Classification of Pteridophytes

Reimers, 1954
 Division Psilophytopsida
• Single order
 Order – Psilophytales
 Family- Rhyniaceae
 Genera – Rhynia, Horneophyton, Cooksonia, Yarravia
 Family – Zosterophyllaceae
 Genus – Zosterophyllum
 Family – Psilophytaceae
 Genus – Psilophytum
 Family – Asteroxylaceae
 Genus –Asteroxylon
Family Rhyniaceae:
🞭 The Rhyniaceae are the simplest of the Psilophytales, often
compared with the sporophyte of Anthoceros. The most
important genera are Rhynia and Homeophyton from the
Middle Devonian in Scotland.

Genus Rhynia:
🞭 The genus Rhynia from the Rhynie chert beds (Middle
Devonian) in Scotland was discovered by Mackie in 1913 and
fully described by Kidston and Land in 1917. This discovery
established the Psilophytales as a separate and distinct taxon.
Three species are known of which Rhynia major and R.
gwynne-vaughani are the better known.

🞭 The plants, apparently, grew in swampy marshes near

volcanoes where the atmosphere contained sulphurous vapour
and the soil was acid. The reconstructions are from silicified
petrifications.
Sporophyte of rhynia
The Sporophyte:

🞭 The plant body was a herbaceous sporophyte with

dichotomously branching horizontal rhizomes bearing
rhizoids on the underside and some of the branches grow-
ing up abruptly forming aerial shoots.

🞭 The aerial shoots of R. major were up to 50 cm long and 6

mm in diameter while those of R. gwynne-vaughani were
shorter and more slender.

🞭 The aerial branches were cylindrical and sparsely

dichotomously branched. These were naked being devoid
of any appendage of leaf and usually tapering upwards
ending in a point or in an erect sporangium.
• R. gwynne-vaughani shows hemispherical, oval or
lenticular protuberances arising from the lower parts
(more mature) of the aerial shoots or from the
rhizomatous parts.

• These are constricted at the points of

and, in mature ones, have their own vascular
attachment
bundles not connected those of the main
with stems.

• They are found to be readily detachable and, possibly;

was a means of vegetative propagation germinating into
new shoots.
🞭 The anatomy of the stem is very simple.

🞭 In the centre is a slender, hadrocentric protostele with a

small central xylem formed of simple annular tracheides
which, in some larger specimens, become smaller towards
the centre.

🞭 This is surrounded by four or five layers of elongated cells

with oblique ends which represent the phloem although
sieve plates have not been observed.

🞭 There is no endodermis or pericycle. All round this

vascular bundle is a massive inner cortex of loose,
rounded, parenchymatous cells with lots of air spaces.
• The vascular bundle fills most of the stem and was
probably the main photosynthetic tissue.

• The outer cortex is formed by one or two layers of

somewhat, larger, comparatively angular, compact
(except below stomata) cells at the hypodermal region.

• The epidermis is a compact layer of cells broken here

and there only by the stomata with pairs of guard cells
as in other vascular plants.

• This is externally covered by a heavy cuticle. Often the

smallest branches show no vascular supply.
🞭 The sporangia are oval or cylindrical structures with pointed
ends at the apices of the dichotomies.

🞭 They may be slightly constricted at the bases though

continuous with the stem and are always wider.

🞭 Those of R. major were rather big (about 12 mm long and 4

mm in diameter).

🞭 The sporangium wall is thick and multi- layered with the outer
cells thick-walled and no method of dehiscence is observed.

🞭 The thinner, inner cells probably represent the tapetum.

🞭 The whole interior is filled with spore tetrads or free spores.

🞭 The spores are spherical, large (up to 65µ in diameter

in
R. major) and covered with a thick cuticle.
The Gametophyte:
• As in all the Psilophytopsida, the gametophytes are not known.
Lyon (1957) found some germinating spores within the same
Rhynie chert beds which show multicellular structures developing
at the ends of germ tubes. These may represent the gametophytic
germination.

• Merker (1959 and 1961) has suggested that it is not possible that
the gametophytes of such a big group were not fossilised while
the large algae had been preserved.

• He argued that the underground creeping parts of Rhynia and

Honuoph) ‘on are the gametophytes and not rhizomes. But no sex
organ has been found on these underground parts and the strong
vascular bundle is not normal in a gametophyte. His view is, till
now, mere speculation.
 Fossil Psilophytales
Zosterophyllum

Asteroxylon

Horneophyton
Psilophyton
Psilotum sporophyte Habit
Tmesipteris
Lycopodium
 Selaginella sp.,

S. kraussiana S. lepidophylla
Isoetes
Class Lycopsida
 General Features of Lycopsida
(i) includes both fossil (e.g.,
It Lepidodendron) and living Pteridophytes
(five living genera e.g., lycopodium,
Phylloglossum, Isoetes, Stylites and
Selaginella).

(ii)Its history indicates that these Pteridophytes

developed during the Devonian period of the
Palaeozoic era.

(iii)The plant body is sporophytic and can be

differentiated into root, stem and leaves.
(iv)The leaves are small (microphyllous), simple with a
single mid vein.

(v)They are usually spirally arranged, sometimes in

opposite fashion and or even in whorls.

(vi)In some cases the leaves are ligulate (e.g.,

Selaginella, Isoetes). The ligule is present at the base of
each leaf.

(vii)The vascular tissue may be either in the form of

plectostele, siphonostele or sometimes even polystele.

(viii) Leaf gaps are absent.

(ix)Sporangia are quite large in size and develop on the
adaxial surface of the leaves (sporophylls). Sporophylls
are loosely arranged and form strobilus.

(x)Some members are homosporous (e.g., Lycopodium)

while others are heterosporous (e.g., Selaginella).

(xi) Antherozoids are biflagellate or multiflagellate.

(xii)Gametophytes which are in the form of prothalli are

formed by the germination of spores.

(xiii)Heterosporous forms have endoscopic gametophytes

while in homosporous forms the gametophyte is
exoscopic.
Protolepidodendron
Lycopodium and Phylloglossum
Lepidodendron
Isoetes coramendalina
S. krausiana S. sinensis

S. rupestris
Class Sphenopsida- Salient Features
1.The stems and branches are jointed with nodes and
internodes. The internodes are with longitudinal-oriented ridges
and furrows.
2.The leaves are extremely reduced and borne in whorls at the
nodes of aerial branches and stems.
3. Branches arise in whorls.
4. The sporangia develop on a peltale appen•dage called
sporangiophore. Sporangial walls are thick.
5.Most of the” members are homosporous including Equisetum.
However, some extinct forms were heterosporous (e.g.,
Calamites casheana).
6. The gametophytes are exosporic and green.
7. Antherozoids are multiflagellated.
8. The embryo is without suspensor and is exo- scopic in nature.
 Order Hyeniales
• Present in the Middle Devonian period(about 398 to 385
million years ago). They lack some significant characters of
Sphenopsida but certain features make them retained in this
Class.
• Protohyenia, Hyenia and Calamophyton
• Hyenia grew as a robust rhizome up to 5 cm (2 inches) in
diameter and parallel to the soil surface.
• Upright branches up to 15 cm (about 6 inches) in height arose
from the rhizome in a low spiral. Some branches divided
several times to form flattened leaflike structures.
• Others bore additional smaller branches tipped with a pair of
elongate sporangia that opened along a lateral slit to
release spores
Hyenia
 Order Sphenophyllales
• Appeared in full swing during Upper
Carboniferous – early Permian until Lower Triassic
era.
• The plant body was sporophytic and the
sporophytes were herbs, shrubs.
• Stem had nodes, internodes and leaves at nodes
in whorls.
• Leaves simple, wedge shaped or dichotomously lobed.
• Stele –actinostelic plectostele.
• Strobilus well organised.
• Sphenophyllum, Sphenophyllostachys, Bowmanites
Sphenophyllum
Order Calamites
Calamites
 Order
Equisetale
s
Equisetu
m
 Class or Division Pteropsida
Sub Class/sub division

Osmundidae Leptosporangiata
Eusporangiatae
Primofilices e
Osmundale Filicales
s Osmunda (19)
Schizaeaceae,
Orders Hymenophyllaceae,
Gleicheniaceae
Marattiale ceae
Cladoxylal Dicksoniaceae Matoniaceae,
s (5)
Asterothecacea Dipteridaceae
es (2)
Cladoxylaceae Angiopteridaceae
e Cyatheaceae,
Pseudosporochnacea Marattiaceae Dennstaedtioideae
e Danaeaceae Pteridoideae,
Christenseni Davallioideae
Coenopteridales aceae
Ophioglossa Oleandroideae,
(3)
Zygopteridaceae
les Ophioglossacea
(1) Onocleoideae
Stauropterid Famil Blechnoideae,
e ies Marsileales
aceae Asplenioideae
(2)
Botryopteridacea Salvinia Pilulariaceae,
Athyrioideae,Dryopteridoideae
e ceae Marsileaceae
Lomariopsidoideae,Adiantaceae
Azollaceae
 Salient Features of Pteropsida
•Generally called as ferns, Represented by 300 genera and 10,000 species, by
megaphyllous pteridophytes.
•Found as back as the Devonian period but less in Carboniferous period, profoundly
evolved in Triassic, Jurassic and Cretaceous eras to present time.
•Plant (Sporophytic) body is distinguished into roots, stem and spirally
arranged
leaves, well developed and vast schlerenchmya is found in roots and stem
•Habitat – moist and shady – humid tropical forests. Mostly land plants, some are
epiphytic (eg. Ophioglossum), aquatic (Azolla, Marsilea, Salvinia)
•Habit- small prostrate herbs (Azolla, Marsilea) to huge tree like (Cyathea)
•Leaves are large with branched veins. Compound, so called as fronds.
In
Ophioglossum, leaves are simple
•Leaf base may be enlarged and functions in starch storage
 Salient Features of Pteropsida
• Stele shows a wide variety of modifications: simple to advanced (Protostele-
siphonostele-solenostele-dictyostele conditions)

• Vegetative propagation – fragmentation, adventitious buds, stem

tubers,
apogamy

• Spores- sporangia are grouped – sorus in marginal or abaxial surface of leaf

blades. Special outgrowth called Indusium is seen

• In most of the genera, leaves are dual in function- photosynthetic

and
reproductive

• Sporangium development may be eusporangiate (from more than one

sporangial initial) or leptosporangiate ( from a single sporangial initial).
Spores –homosporous or heterosporous
Some members of Pteropsida

Marattia sp., Angiopteris sp., Danaea sp.,

 Some members of Pteropsida

Gleichenia sp.,

Ophioglossum sp.,

Lygodium sp.,
 Some members of Pteropsida

Nephrolepis sp.,
Osmunda sp.,
Cyathea sp.,

Pilularia sp., Marsilea sp., Azolla sp., Salvinia sp.,

 Stelar Evolution in Pteridophytes
The term stele has been derived from a Greek word
meaning pillar. Van Tieghem and Douliot (1886)
recognized only three types of steles.
(a) Haplostele:
This is the most primitive type of protostele. Here the central solid smooth
core of xylem remains surrounded by phloem (e.g., in Selaginella spp.).

(b) Actinostele:
This is the modification of the haplostele and somewhat more advanced in
having the central xylem core with radiating ribs (e.g., in Psilotum spp.).

(c) Plectostele:
This is the most advanced type of protostele. Here the central core of
xylem is divided into number of plates arranged parallel to each other. The
phloem alternates the xylem (e.g., in Lycopodium).

(d) Mixed-pith stele:

Here the xylem elements (i.e., tracheids) are mixed with the
parenchymatous cells of the pith. This type is found in primitive fossils and
living ferns. They are treated to be the transitional types in between true
protosteles on the one hand and siphonosteles on the other (e.g., in
Gleichenia spp. and Osmunda spp.).
Siphonostele
This is the modification of protostele. A stele in which the protostele
is medullated is known as siphonostele. Such stele contains a
tubular vascular region and a parenchymatous central region.
Jeffrey (1898) interpreted that the vascular portion of siphonostele
possesses a parenchymatous area known as a gap immediately
above the branch traces only or immediately above leaf and branch
traces.
In one type, however, the leaf gaps are not found and they are
known as cladosiphonic siphonosteles. In the other type both leaf
and branch gaps are present and they are known as phyllosiphonic
siphonosteles.
Ectophloic Siphonostele:
The pith is surrounded by concentric xylem cylinder and next to
xylem the concentric phloem cylinder.

Amphiphloic Siphonostele:
The pith is surrounded by the vascular tissue. The concentric inner
phloem cylinaer surrounds the central pith. Next to the inner
phloem is the concentric xylem cylinder which is immediately
surrounded by outer phloem cylinder (e.g., in Marsilea)

Solenostele:
The vascular plants have been divided into two groups on the basis
of the presence or absence of the leaf gaps. These groups are—
Pteropsida and Lycopsida. The ferns, gymnosperms and
angtosperms are included in Pteropsida, whereas the lycopods,
horse-tails, etc., are included in Lycopsida.
Dictyostele:
In the more advanced siphonosteles of Pteropsida, the
successive gaps may overlap each other. Brebner (1902)
called the siphonosteles with overlapping gaps as
dictyosteles. In such cases the intervening portion of the
vascular tissue between lateral to such leaf gaps is known
as meristele. Each meristele is of protostelic type. The
dictyostele with many meristeles looks like a cylindrical
meshwork.
Polycylic Stele:
This type of stelar organization is the most complex one
amongst all vascular cryptogams (pteridophytes). Such type of
steles are siphonostelic in structure. Each such stele possesses
an internal vascular system connected with an outer
siphonostele. Such connections are always found at the node.

A typical polycyclic stele possesses two or more concentric

rings of vascular tissue. This may be a solenostele or a
dictyostele. Two concentric rings of vascular tissue are found
in Pteridium aquilinum and three in Matonia pectinata
Eustele:

Here the vascular system consists of a ring of collateral or

bicollateral vascular bundles situated on the periphery of
the pith. In such steles, the inter-fascicular areas and the
leaf gaps are not distinguished from each other very
clearly. The example of this type is Equisetum.
HETEROSPORY

 Some Pteridophytes which produce two different types of

spores (differing in size, structure and function).

Such Pteridophytes are known as heterosporous and the

phenomenon is known as heterospory.

 The two types of spores are microspores and megaspores.

Microspores are smaller in size and develop into the male

gametophyte. Megaspores are large and develop into female
gametophyte.

According to Rashid(1976), only 9 genera of

pteridophytes show heterospory,i.e., Selaginella, Isoetes,
Stylites, Marsilea, Pilularia, Regnellidium, Salvinia, Azolla
The origin of heterospory can be better discussed on the
basis of evidences from paleobotany, developmental and
experimental studies.

Palaeobotanical evidences:
It has been suggested that heterospory arose due to degeneration of
some spores in a few sporangia. As more nutrition becomes
available to less number of spores, the surviving spore grow better,
hence increase in their size.

A number of heterosporous genera belonging to the Lycopsida,

Sphenopsida and Pteropsia were known in the late Devonian and
early Carboniferous periods.

Lepidocarpon, Lepidostrobus, Mazocarpon, Plaeuromeia,

Sigillariostrobiis (members Lycopsid) Calamocarpon,
of
Calamostachys, Palaeostachys (members of Sphenosida).
2. Evidences from Developmental Studies:

In heterosporous Pteridophytes, the development of micro and

megasporangia follow the same pattern.
They have identical organization but for their size. While in
megasporangia most of the spore mother cells degenerate but in
microsporangia only a few mother cells are
disorganize.
In Isoetes, there are only 50-300 megaspores in
megasporangium. In Selaginella
megasporangium contains erythropus
only one megaspore which is
functional.
In Marsilea, Salvinia and Azolla the phenomenon of
heterospory becomes distinct after meiosis.
In Marsilea 64 microspores and 64 megaspores are formed after
meiosis in microsporangium and megasporangium respectively.
Biological Significance of Heterospory:
The phenomenon of heterospory is of great biological significance on
account of the following facts:

(i) The development of the female gametophyte starts while the

megaspore is still inside the megasporangium.

(ii) Same is true of microspores i.e., they also start germinating into male
gametophytes while they are still inside microsporangium.

(iii) The female gametophyte derives its nourishment from the sporophyte
i.e., female gametophyte is dependent on sporophyte for its nourishment.

(iv) The dependence of female gametophyte on sporophyte for its nourishment

provides better starting point for the development of new embryo than an
independent green prothallus which has to manufacture its own food.
Seed Habit in Pteridophytes:
The adoption of heterospory and the retention and germination of a single
megaspore within megasporangium to form a female gametophyte, led to
the phenomenon of “seed habit”, a characteristic feature of the
spermatophytes.
A seed is that ovule which contains an embryo developed as a result of
fertilization.

The origin of seed habit is associated with the following:

(i) Production of two types of spores (heterospory).

(ii) Reduction in the number of finally to one per

megaspores
megasporangium.
(iii) Retention and germination of the megaspores and fertilization of the
egg.

(iv) Continued development of the fertilized egg into the embryo while
still in situ.
From the above observations it is concluded that the
life history of Selaginella approaches towards seed
habit because of the following features:

1. The occurrence of the phenomenon of heterospory.

2. Germination of megaspore inside megasporangium.

3.Retention of megaspore inside megasporangium either till

the formation of female gametophyte or even after
fertilization.

4.Development of only one megaspore per megasporangium for

example, in Selaginella monospora, S. rupestris, S.
erythropus etc.
 ECONOMIC IMPORTANCE OF
PTERIDOPHYTES

 The pteridophytes which include the ferns and a group of vascular

plant of ancient or primitive land plant with worldwide distribution.

 They are found in all the continents excepts Antarctica and most
islands, favoring moist temperate and tropical regions.

 The economic value of pteridophytes have been known to men for

more than 2000 years and have been found as an important source
of food and medicine.
 Pteridophytes are usually useful but few are harmful.
 Pteridophytes are used in variousfields---

1. As soil conservation
2. As bio fertilizer
3. As food
4. As ornamental
5. As entertainment
6. As medicinal used
7. As chemical production
8. As manufacturing
9. Metal accumulators
 Usually pteridophytes plants are terrestrial so
they protect the upper part of soil.
 They protect soil from heavy rainfall.
 They help in stopping soil erosion.

e.g. Pteris, Dryopteris, Nephrolepis etc.

Pteris Dryoptris
 Pteridophytesplants are very helpful for the
formation of biofertilisers.

 Azolla spp. are very helpful for the formation

of biofertiliser because root have Anabena help
nitrogen fixation.
Many plants are edible and used in form of vegetable

Ampelopteris prolifera, Isoetes used as food.

Osmunda cinnamomea use as vegetable.
Azolla also used as food production they have
higher carbohydrates and protein values.

Ampelopteris Azolla Osmunda

 Equisetum arevense whole plant are used in
food production.
The tuber of Isoetes are used as food.
Neprolepis biserrata rhizome are
edible.

Equisetum Isoetes
Few pteridophytes are used as ornamental

 Lycopodium obscurum called “Christmas tree”

are used as grassland during Christmas
festival and for purpose of decoration.

 Lycopodium volubile is very commonly used by

decoration.

 Selaginella plant also used during Christmas

festival as grassland and various type of table
decoration.
A few species are grown in pots for their
beautiful colored moss like foliage.
Many ferns are used as decoretary e.g. Pteris and
Dryopteris are used as ornamentally in home.

Pteris Selaginella
Few species of Selaginella such as
Selaginella lepdophylla and
S.pilifera are called resurrection
plant.
As MEDICINE

Plant Medicinal uses

Pteris multifold used in cancer, diarrhoea
hepatitis..
Ophioglassum costatum Used in antiviral, antidote to
snake bite, their rhizome used
in bleeding nose.
Marsilea condensata Leaves are used, diuretic
and plant used in snake bite
diarrhoea.

Lygodium Used for expulsion of intestional

japonicum worms.
 Plants Use
Adiantum capillms Anticancerus and Antibacterial
plants.
Adiantum lunulatum as blood related diseases

Adiantum candatum skin disease

Actinopteris rediata antimalarial

Aspelnium falcatum antihelmintic and tapeworms

reducer
Azolla pinnata Antifungal and antibacterial.

Equisetum ramosissimum diuretic and used in diarrhoea

Selaginella boryoides liver diseases

Dryopteris cochleata used antibacterialin

Pteridium revolutum gastric and intestinal diseases

As CHEMICALS
Plants Yields chemical
Pteris vittata Phenols
Psilotum nudum Psilotic acid, Gibberellin

Pteridium aquilinum Protein, sugar,

starch, H.C.N,beta-
carotene
Azolla pinnata Protein, carotinoids

Diplazinum esculentum Iron ,calcium

Equisetum arvense Oxalic acids, malic acid,

vinilic acid
VITAMINS

Plants Yields vitamins

Diplazinum esculentum Vitamin B

Equisetum arvense Vitamin C

Asplenium yoshinagae Vitamin K3

 OIL
YIELDING
Plants Product
Lycopodium inundataum These are produced a high
amount of fixed oils.

Ophiogloosum vulgatum They produced fixed oils.

 DYE
YIELDING
Plants Obtain Dye

Asplenium ensiformis red dye

Equisetum arvense red dye

Pteridium aquilinum yellow dye

MANUFACTURIN
G
Plants Use
Adiantum pedatum Basket manufacturing.

Lygodium microphyllum Basket manufacturing

Metathelypteris gracilescens Yields fibers.

 METAL
ACCUMULATORS
Plants Metals
Lycopodium clavatum Zinc Arsenic

Lygodium japonicum Arsenic, Calcium, Copper

Equisetum arvense Tin, Cobalt, Zinc

Pteris vittata Arsenic

 Pteridophytes are mostly useful but few are
harmful.

 Few pteridophytes are abnoxious weeds so

they are harmful for animal and for crop
plant.

 Pteridium aquilinum they are cosmopolitan

they are poisonous for Cattle and Horse.