The Hike Alison Farrell

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Copyright © 2019 by Alison Farrell.

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Alison Farrell has a deep and abiding love for wild berries and other
foraged foods. Her favorite items to bring on a hike include: a sketchbook, a
pencil, a map, and a small natural history book. She lives, bikes, and hikes
in Portland, Oregon, and other places in the Pacific Northwest.
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 CHAPTER I
INTRODUCTION—POSITION OF MOLLUSCA IN THE ANIMAL KINGDOM—
CLASSIFICATION—ORIGIN OF LAND AND FRESH-WATER MOLLUSCA

It is the generally accepted opinion among men of science that all

life originated in the sea. Not that all parts of the sea are equally
favourable to the development of forms of life. The ocean surface,
with its entire absence of shelter or resting-place, and the deep sea,
whose abysses are always dark and cold and changeless, offer little
encouragement to plant or animal life, as an original starting-point.
True, both the surface and the depths of the sea have become
colonised by myriads of forms, Mollusca amongst them, but these
quarters are in the truest sense colonised, for the ancestors of those
who inhabit them in all probability migrated from elsewhere.
It was no doubt the littoral region and the shallow waters
immediately below it, a region of changeable currents, of light and
shade, of variation, within definite limits, of temperature and tide
effects, which became the scene of the original development of plant
life, in other words, of the food-supply which rendered possible its
colonisation by higher animals. But the littoral region, besides the
advantages of tenancy which it offers to animal life, has also its
drawbacks. The violence of the surf may beat its inhabitants in
pieces, the retreat of the tide exposes them, not merely to
innumerable enemies in the shape of predatory birds and beasts, but
also to a change in the atmospheric medium by which they are
surrounded. Hence, in all probability, have arisen the various forms
of adaptation which are calculated to bring about the ‘survival of the
fittest’; hence, to narrow our point of view to the Mollusca, the
development of hard shells, or exoskeletons, hence the sand-
burrowing, rock-boring, rock-clinging instincts of various genera and
species.[1]
What was the primitive form of molluscan life is little likely to be
ever positively known, although, on grounds of comparative
anatomy, something approaching to the archi-mollusc is often
constructed, with more or less probability, by careful observers. From
one of the oldest known geological strata, the Cambrian, nearly four
hundred species of Mollusca are known, which include
representatives of nearly all the great Orders as they exist at the
present day, and without the slightest sign of approximation to one
another. With regard to the origin of the land and fresh-water
Mollusca some definite conclusions can be arrived at, which will be
given in their proper place.
Scarcely any portion of the coast-line of the world is destitute of
molluscan life, except in regions where extreme cold forbids its
existence. Thus along the shores of Northern Asia there is no proper
littoral fauna, the constant influence of travelling ice sweeping it all
away; animal life begins at about three fathoms. But in every coast
region not positively hostile to existence Mollusca make their home.
Each description of habitat has its own peculiar species, which there
flourish best, and exist precariously, if at all, elsewhere. Thus the
sandy waste of estuaries, the loose and shingly beaches, the slimy
mud-flats beset with mangroves, the low stretches of jagged rock,
and even the precipitous cliffs, from whose base the sea never
recedes, have all their own special inhabitants. The same is true of
the deep sea, and of the ocean surface. And when we come to
examine the land and fresh-water Mollusca, it is found not merely
that some Mollusca are terrestrial and others fluviatile, but that
certain species haunt the hills and others the valleys, some the
recesses of woods and others the open meadow sides, some prefer
the limestone rocks, others the sandy or clayey districts, some live
only in still or gently moving waters, while others are never found
except where the current is rapid and powerful.
It is within the tropics that the Mollusca become most numerous,
and assume their finest and quaintest forms. A tropical beach,
especially where there is a good tide-fall and considerable variety of
station, abounds in molluscan life to an extent which must literally be
seen to be believed. The beach at Panama, to select an instance
familiar to the present writer, is astonishingly rich in species, which
probably amount in all to several hundreds. This is due to the
immense variety of habitat. On the rocks at high-water mark, and
even above them, occur Truncatella, Melampus, Littorina, and
Siphonaria; where a mangrove-swamp replaces the rock, on the
branches overhead are huge Littorina, while three species of
Cerithidea crawl on the mud, and Cyrena and Arca burrow into it.
Lower down, in the rock pools at half-tide mark are Cerithium,
Purpura, Omphalius, Anachis (2 sp.), Nassa, and several Crepidula.
At low-water mark of ordinary tides, under stones half buried in clean
sand, are Coecum and Vitrinella; under the blocks which rest on
solid rock are Cypraea (4 or 5 sp.), Cantharus, more Anachis,
Columbella (3 sp. including the graceful C. harpiformis), and
Nitidella. Where the blocks of rock are rather muddy, Conus lurks,
and with it Turritella and Latirus. Where the rocks form a flat-topped
platform 2 or 3 feet high, with here and there a deep crack, huge
Chitons 3 inches long conceal themselves, with two species of
Turbo, Purpura, and Clavella. At extreme low-water mark of spring
tides, on the isolated rocks are Monoceros, Leucozonia, and
Vermetus, in them are Pholas and a burrowing Mytilus, under them
are more Conus, Dolium, and huge frilled Murices. Patches of clean
gravelly sand here produce Strombus; on the operculum of the great
Str. galea is sure to be a Crepidula, exactly fitting its breadth. On the
liquid mud-flats to the north glide about Marginella, Nassa, and
Truncaria, in the clean sandstretch to the west Olivella ploughs about
by hundreds with several species of Natica, and Tellina and Donax
bury themselves deep, while farther down are Artemis, Chione, and,
where mud begins to mix with the sand, Mytilus and more Arca.
Each of these species has its own habitat, often circumscribed to a
few square feet at the most, and it would be utterly useless to seek
for it anywhere except in its own special domain.
Equally abundant are the land Mollusca of the tropics. Prof. C. B.
Adams relates that within the limits of a single parish in Jamaica,
named Manchester, which measures no more than four miles long
and one mile broad, he obtained no fewer than one hundred species.
Mr. J. S. Gibbons, in a description of the Mollusca he obtained near
St. Ann’s, Curaçao, gives a lively picture of their abundance in an
exceptionally favoured locality:—[2]
 “Near the outskirts of the town a waste piece of ground supplied
me with occupation for all the time I had to spare. Neither grass nor
water was to be seen, the only vegetation consisting of a few stunted
cacti and still fewer acacia bushes. This, however, was so rich in
shells that of several species enough specimens could have been
collected in a few yards to supply, I should suppose, all the shell
cabinets in the world.... The stones, plants, and ground were
covered with Strophia uva L., Tudora megacheila, P. and M., was in
equal abundance, suspended by its silk-like thread from acacia
boughs, or strewed thickly on the ground underneath. A Bulimulus
(B. multilineatus var. sisalensis) abounded on the smaller boughs,
while under masses of coral Macroceramus inermis Gundl., Pupa
parraiana d’Orb, and P. pellucida Pfr., were abundant. In the loose
soil Cylindrella Raveni Bland, Cistula Raveni Bland, and a curious
Cionella were so numerous that a spade would have been the best
instrument with which to collect them. I wasted a good deal of
valuable time in separating them from the soil, when by simply taking
away a few handfuls of mould, I might have obtained a larger
number of specimens. A species of Stenogyra and a Succinea
complete a list, all of which might have been gathered from almost
any square yard of ground on the hillside.”

Position of Mollusca in the Animal Kingdom.—Up to very

recent times it was usual to regard the Mollusca as one of the four
subdivisions of a great family known as Malacozoa, the subdivisions
being (1) Mollusca, (2) Tunicata, (3) Brachiopoda, (4) Polyzoa or
Bryozoa. This classification is still retained in the leading modern
manual on the subject.[3] The progress, however, of investigation
leads to the belief that the Mollusca are not so closely related to
these other groups as such a classification would seem to imply. The
Tunicata, for instance, appear, from the whole course of their
development, to occupy a position near to the Vertebrata. The
relations of the Brachiopoda and Polyzoa will be more particularly
referred to in that part of this History which deals especially with
those groups. The position of the Mollusca is, in many respects, one
of considerable isolation. Any attempt, therefore, definitely to relate
them to one group or another, is, in all probability, to go further than
the present state of our knowledge warrants. Especially to be
deprecated are systems of classification which confidently derive the
Mollusca in general from this or that group. The first undisputed
traces of animal life, which appear in the Cambrian epoch, exhibit
the same phyletic distinctions as now exist. Sponges, Echinoderms,
Mollusca, and Worms, formed already, in those immeasurably
remote ages, groups apparently as generally distinct from one
another as they are at the present time. It would seem that any
theory of development, which confidently teaches the derivation of
any one of these groups from any other, is, in the present state of the
evidence before us, hazardous in the extreme.
Some indications of relationship, which must not be pushed too
far, may be drawn from a consideration of embryonic resemblance.
An especial characteristic of the Mollusca is the possession of a
particular form of larva, which occurs in one of the stages of
development, known as the trochosphere (see p. 130). This form of
larva is shared with two orders of Annelida, the Chaetopoda and the
Gephyrea armata, and, in all probability, with the Polyzoa as well. It
may also be significant that the adult form in Rotifera bears a close
resemblance to the trochosphere larva in those groups.
Basis of Classification.—The Mollusca are divided into four
great Orders—Cephalopoda, Gasteropoda, Scaphopoda, and
Pelecypoda.[4] Each name, it will be noticed, bears reference to the
‘foot,’ i.e. to the organ of motion which corresponds in function to the
foot in the Vertebrata.
In the Cephalopoda the feet, or, as they are more frequently
termed, the ‘arms,’ are arranged symmetrically round the head or
mouth. The common forms of ‘cuttle-fish’ (Octopus, Loligo) are
familiar examples of Cephalopods.
The Gasteropoda crawl on the flat under-surface or ‘sole’ of the
foot. Snails, slugs, sea-hares, whelks, periwinkles, and coats-of-mail
or chitons are examples of this Order.
The Scaphopoda possess a long tubular shell open at both ends;
with their small and elongated foot they are supposed to dig into the
mud in which they live. The common Dentalium or tusk-shell of our
coasts is a representative of this Order.

Fig. 1.—Examples of the four Orders: A,

Cephalopoda; B, Gasteropoda; C,
Scaphopoda, and D, Pelecypoda.
A, Ommastrephes sagittatus Lam.,
Naples: a, a, arms surrounding the
mouth; f, funnel; t, t, the two ‘tentacular’
arms, × ⅖ . B, Buccinum undatum L.,
Britain: f, foot; pr, proboscis. × ½. C,
Dentalium entalis L., Norway: f, foot. D,
Cardium oblongum Chem., Naples: f,
foot; s, efferent or anal siphon; s’,
efferent or branchial siphon, × ½.
The Pelecypoda[5] are enclosed in a bivalve shell fastened by a
muscular hinge, the adjacent part of the valves being generally more
or less toothed; the foot is as a rule roughly comparable to the shape
of an axe-head.
To these four Orders is frequently added a fifth, the Pteropoda,
whose exact position is at present not absolutely settled. The
Pteropoda[6] are ‘pelagic,’ i.e. they live in the open waters of the
ocean, rising to the surface at night, and sinking into cooler water by
day. They are provided with a pair of wing-like appendages or ‘feet,’
on each side of the head, by means of which they are enabled to
swim. Some authorities regard the Pteropoda as a subdivision of
Gasteropoda, others as forming a separate Order, of equivalent
value to the other four. The question will be further discussed below
(see chap. xv.), but for the present it will be sufficient to state that the
weight of evidence appears to show that the Pteropoda are modified
Gasteropoda, with special adaptations to pelagic life, and are
therefore not entitled to rank as a separate Order.
Some writers conveniently group together the first three of these
Orders, the Cephalopoda, Gasteropoda, and Scaphopoda, under the
title Glossophora,[7] or Mollusca furnished with a radula or ribbon-
shaped ‘tongue,’ set with rows of teeth and situated in something of
the nature of a head, as distinguished from the Aglossa (or
Lipocephala),[8] i.e. those Mollusca which have no radula and no
head. To the latter belong only the fourth Order, the Pelecypoda.
This view postulates, for the primitive ancestral Mollusc, a body with
a more or less developed head, and possibly the rudiments of an
apparatus for grinding or triturating food. This form, it is held, either
developed or degenerated. In the former case, in consequence of
the more active mode of life upon which it may be supposed to have
entered, it gave rise to all the more highly organised forms which are
grouped under the three great Orders. When, on the other hand, the
ancestral form associated itself with an inactive or sedentary life, it
was, we may believe, modified accordingly, and either lost by
atrophy or failed to acquire those special points of organisation
which characterise the highly-developed form. Hence the
Pelecypoda, or bivalves, whose characteristic is the absence of any
definite cephalic region or masticatory apparatus. It is a remarkable
fact in support of this theory of the origin of the Aglossa that certain
of their larvae are known to possess traces of higher organisation,
e.g. an external mouth and eyes, the former of which becomes
covered by the mantle lobes, while the latter disappear long before
the adult stage is reached.
Thus we have

Classification of Gasteropoda.—The Gasteropoda are

numerically very largely in excess of the two other Orders of the
Glossophora, far more complicated as regards classification, and
contain a large proportion of those examples of the Mollusca which
are most familiar to the ordinary observer. It will therefore be
convenient to postpone for the present a fuller discussion of the
subdivisions of the Cephalopoda and Scaphopoda, as well as of the
Aglossa, returning to them again in special chapters (chaps. xiii. and
xvi.), and to devote a few introductory words to the classification and
relations of the Gasteropoda.
The Gasteropoda are divided into four Classes, Amphineura,
Prosobranchiata, Opisthobranchiata, and Pulmonata.
Fig. 2.—An example of the
Polyplacophora: Chiton
spinosus Brug.

Fig. 3.—An example of the

Aplacophora, Neomenia
carinata Tullb.: a, anus; gr,
ventral groove; m, mouth.
 (1) The Amphineura[9] are bilaterally symmetrical Mollusca, i.e.
with organs either single and central, or paired and disposed on
either side of the longer axis of the animal. The shell, when present,
is never spiral, but consists of eight overlapping plates, kept together
by an elliptical girdle. The Amphineura are divided into (a)
Polyplacophora,[10] or Chitons, and (b) Aplacophora (Chaetoderma
and Neomenia).
(2) The Prosobranchiata[11] are so named from the fact that the
breathing organ (branchia or ctenidium[12]) is as a rule situated in
front of the heart, the auricle at the same time being in front of the
ventricle. They are asymmetrical, almost always furnished with a
shell, which is at some time spiral, and with an operculum. The
sexes are separate. They are either marine animals, or can be
shown to be more or less directly derived from genera which are
marine. They are divided into (a) Diotocardia[13] (Haliotis, Fissurella,
Trochus, Nerita, Patella), which have, or whose immediate ancestors
are believed to have had, two auricles to the heart, two sets of
breathing organs, two kidneys, but no proboscis, penis, or siphon,
and (b) Monotocardia,[14] in which the heart has only one auricle, the
true breathing organ is single, and there is a single kidney. To this
division belong the great majority of marine univalve Mollusca, e.g.
Cypraea, Buccinum, Murex, Littorina, Ianthina, all the land and fresh-
water operculates (Cyclostoma, Melania, Paludina, etc.), as well as
the Heteropoda, which are a group of Prosobranchiata which have
betaken themselves to a pelagic life.
 Fig. 4.—Example of a Heteropod, Carinaria
mediterranea Lam., Naples: a, anus; br,
branchia; f, foot; i, intestine; m, mouth; p, penis;
s, sucker; sh, shell; t, tentacles. × ½. The animal
swims foot uppermost.

(3) In the Opisthobranchiata[5] the breathing organs (when

present) are behind the heart, and the auricle of the heart is
consequently behind the ventricle. They are asymmetrical marine
animals; usually, but by no means always, without a shell, scarcely
ever with an operculum in the adult state. The sexes are united in
the same individual. The Opisthobranchiata fall into two divisions: (a)
Tectibranchiata, in which the breathing organ is more or less covered
by the mantle, and a shell is usually present, which is sometimes
rudimentary, e.g. Bulla, Aplysia, Umbrella, and the whole group of
Pteropoda; (b) Nudibranchiata, or sea slugs, which have no shell
and no true ctenidia, but breathe either by the skin, or by ‘cerata’ or
papilliform organs prominently developed on the back: e.g. Doris,
Aeolis, Dendronotus.
Fig. 5.—A, A Tectibranchiate Opisthobranch, Umbrella
mediterranea Lam., Naples: a, anus; br, branchia; f,
foot; m, mouth; rh, rhinophores; sh, shell.
B, A Pteropod, Hyalaea tridentata Forsk., Naples: sh,
shell; l, l, swimming lobes of foot.
C, A Nudibranchiate Opisthobranch, Aeolis peregrina,
Naples: f, foot; c, cerata.

Fig. 6.—Examples of—A, Pulmonata Basommatophora, the

common Limnaea peregra Müll.: e, e, eyes; t, t,
tentacles. B, Pulmonata Stylommatophora, Helix
hortensis Müll.: e, e, eyes; t, t, tentacles; p. o,
pulmonary orifice (the position of the pulmonary orifice
in Limnaea will be seen by reference to Fig. 101).
 (4) The Pulmonata[15] are asymmetrical air-breathing non-marine
Mollusca, generally, but not always, furnished with a shell. The sexes
are always united in the same individual, and the operculum is
always wanting, except in Amphibola. They are conveniently divided
into Stylommatophora,[16] in which the eyes are at the tip of the
upper tentacles, which are retractile (Helix, Limax, Bulimus, and all
true land slugs and snails), and Basommatophora, in which the eyes
are at the base of the tentacles, which are not retractile (Limnaea,
Planorbis, Physa, and all the Auriculidae).
Thus we have
Gasteropoda Amphineura Polyplacophora
Aplacophora
Prosobranchiata Diotocardia
Monotocardia (incl. Heteropoda)
Opisthobranchiata Tectibranchiata (incl. Pteropoda)
Nudibranchiata[17]
Pulmonata Stylommatophora
Basommatophora
The relation of the four great Orders to one another will be better
discussed when we come to deal with each Order separately. The
problem of the origin and mutual relationship of the various forms of
molluscan life is of extreme subtlety, and its solution can only be
approached after a comprehensive survey of many complicated
anatomical details. But there is one branch of the Mollusca—the land
and fresh-water genera—whose origin is, comparatively speaking, of
recent date, and whose relationships are therefore less likely to have
suffered complete obliteration.
Origin of the Land and Fresh-water Mollusca.—The ultimate
derivation of the whole of the land and fresh-water molluscan fauna
must, as has already been remarked, be looked for in the sea. In
certain cases the process of conversion, if it may be so termed, from
a marine to a non-marine genus, is still in progress, and can be
definitely observed; in others the conversion is complete, but the
modification of form has been so slight, or the date of its occurrence
so recent, that the connexion is unmistakable, or at least highly
probable; in others again, the modification has been so great, or the
date of its occurrence so remote, that the actual line of derivation is
obscured or at best only conjectural.

Fig. 7.—A, the common cockle (Cardium edule

L.). B, Adacna plicata Eichw., Caspian
Sea. C, Didacna trigonoides Pall., Caspian
Sea.
This passage from a marine to a non-marine life—in other words,
this direct derivation of non-marine from marine genera—is
illustrated by the faunal phenomena of an inland brackish-water sea
like the Caspian, which is known to have been originally in
connexion with the Mediterranean, and therefore originally supported
a marine fauna. The Mollusca of the Caspian, although without
exception brackish- or fresh-water species, are in their general facies
distinctly marine. Of the 26 univalve species which inhabit it 19
belong to 4 peculiar genera (Micromelania, Caspia, Clessinia,
Nematurella), all of which are modified forms of the marine
Rissoidae. The characteristic bivalves belong to the genera Adacna,
Didacna, and Monodacna, all of which can be shown to be derived
from the common Cardium edule. We have here a case where
complete isolation from the sea, combined no doubt with a gradual
freshening of the water, has resulted in the development of a number
of new genera. The singularly marine facies of several of the fresh-
water genera now inhabiting Lake Tanganyika, has given rise to the
belief, among some authorities, that that lake was at one time an
inlet of the Indian Ocean. In the upper waters of the Baltic, marine
and fresh-water Mollusca flourish side by side. So complete is the
intermixture, that an observer who had lived on no other shores
would probably be unable to separate the one set of species from
the other.[18] Thus between Dagö and Papen-Wiek[19] Mytilus edulis,
Cardium edule, Tellina balthica, Mya arenaria, Littorina rudis, and
Hydrobia balthica are the only true marine species; with these live
Unio, Cyclas, Neritina, Limnaea, and Bithynia. The marine species
and Neritina live down to 15–20 fath., the rest only down to 3 fath.
Under stones close to the shore of the Skärgård at Stockholm[20] are
found young Cardium and Tellina, and at 3 to 6 fath. Limnaea
peregra, and Physa fontinalis. Near Gothland Limnaea is found in
the open sea at 8–12 fath., and with it occur Cardium and Tellina. At
the Frisches Haff[21] Mya arenaria is the only marine species, and
lives in company with 6 sp. Limnaea, 1 Physa, 9 Planorbis, 1
Ancylus, 4 Valvata, 2 Sphaerium. Were the Sound to become
closed, and the waters of the Baltic perfectly fresh, it would be
inevitable that Mya arenaria, and such other marine species as
continued to live under their changed conditions, should in course of
time submit to modifications similar in kind to those experienced by
the quondam marine species of the Caspian.
It seems probable, however, that the origin, at least in a great
part, of the land and fresh-water Mollusca need not be accounted for
by such involuntary changes of environment as the enclosure of
arms of the sea, or the possible drying up of inland lakes. These
cases may be taken as illustrations of the much more gradual
processes of nature by which the land and fresh-water fauna must
have been developed. The ancestry of that fauna must be looked for,
as far as the Gasteropoda are concerned, in the littoral and estuarine
species; for the Pelecypoda, in the estuarine alone. The effect of the
recess of the tide, in the one case, and the effect of the reduced
percentage of salt, in the other, has tended to produce a gradual
adaptation to new surroundings, an adaptation which becomes more
and more perfect. It may be safely asserted that no marine species
could pass into a land or fresh-water species except after a period,
more or less prolonged, of littoral or estuarine existence. Thus we
find no land or fresh-water species exhibiting relationships with such
deep-sea genera as the Volutidae, Cancellariidae, Terebridae, or
even with genera trenching on the lowest part of the littoral zone,
such as the Haliotidae, Conidae, Olividae, Capulidae. The signs of
connexion are rather with the Neritidae, Cerithiidae, and above all
the Littorinidae, which are accustomed to live for hours, and in the
case of Littorina for days or even weeks, without being moistened by
the tide. Similarly the fresh-water Pelecypoda exhibit relationships,
not with genera exclusively marine, but with genera known to inhabit
estuaries, such as the Mytilidae, Corbulidae, Cardiidae.
It would be natural to expect that we should find this process of
conversion still going on, and that we should be able to detect
particular species or groups of species in process of emigration from
sea to land, or from sea to fresh water. Such species will be
intermediate between a marine and a land or fresh-water species,
and difficult to classify distinctly as one or the other. Cases of
Mollusca occupying this intermediate position occur all over the
world. They inhabit brackish swamps, damp places at high-water
mark, and rocks only at intervals visited by the tide. Such are
Potamides, Assiminea, Siphonaria, Melampus, Hydrobia,
Truncatella, among the univalves, and many species of Cyrena and
Arca among the bivalves.

Origin of the Fresh-water Fauna

(a) Pelecypoda.—Estuarine species, which have become

accustomed to a certain admixture of fresh water, have gradually
ascended the streams or been cut off from the sea, and have at last
become habituated to water which is perfectly fresh.
 Fig. 8.—A, The common Mytilus edulis
L., a marine genus and species.
B, Dreissensia, a fresh-water
genus, closely allied to Mytilus.

Fig. 9.—A, Arca navicella Reeve, Philippines, a marine

species. B, Arca (Scaphula) pinna Bens., R.
Tenasserim, a fresh-water species which lives
many miles above the tide-way.
Thus Dreissensia (rivers and canals throughout N. Europe and N.
America) and Mytilopsis (rivers of America) are scarcely modified
Mytili (Fig. 8); Scaphula is a modified Arca, and lives in the Ganges,
the Jumna, and the Tenasserim at a distance of 1600 miles from the
sea (Fig. 9). Pholas rivicola is found imbedded in floating wood on
the R. Pantai many miles from its mouth. Cyrena, Corbicula, and
probably Sphaerium and Pisidium are derived, in different degrees of
removal, from the exclusively marine Veneridae; Potamomya (rivers
of S. America), and Himella (R. Amazon) are forms of Corbula. The
Caspian genera derived from Cardium (Adacna, Didacna,
Monodacna), have already been referred to. Nausitora is a form of
Teredo, which lives in fresh water in Bengal. Rangia, Fischeria, and
Galatea probably share the derivation of the Cyrenidae, while in
Iphigenia we have one of the Donacidae which has not yet mounted
rivers, but is confined to a strictly estuarine life. The familiar
Scrobicularia piperata of our own estuaries is a Tellina, which lives
by preference in brackish water.

Fig. 10.—Trigonia pectinata

Lam., Sydney, N.S.W.

The great family of the Unionidae is regarded by Neumayr[22] as

derived from Trigonia, the points of similarity being the development
of a nacreous shell, the presence of a strong epidermis, and the
arrangement of the muscular scars. It is remarkable, too, that on
many Uniones of Pliocene times there is found shell ornamentation
of such a type as occurs elsewhere among the Pelecypoda only on
Trigonia.
 The genera of fresh-water Pelecypoda are comparatively few in
number, and their origin is far more clearly discernible than that of
any other group. This is perhaps due to the fact that the essential
changes of structure required to convert a marine into a fresh-water
bivalve are but slight. Both animals “breathe water,” and both obtain
their nutriment from matter contained in water. Similar remarks apply
to fresh-water operculate Gasteropoda. But the passage from a
marine to an aerial life involves much profounder changes of
environment, which have to be met by correspondingly important
changes in the organism. This may be in part the reason why the
ancestry of all Pulmonata, whether land or fresh-water, is so difficult
to trace.

Fig. 11.—A, Cominella, a

marine genus, which lives
between tide marks, and
from which is probably
derived B, Clea, a genus
occurring only in fresh
water.
 Fig. 12.—A, Cerithium columna Sowb. (marine). B,
Potamides microptera Kien. (brackish water). C, Io
spinosa Lea, one of the Pleuroceridae (fresh water).
(b) Gasteropoda.—(1) Operculate. Canidia and Clea are closely
allied, with but little modification, to the marine Cominella[23] (Fig.
11), as is also Nassodonta to Nassa. They occur (in fresh water) in
the rivers of India, Indo-China, Java, and Borneo, associated with
essentially fresh-water species. Potamides, with its various sub-
genera (Telescopium, Pyrazus, Pirenella, Cerithidea, etc.), all of
which inhabit swamps and mud-flats just above high-water mark in
all warm countries, are derived from Cerithium (Fig. 12); Assiminea,
Hydrobia, and perhaps Truncatella, from Rissoa. It is a remarkable
fact that in Geomelania (with its sub-genera Chittya and Blandiella)
we have a form of Truncatella which has entirely deserted the
neighbourhood of the sea, and lives in woody mountainous localities
in certain of the West Indies. Cremnoconchus, a remarkable shell
occurring only on wet cliffs in the ghâts of southern India, is a
modified Littorina. Neritina and Nerita form a very interesting case in
illustration of the whole process. Nerita is a purely marine genus,
occurring on rocks in the littoral zone; one species, however, (N.
lineata, Chem.) ascends rivers as far as 25 miles from their mouth,
and others haunt marshes of brackish water. Neritina is the fresh-
water form, some species of which are found in brackish swamps or
even creeping on wet mud between tide marks, while the great