-10
secretory cells; their secretion is in part thread-like and granular and under-
goes but little change on treatment with alcohol, ether or xylol. Here we
may call to mind the fact that Goebel likewise observed the exudatign of a
yellow secretion on cutting fresh stems of Rhyncholacis macrocarpa, luI.; the
organs in 'which the secretion is contained have not yet been demonstrated
in this caSe.
It remains to consider the red bodies' (' Warming's bodies '), which
I

haye been subjected to a careful examination by Mildbraed. They were

fin,t observed by Warming in the spatheUa of Apinagia Riedelis', but occur
also in the wall of the capsule and in the thallus of Castelnavia Lindmaniana.
Judging by the reactions mentioned by Mildbraed it seems as though we were
after all dealing with silica·bodies, which are infiltrated with a substance
resembling anthocyanin, although Mildbraed's statement that these bodies
are not affected by hydrofluoric acid is incomprehensible in this connexion.
In view of the other reactions, however, of what can the ground-substance
consist, if not of silica?
Literature: Hjalmar Moller, Cladopus Nymani, I]OV. gen., Ann. Jardin Buitenzor~, xvi, 1899,
pp. 115-31 and Tab. xii-n.-Warming, Fam. Pod ostemac. , Afb. v, 1899; and Afh. vi, 1901,
Vidensk. Selsk. Skr.-[Willis, Podostemaceae of Ceylon and India, Ann. Roy. Bot. Gardens,
Perade.oiya, i. Part iv, 1902, pp. 267-46!, and pI. iv-xxxviii; detailed abstract in Bot. Centralbl.,
xcii, pp. 193-8.]-Mildbraed, Beitr. z. Kenntnis der Podostemac" Diss., Berlin, 190 4, 4' pp.

NEPENTHACEAE (pp.676-680).
According to Fenner's investigations the digestive glands (p. 677) ot
Nepenthes Raglesiana, Jack are of the nature of emergences, since subepidermal
cells are also concerned in their formation. The same author also states that
the cuticle of the glands is perforated in asieve ..like man~er, and that ~he walls
of the cells composing the uppermost layer of the bod}'. of the gland ' are pro-
vided with membranous ridges. In Nepenthes Ratftesiana the outer wall of the
pitcher bears various types of hairs including peculiar clothing hairs with
dendroid branching.
With reference to the structure of the' axis (p. 679) we may note that,
according to Heinricher, the vessels in Nepenthes melamphora, Reinw. attain
a diameter of '18 mm., that spiral tracheids are found -also in the medullary
rays of the cortex, and that fusiform proteid-bodies occur abundantly in the
cortical parenchyma of the rhizome, but only 41 small numbers in the c~tex
of the climbing stem. .
Literature: Mangin, Cellules spiralees, Boll. Soc. bot. de France, 1882, p. 14 et seq., and AnD.
sc. nat., sch. 6, t. xii, 188:l, pp. :H2-I4 and pl. viii.-Fenner, Anat., Entwicklungsgesch. u. Biologic
der Laubbl. 11. D~sen ein.iger .lnsektivoren, Diss., Zurich, 1904, pp. :.8-33 and Tab. ix, :It; also in
Flora, ~civ~ I904.-Heinricher, Nepenthes, Ann. Jardin bot. Buitenzorg, xx, '2, 1906, pp. :177-98,
and Tab. ~xiv-xxvi.

CYTINACEAE (pp. 680, 681).

Trflcheids have been observed in the myceloid vegetative thallus of Pilo·
styles Ingae (Karst.), by Endriss, while on the lower side of the scale-leaves
borne on the floral shoots of this plant the same author found stomata, which are
formed by a~ingle division in the epidermal cells. Stomata have recently been
observed also by POTsch on the under side of the lowest scale·leaves in Cytinus
Hypocistis, L. Schaar's investigation of RatflesiaRochussenii, Teysm~ et .Binn.
has shown that the vegetative thallus in this species consists only of hyphal
1
I0g6 AJ)lJ£N/).A.-CYTINACEAE

cells and that the cond.ctingtissue, which is differentiated in the floral cushion,
is formed by about 2<1 ~ of vascular bundles exhibiting an annular arrange-
ment.
LiteratlU"e: Schaar, RQ.ffIe~1a R~hu.r~·, Sitz.·Bt'r. Wiener Akad., cvH, Aht. i, 1898, pp.
1039-56 and Tab. i-iii.-Solms-Laubaeh, Ra~esiaceae u. Hydnoraceae, in Pflanzenreich, Heft 5,
'90', Raffles. pp. :3, 3; Hydaor. p. a.-Endrin, A1ost.1les Ingrze, Flora, xci, 190<1, ErgaDZ.-Bd.~
pp. 209-36 and Tab. :u.-Pom:h, Spo.ltoffnungsapparat, Jena, 19°5, pp. 7o-a.

ARISTOLOCHIACEAE (pp.682-688).
2. STRUCTURE OF THE LEAF. A few data regarding the course of de·
velop'ment of the stomata will be ~ound in Montemar~ini's .paper ~loe. dt.,
Paplllae are present on the lower sIde of the leaf also m Anstoloch_~a ellgam •
Mast. (Knothe). The followin~ details regarding the structure of the secretory
cells are based on investigations undertaken by Berthold, Haberlandt and.
R. MUller on Asarum europaeum~ A,istolochia brasiliensis and A. Clematitis.
In the fully differentiated oil·cell the drop of oil is completely enveloped by
a sheath, the upper part of which is formed by the wall of a ' basin-' or funnel·
shaped structure, which arises as a thickening from the cell-membrane, while
the remaining portion of the'sheath is constituted by the.wall of the-vacuole.
(' pouch '), which has undergone a change in substance. In a surface·section
the 'basin' appears in the form of the pit·like structure represented in
Fig. 166, B (on p. 684). R. Muller has also shown that the secretory cells in the
leaf of A,istolochia brasiliensis, although apparently belonging to the integu·
mental tissue, are differentiated in a
subepidermal position, but they soon
come to lie at the surface as the result of sliding growth. Since the statements
hitherto published as to the occurrence of secretory cells in the epidennis
are based only on an examination of the mature leaves, it will be necessary to
undertake a developmental investigation in all these cases. According to Van
Tie~hem the absence of oxalate of lime in the branch and root and the possession
of Oll·cells are features characteristic of a certain group of species of A l'istolochia,
which he comprises as Aristolochia, L. in contrast to a second group of species
(Hocquartia, Dumort.), which are distinguished by having c.lustered crystals
and by the absence of oil-cells (also in the hairs).
3. STRUCTURE OF THE AXIS 1. According to Schellenberg the pith of
A ristolochia SiPho and other twining species is composed of cells with thin
unlignified walls, and undergoes compression so that the stem becomes adapted
to~~sist tensions# Stone-cells occasionally also occur in the pith (Aristolochia
Sipho and A. gigantea).
Literature: Power, Asarum cateiJdlnse, Diss., Strassburg, 1880, pp. ~-Il.--rLa!arski, Asarum,
Pha.rm. Post, [881, n. 3, .oJ; abstr. in Bot. CentralbJ., I88t, ii, p. 49.J-Berthold, Protoplasma-
. mechanlk, ;Leipzig, 1886, p. 26.-[Hooper, Bragantia Wallickii, Amenc. Joorn. of Phann., 1894'-
p. 23 1 .J-Schwabaeh, Mech. Ring, Bot. Centralbl., 1898, pp. 354-7.-&hellenberg, Entwick·
lungsgescb. d. Stammes von AmtolocMa Sipho, Festschr. f. Schwendener, 1899, pp. 301-20 and
Tab. xii.-(Collin, Arisl%ckia Se,.penta1'ia, Jouro. de Pharm. et de Chimie, 1900, p. 309; abstr. in
J~st, t900~ ii, p. 16.J-Van Tiegbem, Hocquartie, Journ. de Bot., 1900, pp. 6s-8.-Pitard,
Pbicycle, These, Bordeaux, 1901, pp. 43-6.-Knotbe, Un~netzb. Epid., Diss., Heidelberg, 19°2,
p. 15.-Montemartini, All8.t. compo delle Aristolocb., Atti dell' 1st. bot. di Pavia, 2& ser., vii, '90:1,
pp. aa~-so and Tab. xii-xvi; $ep. copy, pp. 5-10.-Perrot, Particularite de sthlct. fol. chez cert.
feniUes de I',Anstoloclua SipAo. Bull. Soc. bot. de France.. 1902, pp. 163-6.-Itabedandt, PhysioJ.
Pflanzenanat., 3rd edit., 1904, pp. 46:a-4.-R. MUlleT. Olbehalter, Ber. deutsch. bot. Gesellscb.,
1905, pp. 29 2 -i.-[Holm, Aristolochia Serpentaria, in Merck's Report, xvi, 1907, PP.·,276-9·]

1 Regarding the incorrect statement (in .Perrot, These. 1899, p.. 144) as to the occurrence of
secondary bundles of wood and bast in members of the Aristolocbiache, see the footnote on p. 68'1.
 ADDENDA 1037

PIPERACEAE (pp.688-694).
2. STRUCTURE OF THE LEAF. In A1Iemiopsis cfltifornica, Hook. et Arn.
the stomata are present on both sides of ~ leaf, befng almost more abundant
on the upper than on the lower side; they are surrounded by 4-6 neighbouring
~ells. In the same species a hypoderm composed of a single layer of large
cells is developed on the upper side of the leaf (Holm).
New data regarding the nature of the hypoderm in the Brazilian species of
Peperomia are contained in Jaderholm's paper (IDe. cit.). In the first place we may
note that there is no hypoderm in P. teneTa, Miq. and P. pettuC£da, H.B.K.; in the
latt.er species the large-celled epidermis which is differentiated as aqueous tissue
compensates for the absence of hypoderm. In Peperomia Gardneriana. Miq. the
bypodeml usually consists only of a single layer of cells; in P. diaphana, Miq. it is
I')omposed of two layers, and in P. Caldasiana, C.DC. of several layers of cells,
which in both cases are of large size; in P. trineuroides, Dahlst. the hypoderm is
developed in the same way as in P. pereskiaefolia. In P. reflexa, Dietr. It consists
of very numerous layers, in P. Sellowiana, Miq. of several layers ; in the two species
last named the inner hypodermal cells are arranged in rows at right angles to the
surface of the leaf. Regarding the hypoderm found in the species of Pepet'omia.
see also Jonsson, lac. cit.
According to Duval hypoderm (in some cases situated on both sides of the
!eaves) is found also in Piper ceanothi/olium, H.B.K .• P. citrilolium, Lam., P. coreo-
vadense. C.DC., P. Jaborandi. Vell., P. laetum, C.De., P. molUcomum, Kth., P. ,.eti-
culatum, L., and P. unguiculatum, Ruiz et Pave ; according to Holtermann also in
Piper Thwaitesii.
In the Brazilian species of Peperomia examined by Jaderholm the palisade
tissue consists of a single layer of short funnel-shaped cells. It is typically
differentiated only in P. muscosa, Link (?), a1though even here the cells are of no
great length. In the peculiar leaves of P. dolabritormis, H. B. K. and P. nivalis,
Miq. palisade tissue is completely wanting; for details regarding the structure
of these leaves, see C. de Candolle.loc. cit. According to ] aderholm P. Calda-
siana and P. Sellowiana are distinguished by the fact that the cells of the
uppermost layer of the spongy tissue, which borders on the palisade tissue,
have very thick walls and store up starch, while P. pellucida is characterized
by the walls of the entire spongy tissue being hyaline. Conical papillae occur
on the upper epidermis also in P. increscens, Miq.; in P. diaphana, Miq. some
of the upper epidermal cells are papillose.
'. Noteworthy forms of clothing hairs are consti tuted by the sma1l2-5-celled
trichomes of Peperomia pulchella, A. Dietr., and the small unicellular hairs of
P. ,esedaeflora, Lind. et Andre. These hairs are seated on a large epidermal
cell, which is either vesicular or projects in a convex manner and constitutes
an ocellar, apparatus; in the second of the two species the efficacy of this
organ for the perception of light is further increased by a cell of characteristic
shape, Which is apposed to the inner side of the protruding epidermal cell
and is occasionally differentiated as an oil-cell; this cell and the one above it
are both derived from the same mother-cell (Haberlandt). In this connexion
we may mention the numerous fiat tubercles, which are found on the surface
of the leaf in Peperomia metallica, and according to Haberlandt function as
lenses for the CQIlcentration of light; they are formed by a group of epidermal
cells showing tangential divisions. In this case an oil·cell which is situated
exactly beneath the centre of each group of cells functions as a second eondensing
lens. For 'the diverse forms of trichomes found in Piper, and especially the
clothing hairs, see Plate IX in Duval's paper., lac. cit.
Jliderholm mentions the occurrence of crystals of oxalate of lime re-
sembling raphides also in Peperomia maior, C. DC., ~hile Duval records them
 ADDENDA-PIPERACEAE

in certain species of Pi-pe~. Brownish acicular crystals, which are soluble in

acetic acid (and therefore do not consist of oxalate of lime), were observed by
1aderholm in the axis of PeperomflJ fri1'l!euroides, Dahlst.
Haberlandt has recently demonstrated the importance of the secretory
tells in certain species of Peperomia as organs for the condensation of light. We
may add that in this Order, as in the Aristolochiaceae, the oily secretion is en ..
closed in a pOllch.shaped protrusion of the cell-wall, a feature which is very dis ..
tinctly seen, for eX'Ullple, in Peperomia magnoliaejolia (Berthold and R. Miiller).
r
The nature of the secretory cavities poches secretrices '), recently observed by
Duval side by side with the secretory cells in the mesophyll of Piper hirsutuni,
Sw., still requires further investigation. According to Duval mucilage·canals
are found in the pith also in Piper ceanothifolium, H. B. K., P. citrifolium, Lam.
and P. lepturum., Kth. (in the last of'these species in addition to ruucilage-
cavities, which are·situated in the soft bast of the vascular bundles).
3. STRUCTURE OF THE AXIS. Among the species of Piper, examined by
Duval, P. Jaborandi, Yell. alone has meduUar:y vascular bundles. According
to Jaderholm exceptions to the fourth type (the Peperomia-type) are constituted
by Peperomia delicatula, Hench., which has four vascular bundles arranged
in a ring, and by P. tenera, Miq., in which the vascular system consists of a
single axile strand.
The terrestrial roots of Pt"per nJ'grum contain a pith; in the course of their
.
growth in thickness broad strips of thin-walled tissue constituting primary medullary
rays are alone formed on the outer side of the (5-10) groups of primary trachea~.
I failed to observe the small groups of thin-walled elements, which Keller records
in the secondary wood and regards as being , of the nature of interxylary phloem.
Literature: [Paschkis, Phannakogn. Beitr., Zeitschr. osterreich. Apotheker-Ver., 1880. n. 27.
,S; el-str. in Bot. Centralbl., J88I, i, p. 54-.]-Berthold, Protoplasmamechanik, Leipzig, 1886,
p. "5.-Keller, LuftWllI"Zeln, Diss., Heidelberg, 1889, pp. 50-s.-Went, Luftwurzeln, Ann. Jardin
Buiteazorg, xii, 1895, pp. 47, 48.-]onsson, Anat. Ball des RI., Acta Unil'. LlUld., xxxii, 2, 1896
(PejMromia).-[True, Kava-Kava, Pharrn. Review Milwaukee, xiv, 1896, pp. 28-32 j abstr. in Just,
1896, ii, p. 479.]-Dethan et Bertaut, Nauv. variete de Matico, Jaurn. de Pharm. et de Chimie,
SeT. 6, t. vi, J897, pp. 536-4o.-13jermann, (:)l:tel1en, Diss., .Bern, 1898, pp. 50-3.-Jaderholrn, Anat.
studier ofver sydarnerik. Peperomier, Diss., Upsala, 1898,99 pp. and 2 Tab.; German abstr. in B~
Centralbl., 189i, iv, pp. I90-3.-Siedler, Kawawurzel, Pharmazeut. Zeit., 1903. p. 78I.-Col,
Faisceaux, Ann. se. nat., sb. 8, t. XX, 1904, pp. 168-7o.-Haberlandt, Physiolog. Pflanzenanatomie,
1904, p. 462.-Haberlandt, Lichtsinnesorgane, 1905, pp. I.H-17 and Tab. iii.-Holm, Ammiolsis,
A1lleric. Joum. of Sc .. xix, 1905. PP' 76-82.-R. Miiller, Olbehalter, Ber. deutsch. bot. Gesellscb.,
J9o.,. p. 297.-Duval, Jaborandis, 1905, pp. 95-IIO, and pl. ix; in Perrot, Travaux, iii, 1906.-
C. de Candolle, Sur deux Pejeromia, etc., Arch. sc. phys. et nat. Geneve, 1907, sep. copy, 9 pp., I pI.
-Holterma~ Einfluss des Klimas, 1907, p. 136.

CHLORANTHACEAE (pp. 695,696).

According to H. Schulze's recent investigations the folIowing additional
charaat&s are important for the diagnosis of the Order. The palisade tissue
of the leaf consists of short cells and is often muriform. Arm-palisade cell:t
have been observed in all the species, which have been investigated. "'fhe
stomata are confined to the lower side of the leaf. Secretory cells have now
been demonstrated also in Ascarina, and therefore occur in all the members of
the Order. .
To the earlier statements regarding the secretory recepta~es we may add
that H. Schulze likewise observed the secretory cells of the leaf only in the
mesophyll. The relatively small secretory cells, which are particularly abundant
in the mesophyll of Hedyosfflum racemosum, Don, are exceptional in having
dark brown contents. Mucilage-canals occur in the larger veins of the leaf
also in H edyosmum arborescens, Sw. .
,I The structure of the leaf is in all cases bifaoaI. Typical palisade tissue
 ADDENDA--CHLORANTHACEAE 1039

composed of ]Qng cells is not found in any member of the Order. All th6 species
of Chloranthus, Ascarina and Hedyosmum h;}ve arm-palisade-cells, which vary
in number in the different species and ~ in the first and commonly also
in the second and third layers of the palisade tissue. In H edyosmum arborescens
and H. racemosum the cells of the spongy tissue are slightly sclerosed at certain
points. Schulze also states that distinct subsidiary cells are developed in relation
to the stomata only in certain species. He records two or more subsidiary
cells, which are placed parallel to the pore, for the stomata of Chloranthus
inconspicuus, Sw. and C. officinalis, Bl., and subsidiary cells arranged in the
form of a rosette round the stomata of Ascarina polystachya, Forst. The lateral
walls of the epidermal cells are either straight or undulated. A continuous
one-layered hypoderm beneath the upper epidermis, besides occurring in
Hedyosmum arborescens, is found in H. 1'acemosum and Ascanna polystachya,
while the hypoderm is confined to the neighbourhood of the veins in Chloranthus
brachystachys, Bl., C. inconspicuus, Sw., C. iaponicus, Sieb., C. otJicinalis, Bl.,
Hedyosmum ArlQcarpus, Solms and H. Bonplandianum, H. B. K. In Ascarina
lanceolata" Hook. f. and A. polystachya, Forst. the epidermal cells on the lower
side of the leaf are characterized by their strikingly small size, while in A. 1'ubri-
caulis, Solms they ale distinguished by bearing characteristic papillae, which
project in the form of crests in surface-view. According to Schulze the vas-
cular bundles in the lateral veins of the first order are for t~~ most part sur-
rounded by slightly sclerosed parenchyma. In Ascarina rubricauJis this
sclerenchymatous sheath is incomplete, while in Chloranthus serratus, Roem.
et Schult. and Hedyosmum nutans, Sw. it is absent.
As regards the structure of the wood we may a4d that Engler records
scalariform perforations (with numerous bars) in the vessels also in Ascarina.
The only form of crystals of oxalate of lime observed by Schulze were
small solitary crystals in the upper epidermis of the leaf of Chlo1'anthus tnco,fl,-
sPicuus, Sw.
ApPENDIX: THE ANOMALOUS GENUS Circaeaster.
The genus Circaeaster, which in Durand's Index is appended to the Chior-
anthaceae as a 'genus anomalurn,' differs quite essentially from the Chlorantha-
ceae in the absence of secretory cells. There are no special anatomical features
to give a hint as to the systematic position of the genus.
H: Schulze mentions the following details regarding the structure of the
leaves, which are very thin. The upper 'epidermal cells are elongated in a
direction parallel to the midrib of the leaf and have strongly undulated lateral
walls. The stomata, which are found only on the lower side of the leaf, are small
and not numerous; they have no subsidiary cells. There is no sclerenchyma in
the veins. Both crystals and secretory cells are absen t. Unicellular hairs, which
are bent in the form of a hook at the apex, are present only on the fruit.
With reference_J~t,the structure of the stem, Scott (in Oliver's description
of the genus,.loc; cit.) states that the fibrovascular sys:em is peculiar in being
diarch like that of the main root; only a small amount of secondary wood
and phloem is produced on the two sides of the diarch xylem-plate.
Liter.ature: Oliver, in Hooker, leones, pI. 2366, I 89s.-Harms, in Nachtr. 2U den natiirl.
Pflanzenfam., ii-iv, 1897, p. 333.-H. Schulze, in Beih. z. Bot. Centralbl., ix, 1900, pp. 81-5.

MYRISTICACEAE (pp.696-699)'
Literature: Poulsen, Abnorme rodbygning has en art af 51. Afyrislica, Vidensk. Meddeleis.,
1896, p. 188 and Tab. iii, iv.-Biermann, Olzellen, Diss., Bern, 1898, ,po ... 7.-Pitard, Perieycle,
These, Bordeaux. I90r, p. 93.-Hargagli-Petrucci. Legnami, Malpighia, 1902, p. 296.-Areschoug,
'hop. vaxt. bladbyggn., S.... Vet. Akad. Ha.ndl., 39, D. 2, 190 5. pp. 3!" 36 .
 ADDENDA

MONIMIACEAE (pp. 699-1(2).

2. STRUCTURE OF THE LEAF. The following details are taken from
Perkins' recent anatomical investigations 1, which serve to supplement Hobein's
earlier work. In the large majority of cases the structure of the leaf is bifacial.
In Amborella trichopoda, Baill. typical palisade tissue is wanting, while i.e
T,imenia weinmanniifolia, Seem., Piptocalyx Moor~i, Olivo and Glossocalyx
Staudtii, Engl. the entire mesophyll is composed of spongy tissue. Hypoderm
has been recorded also in A nthobembix, Epl",,"pp'iandra, Levieria, ~1 aC1opeplus,
Macrotor'Us, Nemuaron, Steganthera and Tetrasynandra, while it is wanting in
A tnborella, Glossocalyx, Piptocalyx and Trimenia.
Secretory cells have again been observed in all the genera recently ex·
amined by Perkins; the¥' are situated in the mesophyll. In view of the
statement that the leaves In Chktopatane, Engl. are provid.ed with transparent
dots, secretory cells are probably to be found .also in this genus.
3. STRUCTURE OF THE AXIS. According to Perkins the anatomical dis-
tinction between the Monimieae and Atherospermeae, based by Hohein on the
breadth of the primary medullary .rays, is on the whole supported by the
results of the former·s recent investigations. Thus, am6ng the genera of the
Monimieae recently examined, the medullary rays are: 2-4 seriate in Levieria,
1-4 seriate in Trimenia~ 3-6 seriate in Hennecartia, broad in Macropeplus and
Macrotorus, as in Mollinedia, 4-6 seriate in Steganthera, and Anthobembix, 4-7
seriate in Tetrasynandra, and I-Z seriate alone in A mb orell a ; among the genera
of the Atherospermeae recently investigated, Nemuaron and Glossocalyx have
1-3 seriate medullary rays. .-
Literature: [Hanausek, Folia Boldo, Zeitschr. Osterreich. Apotheker-Ver., 1880, p. ISS; abstr.
in Bot. kit., 1880, p. 4i'.... ]-[Stowell, Boldo leaves, Therapeutic Gazette, :880, p. 257 et seq.;
abstr. in Bot. Centralbl., 1881, i, p. 335.]-Perkins, Beitr. z. Kenntnis d. Monimiac., 1 (MplliQC<iieae),
in Engler, Bot. Jabrb .• xxv, 1898, pp. 549-53.-Perkins, lUollintdia, in Engler, Bot. Jllhrb., xxvii,
1900, pp. 638, 639.-[Neger, Folia Baldo, Pharmaz. Centralhallt:, 190ft n. 31; ahstr. in Just, 1901,
ii, p. 74.]-Perkins. Sipamna., in Engler, Bot. Jahrb., xxviii, 1901, p. 66J.-Perkins and Gilg,
Monimiaceae, in Pfl:mzc:nreich, Heft 4, 1901, pp. 2, 3.-Pitard, Pericycle These, Bordeaux,
1901, p. 66.

LAURINEAE (pp. 702"""106).

I. REVIEW OF THE ANATOMICAL FEATURES. We may add that hypoderm
occurs in the leaf also in species of Bellota, Endlicheria, Hufelandia, Nectandra,
Oco(ea, Persea, and Phoebe, and that the lower epidermis shows papillose
differentiation in species of Acrodiclidium, Aniba, Endlicheria, Nectandra,
Persea and Phoebe.
2 •.STRUCTURE OF THE LEAF. Our knowledge of the structure of the leaf
has been extended especially by Volker Petzold's recent investigations, which
deal with the Americm members of the Order. According to Petzo1d the
leaves, as a general rule, are bifacial in structure; palh:ade tissue is found on
the lower side of the leaf in Silvia and in species of Nectandra and Phoebe. but
it is never as strongly developed as on the upper side. The palisade tissue
consists either of one 2 or several layers ; according to Petzold the occurrence
of relatively large cavities in the palisade tissue is characteristic of most of the

I These deal with the following genera: laieria, Ambor(/la., Tn'mm;a, Pi"ptMa!yx, EIMpJian-
dra, Hennuarlia, .Nemuaron, G'o.rs(Ka~x, Ma(roJePlus, AI(kr%rJIs, Slegan'''~ra, .Anlhohemllix.
and Tttras),m:ndra.
I There is a single layer of palisade tissue in: Aniha (excepting A. rohm/a, Mez and .A.
J..'idleyana, Mez), Btnzoi" odoriftrum, Nees • .lJicypel/illm, Endlulttria (excepting E. impressa .•
.Mez) , Sasso/ras, Silvia (excepting S. polyonlha. Mez), SysftHlIJlUJdapllne, UrMnotUnd"m.
 ADDENDA--LAURINEAE
species of Persea (I.e. with the exception of three closely related species native
to the Andes). The sclerosed paljsade·ceUs previously recorded in a few species
of Ocotea were not observed by Petzold in Ocotea Kunthiana, Mez, although
present in O. ,ubra, Mez, where they show a somewhat different structure;
Holtermann also records stone-cells m the tissue of the leaf in Actinodaphne
molochina and A. speciosa. In some cases the spongy tissue contains large
lacunae, which are filled with stellate tissue; this feature is specially pronounced
in Persea (in contrast to almost all the species of- Phoebe) and is found also in
Systemonodaphne and Urbanodendron, as well as in species oi Acrodiclidium,
A nibt;l, Cryptocarya and Misanteca. The lateral walls of the epidermal cells
are only rarely undulated, and particularly rarely on the upper side of the leaf.
Lateral walls, which are bent in a zig·zag manner with ridge·like processes in
the apicei of the angles, are found in species of Cryptocarya, Hu/elandia, and
Sassafras 1 (for the most part on both sides of the leaf); high epidermal cells
occur in Bellota costaricensis, Mez, E ndlichtria impressa, Mez, and in species
of Aniba, Nectandra, Ocotea, Persea and Phoebe; papillose differentiation of
the lower epidermis has been observed in species of A crodielidium, A niba,
Endlicheria, Nectandra, Persea and Phoebe 2. Gelatinized epidermal cells have
not as yet been recorded in the Laurineae; nor are they present in Cinnamo-
m14m Camphora, Nees et Eberm. as I am able to state on the basis of an investi-
gation of this species, although Tschirch and Shirasawa mention the occurrence
of abundant mucilage in the epidermis of the leaf. In some species of Ocotea.
PeTsea and Phoebe a marking of the cuticle has been observed in the form of
a delicate punctation, but striation of the entire cuticle has not been found
in any case. In the species examined by Petzold the stomata are again con-
fined exclusively to the lower side of the leaf. They are, moreover, invariably
accompanied by subsidiary cells, one of which is placed on either side of. and
parallel to~ the pore (also in Persea iudica, Spreng., in opposition to Clauditz,
loco cit.). According to Petzold peculiar ridge·like process~ are present on
the pairs of guard·cells iIi species of Acrodiclidium, Aiouea, Aniba, Cryptocarya,
Dicypellium~ Endlicheria~ Misanteca, lVectandra, Ocoka and Phoebe, broad
subsIdiary cells showing 2. radial striation in two closely related species of
Phoebe (P. Pittieri, Mez, and P. psychotrioides, Mez), and depressed stomata,
the pores of which are placed at right angles to the slit.shaped vestibule, formed
by the subsidiary cells, in Misanteca capitata, Cham. et Schlecht., and most
of the American species of Cr'Yptocarya. On the stem of Cassytil.a /iU/ormis the
pairs of guard·cells show the same transverse arrangement as in C. ameTicana.
Hypoderm, has in the first place been recorded by Petzold on the upper side
of the leaf in all the species of CryptocaTya and Htlfelandia, as well as in certain
species of Bellota, Endlicheria, Nectandla, Oeotea, Persea and Phoebe 3. In
most cases it consists of a single layer of cells, rarely (Bellota nitida, Persea
boldufolia, Hufetandia rigida, l\Iez) of two layers. Hypoderm has been observed

I ,-it.: C,-yptocarya minutijlora, Mez, C. slI/xot')'lIlll()sa, Mez; Hufeltrlldia ematt;inata, Mez,

11. n"gida, M ez, H. TauiJe1-Hana, Mez: Sassafras van·if()/ium, o. K.
2 ,·iz.: Atrodididium ora.dlieme, Nees; Anilkl firnmla, Mez, A. Gardller', Mc~, A. nUlla, Mez,
A. lIbilleriana, Mez; Endliclzeda anomaltz, Nees, E ••·lIlpressa, Mez; Neetamira ja/ut'tnsis, Nees,
N. tllrbacensis, Nees; Persta klivimsis, Mez, P. tarolinmsis, Nee;, P. cnrysophylla, Mez, P.
c(urulea. Me~, P. cordata, Me:, P. dq11lin~nsis, Mez, P. mityopllylla, Mez, P. rfUt11l0Sa, Mez, P.
vestita, Mez; Plwebe cilmlllllOIllifo/ia, Nets, P. cubensis, Nees, P. he/eropetala, Mez, P. mexicana,
Meissn., P. montalta, Griseb., P. tnplhurvis, Mez.
II These IpeCles are: fltllota Afitrsii, Gay, B. nitida, Phil.; E1Zdlichen'a LhotSN},i, Mez, E.
sniaa, Nees; Nectalldra (lmp/~fo/ia, M(!z; Oeo/co Qunmliotioro, Mez, O. (tllteata, Urb., O.
dapfmifolia r Mez, O• .f.;rmgillca, Mez, O. jloccijera, Mel, O. /omiculacea, Mez, O. NtllwapJlnr,
Mez, O. rufa, Mez, O. Sodir(J(lna, Mez, O. sjat/m/ala, Mez, O. 'lH!N"uclIlosa, Mel, O. Wrt"ghtii,
Mez; Peru4oo!du/(1/i(l, Mez, P. crassi/o/ia, Mez, P. giahrrilllo, Mez, 1'. Lillg7lc, Nc:es, P • .A/utisii.
H. B. K.; Phoebe cOJlm'iemsis, M~z, P. mexi,,zna, Mcissn.
SOLEREDI-:II 3 x:
 ADDENDA-LAURINEAE

on both sides of the leaf only in H ufelandia rigida. In connexion with the
earlier statements on the hypoderm, which were made on Pax's authority, we
nlay note that this author figures hypoderm (in the "Natiirliche Pflanzt:n~
familien ~) in Aydendron serictum, Griseb., and Cryptocarya Boldus, Mol. Holter ~
mann records hypoderm in Cryptocarya Wightiana, and Dubard and Dop
in Ravensara Perrieri sp. nov. According to Petzold the smaller veins are
vertically transcurrent by means of sclerenchyma also in a very large number
of the American members of the Order. In Persea crassi/olia, Mez, P. M 4tisii,
H.B.K., and PO'rufotomentosa, Nees the sheath of sclerenchyma spreads out
beneath the upper epidermis, thus forming a sclerenchymatous hypoderm of
one or two layers.
To the section dealing with the oU· and mucilage-cells (p. 703 et seq.) the
following details may be added. The oil·cells are present in all the genera
and species hitherto examined; in Laurus nobilis (according to Haberlandt)
and in species of Cinnamomum (accordin~ to R. Muller) these elements show
the same features as we have had occasion to notice in'the Aristolochiacea~,&c.,
that is to say the secretion is enclosed in a pouch, which is suspended by means
of a cuticularized' basin: Petzold does not record epidermal secretory cells
in any of the species examined" by 'him. The mucil9-ge-cells, according to
this author, occur either in the palisade tissue only (this being mostly
the case) or in the spongy tissue only (rarely) or both in the palisade
and spongy tissues. According to him they are present in Acrodielidium
pro parte, A niba pro parte, Bellota, Cryptoearya pro parte, EndUcheria
pro parte, Hufelandia, Litsea pro parte, Misanteea pro parte, Nectandra
pro parte, Oeotea pro parte, Persea pro parte, Phoebe pro parte, Pleuro ..
thyrium, Sassafras, Silvia pro parte, Systemonodaphne and Urbanoden ..
dron, while they are wanting in Acrodiclidium pro parte, Aiouea, Aniba pro
parte, Benzoin, Cryptocarya pro parte, Dicypellium, Endlicheria pro parte,
Litsea pro parte, M isanteca pro parte, N ectandra pro parte, Ocotea pro parte,
Persea pro parte, Phoebe pro parte and Silvia pro parte.
With reference to the mode of deposition of .oxalate of lime we may note
that the small crystals occasionally occur also in the epidermis of the leaf.
Petzold, like the earlier observers, found only unicellular clothing hairs
in the hairy covering. A special type of hair occurs in species of Aniba, Nee-
tandra and Ocotea, the body of the hair above the point of its insertion being
prolonged into a lateral crop-like outgrowth. Mention may also be made of
the apparently septate hairs, found in species of A niba, Endlicheria, Nectandra,
Ocotea, Per;sea and Phoebe; the septation depends on the fact that the lumen
ot the cell forming the hair disappears completely at certain points.
3. STRUCTURE OF THE AXIS. The stem of Cassytha filiformis shows about
the same type of structure as that of C. americana (Schmidt, Bowig, Mirande).
According to Mirande the hypocotyl and the young axis of C. {ili/ormis
exhibit a ring of isolated bundles in transverse section; these bundles are
either fully differentiated vascular bundles or consist of phloem only. The
same author states that in very old axes of this species radially elongated bands
of wood are developed by the activity of cambial arcs, arising on the inner side
of the groups of soft bast.,
The.peculiar f celluies de marteau • described by Mirande in the endodermis
of C. fi/ifoYmiS'. and the lysigenous secretory spaces found in C. americana and
C. {ili/ormis, still require a special consideration; the latter resemble canals, are
situated between the groups of pericyclic bast-fibres, and are filled with mu.cilage.
The • celluies de marteau • arc particuJarly prominent in the young axis; they
bear a blunt tip, which projects on the inner side of the endodermis, and in ccmtrast
to the other endodermal cells they contain no starcR, but have abundant proto-
plasmic contents and a large nucleus. Their contents enable one to recognize
 ADDENDA--LAURINEAE 1043

them also in older stems. In C. fili/ormis the mucilage-spaces above mentioned are
formed as the result of the gelatinization of the walls of four or five rows of flattened
cells with wide lumina (Mirande). The mucilage-cells, which occupy a subepidermal
position in the primary cortex of C. fililormis, in some caseS likewise fuse to form
nlucilage-canals. Mucilage-cells are present in the mesophyll of the scale-like leaves
both in C. fililormis and C. americana. .
A c'omposite and continuous ring of sclerencbyma, including U-shaped
stone-cells, is recorded by Pitard in the pericycle in numerous species belonging
to the genera previously enumerated in this connexion, as. well as in Hu/elandia
pendula. According to Hartwich the pericyclic strengthening ring in Cinna-
momum zeylanicum becomes thrown off in older stages owing to the formation of
internal cork, its place being taken by a new ring of stone-cells, which is
formed for the most part from tissue belonging to the phelloderm. According
to Hartwich a secondary ring of stone-cells of this kind is developed also in
Actinodaphne, Caryodaphne (=> Cryptocarya), Haasia (~. Dehaasia), Mespilo.
daphne, Ochnodaphne, and Tetranthera (= Litsea).
Literature; Decaisne, Stmct. anat. de la Cuscute et du Cassytna, Ann. sc. nat., ser. a, t. v, 1846,
pp. 247, 2~.-POUlsen, Hauatorium "Von Cauytka, etc., Flora, t877. p. 501 et ~.-Hohnel,
Gerberinden, Berlin, 188o, p. 95 et seq.-Ptister, Zimmtrinden, in HUger, elc., Forsch.-Ber. f. Lebens-
mittel, i, 1894. pp. 6 and 25 et seq.-[Bastin, Sassafras, Americ. Joum. of Bot., 1895, p. 3 12 et &eej.]-
Biermann, OIzellen, Diss., Bern, 1898, pp. I3-29.-[Sayre, Cinnamon barks, Drugg. Circular, etc.,
r898, D. 9.1-Hartwich, Cotorinde. Archiv d. Pharm., ccxxxvii, 1899. p .•427.eL seq.-Tscbirch,
Harzbild., Feslschrift f. Schwendener, 1899, p. 464 et seq.-Hartwich, CeyloDzimmt, Viertelja.hrschr.
Datun. Geaellsch. ZUrich, I?Oo, pp. 199-l04.-Tschirch, Harze u. Harzbehlilter, 1900, P.·387 et ICq.
-Hartwich, Zimmt, ArchlV d. Pharm., J9C)I, pp. 181-Z0) and Tab.-Pitard, Pericycl~, Th~,
Bordeaux, 1901, PP. 79. 80.-Siedler, Chines. Bandolinenholz, Ber. deutsch. phann. Gesellsch., 19o1,
p. 30: abstr. in Just, 1901, ii, p. 99.-Bargagli.Petrucci, Legnami, Malpighia, 19O1, p. ::197 et seq.-
Claoditz, Blattanat. canar. Gew., Diss., Bal3el, 1902, pp. 17-a3 (Laurus, Oeolea, Persea, PAoelJe).-
Gerhard, Blattanat. v. Gew. d. Knysnawaldes, Diss., Basel, 1902, pp. 2S-30.-A. Th. Schmidt,
Cassytll4 filiformis, Osterreich. bot. Zeitschr., 1901, pp. 173-7 and Tab. vii.-Tschirch and
Sbiruawa, Kampher, Attbiv d. Pharm" 1901, pp. 257-9.-Achner, Fals<:he Chinarindcn, Diss.,
Bern, 1904, p. 30 et seq.-HOwig. CassyfAa filiformi!, Contribut. Pennsylvania Unlv., Philadelphia,
ii, 3,.1904. PP' 399-416, especiaUy P.: 408 et seq. and pI. 33-3+-[Kamiya, Comp. anat. of the ,.ap.
Laonn., Bot. Magaz., Tokyo, XVlII, 1904, pp. 145-56 Gapanese).]-Are&Choug, Trap. vaxt.
bladbyggn., Sv. Vet. Akad. Handl., 39, n. 2, 1905, pp. II, 12, a.nd 71, 12 (Litsea). pp. 8.(-6 and
Tab. x (AcfbU)(/aplme), pp. 87-9 (Cr.1plocotyo).-Mirande, Caisythacees, Ann. &c. nat., ser.9, t: ii,
J905, pp. J81-z8S.-R. Miiller,Olbehalter, Ber. deutscb. bot. Gesellsch., 190f;' p. 297.-Piccioli,
Legnami, Bull. Siena; 1906, p. I 46.-Fr. Weiss, Bark in the Sassafras, Bot. Gazette, 1906, pp.
434-4-4.-Dubard et Dop, Ravensara Perriert', Bull. Soc, bot, de France, J907, p. IS6.-Guttellberg,
Immergriines Laubblatt· dec Mediterranflora, in Engler, 1I0t- Jahrb., xxxviU, 1907, pp. -4 31 - 3
(Laurw nobilis).-Holtermann, EinflllSS d. Klimas, 1907, pp. 119, llo.-PeUold, Syst. anat. 0

Untersuch. uber die Bl. d. am erik. Laurin., Diss., Erlangen,.·1907, 3-4 pp. j sep: copy !rOOl Engler,
Bot. Jahrb., xxxviii.

HERNANDIACEAE (Pp.707-709)'
A one~layered hypoderm is found in the leaf also in Gyrocarpus Jacquini,
Roxb. (HoHermann). In, SParattanthelium 'Tupiniquinorum the pericycle of
the axis contains isolated bundles of bast-fibres, the intervening pericyclic
parenchyma only being sclerosed at certain points (Pitard).
Literature: Pitard. Diagn. anat. des div. esp. de CyroearjiUs, Actes Soc. Linn. Borde.os, h'i,
'901, p. cvii; and Pericyc1e, The!>e, Bordeaux, 190', p. 78.-Areschoug, Trop. vaxt. bladbyw·,
Sv. Vet. Akad. Handl., 39, n. 2, J905, pp. 33-4 (Henzana'ia).-HoltermlJln, EintlUiS des Klimas,
J9071 p. 17 8.

PROTEACEAE (pp. 7°9-7 1 5).

2. STRUCTURE OF THE LEAF. Tassi's paper, which is cited below, contains
data on the structure of the leaf and axis in species of the genera Banksia,
Dryanara, Franklandia, Crevillea, Giuvina, H akea, I sopogon, Leucadendron,
~X2
 A DDENDA--PR 0 TEA CEA E
Macadamia, Protea, Roupala and Stenocarpus 1• We may first mention that,
according to Tassi, hypoderm occurs also in Banksia grandis, WiUd. (composed
of two layers on both sides of the leaf), Guevina A vellana, Mol. (a single layer
on the upper side), and Stenocarpus sinuatus, Endl. (one or two layers on the
upper side, and sometimes also a single layer on the lower side), and solitary
crystals of oxalate of lime in the pith of the axis also in Guevina Avellana and
Leucadendron argenteum, R. Br. and in the epidermal cells of the leaf in H akea
ltJUrina, R. Br. According to Francken sclerenchymatous fibres, which are
connected wjth the sclerenchyma of the veins, are found in the mesophyU also
in species of Banksia and Dryandra (Banksia dryandroides, Baxt., and B. mar-
cescens, R. Br., Dryandra armata, R. Br.).
For the varied structure of the leaf in the heteropbyllous species of Hakea,
see Carlsson and Paoli, 11. cc.
3. STRUCTURE OF THE AXIS. Stenocarpus sinuatus likewise shows sub-
epidermal development of the cork and groups of bast-fibres in the pericycle.
According to Pitard a composite and continuous ring of sclerenchyma is found
in the pericycle in Banksia ericaefolia, Grevillea brevi/olia, G. buxifolia, G. macro-
stachya, Persoo1tia fcrruginea and Rhopala V ieillardi.
Literature: Hohne!, Gerberinden, Berlin, 1880, p. 97 et seq.-Carlsson, llakea Vicloriae, Bot.
Centralbl., 1886, iii, pp. 77-9.-WiJnaendts Francken, Sclereiden, Diss., Utrecht, 1890, pp. +5-50.
-Houlbert, &i5 sec. des Proteac., Assoc. {ran y• Besan~on, 1893, ii, pablo 1894, p. 544 et seq. (with
a table for the determination of the gene ra).-Kn oblauch , Okolog. Anat., etc., Habilitat.-Schr.,
Tiibingen -896, p. 15 et seq.-Tassi, Le Proteacee, in specie dello Stenocarpus sinuatus, Endl.,
studio aniit.·morphol., Bull. Labor. ed Orto bot. Siena, i, 1898, pp. 67-134, 13 tab., especially
pp. IOl-19 aDd Tab. vii-xii.-Pitard, Nricycle, These, Bordeaux, 190I, p. 6s.-Paoli, Eterotillia
(Hakea S1Iaveolms, R. Br.), Nuovo Giom. bot. !tal, xi, 1904, pp. 204-7 and Tab. ii.-Areschoug,
Trop. vut. bladbyggn., Sv. Vet. Akad. Hand!., 39, n',;2, 1905, pp. 41, 42 (1J./acadamz"a).-Ponch,
Spaltoffnungsapparat, Jena, 1905, pp. 126-30 and Tab. i.

THYMELAEACEAE (Pp.715-721).
I. To the REVIEW OF THE ANATOMICAL FEATURES we may add that
interxyl;.uy phloem occurs side by side with intraxylary phloem also in the
genus Brachythalamus, which is closely related to Gyrinops and Gyrinopsis.
2. STRUCTURE OF THE LEAF. I have examined the structure of the leaf
in Brachythalamus caudatus, Gilg (Beccari, n. 911, New Guinea), and may
mention the following details regarding it. The le<1ves are bifacial in structure;
isolated sclerenchymatous fibres branch off from the bast-fibres accompanying
the vascular bundles of the veins and penetrate into the mes0phyll; the
epidermal cells are not gelatinized; the stomata, which are surrounded by
ordinary epidermal cells, are confined to the lower side of the leaf, where they
occur only in small numbers, and are united to form indistinct groups; the
clothing hairs are unicellular; oxalate of lime, lastly, is excreted in the form of
styloids, and the same is the case in thp; .axis of Brachythalamus podocarpus,
Gilg (Beccari, n. 299, New Guinea).
According to Keissler the folia albo-puncticulata' (described by earlier
I

authors as 'glanduloso-punctata ') recorded by him in the species of Daphne,

belonging to the subsections Oleoides, Gnidittm and Cneorum, as well as in
Passerina grandi/lora, L., are due to the fact that each of the stomata is sur-
rounded by a rosette of papillose epidermal cells having a granular cuticle.
3. STRUCTURE OF THE AXIS. In Brachythalamus podocarpus the xylem-
mass contains: (a) uniseriate meduJtlry rays; (b) vessels, which have rather
1 The plant examined by Tassi under the name of 'Perscollia m)'t'lil/ot'cks, Sieb.' is incorrectlr
determjned. According 10 Tassi it has secretory ctnrjljes in the Je,1i and bicollatertll vascu13r bundles
in the petiole, so that it is a member of the Myrtaccae.
 ADDENDA--THYMELAEACEAE
small lumina, are provided with simple perforations, and bear bordered pits
in contact with parenchyma of the medullary rays; (c) wood-fibres, which have
wide lumina and bear rather small bordered pits (bordered pits also with broadly
elliptical, slit-shaped apertures); and (d) interxylary phloem (the presence of
sieve-tubes has been demonstrated I) . .The interxylary phloem occurs abun-
dantly even in young branches in the form of groups of varying shape; in some
cases it is even found on the outer side of the groups of primary vessels; it
does not include bast-fibres. At this point we may also mention that, contr36Y
to Leisering's recent statements 1, the interxylary soft bast develops at the
to
inner margin of the cambium (i.e. internal the latter) in the Thymelaeaceae
examined in detail by Van Tieghem.
To the section dealing with the structure of the cortex we may add that
in Brachythalamus podocarpus I was able to demonstrate epidermal develop-
ment of the cork and the structure of the bast, which is characteristic of the
Thyme1aeaceae. According to Jen~ic 2 the bast-fibres of the Thymelaeaceae
afford an anatomical feature, which is characteristic of the Order; their shape
namely differs from the usual fusiform shape of these elements, since their ends
are pomted only in very rare cases, being for the most part swollen in a clavatE."
manner, although often truncate; enlargements and constrictions and pecu-
liarly shaped indentations and lobes are of almost regular occurrence, being for
the most part confined to one side of the bast-fibre (for details, see loco cit.).
APPENDIX: OCTOLEPIS AND GONYSTYLUS (p. 721).
In the four new species of Octo/epz's, recently established by GiIg. he likewise
demonstrated mucilage-cells in the pith and cortex, and the absence of intraxylary
phloem.
Literature: rThouvenin~ Bois d'aloes et d'aigle, Journ. de Pharm. et de Chim., 1893. D. I, 2.}-
Knoblauch, Okofog. Anatomie, etc., Habilitat.-Schr., Tiibingen, 1896. p. I I et seq.-l3oergesen,
Exkurs. i Sydspanien, Bot. Tidsskrift, xxi, lS97-8, pp. 1.4.3, 1-44 (Tltymtlaea hirsuta).-Keissler,
Dap"m, in Engler, .Bot. Jahrb., xxv, 1898, pp. 34, 35.-J. Moller, Lignum aloes, Pharmaz. Post,
xxxi, 1898, sep. copy, 28 pp.-Gilg, Oclo/epis, in Engler, Bot. Jahrb., xxviii, 1899, p. 141.-
Leiaering, Interxywe5 Leptom, Diss.• Eerlin, 1899. PP' 13, .....-Petersen. Vedanatomi, 19o1, pp.
~3, 54 (.Daplt.m JVezer~m, L.).-Pitard, Perley-cle, These, Borde~ux, 1901, p. 9 2 .-Bargagli-
Petrucci, Legnami, Malpighi4, 1902, p. 3-24 (Gonyst)'Iu.r).-Clanditz, Blattanat. canar. 'Gew., Diss.,
Buel, 1902, pp. 37-9 (Cnidium).-]ent!i(!, Baatfasem der Thymelaeac., Osten. bot. Zeitschr., 1902,
pp. 151-4 and 2l8-3I.-[Armari, Piante della reg. medit., AnnaIi di Bot., i, 1903, p. 17 et seq.
(Dapltne).]-Col, Faisceaux, Ann. se. nat., sa. 8, t. ~xJ 1904, p. 160, etc.-Siisscnguth, BehaarnDgs--
verb. d. Wiirzb. Mnschelkalkpfl., Diss., Wiirzhurg. 1904, p. 5o.-Areschong, Trop. vaxt. bl.dbyggn.,
Sv. Vet. Akad. Handl., 39, n. 2, 1905, pp. 39-41 (Phaleria).-Frommel, Plantas text. chit, 1905,
p. 35.-Giinther, Anat. d. MyrtiB oren , Diss.. Breslaa, 190,5, p. 23.-Boorsma, Al0t3holz, Bull. du
Depart. de l'Agriculture aux lodes neerl4Dd., n. vii, 1907, pp. 6-19.

PENAEACEAE (Pp.722-724).
The thin~walled cells of the cork mentioned on p. 723 consist of cellulose.
Literatllre: Knoblauch, Oko}og. PBanzenanat., etc., Habilitat.-Schr., Tiibingen, 18¢, p. l3 et
seq.-Giinther, ·Anat. d._Myrtitloren, Diss., Breslan, 1905, pp. :u, :1:1.

ELAEAGNACEAE (pp. 724-726).

Litr-rature: Petersen, Vedanatomi, 1901, pp. 55, 56 (H;j~ph"e rllamnoides, L.).-Pitard,
Pericyc1e, These, Bordeaux, 1901, p. go.-Gunther, Anat. d. Myrtiflor(;D, Dis••, Breslau, 190.5, pp.
23, 24.-Piccioli, Legnami, Bull. Siena, 1906, p. 17 8.

~ Leisering maintains that Van Tieghem's resalts are cited incorrectly on p. i20 of this book, but
that is not the case; in Ann. sc. nat., ser. 7, t. xvii, .1893, it is distinctl, stated that: Ices handes
procedent, com me Ie reste du bois, du bord interne de l'assise g~neratrice.
J Species of the following genera were ex:lmined: Cryplaamia, lJais, .Dnphlu, .Daplwojsii,
DilranDILpis, Dj)·.:a, Edgl'Wllrthia, GniJia, LagettIJ, P(JSsuina, Rhamllolltu Y on, Thyme/a"".,
lVi/tslronnia.
10+6 ADDENDA

LORANTHACEAE (pp.726-730).
I.REVIEW OF THE ANATOMICAL FEATURES. In the Visceae the place of
the cork is taken by a • cuticular epithelium.' Anomalous structure of the
axis in the form of concentric rings of vascular bundles occurs also in Loranthus
sp. (Deistel, n. 77, Cameroon).
2. STRUqTURE OF THE LEAF. Van Tieghem's numerous publications on
thjs Order c~tajn scattered observations regarding the distribution of the
sdereids in Jhe mesophyU, a feature which he considers to be of primary im-
portance, further on the distribution of the groups of silicified cells, the storage
trach~ids, the transversely placed stomata, &c.; we may pass over these data
here as well as the detailed statements on the structure of the axis, since we
may expect to find them suml1larized in Van Tieghem's monograph; the
«ritation of the numerous new generic names in connexion with the anatomical
features does not, moreover, appear suited to the purposes of this book. Hypo-
derm (composed of ODe or two layers) has recently been recorded by Holtermann
in the leaf of Loranthus cap:tellatus; it had not hitherto been observed in this
Order.
3. STRUCTURE OF THE AXIS. In the Visceae (Viscum, Arceuthobium,
Dendrophthora, Phoradendron, N otothixos) no cork ,is formed; according to
Damm {for details, see lac. cit.) the epidermis at first keeps pace with the growth
in thickness by stretching and division of its cells, but subsequently a' cuticular
epithelium' appears in place of the cork. This epithelium comprises cells
belonging to the epidermis and primary cortex in which the outer walls have
become thickened by the formation of cuticular layers.
I have casually observed the case of anomalous structure of the uis
referred to above in material of Loranthus, belonging to the Munich herbarium
(cited above). The branch, which is I! em. thick,-has three Or four rings of
growth. The groups of soft bast belonging to the individual rings of bundles
are separated from one another by sclerosed medullary rays, so that in a trans-
Verse section they appear like islands, which show a concentric arrangement.
The secondary vascular bundles are developed in the pericyclic parenchyma
internal to a zone marked by the occurrence of groups of bast-fibres.
Literature: Van Tieghem, NumeroU$ papers in Bull. Soc. bot. de France, 1895-6.-Boergesen
og Paulsen, Veg. paa de dansk. vestind. I:>er, Bot. Tidsskrift, xxii, 1898-9, pp. 107-9 (Lltralllkus
t".ar~nalus, Sw.).-Leisering, InterxyHires Leptom, Diss., Berlin, 1899. p. 13.-[Cannon. Anatomy
of PIzorndendron villosum, Bull. Torrey Bot. Club., xxviii, 19°1, p. 37-4.]-Petersen, Ve<ianatorni,
19o1, pp. ';1, il (Viscum a/bum, L.).-Thiselton-Dyer, Haustorium of Loranthus aphyllus, Annals
of Bot .• 1901, pp. 7-49-57 and pl. xl.-Van Tieghem, Rhizantheme, Joum. de Hot., xv, 190J, pp.
366, 361·-Da lllm, Bau mehrj. Perid., Beih. z. bot. Centralbl., ~i, 1902. pp. 219-60 and Tab. i-iv.-
Van Tieghem, Beccarine, Jouro. de Bot., xvi, 19°::1, pp. 1-5, especially p. ::I.-Reiche, Phrygilantnus .
tI./JII".lItis. Flora, 19(>-4, pp. 271-97 and Tab. v.-Piccioli, Legnami, :Bull. Siena, 1906. p. 175.-
Roltermann, Einfluss d. Klimas, 1907. pp. 72 and I ....o.-[Reiche. PkrygiianIAus·Arten, I"lora.
xc;vii, 1907, pp. 375--401 and Tab. xiii, xiv.]-[Van Tiegbem. Inovulees, Ann. se. oat., ser. 9. t.
vi. 1907.]
SANTALACEAE (pp. 730-737).
The rows of tracheides, mentioned as occurring in the leaf on p. 732. are
found also in Osyris alba, L. (Guttenberg).
APPENDIX: I. MYZODENDRON (p. 733).
The specie~ having medullary vascular bundles form the subsections A.-chi·
I>hyllum and Telophyllum. which are regarded as genera by Van Tieghem.

3· GRUBBIA (p. 737).

We may add that the periderm in Grubbia rosmarini/olia. Berg arises in the
eplderm~. and is composed of tabular cells ,with thin walls, and that in G. stficta.
 ADDENDA--SANTALACEAE
.
DC., which belongs to the section Ophira, stone~cells occur in the primary cortex
and, numerous bast~fibres in the secondary bast (Van Tieghem).
Literature; Van Tieghetn, Myzodeodracees, Bull. Soc. bot. de France, 189!h p. 558.-Van
Tieghem, Grubbiacees, Joam. de .Bot., 1~97, pp. J:Z7-38.-Volkens, Ostafrik. Sandel holz ,
N oti:::bl. Berliner bot. Garten, etc., n. 9, 1897, pp. 2 i ~ - 5.-[ K usano, Haustori 11m of Buc,flep:.
quat/ria!a, Bot. Magaz., Tokyo, xiv, 1900 (Japanese).J-Pitard, P~ricyc1e, 'These, Bordeaux, 1901,
p. 9s.-Kusano, Pl!ra~itism of Bucklo/a 'luadriala, Joum. College of Sc. Imp. Univ. of Tokyo, xvii,
Art. la, 1902, 46 pp., I Tab.-[llarber, Ham:toria of Sandal roots, Indian Forester, xxxi, 1905,
pp. 189-301, pI.; abstr. in Just, 1905, ii, p. 53.]-Fl'aysse, Sur;oirs de !'Osyrt".r alba, Comptes rcoclas
Paris, exl, 19°5, pp. :170, 271; and Parasitisme de l'Osyris, lac. cit., pp. 318, 319.-[Barber, Han-
storh.m of Santalum alhum, Mem. Departm. Agr. Ind., Botany, '906, 30 pp.]-[Pb.zonl, Austorl
dell' Osyris alba, Annali di Bot., iv, 1906, pp. 79-98 and Tab. iiill.]-Gultenberg, Laubbl. d.
Mediterrantlora, in Engler, Bot. Jahrb., xxxviii, 1907, p. 419 (OS)'''';s alba, L.).

BALANOPHOREAE (pp. 738, 739).

We may add that stomata have recently been observed on the scale-leaves,
bracts and certain parts of the flower in Cynomorium coccineum, the ordinary
pairs of guard-cells being accompanied by twin-stomata and stomata, in which
one or both of the guard-cells are transversely divided into two cells (Pirotta
and Longo).
Literature: Pirotta e Longo, Stomi ael CY1l011tQriu",, Rendinc. Accad. dei Lincei, viii, 1899,
pp. 98-1oo.-Baccannl e Cannarelta, CynomDrium r(lccineum, Rendinc. Accad. dei Lincei, viii,
J899, pp. 317-20 and [AUI Accad. Gioenia sc. nat. Cattania, ser. 4, xii, l899, 60 pp., 3 Tab.].-
POrich, Spaltoifnungsapparat,
, . Jena, . pp. 69, 70.-[For fnrther literature,
.1905. . see p. 1169.]

EUPHORBIACEAE (Pp.739-763).
1. The REVIEW OF THE ANATOMICAL FEATURES requires the following
addi tions : .
The cork may occasionally develop 'in the pericyclc. .
Oxalate of lime: Styloids occur also in Claoxylon (incl. J.l1icyococca) and
Erythrococca; some of the idioblasts containing clustered. crystals ~.rel of sub-
epidermal origin, but push their way between the epidermal cells.
Internal secretory system: Elongated secretory sacs (V), like those of
Maliottts, are found also in species of Amanoa and Uapaca, while sac-shaped
secretory cells (VI), which are situated in the epidermis of the leaf and resemble
those of Ricinus, have been observed also in species of Bischogia and J.,fareya.
Trichomes: Branched multicellular clothing hairs occur in specie~ of
Phyllanthus, and peltate hairs a1so in Crotonogyne.
_Anomalous ,structural feat~res of .the axis: Lepidotur'lls (Acalypheae)
likewise has interxylary phloem. Cortical vascular bundles are found in
certain E uphorbias.
The following anatomical features still deserve notice: Cork-cells, tIle
walls of wnich~aie'partly encrusted with small crystals of oxalate of lime (in
species of Croton); fibrous cells in the primary cortex or pith, as the case may
be (in species of Angostylis, Conceveiba~ Euphorb1:a, and Hyaenanche); occur-
rence of palisade tissue or of large lacu::l.ae in the primary cortex (species of
Euphorbia); strong development of the primary cortical parenchyma owing to
the activity of a secondary m~ristem (in the Cactus-like Euphorbias) ; stomata
placed transversely to the longitudinal axis of the stem (in certain Euphorbias).
2 .. STRUCTURE OF THE LEAF 1. \Ve may add the following details to the

1 Among the papers recently pllblishcd on the systematic anatomy of this Order we may, as a
suppiement to the footnote on p. 74[. mention Gauch;:r's investigations. The most important
contribution is that published in Ann. \ie. na.t., str. 8, t. :tv. Unfortunately, however, the literatuft',
 ADDENDA-EUPHORBIACEAE

section dealing wi th the structure of the epidermis (p. 741). The lower epidernlis
of Euphorbia pubescens has jagged lateral walls with projections in the angles,
just as in the epidermis of many petals. Gelatinization of the epidermis of the
leaf is found also in species of Baccaurea (see Rothdauscher and Areschoug).
A thick covering of wax, consisting of sman rods, is found on the leaves in
Euphorbia piscatoria, Ait., the layer of wax being interrupted above the stomata
by gaps, the size of which corresponds to that of the stomata. Incrustations
of wax, which show the same structure, and may be as much as 70 fA, thick,
also cover the epidermis of the axis in the Cactus-like .uphorbias; they are
perforated above the stomata, around which they are occasionally (E. Tirucalli)
elevated to some height in the form of a broad ring of wax. Papillose develop-
ment of the epidermis of the leaf has been observed also in Breynia disticha,
Mill!. Arg. and B. rubra, !lilli. Arg. (Phyllantheae, on the lower side), as well as
in Euphorbia Broteri, Daveau (on both sides), E. dendroides, E. /lavicoma,
E. iacquiniae/lo1'a, Ait., E. ltfyrsinites (on both sides), E. nicaeensis and E.
piscatoria, Ait.; a many-layered hypoderm 'occurs on the upper side of the
cricoid leaves of Mictanthemum ericoides, Desf., M. hexandrum, Hook. f. and
Stachystemon vermicuiaris, Planeh. (Caletieae); a one·layered hypoderm, often
composed of large cells, is present on the lower side of the leaf in the xerophilous
species of Euphorbia (e.g. E. Peptis, L., E. Preslii, E. aegyptiaca). In the
species of Euphorbia belonging to the subgenus A nisophyllum the stomata
arc exceptionally small. Their development in Euphorbia conforms either to
the Ranunculaceous or Rubiaceolls types; the latter type occurs quite generally
among the stomata on the succulent stems of a group of species of Euplwlbia,
which are indigenous in Madagascar, and have been examined by Costantin
and Gallaud. Subsidiary cells, placed parallel to the pore, are present also in
Leptonema venosum, Juss. and Thecacoyis gytnnogyne, Pax (Phyllantheae).
The pairs of guard-cells are arranged transversely to the longitudinal axis of
the succulent stems in Euphorbia Alluaudi, Drake, E. leucadend,on, Drake, and
E. oncoclada, Drake, whilst in the related species the pores of the stomata lie
parallel to the longitudinal axis (Costantin and Gallaud).
With reference to the structure or the leaf (p. 743) we may first mention
that rolled leaves with a furrow to the·right and left of the median· vein occur
also in certain species of Cluytia, e.g. C. poli/oUa, Jacq., C. pterogona, Miill. Arg.,
C. pubescens, Sond. and C. Rustii, Knauf (Knauf). Sclerenchymatous fibres
running freely in the mesophyll are recorded by Gaucher also in species of
Glochidion (Phyllanthus), the names of which are not given; branched scleren- C

chymatous cells traversing the mesophyll in a reticulate manner' are described

by Koorders in Chondrostylis bancana, Boer!.; sclerenchymatous cells resem·
bling rod-cells by Areschoug in the palisade tissue of Excoecaria Agallocha,

apart from Pax'$ paper (in Engler's Jahrb •• v), is not t.ken into consideration by Gaucher, so that it
will be necessary to reinvestigate tnose statements, which are contradictory to the earlier ones,
especially those referring to the occurrence of receptacles for water-storage (loc. cit., pp. :I 17, 218;
lome of these Ilfe probably gela.tiniud epidenntll cells) and the duplication of the epidermis (loc: cit.,
pp• .:119, 220, ta. well as p. 295. gelatinized epidermal cells 1) in the leaf, and statements dealing with
the secretory organs and the intraxylary soft bast. Gaucher also determined the anatomical features
in a large num~r of genera, 'Which had not previoasly been examined, and has thus advanced ODf know-
ledge of the anatomy of the Enphorbiacue. Thesr genera are as follows (arranged under the tribes
distingtlished by Muller.Arg.): i. Caletieae : M;~,.aniJzeum, Stac}l)'slttntm. iii. Ampereae: lI/tJUotIJJ(I"s.
h'. Pbyllantheae: Agyntia, Bunuayia (not enumerated in Miiller·Arg.'s system), Chorio}'hyllulII
(not enumerated in MUller.Arg.'s syst~m), C~"oD"e, Hyaenaluhe (Tox;c~tiron), LtftottetJlo.
Maesobotrya (not en\lmerated in Muller-Arg. s system), Pseudolacltnoslylis, Revercltonia, Siballgea
(not enumerated in Miiller.Arg.'s system), Thecator;s, Uaj{ua. v. Bridelie:te: eleis/an/hus. vii.
Acalypheae: AkluJrneopsis, Angost)'lt"s, Ct'o/01togyne, Erythroco"a. Hasskarlia, Lasio(l'~to".
Leidesia. Ltlidoluros, lUaml;ophY/(J11. Mareya, AJischodMt, j'leohutonia, Platygynt, Py(ll/)(o»za.
,·iii. Hippom~e: Bmnelia (Ga/earia) , Chef/Dsa, Ompllalea. x. Euphorbieae: Altlho-~/wU1.
FeJi/antlms, S),nadmitmt.
 ADDENDA-EUPHORBIACEAE 1 0 49

Miill. Arg. ; and enlarged terminal tracheids by Gaucher in A manoa oblongifolia,

lIiill. Arg. and certain species of Euphorbia (e.g. E. Eroleri, E. Afyrsillites and
E. splendens). In Discocarpus (e.g. D. Spruceanus, Miili. Arg.) and Briduia
micrantha, Mill!. Arg. Gaucher describes and figures sc1erenchymatous plates,
which traverse the mesophyll in the vertical direction and, according to him
(loc. cit., pp. 210, 211), do not include any vascular elements; in all probability,
however, they are merely veins, which are vertically transcurrent by means of
sc1erenchyma.
According to GaU(:her, the sheath of large parenchymatous cells with wide
lumina, previously recorded in certain species of Euphorbia (in the section dealing
with the structure of the veins, p. 744), is a feature characteristic of those
species of the genus which belong to the subgenus A nisophyllum. Similar
sheaths of wide parenchymatous cells containing tannin accompany the scleren·
chyma of the veins in species of Amanoa, Discocarpus and Pseudolachnostylis.
There is a complete absence of sc1erenchymatous tissue accompanying the
veins in the Euphorbias.
According to Gaucher, oxalate of lime (cf. pp. 744-6) is founa in the form
of styloids (in cells which traverse the entire thickness of the mesophyll) also in
Claoxylon (C. affine, Zollo and other species), as well as in Micrococca Mercurialis.
Benth. (Claoxylon Mercurialis, Thw.; this 'record is not in agreement with
Rittershausen's earlier statement) and Erythrococca (E. aculeata, Benth. and
E. capen sis, Milli. Arg.). The results of Knoll's work have shown the necessity
for a reinvestigation of the course of development of the crystal-cells in the
case,of the earlier statements on the ·occurrence of clustered crystals in the
epidermis, or at least in the case of some of them; the object of the reinvestiga-
tion being to determine whether,these elements really belong to the integumental
tissue or whether we are concerned with cells (containing clustered crystals) of
the subepidennallayer, which push their way in between the epidermal cells or
even extend beyond them; the second alternative applies. to Dalechampia
Roezliana, Milll. Arg. (probably also to other Acalypheae) as well as to the
hairs containing clustered crystals found in Plukenetia and Fragariopsis (see
below). I .
Cells, which contain clustered crystals and are either actually or apparently
situated in the epidermis, are recorded by Gaucher also in species of Securinega
(Fliiggea, Phyllantheae), Bridelia (where there are small groups of cells containing
oxalate~ of lime), Crotonogyne and Manniophyton (Acalypheae), and species of
Jatropha (not mentioned in Herbert's paper; Hippomaneae).
According to Gaucher, the crystalline bodies, found in Euphorbia splendem
(see p. 746) and regarded by Pax as crystals of abietinic acid, consist of oxalate
of lime, which is absent in other species of Euphorbia. For the.sphaerites of
calcium phosphate and calcium malophospbate occurring in the Cactus·like
Euphorbias, see also Gaucher; and regarding the colouring-matter (which
produces~ b.!ue t~!»)n dried~plants of ltfercuri~lis, see Fructus, loco cit. T~e
former are found alSo in certaIn Euphorbias, whIch do not have a cactoid habit
(e.g. E. attopurpurM or·E. Lathyris), as well as in Pedilanthus carinatus. The
strongly refractive (silicified ?) bodies; observed by Knauf On the walls of the
epidermal cells in the isothennous species of Cluytia, as well as in other species
of this genus, still require closer investigation.(cf. Herbert, Diss., p. 24).
To the section dealing with the secretory elements of the Euphorbiaceae
(p. 746 et seq.) the following details may be added : -
I. Laticiferous cells. As' regards the distribution of these .elements we
may note that, contrary to Herbert, Gaucher records them in Cluytia; he
likewise mentions their occurrence in Omphalea, which. belongs to the same
Tribe (Hippomaneae), and the genera A nthostema, Pedilanthus and Synademium
(Euphorbieae). The laticiferous tubes have been observed to penetrate into
 ADDENDA-EUPHORBIACEAE

the mesophyll also in J«locroton. In Euphorbia and Macaranga they. often have
very wide lumina and are surrounded by a sheath of small cells containing
starch, so that they appear like secretory canals. The { reticulate anastomoses •
described by Maytl3 in the mesophyll of Euphorbia Lathyris are, no doubt, the
result of incorrect observation 1. MoEsch records rod.shaped starch.grains in
the contents of the laticiferous cel~ also in Hippomane bigland«losa, AubL and
Pedilanthus tithymaloides, Poit., crystalloids, not only in Jatropha Curcas, but
also in J. glauca, Hort .• J. gossyPii/ulia, L. and J. podagrica, Hook., peculiar
spherical elaioplasts in Homalantlms populneus, Pax, and an accumulation of
magnesium in the latex of Euphorbia mammillaris.
IV. The laticiferous or tanniniferous elements comprised in this section
are likewise considered in GallChel"s work. He distinguishes between rows
of secretory cells of equal length (Type' Acalypha » and rows of secretory cells
of unequal length (Type' ,11acarallga digyna, MUll. Arg.' with rather long and
rather short celis, and Type 'Alehurnea cordata, Mull. Arg.' with very long and
very short cells); the division-waHs between the cells mayor may not be
re~orbed. Renewed investigations both on a developmental and on a systematic
anatomical basis are required to enable us once and for all to obtain a clear
conception of the nature and systematic value of the~e secretory organs.
V. According to Gaucher, the wide·lumened secretory saCs of Mallotus
(M. rieinoides, Miill. Arg.). which were referred to under Section V, arise from
a cylindrical complex of irregularly arranged, cells containing secretion; the
same mode of origin is ascri bed to the tanniniferous sacs of a similar type·
which are found in Amanoa I fa'L'Qllica, Miq.' (Zollinger, n. 1662, Java) and
Uapaca IIeudelotii, Baill. i
VI. The tubular secr-etory cells present in the epidermis of the leaf 'in
Ricinus comm1.tttis, Mull. Arg. may be associated with (a) similar epidermal
secretory cells which occur jn Bisclwlfia (according to my own observation,
Hippomaneae), although in this case they are provided with undulated lateral
walls like the remaining epidermal cells, and (b) tanniniferous elements with
wide lumina occurring in the lowf't- epidennis of Mareya brevis, Pax (according
to Gaucher, Acalypheae). In thb connexion we may also note that a few
enlarged tanniniferous cells occasiollully occur in the palisade and spongy tissues
(e.g. in the palisade tissue of Alldraclme cordi/olia, MillL Arg.), that a layer of
tanniniferous cells is situated beneath the typical palisade layer in CrotonogY'l'le
Zenkeri, Pax, and a simllar one in the spongy tissue in Caperonia cordata.
St. Hil.
VII. Intercellular secretory receptacles like those previously recorded
have been found in the following additional cases: in the primary cortex of
Ctuytia hirsutll, ::\1iilI. Arg. (according to Pax); in the primary cortex of Cluytia
alaternoides, Mull. Arg. (d .• however. p. 752) and C. putchella, MUll. Arg. (ac-
cording to Gaucher); and in a subepidermal position in the branches and scale·
leaves of lIura crepitalls, L. (according to Gilles).
To the subsequent paragraphs, \vhkh deal with the hairy covering (p. 752
et seq.) the following data regardjng the clothing hairs (see under I) may first
be added. Little branched clothing hairs occur in species of Phyllanthus, the
names of which are not given. Peltate hairs are found also in Croronogyne, and
sclerenchymatous columns, connecting the trichomes on the two sides of the
leaf, also in Jttlaeroton mOlltevidmsis, Klotzsch. 2
II. Glandular hairs_ The extrafloral nectaries (see p. 755) have recently
-been examined in detail in Excoecaria (by Poulsen), Hevea (by Daguillon and
tOne certaillly does me(t with H·~hapeu con:lexions, which, may be explained as due to
branching, but there are no typicl11 reticulate anastom05es.
3 The plant examined by Areschoug (1905) and described as Bdddia lomentosa, Bl. with a
query, bears unicellular t\\'u-arrued trichomes :lnd probably does not belong to the genus Britkli(J.
 A DDESDA-EUPHORBIACEA£ {OSI

Coupin and also by Parkin) and M acaranga (by Smith)". They resemble the
glands of Ricinus, &c. and the glands occurring on the cyathium (see Gaucher)
in having a secretory palisade epidermis. In Excoecaria biglandtdosa, Milll. Arg.
this epidermis clothes a slit-shaped canal in the peg-shaped nectary, which lies
at the base of the lamina. Both in "Hevca brasilicnsz's, Milll. Arg. and in H.
Spruceana, Milll. Arg. the nectaries aPe situated at the apex of the petioles of
the foliage-leaves and are accompanied by bud-scales, some of which are
entirely transformed into nectaries; the upper epidermis of these bud-scales
is more or less completely modified to form a secretory p~lisade tissue, which
for the most part consists of a single layer Qf cells, but at certain points is
composed of two or three layers. Smith mentions the occurrence of multi~
cellular glandular structures. which are stated to be of the nature of food-bodies,
on the stipules of the myrmecophjlous species M acaranga t"iloba. J

III. The earlier statements regarding the stinging hairs (discussed under
Section III,"pp. 756, 757), which are found in the genera Cnesmone, Leptorhachis,
and Tragia of the Acalypheae, as well as in Dalechampia, require a considerabl~
amount of correction as a result of Knoll's work. This author in the first plas:e
examined the structure and course of development of the stinging hairs, fomi.d
in Dalechampia Roezliana, Mull. Arg. var. rosea; the hairs of this plant corre-
spond to those shown in Fig. 180,.Q on p. 748, although the latter have a more
complicated structure. The investigation has brought to light the fact that the
; outer part of the pedestal in these, hairs consists of 3-5 epidermal cells, which are
elevated above the surface to a considerable extent and envelop an elongated
central cell, which, as its mode of development shows, does not belong to the
epidennis, but arises subepidermally; the pointed end of the central cell, which
contains the crystal of oxalate of lime, projects far beyond the limits of the
pedestal. The' middle cell and' terminal cell' previously mentioned are
J

thus but a single cell, which, moreover, belongs to the subepidennallayer and
not to the epidermis. Similarly, the unicellular stinging hairs, figured for
Tragia cissoides in Fig. 180 t P t are subepidermal cells, which have pushed their
way in between the epidermal cells and thereupon project far b3yond the level
of the epidermis like t~e unicellular (epidermal) hairs; such cases necessitate
a corresponding extension of the definitidn of what is briefly styled a chair.'
Since the lower end of the crystal t which is suspended in the upper part of the
central cell, is provided with a pair of teeth, we are justified in regarding it as a
reduced clustered crystal in which only one'" of the projecting ends of the individual
crystals is strongly developed. Moreover. inasmuch as the crystal-cells (with clus-
tered crystals), which are situated in the epidermis of the leaf of Dalechamp'z"a Roez-
liana, belong to the su bepidermallayer (see above, p. 1049 }-a fact which has likewise
been determined by Knoll-it is more than probable that the crystal-cells (containing
clustered crystals) supposed to haye been demonstrated in the epidermis of other
members of the Acalypheae (and perhaps even of other Euphorbiaceae in which
they occur) are likewise of subepldermal origin; this applies especially: (a) to
the cells containing clustered crystals found in Caperon£a and Argyrothamnia, the
cells ili question projecting somewhat after the manner of a hair; (b) to the cells
containing sphaerites found in Acalypha and Claoxylon, which likewise project
beyond the surface; and (c) to the hairs with clustered crystals occurring in
Fragariopsis and Plukenet1°a (see Fig. 180, S). The cells with clustered crystals
found in Caperonia and A rgyrothamnt'a. and the hairs with clustered crystals ccn-
stitute transitional forms (or, if one prefers. developmental forms) to the typical
stinging hairs of Dalechampia, &c.
In accordance with the preceding statements I agree with Knoll also in
the changed interpretation of the structure of the crystal-hairs (containing
clustered crystals), which. are found in Fragariopsis and Plukenetia. The
'epidern:t.al cell, which is differentiated as a papilla or short hair, and contains a
clustered crystal, the spines of which are all directed outwards' and the' narrow
1052 ~DDENDA--EUPHORBIACEAE

cell, which penetrates 'to a considerable depth in the mesophyll ' (see the previous
description, p. (57), form but a single cell, which is identIcal with the central
cell of the stingmg hair. The division-wall, shown in Fig. 180, S, belongs to
the cellulose-sheath enveloping the clustered crystal.
For the structure of the stipular spines of Euphorbia splendens and E.lactea,
see Mittmann and Barber, 11. cc.; for the prickly structures found on the axes
of Hura crepitans and caused by the prick of an insect, see Didrichsen, loco cit.
The petiole in this Order bas hitherto Peen investigated only in' a very
inadequate manner, especially in view of the large size of the Order. New
details regarding its structure will be found in Gaucher's work (see Ann. sC. nat.,
1902). According to this author, the petiole in the species of Euphorbia
belonging to the subgenus A nisophyUum contains a single vascular bundle,
while in the remaining species of the genus there are three or four· isolated
bundles.
3. STRUCTURE OF THE AxIS. To the list of Acalypheae, which possess
intraxylary llhloem (see p. 757), we may, on Gaucher's authority, add Le-pido-
tu,us (L. 1l1ni/olius, Baili. and L.laxiflorus, Benth.) and Mallotus subulatus, MOO.
Arg.l
CorticaJ vascular bundles are recorded by Kniep in Euphorbia Cyparissias
and E. orientalis.
The following details may be added to.the earlier account of the develop-
ment of the cork. In Euphorbia antiquorum and E. piscatoria' the cork-
cambium arises in the inner layer of the two-layered epidermis; in Phyllanthus
Welwitschianus, Milll. Arg. it develops in the second or third layer, in Mischodon
zeylanicus, Thw. in the third or fourth layer, and in Baccaurea racemosa,.Milll.
Arg. on the inner side of the sclerenchymatqus fibres of the pericycle. In some
cases the place of origin of the cork varies within certain limits in different
parts of the branch of· one and the same species (for details, see Gaucher).
According to Gaucher, the cork in Crown gratissimus, Burch. contains crystals
of oxalate of lime, while in Ditaxis jasciculata, Juss. it includes fibrous cells with
thick walls. Records of the occurrence of crystals of oxalate of lime in the
cork will be found also in the pharmacognostic works of A. Meyer and others,
the data referring to Cortex Cascarillae, which is derived from Croton Eltt-
tena, Benn.; the descriptions, however, are inexact. According to my own
investigation the cork-cells of the Cascarilla-bark, even in the young branches,
have strongly thickened outer tangential walls, wrole the inner tangential walls
and the radial walls are relatively thin; the inner tangential walls and the
adjoining parts of, the radial walls are encrusted with small rhombohedral. or
variously shaped crystals of oxalate of lime, and after the solution of the latter
by means of hydrochloric acid, those parts of the wall, which were previously
encrusted, show a network in relief corresponding to the insertion of the crystals.
I have found an exactly similar type of incrustation, which affects the same parts
of the cell-wall, in the cork-cells (which here have relatively thin walls and wide
lumina) of the Copalche-bark, which is obtained from Croton niveus, Jacq. The
distribution of this highly peculiar type of cork within the genus Croton is worthy
of a more detailed investigation; it-makes its presence noticeable even in the
external aspect of the plant owing to the white colour of the cork. A feature
not hitherto recorded is the occurrence of fibrous cells in the primary cortex and
pith. According to Gaucher they are found in the primary cortex in Hyaenanche
(Toxicodendron) globosa, Lamb. et Vahl (scattered elements with thick walls),

I The specie& of Euphorbia, included by Gaucher (in Ann. &c. nat .. 1902, loco eit., p. 196) in the
'Type Euphorbia.' and 'Type Trag-ia Okanyua, Pax,' are stated by him to have wellkly developed
strand, of internal soft bast; in all these ca~, however, we are probably concerned with uDlignified
eltmentl belonging to the innermost parts of the xy1elTl.
 A DDENDA-EUPHORBIACEAE 1053

Conceveiba guianensis, Aubl. (slightly thickened and arranged in groups),

Angostytis long'ifotia, Benth. (slightly thickened and forming a peripheral zone),
and Euphorbia xylophylloides, which has the habit of a Cactus (isolated elements,
here also in the pith); according to Costantin and Gallaud (see also Fron),
fibrous cells occur also in a group of succulent species of Euphorbia found in
Madagascar (elements with thick stratified walls; confined to the primarycortex
in E. Geayi, Cost. et Gall., E. Intis)" Drake, E. Laro, Drake, E. rhipsaloides, Lem.
and E. Tirucalli, L.; present both in the pith and primary cortex in E. alcicornis,
Bak., E. Alluaudi, Drake, E. arsoides, Cost. et Gall., E. Decorsei, Drake, E.
enterophora, Drake, E.le'Ucadendron, Drake, E. oncoclada, Drake, and E. skmo-
dada, Bail!.). In the leafless species (e.g. Pedilanthus aphyllus, Boiss., Calyco-
peplus ephedroides, P1anch. and species of E14phorbia) the primary cortex con-
tains palisade tissue. In species frequenting a marshy habitat (e.g. Euphorbia
palustris) large lacunae are found both in the primary cortex and in the pith.
In the Cactus·like Euphorbias the primary cortex is particularly strongly deve-
loped, the c;ortical tissue in these forms undergoing continual increase by means
of a peripheral meristematic zone. The pericycle is developed in the -form of
a composite and continuous ring of sclerenchyma, including stone-cells with
U·shaped thickening, also in Sa via erythroxyloides, Griseb.

DAPHNIPHYLLACEAE (p. 760).

The lower epidermis of the leaf includes groups of small cells can taining clustered
crystals of oxalate of lime also in Daphniphyllum bancanum, Kurz and D. laurinum,
Baill.
BUXACEAE (pp. 761-763).
To the second paragraph, summarizing the features which are uniform through-.
out the group, we may add the following details. The cork develops in the pericycle
in later stages also in BUXU5, the superficial formation of the cork in this case being
merely a local phenomenon. Secretory cells are found also in Buxus (Eubuxus).
The anomalous structure of the vascular system~ previously described in Simmondsia
(which Van Tieghem regards as being related to the Tetragonieae). occurs not only
in the stem, but also in the root. Cortical vascular bundles are found in the old.
world species of Buxus (Eubuxus, Baill.) and- in Notobuxus natalensis, Oliv., but
not in the American species of the genus (Tricera, Sw.• as a genus).
The structure of the leaf is bifacial also in SlyloCCYQs. It is a remarkable fact.
that the lower half of the mesophyll of the more or less distinctly bifacialleaves has
a dense structure in all cases, except in the species of Buxus belonging to the section
Eubuxus. Typical spongy tissue with large lacunae, which directly adjoin ~he
assimilating tissue (this latter being either of the nature of palisade or conSisting
of rounded cells) and are the cause of a splitting of the leaf into two halves, is found
in-the species of Eubuxus (e.g. in Buxus sempe1'Virens, L.); the two halves of the
leaf are in ~onnex.ion with one another only at the margin of the lamina; the
process of splitting takes place in the living leaf at a time when it has not yet
attained its full size, and is not merely a result of the drying of the leaf. The earlier
statement"t6"the effect-that oxalate of lime is wanting in the leaf of Buxus sem-
pervirens is incorrect; clustered crystals occur in the mesophyll and ordinary
solitary crystals accompany the vascular bundles of the veins, and in addition .to th~t
one finds a peculiar kind of crystal-sand~ composed of relatively large pnsf:!l3:tlc
crystals. which occasionally have a corroded appearance. The cells contalDmg
this crystal-sand give rise to the small white dots, which ate visible even to the
naked eye along the median vein on the lower side of the leaf. Secretory' cells
are present also in the mesophyll of Buxus semperoirens, and. Van 'Fleghem has
like~se obser~ed th.em in the m~sophyll ~nd primary cortex of B. balwYica. \ViUd.
and In the pnmary cortex of StmmOndsla. The secretory cells -of Buxus semper-
viyens show up prominently in the palisade tissue owing to their large size and in
the spongy tissue owing to a reduction in the development of the arms; the,ir
bright contents. which are strongly refractive (in the living leaf), are insoluble In
water and alcohol 'and assume a deep brown colour on treatment with a solution
 AUDENDA-EUPHORBIACEAE

Qf Iodine. Clothing hairs are found also in Huxus; ill B. scmpervirens they are
uniseriate and composed of two or more cells with thick walls, while in B. acuminata.
Miill. Arg. and B. Hildebrand#z,', Bail1., for instance, thevare unlce-lIular. Boergesen
and Paulsen's. statement as to the occurrcnce of deeply sunk external glands on
the leaf of B. Va hiii , Baill. (' Tricera laevigata, Sw.') requires closer investigation.
In Pachysandra, Sarcoc()cca and Styloccras, the American species of Buxus
(Tricera), Notobuxus natalensis, Olivo and the African spccies of Buxus (B. Hilde·
brandtii, Baill., B. Mac-Owani, Oliv., B. madagascartca, Baill., B. pediceltatus,
Van. Tiegh.) the petiole contains three vascular bundles, viz. a large median
bundle and two smaller lateral ones; in Notobu.rus and in the American 'and Afriqm
species of Buxus these lateral bundles form the margina1.vcin of the leaf. On the
other hand, in the European and Asiatic species of Buxus belonging to the section
Eubuxus (which in acc()rdance with Vall Tieghcl11's investigations is better restricted
to include only the species just mentioned, the above-named African species ,of
Eubuxus being best Classed with Notobuxus) the petiole contains only a sir;gle
vascular bundle, which is accompanied 011 either side by a strand of fibres. The
petiole at Simmondsia contains an arc-::Shapcd vascular bundle, which soon becomes
closed to t'orm a ring.
l'he disb1buhon and general course of the cortical vascular bundles of
j

Buxut; ha~ been determined by Van, Tieghern's recent investigations. Cortical

bundles are found in all the European, A$iatic and African species of Buxus, as well as'
in N :Jtobuxus natalensis, but do not occur in the American species of Buxus (Tt'£cera) •
as itl the case of B. sempervirens the bundles arc four in number and show normal
orielltation of their wood- and bast-groups. In the EU1-opean and Asiatic species,
of Buxus the bundles are supported by a strand of bast~fibrcs situated on their
outer side (i.e. adjoining the bast); whereas groups of mechanical fibres have been
observed on the inner side of ,the bundles (i.e. next to the wood) only in occasional
species (e.g. in B. balearica, WiUd. and B. faponica, Mull. Arg.). In Notobuxus
natalensis and the African species of Btu;us, o.n the other hand, the xylem is invari·
ably surrounded by a group of mechanical fibres, but there arc no fibres adjoining
the bast. The cortical strands branch off from the two bundles, which pass out from
the vascular ring at the node to supply the two opposite leaves; these branches run in
the following internode, and ultimately end blindly. In the European and Asiatic
species of Buxus the groups of fibres situated on the outer side of the cortical
vascular bundles are continued into the petiole. where they appear at the sides of
the single vascular bundle (see above). On the other hand, in Notobuxus and the
African species of Buxus this is not the case; the three vascular bundles present
in the petiole of these species are formed by the branching of the single bundle.
which bends out into the leaf from the vascular ring of the axis; the three vascular
bundles in the petiole of the species of Buxus, belonging to the section TJ'2-cera,
arise in the same way.
The primary cortex contains cells showing U~shaped thickening also in Buxus
acuminata, while slightly sclerosed cells occur quite generally in this tissue in the
African species of Buxus and jn Notobuxtls. 511Ch sclerosed cells arc found in certain
African species of BUXk$ also in the pith, bast and pericycle. The pericycle includes
isolated groups of bast~fibres in Styloceras,. and a few bast-fibres in Buxus longifolia,
Mull. Arg.; in Bu.xlls the pericycJe is generally parenchymatous. Oxalate of lime
occurs in the cortex of Buxus also in the form of clustered crystals.
Literature: Karsten, Zellenkrystallc irn -l\Ii)Cbsa.ft von Ia/njh-a C1m:a.r, extracted from
Poggendorf Ann., 1859, in Karsten, Ges. Abh., i, 186,5, p. 305 et seq.-Uhlworm, Entwicklungs-
gesch. d. I Iaare, Bot. Zeit., 1873, p. 805.-Hohnel, Gerberinden, Berlin, 1880, p. 118 et seq.-
Costantin, Tiges aero et sout., Ann. sc. nat., ser. 6, t. xvi. 188~, p. 107 et seg.-Leitgeb. in Mitten.
bot. Inst. Grat, 1. Heft, 1888 (not 1881, as was previously stated on p. ,62), p. 31+ et seq.-
Mittmann, Anat. d_ Pflanzenstachebl, Verh. bot. Vcr. Mark Brandenburg, 1889, p. 56.-Hansell,
Calciumphosphatausscheid., Flora, 1889, p. 408 ct seq_-Aufrecht, Extraf1o r. Nektarien, Diss.,
Zii rich , 1891, p. 6 et seq. (RicilltlS).-Barber, Corky excresc., Ann. of Bot., 1892, pp. 160 and 166.-
Fmctns, Mercllriales, These, Montpellier, 1894, pp. 29-43.-13randt, \Venig bek. Riuden, Diss.,
Dorpat, 1894, p. 48 et seq.-Groom, Extralloral nectaries of Almrilu, Ann. of 13ot., 1894, pp. :u8-
30.-[Tognini, Stomi, Atti lst. bot_ Pavia, 1894. ]-Didrichsen, 'fornene hos nflra ,repitant, Bot.
Tidsskrift, xix, 1894-5, FP. 189-~oo.-Cocconi, Nettarii estral1Uz. delle Ridmts, MeD'lorie Accad.
Bologna. ser. 5, vol. v, 1895, pp. 423-31, 1 Tab.~Chauvealld, Caract. :mat. des Euphorbia Peplus,
etc_. Jouro. de Bot., 1897, p. 35+-Gnucbcr, Une espece du genre Euphorbe, Joum. de Bot., 1897t
pp_ :u 4-18.- [~elstein, H evea brasilt'ens is, ~l oscow, 1897, Russian.]-Pou Isen, N ektarie~, Vidensk.
Meddelels. natllrh. t·Ol'('U. 1<j0benhavn, 1897. pp. 356-60 and T:tb. i, ii.-Van Tieghem, Elongation
 ADDENDA--EUPHORBIACEAE 1055

de. nc.euds..Ann. sc. nat., ser. 8, t. v, 1897, pp. 159, 160; and, liuxacees, loco cit., pp. 289-338.-
Biermann, Olzelleo, Diss., Bern, 1898, pp. S7, s8.-Gaucher, Et. anat. du genre EupllOrbia, TheSe,
Montpellier, 1898,138 pp.-[Sayre, Stillingia:root, Druggist's Circular, etc., 1898; abstr. in Just, 1898,
ii, p. 49.]-Van Tieghem, Sim1IllJTUisia, Journ. de Bot., 1898, pp. 103-Il.-Boergesen og Paulsen,
Veget. paade dansk.vestind. Oer, Bot. Tidsskiift, xxii, 1898-9, pp. 19, 91 and9J (species of Hippcmane, .
Croton, and Tricera; see also Revue gen. de Bot., 1900).-Gaucber, Racine des Euphorbes cactiformes,
Joum. de Bot., 1899, pp. 173-5; and Glandes du cyathium des Euphorbes, etc.) Ioc. cit., pp. 368-7°.-
Grelot, Caoutchoucs et Gutta-Percha, These, Paris, 1899. p. a5 et seq.-HaHier, Kautscbuklianen,
Jahrb. Hambllrg. naturwiss. Anst., xvii, 1899, pp. 196-8.-Leisering, Interxylires Leptom,Diss., Berlin,
1899, p. 45.-Fron, Euphorbia Intis)" Joum. de Bot., 1900, pp. IS7-63.-Gamper, Angosturarinden,
Diss., Zurich, 1900, pp. 6[, 62.-Gaucher, Role des laticiferes, Ann. sc. nat., ser. 8, t. xii, 1900,
PP.241-60.-Keamy,inContribut. U.S.Nat. Herb., v,n.5, Ig00,p. 296(Croto1Ullaritimus).-Tunmann.
Sekretdriisen, Diss., Bern, 1900, pp. 27-9.-[Hartwich, Rhabarber aus Guatemala (Jatropha podagrica,
Hook.), Schweizer. Wocbenscbr. f. Chemie 11. Phann., 1901, p. 579; abstr. in Just, 1901, ii,
p. 4S·)-Molisch, Milchsaft u. Schleimsaft. Jena, 1901, pp. 4, 12. 17. 3l, 34, 47, 4' and 54·-
Petersen, Vedanatomi, 1"901, PP' 46, 47 (BlIxus).-Pitard, Pericyde, These, Bordeaux, 1901, p. 7:1·-
Areschong, Blattanat. d. MangrovepR., Bibl. bot., Heft S6, 1902, pp. 6 ..-6 and Tab. vii (Ex(oecaria
Agallocna).-Clauditz, Blattanat. canar. Gew., Diss., Basel, 1902, pp. q.-16 (Euphorbia) '.-Gaucher,
Rech. anat. sur les Enphorbiacees, Ann. sc. nat., ser. 8, t. XV, 1902, pp. 161-309 -Knotbe,
Unbenetzbare Blatter, Diss., Heidelberg, 1902, pp. Il-I3.-Penzig, Piante acarofile, Malpigbia, 190'.
p. 439 (Rolt/era).-[Al'mari, Piante della reg. medit., AnnaH di Bot., i. 1903, p. 17 el.leq.]-[Bray.
Plants of the Sotol region, Bull. Torrey Bot. Club, xxx, 1903, p. 611 et seq.]-Daguillon et Coupin,
l'Jectaires extrafloraux des Hroea, Comptes rend us Acad. Paris. cxxxvii, 1903, pp. 767-9.-rGilIes,
Et. morpho et anat. du Sablier (Hura (rep#ans), Ann. lnst. col. Marseille, 1903, pp. 43-1l"O.]_;
Knauf, Geogr. Verbreit. d. Gatt. Cluytia. Diss., 13reslau, 1903, 54 pp.-[Smith, Jlacara1Jga triloba,
a new myrmecophilous plant, New Phytologist, ii, 190~, p. 79, pI. v, vi j abstr. in Bot. CelltN11hl.,
Jtciii, p. 182.]-Acbner, Falscbe Chinarinden, Diss., Rem. 1!J04, p. 58 et seq.-Chrysler, Strand-
plants, Bot. Gazette, xxxvii, 1904, p. 461 et seq. (Euphorbia).-Col, Faisceaux, Ann. SC. nat., ser. 8,
t. xx, J904, pp. 162-7 (C,-ptt?JJ).-Koorders, Ch.JlMroStylis, Ann. ,Jardin Bui!enzorg, xix, 1904-,
PP.48, 49.-Daguillon et Coupin, Glandes pet. d'IIevta bl'asilimsis,' Revue gen. de Bot., xvi, 1904,
pp.8O-90.-Dubard et Viguier, Euphorbia It,tisy) Comptes rendus, Paris, ex xxix, 190 4, pp. 307-9.-
Gilles, Hura crepitallS, These, Montpellier, 1904, go pp.-Paoli, Eterofillia, Nuovo Giorn. Bot.
ltal., xi, 1904, pp. 207-10 and Tab. i.-Parkin, Extrafloral nectaries of Hevea brasiliensis, Ann. of
Bot., 19°4, pp. 217-26.---[Smol,ik, Vielkernige Zellen bei einigen Euphorb., Bull. intern at. Acad.
Sc. Boheme, 1904.]-Solereder, Frostflecken des Buchsblattes, Centralbl. fUr Bakteriologie,
Parasitenkunde u. Infektionskrankh., ii. Abt., xii, 1904, PP' 257, 2S8.-Siissenguth, Behaarungsverh.
d. Wiirzb. Muschelkalkpfl., Diss., Wlit;zburg, 1904, p. 50.-Areschoug, Trap. V3.xt. bladbyggn., Sv.
Vet. Akad. Handl., 39, n. 2, 1905, pp. 2:1-5 and Tab. xvii, xYiii (Trelo/ra, A/writes), pp. 27-9 and
Tab. xxv (Bride/ia, Agynda), p. 34 (Baccaurea), pp. 47::-9 and Tab. xxv (Dalechampia; Acalypha),
p. 52 (Sapium), pp. 77-9 (A Ichornea) , pp. 80-2 (ftIiscllOdOIZ), pp. 9 2, 93 U_atropka) , pp. 95-7
(Trewia), PP' 97-101 (Cyc!ostClIW", Atchonua), pp. 146, 147 and Tab. xi, xii (CroloIJ), pp. 147, 148
and Tab. xxii (DaPhni"phyllum), pp. 151, 152 (Codiattmt).-Bois et Gallaud, Modif. anat., Comptel
rendus Paris, I I Dec. 1905 (Euphorbia).-Costantin et Gall:md, Nouveau groupe du genre
Euphorbia, etc., Ann. sc. nat., ser. 9, t. ii, 1905, pp; 287-312.-Dubard et Viguier, S)st. radie. de
I' EupMrbia Intisy, Revue gen. de Bot., 1905, pp. 260-71.-Kniep, Bedeutung des Milchsaftes,
Flora, 1905, pp. t63-6.-Knoll, Brennhaare cler Euphorb.• Gatt. Dalechampia u. Trag'ia, Sitz.·Ber.
Wiener Akad., cxiv, Abt. i, 1905, pp. 39-48 and Tab. i, ii.-Mayus, Milchr., Beih. z. bot.
Centralbl., xviii, Abt. i, 19°5. pp. 2j8, 279.-~lontemartini, Biologia del Buxus se1llpervirens, Atti
Ist. bot. Pavia, ser. 2, vol. x, 1905, 6 pp. and Tab. xxviii (deals with the same features as Solereder,
1904).-Van Tieghem, Meristeles cort., Ann. sc. nat., ser. 9, t. i, 1905, p. 37.-Piccioli, l..egnami,
Bull. Siena, I!)06, pp. 139 and 155.-Boorsma, Aloeholz, etc., Bull. Depart. Agricult. aux lndes
neerland., n. vii, 1907, p. 19 et seq. (Excl1t'caria).-Holtermann, Einfluss des Klimas, 1907, p. lIO.
..._Solereder, lnkrustation cler Korkzellenwande mit Kalkoxalatkrystallen bei Cortex Cascarillae,
Archiv d. Pharm., ccxlvrl907, pp. 409-1o.~[F()r further literature, see p. 11,0.]

BALANOPSEAE (pp. 763, 76-J.).

A composite and continuous ring of sclerenchyma is found in the pericycle
also in Balanops oliviformis, and has been observed ·in an undetermined species
eveD in .branches of 55 mm. thickness 2.

I The plant described by Clauditz (loc. cit., pp. 40, 41) as ~lIer:t1P'ialis ambigua (which bas
raphides, brown secretory cells, and stomata with parallel subsidiary cells) is incorrectly determined
and does not belong to the Euphorbia.ceae. Judging by the anatomical chatacters it may possibly
·be a member of the Rubiaceae.
~ Pitard, Pericycle, These, Bordeaux, 190 1 ) p. 77.
 ADDENDA

URTICACEAE (pp. 764-779).

To the brief REVIEW OF THE ANATOMICAL FEATURES of the Urticaceaetaken
as a whole (p. 764), we may add that laticiferous cells have been recorded also
in the second genus (Cannabis) of the Cannabineae, that elements resembling
laticiferous tubes occur also in the genus Urera (Urticeae) and that raphides and
styloids have recently been observed in the genus Laportea (Urticeae).

I. ULMACEAE (pp. 764-768).

A to'mposite and continuous ring of sclerenchyma is developed m the
pericyc1e also in Sponia rugosa (= Trema rugosa) (Pitard).

2. CANNABINEAE (pp.;t?9. ;;0).

Laticiferous cells are stated to occur also in Cannabis. Tschirch at least
regards the secretory elements~ which are present in the secondary cortex'as
well aVn the soft bast of the veins of the leaf, as laticiferous cells; they are
tubular elements with rather wide lumina, which in the drug are occupied by
brownish contents. According to Molisch, the latex of the hop is rich in tannin;
it also contains colourless spherical .)r biconvex grains, which recall the protein-
grains found in Cecropia and Brosimum. '
,
3· MORACEAE (pp. 770-775).
We may add the following details to the REVIEW OF THE ANATOMICAL
FEATURES. Laticiferous cells occur in the leaves in all the Artocarpeae, being
found especially in the veins and sometimes even in the mesophylJ itself, whilst
in the Conocephaleae the laticiferous cells are almost confined to the
axis and only rarely penetrate into the lamina of tne leaf. The glandular
hairs found in the Artocarpeae and Conocephaleae vary in structure (see below).
With the glandular hairs we may associate large wax·glands~ which are provided
with a palisade-epithelium, and occur in many species of Ficus; the mucilage:
canaJs of the Conocephaleae are accompanied by rows of mucilage-cells, which
are found in the veins of the leaf. Tanniniferous idioblasts have been observed,
also in genera other than those previously mentioned in this relation. Oxalate
of lime occasionally occurs also in the form of small crystals of varied shape.
To the previous enumeration of the special features presented by the structure
of the leaf (e.g. occurrence of bypoderm and clustered crystals in the epidermis~
which have now likewise been shown to have a wider distribution) we may add:
the occurrence of division ..wa11s in the epidermis; papillose differentiation of
the epidermis; gelatinization of the inner waJIs of epidermal cells; stomatal
groups and stomata situated in pits (Ficus); hydathodes provided with an
epithema; arm-palisade tissue (Parartocarpus); spicular fibres (Balanostreblus,
Ficus, Sahagunia); occurrence of solitary crystals in the epidennis.
A detailed investigation of the STRUCTURE OF THE LEAF (see p. 770) had
not till recently been carried out, but has since been undertaken by Renner 1 in

1 Renner's investigations deal with the following genera.: I. Artocarpeae: Ft'cus, SparatIIJs)'u,
iJammaropsis, Brosimttm, Lanessania, BIJsfjzuia, Sc)'plzosyu, Antiart's, Olmtdia, Pseudolmtdia.
Castilloa, HeliuJslylis, Pereoea. Heliattlhostylis, Clldrania, Poulsenia, Trtculia, PararlocarjJUs
(incl. GymnariocarjJus), Ar/()carpus (incl. Prailua), Brosimopsis, Sahagunio, Ratallos/reMus,
StJrocea; 2. Conocepbaleae: C(cropia, 1I1yrialltJlUs, MIISaltga. Cottssapoa. COIl()C(plzaittl (incl~
Fa/ansaephylum), Fourtmma.
 ADDENDA-UR TICA CEAE l057
the tribes Artocarpeae and ConocephaJeae. The following pages are, in the
main, a summary of the results of his work. The upper epidermis in many
cases consists of a single layer of cells, but it is frequently 2- or 3..layered at
certain points or almost throughout. Only a small number of epidermal cells
divided by tangential walls are found in Antiaris saccidoTa, Dalz., A. toxicaria,
Lesch., Castilloa elastica, Cerv., and Myrianthus arboreus, P.B.; the upper
epidermis is two-layered in Cecropia obtusifolia, Bertol., C. scabra, Mart., and
in numerous species of Ficus, while it is 2-3-layered in Dammaropsis Kingiana,
Warb. The same feature has been observed in the lower epidermis, although
it is of rare occurrence (Conocephalus tonkinensis, Renn" Ficus longi/olia, Schott
and F. gibbosa, Bl.). Regarding the distribution of the hypoderm we may men-
tion the following details on Renner's authority: it occurs on the uppe'£" side of
the leaf in Cecropia and Sahagunia (here one-layered), Artocarpus, Balano-
streblus and PouroumfJ (one- or two-layered), Coussapoa and Musanga(2-layered),
Conocephalus (2-3-layered) and Ficus (I-4-layered); in Ficus it is present also
on the lower side of the leaf, where it consists .of two or three layers. The
species in which hypoderm has been recorded are enumerated below 1 (ex,
cepting the numerous species of Ficus; see also Holtermann, loco cit.). The
hypoderm is almost invariably parenchymatous, but apart from that, its cells
vary in size, in the nature of their lateral walls, and in the general structure of
the walls. A ftbro;.Is hypoderm, which consists of four layers and probably
belongs to the ground tissue, is found beneath the upper epidermis in Artocarpus
l(lnceae/olius, Roxb., this feature going hand in hand with the vertical trans-
currence of the weaker lateral veins by means of sclerenchyma; transitions
to this type of structure occur also in other species of Arl(tcarpus. The develop-
ment of the hypoderm --from the dermatogen has hitherto been demonstrated
only in certain species of Ficus. Other noteworthy features in the structure
of the epidermis are as follows: the septation of epidermal cells by thin
vertical walls (species of Artocarpus, BroSf,'mopsis and Poutsent'a); cuticular
elevations in the form of delicate parallel striae (species of Fic~es and Pararlo-
ca"pus)~ or of coarse crests, showing an irregular arrangement (Ficus c,ocata,
Mart. and allied species), these two characters occurring especially on the
lower side of the leaf; and the presence of one or two peculiar thickenings
shaped like a biconvex lens on the outer walls in Ficus paralnsis, Miq., these
walls perhaps serving for the perception of light. Papillae are not very com-
monly formed on the lower side of the leaf. Variously differentiated papillae
are found in all the species of Brosimum examined by Renner, and also in Ficus
foveolata, Wall., F. pumila, L., Helicostylis Poeppigiana, Tree., and Myrianthus
arboTeus, P.B. Gelatinization of the integumental tissue haS not been demon-
s~ated m any species of Ficus, but cells with mucilaginous inner membranes
occur in the upper epidermis in species of A ntia'l'is, A rtocarpus (in A. V rieseanus.
Miq. in the lower epidermis as well), Bosqueia, Brosimopsis (here the outer wall
is also mucilaginous), Brosimum, Cecropia, Helicostylis, Olmedia, Perebea,
Pseudolmedia, and in the hypoderm on the upper side of the leaf in species of
Balatwstreblus, Cecropia, Conocephalus, Coussapoa and Mu~anga. In Artocarpus
dasyphyltus, Miq. and other species of Arlocarpus belonging to the section Pseu-
tto;aca certain of the upper epidermal cells have strongly thickened inner walls,
which only swell up to a slight extent in water and assume a yellow colour on

I Artocarpus Blumei, Trec., A. Ktmando, Miq., A. lYfaingayi, King, A. Tamaran, Becc.;

Bala1Jl1strelJlus ilidfolia, Knrz; Cecropia adenopus, Mart., C. Humholdtlana, Klotzsch, C. lati/(}"",
Miq., c. kucocoma, Miq.• C. mexicana, Hems!., C. pellata, L., C. sdadophylta, Mart.; Cont)uphalw
ianceDiatuJ, Trec., C.luaveolens, Bl., C. tonllinensis, Renn. i Coussapoa intermedia, Mart., C. nitida,
Miq., C. Scholtii, Mi.q., C. suOincana, Mart .• C. villosa, Poepp. et Endl.; Musanga Smithii,
R. Br. i Pourouma acuminata, Mart., P. bicolor, Mart., P. cecrtJpiaejolia, Mart., P. cinerascerts,
Miq., P.fu/;'g.·nea, Miq., P. lomentosa, Mart., P. velufina, Mart.; Sah!r~mia Pecko/lii, K. Sch.
SOL£R£DER 3y
 ADDENDA-URTICACEAE
treatment with Iodine-solution and Sulphuric acid. Silicification of the walls
of the epidermal cells, especially of the outer walls, is very common.
The stomata in the Artocarpeae and Conocephaleae are found almost
exclusively (exception: Cecropia Humboldtiana, Klotzsch) on the lower side of
the leaf. In some cases (species of Cecropia, Coussapoa, Pourouma) they are
placed on pedestals, formed by the neighbouring cells, while in Brosimopsis
lactescens, Sp.-Moore and certain species of Ficus they are sunk below the
surface. Distinct subsidiary cells, conforming to the Cruciferous type, are
present only in Conocephalus. Apart from that, a rosette of subsidiary cells,
distinguished by having thinner walls, is found in Ficus Binnendijkii, Miq. and'
F. perlusa, L. f. Owing to the projecting network formed by the veins on the
lower side of the leaf, and a corresponding differentiation of the lower epidermal
cells, the stomata with their neighbouring cells are frequently (espec~ally in
many species of Ficus) found to be arranged in groups in the meshes between
the veins; the meshes themselves are often depressed to form small pits.
Typical stomatal groups, however, occur only in Ficus gibbosa, Bl. In this
connexion we may also notice the stomatal pits found in certain species of Ficus,
and visible to the naked eye as spots on the surface of the leaf; they have been
described in detail by Bargagli-Petrucci. In F. callicarpa, Miq. they fonn
small widely open pits, the aperture of which is occupied by hairs; in F. punc-
tata, Thunb. and F. lalcata, Thunb., in which the orifice of the pits likewise
bears hairs, the pits are provided with three recesses, the middle one of which
runs at right angles, while the lateral ones are parallel to the surface of the leaf;
in F. excavata, King the pits are irregularly lobed, and the lobes open directly
to the exterior, no hairs being present. Th~ relatively large stomata, situated
above the veins of the leaf in species of Ficus and in Brosimum Alicast,um, are
probably of the nature of water-pores. Hydathodes, which are provided with
an epithema and are often recognizable with the naked eye as spots or small pits
or warts (mentioned in the earlier part of this book, p. 771), besides occurring
in many species of Cecropia, C01UJcephalus, and Ficus, are found also in Dam-
maropsis Kingiana, Warb. Regarding the structure of these hydathodes, we
may add that the number of water.pores, which are invariably of very small
size, may be as much as 100 in Ficus, a1though generally less, and is only 10--15
in Cecropia. According to Renner, the occurrence of these bydathodes cannot
be recommended for use as a specific character, as they may be present or
absent in one and the same species.
In most cases the structure of the leaf is bifacial, but in numerous species
of Ficus belonging to the section Uros!-,'$ma the lowest layer of the mesophyll is
differentiated as a low palisade, while 111 the genera Antiaris, Brosimum, Ficus
Sect. Urostigma, and Pourouma the entire mesophyll is sometimes found to
consist of palisade-tissue, the lower layers being formed by short conjugate
palisade-cells. Arm-palisade cells have been observed in the first or second
layers of the palisade-tissue or in both these layers in certain species of Pararlo-
carpus. A hypha·like differentiation of the spongy tissue is specially charac-
tenstic of the species of Artocarpus belonging to the section Jaca and of certain
species of Ficus belonging to the section Urostigma. In Pa,artocarpus excelsus,
Beec., the two lowest layers of the spongy tissue have thick pitted walls and
are differentiated like a hypoderm. Spicular fibres, which have a varied course,
are recorded by Renner in the following species: Balanostreblus ilicifolia, Kurz
(only in the veins); Ficus Bonplandiana, Miq., F. c,ocata, Mart., F. dolia,ia,
Mart., F. Gardneriana, Miq., F. longifolia, Schott, F. obscu,a, Bl., F. pisi/era,
WaIl., F. tomentella, Miq.; Sahagunia Peckoltii, K. Sch. The great diversity
presented by the structure of the veins in the Artocarpeae and Conocephaleae
cannot be consideled here; regarding this point, see Renner, loco cit.
For the structure of the leaf in the heterophyllous species, d. Paoli, loco cit.
Renner demonstrated laticiferous tubes (see p. 77I) in the leaves of all the
 ADDENDA-URT1CACEAE

Artocarpeae which he examined. The laticiferous elements present in the

leaves are thin-walled sacs, which are from 8-30 p. wide; they are principally
found traversing the peripheral portion of the tissue accompanying the veins
(sometimes also the medullary tissue of the veins) and in species of many
genera 1 also enter into the mesophyll. They belong to the category of latici-
ferous cells; Mayus's statement as to the occurrence of reticulate anastomoses
in Ficus elastica is doubtless incorrect. According to Renner the latex of the
Artocarpeae is peculiar in containing small spherical masses of caoutchouc
side by side with tannin. The same author showed that the contents of the
laticiferous elements have. a very distinctive character in Ficus Populi/olia, Vahl,
inasmuch as they include albuminous substances. Molisch has recently proved
that the latex in Ficus elastica and in the famous cow-tree (B,osimum Galacto-
dendron, Don) is rich in salts of magnesium, which crystallize out from a drop
of the latex in the form of round or angular sphaerites. In the Conocephaleae
]atkiferous cells have been found in the axis of all the genera examined by
Renner, as wen as in Musanga (according to Engler). They occur chiefly in the
primary cortex, although occasionally present in the phloem as well, but only
rarely (Pourouma acuminata, Mart. and P. 'mollis') enter the leaves (lateral
veins). Their diameter varies between 25 and 85 IA. Their contents, as far as
they are known, never include caoutchouc-bodies, but sometimes contain simple
or composite proteid-bodies, which owe their origin to leucoplasts. These
proteid-bodies were first discovered by Molisch in Cecropia peUata, L. and
Coussapoa Sc/wttii, (' Brosimum microcarputn') and are recorded by Renner
also in Cecropia concolof, Willd" C. obtusa, Tree. and Pou,ouma. tomentosa~ Mart.
Secretory cells, which have brownish contents of the nature of a gum-resin
and which were described for certain species of Artocarpus in the earlier part of
this book, are ra1her widely distributed in the sections Jaca and Prainea of the
genus 2 ; they appear as inflated spherical or ellipsoidal cells of the spongy tissue,
which in these species·has a hypha-like differentiation, and in Artocarpus incisa,
L. f. at least have a suberized wall. In the course of the recent investigations
mucilage-canals have been observed among the Conocephaleae in the axis of
species of the genera Cecropia, ConocepAalus, Coussapoa, ¥usanga and Myrian-
thus, but not in PouroufWl; they occur in the primary cortex and sometimes in
the pith as well. In the leaf, the mucilage-canals of the axis are replaced by
rows of mucilage-cells, which are found in-the lateral veins of the first order in
numbers varying between two and twelve, but only very rarely traverse the
smaller veins as well; such rows of mucilage-cells have been demonstrated in
Cecr,opia, Conocephalus, Coussapoa, Musanga and Myrianthus. In some of the
species of the genus Pourouma there are mucilage-cells in the lateral veins of
the first order. Among the Artocarpeae mucilage-cells are found only in
Olmedia angusti/olia, Poepp.; they are either isolated or united to form ~oups,
and occur in the tissue accompanying the veins. Tannin~idioblasts, Sltuated
in the palis~de-tissue_and differentiated like those of Ficus l'ubiginosa, have been
demonstrated by Renner in species of Antiaris, Arlocarpus, B,osimum, COmJ-
uphalus, Dammaropsis and Ficus 3.

1 viz.: Ar/ocorjJus § Prainea and PuudlJjaca, Bosqueia, Brluil1lo}sis, lJrosi1llUHI, Cudnmia,

Ficus, Hdianlnoslylis, Olllltdia, Parar/pcay/us, P~rebea. Snhagunia, Scyphosyce, Sorocea, SjJarallo-
syee, Treculi"a.
. 2 The species in which they have been observed are as follows: A. anisopltyllus, Miq., A . .dlumti.
Tree., A. CAap/a.rna, Ro:xb., A. eommum's, Forst., A. /rultSUns, Renn., .A. hirsttlu.;, Lam.,
A. A"tmando, Miq., A. lanaae/o/iuJ, Roxb., A. Lim/alo, Miq., A. ftlaingayi, King, A. lIlaruJlum;s,
Tree., A.nooilz's, Thw., A.papuanus, Renn., A. n'gidus, Bl., A. scandells, Renn.,A. Tamara", Ecce.
3 The species are: A,z/ian's sa(cidora~ Dah., A.loximyia, Lesch. j Arlocar;us integrijo/ius, L.f.,
..4. Limpato, Miq., A. Po/yphtma, Pers.; IJrosimllm Aub/~/ii, Poepp. et Endl., B. disclllor, Schott,
B. uhin«arpum, Poepp. tol EndL; CCftQ{lpkalus /altaolatus, Tree., C. sua~(J!ens, Bl., C. tonkitUmis,
R~nn.; Da""narD/sis Killgz'antz, \\'arb. For the rather numerous species of Ficus, which ~long to
the sections Urostigma, ~)notcia, Sycidium, l!.:myce, and Syrotnor1lS, Henner's piper mu!>t be con,ulted.
1. y 2
 1060 ADDENDA-UR TICA CEAE
In most of the Artocarpeae and Conocephaleae clustered crystals constitute
the chief form of excretion of oxalate of lime, although solitary crystals are
also found. In addition to this small, spherical, panduriform Of variously
shaped crystals or crystalline aggregates of the same salt are occasionally
(species of Artocarpus, Cecropia, Ficus, Pararrocarpus) present in the epidermis
of the leaf or in the hypoderm. Oxalate of lime is rarely wanting in the leaf
(e.g. in Helianthostylis Sprucei, Balli. and Lanessania turbinata, Bail!.). Large
clustered crystals, situated in spherical cells of the palisade-tissue, are not
uncommon; the largest idioblasts of this kind are found in Pourouma, where
they occur in the neighbourhood of the vertically transcurrent veins. A more.
important systematic feature is afforded by the occurrence of clustered or
ordinary solitary crystals in the epidermis of the leaf. The clustered crystals
are contained singly in epidennal cells, which are either isolated or u~ited to
fonn groups and occur mainly in the neighbourhood of the veins; in some cases
the cells containing the clustered crystals have a round outline and appear as
idioblasts (e.g. in certain species of Ficus). They have been observed in species
of the following genera: A ntiaris) A rtocarpus (§ Jaca, Prainea and Pseudojaca).
Brosimopsis, Brosimum, CastjUoa, Cecropia, Cudrania, Dammaropsis, Ficus
(§ Urostigma, Palaeomorphe, Sycidium, Coveilia, Eusyce), M usanga, Olmedia.
Perebea, Po urouma , Treculia. Solitary crystals ate found in the -epidermis
of the leaf in species of Balanostreblus, Brosimum, Ficus (§ Synoecia and Uro-
stigma), and Sorocea. In Ficus aurantiaca, Griff., F. callicarpa, Miq. and
F. punctata, Thunb., which belong to the section Synoecia, the ce1ls containing
the crystals combine to form groups; these cells are very small and have
strongly thickened inner walls to which the solitary crystals, which fill the
entire lumen of the cell, are attached. In Brosimum Alicastrum, Sw. the veins
are, as it were, paved with the solitary crystals contained in the epidermal cells.
In the gelatinized epidermal cells of certain species of Conocephalus Renner
demonstrated sphaerocrystalline masses of an unknown chemical substance.
We may add the following details regarding the occurrence of calcification
and silicification of the cell-wall in the Artocarpeae and Conocephaleae.
Silicification of the epidermis is no uncommon feature. In the spongy tissue
of the older leaves of some species of Ficus the walls of the cells bordering
on the respiratory cavities are silicified. Silicified pegs, arising from the
outer walls of the epidermal cells ·and penetrating into the cavity of the cell,
have been recorded in species of Antiaris and Ficus, as well as in Poulsenia
aculeata~ Eggers~ while tuberous protuberances which are silicified are found
in the angles of adjacent cells of the hypoderm in Ficus diversi/olia, Bl. ; lastly.
cells filled with siliceous matter are present in the parenchyma accompanying
the larger veins in Ficus aurantiaca, Griff., F. brevicuspis, Miq., F. clavata, Wall.,
F. obscura, Bl., F. pisifera~ Wall., F. sikkimensis, Miq., Parartocarpus Riedeli'i.
Warb. and SParattosyce dioica, Bur. Regarding silicified trichomes, see
below.
Among the Artocarpeae and Conocephaleae Renner observed typical cysto-
liths (i.e. structures provided with a stalk and a variously.encrusted body,
which shows concentric stratification and in some cases radial striation as
well) only in the genera Conocephalus, Dammaropsis, Ficus, Poulsenia and
Sparattosyce. The cystoliths belong eXclusively to the integumental tissue
and are found both in the upper and lower epidermis; in most cases they
are situated in idioblasts, but frequently occur also in ordinary epidermal
cells, as well as in hypodermal cells or trichomes. In Ficus the lithocysls
are invariably concerned in the formation of the surface of the leaf, the on1y
exception being F. elastica, in which they be]ong to the hypoderm. Only
a small portion of the wall of the lithocyst participates in the formation of
the leaf-surface,. and this.part of the wall is occasionally provided with a small
tip (species of Dammaropsis. F1·cus and P01t/senia) or with a solid attenuated
 ADDENDA-UR TICA CEAL 1061

hair-like peg of greater length (Sparattosyce). The shape of the cystoliths is

particularly striking in the species of Conocephalus, since some of them are
two-armed, the arms either spreading out horizontally or diverging obliquely
with reference to the axis of the cystolith. In some of the species of Ft'-cus
the flattened cells containing the cystoliths are accompanied by cells which
have a similar shape, but are devoid of cystoliths; the latter are related to
the small silicified hairs of Poulsenia, &c. discussed below. Among the struc-
tures resembling cystoliths we may include the silicified pegs mentioned above
as occurring in the epidermis, and the silicified cystotyles, which are· recorded
by Renner as being suspended from the solid tips of the hairs in A ntiaris,
Artocarpus papuanus, Renn. and Parartocarpus Riedelii, Warb. In Cecropia
scabri/olia, A. Richt., lastly, A. Richter figures trichomes, which are com-
pletely filled with silicified matter, showing stratification.
The clothing hairs in the Artocarpeae and Conocephaleae generally
consist of a single cell, and either have a distinct body or appear as short
papillose hairs. Now and then a thin transverse wall occurs, so that the hairs
become bicellular (Cudrania, Myrianthus); in other cases several transverse
(Parartocarpus) or obliquely longitudinal (Ficus aurantiaca, Griff.) walls appear
in the body of the hair. Uniseriate trichomes with several short basal cells
and a long terminal cell, distinguished by the fact that it breaks off at a pre·
determined point, which is situated at the base of the cell, are developed in
Fic'Us pilosa, Reinw. var. chrysocoma, King; ordinary uniseriate hairs, which
have a brown colour, are rich in tannin and consist of twelve or more cens,
are found on the petiolar cushions in Cecropia.
The ordinary unicellular clothing hairs frequently serve in a marked manner
as a protection against excessive transpiration, e.g. in species of Cecropia, Coussapoa,
Ficus. Musanga, Myrianthus and Pourouma (in which they are dIfferentiated as
more or less curled and often filiform woolly hairs). in Af'tocaYpus gla«cus, Bl. and
A. glaucescens, Tree. (in whieh almost all the epidermal cells grow Otlt into clothing
I hairs), and in certain species of Coussapoa. in which the hairs inserted on the veins
cover in the areolae (i.e. the spaces between the veins) on the surface of the leaf.
It remains specially to mention the following types of trichomes: (a) the long
clothing hairs, which are found in species of Castilloa, Ficus and other genera, and
are provided wit}]. a basal pedestal composed of epidermal cells; (b) the trichomes
·of Helicostylis POeppig~'atla, Tree., in which,.the lageniform base of the hair pene-
trates deeply into the mesophyll; and (c) the bracket~hairs, which, besides occurring
in speCies of A rtocarpus and Cecropia, are found also in species of Bt'osimum and
Lanessania. The walls of the clothing hairs are often silicified, and when this
is the case they frequently have a rough surface due to the presence of knobs of
v aryi ng shape.
Papillose hairs are found in the first place among the Artocarpeae, where they
show varied differentiation. They include: (a) the bulbiform tricliomes of Balano-
streblus, PoUisenia and Sahagunia. which have thick walls and very reduced
lumina; (b) the silicified trichomes of Aytocarpu$ Irutescens. Renn., A. Limpato,
Miq. and A ;~scandens,"Renn., "the outer wall of which is fiat, being protruded only
in the middle, "while the lateral walls are delicately undulated and pitted; (c) the
short trichomes of species of A rtocarpus, belonging to the sections]aca and Pseudo-
jaca; the basal portion of these trichomes is enlarged like a spadix .and pitted in
various ways, while the body of the hair merely appears as a solid and silicified
tuberculate knob or as a tip adpressed to one side of the basal portion of the hair.
Among the Conocephaleae papillose hairs, which are more or less strong~y siJicified,
occur in Cecropia, Coussapoa, Musanga and Pourouma. In Cecropia laXiloba, ::\Iiq.,
these hairs occur in large numbers in the upper epidermis and have thin walls,
while in C. me.x£cana, Hemsl. they have a thu;;k silicified outer wall and are occa~
5ionally provided with a thin longitudinal wall. Flat papillose haiTS with a minute
tip inserted at the centre of the cell are found in other species of Cecropia. as well
as in Coussapca v£/losa, Poepp. et Endl. and Musanga Smithi;. With these hairs
we may class the somewhat larger bulbiform trichomes, occurring in certain species
of "'Cecropia and Pourouma; the latter have. a longer tip, and. in Pourotfnta eyen
occasionally exhibit delicate longitudinal walls:
 1062 ADDENDA--UR TICA tEA E
The external glands of the Artocarpeae and Conocephaleae never form
a resinous or oily secretion; they are present in all the genera investigated
and; according to Renner, show a varying type of structure. Among the
Artocarpeae the stalks of the glands are mostly unicellular; the glands occur
isolated on the surface of the leaf and are never arranged in groups. In most
cases the glandular head is ellipsoidal or spherical and divided both by longi-
tudinal and transverse walls. Long uniseriate glands, consisting of three or
four cel1s, have been observed among the Artocarpeae only in Ficus rubigi1tosa,
Desf. and antler~shaped glands (produced by the outgrowth of the cells of
the glandular head into sac-shaped processes) only in SParattosyce. Among
the Conocephaleae, Conocephalus alone has glands arranged in groups of 2-I5 ;
these glands have a unicellular stalk and a head, which consists either of an
irregularly shaped mass of from 3-6 cells (C. ovatus, Tree.) or of three or more
diverging tubular arms composed of 1-3 cells (C. suaveolens, Bl.). 'In the
remaining genera (Cecropia, Coussapoa, M1lsanga, Myrianthus, Pourouma) the
lower side of the leaf as a general rule bears isolated uniseriate glands, which
are long bent structures composed of 4-10 cells, the upper cells being more
or less swollen and forming a kind of head. The upper side of the leaf in
Cecropia bears glands, which have a multiceHular stalk and a head, composed
of numerous cells and divided both by transverse and longitudinal walls;
these glands are either isolated or arranged together in groups comprising
from two to four glands. Jfusauga, Myrt'antkus and Pourouma have groups
of short external glands on the upper side of the leaf, each group consisting
"of from two to seven glands; the latter are broadly club-shaped and have
a stalk, which is made up of short cens and merges gradually into the head,
the latter being divided both by transverse and longitudinal walls. In species
of Cecropia and Pourouma transitional forms between the glands found on
the upper side and those present on the Jower side of the leaf have been demon-
strated, while in certain species of Cecro-bia filiform glands have been observed
also on the upper side of the leaf.
\Ve may add the following details regarding the structure of the head in the
glands of the Artocarpeac. In some cases the heads arc merely unicellular or
divided by a single longitudinal wall (A1'tocarpus § Pseudojaca, Ft"cus § Urostigma.
Parartocarpus, Scyphosyce); in other Cases they are divided into four quadrants
(Ficus clavata, WaII., F. Cunt'a, Ham., and species of Artocal'pus belonging to the
section Prainea) or have a peltate or flabelliform shape with diverse arrangement
of the cells (certain species of Artocarp1es and F£cus).
According to Renner the pearl·glands (Muller'S bodies), which have
already been discussed in the earlier part of this work, are found only in
Cecropia., and the statement (made on Meyen's authority) as to their occurrence
in Pourouma should be cancelled. The pearl-glands, which are situated on
the petiolar cushions, consist of an epidermis of small cells, which inc1ude~
an apical stoma~ and of a complementary tissue, -which is likewise composed
of small cel1s; side by side with the glands of this type the lower surface ot
the leaf bears pearl-glands, which have an epidermis consisting of large ceUs
but not provided with a stoma, and a complementary tissue, composed of very
large cells.
In the main portion of this book no mention was made of the glandular
spots, which OCcur on the leaves in many species of Ficu~, and which in
F. hispida, L. and F. leucantotoma, Poir. are found also on the surface of. the
branch next to the points of insertion of the petioles; according to Renner
they are of the nature of wax-glands and not extrafloral nectaries, as stated
by Mirabella. They are provided with a palisade-epithelium, which is without
stomata, and are present in species belonging to all the sections. They are
found on the lower side~f the leaf. where they either lie sing1y on the midrib
and at the base of the la~ina or in addition to that occur in pairs in the angle~
 ADDENDA-URTICACEAE

between the lateral veins of the first order and the principal vein or even in
the axils of the more delicate lateral veins; in asymmetrical leaves, it may be
noted, the paired glands are found only on that half of the leaf the growth of
which has been promoted. In F. diversifotia, BI., in which the median vein
is forked, a gland occurs in the angle, formed by the bifurcation. As important
from a systematic point of view we may mention that in almost all the species
of Ficus belonging to the section Urostigma (excepting F. nervosa, Heyne
and F. pubinervis, Bl.) there is an unpaired wax-gland at the base of the midrib.
As regards the structure of the cortex (p. 775) it is noteworthy that
","ccording to Auer the bast-fibres in all the investigated genera of the Moraceae
(Antiaris, Artocarpus, Brotessonetia, Cecropia, Cudrania, Ficus, Afaclura,
Morus, Olmedia, Streblus) agree in the fact that the outer thickening-layers
appear as an envelope around the inner layers.
In Artocarpus incisa, L.f. andA. integrijolia~ L.f. the stalks of the fruiting ..
axes show polystelic structure.

4. URTICEAE (pp. 775-77 8

To the REVIEW OF THE ANATOMICAL FEATURES we may add that bundles
of rap hides and styloids occur in Laporiea, secretory organs resembling latici-
ferous tubes in Urera bacci/era, Gaud. and mucilage~canals also in Laporiea.
For the structure of the leaf in Laportea, see Quanjer, lac. cit. In
Laportea the stomata are Hkewise provided with three subsidiary cells
arranged according to the Cruciferous type. According to Nestler hydathodes,
similar to those found in Pilea serpyllifolia and appearing as slight promi~
nences, are scattered over the entire upper surface of the leaf in Bohmeria
tenacissima, Gaud. (=B. nivea, Gaud.). A single layer of hypoderm is present
on the upper side of the leaf in Laportea stimulans, Miq., but not in L. sinuata,
Bl., which is mentioned as a synonym of L. stimulans in the Kew Index.
To the section dealing with the cystoliths we may add that in Laportea
they are situated in the epidermis of the leaf and have a distinct stalk, while
the cells containing them in some cases (L. sinu,ala, Bl.) are provided with
a papillose tip. In L. pel/ala, Gaud. deposits of carbonate of lime have been
observed on the surface of the leaf, an<J in L. sinuata, Bl., they are found in
the mesophyll as well. The new record of the occurrence of raphides and
styloids in Laportea is contained in Qu~jer's paper and is correct.
The glandular hairs in Laportea are provided with a short stalk and
a head divided by vertical walls.
According to Guerin the secretory organs of Urera bacci/eTa above men-
tioned resemble laticiferous tubes; they occur in the periphery of the pith,
the inner portion of the primary cortex and the secondary bast of the stem,
in the cortex of the root and in the neighbourhood of the vascular bundles
in the veins' of the leaf. The: elements found in the pith branch after the
manner oflaticiferous cens. On the basis of my own observation I may add
that the xylem-mass in the stem of U. bacci/era contains tangential bands of
unlignified parenchymatous tissue, which show a concentric arrangement.
Mucilage-canals have been observed in the larger veins of the leaf of
Laportea stimulans, Miq. vaL costata, while in the veins of the leaf of L. sinuata,
Bl. Quanjer merely found an intercellular canal without any secretion.
Literature: Karsten, Cecropia pe/tata, Nova. Acta Leopold.-Carol.. 18.;8; and in Ges. Beitr.,.
i, 1865, pp. 241-52.-Trecul, (.j~mme et tlnnin dans Ie COl1ocephal1lS Ilauc/eijiarm, Ann. sc. nat .•
sera 5, t. ix, 1868. pp. 274-81:-Faivre, Latex du Murier blanci Ann. sc. nat., sec. 5. t. I, (869.
pp. 97-IH.-Costantin, Tiges aer, et sout., Ann. sc. nat., ser. 6, t. xvi, 1883. p. 85 et seq.-[MoUer.
Nesself3.ser. Deutsch. aUg. pol}1echn. Zeit, IS83, n. 34, 35.)-Keller, Luftwurzeln, Dis!>., Heidelberg,
1889. pp. 26-3o.-Lothelier, Epines, These, Paris, 1893. pp.16, 11 a.nd pI. i.-[Bigdow, Gla.nds in.
the Hop tree, Proceed. Iow:1 Acad. of Sc., ii, 189:'. pp. I~8~40, I p1.]-Mirabella. Netlaf!~
 ADDENDA-URTICACEAE
estranuz. nelle vade specie di Ficus, Nuo~o Giorn. bot. Ital., n. s., ii, 1895, pp. 340-7 and Tab.lo..-
Tschirch, Herba Cannabis indica, in Anat. Atlas. i, 1895, Tab, xv and p. 55 et seq.-Went, Haft- u.
Nah~rz:, Ann. Jardin ,Buile~zor~, xii, ~895' pp. 48-54.-[BriOli e Tognini, Anat: dell!- Canapa,
Parte 11, Atti 1st. bot. dl Pa Vla, Ser. :I, IV, I 896, p. 175 et seq., 26 tab.; abstr. In Be1h. z. Bot.
Centrnlb1.. viii, 1898-9, pp. 27-9, find in Just, 1897, i, p. 52;l.]-Biittner,Cortex Murore (l.{roslig-JIIa
cysttJj!Qdutn, Miq.), Diss., Erlangen, 1896, 31 pp.-Jonsson, Anat. Bau d. Blattes. Acta Uni',ets.
Lund., xxxii, :I, 1896 (PelliQnia).-[Baccarini e Buscemi, Nettarii foliari della Ol1llediella Cesatiall4,
BaHl., BuH. Accad. fJioenia Catani.a, Fasc. lvi, 1898, pp. IO-IJ; abs~. in Just, r898, ii, p.. 386.]-
Engler, Moraceae, In Monogr. afrtkan. Plt-Fam. u. -Gatt., 1898, pp. 8-10 and 38, 39,-Mlrabella,
Laticiferi delle radici aeree di Ficus, Contribuz. biolog. veget. 1st. bot. Palermo, ii, 1898, pp. 13~-6.
-Grelot, Caoutchoucs et Guttapercha, These, Paris, 1899. p. 133 et seq.-Haberlandt, Neues Organ
bei C()1l(K4jltaius ovaltn, Festschrift f. Schwendener, J899. pp. lo4-Ig.-HouJbert, Phylogenie des
Ulmacees, Revue gen. de Bot., 1899, pp. 106-19 and pI. 2, 3.-Nestler, Wasserausscheidung an den
Blattem von Biikmeri(J·Arteo, Sitz.-Ber. Wiener Akad., cviii, Abt. i, 1899, pp. 706-8 and Tab.~
Tunmann, Se~retdriisen, Diss., Bern, 1900, pp. 2O-3.-Ursprung, Anat. n. Jahresringbild. trop.
Holzarten, DiS$., Basel, 1900, pp..r;-8 (ArltlCarpUS inl~grijolia, L f.).-BargagH-Petnlcci, Cavita
stomatiCere nel genere Ficus, Nuovo Giom. hot. Ita1., n. 5., viii, 1901, pp. 492....s.-~01isch;
Milchsaft u. Schleimsa(t, I90r, pp. 6, 21, 49 and 5fi.-Petersen, Vedanatomi, 1901, pp. 38-+0
(Ulmus).-Pitard, Pericycle, These, Bordeaux, 1901, pp. 84, 85.-A. Richter, Luftwurz., Bibl. bot.,
Heft 54, 1901, p. 44 and Tab. xiv.-[Baar, Milchrohren, Sif.t.-Her. 'Lotos' in Prague, 1902, n.-f,
5; abstr. in Bot. Centtalbl., xcii, p. 406.]-Bargagli·Petrucci, Legnami. Malpighia. 190:1, p. 290
et seq. CArtocaryus, Sloilia); and Conocephalus. Nuovo Giom. bot. Ital., ix, 190z, p. 213.-
Bouygues, Petiole, These, Paris, 1902, p. Is.-Cas. de Candolle, Hypoascidies de ';;"ims, Bull.
Herbier Boissier. ser. 2, t. ii, 1902, pp. 753-6:1 and 2 pl.-Clauditz, Blattanat. canar. Gew., Diss.,
Basel, 1902, PP' 39, 40 (GeS1louinia).-Copeland, Haberlandt's new organ of COlloceplta/us, Bot.
Gazette, J 902, PP' 30o-S.-Auer, Bastfasern d. Morace~n, Osterreich. bot. Zeitschr., 1903, pp. 353-6.~
Quanjer, Anat. bouw, &c.• Natuurkund. Verhandel. Haarlem, iii, 5, 1903 (Artocarpur, Laportea).--
Solereder, Arl{)(arpus laci1tiata, Hort., Bull. Herbier Boissier, ser. :.I, t. iii, 1903, pp. 518-20.~
TUZion, SpiraIige Struktur d. Zellw. in den Marklltr., Ber. deutsch. bot. Gesellsch., 1903, p. 276.-
Col, Faisceaux, Ann. sc. nat., ser. 8, t. xx, 1904, pp. I3.4-8.-Kniep, Bedeut. d. Milchsaft., :Flora,
1904, pp. 168, J69.-Paoli, Eterofillia, Nuovo Giorn. bot. hal., xi, 1904, PP' J88-204 and Tab. i.
-Sol~reder, Zwei Berichtigungen, Bull. Herbier BoisSier, ser. ;I, t. iv, 190-4, pp.318-23.-Areschong,
Trop. va:Kt. bladbyggu., Sv. Vet. A.kad. Handl" 39,11. 2, 1905, PP' 9-1 J and pp. 113-15, Tab. is. x
and xv (IIrtocaryus) and pp. 112, 1l3t Tab. XV, xvi (An#aris).-Frommel, Plantas chil. text., l~S,
p. 31.-Guerin, Laticiferes de I' Urera bacci/era, Bull. Soc. bot. de France, 1905, p. 406 et seq.~
Mayos, Milehr., Beih. z. Bot. Centralbl., xviii, I. Abt., 1905, pp. 27S, 276.-SartoD, Anat. d. pI.
aRines, Ann. sc. nat., ser. 9, t. ii, 19O5; pp. 60-2 (Panelaria).-Dattphine, Rhizomes, Ann. sc. nat.,
ser. 9, t. iii, 1906, p. 363 et seq.-Perrot, Gilbert, Carnat et Choay, Rech. sur les Cecropia, in Perrot,
Travaux, iii, 1906, 38 pp.; also Bull. d. sc. phannacol., xi, I905.-Piccioli, Gelsolino, sep. copy.
JgOO, 13 pp.-Piccioli, Legnami, Bull. Siena, 1906, pp. 135, 1,0, 17+ and 17 5.-Renner, Anat. n. Syst.
d. Artoc:1rpeen u. ConocephaJeen, insbes. d. Gatt. Ficus, Diss., Munich, 1906 j sep. copy from Engler,
Bot. Jahrb., xxxix, pp. 319-44.8.-Renner, Wachsdrusen a.uf d. BI. u. Zweigen von Fitus, Flora, J90~,
pp. 24-37·-Boorsma, Aloeholz, Bull. du deptut. de l'agricult. aux lndes neerl., n. vii, 1907, p. 33
{Celtis).-lIoltennann, Einflulls d. Klimas, 1907, pp. 17, 106, 135 and 136.-[Schorn Schleimz.
be,; einigen Urtjcaceen 11. Scbldrncystolithen bei Cirardinia pabJUlla, An2eiger Wiener Aknd., l\.f.ath .•
naturw. Kl., 1907, p. 65; and in Sitz.-Ber. Wiener Akad., c)tvi, Abt. 1, 1907, pp. 393-410, 2 Tab.]

ApPENDIX: TRIBE VII. THELYGONEAE (p. 779).

In Thelygonum the upper epidermis of the leaf is characterized by the forma-
tion of papillae.
Literature: Gutenberg, Lichtsinnesorg., Ber. deutsch. bot. Gesellsch., 1905, pp. 269, 270 and
Tab. x, xi.

PLATANACEAE (pp. 779-781).

According to Bouygues, the vascular bundles of the petiole are what he
calls 'pseudo~faisceaux rayonnes.' In the mature condition they show an
annular arrangement, although they do not arise from a single procambia]
strand; in correspondence with this their conjunctive tissue resembles the
ground tissue.
Literature: Pitard, Peri cycle, Th~. Bordeaux, 190', pp. 81, 8z.-Bol.lygnes, Celt. (ocme.;; va%c.
anorm. du petiole, Act. Soc. Linn. Bordeaux, lvii, 1902, pp. 91, 9',-TU2son, Spir:tlige Struktl.lr d.
Zellw. in den Markstr., Ber. deutsch. bot. Gesellsch., 1903, p. :Z76.-C()t, Ji'aisceaux, Ann, sc. nat.,
~r .. 8, t. :xx, 1904. p. 1.4J.-Piccioli. Legnami, Bult. Siena, 1906, p. 125.
 ADDENDA

JDGLANDEAE (pp. 783-785).

Literature: Kassner, Mark einiger Holzpfl., Djs~., Basel, 1884, pp. J 1-15 and Tab. i.-Pitard,
Pericycle, These, Bordeaux, 1901, pp. 55-7.-Co), Faisceaux, Ann. fiC. nat., set'. 8, t :n, 1904.
p. 1-46.-Piccioli, Legnami. Bull. Siena, 1906, pp. 131 and 17 I.

)IYRICACEAE (pp. 785, 786).

The ANATOMICAL CHARACTERS OF THE ORDER, which were enumerated in
the earlier part of this book, are confirmed by the results of Chevalier's recent
researches. It only remains to point out that in addition to the peltate external
glands one finds stalked capitate glands, which are either uniseriate or unicellular,
and that uniseriate clothing hairs occur side by side with the unicellular ones.
The STRUCTURE OF THE LEAF is now adequately known as a result of
Chevalier's investigations. The leaves are almost always bifacial in structure,
the palisade~tissue consisting of one or two or sometimes of several layers.
The stomata are confined to the lower side of the leaf, and are either raised
above the level of the epidermis (e.g. in Jfyrica Faya, Ait.) or covered in by
the neighbouring cells (as in the sections Comptonia and Gale). They develop
according to the Ranunculaceous type and have 5-10 neighhounng cells;
in Myrica co rdifolia t L., they are restricted to deep pits in the surface of the
leaf, while in M. Gale, L., they are distinguished by the papillose character of
the neighbouring cens. The epidermal cells are for the most part small
and have straight or undulated lateral walls; the cuticle is commonly granular,
more rarely striated. Papillose differentiation of the lower epidermal cells,
like that observed in M. Gale, appears to occur rather frequently. M'1'avanica,
Bl. is alone characterized by the possession of a hypoderm, which is situated
on the upper side of the leaf and consists of two layers. The peltate glaDds
mentioned in the earlier part of this book are present in almost. all the species ;
their shield, as we may notice in passing, may be divided both by tangential
and radial wans and the"component cells may eventually even show an irre-
gular arrangement. Side by side with these glands other types of glandular
hairs occur. In M. Gale, L., M. aspleni/olia, L. and M. esculema, Buch.-Ham.
isolated epidermal cells of the ordinary kind occasionally function as glands ;
they become filled with oil, and their walls remain thin, while the outer walls
become slightly arched outwards. In species of the sections Comptonia
and Gale (e.g. Myrica Gale) one also finds uniseriate external glands, which
are composed of three or four cells; of these either' the terminal cell or one
of the middle cells secretes oil. It is no great step from these forms to the
stalked external glands, which, for instance, are likewise found in M. aspleni-
folia and M. esculenta and have heads composed of a small number of cells.
It remains to mention those types of peltate glands, in which the shield is
provided.)\jth.aojagged margin_ (e.g. in M. pennsylvanica, Lois.-Desl.). Uni-
cellular- sclerotic clothing hairs, which have either a smooth or a verrucose
surface, are found in varying abundance in all the species. In M. Zeyheri,
c.nc. and M. asplenifolia these trichomes occasionally occur in pairs, while
in M. pennsylvanica and }.II. 9uercilolia, L., their basal portion is surrounded
by several rows of small Subsldiary cells; in M. asplenifolia small peg~shaped
outgrowths, which penetrate in between the neighbouring epidermal cells,
are often developed on the base of the hair. In the hairs of M. esculenta,
Buch.-Ham. var. longi/oUa (Teysm. et Binn.) Chevalier figures a few relatively
thin transverse walls.
Three vascular bundles enter the petiole; in their further course these
bundles approach one another and ultimately fuse to form an arc of wood
and bast. In rare cases the lateral bundles undergo subdivision, so that five
(066 ADDENDA-lt/YRICACEAE

vascular bundles are to be found at certain points (e.g. occasionally in M. Nagi,

Thunb.).
As regards the structure of the cortex we may add the. following details.
The primary medullary rays between the groups of bast broaden out towards
the exterior in the shape of a wedge. There are no sclerenchymatous elements
in the secondary bast in M. Gale, in contrast to M. aspleni/olia (where they are
generally isolated) and other species (where they fonn groups). Secretory
cells (' cellules mortes de lignine gommeuse ' according to Chevalier; (Zellen
mit rotgeHirbtem Milchsaft' according to Beringer) occasionally (e.g. i~ M.
aspleni/olia) occur in the cortex. The statements in the literature as to the
occurrence of laticiferous tubes or laticiferous vessels (see Hooper and Dymock)
have as little foundation as the recorded presence of resin-canals 1, discussed
in the earlier part of this book.
Literatnre: [Dymock, Indian Drugs, Pharm. Joum. and Transact., 188o, p. ,s81 et seq.; abstr.
in. Bot. Centralbl., J 880, iii, p. 977.]-lHooper, Myrica Nagi, Americ. Joum . .Pharm., 1894. p. 209;
abstr. in Jnst, 1894, ii, p. ,po.]-[Berillger, lI-fyn'ca, Americ. Jonrn. Pharm., 1894, p. :uo; abstr. in
Just, 1894, ii. p. 401.]-[Planchon et Collin, Drogues simples, Paris, 1895 (.Myrica urifera).]-
Chevalier, Appareil vegthatif des Myric., Comptes rendus Assoc. fran't. pour l'avancement des sc.,
Nantes, 1898, i, publ. 1899, p. 16... , and ii, 1899. pp. 457-66.-Kearny, in Contribut. U. S. Nat.
Herb., v, n. 5, 1900, p. 294.-Chevalier, Monographie des Myricacees, These. Paris, 190 1 , ""2.57 pp.,
8 pi., 1 carte; also in Mem. Soc. Sc. 'de Cherbourg, xxxii.-[Kremhs and Denniston, Structure of the
~tem of Myrica Gale and M. cerift1'a, Proceed. Americ. Pbarm. Assoc., 1901, pp. 414-:I3.1-Petersen,
Vedanatomi, 1901, pp. 37, 38.-Pitard, Pericycle, These, Bordeaux, 1901, pp. 78, 79.-Clauditz.
Blattanat. canar. Gew., Diss., Basel, 1902, pp. 44-6.

CASUARINEAE (pp. 786-790).

For the detailed structure of the stoDlatal apparatus in CaEuarina, see Porsch.
ll. cc. According to this author a particularly noteworthy feature from the phylo~
genetic standpoint, considered in relation to the indications of affinity between the
Casuarineae and the Gymnosperms. lies in the intercalation of cutine-lamellae in
the cellulose-membrane of the dorsal and ventral walls of the guard-cells, which
are rather deeply sunk. These lamellae completely coincide with the • xylem-
lamellae' of Gymnospermous stomata both in their shape and in the extent of their
development.
Literature: Hohnel, Gerberinden, Berlin, [SSO, p. So.-[Monni, Anat. della radice delle
Casuarin., Mem. Accad. Bologna, Ser. 5. \'01. 6, pp. 201-24,2 tab.; abstr. in Just. 1897, i, p. 507']-
Hargagli.Petrucci, Legnami, Malpighia. 19°2, p. 288.-Porsch, Spaltoffnungsapparat von Casuarilla
u. seine pbyletilche Bedeut., Oster reich. bot. Zeitschr., [9°4, n. 1 et seq.; sep. copy, 21 pp. and Tab.
iii.-[Maiden, Casuar;lIa in()plzll1ia. Proc. Linn. Soc. New S. Wales, 1905, p. m.]-porscb, Del
Spaltoffnungsapparat im Lichte der Phylogenie. Jena, 19051 p. 17 et seq.

CUPULIFERAE (PP.791-797).
2. STRUCTURE OF THF: LEAF. The following supplementary facts refe[
mainly to the structure of the leaf in Quercus (excI. Pasania), which ha:"
recently been examined in detail especially by Schott, Kuster and Brenner.
IsolateraI structure~ like that found in Q. !lex, occurs in the leaves of many
species, e.g.: Q. agri/olia, Nee, Q. alni/olia, Poech, Q. calUprinos, Webb~
Q. chrysolepis. L:ebm., Q. cinerea, Michx., Q. cocci/era. L., Q. crassipes, Humb.
et Bonpl., Q. incana, Roxb., Q. macrolepis, Kotschy, Q. persica, Jaub. et Spach.
Q. Vallonea. Kotschy, and Q. xalapensis, Humb. et Bonpl. (according to
Schott). Hypoderm is present in a few species of Quercus (e.g. in Q. xala-
pensis according to Schott, in Q. de1lsi/lora, Q. glabra and Q. pachyphylla accord-
ing to Kiister, and in Q. virginia1la. L. according to Kearny). A tendency
to form papillae on the 10wer epidermis is recorded by Kiister in Q. Ballota,
l According to Chevalier the plant, which was examined by Hohnel under the name of Myri(~
sapida and wai lonnd to be pro\'ided with re!iin-can.1b. ii probablr r()d~(a,../,us /I.'i7§i.
 ADDE!,/DA-CUPULIFERAE

Q. cuspidata, and Q. glabra, by SchoU in Q. chrysophylla, and by J{ohne also

in Alnus incana, WiUd. and A. rubra, Bong. In all the species of Quercus
that have been examined, the stomata are confined to the lower surface of
the leaf. For the detailed structure of the stomata in Q. Ilex, see Guttenberg,
loco cit.; a particularly noteworthy feature is that the guard-cells develop
peculiar longitudinal cuticular folds on the upper portion of the ventral walls.
which surround the front cavity; in a transverse section these folds appear
as small projecting horn-like outgrowths, lying opposite to one another. Less
strongly developed ridges of the same type have been observed also jn Q. Suber.
L. In the majority of the species of Qu.ercus three arcs of wood and bast
are found between the base and the middle of the midrib of the leaf, and the
same is the case in the petiole; the upper arc is inversely orientated (with
the phloem on the upper side), while the middle and lower arcs are normally
orientated. The middle one arises from the upper one by the inrolling of
its ends (i.e. by a process of twisting through an angle of 180°); for the dis-
tribution of this median arc of wood and bac;t (which was first observed by
Frank) in the genus Q1,ercus, :see C. de Candolle and Schott, n. cc. For details
as to the structure of the margin and teeth of the leaf in the species of Quercus,
see likewise Schott. -
To the previous discussion of the clothing hairs (p. 792) we may add
the following details. Simple unicellular clothing hairs with walls of varying
thickness also occur in Quercus, although not very commonly (e.g. in Q. annu-
lata. Q. dentata, Thunb., Q. glandulifera, Bl., Q. hungarica, Q. mongolica, Fisch.•
Q. robur, L., Q. sessilijlora, &c.). On the other hand stellate and tufted hairs
are-widely distributed in the genus Quercus, the number of ray-cells occasionally
being small (2-4) and large in one and the same species. The following types
require special mention: (a) in the stellate hairs, which cover in the stomata
on the lower surface of the leaf in Q. leucocarpa, Hook. f. et Th. and Q. Henryi,
Seem., the 3-5 rays show unilateral development, and the hairs, as seen from
the surface, resemble a hand with outspread fingers (according to Brenner);
(b) the stellate hairs of Q. crassipes, Humb. et Bonpl.. Q. fulva, Liebm.,
Q. imbricaria, Michx. and Q. nitens, Mart. et Gal., in which the lov/er parts
of the ray-cells are fused to form a stalk to the hair (according to Schott) ;
(c) the tufted hairs of Q. Brantii, Lindl., which are composed of a particularly
large number of cells (forming as many aS~20 rays), and in which the ray-cells
are arranged approximately in two tiers (according to Schott); (d) lastly hairs,
which resemble stel1ate trichomes or small scales and have thin cell·walls;
these are described by Brenner, who figures them in Q: (Sieboldii' and records
them also in Q. cuspidata, Thunb., Q. ]unghuhnii, Miq., Q. lanceaefolia, Roxb.
and Q. sclerophylla, Lindl. et Paxt. According to Schott true peltate hairs
occur in Q. nigra, L., which is not included in the s~ction Pasania, but in the
section Erythrobalanus; the shield has a jagged margi!l, and the 8-14 ray-
cells of_w]li~h_it_is _~on1posed meet in the centre. We may add that according
to Kuster brown stellate hairs are found on the Len#cztiaris-galls of Q. pedun-
cutata, and two-armed hairs on the N umismatis-galls of the same species;
the type of hair first named is not normally present in Q. peduncttlata, while
two-armed hairs have not as yet been observed as a normal feature in any
species of Quercus. The two types of glandular hairs, previously described
by me in Q. Farnetio, occur also in other species of Quercus. ~hus according
to Kiister nniseriate glandular hairs (Fig. 187, J) are found also :in Q. alba.
Q. aquatica, Q. glaucoides. Q. magnoliae/olia, Q. nigra, Q. oblongifolia, Q. reti-
culata. Q. Sartorit, Q. undttlala, and according to Schott in Q. alnifolia, Poech
and Q. imbricaria, Michx.; according to Kuster capitate glandular hairs are
present in Q. pedunculata, Q. ilicifolia, Q. lyrata, Q. macrocarpa. &c. Brenner
(loc. cit., p. 154) also~ 'records glandular hairs, which probably function as
 ADDENDA-CUPULIFERAE

hydathodes, in Q. 1'ejlexa, King and other species, but the details of their
structure are not recognizabJe in his figure (30 a, a).
3. STRUCTURE OF THE AXIS. For the strongly developed cork-wings
found on the axes of Q. macrocarpa, Michx., see Gregory, loco cit. Regarding
a peculiar spiral structure in the medulJary rays, see Tuzson, loco cit.; this
feature is only noticed at points at which the xylem-mass is ruptured and is
particularly well shown by Fagus silvatica, but also by Alnus incana, Betula
vc"ucosa, Carpinus Betulus, Ostrya carpinifolia, QucrcusCerris and Q. sessiliflora.
Literature: Hahnel, Gerberinden, Berlin, 1880. p. 53 et seq.-Kiigler, Kork von Quercus mOe"
Archlv d. Ph ann.• 1884. pp. :z 17-30.-Gregory, Cork-wings, Bot. Gazette, 1888, l'P' 249, ~so.­
Hartig, Eichenhob:, Sitz.-.l:Jer. Miinch. Akad., 189 ... p. 385 et seq. and elsewhere.-rLamey, Chene
liege, Paris, 189.f..]-Kohne, Papillen n. obers. Spaltoffn., Mitteil. deutsch. dendrofog. Gesellscb.,
18w, p. 53.-Kearny, in eontribut. U. S. Nat. Herb., Y, n. 5, 1900, p. 29.....-Kiister, .Bemerk. iiber
die Anat. d. Eichen, Bot. Centralbl., 1900, iii, pp. 177-8S.-Eug. Ant. Muller, Korkeiche. Abh. k.
k. geogr. GeseUsch. Wien, ii, J9°o.-SchoU, Anat. Hau d. Blattes d. Gatt. Querros, Diss., Heidel-
berg, J9OO, 53 pp., 3 Tab.-Tnnmann, Sekrddriisen, Diss" Bern, 1900. Pp. 17-20.-Peter~,
Vedanatomi, I~I, pp. a8-37.-Pitnrd, Pericycle, ',I'hese, 1901. pp. 62-+-Bargagli-Petrucci,
Lcgnami, Malplgbia, J902, p. 289 (QueJ"(us):-W. Brenner, Klima u. Blatt Lei Q"n'"(US, "lora, 1901,
pro J 1....-6o.-[Piccioli, Monografia tIel Castagno, Firenze, 1901.]-Simon. Holzkorper sommer· n.
Wlntergriiner Gew., Ber. deutsch. bot. Gescllsch., J90~. pp. 2 .... 2-.... and Tab. xi.-Tllzson, Spiralige
Struktur d. Zellwand in den Markstr. des Rotbucbenhol.zes, Ber. deutsch. bot. Gesellsch., 1903,
pp. '7~.-Col, Faisceallx. Ann. sc. nat., aer. 8, t. xx, l!;lo4, pp. 146-9.-H. Winkler, Betulaceae,
ID Pflanzenreicb, Heft 19, 1904, pp. 3, .....-Siissengnth, Bebaarungsverh. d. Wiirzb. Muschelkalkpfl.,
Vi 15. , Wiirzborg, 1904, p. 5 I .-Haberlandt. LichtsiJlnesorg., 190!', p. 97 aod Tab. i.-Piccioli,
Qaerci Italiane, Fireoze, 1906 ,60 pp.-Piccioli, Legnami. Hull. Siena, 1906, pp. 133, '37, 139, 1...3,
168, 169, J76.-Guttenberg, Immergr. Laubbl. d. Mediterranflora, in Engler, Hot. Jahrb., xxxviii.
1907, pp..... ' 7- J 9 (QutrcltS Iltx).- [For fn rther literature, see p. I I 70.]

SALICINEAE (PP.797-799).
The following further points of distinction between the two genera of
this Order may be added: Populus alone has bundles of sclerenchymatous
fibres in the rnedulJary sheath, and invariably has stone-cells in the cortex,
while stone-cells are never present in the cortex in Sali:x; the cork in Salix
always contains cells with rather strongly thickened outer tangential walls,
whereas such cells do not occur in the cork in Populus. As regards the
mode of origin of the cork, which in the earlier part of this bookwas mentioned
as a means of distinguishing the two genera, recent investigations have con-
firmed the statement that epidermal development of the cork is characteristic
of Salix; in Populus the cork generany arises in a subepidermal position and in
P.Premontiialone(according to Perredes)in the third layer of the primary cortex.
The following additions on the structure of the leaf deal principally with
Salix and are based on Camus's observations. The type of stoma, previously
described in Salix, is present throughout the genus; the stomata in Salix
are always found on both sides of the leaf. Gelatinized epidermal cells
have been recorded also in S. babylonica, S. caesia t S. glabta, S. glauca, S.
hastata, S. helvetica, S. Lapponum, S. pentandra, S. Phylici/olia, S. pyrenaica.
S. repens, S. reticulata and S. retusa. In all probability, however, they are
even more widely distributed, for the 1-3 tangential walls described by Camus
are probably in aU cases (this is certainly so at least in S. nigricans, as I have
found by a reinvestigation of this species) merely the cellulose-lamellae of
gelatinized cell·walls. Wax (in the form of small rods) is excreted on the
surface of the leaf in many species of Salix. The mesopbyU is differentiated
after the manner of a hypoderm (just as in S. alba) also in S. babylonica. In
alcohol-material Jf the leaves of S. cinerea, S. daphnoides and other species
Camus observed sphaerocrystalline excretions (d. also the abstract of Dobrow-
lianskij's earlier paper in Just, 1893, I, p. 560 et seq.). In connexion with
the glandular Jeaf-teeth and the glandular spots on the bud-scales, mentioned
 ADDENDA-SALICINEAE

on p. 798 as occurring in PlJj>ulus, we may point out that in many species of

this genus extraftoral nectaries are found on the leaves formed in spring,
or even on all the leaves; the structure of these nectaries, which are situated
at the apex of the petiole, is similar to that of the leaf-teeth (Trelease).
To the section dealing with the structure of the wood we may add that
in Salix aurita and S. cinerea (but not in S. Caprea) portions of the wood pro-
ject like ridges into the bast; this feature occurs also in otherspecif's of Salix,
but is less distinct (Camus).
In turning our attention to the structure of the cortex we may once more
refer to the above-mentioned features distinguishing Populus and Salix (pre-
sence or absence of stone-cells and structure of the periderm); they were
determined by So~tarj~ and Perredes. According to Camus the pericyc1e in
Salix usually contains a ring- of fibres, which subsequently becomes burst open,
while in other cases isolated groups of fibres are present from the first; in
the creeping and procumbent" axes of S. herbacea and S. polaris and in those
branches of other specie!;, which show the same habit, there are no pericyclic
fibres. The groups of fibres in the secondary bast vary in abu~dance in Salix;
in S. reticulata no secondary hard bast was found, even in thick axes.
Regarding the pith the. following details may be added on Sostari~'s
authority... In most of the species of Populus ,the inner cells of the pith are
partly lignified and partly unlignified; in P. euphratica alone all the cells have
thick lignified waIls. As regards the genus Salix the medullary cells are lignified
in the shrubby willows, .while in the dwarf and glacial willows only isolated
cells of the pith are Jigriified; or (5. ret usa, L~) lignified cells are completely
absent. In the case of Populus euphratica the 'groups of sc1erenchymatous
fibres, characteristic of Populus (see above), are present not only at the margin
of the pith, but are scattered throughout its entire mass. According to Camus
carbonate of lime is deposited also in the pith of S. triandra.
Literature: Hobnel, Gerberinden, Berlin. 1880, p. 87 et seq.-rTrelease, Nectar-glands of
Populus, Bot. Gazette, 188t, pp. 28';-90.J-[Theorin, Om bladstendsgfandema has en del Salices,
Stockholm, 188l.]-Boergesen, Arkt. pI. bladbygn., Bot. Tidsskrift, xix, 1895, p. 219 et scq.-
SoAtari~, Ban des Stammes d. Salicin., Sitz.-Ber. Wiener Akad., cvii, Abt. i, 1898, pp.!I210-I9; and
Osterreich. bot. Zeitschr., 1899, p. 117.-Kiihne, Papillen u. oberseit. S~ltiiffn., Mitteil. deutsch.
dendrolog. Gesellsch., [899, p. 52.-Paule5co, Strnct. and. des hybrides, fhese, Geneve, 1900, p. <)0
(Sali.... ).-Tunmann, Sekrctdriisen, Dlss.,. Bern,- J900, pp. 14-17.-Petersen, Vedanatomi, 1901,
pp. ~4-7.-Pitard, Pericycle, These, Bordeaux, 1901 , pp. 94, 95.-Perredes, Compo anat. of the
barks of Salicin., Phann. lOllm., 1903; sep. copy from Wellcome Chern. Research Labor., n. 39.
34 pp., 6 pl.-Col, Faisceau.x, Ann. sc. nat., ser. 8, t. XX, 1904, p. l49.-Camus, Classi6cat. des
Sauies d'Europe et monogr. des Saules de France, Joum. de Bot., '9°4, p. 175 et seq., and 1905,

~
. [I] et seq.; especially 1904, pp. 184-:n3 (with anatomical diagnoses of the individual specles).-
Camus, Atlas de la monogr. des SauIes, Paris, '904, with eight anatomical platea.HPenhallow,
yst. Study of the Salic., Americ. Naturalist, xxxix, 1905, pp. 509-35 and 797-838 j abstr. in Just,
1905, ii, p. 13 etseq.J-PiCclOli, Legnami, Bull. Siena, 1906, p. 1 .... 3.-[Gaertner,Vergl. Blattanat. z.
SysL d. Gatt. Sa/ix. Diss., Giittingen, 1907, 59 pp.]

c... _ EMPETRACEAE (pp. 800, 801).

Literature :-Boergesell, Arkt. pl. bladbygn., Bot. Tidsskrift, xix, 1895, p. ll9 et seq.-Petenen.
VedaDatomi, 1901, pp. 46, 47.-Theorin, Viixltrichom., Arkiv for .Hot., i, 1903, p. 161.

CERATOPHYLLEAE (pp. 801-803).

The oily contents found in young stages in the shaggy structures consist
of myriophyllin; cells containing myriophyllin. moreover, are also found dis~
tributed through the tissues of the plant (Strasburger).
Literature: Weinrowsky Scheiteli;ifl'n. bei \Vasserptl., Dis!., Berlin, 1898, p. 22 (also in
FUnfstiiclc, Beitr., iii).-RoedIer, Assimilator. Gewebesyst., Diss., Freiburg i. d. Schw., 1898-9,
p. 37.--Strdbttrger, CerattJphylluIII aemermm, in Pringshcim Jahrb., xxx\'ii, 1902, p. 500 et seq.
and Tab. x, xi.
 CONCLUDING REMARKS
THE following pages are devoted to a review of those anatomical cha-
racters which have proved to be of taxonomic value, with reference to their
occurrence 1 in particular Orders, genera, or species. In the course of our
survey the varying systematic value, which, as experience has shown, attaches
to the individual anatomical characters, will become evident. These concluding
remarks may therefore serve both as an ai~ in the determination of a, plant by
means of its anatomical characters, and (in conjunction with the Introduction
on p. I et seq.) as a guide to the anatomical method.
In this review, however, it is not my intention to propOU':1d any definite
doctrine, nor to forestall in any way the results of free and independent re-
search. The facts and views presented in these pages are merely relative to
the present state of our knowledge and refer only to the plant-material hitherto
investigated, the precise nature of which is mentioned under the individual
Orders in the main part of this book; as the result of further investigations
these statements will often enough require amplification or restriction. Let
the guiding principle in systematic~anatomical investigations ,be invariably to
take all the ana.tomical features into' consideration and to test their systematic
value in each individual case.
The anatomical characters are discussed in the following seven chapters:-
1. Structure of the l~mina of the leaf; II. Structure of the petiole; III. Secre-
tory and excretory receptacles; IV. Hairy covering; V. Normal structure of
the axis; VI. Anomalous structure of the axis; VII. Structure of the root.

1. STRUCTURE OF THE LAMINA OF THE LEAF'!.

§ 1. SIMPLE EPIDERMIS. The epidermis of the leaf exhibits a large

number of structural variations, which for the most part constitute biological
characters and are generally only, of value for specific diagnosis.
These varia.tions are primarily connected with the shape of the cells in
surface.view. As a rule the epidermal cells, when seen from the surface, are
approximately isodiametric, the lateral walls ,being either straight or undulated.
It is chiefly the extreme types of structure shown by the lateral walls on any
one surface of the leaf that are of value for taxonomic purposes, and more
especially for specific diagnosis, whilst variation of these characters within
narrow limits (e.g. straight and slightly curved lateral walls, Of lateral walls
curved to a varying extent) occurs frequently enough in one and the same
species. Epidermal cells with curved lateral walls, moreover, are found
especially in species growing in damp habitats, while epidermal cells with

1 In reviewing the distribution of those anatomical features which are of frequent occurrence,
:l!;a rule only Order5 and anomalous genera are cited.
) As in the descriptions of the individual Orders (under • Structure of the Leaf') certain
:-.natomical characters of the axi.s. (such as spicular cells, &C')I whi.ch correspond to a.nalogous featuyes
in the leaf. are likewise taken into consideration.
For the nnatomical structure of subterranean leaves, we may refer to the paper by Thomas
(Anal. compo et exp. des feniiles sout., These, P!\ri~, IQOO, 106 pp.; see also Revue gen. de Bot.,
l~OOI p. 394 et seq.).
 STRUCTURE OF THE LAl.lIINA OF THE LEAF

straight :lateral walls are com'mon in species occupying a dry habitat; the
former type of cell in general occurs more frequently on the lower than on the
upper side of the leaf
The following special forms of epi~ermal cells require mention. In certain
species belonging to a large number of different Orders (e.g. Ranunculaceae,
Papilionaceae, Laurineae, Euphorbiaceae, &c.) one finds epidermal cells with their
lateral walls more or less bent in a zigzag fashion, while the apices of the angles
exhibit ridge-like projections, the ends of which are swollen in a nodose manncr
or in the shape of a T. much as in the petals of many plants. A local separation
of the lateral walls has been observed in species of Ranunculus. In narrow leaves
(Caryophylleae, Papilionaceae, Epacrideae, Polernoniaceae, &c.) elongated epidermal
cells are not uncommon, the cens in some cases {species of LathyYHs, Candollea}
being almost prosenchymatous. The long axis of these cells is usually parallel to
the median vem of the leaf, although occasionally (Sitene /yuticosa. species of Eutaxia
and Trifolium~ transverse to it.
The size 1 of the epidermal cells often varies very considerably in the three
directions of space. As a general rule, in the same species: the cells of the
upper epidermis are larger than those of the lower epidermis. The extreme
limits (i.e. very large or very small dimensions) in the size of the cells on
one and the same surface of the leaf are at least of specific value, while varia-
tions of this feature within narrower limits occur in individuals of the same
species, occupying different habitats (see Introduction, p. 9)' A considerable
development in the height of the epidermal cells is occasionally connected with
both extremes of size.
A small-celled epidermis, as seen in surface-view, has been described in certain
Capparideae. Vochysiaceae, Chlaenaceae, Malpighiaceae, Hippocrateaceae. Salva-
doraceae. Chloranthaceae, Myristicaceae and Buxaceae; a large·celled epidermis,
as regards all three dimensions, this feature being occasionally combined with a
special development in the height of the cells, is found in certam Menispermaceae,
Clstineae, Violarieae. Tremandreae, Elatineae, Sterculiaceae, Malpighiaceae,
Geraniaceat, Rosaceae. Bruniaceae, .Myrtaceae sens. str., Melastomaceae. Cucurbi-
taceae. Begoniaceae, Plurnbagineae, Styraceae, Apocynaceae, Asclepiadeae, Gesnera-
ceae, Verbenaceae and Illecebraceae; epidermal cells of considerable height. gene-
rally combined with a palisade-like shape of the cells, when seen in a transverse
section of the leaf, have been observed in certain Anonaceae, Guttiferae, Diptero-
carpeae, Rhaptopetalaceae, Malpighiaceae, I1icineae, Celastrineae, Hippocrateaceae,
Sapindaceae, Caesalpinieae, Chrysobalaneae, ..rSaxifragaceae, Hamarnelideae. Mela-
stomaceae. Cornaceae, Candolleaceae, Vacciniaceae, Ericaceae, Epacrideae, Primu-
laceae. Bignoniaceae, Illecebraceae, Laurineae, Thymelaeaceae, Santalaceae a"d
Buxaceae. In a few species of Candollea (Candolleaceae) tIie high epidermal cells,
which have a rhombic outline or assume a fibrous form, are placed obliquely to the
surface, and as a consequence the epidermis is • apparently many-layered' in a
transverse section of the leaf.
Further differences in the epidermis are connected with the structure
of the cuticle" and ~'aJ)s of the. epidermal cells, and with the excretion of wax
on the surface", -
The cuticle varies in thickness and, as seen in surface-view, is either smooth
or provided with granular or verrucose thickenings, or striated. The thickness
of the c~ticIe as well as the degree of marking on its surface may in extreme
cases be utilized, for syst~matic purposes; in other cases a certain amount of
discretion is necessary, i.e. these f~atures should not be employed until abundant
material of the species in question has been examined; the two features,

I The vnriations in the sizes of the cell:; on the corresponding organs of one and the !lD.lne genII";
GUflJura) are often inconsiderable, even when there are great difference!! in the dimemions o( the
organs in question; (or details, see Scbnegg lin Flor::l. xc, 1 !.10], p. 20U el seq.) 4nd Amelung (in
Vlor:)., Jfol93, p. l08 et 'Stf).).
 CONCLUDING REMARKS
moreover, 'are not always developed in the same way on the two surfaces of
the leaf. A more important character from the systematic point .of view is.
the.kind of marking on the cuticle. This applies particularly to the cuticular
ridges or crests which are often of considerable height and occur on the flat
y

surface of the epidermis (especially oh the lower'side of the leaf) 'in certain
Capparideae, Dipterocarpeae, .Malvaceae, Cyrilleae Mimoseae, JRosaceae,
y

Lythrarieae, Cacteae, Araliaceae, Vacciniaceae, Ericaceae, Sapotaceae (Fig. 117"

p. 513), Styraceae, ,Apocynaceae, Euphorbiaceae and .ulmaceae; .they are
frequently the cause of the dull appearance of the surface of the leaf. Similar·
high cuticular ridges occasionally unite the papillae developed from the epi.)
dermal cells of the leaf, such papillae being provided with cuticular outgrowths
in the form of a small crown or ,ridge (see the discussion of. the papillae below).,
~t rep:lains to consider: (a) the occun'enc~ of the' cuticular layers,' which are'
more or less cuticularized and apposed to the cuticle on its iriner sjde,· (b) the
penetration of the cuticle or of the cuticular layers into the lateral walls of the
epidermal cells in the form of lamellae or pegs, and (c) similarly, the penetration
of the cellulose· membrane 'into the cuticularized .portion of the outer wall in
the form of lamellae or pegs. The structural feature last named has been
recorded chiefly in certain Papilionaceae belonging to the tribes Podalyrieae
and Genisteae, as well as in certain Lythrarieae and Proteaceae, and commonly,
causes • false pitting or internal striation' of the outer walls of the epidermal
cells, ~hen seen from the surface (see pp. 896, 897, &c.).
The thickness of the entire outer wall· and of the lateral and inner
walls varies in the same way as does the thickness of the cuticle. It· is a
familiar· fact that strongly thickened wal,ls constitute a peculiar~ty of many
xerophilous species. Exceptionally.strong thickening of the inner walls of
the upper epidermal cells has been observed in certain Epacrideae and Euphor.,
biaceae j uniformly -sclerosed epidermal cells occur in 'certain Menisperma.
ceae, Capparideae, Ochnaceae, Sapindaceae, Melastomaceae, Candolleaceae
(fibrous), Epacrideae, Thymelaeaceae and species of Gonystylus, while in certain'
Caesalpinieae, Vacciniaceae and Epacrideae the epidermal cells have ,the
appearance of ston.e-cells only when seen, from the surface. The lateral and
inner walls, when strongly thickened, are for the most part pitted. A feature:
deserving quite special notice is the occurrence of distinct bordered pits side,
',by side wit~r simple pits on th.e lateral walls in certain species of Candollea
(Candolleac'eae). Pitting of the entire outer wall I 'is rare (Capparideae, Cela·
strineae~ Hippocrateaceae, Sapindaceae); peculiar long, branched pit·canals·
have been observed in the outer walls in the genus Mortonia (Celastrineae)~
A co~moner feature, found in species having undulated ~ateral walls, is th~
occurrence of so-called marginal pits, i.e. pits which traverse the outer walls,
but are' situated in the bays and often run obliquely towards the outside;
Lignified thickening bands have 1?een observed on the lateral walls in
Notouratea i·nun'data, Van Tieghem (Ochnaceae), spiral or reticulate thickening
of the epidermal cells in the Order Vacciniaceae, and reticulate thickening of the
outer walls in certain Ericaceae and Epacrideae., The exact chemical nature
of the inner land lateral walls (regarding gelatinization, see § 2, p~ 1074) may
likewise be taken into consideration in systematic-anatomical investigations.
In many species the protective action of the cuticle is augmented by
a considerable excreUon of wax, which gives t~ose particular parts of the plant
(both leaves and stems) a glaucous oj' hoary ai)pearance.
Such an excretion of wax is found in certain Magnoliaceae, Menispermaceae,

1 Ambronn, Poren in den AussenwandeI_l von Epidermiszellen. in Pringsheim Jahrb., xiv. ISS ••
p. 82' et seq. and Tab. viii. Regarding sensitive pits in the tendrils of Cucurbitaceae, &c .• see
lIa~rlandt. Physiolog. Ptianzenanatomie, 1904, ~. 509 ct ~eq.; also under Cllcurbitaceae. p. 939-
 STRUCTURE OF THE LAMINA OF lTHE LEAF 1073.,

Papaveraceae, Cruciferae, Violarieae. Bixineae, Tamariscineae, Malvaceae, Zygo-

phylleae, Aceraceae. Mimoseae, Rosaceae. Crassulaceae, Myrtaceae, Passifioraceae,
Cacteae, Ficoideae. Compositae, Ericaceae, Vacciniaceae. Epacrideae. Asclepiadeae,
Gentianeae, Solanaceae, Nyctagineae, Illecebraceae, Chenopodiaceae. Laurineae,
Proteaceae, Thyinelaeaceae, Euphorbiaceae and Salicineae. Concerning the varied
nature of -this excretioo, see under the Orders cited, and De. Bary, Vergl. Anat.,
p. 87 et seq.
A valuable specific character is afforded by the papillose differentiation
of the epidermal cells 1, This phenomenon is particularly common in the cells
of the lower epidermis; but 'rarer in those of the upper epidermis; it is occa-
sionally found on both, sides of the leaf in ol}e and the same species. The
differentiation of papillae generally causes the surface of the leaf to have a dull
appearance. In some cases the papillae are confined to the cells immediately
surrounding the stomata or to the margin of the leaf. The papillae vary in
the extent of their deVelopment and in their differentiation. One' finds all
stages, from'the formation of distinct papillae down to a simple convex protru-
sion of the outer walls on the one hand, and culminating in simple unicellular
hairs on the other. Epidermal cells, ill which the outer walls are strongly arched,
outwards, may be described as subpapillose. The development of a typical
papilla affects either the entire outer wall of the epidermal cell or merely its
central portion. The length and shape of the papillae vary. Their walls may be
either thin or'thick; the short (arid more rarely the long) papillae occasionally
develop into soHd pegs owing to secondary thickening of the cell-wall (in certain
Polygaleae, ROsaceae, Ebenaceae (Fig. lIB, p. 517), Proteaceae, Penaeaceae,
Geissoloma, &c.). In some cases also the papillae form the centres for a radial
striation of the epidermal celJs. Special mention may be made of the relatively
long and striatea papillae, which are provided with a small cuticular crown and
stand in connexion with 'one another by means of cuticular ridges, placed at
right angles to' the surface of the leaf (e.g. ih certain Anonaceae, Sapindaceae,
Papilionaceae,-Araliaceae, Comaceae, Ebenaceae, Fig. 118, p. 517, Styraceae,
Oleaceae). .
Papillose or subpapillose differentiation of the epidermal cells has been observed
in certain species belonging ,to the following Orders or genera: Ra"nunculaceae,
Magnoliaceae, Trochodendraceae, Lactoridaceae, .Anonaceae, Menispermaceae,
Berberideae, Nymphaeaceae, Papav{"raceae, Capparideae, Violarieae, Bixineae, Pitto..
sporeae, Polygaleae. Vochysiaceae. Caryophylleae, Tamariscineae, Hypericineae.
Guttiferae, Ternstroemiaceae, Lineae, Malpigbiaceae, Geraniaceae, Rutacea{",
Simambaceae, Ochnaceae, Burseraceae, Meliaceae. Olacineae, Celastrineae. Rham-
neae, Ampelidaceae, Sapindaceae, Bretsch:neiderll. Hippocastanaceae, Aceraceae,
Staphyleaceae, Anacardiaceae. Connaraceae, Papilionaceae, Caesalpinieae, Mimoseae,
Rosaceae, Crassulaceae, Hamamelideae, Bruniaceae, Halorageae, Combretaceae.
Myrtaceae sens. str., Lecythidaceae, Melastomaceae, Lythrarieae, Passifioraceae.
Cucurbitaceae. Begoniaceae, Ficoideae, Umbelliferae. Araliaceae, Comaceae, Capri-
foliaceae, R u biaceae. Compositae, Cam pan ulaceae (only isolated papillae). Ericaceae.
Epacrideae, Diapensiaceae, Ebeo<l:ceae (Fig. 118). Styraceae, Oleaceae, Apocynaceae;
Loganiac-eae, Polemoniaceae (only isolated papillae), Boragineae, Convolvulaceae,
Solanaceae;- Z ombiana, Gesneraceae, Bignoniaceae, Acan thaceae, Aristolochiaceae,
Chloranthaceae, Piperaceae, Myristicaceae, Laurineae, Hernandiaceae, Proteaceae,
Thymelaeaceae, Penaeaceae, Gez'ssoloma, Santalaceae, Euphorbiaceae, Daphniphyl-
laceae, Ulmaceae, Moraceae, Thelygoneae, Myricaceae, Cupuliferae. For certain
epidermal cells of the leaf, which are developed like bladders or hairs, see also
§§ 2 and 31.
A feature, which can generally likewise be employed only for specific
diagnosis, is afforded by the occurrence of thin horizontal or vertical

1 Kohne, Vorkommen von Papillen u. oberseit. Spaltoffn., Mitteil. deutsch. dendrolog. Gesellsch.,
18 99, pp. 47-67 j see also Kohne, ,Frl7xi,lus·Arler,t, in Regel, G:r.rtenflora, 1899, pp. :184-8.
SO Lit R RrWp. 3 Z
 J074 CONCLUDING REAIARKS

divisIon-walls in the epidermal cells in certain Cehistrineae, Sapindaceae,

Hippocastanaceae, Papilionaceae, Caesalpinieae, Mimoseae, Araliaceae, Loga-
niaceae and Moraceae; see also § 3, p. 1076.
Of special contents found in the epidermal cells we mayin the first place
mention the presence of chlorophyll 1, which occurs chiefly in species growing
in shady or damp localities.
The species in <l!lestion belong to the following Orders: Ranunculaceae, Gera-
niaceae {Impatiens, Tropaeolum}, llicineae, Staphyleaceae (Staphylea), Droseraceae,
Onagrarieae (Epz'lobium), Diapensiaceae, Solanaceae (Datura § Brugmanst'a), Sero-
phularineae, Lentibularieae, Pedalineae, Selagineae, Verbenaceae. Labiatae. Phyto-
laccaceae, Podostemaceae, Euphorbiaeeae (Mercurialis). Thelygoneae, Cerato-
phyneae.
The occurrence of anthocyanin (widely distributed in the Begoniaceae and
Gesrieraceae) and tannin in the epidermis also requires to be taken into account.
For the occurrence of oxalate of lime, see § 2; for sphaerocrystalline masses
and other contents composed of organic substances, see §§ 23 and 25; for
mucilage, see § 2; for other kinds of secretions, see § 14; and for cystolit~
see § 28.
Regarding incrust~t~on of the walls of the epidermal cells, see §§ 26 and 27.
§ 2. DIFFERENTIATION OF THE SIMPLE EPIDERMIS OF THE LEAF. In this
section we shall discuss especially the differentiation of the epidermis of the
leaf in so far as it is brought about by the occurrence of mucilaginous epidermal
cells, of cells containing oxalate of lime, and of large epidermal cells, which
either serve for the storage of water or are ,filled with brown contents in the dry
leaf. Regarding epidermal idioblasts, which contain other kinds of secretion
or are occupied by cystoliths, see §§ 14 and 28 respectively.
1. The gelatinization of the epidermis of the leaf 2, which serves for the
storage of water, but occasionally occurs also in species from damp habitats,
is a good specific character. Gelatinization usually affects only the inner walls
of the epidermal cells, rarely the outer walls as well, and as a general rule is found
more commonly on the upper than on the lower side of the leaf. Gelatinized
epidermal cells are often distinguished by their large size; in such cases the
mucilaginous portions of the inner walls, which are separated from the lumina-of
the cells by a cellulose-lamella and occasionally include further internal cellulose-
lamellae, commonly penetrate into the mesoph yll in a hemispherical or conical
form, sometimes giving rise to transparent dots in the leaf. As a rule only
isolated cf'lls of the epidermis are mucilaginous, but occasionally there are groups
of gelatinized cells of varying size or the entire epidermis is affected. In the cases
last mentioned a regular layer of mucilage is produced below the upper
epidermis, in the formation of which the adjacent walls of the subepidermal
layer of cells may also playa part (in certain Menispermaceae, Rutaceae,
Fig. 40, p. 175. Loganiaceae. Gentianeae and Euphorbiaceae; in the Gentianeae
also beneath the epidermis of the stem).
CeUs with gelatinized innet' membranes have been observed in the integu.
mental tissue (epidermis and hypoderm) in species of the following Orders or genera.
as the case may be ~ Magnoliaceae. Anonaceae. Cruciferae. Resedaceae, Cistineae,

I Stohr. ill Sitz.-Ber. Wiener Akad., lxxix, Abt. J, 1879; and De Bary, Vergl. Anat., 1877.
p. 7~. Radlkofer, Monogr. Sujant"a. 18 75, p. 100: Walliczek, Membmnscbleime, in Pringshtim
Jahrb., xxv, J893. p. 209; Kruch, Epid. mucH., Ann. R. 1st. bot. di Roma, vi, 1896, 86 pp. and
a T ..b. .
It may be well at this point to warn the reader against the confusion between gelatinized
portions of the walls 01 the epioermis and the cells of a hypoderm,-an error which oCCUrS over and
over again in ,he literatnre.
 STRUCTURE OF THE LAir/INA OF THE LEAF )075
Violarieae, Bixineae, Tremand reae, Vochysiaceae, Elatineae, Ternstroemiaceae,
Dipterocarpeae, Monotes, Chlaenaceae, Malvaceae, Sterculiaceae, Tiliaceae, Lineae,
~alpighiaceae, Rutaceae, Simarubaceae, Ochnaceae, Luxemburgiaceae, Bursera-
ceae. Meliaceae, Chailletiaceae, Olacineae, Ilicineae, CyriUeae, Celastrineae, Penta·
phylacaceae, Rhamneae, Sapindaceae, Aceraceae, Staphyleaceae, Moringeae, Con-
naraceae, Papilionaceae, Caesalpinieae, Mimoseae, Rosaceae, Saxifragaceae, Hama-
melideae, ,Rhizophoraceae, Melastomaceae, Lythrarieae, Onagtarieae, Samydaceae
(only in Gerrardlna), Turneraceae, Passifioraceae, Comaceac, Ericaceae, Myrsineae,
Sapotaceae, Gentianeae, Phytolaccaceae, Polygonaceae. Thymelaeaceae (Fig. 174,
p. 717), Gonyslylus, Ge~'ss()/oma, Euphorbiaceae, UJmaceae, Moracea~, Cupuliferae,
Salicineae, Empetraceae.
Epidermal cells, in which the inner walls are swollen (e.g. in certain Samydaceae),
must not be confused with gelatinized epidermal cells, even though in some respects
they constitute a transition to the latter.
II. Epidermal cells containing oxalate of lime are rarely the cause of a
differentiation of the epidermis of the leaf, this being the case only when the
crystal-cells are distinguished from the remaining epidermal cells by their
larger or smaller size and by their shape. The small crystal.jdioblasts mostly
appear round, when seen from the surface, and contain either a clustered or
a solitary crystal; they are rarely isolated, being more commonly paired or
united to form small groups; in the Papilionaceous genus Stylosanthes they
have a polygonal outline and enclose a single rod-shaped or geniculate solitary
crystal, the entire superficial layer of the integumental tissue on .the lower side
of the leaf (except for the stomata and their subsidiary cells) in this case being
composed of such crystal-cells (Fig. 58, B, p. 265).
The large crystal-idioblasts, which in the' Schlussbemerkungen ' of the
original German edition of this book (p. g08) were mentioned as occurring in
certain Rutaceae and Euphorhiaceae, and are perhaps found in the same
position also in other Orders, urgently require an investigation of their mode
of development, in order to determine whether they really, or only apparently,
belong to the epidermis.
The well-known crystal-cells of Citrus, which are wedged in betWeen the
epidermal cells on the two surfaces of the leaf, are subepidermal cells,iwhich have
penetrated into the epidermis by a process of sliding growth. This is likewise true
of (a) the large cells with clustered crystals, found in Dalechampia Roezliana, and
probab1yalso the cells with clustered crysta1s in other Acalypheae; (b} the cells
with clustered crystals, which occur in Capeyonia and Af'cyyothamn~'a, the cells in
this case already projecting somewhat- after the manner of a hair; (c) the cells
containing sphaerites, which are found in AcalYPha and Claoxylon. the cells in
question likewise projecting beyond the surface; and (d) the hairs with clustered
crystals, recorded in Fragariopsis and Plukenetia (d. pp. 1049, 105 I, and § 3 1 ) 1.
In the following synopsis all the forms, in which oxalate of 'lime occurs
in the epidermis of the leaf, are taken into consideration, whether the cells
containing the crystals appear as idioblasts or not. The Orders, in which
relatively.small_ q-ystal-jdioblasts have been observed, are provided with a •
preceding the mime of the Order; cr. indicates ordinary solitary crystals;
clust. cr. = clustered crystals j S = crystal-sand; R = raphides; A = acicular
crystals; p = small prismatic or rod-shaped crystals.
Oxalate of lime. has been observed in the epidermis in the following Orders :
Magnoliaceae? (cr.). Anonaceae (cr., dust. cr.), Fig. 6. A (p. 36). *Meni,spermC!Ceae
(A, p. cr.), Capparideae (p). *CaneHaceae (cr., dust. cr.), *Bixineae (cr.• dust. cr.},
Guttiferae (dust. cr., pl. Fig. 27, C (p. 122), *Temstroemiaceae (cr., dust. cr.), Tilia-
ceae (dust. cr.). Lineae (cr.), Rutaceae (cr.). Simarubaceae (p), Olacineae (cr., clust.

Guttenberg, KrystalluUen im Blatt von Citrus, Sitz.-lJer. Wiener Akad., exi, Abt. I, 1903,
18 pp., 1 Tab.; Knoll, Brenohaare der Euphorbiaceen-Gattungen .lJaledzampia und Tragi4, Siu.-
Ber. Wiener Akad.) cxiv, Abt. " 1905. 20 pp., 2 Tnb. -
'\ Z 2_
 CONCLUDING REIJARKS

cr.}, *Celastrineac (cr., clust. cr.), ·Hippocrateaceae (cr., clust. cr.), ·Rhamneae (cr.),
Sapindaceae (cr., clust. cr., s), Didiereae (clust. cr.), Aceraceae (cr.), ·Papilionaceae
(cr.). Fig. 58, B (p; 265), ·Caesalpinieae (cr., clust. cr.). Mimoseae (cr.), Rhizophora-
ceae (cr., clust. cr.), Lecythidaceae, Melastomaceae (clust. cr.), Lythrarieae ,(cr.),
·Samydaceae (cr., clust. cr.), Araliaceae (cr., clust. cr., p), Cornaceae (clust. cr.).
Rubiaceae (R), Compositae (A). Epa~rideae (cr.), Myrsineae (clust. cr., A, p),
Sapotaceae (cr., dnst. cr.), Styraceae (cr., clust. cr.), Oleaceae (A, p), Apocynaceae
(cr.), Polemoniaceae (A), Solanaceae (s), Pedalineae, Acanthaceae (A), Myoporineae
(clust., cr., A), Selagineae (p), *Piperaceae (Symb1'yon. clust. cr:), Chloranthaceae
(small cr.), Laurlneae (small cr.), Hernandiaceae (A, p), Proteaceae (cr.). Elaeagna-
ceae CA), Santalaceae (cr.), *Euphorbiaceae (cr., dust. cr.). *Daphniphyllaceae
(clust. cr.), *Moraceae (cr., clust. cr., small cr.).
In certain Capparideae small crystals of gypsum are .found in the epidermis
of the leaf; these crystals can readily be distinguished from those of oxalate of
lime by their chemical reactions (cf. § 25).
The mode of excretion of oxalate ot" lime in the epidermis is of varying
systematic value. Special crystal-idioblasts are characteristic of species or
genera. Among the Anonaceae oxalate of lime (either in the form of isolated
or clustered crystals) is found in the epidermis in most of the species and there-
fore furnishes a useful character for the recognition' of -the Order. The excre-
tion. of oxalate of lime in the form of acicular crystals .or of small prismatic
or variously shaped crystals is, as experience has shown, of trifling systematic
value..
HI. In the third case a differentiation of the epidermis is. brought about
by the presence of epidermal cells with wide lumina among cells ,of the
ordihary size; the large cells either store up water solely or are filled with
~peci41 contents, which for the most part have a brown colour in the dry leaf.
9ucll large epidermal cells, serving the purpose of water-storage, commonly
form rows or are even arranged in a reticulate manner (Cruciferae, Fig. 14, p. 59);
in certain Orders they are differentiated like bladders or hairs and occasionally
give the living plant the appearance of being covered with drops of dew.
Epidermal cells, which have large lumina and store up water, but are not
vesicular, are found among cells of the normal size in certain Cruciferae. Resedaceae,
Elatineae and Malpighiaceae; vesicular epidermal cells (see also § 3 I) occur in
certain Resedaceae. Caryophylleae, Portulaceae, Crassulaceae (Fig. 70, p. 321) and
Ficoideae (Fig. 94, p. 416); tubular epidermal cells. filled with brown contents
(see also § 14), are present in certain Violarieae, Geraniaceae, Saxifragaceae. Crassu-
laceae and Euphorbiaceae; elongated epidermal cells, described as mucilage-cells
(see § 14), are found in Tyopaeolum (Geraniaceae).
The following anatomical features remain to be mentioned at this point:
the special epidermal cells, found in A namirta and A rcangelisia (Menispermaceae,
Fig. 7, B, p. 4 I ) and in Gonocaryum (Olacineae, Fig. 48, p. 204), these cells
functioning as hydathodes; the epidermis of the leaf of Phyllachne (Can-
do])eaceae), the median and marginal portions of which are composed of
prosenchymatous cells; and the peculiar groups of cells in th~ lower epidermis
of species of Limnanthemum and Villarsia (Gentianeae).
§ 3. HYPODERM 1. The water-storing epidermis is very frequently
(especially on the upper side of the leaf) supplemented by a hypoderm, which
likewise serves for the storage of water. A hypoderm may originate in two
ways, viz. in some cases from the dermatogen, in other cases from the ground
tissue. Our discussion should really be confined here to the hypoderm
which is developed from the integumental tissue; such a hypoderm, together
with the epidermis, is styled a 'many-layeredepidennis 'by Pfitzer, who reserves
the term 'hypoderm' for the aqueous tissue arising from the mesophyll.

1 See Pfitler, Hautgewebe, in Pringsheim Jahrb •• viii, 187:1, p. 16 and Tab. vi.
 STRUCTURE OF THE LAMINA OF THE LEAF 1077

Since, however, the source of the hypoderm' is ·usually' not'recognizable in the

fully develope,d' ]e':if· (as is' 'shown, for example, by th,e close·~greeme~t..b'etween
the hypoderm of 7lex Aque'joHum, whiCh)s formed from the 'mesophyll,
and, that :0£ Ficus .ela~tica,' which~ is 'developed 'from. 'the epidermis),' both
kinds of hypoderm ·will be considered together here. The, .derivation ~of the
,bypoderIl}. from t~e epidermis cannot be certainly established.without a develop-
menta!', investigation, except· in those, cases in· which the·lateral_"walls. of :the
epidermal and hypodermal cells corresp'ond with oneanotlier'even in the mature
leaf.le.g. in certain. ~nner~ce~e and. Piperaceae, .which ha~e a typical-two. or
many-layered epi9.ermis),. .
The extent ofdev:elopment .. of the hYP9derm, yaries. In the, simplest
case there !S, merely ,a .tendency .!ow~rcl~ the. torm,ation of a hypoderm, ·as
shown by·the appearance of division-walls, parallel tq the surface of theJeaf,
in a varying .f:l~~b~r of the epidermal cells. When ,a p'~r~n~!lymato~ hypo·
derm of one or more layers is developed, its ~el1s o~ten _contrast with the epi-
,dermal cells by their ~onsiderable. size... Hypodermal cells of: the, parenchyma·
tous type have a polygonal or undulated outline in surface-view.and walls, of
varying t~~c~ess; in those c~es in which the Ja~ter_ are deJi~ate, a loss of watet
results in a "bellows-like folding ,of t~~ ,lateF~. walls; i,n o~her. cases the hypo-
dermll ce~,s are :prismati~, .the e~g~~, whi,ch ~r~ placed at right angles to the
surface of the leaf, being relatively strongly thIckened. , All these differences
.in ,the char3:cter qf the ~ypodermal cells may p,~ ~w-pl<?y~9._for systematic
purpps~ (~f. ,the Jn~roducti~n).. Th~.sam,~ applies to ~he.,developmc.nt of the
h ypod~rm on. b.'?th sld~~ Of, onl y on th~ uppe~,.or }O\yer sid~. of the le.ai. Fe'ltures
<

deservmg speCIal mentIon are: the dlfferenflatlo1':1,'of a collenchymatoUS hypo·

derm in the axis 9,f ~,ert~in Cacteae (Fig ..90, p. 4°7); the p'r~senc_e o.f a )1ypode.nn,
developed. like. spongy' tissue, in the l~!'!-t' ,qf Oedematopus: oboValus. l'I~ et Pl
(~uttifera.e~ Fig. 27, B; .p.; .122)'; and the occurrence of(a sc1ero.ti.~:hYp'oderm,
composed of fibrous gr r04~shaped cells and occasionally con:lle~ted with the
sclerenchyma of the veins, in the' leaves of cert~in spe~ies of Berberis and
Mahonia (B_erberideae), ,Ochrocarpu.~ (Gu$tiferae), . Elyasioideae, Van Tiegh.
.(Ochnaceae, the fibrous cells j.n th!s case oell'!g arranged transversely), Eu:Phoria
and Litchi (Sapindaceae), Connarus. (Connaraceae),. Cephatostigma, Light/ootia,
.J.l,.f erciera, Mi.c:ro~on, PrismatocarpHs, .Roella. .and W. (fhle1Jbergia (Campanulaceae),
Theophrasteae (her~ in almost, all tp.~ ,sp~cies)' and TVeigeltia (Myrsineae)
Persea t~~l!rin~ae)" Dryattd.rf;l (proteaceae), and' Artocarpus (Moraceae).
The appearance, of horizont'al walls in tlie 'epidermal celis, which leads to the
differentiation of a tWo- or· many-layered epiderinis in the Orders marked with a .,
has been observed in certain *Anonaceae, Menispermaceae,- Bixineaet *Pittosporeae,
Vochysiaceae, 'Malvacea.e., .Rhaptopetalaceae, Rutaceae" Ilicineae, *Celastrineae,
*Rhamneae. Sapindaceae, Anacardiaceae ? *Connaraceae, Saxifragaceae. "Crassu-
laceae, Rhizophorac'eae, MelaSt6iriaceae, Lythrarie!le,' Cacteae (axis), Araliaceae,
Cornaceae, Ru biaceae. Ericaceae~Epac?~eae, ~yrs~neae,. SaI?otaceae, Salva?-o~aceae.
A pocynaceae, .~onvolyul?-ceae~ .~lgnQmaceae; Selagmeae, *Piperaceae, Momrmaceae,
Hernandiaceae, Moraceae, *Santalaceae.
Hypoderm is found in certain species of the following Orders or genera: Dille-
niaceae, Magnoliaceae, Anonaceae, Menispermaceae, Berberideae, Capparideae,
Violarieae,( Canellaceae, Bixineae, Pittosporeae, Polygaleae, Vochysiaceae. Caryo-
phylleae (observed only on the'lower·side),-.Portulaceae (LetlZia, ,only'qn the~lower
side), H ypericineae, Guttiferae, Terns,troemiaceae, Stl'asbuyge'ria~ A n.c_£styocladu5, Lo·
phira, Monotes, Chlaenaceae', Malvaceae, Sterculiad~ae, Ti1iace3:~, Lineae; Malpigma-
ceae, Gera'niaceae, Rutaceae, Simarubaceae, Ochnaceae, Burseraceae, l\~eliaceae,
Chai11etiaceae, Olacineae, Ilicineae~ Celastrineae, Hippocrateaceae:, CorYl!ocarpaceae,
Rhamneae, Sapindaceae, Didiereae, Anacardiaceae; CoImaraceae, Papilionaceae,
Caesalpini~ae. ,Rosact:.ae, Saxifragaceae, _~~mam~li~eae, Rhizophora:ea~. Combre-
taceae, Myrtaceae sens. str., Lecythidaceae, Melastomaceae, .Lythrane~e, Sam:yda-
ceae, Passifioraceae, Cucurbitaceae, .Begoniaceae, :Datisc~~e, .Cacteae' (axIS I),
 CONCLUDING REMARKS
Umbelliferc1.C, Araliaceae, Cornaceae, Rubiaceae, Dipsqceae, Compositae, Campanu-
laceae, Vacciniaceae, Ericaceae. Epacrideae, Myrsineae, Sa,potaceae, EbeDaceae~
Styraceae, Oleaceae, Salvadoraceae, Apocynaceae, Asclept.adeae, Loganiaceae.,
Scrophularineae, Columellia.ceae, Gesneraceae. Bignoniaceae. Acanthaceae, Verbena-
ceae, Labiatae, Nyctagineae, Amarantaceae, Phytolaccaceae, Polygonaceae, Nepen-
thaceae, Aristolochiaceae, Piperaceae (Fig. 168, p. 689), Cbloranthaceae, Myristica-
ceae, Monimiaceae, :cau.neae, Hernandiaceae, Gomortegaceae, Proteaceae,
Elaeagnaceae, Loranthaceae. Santalaceae. Euphorbiaceae, Balanopseae, Ulmaceae,.
Moraceae, Urticeae, Leitnerieae, Myricaceae, Cupuliferae, Salicineae.
§ 4. STOMATA 1. Among the manifold structural differences, which accom-
pany the appearance of stomata, the mode of attachment of the epidermal cells
surrounding the stomata to the pairs of guard-celis has proved to be of the
greatest systematic importance. This feature is intimately connected with the
course of development of the stomata from the cells of the dermatogen.
As regards the mode of development we may, in agreement with Prantl,
especially distinguish the following cases:
I. The mother-cell of the pair of guard-cells i~ formed by the first division~
wall 2.
(a) The stomatal apparatus is completed, as soon as the pair of guard-
cells is differentiated (Ranunculaceous type).
(b) In other cases accessory subsidiary cells 'are formed by secondary
divisions in the surrounding cells.
(II) These divisions are confined to two neighbouring cells, situated
at the sides of the mother· cell of the guard· cells (false Rubia-
ceous type; in many Monocotyledons, certain Portulaceae
and Proteaceae), or
(P) they take place in four or 'more of the neighbouring cells (in
Coniferae, Cycadeae, Monocotyledons, but also in Ficus).
II. The formation of the mother·cell of the guard-cells is preceded by the
development of cells, subsidiary to the pair of guard-cells. The number of
preparatory divisions varies.
(a) The preparatory divisions take place in two directions.
(~) The diVlsion-wall in the mother-cell of the guard-cells runs
parallel to the walls, formed during the preparatory divisions.
In this case the pairs of guard-cells are accompanied on
either side (i.e. to the right and left) by one or more subsidiary
cells t which are placed paraliel to the pore (Rubiaceous type);
the subsidiary cells in stomata of this type are either merely
arranged parillel to the guard-cells (and also to one another,
when more than two subsidiary cells are present), or partly
surround one another; if, in the latter case, there are several
subsidiary cells, a right-hand or left-hand subsidiary cell,
as seen in surface-view, is alternately in part enveloped by
one situated to'the left or to the right respectively

I Strasburger, in Pringsheim Jahrb., v, 1866-7, p. 297 and Tab. xxxv-xlii; Prantl, in Flora,
187a, p. 30 5 and Tab. vi; De Bary, Vergl. Anat., 1877. p. 42; Immich, in Flora, 188 7, p. 435;
Vesque, in !lull. Soc. bot. de France, 1889, p. lxiii; Benecke, in Bot. Zeit., 1892, p. 5:31 and
Tab. viii; [Tognini, in Atti 1st. bot. Pavia, set. ii. vol. iv, 1894,.P pp., 3 Tab.; abstr. in Just, 1894.
p. 466, and Bot. Centralbl., Beih., 1894, p. 423] ; W estemlaier, in Festschrift flir Schwendener, 1899.
p. 63; Porscb, Der Spaltoffnungsapparat im Lichte der Phylogenie, lena, 1905.
The term ' stomata • is used in this book to mean the same as ' pair of guard-cells.' When the
epidermal cells sOrr1)uoding the stomata are distinguished (rom the remaining cells of the epidermis
by having a special shape or by their arrangement, they are spoken of as 8ubsidiary cells, but in all
other cases as neighbouring cells. .
I In the following synopsis I have not includ~ the «!aSe, in wbich the mother-cell of the pair of
guard~lls arises directly (rom a cell of the dennatogen (regarding this point, lee Tognini, loco cit.) ;
tbis has been recorded by Briosi and Tognini, but occurs very rarely.
 STRUCTURE OF THE LAMINA OF THE LEAF 1079

(f3) The division-wall in the mother-cell of the guard-cells is placed

at ri~ht angles to the walls, formed during the preparatory
divisIons. The guard-cells are then accompanied by two or
more subsidiary cells, which are arranged transversely with
reference to the pore (CaryophyUeous type). The mode of
att~chment of the subsidiary celli v4tries in the same way
as IDa.
(b) The preparatory divisions take place in three directions. This
results in a stomatal apparatus, in which the guard-cells are
surrounded by one or more rings, each consisting of three sub-
sidiary cells (Cruciferous type).
The types just mentioned are occasionally characteristic of certain Orders,
as indicated already by the names given to them by Vesque, while in other cases
they are constant only for a genus or a group of species, or merely for a single
species. Species are, moreover, known, in which two different types of stomata
occur side by side on one and the same surface-section (see Selagineae, Verbe-
naceae, Labiatae, &c.). The combination of two types in the same stomatal
apparatus is very rare (Silvianthu-s, Caprifoliaceae). Lastly, in employing
these types for systematic purposes, it is important to bear in mind that,
according to Tognini's investigations, the development of the stomata on the
various organs (e.g. foliage-leaf, cotyledon, petal, stem) of the same plant~
species may either be identical or may vary.
The utilization of the different types of stomata for systematic pur-
poses is involved in great difficulties in practice, since these types can be
determined with certainty only by a study of the course of development of the
stomata, and are very commonly altogether unrecognizable in the mature leaf. In
a fully grown leaf the true Rubiaceous type is often indistinguishable from the false
one, and similarly the Ranunculaceous type can frequently not be distinguished
from the Cruciferous type; a stomatal apparatus of the Ranunculaceous type,
in which the edges of the V-shaped division-wall touch the common wall of
contact between two epidermal cells, looks like a stomatal apparatus of the
Caryophylleous type (false Caryophylleous type); a stomatal appara~tis of the
Caryophylleous type, in which the breadth of the pair of guard-cells is equivalent
to that of the mother~cell of the whole stomatal apparatus, or in which the
guard-cells have come to lie next to one of the longitUdinal walls of the mother..
cell, shows more than two cells in the immediate neighbourhood of the pair of
guard-cells, and so on. In the following review it is therefore not possible to
distinguish more than the following types: (I) Stomata without subsidiary
cells, and having three or more neighbouring cells, arranged as in the Cruciferous
or Ranuncu1aceous types; (2) Stomata with subsidiary cells, which are placed
parallel to the pore (true and false Rubiaceous types); (3) Stomata with sub-
sidiary cells lying transversely to the pore (true and false Caryophylleous types);
(4) Stomata ill which the Cruciferous type is prominent in the mature leaf.
The discrimination of the types of stomata themselves is a task, which is
far from being completed; its solution, even though of no practical importance
in the determination of plants, will nevertheless be of great value in connexion
with the natural system of classification.
I. Stomata having no subsidiary cells and conforming to the Cruciferous or
Ranunculaceous types 1 are found in: Ranunculaceae, Dilleniaceae pro parte,
Trochodendraceae, Menispermaceae pro parte, Berberideae (subsidiary cells arranged
in the form of a rosette in Berbet'idopsis), Nymphaeaceae, Sarraceniaceae, Papa-
veraceae, Fumariaceae, Cruciferae pro parte, Capparideae, Resedaceae, Cistineae,

1 Under I I have also taken into consideration the few cases, in which more than three subsidiary
cells are knowD to occur in contact with the pair of glUud·cells.
 'CONCLUDING RE.MARKS

Violarieae pro parte, Bixineae. pro parte, Tremandreae, :Polygaleae pro parte,
V~chysiaceae pro. -p~rte,. Frankeniaceae, ~aryophyneae., 'pr,~ parte, Portulaceae
pro parte" Tamanscmeae pro parte, Elatmeae, Hypencmeae pro parte, Tern-
stroemiaceae pro 'parte (sometimes 'with subsidiary cells), Strasburgeria, Diptero-
carpeae (in'inost cases),' A nci"stroctadus, La phira, Mimotes, Chlaenaceae, Malvaceae,
Triplochitonaceae, Sterculiaceae; Tiliaceae; Humiriaceae pro parte, Zygophylleae,
Geraniaceae pro parte,J Rutaceae (in most cases, also,subsid. cells in a rosette),
Simarubaceae (in most cases). Burseraceae, Meliaceae, OIacineae pro parte,
Octocnemaceae, ,Ilicineae, Cyrilleae, ,Celastrineae pro parte (also subsid. cells in a
rosette), Hippocrateaceae, Stackhousieae, 'Rh;~.mneae pro parte; Am pelidaceae,'Sapin-
daceae (in most" cases), ,Hi ppocastanaceae, Aceraceae, Melianthaceae, Staphy leaceae.
Sabiaceae, Anacardiaceae, Moringeae, Connaraceae pro parte,'Papilioriaceae pro parte
(also subsid. cells in a rosette), Caesalpinieae prorparte,:Rosaceae pro parte, Crosso-
somataceae, Saxifragaceae pro parte, Crassulaceae (rare), Droseraceae (excl. 'Bybl£s),
Myrothamnus, Bruniaceae, Halorageae, Rhizophoraceae, Combretaceae, Myrtaceae
sens. str., Lecythidaceae,pro parte, Melastomaceae pro p~rte, Lythrarieae pro parte
(also, subsid. cells,in a rosette), Onagrarieae, Samydaceae pro parte, Loaseae,
Turneraceae pro .. paite~ Passifioraceae1 Pa:payaceae,' Cucurbi hlceae, Datisceae,
Ficoideae pro parte, U mbelliferae pro parte, Araliaceae 'pro parte',- Cornaceae; Capri-
foliaceae (excl. S#vianthus), Valerianeae; Dipsaceae pro parte, Calycereae, Compo~
sitae, Candolleaceae pro parte, Goodeniaceae pro parte, Campanulaceae' incl:
Lobeliaceae (excl. Pentaphragma), Ericaceae (in most cases), Epacrideae, Diapensia-
ceae, Lennoaceae, Plumbagineae pr9 parte, 'f:.;imula~ce;:te, Myrsineae, Sapotace~e pro
parte, Ebenaceae, Styraceae pro parte, Oleaceae (in most cases). Salvadoraceae pro
parte, Asclepiadeae pro' parte, Loganiaceae 'pro parte, Desjontainea, .E~tocosperma.
Gentianeae, Polemoniaceae, HydrophyUaceae, Boragineae pro parte, Convolvu-
laceae' pro 'parte, Solanaceae 'pro parte. Lonchostoma, Scrophularineae, Orobancha-
ceae, Lentibularieae pro parte, ColumelIiaceae, Gesneraceae pro parte, Bignoniaceae
pro parte, .Pedalineae pro 'parte, Myoporineae pro parte, Zombiana, Selagineae
(on the. same surface-section so~etimes, also subsid. cells" placed parallel' or trans-
versely to the pore), Verbenaceae pro parte, Labiatae ,pro parte, Plantagineae pro
parte, ~yc~aginea~ pro parte,Il~ecebraceae 'pro parte,',Amarantaceae, Chenopo~ia­
ceae pro parte, Phytolaccaceae pro parte, Polygonaceae pro parte~ Nepenthaceae,
Aristolochiaceae, Piperaceae 'pro parte; Chloranthaceae pro 'parte (sometimes :with'
subsid. cells arranged in a rosette, or with one or other subsidiary cell parallel to
the pore), Monimiaceae pro parte, Hernanaiaceae pro parte, Thymelaeaceae (papillose
subsid~ cells" arranged .like a, rosette,. in .certain,. species of_ Daphne), ,Penaeaceae,
Elaeagnaceae, Santalaceae (sometimes side by side with stomata ~ith parallel subsid.
cells), Myzodendron, Grubbia, Euphorbiaceae pro parte, Buxaceae (also subsid. 'cells
arranged in a rosette), Balanopseae, Ulmaceae pro parte,' Canmibineae, Moraceae pro
parte (rarely subsid. cells in'a rosette), Urticeae pro parte, 'Platanaceae; 'JugHmdeac;
l\Iyricaceae, Cupulifeiae~' Lacistemaceae, Empetraceae. .
2: Stomata with subsidiary cells, lying parallel to the pore; occur in: Dillenia*
ceae pro parte, Calycanthaceae (Fig. 3. B, p. 25), Magnoliaceae (occasionally indis~inct.
among the Schizandreae), ,Anonaceae, Menispermaceae (rar~),~Violar;ieae pro parte
(often obscured), ~anellaceae (partly obscured), 'Bi-'4n~ae pro parte, ~~,tosporeae,
Polygaleae:pro parte, Vochysiaceae pro p,arte, Portulaceae (almost .of genera[ occur-
rence,true and false Rubiaceous types;flg. 26, E, p. 'I 12), Tarriariscineae pro,parte (?),
HyPeril?i~ea~ pro 'parte, Guttifer<l;e ~Fig. 27, C; P: 122), Ternstroem~ac~ae.p'ro parte,
Dlpterocarpeae pro :parte (rare), "Lmeae; Hunnnaceae 'pro parte (mdlstinct), 'Mal-,
pighiaceae. Geraniaceae pro parte, Rutaceae pro ,parte' (rare}; Simarubacdle (riire),
Och naceae (always ?). Chailletiaceae (Fig. 46, p. 198), Olacineae pro parte, Celastrineae
pro partel .Corynocarpaceae, Rhamneae pro parte; Sapindaceae pro parte (yery rare).
Coriarieae, Connaraceae pro parte, Papilionaceae pro parte, Caesalpinieae pro parte,
Mimoseae, Rosaceae pro parte (ChrysobaIaneae), Saxifragaceae pro parte~ B-ybUs.
Hamamelideae, Ostrearia, Myrtaceae sens.' str. pro parte" Lecythidaceae pro, parte,
Melastomaceae pro parte, Samydaceae pro parte, Turneraceae· pro. parte, Cacteae.
Ficoideae pro p~tet Umb~Jlifer~e pro parte (with transitions to the. CaryophyUeous
type), Araliac~ae pro parte, 'Rubiacea~ (Fig. 101, p. 446). Candolleaceae. pro
parte, Goodeniaceae pro parte (four subsidiary cells, as in T1'adescantia), Vacciniaceae,.
Ericaceae,pro parte (rare), ,Plumbagineae pro parte, Sapotaceae pro parte, Styra*
ceae pro parte, Oleaceae pro parte (rare), Salvadoraceae pro parte, Apocynaceae (in
most cases)" Ascl~piadeae (in most cases), Loganiaceae pro parte •.Convolvulaceae
pro parte, Blgnomaceae pro parte~ Selagineae (see under I), Nyctagmeae 'pro parte,
 STRUCTURE ,0£ THE ,LAMII'{A' _OF THE LEAF loBI

Chenopodiaceae pro parte, BaselIaceae, Phytolaccaceae :pro parte, ,Batideae. Poly-

gonaceae ',pro parte, Chloranthaceae pro ,parte,. Myristioo.ceae, Monimiaceae pro
parte, Laurineae, Hernandiaceae pro parte, (i()monegaceae, Proteaceae (false
Rubiaceous type), Loranthaceae;,Santalaceae (in mostcases), ChaW}-pereia, Euphor-
biaceae pro parte; Da:phniphyllaceae, Ulmaceae pro parte, Thelygonea~, Casuarineae
(Fig. 186, p. 788), Salicineae. ,
'3. Stomata with subsidiary cells, lying transversely to the pore, occur ,in:
Caryoplcylleae pro parte (with exceptions, Fig. 25, C, p. 108), Melastomaceae pro parte
(in part at least false Caryophylleous type, gua!Q..cells occasionally almostcompfetely
surrounded by. a single epidenn~al .cell), ~ Solanaceae pro parte (side. by side"with
other. types), Lentibularieae pro parte, Bignoniaceae pro ,parte. Acanthaceae.
(Fig. 144 •. A, p .. 615), Verbenacea~ pro ,parte, Labiatae pro parte (Fig• .15 2 , P.
p. 638), Plantagineae, pro parte, Illecebraceae pro parte. i' ,
4. Stomata, in which the pronounced Cruciferous type, is' still recognizable in
the mature.leaf,:are found in I: Cruciferae pro parte, Bixineae pro parte, .Rhapto--
petalaceae pro ,parte, ,Rhamneae pro parte, Staphyleaceae pro 'parte, Papilionaceae
pro parte,Crassulaceae (in most cases, Fig. 70, A, p.,,32I), Lecythidaceae pro parte,
Melastomaceae pro parte, Lythrarieae pro parte•. Samydaceae.pro parte, Begonia-
ceae, Araliaceae pro, parte, Dipsaceae pro parte, .Pentaphragma (Campanul~c~e).
Bt'achynema, Apocynaceae pro parte, Asclepiadeae pro parte, Log~niaceae pro parte,
Boragineae,pro parte, Convolvulaceae pro part~, Solanaceae pro parte, Gesneraceae
pro parte, Bignoniaceae ,pro parte, Pedalineae pro parte, Myoporineae pro parte,
Verbenaceae pro parte, Piperaceae pro parte, Euphorbiaceae pro parte, Moraceae
pro parte, Urticeae ,pro parte.
The shape of the gUard.cells· and the 'detailed' structure of their walls
likeWise appear to be features 'of great systematic importance, although hitherto
little attention has'been'pa~d. to t~em in papers dealing With systemati~ ap.~tomy
or, for the matter, of that,:m DIcotyledons general).y. In this connexion the
following features chiefly require to be considered: the contour of the pairs of
guard·cells and the shape of·the front cavity, when seen from the surface; the
structure of the back cavity; the v~ried character and chemical nature of the
unequal thickening of. the walls of the ,'guard-'cells, and the corresponding
differences iri the shape 'of their lumina; lastly; the thickening ridges which
arch over the:front; and' back cayities" these ,ridges bei#g f<?r~ the most part
strongly cuticularized, and the more or less ,strongly developed epidermal
joints, found on either side of the guard-cells 2.. '

These features. have ,bee~ 'examined chiefly.in the ~onocotyledons. GymnO-

sperms and Mosses. The' Gramineous type,~:first demonstrated by Schwendener
in the Gramineae and Cyperaceae, is primarily characterized by the fact that the
two guard-cells ar~ duml;>bell-shaped. i.e. they are low and flat in the ,middle region
and enlarged at bo~h ends. The outer and inner walls of ,the middle portion of
the guard·cells are distinguished ,by having. strong thickening ridges, which are
not, .as is elsewhere,the case, approximated to the ventral side, but take up the
entire breadth of the miqdl~, part and merge gr~du~y' into 'the thin walls of ,the
enlarged ends' of the guard-cells; as a result the middle part' of the guard·cell,
when seen in it'transverse section. shows a slit-shaped lumen; ',which is placed trans·
versely. The front cavity, as seen in surface-view, has an hexagonal outline with
two longer sides, which are, placed- parallel to the, direction of the pore. In
at.l4iti,on, there -are two supsidiary cells, which have, thin 'walls and lie parau.el to .Hie
pore.-'The Gymno~permous type is particular~y characterized by the fact that the

l The followJng "enumeration is not exhaustive, among other reasons, because, 'In .'the case of the
material' hitherto investigated, there are frequently enough no precise data, as to whether' the three
epidermal cells, surrounding the pair of guard-cells, are differentiated as subsidiary ceIIs or not•
. I Schwendener, Ball u. Mecbanik d. Spaltoffn., Monatsber. Berliner Akad., 1881, p. 833;
Mahlert, Anat: d, Laubbl. der Coniferen, &c" Bot.' Centralbl., 1885, iv, p. 54; Haberlan'dt; 'Anat~ u.
Phys. der Laubmoose, in Pringsheim Jahrb., xvii, 1886, p. 359 and Tab: xxvi;' Schwendener,
Spaltoffn. d. Gramineen n, Cyperaceen, Sitz.·Ber. Berliner Akad., 1889, p. 6:; ; 'Copeland, Mechanism;"
of stomata, Ann. of Bot., xvi, 1902, p. 327; Buck, Vergl. Anat. des Durchliiftungssyst., Diss.,
Freiburg i. Br., 1902; see especially Porscb, loc, cit., 'pp. 5, 2J, and 33 et.seq. and the literature cited
in this paper. . ,
 CONCLUDING REMARKS
guard-cells, as seen in a median transverse section, have the shape of an ellipse, the
longitudinal axis of which is inclined approximately at an angle of 45° to the surface
of the organ; other characteristic features are that the outer walls are mostly more
strongly thickened than the inner walls, and that as a rule both walls-rarely the
outer walls alone-include a lignified lamella.-The chief points to mention regarding
the Mu.cineou. type are that the thickening ridges on the front and back cavities
are not typically differentiated, and that nQt uncommonly the guard-cells undergo
fusion or~the stomatal apparatus consists of three or four cells.
As has already been stated above, there are very few observations dealing with
Dicotyledons from this point of view. The outline of the pair of guard~cells is
generally elliptical or circular, the angular outline, found in the Papaveraceae and
Fumariaceae, being remarkable. Thickening ridges, like the flaps of an envelope
and resembling those present in CY'peraceae and Gramineae, are not at all rare in the
guard.cclls of Dicotyledons (e.g. In the Celastrineae according to Metz). Similar
types of structure are produced by the presence of thin areas in the walls of the
guard-cells, these thin portions being round or of other shapes and occupying'the
same polar position as in Gramineae, &c.; they occur in the Hippocrateaceous
genera Hippocratea and Salacia (according to F. E. Fritsch), in many Ericaceae
(according to Copeland), and in the Solanaceous genu's Tf'~·anaea (here comma-
shaped, according to Solereder). Other noteworthy features are constituted by
the splitting of the outer appendicular ridqes (' horns ') on the guard-cells into two
ridges, so that the front cavity becomes diVIded into an outer and an inner compart-
ment (in certain Rhizophoraceae), and the deposition of' cutine-Iamellae, which
correspond in all respects (see Porsch. Ioc. cit.) to the lignified lamellae of Gymno-
sperms (see above), in the outer and inner walls of the guard-cells in the Casuarineae.
Schwendener refers also to a well-known type of structure. which is shown by the
guard·cells of BelJehorus and is figured in Sachs's and Haberlandt's textbooks;
similarly Porsch, in the work cited in the footnote on p. 1079, describes a certain
type of structure, exhibited by the guard-cells in the phyllodineous Acacias and in
~U&alyPtus. '
At this point mention may also be made of peculiar processes on the walls
of the guard-cells, as seen in surface-view; these processes take the form of lobes
or have some other shape. and are found in certain Trochodendraceae, Berberideae,
Geraniaceae (Fig. 39, A, p. 170). Sabiaceae (according to Dihm in Beih. z. Bot.
Centralbl. xxi, Abt. It 1907), Laurineae and Penaeaceae (Fig. 175, p. 723); they
occasionally occur also when the ordinary epidermal cells have jagged lateral walls
with ridge-like thicket;rings in the angles. The distinctive shape of the guard-cells
in certain floating plants (Nymphaeaceae, Trapa), when seen in a transverse section,
is a biological feature; the closure of the pore in these forms is not brought about
by contact of the protruding ventral walls. but by means of the outer cuticular
ridges, which are much broadened.
The size and number of the stomata, especially in extreme cases (very
large or very smaH stomata, occasionally even stomata of two sizes on the same
leaf-surface; further, very many or very few stomata, which in the former case
together with the neighbouring or subsidiary cells form almost the entire
epidermal surface). may be employed for the diagnosis of species and occasion-
ally even of more extensi.ve taxonomic groups. A complete reduction of the
stomata on the leaves or on other organs as well is found only in certain (not
all) plants, which are submerged or lead a saprophytic or holoparasitic life.
When stomata occur in such plants, they are mostly present only in small
numbers, and as a rule merely owe their presence to inheritance; they are
functionless, and in correspondence with this they often show a reduced type
of structure and are occasionally provided with special arrangements for the
closure of the pore 1.
Complete absence of stomata has been recorded: in the submerged leaves of
Ranunculus § Batrachium, Nymphaeaceae pro parte and Halorageae pro parte; in
Aldrovanda, Podostemaceae, and Ceratophylleae; in some of the saprophytic
1 Ponch, Spaltoffnungsapparat, pp. 47-90 i Porsch, SpOlltoffnungsapparate submerser Pflanzen.
leitE', Sitz.-Ber. Wiener Akad., cxii, Abt.
I, 19 0 3, p. 97.
 STRUCTURE OF THE LAMINA OF THE LEAF 1083

Monotropeae; and lastly, in the parasitic Orobanchaceae pro parte, Cytinaceae

pro parte and Balanophoreae pro parte.
The distribution of the stomata on the two surfaces of the leaf! varies.
It generally depends on the structure of the leaf, since leaves showing centric
or homogeneous structure (especially those which are centric and are placed
in a vertical position) in most cases bear stomata on both surfaces, while in
leaves with typical bifacial structure the stomata are generally confined to the
lower side. When stomata are present on both sides of the leaf, they are
mostly more numerous on the lower than on the upper side~ although the reverse
is sometimes the case. Even though as a general rule these features are only
of value for specific diagnosis, the exclusive occurrence of stomata on the
lower side of the leaf is occasionally characteristic of entire Orders of varying·
magnitude (Dipterocarpeae, Hamamelideae, Begoniaceae). No great systematic
value can be attributed to the occurrence of isolated stomata in the neighbour-
hood of the veins on the upper side of the leaf. The floating leaves found in
some species of Rammculus, the Nymphaeaceae (also on the leaves of Nelum-
bium, which are not floating, see Introduction~ p. 9, footnote I), CaUitriche
(Halorageae), Trapa {On agrarieae) , Limnanthemum (Gentianeae), TrapeUa
(Pedalineae) and Polygonum naturally bear stomata onlyon the upper side. Even
in terrestrial plants, however, the stomata are sometimes confined to the upper
side of the leaf. In some cases (e.g. in the leaves of Lepidophyllum or Passerina,
which are adpressed to the stem, or in the in duplicate leaves ot certain Podalyrieae
and of H emzphragma) this peculiar distribution is connected with the formation
of spaces, which contain the stomata and are not reached by the wind; in other
cases, as in the leaves of certain species of Lythrum, PhiloxeTus, Pitea and
Euphorbia, it is related to the differentiation of aqueous tissue in the lower part
of the leaf; in other cases again (species of Geoffroya) the restriction of the
stomata to the upper side is inexplicable. The same arrangement of the stomata,
combined with other remarkable peculiarities in their distribution, is found
also in certain Mimoseae, where these features are connected ~ith the assumption
of the' sleep-position. The curious distribution of the stomata in certain
saxifrages may a1so be briefly referred'to at this point. ,
Stomata, which are situated exclusively on the upper side, are found (apart
from the floating leaves already discussed) in the following land-plants: Antarctic
species of Caltha (which have curiously shaped leaves), species of V~·ola, species of
Myricaria and Tamarix (Tamariscineae), Geo/Jroya spinosa, Jacq. and species of
Dillwynia, Eutaxia, Pultenaea, Coelldium (Papilionaceae), certain Mimoseae.
species of Brunia f. Lonchostoma. Pseudobaeckea and Raspalia (Bruniaceae), Epi-
lobium crassum. Hook. f. (Onagrarieae). species of Candollea (Candolleaceae).
Hedraeanthus PumiUo, Porta (Campanulaceae). Lepidophyllum (Compositae).
Cassiope (Ericaceae), Leucopogon (Epacrideae). Hemiphragma and Lathraea (in the
latter. genus on the' scale-leaves: Scrophularineae). Philoxcrus (Amarantaceae),
Passerina (Thymelaeaceae), Euphorbia buxitoUa, Pilea serpylli/ol£a (Urticeae).
The arrangement of the stomata with respect to one another, as seen
in surface-view; is wellJmown ~to be quite irregular in most of the Dicotyledons.
In exceptional cases the stomata lie with their pores directed parallel to one
another, and at the same time parallel to the median vein of the leaf, as is the
general rule among Monocotyledons; such an arrangement occurs chiefly in
species having narrow leaves or leaflets. Another specially remarkable case
is obtained, when the pores of the stomata are placed parallel to one another,
but at the same time lie transversely to the midrib of the leaf; and it may
at once be added, that, when stomata are present on the axis of such plants,
they are placed transversely to the vertica1 direction.
1 Fr. Darwin, Bloom on leaves and distribution of the ~tomata, Joum. Linn. Soc., xxltii, 1887,
p. 99; Kohne, Oberseit. Spaltoffn. Sluf Laubholzgew., Mitteil. deutsch. dendrolog. Ge:;ellsch., 1899,
P·47·
 CONCLUDING REMARKS

Stomata, which have.their pores directed parallel to one'another and to the

midrib of the leaf, have been observed in certain: Mimoseae (cominon), Bruniaceae,
Melastomaceae, Cacteae ,(on the stem parallel.to the longitudinal axis of the shoot),
CandoUeaceae,' Cainpanulaceae. Epacrideae' (in almost an the' species);~lumbagineae,
Loganiaceae, Polcmoniaceae, Chenopodiaceae, Proteaceae, Santalaceae, Myzoden-
dron; stomata, which are arranged parallel to one another; but lie transvenely to
the midrib of the leaf or to the vertical direction in the axis,'as the case may be, 'are
found in certain: Polygaleae (K"amel't'a pro 'parte, leaf), Tamariscineae (leaf); Bala·
nites (axis), Rharnneae (axis of Colletia, according to Pfitzer), Staphyleaceae (axis 'of
Staphylea pinnata, acc~rding to De Bary), PapilioD?-ceae (leaf ,in A nartM:op"yll~m,
EutaXia and Latyobea, axis in Daviesia, phylloc1ades and branches respectively
in Carm'ichaelia' and Alhagi), BiUniaceae (Brunia' and Staavia,'leaf), Cacteae' (aXis),
Ficoideae (Mesembryanthemum, leaf), Epacrideae (Lysinema, leaf), Chenopodiaceae
'(leaf in Suaeda, Salsala, Camphorosma, Echinopsilon, ·Halogeton; T"aganum. and axis
in Camphorosma, Salicornia, Suaeda),. Batideae (leaf and axis), Nepentha'ceae pro
parte, Laurineae (leaf and axis of Cassytha), Loranthaceae (leaf and axis of Nuytsia,
and axis of other Loranthaceous genera), Santalaceae (commonly on ·the branches
"and also on the ,leaves 1), Euphorbiac'eae (axis, of succulent Euphorbias). Casuarineae
(~heaths an,d b~anches. Fig. i86, P.,7 88 ),' I

The combination of stomata' to form groups is' not' very common. ~uch
groups are found in the. first place in certain Cruciferae, .where -they are due to
the differentiation ~f the' epidermis of the leaf"into cells with large' 1umina
and a reticulate distribution, and 'cells of the normal siie, which constitute
the 'subsidiary cells of the stomata. Stomatal groups are
likewise present on
the surface of the leaf in M acrO co cculus (Menispermaceae), speCies of So~,la1nea
and Castela (Simarubaceae), species of Euchresta '(Papilionaceae), species' of
Saxi/raga and Chrysosplenium, species of Calycogonium, J;,eiind,a and Ossaea
(Melastomaceae),' 'certain Begonias (Fig. 87; A,- 'po 400), all the species, ,of
Pagamea (Rubiaceae) 'and Napeanthus (Gesneraceae), ana in ficus gibbosa,
BI:; and in small pits 'in the surface of the leaf in Sarcolaeit'a aridSchizoliuma
(Chlaenaceae, Fig: :33~" p. I45), S01.tlamea Pancheri, Brongn. et Gris (Simaiu·
baceae), Trichouratea'subvelutina, Van'Tiegh. (Ochnaceae), Akania' (Staphy·
leaceae), Mouriria (Melastomaceae, Fig: 78,. 'p. 359), Nerium (Ap6cynacea'e),
species of Banksia and Dryandra' (Proteaceae), and species·of Ficus 2. Lastly,
we can 'also speak of ' stomatal groups'when the network ,of "the veins projects
strongly on the lower side of the leaf and the stomata are confined to the' de-
pr~ssed p,ortions 'of the surface. , ;In connexion with our discussion of the stomatal
pits (crypts), we may notice the' occurrence of furrows occypied'by stomata)n
Tolled leaves ~d on the assimilating stems of._plants 'having reduced le~ves
(e.g. in certain Papilionaceae~ Casuarineae, &<;:), whilst with. the stomatal groups
situated on the surface, ·of the leaf we may class the arrangement of ,the
stomata in longitudinal zones on- the leaves of certaIn species of Cand~llea
(Candolleaceae). ',
uther points 6f difference are connected with: the varying position of the
guard-cells, with, respect to the' neighbouring epidermal 'cells, 'a feature
which in most cases may be regarded as 'a measll:re of the 'humidity of
the habitat in which the species grows~ The' guard:cel1s 'may' either lie on
a: level with the epidermal cells or' project to a marked extent, the ;latter being
especially the case in species from damp localities, in species in which the stomata
are contained 'in srnall 1pits, or in species which are protected from desiccation
by the possession of a: 'very dense hairy covering; 'in other·· cases again' the
'guard-cells are sunk below the surface, this feature being found'chiefly in species
from dry habitats. The' elevation of. the stomata is particularly marked in

·1 See also De Bary, Vergl. Anat., 1877, p. 48. , ,

2 Haberlandt (Physiol. ]lllan:zenanat" 1904) incorrectly ascribes small stomatal pits~·like those
of the Oleander, also to CO!(illillln (Menispermaceae).
 STRUCTURE OF THE LAMINA OF THE LEAF 1085

the veins of the leaf of Santiria (Burseraceae, Fig. 43, p. 191), and in Cineraria
cruenta (Compositae), certain species of Cordia, in the leaves of Fab~·ana (Sola-
naceae), which are covered with varnish, and on the stems of certain Cucur-
bitaceae. The depression of the guard-cells is effected either (a) by the diffe·
rentiation of a thick outer wall on the epidermis, which is accompanied by a
correspondingly strong development of cuticular crests, or (b) simply by the
depression of the guard-cells (in some cases together with their subsidiary cells)
beneath the neighbouring epidermal cells, or (c) by a more or less pronounced
protrusion of the epidermal cells adjoining the guard-cells or subsidiary cells,
which leads to the formation of a chimney-shaped outer respiratory cavity of
varying shape and often of great depth 1. The depression of the stomata in
certain Thymelaeaceae and certain species of Jacksonia (Papilionaceae) is of a
very peculiar character, and may therefore find special mention; in the former
case the stomata occur singly at the base of lageniform pits, the wall of which
is formed by the elongation of the 6-ro neighbouring cells surrounding the guard-
cells (Fig. 174, p. 717), while in the species of Jacksonia the stomata together
with their subsidiary cells are sunk in deep pits on the surface of the stem, each
pit being closed by a clothing hair. Regarding certain Sapindaceae and Umbel·
liferae, see pp. 228 and 420.
4 Plugged' stomata, which have long been known to occur in the Coniferae,

have recently 7. been demonstrated in the Monocotyledons and Dicotyledons.

They do not, however, possess great systematic value, since they are not always
of const~nt occurrence in one and the same species, and are often found only on
certain parts of the plant. In stomata of this kind the outer respiratory cavity
(vestibule) is filled with a resinous mass, which interferes with the process of
transpiration and the ~xchange of gases as a whole.
The same effect is produced by the peculiar thickenings found in certain
Epacrideae (Fig. III, p. 492) on those walls of the subsidiary cells, which border
on the inner respiratory cavity. Finally, stomata may be put completely out of
action by a local development of cork beneath the pairs of guard-cells; this
ultimately leads to the formation of what are called cork-warts; for the cork-
warts on the leaf, see § 39. I
Twin-stoJp.ata (i.e. pairs of stomata lying side by side and touching one
another by one of their longitudinal walls) now and then occur together with
solitary stomata on the same surface of the leaf, but they are no doubt merely
casual structures and have no systematic value.
§ 5. WATER-PORES 3. Water-pores (i.e. open stomata, which function as
hydathodes) occur singly or in groups on the leaf-teeth of many plants (d. § 35,
and the descriptions of the Orders there enumerated), or are found near the
margin of the leaf (as in certain GeraniaceaeJ Begoniaceae, and Campanulaceae)
or above the points of intersection of the veins of the leaf (as in certain Nym-
phaeaceae). They are distinguished sometimes by small dimensions, sometimes
by considerable size 4• The water-pores of the Lobeliaceae are particularly
remarkable in that their pore is closed by the cuticle, a rather thick and
cuticularized ridge penetrating from the latter into the cavity of the pore

1 See Tscbirch, Bel. des anat. H!lnes, &c., Linnaea, xliii, 1881. p. 223 and Tab. ii.
: Wulff, Verstopfte Spaltoffn., Osterreich. bot. Zeitschr., 1898, pp. :4101, 25:2 aJ?d :298, Tab. viii.
S Volkens, Wasserausscbeidnng, J ahrb. Berliner Garten, ii, 1883, p. 166 i"'_ Haberland t,
Wassersec. u. absorb. Org., Sitz.·Ber. 'Wiener Akad., cHi and civ, Abt. 1, 1894 and •1895, pp. 489
and S5 respectively; Haberlandt, Hydathoden. Her. deutsch. bot. Gesellsch., 189., p. 367; Nestler,
Wasserspalten. Nova Acta Leopold., lldv, I89<f, p. loB: Spanjer. Wasserapparate, Bot. Zeit., 1898,
i, p. 35; Minden, Wassersec. Org., Bibl. Bot .• Heft 46, 1899.
" It remains an open qurstioD, whether the pain of guard.~lls, which occur in many plants, and
are distinguished from other stomau. on the same surface of the leaf by their often considerable size,
are invariably of the nature of water-pores.
 I 086 CONCLVDING RE}'IARKS

(' stomates aquiferes septes ') 1. Beneath the water-pores one finds either the
ordinary tissue of the mesophyll containing the termination of a vascular bundle,
or a special thin-walled tissue (epithema), provided with intercellular spaces and
enveloping the termination of the bundle. From the systematic point of view we
may notice that the presen<:e of water-pores constitutes a feature characteristic
of the Menyanthoideae in contrast to the Gentianoideae. The water-pores;
found in certain Crassulaceae, Moraceae and Urticeae, deserve special mention;
they are united to form groups and are recognizable with the naked ~ye or with
the help of a lens as small spots on the surface of the leaf. We may also refer
to the groups of water-potes, found in certain species of Saxilraga (Fig. 66,
p. 313), where they function as chalk-glands and have long been employed as
a specific character by systematists.
With the water-pores We may dass the apical pores 2 occurring at the tip
of the leaf in various Dicotyledonous water-plants. They serve for the.excre e

tion of water and arise by the breaking down of the cells, situated above the
terminations of the veins (mainly epidermal cells~ but occasionally waterepores
as well). .
Small pits (which do not contain water-pores, but are formed by special epi-
dermal cells surmounting the termination of a vascular bundle), such as occur in
certain Ferns, have not been observed either in Dicoiyledons or Monocotyledons.
Regarding special epidermal cells, functioning as hydathodes and found in certain
Menispermaceae and Olacineae. see the end of § 2, p. 1076; for the epidermal chalk-
and salt-glands of the Frankeniaceae. Tamariscineae and Plumbaginaceae, see § 37 ;
for peltate glands. secreting 'Water and occurring in certain Rhinanthaceae and in
Monophyllaea (Gesneraceae), see § 34 under I, b. Trichomes. 'Whic~ in correspon-
dence with their outward form, are described as clothing or glandular hairs in
descriptive anatomy. may also function as hydathodes (see § 29).
§ 6. MESOPHYLL, PALISADE AND SPONGY TISSUES 3. The influence of the
surrounding medium on the structure of the leaf (i.e. the bifacial (dorsiventral)
or centric (isolateral) development of the mesophyll and the detailed differentia-
tion of its cells) has already been discussed in the Introduction, and in this
cannexion mention was likewise made of the variability of these structural
features in one and the same species (see Introduction, pp.6 and 10). In spite
of this fact, these features may be employed for systematic purJX?ses, and in the
first place for specific diagnosis, provided their constancy is sufficiently estab-
lished. This applies especially to the structure of the leaf, which we may dis·
tinguish as bifacial (palisade-tIssue On the upper, spongy tissue on the lower side),
subcentric to centric (palisade-tissue on both sides, occasionally formin&, 'the whole
of the mesophyll) and homogeneous (mesophyll showing no differentIation into
palisade and spongy tissues); occa$ionally the structure of the leaf is constant
even in all the members of a small Order (Hamamelideae with bifacial structure).
The most important features to notke in the structure of the paUsade-tissue are
the following: the regular or irregular stratification of the layers of palisade-cells,
the cells of the same layer in the latter case not being of equal length, and super-
posed cells occasionally appearing to be derived from a single much elongated
palisade-cell by transverse division; the number of layers constituting the

1Tswett, Hydatbodes, &c., 100m. de bOt., J907, p. 305.

tWeinrowsky, Scheiteloffn. bei Wasserp6., D.iss., Berlin l J 898 (also in Fiinfstiick, Beitl., iii); Re
also Minden, loco cit.
S Haberlandt, Assirnilat.. Gewebesyst., in Pringsheim Jabrb., xiii, J882, p. 7.. and Tab. iii-viii,
and Ber. deutlch. bot. Gesellsch., 1886, p. ~06; Sta.hl, Einti. d. sonn. und schatt. Standorts, &1.:.,
Zeitschr. f. Natnrw., lena, 1883 j Heinrich~r,. lsolat. Blattbau, in Pringsheim Jahrb., x1/, I8S.,
Po [,0; and Tab. xxvii-xxxi j Loebel, Anat. d. Laubbl., loco cit.,.xx, .BSg, p. 38 and Tab. i-iii j
Ruck. Vergl. Anat. d. Durchliiftangssyst., Diss.• Freibnrg i. Br., 1901, p. i-4; Schwartz.Clements,
Relat. of leaf.strnct. to phys. (acton, Transa.ct. Americ. Microscop. Soc., 1905, p. 19.
 STRUCTURE OF THE LAMINA OF THE LEAF lolS7

palisade-tissue; the shape of 'the palisade-cells (IQng and narrow cells, short
and broad cells, 'funnel-cells,' and arm-palisade cells; for the latter, see also
below); the relative extent to which the palisade-tissue, as compared with the
spongy tissue, participates in the formation of the mesopbyll. In the structure
of the spongy tissue important characters are afforded by the varied arrange-
ment (stratified or irregular) and shape (rounded cells, cells provided with several
arms of varying length, cells with flat or with spreading arms, i.e. with arms
developed para1le] to the surface of the leaf only or radiating out in all directions)
of the cells, and the consequent variations in the nature and size of the
intercellular spaces.
Rolled leaves 1 generally show a distribution of palisade and spongy tjssue
which differs from that normally found, in so far as palisade-tissue is mostly
developed in all those parts of the leaf which are turned towards the light, while
spongy tissue occurs on the surface of the furrows, present on the upper Of lower
side of the leaf. The leaves of certain Tamariscineae, which are reduced to leaf-
sheaths, and the leaves of certain Compositae, &c., which are adpressed to the
axis and generally have their stomata restricted to the upper epidermis
(ct. § 4), resemble those ,rolled leaves, which have a furrow on their upper side,
in the development of palisade parenchyma only on the lower side of the leaf,
which is turned towards the light.-
In addition to the palisade and spongy tissues, which contain chlorophyll;
aqueous tissue with little or no chlorophyll plays a part in the formation of the
mesophyll, especially in thick and fleshy leaves. The differentiation of hypo·
dermal aqueous tissue of varying thickness, but belonging to the mesophyll, as
shown by a study'of its development, has already been discussed in § 3. In
other cases the aqueous tissue, the cells of which may have thin or thick walls,
forms a middle layer (sometimes differentiated like a pith) in the leaf, or com-
poses almost the entire mesophyll, so that the assimilatory tissue is cr9_wded
into the middle of the leaf; in the latter case the assimilatory tissue sometimes
merely envelops the sheaths of the veins in the form of a rosette (Portulaceae,
Fig. 26, A, p. II2, Chenopodiaceae, Fig. 158, p. 656). The occurrence of' a tanni-
niferous middle layer, which therefore has a brown colour in the dry leaf, is
characteristic of certain plants (Anonaceae, Sapindaceae, Papilionaceae,
Mimoseae, Rubiaceae, and no doubt other Orders as wen).
As regards the special structural features presented by the palisade-
tissue, we mat first deal with the occurrence of funnel-cells, arm-palisade cells,
and conjugate cells as component elements of this tissue. Funnel-cells are short
}Jalisade-cells, which are narrowed down at their lower end in the form of a
blunt cone; they are characteristic of plants inhabiting moist localities (occur-
ring for example in certain Gesneraceae and Piperaceae). Cells having a
similar shape (' collecting cells') are also found in the interior of the mesophyll,
adjacent to the typical palisade-parenchyma. A more important systematic
feature appears to be constituted by arm-palisade parenchyma, in which the
elements,_ as _·seen ill a transverse section of the leaf, are .not single cells
independent of one another, but the arms of what are called arm-palisade cells.
The latter, or rather their individual branches, may be either long or (as in
most cases) short, while the number of the branches is two or more. In con-
jugate palisade-cells the longitudinal walls are provided with two or more
small papillose protusions, which enter into connexion (conjugate) with one
another in neighbouring cells. This type of structure has been observed mainly

1 Rolled leaves have been observed in tbe following Orders: Berberideae, Frankeni.a.ceae',
Tremandre~, Polygaleae, Hypericineae, Lineae, Geranilceae, Rhamneae, Papilionaceae, Rosaceae,
Rubiauae, Compositae. Ericaceae, Epac::rideae, Primulaceae, Scrophularineae, Empetraceae (Fig. 188,
p. 800), Euphorbiaceae; see also Meigen, in Engler, Bot. J..brb., xviii, l~.
.
J088 CONCLUDING REMARKS

in the short palisade-tissue found on the lower side of subcentric leaves and
occasionally extends also to layers of the adjoining spongy tissue.
Arm-pali~ade tissue is found not only in the Filices, Equisetaceae, Coniferae
and Gramineae, but also in a number of Dicotyledons, viz.: Aconitum, Adonis,
Anemone, Caltha, Clematis, Delphinium, Nigella, Paeonia and Trollius (Ranuncu-'
laceae), Euptelea (Trochodendraceae), Discipham'a (Menispermaceae). Sauyauja (Tern-
stroemiaceae, Fig. 29. A, p. 129), Meliosma (Sabiaceae), Casearia (Samydaceae.
only faint indications), Acanthopanax, Cussonia, Gt'libertia, and Pseudopanax (Aralia.
ceae), Ado:ra, Sambucus and Vibttrnum (Caprifoliaceae), Candollea (Candolleaceae),
numerous Campanulaceae, Lysimachia and Trientalis (Primulaceae), Symplocos
(Styraceae). Protoschwenkia and Schwenkia (Solanaceae), Ascarina, Chlof'anthus,
and Hedyosmum (Chloranthaceae), Phyllanthus (Euphorbiaceae) and Parartocaypus
(Moraceae). The systematic value of this feature is either restricted to species
(e.g. in Anemone and Phyllanthus) or to genera (e.g. Meliosma).

The fine bellows-like folding, which may commonly be observed in her-

barium-material on the lateral walls of the palisade-tissue, and is met with also
in the hypoderm, is a result of the participation of these tissues in the storage of
water; the same explanation applies to the occurrence of ridge-like or reticulate
thickenings on the walls of the palisade-cells in certain plants, e.g. species of
Clusia (Guttiferae, Fig. 27, A, p. 122), Itleriania and GraiJenrt·eda (Melastoma-
ceae), Candollea (Candolleaceae). Sophoclesia (Vacciniaceae). Dyssochroma,
juanulloa, and Markea (Solanaceae).
In certain species of Reynosia and Sarcomphalus (Rhamneae), Light/oolia
(Campanulaceae) and Artocarpus and Ficus (Moraceae) the cells of the spongy
tissue have a very remarkable hypha-like form.
In the species of Buxus, belonging to the section Eubuxus. the sudden tran-
sition from assimilatory tissue with rounded lumina (which adjoins the palisade-
tissue on its lower side) to spongy tissue with large lacunae leads to the splitting
of the leaf into two corresponding halves.
Another structural feature of the mesophyll, which may be mentioned at
this point and can be employed for systematic purposes) is the occurrence of
peculiar thickenings or swellings, which are restricted to certain points of the
cell-wall~ and are found~ especially in the spongy tissue~ but also on the upper
and lower walls of the palisade-cells, in many Menispermaceae and Melasto-
maceae, in certain Loganiaceae and in M elananthus guatemalensis, Solered.
(Solanaceae); these thickenings have a mucilaginous appearance and more or
less recall the structure of the wall in collenchymatous tissue. They certainly do
not differ very materially from the gelatinized portions of a cell-wall, especlally
as subepidermal layers of mucilage occur side by side with them in the same
species (e.g. in certain Menispermaceae and Loganlaceae); both the inner walls
of the epidermal cells and the adjoining walls of the palisade-tissue are concerned
in the formation of these mucilage-layers. Gelatinization of the spongy tissue
(differing from the swellings just discussed ?) has been recorded in numerous
Apocynaceae and in Sclerophylax (Solanaceae), and gelatinization of the entire
mesophyll in certain species of Gentiana. It remains to mention the cracks
due to drying 1, which are formed in certain species by a process of fissuring
owing to the drying of the leaf, and frequently give rise to transparent dots
(Capparideae, BUfseraceae, Sapindaceae, Connaraceae, Cornaceae, Bignoniaceae,
Verbenaceae ).
For the occurrence of sclerosed cells in the mesophyll, see § 9, p. lOgo; for
the occurrence of secretory organs, see § 14 et seq., and of oxalate of lime and
other contents, see § 22 et seq.

1 See Blenk, in Flora, 18S .. ; ~nd Radlkofer, in Sitz.·Ber. Munch. Akad., 1886, p. 341.
 STRUCTURE OF THE LAMINA OF THE LEAF I<>B9
§ 7. STRUCTURE OF THE VEINS OF THE LEAF. The following features can
be used for systematic purposes: the appearance of the fibrovascular system,
as seen in transverse section; the presence or absence of mechanical tissue or of
a protective sheath in connexion with the vascular system; and the structure
of the tissue, developed above and below the vascular system. When the
vascular system of the veins is surrounded on all sides by assimilatory tissue,
we may speak of 'embedded veins,' in contrast to 'vertically transcurrent
r
veins,' in which a special colourless tissue Begleitgewebe ') adjoins the vascular
system on its upper and lower side.
When undertaking a comparative investigation, the observations should, in the
first place, invariably be based on veins of the same order, and also on a transverse
section taken through a definite part of the vein in question. For, in the case of the
principal vein and all the larger lateral veins, the appearance of the fibrovascular
system in a transverse section and the differentiation of the mechanical tissue
accompanying the vascular bundles generally differ according as the vein is examined
at.its base or in its jurther course; and in the case of the smaller veins this applies
at! least to the latter feature. A further point to be taken into account IS the
thickness of the veins. Lateral veins of the same order may vary in thickness and
may conseq_uently show a different structure at corresponding points in allied species,
especially If the latter have leaves of different sizes. The systematic employment
of the features in question requires a critical judgement, and has on the whole been
little attempted hitherto, mainly o'wing to the amount of labour involved, and in
view of the fact that the results to be obtained are not very considerable.
Among features, which are more easily determined, the following chiefly
come into consideration for the purposes of anatomical diagnosis: the embedded
character or vertical transcurrence of the medium-sized and smaller veins; the
presence or absence of a strongly developed mass 'of mechanical tissue (scleren-
chyma) in connexion with the vascular bundles; and the occurrence of a sheath
of large cells around the vascular bundles. Vertical transcurrence of the
smaller veins is often characteristic of a genus (see for instance Epacrideae,
p. 492) and occasionally even of an Order (Dipterocarpeae); such ve~s give rise
to a vascular network, which is visible even to the naked eye. The diverse ways,
in which transcurrence is effected, are generally only of specific value/; the veins
may be vertically transcurrent by means of colourless parenchyma, the walls of
which are either thin or slightly lignified or sclerosed or collenchymatous, or
by means of plates of sclerenchymatouS fibres, which sometimes spread out
beneath the epidermis of the leaf like a T.girder or may resemble a hypoderm.
When sclerenchymatous tissue is present, the extreme cases (on the one hand
a more or less closed ring or: at least well-developed arcs of sclerenchyma, on
the other hand occurrence of mere isolated sclerenchymatous cells or complete
absence of this tissue) are chiefly to be considered. In the majority of cases
this biological feature is only of specific value (see the Introduction, p. 8 et seq.),
but there are entire Orders (Begoniaceae), in which sclerenchyma is either
completely wanting in the veinsor is found only in a few species, and then mostly
in small quantity. Sheaths of large parenchymatous cells 1 occur more
commonly in Monocotyledons than in Dicotyledons; they function as a pro-
tective envelope and to a less extent for the conduction of food-substances; such
sheaths have been observed in the following Dicotyledons: Portulaca (Portula-
ceae), Tribtdtts (Zygophylleae, here with thick pitted walls, Fig. 38, D, p. r68),
certain Papilionaceae, Pedis humitusa, Sw. (Compositae), HeUotropium frutico-
sum, L. (Boragineae), certain Polemoniaceae (cells thickened in aU-shaped
manner in Gilia pungens, Hook., suberized cells in other cases), Nyctagineae,
Amarantaceae (Fig. 156, A, B, p. 652), Chenopodiaceae (Fig. 158, p. 656) and

1 SchUbert, Parenchymscheiden in den Bl., Bot. Centralbl., 1897. iii, p. 307.

SOLF.REOI'P. 4 A
logo CONCLUDING REMARKS

Euphorbiaceae. In some of these plants (Portulaca, Pedis, Amarantaceae,

Euphorbiaceae) the sh~ath is enveloped by radially arranged palisade-tissue.
In connexion with our discussion of the pan~nchyma-sheaths we may mention
the occurrence of a special layer of cells, observed by Van Tieghem in the veins
of the leaf (chiefly in the Ochnaceae and Luxemburgiaceae~ but also in the
Lineae) and styled by him 'crlstarque 1;' the' cristarque is separated fromI

the epidermis by a single cell~layer and is composed of cells, which are thickened
in the shape of a horseshoe, and enclose a clustered or solitary crystal of oxalate
of lime; in the lateral veins the cristarque' is in direct contact with the
I

sclerenchyma and corresponds to the endodermis. The cristarque ' is found

I

also in the petiole and axis.

§ 8. STRUCTURE OF THE MARGIN OF THE LEAF2. The peculiarities in the
structure of the margin of the leaf may be used especially for the purposes of
specific diagnosis, more rarely (see Berberideae, pp. 45, 820) for generic diagnosis.
The margin of the leaf is diStinguished either by a stronger development of the
leaf· tissue or (very commonly) by the considerable size of its epidermal cells
or by the development of a hypoderm, which is differentiated either as collen-
chyma or resembles hard bast (marginal bast); in the latter case the hypoderm
often enters into connexion with the marginal vascular bundles. Marginal
bast has been observed, for example, in certain Berberideae, Frankeniaceaet
llicineae, Melastomaceae, Vacciniaceae, Ericac~ae and· Myrsineae.
§ 9. SPICULAR CELLS (including similar elements occurring in the axis).
The presence of spicular cells (Le. mechanical cells having a parenchymatous or
prosenchymatous (fibJ;ous) shape) in the: mesophyll is a widely distributed
feature, which is in general only of value for specific diagnosis. In most cases
the spicular cells arise from cells of the mesophyll and only very rarely from
epidermal cells. The latter applies to the spicular cells, found in certain species
oJ. Capparis; the greater part of the body of these elemen;s is situated in the
mesophyllt but the upper part of each spicular cell is wedged in between the
epidermal cells or even projects beyond the level of the epidermis like a hair
(Fig. 18, C, p. 68).
The differentiation of the spicular cells varies very considerably. In the
simplest case they are ordinary stone-cells, those situated in the palisade.tissue
being elongated like rod-cells; in other cases they give off branches in all
directions or are elongated in a columnar manner, spicular cells of the latter
type often extending from one epidermis to the other and even branching
beneath the epidermis. In other cases again they are fibrous and then generally
branch off from the sclerenchymatous sheaths of the veins; such spicular cells
traverse the mesophyll in VarIOUS ways (their course often being very irregular)
and frequently form a complete plexus beneath the epidennis on the two sides
of the leaf. The spicular cells and especially those which are differentiated like
9

fibres, for the most part have thick walls; in certain Euphorbiaceae the walls
show well-marked stratification. The following are special forms of spicular
cells: (a) the fibre~like spicular cells of Pelliciera (Ternstroemiaceae), which have
pointed ends and run parallel to one another in the mesophyU in two planes,
parallel to the surface of the leaf; (b) the spicular cells of Hemiboea Henryi,
Clarke (Gesneraceae), which resemble rod-cells, are placed parallel to the surface
of the leaf and are covered by a group of upper epidermal cells of relatively
small size; (c) the sac~like spicular cells of LinocieT€l glomeyata, Pohl (Oleaceae),

l See Van Tieghem, Cristarqu~, &c., Boll. Mas. d'hist. nat., '902, p. 266, and the rem2illing
papers by thil author, ~ited in the list of literature (ot the Ochna.cel.e, p. 869-
:II Hintz, Meehall. Bau d. Blataandes, Nova Acta Acad. Leopold.~Carol •• liv, 189C, P.97 and
tab. v-vii.
 STRUCTURE OF THE LAMINA OF THE LEAF 1"91

which frequently fork and have wide lumina and relatively thin walls, the latter
being wrinkled owing to the presence of transverse folds; (d) tl~e branched
spicular cells of Heptacyclum Zenkeri, Eng). (Menispermaceae), which are likewise
sac·shaped and have wide lumina and relatively thin walls; (e) the bundles of
sclerenchymatous fibres running in a subepidermal position in the leaf of the
Tbeophrasteae and Polygonum equisetilorme; (I) the groups of sclerenchy·
matous fibres, developed beneath the hypoderm of the leaf in the E"yngias
having a Monocotyledonous habit; (g) the sclerenchymatous fibres of certain
Sapotaceae and Penaeaceae (Fig. 175, p. 723), which are provided with spiral
thickening; (h) the parenchymatous or prosenchymatous spicular cells of certain
Mimoseae(Fig. 6S,A,B, p. 295), Melastomaceae (Fig. 79, E,F, P.362),Compositae
and Euphorbiaceae (Fig. 180, p. 748), which occur in connexion with simple tri.
chomes or shaggy hairs of complicated structure and form a foot to the hair
(see also § 33); (i) the spicular fibres penetrating into the glandular shagl?}'
hairs of Begonia imperialis; (i) the sclerenchymatous rod-cells of certam
Melastomaceae, which lie in the parenchyma, accompanying the larger veins,
and are placed parallel to the surface of the leaf; (k) the branched spicular cells
of Cynometra (Caesalpinieae), which are filled with siliceous contents; (1) the
branched spicular cells of certain Begonias (Fig. 87, B, p. 400), which have wide
lumina and mostly include solitary crystals of oxalr..te of lime; (m) the spicular
cells which have been observed in many Loranthaceae; these elements are
mostly branched, and their walls are thickened to such an extent that only one
or more peripheral portions of the lumina remain, these being filled by solitary
crystals of oxalate of lime (crystal-sclerenchyma, Fig. 177, p. 728); (n) lastly,
the branched spicular cells, which are differentiated as 'internal hairs.' In the
Nymphaeaceae (Fig. II, A, p. 47) the elements last named (the occurrence of
which is indicated in the following enumeration by the insertion of a *) are fre·
quently distinguished by being encru5ted with small crystals of oxalate of lime;
in Rhizophora (Fig. 74, p. 340) and Nymphaea they sometimes have the shape
of an H.
The special forms of spicular cells are by no means characteristic of certain
Orders or genera. In Orders like the Capparideae or Ternstroemiaceae, or genera
like Linociera (Oleaceae) or M ouriria and M emecylon (Melastomaceae), in which
spicular cells are of frequent occurrence, different forms of these elements are
found from species to species. We qlay add that spicular cells, which traverse
the leaf in a vertical direction, may give rise to transparent dots in the Saple.
With the spicular cells we may class the I cristarque '-cells (see the -end of
§ 1, p. 1090), observed in the mesophyll of certain Ochnaceae and Lineae, and the
peculiar groups or rows of cells, which. occur in the mesophyll and cortical
parenChyma of certain Convolvulaceae (Fig. I28, p. 564); the function of these
groups of cells is still quite unknown.
Spicular cella have been observed in the mesophyll in certain species of the
following, Orders.,. and genera respectively: Dilleniaceae, Magnoliaceae, *Trocho-
dendraceae, Anonaceae, *Menispermaceae, *Nymphaeaceae. Capparideae (Fig. 18,
p. 68), Bixineae, Polygaleae, Vochysiaceae. Frankeniaceae. Tamariscineae, Gutti-
ferae, *Temstroemiaceae (Fig. 29. p. 129). Microsemma, Dipterocarpeae. Malva-
ceae, Rhaptopetalaceae. Lineae, Humiriaceae, Malpighiaceae. Zygophylleae, Ruta·
ceae, Simarubaceae,Ochnaceae sens. str., Luxemburgiaceae, Meliaceae, Chailletiaceae,
Olacineae. Ilicineae, Celastrineae, Hip~crateaceae, Rhamneae, Sapindaceae, Hippo-
castanaceae, Papilionaceae, Caesalplnieae. Mimoseae, Rosaceae,' S~fragaceae,
Hamamelideae. Ostrearia, Bruniaceae, ·Rhizophoraceae (Fig. 7-4, p. 34'0), Cc¥nbreta-
ceae, Myrtaceae sens. str., Lecythidaceae. Melastomaceae, Lythririeae. Samyda.
ceae, Passifloraceae. Begoniaceae (Fig. 87, p. 400), Datisceae. Cornaceae (Fig. 99,
p. 435). Rubiaceae, Candolleaceae, ·Goodeniaceae, Vacciniaceae, Ericaceae, Epacri-
deae. IJIPlumbagineae, Myrsineae, Sapotaceae, Ebenaceae, Styraceae, Oleaceae
(Fig. 119, p. 523), Apocynaceae, Asclepiadeae, ·Loganiaceae, .Gentianeae, Bora·
4A2
 CONCLUDING REJfARKS

gineae, Convolvulaceae, Solanaceae (not typical), Gesneraceae, Bignoniaceae~ Sela·

gineae, Polygoneae, *Myristicaceae, Monimiaceae, Laurineae, ·Proteaceae (Fig. 173.
p. 7 1 3), Thymelaeaceae, Penaeaceae (Fig. 175. p. 723). Loranthaccae (Fig. 177,.
p. 728), Euphorbiaceae (Fig. 180, p. 748), l\:(oraccae, Didymeles.
In connexion with this enumeration we may mention that extensive groups of
sclerosed or at least thick-walled tissue occur in the mesophyU (especially in the
spongy tissue) in certain Magnoliaceae, Menisperma.ceae, Berberideae, Capparideae,
Polygaleae, Tamariscineae. Guttiferae, Lineae, Celastrineae, Sapindaceae, Connara~
ceae, Melastomaceae, Vacciniaceae, Styraceae, Orobanchaceae, Pip eraceae , and
Chloranthaceae.
Spicular cells, similar to those found in the mesophyll, occur also in the
pith, primary cortex, and (more rarely) bast of the axis, and may therefore be
discussed here, at once. Branched spicular cells, some of which are differentiated
as internal hairs, are present in the axis in certain species of the Magnoliaceae~
Trochodendraceae. Nymphaeaceae, Ternstroemiaceae. Lopkira, Rutaceae, Meliaceae,
Hippocrateaceae, Rhizophoraceae, Lythrarieae, Datisceae, Primulaceae, Loga~
niaceae. Gentianeae, Chenopodiaceae, and Gomortegaceae; as a rule their presence
corresponds to the occurrence of spicular cells in the mesophyll, and this applies also
to the crystal-sc1erenchyma found in th~ cortex I of certain Loranthaceae. It is
an interesting fact that the sclerenchymatous fibres, which are found running
irregularly in the mesophyll in many species-, do not occur in the cortex in these same
species, such elements being extraordinarily rare in this part of the plant; itis only
in the mesophyll, of course,that they are of function;il imporlance. It remains to
mention the following special types of spicular cells observed in the axis ~
(a) the crystal-sclerenchyma. found in certain genera of the Schizandreae (Fig. 4p
p. 29) and Rubiaceae. as well as in the Combretaceous genus Macl'optc1'anthes; the
crystal-sclerenchyma in these forms develops in the same way as in the Loranthaceae
(see above), but the shape of the elements is difierent and subject to variation;
(b) the elongated sclerenchymatous cells. occurring in the pith in certain Dilleniaceae ..
Combretaceae, and Loganiaceae; (c) the sClerenchymatous fibres. which run in
various directions in the primary cortex of species of Pl'unus {Rosaceae 1) ; (d) the
well-known spicular cells. found in the bast of the Cinchona-bark, and the elements
of a similar shape, occurring in Ancz"stl'ocladus and Styrax (Styraceae); (e) the long
sclerenchymatous cells present in certain Apocynaceae. these elements having thick
walls and a tuberculate surface; (/) the bast-fibres of the Quebracho-bark (Apocyna-
ceae), which are completely enveloped by a sheath composed of a single layer of
~Jls, each containing a solitary crystal. For the occurrence of ordinary stone~cells,
groups of stone-cells, 'cristarque '....cells, and groups of sclerenchymatous fibres in the
pith or primary cortex, cf. §§ 40 and 5I; see also § 53. .

§ 10. ENLARGED TERMINAL TRACHEIDES 2. Enlarged tenninal tracheides

(i.e. cells, which have wide lumina and are either spirally thickened or have thick
pitted walls) are in general only of specific value; they are found at th~ ends-of
the vascular bundles of the veins, and function as water-reservoirs.
They have been observed in certain Capparideae (Fig. 18, p. 68). Polygaleae p

Tamariscineae~ Hypericineae, Guttiferae. Ternstroemiaceae, Geraniaceae, Rutaceae,

Olacineae, Celastrineae, Papilionaceae, Mimoseae, Rosaceae, Rhizophoraceae, Com-
breta.ceae, Lythrarieae, CoI1!Positae. Salvadoraceae, Gentianeae - Menyanthoideae,
Loranthaceae, Santalaceae, Euphorbiaceae and BalanopSeae 21.

§ II. STORAGE-TRACHEIDES IN THE GROUND TISSUE OF THE LEAF AND AXIS.

The storage-tracheides found in the ground tissue have the same physiological
function and the sa.me systematic value as the terminal tracheides. In some cases
entire groups· of tissue are composed of such elements, c.g. in the axis of Oudneya
africaM, R. Br. (Cruciferae) and in the root of certain specie, of Drosera, in

1 Moller, RinderutJlatomie, p. 369 et seq.

IIHeinricher, in Bot. Centralbl., J885, iii. p. 2'5.
S The above enumeration is but an incomplete one, owing to the fact that the term I enh.rged
terminal tracheides' has Dot always been interpreted in the same sense by different authors.
 STRUCTURE OF THE LAMINA OF THE LEAF 1093

which the outer cortical tissue is formed by cells with reticulate or spiral
thickening; in this relation we may once more recall the reticulate thickening
of the palisade-tissue of Clusia rosea and other features, already discussed in
§ 6. In the majority of cases the storage-tracheides are more or less isolated,
occurring either in the middle of the ground tissue or in connexion with the
vascular system or the secretory organs.
Special storage-tracheides, in the form of narrow sac-shaped elements with spiral
'thickening, adjoin the ends of the veins in Vochysia ~ulaJ Mart., and penetrate from
these points as far as the epidermis of the leaf; short spiral tracheides are found in
the pith of Anacampseros (Portulaceae); isolated spiral and annular tracheae are
present in the pith of. Impatiens (Geraniaceae). Wide sac-shaped spiral tracheid«:s
(occasionally side by side with sclerenchyma) accompany the secretory canals of
CaJophyllum (Guttiferae, Fig. 27, D, p. 122), which run freely in the mesophyll; a
similar relatigD to secretory cavities is shown by the spiral tracheides. which branch
off from the vascular bundles of the veins in Caraipa (Ternstroermaceae, Fig. 29, E,
p. 129) and Haplophyllum (Rutaceae). The spirally thickened spicular fibres,
running irregularly in the mesophyU in certain }:"'enaeaceae (Fig. 175, p. 723), and the
spirally thickened spicular fibres of certain species of MicfOPholis (Sapotaceae) have
already been briefly referred to in § 9. The spiral fibres of A-!icropholis are closely
related to the tubular spiral tracheides, developed in the mesophyll of Ochthocosmos
(Lineae), Aciotis annua, Tr. (Melastomaceae). Macyocnemum and Pentagonia (Rubia-
ceae), the Nepenthaceae (Fig. 164, A, B, p. 678) and Pogonophora (Euphorbiaceae),
as well as in the pith and cortex' of the Nepenthaceae, and in the palisade-tissue,
found in the primary cortex of species of A ythrocnemum and Sal£cornia (Chenopo-
diaceae). Spiral tracheides of exactly the same type accompany the vascular bundles
of the veins in Sommera (Rubiaceae), while transitional forms between such tracheides
and ordinary sclerenchymatous fibres are found in the same position in Emmotum
(Olacineae) and Chomelia (Rubiaceae). It remains to mention the systems of tra-
~heides, running freely in the mesophyll and independently of the vascular bundles
of the veins in Petalonyx Thuyberi (Loaseae), the Opilieae (Olacineae), Nwytsia and
species of Loranthus (Loranthaceae), species of Osyris, Quinchamalium, Thesium,
and MyzodendYon (Santalaceae), and the rows of tracheides, situated on either side
of, and parallel to, the cortical vascular bundles in the axis of the Casuarineae
(Fig. 186, p. 788). Regarding the nature of the storage-tracheides found in Boscia
(Capparideae) and spec;ies of Bellucia, Henriettea, and Sonerila (Melastomaceae).
see under the Orders named.

II. STRUCTURE OF THE PETIOLE.

§ 12. The STRUCTURE OF THE PETIOLE was already employed for

systematic purposes 1 at an early date, out its systematic value has frequently
been overestimated. At the outset it was to be presumed that the number of
vascular bundles in the petiole and to some extent also their arrangement
(including the occurrence of medullary and cortical vascular bundles) would
depend on the dimensions of the leaf; it has moreover been shown that as a
general fule the vascular bundles of the petiole are isolated in herbaceous species
and fused in various ways in woody species.
-.-.~----------------

1 C<J~. de Candolle, in Mem. Soc. de phys. et d'hist. nat. de Geneve, t. xxvi,;I' part., 1879. p. 427
and pl. I, 2; Plitt, Blattstiel d. Dicot., Diss., Marburg, 1886, 52 pp. and 1 Tab. i Petit, in Mem. Soc.
se. phys. et nat. de Bordeaux, Set. 3, t. iii, 1887, p. 218, and 6 pl., and in Act. Soc. Linn. de Bordeaux,
t. xliii, 1R89, p. It, see also in Comptes rendus, t. civ, 1886, and Ann. IC. nat., ~er.,i, t. vi, 1887.
p. 342; Lignier, in Bull. Soc. Linn. de Norm andie, 1888, p. 81, and Comjt~s rendus", L cvii, 1888,
p ....02; Acqua, in [Ann. 1st. bot. di Roma, Vol. iii, fase. I, 1887, 3S pp. and Malpigbia, i, 1887,
p. 277; Chatin, in Bull. Soc. bot. de France, 1897, p. 464, and 1898, pp. 137, 1..f.5, 165. :.a ..P. and
310; Bouyglles, Various pa.pers in Act. Soc. Linn. :Bordeaux, lvi, lvii, and lviii, 1901-3, especially
the 'These, Paris, 1902 (Cert. formes vasco anomales OU petiole des Dicotyltd.)' contained in t. lvii;
Col, in Ann. sc. nat., sC:r. 8, t. xx, 1904, p. I, and Comptes rend us Acad. Paris, cxxsv~ 1903,
p. 516; finally, Van Tieghem's numerous sy~tematic paprrs, in which the structure o( the petiole is
-often profitably employed for systematic PQrpo;;e~.
1094 CONCLUDING REMARKS

In most cases the structural features just mentioned. can be used only for
purposes of specific diagnosis, but they are occasionally of greater systematic
vaJue. Thus, a transverse sect jon through the apex of the petiole of any of the
Dipterocarpeae shows a closed ring of wood and bast, surrounding a tissue
which resembles a pith and encloses a vascular system of varied structure;
again, the genera of the Dipterocarpeae, the subdivisions of the Lecythidaceae,
the genera of the Salicineae, the genera and generic groups of the Cupuliferae,..
can all be rustinguished with the help of the structure of their petiole.
In an investigation of the structure of the petiole it is of course requisite
that analogous portions be compared with one another (unless the entire course
of the vascular system is followed up,-an arduous task, which generally does not
repay the amount of labour involved and cannot always be undertaken owing to
lack of the necessary material), since the appearance of the transverse section
may vary at different points in the course of the petiole. Petit's investigations
in particul~ have helped to show that the most suitable points at which tc)
examine the petiole are its base and especially its apex; we may follow Petit in
describing the transverse sections taken through these particular regions respec-
tively as the' initial' (' initiale ') and ' characteristic' (' caractt~ristique '). A
point of special importance- appears to be the number of vascular bundles 1,
passing out from the stem into the' leaf or petiole, as the case may be.
An anomalous differentiation of the fibrovascular system of the petiol~
is connected with the occurrence of what are called 'rayed bundles' ('Faisceaux
rayonnes ') and I true concentric or hemiconcentric bundles' CFaisceaux con-
centriques ou hemiconcentriques vrais ')2. The first anomaly has been recorded
in the Cruciferae J. A transverse section through the petiole in this Order shows·
a tissue-system, which has a lobed outline and is composed of radially arranged
vascular bundles with intervening tissue consisting of small thin-walled cells, the
whole being surrounded by a tissue made up of large cells with thin walls; the
whole of the lobed tissue-system arises from a single procambial strand. In the
case of the second anomaly, the petiolar tissue contains several vascular bundles,.
which are completely or mcompletely concentric (with a central xylem-mass) •.
and in most cases are provided with an endodermis, distinguished by the presence
of Caspary's dots; in other words, we have a kind of polystely (d. § 55).
The second anomaly has been observed in certain Rosaceae, Saxifragaceae,
Hamamelideae, Halorageae (Gunnera pro parte), Valerianeae, Campanulaceae,.
PrimuIaceae, Gesneraceae, Acanthaceae and Labiatae.
The pulvini (l Schwellpolster '), which occur at the bases of the petioles or
of the stalks of the individual pinnae, for example in many Menispermaceae,
Leguminosae, Oxalideae and Zygophylleae, show few noteworthy points of
difference in structure 4.
On the other hand, from the syStematic point of view, it remains to consider
the sclerenchymatous tissue, accompanying the fibrovascular system on its
outer side, and perhaps also the pith-like ground tissue enclosed by the vascular
system. The sclerenchyma, accompanying the vascular tissue, shows features
SImilar to those exhibited by the sclerenchyma found in the larger veins. The
outer ground tissue, which forms a continuation of the primary cortex of the
stem, commonly repeats the structural features presented by the latter, while
the inner ground tissue shows the same type of structure as the pith.

t See also especia.lly Chatin, 11. cc. I Bouygues, loc. cit.

I Similar types of structure, shown by the fibrovascular system of the petiole, are termed
jPseudo-faisceaux rayonnes' by Bouygues; they differ (rom the 'Faisceaux rayonnes' in their
cou~ of development.
'See Haberlandt, Pbysiol. Pfianzenanat., 1904-, p. -499, and the papers by Schwendener,
Haberlandt, Mobius and Pantanelli there cited (under II); also Preuss, Bezieb. zwischen dem anat.
B:lu u. der physiol. Funktion d. Blattstiele 0.. Gelenkpolster, Diss., Berlin, r885. 38 pp.
 1095

III. SECRETORY AND EXCRETORY RECEPTACLES.

§ 13. GENERAL POINTS ABOUT SECRETORY RECEPTACLES 1. The great

systematic value of the secretory receptacles was recognized at an early date.
In employing these elements for systematic purposes we have ill the first place
to determine their nature (secretory cells, secretory cell-fusions, secretory
cavities, secretory canals), but, apart from this, their contents 2 and their position
within the plant also come into consideration. All these features are of great
importance; they are often characteristic of entire Orders or at least of tribes
or genera, although in some cases they only serve to distinguish species.
As regards the occurrence of the secretory receptacles in the different parts
of one and the same. species, we may note at this point that in most cases the
same types o{secretory receptacles are present in all the vegetative and repro-
ductive organs of the plant. We do, however, sometimes find that the secretory
receptacles are confined to certain organs of the plant (such as stem, leaf, root
or flower), or that different types of secretory receptacles are present in the
different parts of the plant. Thus in the Hypericineae, for example, the secre-
fory cavities of the leaf are replaced by secretory can~ in the axis, and similarly
in the Rutaceae the secretory cavities of the leaf give way to secretory cells
in the bast of the axis. In the same way as different types of secretory organs
may occur in one and the same species, various kinds of internal secretory
organs may be found in different species or genera belonging to the same Order.
Apart from those secretory receptacles which are typically differentiated,
attention should also be paid to the distribution of tannin in the ordinary cells
of a tissue. Very striking features are afforded by the way in which the dry
cortex or mesophyll is filled with reddish-brown phlobaphenes in herbarium-
material-of a certain species. genus or Order, while such substances are wanting
in other species, &c.
The following is an enumeration of the Orders and genera in which special
internal secretory organs I have not hitherto been observed: Ranunculaceae,
Dilleniacea.e?, Berberideae, Sarraceniaceae, Cistineae, Tremandreae, Polygaleae?,
Frankeniaceae, Caryophylleae. Tamariscineae. Lophi~a, Rhaptopetalaceae ?, Lineae,
Humiriaceae, Malpigbiaceae ?, Ochnaceae incl. Luxemburgiaceae, Chailletiaceae ?,
Octocnemaceae, llicineae, Cyrilleae, Pentaphylacaceae, COrynocarpaceae, Sta.ck-
housieae ?, Melianthaceae, Staphyleaceae, Sabiaceae, Coriarieae, Crossosomataceae,
Droseraceae, Ostrearia, Bruniaceae. Halorageae, Rhizophoraceae, Melastomaceae
{apart from tannin-idioblasts). Onagrarieae, Loaseae, Turneraceae? (apart from
tannin-cells), Cucurbitaceae?, Begoniaceae? Datisceae, Calycereae, Candolleaceae?,
Vacciniaceae, Ericaceae, Epacrideae. Diapensiaceae. Lennoaceae. Plumbagineae.
Ebenaceae, Oleaceae, Salvadoraceae. Loganiaceae, Desfontaz'nea, Plocosperma, Gentia-
neae? (apart from secretion contained in intercellular spaces). Polemoniaceae (apart
from secretion contained in intercellular spaces), Hydrophyllaceae, Boragineae"
Nolaneae, Solanaceae ?, Retzia, Scrophularineae (apart from tar~ain-idioblasts).
Orobanchaceae; "Leiltibulaiieae, Columelliaceae, Bignoniaceae, Pedalineae, Acan-
thaceae ?, Zombiana, Selagineae, Plantagineae, Illecebraceae, Amarantaceae, Cheno-
EO<liaceae. Phytolaccaceae, Batideae, Nepenthaceae, Thymelaeaceae, Penaeaceae,
Geissoloma, Elaeagnaceae, Santalaceae, MyzodendYon, Champereia, Grubbia, Balano-

I The term • secretory receptac:les' is here understood to comprise cells, cell-fusions, or inter-
ce:1ular spaces, which are filled with a secretion.
I Mangin. Classification des mucilages, Bull. Soc. bot. de France, 1894. p. xl; Tschirch, Die
Hane u. die Harz,behalteT, Leipzig, 1900; Moliscb, Milcbsaft u. Scbleimsaft, Jena, 1901.
:I In this enumeration gelatinized epidermal cells and tne!!ophyll-cells provided with local
swellings (for these d. U :I and 6) are not taken into consideration; ordinary tanniniferous cells,
that is to say cells, which are not distinguished by their shape or Si1.e, and 3rt filled with brown
contents in herbarium-material, are likewise not taken into account.
10g6 CONCLUDING REMARRS

phoreae, Daphniphyllaceae, Balanopseae, T~elygoneae, Platanaceae~ Juglande~,

M yricaceae ?, Casuarineae, Cupuliferae, Salicineae, Lacistemaceae, Empetraceae,
Ceratophylleae (apart from tannin-cells).
§ 14. SECRETORY CELLS 1. This term is taken to include all such cells as
contain secretion and appear as idioblasts,owing to their shape or their contents,
or both shape and contents, with the exception of mucilage-cells and laticiferous
cells, which will be discussed separately in §§ IS and 20.
The shape of the secretory cells varies. In some cases they appear as dis-
tinct idioblasts, which are spherical, ellipsoidal, branched or sac~shaped, and
vary in length, while commonly giving rise to pellucid dots in the leaf; in other
cases, however, they differ but little from the neighbouring cells or are distin-
guished merely by their contents. The sac-shaped secretory cells are frequently
arranged in rows, while the spherical secretory cells of the leaf are often replaced
by elongated cells in the axis (especially in the bast) of the same species. The
contents are resinous, oily, gummy, or resemble latex or caoutchouc; in other
cases they are tanniniferous, and are t~en coloured brown in herbariumMmaterial.
The cell-wall is occasionally suberized or provided with a suberized lamella.
The detailed structure of the secretory cells of certain Magnoliaceae, Canellaceae.
Aristolochiaceae, Piperaceae and Laurineae, which are filled with a resinous or ~ily
5ecretion, is specially noteworthy; their secretory contents are enveloped by
a thin-walled sheath (' pouch '), which is formed from the wall of the vacuole
by a change in its substance, and is· connected with the cell-wall by means of a
stalk (' basin' or • funnel '), which constitutes a thickening of the membrane'.
These secretory cells are doubtless very closely related to the (paired) cystospheres
of certain Begoniaceae a, which are dealt with In § 28. The secretory cells of Pogo-
stem on Patchouli (Labiatae), which are differentiated as true internal glandular hairs,
may also receive special notice at this point; they are provided with a short stalk,
mostly composed of two or three low cells having suberized walls, and project
into the intercellular spaces; the abundant secretion accumulates chiefly between
the cuticle, which is raised like a bladder, and the cellulose-membrane.
As regards the position of the secretory cells in the different tissue-systems it
may be pointed out that the occurrence of these elements in the epidermis (especially
in the leaf) has proved to be of systematic value. I t must, however, be borne in mind
that secretory cells. which are apparently situated in the epidermis, but belong to
the subepidermal tissue (having pushed their way secondarily between the epidermal
cells by a process of sliding growth), can in some cases be distinguished with certain~
from true epidermal secretory cells only by a study of their course of development.

Secretory cells sometimes characterize entire Orders, these being indicated

by a * in the following synopsis; in other cases they are at least distinctive of
genera or species.
I. Secretory cells, of various shapes, with resinous (oily). latex·like or
other contents (apart from very much elongated secretory cells. filled with similar
contents, which are enumerated under II). have been observed in the following
Orders and genera respectively: *Calycanthaceae (resin-cells, rounded in the leaf),
·Magnoliaceae (resin-cellS, rounded in the leaf and occasionally also in the wood),
Trochodendraceae (branched and elongated resin-cells), *Lactoridaceae (resin-cells,
rounded in the leaf), • Anonaceae (resin-cells, rounded in the leaf and occasionally
also in the wood), Menispermaceae (sac-shaped secretory cells of varied length and

1 De Bary, Vergl. Anat., 1877, p. 152; Zacharias, in Bot. Zeit., 1879, pp. 611 and 633;
Biermann, Olzellen, Diss., Bern, 1898, also in Archiv d. Phnnn., 1898, p. 74.
S Berthold, Protoplasmamechanik, Leipzig, 1886, p. 25; Haberlandt, Physiol. PflanzenanAt.,
190... , p. 462; R. Muller, in Ber. deutsch. bot. Gesellsch., 1905, p. 292.
S Cf. the detailed statements on this point in Fellelel', Beitr. z. Anat. u. Syst. d. Begoniaceae,
Diss., MOOchen, 1892, p ..... r et seq. and Tab. i; this paper also deals with the course of development
()f these structures.
, See R. Miiller, in Ber. deutsch. bot. Cesellscb., 190:'. p. 293.
 SECRETORY AND EXCRETORY RECEPTACLES 1097

provided with brown, gummy or other contents in the leaf and axis), ·Nymphaeaceae
(isodiametric or elongated laticiferous sacs, occasionally arranged in rows), Papa~
veraceae (laticiferous sacs, as in the previous case), Fumariaceae (sac-sha~d cells.
often very long and likewise arranged in rows). Capparideae (Cleomeae WIth secre~
tory cells in the neighbourhood of the vascular bundles of the veins), *Canellaceae
(resin-cells, rounded in the leaf), Bixineae (sac-shaped and branched resin-cells),
Polygaleae. (oil-cells only in • Radix Senegae '), Elatineae (resin-cells), Ternstroe-
miaceae, Ancistrocladus (cortex), Tiliaceae (rounded resin-cells), Lineae (pith and
cortex of Erythroxylon), Geraniaceae (rounded secretory cells), Rutaceae (resin-cells.
only in the axis), Simarubaceae (resin-cells, in part only in the axis), "'Meliaceae
(spherical to sac-shaped or branched resin-cells, occasionally arranged in rows).
Olacineae, Sapindaceae (often laticiferous, isolated and of varied shape or arranged
jn rows, very widely distributed), Hippocastanaceae (resin-cells, rounded in the leai),
*Aceraceae (with latex-like or other contents, probably always present in the bast
of the veins, but also occurring freely in the mesophyl1 and in the axis), Connaraceae,
Papilionaceae, Caesalpinieae, Mimoseae (various types, occasionally with gelatinized
membranes), Saxifragaceae (elongated secretory cells only in the pericycle of AbYo-
phyllum), Hamamelideae (Altingieae, not differentiated as idioblasts), Myrothamnu.c:
(spherical resin-cells), Myrtaceae-Lecythidaceae ? (Napo/eonea), Samydaceae (very
rare, in Casearia, here side by side with secretory cavities), Cornaceae (only in the veins
of the leaves of species of Nyssa), Rubiaceae(resin-cells),Valerianeae (only in the root),
Compositae, Candolleaceae (indistinctly differen tiated), Plumbagineae (cells in the stem
·and root filled with plumbagin), Primulaceae (red contents, in part only in the root),
Myrsineae (red contents), "'Sapotaceae (laticiferous, the latex occasionally containing
crystal-sand; elongated cells, sometirdes arranged in rows), Bf'achynema (blood-red
contents in rows of cells, situated in the primary cortex), Apocynaceae (rare, in the
cortex; also a layer of secretory cells beneath the palisade-tissue in certain species).
Asclepiadeae (Solenostemma), Convol vulaceae (frequently laticiferous, solitary secre-
tory cells and rows of secretory cells, the latter almost of general distribution),
"Cuscuteae (resin-cells), Verbenaceae (Confea and Symphorema), Labiatae (sac-shaped
resin-cells; in Pogostemon in the form 0 typical internal glandular hairs), Polygo-
naceae? (probably merely secretory cavities), "'Aristolochiaceae (rounded resin-cells,
of frequent OCCurrence in the epidermis of the leaf, occasionally as basal cells of the
hairs, not always present in the leaf), *Piperaceae (rounded resin-cells), *Chlorantha-
ceae (as in the previous case), *Myristicaceae (as in the Piperaceae, occasionally
with gelatinized membranes), "'Monimiaceae (as in the Piperaceae), ·Laurineae (as
in the Piperaceae, occasionally also in the wood), "'Hernandiaceae (as in the Pipera-
ceae), "'Gomortegaceae (as in the Piperaceae). Euphorbiaceae (rows of laticiferous
sacs, and oil-cells~ the latter occasionally differentiated like hairs, Fig. 180, p. 748),
Buxaceae {not very typically differentiated), Moraceae (spherical secretory cells),
Myricaceae (rare). We may add that the rows of laticiferous sacs found in certain
Sapotaceae and Convolvulaceae show transitions to laticiferous vessels (see § 21).
We may also mention the following elements at this point: (a) cells, containing
myrosin (myrosin-cells 1) and differing more or less from the neighbouring cells,
probably occur in all Cruciferae, and have been recorded also in certain Capparideae,
Resedaceae. and Geraniaceae. as well as in Bt'eischneidet'a, M01'inga, and the genus
Scorodophloeus (Caesalpinieae); (b) elongated spicular cells,·some of which at least
are filled with contents, are found in the pith of certain Dilleniaceae and species of
CombYetum; (c) peculiar strands of secretory cells, which appear as ribs even to the
naked eye, occur in.the axis of Stocksia (Sapindaceae), and similar strands of secre~
tory cells. in- which the inner cells subsequently become resorbed and replaced by a
secretory canal, are present in Pl'osopanche (Cytinaceae); lastly (d) groups of glan-
dular cells have been observed in the mesophyll in Heterophyllaea (Rubiaceae).
II. Elongated secretory sacs have been observed in the pith, bast, or pericycle
of the stem, and occasionally also in the larger veins of the leaf in certain members

1 In addition to the papers oC Heinricher, Gllignard, Spanier, Raillkofer, &c., which are cited
in the lists of literature belonging to the respective Orders, see especially Hartwich. Bubimbirinde,
Zeit. d. deutsch. Apothekerver., 1902, p. 339. In this paper sulphur-containing oils are stated to
occur in certain members of the followin~ Orders: Bombaceae (Malvaceae). Caesalpinieae,
Capparideae, Caricaceae (Passifioraceae). Cmcjferae. Enpborbiace.ae, Labjatae~ Limn3nlbaceae
(Geraniaceae), Meliaceae, Moringeae, Phytolaccaceae, Resedaceae. Rubiaceae, Tropaeolaceae
(Geraniaceae). Urnbelli{erae.
 CONCLUDING REMARKS

of the Orders enumerated below; most of these elements resemble the well-known
tannin-sacs, found in the pith of the elder, but in some cases they are shorter, anct.a.re
then often arranged in rows; their contents are mostly brown and tanniniferoU5
or latex-like or resinous. The Orders in question are: Trochodendracea.e,
Menispermaceae, Berberideae? J, Hippocrateaceae, Anacardiaceae, Connaraceae,
Papilionaceae (very widely distributed, Fig. 56. p. 259), Mimoseae (with diverse
contents), Chrysobalaneae, Crassulaceae, Lecythidaceae, Passifioraceae, Capri folia-
ceae, Rubiaceae{in part resembling bast-fibres), Dipsaceae, Compositae, Polygona-
ceae, Myristicaceae. Monimiaceae, and Euphorbiaceae. In certain Compositae
(Gazania, Atracty/is, and Carlina) one finds transitional forms between the elements.
in question and laticiferous vessels; in the group Compositae-Cynaroideae the
secretory sacs and secretory canals are vicarious in their occurrence in one and the
same sfccies (for details, see under Composi tae. pp. 959, 960). See also § 20, p. I l o
under secretory organs resembling laticiferous cells.'
t
Similar long secretory sacs with brown contents have been observed in the
epidermis of the leaf (cf. § 2) in certain Violarieae, Geraniaceae, Saxifragaceae.
Crassulaceae and Euphorbiaceae~ and in the uppermost layers of the mesophyll in
certain Crassulaceae and Ficoideae, while sac-shaped mucilage-cells occur in the
epidermis of the leaf of Tt'opaeolum (Geraniaceae). .
Tannjn-idioblasts, the shape of which is similar to or differs from that of the ele-
ments previously discussed, are found in the leaf (h.ere oommonly in the form of en-
larged cells of the palisade-tissue) or axis. in representatives of the following Orders :
Violariea.e (mesophyU), Zygophylleae(mesophyll), Ge.raniaceae (mesophyU), Chaille-
tiaceae (mesophyll), Celastrineae (mesophyll), Hippocrateaceae (mesophyll and bast
of th~ axis), Stackhousieae (leaf and axis). Rhamneae (mesophyU), Didiereae (leaf
and axis), Papilionaceae (leaf and axis), Melastomaceae (leaf), Turneraceae (scattered
groups of tannin-cells in the leaf, idioblasts in the cortex), Passifloraceae (mesophyll),
Rublaceae (mesophyll and bast). Monotropeae (axis). Gentianeae (rhizome), Sola-
naceae (root). Scrophularineae (mesophyll), ,Nyctagineae (enlarged cells in the epi-
dermis of the leaf), Polygonaceae (branched cells in -the cortex), Aristolochiaceae
(mesophyll), Piperaceae (mesophyll). Euphorbiaceae (leaf and axis), Moraceae
(mesophyll). Ceratophylleae (cortex and mesophyll).
§ IS. MUCILAGE-CELLS. The gelatinization of epidermal and hypodermal
cells has already been discussed in §2 (p. 1074), and at this point I propose to con-
sider only those mucilage-cells which occur outside the integumental tissue. The
mucilage contained in these elements is in almost all cases membrane.mucilage.
which arises by the gelatinization either of the entire cell-wall or only of a part
of it. The mucilage appears to be derived from the contents only in the Ampe-
lideae, Onagrarieae, and Rubiaceae, in which the receptacles in question can be
interpreted as incompletely differentiated raphide-sacs (i.e. without raphides) ;
the much-discussed mucilage of the Cacteae is probably membrane-mucilage.,
Mucilage-cens have been demonstrated in the tissue of the leafin certain species
of the followiD~ Orders and genera: Magnoliaceae, Anonaceae. Cappariaeae?,
Resedaceae, Bixmeae, Portulaceae, Ternstroemiaceae, Strasbuygeria, Dipterocarpeae
(alsoceJls having a gelatinized membrane and containing a solitary crystal), Chlaena-
ceae, Malvaceae. Sterculiaceae, ZygophyUeae, Simarubaceae, Eutiiem~s, Burseraceae,
Meliaceae, Olacineae, Hippocrateaceae. Rhamneae~ Ampelideae, Didierea, Aceraceae.
Connaraceae ? Papilionaceae, Caesalpinieae, Rosaceae, -Rhlzophoraceae (Fig. 74,
p. 340 ), Melastomaceae, Lythrarieae, Ona.grari.eae~ Rubia.ceae, Apocynaceae,
Gentianeae, Acanthaceae, Basellaceae, Phytolaccaceae, Laurineae (very widely dis·
tributed, absent in the related Monimiaceae), Hernandiaceae, Gonystylus, Octolepis,
Ulmaceae, and Moraceae. Regarding gelatinization of the spongy tissue and peculiar
thickenings of the cells of the mesophyll, which are restricted to certain parts of the
cell-wall and have a mucilaginous appearance, see § 6, p. 1088.
Mucilage-cells are present in the axis (pith, bast, or primary cortex) in certain
species of the following Orders or genera: Schizandreae. species of Tovaria (?),
Resedaceae, Violarieae, Bixineae. Vochysia.ceae, Portulaceae, Stt'Qsbut'geria, Diptero·
carpeae, Monotes, Chlaenaceae, Malvaceae, Sterculiaceae, Tiliaceae, Simarubaceae,.
Euthemis, Burseraceae. Chailletiaceae, Pentaphylacaceae, Rhamneae, Ampelidaceae ..
1 Holm has lately described • secretory ducts' in CattlopkJlllum Ikaliciroitk!, Michx. (see
Mt't"clc's R("port xvi. 1907, pp. 9.-6.)
 SECRETORY AND EXCRETORY RECEPTACLES 1099
Didiereae,- Staphyleaceae, Papilionaceae, Rosaceae, Saxifragaceae, Cacteae. Apocy-
naceae, Basellaceae, Laurineae, Thymelaeaceae (Gonystylus, Octolepis), Euphorbiaceae,
Ulmaceae.
§ 16. MUCILAGE-CAVITIES. Mucilage-cavities generally arise from groups
of gelatinized cells, and it is frequently impossible to draw a sharp limit between
the former and the latter; similarly, one finds transitions between mucilage-
cavities and. mucilage-canals. It remains doubtful whether the rnu~e­
cavities, which are situated above the xylem in the veins of certain Specl~
of Terminalia (Combretaceae) and are provided with an epithelium, are schizo-
genous in origin.
Mucilage-cavities have been observed in certain: Magnoliaceae? (veins of the
leaf). Dipteroca~ae (primary cortex, petiole), Monotes, Chlaenaceae, Malvaceae
(pith, cortex, petiole), Triplochitonaceae (cortex, veins of the leaf), Sterculiaceae (as
in the Malvaceae), Tiliaceae (as in the Malvaceae, the cavities sometimes resembling
canals), Zygophylleae (cortex. the contents including crystals), Rutaceae (primary
cortex; in the wood in Evodia 1'utaecarpa, Hook. f. et Th. and Zanthoxylon Budyunga,
Wall.), Simarubaceae (pith, prim. cortex, petiole, and veins of the leaf). Olacineae (in
the soft tissue of the axis of Phytacrene). Rhamneae (widely distributed), Didie,ea
(prim. cortex), Alluaudia (prim. cortex), Anacardiaceae (prim. cortex). Morin~eae
(cortex), Connaraceae (prim. cortex), Papilionaceae (secondary cortex and pith),
Caesalpinieae (gum.lacunae in the wood of Burkea atricana) 1, Combretaceae (~hloem
of the axis and veins of the leaf), Apocynaceae (various tissues of the axIS, also
in the leaf), Asclepiadeae (cortex of the root), Euphorbiaceae (cortex, see f IS).
§ 17. MUCILAGE-CANALS. The mucilage-canals develop schizogenously or
(as in most cases) lysigenously. The presence of a distinct epithelium in the
mucilage-canals is no more an absolutely certain criterion of their schizogenous
origin, than it is in the case of the secretory cavities and secretory canals, to be
discussed in §§ 18 and 19. The mucilage-canals occur in the pith and primary
cortex; in the older axes of certain Sterculiaceae and of the PapHionaceous
genus Herminiera they are found in the wood. In some cases they are present
also in the veins of the leaf.
The occurrence of mucilage-canals is confined to certain: Violarieae, . Bixineae,
Vochysiaceae, Guttiferae, Quiineae, Malvaceae. Sterculiaceae (sometimes in the
wood),Olacineae, Rhamneae, Ampelidaceae. Moringeae, Papilionaceae (in the wood),
Rosaceae, Combretaceae, Lecythidaceae, G.acteae. Piperaceae, Chloranthaceae,
Laurineae (Cassytha). Loranthaceae, Moraceae and Urticeae.
I 18. SECRETORY CAVITIES 2, The term secretory cavity is here used to
comprise intercellular secretory spaces, which are spherical or ellipsoidal in
shape and have contents, which are not mucilaginous (see § 16), but in other
respects vary in character (being resinous, oily, gummy, crystalline or brown and
tanniniferous); the mode of development of the cavities (whether schizogenous,
lysigenous or schizolysigenous) is left out of consideration. The secretory
cavities often "give rise to transparent dots in the leaf, or more rarely to what
are called' opaque' dots. They are not always equally abundant in the
different parts of the leaf, and are occasionally confined to its margin. In the
axis, they are generally found in the pith and primary cortex, and have hitheJiq
been observed in the bast, only in Casea,ia g,andi/lora (Samydaceae) and a few
species of Eucalyptus (Myrtaceae); in some cases, moreover, their place in the
axis, and especially in the bast, is taken by secretory cells (e.g. in the Rutaceae)
or secretory canals (e.g. in the Hypericineae). Conversely" secretory cavities
sometimes also occur vicariously in place of secretory canals, this being the
case in the secondary tissues of the vascular system in tlle axis of certain Com ..

t See PelTot et Gerard, in Bull. Soc. bot: de France, M~m. 6, 1901.

S De Bary. Vcrgl. Annt., 1877 p. 210.
1100 CONCLUDING REMARKS

positae. As regards the systematic value of the secretory caviti{(s, they mostly
occur in all the species of a genus and occasionally even (in the cases indicated
by a * in the enumeration below) in all the members ot an Order, and are
rarely (as in the genus Couepia of the Chrysobalaneae and the genus Cuphea
of the Lythrarieae) characteristic of species only. The determination of the
mode of origin of the secretory cavities invariably requires an investigation
of their course of development. The presence of an epithelium, lining the
fu~~ developed cavity, is not in itself a conclusive proof of its schizogenous
ongm.
The following are special forms of secretory cavities: (a) the secretory
cavities of certain Rhamneae and Papilionaceae (Fig. 57, B, p. 262), which are
provided with a papillose epithelium; (b) the secretory cavities of Polygonum
Hyd1'opiper, L., which are enveloped by ~our epidermal cells; (c) the secretory
cavities of certain Papilionaceae (Fig. 57, C, p. 262), Mimoseae and Euphorbia·
ceae, ~hich are surrounded by ~ bracket·ceIls,' and the open secretory cavities
of certain Menispermaceae and Papilionaceae, which may be classed with the
former; (d) the intramural glands of the Papilionaceous genus Pso1'alea (Fig. 57t
A, p. 262), and the glands of similar structure, found at the margin of the leaf
and in the leaf-teeth in certain species of Symplocos (Styraceae); (e) the secretory
cavities of the Proteaceous genus Franklandia (Fig. 173, C, p. 7I3), which are
traversed by rows of cells and, so far as their development is concerned, are no
doubt related to the intramural glands; (f) the fused secretory sacs of certain
Myrsineae, which arise from groups of cells, the peripheral walls of which
remain intact; and (g) the albumen·containing glands, found in the Myrsineae ;
they probably develop after the manner ofj the intramural glands and recall the
structure of the glands of F1'anklandia (see above). Secretory cavities have been
observed in hairs or emergences in certainRutaceae,Papilionaceae,Caesalpinieae,
Myrtaceae, and Lythrarieae (d. also § 34, under It a and II).
In the secretory cavities of the Rutaceae and Myrtaceae, which occupy a
superficial position, a special mechanism of a characteristic kind serving for
the emission of the secretion has been demonstrated; for details on this point,
see the Orders named, pp. 856 and 920.
Secretory cavities have been recorded in the following Orders and genera
respectively: Menispermaceae (open secretory cavities), Bixineae, Polygale~ ?,
*Hypericineae (resin~canals in the axis), Guttiferae, Ternstroemiaceae (all Bonne-
tieae excepting Arckytaea and Bonnetia. Fig. 29. p. 129). Malvaceae. Triplochitona-
ceae(secretory cavities?),l\Ialpighiaceae? (Asp£dopterys), Geraniaceae (with sphaero-
crystalline contents). *Rutaceae (secretory cells common in the bast), Simarubaceae
(in genera, which should rather be referred to the Rutaceae), Meliaceae (Ckloroxylon,
Flindersia). Olacineae, Rhamneae (secretory cavities with a papil10se epithelium).
Connaraceae (with sphaerocrystalline contents), Papilionaceae (also Intramural
glands, open secretory cavities, and cavities with a papillose epithelium and a
bracket-epithelium, Fig. 57, p. 262), Caesalpinieaef Mimoseae (secretory cavities
with bracket-epithelium), Rosaceae, *Myrtaceae sens. str. (in T'1istania exceptionally
with mucilaginous contents), Lythrarieae, Samydaceae, Passitloraceae, Araliaceae,
Rubiaceae, Compositae. Primulaceae (occasionally with red crystalline contents),
Myrsineae (Eumyrsineae and Maeseae. frequently with red crystalline contents;
like canals in the axis, see also above), Styraceae (secretory cavities resembling in·
tram ural glands). *Myoporineae (excl. Oltia and Zombiana), Polygonaceae (secretory
cavities sometimes formed by four epidermal cells), Podostemaceae, Piperaceae (side
by side with secretory cells), Proteaceae (Fig. 17J' p. 7I3. intramural glands),
Gonystylus, Euphorbiaceae (only secretory cavities Wlth bracket-epithelium) ; ·lastly,
also in the genus Panda, which has not yet been assigned a definite position in the
Natural System (see Pierre, in Bull. Soc. Linn. de Paris, 189(5, p. 1256).
The development of irregular secretory cavities by the disorganization of
portions of the tissue of the wood has be~n recorded in Evodia and Zanthoxylon
 SECRETORY AND EXCRETORY RECEPTACLES IIOI

(Rutaceae, d. § 16), Carapa (Meliaceae), Dilodendron (Sapindaceae), Bu,kea

(Caesalpinieae) and Terminalia (Combretaceae). With this feature we may
associate the formation of tragacanth in the species of Astragalus and the
occurrence of gummosis in the species of Acacia and in the Pruneae. Lastly, the
presence of mucilaginous or resinous or latex-like secl"etions in the ordinary
intercellu~ar spaces of the leaf or axis may likewise be mentioned here.
This feattire has been demonstrated in Peganum, the genus Vahlia (Fig. 07.
p. 314), Donatia and Roussea (Saxifragaceae), Lys£machia and Primula, Gentiana,
and Cobaea (Polemoniaceae), and probably occurs also in Humulus and some
Acanthaceae.
§ 19. SECRETORY CANALS 1. The secretory canals are here understood in
an analogous sense to the secretory cavities discussed in § 18. They resemble
the latter in their diverse contents and their varied mode of origin and have
the same systematic value, which is often very considerable (characteristic of
all the members in the Orders indicated by a * in the enumeration below).
In determining their distribution the axis should first be examined, since i.
some cases the secretory canals do not enter the lamina of the leaf at all (e.g. in
the HyPericineae, certain Araliaceae, &c.). The distribution of the secretory
cana]s m the a.xis varies and is sometimes characteristic of certain taxonomic
groups. They may occur in the pith~ bast, pericycle, endodermis and primary
cortex, rarely in the wood as well! (only in the Dipterocarpeae-but here their
occurrence in the wood is perhaps quite general in axes of a certain thick·
ness-and in certain Caesalpinieae). In the Altingieae, for example, the
secretory canals are found only in the pith; in the Burseraceae or Anacardia-
ceae it is the secretory canals in the bast which are characteristic of their
respective Orders, but in certain species of these Orders 3 canals are present also
in the pith and primary cortex; in the Compositae the endodermal secretory
canals are characteristic, &c. When the secretory canals penetrate into the
leaf, they generally occupy the same position with reference to the vascular
system as in the axis of the species in question, i.e. secretory canals situated in
the bast of the vascular system of the veins correspond to those found in the
bast of the axis. canals lying above the fibrovascular system of the ,veins to the
medullary canals of the axis, and so on; but occasionally (Guttiferae pro parte,
Fig. 27, D, p. I22) the canals also run freely in the mesopbyll ". In some cases"
moreover, an investigation of the secretory canals in the root is indispensable,
although such an investigation can rarely be carried out systematically owing
to lack of material. It will suffice to point out that the secretory canals in
some of the Tubuliflorae (Compositae) are absent above the level of the coty-
ledons, and that the secretory canals, situated in the root in the Pittosporeae.
Araliaceae, and Umbelliferae, occur in a characteristic position, viz. in the
pericambium opposite the groups of wood arid bast, belonging to the primary
fibrovascular system.
_FQrJhe vicaripus ocC!1~e_nce of other types of secretory organs in plaQe of
the secretory canals of the Compositae, see under that Order in the following
enumeration, and also § IS (p. 1099).
Secretory canals have been observed in the *Hypericineae (only in the axis;

J De Bary, Verg1. Anat., 18n, pp. 210 and 4;5.

1 Interxylary secretory canals occnr more frequently in the root (Umbelliferae. C()mpolitae).
For the occurrence of mucilage-canals or mucilage-cavities and other kinds of SttCretory caviti~ in
the secondary wood, see §§ 16, 17.18 .
• In such cases the occurrence of secretory canals 10 the pith and primary eortex is getlen1t}
ooly a specific character.
• The secretory canals running freely in the mesopbyll occasionaUy. give rise to transparent Jines
in the leaves, while those accompanying the vascular bundles or the veins somdime'i pco«ucc
translucent veins.
 CONCLUDING REMARKS
always in the bast. sometimes also in the pith, prim. cortex and pericycle), ·Pitto-
sporeae (pericycle, sometimes also in the bast), *Guttiferae (always in the pith and
prim. cortex of the axis, in some cases also in the bast; in the leaf, either running
freely in the mesophyll or in the veins~ sometimes replaced by secret. cavities), Tern-
stroemiaceae (Bonnetieae, excepting Archytaea and Bonnetia; in the axis in the
pith and prim. cortex, in some cases also in the bast), *Dipterocarpeae (always
lD the pith and? second. wood, rarely? also in the bast), Rutaceae (in the centre of
the pith in Clausena punc.tata). SiIIlaru baceae (pi th). K oeberlinz'a (bast), ·Burseraceae
(secret. canals always present in the bast, and side by side with them sometimes
canals in the pith and prim. cortex), Celastrineae (veins of the leaf of Martonia and
margin of the 1eaf in Pachystima), Rhamneae (veins of the leaf in Reynosia pro parte,
the canals being provided with a papillose epithelium), *Anacardiaceae (typical
utembers of the Order alwa.ys provided with secretory canals situated in the bast,
but besides these there may be canals also in the pith a.nd prim. cortex), ]uUania (as
in the previous case), Papilionaceae (prim. cortex in Cot'dyla, M#leUia pro parte
and M)'!'oxylon), Caesalpinieae (in the secondary 'W'OOd of Copaifera, Daniella, Ep't:f'Ua,
Kingiodend7O'n, Oxystigma and Prioria, in these genera sometimes also in the pith or
cortex, as the case may be), Hamamelideae (margin of the pith in the Altingieae),
Cactea.e (in the cortex, the contents resembling late~ in Mammillaria pro parte and
Leuchtenbergia), ·Umbelliferae (above all in the pericycle, but also in the pith, bast,
and prim. cortex). ·Araliaceae (excl. AJ'alid'ium, otherwise as in the Umbelliferae),
Cornaceae (pith of Mastixitl), Compositae (in members of aU the thirteen tribes of
Bentham and Hooker's system of ~la.ssification; typically present in the root, and
occasionally absent in the shoot, the canals in the latter case being replaced in the
shoot by other types of secretory organs, viz. secretory sacs or laticiferous vessels' ;
in the root there ate always canals, which arise from the endodermis, but in addition
to this, canals may be found in the primary cortex, the periphery of the pith, the
secondary wood and bast of the root, or in the pericycle, the place of the canals
being sometimes taken by secretory cavities), Myrsineae (see § 18), Gesneraceae
(internal to the vasco bundles of the stem and in an analogous position in the veins
of the leaf, in Klugia and RI/Ynchoglossum), Podostemaceae (Weddellina, intercell.
spaces resembling resin~canals ?), "Leimerieae (at the margin of the pith).

f 20. LATICIFEROUS CELLS 1 and secretory organs of a similar type. It is

a familiar fact that the laticiferous cells (non-articulated laticiferous tubes) are
long tubular branching elements, which are filled with various contents, the
latter often being of the nature of latex or caoutchouc; the laticiferous cells
are already differentiated as initial cells in the hypoco~yl of the embryo, and
in later stages their branches often traverse all the organs of the plant. The
term ' non-articulated laticiferous tubes thus refers to the course of develop-
J

ment of these elements. It is not, however possible to undertake a develop.

9

mental investigation in each individual case ; according toChauveaud s~ moreover,

it is not out of the question that, to take the case of the Apocynaceae for
instance, some of the secretory organs, there described as laticiferous cells, are
differentiated only at a later stage, and develop in a different way from that
mentioned above, and according to F. E. Weiss a this is really true of the
caoutchouc-cells of Eucommia, which cannot be distinguished from true latici·
ferous cells, unless their mode of development is studied. For these reasons
<>nly those Orders or genera are enumerated under the heading 41aticiferous
-cells' in the following review, some at least of the species of which have been
investigated developmentally with positive results; the remainder are com·
prised Wlder the heading of 'secretory organs resembling laticiferous cells.'
Thp. detailed structure of the laticiferous tubes, viz. the size of their lumina
and the nature of their walls and contents, is only of trilling systematic value.
It is a]so necessary to ascertain, whether the elements in question occur in all

, Cf. De Buy, Vergl. Anat., 1817. p. 199.

• In Ann. IC. nat., aer. 7t t. xiv, 1891.
• In Transact. Linne:tn Soc. London,.2 seT" vol. iii, Bot., 18g2.
 SECRETORY AND EXCRETORY RECEPTACLES Ilo3
the vegetative organs, for in some cases they do not enter the leaf (see e.g.
under Moraceae, p. 1059). For transitions to laticiferous vessels (in the
Apocynaceae), see § 2I.
Laticiferous cells, the branches of which may run in the pith, bast, and
primary co~exl, in the veins of th~ leaf, and occasionally also fttely in the mesophyll,
but are chiefly found accompanymg the bast-groups, occur in all ( ?) Apocynaceae
and Asclepiadeae, in many Euphorbiaceae. in the genera Humulus and (?) Cannabis
(Cannabineae), and lastly in many, perhaps all, Moraceae.
Secreto~y. organs resembling laticiferous. cells,tsome of them perhaps of
secondary ongm, as has ~n proyed to be t~~ case m Eucommia (Trocho~end~aceae),
ate alsoknown to OCCur m Plagtopteron (TIllaceae). Coula, Eganthus, /{ezstuJa, Min.
quat'tia and Ochanostachys (Olacineae), Celastrus, .Euonymus, Gymnosporia pro
parte', Myst'Yoxylon and Wimmeyia (Celastrineae), Campylostemon, H£ppoCfatea pro
parte and Salacia pro farte (Hippocrateaceae) and Urtt'a (Urticaceae). Caout-
chouc~cells, like those 0 Eucomm'(a, are closely related to the elongated secretory
sa.cs, mentioned under II in § 14 (p. 1097).
The secretory organs discussed in this section are sometimes visible even to
the naked eye or with the help of a lens; this is the case, when their caoutchouc-Uk,
contents appear in the form of delicate elastic threads' on breaking through the
leaves and branches, or when the laticiferous cells give rise to transparent lines in
the leaves.
§ 21. LATICIFEROUS VESSELS. These elements arise by the absorption of
the transverse walls in rows of cells, which are either simple or arranged to form
a network; they occur in the same tissues of the axis and leaf'as the laticiferous
cells, but are chiefly associated with the phloem. In some cases they also run
freely in the mesophyll, while in the Papayaceae they are even found traversing
the soft xylem-mass.
Laticiferous vessels have been demonstrated: among the Pap ave raceae , in the
genera Papaver, Argemone, Roemeria and Chelidonium (? also in Meconopsis and
Platystemon); in the tubers of T1'opaeolum ? (Geraniaceae); among the Olacineae,
in the genera Endusa and Cardiopteris ; in all the Papayaceae; in all the Cichoria-
ceae, and, outside the limits of: this tribe, in the genera Gundelia, Gazania, Caflina
and Atractylis (Compositae); in all Campanulaceae and Lobeliaceae with the ex-
ception of Sphenoclea; among the Convolvulaceae, in the genera Dic?ondra and
Falkia (in the latter genus as yet demonstrated only in the floral organs); and
among the Euphorbiaceae, in the genera Hevea and Manihot.
Transi tions between rows of laticiferous sacs and la ticiferou~ vessels, as evidenced
bv the occurrence of perforations on the transverse walls of the former, have been
observed in the Orders Sapotaceae and? Convolvulaceae, while transitions between
elongated secretory sacs (see § 14) and laticiferous vessels have been met with in
certain Compositae; lastly, transitions between laticiferous cells and laticiferous
vessels are found in the petals of cer1::ain Apocynaceae. All these features require
careful reinvestigation.
Trichomes, which enter into connexion with the system of laticiferous vessels,
are found in some of the Cicboriaceae (Fig. 103, 0, p. 458), while papillose termina-
tions of the laticiferous tubes occur in Sz'phocampylus (Lobe~aceae).
§ 22. OXALATE OF LIME -t. The diverse forms of excretion of oxalate of
lime, as "also'-its absence and its distribution in the various organs and tissues of
a plant, afford a whole series of valuable anatomical features,
1 Lat'iciferous cells occur in the wood only in so far as bunches of the corticallaticiferous tubes
sometimes traverse the medullary rays of the wood and finally penetrate into the pith.
a· See Loesener, in Notizbl. Berliner Garten n. ,p, 1908, p. 64. .
• Such cases of ' spinning' on the part of the caoutchouc-like contents have been observed in the
secretory organs of Eu(ommia, Plagiopteron, Ce!artntS, Euollymus, MJlstr{)xy~n, IVimmen"a,
Hi,!p()(ratea, and Salada (mentioned above under the heading of 'secretory organs reSembling
lauciferous cells '), in the Jaticiferous cells of Parameria (Apocynaceae) and in the' elongated
secretory sacs' of Tinoml'.uiUfJl (Menispermllceae, included under II in § 14); ,
• 4 Cf. Kohl, Kalksalze und Kieselsaure in der Pflanze, Marburg, 1889; Poli, CristaUi di ossalato
calcico, Roma, 1882 (ab!itr. in Bot. Centralbl., 1882, ii, p. 311); and also Busc::"lioni, in MlIlpighia,
x, I¥, where a general idea. of the literature dealing with oxalate of lime can be obtained.
 CONCLUDING REMARKS

We may distinguish the following modes of excretion of oxalate of lime:

I. Ordinary solitary crystals (i.e. rhombohedral crystals (' Hendyoeder ') be-
longing to the monoclinic system}, which are of large but variable size, and
are very widely distributed) and their modifications and twin-crystals; 2. Sty-
loids 1 or columnar crystals (i.e. elongated prismatic crystals belonging to the
monoclinic system) or twin-forms of such crystals, the latter, when typically
differentiated, recalling the well~known crystals characteristic of gypsum;
r~arded from the systematic point of view, styloids commonly replace raphides
and occasionally show transitions to the forms of crystals, discussed under 4 ;
3. Octahedml or prismatic crystals, belonging to the quadratic or monoclinic
systems and mostly of rather small size; 4. Acicular crystals, i.e. small needle-
shaped or fusiform crystals, which sometimes merge into small crystals of
prismatic shape or crystals, which are almost of the nature of styloids; when
elongated, the acicular crystals come to resemble raphides 2, whilst, when they
are very small, they appear like sand; in most cases large numbers of acicular
crystals occur in one and the same cell; 5. Clustered crystals, which are com-
oosed of individual crystals belonging to the quadratic or monoclinic systems ;
l>. Sphaerltes (Sphere-crystals), i.e. bodies resembling clustered crystals and
having a sphaerocrystalline structure with a smooth or rather smooth surface;
7. Rapbides 3, Le. long acicular crystals, large numbers of which lie parallel to
one another in a cell containing mucilage; 8. Crystal-sand (Fig. I34r p. 579),
which is composed of numberless small crystals, completely filling the cells and,
in the case of typical crystal-sand (' sable t~traedrique ')"having a tetrahedral
form '. The systematic value of these forms of crystals varies very considerably.
In- some cases a certain form of crystal is characteristic of an entire Order,
while in other cases it serves to distinguish only genera or species; and this
applies to all the different kinds of crystals above enumerated, even to those
which are specially typical and have the greatest systematic value (viz. raphides,
styloids, and crystal~sand). The three forms of crystals just named, for ex-
ample, are of strikingly small systematic value in the Order Thymelaeaceae.
Several forms of crystals may occasionally occur side by side in one and
the same plant, the chief combinations being solitary and clustered crystals,
raphides and clustered crystals, raphides and styloids, clustered crystals and
crystal~sand (the clustered crystals sometimes even being enclosed in the cells
containing the crystal.sand), and so on; in such cases the various forms of
crystals are found either in the same or in different tissues, or even in different
organs of the plant. In other species or genera, or even Orders, however,
only one form of crystal is present. In employing the occurrence uf oxalate
of lime for systematic purposes it must be borne in mind that this salt has
been shown to be capable of re-entering into the metabolism of the plant,
and that clustered crystals often occur alone in the primary cortex in early
sta«es, while solitary crystals only appear subsequently.
Oxalate of lime has hitherto' not been demonstrated in the: Fumariaceae.
Cruciferae. Stackhousieae. Crossosomataceae, Lobeliaceae, Monottopeae, Lennoaceae,
Primulaeeae, Salvadoraceae. D~sIDntain8a, Gentianeae-Menyanthoideae, Cuscuteae?

1 Radlkofer. in Sib.• Ber. Munch. Akad., 1890, p. 114; see also Rothert and Zalenski, Bes.
Xategorie von KrYltallbeh., Bot. Centralb1., J899. iv, p. 1 et seq.
• The!le shorter acicular crystals are rarely (see Gentianeae) enclosed in cells containing mucilage .
• Seoe a150 A_ Fuchs, Ban der Raphidenzelle, (ht. bot. Zeitschr., 1898, p. 32-4 et seq.; and Kohl,
Raphldenzelle, Bot. Centrll.l!>1., 1899. iii, p. 373 et seq.
a See Arcangeli, in Noo? Giorn. bot. Ital., xxiii, 1891.
, The following statements refer solely to the vegetative organs (especially the axis and the
leaf). For it frequntly happens that crystalline elements occu'r in the reproductive organs
(telta,etc." although it haa been found impo!lSible to demonstrate them in the \"egetative organs
., the IIlll\C pl.nt (e. g. in certain Caryophylleae, Valerianeae, and Primulaceae).
 SECRETORY AND EXCRETORY RECEPTACLES 1 105

Orobanchaceae and Plantagineae ?, besides numerous genera and species belonging

to other Orders.
Ordinary solitary crystals and clustered crystals (sometimes side by side with
a third form) have been observed in the same or in different species of the follow-
ing Orders and genera respectively: Trochodendraceae, Anonaceae, Menispermaceae
(solitary crystals not very abundant, clustered crystals rare), Berberideae, Cappari-
deae, Cistineae (solit. cryst. rare), Violarieae, Canellaccae, Bixineae, Pittosporeae,
Tremandreae, Polygaleae, Vochysiaceae, Caryo£hylleae ~solit. cryst. very rare}.
Portulaceae, Tamariscineae, Hypericineae, Guttiierae, Ternstroemiaceae, Diptero~
carpeae, Lophira, Malvaceae, Triplocmtonaceae, Sterculiaceae, Tiliaceae, Lineae
(clust. cryst. rare), Humiriaceae, Malpighiaceae, Zygophylleae, Geraniaceae, Rutaceae,
Si.marubacea.e, Balanites 1, Ochnaceae, Luxemburgiaceae, Wallacea, Burseraceae.
Meliaceae. Chailletiaceae. Olacineae, Ilicineae, Cyrilleae. Celastrineae, Hippocratea-
ceae. Pentaphylacaceae, Corynocarpaceae, Rhamneae, Ampelidaceae, Sapmdaceae,
Hippoca.stanaceae, Aceraceae, Staphyleaceae, Sabiaceae, Anacardiaceae, Coriarieae
(dust. cryst. only in the rhizome), Moringeae, Caesalpinieac, .l\tlimoseae (dust. cryst.
rare), Rosaceae, Saxifragaceae. Hamamelideae, Ostrearia, Bruniaceae, RhizoJ?ho~
raceae. Combretaceae (chiefly clust. cryst.), Myrtaceae, Lecythidaceae, Lythraneae,
Samydaceae, Passifioraceae, Cacteae, Umbellifcrae (solit. eryst. rare), Araliaeeae,
Comaceae, Caprifoliaceae. Rubiaceae (solit. cryst. rare), Compositae (rare). Vaccinia-
ceae, Ericaceae, Epacrideae, Diapensiaceae, Plumbagineae, Myrsineae. Sapotaceae,
Ebenaceae, Styraceae, Apocynaceae, AscIepiadeae, Loganiaceae, Convolvulaceae,
Solanaceae, Scrophularineae (rare), Columelliaceae, Bignoniaceae. Pedalineae,
Myoporineae, Verbenaceae (solit. and clust. cryst. rare), Labiatae (rare in the vege-
tativeorgans), Amarantaceae (solit. eryst. rare), Chenopo<iiaeeae (solit. eryst. rare),
Basellaceae, Batideae, Polygonaceae, Podostemaceae. Aristolochiaceae (solit. cryst.
rare), Piperaceae (Symbryon). Proteaceae (not abundant). Thymelaeaceae. Gonyslylus.
Geissoloma. Loraniliaceae, Santalaceae, Myzodend"on, Grubbt"a. Balanophoreae,
Euphorbiaceae. Buxaceae. Balanopseae) Ulmaceae, Moraceae. Urticeae, Platanaceae,
Juglandeae, Myricaceae, Casuarineae, Cupuliferae, Salicineae. Lacistemaceae.
Clustered crystals alone. Or accompanying some other form of crystal (with
the exception of ordinary solitary crystals),-in the latter case in the same or in
different species-have been observed in the following Orders and genera respectively:
Ranunculaceae (rare), Magnoliaceae, Nymphaeaceae, Sarraceniaceae, Papaveraceae,
Frankeniaceae. Elatineae, Lophira, M onoles, Chlaenaceae. Melianthaceae, Penthorum,
Myrothamnus, Halorageae (Fig. 72, A, p. 337), Melastomaceae (elust. cryst. occa·
sionally reduced to short and thick solit. cryst.), Onagrarieae, Loaseae, Tumeraceae.
Papayaceae. Begoniaceae. Ficoideae, Valerianeae (rare), Dipsaceae, Calycertae,
Candolleaceae. Campanulaceae (only in Sphenoclea), Plocosperma, Gentianeae (only
small clust. cryst;). Polemoniaceae (clust. cryst. rare and of small size), Hydro.
phyUaceae, Baragineae. Gesneraceae, Acanthaceae, 4ombiana, Labiatae, N yctagineae,
Illecebraceae~ Chenopodiaceae, Phytolaccaceae (rare), Nepenthaceae, Piperaceae
(excl. Symiwyon). Chloranthaceae. Myristicaceae, Octotepis, Penaeaceae, Daphni-
phyllaceae, Cannabineae. Leitnerieae, Empetraceae, but also in _certain genera or
species belonging to other Orders.
Rhombohedral solitary crystals are found alone or accompanied by some
other form of crystal (excluding clustered crystals)-in the latter case in the same or
in different specie~-in the Rhaptopetalaceae, Koebe,.linia~ Octocnemaceae. Co:'lnara·
ceae, Crassulaceae; Cucurbitaceae (rare), Hernandiaceae (only in Iltigera obtusa), and in
most Papilionaceae and Mimoseae, and in numerous genera or species of other Orders.
As regards-the form"of·the-clustered crystals, we may note at this point that
the well-known star-shape is the commonest. Clustered crystals, which are made
up of acicular or quadratic (Begoniaceae, Lecythidaceae. Proteaceae, Fig. 173, p. 7 I 3)
crystals, are more rarely found. Special forms requiring mention are the clustered
crystals of certain Combretaceae, which are like stars with unequal rays (~ krippen-
stemartig '), and the clustered crystals of species of Centrati&nia (Melastomaceae,
Fig. 79, A, p. 362), which are found combined with styloids.
Sphaerites have been observed only in certain Berberideae, Tamariscineae
(Fouqtu'era), Geraniaceae, Aceraceae, Sabiaceae I, Papilionaceae. Rosaceae, Crassula-
ceae, Combretaceae (according to Hahnel, in Bot. Zeit., 1882), Melastomaceae.
Lytbrarieae. Cacteae (Fig. 93. p. 413), Rubiaceae, Asclepiadeae, Solanaceae, Phyto-
I According to Van Tieghem, Ann. sC. nat., ser. 9, t. iv, 1906.
2 According to Dihm, Beih. Z. Bot. CentralbJ., xxi, Ab. i. 1907.
4 B
 1106 CONCLUDING REMARKS

laccaceae, Balanophoreae, Euphorbiaeeae and Empetraceae; according to Hegel-

maier, they also occur in certaIn Caryophylleae, but are there confined to the testa.
Octahedral or
prismatic crystals of the ~uadratic system fq) (which in most
c?Ses are at once distinguishable from monoclinic crystals by their smaller size),
as well as twin-forms of such crystals, ~ small prismatiGt or fusiform to acicular
crystals (a) (which pccasionally, when of sufficient length" resemble raphides), or
small crystalline bodles of diverse other shapes (rounded, &c_) t, have been met with
in the following Orders: Ranunculaceae (q), Calycanthaceae (q), Magnoliaceae (q),
Menispermacea.e (a), (also transitions to crystal-sand and styloids), Capparideae,
Resedaeeae (rare), Canellaceae (q), Guttiferae, Sterculiaceae, Zygophylleae(a), Pega-
num (a), Simarubaceae (Fig. 4"";, B, p. 183), Ampelidaceae (a), Aceraceae, Papiliona-
ceae, Caesalpinieae a, Saxifragaceae (a), Melastomaceae (a), Lythrarieae, Onagrarieae,
Begoniaceae (q), Datisceae (a), Cacteae (q), Ficoideae, Arali~ae, R~biaceae (a),
Compositae (q, al, Goodeniaceae, Campanulaceae (q, a), Myrsineae-Theophrasteae,
Styraceae, Oleaceae (a), Apocynaceae (a), Loganiaceae (q, a), Gentianeae (a), Pole-
moniaceae (a), Boragineae (a), Convolvulaceae (a), Solanaceae (q), Retna (a),
Scrophularineae (q. a), Lentibularieae. Gesneraceae (q, a), Bignoniaeeae (a), Pedali-
neae, Acanthaceae (a), Myoporineae (a), Selagineae, Verbenaceae (q, a), Labiatae
(q, a), Nyctaginea.e (a), Chenopodiaceae (q), Basellaceae (q), Phytolaccaceae (rare
and observed only in the ovary), Cytinaceae (q), Aristolochiaceae (a), Piperaceae (a),
Chloranthaceae (very rare), Myristicaceae (a), Monimiaceae (a), Laurineae (a),
Hernandiaceae (a), Gomortegaceae (a), Elaeagnaceae (al, Moraceae.
Bundles of raphides are found in the following Orders, being of constant occur-
rence in those marked with a *: *Dilleniaceae, Ternstroemiaceae (Marcgravieae
and Sauraujeae exc). Stachyurus and Pelliciera, Fig. 29, A, p. 129), Geraniaceae,
Rutaceae, ZygophyUeae (Peganum). Ochnaeeae (Tetl'amerista), *Ampelidaceae
(Fig. 52, E, p. 223), Melianthaceae, *Hydrangeae (Saxifragaceae), *Onagrarieae
(exc1. Tf'apa), Ficoideae, Rubiaceae, Gesneraceae (N apeanthus f'epens, J. D. Smith!),
Nyctagineae, Phytolaccaeeae (Euphytolaec~ae and Agdestis), Urticeae (Lapcwtea),
Thelygoneae. I

Typical styloids have been recorded in the following Orders or genera, as the
case may be: Pittosporeae (constant), Ternstroemiaceae, Zygophylleae, Geraniaceae,
Rutaceae, Simarubaceae, llicineae. Rhamneae, Sapindaceae, Melianthaceae. Rosaceae,
Saxifragaceae, Melastomaceae (often distinctive of groups of genera, not constant
in Centradenia). Lythrarieae, Onagrarieae, Ficoideae, Caprifoliaceae, Rubiaceae,
Apocynaceae, Asc1epiadeae, Loganiaceae, Plocosperma, Nyctagineae, Phytolaccaceae
CRiviDeae), Thymelaeaceae (Fig. 174, D, p. 717), Eupborhiaceae (not constant in
Phyllanthus), Urticeae (Laportea). We have already noticed above that styloids
often take the place of the bundles of raphides. The styloids are commonly confined
to the bast, while some other form of crystal occurs in the remaining tissues
of the species in question. At this point we may also include: (a) crystals
resembling styloids, but shorter than typical styloids, such crystals being found
in certain Menispermaceae, Tiliaceae, ZygophyUeae, Celastrineae, Papilionaceae,
Ficoideae, Columelliaceae, Nyctagineae, Phytolaccaceae. Thymela.eaceae, Euphor-
biaceae, Buxaceae, Ulmaceae and Juglandeae; (b) small rod-shaped crystals, which
likewise resemble styloids, and are combined with one another so as to have the
shape of a widely open V or W, or the form of an I; crystals of this type occur in
certain Menispermaceae, Malpighiaceae, Sapindaceae, Papilionaceae (Fig. 58, p. 26S),
Mimoseae~ Rosaceae, Loganiaceae and Euphorbiaceae; (e) crystals, which are
shaped like styloids, but vary in length, and occur to the number of several in the
same cell; they have been observed in Krameria (Poly_galeaeJ. certain Cacteae,
Pisonia (Nyctagineae. Fig. 154, C, p. 646). Rivina and Vdlamilla (Phytolaccaceae).
Typical cTystal~sand (' sable tetrai;!drique J). which occasionally includes
clustered crystals Ccl. cr.) or even solitary crystals (sol. cr.), has been observed in
the Rutaceae (el. cr.), Olacineae (sol. cr.), Sapmdaceae, Saxifragaceae, Crassulaceae,
Araliaceae, Comaceae, Caprifoliaceae, Rublaceae, Sapotaceae (only in the latici-

1 All these forms of crystals are considered jointly here, because they often occur side by side in
different s~cies of an Order or even in the same species, and because it is ~ometimes difficult to draw
a sharp hmit between the different forms. The addition of the letter 9 or a in the subsequent
enumeration is Dot intended to indicate that the fonns of crystals, represented by these letters, are the
only ones occurring in the Order in question, but testifies solely to the occurrence of such crystals, in
so (ar u it has been definitely established.
, In DeiariuTlt, according to recent observations of my own.
 SECRETORY AND EXCRETORY RECEPTACLES 1107

ferous sacs), Loganiaceae, Boragineae, Nolaneae, Solanaceae (cl. cr.), Amarantaceae,

Chenopodiaceae, Thymelaeaceae. Buxaceae; powdery or finely granuIat.-cry.tal-
sud has been obstryed in A canthophyllum (Cary~phylleae). Dysphans'a, Gymnocarpos
and Halwosia (Illeceoraceae), and Saruma (Aristolochiaceae) ; in the last of these
genera it is present only in the tennina.Pcells of the trichomes. We may include
here the following spe~d features: (a) the occunence of a kind ?f crystal-sand above
the gelatinized'membranes of the epidermal cells in the leaves of certain species of
Rou'I{Q (Connaraceae); (b) crystal-sand, consisting of tetrahedral granules and small
acicufar and prismatic crystals, in Leucaster (Nyctagineae); (c) cry~ta1-sand,
compoSed of sma)l acicular and prismatic crystals, in CaJandf'initJ-(Portulaceae),
and crystal-sand, showing a similar composition. in Protoschwenks'a (Solanaceae) and
Buxus (Buxaceae); (d) coarsely granular crystal-sand, consisting of quadratic
or short prismatig crystals, in DunaJia (Solanaceae), Galtesia (Pbytolacca.ceae,
Fig. 160, p. 666) and members of the tribe Monimieae; lastly, (e) a kind of crystal-
sand (?), also in two species of Meliosma (Sabiaceae). When large numbers of
acicular crystals occur in the same cell, they are occasionally reduced so as to :form
a kind of sand (e.g. in certain Menispennaceae, Melastom~ae. Gentianeae, Pipera-
ceae and Elaeagnaceae). A number of small crystals are commonly found accom-
panying solitary or clustered crystals, especially in the large crystal-ldioblasts of, the
leaf, but such cases have not been taken into consideration in the preceding review
of the distribution of crystal-sand.
As a rule the crystalline elements are found in the lumina of the cells,
but in some cases (and this applies especially to solitary and clustered crystals)
they are embedded in a thickened part of the: cell·wall; in the latter case they
are either completely enveloped by the cell-membrane or lie in the middle
of the lumen of the cell enclosed in a sack of cellulose, after the manner of
Rosanoff's clustered crystals, this sack being connected with the cell-wall by
means of one or more beams of cellulose 1. We can only attribute systematic
value to these features, when the differentiation of the structure in question
is particularly typical (e.g. in the Papilionaceae, Citrus or the Rhizophoraceae).
Features of greater systematic importance are afforded by the deposition of
numerous small crystaIline granules resembling crystal-sand in the walls of the
cells of various tissues in certain Crassulaceae (Fig. 70, D, p. 321), Cadeae,
Ficoideae, Nyctagineae (Fig. I54, A, p. 646), and Euphorbiaceae (in Croton/here
in the tissue of the cork), and the deposition of large crystals in and on the
cell-Walls in the Nymphaeaceae (Fig. II, p. 47). An apparent embedding of
rather large crystals is found in the crystal.§c1erenchyma of certain Magno-
liaceae (Fig. 4, p. 29), Combretaceae, Rubiaceae and Loranthaceae (Fig. 177,
p. 728), and in the crystal-hairs of the Guettardeae (Fig. IOI, B, C, p. 446) ;
the former has already been discussed in § 9 (PP. 1090, 1°91); in the 'latter the
ceIl·wall rmdergoes thickening to such an extent, that really nothing but the
spaces filled by the crystab remains to represent the lumina of the hairs.
The various fonns of crystals described in the preceding pages occur in the
different tissues of the leaf and axis, although sometimes confined to a definite
tissue. The shape and size of the celIs, containing the crystals, correspond to
the shape and size of the crystalline deposits themselves. Cells containing crystals
are often not distinguished from the neighbouring cells, but in other cases they
appear as idioblasts. The following special types of crystal-receptacles are
of systematic importance: (a) crystal·idioblasts actually or apparently situ·
ated in the epidermis (see § 2, p. 1075); (b) celIs containing clustered cTtals
or sphaerites and projecting towards the exterior in the form of a papilla or
hair (in certain Euphorbiaceae, Fig. lBo, R, S, p. 748); (c) small cells, contain·
ing clustered crystals and bordering on the intercellular spaces into which they
often project like papillae, in certain Nymphaeaceae (Fig. II, B, p. 47), Halora-
geae (Fig. 72, A, p. 337), and Onagrarieae (Trapa); (d) crystal·idioblasts,
1 See also Wittlin, Kalkoxalattaschen, Bot. Centralbl., 1896, iii, p. 33 et seq. and Mobius,
Festlegung dcr Kalksalze D. Kiesclsanre, Ber. deutsch. bot. Gesellsch., 1908, p. 39 et seq.
4 B '2
ll08 CONCLUDING REMARKS

provided with a gelatinized or merely thickened inner wall, in the mesophyll

of certain Dipterocarpeae; (e) small cells bearing clustered crystals, which
bring about a blocking up of the respiratory cavities (Pilocarpus, Rutaceae); (I)
transversely septate palisade-cells with a single clustered or solitary crystal in
the compartments; .. (g) idioblasts in the tissue of the leaf, filled with a solitary
or clustered crystal, such idioblasts having wide lumina and often being dis-
tinguished by their spherical shape; (h) groups of cells with clustered crystals
in the mesophyll (Pemphis, Lythrarieae); (i) entire layers of crystal-cells, lying
parallel to the surface of the leaf (e.g. in certain Celastrineae, Melastomaceae or
Samydaceae); (i) sheaths composed of crystal-cells in the veins of the leaf; (k)
the 'cristarque -cells of the leaf and axis (see §§ 7 and 51) ; and (1) chambered
parenchyma with solitary crystals (in the wood and bast) or clustered crystals
(in the bast). A special and peculiar mode of occurrence of acicular crystals is
iound in the fluke-cells of the anchor-like shaggy hairs of Cranocarpus (Papilio-
naceae, Fig. 59, G, p. ~69) and in the terminal cells of the stinging hairs of
T,agia and other Euphorbiaceae (Fig., 180, P, Q, p. 748).
It remains to mention that the crystalline forms of oxalate of lime (and
especially large clustered or solitary crystals, styloids, which often traverse
the entire thickness of the leaf at right angles to its surface, and bundles of
raphides, orientated in the same direction) freq,uently give rise to delicate or
distinct transparent dots in the leaf, while raphldes, when lying parallel to, the
surface of the leaf, may produce transparent striulae.
§ 23. SPHAEROCRYSTALS (sphere-crystals) 1. Sphaerocrystals which do not
consist of oxalate of lime have been observed in herbarium- or alcohol-material
of a number of plants. Only in a few cases is their chemical nature at all clearly
known. In some Orders they consist of inulin, in the Ampelidaceae perhaps of
cissose, in certain Cruciferae and Rutaceae of a substance resembling hesperidin,
in certain EuphoTbias of calcium phosphate and calcium malophosphate,. &c.
Inulin· has been demonstrated in: Violarieae, Malpighiaceae I, Droseraceae.
Compositae, Candolleaceae, Goodeniaceae. Campanulaceae, Lobeliaceae and Myopo-
rineae; sphaerocrystals of a different chemical composition in: Ranunculaceae.
Menispermaceae~ Cruciferae, Cap:parideae. Violarieae, Temstroemiaceae, Stras-
burgeria, Geraniaceae. Rutaceae.llicineae. Ampelidaceae, Papilionaceae, Crossoso-
mataceae. Lythia.rieae~ Caprifoliaceae (Adoxa) , Passifloraceae, Cacteae, Ficoideae,
Rubiaceae, Valerianeae, Calycereae. Compositae~ Campanulaceae, Asclepiadeae,
Nolaneae, Bignoniaceae, Verbenaceae, LabIatae,. Basellaceae, NeJ?Cnthaceae, ChIo-
ranthaceae, Thymelaeaceae, Santalaceae, Myzodendran, Euphorblaceae. Moraceae,
Urticeae, Salicineae. For sphaerites of oxalate of lime, see § 22, p. 1105. .

I 24. CRYSTALLOIDS 6. Crystalloids, which occur in the nuclei, the cllfO~

matophores or the cell-sap, and are either crystalline or of some other shape,
have a wider distribution than has generally been supposed.
They are found in certain Berberideae, N:ymphae.aceae, Capparideae, Caryo-
phylleae, Lineae, Geraniaceae, Aceraceae, Papllionaceae (according to Baccanni,
In the floral organs), Halorageae, Passifioraceae (spindle-shaped), Cacteae (spindle-
Hansen. in Arb. bot. lnst. Wiirzburg. iii, 1884 i Leitgeh, in Mitteil. bot. lnst. Graz, 1888,
1
Heft 2, p. 257 et seq.; Baccarini, in Malpighia. 1888, p. I et seq.; I:!ee also Schaarschmidt, in
Mag~ar novenyt. lapok, 1881, p. 134 et seq. (abstr. in Bot. CentralbL, 1883, i, p. 46; and Just,
1882, i, p. 412). .
t Gr. Kraus, in Bot. Zeit" 1875, p. 171, and 1877, p. 329; H. Fischer, iu Cohn, Beitr., viii,
1898, p. 53 et seq.# Bay, Materials for a monograph of Inulin, Transact. Acad. S. Louis, vi, 1895,
p. [51 et seq.
~ See the papers by Hartwich, &c.• cited in the literature--supplement, p. 1171.
, According to Eichinger, Diss., Munich, 1907.
I Leitgeb, KrystaUoide in Zellkernen, MitteH. bot. Instit. Graz, 1888. p. 113 ~t seq.; Zimmer-
mann, Heitr. z. Morph. 'LI.. BioI. d. PRanzenzelle, Tiibingt'n, Heft 1, 1890, p. 54- et seq., and Heft lI,
t891, p. r u ct seq.; Stock, Prot~inkrystalle, in Cohn" Beitr., vi, 1893, p. 313 et seq.
 SECRETORY AND EXCRETORY RECEPTACLES lIog

shaped, annular, or filiform, Fig. 93. p. 413), Droseraceae (as in the Cacteae). Aralia-
ceae, CandoHeaceae~ Campanulaceae, Ericaceae, Oleaceae, Gentianeae. Convolvula-
ce~, Scrophularineae, Lentibularieae, Gesneraceae, Bigno~aceae, Verbenaceae.
AI1!arantaceae, Phytolaccaceae, Nepenthaceae, Euphorbiaceae. Urticeae.

§ 25. OTHER CELL-CONTENTS. The following enumer~tion deals with

e'ell-contents other than those hitherto mentioned; the deposits in question
for the most part occur only in herbarium-material and are rardy found
in the living plant as well; the latter is the case with the deposits of gypsum,
occurring in the epidermis of certain Capparid.ae (Fig. 18, E, F, p. 68), and the
characteristic clustered and solitary crystals of the Salvadoraceae, which con·
sist of an unlo1own organic salt of lime (Fig. 120, A, B, p. 527) .
.Apart from the prismatic crystals of gypsum, found in the epidermis of the
leaf of certain Cappandeae, and the peculiar crystalline elements of the Salvadora-
ceae, the following special contents have been observed; (a) crystalline bodies of
diverse kinds and of unknown chemicai composition, in certain Berberideae (Fig_ 9,
p. 44), 'Nymphaeaceae, Polygaleae, Terns troemi ace ae. Malvaceae. Zygophylle~
(herejn part also present in the intercellular spaces), GU1tnera, Onagrarieae, Umbel-
li ferae , Myoporineae, Selagineae. Polygonaceae and Piperaceae; (b) crystal! of
hesperidin in certain Rutaceae (Fig, 40, p, 175); (c) crystals of carotin in certain
Onagrarieae and Scrophularineae; (d) berberin, which gives a yellow coloration to
the wood, in the Ranunculaceae, Menispermaceae, and Berberideae j (e) rhamno·
cathartin, filling the medullary rays of the cortex with yellow contents, in the
Rhamneae; (I) cissose in the Amr.elidaceae; (g) saponin.like substances 1 in the
Caryophylleae;, Sapindaceae, Papilionaceae, Rosaceae {Quillaja) and Cactea.e;
(h) crystals of quinme in the Cinchonapbarks i (i) crystals of ' cumarin in the leaves
of Liatris odo,.atissima (Camp-ositae); (i) scutellarin III the Labiatae; (A) oroxylin,
in the form of a green wax-like substance, in Oroxylum (Bignoni~ae); (I} balano-
phorin, a body resembling wax. in the Balanophoreae; tm) crystals of indigo or bodies,
consisting of indican, in -many Papilionaceae (secretions containing indican also in
the secretory cells of Ayistolochia Lindeniana var., and in the secretory cavities of
certain Olacineae, these secretions taking on a blue colour with eau de Javell~;
for blue -cystoliths. see under Acanthaceae, p. 1019); (nJ lastIy~ alumina-bodies·
in the mesophyll and cortex of the species of.Symp/ocos (Styraceae).
As a general rule no very great degree of systematic value can be 'attributed
to the occurrence of fat-bodies', which are found in the assimilatory tissui (especially
the palisade-tissue) in herbarium-material of species belonging to numerous Orders,
and are occasionally doubly refractive. For the distribution of these bodies, !ee
chiefly Petit, who points out that they are-rare in the Apetalae and in the Mono-
cotyledons. With the fat-bodies ~e may class the sI11;all caoutchouc-bodies,
WhlCh occur side by side-with the former in the Celastrineae, p_nd are described as
characteristic of the Hippocrateaceae and Sapotaceae.
For the formation of tragacanth and the occurrence of gummosis, see. f 18,
p. 110 I.
§ 26. SILICA C, It is a familiar fact that silica occurs both as an incrusta~
tion on the cell-wall 'and as a mass filling the lumen of the cell. Silicification
or- the 'ceU;membrane is_a very widely distributed phenomenon, which more
particularly affects'the integumental tissue, and ,chiefly its outer walls. Silicifica-
tion ,of the .walls of hairs is of very frequent occurrence, and silicification of
groups of epidermal cells is likewise a common feature; the silicified cells in

. I Regarding th~se bodies, see Kunk~l, ill Arb. d. pharmakolog. In!;t. Dorpat, vi, 189 1, P. 5;
Th. Waage, in ·Pharm. Ce1ltral~1111e, 1892, p. 65i; HeyJ, in Archil' d. }'harm., 190 1• p. 45 1 j ~
Weil, Sa.poninsubst., Dis~, Strassburg, 1901, also in Archiv d. Ph:mn., 1901, p. 363~
II. Radlkofer, in·Ber. -deutsch. bot. Gesell.ich., 190-4, p. 18B et seq.
! Radlko(er, in Sitz .&r. Miinch. Aknd., 1889, p. 26'1 and 1890, p. Il-4; Petit, Globulesrefringenr.,
Comptes rendns Acrtd. Paris, cx:'txiii, 2 e sem., 19c1l, p. 1250 and Sptll!rnlins, loc. cit., ex-xxv, :lie !tm .•
1902 , p. 99 1 • ,
4 De Rary, Vergl. Anat., 18i7, p~ 108 j Kohl, Kalksalze u. Kitteluure,.1889; p. 197 et Sleq.
1110 CONCLUDING REMARKS

the latter cue are often grouped round about silicified hairs or function as
basal cells bearing such hairs, and are not uncommonly visible to the naked
eye as small rough knobs on the surface of the dry leaf. In other cases spherical
or hemispherical groups of mesophyll-cells, or groups composed of epidermal
cells and the subjacent cells of the palisade-tissue undergo silicification, those
pats of the walls, which abut on one another, being affected; these walls are
often considerably thickened or may even be provided with cystolith-like
protuberances (see Fig. 166, G, p. 684). In other cases again, isolated cells of the
mesopbyU are silicified. The features last mentioned are sometimes of relatively
great systematic value (Oladneae). Apart from that, the tendency to form
silicified cells is 'tharacteristic of certain Orders (e:g. Boragineae and Aristo-
lochiaceae). Groups of sqicified cells, consisting of a malformed stoma and the
neighbouring epidermal cells, may commonly be observed, but their occurrence
is of no value for systematic purposes. It remains to mention that carbonate
of lime often occurs side by side with the silica in the silicified cell·waHs ~.
Cells with strongly.silicified membranes have been observed in the following
Orders: Dilleniaceae (epidermal cells, mesophyll. and spiny hairs), Calycanthaceae
(hairs and their subsidiary cells), Magnoliaceae (epid. cells), Trochodendraceae
(epid. cells + palisade.cel~s). Menispermaceae (special groups of epid. cells),
Burseraceae (epidermis). Olacineae (cells of the mesophyll, solitary or in groups.
Tribe Olaceae), Philadelpheae (epidermis), Caesalpinieae (cells of the mesophyll),
Mimoseae (epid. cells + pal.·cells). Chrysobalaneae (e~dermis, cells of the meso--
phyll, small knobs due to the hairs), Bruniaceae (epldermis), Halorageae (small
knobs due to the hairs), Combretaceae (epidermis), Cucurbitaceae (cells of the
mesophyll, small knobs due to the hairs), Rubiaceae (epidermis), Compositae (small
knobs due to the hairs), Goodeniaceaer(epidermis + cells of the me!ophyll).
Campanulaceae (reduced hairs),. Oleaceae (small knobs due to the hairs). Hydro-
phyllaceae (small knobs due to the hairs). Boragineae incI. Cordiaceae (small knobs
due to :the hairs). Gesneraceae (walls of the hairskBignoniaceae (cells of the meso-
phyll). Acanthaceae (epidermis). Verbenaceae (small, knobs due to the hairs)"
Aristolochiaceae (epid. cells + palisade-cells;' cells of the mesophyll, Fig. 166"
p. 684), Piperaceae (epidermis),.Cliloranthaceae (epidermis +·cells of the mesophyll.
cortex), Proteaceae (epidermis), Loranthaceae (cells of the mesophyll). Santalaceae
(cells of the mesophyll). Myzodendron (cells of the mesophyll) .. C"ampereia (cells of
the mesophyll). Euphorbiaceae (small knobs due to the hairs. epidermis), Urticaceae
(small knobs due to the hairs. epidermis,. cells of the mesophyll, fibrouS cells of the
bast).
. f' .
There are far fewer records of the occurrence of siliceous matter, in the
lumina of the cells than of silicification of the cell·wall. The siliceous matter
appears in the form either of silica-bodies, which lie freely' in the lumina
of the cells and arise freely in the protoplasm, or of silica-plugs; the latter are
either formed freely in the protoplasm or are due to the apposition of siliceous
matter upon the cell-wall~ and ultim.at~ly ·fill·the entir~ lum!~~ of the, ce~.
Tothiscategorybelon~: the siliceous excretions, found in the mesoph:yll of certain.
Dilleniaceae and in the epidermis of the leaf of a member of the Magnoliaceae; the
silica-bodies.and silica-plugs ·in the cortex of Eucommia (Tr6chOdendtaceae);' the
silica,-l;lodies. in the mesophyU of species. of Melios",a (Sabiaceae); the siliceous
masses, filling (GI) the sac-shaped cells found near the terminations of,the.veiM in
A,.can~eJisia (M~permaceae). (b) some of the cells of the parenchyma accompanying
the vems in Ficus, Pararlotaf"/"US. and S{itwattoryu (Moraceae), and (c) the spicular
cells of Cenwapl4C'Us (Bixineae) and Cyncriuwa (Caesalpinieae); the silica-bodies
and silica:'plugs, occurring in numerous ·ChrysobaJanea.e ; "the silica:bOdies of the
PodOBtemaceae (Fig. 163. p. 675); the silica-plu{S in the hypoderm of the leaf of
Thottea and Apama (Aristolochiaceae. Fig. 166. H, i, p. 684); :tne silica-bodies. found
in the Wood·parenchyma' in certain DipterocafPE:.ae, Malvaceae. Sterculiaceae•. ;rilia-

1 For the occurl'alte of silica in cystolitbs

----
and 'cystolith-like structures, see' ,l8.,
II 'Bargagli-Petrucci. in Malplghia, 1902 • ' .
 SECRETORY AND EXCRETORY RECEPTACLES IIII

ccae. Burseraceae, Anacardiaceae and Sapotaceae. Silica-plugs, moreover, ap~ar

to occur 'quite frequently in some of the cells composing cell-groups with silicified
walls. The occurrence of silica-plugs in the vessels or other cells of the heart-
wood has only been recorded in the plant from which • cauto' -bark is obtained
(Chrysobalaneae) and in some of the Verbenaceae; silica-plugs have been observed
in intercellular spaces only in cauto '-bark.
I

§ 27. CARBONATE ~F LIME. Calcification of the cell-wall is no common

phenomenon. The parts chiefly affected are the walls of certain forms of
hairs, which in thes~ cases often have a verrucose surface (the most important
being the 'cystolith~hairs '), and the cystoliths and cystolith-like structures
(see § 28)., Carbonate of lime, however, iUso occurs side by side with the
silica in the walls of the small knobs, formed by the hairs of certain Com-
positae, &c. A feature deserving special mention is shown by the groups of
enlarged epidermal cells, found in the leaf of Hanburia (Cucurbitaceae, see
Fig. 86, p. 393); the common walls of contact between the cells are encrusted
with carbonate of lime, while at the same time well-differentiated crystals of
the same salt ~re deposited on the walls.
An incrustation of the cell-wall with carbonate of lime has been observed
in the trichomes of the Cruciferae, Papilionaceae, Saxifragaceae (quite generally
in the Philadelpbeae and Hydrangeae), Loaseae. Cucurbitaceae, Urnbelliier.e,
Cornaceae, Compositae, Campanulaceae, P!ocosperma, Hydrophyllaceae, Boragineae.
Scrophularineae, Gesneraceae, Verbenaceae and Urticaceae (Figs. 181 and 182,
pp. 769 and 773).
In some woody plants carbonate of lime appears in the form of cell-
contents, filling the vessels and other elements of the wood (especially of the
heart-wood), as well as cells of the pith 1. In order to avoid confusion with·
resino~s substances, which very frequently block up the lumina of the vessels,
it is necessary to apply the well·known reactions demonstrating the presence
of carbonate of·lime.
Carbonate of lime has been shown to occur in the wood or pith in the following
Orders: Anonaceae. Zygophylleae, Aceraceae. Rosaceae, Cornaceae. Epacride.ae.
S~p<>taceae. Urticaceae. Cupuliferae. Salicineae.

§ 28. CYSTOLITHS AND STRUCTURES. RESEMBLING CYSTOLlTHS 2. Under

cystoliths in the narrower sense only· those protuberances of the cell-wall
are understood, which, like the familiar cystoliths of Ficus, show differentiation
into a stalk and a body. As a rule both carbonate of lime and silica play
a part in the incrustation of the cystolith, the lime being deposited chiefly
in and on the body, while the silica is mostly confined to the stalk and the
nucleus of the cystolith. These structures are still described as cystoliths,
when carbonate of lime is absent (un calcified cystoliths, • cystotyles ' of Radl-
kofer), the body of the cystolith in such a case occasionally becoming suberized
or lignified., _They_.even remain cystoliths, when they consist throughout of
silicified cellulose. We may at once note that these differences in the nature
of the incrustation are of no very great systematic value, since cystoliths with
and without incrustation, and cystoliths showing diverse types of incrustation
may occur side by side even in one and the same species.
Cystoliths are found in the irtegumental tissue and mesopbyll of
the leaf and in the pith and cortex a the axis; they have not hitherto been
observed in the wood. In the majority of cases they occur chiefly in the
epidermis of the leaft where they fill cells which are often of large size and

I See Molisch, in Sitz.-Eer. Wiener Akad.,l~ltK.iv, Abt. I, 1881.

2 S~ especially Kohl, Kalksalze u. Kieselsiiure, 1889, p. 115 et seq.
 111:2 CONCLUDING REMARKS

have a shape adapted to that of the cystolith; these cells penetrate into the
inner tissues and mostly participate in 'he formation of the surface of the
leaf only with a small part of their wall. At this point the stalk of the cystolith
is generally attached; however, not uncommonly (Acanthaceae), the stalk
appeat;j to become resorbed secondarily. \Vell-developed cystoliths are occa-
sionally found also in .epidermal cells, which are differentiated as papillae or
short hairs. The sha:f'e of the cystoliths varies, being spherical, ellipsoidal,
fusiform, &c.; they are mostly unbranched, branched cystoliths being rare
(~rtain Acanthaceae, Hernanruaceae and Urticaceae). The sha,:pe of the
cystoliths can often be determined with the help of; a lens, since In the dry
leaf they frequently appear in the form of dots, striulae or small star-like
structures. By the combination of two or more cystoliths ·Qr of the cells
e
containing them lithocysts ' of Radlkofer) there result the double cystoliths,
or groups of ~toliths) which occur in the Opilieae (Fig. 47, p. 203), in.certain
Cucurbitaceal (Fig. 85, p. 391) and Acanthaceae (Fig. 144, p. 6IS), and iq
Champe1'eia (Fig. 179, p. 736). In these struct~res the stalks of the cystoliths
are attached to the common wall of contact, or, in the case of groups of cysto-
liths, to the walls, which abut on one another.
Cystoliths have been observed in the Olacineae (all Opilieae, Fig. 47, p. 203).
Cucurbitaceae (Momordica and Coccinia, Fig. 85. p. 391), Boragineae (Tou'Yne/ortia
and Cordia, Fig. 127, A, p. 559). Gesneraceae (Klugia and Rhynchoglossump, Acantha-
.ceae (very widely distributed, Fig. 144, p. 615; regarding blue cystoliths, see
-under Acanthaceae, p_ JOI9), Hernandiaceae (Gyrocarpus and Spat'attantkel";um,
, Fig. 172, p. 708). Champereia (only in the axis, Fig. 179. p. 736), and Urticaceae
(cerlain_ Ulmaceae, Cannabineae, Moreae and Urticeae, Figs. 182 and 183. A, pp. 773
I and 776 ).
The structures resembling cystoliths differ from true cystoliths in the
fact that the stalk is absent, or at least not distinctly differentiated. As in
the case of the typical cystoliths, the cystOlith-like structures are encrusted
either with carbonate of lime and silica, or only with the one or the other sub-
stance. Structures resembling cystoliths are, in the first place, found in tri-
chomes, and are then known as hair-cystoliths (Figs. 82, 127, r8I, G, and 182,
pp. 379, 559, 769, 773)· These hair-cystoliths either arise from the lateral
wall at the base of the hair or form a continuation of the body of the hair,
which is filled for a varying distance with caps of cellulose, the latter being
either calcified or silicified, or both calcified and silicified. The differentiation
of the hair-cystoliths shows an antagonistic relation between the development
of hairs on the one hand and of cystoliths on the other, inasmuch as the
greater the reduction in the length of the hair, the more strongly developed
are the bodies of the cystoliths, and the more closely do they resemble the
typical cystoliths (see Figs. rSI, rSz, pp. 769, 773). Papillose epidermal
cells are frequently seen to contain typical cystoliths (see above). So that,
whilst the hair·cystoliths on the one hand are connected by transitional forms
with typical cystoliths, we find on the other hand all transitions from hair-
cystoliths to hairs showing complete reduction of the bodies of the cystoli ths, cul-
minating in hairs, which are simply calcified. With the cystolith·like structures
we rna) also class thickenings of the cell-wall (briefly described as protuber-
ances in the synopsis, which follows below), which are found projecting in the
form of humps into the lumina of the subsidiary cells of certain kinds of
trichomes (the latter being calcified or silicified or provided with hair~cysto­
liths, Fig. I27, C, D, p. 559), and now and then also occur independently of the

The structnres, observed by Clarke (Hooker, leones; Plate I i98, eel. 1 888) in the leaf of the
Gesneraceous species HemihQea Hmr)'i, Clarke and interpreted by him as cystoliths, are not
cystoliths, bnt rod-shaped spicular ceUs, as I am ahle to state on the basis of a recent investigation.
 SECRETORY AND EXCRETORY RECEPTACLES rIl3

hairs in groups of epidermal cells (see Fig. 85, p. 391) or in the ground tissue
of the leaf (Fig. 179, p. 736) ; we may also include here the local peg-shaped
thickenings found in certain species. The cystolith-like structures found in
the subsidiary cells of the hairs arise from those parts of the walls which abut
on the trichomes; the latter are invariably unicellular. The lithocysts form
small knobs at the bases of the hairs, sometimes together with calcified and
silicified cells devoid of cystolith-like structures (d. § 26)
Structures resembling cystoliths ha.ve been observed in: Cistineae (species of
Cistus, with silicified protuberances in epidermal cells, &c!), Papilionaceae (species
of Cyamopsis a.nd Indigo/era, with a calcified body resembling a cystolith in the
terminal cells of the two-armed hairs), Mimoseae (protuberances in the subsidiary
cells of the hairs, calcified in AtJonsea), Samydaceae (Homalium donquaiense, \\ith
silicified protuberances in the roesophyJI), Loaseae (hair-cystoliths, protuberances
in the subsidiary cells of the hairs, Fig. 82, p. 379), Cucurbitaceae (as in the Loaseae,
see also Fig. 85, p. 391), Compositae (calcified protuberances sometimes present
in the subsidiary cells of the hairs), Campanulaceae (protuberances in reduced
trichomes, in the epidermal cells of the margm of the leaf and in the subsidiary cells
of the hairs, occasionally calcified), Myrsineae (protuberances in the epidermis of the
leaf in Claviia boliviensis). Oleaceae (silicified protuberances in the subsidiary cells of
the hairs of Nyctanthes), Plocosperma (calcified protuberances in the subsidiary cells
of calcified hairs), Polemoniaceae (silicified protuberances in papillose epidermal cells
and hair-cells), Hydrophyllaceae (hair-cystoliths and l?rotuberances in subsidiary
cells of the hairs, calcified), Boragineae (as in the prevIous case, Fig. 12 7, p. 559),
Scrophularineae (hair-cystoliths and proh:berances in the subsidiary cells of the
hairs, calcified), Verbenaceae (as in the previous Order), Loranthaceae (silicified pro-
tuberances in the epidermis and mesopliyll), Santalaceae (silicified protuberances in
the ground tissue), Champerda (as in the :previous case, Fig. 1.79. p. 736), Euphorbia-
ceae (silicified protuberances in the subsidIary cells of the hairs of BC1'na1'dia), Urtica-
ceae (hair-cystoliths, protuberances, some oi them peg-shaped, in the subsidiary
cells of the hairs, and in epidermal cells, hypodermal cells and cells of the mesophyll,
calcified or silicified, Figs. 181, 182, pp. 769, 773)'
We may also include here the small peg-shaped structures arising from the
outer walls of the epidermal cells in Erythroxylon obtusum (Lineae), and the peg-
shaped bodies found in special epidermal cells in the leaf of Anami1'ta and Af'can-
gelisia (Menispennaceae), the cells in question functioning as hydathodcs:

Quite a special form of the structures resembling cystoliths is constituted

by the un calcified cystotyles and cystosp_heres, which occur in certain Begonia-
ceae, and are arranged in pairs after the manner of the double cystoliths;
they' have an approximately hemispherical shape and are inserted on the
common wall of contact between their lithocysts by means of a small, scarcely
marked stalk. The cystotyles of the Begoniaceae are distinguished by the
fact that they consist of a mucilaginous ground-substance, which after treat-
ment with alcohol shows concentric stratification and radial striation, while
in some of the cystotyles the ground-mass is impregnated with a resinous
substance. The distinctive character of the cystospheres lies in their being
composed of a-mass of secretion~ which is enclosed in a kind of sac; for details
see p.p. 402; 403J and Fig. 87. The cystospheres are closely related to the
secretory cells of certain Magnoliaceae, Canellaceae, &c., in which the secretion
is enclosed in a peculiar pouch; d. § 14, p. IO¢.
We may mention the following facts regarding the systematic impor-
tance of the cystoliths and cystolith-like structures. The cystoliths in the
strict sense have proved to be of most value, for they occur only in a few Orders
and are characteristic of whole tribes or genera. How far the shape and
position of the cystoliths can be employed for taxonomic purposes is shown

J According to Gard, in Comptes rendus Acad. Paris, cxlv, 1907.

J1I4 CONCLUDING REMARKS

especially by the systematic papers dealing with the cystoliths of the Acan-
thaceae and Urticaceae. The cystotyles and cystospheres of the Begoniaceae
are not of general occurrence throughout the Order, but are often character-
istic of certain sections of the genus BegoftJ'a Data as to the extent to which
the cystolith-like structures can be 'employed in distinguishing species are
contained chiefly in Mez's and Priemer's papers, which deal with the cystoliths
of the Cordiaceae and Ulmaceae respectively.

IV. HAIRY COVERING 1.

§ 29. GENERAL REVIEW. in this ·chapter I propose to deal both with

the clothing hairs (§§ 30-33) and the glandular hairs (§ 34); in addition,
we may append to our discussion of the laUer a consideration of glandular
leaf-teeth (§ 35), large glandular mechanisms, which mostly excr·ete nectar
and are therefore generally described as extrafloral nectaries (§ 36), the lime-
and salt-glands of the Frankeniaceae, Tamariscineae, and Plumbagineae (§ 37)~
and special forms of glands found in insectivorous plants (§ 38).
As is well known, clothing and glandular hairs are distinguished from
one another by the absence and presence respectively of secretion. The
product of the secretion is, however, not always of an oily, resinous or muci-
laginous nature, but may sometimes (in the case of bydathodes) be .nothing but
water, nor does the process of secretion always continue throughout the life
of the plant; very often, moreover, the plants are only available for investiga.
lion in, the form of herbarium·material. Under' these circumstances the
recognition of a trichome as a clothing or glandular hair would be attended
with difficulties were it not for the fact that 'the glandular hairs are in almost
all cases distinguished morphologically as well as by their function, inasmuch
as the upper secreting portion is more or less sharply marked off (frequently
in the form of a head) from a basal portion resembling'a stalk, or at least has
thinner walls than the basal portion. When a systematic anatomical investiga-
tion is being undertaken on herbarium·material, it is necessary to pay special
attention to these features. In some cases living material is indispensable, if
the nature of a trichome is to be properly determined.
The systematic value of the hairy covering is very great. In the
first place the presence or absence of glandular hairs affords an important
feature, which is sometimes characteristic of entire Orders, while in other
Orders it only serves to distinguish genera or species. 'In the second place the
morphological structure of the hairs is to be taken into consideration. As
regards the clothing hairs, a first point of great importance is whether they are
unicellular or uniseriate. Although these two forms of hairs are not at all
uncommonly found side by side in the same species (often even on the same
organ of the plant), in certain Orders we find that without exception, or ahnost
without exception, only unicellular or only uniseriate, hairs are present either
on all the organs of the plant or at least on the vegetative organs. In the
Papilionaceae and Compositae t for instance, nearly all the clothing hairs hitherto
observed are uniseriate, while in the Cruciferae only unicellular clothing hairs
have been recorded. In the .case of multicellular glandular hairs, a feature
which is quite especially valuable for taxonomic purposes has been shown to
lie in the mode of division of the head; in some cases the head is divided

I See A. Weiss, Pflanzenh., Berlin, 1867. also in Karsten, Bot. Untersnch., Bd. I; Martinet, 1[1
Ann. sc. nat., ser. 5, t. xiv. 1872; De Baty, Vergl. Anat., 1871 i Theorin, Vaxttrichom., Archiv for
Bot., i, iii, and iv, 1903-5; Hinch, Untersnch. liber d. Entwickl. d. Haare bei den PlI., Diss., Berlin,
18 99. -41 pp., also in Fiinfstiick, Beitr., iv.
 HAIRY COVERING HIS

exclusively by means of vertical waDs, while in other cases the division-walls

are horizontal, or both horizontal and vertidll. In this way, for example, it
is possible to distinguish ,the Scroehularineae from the Solanaceae, since the
Scrophularineae have glandular hairs of the first type, while the Solanaceae
have those of the second type. In the third place an abundance of useful
characters, serving to distinguish taxonomic groups of varying magnitudQ
doym to species, is afforded by the numerous modifications in the form and
size of the hairs ; .in the case of the clothing hairs similar characters are pro-
vided 'by the shape of the cells composing the h:Jrs, as well as by the nature
of their walls and lumina, and, iii the case of the glandular hairs, by the struc-
ture of the secreting portion and stalk, and the localization and nature of the
secretion.
In some Orders, genera, and species only one form of hair is found, but
in other cases two or more types of hairs are present. In the latter case a
certain definite plan. of ~tructure frequen~ly reappears in all the different forms
of hairs belonging to the same Order (see Cruciferae,' Droseraceae, ,Loganiaceae-
Buddleioideaet Convolvulaceae, &c,), genus or species.. .
,In investigating the haii'y covering with a view to its employmen't for
systematic purposes it is advisable to make the investigation a comparative
one, and, wherever· possible, always to examine the same organs of the plant,
viz: in the·1irst place the leaf;then the axis, and only as a last resort (na~ely,
when no trichomes are to be found on the vegetative organs)·the reproductive
organs 1,
. § ,30. CLASSIFICATION OF THE CLOTHING HAIRS. The tyPical forms of
clothing hairs may be distinguished as follows :-,.I. Simple clothing hairs
or briefly hairs, which· are' formed by a single cell (not developed as a flat
structure), or by a row of two or more cells. .II. Peltate hairs, scales or lepides,
i.e. sessile or stalked structures, which consist in the main either of a single cell,
developed as a fiat surface"(in the formation of which the stalk may participate),
or of. a varying number of cells arranged in one or more layers; with tho peltate
hairs we may associate the s.~ellate and candelabra.hairs, discussed III §§ 3I
and 32. III. Shaggy hairs, i.e. :filiform structures, the base of which at
least consists of two or more rows of cells, such hairs being either of epidermal
origin or of the n~tur~ of emergences; and with them we may class warts
and spines, i.e, massive multicellular structures, which either ,have blunt or
poiilted ends _and are likewise partly of the nature of emergences.
§ 3r. SIMPLE CiO'fllING HAIRS. This section first contains a synopsis
of the distribution of the ordinary (mostly filiform) unicellular and uni-
seriate clothing hairs,in the different Orders; unicellular hairs, which show a
vesicular or papillose differentiation, are summarized under a sep'arate heading.
It is impossible to -enter into details here regarding the mamfold individual
differences~in- structure shown by the· simple clothing hairs (already in part
referred~to 'in -f 29); these differences, which concern the length, shape, and
structure of the walls and lumina' of the cells composing the hairs, and (more
especially as regardS the uniseriate trichomes) also the structure ~nd pqsition
of the division-walls and the mode of demarcation of the cells; ·can all be
employed for systematic purposes, when .their constancy has been sufficiently
established. Only a' few.forms of hairs, -which are quite parti.cularly note-
worthy (e,g. especially bracket-hairs, two- and one-armed hairs, and branched
----------
1 The hairs (~nd es~ciany the glandular hairs), found in the floral region, occ•• ion:t11y show
.. reduced form or even an altered plan of stmcture as compared with the hairs present on the leaves.
of the same specie!!; see, for instance, under \' erben:tce~e, p. IOU.
1116 CONCLUDING R'CMARKS

hairs of the stellate and candel~bra·types, which can be derived from unicellular
and uniseriate hairs), will be conSJdered in greater detail and with special refer-
ence to their distribution.
I. Unicellular clothing hairs of a papUl08e or vesicular type (d. also § 2,
p. 1076) have been observed in the: Resedaceae, Fumariaceae, Caryophylleae,
Portulaceae, Geraniaceae, Celastrineae, Hippocrateaceae, Rhamneae, Aceracea.e.
Coriar~eae, Papilionaceae (rare and only occurring in those species. which have a
papillose epidermis), Crassulaceae (Fig. 70, B. C. p. 32 I), Halorageae, Melastomaceae,
Lythrarieae Ficoideae. Umbelliferae, Rubiaceae, Campanulaceae, Loganiaceae,
Gentianeae, Polemoniaceae, Plantagineae, Illecebraceae, Amarantaceae, and Mora-
ceae. For the development of ordinary papillae on the epidermis, see § I, p. 1073. ,
II. Ordinary unicellular clothing hairs 1 of ~reater length have been observed
in: Ranunculaceae (Fig. I, p. 16, see also III), Dilleniaceae, Calycanthaceae (Fig. 3.
p. 25), Magnoliaceae (see III), Trochodendraceae (see III), Anonaceae (see III),
Menisperrilaceae (see III), Berberideae (see III), Sarraceniaceae, Cruciferae (Fig. 15,
p. 63), Capparideae (Fig. 19, p. 75. see' III), Resedaceae, Cistineae (Fig. 2 I. p. 8 I),
Violarieae (see III), Bixineae (see Ill), Tremandreae, Polygaleae (see III), Vochy-
siaceae, Frankeniaceae, Portulaceae, Tamariscineae. Elatineae (see II I), H ypericineae
(see III), Guttiferae (see III), Ternstroemiaceae (see Ill).. Dipterocarpeae, Monotes,
Chlaenaceae, Malvaceae (see III), Sterculiaceae (see III), Tiliaceae (see III), Rhapto-
petaIaceae. Lineae, Humiriaceae (seelII), Zygophylleae (Fig. 38, p. 168), Geraniaceae
(see III), Rutaceae (see III), Simarubaceae(see III). Koeberlinia, Balanites, Ochnaceae
(see III), Burseraceae (see III). Meliaceae (see III), Chailletiaceae (Fig. 46, p. 198),
Olacineae (sec III), Ilidneae (see III), Cela.~trineae (see III), Pentaphylacaceae,
Stackhousieae, Rhamneae (see III), Ampelidaceae (see III), Sapindaceae (see IU),
Hippocastanaceae (see III), Aceraceae (see III). Staphyleaceae, Sabiaceae (see III),
Anacardiaceae (see III), Moringeae. ~nnaraceae. Caesalpinieae (see III), Mimoseae
(see III) Rosaceae, Saxifragaceae (see IJI). iDroseraceae, Hamamelideae, Brunia.-
ceae, Halorageae (see III. Gunneroideae, Scl1indl.), Rhizophoraceae, Combretaceae
(Fig. 76, p. 346), Myrtaceae (sens. str.), Lecythidaceae (see III), Melastomaceae (rare,
see Ill), Lythrarieae (see nil. Onagrarieae (see III), Samydaceae (see III). Loaseae
(Fjg. 82, p. 379, see III), Turneraceae (see III). Passifloraceae (see III), CucurbHa-
ceae (see III), Cacteae (see' III), Ficoideae. Umbelliierae (see III), Cornaceae, Capri-
foliaceae., Rubiaceae (Fig. IOI, p. 446, see III), Valerianeae (see III), Dipsaceae,
Candolleaceae, Goodeniaceae."(see III). Campanulaceae (see III), Lobeliaceae (see
III), Vacciniaceae (see III). Ericaceae (see III), Monotropeae, Epacrideae (see III),
piapensiaceae, Plumbagineae, Sapotaceae (Delpydot'a), Ebenaceae, Oleaceae (see III),
Salvadoraceae. Apocynaceae (see III), Asclepiadeae (see III), Loganiaceae (see III),
Plocospeyma, Gentianeae (see III), Polemoniaceae (see III), Hydrophyllaceae (see
III), Boragineae (Fig. 127. p. 559. see-III), Cuscuteae (see III). Nolaneae (see III),
Solanaceae (rare, see III), Scrophularineae (see III), Orobanchaceae (see III), Colu-
melliaceae, Bignoniaceae (~ee III), Pedalineae (see III), Acanthaceae (see III), Ver-
benaceae (see III), Labiatae (see III), Illecebraceae (see III), Amarantaceae (see III),
Chenopodiaceae (very rare, see III), Polygonaceae (see III), Myristicaceae, Monimia-
ceae, Laurineae. Hernandiaceae. Proteaceae (see III). Thymelaeace~e, Octolepis,
Go ny s tvl us, Penaeaceae. Geissoloma, Santalaceae (see I I I ). My zodendron. Grubbia,
Balanophoreae, Euphorbiaceae (see III), Buxaceae (see III), Ulmaceae (mostly, see
III), Cannabineae (see III), Moraceae (see III), Urticeae, Thelygoneae, Juglandeae,
Myricaceae (see III), Cupuliferae (see III), Salicineae, Lacistemaceae (see III),
Empetraceae, Ceratophylleae; unicellular clothing hairs, but showing a special type
of structure throughout (see p. 1 I 17 et seq,), are found also in. the Orders Malpighia-
ceae, Papilionaceae, Composltae, Sapotaceae (excl. Dtlpydora) and Styraceae. _
III. Ordinary simple uniseriate clothing hair. or trichomes differing but little
from the ordinary type (see p. I I 18 under trichomes with long terminal cells) have
b~n observed in: Ranunculaceae (see also II), Magnoliaceae (see II), Trochodendra-
ceae (see -II). Anonaceae (see II), Menispermaceae (see II), Berberideae (see II),
Nympbaeaceae, Papaveraceae, Capparideae (see II), Violarieae (see II), Bixineae·(see
11), pjttosporeae (Fig. 22, p. 93), Polygaleae (see II), Caryophylleae, Elatineae (see

I The ngur~ cit~d under II and III in ~rt refer to special forms of nnieelltllar or uniseriate
clothing hair;; (see p. J II 7 et seq.). -
 HAIRY COVERING

II), Hypericineae (see II), Guttiferae (Fig. 26, p; l~, gee II), Ternstroemiaceae (see
II). Malvaceae (see II), Sterculiaceae (see II), Tiliaceae (see II), Humiriaceae (see II).
Geraniaceae (see II). Rutaceae (see II), SimarulJt\ceae (see II). Ochnaceae (see II):
Burseraceae (see II), Meliaceae (see II), Olacineae ? (see II), Ilicineae (see II). Cela-
strineae (see II), Hippocrateaceae (see I), Rhamneae (see II), Ampelidaceae (see II).
Sapindaceae (see II), Hippocastana~eae (see II), Aceraceae (see II). Sabiaceae (see
II). Anacardiaceae (see II), Papilionaceae (rare, F1g. 59, p. 268). Caesalpinieae (rare.
see II), Mimoseae (very rare, see II), Saxifragaceae (see II), halorageae (see II,
Haloragoideae, Schindl.), Lecythidaceae (see Ill, Melastomaceae (rare, see II).
Lythrarieae (see II), Onagrarieae (see II), Samydaceae (see II), Loaseae (see II),
Tumeraceae (see II), Passifloraceae (see II). Cucurbitaceae (see II) ~goniaceae,
Cacteae (see II). Umbelliferae (se~ II):.Araliaceae. Rubiaceae (see 1l), Valerianeae
(see II), Compositae, Goodeniaceae (see II), Campanulaceae (rare, see II), Lobeliaceae
(see II), Vacciniaceae (see II), Ericaceae (see II), Epacrideae (see II), Primulaceae
(Fig. 114. p. 502), Myrsineae, Styraceae, Oleaceae (see II). Apocynaceae (Fig. 12 I,
p. 530, see II), Asclepiadeae (see II), Loganiaceae (see II), Gentianeae (see II),
Polemoniaceae (see II), Hydrophyllaceae (see II). Boragineae (see II), Convolvu-
laceae (Fig. 129, p. 567), Cuscuteae (see II), Nolaneae (see II), Solanaceae (see II).
Retna, Scrophularincae (see II). Orobanchaceae (see II), Lentibularieae, Gesnera-
ceae (Fig. 140, p. 599). Bi~noniaceae (see II). Pedalineae (see II), Acanthaceae
(see II), Myoporineae, Zomb:ana, Selagineae, Verbenaceae (see II), Labiatae (see II).
Plantagineae (Fig. 153. p. 643), Nyctagineae, Illecebraceae (see II). Amarantaceae
(Fig. 156, p. 652, see II), Chenopodiaceae (Fig. 159, p. 659, see II), Polygonaceae
(see II), Phytolaccaceae, Nepenthaceae (Fig. 165, p. 679), Aristolochiaceae (Fig.
166, p. 684), Piperaceae, Proteaceae (Fig. 173, p. ,13, see II). Santalaceae (see II),
Euphorhiaceae (see II). Buxaceae (see II) .. Ulmaceae (rare, see II), Cannabineae (see
II), Moraceae (see II), Leitnerieae. Myricaceae (see II), Casuarineae (Fig. r86, p. 788).
Cupuliferae (see II), Lacistemaceae (see II); uniseriate clothing hairs, which are,
however, distinguished by a special shape (see p. 1118 et seq.), occur also in the
Chlaenaceae, Connaraceae, Caesalpinieae, Cornaceae, Myristicaceae and Loran thaceae.

The following lines in the first place contain an enumeration of the special
forms of unicellular hairs, which require mention; these hairs are. distinguished
by their shape, their size, the structure of their wall or their contents. Very
small silicified trichomes, which are inserted in the outer wall of the epidennis,
are found in Petraea (Verbenaceae, Fig. 151, A, B, p. 632); small silicified tri·
chomes resembling papillae occur in certain Moraceae; short hairs, provided
with a spherical base showing spiral thickening, in certain Sapindaceae, and
hairs ~aving a similar structure in a certain specie." of Zizyphus (Rhamnaceae).>
and -in species of Artocarpus (Moraceae); short unicellular laticiferous hairs,
'which enter into connexion with the laticiferous system, in some Cichoriaceae
(Fig. r03, 0, p. 458); small capitate hairs with a spherical or pyriform
(glandular?) head in Cardiopteris and Platea (Olacineae, Fig. 48, B, C, p. 204)
and hairs having a similar structure in Symplocos (Styraceae) ; small trichomes,
-.shaped like a clustered crystal, in Peixotoa (Malpighiaceae, Fig. 36, p. 164) ;
longer trichomes, the apex of which is bent in the form of a hook (bracket.
hairs, climbing hairs, or hooked hairs, see Fig. 182, D, p. 773), in certain Bixi-
neae, Burseraceae," Caesalpinieae, Mimoseae, Passifioraceae, R ubiaceae, Boragi·
neae, Labiatae, Circaeaster (only on the fruit), Hemandiaceae, Moraceae, and
Urticeae, and short hairs, shaped like an angler's hook, in certain Cornaceae
(Fig. 99, G, p. 435); anchor-hairs and trichomes, provided with spines, which
are directed either forwards or backwards, in the Loaseae (Fig. 82, p. 379).
At this point we may also mention the following types of hair~: the two-
armed hairs (the arms being of equal or of unequal length), found·ln certain
Cruciferae (Fig. 15, C, p. 63), Capparideae (Fig. 19, B, p. 75),Vochysiaceae, Tern-
stroemiaceae (Microsemma ?), Chlaenaceae, Malpighiaceae (of almost constant
occurrence, Fig. 36, p. 164), Zygophylleae, Simarubaceae, Burseraceae (Fig. 43,
E, p. 191), Meliaceae, Celastrineae, Ampelidaceae, Connaraceae (Fig. 55, A-C,
p. 252), Caesalpinieae, Rosaceae, Com~r~~aceae, Myrtaceae sens. str., Lythrarieae
 lu8 CONCLUDING REMARKS

(CupJua pro parte! Fjg. So, C, p. -'7°), Samydaceae (Ba1taTa, Homalium J),
Ficoideae, Cornaceae (Fig. 99, B, p. 435), Sapotaceae (almost:constant, in part
glandular), Boragineae, Acanthaceae, Verbenaceae, Monimiaceae, Hernandia..
ceae (only in Illigera obtusa), Thymelaeaceae, Euphorbiaceae and Cannabineae
(Fig. 181, p. 769); the one-armed hairs (which sometimes show transitions
to two~anned hairs, while in othelf tales they are m~ly provided with a faint
crop-like protrusion on otie side of the ttalk), occurritlg in certain Vochysiaceae,
Malpighiaceae (Fig. 36, E, p. 164), ZygophyUeae, Meliaceae, Celastrineae, Sapin~
daceae, Connaraceae, Mimoseae, Saxifragaceae, Combretaceae, Myrtaceae sens.
str., Sapotaceae, Ebenaceae, Boragineae lFig. 126, D, p. 557), Acanthaceae and
Laurineae; the stellate hairs present in Certain Cruciferae (Fig. IS, D, E, p.63),
Capparideae (Fig. 19, Ct p. 75), Ternstrocmiaceae (Fig. 30, B, p. I33; in this
Order the stalk is sometimes demarcated, while the shield is now and then pro~
vided with a division~waU) and Saxifragaceae (Fig. 68, B, C, p. 316); the peltate
hairs, occurring in certain Cruciferae (Fig. IS, F, p. 63); and the dendroid hairs,
found in certain Cruciferae (Fig. IS, G, p. 63), Papilionaceae (Dipteryx, Fig. 59, F,
p. 268) and Boragineae (Cordia, Fig. 126, C, p. 557)1. A kind of doubling of the
unicellular body of the hair in the longitUdinal direction owing to the formation
of a cap of cellulose is found in the Cis~eae (Fig. 21, A, p. 81), Combretaceae
(Fig. 76, A, 13, p. 346), and Myrtaceae sens. str. The following hairs are distin·
guished by having special contents: the stinging hairs of the Loaseae, Euphor-
biaceae and Urticeae (Fig. 183:P. 776),-these hairs being occasionally inserted on
a pedestal composed of a large number of cells; the crystal-containing hairs
of certain Euphorbiaceae (Fig. 180, R, S, p. 748) 1, which are differentiated as
papillae and enclose either a clustered crystal ,or a sphaerite; the stinging hairs
of certain Euphorbiaceae, which are occasion'ally situated on a pedestal resem·
bling a shaggy hair, and con tain an acicular crystal of oxalate of lime, suspended
in the cavity of the cell by means of beams of cellulose (Fig. 180, P, Q, p. 748) 1 ;
and the crystal-hairs (Fig. 101, B, C, p. 446) of the Guettardeae (a tribe of
the Rubiaceae), which are paved with one or more rows of small crystals
of oxalate of lime. The formation of more or less distinct papillae on the
walls of the hairs, in which case the lumina of the hairs mayor may not pene~
trate into the papillae~ has been recorded in the unicellular trichomes of certain
Crudferae, Bixineae, Geraniaceae, Chailletiaceae (Fig. 46, p. 198), Rosaceae,
Saxifragaceae (Fig. 68, A, p. 316) and Cornaceae. Regarding calcification of the
cell-wall in unicellular hairs and regarding cystolith-hairs, see §§ 27 and 28.
The following noteworthy modifications of uniserlate trlchomes may be
mentioned. The uniseriate clothing hairs are sometimes characterized by the
fact that the lower or lowest cells of the hair are short, while the terminal cell
is long and forms the greater pru-t of the trichome. Such trichomes with
long terminal cells have been observed in the following Orders : Magnoliaceae,
Trochodendraceae, Anonaceae, Menispermaceae, Berberideae, Nymphaeaceae,
Pittosporeae (Fig. 22, B, p. 93), CaryophyUeae, Sabiaceae, Papilionaceae (Fig.
59, A-C, p. 268), Caesalpinieae (very rare), Umbelliferae, Compositae (Fig. 103,
A, B, p. 458), Polemonjaceae, Boragineae, Convolvulaceae (in all the members
'of the Order! Fig. 129,A, p. 567), Solanaceae, Retzia, Acanthaceae, Verbenaceae,
Labiatae, Plantagineae, Amarantaceae, Chenopodiaceae, Aristolochiaceae, Pro~
teaceae (Fig. 173, E, p. 713), Casuarineae (Fig. 186, E, p. 788). Uniseriate hairs

I It may be pointed out in passing that the crystal. hairs and stinging hairs mentioned above as ~
occurring in certain Euphorbiaceae. although they appear like trichomes, are "",t hairs in the strict
sense (i. e. according to a scientific interpretation of their morphology), inasmuch as they arise (rom
subepidermal and not (roUl ef,idermal cells; only secondarily do they push their way between the
epidermal cells, thus ultimate y extending beyond the level of the epidermiS. For details, see under
Enphorbiaceae, p. I05J.
 ,HAIRY COVERING 11[9

with a capitate terminal cell, which, however, has no secretory function, have
l
been observed in many Amarantaceae and Chenopodiaceae (Fig. 159, C, p. 659),
as well as in certain Papaveraceae, Compositae (Fig. 103, E, p. 458), Nyctagineae
(here glandular? Fig. 154, A, p. 646) and Illecebraceae. Other special1:Y note-
worthy forms are constituted by the urn-shaped trichomes and the paired uni-
seriate hairs, with fused ba~ portions, found in species of Gilia (Polemoniaceae),
and the uniseriate hairs with a sunken ~sal portion, divided by closely placed
transverse septa, occurring in certain Hippocrateaceae and Polemoniaceae. The
unicellular bracket-hairs, two-armed hairs, stellate and pelt ate hairs find their
equivalents in uniseriate trichomes, iJl which the terminal cell shows a corre-
sponding shape. Uniseriate trichomes with a terminal cell bent in the
form of a hook are found in certain Papilionaceae, Gesneraceae, Verbenaceae
and Aristolochiaceae (Fig. I66, K, p. 684; in the Order last named the cells of
the stalk occasionally also show division-walls running parallel to the length of
the hair); two-armed hairs, in which the stalk is unicellular or uniseriate,
and the terminal portion for the most part unicellular, occur in certain Pitto-
sporeae (Fig. 22, A, p. 93), Rhamneae, Aceraceae, Papilionaceae, Caesalpinieae,
Sa:xifraga~ae, Araliaceae, Cornaceae (Fig. 99, D, p. 435), Compositae (Fig. 103.
F, p. 458), Boragineae (Fig. 126, E, p. 557; in Cordia the two arms are occasion-
ally septate), Convolvulaceae (Fig. 129, E, C, p. 567; occasionallyone-a.rmed, the
long arm being septate), Amarantaceae (Fig. 156, F, p. 652), Chenopodiaceae
(Fig. 159, B, p. 659), Myristicaceae and Proteaceae (Fig. 173, F, p. 713); uni-
seriate trlchomes with a stellate or peJtate terminal cell are found in certain
Chlaenaceae (Fig. 33, p. 145), Papilionaceae (Fig. 59, E, p. 268), Ficoideae
(Fig. % C, p. 416), Compositae (Fig. 103, D; p. ~·s8),:Convolvulaceae (Fig. 129,
F, G, p. 567), Amarantaceae (in Iresine the terminal cell shows transitions to a
two.;armed differentiation, Fig. 156, E, po 652), Chenopodiaceae (Fig. 159, G, H,
p. 659) and Loranthaceae (?). With the trichomes last menti(j~ed we may class
uniseriate candelabra-hairs, in which the individual tiers are composed either
of two-armed cells-the arms of which point in different directions (Dampiera,
Goodeniaceae, Fig. 106, p. 471) or lie in the same vertical plane (Myristicaceae,
Fig. 170, D-F, p. 698)-or of cells, developed after the manner of i stellate hair
(Compositae, Amarantaceae, Fig. 156, D, p. 652, Loranthaceae, Fig. 177. C-E,
p. 728). Uniseriate hairs, in which the division-walls aTf~ oblique, whil~ the
ends of some or all of the component cells are drawn out mto lateral papillae,
or; in other words, push their way laterally along the basal portion of the cell
immediately above, are found in certain Polemoniaceae, Plantagineae (Fig. 153,
p. 643), Nepenthaceae (Fig. 165). and Loranthaceae (Fig. 177, B~ p. 728) j such
hairs constitute transitions to trichomes with sympodial structure (see below).
-Multicellular hairs, which vary in shape and are branched in a dendroid
or dichotomous manner, have been observed in the Orders Guttiferae (Fig.
28, p. 123), Ternstroemiaceae (Fig. 30, A, p. 133), Olacineae (Fig. 48, A, p. 204),
Lecythidaceae,-Melastomaceae, Lythrarieae, Araliaceae, Campanulaceae, Primu-
laceae (Fig. 114. A, p. 502), Myrsineae, Apocynaceae (Fig. I21, B, p. 530), Logania-
ceae, Polemoniaceae, Boragineae (Fig. 126, B, p. 557), Nolaneae (Fig. I3 2, p. 574),
Solanaceae (Fig. 133, p. 577), Scrophularineae, Gesneraceae, Bignoniaceae
(Fig. 141, A. p. 603), Myoporineae, Verbenaceae, Labiatae (Fig. 152, p. 63 8),
Nyctagineae (Fig. 154, B, p. 646, glandular ?), Illecebraceae, Amarantaceae,
Euphorbiaceae (rare) and Casuarineae; it is possible to derive such hairs
from uniseriate trichomes by assuming a protrusion of the component cells
into branches, demarcation of these branches, and the appearance of division-
walls in them, this being followed by the protrusion of branches of the second
Order, and so on. Special forms of these trichomes are constituted by the
peculiar hairs of 1acquinia (Myrsineae, Fig. 116, C, p. 510) and by branched or
tufted hairs, which in typical casea consist of flattened cells, each of which is
lI20 CONCLUDING REMARKS

protruded to form a single branch (i.e. the component cells are one-armed).
Since the axial portion of such trichomes shows a sympodial relation to the
branches, they may be described as sympodially branched hairs; they an.
more or less markedly differentiated in certain CaryophyIIeae, Celastrineae
(very faintly indicated), Connaraceae (Fig. 55, D-F, p. 252), Bignoniaceae.
Myoporineae, Nepenthaceae (Fig. 1.65, C, p. 679). Myristicaceae (Fig. I70,
p. 698), and Casuarineae (Fig. 186, D, p. 788). A brief reference may still be
made to the manifold transitions between branched trichomes, derived from
uniseriate hairs, and the trichomes dealt with in the following paragraphs.,
The wall of the uniseria te trichomes, as in the case of the unicellular hairs.
is occasionally (Papilionaceae.-here th~ wall of the terminal cell,-Polemonia-
ceae, Amarantaceae, Fig. 156, p. 652, and Chenopodiaceae, Fig. 159, p. 659)
provided with small papillose protrusions, while in certain Compositae (Fig. 103,
C, p. 458), Plantagineae (Fig. 153, B, p. 643). and Amarantaceae (Fig. 156,
C, p. 652) the upper ends of the cells composing the hairs bear papillose processes.
which establish a firm connexion between the ceUs. Regarding calcification
of the walls of the hairs, see § 27.
§ 32. PELTATE, STELLATE AND CANDELABRA-l!AIRS. In this section our
summary will be confined to those pelt ate, stellate and candelabra-hairs, in
which the shield, stellate portion and radiating tiers respectively are composed
of two or more cells." Pelt ate and stellate hairs, which with or without the
stalk consist of a single cell. and candelabra~hairs, the tiers of which are com-
posed of single cells drawn out into rays, have already been dealt with in § 3I.
The three forms of hairs in question are:very closely related to one another.
From a stellate hair, in which the ray-cells are spre~,d out ,horizontally. we can
derive a pelt ate hair by the coalescence of the '.r.ay-ceUs~~ile a candelabra-
hair is obtained by the repeated development of ray-cells (a"'rl:d~ometimes of
stalk-cells as well) in the longitudinal direction. We accordingly, find that
stellate and peltate hairs, or stellate and candelabra-hairs, not unCommonly
occur side by side with one another in the same taxonomic group' or even in
the same species; in such cases the two forms of hairs may show the same
special type of structure, while transitional forms between them are of frequent
occurrence.
The peUate hairs 1 (the delimitation of which from the peltate glands
dealt with under I in § 34 occasionally-when an investigation of living material
is impossible-presents some difficulties) show a very diverse structure, and
the differences can be employed for systematic purposes in the same way as
the presence or absence of these trichomes; the latter are sometimes (Elaeag-
naceae) of constant occurrence throughout an Order, although generally can·
fined to certain species. The differences in the structure of the peltate hairs
are afforded either by the shield or by the stalk. The former for the most part
consists of a single layer of cells and varies in size. In the ordinary and most
widely distributed type of peltate hairs the shield is composed of radially
arranged cells, of which all or only some reach the centre of the shield; these
ray~cells have walls of varying thickness and mostly project at the margin
of the shield in the form of short rays. In other cases the shield has an entire
margin, the ray-cells being broadened towards the outside in the shape of
a wedge. The shield presents still a different appearance, w~en the radially
arranged ray-cells undergo tangential divisions. If irregular divisions follow
the appearance of the first radial division-walls, the shield seems to be com-
posed of polygonal cells, when seen in surface-view. In the following cases
the shield shows quite a special type of structure. In certain Buddleioideae

A See O. Bnchmann, Schilclh3.are, Flora, 1886.

 HAIRY COVERING II21

(Fig. 124, p. 542) we find peltate hairs with a bicellular shield. In the specie$
of Solanum (Fig. 133, p. 577) a spherical or otherwise shaped cell is seated on
the centre of the shield of the peltate hairs, which in other respects show the
ordinary type of structure; in certain Capparideae (Fig. Ig, p. 75), Malvaceae
and Sterculiaceae there is a so-called' tipper scale in the form of a rosette of
J

small cells, while in certain species of Croton (Fig. 180, p. 748) a similar rosette
of small cells is situa ted beneath the shield, constituting a so-called 'lower scale,'
the shield in the latter case being provided with a central cell. In certain species
of Miconia (Melastomaceae) there is a small upper scale composed of short ray·
cells, and in some cases also a small lower scale. A doubling of the shield in the
radial direction is found in Phebalium (Rutaceae, Fig. 41, p. 177) and in certain
Bombaceae (Malvaceae, Fig.34,P. 149); in these cases most of the cells extending
outwards from the centre of the shield stop short of the margin, while those
forming the margin do not reach the centre, so that the marginal cells can·
stitute, as it were, a duplication of the central cells, which in themselves are
already united to form a scale. It remains to mention the pelt ate hairs of
Clerodendron (Verbenaceae, Fig. l5I, p. 632), in which the shield is composed of
several layers of cells showing a polygonal outline, when seen from the surface.
The stalk of the peltate hairs is formed either by a single cell or by a row of cells,
or by a multiseriate complex of cells, its length often varying very considerably
on the surface of one and the same organ ; in some cases (Dilleniaceae, Logania e

ceae, Euphorbiaceae pro parte) the cells of the shield are themselves either
exclusively or partially concerned in the formation of the stalk.
Peltate hairs with a shield, composed of two or more 'cells, have been observed
in the following Orders and genera respectively: Dilleniaceae (Fig. 2, p. 22), Ano-
naceae (Fig. 6, p. 36), Capparideae (Fig. 19, p. 75), Cistineae (Fig. 2 I, p. 8 I). Bixineae,
Malvaceae (Fig. 34, p. 149), Sterculiaceae (-Fig. 35. p. 154). Tiliaceae, Rutaceae
(Fig. 41, p. 177), Burseraceae (Zanha, Hiern), Meliaceae (Fig. 45. p. 197), Olacineae
(Fig. 48, p. 204), Rosaceae, Halorageae (Fi~. 72, p. 337, Callitriche, Hippur,'s),
Melastomaceae, Begoniaceae, Datisceae, Araliaceae (Fig. 98, p. 429), Caprifoliaceae
(Fig. 100. p. 441), Ericaceae (Fig. 110, p. 485). Myrsineae (Fig. 116, p. 510), Styraceae,
Oleaceae (Fig. 119. p. 523), Salvadoraceae (in this Order there is only a tendency to
form small scales, Fig. 120, p. 527), Loganiaceae (Buddleioideae, Fig. 124, p. 542),
Solanaceae (Fig. 133. p. 577), Verbenaceae (Fig. 151, p. 63Z), Monimiaceae (Fig. 171,
p. 701), Elaeagnaceae (Fig. 176, p. 725), Euphorbiaceae (Fig. 180, p. 748), Cupuli-
ferae (Fig~ 187, p. 793). For glandular peltate hairs, see under Ib in § 34. p. IIz8.
The stellate hairs, according to the direction of their rays, which generally
consist of single cells, are either stellate hairs in the narrow sense or tufted
hairs; in the former case the rays are spread out in,a plane, which is roughly
parallel to the surface of the organ, while in the case of the tufted hairs the
rays diverge. in various directions or are all placed approximately at right
angles to the surface. In the simplest case, the ray-portion, consisting of
unicellular rays, is sunk directlyjn the epidermis, so that the trichome is formed
merely by a -group of epidermal cells, developed as hairs; the neighbouring
epidermal cells are frequently prolonged onto the body of the hair, thus forming
a pedestal of varying height. In other cases the rays are borne on a stalk,
which varies in length, and is composed of one or more rows of cells;
it occasionally resembles a shaggy hair. The rays are rarely uniseriate.
Other differences in the structure of the stellate hairs, which can be employed
for systematic purposes, are found in the length and number-of the rays, and
in the nature of their walls and lumina. A considerable number of transitions
have been observed between the branched multicellular hairs, dealt with in
§ 31 (p. 1II9) and the hairs under discussion.
Stellate hairs have been met with in the following Orders and genera respec-
tively: Dilleniaceae (Fig. 2, p. 22), Magnoliaceae ?, Anonaceae. Capparideae (Fig. 19,
p. 75), Cistineae (Fig. 21, p. 81), Bixineae, Tremandreae. Vochysiaceae. Frankenia-
SOLUEDU 4 C
 Il22 CONCLUDING REMARKS
ceae, Hypericineae, Ternstroemiaceae (Fig. 30, p. 133), Dipterocarpeae, Monous,
Malvaceae (Fig. 34, p. 149), Sterculiaceae, Tiliaceae, Rutaceae, Ochnaceae, Bursera-
ceae, Meliaceae, Olacineae, Octocnemaceae, Hippocrateaceae, Rhamneae, ~pin'­
daceae, Melianthaceae, Connaraceae, Caesalpinieae, Mimoseae, Rosaceae, Saxifraga-
ceae, Droseraceae (special forms of stellate hairs, see Fig. 7 I, p. 325), HamameIideae,
Rhizophoraceae, Myrtaceae, Melastomaceae (Fig. 79, p. 362), Lythrarieae, Samy-
daceae, Turneraceae, Begoniaceae, Umbelliferae, Araliaceae (Fig. 98, p. 429),
Cofll!B.Ceae, Caprifoliaceae, Rubiaceae, Lobeliaceae, Ericaceae (Fig. 110, p. 485),
EbeIiaceae, Styraceae, Loganiaceae (Fig. 124, p. 542), Hydrophyllaceae, Boragineae,
Convolvulaceae (Fig. 129. p. $67), Solanaceae, Scrophularineae, Verbenaceae, Labia a

tae (Fig. 152, p. 638), N yctagmeae, Chenopodiaceae (Fig. 159. p. 659). Nepenthaceae,
Monimiaceae, Elaeagnaceae, Santalaceae, Euphorbiaceae (Fig. 180, p. 748), JugJan 4

deae, Cupuliferae.
Tl;le following types of stellate and tufted hairs deserve special notice.
The spiny hairs (Fig. 2, B, C, p. 22) of the Dilleniaceae are reduced -tufted
hairs, consisting of a group of strongly silicified epidermal cells, which are
developed as short hairs; if we imagine the marginal ce)ls of the group to be
differentiated as rays, we obtain th~ stellate trichomes of the Dilleniaceae.
Cushion-shaped structures, which are similar to the spiny hairs of the Dillenia-
ceae, although they are not. silicified, occur also in the Cistineae (Fig. 21, p. 81)
and Melastomaceae. The stellate hairs of the genus Stet'iphoma (Capparideae,
Fig. 19, p. 75) resemble th~ peltate hairs of the Capparideae in being provided
with a small upper scale, while some of the stellate hairs found in the Cistineae
show the same apparent doubling of the ray-cells'in the longitudinal direction
(Fig. 21, p. 81), as is seen Wthe simple trichomes occurring in this Order.
Stellate or tufted hairs witli uniseriate ;rays have been observed in certain
Magnoliaceae?, Anonaceae, Malvaceae (Fig. 3~ p. 149), Sterculiaceae, Tiliaceae,
Hippocrateaceae, Rhamneae, Lecythidaceae and Ericaceae (Fig. 110, p. 485).
while in the stellate hairs of Axyris (Chenopodiaceae, Fig. 159, p. 659) the
principal ray alone is uniseriate. The two-armed hairs of Ald1'ovanda (Fig.
71, p. 325), which show transverse division into two cells, may be regarded
as a reduced form of the stellate hairs of certain of the Droseraceae, in which
the ray-portion is composed of four or more thin-walled. rays, expanded in
one plane. The stellate hairs of the Loganiaceae-Buddleioideae are likewise
specially characteristic~ since the ray-portion consists of two (mostly two-rayed)
cells, which also contribute to the formation of the stalk (Fig. I24, p. 542).
Mention may lastly be made of: (a) the tufted hairs of SauTauja (Temstroemia'-
ceae, Fig. 30, p. 133) and certain Melastomaceae, which resemble small scales ;
(b) the tufted hairs of Santiria (Burseraceae, Fig. 43, p. 19I) and Rhus (Ana.;
cardiaceae), which may possibly have a glandular function; and (c) the peculiar
stellate hairs of some Araliaceae (Fig. 98, p. 429), which are provided with
uniseriate rays having thin walls.
The simplest kinds of candelabra- or abietiform hairs are those which
have a uniseriate main axis, interrupted at intervals by whorls of ray-cells.
Such hairs are occasionally connected by means of transitional forms with the
stellate and tufted hairs just discussed, hairs of the latter type not uncommonly
3:ppearing as reduced forms of the candelabra-hairs and then often occurring
side by side with them in the same species. The candelabra-hairs of the
second type present a different appearance, owing to the fact that the ray-cells
follow upon one another in the longitudinal direction without intenuption,
there being no special cells of the main axis separating. the rays from one
another. These candelabraahairs are often difficult to distinguish from the
branched sympodial hairs discussed in § 31, p. lI20; and similarly, if the stalks
of such candelabra·hairs are multiseriate, It is not easy ~o distiriguish them
from candelabra-hairs of the shaggy type, so that the latter may be considered
together with the ordinary candelabra-hairs. Candelabra..hairs of the shaggy
type may be derived from ordinary shaggy h~ir.s (I 33) by the development of
 HAIRY COVERING

superficial cells of the latter in the form of hairs (this process occasionally
taking place in tiers).
Candelabra-bairs of the first type have been observed in certain Lythrarieae,
Ericaceae (Fig. II 0, p. 485), BuddIeioideae (Loga.niaceae, Fig. 124, p.,S42, the
tiers here consisting of two (mostly two-rayed) cells, as in the case of the analogous
peltate and stellate hairs, see above), Scrophularineae (Fig. 135. p. 586).Verbon~,
Chenopodiaceae (Fig. 159, p. 659), and Platanaceae (Fig. 184, p. 780); candel<lbra-
hairs of the second type occur in certain Capparideae (Fig. 19, p. 7 S)~ Melastomaceae
(Fig. 79. p. 362), Uinbelliferae (Fig. 95, p. 4 21 ), Solanaceae (Fig. 133, p. 577), and
Acanthaceae; candel~bra-hain ot the shallY type, lastly, are_ found in certain
Ciesalpinieae, Mimoseae (Fig. 65, p. 295), Melastomac.eae (Fig. 79, p. 362). Samy-
daceae, Labiatae (Fig. 152, p. 638), and Euphorbiaceae (Fig. lBo, p. 748,.
§ 33. SHAGGY HAIRS (Le. clothing hairs of the shaggy type, villi). Simple
ii.e. unbranched) shaggy hairs are widely distributed in many Orders. They
vary in length and thickness, and consist of parenchymatous or prosenchyma-
tous cells with thin or thick (sclerenchyanatous) walls. The filiform shaggy
hairs are freq.ently distin~hed by the fact that the apices of the superficial
cells are drawn out into papillae, which point towards the end of the shaggy
hair; a more extensive development of the apices of the cells results in the
production of trichomes, which are transitional to the candelabra-hairs of the
shaggy type, already discussed at the end of § 32.
Simple shaggy hairs, which otherwise vary in shape. are found in the following
Orders: -Papaveraceae, Capparideae (Fi~. 19, p. 7S).._ "Portulaceae (Fig. 26, p. I d),
Temstroermaceae (Fig. 3D, p. I ~3), Papilionaceae (Pig. 59, p. 268), Caesalpinieae,
Mimoseae (Fig. 6S. p. 295), Saxlfragaceae, Crassulaceae, Droseraceae, Halorageae,
Mvrta,ceae sens. str., Melastomaceae (Fig. 79, p. 362), Lythrariea.e; Begoniaceae
(Ag. 88, p. 401), Cacteae, Umbelliferae, Araliaceae (Fig. 98, p. 429), Compositae
• Fig. 103. p. 458), Ericaceae (Fig. 110, p. 485). Convolvulaceae, Gesneraceae,
Polygonaceae and Euphorbiaceae.
In the following cases the shaggy hairs are- speclally dlfl'e.rentiated.
Silicified wart.shaped shaggy hairs are found in Gunnera; in C,anoca,pus
(Papilionaceae) t~e tip of the shaggy hair, has the form of.an ~chor, the/flukes
Qf the anchor bemg fonned by pomted cells, each of which mcludes a crystal
of oxalate of lime (Fig. 59, G_ p. 268). which is pointed like the cell containing it;
in certain Melastomaceae, which exhibit a great degree of diversity in the
-forms' of the shaggy hairs, the latter occasionally show surface-development,
so that they come to resemble the leaf of a Moss (Fig. 79, p. 362), while in
certaiD Begonias they are twoearmed or star-like (Fig. 88, p. 40I); in the
genus PiC1'is (Compositae, Fig. 103, p. 458) the shaggy hair is crowned by
a'two-armed cell, while the forked shaggy hairs of Leontodon, which are well
1mown to systematists, are produced by two or more of the terminal cells of
the trichome 'being differentiated as hairs. The shaggy hairs of Picris are
closely related to the emergences of Humulus, each of which bears a two-
anned trichome.- Rttiuced forms of shaggy hairs are found on the floral
organs of the Compositae (Fig. 103, P: 458); they cortsist either of t.wo b~l
cells and two longer cells, the latter lymg parallel to one another and (m Sphae e

1'alUhus) embracing one another spirally, or merely of two cells, which are placed
parallel to one another (as in the sensitive hairs of the Cynareae). The peculiar
hollow structures, fow::nd in Heterotrichum (Melastomaceae), must also be "in-
~ded among the shaggy hairs; their wall is formed by a prosenchymatous
epidermis and a layer of red\1ced palisadeacells. With the· shaggy hairs of
PScris and Humulus we may 'associate trichomes having (1 stalIc. of varying
length and a stellate or tuft~ terminal portion; these structures have already
been dealt with under stellate and'tufted hairs in § 32. The· candelabra-
hairs of the shaggy type' have likewise been considered in the same section.
Sheggy hairs, which are provided with' glandular heads,.will be discussed und:er
4C2
1124 CONCLUDING REMARKS

the heading of glandular shaggy hairs in the section dealing with the external
glands (see § 34 under II, p. II29).
Branched shaggy hairs have been observed in Calandrinia (Portulaceae ;
these are the pili plumosi/ mentioned by different authors) and in certain
I

Melasto&aceae (the hairs here being penicillate or branched in a dendroid

manner and at the same time parenchymatous).
The shaggy hairs, as already stated in § 30, are partly epidermal structures
and partly of the nature of emergences. To the latter category there belong.
among other types of trichomes, the shaggy hairs of certain Begonias and
Melastomaceae, which are stiffened by means of isolated sclerenchymatous
fibres or by whole bundles of such fibres and (in the Melastomaceae) occasion-
ally include a vascular bundle in their basal portion. In the Melastomaceae
the bundle of sclerenchyma in these shaggy hairs is frequently connected
with cells of the mesophyll, which are differentiated like fibres; in those
shaggy hairs, which are placed obliquely to the surface of the organ, these
fibres fonn, an anchoring foot, consisting of a bundle of sclerenchymatous
fibres, which spread out for some distance beneath the epidermis; on the
other hand, when the shaggy hair is placed at right angles to the surface, the
fibrous foot is branched after the manner of a root (Fig. 79, p. 362); the shaggy
hairs of Cirsium (Compositae) show similar features. We may add that
a similar sclerenchymatous foot has been observed also in the Mimoseae (Fig. 65,
p. 295) and Euphorbiaceae (Fig. 180, p. 748) in connexion with trichomes of
a more complicated type, such as candelabra-, peitate and stellate hairs.
Certain species of Croton are quite particularly noteworthy in this respect; in
these species the bases of the hairs are formed by a bundle of sclerenchymatous
cells, which traverse the leaf in the vertical direction, while the bases of two
trichomes, situated opposite one another on the upper and lower surface of
the Jeaf, are fused to form a sclerenchymatous column, traversing the entire
thickness of the leaf (Fig. 180, p. 748).
At this point we may also refer to the branched or unbranched structures resem
bling shaggy hairs found in the intercellular spaces of Nelumbium (Nymphaeaceae).
§ 34. HAIR-LIKE EXTERNAL GLANDS l~ The demonstration of external
glands and the investigation of their structure in herbarium-material often pre·
sents great difficulties. It is a familiar fact that glandular hairs are often
functional only on the young organs of a plant, and in this case fall off in later
stages or shrivel up, so that their structure becomes unrecognizable. As a
consequence external glands are best sought after and examined on the young
leaves, found on the branches of herbarium-material, or even on the young
branches themselves (especially those of the floral region); but, whenever it is
at all possible, living material should also be employed in determining the pre·
sence of external glands and in the elucidation of their structure and function.
In certain Orders (e.g. Sapindaceae, Oleaceae, &c.) hair-like external
glands are of general or almost general distribution; in other' Orders they are
restricted to certain genera, while in others again glandular hairs are completely
absent. It is worthy of note that the subsequent enumeration of Orders, in
which external glands have not been observed, includes whole series of closely
allied Orders, as well as many Orders, in which the secretory requirements of
the plant are apparently sufficiently satisfied by internal secretory organs.
As a rule the hair-like external glands occur on all the organs of the plant,
although in varying abundance; in other cases-and this holds good for
species, as well as for genera and Orders-they are found only on certain .parts
of the plant. Thus in the Rubiaceae, for instance, glandular hairs are almost
completely wanting on the surfaces of the leaves and on the branches, while
1 For the hair·like iotemal glands ~internal glandular hairs) found in P~gtl;te"tlm (Labi4tae).
~e under secretory cells in § [4 (p. 1096). .
 HAIRY COVERING 1125

external glands, having a complicated structure and serving for the protection
of the buds, occur OIl the stipules, and glands t excreting nectar and showing
the same· type of structure, .He found.in the flowers. It is necessary to hike
such features into account in an investigation of the hairy covering.
Another difficulty, with which one is confronted in employing the occurrence
of external glands for systematic purposes, was already briefly referred to in
§ 29, and lies in the fact that the glandular hairs are not in all cases distinguished
morphologically, and that the presence of secretion cannot always be· deter-
mined in herbarium-material with as much certainty a.s is desirable. In
addition to that the ordinary clothing hairs may exceptionally take on a. secre-
tory function in certain Orders (e.g. the uniseriate.hairs of the Nymphaeaceae
and Gentianeae, and the two-armed trichomes of the Sapotaceae 1).
HaiT-like external gJa,nds have not been ob2ioerved in any form in the follow-
ing Orders and genera respectively: Dilleniaceae, Calycanthaceae. Magnoliaceae.
Trochodendraceae. Lactoridaceae, Anonaceae, Sarraceniaceae (with external glands~
which are not of the nature of hairs. see § 38), Papaveraceae, Fumariaceae, Reseda-
ceae, Violarieae (apart from glandular shaggy hairs on the margin of the leaf),
Canellaceae, Polygaleae, Vochysiaceae (apart from nectaries, see § 36), Frankeniaceae
(apart from chalk-glands, see § 37), Tamariscineae (apart from chalk-glands, see § 37),
Hypericineae, Guttiferae, Ternstroemiaceae? (apart from nectaries, see § 36).
Humiriaceae (with the exception of glandular shaggy hairs on the margin of the leaf),
Malpighiaceae (apart from nectaries. see § 36), KoebeYlinia, Ochnacea.e, Luxemburgia-
ceae (excepting glandular shaggy hairs found on the stipules and sepals of the
Godoyeae), Wallacea. Chailletiaceae. Olacineae ? (see p. I t 17), Octocnemaccae,
Ilicineae. Cyrilleae, Celastrineae, Hippocrateaceae, Pentaphyiacaceae, Corynocarpa-
ceae, Stackhousieae, Rhamneae (apart from glandular shaggy hairs situated on or
near the margin of tb.e leaf and (see § 36) the nectaries). Didiereae, Melianthaceae~
Staphyleaceae (apart from nectaries, see § 36), Coriarieae, Moriageae (apart from
nectaries, see § 36), Crossosomataceae, Hamamelideae, Ost1'earia, MYYQthamnus,
Bruniaceae. Rbizophoraceae (apart from glandular shaggy hairs 011 the stipules).
Myrtaceae sens. str., Samydaceae (apart from nectaries, see § 36). Papayaceae. Cacteae
(see § 36). Ficoideae, Umbelliferae? Araliaceae (almost always), Rubiaceae (apart
from gland ular shaggy hairs on the sti pules, &c.). Calycereae. Cam pan ulaceae, Lobelia-
ceae. Epacrideae. Diapensiaceae. Sapotaceae (two-armed clothing hairs occasionally
glandular). Brackynema. Styraceae ? (see p. 1117), Salvadoraceae. Apocynaceae and
Asclepiadeae (apart from glandular shaggy hairs at the base of the petiole and the
gland ular hairs of Disch £dia ), Loganioideae (Loganiaceae, a part from gland ular shaggy
hairs at the base of the petiole), Desjontainea, Gentianeae (apart from uniseriatc
glandular trichomes in the aquatic forms and glandular bodies, composed of nume-
rous cells, in saprophytes), Cuscuteae (apart from glandular shaggy hairs on the
flower),' Columelliaceae. Nyctagineae? Amarantaceae? Phytolaccaceae, Bati-
deae, Aristolochiace~e. Chloranthaceae, M yristicaceae ?, Moniruiaceae, Laurineae.
Gomortegaceae, Proteaceae?, Thymelaeaceae, Octolepis, Gonystylus. Penaeaceae.
Geissoloma, Elaeagnaceae, Lorantbacea.e, Santalaceae. Myzodendyon, Champe,.eia,
Gyubbia, Balanophoreae, Daphniphyllaceae, Buxaceae ?, Balanopseae, Thelygoneae,
Casuarineae. Sa.1idneae (apart from nectaries. see § 36), Lacistemaceae; but also in
certain genera of numerous other Orders.
,~

The hair-like external glands show numerous points of difference as regards

their size, shape, and structure (number, arrangement, and structure of the
cells,' and localization of the secretion). and it is almost impossible to comprise
these in a single system, applicable to all the various Orders. For, one and the
same form of trichome occurring in two different Orders may be of <hfferent
I A combination of the glandular and clothing functions in simple hairs., ,as well as in tricbome~
of more complicated stnlctlJre. has been observed in the Orders Portulaceae, CaesalpUtieae, Metasto-
maceae (particularly frequent in the tribe Miconieae, in which combinations of glaodulat hairs with.
stel1ate, tufted. candelabra- or peltate hairs occnr in almost all the genera), Loganiaceae, Solanaceae,
Myoporineae, Nyctagineae (see I"ig. 154, p. 646) and Platanaceae; in hairs oUbis type the glandular
portion ma.y be restricted to a sirJgle ceil or may be differentia.ted as a multicellular glaodnlar body,
which, as it were, is independent of the rest o( the hair. For details, ~e the anatomical descriptions of
the OHlers name.].
 JJ26 CONCLUDllrG REMARKS

phylogenetic. importance in the two cases; thus the uniseriate glandular hairs
of the Solanaceae, in which the head is unicellular or divided by a vertical
wall, form the starting-point for external glands with a uniseriate stalk and
a variously shaped head, divided by horizontal or both by horizontal and vertical
walls, while the glandular hairs of a similar structure, found in the Scrophula-
rineae, constitute a starting-point for external glands, the head of which is
divided by vertical walls only, that is to say for quite another type of hair.
For the purpose of the practical determination of a plant, we may in the
following paragraphs first (under I) summarize especially the small forms of
glandular hairs, which have a simple structure, in contrast to other types of
glandular hairs, which are mostly of large size and generally have a com-
plicated structure (see under II); it must, however, be pointed out that the
boundary-line between the two kinds of hairs is artificial and therefore not
sharply marked. The small glandular hairs may then in the first place be
classified in two further group:=; in accordance with the remarks made in § 29 ;
the second of these groups (I b) includes tho!Se forms of glandular hairs, in
which the head is divided solely by vertical walls, while the other (I a) com-
prises all the remaining forms.
I a. Unicellular gla1!dular hairs, which for the most part have a tubular
·shape. have been observed in the following Orders: Ranunculaceae (Fig. I~
p. 16). l\Ienispermaceae, Malvaceae, Tiliaceae, Zygophylleae (Fig. 38, p. 168),
Geraniaceae. Rutaceae, Olacineae ? (Fig. 48, p. 204), Ampelidaceae, Anacardia~
ceae, Papilionaceae, Onagrarieae, Cornaceae (Fig. 99, p. 435), Compositae (latici ..
ferous hairs; which stand in connexion with the laticiferous vessels), Dischidia
(Asclepiadeae), Monotropeae, Symplocos ? (Styraceae), Podostemaceae, Pipera~
ceae (in the form of large pearl-glands, Fig. r68, p. 689), Euphorbiaceae (secre~
tory cells of Croton, which are differentiated like papillae, see § 14 and Fig.
180, p. 748).
The multicellular glandular hairs, to be discussed here, are very diverse.
In the simplest case they consist merely of a row of cells, which is either filiform
or broadened towards the upper end in a clavate manner. The appearance
of longitudinal divisionawalls in most of the cells, or only in the terminal portion
of the hair, results in the formation of complicated, filiform or club-shaped glan A

dular hairs, which are occasionally curved or involuted in a peculiar manner.

Other types of glandular hairs are differentiated into a stalk and a head.
The stalk varies in length, and is unicellular, uniseriate or composed of a few
rows of cells, while the head~ which varies in size, is spherical or ellipsoidal, or
otherw.ise shaped, and consists of one or many cells; in those cases, in which the
head is multicellular~ it may be divided by horizontal walls only, or both by
horizontal and vertical walls or quite irregularly.
The multicellularglandular hairs in question have been observed in the following
Orders and genera:· Menispermaceae (uIDseriate), Berberideae (rare), Nymphaeaceae
(uniseriate. in part very short, Fig. 12, p. 49), Cruci;erae (uniseriate, head uni· or
bicellular, Fig. IS, p. 63). Capparideae (diverse types, head in part unicellular),
Cistineae (uniseriate, glandular hairs in part very long and filiform, Fig. 21. p. 81),
Bixioeae (various types), Pittosporeae (uniseriate). Caryophylleae (uniseriate, head
unicellular. Fig. 25, p_ 108). Portulaceae (uniseria.te, Fig. 26. p. 112), Ternstroemia-
ceae ? Dipterocarpeae (unicellular peltate head. Fig. 3 I. p. 137; see I b), M onotes
(uniseriate), Chlaenaceae (various types), Malvaceae (various types, inci. uniseriate
~lands with a unicellular head; see I b). Sterculiaceae (various types), Tiliaceae (as
1U the Sterculiaceae), Geraniaceae (uniseriate with a unicellular, rarely multicellular
head. Fig. 39, p. 170). Rutaceae (various types, Fig. 41, p. 177), Simarubaceae
(diverse types. Fig. 42, p. 183), Burseraceae (varioUs types, Fig. 43. p. 191), Melia·
ceae (various types, Fig. 45. p. 197), Sapindaceae (various types; see I b). Hippo-
castanaceae (as In the Sapindaceae), Aceraceae (various types), Sabiaceae (head
unicellular; see I b). Anacardiaceae (various types; see I b): J1tlz"ania t (head multi·
1 According to F. K Fritsch, Trt\n~. Linn. ~oc. London, vii. l908.
 HAlRY COVERING
cellular), Connaraceae (only on the floral organs), Papilionaceae (various types, heads
in part unicellular, Fig. 59, p. 268; see I b), Caesalpinieae (diverse types, Fig. 63,
p. 287), Mimoseae (various types, Fig. 65, p. 295; see I b), Rosaceae (various !ypes).
Saxifragaceae (various types, heads in part unicellular; see I b), Crassulaceae (di verse
types). Combretaceae (various types j see I b), Lecythidaceae (groups of uniseriate
glands ?). Melastomaceae (various types; see I b), Loaseae (uniseriate). Turneraceae
(various types; see I b), Cucurbi taceae (head uni - or m ul ticellular, its cells occasionally
drawn out mto lappets; see I b), Begoniaceae (various types, Fig. 88, p. 401 I head in
part hammer-shaped; multicellular pearl-glands), Umbelliferae (??; certainly very
rare). Araliaceae (uniseriate), Cornaceae (various types), Caprifoliaceae (various types;
see I b), Valerianeae (multicellular heads; see I b), Dipsaceae (as in the Valerianeae),
Compositae (various types, commonly biseriate vesicular glandularhairs, Fig. 103,
p. 458; see I b), Candolleaceae (heads of diverse types; see I b), Goodeniaceae (multi.
cellular heads; see I b),Vacciniaceae (various types; in some cases biserlate ligulate
glandular hairs, Fig. lOS. p. 47S), Ericaceae (various types, occasionally uniseriate,
Fig. 110, p. 485; see I b), Monotropeae (various types, see I b). Lennoaceae (uni-
seriate glands with a unicellular head). Primulaceae (uniseriate glands with a uni-
cellular head, Fig. 114, p. 502; see I b). Myrsineae (head unicellular; see I b), Ebena·
ceae (uni- or multicellular heads; see I b), Plocosperma, Polemoniaceae (heads uni-
cellular or divided by means of horizontal walls into several tiers, which are themselves
divided by vertical walls; see I b), Hydrophyllaceae (various types, in part with
unicellular heads'; see Ib), Boragineae (heads unicellular and variously shaped,
Fig. 126, p. 557), Convolvulaceae (various types, in part uniseriate, Fig. 12 9, p. 567;
see I b), N olaneae (heads of di verse types, also unkell ular), Solanaceae (various types,
heads in part unicell ula.r, Fig. 133, p. 577; see I b). R etzia (diverse types), Scrophu-
larineae (unicellular heads; see I b), Orobanchaceae (as in the Scrophularineae),
Lentibularieae (as in the Scrophularineae). Gesnel'aceae {as in the Scrophularineae},
Acanthaceae (as in the Scropnularineae), Myoporineae (as in the Scrophularineae),
Verbenaceae(as in the Scrophularineae). Labiatae (rarely unicellular heads, which are
of large size and have a cuticle, which is strongly elevated in a vesicular manner,
otherwise as in the Scrophularineae), Plantagineae (various types; see I b). Nycta-
gineae (uniseriate, in part branched, with unicellular heads, Fig. 154, p. 646; hairs
really glandular ?). Illecebraceae (uniseriate with unicellular heads), Amarantaceae
(as in the Illecebraceae. glandular ?), Chenopodiaceae (uniseriate, Fig. I 59, p. 659),
Polygonaceae (unicellular heads; see I b), Pi peraceae (unicellular heads; hydathodes,
Fig. 168, p. 689) Myristicaceae (see p. 699). Hernandiaceae(IIUgera with a transversely
septate. bicellular head. Fig. 172, p. 708). Proteaceae (Fig. 173. p. 7 I 3), E u phor biaceae
(various types. heads in part unicellular; see I b), Ulmaceae (various types), Canna·
bineae (Fig. 18 I , p. 769 ;. see I b), Moraceae (various types, in part u niseria te; see I b),
Urticeae (see Ib), Platanaceae (unicellular h!!ads), Leitnerieae, Myricaceae (various
types, in part uniseriate; see I b), Cupuliferae (diverse types, in part uniseri ate ,
Fig. 187. p. 793; see I b), Empetraceae (various types, heads in part unicellular,
Fig~ 188, p. 800) .
. It remains to mention the following special forms of the multicellular
glands (apart from those already indicated in the preceding review by the
use of heavy type): (a) the pelt ate glandular hairs of certain Papilionaceae
(Fig. S9~ p .. 268), Mimoseae and Droseraceae (Fig. 7r, p. 325). which have a
shield composed of two layers of cells; (b) the spherical .glands of certain
Papilionaceae' (especially .Phaseoleae) , in which abundant secretion is deposited
between the cells of the head (which are in loose connexion with one another>.
and in a subcuticular position (Fig. 59, p. 268); (c) the peltate or spherical
glands of certain Papilionaceae (Fig. 59), Caesalpinieae (Fig. 63, p. 281)
and Lythrarieae (Fig. 80, p. 370), which enclose an intercellular secretory space;
(d) the branched external glands of certain species of Santiria (Burseraceae,
Fig. 43~ p. 191), Rhus (Anacardiaceae) and Mimosa (Mimoseae), and of Cono-
cephalus and Sparattosyce (Moraceae); (e) the penicillate mucilage·hairs of
certain species of Candollea (Candolleaceae), which show special peculiarities
in certain stages of their development, and are proviqed with two or more
uniseriate rays, which are bi- or multicellular; (f) lastly, the external glands
of certain Melastomaceae, which have two or four, or even more heads situated
on a common stalk, and the paired glands, found in species of Acer (~ceraceae).
 CONCLUDING REltiARKS
in which the unicellular stalks are fused lengthwise with one another. Finally,
brief mention may be made of the occasional combination of glandular hairs
in groups (Compositae, Ficoideae 1, Moraceae, see also § 36 under nectaries),
and the outgrowth of cells of the stalks of the glandular hairs in a papillose
or hair·like manner (in certain Melastomaceae).
I b. The glandular hairs, the heads of which are -divided solely by
means of vertical walls 2, may be divided into those which have peltate heads,
and those which have spherical heads. The former for the most part have
short stalks, while the latter generally have longer stalks, the stalks often
being of considerable length. The number and mode of arrangement of the
cells forming the head (which is of variable size) varies, occasionally even in
the trichomes of one and the same species. In most cases there is only a slight
accumulation of secretion beneath the cuticle. Vesicular integumental glands
of the peltate type, in which the cuticle is raised like a bladder owing to the
formation of abundant secretion, have been recorded only in a few Orders
(Mimoseae, Combreta,ceae, Fig. 76, p. 346, Melastomaceae, Turneraceae?,
Fig. 83, p. 382, Compositae?, Anthotroche, Bignoniaceae, Fig. 141, p. 603,
f'
Verbenaceae, Labiatae, Fig. 15 2 , 638, Euphorbiaceae, Cannabineae, Fig. 181,
p. 769, and Myricaceae). Specia forms of these glands are constituted by the
mucilage-hairs of certain species of C,andollea, in which the cells of the head are
rod-shaped, and the well-known kamala-glands of Mallotus (Euphorbiaceae) with
a head, composed of dub-shaped cells. Other special forms of the glandular
hairs in question are as follows. Large peltate glands, which are differentiated
after the manner of the intra·mural glands, are found in Rhododendron and Ledttm
(Ericaceae, Fig. IIO, p. 485), while specia~ types of ·glands, the heads of which
are divided solely by a single vertical wall, occur in some of the Buddleioideae
(Loganiaceae, Fig. 124, p. 542). Characteristic peltate glands -(Fig. 135,
p. 586) are widely distributed in the Rhinanthaceae; these glands consist of
a uru- or multicellular basal portion, a middle cell and a group of 2-4 lid·cells,
covering the middle cell. Peltate glands of quite a similar type are found
also in Monophyltaea (Gesneraceae), the glands here excreting chalk and having
two lid-cells. Other particularly noteworthy forms are: (a) the glandular
hairs of certain LentibuIarieae (Fig. 139, p. 596), which are provided with
a stellate or two-armed terminal portion, composed of four and two cells
respectively, these glands resembling the stellate hairs of certain Droseraceae,
already mentioned in § 32; (b) the curious forked glands of Primulina (Gesnera-
ceae); (c) the glandular hairs of certain Gesneraceae, which are provided with
a stellate head with 3-4 rays; (d) the mucilage-glands of many Pedalineae, in
which the outer walls of the glandular head, which consists of four or more
cells, become mucilaginous (Fig. 143, p. 612); (e) the peculiar peltate glands
of the Myoporineae (Fig. 149, p. 626), in which the stalk is excentric; (I) the
shortly stalked chalk-glands of the Selagineae (Fig. ISO, p. 629), the heads of
which consist of two cells and mostly have two knobs; (g) the external glands
of certain Gesneraceae and of the Thunbergieae (Acanthaceae, Fig. 144;P. 615),
which are provided with a hammer-shaped or biscuit-shaped head, divided by a
t'ransverse wall; and (h) the peculiar external glands of Littorella (Plantagineae,
Fig. 153, p.643).
Glandular hairs with a spherical or pelt ate head, divided e::scclusivcly by one
or morC vertical walls, and consisting of two or many cells. are found in the following
Orders and genera: Bixineae, Dipterocarpeae (Fig. 3I, p. ] 37 j see I a), AnC£slrocladus
(Fig. 3z, p. 143), Malvaceae (scarcely typical; see I a), Burseraceae, Sap.illdaceae

I In Gli'scarothamntts, according to Pilger, in-Engler, Bot. Jabrb., xl, 1908, p. 397.

2 The term' vertical wall' is not always to be interpreted in the exact geometrical sense. When
the heads of the glandular haiJ:s consist of numerous cells and aJe strongly arched outwards, the
division-walls namely converge towards the stalk, the cells of the head in such cases being generally
elongated in a palii:'ade-like mAnner,
 HAIRY COVERiNG

(see I a), Hippocastanaceae (see I a), Sabiaceae (see la), Anacardiaceae (see I a),
Papilionaceae (very rare; see I a), Mimoseae (Fig. 65, p. 295; see I a), Saxifragaceae
(see I a), Droseraceae (Byblis), Halorageae (Fig. 72, p. 337), Combretaceae (Fig. 76,
p. 346; see I a), Melastomaceae (see I a), Turneraceae (Fig. 83, p. 382; see I a),
Cucurbitaceae (see I a), Caprifoliaceae (Fig. 100, p. 441 ; see 1 a), Valerianeae (see
I a). Compositae ? (see 1 a), Dipsaceae (see 1 a), Candolleaceae (see I a), Goodeniaceae
(see 1 a), Ericaceae (Fig. 1 10, p. 485; see I a), Monotropeae (see 1 a). Primulaceae (see
la), Myrsineae (Fig. Il6, p. 510; see la), Ebenaceae (see la). Oleaceae (Fig. IJ9,
.
523), Loganiaceae (Fig. 1 2 4, p. 54 2 ), Polemoniaceae (s~ 1 a), Hydrophyllaceae (see
f a), Convolvulaceae (Fig. 129, p. 567; see I a), Solanaceae (Fig. 133. p. 577; see I a),
Scrophularineae (Fig. 135, p. 586; see 1 a), Oroban~haceae (see I a). Lentibularieae
(Fig. q6, p. 592; see J a), Gesneraceae (Fig. 140, p. 599; see I a), Bignoniaceae
(Fig. 141, p. 603),·Pedalineae (Fig. 143, p. 612), Acanthaceae (Fig. I44. p. 615 ;--see
I a), Myoporineq.c (Fig. 149. p. 626; see I a). Selagineae (Fig. 150, p. 629). Verbenaceae
(Fig. 151. p. 632; see I a), Labiatae (Fig. 152, p. 638; see la), Plantagineae (Fig: .[ 53,
p. 643; see I a), Polygonaceae (see I a), Nepenthaceae {Fig. 165. p. t579}, Cytinaceae,
Euphorbiaceae (see I a). Canna.bineae (Fig. 18 I, p. 769; see I a), Moraceae (see I a),
Urticeae (see I a), Juglandeae, Myric::)..ceac (see I a). Juglandeae, Myricaceae (see I a),
Cupuliferae (Fig. 187. p. 793 ; see ,I a).
II. Hair-like external glands 1, which are generally of large size and
in most cases have a complicated structure. We ~ay first mention the.
following forms. Multicellular lageniform or urceolate glands (Fig. 34, p. 149)
occur in certain Malvaceae. Pearl~glands. which are compOsed of many (some-
times very many) cells. are found in certain Sterculiaceae, Ampelidaceae (Fig. 52.
p. 223), Caesalpinieae, Begoniaceae, Moraceae, and Urticeae; some of these
glands have an epidermis of small cells, which includes a stoma situated on
the side opposite to the stalk. Spherical glands with a palisade~epidermis and
a pair of guard-cells, situated on the side opposite to the short stalk, occur in Leea
(Ampelidaceae, Fig. 52, p. 223). Small. almost sessile, cucullate glands are found
at the base of a depression (which is provided with a narrow aperture) in the sur-
face of the plant in Laguncularia(Fig. 76, p. 346) and Conocarpus (Combretaceae).
Other forms requiring mention are the lageniform external glands of Dictamnus
(Rutaceae, Fig. 41, p. 177), the basal portion of which includes a secretory cavity,
and the cylindrical emergences of Eucalyptus (Myrtaceae, Fig. 77, p. 353), which
likewise contain a secretory cavity. The external glands of Dictamnus are closely
related on the one hand to the conical glandular shaggy hairs of Cuphea lanceolata
(Lythrarieae, Fig. So, p. 370), which are composed of numerous cells and contain
a grollP of secretory cells in their basal portion, and on the other hand to the
bulbous hairs of certain Papilionaceae, Caesalpinieae (Fig. 59, p. 268, and Fig.
63, A, p. 287) and Turneraceae (Fig. 83, p. 382), in which there is no secretory
cavity in the swollen base of the gland. The retort-shaped glands of certain
Caesalpinieae (Fig. 63, D, p. 287), lastly, constitute somewhat reduced forms of
bulbous h~rs, the basal portion of which is sunk into the tissue of the organ
bearing them; with them we may class the retort~shaped hairs of Bellucia
(Melastomaceae), which are merely uniseriate and have a basal portion, divided
by numerous:septa~":-The remaining external glands, to be included here, are
glandular shaggy hairs, having a multiseriate stalk of varying length and a
head, which is either (rarely, in Menispermaceae, Fig. 7, p. 41, Papilionaceae,
Con volvulace ae, Cuscuteae) unicellular or irregularly multicellular or (in very
many cases, see Fig. 101, p. 446) consists of a central core of cells, which are
elongated in the direction of the length of the stalk and are enveloped by one
or more layers of secretory palisade~cells. The stalk (and sometimes also the
core in those glands, the heads of which are provided with a palisade-epithelium)
often contains a vascular bundle or the termination of a vascular bundle.
Glandular shaggy hairs, showing the type of structure just described, are
occasionally (e.g. in the Rosaceae and Euphorbiaceae) connected by transi-
tions with glandular spots, having an identical structure (see § 36).
I For the e:cternal glands found in insectivorous plants, see § 38, p. I133.
 CONCLUDING REMARKS

Glandular shaggy hairs have been observed in the following Orders and
genera: Menisperinaceae (Fig. 7. p. 41). Cruciferae (Fig. IS. p. 63), Capparideae,
Tremandreae, Elatineae. Tiliaceae. Lineae. Geraniaceae. Simarubaceae (Fig. 42.
p. 183), Luxemburgiaceae (Godoyeae. on the stipules and sepals), Rhamneae (for
the most part only on the margin of the leaf), Sapindaceae. Aceraceae (occa-
sionally separating into two uniseriate hair-like filaments at the apex, see p. 893),
Papilionaceae, Mimoseae, Rosaceae, Saxifragaceae (Fig. 68, p. 316), Penthorum,
Crassulaceae, Droseraceae (digestive glands, showing a uniform type of structure,
Fig. 7 I. p. 325), Halorageae (Fig. 72, p. 337), Rhizophoraceae (generally on the stipules
only), Melastomaceae, Lythrarieae, Turneraceae (Fig. 83. p. 382), Passiftoraceae (Fig.
84, p. 385), Begoniaceae, Datisceae, Araliaceae, Rubiaceae (only on the stipules and
floral organs, Fig. 101, p. 446). Compositae. Candolleaceae, Vacciniaceae, Ericaceae.
Monotropeae, Plumbagineae, Apocynaceae (on the petioles, &c.), Asclepiadeae
(for the most part as in the Apocynaceae), Loganiaceae (as in the Asclepiadeae).
Gentianeae (rare, in the axils or on the surface of the leaves), Convolvulaceae, Cuscu-
teae (in the dower). Solanaceae (Fig. 133. p. 577 ), Polygonaceae, N epenthaceae (pitcher.
and nectarial glands), Euphorbiaceae. Thelygoneae. Ceratophylleae (apices of the
leaves). For details as to the structure of the glandular shaggy hairs, see the
individual Orders.
The varied nature of the secretion, produced by the hair·like external
glands, has already been repeatedly referred to (see also § 29), and it is only
necessary to add that excretion of. carbonate of lime or of some other salt
has been observed in certain Papilionaceae, Convolvulaceae, Scrophularineae,
Gesneraceae, Bignoniaceae and Selagineae. In some cases the secretion of
resin is so considerable that the leaves become covered with a perfect layer of
varnish 1.
External glands, which are sunk in the surface of the leaf, occasionally
give rise to transparent dots. I

§ 35. GLANDULAR LEAF-TEETH 2; are found in species belonging to a large

number of Orders. According to the type of structure they show they may
be grouped in three categories. In the first case the leaf·teeth bear the same
kinds of glandular hairs as occur on the surface of the leaf; in the second case
they have a secretory epidermis, which is differentiated like palisade, or they
are completely transformed into glandular shaggy hairs, the epidermis of
which shows the same differentiation; and in the third case they include the
termination of a vascular bundle, which is associated with an epithema (often
containing mucilage), the epidermis above the latter containing one or more_
water-pores. Leaf·teeth of the first and second types secrete resin, while
those of the third type secrete mucilage or water (the latter in some cases
contains chal~ which becomes deposited on the surface of the plant, e.g. in
the Violarieae and Saxifragaceae). The process of secretion frequently takes
place only in the young leaf. The glandular leaf·teeth in some cases no doubt
function also as extrafloral nectaries.
The structure of the glandular leaf-teeth has been examined in the following
Orders: Ranunculaceae, Trochodendraceae, Cruciferae, Violarieae, CaryophyUeae.
Temstroemiaceae, Tiliaceae. Humiriaceae, Geraniaceae. Celastrineae, Rhamneae,
Aceraceae, Staphyleaceae. Ampelidaceae, Papilionaceae, Rosaceae, Saxifragaceae.
Hamamelideae, Halorageae. Onagrarieae, Cucurbitaceae, Caprifoliaceae, Dipsaceae,
Compositae, Campanulaceae, Vacciniaceae, Primulaceae. Gentianeae, Polemoniaceae,
Scrophularineae. Verbenaceae, Labiatae, Ulmaceae, Cannabineae, Moraceae,
Juglandeae, Cupuliferae, Salicineae. For details, see unuer the individual Orders.
§ 36. LARGE GLANDULAR MECHA~ISMS (NECTARIES) 3. At this point
I Volkens, Lack. Bl., Sitz.·Ber. deutsch. bot. Geselbch. 1890, p. 120.
, See Reinke, in Pringsbeim Jahrb., x, 1876, p. (19; and Virchow, Bl:tttzahoe, Archi,
d. Pharm., 1896.
, See espedlllly Poulsen, Trik. og Nekbr., Vidensk. Meddel. paturh. For. Kjt<thenb3.m, 187Fo.
abstr. in Just, 1875, p. 1013: Bonnier, Nect .• Ann. sc. nat., ser. 6, t. viii, 1878: lPa\'is, Nect:lr·
 HAIRY COVERING 113 1

I propose to give a collective account of the large glandular mechanisms,

generally described as nectarial glands or nectaries. The glands in question
vary very much in structure. They do not always form a secretion, which
contains sugar or is of the nature of honey, for the secretion is sometimes
mucilaginous or like wax, while in other cases {in the hydathodes} it is no
doubt merely water ; so that the term 'nectary' is not always suitable.
Owing to their large size these glands are visible even to the naked eye, and
have therefore been recorded in many Orders by systematists. In sonw
Orders they are of such general occurrence that they constitute a diagnostil:
character aiding in the recognition of a member of the Order, while in
other Orders they are found either in all (e. g. Qual~a. Vochysiaceae) or only
in certain species of a genus. Only those nectaries, which appear on the
vegetative organs, will be considered here; very commonly, however, nectaries
occur on the inflorescence and on the sepals, either side by side with those on
the vegetative organs or to the exclusion of the latter. The position, in which
they occur on the leaf, varies. They may be present to the number ,of one
or more on either side of the base of the petiole in the form of disc-shaped
or wart·like glands; in other cases they are shifted up on to the petiole, where
they constitute wart-like appendages, or are developed in the form of depressed
callosities on its upper side, while not uncommonly they are situated at the
limit of lamina and petiole, appearing as discoid, wart-shaped or pit-like struc-
tures. In other cases again they are found on the lower side of the lamina,
although sometimes restricted to the basal portion; the nectaries on the
lamina are developed as pit-like callosities on the midrib or apex of the leaf,
as disc-shaped structures on the leaf-margin, and as shallow pit·like or disc-
shaped glands, or as mere spots on the lower surface of the leaf, where they
occur in varying numbers and occasionally show a definite arrangement with
reference to the veins. Nectaries are rarely found also on the upper surface
of the leaf. They have, however, also been observed on the stipules, and in
some cases they are metamorphosed stipules or stipels (Sterculiaceae, C-apri.
foliaceae, Leguminosae, Combretaceae); in the species of Capparis, belonging
to the section Cynophalla, DC., moreover, they are transformed foliage-shoots.
The structure of the nectaries is very diverse, and on the -basis of these differ-
ences in structure the majority of the nectaries may be classified in two series,
which are connected with one another by". transitional forms. The nectaries
of the ,first series are composed of groups of small glandular hairs, which are
fou~d either (a) on the surface of the organ, or (b) in shallow depressions in the
surface, or (c) clothing deep pits of diverse structure in the surface of the
organ, these pits often being provided with a narrow ostiole, or (d) lining the
inner wall of hollow glandular bodies, which have an ovoid form (Olacineae).
The structure of the glandular hairs forming these nectaries is, as a general
rule, identical with that of the external glands, which are found as isolated
trichomes on the vegetative o~gans of the species in question, although the
secretory portion is"usually more strongly developed. To this series belong
the nectaries found in certain Menispermaceae, Malvaceae, Sterculiaceae,
Olacineae, Papilionaceae (Fig. 59, p. 268), Oleaceae, Convolvulaceae and
Polygonaceae, as well as the glandular zones on the stems of certain Sileneae
(Fig. 25, p. 108). The simpler forms of the nectaries of the first series are
related to the groups of glandular: hairs, mentioned in § 34 (p. 1128), and the
latter can to some extent be regarded as initial stages in the d'evelopment of

gland. on leaves, Hot. Gllzette, 1883. p. 339]; Delpino, Funz. mirmecof., Mem. Ace. Bologna, 1886.
p. uS t and 1888. p. 601 ; Morini, Nett. estranuz., Mem. Ace. Bologna, 1886. p. 3:ZS, and 6 tah.,
here also the older literature i Haupt. Sekr~tionsmechanik d. extranupt. N ekt., Flora, '90l, p. "
also Dils., Leipzig; Sc~wendt, ExtraAorale Nektarien, Beib. '1.. Bot. Centralbl., uii. Abt. I, 190;.
p. 245 et seq. and ttlb. IX..
 1132 CONCLUDING REMARKS

such nectaries. The forms of nectaries, comprised in the second series, are
more widely distributed than those of the first series. The former in the
first place show all transitions between bodies, which project in the form of
a head, wart, or disc, and may occasionally even be stalked, and structures
formed solely by a group of superficial cells on the organ bearing the gland.
In the two extreme types of the nectaries belonging to this series, as well as
in their intermediate forms, th~ glandular tissue may consist throughout of
small polyhedral cells with thin wails, or the epidermis of the gland may be
differentiated as a palisade~epithelium; when the latter is strongly elongated,
each of its prismatiG cells is frequently divided by a transverse wall. Pit-like
nectaries, which occasionally have a number of subsidiary excavations and
open to the exterior by means of a wide -or narrow orifice, are very rare, but
have been observed, for example, in Ma'tcgravia (Ternstroemiaceae), Cf}prosma
(Rubiace~e), Fagraea (Loganiaceae), and in the Nepenthaceae (Fig_ 164, p. 678);
the pits in these nectaries are clothed by one or several layers of secretory
palisade-cells. I

In the f611owingsynopsis of the Orders, in which' exirafioral nectaries' occur,

details as to their shape and distribution are added only in those cases in which the
structure of the glands has not yet been investigated, so that, as a general rule, they
have not been considered in the earlier part of this book: Menispermaceae, Cappari-
deae (axillary nectaries in species of Capparis belonging to the section Cynophtllla).
Bixineae (two large nectaries at the base of the leaf in Scolopia and I desia. also
glandular leaf-teeth), Vochysiaceae (impressed nectaries on either side of the base
of the petiole in QuaJea), Caryophylleae, Ternstrcemiaceae (Marcgravieae), Diptero-
ca.rpeae, Malvaceae, Sterculiaceae, Tiliaceae (glandular leaf-teeth in Grewia and
Triwntetta), Humiriaceae, Malpiglliaceae (Fig.; 36, p. 164), Geraniaceae (glandular
warts on the petiole in Impatians), Rutaceae (in Boronia, also discoid or wart-shaped
nectaries on the lower side of the leaf in spedeg of Z anth'oxylon), Simaro baceae
(in Ailanthus and Cadell-I:a, also wart-like nectaries on both surfaces of the leaf in
species of Samaaera). Olacineae, Rhamneae, Staphyleaceae (two glands on the base
of the leaf in HUfirtea), Anacardiaceae (wart-like nectaries on the uP1?er part of the
petiole or in place of the stipels in Hohgarna). Moringeae (nectanes, which are
occasionally stalked, at the base of the pinnules and on the petiole in Moringa),
Papilionaceae (Fig. 59, p. 268), Caesalpinieae, Mimoseae, Rosaceae, Combretaceae,
Lecythidaceae, Lythrarieae (thick bodies, provided with an open pore and situCtted
at the end of the midrib of the leaf in Lajoertsia, acarodomatia ?), Samydaceae
(nectaries ...t the apex of the petiole or on the margin of the leaf in Banara), Turnera·
ceac, Passifloraceae. Cucurbitaceae (nectaries of diverse shape, often discoid,
situated either en the entire lower side of the leaf or confined to the base of the leaf
in species of Abob1'a, A den opus. A lsom it1'a , Bryonia, Cephaland'f'a, Cucu1'bita, Feuillea,
Lagenaria, LuOa, Momord'lca, Sphaeros'icyos, Trlanosperma, Trichosanthes). Cacteae
(3ee Delpino), Caprifoliaceae. Rubiaceae, Compositae (at the base of the foliage-
leaves in th~ floral region of Helianthus), Vo.cciniaceae, Ebenaceae (nectaries on the
lower side of the leaf in specieE of Diospyros and Royena), Olea~eae. Loganiaceae,
Convolvulace ae. Scrophnlarin.eae, _Bignoniaceae (Fig. 1419 p, 603), Verbellaceae,
Polygonacea.e. Nepenthaceae: Euphorbiaceae (Fig. 180, p. 748), Moraceae, Sali-
cineae.
For groups of water-pores (hydathodes) appearing as small pits, warts or spots
on the surface of the leaf, see § 5, p. 1086.
Before leaving this subject '\Ie may point out that the glandular spots and pits,
which have been included among the nectaries, must not be confused with so-called
acarodomatia 1. The latter are commonly differentiated as small pits or pockets,
and have been recorded in the following Orders: Ma~cliaceae, Anonaceae 9 Meni-
spermaceae. Violarieae, Bixineae, Ternstroemiaceae (mel. Marcgravieae), Diptero-
carpeae. Stercuhaceae. Tiliaceae, Meliaceae, Ilicineae, Rhamneae, Sapindaceae,
Aceraceae, Anacardiaceae, Rosaceae, Saxifragac::eae (Ribesieae), Hamamelideae,
Combretaceae. :L\lyrtaceae, Melastomaceae, Lythrarieae, Cornaceae. Caprifoliaceae,
Rubiaceae. Compositae, Sapotaceae.Oleaceae, Apocynaceae, Asclepiadeae, Logania e

I See especially Penzig e Chiaberra. Piante acarofile, Malpig~ia, 1902, p. 413 et seq.
 HAIRY COVERING II33
ceae, Boragincac, Solanaceae, Bignoniaceae, Verbenaccae, Piperaccae, Laurineae,
Euphorbiaceae, Urticaceae (incl. Ulmaceae), Platanaceae, Juglandeae. Cupuliferae.
§ 37. CHALK- A~D SALT-GLANDS of the Frankeniaceae, Tamariscineae and
Plumbagineae. These glands are distinguished by the fact that they are
not differentiated as hairs. Their structure is simple and identical in the
Frankeniaceae (Fig. 24. p. 106) and Tamariscineae, while in the Plumbagineae
(Fig. 1I3. P..: 497) it is of a more complicated type. In this Order one also
finds mucilage-glands. which show a similar~structure. apart from the fact
that they consist of still more numerous cells (Fig. 113). Mucilage-glands,
which are differentiated as hair~J have also been observed in the Plumbagineae,
while glandular shaggy hairs occur on the inflorescence, and in respect of their
structure both kinds of trichomes can be derived from the above-mentioned
mucilage-glands. which are not of the nature of hairs.
For glandular hairs. excreting chalk or other salts, see the end of § 34
(p. 1130); regarding leaf.teeth, which excrete chalk, see § 35 (p. 1130).
§ 38. SPECIAL FORMS OF. EXTERNAL GLANDS OCCURRING IN INSECTIVOROUS
PLANTS. The glands of the Sarraceniaceae are not hair-like, although in
other respects they show diverse structure (see p. 53 et seq. and Fig. 13);
glands of a similar kind are found also in Cephalotus (see p. 3I9 and Fig. 69).
The Nepenthaceae have nectarial glands, which are not of the nature of hairs,
but take the form of small pits, the wall of. which is lined by three layers of
secretory cells (see § 36); in this Order one also finds nectarial and digestive
glands. which are differentiated as hairs and have 2-3 layers of secretory
cells (see p. 677 et seq. and Fig. I64). The glands of the Droseraceae (Fig. 7I,
p. 325) are always developed as hairs and in all the genera (with the sale
exception of the genus Byblis, which, according to Lang, must be transferred
from the Droseraceae to the Len tibularieae) are distinguished by having two
layers of secretory cells and a suberized middle layer.
§ 39. CORK- \V ARTS on the leaves. In certain species the lower sides
of the leaves bear numerous brown dots, which are often styled glands in
anatomical descriptions, and may therefore be discussed at this point! These
dots, however, as shown by a microscopical examination, are not glandular,
but represent'local formations of cork-so·called cork·warts. Mention may
also be.made here of the constant occurrence of corky tissue at the apices of
the leaves in the Bruniaceae .
. Cork-warts have been observed in the following Orders: Berberideae, Gutti-
ferae, Ternstroemiaceae, llicineae (Fig. 50, p. 210), Celastrineae, Hippocrateaceae,
Chrysobalaneae, Saxifragaceae. Hamarnelideae, Rhizophoraceae, Myrtaceae sens.
str., Melastomaceae. Lythrarieae, Araliaceae (only on the petiole), Apocynaceae,
Loganiaceae•. Gesneraceae, Piperaceae, Laurineae ? Loranthaceae. Euphorbiaceae.

V.- NORMAL STRUCTURE OF THE AXIS.

§ 40. MEDULLARY TISSUE. Gris 1 was the first to show that the presence
or absence of starch in the pith, as in the case of the endosperm of the seed,
is a character of systematic importance. He distinguishes (i) active medullary
cells, i. e. cells storing starch and generally having rather thick walls, (ii) empty
medullary cells, i. e. dead cells filled with air and for the most part having-
1 Gris, Moell~, Nouv. Arch. du Mus. d'hist. nat., t. vi, 197~, p. :20l and pI. xi~-xx, and also Ann.
se. na.t., ser. 5. t. XIV, 1872. p. 34 and pl. 4-7 ; Kassner, Mark etnlger Holzpfi., DISS., Breslau; 1884,
38 pp., :I Tab.; Magoscy-Dietz, Diaphragma in dem Mark d. dikotyl. Holzgew., Math. u. natllrwisSi.
Ber. auSi Ungarn, xvii, 1901, p. 181 et seq. (this paper deals with nodal, and not with internodal
diaphragms); [Fox-worthy, Discoid pith in woody plants, Proc. Indiana Acad. SC. IQO~ (1904).
pp. 19 1 -4·J
 CONCLUDING REMARKS

relatively thin walls, and (ill) cells containing crystalline elements or some kind
of secretion. The occurrence of an empty or active pith, consisting of empty
cells only, or of active cells only, as the case may be (secretory cells or cells con-
taining crystals being present or absent), or the occurrence of a heterogeneous
pith, formed both by empty and active cells, is characteristic of taxonomic
groups of varying magnitude. The varied distribution of the empty and
active cells in a heterogeneous pith may, moreover, also be employed as a
systematic character. The active cells either form a network between the
empty cells or are confined to the periphery of the pith, the central portion
of which in the latter case is occupied by empty-cells. The fact that the active
cells do not contain starch all the year round, so that they cannot always be
recognized in herbarium·material with as much certainty as is desirable, COD·
stitutes an obstacle to the extensive practical use of all these features.
The occurrence of spicular cells and of special secretory elements in the
pith has already been dea_It with in the previous sections (see § 9 and § 14
et seq.). It remains to mention the following features: (a) the pith of certain
species of Myzodendron, which consists throughout of prosenchymatous cells ;
(b) the groups of sc1erenchymatous fibres. situated at the outer margin of
the pith and in direct cont{l~t with the primary xylem·groups of the vascular
bundles, in certain Menispermaceae, Malvaceae, Corynocarpaceae, Araliaceae,
Polygonaceae, Loranthaceae, species of Myzodendron, Proteaceae, Platanaceae,
Salicineae, &c., and the ring of sclerenchymatous fibres, occupying the same
position in the young branches of species of Piper; (c) lastly, the occurrence
of isolated scIerenchymatous fibres or of groups of ,such fibres in the interior
of the pith in Lophira and in certain Meliaceae, Geraniaceae, Rutaceae, Lythra·
rieae, Araliaceae (Aralidium), Plumbagineae, Asclepiadeae, Euphorbiaceae, and
Salicineae.
The occurrence of groups of stone-cells in the pith is generally only of
importance for specific diagnosis. Horizontal diaphragms, composed of stone-
cells and forming transverse septa in the pith, possess greater systematic value;
they have been observed in many Magnoliaceae, Anonaceae (Fig. 5, p. 35),
and Ternstroemiaceae, as well as in Brachynema and certain Convolvulaceae.
Other features of systematic value are the complete or partial disappear-
ance of the pith-the fistular character of the stem in older internodes (Umbelli-
ferae !), and especially the septation of the pith without sclerosis (Fig. 185,
p. 784), which goes hand in hand with a partial disappearance of the pith. The
last feature has been observed in the following genera, of which it is generally
(but not always, e. g. Ja.sminum) characteristic: Wormia (Dillen iaceae) ,
Diplotaxis (Cruciferae), Fouquiera (Tamariscineae), Prinsepia (Chrysobalaneae),
A ucuba (Cornaceae, only in herbarium·material !), Senecio (Compositae),
Halesia (Styraceae), Jasminum (Oleaceae), Paulownia (Scrophularineae),
Pedalitlm (Pedalineae), Phytolacca decandra (Phytolaccaceae) 1, Daphniphyllum
(Daphniphyllaceae), Juglans and Pterocarya (Juglandeae).
§ 41. ApPEARANCE OF THE TRANSVERSE SECTION OF THE STEM IN HER·
BACEOUS PLANTS. The great diversity, shown by the appearance of the trans-
verse section of the stem in herbaceous plants, is mainly due to the varied
extent of development or to the complete or partial absence of mechanical
tissue accompanying the vascular bundles on their inner and outer side, and
to the diverse $tructure of the secondary wood, formed within the vascular
bundles, and of the interfascicular tissue (presence or absence of wood·fibres,
vt"ssels and medullary rays) 2. With the help of these features we are able to

~ According to Mirbe\ and Holm, see Holm, in Merck's Report, xvi, 1907.
~ See S~hwt"ndenerl Mech. Prinz.ip, Leipzig.· 1874, p. li3 et seq. nnd Tab. xiv.
 NORJIAL STRUCTURE OF THE AXIS 1135
distinguish a number of different types of transverse section, but it would
be out of place to enter into a detailed description of them at this point, since
it has been shown (see Ranunculaceae, Cruciferae, &c.) that they are only of
very limited systematic value (in most cases they can be employed only for
specific diagnosis); and, apart from that, the occurrence of intermediate
forms often renders distinction between the principal types a difficult matter.
In certain Ranunculaceae .(~ig. I, p. 16), Berberideae 1, and Fumariaceae.
and according to Schwendener also in certain Plumbagineae and Compositae,
the line of demarcation between the phloem and xylem of the vascular bundles,
as seen in a transverse section of the stem, has the shape of an arc, the concave
side of which (and not the convex side, as is generally the case) is directed
outwards 2.
§ 42. SYSTEMATIC IMPORTANCE OF THE STRUCTURE OF THE WOOD in the
woody Dicotyledons 3. The structure of the wood affords a whole series of
anatomical characters, which are generally distinctive of genera or even of
tribes or Orders. The most important of these characters are as follows :
the kinds of perforations found in the~ vessels; the structure of the walls of
the vessels in contact with parenchyma of the medullary rays or with wood-
parenchyma; the nature of the pits (simple or distinctly bordered) on the
walls of the wood-prosenchyma (wood-fibres), which forms the ground-mass

I In Caukphyllum tlzalictroMes, MichX., according to a recent observation of my own.

I For the structure of subterranean stems, see Costantin, Tiges aero et souto des Dicotyl., Ann.
sc. nat., ser. 6, t_ xvi, 1883, p. 5 et seq., and Noelle, Vergl. anat. Untersuch. der AusHiufer, n1s5.,
.Freiburg i. Br., 1892,72 pp.
S Sanio, in Bot. Zeit., 1863, pp. 85 and .lSi et seq.; Rossmann, Bau des Holzes, Frankfurt a. M.,
1865, 100 pp.; De Bary, Vergl. Anat•• • 871. p. 493 et seq.; Moller, Beitr. z. vergl. Anat. des Holzes,
Denkscbr. Wiener Akad., xxxvi, 1876, pp. 297-426 and Tab. i-vi; Solereder, Systemat. Wert der
Holzstruktur, Miinchen, 1885, 26. pp. j Vesq\le, in Bull. Soc. bot de France, 1889, p. lxv; Houlbert,
Eois sec. dans lea Apetales, Ann. $C. nat., ser. 7, t. xvii, 1893, also These, Paris, 1893; Petersen,
Diagnostisk Vedanatomi, Kjta'benhavn, 1901, 93 pp.; Stone, Anatomical characters in the identi~cat.
of wood, Nature, lxv, 1901-2. pp. 379. 380; Wiesner, Rohstoffe, ~. Aufl., Bd. ii, 1903, P: I et seq.;
Moll and Janssonius, Mikro~raphie des Holzes der auf Java vorkommenden Baumarten, Heft I.
Leiden, 1906, 368 pp. (Di11emacea.e to Dipterocarpeae, in the sequence of Durand's Index), especially
Allgem. Teil, pp••0-62; Perrot et Gerard, Anat. du tis!lu ligneux d3.DS ses rapports avec 1& diagnose
des bois, Bell. Soc. bot. de France, Mem. 6, 1907, +3 pp., 6 pL
As regards the nomenclature of the elements of the wood, 1 IWly point out that practical con-
siderations lead me in the fiNt place to distinguish only three types of elements (apart from the
parenchyma of the medullary rays), the distinctions being based on purely morphological character~.
since this seems most suitable for the purposes of systematic-anatomical investigatior.s; the three
kinds of elements are: (4) the wood-vessels (cell-fusions, serving for the conduction of water); (6) the
wood-parenchyma (elements, which conduct and store up carbohydrates); and (c) the wood-
prosenchyma or the wood-fibres (elements, which always have a more or less pronounced mechanical
function, but at the same time occasionally also participate in the functions of the vessels or of
thewood-parenchyma~_ In this terminology the following elementary organs oeSanio's nomenclature
are comprised under the term' wood-prosenchyma': (a) the simple- or bordered-pitted libri{orm
and the septate libriform, which according to Sanio constitute a system in the wood, comparable
to the bast~fibres;- (0) the~ intermediate "fibres, which are shorter than the libriform fibres. store
lip starch, and function as a substitute for the wood-parenchyma; for this renson they are included in
Sarno's parenchymatous system along with the wood-parenchyma; lastly (c) the tracheides
:included in Sanio's tracheal system together with the vessels), which, when they form the grotmd-
mass of the wood, can only be distinguished with difficulty from libriform bearing bordered pits,
while in other cases such a distinction is scarcely possible. The use of the term • tracheides' may
be restricted to such xylem-elements as bear bordered p'its and are of the nature of fibres; they occur
side by side with simple-pitted wood-prosenchyma (hbrifonn) :md show distinct transitions to the
,,·estels in the detailed structure of their walls and in the occasional presence of perforations at
the ends of the cel1s. The exact nature of the wood-prosenchyma within the prescribed limits must
naturally be taken into consideration in a detailed description. See also Caspary, in Pringsheim
Jahrb., iv, J86.., p. 121. and Schriften d. physikal.-okonom. Gese1lsch. Konigsberg, nviii. 1887,
p. 27 et seq.; Moller, loco cit., p. 301; and Petersen, Trakeide, Oversigt Danske Videnslc. Selsk..
Forb., 190J, n. J, pp. 95-108 (with a French re,,~me).
 CONCLUDING RE!l-IARKS

of the wood; the abundant or scanty development of wood·parenchyma;

the breadth and structure of the medullary rays. Those structural features.
which vary with the amount of moisture in the soil and air, and the degree
of transpiration on the part of the plant (e.g. chiefly,the relative abundance
elf the vessels and the width of their lumina), are for the, most part of minor
systematic value (d. the Introduction). This statement applies also to the
presence or absence of those xylem-elements, which we have distinguished as
tracheides, intermediate fibres and septate fibrous cells (cf. foot·note 3 on
p. II3S). and which are developed side by side with the wood-prosenchyma (the
latter forming the ground-mass of the wood); the relatively small systematic,
value of these elements corresponds with their function as forms physiologically
intermediate between the wood·prosenchyma and the vessels. or between the
wood-prosenchyma and the wood·parenchyma, as the case may be.
In practice, the determination of a piece of wood is based, primarily, on the
coarser features, and especially on those which are recognizable in a transverse section
with the help of a lens (e.g. the breadth of the medullary rays j the size of the lumina,
the mode of arrangement and the relative abundance bf the vessels; and the forma-
tion of annual rings, which depends on the features last named, and is found also in
the wood of tropical plan ts). as well as on certain physical properties (such as hardness,
colour, and occurrence of heart- and splint-wood) 1. These features can only be
employed to a very limited extent for systematic-anatomical pllrposes, since some of
them can be detected only in pieces of wood, which have attained a certain degree
of thickness; at the present time. moreover, a systematic investigation of the wood
of foreign plants can only be undertaken on branches of, herbarium-mat~rial, Q'wing
to the difficulty in obtaining correctly determined material of exotic stems. For
the same reason we will only briefly refer to the numerical methods, employed
by various authors for the purpose of specific diagnosis; these methods consist in
the determination of the number of vessels found in a certain surface-area (one
square millimetre) of the transverse section, or of the number of medullary rays
(which are approximately parallel to one another in older stems) situated along a
line, which is a definite number of millimetres in length, and is perpendicular to the
direction of the rays; in certain species these numbers are constants.

§ 43. WOOD~VESSELS. ~he absence of xylem-vessels in woody Dicotyle-

dons is a very rare phenom'enon, which has been recorded only in the genera
Drimys and ZygogynwI'n (Magnoliaceae) and the genera Tetracentron and
Trochodendron (Trochodendraceae); it is possible that the absence of vessels
in these genera may, be explained on phylogenetic grounds, the structure ot
their wood being altogether like that of the Coniferae. The absence of vessels
is of more frequent occurrence in certain succulents, water-plants. and parasites,
in which it is to be regarded as an indication of reduction. This is the case
in the following forms: the Nymphaeaceae; Aldrovanda (Droseraceae); the
Cacteae, in which the secondary wood is provided with peculiar annular and
spiral tracheides; certain of the Monotropeae; the Cuscuteae; the Podostema-
ceae and Ceratophylleae 2; and.lastly, the thorns of Balanites (Simarubaceae),
when they are destitute of leaves.
The anatomical features, which- require to be taken into special con-
sid~ration in examining the vessels of the secondary wood (briefly styled
• vessels' in the diagnoses of the Orders) with a view to their employment
for systematic purposes, are as follows. In the first place it is necessary to
notice the mode of arrangement and the width of the vessels: The latter
are either isolated or lie in radial or tangential groups comprising 1\vo or more
----------------
t See R. Hartig, Unterscheidungsmerkmale der wichtigeren, in Deutschlaod wachsenden Holzer.
Munchen (1St edit., 1878, &c.); Nordling-er. Querschnitte i-xi, 1852-88 (including also transverse
sections of the wood of exotic plants): and other authors. -
, See Caspary, in Monatsber. Berliner Akad., 1862, p. 448 et seq.
 NORMAL STRUCTURE OF THE AXIS 1137

vessels. The width of the lumina of the vessels varies greatly, being very
considerable, especially in lianes, but small in species having an ericoid habit.
and altogether in plants inhabiting dry localities. For taxonomic purposes
it is important to know the I medium diameter' and the' maximum diameter;'
The'abundance of the vessels is likewise a very variable feature. The diverse
appearance' of the annual rings 1, especially in the woody plants of our zones,
is related to the structural differences just mentioned. All these features are
epharmonic, and therefore in most cases merely constitute specific characters.
and it is only when all the species of a certain group live under identical con-
ditions that they are characteristic of more extensive taxonomic groups,-in some
cases even of Orders. The systematic employment of these features is rendered
difficult by the fact that they are occasionally developed in a characteristic
manner only in the later growth of the wood. so that it is impossible to obtain
an adequate knowledge of them from an investigation of the branches of
herbarium-material. For the purpose of distinguishing between the wood
from the trunk of different trees they have been in practical use for a long
time (d. § 42).
Other characters, to be noticed in the cOllfse of an examination of the
transverse section of a branch, are the round or angular outline of the vessels
and the thickness of their walls. The plugging of the vessels of the heart-
wood with resinous or gummy substances is a widely distributed phenomenon.
but is occasionally found also in the wood of perfectly young branches, especially
in such plants as have secretory organs in the ground tissue; regarding plugs
of silica or carbonate of lime, see §§ 26 and 27 (p. IIIl). Details are at
pr~sent wanting as to the systematic value of tyloses 2, but the latter appear
to occur only in certain species.
A further character to be kept in view IS the type of perforation (' Gefass-
durchbrechung'r\ found on the division-walls between the segments of the
vessels; these perforations are either simple or scalariform. Simple perfora-
tions are found both on horizontal and on more or less strongly inclined division-
walls, their outline being circular on the horizontal walls, and varying from
circular to elliptical (occasionally even elongated-elliptical) on the/inclined
walls. Scalariform perforations are in most cases met with only on strongly
inclined division-walls; the bars, which vary in number (few or very many
bars) and thickness, are generally placed transversely to the longitudinal axis
of the vessel, while the outline of the perforation is for the most part elliptical
(the ellipse occasionally being drawn out to a very considerable extent). Special
forms of scalariform perforations (which, we may notice at once, are not of any
great systematic interest and generally occur in the same piece of wood side
by side with one of the two chief types of perforations) are as follows: (a) per-
forations. in which the two halves of the bars (i.e. those parts of the bars, which
belong to the adjoining end-walls of two contiguous segments of the vessel)
are shifted parallel to one another or intersect one another (in certain Com-
positae); (b) 'perforations. in which the bars are arranged obliquely or parallel
to the longitudinal axis of the perforation; (c) malformed scalariform perfora-
tions, in which only a small number of bars or only imperfectly differentiated
bars are present; (d) perforations, in which the barsform an elegant network
as the result of anastomosis (in certain Compositae. Rosaceae, Nyctagineae) :
(e) the peculiar perforations of certain species of Epacris (Fig. 112, p. 493)
1 Urspnmg. Anat. u. Jahresringbild. trop. IIoizarten, Di~s., llasd, l(jOO, 82 pp.; Moll and
J:1nssonius, Mikrographie d. Holle~, Heft I, Leidell, 1906.
~ cr. De Bary, Vergl. Anat., 187i, p. 179.
, See Tangl, Perforationen an Ptl:1nzengef., Sitz.· Ber. \\"ient'~ Ak3.d •• lxiii, Abt. I, 187 I, 12 pp.
and Tab., and lxvii, Abt. I, J8n, t4 pp. and Tab.; Solcredcr, Ubcr den $}st. u. phylogenet. Wert
def Gefassdurchbrech., Bot. Centralbl., 1888, i, pp. ~1:::-19.
SOLE.Rt!:DF.K 4- 0
 CONCLUDING REMARKS

(I) perforations, in which the division-walls of the vessels are traversed by

rather a large number of circular or elliptical holes of varying size (' foraminate
perforations ') (in Myrothamnus and certain Dilleniaceae, Ericaceae and Vac·
ciniaceae, Fig. log, p. 479); (g) lastly, the combinations of simple and scalari·
form perforations, which have been observed in certa.in Begonias (Fig. 89,
p. 405) and Quillaia (Rosaceae). As a rule the mode of origin of the perforated
areas on the division-walls (either from simple pits or from bordered pits of
Jarge size) is of no great systematic value, since their development often varies
in one and the same longitudinal section 1.
In a given species the perforations of the vessels are either exclusively
simple or exclusively scala.riform, or both simple and scalariform. In the
latter case the two kinds ()f perforations either occur side by side with one
another in the vessels of the secondary wood (occasionally we even find a
simple perforation at one end, and a scalariform perforation at the other end
of the same segment), or the scalariform perforations are confined to the primary
wood and its immediate neighbourhood. When simple perforations are found
in the same plant side by side with those of the scalariform type, the outline
of the former is more commonly elliptical than circular, and in some cases
even elongated-eHi ptical.
The presence or absence of scalariform perforations has mostly proved
to be of value for the diagnosis of extensive taxonomic groups (Orders, tribes,
genera). Certain groups have only simple or only scalariform perforations in
the vessels, while others show a tendency to form scalariform perforations;
this tendency betrays itself in the fact that different genera or different species
of the same genus may in one case have exclusively scalariform. perforations,
and in another case both simple and scal;,triform perforations, although the
perforations are never eXClusively simple (scalariform perforations occurring
at least in or near the primary wood, even if they are wanting in the secondary
wood). Simple perforations are more widely distributed than those of the
scalariform type. The constant occurrence of scalariform perforations in all
the members of an Order i!; a rare phenomenon, found only in a very limited
number of Orders, which aI"e all of small size.
It remains to mention the striking fact that the occurrence of scalarifonn
perforations in the vessels often goes hand in hand with small lumina in the
vessels and the presence of bordered pits on the wood-prosenchyma.
Exclusively simple perforations are found in the saIbe species il\: Ranuntu-
1aceae pro parte, Calycanthaceae. -Lactoridaceae. Anonaceae, Menispermaceae,
Berberideae pro parte. Papav~raceae, Cruciferae, Capparidea.e, Resedaceae, Cistineae,
Violarieae pro parte, Bixineae pro parte, Pittosporeae, l'remandreae pro parte,
Polygaleae,Vocliysiaceae, Frankeniaceae, Caryophylleae, Portulaceae, Tamariscineae
pro parte, Elatineae, Hypericineae. Guttiferae, Ternstroemiaceae pro parte, Diptero-
carpeae, Ancistrocladus, Chlaenaceae, Malvaceae, Triplochitonaceae, Sterculiaceae
pro parte, Tiliaceae pro par-teo Lineae pro parte, Malpighiaceae, Zygophylleae,
Geraniaceae. Rutaceae pro p'arte, Simarubaceae pro :parte, Koeberlinta, Balanites,
Burseraceae, Meliaceae, Chrulletiaceae pro parte, Olacmeae pro parte, Celastrineae
pro parte, Hippocrateaceae pro parte, Corynocarpaceae, Stackhousieae, Rhamneae
pro parte, Ampclidaceae pro parte, Sapindaceae (almost always), Didiereae, Hippo-
castanaceae pro parte, Aceraceae, Melianthaceae, Anacardiaceae pro parte, Corianeae,
Moringeae. Connaraceae. Papi!ionaceae, Caesalpinieae, Mitnoseae, Chrysobalaneae,
Crassulac eae, Droseraceae pro parte, Halorageae, Combretaceae, Myrtaceae sens.
str. pro parte, Lecythidaceae, l\Ielastomaceae, Lythrarieae, Onagrarieae pro parte,
Samydaceae pro parte, Loas(!ae, Passifloraceae pro parte, Papayaceae, Cucurbita-
ceae, Datisceae, Cacteae, Ficoideae, Umbelliferae pro parte, Araliaceae pro parte,
Carlemannia (Caprifoliaccae), Rubiaceae pro parte, Calycereae, Composi-tae pro
parte, Candolleaceae, Goodcniaceae. Lobcliaceae, Ericaceae pro parte ?, Epacrideae

I See SolereUer, Hollstruktur, 1885, p. IS,

 NORJIAL STRUCTURE OF THE AX1S I139
pro parte?, Lennoaceae, Plumbagineae, Primulaceae. Myrsineae pro parte, Sapota-
ceae pro parte, Ebena.ceae, Styraceae prQ parte, Oleaceae pro parte, Salvadoraceae,
AIX>Cynaceae, Asclepiadeae, Loganiaceae pro parte, Gentianeae pro parte, Pole-
ntoniaceae pro parte, Hydrophyllaceae, Boragineae, Convolvulaceae, Solanaceae,
Retzia, Scrophularineae, Orobanchaceae, Gesneraceae, Bignoniaceae, Pedalineae,
Acanthac~ae, Myoporineae, Sel~neae, Verbenaceae pro parte, Labiatae pro parte,-
PI antagineae, Nyctagineae (diVIsion-walls very rarely perforated in a reticilla.te
manner), Illecebraceae, Amarantaceae, Chenopodiaceae pro parte (almost-always),
Basellaceae, 'Phytolaccaceae, Batideae, Polygonaceae, Nepenthaceae, Cytinacea.e,
Aristolochiaceae, Piperaceae pro parte, Lallrineae pro parte, Hernandiaceae pro
parte, Proteaceae, Thymelaeaceae. Gonystylus, Penaeaceae, Elaeagnaceae, Lorantha-
ceae, Santalac~ae, Myzodendl'on, Champel'eia, Balanophoreae?, Euphorbiaceae pro
parte, Buxaceae pro parte, Ulmaceae pro parte, Cannabineae, Moraceae, Urticeae,
Leitnerieae, Juglandeae pro parte, Sahcincae.
Exclusively scalariform perforations are found in the same species in: Ranun·
culaceae pro parte, Dilleniaceae pro parte, Magnoliaceae pro parte, Trochoden-
draceae pro parte, Violarieae pro parte, Canellaceae, Bixineae pro parte, Ternstroe-
miaceaepro parte, Rhaptopetalaceae, Humiriaceae, Olacineae pro parte, Scytopetatutn.
llicineae. Cyrilleae. Celastrineae pro parte, Staphyleaceae pro parte, Saxifragaceae
pro parte, Penthol'um. Droseraceae pro parte. Hamamelideae, Myrothamnus, Ostreal'ia,
Bruniaceae; Rhizophoraceae pro parte {RhizQphoreae}, Myrtaceae pro parte, Samy-
daceae pro parte, Passilloraceae pro parte, Cornaceae :pro parte, Caprifoliaceae pro
parte, Dipsaceae pro parte?, Vacciniaceae pro par~e, Encaceae pro parte, Epacrideae
pro parte, Myrsineae pro parte, Bl'achynema, Styraceae pro parte, Desfontainea.
Columelliaceae, Piperaceae pro parte, CWoranthaceae, Monimiaceae pro parte,
Laurineae pro parte, Go mortegaceae , Geissoloma, Grubbia, Euphorbiaceae pro
parte, Dapbniphyllaceae, Buxaceae pro parte, Balanopseae, Myricaceae pro parte.
Cupuliferae pro parte, Lacistemaceae, Empetraceae pro parte.
Simple and scalariform perforations are found side by side in the same species
in: Dilleniaceae pro parte, Magnoliaceae pro parte, Trochodendraceae pro parte, Ber-
berideae pro parte, Violarieae pro parte, Bixineae pro parte, Tremandreae pro parte,
Tamariscineae pro parte, Ternstroemiaceae pro parte, Sterculiaceae pro parte,
Tiliaceae pro parte, Lineae pro parte, Rutaceae pro parte, Ochnaceae, Simarubaceae
pro parte, Chailletiaceae pro parte, Olacineae pro parte, Celastrineae pro parte,.
Hippocrateaceae pro parte, 'Rhamneae pro parte, Ampelidaceae pro parte, Sapin-
daceae pro parte, Hippocastanaceae pro parte, Staphyleaceae pro parte, Sabiaceae.
Anacardiaceae pro parte, Rosaceae (exc1. Chrysobalaneae), Crossosomataceae,
Saxifragaceae pro parte, Rhizophoraceae pro parte (Legnotideae), Onagrarieae
pro parte, Samydaceae pro parte, Turneraceae, Passifloraceae pro parte, Begonia-
ceae, U mbelliferae pro :parte, Araliaceae pro -' parte, Cornaceae prQ parte, Capri-
foliaceae pro parte, Rublaceae pro parte, Valerianeae, Dipsaceae pro parte, Compo-
sitae pio parte, Campanulaceae, Vacciniaceae pro parte, Ericaceae pro parte,
Epacndeae pro parte, Diapensiaceae, Myrsineae {'ro parte, Sapotaceae pro parte.
Oleaceae pro parte, Loganiaceae pro parte, Gentianeae pro parte. Polemomaceae
pro parte, Verbenaceae pro :parte, Labiatae pro parte, Chenopodiaceae pro parte.
J?iperaceae pro parte, Myristlcaceae, Monimiaceae pro parte, Laurineae pro parte,
Hemandiaceae pro parte, Euphorbiaceae pro parte, Ulmaceae pro parte, Platana-
cea.e, Juglandeae pro parte, l\Iyricaceae pro parte, Casuarineae, Cupuliferae pro
parte, Empetraceae pro parte.
As regaros the structure of the walls of the vessels the following features
are chiefly to be taken into consideration: the occurrence of spiral thickening
on the walls of the pitted vessels of the secondary wood; the size and nature
(outline of the border and of the aperture of the pit) of the bordered pits (occa..
sionally scalariform bordered pits), found on the walls of conta.ct between
two vessels; the occurrence of a 'sieve-like' structure (first demonstrated
by Bengt Jonsson 1) on the pit ..membranes of the bordered pits of the vessels ;
lastly, the varying structure of the walls of the vessels in contact with paren ..
chyma of the medullary rays or with wood-parenchyma. These walls are

1 Ber. deutsch. bot. Gesellsch., 189:11, p. 49.f, and Tab. n :ii; ;ee also Heiden, in l!ot. Ccntrs.lbJ.,
18 93, iv, p. 3.
4 D :2
 CONCLUDING REIJfARKS

occupied (a) either by one-sided bordered pits, one or more of which correspond,
to large one-sided simple pits on the adjoining parenchyma~wall, or (b) by
one-sided simple pits of varying size, which correspond to pits of equal dimen-
sions on the wall of the adjoining parenchyma-cell; in the latter case transi··
tional forms to distinct one-sided bordered pits frequently occur side by side
with the simple ones. The systematic value of these features varies and·
requires to be tested in each individual case; the most valuable characters
are afforded by the structure of the walls of the vessels in contact with paren-
chyma, which is occasionally constant throughout all the members of an Order
(Burseraceae, Hamamelideae).
Spiral thicken in g of the walls of the vessels is generally only a specific character ~
and has been observed in the following Orders: Ranunculaceae, Calycanthaceae,
Magnoliaceae, Trochodendraceae. Berberideae, Papaveraceae, Cruciferae, Cappari-
deae, Bixineae, Pittosporeae (universally?), Tremandreae, Polygaleae, Portulaceae,
Ternstroemiaceae. Malvaceae. Sterculiaceae, Tiliaceae, Lineae, Malpighiaceae.
Geraniaceae, Rutaceae, Meliaceae. Olacineae, Ilicineae, Celastrineae. Rhamneae.
Sapindaceae, Hippocastanaceae,' Aceraceae, Melianthaceae, Staphyleaceae, Ana-
cardiaceae, Papilionaceae, Caesalpinieae, Rosaceae, Droseraceae. Hamamelideae
(only on the ends of the segments of the vessels), Samydaceae, Ficoideae, Aralia-
ceae, Cornaceae. Caprifoliaceae, Rubiaceae, Compositae, Vacciniaceae, Ericaceae,
Epacrideae, Myrsineae, Oleaceae. Apocynaceae, Loganiaceae, Boragineae, Solana-
ceae. Scrophularineae, Bignoniaceae, hI yoporineae, Se lagineae. Verbenaceae,
Labiatae, Illecebraceae. Chenopodiaceae, Phytolaccaceae, Polygonaceae, Aristolo-
chiaceae, Monimiaceae, Laurineae, Proteaceae, Thymelaeaceae, Loranthaceae,
Santalaceae, Buxaceae, Ulmaceae, Moraceae, Juglandeae. Casuarineae, Cupuliferae.
Species, in which the walls of the vessels show spiral thickening, generally have
spirally thickened tracheides as well. For the occurrence of special spirally
thickened vessels or cells in the wood, see § 47 (p. 1145).
Scalariform bordered pits have been observed on the vessels of the secondary
wood in certain Magnoliaceae. Violarieae, llicineae, Ampelidaceae, Bruniaceae,
Rhizophoraceae, Begoniaceae, Apocynaceae. Piperaceae, Monimiaceae, Euphor-
biaceae, Cupuliferae and in JI,[yzodendron. The simple pitting of the walls of the
vessels in the Papayaceae and Crassulaceae is due to their being embedded in
unlignified tissue (see § 47, p_ 1144).
Sieve·like structure of the pit-membranes in the bordered pits of the vessels
has been recorded in the following Orders: Vochysiaceae, Guttiferae, Rhamneae, I
Hipp<?castanaceae, Papilionaceae, Caesalpinieae, l\hmoseae, Amygdaleae, Combreta-
ceae, Myrtaceae, Melastomaceae, Onagrarieae,Araliaceae. Compositae. Asc1epiadeae,
Oleaceae, Scrophularineae, Cupuliferae.
The occurrence of simple and occasionally large pits on the walls of the
vessels in contact with parenchyma (sometimes side by side with bordered pits)
has been recorded in the following Orders and genera respectively: Calycanthaceae,
Magnoliaceae, Trochodendraceae pro parte, Vioiarieae, Bixineae pro parte, Tre-
mandreae, Vochysiaceae pro parte, Tamariscineae pro parte, Hypericineae pro,
parte, Guttiferae pro parte, Ternstroemiaceap, pro parte, Dipterocarpeae, Chlaena-
ceae, Malvaceae pro parte, Sterculiaceae pro parte, Tiliaceae pro parte, Lineae
pro parte, Humiriaceae pro parte, Malpighlaceae pro parte, Geraniaceae, Rutaceae
pro parte, Simarubaceae pro parte, Burseraceae, Olacineae pro parte, Rhamneae
pro parte, Ampelidaceae, Hippocastanaceae, Staphyleaceae pro parte, Sabiaceae,
Anacardiaceae, Moringeae, Connaraceae. Papilionaceae pro parte (rare), Rosaceae
pro parte, Saxifragaceae pro parte, Hamamelideae. Ostrearia, Rhizophoraceae
pro parte, Myrtaceae pro parte, Melastomaceae pro parte, Lythrarieae pro parte,
Onagrarieae. Passifloraceae pro parte, Datisceae. Umbelliferae pro parte, Araliaceae,
Cornaceae pro parte, Vacciniaceae pro parte, Myrsineae pro parte, Sa.potaceae,
BrQchynema, Styraceae pro parte. Apocynaceae pro parte, Loganiaceae pro parte,
Convolvulaceae, Solanaceae pro parte, Gesneraceae pro parte, Bignoniaceae pro
parte, Acanthaceae pro parte, Verbenaceae pro parte, Labiatae pro parte, Phytolac-
caceae pro parte, Piperaceae, Myristicaceae, Monimiaceae, Laurineae, Hernandia-
ceae, Gomortegaceae, Geissoloma, Santalaceae pro parte, Grubbia. Euphorbiaceae
pro parte, Balanopseae, Moraceae pro partc'. Urticeae, Leitnerieae, Juglandeae,
Cupuliferae pro parte, Salicineae.
 NORMAL STRUCTURE OF THE AXIS
:Bordered pits alone have been observed on the wan. of contact between veud.
and parenchyma in the followin~ Orders and genera respect!vely: Trochodend,ra.
,~ pro p.¥te, Anonaceae, Memspermaceae, Capparideae, CaneUaceae, Bixineae
Vro parte, Pittosporeae, Polygaleae, Vochysiaceae pro parte. Tamariscinea.e pro
parte. Elatineae, Hypcricineae pro parte. Guttiferae pro parte, Ternst:rOemiaceae
pro parte, Ancistroclaau$, Malvaceae pro parte, Triplochitonacea.e, StercUliaceae pro
parte; Tiliaceae pro parte, Lineae pro parte, H umiriaceae pro parte. Malpigbiacea.e
pro parte, ZygopnyUeae, Rutaceae pro parte (in most -cases), Simarubaceae pro parte
(almost always), KQcberlinia, BaJaniu:s, Ochnaceae, Meliaceae, Olacineae pro~,
llicineae, Cyrilleae ?, Celastrineae, Hippocrateaceae, Rhamneae pro parte, Sapinda-
ceae, Aceraceae, Melianthaceae, Staphyleaceae pro parte, Papilionaceae pro pa.r!e (in
most cases), Caesalpinieae, Mimoseae, Rosaceae pro p;ute, Crossosomataceae, Saxi·
fr~aceae pro parte. Pentho,um, Droseraceae, M yt'othamnus, Brunia.ceae, Halomgeae,
Rbizo~horaceae pro parte, Combretaceae, Myrtacea.e pro parte, M~maceae pro
parte, Lythrarieae pro parte, Samydaceae, Loa.seae, Turneraceae, Pa.ssi1loraceae pro
parte, Umbelliferae pro parte, Comaceae pro parte, Rubiaceae, Calycereae, COm-
positae, Goodeniaceae, ca.mpanulaceae, Lobeliaceae, Vacciniaceae pro parte,.Erica-
ceae, Epacrideae, Myrsineae prQ parte, Ebenacea.e, Styra.cea.e pro parte, Oleaceoe,
Salvadoraceae, Apocynaceae pro parte, Asclepiadeae, Loganiaceae pro parte, Dn-
/ontainea, Gentianeae, Hydrophyllace ae, Solanaceae pro parte, Retna, Scr~:u1a­
lineae, Columelliaceae, Gesneraceae pro parte, Bignoniaceae pro parte, Ped . eae,
Acanthaceae pro parle, Myoporineae, Selagineae, Verbenaceae _pro ~, Labiatae
pro parte, Phytolaccaceae pro parte, Batidea.e, Polygonacea.e, Chlora.ntha.<:eae, Pr0-
teaceae. Thymelaeaceae, Penaeaceae, E1aeagnaceae, Santalaceae pro ~t C.kHJ.
pereia, Euphorbiaceae pro parte, Daphniphyllaceae, Buxa.ceae, Ulmaceae, Mora.ceae
pro parte. Myricaceae, Casuarineae, Cupuliferae pro parte, Lacistemaceae?

§ 44. WOOD"PROSENCHntA. The most valuable systematic feature,

afforded by the structure of the wood·prosenchyma, has proved to be the
nature of the pitting of the, prosenchymatous ground~ma8S 1 of the wood.
The presence of distinct simple pits (i.e. without a border) or of distinct
bordered pits (alth9ugb the borders are sometimes small) on the Wood·
prosenchyrna is often characteristic of entire Orders, tribes, genera or generic
sections; in utilizing these features it is necessary to neglect the occurrence
of small numbers of fibrous cells, provided with bordered pits (i.e. 4 tracheides'
which are transitional to the vessels), side by side with the wood~prosenchyma
-bearing simple· pits, or of simple-pitted fibrous cells side by side with the
bordered-pitted wood-prosenchyma. A more detailed classification of the wood-
prosenchyma (ci. § 42, p. 1135, footnote 3) is not of much value in systematic-
anatomical researches, as far as can be gathered from the experience hitherto
obtained, among other reasons because a classification of this kind is partly based
on phy'siological characters (viz. the cell-contents), so that it is impossible to
apply it with the requisite precision, when using herbarium-material. Apart
from -those already mentioned, a few other structural features of the wOOd·
prosenchyma deserve to be taken into consideration; the most important
of these _are th~ followiI}g: the. varying size of the lumina and the divet'Se
thickness of the walls';"" the occurrence of thin transverse walls in the wood-
prosenchymir (septate wood.prosenchyma), a feature connected with the
storage of starch in the lumina of the cells; the presence of starch in fibrous
cells, which are not septate; the occurrence of a so-called gelatinous layer,
viz. a- cartilaginous thickening-layer of the cell-wall, consisting of hemicellulose
and occasionally disappearing in the course of the metabolism of the plant;
the detailed structure and abundance of the pits; the length of the cells and
their mode of arrangement in a transverse section; the appearance of the
cells in transverse section (outline and siz;e~ the latter occasionally varying

<, See also Gregory, Pores of the Ubrif. tissue, BulL Torrey Bot. Club, New York J J 887; abltt.
1ft Bot. Centrall>l., \S87. iii, p. '2.
 CONCLUDI~G REMARKS

in one and the same section, owing to the fact that the fibrous cells are cut
through at different points in the course of their length), &c.
Although the distinction between wood-prosenchyma bearing bordered'
pits and that bearing simple pits is so simple in itself, the maintenance of this.
distinction, which is very important for systematic purposes, becomes very
difficult, when the borders of the pits are very small; for, simple slit-shaped
pits occasionally appear to be provided with a small border, when seen in
surface-view. In such a case the appearance of the pit in transverse section
can alone decide whether the pit is really bordered or not.
The prosenchymatous ground·mass of the wood bears silllple pits in the {ol.
lowing Orders and genera respectively: Ranunculaceae, Anonaceae, Berberideae
pro parte. Papaveraceae, Cruciferae. Capparideae. Resedaceae. Cistineae pro parte.
Violarieae. Bixineae pro parte. Pittosporeae. Tremandreae pro parte, Vochyslaceae
(excI. Trigoniaceae), Frcinkeniaceae. Caryophylleae. Portulaceae. Tamariscineae
pro parte, Hypericineae pro parte, Guttiferae pro parte, Ternstroemiaceae pro parte,
Dipterocarpeae pro parte. Malvaceae. Triplochiton~ceae, Sterculiaceae pro parte ..
Tiliaceae, Rhaptopetalaceae, Lineae pro parte, Malpighiaceae, Geraniaceae, Rutaceae,
Simarubaceae pro parte {almost always). Ochnaceae pro parte, Burseraceae, Meliaceae.
Corynocarpaceae.1lliamneae, Ampelidaceae, Sapindaceae, Didiereae, Hippocastana-
ceae. Aceraceae. Melianthaceae, Staphyleaceae pro parte, Sabiaceae pro parte, Anacar-
diaceae. Coriarieae, Moringeae, Connaraceae, Papilionaceae, Caesalpinieae, Mimoseae~
Rosaceae pro parte (rare). Saxifragaceae pro parte. Crassulaceae. Halorageae,
Rhizophoraceae pro parte (tribe Rhizophoreae). COInbretaceae pro ~ (in most
cases), Lecythidaceae, Melastomaceae pro parte, Lythrarieae, Onagrarieae. Samy..
daceae. Loaseae ;pro parte, Turneraceae pro parte, Passifloraceae pro parte, Cucur-
bitaceae, Begomaceae, Datisceae, Cacteae, Ficoideae, Umbelliferae, Araliaceae.
Cornaceae pro parte, Caprifoliaceae pto part~, Rubiaceae pro parte, Valerianeae,.
Dipsaceae pro parte. Calycereae, Compositae, Lobeliaceae, Diapensiaceae pro parte.
Plumbagineae. Myrsineae. Sapotaceae, Oleaceae pro parte, Salvadoraceae. Apocyna-
ceae pro parte, Asclepiadeae pro parte, Loganiaceae pro parte, Gentianeae pro parte.
Polemoniaceae. Boragineae pro parte. Solanaceae pro parte,Retlia, Scrophulanneae,
Orobanchaceae. Gesneraceae, Bignoniaceae, Pedalineae, Acantbaceae, M yoporineae.
Zombt"ana, Selagineae pro parte, Verbenaceae, Labiatae, Plantagineae, Nyctagineae,
IUecebraceae pro parte (rare), Amarantaceae, Chenopodiaceae, Phytolaccaceae pro
parte, Polygonaceae. Piperaceae, Chloranthaceae pro parte, Myrlsticaceae, Monimia-
ceae pro parte, Laurineae, Hernandiaceae, Thyrnelaeaceae pro parte (rare), Lorantha~
ceae pro parte. Myzodendron (short prosenchyma resembling rod-cells), Euphorbia-
ceae pro parte, Balanopseae pro parte. Ulmaceae pro parte, Cannabineae, Moraceae.
Urticeae, Leitnerieae. Juglandeae pro parte. Cupuliferae pro parte, Salicinea.e.
In the ensuing list of Orders and genera, the prosenchymatous ground-mass
of the wood is provided with distinct bordered pits: Dillenia'ceae, Calrcanthaceae,
Magnoliaceae, Trochodendraceae, Lactoridaceae, Eupomatia, Meruspermaceae,.
Berberideae pro parte, Cistin.eae pro parte, Canellaceae. Bixineae pro parte, Tre~
mandreae pro parte, Polygaleae. Trigoniaceae, Tamariscincae pro parte, Elatineae,.
Hypericineae pro pa.rte, Guttiferae pro parie;Temstroemiaceae pro parte, Diptero-
carpeae pro parte, Ancistrociadus, Chlaenaceae, Sterculiaceae pro parte, Lineae
pro parte, Humiriaceae. ZygophyUeae, Simarubaceae pro ~rte, Koebet'linia,
Ochnaceae pro parte, Chailletiaceae, Olacineae, Ilicineae, Cyrilleae, Celastrineae
(occasionally side by side with prosenchyma bearing simple pits), Hippocratea-
ceae, Stackhousieae (side by side with. prosencbyma bearing simple pits), Staphy-
leaceae pro parte, Sabiaceae pro parte, Rosaceae pro parte (in most cases), Crossoso·
mataceae. Sa,nfragaceae pro parte, Penthorum, Droseraceae, Hamamelideae.
Myrothamnus, Ost,.eayia, Bruniaceae, Rhizophoraceae pro parte (Legnotideae).
Combretaceae pro parte, Myrtaceae sens. str., Melastomaceae pro parte, Loaseae
pro parte, Turneraceae pro parte, Passifloraceae pro parte, Comaceae pro parte.
Caprifoliaceae pro parte, Rubiaceae pro parte. Dipsaceae pro parte, Goodeniaceae.
Campanulaceae. Vacciniaceae (side by side with prosenchyma bearing simple pits),.
Ericaceae (in part side by side with prosenchyma bearing simple pits), Epacrideae.
Diapensiaceae pro parte, Ebenaceae. Brachynema. Styraceae. Oleaceae pro parte.
Apocynaceae pro parte, Asclepiadeae pro parte, Loganiaceae pro parte, Desfon-
taitre(J~ Gentianeae pro parte, Hydtophyllaceae. Boragineae pro parte, Convolvula.ceae~
 NORMAL STRUCTURE OF THE AXIS
Solanaceae pro parte, Columelliaceae, Selagineae pro parte, Illecebraceae (in most
cases), Phytola.ccaceae pro parte, Batideae. Nepenthaceae, Aristolochiaceae,Chloran-
thaceae _pro parte, Monimiaceae pro parte, Gomortegaceae, Proteaceae, Thymelaea.
ceae (in most cases), Gonystylus, Penaeaceae, Geissoloma, Elaeagnaceae, Lorantha-
ceae pro parte, Santalaceae, Champeret"a l Grubbia, Euphorbiaceae pro parte, Daphni-
phyllaceae, Buxaceae. Balanopseae pro parte, Ulmaceae pro parte, Platanaceae,
Juglandeae pro parte, Myricaceae, Casuarineae, Cupulifetae pro parte (in most
cases), Lacistemaceae, Empetraceae. _
Septatewood·prosenchyma has been observed in certain Berberideae, Violarieae,
Bixineae, Pittosporeae, Vochysiaceae, Ternstroemiaceae, Dipterocarpeae, Malvaceae,
Tiliaceae, Lineae, Malpighiaceae, Geraniaceae. Simarubaceae, Burseraceae, Melia-
ceae, Celastrineae, Ampelidaceae, Sapindaceae, Staphyleaceae. Sabiaceae, Anacar-
diaceae, Connaraceae, Papilionaceae, Caesalpinieae. Mimoseae, Rosaceae, Saxi-
fragaceae, Combretaceae. Melastomaceae, Lythrarieae, Onagrarieae, Samydaceae.
Tumeraceae, Passifioraceae, Begoniaceae, Cacteae, Araliaceae, Cornaceae, Capri~
foliaceae, Rubiaceae, Compositae, Lobeliaceae, Vacciniaceae, Myrsineae, Oleaceae.
Loganiaceae, Polemoniaceae, Solanaceae, Scrophularineae, Gesneraceae, Bignonia-
ceae, Acanthaceae, Z ombiana, Verbenaceae, Lablatae, Phytolaccaceae, Polygc;maceae.
Piperaceac, Chloranthaceae, Myristicaceae, Monimiaceae. Laurineae, Euphorbia-
ceae, Moraceae, Cupuliferae, Salicineae.
A gelatinous layer has been recorded in certain Calycanthaceae, Violarieae,
Bixineae, Lineae, Simarubaceae, Hippocastanaceae, Papilionaceae, Caesalpinieae.
Mimoseae, Rosaceae (All1ygdaleae), Myrtaceae, Onagrarieae, Ebenaceae, Laurineae.
Piperaceae, Proteaceae, Euphorbiaceae. Ulmaceae, Moraceae, Casuarineae, Cupuli-
ferae and Salicineae; this feature is often found only in a small number of the
wood-fibres composing a piece of wood (d. also De Bary, Vergl. Anat., p. 497; and
Potter, in Annals of Bot., 1904. p. 121 et seq.)•
.§ 45. WOOD-PARENCHYMA. The scanty or abundant occurrence -of wood-
parenchyma is of considerable systematic value, sometimes even constituting
an ordinal character; when present in large quantities, the wood-parenchyma
generally forms tangential bands (known as the' metatracheal ' parenchyma
in contrast to th~ 'paratracheal' parenchyma, aggregated round about the
vessels), as seen in ~ transverse section of the branch. For the purposes of
specific diagnosis it is also well to take note of the occurrence- of scattered
wood-parenchyma between the wood-fibres and in the autumn-wood of species
forming annual rings. The occurrence of 'chambered crystal.parenchyma'
(i.e. parenchymatous strands, divided by means of transverse walls into
chambers, which are for the most part of approximately equal size and contain
solitary crystals 1) in the wood is generally only of specific value.
Abundant development of the wood·parenchyma has been observed in the
folloWing Orders and genera respectively: Dilleniaceae, Magnoliaceae, Anonaceae.
Vochysiaceae, Hypericineae, Gutti ferae, Dipterocarpe<i1e, Ancistrocladus, Chlaena-
ceae, Malvaceae, Sterculiaceae, Tiliaceae, Rhaptopetalaceae, Lineae, Malpighiaceae,
Simarubaceae, Balanites I, Olacineae, Celastrineae,. Sapindaceae, Papilionaceae,
Caesalpinieae, Chrysobalaneae, Combretaceae. Lecythidaceae, Rubiaceae, Sapota-
ceae, Ebenaceae, Boragineae~ Verbenaceae, Aristolochiaceae. Laurineae. Proteaceae,
Thymela.eaceae, EuphQwiaceae, Moraceae, Juglandeae. Casuarineae. Cupuliferae.
§ 46. MEDULLARY RAYS OF THE WOOD 3. The most important systematic
characters are the appearance of the medullary rays in a tangential section,
and the breadth of the rays; in determining the latter feature it is necessary
to examine both transverse and tangential sections. The varied appearance
1 Clustered crystals rarely occur in the wood-parenchyma (Guui[erae: Garcinia d£oica, Bi."
while raphides have been observed in this tissue in members of the Dilleniaceae and Ternstroemiaceae.
i According to Van Tieghem, Ann. sc. nat., ~er. 9. t. iv. 1906.
IS Essner, Diagnost. Wert der Ani. u. Hohe d. Markstr. bei d. Coniferen, Halle. 188i; Schnl/,
Markstrablgewebe. Jahrb. Berliner Gatt., ii, 1883, p. 224 and Tab. vii; Michael, Bau d. Holzes def
Compositen, etc._ Diss., Leipzig, 1885; Zache, Anz. u. Gr. d. Markstr. bei einig. Laubh., Zeitschr.
f. Naturw. C. Sachsen u. Thur., 1886, p. 1; Kny, Markstr. dicotyl. Gew., Sitz.~Ber. dentsch. bot
Gesellsch., 1890, p. 176; Herbst, Mark~tr., Hot. Centralbl., 1894, i, p. 2~7 et seq., especially p. 409.
 CONCLUDING REMARKS
of the medullary rays in a tangential section is due to -the fact that the com..
ponent cells either show palisade-like elongation in the vertical direction
(upright cells) or radial elongation (horizontal cells), the rays being composed
either of goth or only of one of these two forms of cells. The degree of elonga-
tion of the cells in the vertical or radial direction varies. I t is very rare
(certain Malvaceae and Passifloraceae) to find medullary rays, in which the
cells are compressed in the radial direction, so that they appear strikingly
broad (in the tangential direction) in a transverse section of the branch, but
narrow and upright in a radial longitudinal section. Trachcidal cells provided
with bordered pits have not been observed in the medullary rays of Dicotyle-
dons, being confined to Conifers. An exceptional development in the height
of the entire complex of medullary rays often goes hand in hand with the
differentiation of broad rays, especIally among llanes, but. occasionally also ill
plants which are not lianes (e.g. Urticaceae}. For the systematic valu~. of the
number of the medullary rays, see § 42, p. I136. Tne absence of medullary rays
is no u.ncommon feature among herbaceous plants, but occurs also in certain
woody plants. In the latter case it is sometimes only rays of the parenchy-
matous t)'Pe that are wanting, the cells of the medullary rays being differen·
Hated as short fibrous cells, so that the resulting rays are not very prominent.
It remains to mention the .penetration of scler~ed tissue, belonging to the
medullary rays of the cortex, into the medullary rays of the wood in certain
Menisperrnaceae, Hippocrateaceae, Platanaceae, Casuari.,eae and Cupuliferae;
this phenomenon goes hand in hand with a sinuation of the edge of the cam·
bium, the convexities of which are directed towards the pith.
_Broad medullary rays have been observed in the wood in certain species o'f
the following Orders and genera: RanuDculaceac, DiIleniaceae, Magnoliaceae,
·Menispermaceae, *Berberideae, Cap~deae. "'Violarieae, Portulaceae~ Ternstroe·
miaceae, Malvaceae, Sterculiaceae, Tdiaceae, Simarubaceae, Koebe"linia, Balanites,
Chailletiaceae, llicineae, *Hippo<;-rateaceae, Corynocarpaceae, *Ampelidaceae, Sabia·
ceae, Coriariea.e, Papilionaceae, Caesalpinieae, Rosaceae, Saxifragace~, Rhizophora-
ceae, Passifioraceae, Papayaceae, *Cucurbitaceae, "'Begoniaceae, Aralia.ceae, Compo-
sitae, Myrsineae, Salvadoraceae, Verbenaceae, Labiatae, Phytolaccaceae, Batideae,
• Aristoiochiaceae, ·Piperaceae, Chloranthaceae, Monimiaceae, Proteaceae, Lorantha-
ceae, Ulmaceae, Moraceae, Urticeae, Platanaceae, Casuarineae, Cupullferae. Those
Orders, which are provided with a *, have bload primary and secondary rays,
traversill~ the entire length of the internodes in the form of continuous plates
and dividing up the xylem-mass into narrow segments (H. Schenck's' An·stoloch·ia-
type ').
~ The absence of parenchymatous medullary rays has been demonstra.ted in
certain woody species belonging to the following Orders and genera: Cruciferae~
Cistineae, Fr3.nkeniaceae, Caryophylleae, Portulaceae, Tamariscineae. Geraniaceae,
Stackhousieae, Crassulacea.." Flcoideae, Compositae. Epacrideae, Diapensiaceae,
Gentianeae, Boragineae, Scrophularineae, Orobanchaceae, Plantagincae (secondary
medullary rays), Nyctagineae, Illecebraceae, Amarantaceae, Chenopodiacea.e, Phyto.
laccaceae, Polygonaceae, Laurineae (Cassyeha). llfyzotlendyon (Sect. Gymnophyton).
See also Schwendener, Mech. Prinzip, p. 148 et seq.

§ 47. SPECIAL ELEMENTS FOUND IN THE NORMALLY DEVELOPED X"l'1.EM"

MASS. UnUgnified tissue composed of thin-walled cells very commonly
accompanies the primary tracheae in the protoxylem 1, and must not be con-
founded with intraxylary soft bast (see § 57). A more important systematic
feature lies in the occurrence of unlignified tissue in the sc(:ondary xylem of

, f Int:-axylary Cambiform ' according to Raimann, in Si tz.-Ber. Wiener Akad., xcvi ii, A!:>t. I, ,889;

primaftS Vasalparenchym' according to Strasburger, Leitungsb., 1891; 'parenchyme intraligncllx'

••:cording to Prunet, in .AnD. sc. nat., ser. 7, t. xiii, 1891, p. 362; c endo!tyle' or' r~gii)n endoxylaire'
according to Briquet, in Atti CO!]~O bot. intemaz., 1892.
 NORMAL STRUCTURE OF THE AXIS I I 45

woody pla.1'lts. In certain [;pecies of Vella and Brassica (Cruci ferae , Fig. ::::6,
p. 65), AcanthophyUum (Caryophylleae) and Astragalus (Papilionaceae) it is
differentiated (like the soft bast in species exhibiting the anomaly known as
successive zones of growth) in the form of concentric rings~ and contains nlL"1lCr-
OllS tracheae, having small lumina and showing spiral to reticulate thickening.
Similar concentric layers of unlignified tissue, which surround the vessels and
alternate with prosenchymatous layers devoid of vessels, are developed in,
certain Crassulaceae; in other members of this Order the unlig"'..jfi~d tissue
enveloping the vessels forms islands. which occasionally show a concentric
arrangement in the fibro'Us ground-mass of the wood (in w'bjch no vessels are
found), or composes the entire tissue of the xyiem. The1atter type of structure
is characteristic also of the soft stem'S cf the Papayaceae. The small groups
of thin-walled cells in the wood of Olinia (Lythrarieae) include crystal-sacs.
It remains to make special mention of the appearance of the transverse section
through the wood in the species of 111yzodendron, belonging to the section
Eumyzodel".dr01s (Fig. 178, p. 735), the wood in these species shov.>ing a peculiar
di~tribution of the thin·walled tissue, the tracheae, and the wood-prosenchyma.
Apart from the cases already mentioned, unlignified tissue has been observed
in the xylem of certain woody species (which exhibit normal structure), belong·
ing to the followmg Orders: Portulaceae, Malpighiaceae, Ampelidaceae)
Papilionaceae, CaesalpL.,ieae, Combretaceac, Apocynaceae, Asclepiadeac, Con~
volvulaceae, Phytolaccaceae, Urticaceae; unlignified tissue is very commorJy
developed in lianes, showi.l1g anomalolls structure. Other kinds of special
elements, which have been observed in .the wood, ar~: the short pro~nchy­
matous cells of certain .Papilionaceae (Fig. 60, p. 275), which have a special
shape, and are provided WIth wide lumina and sieve·like E:r..d-walls; and the
special barrel- or spindle-shaped cells of certain Cactcae (Fig. 9I, p. 410),
which show annular thickenings or are provided with a spiral band. .
For the rare occurrence of secretory receptacles in the 'Wood, see § 14
(secretory celIs), § 16 (mucilage-cavities), § 17 (mucilage-canals) and § 19
{secretory canals).

§ 48. AxIAL WOOD AND \VOOD SHOWING A TIER-LIKE STRUcr£RE. Axial

wood (central wood or ring wood) is widely distributed in lianes, belonging
to a large number of different Orders; it forms the innermost anllUlar portion
of the secondary xyiem, and contains ·vessels, which only have small lumina,
so that it is distinctly marked off from theJater growth of the wood (periaxial
wood) 1.
t A xylem-mas~ showing a tier-like str~ctureJ has been recorded by
Hohnel 2 and other authorities in certain Malvaceae, Sterculiaceae, Zygophylleae,
Simarubaceae, Meliaceae, Sapindaceae, Papilionaceae, Caesalpinie3.e, Mimoseae,
Rosaceae, Combretaceae, Ebenaceae, and Bignoniaceae; in the Zygophylleae
and CaesaJpinieae it is of alm05t lll1iversal occurrence. In the wood from the
stem of these species delicate transverse lines, which are occasionally somewhat
undulated; 'can be recognized on the surface of a tangential section even with
the naked eye or with the help of a lens. The cause of this phenomenon is
mostly to be found jn the horizontal seriation of the medullary rays, which are
all of approximately equal height, each ray arising from a single ceil of the
cambium. In other cases it is due to a tier-like arrangement of the fibrous
cells of the wood or of the pitted areas on these elements, or both of these
causes may be combined (for details, see loco cit.).

1 cr. H. Schenck, Anat. d. Lianen, J 893, p. 3-

, Hahnel, in Sitz.-&r. Wimer A.kad.) lxxxix, Abt. J I 1884, p. 30, and Sitz.-Ber. deutscb. boL
Gesellsch., 188.... p. 2.
 CONCLUDING REMARKS

§ 49. PERIDERM WITH SPECIAL REFERENCE TO- THE In employjng

CORK 1.
the characters of the cork for taxonomic purposes we have firstly to take into
consideration the place of origin of the first layer of cork, and secondly the
detailed structure of the cork·cells. The place of development of the cork
in the subaerial parts of the axis 2 is occasionally (although there are generally
a few exceptions) characteristic of entire Orders or even of groups of Orders,
while in other cases it at least serves to distinguish genera; the cork arises
either in a superficial position (i.e. in the epidermis or in the subepidermal
layer, or at least in a superficial layer of the primary cortex) or in the interior
(Le. in a deeper cell·layer) of the primary cortex or in the pericyc1e (when primary
hard bast is present, on its inner side). When the cork is formed in a super-
ficial position, its exact place of origin (the epidermal or subepidermal layer
of cells, the subepidermal layer or a somewhat deeper layer of primary cortical
cells) generally varies from species to species; in some cases, however, .the
development of the cork from a definite layer is characteristic of the genus
(e.g. in Salix and Papulus, see also the Thymelae'l-ceae), while in other cases
it differs even in different specimens of one; and the same species. The forma-
tion of the cork is particularly irregular in certain species of the Papilionaceae,
in which the cork-cambium may develop in the first to the sixth cell-layer
of the primary cortex. A further irregularity in the development of the cork
has been observed in certain Menispermaceae and Convolvu!aceac; here the
different parts of the ring of phellogen arise in different positions along the
circumference of the branch, viz. at some points in the epidermis, at other
points in a deeper cell-layer of the primary cortex, and so on. A similar
·phenomenon is shown by many plants with .reduced leaves 3, where it allows
of the retention of the assimilatory tissue of the stem for a considerable length
of time. It remains to mention the occurrence of rings of cork in the wood of
Sedum papulifolium, and in the pith of species of Campanula and Phyteuma 4.
Superficial development of the cork has been observed in the following Orders
and genera respectively: Di11eniaceae pro parte, Calycanthaceae, Magnoliaceae,
Trochodendraceae, Anonaceae, Menispermaceae pro parte, Berberideae pro parte,
Capparideae pro parte, Cistineae pro parte, Canellaceae, Bixineae pro parte, Pitto-
~reae pro parte, Tremandreae, Polygaleae, Vochysiaceae pro parte, Frankeniaceae,
caryophylleae (rare),Tamariscineae pro parte, H ypericineae, Guttiferae,Ternstroemia~
ceae ]?ro parte, Strasburgeria, Microsemma, Dipterocarpeae, Ancistrocladus pro parte,
Lof>bJ~a. Mal vaceae, Sterculiaceae, Tiliaceae, Rhaptopetalaceae, Lineae, H umiriaceae,
Malpighiaceae. Zygophylleae pro parte, Geraniaceae pro parte, Rutaceae, Simaru·
baceae (excl. Suriana). Ochnaceae, Luxemburgiaceae, Wallacea, Burseraceae, Melia4
ceae, Chailletiaceae, Olacineae, Octocnemaceae, llicineae pro parte, Celastrineae pro
parte, Hippocrateaceae pro parte, Pentaphylacaceae, Corynocarpaceae, Rhamneae,
Ampelidaceae pro parte, Sapindaceae pro parte (almost always), Didiereae, HipP04
castanaceae. Aceraceae pro parte, Staphyleateae, Sabiaceae, Anacardiaceae, Coria..
rieae, Connaraceae, Papilionaceae pro parte, Caesalpinieae, Mimoseae, Rosaceae pro
parte, Saxifragaceae pro parte. Crassulaceae, Hamamelideae, Bruniaceae, Rhizophora·
1 Sanio, in Prillgsru:im Jahtb •• ii. 1860, p. 39 et .eq.; De Bary, Vergl. Anat., 1877, pp. 114
and 560 et sc:q.; Hohnel, in Sitz.·Ber. Wiener Akad., lxxvi, Abt. I, 1877, p. 507 et seq.; Ross,
?eriderma, Malpighia, iii and iv, sep. COP}:, 1~90, 67 pp.; Moller, Rindenanat., 1883; Doulio~,
m Ann. sc. nat., ser. 7. t. x, 1889; J: E. WeiSS, In Denkscbr. bayer. bot. Gesellsch., Regensbnrg, VI,
1890. 69 pp., I Tab.; Segerstedt, in Bibang S". Vel. Akad. Hand1., xix, Afd. iii, D. 4-, 86 pp.,
3 Tab.• abstr. in Bot. Centralbl., 18(}6, i, p. 154-.
, We may point out here that in one and the same species the place of development of the cork
is frequently different in the subaerial parts of the axis and in the subterranean organs (rhizomes and
roots). Not uncommonly the cork arises superficially in the subaerial parts, while in the subterranean
parts its development is pericyclic; the latter is Altogether the rule in roots;
S Ross, in Ber. deutsch. bot. Gesellsch., 1886, p. 361, and in Nuovo Giom. bot. ltaL, xxi, 1889;
see also Ross, Asa. Gew., etc., Diss., Freiborg i. Br., 1887.
• For similar formations of cork in the root of species of C.1psophila, TluzpsitJ, R.",ux,
T~grm, Bec., see Maheu et Combes, Formations snhtro-pbeUodermiques anonnales, Bull. Soc.
bot. de France, 190 7, pp. 4-29-4-2.
 NORMAL STRUCTURE OF THE AXIS 1147
ceae, Combretaceae pro parte, M yrtaceae pro parte, Lecythidaceae, Melastomaceae
pro parte, L ytbrarieae (very rare), T,.af.' a, Sam ydaceae, Turneraceae. Passifloraceae pro
parte, Begoniaceae, Cacteae, Urnbelhferae pro parte, Araliaceae, Cornaceae, Capri-
foliaceae pro parte, Rubiaceae pro parte, Valerianeae pro parte, Dipsaceae pro parte,
Conipositae, Goodeniaceae pro parte, Lobeliaceae, Diapensiaceae pro parte, Plum-
bagineae, Primulaceae pro parte, Myrsineae, Sapotaceae, Ebenaceae pro parte,
S~ae pro parte, Oleaceae pro parte, Salvadoraceae, Apocynaceae, Ascle-
piadeae, <Logamaceae pro parte (Loganioideae, Polyp,.emum and Peltanthet'tJ).
Hydro£hyllacea.e, Boragineae pro parte, Convolvulaceae, Nolaneae, Solanaceae pro
parte, Scrophularineae pro parte, Gesneraceae pro parte, BifQloniaceae pro parte,
Pedalineae, Acanthaceae pro parte, Myoporineae, Zombiana, Selagineae pro parte,
Verbenaceae pro parte, Labiatae pro parte, Plantagjneae pro parte, Nyctagineae
pro parte, Illecebraceae pro parte, Amarantaceae, ChenOpodiaceae pro parte,
Basellaceae, Phytolaccaceae, Polygonaceae pro parte, Aristolochiaceae, Piperaceae,
Myristicaceae, Monimiaceae, Laurineae, Hernandiaceae, Proteaceae. Thymelaeaceae,
OaoJepis, Gonystylus, Geissoloma~ Elaeagoaceae, Loranthaceae, Santalaceae, Myzo·
dendron, Champereia, Grubbia, Euphorbiaceae pro parte, Buxaceae pro parte,
Balanopseae, Ulmaceae, Moraceae, Urticeae prq parte, Platan ace ae, Leitnerieae,
Juglandeae, Myricaceae, Cupuliferae. Salicineae, Lacistemaceae.
Internal development of the cork, i.e., from a deeper layer of cells, which is still,
however, situated)n the primary cortex, is found in the following Orders and genera:
Dilleniaceae pro parte, Menispermaceae pro parte, Berberideae pro parte, Cruciferae
pro parte, Capparideae pro parte, Cistineae pro parte, Bixineae pro parte, Pittosporeae
pro parte, Vochysiaceae pro parte, Tamariscineae pro parte, Ternstroemiaceae pro
parte,Ancist1'oc;ladus pro parte, Geraniaceae (T1'Dpaeolum), Suriana (Simarubaceae),
llicineae pro parte, Cyrilleae (pericyclic ?), Celastrineae pro parte, Hippocrateaceae
pro parte, Aceraceae pro parte. Melianthaceae. Papilionaceae pro parte, Rosaceae pro
parte. Combretaceae pro parte ?, Myrtaceae pro parte, Lythrarieae pro parte, Passi·
fioraceae pro parte, Cucurbitaceae, Ficoideae, Rubiaceae pro parte,Oleaceae pro
parte, Des/ontainea, Gentianeae (pericyclic ?). Folemoniaceae pro parte?, Bora·
gineae pro parte, Bignoniaceae pro parte, Verbenaceae pro parte, Labiatae pro parte.
Plantagineae pro parte, Nyctagineae pro parte, Chenopodiaceae pro parte, Euphor-
biaceae pro parte, Urticeae pro p a r t e . ' .
Pericycllc "development of the cork has been observed in: Ranunculaceae,
Dilleniaceae pro parte, Berberideae pro parte, Cruciferae J>~o parte, Cistineae pro
parte, Bixineae pro parte, Pittosporeae pro p~e, VochYSlaceae pro parte, Caryo~
phylleae pro parte, Hypericineae pro parte. Ternstroewaceae pro parte, Ancist,.o-
claaus pro parte, ZygophyUeae pro parte, KoebeYlinia, Ampelidaceae pro parte (in
the bast ?), Sapindaceae pro parte. Papilionaceae pro parte, Rosaceae pro parte,
Saxifragaceae pro parte, Combretaceae pro parte, Myrtaceae pro parte, Melastomaceae
pro parte, Lythrarieae pro parte, Onagrarieae (excl. T1'apa), Loaseae, Umbelliferae
pro parte, Caprifoliaceae pro parte. Rubiaceae pro parte, Valerianeae pro parte, Dipsa·
ceae pro parte. Goodeniaceae pro parte, Campanulaceae"Lobeliacea.e, Vacciniaceae,
Ericaceae, Epacridea.e, Diapensiaceae pro parte, Primulaceae pro~. Ebenaceae
pro parte, Styraceae pro parte, Loganiaceae (most Buddleioideae), Polemoniaceae
pro parte, Boraginea.e pro parte, Solanaceae pro parte, Scrophularineae pro parte.
Columelliaceae, Gesneraceae pro parte, Acanthaceae pro parte (really endodermal),
Selagineae' pro parte. Labiatae pro parte, IUecebraceae pro parte, Chenopodiaceae
pro parte (see p. 1029). Batideae, Polygona.ceae pro parte, Nepenthaceae, Penaea·
ceae, Euphorbiaceae-pro parte, Buxaceae pro parte, Empetraceae 1 •
In the case. of repeated development of cock on the same axis, the
arrangement of the successive cork-cambia with reference to one another,
and the resulting mode of formation of the bark (whether scale.. or ring-bark),
afford characters, which are of systematic value; as a rule, however, these
features can only be determined in axes of some thiclmess, and not in h,erbarium-
material_ Ring-bark is found only in species, in which tlIe firsf·phellogen
appears.in a relatively deep layer of cells. The peculiar repeated dbvelopment
of cork in the Sapindaceous genera Dodonaea and Distichostemon and some

1 Peric),clic de,·e)opment of th~ cork is prevalent in herbaceous plants and in plants baving aD
cricoid habit.
 CONCLUDING REMARKS

other plants, and the lamella ted or tier-like cork of certain Saxifragaceae, Rubia ..
ceae, Loganiaceae (Fig. 125, p. 544), and Labiatae (which is due to "J'epeated
development of cork), still require a more detailed discussion; both .of these
features can be recognized already in the branches of herbarium-material. In
Dodonaea and Distichostemon the development of the cork commences in the
pericyc1e ~m the inner side of the ring of pericyclic sclerenchyma, and is accom-
panied by the formation of a many-layered phelloderm; subsequently, a fresh
development of cork takes place internal to a second ring of sclerenchyma, which
arises at the inner limit of the phellodenn, and so on; similar features are
found also in species of ColumeUia (Columelliaceae) and in species of Rosmarinus
and Salvia (Labiatae). In the formation of the lamellated cork consecutive
layers of the primary c6rtex develop from without inwards into cork-cambia,
each of which gives rise only to a very small number of layers of cork-cells ;
as a consequence, the rows of cork-cells belonging to the· individual Hers do
not correspond with one another in the radial direction.
The detaiJed structure of the cork-cells varies, but is frequently only
of value for specific diagnosis. The walls of the cells are either (a) thm, the
cells in this case frequently having very wide lumina (spongy cork), and at the
same time often being considerably elongated in the radial direction, or (b)
thick, in which case the cells are compressed in various ways, .often showing
a very marked radial compression (tabular cork), or (c) sclerosed (stone-cork).
In the latter case the corkAcells are either uniformly sclerosed, or the process
of sclerosis affects only one side of the cell, or merely a horseshoe-shaped
piece of the wall; in the second case sclerosis is confined either to the
outer or to the inner tangential walls. The sclerosed cells either compose
the entire cork or are confined to certain layers or occur as isolated elements
in the cork. In certain species of Croton (Euphorbiaceae) the structure of the
walls of the cork-cells is particularly noteworthy, since the inner tangential
walls are encrusted with small crystals of oxalate of Jime, while in LitJ.uidamba,
(Hamamelideae) the cells of the cork are partly silicified. In certain plants,
moreover, the cork contains unsuberized cells having walls, which consist
r
of cellulose or are even more or less lignified phelloid-ce1ts ') 1,
The occurrence of layers of phelloid-cells (' Trcnnungsphelloide' of Hohnel),
alternating with one or more, sometimes even a large number, of layers of cork-cells,
is of considerable taxonomic value; smaller systematic value is to be attributed
to the often abundant occurrence of phelloid-cells (' Massen-' or • Ersatzphelloid ')
in thick ma.s...~ of cork, the strong development of which is frequently only a local
phenomenon. Layers of phelloid-ceJls have been recorded in the following Orders:
Hypericineae, Burseraceae (phelloid-cells with silicified inner tangential walls I),
Rosaceae, Combretaceae, Myrtaceae, Melastomaceae, Lythrarieae, Onagrarieae,
CaprifoUaceae', Penaeaceae. The species involved are, in almost all cases, such as
show internal development of the cork.
A cork containing phelloid-cells is related to the mucilaginous cork/ I

which has been observed in certain desertaplants belonging to the Orders

Papilionaceae, Chenopodiaceae, and Polygonaceae 3
. At this point we may also refer to the structure of the cork in the
Epacrideae, ·in which the cells do not show a distinct radial arrangement.
The development of the cork in this case still requires further investigation.
Contents of a special kind are found·in the cells of the cork in Adesmia
(Papilionaceae), Sarcocaulon (Geraniaceae) and Betula (Cupuliferae). CArk-
cells having wide lumina often contain nothing but air; in other cases one
finds remnants of the dead cell-contents, which are frequently coloured brown
by phlobaphenes. .
J See HOhnel, loco cit., p. 595, and J. E. Weiss, loe. cit., p. 6. 9 J See Hohnel, Joe. cit.
• See Jo.won, Anat. Ban der Wiistenpflanzetl, Lunds Universitets arsskrift, xxxviii, Ald. "
n. 6, 1902.
 NORMAL STRUCTURE OF THE AXIS 1149

The thickness of the cork can also to a certain extent be employed for
systematic purposes. With regard to this point, we may note that strong
local proliferations of cork in the form of tubercules, ridges, &c., which are
visible to the naked eye 1, have long been taken into consideration.
The development of phelloderm 2 is a very widely distributed feature,
although as a general rule of little taxonomic interest. In many cases the
pheUoderm does not differ in any way or only to a slight extent from th~ tissue-
of the primary cortex, and under these circumstances it can only be recog~
nized as phelloderm, if its course of development is studied. In other cases,
however, it is readily identified owing to the radial arrangement of its cells.
The phelloderm is sometimes completely or parJ:ially sclerosed. That of
CaneUa, for instance, is very characteristic, being compOsed of radially arranged
cells, the inner tangential and radial walls of which are sclerosed.
A point, which has hitherto received too little attention, as far as its
taxonomic application is concerned, is the succession of divisions in the
cork~cambium; these divisions, which are parallel to the surface of the cortex,
lead to the formation of the cork and phelloderm; Sanio 3 distinguishes the
most important types as ' centripetal,' , centrifugal,' and' reciprocal.'
The structure of the lenticels" (cortical pores) .likewise requires further
attention; they are often present on the branches of herbarium-material and
have already been used by systematists in their diagnoses. Two types of
lenticels may be distinguished. In those of the first type the tissue of the
lenticel consists solely of complementary cells, which remain in rather firm
connexion, although separating from one another to a varying extent by the
rounding off of their edges. In the lenticels of the second type layers of loosely
arranged complementary cells, which are quite distinct from one another
and frequently fonn a powdery mass, alternate with firm and compact layers
of thick-walled cells, which are often of the nature of cork-cells.
The development of periderm, lastly, may be postponed for an often
considerable length of time 5, and this phenomenon is also of systematic impor-
tance. Late formation of the periderm occurs in the Orders enumerated
below, and is also found prevalently in plants with reduced leaves, which, owing
to the reduction of their foliage-leaves, are dependent upon the assimilatory
tissue in the primary cortex. In the Visceae (Loranthaceae) and certain
,Menispermaceae and Papilionaceae (Oxylobium) , in which cork only develops
at a late stage or, as in the case of the Visceae, is never formed at all, its
place is taken by what is called a cuticular epithelium 6'; the latter is
I

constituted by cells of the epidermis and primary cortex, in which the outer

1 A synopsis of proliferations of this kind will be found in Barber, in Annals of Bot., vi, .sta,
p: 163; the species named in this synopsis belong to the Malvaceae (Bombaceae), Rutaceae,
Simarubaceae, Rhamneae, Leguminosae(Papilionaceae, Caesalpinieae, Mimoseae), Rosaceae, Cacteae,
Araliaceae, and Enphorbiaceae.
3 K~I~,_~~t8teh. D...~erbr~it. des Phelloderms, Bot. Centralb1., 1891, iii. p. 81 et seq.
a Sanio, loc. cit., p. -44 et seq.; see also J. E. Weiss, loco cit., p. 38. and p ...8 under 9 and 10.
t Stahl, Entwicklungsgesch. u. Anat. d. Lentizellen, Bot. Zeit., 1863, p. 56t et seq.; Klebabn,
Struktnr u. Funkt. d. Lentiz., Ber. deutsch. bot. GeseHsch., 1883. p. 113 et seq., and Rindenporeo,
Diss., Jena, 188... also in Zeitschr. f. Natnrw., xvii, Neue Folge, x, p. 537 et seq.; Devaux, Lenticclles•.
Ann. Be. nat., ser. 8, t. xii, 1900, p. J et seq. '
I The genera and species, which are mentioned in Moller's' Rindenanatomie' and in the
respective papers by Ross and Damm as showing late development of periderm, belong to the
following Orders: Menispermaceae, Polygaleae, Gtlttiferae, Temstroemiaceae, Malvaceae, Geraniaceae,
Rutac;eae, Ilicineae, Celastrineae, Rhamneae. Aceraceae, Staphyleaceae, Papiliona.ceae, Caesalpinieae,
Mimoseae. Rosaceae, Myrtaceae, Arali:l.Ceae, Comaceae, Compositae, Sapotaceae, OIeaceae, Loga-
niaceae, Scrophularineae, Polygonaceae, Aristolochiaceae, Laurineae, Proteace!le, Loranth~ceae,
Euphorbiaceae, Buxaceae, Urticnceae. .
II Damm, Bau mehrjahr. Epid. bei den Dicotyled., Beih. :t. bot•. Centralbl., xi, 1902, p. f19
el seq. and Tab.
 CONCLUDING REMARKS

wa1ls become thickened by the formation of cuticular layers. The superficial

layer of wax, differentiated in Acer pennsylvanicum, L., is closely related to
such a cuticular epithelium; in this species of Acer the outer epidermal walls
and thereupon the lateral and inner walls of the epidermal cells, and ultimately
also the walls of the layers of cortical cells, situated internal to the epidennis, in
turn become thickened and changed into a wax-like substance, while in other
species of Acer the epidermis alone undergoes transformation into wax 1.
The peculiar gelatiniiation of the outer portions of the walls of cells of the
epidermis and hypoderm in the stem of Calligonum sp. (Polygonaceae)2 is like ..
wise similar to a cuticular epithelium.
The formation of suberized tissue without the help of a phellogen is
found in certain Labiatae, and (see above) apparently also in the Epacrideae (in
which the cells of the bast undergo suberization from without inwards), as well as
in Asa,um (Aristolochiaceae, in which the primary cortex becomes suberized).
Development of cork in the leaf is rare (Fabiana, Solanaceae); for
cork·warts on the leaf, see § 39 (p. 1133).
§ 50. AERENCHYMA 3. In certain plants aerenchyma (i.e. a lacunar
tissue, composed of living cells with thin unsuberized walls, Fig. 81, p. 374) is
produced by the phellogen in place of cork on those parts of the stem and of
. the older roots, which are submerged or are embedded in wet soil. Aerencbyma
is formed only in a moist habitat, and it is possible to induce its development
by artificial means. It has been observed in the following Orders: CaPJ?ari..
deae, HyPCricineae, Papilionaceae, Mimoseae, Melastomaceae, Lythraneae,
Onagraneae, Labiatae, and Euphorbiaceae. The groups of phelloid-cells,
which are found in the cork of certain plants (see § 49, p. 1148) and are provided
with intercellular spaces, may be regarded as a simple type of aerenchyma.
§ 51. PRIMARY CORTEX"'. The differentiation of the assimilatory tissue
and endodermis, and the occurrence of collenchyma and sclerenchyma provide
a number of structural differences, which may be employed for systematic
purposes, althJugh only of value for special diagnosis. A palisade-like differen-
tiation of the hypodermal tissue is found chiefly among plants with reduced
leaves s, and occasionally goes hand in hand with the development of furrows
on th~ surface of the stem and the restriction of the stomata to the epidermis
of these furrows (see Fig. 61, p. 278, and Fig. 186, p. 788); the same type of
differentiation is shown also by the wing·like appendages, found on the stems
of plants with decurrent leaves. Ordinary collenchymatous tissue is Qf
frequent occurrence in the cortex and is restricted either to a hypodermal layer
or to a median zone in the primary cortex (see Moller, loco cit., p. 416 et seq.).
Typical collenchyma (of the well-known form, as found in Begonia or Cucflrbita)
is rarer; it is found in the same position as the ordinary collenchyma, but
may also be developed in the form of strands on the outer side of the bast-
groups. Subepidermal collenchyma of the typical kind frequently participates
in the formation of the ribs on the stem. In some cases (e.g. in certain Labiatae
and Piperaceae) cells of the typical collenchyma may secondarily become
sclerosed and thus transformed into elements resembling bast-fibres. As

1 Uloth, in I<lora, 1867, p. 385 et seq.

\IJonsson, in Londs Univers. .Arsskr., xxxviii, A(d. 2, n. 6, 1902, p. 20.
S H. Schenck, Aercnchym, in Pringsheirn Jabrb., xx, 188 9. R' 525.
~ R~garding tlre cortex in general, see Vesque, Anat. de 1 ecorce, Ann. sc. unt., st!r. 6, t. ii.
1875, P. 82; Hahnel, Gcberinden, Berlin, 1880; Milller, Rinde unserer-Laubh., Diss., Breslau,
J875; Moller, Rindenanat., 1882. .
_. See Schube, Blattanne Pit, Breslall, 1885; Pick, Ass. Gewebe annlanb. Gewachre., Diss.,
Bonn, 1881 ; Nilsson, StudieT aev. stammen aAsom assimiler. org., GOteborgs k. Vetensk. och Vitter.
beUs Samhall. Handl., 1887; Ross, in Nuov. Giom. bot. Ita1., xxi, 1889, etc.
 NORMAL STRUCTURE OF THE AXIS 115 1

regards the occurrence of sclerenchyma, it is necessary to bear in mind the fact

that sclerosis of the primary corteJC often sets in only at a late stage, so that
it is sometimes impossible to determine this feature adequately in the branches
of herbarium-material. Features deserving special consideration are: (a) the
occurrence of bundles of sclerenchymatous fibres· (b) the presence of.a ring
of stone-cells; (c) the presence of cells, which (in a transverse section) are
sclerosed,in the shape of aU; (d) the occurrence of l cristarque '-cells; and
(e) the spicular cells and stomge-tracheides, already discussed. in §§ 9 and II.
We may note that the l cristarque '-~ (Le. cells, which are thickened in
a U-shaped manner, each of them closely enveloping a single solitary or clus-
tered crystal) 1 occasionally form complete zones (' cristarque '), which are
more or less interrupted and coincide either with the first or second cell-layers
of the primary cortex or with the endodennis; when such a cristarque ' l

is present in the body of the corte~ it sometimes occurs in a corresponding

position also in the petiole and the veins of the leaf of the same species. In
plants inhabiting damp localities and especially in water-plants the primary
cortex contains a system of intercellular spaces, which is often strongly
developed.
The endodermis (protective sheath, _phloeoterma) 2, i.e. the innermost
cell~layer of the primary cortex, affords but few anatomical characters. In
woody species it is at the best developed only in the young parts of the axis,
being differentiated as a starc1;t-sheath and containing amylon. In the different
parts of the stem of herbaceous plants it often shows up distinctly owing to the
suberization of its radial walls, which are provided with Caspary's dots or
lines. The divisions in the endodermal cells (for the most part by means of
radial walls), characteristic of many Gentianeae-Gentianoideae, are found not
only in the root, but sometimes also in the stem of this group of plants.
The secretions and excretions found_ in the primary cortex have already
been discussed in §-I3 et seq.
TyPical collenchyma has been observed in certain Malvaceae, Cucurbitaceae.
Begomaceae, Araliaceae, Umbelliferae, Dipsaceae, Compositae, Labiatae (four
strands of collenchyma in the four angles of the stem), Amarantaceae, Chenopodia-
ceae, Phytolaccaceae, Polygonaceae and Piperaceae a; quite a special form of collen-
chyma occurs in the Cacteae (Fig. 90, p. 407). The primary cortex contains com-
plete zones or groups of sclerenchymatolls fibres or isolated sclerenchymatous
fibres, some of which may have developed/secondarily from collenchymatous cells,
in certain Tremandreae, Pittosporeae"Tamariscineae (m Fouqu£era there is a peculiar
sclerenchymatous zone, which arises in the subepidermal layer of cells and consists
of many layers), Polygaleae, Hypericineae, species of Anc£strocladus, species of
Lophira. Simarubaceae, Luxemburgiaceae, Stackhousieae, Papilionaceae, Rosaceae
(Prunus with fibrous cel~s, some of which have a horizontal course), Halorageae,
Umbelliferae. Plumbagineae. Oleaceae, Asclepiadeae, Loganiaceae, Scrophularineae,
Bignoniaceae. Acanthaceae. Verbenaceae, Labiatae, Polygonaceae, Piperaceae,
Santalaceae, Euphorbiaceae, and Casuarineae (Fig. 186, p. 788) ; in :Plants with re-
duced leaves (see Fig. 6 r, p. 278). the scleren!=hymatous fibres occasionally form plates,
which traverse- the entire primary cortex, extending from the epidermIS to the bast-
groups of the vascular bundles. A ring of stone·cells has been observed in the primary
cortex in certain Capparideae, Guttiferae, Simarubaceae, Burseraceae, Meliaceae,
Anacardiaceae, Saxifragaceae, Melastomaceae, Asclepiadeae, Loganiaceae (Strychnos)f
Bignoniaceae. Monimiaceae, species of Myzodendron, Euphorbiaceae. Balanopseae.
Moraceae. Salicineae, as well as in Brachynema, and (according to Moller, p. 419)

1 See especially Van Tieghem, Le cristarque dans la tige et la feuille des Ochnactes, Bull. Mus.
d'hist. nat., xvi, 1902, p. 266 et seq. j and Van Tieghem, in Ann. sc. nat., ser. 8, t. xvi,
J 902, p. 166 et seq., etc.
I For the distribution of the endodermis among the Dicotyledons, see SchoJte, Stehirtheorie,
Jena and Groningeo. 1903, pp. 108-32.
~ See also Schweodener, :\Iecban. Prinzip, p. 15i.
 CONCLUDING REMARKS

also in certain A pocynaccae and Asclepiadeae; in some of these cases, the ring of
stone~cellsarises in the phelloderm. Cells, sclerosed in a U-sbaped. manner, are
present in the primary cortex in certain Calycanthaceae, Canellaceae, B.ixineae,
Dipterocarpeae, Simarubaceae, Melastomaceae, Gesneraceae, Buxaceae and Empe~
traceae. Cristarque-cells occur in the Rhaptopetalaceae, Lineae, Simarubaceae
(Irvingieae), Ochnaceae (with clustered crystals), Luxemburgiaceae (apparently
only in the-veins of the leaf), and Octocnemaceae. Parenchymatous cells, provided
with ridge-like thickenings, are found in the lacunar cortical parenchyma of Herpestis
Monnieria (Scrophularineae). Regarding the transformation of the outer cells of
the primary cortex (together with the epidermal cells) into a cuticular epithelium
or a COIJlplete layer of wax, sec § 49 (pp. 1149-1 150).

§ 52. PERICYCLE 1. The term 'pericycle' is on the whole a practical

designation, which is·not altogether founded on a developmental basis; it is
used to indicate that region of the cortex of the stem, which lies between the
innermost cell-layer of the primary cortex (which is sometimes differentiated
as a distinct endodermis) and the vascular system (or, to put it more exactly,
the bast-groups and the intervening primary medullary rays of the cortex).
The pericycle is differentiated in a variety of ways. In some cases it is paren-
chymatous throughout. When this is the case, the pericyclic region can
occasionally (and especially in the cortex of the la:rger branches) be deter-
mined only by tracing the primary medullary rays of the cortex (in a trans-
verse section) outwards in the radial direction up to the point, at which they
merge into the tissue of the primary cortex. In other cases the pericyc1e is
distinguished by the development of sc1erenchymatous elements and is diffe-
rentiated as an interrupted or continuous mechanical strengthening ring;
under these circumstances the pericycle is.: generally easily recognized. In
a pericycle of this kind, a parenchymatous zone, which resembles the ground
tissue, 15 'frequently intercalated between the pericyclic sclerenchyma and the
vascular system or rather the bast-groups of the latter; this zone either con-
sists only of a few layers of cells or is strongly developed (e.g. in the Meni~
spermaceae or Cucurbitaceae), and is described as the parenchymatous pericycle;
when it is wanting, the sclerenchymatous elements of the peri cycle are in
direct contact with the bast-groups. The sclerenchymatous pericycle consists
either (a) of a non-sclerosed parenchymatous region, containmg only isolated
bast-fibres (primary bast-fibres 2) or groups of bast-fibres of varying dimensions,
or (b) of a completely or more or less completely closed ring of bast·fibres,
or (c) of a completely or more or less completely continuous and composite
(i.e. composed of bast-fibres and stone-cells) ring of sclerenchyma.
The differentiation of the sderenchymatous pericycle may vary in branches
of different thickness in one and the same species. Thus in young branches
the pericycle is occasionally found to contain a closed ring of bast-fibres, which
during the subsequent growth in thickness of the branch becomes broken
up into isolated groups of fibres (the latter decreasing in size as secondary

1 Vau Tieghem, in Bull. Soc. bot. de France, I88~. p. 380; Morot, in Ann. sc. nat., aer. 6,
t. xx, 1884, p. 211. and Bull. Soc. bot. de France, 1886. p. 2°3; d'Arbautllont, in Bull. Soc. bot.
de France. 1886, p. I4t; H. Fischer, Der Pericykel. in Pringsheim· 1ahrb., XXXY, 1900, pp. 1-27
and Tab. i; Pitard, L'evolution et la valeur anat. et taxinom. du peri cycle des Angiospermes, These,
Bordeaux, 1901, 197 pp., 5 pl., also in Mem. Soc. sc. phys. et nat. Bordeaux, l lvi, 1901; see also
the fwther papers by Pitard (in M~m. Bordeaux, Iv and lvi, 1900-1) wbich are cited in this thesis.
I Tbe bast-fibres of the sclereochymatotl5 pericycle (primary bast-fibres) are frequently dis-
tinguished from the ~at-fibres of the secondary bast (secondary bast-fibta) by their atructure
(the outline and coloar ~ing different, while the walls show a slightly different chemical com-
position, which is probably c:onnrcted with the difference in colour). To mention examples,
with which everyone i, familiar, such points of difference arc shown by the oost-fibres, which
compose the ontermost layers 0[ hard ba$t in the phloem-groups of the lime (which are narrowed
outwards in the form of wedges), and which are to be regarded as pericyclic elements, by the isolated
groups of bast-fibres in the pericyc1e of • Cortex Frangulae,' etc.
 NORMAL STRUCTURE OF THE AXIS .1153

growth continues), while parenchymatous tissue penetrates into the gaps

thus formed both from the outer and inner sides; this parenchyma is either
not sclerosed or becomes more or less completely sclerosed. In this way
the thicker axes of one and the same species contain either isolated groups
of bast-fibres of varying size or a composite ring of sclerenchyma, which is
either intenupted or continuous; in thick branches, moreover, the composite
ring of sclerenchyma naturally includes a far larger number of stone-cells
than in those which are slightly thinner, while the proportion of bast-fibres
as compared with stone-cells decreases more and more. In the course of
further growth in thickness, such a composite and continuous rin9' of scleren-
cbyma may again be burst open, unless prior to that it falls a Victim to the
formation of bark. It remains to mention that the parenchymatous pericycle,
found on the inner side~ of the sclerenchyrnatous peri cycle in some plants,
commences to develop only at a certain stage.
In employing the pencycle for systematic purposes, it is of the utmost
importance to take due consideration of its progressive development and of the
transformations, which it undergoes. The most valuable feature from this point
of view has proved to be the composite and continuous ring of scleren-
chyma, which is frequently characteristic of entire Orders or genera.
In utilizing the pericyclic sclerenchyma for taxonomic purposes it is, however,
necessary to keep in view its physiological importance as a mechanical tissue.
Although the ring of pericyclic sclerenchyma dOes not occur i.n all Primulaceae,
it is nevertheless characteristic of this Order, since it is absent only in those species,
which have no need of flexile strength (Westennaier). Among the Cucurbitaceae
a strengthening ring is found in the pericycle only in those species, which show no
growth in thickness. In succulent plants there is no peric"yclic sclerenchyma. '
A composite and continuous ring of sclerencbyma is found in the perlcy-ele
in the following Orders and genera respectively: Ranunculaceae pro ~arte, Dille-
niaceae pro parte. Calycanthaceae pro parte, Magnoliaceae pro parte, Trochoden· ~
draceae pro ,Parte, Menispennaceae pro parte, Berberideae pro parte, Capparideae
pro parte, VlOlarieae pro parte, Bixineae pro parte, Tremandreae pro parte, Vochy-
siaceae pro parte~ Tamanscineae pro parte, H~ricineae pro parte, Guttiferae pro
parte, Ternstroemiaceae pro parte, St1'asbuf'ceria, LopM1'4, Tiliaceae pro parte, Lineae
pro parte, Humiriaceae, MaJpighiaceae pro parte, Zygophylleae pro parte, Gerania-
ecae pro parte, Ruta.Geae pro parte (rare), Simarubaceae pro parte, Koebtf'linia,
Ocbnaceae pro parte, Burseraceae, Meliaceae (very rare), Olacineae pro parte,
Octocnemaceae,llicineae pro parte. Celastrinea.e pro parte, Hippocrateaceae pro
parte, PentaphyLacaceae. Rhamneae pro parte. Ampelidaceae pro parte, Sapindaceae
(almost always). Hippocastanaceae, Aceraceae pro parte. Sabiaceae pro parte,
Anacard.iaceae pro parte, Moringeae pro parte, Connaraceae, Papilionaceae pro
parte, Caesalpinieae (almost always), Mimos.eae pro parte, Rosaceae pro parte,
Saxifraga.ceae pro parte. Hamamelideae, Ost"earJa. Rhlzophoraceae pro parte, Samy-
daceae pro parte, Passi1loraceae pro parte, Datisceae pro parte, Cornaceae pro parte,
Caprifoliaeeae pro parte, Epacrideae pro parte, Myrsmeae pro part~, Ebenaceae pro
parte, Styraeeae pro parte._Oleaceae pro parte. Salvadoraceae pro parte, Hy~ro.
phyUaceae pro ~. Boragineae.~ parte, Acanthaeeae pro~. Verbenaceae
pro parte, "Illecebraceae pro parte, Phytolacca.ceae pro parte. Polygonaceae pro
parte, Aristolochiaceae pro parte, Piperaceae pro parte. Chloranthaceae pro parte,
Monimiaceae pro parte~ Laurineae pro parte~ Hernandiaceae pro parte. Gomortega.
ceae, Proteaceae pro parte, Geissoloma, Santalaceae pro parte, Clulmpe,.eia, GndIbia
pro parte~ Euphorbiaceae pro parte. Balanopseae, Ulmaceae pro parte, Platanaceae.
Jugfandeae pro parte, Myncaceae pro parte, Casuarineae, Cupuliferae, Lacistemaceae.
The exact nature of the elements composing the pericyclic sclerenchyma,
and especially the detailed structure of the bast-fibres (length, structure of the
walls and Jumina,.ihe kind of pitting t, and the occtlrrence of septation). also requires
to be taken into account in systematic anatomical researches. In the following
cases the arrangement or structure of the elements forming the sclerenchyma-ring
is particularly noteworthy: in Balanites (Simarubaceae •. p. 185), the ring of

J Bordered pits occur on the bast-fibres in the EpacrideaC'.

sou.altDl!.R 4E
1154 CONCLUDING RE}t[ARKS
sclerenchyma consists of a ring of stone·cells, at the inner margin of which groups
of bast-fibres are inserted; in-certain Calycanthaceae, Guttiferae, Ternstroemiaceae,
Simarubacea.e (Irvingieae), Chrysobalaneae, Rhlzophoraceae, SaDlydaceae, Salva..
doraceae, Monimiaceae, Laurineae, Hernandiaceae, Gomortegaceae aIld Euphor-
biaceae the sclerencbymatous ring contains stone-eells, which are thickened in a
U-shaped manner.
§ 53. BAST-GROUPS AND ESPECIALLY TilE SECONDARY BAST. The struc ..
ture of the bast likewise affords an abundance of anatomical characters. These
are as follows: (a) the occasional prominent demarcation of the bast-groups of
the individual vascular bundles (their outline in transverse section in this case.
resembling that of a symmetrical trapezium, or being Convex on its outer side),
a featurewmch goes hand in hand with the OCCurrence of broad primary (cortical)
medullary rays, which often widen out towards the exterior to a marked extent;
(b) the presence or absence of secondary hard bast, which, when present, is deve- 1

loped in varying amount and shows diverse distribution (solitary b ast.. fibres ;
groups of bast-fibres with a scattered, reticulate or stratified arrangement) and
differentiation (chiefly as regards the length,. the appearance in transverse sec-
tion, the mode of thickening, the'thickness of the walls, the size of the lumina and
the kind of pitting of the fibres; the latter occasionally also septate or merging
into sclerenchymatous parenchyma) of its elements; (c) the occurrence of
ordinary stone-cells or spicular cells (see § 9); (d) the structure of the sieve-tubes
(the width of their lumina; the diverse differentiation and arrangement of the
sieve-plates, viz. either (i) simple sieve-plates (i.e. prOvided with a single sie\re-
area), which are mostly situated on horizontal cross-walls, or (ii) scalarifonn
sieve-plates (having from two to many sieve-areas), which are found on strongly
inclined dlvision-walls, the latter being seeR from the surface in a radiallongi-
tudinal section, or (iii) both simple and scalariform sieve-plates side by side; the
occurrence of sieve-plates on the lateral walls; and finally, the delicate or coarse
nature of the pores in the sieve-plates); (e) the breadth of the secondary
medullary rays and the sclerosis of the parenchyJIla of the secondary rays
between the groups of secondary hard· bast, which occasionally leads to the
formation of complete sclerenchymatous rings; (I) lastly. the occurrence and
tmtribution of the manifold excretions and secretions (see § I3 et seq.~ especially
the occurrenCe of chambered crystal-fibres, bearing solitary or clustered c~talst
i.e. fibre-like groups of cells, with numerous transverse septa, the individual
chambers containing deposits of oxalate of lime).
The conditions for the determination of many of these features 1 are
frequently very unfavoura~le in the case of herbarium-material, whl,h is
generally alone available for the purposes of a systematic investigation, since
the features in question can often only be reco~ed, when the secondary
bast has attained a certain degree of thickness. It is therefore impossible to
deal fully with all of the above ..mentioned characters.
The follo"Wing lines, in the first place. contain a list of the Orders and Cinomalous
genera, in which, unless there is a statement to the contrary. typical ~eond8l"J'
hard ba8t (i.e. composed of bast-fibres), has been observed; the OrC:lers and genera,
in.which some of the members at least show a distinct stratification of the phloem
into soft and hard bast, are indicated by a·: ·Ranunculaceae, Dilleniacea.e, *:Mag..
noliaceae, Trochodendraceae, • Anonaceae, Becberideae, Crucifen1e, C3nellaceae,
*Bixineae, POlygaleae. Vochysiaceae ?, *Tamariscineae. *Guttiferae, Ter1lstraemia-
c~, ·Mict'Osemma, ~Dipteroc~ae, *Monotes, Chlaepaceae, *.Malvaceae, .*Triplo-
chitonaceae. ·Sterculiaceae, *Tiliaceae. - Rhaptopetala.ccae, ·Lineae, H umuiaceae,
-Malpighiaceae, Geraniaceae. *Ruta.ceae. ·Simarubaceae. • Balanites I ~ Ochnaceae,
1 See Moller. Rilldenanat., 1882, especially p. -423 et seq.; Lecomte, Liber, .A.nD. Ie; nat., ~r. 7,
t. ,.:, ]889. p. 192; Perrot, Tissu crible. These, Paris, 1899. 243 pp.; Hill, Sieve.-tubes, Annals
of Bot., 1903, pp. 265-7.
J ACCOrding to Van Tieghem j Ann. sc. nat., IC!r. 9. t. iv, 1906.
 NORJ,[AL STRUCTURE OF THE AXIS 1155
*Luxemburgiaceae, Wallacea, Burseraceae, ·Meliaceae, Olacineae, Octocnema·
ceae, Ilicinea.e (rare), Celastrineae, $Hippocrateaceae, *Rhamneae, ...Ampelid~,
ceae, Sapindaceae, Hippocastanaceae. • Aceraceae, Staphyleaceae, *Anacardiaceae,
Connaraceae, *Papilionaceae, Caesalpinieae, ·Mimoseae, -Rosaceae, Saxifragaceae,
Hamamelideae, COmbretaceae, ·Myrtaceae, Lythrarieae, Onagrarieae, Passifloraceae,
.Papa,r,aC(!ae. Datisceae, ·Comaceae, ·Caprifoliaceae, Rubiaceae, Compositae, ~rica­
~eae, Epacrideae, Sapotaceae, Styraceae, ·0 leaceae , Apocynaceae, Bor~nea.e,
.Bignomaceae, Acanthaceae, Verbenaceae, Batideae, *Myristicaceae, Moninuaceae.
Laurineae, ·Proteaceae, ·Thymelaeaceae with Octoiepis and Gonystylus (bas~fibres in
this case projecting like threads of silk on the ruptured surface, when the dried
branches are broken in two); ·Elaeagnaceae, Santalaceae, G1'ubl»a, ·Euphorbiaceae,
Buxaceae, ·Ulmaceae, Cannabineae, ·Moraceae•.·Pla.tanaceae (sclerotic ~enchyma!),
• Leitnerieae, • J uglandeae. M yricaceae, Casuarineae, *Cupuliferae, ·Salicineae.
The following types of bast· fibres deserve special mention (apart from the
special spicular cells occurring in the bast of the axis, and referred to at the end of
§9): (a) the long bast-fibres, found in many Lineae and Urticaceae, and the strikingly
short spindle-shaped bast-fibres of Berberis; (b) the bast· fibres of many Apocyna-
ceae and Asclepiadeae, which are provided with local enlargements; (e) the bast-
fibres of certain Vacciniaceae, Ericaceae and Epacrideae (FIg. II I, p. 492), which
bear bordered pits; Cd) the bast-fibres of many Euphorbiaceae, which show a well-
marked stratification of their wall; (e) the septate bast-fibres, found in many
Orders; and lastly, U) the acicular fibres or rapbidines, occurrin6' in many Acantha-
ceae. The raphidines are fibrous cells. which are of small dimeUSlons in a transverse
section, and, like the raphides in a raphide-sac, are found.in large numbers in long
sac~shaped cells of the soft bast. from which .they are originally derived by cell-
division (Fig. 146. p. 619). Rod--cells, i.e. elongated parenchymatous cells, are
frequently developed in the bast in place of bast-fibres. Closed rings of stone-
cells have been observed only in the outermost portion of the bast of certain Logania-
ceae (species of Strychnos) and Asclepiadeae. The stnlcture of the soft bast shows
noteworthy features in Podophyllum (Berberideae), in which, as in the case of
Monocotyledons, it consists only of sieve-tubes and companion cells, and in almost
all the Gentianeae-Menyanthoideae, which have characteristic groups of small
sieve-tubes, these groups being equal in size to a single cell of the neighbouring
parenchyma. Sieve-tubes with wide lumina are found chiefly in lianes, while in
laticiferous and succulent plants the sieve-tubes are extremely narrow and insig·
nificant. A tendency towards a collenchymatous differentiation of the soft bast
has been recorded in the Caryophylleae and allied Orders, as well as in the Pitta.
sporeae and Plantagineae.
Stratification of the bast by means of tan~ential layers of c.hambered fi.bres
containi~g clustered crystals is found in c,ertain Combretaceae~ Alyrtaceae and
Lythraneae.
VI. ANOMALOUS STRUCTURE OF THE AxIS 1.
§ 54. CONTRACTED VASCULAR SYSTEMS of submerged plants and certain
other Dicotyledons, AND DISSOCIATION OF THE RING OF VASCULAR BUNDLES.
The vascular bundles. found in the different parts of the stem of submerged
plants t, fuse to form axile strands. in which the individual bundles are
gener.ally no .longer ~tinguishable from one ano~~er; the only known exception
to this rule-IS constituted by Ranunculus aquatilu. In those fonns (~peCles of
Peptis and Elatine), which have a type of structure not far removed from
that normally found in Dicotyledons, a pith is present within the ring of
wood and bast. while in Peptis, which belongs to an Order 1i.aving intraxylary
phloem, the latter is likewise found. In CaUitriche there is a reduced pith,
composed only of a small number of cells, which subsequently undergo resorp- .
tion. Hippuyis, MyriophyUum, Hottonia, Aldrovanda and CeratophyUutn do
not possess a true pith; the vascular system in these genera is concentric
1 The anomAlies presented by the atrncture of the root in part correspond to analogous feAtur~
in the·stem, and, when this iii the case, they will be discussed in common with the latter; for the
femaining (ea.ttlJ'eS, ~ • 6.... p. u68.
S H. Schenck, Vergl. ~nat. d. subm. Gew., Bibl. bot., Heft J, 1886,67 pp. and 10 Tab.

4E2
 CONCLUDING REMARKS

with central xylem. In Hippuris, Myriophyllum and Hottonia the xylem

of the concentric vascular strand consists of an apparent pith and of tracheae
with intenningled wood·parenchyma; the former is homologous with wood-
parenchyma, since the first tracheae, which subsequentlY become resorbed,
develop in the centre of this tissue. In Aldrovanda and CeratophyUum (Fig.
I~, p. 80:2) the xylem consists solely of pith-like wood·parenchyma, enclosing
a centJal air-canal, which arises by the resorption of a group of pnmary tracheae
(AUrovanda) or of procambial cells devOId of annular or spiral thickening
(CeratophyUum ).
Contratted vascular systems, taking the form of a concentric vascular
bundle with central xylem, are found also in an undetermined species of Lei-
pha,mos (a saprophytic member of the Gentianeae, See p. 550), in certain
Cuscuteae (parasitic fonns, see p. 573), and in Peperomia tenera, Miq., .and
Verhuellia (Piperaceae, the species in question being neither aquatics nor
parasites nor saprophytes).
Diasodatfon or the fibrovascular system and mutual independence of
the wood and bast has been observed among the Lentibularieae in Genlisea
(in the stem and peduncle), Polypompholyx (in the vegetative and floral axes)
and Ut,icula,ia (m the axis o~ inflorescence, Fig. 138, p. 595); the groups
of wood and bast in these genera are either irregularly scattered or show an
annular arrangement, the latter being the case in PolypomPholyx, in which
the groups of soft bast are placed in a ring and alternate with solitary vessels.
Here we may no doubt also include those cases, in which the vascular ring
contains incomplete bundles in the form of groups of soft bast or (Barlonia
verna), in addition to these, of groups of vessels side by side with complete
bundles (provided with xylem and phloem); this feature has been recorded
in certain Cuscuteae and in a saprophytic member of the Gentianeae (Barlonia
v~). We may finally mention the anomalous structure of the stem in the
anornalous genus Ci1Caeaster, which is appended to the Chloranthaceae in
Bentham and Hooker's Genera Plantarum; the delicate stem in this genus
resembles-the root in containing a diarch vascular system.
155. AXEs SHOWING POLYSTE:LIC STRUCTURE 1. Van Tieghem has intro-
duced the term 4 stele' to designate the ring of vascular bundles, found in the
axis of a normal Dicotyledon, together with the pith enveloped by the vascular
ring; so that the axis of Dicotyledons as a rule has a monostelic structure.
When the axis shows several steles in a transverse section, it is said to be
poI~telic. This structure arises in the following way. The vascular system
at the base of the axis is devoid of a pith; in its further course up the stem
it becomes band-shaped and divides by a median constriction into two systems ;
at a higher ;t?Oint in the axis the latter again become band-shaped and the
process of division is repeated, and so on. In an axis showing polystelic
structure the individual steles are pro\Tided with an endodermis and pericycle
of their own, While a pith mayor may not be present. In a transverse section
the steles are either irregularly scattered or show an antlu1ar arrangement p

while in the vertical direction they anastomose to form a network. In some

cases, moreover, they are fused in a transverse section so as to form a ring,
which is interrupted at certain points (gamostelic structure1 in contrast to the
dial~telic arrangement, in which the steles lie isolated in a trans"Verse section).
Whereas polystelic structure is very widely distributed in the axes of
I Van Tfeghem et Douliot, polysttme, Ann. sc. nat., iii, 1886, p. }7s. see .Iso Bull.
Set. i, t.
,&e. bot. de France, 1886, p. 213: Scott, in Ann. of Bot., Y. 1891,p. St.; S<:h()tlte, Sttla.rthe<lrie,
GroniageD and Jena, 190 2 , 175 pp.; [Worsdell, Stelar theory, in New Phytologist, 1903, I!p.I~+] ;
Schindler, in Engler, Bot. Jahrb., xuiv, 190.., Beibl. n, p. 67 et seq.; [Hill, Stelar theortet, Science
~r()~ i, 19OO, pp. aIS-i2].
 ANOl'rIALOUS STRUCTURE OF THE AXIS 1157
Vascular Cryptogams (among the Ferns and inSela.gineUa, seeLeclerc du Sablon,
in Ann. sc. nat., ser. 7, t. xi, 1890), it is rare among Dicotyledons. It has
been observed in the following cases: in Victoria regia (locally) and- in the
rhizome of certain species of Nymphaea (Nymphaeaceae); in the nodes of
the stem of Parnassia (Saxifragaceae); in the stolons of species of Gunnera
(Halorageae); in certain Primulaceae (species of Primuia, hr~longing to the
section Auricula, Fig. 115, p. 505, Bryocarpum and Dodecatheon) and Acan·
thaceae (species of Dianthera) 1; finally, also in the fruit-stalks of certain
Guttiferae, Ternstroemiaceae, Malvaceae, Sterculiaceae. Meliaceae and Mora-
ceae 2; regarding Pinguicula (Lentibularieae), see p. 59! 3.
A similar type of vascular structure, which, however, does not fall under
the heading of polystely (among other reasons because of the absence of an
endoderrnis around the individual bundles) and might perhaps be described
as apparent polystely, occurs in certain species of Leiphaimos (Gentianeae)
and Christisania (Orobanchaceae); in these genera concentric vascular bundles
with central xylem are found in the vascular ring. For a similar type of
structure, viz. divided xylem-masses, see § 60.
§ 56. MEDULLARY AND CORTICAL VASCULAR BUNDLES 4. A considerable
number of Dicotyledons depart from the ordinary type of structure, in the
fact that the vascular bundles are not arranged in a simple ring. In som~
of these plants there are medullary bundles or cortical bundles or both, in addi-
tion to the normal ring of vascular bundles; in other cases all the vascular
bundles of the axis are arranged in two or more rings, which are sometimes
not distinctly marked off from one another, or the bundles, as seen in: a trans-
verse section, lie irregularly scattered in the ground tissue. We cannot enter
into details here as to the nature (leaf-trace bundles or (rarely) cauline bundles)
and course of these vascular strands. From the systematic point of view the
features in question are for the most part characteristic of speCIes only and
rarely of genera, tribes, or even Orders.
Vascular bundles, which are irregularly scattered in a transver&e section of
the stem or are arranged in more or less distinctly marked rin~s, are found in the
follOwing genera and Orders respectively: Actaea, Cimicijuga, ThlJ'/ictrum (Ranun-
culaceae); Podophyllum (Fig. 10, p. 46). JeOersonla. Dip_hylteia, Leon/ice, etc., see
p. 821 (Berberideae); Nymphaeaceae (d. § 55); Papaver (Papaveraceae); Geranium
(Geraniaceae); Cucurbitaceae; UmbeUiferac; Cet'tfaurea, Scorzonera (Compositae) ;
Candollea (Candolleaceae); Limnanthemum (Gentianeae. in the axis of inflorescence
of the species belonging to the section Nympkaeanthe); Christi§onia. Con opkolis,
Epiphegus (Orobanchaceaeh MOut'era (Podostemaceae) j Hydnoya (Cytinaceae, in
the rhizoHi-shoots); Peperomia (Piperaceae, Fig. 169. p. 69;) ; Jl.fyzodendfon; Balano-
phoreae. See also the list of plants with medullary or Cortical vascular bundles.
The medullary vascular bundles, as seen in a transverse section, are, either
arranged in one or several rings or are irregularly scattered; it is very rare to find
only a single vascular bundle in the centre of the pith. The medullary bundles are
either collateral or concentric 6 in stru~tlJre and are occasionally reduced to small
1
- - =-~;;::"'==-.--------~-----
According to HOTm, B~t. Gaz. xliii, 1907. _
2 These anomalies ShOllld perhaps be included in the category oC 'divided xylem-masses' (see § 60).
3 In addition to mono- and polystelic strncture of the stem Van Tiegbem also distinguishes
'astelic' structure. The structure of the stem is said to be astelic, when the vascular system, found
at the base of the stem, becomes di~lved into individual vascular bundles at a higher level, these
bundles showing an annubr or scattered arrangement in the transverse section and each having
its own endoderm is and pericycle. Astelic structure has been demonstrated in: Ranunculaceae
(Anemotu, Eranthis, Oxygraphis, Ranunculm incl. Ficaria), Nymphaeaceae, Chelidtmium maius
(PApaveraceae), Byblis (Droseraceae, peduncle), and Trifolium (Papilionaceae).
, De Bary, VergI. Anat., 1877; J. E. Weiss, Markstandiges Gef:issbiindelsyst., Bot. Centralbl.,
1883, iii, p, 280 et seq.; Col, Faiscennx mcdullaires, Joom. de bot., xvi, J90 2• pp. 23+-55; Col,
Disposition des faisceaux, Ann. sc. nat., ser. 8, t. xx, 19°4, pp. 1-288 j Van Tieghem, MeristeIes
cort., Ann. sc. nat., sc!r. 9, l. i, 1905, pp. 33-44'
~ Mobius. in Ber. deutsch. bot. Gesellsch., 1887, p. 2 et seq
 I IsS CONCLUDING REMARKS
groups of soft bast, bundles of fibres, etc. Medullary bundles showing collateral
structure are generally normally orientated (i. e. their xylem lies on the inner, their
phloem on the outer side), but in some cases they are inversely orientated (i. e. the
phloem lies on the inner side). The concentric medullary bundles have either central
xyleIIl.or central phloem. The apparent bicollateral bundles of the pith are pro-
duced by the fUSlon of the contiguous xylem-groups of two collateral vascular
bundles. .
In most cas~ the medullary vascular bundles are of the nature of leaf-traces;
for dettils, see ]. E. Weiss, loco cit. The vascular bundles or bundles of soft bast.
which occur in the pith in members of Orders having intraxylary phloem and are
included in the following list, do not, strictly speaking, belong to this category
(see also § 57). Some of these medullary bundles are branches of the intraxylary
phloem, situated at the margin of the pith, the primary tracheae occasionally
(especially in the Melastomaceae) accompanying the ·phloem in its passage into
the pith; in other cases they arise from the intraxylarysoft bast. owing tothe fact
that the cambium of the latter not only produces bast internally, but also'woody
tissue externally, so that the bundles of soft bast at the margin of the pith become
transformed into inversely orientated vascular bundles. For the apparent medul~
lary vascular bundles, which occur in certain Nyctagineae, Amarantaceae and
Chenopodiaceae with anomalous structure, see § 59 (p. 1165).
Medullary vascular bundles have been observed in the following Orders and
genera respectively: Ranunculaceae (A nemone. Delphinium, Hydrastis, Glauc~·dium);
Cruciferae (rhizome of the kohl-rabi and of Cochlearia Al'moracia, concentric);
Vochysiaceae (in T"igoniastyum, inversely orientated,; phloem-bundles in other
members of the Order); Caryophylleae (A can th ophyllum , inversely orientated) ;
Sterculia.ceae (Leptonychia, concentric with central phloem); Luxemburgiaceae
(Godoyeae. composed of vessels and fibrous cells or of a phloem-group and fibrous
cells); Burseraceae (Canarium. inv. orient.); Olacineae (lodes, inv. orient.);
- Sapindaceae (Toulicia, Guioa, and Mischocarpus, often reduced to' bundles of
fibres) -; Meliantbaceae (Melianthus and Bersama, concentric with central phloem;
sometimes reduced to bundIes of fibres, Fig. 54, p. 241); Mimoseae (Elephantorrhiza,
concentric with central phloem); SaJdfragaceae (Saxi/raga. Rodgersia, Peltiphyllum);
Melastomaceae (very comm.on, in connexion with the intraxylary phloem; occa-
sionally reduced to phloem-bundles); OnaS!arieae (phloem-bundles, in connexion
with the intraxylary phloem) ; Halorageae (Gunnera, inv. orient.); Passifloraceae?
(Passiflora p); Cucurbitaceae (Coccinia, Cucufbita, Kedrostis, Melolhria, Wilbrandia,
mv. orient.. in connexion with the intraxyl. phl.-Siolmatra;inv. orient., here a~sing
subsequently on the inner side of originally collateral vascular bundles); Begoniaceae
(Begonia, sometimes concentr. with central phloem); Cacteae (Mammillaria, Echino-
&fI&tus, Echinopsis, Cereus); Umbelliferae (common, in partinv. orient., coneentr. with
central phloem or apparently bicollateral); Araliaceae (Aralia and Arthrophyllum
with varied orientation of wood and bast, Aralidium with bundles of fibres; for the
rest, see pp. 431, 946); Compositae (very widely distributed in the Cichoriaceae,
here in part reduced to phloem-bundles; rare in other Compositae); Candolleaceae
(Candollea, see above); Goodeniaceae (Gooden£a, SeJJiera, Disty/is, penetrating into
the normal ring of vascular bundles, Fig. 107, p. 472); Campanulaceae (phloem-
bundles common, in part at the margin of the pith; also inv. orient. and concentr.
vascular bundles. the latter with central pliloem); Plumbagineae (Sta#ce and
Acantholimon, coneentr. or inv. orient.); Apocrnaceae (in WillougJabeia and species
of Apocynum an inv. orient. ring of vascular bundles, arising from the intraxyl.
phloem; not uncommonly also medullary phloem-bundles, as branches of the intraxyl.
phloem); Asclepiadeae (PeYiploca with formation of wood from the cambium of the
mtra."xyl.phloem; medullary phloem-bundles. as in the Apocynaceae); Loganiaceae
(Anthoc/e'tsta with medullary vascular bundIes,-species of Gelsemz·um and SpigeUa
with inv. orient. vascular bundles, arising from the intraxyl. phloem,-in·addition tQ
that phlOem-bundles, as in the Apocynaceae) ; Gentianeae (Gentiana ·with medullary
vascular bundles or phloem~bund.1es,-in addition to that phloem-bundles, as in the
Apocynaceae) ; ..convolvulaceae (in A1'gyyeia, Erycibe. Evo/vulus, Hewittia, NeuYo-
peltis, Prevostea and SU&tocardia an inv. orient. ring of vascular bundles~ formed
from the intraxyl. phloem; normal medullary vascular bundles in Rivea); Solanaceae
(Anthotroche with a slight formation of wood from the cambium of the intraxyl.
phloem; medullary phloem-bundles, as in the Apocynaceae); Orobanchaceae
(Orobanche, Oistan&he); Gesneraceae (Klugia, Rhynchoglossutn. Monophyllaea, also
the tubers of Coleus); Bignoniaceae (Campsis~ inv. o.rient.); Acanthaceae (AcanthUS,
 ANOMALOUS STRUCTURE OF THE AXIS 1J59
Mendoncia, Pseudocalyx and Thunbe,gia, inv. orient.); Verbenaceae (Teiismannio.
dendron); Plantagineae (Plantago, eambiform- and phloem-bundles); Nyctagineae
(apparent and ? true medullary vascular bundles, common); Amarantaceae (true
and apparent medullary vascular bundles); Chenopodiaceae (as in the Amaranta.
ceae); Phytolaccace~ (Phyt~/acca, coneentr. with cent~al phlo~m); Poly~onaceae
(Rheum and Rumex, mv. onent., coneentr., etc.); Anstolochiaceae (A1'zstoloc;hia
t,iangularis, !,i~. 167, p. 686, h~re as a secondary formation,. which only ~evelops
after tf?e splitting up. of the n~g of vascular bundles); Piperaceae (pzpe, incL..
Hecke,za and Macyopzpe'l', see FIg. 169, p. 692); Thymelaeacea.e (Aqu£laYia with
secondary development of wood from the serial cambium of the intraxyl.. phloem) ;
MyzodendYDn (species of the section Eumyzodendyon, Fig. 178, p. 735); Balanopho-
reae; Euphorhlaceae (Ricinus); Cupuliferae (Alnus, cauline bundles, which are
only developed as an exception).
The cortical vascular bundles 1 generally have collateral structure, and are
then either normally or inversely orientated; cortical bundles with concentric
structure are rare, and the same is true of the reduction of cortical bundles to
phloem-bundles. Cortical bundles have been found in the following Orders or genera:
Ranunculaceae (Paeonia); Calycanthaceae (inv. orient. and showing secondary
growth, universally distributed, Fig. 3. p. 25); Crud ferae (Eruca. Lepidium);
Violarieae (all Sauvagesieae); Tremandreae (Tetratheca with winged band-shaped
axes); Stl'asburgeria; Dipterocarpeae (universally distributed) ; Lophiya; Monotes;
Rhaptopetalaceae (two cortical vascular bundles of quite general occurrence);
Lineae (Aneulophus, E1'yth"oxylon); Ochnaceae sense str. (universally distributed) ;
Luxemburgiaceae (universally distributed); Wallacea; Papilionaceae (species with
decurrent foliar wings on their axes; species of Cytisus, Genista and Retama
with reduced leaves and furrowed stems; B01'bonia and Viboygia; Vicieae);
Saxifragaceae (Peltiphyllum); Crassulaceae (Sempervivum, Sedum and Rochea.
coneentr. with central xylem); Droseraceae (Drosophyllum, inv. orient.); Lecythi-
daceae (inv. orient. in the Barringtonieae, normally orient. in the remaining members
of the Order); Melastomaceae (common); Turneraceae (Piriqueta); Begoniaceae;
Cacteae (Rhips~/is, Lepismium, Pftilfera, and Phyllocactus, forming a plexus and
showing growth in thickness); Ficoioeae (MesembYyanthemum, reticulate); Umbel-
liferae (Mulinum, Siler and Eyyngium, in the last genus forming contracted vascular
systems composed of several bundles, which have a certain similarity'to the peri-
pheral xylem-masses of certain Serjan£as); Araliaceae (O,eopanax and the anoma-
lous genus AYalidz'um, see p._ 946); Cornaceae (Mast£xia); Rubiacea~' (Sickingia) ;
Compositae (Achillea, Ammobium, Anthemis, Atractylis, Centau1'ea, Cynara, Gun-
delia. Hei!!nt"um, Madia, Senecio); Candolleaceae (Candollea ?, rightly included here?
see P.963t and § 58); Campanulaceae (species of Campanula, here concentt. and like
those of Eyyngium; Lobelia Rhynchopetalum with coneentr. cortical vascular bundles.
resembling those of a Cycas); Lennoaceae (AmmobYoma, Lennoa); Plumbagineae
(Aegialitis, Armeria, Limoniastyum, Statice); Primulaceae (' reseau radicifere' in
the root-stock of the species of Primula of the sections A u,icula and 0 fficinales
and of Bryocaypum and Dodecatheon); Asclepiadeae (rhizome of Tylophora);
Loganiaceae (Anthoclcz'sta, concentr.); Gentianeae (Menyanthoideae, viz. Limnan·
themum. Menyanthes and Villarsia); Lentibularieae (root-stock of Pinguicula with a
• reseau radicifere '); Gesneraceae (Rhynchoglossum); Verbenaceae (species with
winged stems); Plantagineae (Plantago, cambiform-bundles and vascular bundles) ;
Chenopodiaceae (Salsoleae and ~alicornieae. in part normally, in part inv. orient.) ;
Polygonaceae (Calligonum, Fig. 162, p. 673; Rumex, here in the pericycIe);
Nepenthaceae (Nepenthes); Penaeaceae (angles of the stem of Endonema); Bala·
nophoreae (see p. 738) ; Euphorbiaceae (Euphorbia); Buxaceae (species of Buxus,
belonging to the sections Eubuxus and Notobuxus); Casuarineae (Fig. 186, p. 788).
For the occurrence of sieve-tubes, which are situated outside the limits ot the
vascular system, in the Cucurbitaceae, see p. 396.
§ 57. INTRAXYLARY PHLOEM 2. The term' intraxylary phloem' is here
1 For their course, see especially De Bary and J. E. Weiss, 11. C~.
~ Petersen, Bicoll. Gef:i.ssb., in Engler, Bot. Jahrb., iii, 1882, pp. ~S9-400 and Tab. iv':'viii, see
also Bicoll. Karbundt, etc., Diss." Copenhagen, 1882 i J. E. Weiss, Markst. GeraSsbiinde1syste~,
Bot. Centralbl., 1883, iii; Solereder, Holzstr., 1885, p. 27 j Herail, in Ann .. sc. nat., ser. 7, t.ll,
1885, p. 267; Lamonnette, in Ann. sc. nat., ser. 7, t. x, 1890, p. 193 i Scott and B.rebne~. in Ann: of
Bot., v, 1891, p. 259; Van Tieghem, in Jouro. de bot., 1891, p. IJ7; Perrot, TlSSU cnbIe, These,
 1160 CONCLUDING REMARKS

taken in a general sense to include those groups of soft bast, which are situated
internally to the ring of vascular bundles; in their further course these phloem~
groups bend out into the leaves together with the leaf·trace bundles, on the
inner side of which they are situated, so that it is possible to trace them in
a corresponding position into the petiole and commonly also into the larger
or even the finer veins of the leaf. The groups of internal phloem, moreovef,
I appear to be characterited by the fact that they are generally differentiated
only at a slightly later stage than the outer phloem-groups. .
As a general rule the course of the bundles of iotraxylary phloem serves
to distinJUish them sharply from medullary vascular bundles and medullary
phloem-bundles. The medullary phloem-bundles, which are found in the
Cichoriaceae and Lobeliaceae and which, in the absence of an investigation
of their course, might be regarded as intraxylary phloem, are of an essentially
different nature; they belong to the normally constructed ring of vascular
bundles from which groups of phloem bend out secondarily into the pith 1.
The differentiation of the intraxylary phloem varies. If either forms
a ring of tissue at the margin of tbe pIth, Of appears in the shape of isolated
bundles, which vary very much in size (massive or very small) and lie opposite
the primary portions of the vas_cular bundles. In tlle latter case the intraxylary
phloem is eithor closely apposed to the xylem of the vascular ring, the bundles
of which are then said to be bicollateral, or it is separated from the xylem
by one or more layers of parenchymatous cells. ,Portions of the intraxylary
phloem may be given off as branches, and these,strands of phloem are occa-
sionally (e.g. in many Melastomaeeae) accompanied by the primary ·tracheae,
su that phloem-bundles or vascular bundles appear in the middle of the pith
(d. § 56, p. 1158). Hard bast is frequently present at the inner margin of the
internal phloem. In many cases, moreover, a cambium develops at the outer
boundary of the intraxylary phloem: this cambium produces secondary soft
bast on its inner side and occasionally leads to such an extensive increase of
the intraxylary phloem that the innermost portions of the latter become com-
pressed so as to resemble horn-bast. In-a few cases (in certain Cucurbitaceae,
Apocynaceae, Asclepiadeae, Loganiaceae, Convolv--ulaceae, Solanaceae, Thyme-
laeaceae; regardmg this point d. § 56, p. 1158) this cambium produces xylem
on its outer side, so that medullary vascular bundles with inverse orientation
of the wood and bast are formed; these bundles must be distinguished from
the medullary vascular bundles of Campsis rad£cans, &c. (see § 56).
Intraxylary phloem is found in the following Orders, being constantly present as
a rule in those indicated by a"': *VQchysiaceae (excl. Trigoniaceae), *Combrctaceae
(excl. Macropteranthes and the Gyrocarpeae), *Myrtaceae sens. str., *Melastomaceae,
*Lythrarieae (incl. Punica and OUnia), *Onagrarieae (incl. Trapa), ·Cucurbitaceae
(excl ..the tri~e~ Gynostemmeae, Zanonieae ? and Feuilleae; tbe.intraxylary phloem
sometimes anslng only at a late stage), ·Apocynaceae, "'As clepladeae , Loganiaceae

Paris, 1899, p. 167; Baranetzky, Faisc. bicoll., Ann. sc. nat... Str. 8, t. xii, 1900, pp. 26r-321 and
pI. Tii-x; Viret, Liaisons du phlol-me 10M., perimed. et interligneux avec Ie liber normal, Travaux
last. but. Univ. Geneve, ser.6, Fasc. vi, 1904, 100 pp.
1 The delimitAtion of what is known as • intraxylary phloem from tbe groups of meduUary soft
I

but, discussed in the previous ~ction. has not yet been quite finally accomplished, owing to the fact
that the coarse of the intraxylary soft bast. in the various Orders in wbich it occurs, has not by any
m~ans been sufficiently im.. estignted up to the present. In the case of some of the Thymelaeaceae
Van Tiegbem has shown that the groups of intraxylary $oft bast do not pass out into the leaf. The
necessity of investigating the course of these stra.ndll of phloem ts ~own by the cases (Camj(utula
irf'tu1n'th:lb and Locluca jertnm's), mentioned by J. E. Weiss; h~e one finds medullary phloem-
oondln. which, Ib; shown by their course, correspond in part to intruylary phloem and in 'Part to
the ordinary medullary phloem-bundles of the Campnno1:tceae and Cichoriaceae (see above). In
the su~aent lilt of Orden having intraxylary phloem these two cases are lIot further considereo for
the sake of convenience.
 ANOMALOUS STRUCTURE OF THE AXiS JI61

('!Loganioideae), *Gentianeae (excl. ~renyantheae, for the apparent exception' Lei-

phaimos,' see p. 994), "Convolvulaceae (incl. Nolaneae and excl. Humberlia and
the Cuscuteae). ·Solanaceae (excl. Retzia). Acanthaceae ("'Barlerieae. Nelsonia),
Myoporineae (Ojtia). "'Basellaceae (correctly, included here ?, intraxylary phloem
namely developing at a late stage), Polygonaceae (Emex, etc., really intraxylary
phloem ?), .$Thymelaeaceae (excl. Drapetes), *Penaeaceae, Euphorbtaceae (·Eucro~
toDeae excl. species of Croton of the section Astraea; Alchornea. Conceveiba, Lepi.
doturus, Pera, M altotus pro parte; Dactylostemon, M abea, Sene/eldera, Sebastiania
pro parte).
The great systematic value of the intraxylary phloem is sufficiently evident
from the preceding table. Thus, it often constitutes an ordinal character; it is fre·
quently found in groups of allied Orders; the Scrophularineae and Solanaceae are
sharply distinguished by the absence and presence respectively of intraxylary
phloem. &c.
We may add here that intraxylary phloem only rarely occurs in the
root (in certain species of Oenothera, Vinca, Strychnos and Chironia, and in
those Cucurbitaceae, which have a pith). In most cases we· find the internal
phloem passing to the outside (simultaneously with the primary vessels) already
in the region of the bypocotyl, the intraxylary phloem then entering into con-
nexion with the outer phloem of the root. In other cases the intraxylary
phloem is continued into the root. as interxylary phloem (e. g. in species of
Oenothe~, d. § 58, p. 1162)

§ 58. INTERXYLARY PHLOEM 1. The term • mterxylary phloem ~ is used

to denote groups of soft bast, which arc enclosed in the wood of the vascular
ring. In the previous summary (see I Systematische Anatomie,' German
Edition, r899, pp. 972 et seq.) we followed the example of H. ·Schenck in dis-
tinguishing three types, but in view of Leisering's researches these are better
comprised in the following two types. In the first type, which may retain
the earlier nam~ of • Strychnos-type: the groups of interxylary soft bast are
given off by the cambium on its outer side in a perfectly normal manner, as is
generally the case with the soft bast; subsequently, however, these groups
Qf phloem become bridged over by a cambial are, which develops itt the peri-
cycle on their outer side; as a result of the activity of this cambial arc, which
produces wood internally and bast externally in the ordinary way, the groups
of interxylary phloem become sunk in _the xylem·mass. A modification of
this type is found in the cases mentioned below as occurring in certain
Acanthaceae, Apocynaceae and Bignoniaceae j here the groups of soft bast
(which playa part in the formation of the furrows (see § 6r) in the xylem) are
not embedded in the wood by the agency of new cambial arcs, but secondarily
by the production of tissue formed from a proliferation of the strips of cambium,
~ituated on either side of the furrows; a modification of the first type is like-
wise .constituted by the anomaly, found in CUSC1tta iaponica (see p. lOO4}. In
the second type~ which includes both the other cases previously distinguished,
the islands·_·of _intetxylary soft oast arise by the subsequent differentiation of
in
sieve-tubes groups of thin-walled tissue, which are given off by the cambium
on its inner side, together with the secondary wood. These groups of tissue
are c.omposed of cells, which at first have the character of unlignified elements
of the wood-parenchyma. Their transformation into phloem may commence
at a rather early stage, viz. immediately after they are produced by the cam-
bium, in which case the groups of interxylary phloem may be said to be
1 Solereder, Holzstr., 1885, p. 3z; lUrail, in Ann. sc. nat., ser. 7, t. ii, 1885, p. 256;
Scott and Brebner, in Ann. of bot., iii, 1889, p. 275; Van Tieghem, in Journ. de bot•• J891 ;
Chodat, in Arch. sc. phys. et nat. Geneve, 1892, p. 229, nod Report of ConJ,,'Tess, Genoa, 1893,
p. 1.4-4; H. ~chenck, Annt. d. Lianen, 1893. p. 17 i Perrot, Tissu cribte, These, Pads, 18 99, p. 177 ;
Leiseriog, Entwicklungsgeschichte des interxyl. Leptoms bei den Dicotyl., Diss., Berlin, 1899,
.:,0 pp.• :2 Tab., also in Bot. Centralbt., 1899, iv.
 CONCLUDING REMARKS

given off by the cambium on its inner side (i. e. the Mucuna-type previously
distinguished), or it may be postponed until a relatively late stage (this being
the case formerly distinguished as 'development of interxylary phloem by the
subsequent differentiation of sieve-tubes in islands of unlignified tissue, which
are situated in the xylem-mass '). These two cases are connected with one
another by transitions, and it is only the extreme forms that can be properly
distinguished from each other.
It may once more be emphasized that the two types of interxylaiy phloem
above described are perfectly different structures. Practical considerations.
alone lead me to discuss them m the same paragraph and to class them together
as interxylary phloem, since it is not always possible to determine their course
of development in the material, which suffices for the mere demonstration
of the anatomical features in question; moreover, the development of the
interxylary phloem has not been investigated in all cases. Axes having inter-
xylary phloem of the Strychnos-type are very closely related to the axes show-
ing the anomaly discussed in § 59, in which successive annular or arc-shaped
I

groups of wood and bast arise in the pericycle. The only essential difference
between the two types of anomalies lies in the fact that in Strychnos, &c., the
successive extinction and renewal of the cambium is confined to very small
portions of the original cambial rings, while in the second case.it affects the
entire cambial ring or, at least, relatively extensive arcs of cambium. The-
occurrence of transitional forms between the two 'kinds of anomalies- (see
Hippocrateaceae) is in agreement with this close relationship.
Considered from the systematic point of ,view, the occurrence of interxylary
phloem is mostly constant throughout all the species of a genus. It generally
appears already in the first-formed mass of secondary wood, and is consequently
developed in large quantity in the branches of herbarium-material. In other
species, however, it arises only at a late stage and in such a case is not to be
found in herbarium·material. It follows that if interxylary phloem can be
demonstrated only in some of the species of a genus, when herbarium-material
is being examined, it need not necessarily be absent in the remaining species.
Judging by the experience gained in the investigation of the species of Strychnos
(see p. 546), it is far more probable that interxylary phloem occurs also in
those species, in which it is not developed in herbarium-material, but that
it arises only in the later growth of secondary wood. Interxylary phloem
is particularly widely distributed in genera belonging to Orders, which are
distinguished by the possession of intraxylary phloem. The species provided
with interxylary phloem mayor may not be lianes. In those Orders, in which
several genera have interxylary phloem, its mode of origin, as far as the
hitherto published investigations show, is generally the same in all cases.
The islands of interxylary soft bast vary in size, sometimes (e.g. in many
Barlerieae) even being very small; in a transverse section they are (mostly)
rounded or (more rarely) elongated in the form of tangential bands and are
commonly arranged in concentric rings. The groups of interxylary soft bast,
found in Condylocarpon (Apocynaceae, Fig. 123, p. 533) and certain Bignonia-
ceae, are peculiar in being elongated in the radial direction; the development
of phloem-islands in these cases is connected with the formation of furrows
in the wood (see above, and pp. 533 and 605). Bast-fibres are rarely (Aqui-
larieae) found in the groups of interxylary phloem.
Interxylary phloem occurs not only in the stem, but also in the root.
The development of the interxylary phloem in the latter conforms to the same
two types, as have been distinguished in the stem. In some cases the islands
of soft bast found in the wood of the root appear as downward prolongations
of the strands of intraxylary phloem occurring in the stem (d. § 57, p. 1161) .
•
Interxylary phloem has been found in the stem or root, or in both, in the follow-
 ANOMALOUS STRUCTURE OF THE AXIS u63
ing Orders and genera respectivelyl: Vochysiaceae (E,,"sma, subseq. diff.); H~ri­
cineae (Endodesmia); Malpighiaceae (Dicella and Stigmaphytton, subseq. diil.);
Olacineae (Sarcostigma, Str.-type; Chlamydocarya); Hippocrateaceae (Sa/acia,
Str.-type, Fig. 51, p. 217); Papilionaceae (Dioclea, Mucuna and Phaseolus, subseq.
diff.); Mimoseae (Entada, subseq. diff.); Combretaceae (Calycopteris, Cambretu",
and Guiera, Str•.type; Thiloa); Melastornaceae (Memecylon. Mouriria. KJ"besn"a,
Pternandra, Str.-type ; wood of the root of Memecylon); Lythrarieae (wood of thE'
root of Lythrum, subseq. diff.); Onagrarieae (wood of the stem and root Qf Oenothera
and Epilobium, and wood of the root of Gaura and Lopezia. probabty always subseq.
diff.); Cucurbitaceae (wood of the root of Cucurln;ta and ThladJ"antha. subseq. diff.,
in part as a continuation of the intraxylary phloem of the stem) ; Candolleaceae?
(Candollea, really interxylary phloem ?, see p. 963 and § 56 under cortical bundles ');
4

Plumbagineae (m the anomalous growth of the wood bf Aegl,"al";#s, d. § 59); Salvado-

raceae (Salvado1'a, s ubseq. iliff. ; Dobera, Platymitium ; wood of the root of Salvadora);
Apocynaceae (Condylocarp,on, in connexion with the formation of furrows in the
wood; Lyonsia, subseq. dIff.); Asclepiadeae (' Ceropegia macrocarpa,' subseq. diff.;
lower part of the stem and root of Asclep~"as obtusi/olia, Michx., subseq. diff., in
the root also as a continuation of the intraxylary fhloem of the stem; wood of
the root of Asclepias syriaca, subseq. diff.; wood 0 the root of MOffenia b1'acny-
stephana); Loganiaceae (Strychnos, Str.-type; Antonia, Norrisia, Bonyunia,
Logania); Gentianeae (Chi,.onia, Craw/urdia, Ixanthus and Orphium; wood of the
root of very many Gentianoideae, see p. 995, probably in all cases subseq. diff.);
Convolvulaceae (hypocotyl of Ipomoea versicolor, subseq. diff.) ; Cuscuteae (Cuscuta
iaponica, see p. 1004); Solanaceae (wood of the root of Atropa, Datura and Scopolia.
subseq. iliH. ; wood of the root of Browallia); Bignoniaceae (embedded bast-wedges
in Distictis,Haplolophium? and PUhecoctenium); Acanthaceae (Barlerieae, in BarlBria
subseq. iliff.; Thunberg";a, subseq. diff., Fig. 147, p. 622, sometimes developing from
wedges of .soft bast); Thymelaeaceae (Aquilarla, Gyrinops, Gyrinops~"s and Litw-
stoma, subseq. diff.; Brachythalamus, Lophostoma, Synaptolepis); Loranthaceae
(Nuytsia, Str.-type); Euphorbiaceae (Dalechampia. subseq. diff.).
In connexion with the groups of interxylary soft bast of the second type
(i. e. those arising secondarily from groups of unlignified wood-parenchyma),
we may notice the occurrence of interxylary vascular bundles, showing con~
centric structure with central phloem, in the rhizome of Cochlea ria A,moTacia
and in the roots of certain other Cruciferae (J. E. Weiss). These bundles take
their origin from groups of phloem, which develop secondarily in the wood·
parenchyma and subsequently become enveloped by a ring of cambium, which
gives off wood on its outer side.
. ./
§ 59- SUCCESSIVE DEVELOPMENT OF SECONDARY GROUPS OF WOOD AND
BAST 2. The development of successive secondary groups of wood and bast
has been observed both in lianes and in non·lianes and is either characteristic
of species only or serves to distinguish genera or even Orders. The diverse
appearance presented by the axes,_ which show this anomaly, depends on the
tIme at which the latter appears, a.§ well as on the place of origin, the shape
and the mode of differentiation of the secondary groups of wood and bast.
The formation of the secondary groups sometimes takes place at a very early
period, while in other cases they only arise at a very late stage; it is only in the
former case that" the anomaly can be demonstrated in herbarium·material.
The secondary groups of wood and bast either form (a) complete rings (successive
zones of growth)~ which are arranged concentrically with reference to the ring
of primary vascular bundles, or (b) segments of such rings, or (c) mere strands,
which occasionally project towards the exterior in the form of weals or ribs.
1 Unless there is a statement to the contrary. these data refer to the occurrence of interxylary
phloem in the stem; the abbreviations' Str.-type· (i•. e. Strychnos-type) and' subseq" diff.· (i. e.
subsequent differentiation-at an earlier or later stage-of phloem from groups of thin-walled tissue,
given off by the cambium on its inner side) are used to indicate the two chief types. For details as
to course of development, see the descriptions of the individual Orders"
2 See especially H" Schenck's • Anatomic der Lianen • (1893). on which the description of the
anomalously constructed axes in the special part of this book is altogether mainly based.
 CONCLUDING REMARKS

In addition to that, the secondary rings of wood and bast or segments of wood
and bast, as in the case of the primary vascular ring, either form closed zones
traversed only by narrow medullary rays, or are split up into so·called secondary
vascular bundles by broad plate·like medullary ra~, which mayor may cot
be lignified. I shall return below to certain specIal cases presented by this
anomaly. The place of origin of the secondary groups of wood and bast
varies. The secondary meristems, from which they arise, originate either in
the inner cell-layers of the primary cortex (occasionally in the ,innermost
(endodermaI) layer of the latter) or in the parenchymatous pericycle, or in the
phloem of the original vascular ring. As a matter of practical convenience
the last two modes of development are not kept distinct in the following
enumeration ot Orders and genera, which show this anomaly, since it is otten
enough (and especially in the case of herbarium-material) difficult to determine,
whether the secondary structures develop from the pericycle or from the outer·
most cell-layers of the soft bast. Nor is it always easy to settle, whether 'the
pericycle or the inner layers of the primary cortex are the seat of the anomalous
growth., this being the case when peri cyclic sclerenchyma or a distinctly diffe-
rentiated endodennis are absent.
The point of origin of the secondary structures is generally constant
throughout all the species 'of a. genus, but this is not always the case (see
Forchhamme,ia and SPatlwZobus); in the same Order it may vary from genus
to genus, or may be identical in all the species shoWIng anomalous structure.
The development of secondary groups of wood· and bast from ·meristems,
which arise in a distinct extrafascicular position in ilie primary cortex, has been
obseryed in the following Orders and gen«;ra respectively 1: Menispermaceae
(Abuta, Anomospermum, FIg. 8, f'
43. Chondrodendron. Cocculus, Pachygone, Pe"i-
campylus j also in the root 0 A buta. Chondrodendron, Cocculus, Cissampelos
and Clypea); Capparideae (Cadaba, FOfchhatnmeria); Ca,ryophyUeae (in the root or
also in the stem of Spergularia and Polycarpaea, Fig. 25. J>. 108 j here we may
probably include also the anomaly found in the root of Cefd,a, Orlegia, Polycarpo'll.
Pycnophyllum, Silene l , Spet'gula and St,·pulicida); Papilionaceae (Defr~·s, Rhynchosia.
Spatholobus, Wistaria); Cucurbitaceae (Ac#nostemma, Momof'dica); Umbelliferae
(E,yngium); Compositae (Coreopsis); Plumbagineae (AegiaUNs, see § 58, p. U63) j
Verbenaceae (A vicennia)~
The development of secondary groups of wood and bast from meristems.
which arise in the parenchymatous pericycle or (more rarely) in the bast of the
original vascular ring, has been observed in the following Orders and genera re·
spectively: Dilleniaceae (Doliocaypus); Capparideae tForchhammeria. Maerua,
Roydsia); Polygaleae (B,edemeyera, Co mespet'ma , Moutabea, Securidaca, Fig. 23.
p. 991; Olacineae (Phytocrene with successive rings, consisting of plates of wood
and bast. Fig. 49. p. 208; T,ematospcy_ma, undetermined member of the tribe
Icacineae); Hippocrateaceae (Salacia with transitions to interxylary phloem,
see § 58, and Fig. 5 I, p. 217); Rhamneae? (Gouania?); Ampelidaceae (Tetra-
stigma), Sapindaceae (corded xylem-mass in PaulUnia, Serjania and Thino",a,
Fig. 53. p. 235); Papilionaceae. (Derris I, Macnaerium, Mucuna, Pacnyr}u·zus.
Puera ria , Spatholobus, Stt'Dngylodon); Caesalpinieae (Ba~hinia); Cucurbitaceae
(only in the root of Ecballium elate,.ium); Ficoideae (as far as is known. in all the
woody genera except Polpoda, also in the root); Rubiaceae (Chiococca); Compositae
(Mikan'ia, Kleinia. Dahlia); Candolleaceae ? (Candollea, really belonging to this
category ?) j Plumbagiileae (Acantholimon); Convolvulaceae (AYgyt'eia, Calonyction,
Hewitti,,?, Ipomoea, Maripa, Me rl'emia , Parana, Fig. 130, p. 571, Rivea); Bigno-
niaceae (Callichlamys, Distictis, Doxantha, Glaziovia, Haplolophium); Labiatae
(Thymus); Nycta&ineae (quite generally in the woody species, Fig. 155, p. 648;
1 The above data as a rule refer to the stem, except when there is a statement to the contrary; .in
those species, in which the anomaly occurs in the stem, it is, however, very frequently found also in the
mot (for details on this point, see thedjagnoses of the individual Orden, and Schenck,loc. cit., pp. 251-6).
~ In Silm~ acaulis, L. according to my own investigation.
S Derris is included also at thi£ point on th~ authority of 1{. Schenck (loc. ctt' l p. 176),
who mentions the • secondary cortex' 3.3 the seat 'of the formntion of the anomalous growth.
 ANOMALOUS STRUCTURE OF THE AXIS 116s
also in tbe root); Illecebraceae (Pollichia; here we may probably include the
anomaly found in the stem of Cometes and CO"iliola, and in the root of Aca'H,hony-
chiao Achy"onychia. Cometes. Co"",,'giola, Dysphan~a, Haya, and Pollichia); Amaranta-
ceae (of almost general occurrence in the woody species, Fig, 157, p. 654, also in the
root; wanting in Achatoca1'pus and Cladoth"ix); Chenopodiaceae (quite generally
in the woody species, also in the root); Phytolaccaceae (Agdestis, EfCilla, Gallesia.
Phytolacca, Segui"ia, in A gdestis and Pkytolacca also in the root; here we may prob-
ably also include the anomaly found in the stem of Barbeuia. Pdiveria and Rivina, and
in the root of Ani.wmeria); Polygonaceae (Antigonon); Loranthaceae (Lo1'an'hus;
Nuytsia.see § 58); Euphorbiaceae (Dalechampia and Fraga"iopsis. secondary strands
of wood and bast, appearing like ribs); Buxaceae (Simmonasia, also in the root).
The place of ongin of the secondary groups of wood and bast in the axis still
remain!] to be determined in Dalbe1'gia (Papilionaceae). E"ycibe (Convolvulaceae.
here developed in connexion with a cleavage of the xylem.mass), and in some of the
genera of the Caryophylleae, Illecebraceae and Phytolaccaceae already enumerated
under their respective Orders in the ,two previous paragraphs.
We may lastly add, that- according to Maheu, the anomalous growth in the
Menispermaceae shows a varied mode of development, and is found also in
Cluumantlera, 'Menispe1'mum,' and Stephania (see p. 818); Maheu's statements,
however, still require to be verified.
The structure of the anomalous growth deserves special mentloh in the
case of the Sapindaceae, Chiococca, Aegialitis, Phytoaene, andin certain Ficoideae,
Nyctagineae, Amarantaceae and Chenopodiaceae. The secondary masses of
wood and bast, which give rise to the corded xylem-mass of the Sapindaceae
(Fig. 53. p. 235), are always annular and completely enveloped by cambium,
but apart from that they are either cylindrical or flattened. They are related
to the strands of wood and bast, which are found in Chiococca (Fig. 102, p. 453)
and which are likewise provided with a ring of cambium. Aegialitis is distin·
guished by the fact that only one secondary meristem is formed; this produces a
ring of wood including large groups of interxylary soft bast on its inner side, but
there is no formation of bast on the outer side. The successive rings of growth
found in Phytoc1'ene are peculiar, owing to the fact that they consist of grou~
of wood and plates of bast, which alternate with one ancther in the tangential
direction (see Fig. 49, p. 208). In many Ficoideae, Nyctagineae (Fig. ISS,
p. 648), Amarantaceae (Fig. 157, p. 654), and Chenopodiaceae, lastly, the
secondary bundles of wood and bast are enveloped by prosenchymatous inter-
fascicular tissue, which is likewise formed by the secondary meristems; this
tissue is sometimes traversed by medullary rays or may include vessels. In
the Nyctagineae, Amarantaceae, and Chenopodiaceae, the ground tissue
situated between the primary vascular bundles and the tissue formed at the
commencement of the activity of the secondary meristem, is occasionally
difierentiated like a pith, and in such cases the primary vascular bundles appear
as apparent medullary bundles (see § 56).
As regards the composition of the secondary strands of wood and bast we
may a4~ that primary tracheae with spiral and annular thic.kenings, such as
occur quite generallY'in the leaf·trace bundles, are almost invariably wanting
in the secondary strands.
160. COMPOUND AND DIVIDED XVLEM·MASSES 1. Both types of anomalies
depend on a special arrangement of the vascular bundles at the time of their
development. In the case of the compound xylem,:" mass, which occurs in
certain species of Seriania (Fig. 53, p. 235) and Paullinia, and ,in ,an undeter-
mined member of the Leguminosae (Fig. 62, p. 281), one finds in"addition to
a central ring of vascular bundles a number of peripheral rings of wood and bast;
each of the latter includes a pith, is provided with primary spiral tracheae and
grows in thickness by means of a ring of cambium. The divided xylem-mass
1 Cf. Radlkofer', papen cited on p. 236t and H, Schenck, Anat, d. Lianen, 1893. p. 81 et teq.
 I I 66 CONCLUDING REMARKS

(Fig. 53, p. 235) lacks the central ring of vascular bundles found in the case
of the compound xylem-mass. It takes its origin from a rmg of horseshoe-
shaped groups of vascular bundles, each of these groups being open on its
inner side and having a pith, which in early stages still communicates with the
central pith; subsequently, however, the rings become closed. A divided
xylem-mass is found only in Serjania cOTTlegata, Radlk., and a few allied species.
For details, see p. 234.
§ 61. UNEQUAL THICKENING OF THE XYLEM-MASS 1. Among the axes
showing anomalous growth it is usual to include a number of structures, which
appear in certam lianes (provided with a normal ring of vascular bundles)
in the course of secondary growth, and which are generally to be found only
in axes 9f a certain degree of thickness (only rarely occurring in the. branches-
of herbarium-material). These anomalies are due either to unequal growth in
thickness of the wood alone or both of the wood and bast. In the former case,
according to the manner in which the growth of the wood is accelerated, we
obtain axes, which are flattened or band-shaped, or are provided with superficial
furrows of'varying breadth (ribbed or winged axes). In the case of strongly
winged stems, as may be noticed in p~sing, the anomaly occasionally (Sabicea,
Lantana) leads to an ultimate cleavage into longitudinal segments, correspond-
ing to the wings. When both the wood and bast show irregular growth in
thickness, the irregularity is not noticeable-or only faintly indicated-on the
outer surface of the axis, owing to the fact that at those points at which
the formation of the wood is retarded, there is a corresponding increase in
the production of. bast on the part of the cambium. In this way the xyle~
mass, as seen in a transverse section, has a lobed outline, the furrows between
the lobes being filled with phloem in the shape of bast-wedges. The ring of
cambium in these cases is either continuous, so that it forms a complete lining
to the furrows, or interrupted (Bignonieous type according to H. Schenck I).
the cambial tissue being confined to the outer side of the projecting portions of
the wood and to the base of the furrows (i.e. it is absent on the radial surfaces
of the bast-wedges).
Flattened or band-shaped axes have been recorded in the following Orders
and genera respectively: Papilionaceae (Abrus, Machaerium, Rhynchosia) ; Caesal-
pinieae (Bauh-inia, Fig.. 64, p. 290); Asclepiadeae (Cef'opegia, lbatia. e~); Conw
volvulaceae (Merremz·a); Polygonaceae (Coccoloba); Ulmaceae (Celtis). Ribbed-
or winged axes', the outer surface of which is furrowed, ha.ve been observed in:
Malpighiaceae (Hetcropteris, Fig. 37. p. 166); Celastrineae (Euonymus, Celastrus);
Sapmdaceae (Seriania, Urvillea); Caesalpinieae (Bauhinia, Fig. 64, p. 290, Cassia) ;
Mimoseae (AcacIa, Piptadenia); Rubiaceae (Sabicea); Verbenaceae (LantalUl);
Polygonaceae (Atraphaxis, ~nghtl.y included in this category ?). Axes with but:
wedges 'are found in: Anonaceae (Melodorum); MalJ?ighiaceae (Hetef'opteris, Fig. 37.
f. 166, P~ixotoa. Tet!,aptef'is); Chailletiact;ae (Chaitlella); Phytocreneae (*Phytocrene,
Pyrenacantha); Hlppocrateacea~ (spec. mdet.); Rubiaceae (Lygodysodea, Maneltia,
Sabicea); Compositae (*Bidens, ·Mikania); Apocynaceae (Al/amanda, Alstonia,
Condylocarpon, Fig. 123, p. 533, Echites, Lyonsia, Parsonsia, Tabernaemontana);
Asclepiadeae (Gymnema); Boragineae (Tourne/ortt"a); Convolvulaceae (MeYremia);
Bignoniaceae (numerous ·Bignonieae with bast-wedges, which in transverse section
are separated from the xylem-mass by lines, which are straight or resemble a stair-
case, Pandorea) ; Acanthaceae (A /romendoncia, M endoncia, Pseudocalyx, Thunb~rgia).
In some of the cases, named in connexion with the last of these anomalies. the
suhsequent cleavage of the xylem-mass is initiated by the appearance of the bast-

J See H. SchenckJ Anat. der Lianen, 1893, p. 15.

I The formation offurrows and wings in plants with reduced leaves (formation of 'micropteres'
and • macroptere&' of Briquet) does not belong here and is therefore not considered at this point.
S The occurrence of but-wedges of the Bignonieous type is indicated by a * in the following
synopsis.
 ANOMALOUS STRUCTURE OF THE AXiS

wedges (see § 62); in Phytocrene (see § 59), moreover, t,he development of successive
zones of wood and bast having a characteristic structure, is combined with the
anomaly in question; in a few cases (Gondylocarpon and certain Bignonieae), finally,
bast-wedges, which are formed successively in the radial direction, subsequently
come to be enclosed in the xylem-mass (cf. § 58, p. 1162).
§ 62. CLEAVAGE OF THE XYLEM-MASS I, Very complicated structures are
produced by the cleavage of the xylem-mass of the stem. It commences
with the splitting up of the ring of wood into a number of separate strands by
a -process of dilatation, i. e. ceU-division in the parenchyma of the wood, the
pith and the medullary rays. Subsequently meristems, which produce wood
and bast, are frequently developed in connexion with these strands.
A clen xylem·mass has been found in the following Orders and genera respec-
tively: Caryophylleae (Acanthophyllum); Ma.lpiglUaceae (Banisteria, Mascagnia,
Mena. Stigmapkytlon, Tetraj:Jterys, Fig. 37, p. 1(6) ; Sapindaceae (Seriania, Urvillea) ;
Caesalpinieae (Baukinia, Fig. 64, p. 290); Umbelliferae (Azore/la); Asclepiadeae
(Galotropis); Convolvulaceae (El'ycibe, Merl'emia); Bignoniaceae (B'gno-nia,
Fig. 142, p. 606, Doxantha, Dolichandra It Macfadyena. Melloa. Parabignon'ia I) ;
Acanthaceae (Afromendoncia, Mendonda. Pseudocalyx!. Thunberg,'a); Aristolochia-
ceae (Aris/olockia, Fig. 167. p. 686).

VII. STRUCTURE OF THE ROOT.

§ 63. GENERAL STRUCTURE OF THE ROOT 2. The structure of the root

in the Orders of Dicotyledons has not yet been methodically investigated to
any consid~rable extent, because the requisite material is generally wanting,
and the anatomical investigation of the leaf and axis, which are more easily
obtained and show a greater diversity of structural features, still affords
abundant scope for research. The more important characters, to be taken
into account in an investigation of the root, are as follows: (a) the. number
of xylem- and phloem-groups in the primary fibrovascular system (exception-
ally very large,-as in the case of Monocotyledons,-in Lophira, or reduced
to a monarch condition in the delicate lateral roots of Trapa), a feature which
is occasionally of considerable systematic value (e. g. march roots in the Gen-
tianeae-Gentianoideae, 5-9-arch roots in the Menyanthoideae), although some-
times varying within certain limits in onellnd the same species; (b) the mode
of differentiation of the secondary xylem-mass, the ground-mass of the wood
being either (I) unlignified and in this case containing vessels only or vessels
together with scattered groups of wood-fibres, or (2) lignified, in which case
it is composed for the most part of wood-fibres; these features are connected
with the physiological functions of the root (as a nutritive or attaching organ,
as a respiratory root, &c.), and are therefore subject to variation within certain
limits in one and the same species; (c) the rare occurrence of hard bast in the
phloem-groups ,(Anonaceae, Malvaceae, Sterculiaceae, Tiliaceae, Papilionaceae,
Urticaceae); (d) the structure of the en do dermis , including among other
features the occurrence of secondary division-walls in this layer (Valerianeae,
Compositae. Gentianeae-Gentianoideae); (e) the structure of the hypodermal

1 See H. Schenck, loc. cit., 1893, po. 21, and in Pringshebn Jahrb., xx:vii, 1895, p. 58 I.
, Van Tieghem, Trait~ de bot., 1891, p. 6i3, and Van Tiegbem et DOllliot, in Ann. &C. nat.,
sCr. " t. viii, 1888 j Kramer, Wurzelbaut, Hypodermis n. Endodermis der Angiospermenwurzel.
Bibl. bot., Heft 59, 1903, also Diss., Marburg, 151 pp. and 6 Tab.; Freidenfeldt, Anat. Ball
d. Wurzel, Bibl. bot., Heft 61, 190..., u8 pp. and 5 Tab. (see a\so in Botaniska Notiser, 1900,
Helt 5, and Flora, 190 3, p. 115 et seq.); Leavitt, TrichOdlel of the root, Proceed. Boston Soc. Nat.
, Hiat., xxxi, 1904. pp. :173-313; Neuber, Vergl. Anat. d. Wuneln mit bts. Beriicksichtigung der
Heterorhizie bei Dicotyl., Diss., Bern, 190.,70 pp.; Tschircb, Heterorbizie, Flora, 1905, pp. 68-78;
Biisgen, WtIJu15ylteme einiger dicotyler HobpfiuzeD, Flora, 1905, Ergau.-Band, PP.58-9. and
Tab. i-iv, especially p. 91.
 u68 CONCLUDING REMARKS

cell·layers, which are frequ~ntly marked off from the remaining ground-tissue;
(I) the occurrence of peculiar ridge·like or otherwise-shaped thickenin~ (Van
Tieghem's reseau de soutien ') in the s~bepidermal layer of cells or In more
f

deeply situated cells of the primary cortex (especially in the layer of cells
external to the endodermis); (g) the structure of the root-hairs (branched in
B,assica, Saxi/raga, &c., in other cases grouped together in tufts, in the
Nymphaeaceae originating from idioblasts in the dermatogen); (n) the
presence of secretory organs, their position with reference to the primary
fibrovascular system of the root in some· cases being, of importance in the
diagnosis of an Order, e. g. the dorsal position of the resin-canals with reg_ard
to the primary phloem-f?l:oups in the Pittosporeae, ¥aliaceae and Umbellifelae,
which is connected WIth a doubling of the usual number of young lateral
roots, the endodemial origin of the resin-canals in the Compositae, &c. lcl. § 13
et seq.); lastly, (J) the occurrence of mycorrhiza 1.
A strengthening net-work (' reseau de souUen ')' has been observed in the
following Orders: Berberideae, Cruciferae (Fig. 17. p. 66). Geraniaceae (Fig. 39,
p. 170). Rutaceae, Sapindaceae, Papilionaceae, Caesalpinieae, Rosaceae, Rhizo·
phoraceae, Lythrarieae 1, Caprifoliaceae, Ericaceae, Myrsineae, Verbenaceae. In
the plants, which form the landscape of the mangrove, the ridge-like thickenings
are found only in some parts of the root-system, which is here adal?ted to carry out
diverse functions. Regarding similar thickenings in Acanthus dicilolius. which
likewise grows in the mangrove-formation, see under Acanthaceae, p. 1020.

§ 64. ANOMALOUS STRUCTURE OF THE ROOT. The occurrence of intra-

xylary and interxylary phloem and of successive secondary zones of wood
and bast in the root has already been dealt with in §§ 57, 58 and 59 in
connexion with the analogous features found in the stem 3. In this paragraph
a fe.w other cases of-anomalous root-structure may be briefly mentioned, viz. :
(a) the development of concentric vascular bundles with central phloem in
the wood of the root in species of Brassica and Raphanus; (b) the formation
of concentric vascular bundles with central xylem or rarely (Drosera) phloem
in the wood of the root, which frequently leads to a separation of the fibro-
vascular system into concentric vascular'bundles, arranged in one or more rings
(in certain Crassulaceae, species of Drosera, Cucurbitaceae, Umbelliferae, Fig. 97,
P.425, Compositae and Convolvulaceae, Fig. 131, p. 572); (c) the occurrence of
similar structures, which, however, show a different course of development, in
species of Aconitum (Ranunculaceae); (d) the complicated structure of the roots,
of Myrrhis (Umbelliferae), AtractyUs (Compositae) and certain Convolvulaceae,
which is produced by the appearance of secondary meristems in the wood or
sometimes (Convolvulaceae) in the secondary bast and the formation of
secondary groups or rings (the latter occasionally being inversely orientated
in the three Orders named) of wood and bast; (e) the occurrence of cortical
vascular bundles in the tuberous roots of Mynnecodia (Rubiaceae). For
details, particularly as regards t~e course of development of these anomalous
structures, see the descriptions of the individual Orders.
For a process of cleavage, which aets In in root. and rhizomes in the course
of growth in thickness (Ranunculaceae, Fumariaceae, Crassulaceae, Gentianeae.
Labiatae; also Plantagineae, according to Pilger), see especially JO5t, in Bot. Zeit..
1890, p. 433 et seq. and Tab. vi; the cleavage in these cases is due to.the removal
of groups of dying tissue, while the intervening living tissue continues its growth.

1 In the root internal de\·elopment of tbe cork is of general OCCUlTence, the' outermost cell-layer
of the pericamblum giving rue to the lint pbello~.
• Van Tieghem, in Ann. Ie. nat., ~r. J, t. Vii, 1888, P. 375-
I See H. Schenck, Anat. der Lfanen, 1S,3, pp. 251-6.
 LITERATURE SUPPLEl\1ENT
(Papen which c:a.me to the notice of the' author after the Addenda had gone to press).
AMARANTACE ..\E (p'p. 655, I028): Gravis, Contrib, a l'anat. des Amarant., Mem. soc.
roy. d. sc. de Li~ge. ser. 3, t. vii, 1907. 65 pp. and 14 pI.
AMPELIDACEAE {pp. 226, 890~: Szigethi-Gyula, Anat. d. Weinwurzeln, etc., NOvenyt
Kozlem{myek, IV, 1905, p. 45 et seq. (Hungarian, with a French r~uml!).-Baker
and Smith, Vitis opaca, Proc. Roy. SOC. New South \Vales, 1906; author's abstr. in
Bot. Centralbl., crv, p. 661 .-Hoitermann, Einfluss d. Klimas, 1907, p. I 10 (Vitis,.
ANACARDIACEAE (pp. 248, 894): Dubard et Dop, Protorhus Hecke{ii, Bull. Soc.
bot. de France, 1907, p. 155.-Guttenberg, Immergr. Laubbl. d. Mediterranfiora,
in ~ngIer, Bot.Jahrb., XXXVIII, 1907, p. 425 (Pistacia).-Hemsley.lulianiaceae,
Phil. Trans. Roy. Soc. London, Ser. B, CIC, 1907, pp. 169-97; and Proc. Roy.
Soc. London, LXXVII, 1906, pp. 231-6.-Holtermann, Einfluss d. Klimas,
I'P7, p. 118 (Sem~ca,.pus).-Lecomte, Phtebochiton, Bull. SOc. bot. de France.
1907. p. 525 et seq.-F. E. Fritsch. Anatomy of the Julianiaceae, Trans. Linn.
,Soc. London, VII, 1908. pp. 129-51, pl. zo. 21.-Hallier, juliania, Beih. z. bot.
Centralbl., XXIII. 2. Abt., 1908. p. 82 et seq.
ANoNACEAE (pp. 39. 8Il) : Moll and Janssonius, Mikrographie des Hones, Heft I,
Leiden, 1906. pp. 106-73 (Stelechoca,.pus, Cyathocalyx, Canangiu"" Meiogyne. Poly-
tJlUaia. T,.iva/varia, Popowia,llJitrepho,.a, Platymitra, A nona, Saccopeealum, Orof'lttJ,
Alplwnsea).
APoCYNACEAE (pp. 533, 988): Cantoni, Produz. sugh. della Strophanth#s 'hispuus,
Malpighia. 1906, pp. J7I-9. and tav. II, III; abstr. in Bot. Centralbl., CIV.
p. 194.-Perrot et Burrier, Mat. m~d. et Phannac. sino~annamites, pp. 168, 169
(Apocynum iuventas, Lour.). in Perrot, Travaux, V, I9C?7.
ARALIACEAE (pp. 432, 948): Hurrier et Perrot, Ginseng, Bull. sc. pharmacol., 1906,
p. 660; also in Perrot, Travaux, IV.-Sperlich. Opt. Verh. in der oberseit.
BIattepid., Sitz.-Ber. Wiener Akad.,.CXVI, Abt. I, I'P7, p. 6c)8.
ASCLEPJADEAE (pp. 537.989): Pearson. Sp. of Dischidia with double pitchers, Joum.
Unn. Soc. London, XXXV, 19(>2, Pt' 375-9(>, pI. 9.-Glabisz, Ceropegia Woodii,
Schlecht., Beilt. z. Bot. CentralbL, XX II,!. Abt., 19Q8, pp.6S-136 and Tab, IX-XI.
BALANOPHOREAE (pp. 739, 1047): Heinricher, BaJanop_hora, Sitz.-Ber. Wiener
Akad., CXVI. Abt. I. 1907, pp. 439-65. 1 Tab.-Van iieghem, Inovulees, Ann.
!c. nat., s~r. 9, t. vi, 1907. n. 3, 4.
BERBltRIDEAE (pp. 46, 822): Frommel, Plant. text. chil., 1905. pp. 38~.·-Ruu­
bourg, Et. org. et anat. de la famille des LardizabaIees, These. Pans, 1900.
127 pp.-Holm. CauIophyllum tha/ictroides, ill Merck's Report. XVI, 1907,
pp. 94-6.-Holm, jeUet'sonia diph'llla. in Merck's Report, XVI, 1907. pp. 125-7.
-Holm. Podophyllum peltatum, m Merck's Report, XVI, 1907. pp. 250-2.-
R~aubourg. Boquila tyi/oliata, Bull. Soc. bot. de France, 1907, pp. 8-10.
BIXINEAE (pp. 91, 8jl): Calderera. Foglie della Kiggelana a/ricana, Contrib. BioI.
veg., Palermo. II , 1904, pp. 273-92.-Moll' and Janssoni us, Mikrographie des
Holzes, Heft I, Leiden. I9CJ6,pp. 197-223 (Scolopia,Flacourtia, Bennetlia, Pangi"""
Bergsmia. Taraktogenos, Ryparosa).
CAMPANULACEAE, incI. LOBELIACEAE (pp. 475, g68): Flachsbers-er, in Sitz.-Ber.
naturf. Gesellsch. Dorpat, 1906, n. 3.-Hurrier et Perrot, GInseng. Bull. SC.
pharmacol.. 1906, p. 661 et seq.-Tswett, Hydathodes des Lobel., Revue g~n. de
bot., XIX. 1907, pp. 305-16 and pl. I4--Perrot et Humer, Mat. med. et Ph.a.qn.ac.
sino-annamites, 'p: 184 et'seq. (Adenopho,.a, Campanula, Platyeodon), in Perrot,
Travaux. V, 1907.
CAPRIFOLIACEAE (pP. 444. 949): Hohnel, Kork, Sitz.-Ber. Wiener Akad., LXXVI,
Abt. I, 1877. p. 607.-Kanngiesser, Holz von Lonicera PCl'icJymenum, in Tubeuf,
Naturw. Zeitschr., I9Q6, pp. 404-8.-Eichinger, Vergl. Entwicklungsgesch. von
A.doxa u. Chyysosplenium. Diss., Munich, 1907, 27 pp.; also in Mitteil. Bayer.
bot. Gesellsch., I907.-Guttenberg, Immergr. Laubbl. d. Mediterranfl., in Engler.
Bot. Jabrb., XXXVIII, 1907, p. 440 (Viburnum Tinus).
CELASTRINEAE (pp. 214. 879): Loesener, Kautschuk bei ein.ostafrikan. Gymnosporia-
Arlen, N otizblatt Berliner Garten, n, 42, 1908, p. 64 et seq. (as in Celastrus. etc.).
CISTINEAE (pp. 82, 827): Roche, Anat. compo de la feuille des Cistin., These, Paris,
1906, 108 pp.; also in Perrot, Travaux, IV.-Gard, Anat. compo des Cistes,
Comptes rendus Acad. Paris, CXLIVJ I'P7, pp. 1229-32.-Guttenberg. Immergr.
Laubbl. d. Mediterranfiora, in Engler, Bot. Jahrb., XXXVIII, 1907, pp. 426-32
SOLEUDU 4F
1170 LITERATURE SUPPLEMh."N"F
(Cistus).-Gard, Format. cystolithiques des Cistes, Com.Jlte5 rendus Acad. Paris,
eLV, 1907, pp. 136-7.
COMVOSITAE (PJ>. 468, 962): Drobnig, WurzelknoUen. Diss•• Rostock, 189:l, p: sz
(DaMia. Cif'S"um ).-Herriott, Leaf structure, Trans. New Zealand Inst., XXXVII~
1906, p. 377 et seq. - Dauphin6, Struct. du rhizome de l'Artem(sia vulgaris,
Revue gen. de bot., XIX, IfP7, p. zg6 et seq.; abstr. in Bot. Centralbl., CV,
p. 4z1.-Maheu et Combes. Format. sub&'o--pheUodenn. anorm., Bull. Soc. bot.
(Ie France, IC)07, p. 436 et seq. (Trtlgoptltrm).-Meinheit, Anat. Ba'!l d Sten~els
bei den ComPQS.itae-Cynareae, Diss.• Gottingen. 1907, 1I8 pp.-Perrot et Humer,
Mat. med. etllharmac. sino-annamites. p. 196 {(;ynuYQ pinnalifida, DC.), in Perrot,
Travaux, V, 1907.
CRtJCIFERAE (pp. 67, 82$): Thellung, LePidium, Neue Denkschr. allg. schweizer.
GeseUsch. f. d. ges. Naturw.., XLI, Abt.l, 1906.-Gerber, L'arc renverse de l'Au-
brielia delloidea. Com_ptes rendus Soc. BioI. Paris, LXII, 1907, pp. 976-8.-Gerber.
Faisceau inverse de Zilla macf'opttJ'a, Comptes rendus Acad. Paris, CXLIV, 1907.
pp. I374-6.-Muschler. OOJ'onopus, in Engler, Bot. Jahrb., XLI, 1907. p. 118.
CUCURBITACEAIt (pp. 397, 939): Bernet, Obs. ana.t. nouv. sur la tige des Cucurbit..
Bull Herb. Boissier, ser. 2, t. v. 1905_ p" 312. '
CUPULlFERAE (pp. 796. 1068): Gallagher, Root~anat. of the CupuUf. and of tlie
Me1ia.ceae, Rep. Brit. Ass., York. 1906, pp. 74s>-so.-Kramer, Mikr ••pharm.
Beitr., Diss., Wiirzburg. 1907, p. 12 (Castanea).
DIPTltROCARPEAE (pp. 144. 842): Moll and Janssonius. Mikrograpbie d. Holzes,
Heft I~ Leiden. 1906. pp. 343-68 (Dipteroca,pus. Vatka. Shorea. Hopta}.-Gu~riD.
Et. anat. de Ia tige et de la feuille des Dipteroca.rp., Bull. Soc. bot. de France,
M~m. II, 1907, 93 pp.-Sperlich, Opt. Verh. in der oberseit. Blattepidermis, Sitz.-
Ber. Wiener Akad., CXVI. Abt. I. 1907, p. 702. ' "
EUPHORBIACEAE tpp. 76z. 1°54): IBalfour. Cutic. struct. of Eupho,bia Abdelkuri,
Rep. Brit. Ass •• 1901, p. 8S4.-Woodhead. ,Reol. of Woodland Pl., Journ. Linn.
soc. London. XXXVII. 1906, pp. 394-5 (Me,cu"ialis pe"ennis).-Perrotet Burrier,
Mat. med. et Pharma.c. sino-annamites, pp. 104-5. in Perrot, Tra.va.ux, V. 1907
(AcalyphQ f,u#cosa, Forsk. }.-Sperlic~ Op~ Verh.. in der oberseit. Blattepid.•
Sitz.~Ber. Wiener Akad., CXVI, Abt. I, 1907. p. 708 et seq.
GUTTIFERAE (pp. 126, 836): Moll and Janssonius, Mikrographie d. Holzes, Leiden,
Heft I, I906, pp. 250-82 (Gaycim'a, Calophyllum).-Holm. Ga,cinia coch.jnchinensis.
in Merck~s Report, XVI, IC)07, pp. 1-4; abstr. in Bot. Ceqtralbl., CV, p. 39--
Holtermann, Einfiuss d. Klimas, etc., 1907. pp. 114-15 (CaJophyllum).
lLlClNEAE (pp. 209. 874): Thevenard, Rech. mstal. sur les lUciD., lhese, Paris.
190(>, 150 pp., 6 pl.,; also in Perrot, Travaux, IV, 1907.-Holtermann. Eintluss
d. Klimas. etc., 1907. pp. 116-17. •
LABIATAE (pp. 641, 1023): Kimpfiin. Affinites des Bora.gin~es et des Lamiacee!.,-
Assoc. Avanc., 3S1!1 Sess., Lyon, 1906, pp. 428-3 I.-Woodhead, EcoI. of Woodland
Pl.. loum. Linn. Soc. London, XXXVII. 1906. pp. 393-4 (Lanu'um Galtobdolon).-
Holm, Cun'ila Mariana, in Merck's Report, XVI. H}OJ, pp. I88-9.-Mitla.cher.
Anat. Verb. d. Labiaten, Zeitschr. osterr. Apoth.~Ver., 1908, pp. I. 17, 33, 4S
et seq. and 4 Tab.
LEGUMIl'iOSAE (pp. 300, 905): Cordemoy, Appareil secret. de l'Epeyua jalu:da, Ann.
lnst. col. Ma.rseille, 1906, pp. 1-22.-Decrock et Riba-ut. Appareil secret. du
Vatairea guianensis et d u 1\-facliaerium /et't'ugine'Um: Ann. Inst.. Marseille, XIV, 1906.
-Boorsma, Alo~holz, Bull. Depart. de l'Agric. aux Iudes u~rland•• VII. 19:J7.
p. 25 (Dalbergia).-Dubard et Dop, Esp. DOUV. de Madagascar, Bull. Soc. bot. de
France, 1907, p. IS7 et seq. (Mundulta. ChadS1'a).-Guttenberg, lmmergr. Laubbl.
cL Meditcrrandora_ in Engler, Bot. Jahrb., XXXVIII, 1907, pp. 423-5 (CeratofJitl
5iliqua and Spa,.lium junuum).-HoltermanD, Einfiuss d. Klimas, [907, p. 91
(Indigo/eya 4spalathoius).-Perrot et Gerard, Anat. du tissu ligneux, Bull. Soc.
bot. de France, M~m. 6, 1907. 43 pp .• 6 pl.-Perrot et ~rard. Bois de clift. esp. de
Ugum. Afric., Pa.ris, 1907, ISS pp. and Tab., in Perrot, Travaux, V, 1907.-Perrot
et H umer, Ma.t. med. et Pharmac. sino-a.nnamites, p. I 52 (Sophcwa .angusti/oli4
S. et Z.l. in Perrot, Travaux. V, 1907.-Schwendt, ExtraJior. Nektar., Beih. z. bot.
Centralbl., XXII, I. Abt.. 1907, ~p. 264--9 (Acac;'Q ).-Severini, Radici di H~d'Ys(J,ru".
corona"ium, Atti R. Accad. Lincei. XVI, 2, 1907, pp. I4S-8.-Sperllch. Opt.
Verh. d. oberseit. Epid .• Sitz. Ber. Wiener Akad., CXVI. Abt. I, 1907, p. ;719
et seq.-S1:scherbatschetf, Entwicklungsgesch.. ein. offiz. Ptt, Archiv d. Pharm.,
IfP7,;P. 48 et seq. (Glycyy,hiza). -
LVTBRARIEAE (pp. 373. 931): Pascbkis; in Zeitschr. Osterreicb. A~theker.Ver.,
1879. p. 433 (Lawsonia).-Blatter, Mangrove of the Bombay PreSIdency, Joum.
Bomba.y Nat. Hist. Soc., X'v'I, 1905: pp. 644-56 and pl.
 LITERATURE SUPPLEMENT
MALPIGHIACEAE (pp. 167,850): Hartwich, lpecacuanhawurzeln, Archiv d. PbarnJ ••
CCXLII, 1904, p. 666 et seq. and Tab.-Mannich and Brandt. H.teropuri$
paueiflor(J, Ber. deutsch. pharm. Gesellsch., 1904, pp. 297-302.-Lutz, lJluline
dans quo Malpighiac~, Bull. Soc. bot. de' France, 1907, pp. 449-S2.-Dubard et
Dop, Observe sur l'ana.t. et les aftiniUs des Malpigh. de Madagascar, Comptes
rendus Acad. Paris, CXLVI, 1908, pp. 355-7.
MALVACEAB (pp. 152,843): Frommel,-Plantas text. chit, 1905, p. 33.-Ho1termann,.
EinBuss d. Klimas, 1907, p. 14 (Du,io).-Schwendt, Ex:traflO~e Nektar.,. Dem.
Z. Bot. Centralbl., XXII, Abt. I, 1907, pp. 2S4-6.--Stseherbauchefi, Eutwiek-
I u~gsgesch. ein!g. offiz: Pft, Arch,i v ~. Pharm., 1907, p. 48 et seq. (A IlAaeti ).--CaJvet,
Rist. bot. des Kapokiers. Montpellier, 153 pp.; abstr. Bot. CentralbL. CV, p. 420 ..
MELASTOMACRAB (pp. 368, 928) : Rohnel, KorJr., Siu.-Ber. Wiener Akad., LXXVI,
Abt. I, 1877, p. 6JO.-Clark, Sec. thickening in Kmd,ickitJ Walkeri, Hook. fiL,
Ann. of Bot., XXI. 11)07. PP. 361-7 and pl.-HOlm. Morph. and ana~ stud. of
the vegetative organs of Rhexia, Bot. Gaz., XLIV. 1907, pp. 22-33. 2 pl.
MELlACEAE (pp. 198, 870): Courcbet, Katafa (Ctdrelopsis). Ann. Inst.: Marseille.
XIV, 19Q6, pp. 29-II8.-GaUagher, Root-anat. of the CUJ?uliferae and of the
Meliaceae. Rep. Brit. Ass•• ]906. pp. 749. i'50.-Radlkofer,lD Natiirl. P1ianzen-
hm., Erg.-Heft II, 1907. p. 204 (Rhetinosperma).
Congres internat.• Paris. 1900-1, p. 218 et seq.-Maheu, 0=,
MltNlSPERMACEAE (pp. 43. 818): Maheu, Et. aur les Menisperm.. Compte rendu
s~cr~t. des M~nisperm. ••
Bull. Soc. bot. de France~ 1906~ pp. 6S1-63.-Molland anssonius.Mikro~phie
d. Holzes, Heft I. Leiden, IQ06• .l'P. 173-5 (Coceulus).- Syst.-a.nat. Unter·
such. d. Blattstruktur bei den Menisperm., Diss., Erlangen, 1907,92 pp.,_1 Tab.-
SperIich, Opt. Verh. in d. oberseit. Blattepid.• Sitz...Ber. Wiener Akad., CXVI,.
Abt. I, 1907, p. 712 et seq.
MVRTACEAE (pp. 357'-920): Hohnel, Kork, Sitz.-Ber. Wiener Aka.d., LXXVI, Abt. t.
1877, p. 6IJ et seq.-Baker and Smith, Australian Melaleu&as. Proc. Roy. Soc.
New S. Wales, 1906; authors' abstr. in Bot. Centralbl., CIV, p. 66z.-Guttenberg,
Immergr. Laubbl. d. Mediterranfiora, in Engler, Bot. Jahrb .• XXXVIII, 1907,
p. 432 (My,tus italica).-Kramer, Mikr.-pharm. Britt Diss., Wurzburg, 1907.
e. 16 (Eucalyptus).-Solereder. Deckhaare der Pimeutiriichte u. der Myrtac.
uberhaupt, Archlv d. Pharm•• CCXLV, 1907, pp. 410-14.
QLEACEAE (Pp. 525,982): Alquati, Studi anat. e mori. sull Ullvo, Atti Soc~Ligtv"'
l

Sc. nat., 1906, pp. I28-48.-Guttenberg, lmmergr. Laubbl. d. MediteriantL. lU

Engler, Bot. Jahrb., XXXVIII, 1907, pp. 435-40 (Olea eur<>/Jaea and PhillyretJ
lati/olia).-I..ingelsheim, Vorarb. zu einer Monogr. d. Gatt. PrlUs'"us, in Engler,
Bot. _lahrb .• XL, 1907, pp. 193-5.-Schwendt, Extra1lor. Nektar., Bot. Centralbl.,
xxII, Abt. I. 1907, pp. 259-61. "
()NA.GR.ARIEAE (pp. 315. 932): Hohnel, Kork, Sitz. Ber. Wiener Akad., LXXVI,
6

Abt. I, .1877. p. 609.-Blatter, Pectinate organs ~f Tf'apa bicornis, loum. Bombay

Nat. Hist. Soc., XVII, 1906, _pp. ·84-8; abstr. m Bot. Centralbl .• eIV. p. 467.
PIPJtRACBAB (pp. 694. 1038): Schiirhofl, Oullen u. Lichtkondensatoren 6ei ein.
PePeromien. Beih. z. Bot. CentralbL. XXIII, Abt. I, IgoS, pp. 14-26,
PInosPoRBAE (pp. 94. 831): Guenot, Et. anat. des Pittos~r., Th~se, Part$, 1906,
1 8 pp.-Moll & ]anssonius, Mikrogr. d. Holzes, Heft I, Lciden, 1906. pp. 223-32.
POLYGALEAE (pp. 100, 83r): Moll and Janssonius, Mikrographie d. HoIzes, Heft I,
Leiden, 1906, pp. 232-9 (XantJaophyl1um).-Holm, PoJygala Senega, in Merck's
Report, XVI,_1~7, pp. 155-7.
POLYGONACEAE (pp. 674. 1°34): Porsch, SpaltOfinungsapp. submers. Pfianzenteile.
Sitz.-Ber. Wiener Akad•• CXII. Abt. 1,1903, p.128(Polygonumamphibium).-Goris
~t Crete, Rhubarbe de Chine, Bull. sc. pharmacaL, 1907, pp. 93-104; also in
Perrot. Travaux, V.-Goris et Cr~te. PoljIgonum cuspida/;:m, S. et Z., Bull. ~.
pharmacol., 1907, p. 698 et seq.; also in Perrot, Travaux, V.-Maheu et Combes,
Format. subero-pnelloderm. anorm., Bull. Soc.. bot:. de France. 1907, p- 434
(Rumex).- J. Schuster. Polygonum lapathi/olium, Mitteil. bayer. bot. Oesellsch.,
II, n.4, 1907. p. SO et seq.-'Schwendt, ,ExtrafiQrale Nektar., Beih. z. bot. Cen-
tralbL. XXII. Abt. l, H)07, pp. 249-53 and Tab. IX (Polyg(»J~m, MilJilenbeclria).
PRIMULACEAE (pp. 506, 976): Ricome, Auricule, Comptes rendus Acad. Paris,
CXXXIX. 1904. pp. 46S-io.-Hildebrand, Die Cyc;lamen-Arten, Beih. z. bot..
Centralbl.. XXII. Abt. 2, 1907, p. 143 et seq. and Tab. II.
RASUNCULACEAE (pp. 19. 806): Drobnig. Wurzelknollen. Diss .• Rostock, 1892.
pp. 10, 39 and 62 et seq. (Fica,ia. Acon£tum. Paeonia).-Tonkoff, Blattstielan-
schwell. von At,agene. Ber. deutsch. bot. Gesellsch., 1894. p. 40 et seq.-Goris et
WaJIatt, Hydl'astt"s canadensis, Bull. sc. pharmacol., 1906, p. 6~4 et seq.; also in
Perrot, Travaux, IV.-Haberlandt, Sinnesorg .• 2nd Edition, 1906.-Herriott,
~ ,.. ~
 LITERATURE SUPPLEMENT
Leaf struct.., Trans. New Zealand Inst.. XXXVIII. 1906, p. 377 et seq.-Holm,
Anat. method, Americ. JO\U1l. of Pha.rm., 1907, p. ?9.-Holm, ,A,cmifum 'lm&i~.
ill Merck's Report, xvI, rep7, pp. 65-7; abstr. in Bot. Centralbl.. CV, p. 239.-
Ho~ AnemtJfUUill"tJJi~s. Spacb. Americ.. Joum. Sc., XXIV, I~, pp. 343-8.
-Kramer. Mikr.-pharm. &itr., Diss.., Wilrzburg, 1907, P.7 (Adonis).-Perrot et
Burrier, Mat. med. et Pharm. sino-annami tes, Perrot, Travaux. V, 1907, pp. 1 I S.r9
lbt.uiNEAE (pp. 221, 889): Holm, Ceanolhu$ amencanus, Americ. Journ. of Se.,
XXII, 1906, pp. S23-30.-TichomUow, Inclusions intraceU. de Ia feuiUe du
Nerprun purgati£. Comptes rendus Acad. Paris, CXL II I, 19Q6, pp. 92~-4-
-Gutten~, IUlDl~. Laubbl. d. Meditenantl., in Engler, Bot. Jahrb~.
XXXVIII. rt;1J7, p. 426 (Rhamnus Alalernus).
ROSACEAE (pp. 309. 908): Ho~l. KOJ:'k, Sitz.-Ber. Wiener Akad., LXXVI, Aht. I.
1877. p. 606.-Drobnig, Wurzelknollen. Diss.. Rostock, 1892. p, 68 (SplrlUlJ
Filspmdtlla).-MiJroscb, Kirschgummj. Sitz.-Ber. Wiener Akad., CXV, Aht.' I.
1906. pp. 91I-6I.-Kramer, Mikr.-pharm. Beitr., Diss., Wnrzburg, 1907, p. 22 •.
-Viguier. Anat. du Geum ..if/ale. Revue gen. de bot.~ XIX, 1907~ pp. 22[-5- '
RUBIACEAE (pp. 454, 9S I): Hartwich, lpecacuanhawurzeln, Arcbiv d. Pharm.,
CCXLII, 1904. pp. _649-74 and 2 Tab.-Herriott,i Leaf structure, Trans. N~w
Zealand' Inst., XXXVIII~ I~ p. 377 et se_q..-Holm, Anat. stud. of North
Americ. represent- of Cepbalanlhus, Vldmland1a, Houstonia, Mi~1uU(l, Diodill,
aDd Gal."",. Bot. Gazette, XLIII, 1907, pp. 153-86, 3 pi.
RUl'ACEAB (pp. 182. 857): Holmes, A new vu. of Buchu leaves, Pharm. Tourn.,
_1901, p. 598.-Holtl!rma nn. Einfluss d. Klimas, 1907. po., 106 (Fe,onia).-lUa.mer ..
Mikr.-pbarm. Beitr., Diss., Wiirzburg, 1 907, p. 9 (Barosma).
SAPOTACBAE (pp. 51;, 980): Perrot, Le KariM, I'Argan et quo autres Sa,P.Ot., in '
Perrot. Travaux, V. I907.--Sperlich, Opt. Verh. in d. oberseit. Blattep.dermis.
Sitz.-Ber. Wiener Akad., CXVI. Abt. I, 1 907. pp. 704-6.
SAXIFRAGACEAlt (pp. 319. 91 I): H6hnel, Kork. Sitz.~Ber.· Wiener Akad., LXXVI •.
~ I, 1877. p. 607 •._.Hamilton, West Austral. pitcher-plant (Cephalotus/olli&,!,-
141"'$). Proc. Linn. Soc. NeW' S. Wa1es~ XXIX. 19(14, pp. 36-53~ pI. It II; abstr.lO
Just. 1905, II, p. I8.-Montemartini, Ascidi anomali nelle foglie di Saxifr(Jga
ertUst/olia, Atti lat. bot. Pavia, 1904.-Rasendahl. Die nordamerik. Saxifra.~nae,..
in Engler. Bot- Jahrb., XXXVII, Beibl. n. 83, 1905, p. 23 et seq.- Eichinger,
Vergl. Entwic~un~g~ch. von Adoxa u. Chrysosplemum f Diss., Munich, 1907 ••
27 pp.; also In Mitteil. Bayer. bot. Gesellsch., 1907.-Kudelka, Anat. compo
d'org. v~get. des Groseilliers, Bull. internat. Ac. Sc. Cracovie, 1907, I.
SoLAN4CEAE (pp. 582, 1007): Terras. Relat. hetw. lenticels and adv. roots of
Solanum DuJcama,.a. Tr~s. Bot. Soc. Edinburgh. 1900, pp. 341-S3.-Wallis,.
Struct. of Capsicum minimum, Pharm. Jauro. London, 1901, pp. 552-7.-Chrysler,
Strand. plants, Bot. Gazette. XXXVII. 1904, p. 461 (Soltmum).-Stscherbatscheft.
Entwicklungsgesch. einig. o:ffi.z. Pfl., Archlv d. Phann., 1907, p. 48 et seq.(Ah'oj>a).
TBRNSTROBMIACltAB (pp. 135. 839): Moll and Janssonius, Mikro~aphie d. Hofzes.
Leiden, Heft I. 1906, pp. 282:-.342 (T"",stroemia, Adinandra, Eurya, Savrlltlill,
ScMma, Pyrenaria, GOf'dmia, Haemocharis, Camellia).-Pilger, in Natiirl. Pilani
zenIam., Erg.-Heft II, 1907. p. 226.
TSYYBLAEACEA2 (pp. 721,1°45): Nitsche, Britr. z. Kenntnis d. Gatt. Daphn~. Diss.,
Breslau. 19CYl, 34 pp.
TUMANDREAE (p. 96): Van Tieghem, QU. rem. sur les Tremand., Ann. sc. nat..
sh. 9, t. iv, 1906, especially pp. 378-82.
UMBELLIFR~Alt (PP. 426.941): Drobnig, WurzelknoUen, Diss., Rostock, 1892. p. 22
et seq. (Oenanthil).-Schinz. Beitr. z. afrik. Flora, Bull. Herb. &iss. 1894, pi. 4
(Pitu,.anthos). - Porsch, SpaltOffnungsapp. submers. Pfianzenteile, Sitz.-Ber.
Wiener Akad .• cxn, Aht. I, 1903. pp. 112 and II4 (Oenanthe aquatica).-Tun-
mann, KristaUe, Phann."2.eit., 1905, pp. IOS5-7.-Herriott. Leaf struct., Trans.
New Zealand Inst., XXXVIII, I9Q6, p. 377 et seq.-Hurrieret Perrot, Ginseng.
Bull. sc. ptiarmacol., 19Q6. f. 664 (Angelica); also in Perrot, Travaux. IV.-
Woodheaa, Woodland Pl., ~Journ. Linn. Soc. London, XXXVII, I9Q6. pp. 39 1,
392 (Heracleum Sphondylium).-Maheu et Combes. Format. subero-phellOderm.
anorm., Bull. Soc. bot. de France, 1907, p. 432 (Thapsia).-Perrot et Hurrier,
p. 160 (Angelica polyclada); in Perrot, Travaux, V, 1907.
VACCUlIACEAE (pp._ 479. 969): Woodhead, Eeol. of Woodland Pl., loum. Lin".
Soc, London, XXXVII. 1906, pp. 387-91 (Vaccinium Myrt1.'lIus).
Papen tIeldi", with a numbe,. 01 Ordet'S 0,. tV1~h defi'l1d~ atla~o",icl1llea.tu,.es. Hill, Sieve·
tubes of Angiosperms, Part I, Ann. ot Bot., XXII, I90 8, pr.. 245-90.-0no, Some extra·
nuptial nectaries, 10um. Coli. Se. imp. Univ. Tokyo, XXII, 1907, 28 pp. and 3 pl.-HoU-
stein, Vergl. Anat. d. Stengel u. Rhiz. von dicotylen Alpenpflanzen, Diss., Gottingen. 1907·
 INDEX TO ORDERS AND ANOMALOUS GENERA
(for Vols, I and II)
A Colurnelliaceae 598. 101 I. Geraniaceae 169, 85 I.
Acanthaceae 613, 101 S. Combretaceae 343,918. Gesnera.ceae 599, to 12.
Aceracea.e 238. 891. ComJx>sitae 45 6, 953. Gomortegaceae 709.
Amarantaceae 651, 1027. .. (Literature) 468, Gonystylus 72 I, 1045.
Ampelidaceae 221,889. 962. Goodeniaceae 47 1 , 965.
Anacardiaceae 244, 893· Connaraceac 250, 895. Gyubbia 737, 1046.
Ancistrocladus 143, 84 1 • Convolvulaceae 562, 1002. Guttiferae 120, 836.
Anonaceae 34, 810. Coriarieae 249, 894- ..
Comaceae 432, 948. H
Apocynaceae 5 28 , 983.
Araliaceae 426. 942. Corynocarpaceae 247, 884· Halorageae 335, 9 1 5.
Aristolochiaceae 6Sz, J036. Crassulaceae 320, 91 I. Hamamelideae 328. 914.
AscJepiadeae 534.988. Crossosomataceae 909. Hernandiaceae 70 7. 1043.
Cruciferae 58, 824. Hippocastanaceae 23 6,891.
Cucurbitaceae 389.936. Hippocrateaceae 214. 880.
B • Cupuliferae 791, 1066. H umiriaceae 160. 849.
Balanophoreae 738, 1047. Cuscuteae 573. 1003· Hydrovhyllaceae 552, lOOt.
Balanopseae 763. 1055. Cyrilleae 2 I I. Hypencineae 117.834-
Basellaceae 663. 1030. Cytinaceae 680, 1035.
Batideae 668, 1°32. I
Begoniaceae 398.939. D llicineae 2<>9, 873.
Berberideae 44. 81 9. Illecebraceae 649. loz6.
Bignoniacea.e 60 I. 10 16. Dapbniphyllaceae 760,
Bixineae 87.828. 1°53· J
Boragineae 5-54. 1001. Datisceae 406.939.
Bt'luhynema 516. -_ Desfontainea 547. Juglandeae 783, 1065.
Diapensiaceae 494, 973. ] uliam'a 244. Il{9.
Brunia.ceae 333, 9 1 4-
Burseraceae 190. 869. Dicbapetaleae 198, 87 I. K
Buxaceae 761, 1053. Didiereae 8go.
Didymeles 783. Koebeylinia ]82, B6,.
c, Dilleniaceae 20. 807. L
Dipsaceae 455. 953·
Cacteae 406.939. Dipterocarpeae 136:840. Labiatae 636. 1022.
Caesalpinieae 28 I. 904- Droseraceae 324. 912. Lacistemaceae ·799.
Calycanthaceae 25.807. Lactoridaceae 34-
Calycereae 45 6 , 953. E Laurineae 702. 1040.
Campanulaceae 473.965. Lecytbidaceae 355.920.
Candolleaceae 469. 963. Ebenaceae 516, 980. Leguminosae 253. 895·
Canellaceae 86, 828.. Elaeagnaceae 724. 1045. II (Literature) 3001
Elatineae 116.
Cannabineae 769. 1056• . Empetraceae 800, 1069. 905·
Capparideae 67.825. Leitnerieae 782.
Epacrideae 490. 97 [. Lennoaceae 494-
Caprifoliaceae 439. 949· Ericaceae 479. 9(59.
Caiyophylleae- 107. 83 I: -- . Lentibularieae 59 1 , 1009.
Casuarineae 786, 1066. Euphorbiaceae 739. 1047· Ltnzia833:
Celastrineae 212,874. " (Literature) 762. Lineae 159. 847.
Cephalotus 319. 10 54- Loaseae 379.
Ceratophylleae 801, 1069. Eupomatia 34. Lobeliaceae 473,965.
Chailletiaceae 198, 871. Euthemis 869. Loganiaceae 538,989.
Champueia 736. F Lonchostoma 581,914.
Chenopodiaceae 655. 1028. Lophiya 144. 841.
Chlaenaceae 145. 842. Ficoideae 4 I 5, 940 • Loranthaceae 726, 1046.
Chloranthaceae 695. 10 38. Frankenia.ceae 104. Luxemburgiaceae ]88, 867.
Chrysobalaneae 30 I. 907. Fumaria.ceae 56. 824- Lythrarieae 369, 928.
Circaeasln 1039.
. Cistineae 79.827. G M
Clusiaceae 120, 836. Geissoloma 724. Magnoliaceae 27. 807.
CtUOJ'um 182. 858. Gentianeae 548.991, Malpjghiaceae 161.849-
1174 lNDEX TO ORDERS A.ND A.NOMA.LOUS GENERA

Malvaceae 146,842. Pedalineae 61 I. 1017. Selagineae 628, 1020.

Melastomaceae 3S8, 92 I. Peganum 178, 855. Simarubaceae 182.857.
Meliaceae 194, 870. Penaeaceae 722, 1045. Solanaceae 575, 1006.
Melianthaceae 240. Pentaphylacaceae 883. Stachyurus 127,838.
Menispermace ae 39, 812. Penthorum 910. Stackhousieae 2 17, 884.
Microsemma 135. Phytolaccaceae 664, 1030. Staphyleaceae 242, 893.
Mimoseae 29 1 , sos. Piperacea.e 6aS. 1037. Sterculiaceae t 52, 844.
Monimiaceae 699, 1040· Pittosporeae 91, 831. Strasburgeria 839.
Monotes 144.842. Plantagineae 642, IOl3. Stylidiaceae 469. 963.
Monotropeae 489, 971. Pla.tanaceae 779. 1064. Styraceae 51.9.980.
Moraceae 770, 1056. PloCQspef'ma 547.
Moringeae z49, 895- Plu1l1b~gineae 495. 973· T
Myo,porinea.e 624. l02Q. Podostemaceae 674, 10 34. Tamariscineae II3, 8J3'
Myncaceae 785, I06S. Polemoniaooae 550.999. Ternstroetoiaceae 127.837-
Myristicaceae 696, 1039· Polygaleae 96. 83 1 • T e/t'ame1'isla 189.
M",.othamnus 332. Polygonaceae 669. 1°32. Thelygoneae 779, 1604-
Myrsineae 5°7,.976. Portulaeeae I I I, 833' Thymelaeaceae JI5, 1044·
Myrtaccae 350) 919. Prlmulaceae-SOl,974· \ Tiliaceae 1 S5. 846 .
M yrtaceae sense str. 35:Z, Proteaceae 709. 1Q43· Tovaria 77.
9 1 9. Tremandreae 94.
Myzolknd,.on 733, 10 46. Q Trigoniaceae 100.
Quiineae 120. Triplocbitonaceae 843.
N Trochodendraceae 3 J, 8og.
Nepentbaceae 676, 1035. R Turneraceae 381,936.
Nolaneae 573. 100 5. Ranunculaceae 14,8°5.
Nyctagineae 645, 1025. Resedaceae 77. 826. U
Nympbaeaceae 47.822. Retna 581. Ulmaceae 764. 1056.
Rhamneae 218. 885. Umbelliferae 419. 940.
o Rhaptopetalacea~ 208,846. Urticaceae 764, 1056.
Ochnaceae 188,862. Rhizoplioracefl,e 339. 9I7. 7f (literature) 778,
Ochnaceae sense su. 863. Rosaceae 301 , 907. 1064·
Octocnemacea.e 872. Rubiaceae 444,950. Urticeae 77 5. 1063·
Octclepis 721t 1045. Rutaceae 174.854.
Olacineae 200, 87 t. V
Oleaceae 521,982. S Vacciniaceae 47 6 , 969.
Onagrarieae 373,93 1 . Sabiaceae 243,893. Valerianeae 454, 95 2 •
Orobanchaceae 589. 1009. ·Salicineae 797; 1068. Verbenaceae 630, 1020.
Ost,ea,ia 332. Salvadoraceae 526,982. Violarieae 82, SZ7.
Oxalideae 169, 851. Samydaceae 376 • 933- Vochysiaceae roo, 83 I.
SantaIaceae 730, 1046.
P -Sapindaceae 226, 890. W
Pa;vaeusa 190. Sapotaceae 512,979. Wallacea 800.
Papaveraceae 54. 824. Sarra.ceniaceae 5 I, 823.
Papayaceae 388, 936. Saxifragaceae 310, 910. Z
Papilionaceae 253. 895. Scrophularineae 583, 1008. Zombiana 62.7.
Passifioraceae 383. 936. Scytopetalaceae 208, 846. ZygophyUeae 167. 8so.
 GLOSSAR Y
[The main object or. this glossary is to give a synopsis of the chief terms used in. anatomical
description (including a considerable number used.in this work for' the first time) together with
their Gennau equivalents. A few definitions are iupplied, but more generally a reference is given to
a page in the Condtulinff Remarks or in the main Portion of the book, on which the term in ques-
tioll will be fonnd to be more or less fully explained. The glossary may also serve as a rough
index to the Coaclutling Remarks, and, with this object in view, rather more tems have been
included thaq would have been necessary solely for the purposes of the glossary. In a few cases,
where the English and German terms are practically identical, the latter have been omitted.
F. E. FRITSCH.]

A A ttenualed (i. e. gradually drawn out into a

Abi~tifqrm lzair~ (i. e. hairs branched like a fir- point, a.ppUed to haits), sich verj\;.ng'!nd, aus-
tree), Tannenbaumhaare (p. lJu). lanfend verjungt.
Acarodumalia (p. 1I3:Z). A urides, bbrchen; auriculiform (i. e. ear-shsped),
A dcular crystals, Kristallnadelchen. nadettonnige ohrc 4enartig. .
Kristalle (pp. 1104, Il 06); aekfllar fibres (ra- Axial wood, Axiales Ho~z (p. IT 45).
plzidine.r), Nadelfasem (p. lIS:;)'
Active cells (of the pith). Aktive Zellen (p. 1133). B
Adjacent ull.s (of stomata), Nacbbarzellen (see BQ(k-ca'llily (of stomata), Hinterbof (p. 1081).
neigkbquring cells). Band-skaped (jIattemd) axes~ bandtQrmige
Airenckyma, Aerenchym (p. Il50). (abgeflachte) Axen (p. 1166).
Air~anal, air.passa~, Luftgang; air.pores (= Barbed hairs (d. p. 379), Widerhakenhaare;
stomata), Lutlspalten. .. barbs, Widerha1:en.
Alumitl4·00dies, Toneroekorper (p. 1109). Bark, Borke (p. J 14-7).
Amj>ulliform (depressions, &c., i. e. such as are Barrel-shaped, tonnenfdrmig.
shaped like a flask), krogformig. Bars (the persistent portions in a scalariform
Anchor-kairs (cf. p. 379). Ankerhaare (p. III1); perforation), Speidun.
anchor-like skaggy hairs, Ankerzotten. Base (foot) ~ kair, HAarfnss (pp. logl, H24);
Allnual rinp (in the wood), Jabresringe (pp. base of sbaggy hair, Zotten{uss.
II36 , JI3i)· Baiin, Napf (apl,lied to part of the complicated
Annular, ringformig, ringartig; annular tra· secretory cells of Magnoliacea.e, Canellaceae,
clzddes (i. c. provided witb annular thickenings), &c.,see p. 1096); basin-shaped, schalenfOrmig,
Ringtracheiden. .schiisselformig.
Anthocyanin, Anthocyan (p. 1074). Bast:filwt.!, Bastfasem (p. 1152); IHzst-groups,
Antler·/ih, antle,-shaped (generally nppHed to Bastteile (p. 1 154); bast-wedgts, Bastkeite. (p.
trichomes), hirschgeweihartig, gcweibartig. u66).
Apical pres (of water·plants), Apikaloffnungen Bays (of nndnlated walls), Bachten..
(see Po 1086). Beam-like, balkenartig; transverse hams, Quer.
APiculus, Spitzchen •• _____ balken i beams of celll4lose, Ze1I8toflbalken~
4Pparent- medullary bundles, scbeinbar mark· Bel/Qws·lilu, blase balgartig.
standige GefassbUudel (p. 1165). Buollateralbundles (p. 1160).
Appendicular ridges (of stomata), Anhangsleisten oifaria! (structure of leaf). biracial, bifazial (see
(see horns) (p. 1082). p. 1086).
Aptous tzssue, Wassergewebe (see pp. 1077. Bi/4bed, zweilcnopfig, zweilappig.
108 7). Bladder-nairs, Blasenhaare (p. 1116); olCJdder-
Arcs (of fibres, &:c.), Bogen; arc·sJUzped, arched, like, blasig; raised like a bladder, blasig ab-
bogenformig; arched outwards, vorgewolbt. geboben; sw"llen Hke bladders, blasig aufge.
Arm-palisafk,.cells, Armpallisadenzellen (see p. trieben.
1087); arm-porlum (of two-armed hairs), Solryuidal (i. e. shaped like a bunch of grapes
Armteil. traubig.
Articulated, abgegliedert, gegliedert; artimlatt!d Bracket·ulls, Klammer~llen (p. 1100); Orathl-
hairs (cr. p. '51), Gliederhaare. epithelium, Klammerepitbel (p~ 1100) ;
Asddia, asddiftWm leaves, Schlauchblatter. Bracke/-hairs, Klamlllerhaare (pp. I I 17,1119);
A stefy, Astelie (p. II 57, footnote). bradu/-shaped, klammerartig,
SOLI!:REDER
 GLOSSARY

Brick·like, b&cbteinartig. Columnar, Auleo(ijrtnig. Aulenartig; (o/unwa,.

Bn"ltle, .8orst~; 6""'slle-l,ai,,s, Bontetlhur~: crystals (i. e. Jty~), Siillienkristalle (p. ll~).
!Jri.ttk-lik~, boning, bQrsteaf6llPj~. Combined d~/hif'lff and ,riaRdu/ur 1uzi17 (po 1]:15,
ErIlJR-Jlulped, piQst:lforznig; DNm,-It/le S4tJIW footnote). .
"airs, Pin.eltotten. ~mmissuresJ Kom.JXlisSuren; Clmlnissural sieve-
Bu/!Jow J,airs, Zwiebelhaare (p. I I '9); bu/~us tUDes (d. p. 396). Kommissutalaiebri:ihren i
a"a SfDQllm, zwic:belig angtSChwolltJ). a)tn"limmd strands (cr. p .... .f3), Straogvtr-
bindt11'lgec.
C C"m~rImPf'.r (of. palisade-ceIl, etc.). FliCber.
CtUdfoatitm, VetblkQJlg (p. JIll); falCijiedl CgmpltmenJ4ry (elts (of a. lenticel), FUltz,ellen
, verkallct. (:p. 1149)'
Clll/l)s~,?, Sehwiele. CI1"'POnenJ hairs (of tufted hairs, &:c.) , Teilhaarc:.
Ca.",{Jij()l"m strfJ1f4s (pp. 11,&8, I I ~9), C,.rnbiforDl- CDf"pos#e (I\pplied t.o pericyclic strengtbening
hiindel. ring), gern'idlt (p. uS',; cilIISjJqs#e (C4flt-
Ca,uklalm~.llairs, lCalldeJaberhaare (pp. I I l~, /'oWtd) -91em.-I1JI1..rS, zusalll!I1engeseuter H ob-
11I9, I I H); ta'lUieIIJora-lwi,.s f!! tlie sham korper (p. u6S).
ty~t Ka.ndela~otten (p. lUl). Cotn/Cllnd $)IIern-miUs, see c~mposile xy/n,,·
Cauut&MUl: /JotiuJ:, Kaut&'lhnkki:)rper (p. 1I09). mass; CO'lllpqund dO/hing IltZ;"I, zuummen-
Capitate, ko~&" kapfforroig, kopfchenartig; gese\zte Deckhaare.
capifate IIalrs (i. e· hairs ending tn a swollen C(J1tCrelcNtl, vecWachsen.
b~d). ]\opfchenhalttt (s« p. IJJ9 and ullder Co"kal, k~lfOrllli~.
.glandula, Aairs). Conjugate (e14, conJugierte ZeUen (see p. lo8j) •
Caps of cell~/C)se, Celh:l1o;ekappen. Conjunctive tissue (.i. e. that between tUld on tbe
Carlilaginqus, knorpeUg. inner &ide of the vascular b1llldles of the stele),
CtZU,.tU (bundles, t e, bllodles ~lollgitlg to the Zwischengewehe.
stem ooly), 5t.am.1Jle~getJ. C""na#nc !irme (in the veins), Begleitgewebe
C,U;[usiImJ, Zellfusiotl~, (p, 1089).
Cellu/IJy fro11Jinenc(S, ZellMcktr, Ztllhiigel. Constriction. EinsehnUrung.
Cellulose, Ztllstofl'; Ccllu/osf-(qjs I !ice 1llldtr C01Ua;ninr~ls (of cy;.toUtbJ), Triigettellen
cail; strands (Clams) Df (ellts/()st, Ztllstoff~ (see fitf1«J'Jls).
b.,lken. ' Ctmfffl/s (of a cdl), Inbaltsko~r.
C~nlric (leat-stl1lctute), cetltnSch (see pp. 1083, C(m4'fJ.UdUt (applied to pericyclic lttre.ogthening
1(86). ring). kontinuicdich (p. 1153).
C"(J~-llandz, Kalkdtiisen (sec liP. 1 ()S6, lJ28, C",uorltd (applied to ht.in, &:c.), ~wu,.den;
1133). irregularly (ontorted, }cRUS gebogcn.
CluztnlJerttl, gekamDl.~rt: Oht-, f'UJO-. nldfly-c!,aflt- Contrt.icted l.HUCN/lJr S7stems, kontrahierte Leftbiiu.-
hmi, ein-, .twti-, vieUcallJrnerig j (h<zm6e~ delsysteme (p. I:r5S).
.Ji[1re.s (r/J.IJ"wtrea o,,),staJfio,w). K.aJ::omerfastJ"n C"rd,d XfUl1NI14fS, uJDitrickter HoWtorptr
(Kryst.a.llkaD).mtrt'ascrn) (p. lIM); (namlJered (p. 1165)'
crystal..parendzJ'flUl. (chambered pareDchytlla Core (of a shaggy hair, &c.), Kern.
containing cry.tals). lCrystallkUllmerparen- Ct}r/t, KOl'k (p. I t46) i (Drll-warls, Korkwatzen
cbYIn (pp. 1108. 11,,:\), (p. 1133)·
Cil4/'(Zd~H·.ftk r~~~ (of the petiole), 'caT4c- COr0t/4/4 (applied to pApillae), mit Kronchen
teriatiqde I (p. 109.), vtr~en. _
Chi",~ (applied to "tomab.). Kamin (]>. 1085). Corlicq/ 'Oos('Ular /rultd!eS, rindenstandige Gefass-
Cla.114le, see duh·sA(Jllli. biindel (p, 1159).
CleafJfJgt (of xyletn-lllau), Z«:r kliitbmg, Spaltung Cracls a41 to drying, Trockenrisse (p. 10S8).
(pp. n66, IJ61, u68). Crat~ri/orml bech(:rionnig; qaleriJ""" gland;,
CI¢I ,s;*/~»I-"lIl.JJ, teTlclilftetet Hobkorptt (p. BecherdriiseJ:l (cE. p. 'no).
11 6 1). Cre,uUt (Dlargin of shield of pelt..te hair),
ClimMng hai,.s, l{limllJhUJt (p. 1J 1'1)' ansgerandc:t.
Closed (applitd to peticyclic strengthening ring CrtJ(et¢~s!tajld, tiichelfonnig.
and "ascwar rirlg), geschlossen (p. US'); Crest: (on Cllticle, guaru-cc::lls, &e.), Ka.JntJle
(hsin.! NmbralU ~!::orz:tt1t .. ). Schli~~ut. (pp. 1072, 1085); ~.r/;li.te, Jcammanig.
~/p!kjn.y hai,., ))e~)c (p.. 1 Jl$ et seq.: of a 'C."utar9U11· (see Pi J090); '~ri!tlU''lUi '.cel/s,
papUlo~ Ot v~cub.r ~,. p. I J 16); clqtA,'nK Cristarque-Zellen (see pp. 1091 and 1f.51).
flUrnl>,a)le (of intercefltl.lal' IIpace:s). l{,nt- Crt?/.·hai,s. Kropfhure (see pp. 557; 558) ; crop-
"U$kleiduDg; dQlhinr sIJaw hairs. ~. like, kropfartig.
zottCl'l (p. H:J3). Cr"dal~, kreut:fdrtnig.
Club~.1Mped. davaJe, lcelllenf6rmig. C",jI.f (&totnatAl), KryVten (gee p. r08,,).
Clusters, b~ko~; dust~rl(J crystak, Dr'Ll- C':1 st(l./--cells,1\ ristallzellen ; (r)lstal.(IIHKIQ1Ju"4I~ ),',
sen. KtysWldruse n (pp. 1.04. nos) i Aairs I{ristallc:ongloOlerAte; e'Ystal·A(#YJ, Kristall~
conJa.ining ~/uSlmuJ &'l'SIQ/I (cl. p. 740), haare (pp. Jl 07 I 1 tt S); crystals tJ/ gypsum,
Drusenhaare. Gipskristlllle (see p. 1076); crystal-,.euptacfes,
C,'kcl",,~,t//J. Sabl~eLtdJen (see p. 1081). Kristallbchalter (lk.( r07); crysJtJI~stJCs, Kri-
Cp/jpu./JJI"""- Ko1len~ym (po JlSo). 5tallsclllauche j c')'stal-Jand, Kz1stallsancJ (pp.
CoJ4lbl (I. e. large h&in secreting mllci1Age no.> 1106); c,,),l/(ZI.sc/ertHC/v"I'1(z, Kristall-
&tid found OD bud,), Colleteren. &klet'tnchytn (see' pp. 1091-" J 107).
I
 GLOSSARY 1:1'

C"7sttdlim pan""!} Kristallkomehen. EndDdtNnis, SchuU&cl1eide, Phloeoterma

C'7SltJllqids, Kristauoide (p. 1108). 1151).
CWM1Wl, kap~nfOnnig. ErJtOC7dic sitw-tuHs (d. p. 396)-
CfI"'7 (applied to trichomes), kraa5, krilp. Epi~nd«krfna/ rId1rIf1rk (d. p. 66), 'reseao "
CI/I'Wd (appJled to hteral walls of epjdmnal eooodermjque. '
cells in surfac::e-view), ~~ (p. IOio). EpiJluma, Epithem (p. 1086).
Cldicuiar htMb, Cuticularperlea; (small) (uti- Excrescence, Wucbe1'1lllg.
crU4r crDWn (of papillae), CuUculatktOnchen E~reIWn ofWQ.T, W~ulSCbeidung(p. 10?'!
(see p. 1°73); {uticular crests, Cuticular- excretory r«eptlkks, Exkretbehii.lter (p. 1095
kimme (lief: p. rOT::); ~uticular eli/MliNM, Exl,rnQI pandt, Aussendriisen (p. 1124); I,
CaticularrpitJiel (p. I J49); tutitultzr layers, lernaljore (of 5tomata), Zufiib.runp;~t.
CuticuiarlChlchten (p. 1013); cuticular pip. ExtrtJjiDral (exl1-an"ptial) n({t4rin, eltn
Cuticularzapfen (p. 1073); culicuilJr pro?{- Duptiale Nektarien (p. 1131).
tUms t Cuticularvonpriinge; cull'CU{Qr ddglS,
ClIticnlaclei5ten (p. I01~). F
C,fSlPlilAl, CYltolitben (p. II J I); fJ'S/~/ilhic. Fa<~'1 <of ayJtab), Flacben.
cystolitbisch; cyst6Wh . like, cystolithenahnlicb; . False jJttJins (of epidennal cell.), Schdntiipfelun,
t7sto/i/lJ ..cells, Cystolitbenzel1en ; C)'slolitll·lJod)', (p. 107a); false stales (d. p. 380), Scheill
CY5tolithtnkorper; (yslolith-jurmaiium, Cys~o­ achiilferchen.
lithmbildungen; cystolith.. flair, C)'Itotitben- Fat bot:iies.faJty bodies, Fettkorper (p. 1109).
hau, CYlI"toJiOlische Haare (p. 1 JI J). Feltta kairs, FUzhaan:; foil-like, filZie-.
C?.lt#spluws, Cystosphliten (p. J 113). Frw-}Jarred (applied to scalariform perforations
C,-stDo/ltl (i. e. uncalci6ed cystolith,), (pp. I I I I, armspangig", wenigspangig.
1113)· Fibre-Uke,jibrous, faserartig; filJrous (ells, Fasc
D tellen.•
DeU",itetl, &bgegremt. Filamen/(!US,jilij'onn, fadenformig, {adenartig.
Dendrift1t"n (applied to bairs) , baumartfg; FislWn, Zerklilftnng (Ie~ (/eavore); fonl,e
lJnv«/ted in fJ dendritic manllU, baWDartig terklU.I'tet.
...rnwdgt. Fissures due t(l d"..nK', Trocktnrisse (see era,
Dnulroid. baumarUg; DtttdrrnJ hairs, Baum- due to drying).
buret balllnartig verastelte HUTe (pp. 1118, Fislular (applied to the pith), robrig.
1l1.9). Flag-eIli/t1f"m (of haICI), peitscb<!lformig.
DmlUUlate (oC ~r(orations), gtriihnelt. Flat-armed (applied to cells of the spongy tisiQ,
·J)efreued, vertlefi, eiDgesenit; Je/f'tSJun (of flacharmig (p. 108?); flatl:! areAd, flac.bJ
pam-cells, &c.), Vcrtiefung, EinscnkllDg (p. wOlbt; flattened axes, see under IJand-sIJape.
Fkrilt strmgt", Bie~sfestig1t.eit.
108S)·
DialysttHc (p. 11 56). .lluke-all (of anchor·hatrs. d. p. :171), Zinkl
Digest.·w elaMs, nlgestionsdrasen (po I I 30). zelle. I
/)il"_ate, fingerfonnig. Foot (of a hair). Haarlusli (p. 1114).
Di.J4Ullitm (p. 1167). FOnlPHitttlte perforations, LOcbcrperforatiOi
Disc, Scheibe; dire-slurped, discoid, scbelben- (p. 1138).
formig; du~f1id glands (d. p. 710), Scbeiben .. l"'or~ea /Jai,s (d. p. 63), GabeIhure; {fir
driisci. r shagu llain, GabeJzotten (p. 112 3); f'"
DisSP(jatitm (oC the vascular £yStem), Zctspal- glands, Gabeldriisen (p. JUS).
IDDi', AuflOsnDg (des uitbiindeuyslemlJ) (p. FriaIJ/~. krnmOs.
ll!S)'
Front (Ow)' (of a. stoma), Vorbof (p. loSl),
DislDrled fwms (of acala.ri(orm perforations), Ftmne/--(t!!s, TrichtenetIen (l5ee p. ~087).
Kriippelformen (p. Jl37). Furftmzc~s (5Cttrfy, applied to hAiry comi
Divided xylem·mass, getei1ter Hobkorper (p~ kfdig.
t 165) j divilim-walls, Teilwlind~. Furrows (in the xylem-masa), Furchen (p. III
DMIU-shajta, kupptnfOrmig. juN'tJ't11ta xylem-mass; gefurchter Hohlti::
DtlUhli7fg, dup/icalim (of epidermis, hair.). Ver~ (p. 1166); .f"rruwing (of the qlem-m
doppelung; dt>.uhle cyslolilM" (tUt;,.-rystDlitlrs). Fa rehung.
Doppelcystolitnen (po 11 12). FUlij(Jfflt, spindelig; spindeJformig.
f)um};hl/~Il"pd, bantelformig.
G
Call1osle/ic (p. J 156).
E GtiaiinJUz/ion, Venchlebnung (pp. IOH, I J
Erutydif sin.~-Iulw (d. po 396), geJalinizeti, verschleimt.
EisMiai ridgu (i. e. on the outer 51lrraCt of the Gelalinow layer (of wood~prosenchyma),
guard-cella), Eisodialleisten. lertschicht (pp. I J.p, 1143); ~Jatinoru
EIt1!aJions of flie culide, Kutikutarerhebungen; bra.u (or wOod-prosenchyma). Gallert
,/evaliu,. (ohtomata), Emporhebung (p. J08 .. ). bran.
EtnQrgina/~. 4usgebocbttt:. Cmi'culaN (of crystals, i.e. bent ljk~ II. knee-j
amlKdtkd (of 'lie ins), eingebettet (p. 1089). geknickt.
E"'Jd)' (tlls (af pith), leere ~llen (p. 1133). Girder-scln"tnd,yllla, Tragtnklercnchym
Elfdrcled slomata (cC. p. 350). tlmkranzte Spall- sn.aped girders, I-Trager j gi1'iUr-sMje<J
tiffn llD gen. ger1ormig.
 GLOSSARY

Ginlk (of I1IbsidJuy celIs), GiitteJ ; girdk-lih, buiul/icale (margins of lea\'es)_ n.ch oben etn ..
KJ'nIIl'lllapN, giirtelfOrmig. geschla~n.
Glalld, DrUse; iland·II'kl, driisePJutjg. !tulualitJes, J'Yf'~gularit"es (of the cuticle, &c.),
Clandular d.lJdralUS (mrd,anl'sfn), DriiseDap- U nebenheiten.
parat: - dUc, Dril$eu5Cheibe' - ~,iI}Uli~m, Injilt,.ation (01 silica, &c. in ce1t..waU,), Einb.•
Drilsenepitbel ; - Izain, DtUsenbaare (p. ~el1lng.
H2o(); -ltaf-JUIIt., dru!lige lJlatWibne (p. bifolded (margins of leaves), eingeseh1Agen.
1130); -scales, Dri1senschuppen, -slzam b,ilial reciM (of petiole), Initiale (p. J094).
hin, DriUenzotten (p. J 1 ~9); - sputs, Drii~n~ Integummlal .flamr, HautdrUllen (p. tIl8);
fide (~ ll~p). integumtnlal (intt.(Z111lentt)ry) tissue (i. e.
Gmsl"fed (of ~pieulat ctlb), knorrlg. epidermis and hypodenn), Hnutgewebe. (p.
Grt$nu{ar (applied to the cuticle), ki)rnig (p. 1°74)·
1071)jgmnulaled, knunOs(appUed tGCJystals), InterjfJUar side$ (see p. 207), Zwischenblatt ..
gekQrnt; granuta(io" (of the eutide), Korne~ teiten.
lung. [,,(e"mediate 6u1klIu, Zwi&c.benbiindel ; -forti,
Cf"QQWa $:yiettNtt(MS, 6I!e/ur'rQ'fHtd xylem-tIlaI.1. l!:rse.tda.sem (p. Il35. footnote) '; _._ tissue {-
CrlJund-maJ,f, grlJund-wwk' (of the wood). c.onjunctive tisSue), ZwUchengeweb4::
Grundnta5se (p. lJ-4-I). /n(trna/ g/a.ntir, innete Drii$en, Innendrii.s.cD.
Cumfltl)s-es (p. 1101). (p, 1095 et seq.); internal KiaM14lar hairs,
Gum..,.esin, Gummiharz. itlnere, Driistlnhaare (p. 1096); ,'nkrna/ /Jairs
GypsUIn-nyslalt, Gypskristalle (p. 1016); rJlp- (Iri.ACJ6Ia.rls), innere Hll.R.re (pp. logl, 109:1) ;
SN"Np/te~s, Gypskugeln (cl. p. 7'). internal soft ~lUI. innerer Weicbbast (po 1159);
;",Iemal stri4licII (of epiderm&! wAIls), innc:re
H StreUutlg (p. "'72).
Interrupted (applied to ring or ~ricydic scleren..
Ha;r, base (l0P/) of, see under B; lJair-rells, chyma), Qnttrbrochen (p. II:;;').
Haarzellen j }'air-Q's/IJlit/t.s, Haa.rcptoHthen InJlr'Wt!flen likl/ell CoChairs), tilzig ver.6ochten.
(p. llU); ltair"'/ike, baarartig; hairy cquer- 1"ltl'xyllV)' }AI«m, holzstiindi~ (intenylares)
inl. Debaa1"UJlg (p. 1114 et Jeoq.). Phloem (p; U61) j inlerX)'lary vascular
HtarlJt»DOd, Kernhou (p. 1136). Impales, holzstandige Gefassbiindel (p. n63).
HeluoilJ (applied to shape of ~ystolithl), sclme- lnirlUambial (i. e. on the inna &ide of the
~kenforn\ig.
:catnbium. liCe p. 113). '"
Hemuolf€etUn"c 6undles (o( the petiole), f Fa.i~ Itt.lramdulia", (bundles, ct. p. 5g i), intrame..
sceaux Mmioonoentriques' (Bo~ygu.e.) (see p. dullate (BUndel).
1094)· Intramura.l.(fa"as (cf. pp. '54 .....86), Zwischen-
HemitrDpic ",stall, Hemitropieen. wnndiiriisen (p. 1100).
Hettrr1~dU.r (applied to the pith), heterogen Inlraxylary (1. e. on the inner sidt. of the ~ylem)
(p. 1134)' plr.l«tIl, &c., inttaxylires Phloem. (p. n59),
nmgNtU )!f~airs. &c.). Gelenktelle. &c.
nip.ftx':e/i (i.e. ho.rseihoe-&haped), hufeisen- lnva'u f)tuntation, nmgekebrte Orientierung .
fomug. .{trboluttd (of gl.a.n.dular hairs), eingeroUt.
HIJf1tIJ,ft7UtnJl (strtlcture of the teaf,. see PI" 1083, Irregu/ari#u, &ee illequali#es.
J 086). hornogen; "MI/DC-meNS
consisting of empty Of active cells only),
(pith, 1. e. oDe EsI<zntis of 10(1 ~a.rl (applied
phloem), W'eichbastinseIu (p. u61).
'0 inter.J:yl.ary
bomogen (p. II3.f.).
Om·like, kApttunfOrmig.
H~. htn/like a (applied to hairs, &c.). baken- J
fOrmlg geloriimPlt j IzcoJud hairs, Hakenhaare Jagged (nmm, small (at the end of a leru-~piUa),
(p. 1117); !wok-shaped, bakenformig. u.cl:iges Krol'lchen; jagged jJro1!ltfUlUtJ,
Borium/at (llpplied to i:t:lts of medullary raya), Zacken; jagceJ 11Iargins (of epja~al cells,
liegeJ1d (p. 11 .....). shields of peltate bain), geuckte Randel.
Horm (eqw'1alent to o\1ter appendicuhu ridges Joint (of hairs), Gelenk; jlJintoo«lls (t-. g. ()f
on guard~lb), Horner, Hornchen (p. 108:a); sympodial hain), GU~enellell (p. t 120).
korn-&ut (1. e. gtoupt. of thicken.ed and.ob-
li~nLted si.eve~tubes.). Hornbast; Mm, CUNJd K
Iii!( 411 homronnig gektiirnmt.
HurtiJ$ (projections on cell~w.l1s), Bw:keln; }.;~z (on cuticle. \t"aUa of haj", Sec.). Knotchen ;
"""'1. pt'ojulinc Ilk
'pritlgend.
41, buckelfonnig yor. ~-lih knopfardg, kootchen:utig j huNJ-
s/uJprJ, koopffotnlig'; lm~1 title II) Mirs,
U"tlaJitAiu (i. e. s~ial organs excreting Jiqwd Haarknotcben (pp. I f 10, U II).
water), H)'dathOden (p. 1016).
H),JIIdt,.", (p. 10 76). L
HYs/vW_ynrmcuJ, hysteroly.igcn. Laci"," (of ~a~, &:c.)) Lacin[en. ZipfcJ;
glandula,. 'Iuinw, Dtiisenzip[el (d. po 337);
1 /ac"niale (dh·fded into narrow tabes), ter..
Incnnli"c. knutig. &cbUtzt.
.l*llrwa (OUillod of oeUs), gebachtet; ;murvea Lluunar (applied tc) rpongy tissue, &:c.), liIckig,
(tips 01 leaves), eingebogen. lakuno~.
 GLOSSARY (179
Lagt!!Iij'orm, flaschenformig. lI/em6ranous, hiiutiC'
Lamllla, hafling lhe /qrm - ~I a, Jamellenformig, lI/tsh·liRe spaces (in the spongy tissue, cf. p.
lappcnformig; Jamellaltd (composed of thin 2'l9), Maschenrliume.
platei), blitterig, i:etcbkhtet; (a.ml/laid cork, }.{etalradual ptJrtnCh)l11w. (p. 11.3).
LamelIenkork (p. 11048) i /rallS"WI'se lanulla· Middk /arer (of leaves and glands), Mittel-
lion, QuerlamelDellUlg Ca. p. 532'). schicht (p. 108i).
Ltuh·like, pfriemlich. Mmodinu crystals, monokline Kristalle (p.
Latex-receptacles, Milcllsaftbdlaltel' (pp. 1096, 11 °4).
1102, lJ03): /alu-sQ.(s (- latidferoul sacs), "Ionostelk (p. 11,56).
Milcb.saft:scbIauche (pp. 1096, 1097). MoruUJse g-laluis (cC. p. i4-S). Maulbeerdru8eo.
Lati4/ertIUS can(J/s, Milchsaftgaoge lei. p. 4(2) : Mua·lage..canah, Schlei~ge (p. 1099) ; -<(rlJi,
-&eI4 (non-articulaled ~i&iferou$ (uk:), un- tus, Schkiml\icken (p. 1°99); -cdlt)Sd;Lleim.~
gegliederte MiIchSAftrobren (p. J 10:1) ; - hairs, zellen (p. I~) i -rlands, Schlei_mdriisen (pp.
Milcbsafthaare (pp. I JIt. II 26); - sacs,milch. II 28, 1 133); ·"ai'rJJ ..8chJcimbaare (pp. 1127,
saftfubrende Zellen. Milchsaftzellen (pp. 1096, 1128); ../ocunae (cr. p. 752), Schleimlalnmen;
1(97) ; - t~e.t. ~lilcb.rohreu, Milchsaftrohr~ ; ·pores (cf. p. 336), Schleimspalten.
-1J(ssels (arliculated fatirifi'-ous lubes), geg-lie· Jlmi!agilWUs epiaermal (ells, verschleirote
derte MilchsaftroMeQ (p. 1 103), EpidermiszelIen (p. 10,4); m#Cilag;mmscells,
Loyers of mucilage (subepidermal), Scbleim- veTSChleimte Zellen; mucilaginous cork,
schichten (p. 1088). Schleimkork (p. I J 48).
.untice!1 . (cortical jXlres), Lentizdlen (Rinden- Nultlple cryslals (cf. p. 9;;), Verwachsangen
poren) (p. 1149)' von Eiru:elkristallcn.
Lmii&ular (i.e. lens-shaped), linsenfoJmig. Afultisviate (i. e. compo&ed of 5e,·etal ro\\'111 Of
Lepi,Ju (small scales, q. v.). , layers of cells), mehrzellreihig (applied to
Li6riform tiswe (Le. composed of narrow cells hairs), mebrschiclttig, mebrreibig (applied to
with thick woody walIa), Librifonngewebe medullary rays, epidermis, &:c.).
(p. l135, footnote). Afuri/onn parench>'"14 (i. e. made up of brick-
Lia-cdls (of glands, e.g. p. 585', DeckeJzeIlen. shaped cells, d. p. 68:1), Mauerpart1lchym.
Lignified lamella (in wall of guard-cells), Holz· llZYnuz'n.cells, Myrosinztflen (p. ICl9i).
lamelle (p. 108:1).
Lil"(ate (i.e. tongue-sbaped), zungtmf,-}Tmig. N
LilAQC)',s1 (i.e. the; cell containing a cystolith)
Lithoqlt, Trag~rzelle (p. nul. Neck-cell (or hairs, cl. pp. [053. 585), H.1szel\e;
Lobed X)'lem-ma.rs, gelappter Holzkorper (p. neck-/C!1,'()ft (of gl&nds, d. p. 287), HalsteiL
1166); lo6e-like, tappenanig, lappenformig. il/eetanal glands, Nektardriisen (p. 1131); "eel·-
L«ttli (of chambered aystal-fibres, see under an'al spots, Nektarflecke (p. J 13.2).
duzmbered), i"acher. Nataries, NelctaritD (p. IIJo). ,
LtmriludiNlI fiJ.fUJn (of the stem in anomalous Nt1'gft.60uring ce-lis (of stomata), N!chbanellen
growth), Lt.ngupaltung (p. 1166)~ (p. 1018).
L""1e (applied to spongy tissue). loclcer. ,Nodosl, "Mula,- (applied to thickening), knotig.
LlIWtr scale (of peltate bairs, Sec.), nnteres Non-arlkulaJea, ungegliedert.
SchUlfercben (p. 1121). Jtuc/tar cryrtall4ids, ZellkcrnkrystatIoide (p.
Lumps (on cell-walls. &:c.), HOcker. 1109)·
Lyra/e (applied to shape of two--armed hail'S), o
~ geataltet. Okpltt'cQl, verkebrt (umgekelut) kegelfonnig.
M ODlale-SjJheri(a/, abgetJacbt kugelig.
OctaAetfra/ crystals (p. 110i).
Mal/ormed, distorted (perforations), Kriippelfor- O,u-DNlUd hairs, elnannige Hs.a.re (pp. 1118,
men (p. 1137); maiformed stoma/a, verbildete II 19); one,sided f;(meS "1 ,f'f'1IU"th (d. p. 608),
Spaltoft"nungen (p. lUO). einseltige Zuwach!lzoncn.
Ma/piglJian "a;rJ (i. c. two ..armtd hairs, q. v.), OplllJue dots (in the leaf), undulchsicbtige I'un~te
Malp~~bjscbe Haare.
Mammijo1'ln (1. e. teat-sbaped). mammilliform, (p. 1°99)·
titzenfonnrg.~ ~ - Oten slcrelory cavilies, nngeschlossene Sekret·
Hicken (p. 1100). .
Maflf"jltikea (applied to ancbor·hairs), me!!r- Opmt'ngs (equivalent to peiftwatioHS, q.v.).
dnkl/l. Ort'jUe, o.ftio/e (of a nectary, &c:.). Miindung.
MIZI'ginal-[w/, Randbast (p. 1090); marginal Ostium.
Ittl. Randtupfel <,P' 107a), Outer respiratory (O'l'ily (of stomata), aussete
AftZrking (of the cuticle), Zeichntmg (p. 107J). A\emhUl\le (p. 108S).
Meckanical sup}"'" (of a stem, &c.), .I'"estigung.
Medullary bundleJ, &e., markstandige Bunde1
(p. I 151), &c:.; medullary rays, Markstrahlen P
(p. 11 ..3); medullary s"ea/A (i. e. tracheides Palisade_-flands (d. p. 306), Pallisadcndrlisen.
lanning a circle rOlUld the periphery of the Panalll'iJorm (biscuit - sbaped, fiddle - shaped),
pith), MarUcheido; medullary rQ)I·paren- bis\;.tlitf6nnig, geigeJllonnig.
cnJ1ma. Markstrahlpa.renchym ; Pkdullary PapiUiftwm (applied to hairl), papillenartig.
/iss_e, Markgewebe (p. J '33). PalillPse dijfireNialim (of the !pidermis of tbe
Me1lt6ratU·mudla~, Membrauschlrim (p. 1098). lcaf).. papillOse Ausbildung (p. 1013).
JJ80 GLOSSARY

Partlinzd"al Jartnd,ytna (p. 1143). R

P~"~s}/~alll (of veins), Parenchymscheide J?Qdiat. (structure). Jtrahlig.
(p. 1(89)' , Roised doIs (cf. p. 635), ~orspringcnde P11n1ste.
Partm"-~fII4I""1 jeriLyrie (p. tJ ~ :I ). Rumpart (o! cella, &c::.) , Wan; ,-ampart ¢
Pu.r#iJ/ ~atUh (of IplNJ thickening). Teilbander. cutick, Coticu1arwall.
Pottllifrwm. bec.htrl'onnlg; patelli/om. ZlMuls, }?a/~iJes, Rbaphiden (p. 1104); rQjJAitk-celb.
&huueldrUsen (cl. pp. 60:1, 60.. ). Raphidenullen ; ·nealles. Rhaphidennadeln j
Pearl-g(antis, Perldriiscn (pp- 11 :110, I J :19). .sacs, RhApJaiden$chlan<:he (pp. lC?98. 11°4);
PedtJtQI (of 11; hair), Postament, ~ke\. bndks fl rapMdts, Rh.tpbid~bilnde} (p.
Peg1, ~p{tn; PlC../w, zapfenaJtig; Jle("sllajed. 1106).
tapfcmformlg. R"phidines, see adcularjiDres.
Pelludd tl#l1, pe\111ciae Pankte (p. I~): pel. R4Jfed bundks (in th~ petiole), • Fa1sc~au rayon-
",dd, 'W.saetbe~. • . bes' (of Bouygues, p. 1094)'
Ptltatt &chlldJOnDlg; -glands, SdulddriJlIeh (p. R~yJ (or stellilte or peltate balrs. &c.), Strahlen;
IIJS); - }'"irs (seaks, It/ian). Schildhaare ra7..(ells I Strahlent.ellen; ra.r:/onnflliDn, Strah-
(Pp. II I~, 1118, II 19.11:110); -uaks, SchUl-
teiehen; false ~lIaie scaks, ScheiDIChiUfer-
lenbildung; rar-I"rti:"n, S~lteil;
rayJ. melmttalilig~ \'lelstrahhg.
"'an,-
eben. Buurved (of hairs, &:c.)~ umgebo£en.
Pmidllale (applied to hairs), pinstlformig. }?e"i/tIf"'In (Jddney..ahaped), D~rormig. (~
Perf~atiMIS. Perforationen, Getassperl'orationetl, Rluau de SMUUtf. strengthening aetwork r
~h1mchungen (p. \137). 1168).
Periaxial fj}()(}(/, ptriaxiales Hob (p. 1 '45). Bes~-/r(I(Aeidts, Speichertracheideo (sec
Pmo/ck, J'cricyktl (p. II53). sttW~Nm'Aeidts).
PerilfUdulIP'7. Le. in the outermost rone of tbe , Resin-iJlt"Ous (t e. containing resin), harzfUhtend.
pith. RninoC)'lis (d. p. "Q:I), ResinoC::Yl5ten.
Pdula,. g/pl(/s. BlatllticldrUsen (p. 1131).
PJull«ler", (p. 1149)'
Rupir/Jtrny '"vity (o($torn..ta), Atemho'ble.
Reticwlatt, neturtig; rtlkulflle perfin-atil,!s (cf.
PMI/Md-UPs, Phelloidsellen (p. 11+8). P.'561), Nettperforationeu. (p. 1137); rdinda/(
Pi[qtm·ul6nds, Ph1.oemiu.seltl (p~ (161). Jeyktn-masr (d. p. 2'1), 1l1tlstriclttcr Holz-
.Pit smail P,'t (iD leaf, &c.), Grilbchell; piJ (in kiirpeJ; rttUlilaIllJIluft1'(J~ /iii" (i. e. in
c:dJ-walf). Tiipfel; /JiI-<anals, Tilpfelkanal<; which the active ctlls (orm a network betl¥eCn
Porcnkanaie; pil-lW, grubig. £tii\icll.ena.rtig; the empty ones), neufQrmig heterogeaes Mark
piI,pu/o"ati4nJ (cr. p. 333). "I'iip{clperfora· (d. p. lJ3-4).
t!oPCIl. R etijtWtn, gitterformig.
PlIIcenti/orl' (r.b.a.ped like a S.,t ake), kue'hen· Pe/(J1'I.JAaptti nain, Retortenh.ure (p. 11'9).
fOrmJg. Reversed orimta/itm, verkehrte Orlentienmg.
P/utJs (e.t:. of fibres), Geflec.ht. J?~lule (margins of leaves), tuugeroUt.
Pl».l ,[1J1fS', Kallu:Dsru.l.lung (p. 1111); plug /{""'ulUt/ra, Rhoillboeder t Rendyoed~, (pp.
0/ silica, KieseLa.uafiiUung (p. 1110); plugpd 110.., nos); rMm!J4hedral, rhomboedrisch,
slqr1t4la, ftI'Itopfte Spaltofi'nungen (p. 108S). hendyoedrisch-
PtJl)tstelic (p. ll56); "/Jartnt jolysilly (po .Ki/J/Ied tutl, gtrippt~ Achsen. (p. 11(6).. .
IlS7). BiIJbmw},aftd (apphed to bans). bandfonJU2'_
Pt1f«" (applied to part of the complicated fUdze, Leiste; ridg~/ile (of thkkeningsh lei-
secretory cells of MAgnoliaeeac, Cane11aceae, &tenfOnmg.
&CO, cf. po 1096), Beutel • /(ing-WA, Ricgelborke (p. J I.n); ring-wtJOd
PflrlJdlr;y (applied to crystal-sand), meblartig. (d. p. :.06)1 Ringholz (p. U-45)-
Pri~ cttrlex (p. 1150). Binp 4"grO'lJ)III. Zuwa.chsringc (p. 1163)•
.Pro/ecliqm (of hairs, &c.). VorspriiDge. r.~s. small, SUi.bchen; rod-tellI, StabuUen (p.
PtWifn-atitJm, Wucherttngen (cf. excresunu). I 155); rllil-cell-sc!erntcAjIfM, Stab&ellenskle~
Pr_INmCeJ (on tell-waUs, &c.), Vorspriinge, retlchytn; nki"/ile, Itabc::benutig; rod-
Hader. sAa}ed cr')'slals, stii.bchenfOnnige Krlstalle (p-
PrtJnr-c.lls (cr. p. 304). ZAckt;ttel1en; prongs 1(06).
(of anchor-hairs, &:c.), Zacken. ,R,,/{d llatlts, Rollblattc:r (pp. loS4, 1087).
~e1Js (ct. p. 710), Strebez.ellcn. Gosette (0£ epidermal t;ells around stomata, Sec.),
Protective shtatk (of vaacDlar ')'item), Schutz- l<tanz, Rosette; likt. a. f'OSttte (I'O$t.tU- l iA,t),
ICbdde(cf. pp.l089,U51) (.ee also nuibliermiJ). kranzlOrmig.
PrDlrusiOtU (of w&lIs of hairs, &c·.) , Ausbuch-
tnDgen. Auutiilptmgen.
PTDluhertm(ts (of cell-walls, &c.), Protnbe-
raDteIl t A ttSstiilp1U1geJl.
P.stuti«dlll/tu"t ~knahnlicll..
Pulvim", Gelenkpolster, Sch well polster (p. 1094J.-
Pu"ctat( (of cell-walls, &c.). punktiert; punell-
j()rtIC, pttnktformig.
PynJQrm (pear-shaped), bimformig.

Q
Quaelnliu cry:tn/s (p. I ro.,).
 GLOSSARY u8t

penhoftupfel, treppena.rtige Hoftupfel (pp. 1 139. Speckled (appearance of lea'Ve1), gtsprenkelt.

J .-4°): - sieve"jJlaUs, leitafonnige Sieb- Spllturite:r, splure-cryslals, Spbaerire (pp. 1 r04.
platten (p. tlS4). .105) ; sphaerocrystalli1Ut spb.a.erokristal-
SciJIei, Imail (leJide.s), Scbulfercheo (see under lioiJch ; Ipluze1TJC1')'stall, sphere - crystals,
peltate hairs); scales of lime, Kalk&chiippchen; Spbaerokristalle (p. II 08).
scale-wk, Scbappenborke (p. J 1+7); scale- Spicular cells, 'Spilcnlarzelleo (p. 10(1).
hairs, &:hnppenhaare; sctJIy, schuppig; scale- Spines, Stacheln (p. n 15).
like, ilChiilfetcheruntig. Spini/()rm, spiessfonnig; spif17 hairs (d. p. 22),
SlIlir;o/ysilelWus (i.e. arising tirst by separation St8.chelbaare (p. Il33); splny (ol clustered
of <::.ells, bu.t then enlarging by the actual crystals), stacheUg.
breaking down of celli), &ehizolysigeo. Spl;,"·wtXXI, Splint ~p. I I 36).
Sdlrnun},m.a/pus inveJtmmJ (of vascular Split (cleft) xylem-mass, zerkliifteter Holzkorper
buudIes), Sklerenchymbeleg(p. 1089); sdn-en- (p. 1167); spHttinC up (ot xylem-mus, &c.,
cllymt:/(IUS pericyde (p. IJ53). i.e. cleavage), Zerli::liiftung (p. 1166 et seq.) ;
SnttiJorm (peltate), schudformlg. splitting (of the stem, !ce.), Spaltung_
Secondary vascular /JUndks (p. 1164); secondary Sponq cork, scbwammiger ]{ork, Schwammkork
grO"PS cf 'WQ(J(/ awl &ut, Itkunda.re Holz.balt· (po. 1148).
gruppen (p. u6g). SpreaJing-anned (spongy celIs). gespreiztatmig
Secrelory canals (passages), Sekretgaoge (p. (p. 108 7).
I101); - cavities, SekretlUcken (p. :£099); Starch·sheath, Stilrkescheide (p. 1151).
- ctlls, Sekret%ellen (p. 10g6) ; -JfKUnae (cC. Stele (p. IIS6).
p. 7{0), Sektetlakunen; - ,·eclptacles, Sekret- Slellal~, sternformig; slellate "airs, Sternhs.are
belWter (po 1095); - sQts, S:eJcretschlanche (pp. 1115, 1118, 1119, lUI).
(p.lo¢). St"eilks (i.~. a lignified &lrenglhening cell),
Semilunar, balbmondfonnig. Stereiden.
Semi-Malpighian (".e. me-a,.."w) "airs. haIb- Sterel)m-sneal4 (i.e. strengthening sheath ohelna r
malpighische Haare (pp. IllS, 1119). leaf· margin , &c.), Stereoinbeleg.
Sensitive luJin, Fiihlhaare; sensiH-w pz'ls (on Stinging bristles (d. p. 379), Breonbouten;
tendrils), Fubltiipfel. stinginc hairs, BrcDnhaare (p. 1118).
Septate (of fibres, &::c.), gef<icherl, septiert; Stomatal apfi1.ratus (i. e. the gua.rd-celis together
septation, Fachemng; septation of the pith with the subsidiary cells, when present), Spalt-
UJillwut sclerosis, Facherung des Markes obne offnuogsapparat (p. (078); - groul.s, Spalt-
Skler~ (p. 11M)' offnnngsgruppell (p. (084); - jlts, Spalt-
Se",'-aJed, gereiht. offnnngsgriibchen (p. loS...) j regarding. other
Serpenlim (applied to hairs), s".hla.ngeoartig, terms applied to stomata, see p. 399-
scblangenformig. Sione-cells, SteintelleD (p. 1°90); 11000-4lled (wi
Serra/ed CUlicular crestl, gebirgskammlibnliche (. c()7'k made up oj slone.cells '), SteiDzel1enkork
C uticulnrleisteD. (p. II48); stonl-C{lrk, Steinkork (p. HiS).
Shaft (applied to stomata, d. p. 493), Schacht. St()rage·tracheicies, Speichertracheiden. '(po I 09 ~).
SkaEgy hairs (fJill,), Zottel\ (pp. nIS, Ita3); Straight (applied to WjlUS of epidernu.l ceU. in
d'iuu·nc kairs '!f IhI slJaggy tyjJe) Deckzotten surface-view), gerade, geradlinig (p. 1070).
(p. II2l)· St,-alifi:d, geschichtet; stratiform (arrangement),
SlutJIlling (of the veins with crystau), PHa. schlcbtenweise (Anordnnng).
sternng (d p. 1108). Sti-englnenz'n,f ,,"n,f, Festigungsring.
SMeld (of a peltate hair), Sclilld. Stretclzing. Zerrnng.
S"",-11UmDered(sclerenchyma,&c.),kurzgliedrig. Striat,Strichdchen; striate (thickening), strei6~;
Sickle-liRe n.airs (cf. p. 379), Skhelhaare. striated, gestreift; striation (of cuticle, &e.),
Sieve1ie/ds, Siebfelder (p. 1154); st·eve,,/it·/ilu Streifang.
strudure, sieve-plale structure (of bOrdered Sln"gae (= bristle-hairs), Striegelhaare.
pits), Siebttipfelstruktur (pp. 1139, I I.~o). Sln·ulations, striulae (in leaves, &c.), Sm·
Silica, Kieselsliure (p. 1109); silica·bodiu, chelchen.
Kieselkorper (p. 1110); silica-plugs, Kiesel. Stj'/()ids. Styloiden (pp. 1004, 1106).
fullnngen (p. II 10); silica-sacs, Kiesel- Subcentric (leaC·strtlctnre), snbzentrisch (p,.1086).
sthlliuche. . ' .. - Suhpapillose (epidermal cells), snbpapl11iis (p.
Silin:JicaiUm. Verkieselnng (p. 11°9): lilid/Ud, J07.~).
verkieselt; nlutIIUS c01W'eticns (cC. p. 675), SulJsidiary c~lu (of ltomata. and hAirs), Neben-
Kieselkonkretionen j siliceous Ix(retilms, Kie· tellen (po 1078).
selansscheidnngen (p. II J 0). Suhulat~, pfriemenronnig. pfritmenartiggestalttt.
Simple clothing- hairs, einfache Deckbaare Successive rings 0/ bulzdles, successive Biindel-
(p. I I 15); simple-pitied (vessels, &c.). eiofach tinge: sucttssivt ringJ of grnJtlc, iticCessi.«
getUpfelt. Zuwachsringe (p. 1163); JU(ctssi:Je unes'l!/
Slidinc'grMlJth (i. e. growth leading to the inter- gr()Wt", successive Zuwachszoneo (p. u63)·.
cala.tion of new elements between those a.lready Slink (of stomata, &'c.),eingef.enkt (see p. 1085).
present), gleitendes WacbstaIn. Supporting network,' resean de soutien' (p.1168).
SIit$, Spalten; Ilil-lilte (-slulped) #11, Spalt- SupratJasal (i. e. sitnated opposite the xylem-
tiipfel; slit-shaped, Bpalten1brmig. groups in the root).
Solita'7 €rystals, Einzelkristalle (pp. 1104. IIOS). SW~II;'IP, Verquellungen; S"llJlIlIen, verquollen.
Spear-shaped, spiessfonnig, spiessig. S)'JlljDdja/l)' branched lIairr (p. 1120).
, . 'I~ I 1
l •
 1]82 GLOSSARY

T Unilateral (one~sided) 6Qrdered pils. einseitige

Tabular (applied to !)hape of cells, &c.), tafel· Hoftiipfel (p. 11 <fo).
formig, plattenfonnig; tabular cork, Platten· Uniseriate (i. e. composed of a single rQW of
kork (po 11 ... 8). cells). einzellreihig, einreihig; /Ji-seriate~c.,
Tannin·cclls, Gerbstoffzellen; ·contents, Tannin- zweheJJreihig, zweireihig, &c.
gebalt, Gerbstoffgehalt; • idi'obla .•ts , Gerb· Upper scale (of pel tate hairs, &c.), oberes
stoffidioblasten (pp. 1095, 1098); ·S(US, Gerlr ~chii1ferchen (pp. 1121, lI~l).
stoffscbHiuche. Upright (cells in medullary rays), stehend (p.
Tanniniferous (i. e. containing tannin), gerb· 11-1-4).
stoffhaltig. Urceolate, krugformig.
Teeth. spines (of a crystal), Zacken. Urn-skaped (trichomes). umenfOrmige Trichome
(p. 11l9)·
Temile strmgtk, Zugfestigkeit.
Tertiary thickening·layer (in vessels, tracheides,
&c.), tertiiire Verdickungsschicbt.
Tessellated epitkelium, Pflasterepithel.
. v ,

Ve/ltral (ventricose) part (of glands, d. -po 319),

Bauchteil. .
Tdragona! (crystals), quadratiscb; tetragonal Ventricose (applied to shape of cells), bauchig.
octahedra, Quadratoktaeder (cf. p. 110.). Vermiform (i. e. worm-shaped). wurmformig.
Tiers (of candelabra·bairs, &c.), Etagen, Stock. wurmartig.
werke; tiff-lik~. etagenformig; lier..Jih slrm- Verruccse (wart-like), warzig.
ture of the w()()ti, Etagenbau des Holzkorpers VertiCal d,'recliM (in the axis), axile Richtung,
(p. J J45); lier-like cork, Etagenkork (p. 1148). Langsrichtung. .
Tomentum, Filz. Vertically transcurrent (applied to veins). durch-
Torose (thickening), wulstig. gehend; vertical transcurrena, Durchgehen
TQrlu~j (of .ha.irs), hin nnd hergebogen, ge-
(p. 1089).
schlangelt. Vesicles, Blaschen; vesicular, blasig. bla.senartig;
Tradteirks. Trll.cheiden p. 1135, footnote); vesicular' hairs, Blasenhaare (pp_ 1076, I 116).
bruadmed (enlayged) terminal tracheides, er- Vmel-petforatibtts, Gef&ssdurchbrechnngen (p.
weiterte Endtracheiden (p. 1093); tracheide-
sder'endtyma (cf. p. 446), Tracheidenskleren- 1137)·
Vestibule (of stomata, equivalent to outer respira-
chym_
tory cavity), Kamin (p. 1085).
Translucmt. durcbscheinend.
Vitreous (i.e. transparent as glass), glasartig.
Tramparent dols, dnrchsichtige Punkte (pp. 1074,
1088,1091,1096,1108, 1130); -lines,dnrcb
sichtige Linien (pp. JJ01, HO",); - ·stri'uI(U, w
durchsicbtige Strichelcben (p. 1 J08). Walts, Warzen (p. 1115); 'Wart-like (see verru-
Tr'ullrated. abgestumpft_ (Ou) , warzenartig; wart-shaped, warzenformig.
Tubercula/e, hockrig; tubercles, Hocker. Waterpores, Wasserspalten (p. 1085): -rtc~­
Tllberous, knollig. lacIes, Wasserbehii.lter j ·rnervoirs, Wasser-
Tubular (applied to cells, hairs, &c.), rohren- speicher, Wasserreservoire (p. 1091); -storing,
artig. schlauchformig, schlauchartig. wasserspeichernd_
Tuft, Buschel; tuft of rays, Strahlenbiischel; Wavily folded (walls), wellig gefaltet; wavily
tufled, biischelig ansgebildet; tufted hairs, nodular strlatitm (of the cuticle, cr. p_ 70),
Biischelbaare (p. II 2 I). wellig-wulstige Streifung; wavy (applied to
Turbinate, kreiselfOrmig. hairs). krans.
:Twill-crystals. Zwillingskristalle (p_ 110...) i IVeal, Wulst.
twi1l10r'ms, Zwillingsbildnngen; -stomata, Wedges of bast, Bastkeile (p. 1I66); 'Wedged i,l
Zwillir.gsspaltoffnnngen (p. 1085); -trichomes (applied to position of hairs, &c_. with refer-
(cf.p. 16), Zwillingstrichome; -glandular "airs, ence to other cells), eingeklemmt, eingekeilt;
(d. p. 484), Driisenhaarzwillinge. Wedge-shaped (in tke fot'm of wedges), keil-
Two-armed hairs (Malpigbian hairs), zweiannige fonnig.
Haare(pp.1I17, 1119, uu). IYhip-l£ke hairs, Peitschenhaare (cr. p. 401).
Tyloses, Thyllen (p. 1137). Winged axes, gefliigelte Achsen (p. 1166).
lPood-jibres, Holzfasern j wood-parenchyma,
u HQ)zparenchym (p. 1J43); 'Wo()(/.prosenckyma,
Hobprosenchym (pp. J13S, footnote, n"p);
wood-vessels, Holzgefasse (p. 1136).
JVrinkled (applied to cell-walls), runzelig.
 n ERRA'TA
PAGE 17, line 7 from bottom, first half should read from being wetted, and to form an
I

air-chamber '.
::!4, thirteenth line of third paragraph, for' Bejuco de aqua' read' Bejuco de agua '.
57 ,~line 2 of last paragraph, insert a semicolon after' Leger '.
64, top line, for • place' read • plane '.
7 r, line 9 of first paragraph, for ' spread ' read • spreading '.
75, description of Fig. 19, c shoul.d read' C. teylanita, L.'.
95, last line, for' resent' read' present '.
I IS, last line of first paragraph, before the word' recorded' insert' has been "
136, first line of second paragraph, for . developed' l'ead ' deposited'.
141, last line of fourth paragraph (under 5). for • Dipterocarpous' read • Diptero-
carpecus '.
192, fifth line of last paragraph, for • see note 2, p. 21$ , read • see note 1, p. 190'.
201, fifth line of third paragraph, insert a semicolon after • hairs'.
204, description of Fig. 48, E-F, for' Goniocaryum ' read' Gonocaryum '.
20" second line of first paragraph, for ' Ganiocaryum ' read ' Gonocaryum '.
zoo, last line but one of first paragraph, for' Chtamydcarya' read' Chlamydocaf'ya'.
2 14, for ' Hippocra teacea ' read ' Hippocrateaceae '.
255, ninth line of first paragraph, after' Platymiscium' insert a comma.
280, twenty-second line from bottom, for' Spalky/obus ' read • SPatholobus "
334, footnote, for' Brunai ' read' Bruni(l '.
343, footnote, first word on second line should read' Gyrocarpeae '.
366, last line but six of second paragraph, for' Ptermand1'a ,. read • Pternandra '.
366, last line, for' Dyssochaete ' read • Dissochaeta '.
374, first line, after' Trapa' remove the semicolon.
339, fourth line, iQr ' laticiferous cells' read' laticiferous sacs '.
391, fifth line from bottom, read' cystoliths '.
396, fourth line, for • vgtice/lata ' read' verticil/ata '.
399, third line from bottom, for • C. Urticae, A, DC. B. hispida' read • C. Urticae,
A. DC. /3 hispida '.
408, eleventh line from end of first paragraph, for' Opunita ' read • Opuntia '.
420, third line, for' offlcinale • read' officinale '.
439, fifth line from end of first paragraph of Caprifoliaceae, far • stufted ' read
'tufted '. I

500, ninth line, for' GOlliolmmt • read' GOlliolimon "

504, twelfth line, for' rigidity' read 'flexile strength "
S08, footnote I, for 'Sapotaceae' read 'Ilicineae', and for 'Ilicineae' read 'Sapo-
taceae '.
S34, footnote, for ' A rgel ' read ' A rg hel " and for ' described a ' read 'described as '.
535. footnote 4, first word in line 2 should read • intraxylary '.
563, first line of third paragraph. after' within certain limits' insert' it '.
566, last line of first paragraph, for' Ipomoceae ' read' Ipomoeae '.
S68, hvelfth line from bottom, after' Fig. I 29, ] ' insert a comma in place of the
full stop.
573, second line of Cuscuteae should read' poor in chlorophyll '.
633, last word of fourth line should read' thin "
647, fourth line of nrst pa.agraph, for ' De Barry' read • De Bary '.
694, second line of first paragraph, for ' Fig. I 59, C . read ' Fig. 169, C'.
from
7 J 2;· ten th. line- bottom, for • vesti bu Ie • read 'fran t cavity '.
7~~ hlthe second line read • Gonystylus, Asclemm and Amyxa '.
764, fourth line of Urticaceae, for' Arctocarpeae . read • Artocarpeae '.
766, second line of second paragraph, for' C. ]{raussiana . read • Celtis Kraussiana "
779, last line but one of Thelygoneae. for • or ans ' read ' organs'.
823, first line of third paragraph, for • Hewitt' read' Leavitt "
835. last line but one of second paragraph, for • simpler' read' simple '.
841, fourth line, tor' Tsoptera read' [soptera '.
I

845, footnote, for' latiferaus ' read laticiferous "

I

_Note. It wa.s statet1 in the Editorial Preface to Vol. 1 that the English edition had
receIved from the author's hand all the emendations embodied in the new German edition.
Since this preface was written it has been ascertained that some small additions were
made to the German edition while in the press, and too late to be incorporated in the
translation.