their almost infinite variety in form and coloring, has given to
them an interest among collectors that is very great. There are
many wonderful conchological collections in public museums and in
private cabinets. It would well repay the lover of beauty as well
as the more serious student of nature to examine carefully such
collections when opportunity offers, for nowhere in the realm of
nature can more exquisite coloring and modeling be found. The fact
that shells may be preserved for all time without the expense and the
vexations of preservative fluids has no doubt induced many [pg307]
to choose this field of natural history for special study. But the
true student of zoölogy does not confine his investigations in this
phylum to considerations of the shell only, for the protective
armor constitutes only a portion, although an important portion, of
the anatomy of the mollusk. On the same principle, it would hardly
suffice to examine only the skeleton of vertebrate animals in order
to become acquainted with the fishes or mammals. In a sense, the
testaceous covering of a mollusk may be likened to an outer skeleton.
The older zoölogists were inclined to ignore almost entirely the
animal or "soft parts" of mollusks, and their classification of
the phylum into "univalve," "bivalve," and "multivalve," based
upon shell-characters alone, has since been proved to be wholly
artificial. The terms "univalve" and "bivalve" are, of course, often
and correctly used to-day to designate mollusks possessing a single
or a double shell, such as the common garden-snail, which carries
upon its back a single spiral shell, and, on the other hand, the
oyster or clam, which is inclosed within two valves which fit closely
together. But these terms are not /technically/ used, although
they do indicate in a general way the more modern classification.
Even among some comparatively recent conchologists a tendency to
belittle the value of the "slimy creature" within the shell has led
to some curious errors of grouping--mistakes which more thorough
and scientific investigation has brought to light. In general, the
higher classification into classes, orders, suborders, and families
is exclusively based upon anatomical features, and to a large extent,
also, is that of the genera; but species are always determined by
their shell or conchological characteristics.
When a certain familiarity with the forms and general appearance of
shells is gained, a glance at the shell alone will almost always
suffice to place it at once in its true generic position, for, along
with modifications of the animal itself, corresponding modifications
of the shell are quite certain to take place. Hence a fairly expert
zoölogist should be able to determine with considerable accuracy from
an empty and dead shell the anatomical features of the animal that at
one time inhabited it. [pg308]
One of the main reasons for the special interest that seems to have
attached to the study of conchology is derived from the fact that the
durability of shells has caused their almost perfect preservation
as fossils from the very earliest periods of geological time, thus
furnishing the key to the solution of many problems of evolution.
Upon this page of the earth's history the letters are sharp and
clear, and geologists possessing a knowledge of recent forms of
mollusks as well as of their geographical distribution have been
enabled to read some wonderful stories of the cosmic history of
islands and continents.
GEOGRAPHICAL DISTRIBUTION
Mollusks are found in every part of the world. The arctic seas
possess their own characteristic faunæ, the more temperate waters of
Europe and of America contain their own peculiar genera and species,
and the warm waters of the tropic seas furnish the conditions
favorable to the life of an immense number of characteristic forms.
Again, there are the littoral species, that live only between
tide-marks and are therefore exposed to the air for a number of
hours each day; some of these live just about high-tide mark and
have become almost terrestrial in their habits, while others must be
sought at the point of lowest tide, where for only a short time each
day they are deprived of their natural element. Then there are the
shallow-water forms, which never appear above low-tide mark save when
a neap tide surprises them; their range in depth extends to about
the hundred-fathom line, more or less. Beyond this depth to several
hundred fathoms other characteristic forms appear, and from the more
profound depths of mid-ocean the dredge has brought to light a host
of curious and interesting species.
It is the temperature of the water rather than the depth that appears
to influence the distribution of marine mollusks. Thus, certain
species whose natural home is in the shallow waters of the Arctic
Ocean have been taken in very deep waters off the southern coast of
the United States, the temperature conditions in both stations being
substantially the same. This fact, however, must not be too freely
accepted as establishing a principle. Some [pg309] shallow-water
forms have been taken at great depths, but in general the abyssal
fauna is a peculiar one that cannot well be marked off into
geographical provinces.
THE ARCTIC PROVINCE
The east coast of America is divided into several molluscan
faunal regions. A series of very-cold-water forms belonging to a
circumpolar region, called the "arctic province," are found as far
south as Newfoundland. On the New England coast a number of these
arctic species are also found, urged south by the influence of the
cold Labrador current. The most characteristic genera belonging to
this arctic fauna which are found upon the Maine and Massachusetts
coasts are /Buccinum/, /Chrysodomus/, /Sipho/, /Trophon/, /Bela/,
/Velutina/, /Trichotropis/, /Lacuna/, /Margarita/, /Pecten/, /Leda/,
/Yoldia/, /Astarte/, and /Mya/. Examples of all these genera are
encountered as far south as Cape Cod.
THE BOREAL PROVINCE
A "boreal province" corresponds with a similar faunal region upon
the European shore. The mollusks which compose this fauna are about
three hundred in number, and range along the New England coast from
the Gulf of St. Lawrence to Cape Cod. It is a somewhat remarkable
fact that many of these species are identical with English and French
forms. The striking genera upon the American side are /Purpura/,
/Littorina/, /Polynices/ (/Lunatia/ and /Neverita/), /Acmæa/,
/Margarita/, /Chiton/, /Doris/, /Æolis/, /Mytilus/, /Modiola/,
/Thracia/, and /Nucula/.
THE TRANSATLANTIC PROVINCE
Cape Cod has been regarded, until very recently, as a sharp
divisional point between the boreal and the transatlantic provinces,
the latter faunal area extending down the Atlantic coast of the
United States to southern Florida. At Cape Cod the Labrador current
is deflected from the coast, and the warmer shore waters south of
that point are unfitted for the development of the boreal forms,
though some of them, as we shall see, have passed the [pg310]
barrier and maintain themselves very successfully at Martha's
Vineyard and in Long Island Sound. Some of the species whose natural
habitat is south of Cape Cod are also to be found north of that
point, but they do not reach their full development in the colder
waters of Massachusetts Bay.
The long stretch of coast-line from Cape Cod to Florida is easily
susceptible of faunal subdivision at Hatteras, below which locality a
decided mixture of Antillean species is apparent. Between Long Island
Sound and Hatteras the littoral molluscan fauna is disappointing
to the collector, for the number of species is relatively small,
and few even of these are entitled to any claims to beauty of
shell. The nature of this coast, virtually a great sweep of exposed
sand-beach, is not conducive to a varied or rich fauna. But if this
portion of the transatlantic province is lacking in interest to
the shell-collector, it is anything but disappointing to the army
of men employed in the oyster- and clam-fishery. It is only within
this faunal area that the American oyster, /Ostrea virginica/, the
best of the edible shell-fish, finds its natural home, and here the
oyster-culture is most extensively carried on.
After storms the Jersey beaches are frequently strewn with /Mactra/,
/Tagelus/, /Arca/, and /Ensis/; in more sheltered places /Fulgur/ and
/Polynices/ (/Lunatia/) are commonly met. These may be accepted as
the most characteristic genera of this province.
Just south of Cape Hatteras an observer is struck with the sudden
change in the appearance of the shells on the beach. /Cardium/,
/Cassis/, /Dolium/, /Arca/, and /Cancellaria/ at once suggest
the West Indies. About the vicinity of Hatteras the Gulf Stream
approaches very near the land, bringing with its warm waters many
wanderers from tropical homes. From Beaufort to Florida there is
another long stretch of exposed and shifting sand-beach, which offers
a scant return to the collector of mollusks.
THE CARIBBEAN PROVINCE
When Florida is reached a new world is opened to the naturalist,
for there a better acquaintance is made with the great Caribbean
province, which, extending from Florida to the northern shores of
South America, embraces all the Bahamas, the West [pg311] Indies,
the Gulf of Mexico, and the coast of Central America. Strangely
enough, the mollusks of the west coast of Florida are more nearly
related to those of the transatlantic province than are the shells
of the east Florida coast. This remarkable fact in geographical
distribution can only be explained by assuming that an open waterway
once existed across the northern part of the Florida peninsula,
connecting the Atlantic and the Gulf, through which a mingling of the
shallow-water forms from Georgia, the Carolinas, West Florida, and
the Gulf coast took place. Thus it would appear that when Florida
was an island the fauna of its east coast, originally largely
Antillean, was less disturbed by the southern migration of American
species. The Florida Keys are essentially Antillean. The Caribbean
or Antillean fauna is an exceedingly rich and varied one. Among the
most prominent genera which may be encountered upon the Florida coast
are /Strombus/, /Fasciolaria/, /Oliva/, /Marginella/, /Natica/,
/Sigaretus/, /Littorina/, /Tectarius/, /Neritina/, /Melongena/,
/Cardium/, /Callista/, /Tellina/, /Lucina/, and /Cyrena/. In such an
extensive and rich faunal province one must not expect to find in any
given locality a representation of all the characteristic genera. For
instance, collections made at Havana, at Nassau, and at Vera Cruz
would each contain forms more or less peculiar and local, but certain
species would be found in each collection.
* * * * *
On the west coast of America quite the same faunal division into more
or less well-marked provinces is to be found. Bering Sea belongs to
the arctic province, and contains, with Labrador and Greenland, many
identical forms.
THE ALEUTIAN PROVINCE
The shore waters from the southern peninsula of Alaska down to about
Vancouver comprise the Aleutian province. Within this area occur
some arctic species, notably the soft-shell clam, /Mya arenaria/,
which the fishermen gather in such large quantities for bait along
the Maine coast. An increasing number of species common to this
west-coast faunal region and to the boreal province of the east coast
are being recorded. Some of [pg312] these have been introduced from
the East by the agency of man, but others no doubt have a natural
range through the icy waters north of America and down through the
Bering Sea into the Pacific.
THE CALIFORNIAN PROVINCE
The Californian province extends from the neighborhood of Vancouver
to Cape St. Lucas and has about five hundred species. The northern
portion of this area contains some very characteristic forms,
together with a good mixture of species of more northern habitat;
it might properly be regarded as a subprovince. From San Diego
south to the cape the character of the fauna gradually changes;
this coast-line of southern California has also been considered a
subregion. The main characteristic of the Californian province is a
large development of /Haliotis/, of the patelliform mollusks, and
of the chitons. A very few minutes on any good collecting-beach
of California would convince the Eastern conchologist that he had
entered upon an entirely new field of research. The most notable
genera are /Calliostoma/, /Chlorostoma/, /Scurria/, /Acmæa/, various
genera of the /Chitonidæ/, /Purpura/, /Monoceras/, /Amphissa/,
/Norrisia/, /Olivella/, /Chorus/, and /Haliotis/.
THE PANAMIC PROVINCE
The warmer waters of the Gulf of California belong to the Panamic
province, which extends down to South America. This is one of the
richest and most interesting faunas in the world. At Panama alone,
Mr. C. B. Adams, a noted collector and student of conchology, took
upward of four hundred species. About eight hundred species are known
in the Gulf of California alone.
* * * * *
It must be borne in mind that these faunal provinces relate to
littoral and shallow-water species, for the present knowledge of
deep-water forms is not sufficient to permit of generalization upon
their distribution.
We review here only the conspicuous forms along our own coasts. It
may be well to note, however, that in some other parts of the world
the development of molluscan life is far [pg313] in excess of
anything to be found upon our shores. The veritable paradise of the
naturalist is the East Indies. There the "aristocratic" genera, so
called on account of their marvelous beauty, occur in their highest
development--/Voluta/, /Mitra/, /Oliva/, /Conus/, and the various
murices. There also are to be found the pearl-oysters, /Meleagrina
margaritifera/, that yield their valuable harvest, and the giant
clam, /Tridacna gigas/, which measures sometimes five feet in
length. Over eight thousand species of mollusks are described from
this surpassingly rich region, yet this vast province, as compared
with the American and European shores, has been but superficially
exploited by the naturalist. In whatever part of the world a
naturalist may find himself, there is always a tempting array of
molluscan life to attract him. Each fauna possesses features peculiar
to itself, and from the point of view of the true naturalist, the
more somber-hued and conventionally formed mollusks of Northern
shores are no less interesting than the gorgeously tinted and
fantastically shaped species of the tropics. From any faunal province
of our own country one may readily gather all forms necessary to
furnish ample material for study from which one may acquire an
excellent idea of the biological features of the entire phylum.
STATION AND HABITS OF THE MOLLUSCA
The word "station" is used to indicate the nature of the surroundings
which an animal chooses as most suited to his well-being. Some groups
of mollusks, like the littorinas, the trochids, the purpuras, and the
majority of those having patelliform shells, generally live on rocks
above low-tide mark; other genera, like /Buccinum/, /Sipho/, and
/Chrysodomus/, prefer rocky or gravelly ground below low-tide mark.
Other mollusks burrow deep in the mud, many prefer sandy bottoms,
while a host of other species seek homes upon the tangled masses of
seaweed, living like arboreal creatures in the submarine forests.
Nearly every conceivable character of sea-bottom or shore-line
between tides harbors its own peculiar types of molluscan life. There
are some very curious genera of bivalves that bore their way into the
hardest rock and [pg314] there find a comfortable shelter for life.
A numerous class of pelagic or free-swimming mollusks supposedly
never go to the bottom at all, but spend the whole of their existence
on or near the surface of the sea, always in open water, where
their fragile shells may not be injured by rough contact with solid
substances.
The food of some is vegetable, of others, animal. The bivalves, like
the clams, oysters, cockles, and mussels, feed only on microscopic
organisms. They create a current of water through their siphons, or
mantle openings, and then, by a process best known to themselves,
catch and swallow all the animalculæ thus brought to their mouths.
The univalves which possess a siphon are, for the most part,
carnivorous, and are often most voracious creatures. They feed upon
any animal matter they can find, while some of them are enabled by
means of a sharply toothed tongue to bore through the solid shells
of other mollusks and extract the succulent vitals from within. One
energetic little mollusk in particular, /Urosalpinx cinerea/, is
for this reason a great pest upon the oyster-beds. Univalves not
possessed of a siphon may generally be considered herbivorous; they
pass most of their time peacefully browsing upon algæ.
Mollusks are all oviparous or ovoviviparous; that is, they lay eggs,
or, laying eggs, they retain them within their shells until the
young are hatched out. As a rule, each species of mollusk has its
own particular method of protecting its eggs from external injury.
Some construct tough, leathery capsules which are strung together in
various patterns. The egg-capsules of /Purpura lapillus/, resembling
little pinkish or yellowish club-shaped stalks, may frequently
be found in the crevices of rocks and under the rockweeds. The
egg-cases of /Polynices/ (/Lunatia/) are most peculiar, resembling
inverted gelatine-bowls with the bottom knocked out; when wet they
are semi-elastic translucent masses in which may be seen myriads of
eggs. /Buccinum undatum/ arranges its hemispherical egg-capsules
in layers one above the other. The number of eggs so deposited is
often very great, running well into the thousands. Egg-capsules of
/Fulgur/ are leathery coils of angular disks adhering by one edge to
a connecting-band of a similar texture. (See Plate I.) [pg315]
In bivalves the eggs are retained within the parent shell until
hatched, and the young, which then escape into the water, are very
different-looking creatures from the parent. Generally the young
bivalve is free-swimming and exceedingly lively in its movements;
sometimes by means of cilia it is enabled to attach itself to any
passing host, and thus the species is widely disseminated. The fry
soon lose their embryonic form and take on the characteristics of the
adult.
The longevity of mollusks varies greatly with the species. Some
attain an age of from ten to fifteen years, while some are only
annuals. Oysters continue to grow for four or five years, after
which they may survive many years more. In truth, little is known
concerning the duration of life even of our better-known shore
species.
In point of intelligence, mollusks must be relegated to a very
low position. They give evidence of possessing no more than
the most primary instincts, those of self-preservation and of
reproduction. There are some forms that exercise a high degree of
protective mimicry. The highest class of mollusks, the /Cephalopoda/
(cuttlefishes, octopi, etc.), is widely separated from the other
classes in the development of faculties that appear to correspond
with intelligence. They are exceedingly crafty in the pursuit of
their prey as well as in eluding capture.
STRUCTURE OF MOLLUSKS
Among the invertebrate animals, especially in the lower orders, wide
physiological departures from central or typical forms are frequent.
It often happens that a whole group or even suborder will differ
so materially in its general characteristics from another group
or suborder belonging to the same phylum that the student will be
puzzled at first to understand how the two can be nearly related. In
many such cases it is only when the essential features of the phylum
are thoroughly understood that the relationships of its suborders
can be appreciated. An example of widely divergent groups within the
same phylum has already been shown in the /Echinodermata/. Who would
at first suppose that the graceful /Astrophyton/ could bear kinship
to the inert [pg316] holothurian, or that both of these have the
closest family ties with the sea-urchins?
The probable reason for these startling departures from a typical
form is that in the lower orders of life, where the organs and their
functions are comparatively simple, the laws of evolution operate
far more quickly in the adaptation of an organism to environment and
changed conditions of life. A decided tendency to this departure from
the type is a feature of the mollusks. It would be difficult, if
not impossible, to give a general description that would fit both a
common garden-snail and the common oyster, yet both are mollusks. A
concise definition of the phylum is therefore practically impossible,
and it seems here as if nature resented all attempts to circumscribe
her living forms by rigid rules.
The most persistent characteristic of the /Mollusca/ is the
possession of the "mantle," a sort of outer skin which, like a bag or
sack, envelops the creature above and about the sides, but is open
below, permitting the animal at will to protrude or withdraw its
foot. From the outer surface of this sheath-like mantle, or fold, a
calcareous shell is secreted. The modifications of this mantle are
infinite; indeed, in some well-marked groups it seems to have wholly
disappeared, having in reality become fused to the sides of the
animal, and having become therefore a mere outer skin, and no longer
a tunic or cloak, as originally intended. As already noted, the
mantle does not always secrete a shell, a fact which again reminds us
that we cannot give a good comprehensive description of the phylum
that always holds true throughout its subdivisions. The best that can
be done is to give in general terms the characters which appear in
a majority of the forms. A convenient method of going about this is
to create, for the sake of clearness, an ideal, model, or fanciful
mollusk and clothe it with all the most salient molluscan features,
and from such a basis to develop, as we go on, the various types of
the actual orders and suborders. There is one class of mollusks,
the /Amphineura/, which conforms reasonably well in most of its
features to our comprehension of the schematic mollusk. This class is
sometimes referred to by biologists as the "primitive" or "original"
mollusks, from which all the other classes have since been derived;
but the [pg317] evolutionary history of the development of the
various molluscan orders through geological time does not seem to
bear out the fact that an "ideal" mollusk, as we shall construct him,
ever actually existed, or that his prototype, the /Amphineura/, was
the first and consequently the ancestral type of the phylum. We must
therefore only accept this ideal mollusk theoretically, to serve as
a guide for the time being. We may profitably use it only as a lay
figure--a sort of manikin, which will introduce us to the phylum.
After our introduction we must forget our imaginary type and abandon
any notion we may have conceived of the parental position of the
/Amphineura/, its closest imitator in actual life.
THE IDEAL MOLLUSK
The ideal mollusk has a compact, somewhat elongated body, the under
or ventral portion of which is thickened into a muscular disk, upon
which it creeps along. This disk is called the /foot/. (Note a snail
thus crawling upon his "foot.") The foot is rounded or obtusely
pointed posteriorly. Forward, the body is slightly narrowed into
a neck, and then expanded again into a head. Upon this head are
carried two projecting, mobile, fleshy tentacles, one upon each
side, that superficially resemble two horns. At the base of each
tentacle, upon the inner side, is an eye. Just below and between the
tentacles is a small horizontal slit,--the mouth,--the lips being
slightly thickened. The visceral portion of the animal is above the
foot, or /dorsal/ to it, and extends the whole length of the animal
from the mouth to the excretory opening, the latter being situated
posteriorly. The visceral cavity is inclosed above by a thin skin;
indeed, it is but the great thickening of this skin below that
constitutes the foot.
[Illustration: Ideal or schematic mollusk.
1, dorsal view: /h/, head; /t/, cephalic tentacle; /sh./, shell
covering visceral mass; /b.m./, outline of body-mass; /r.m.e./,
edge of mantle; /c.t./, ctenidia, or gills; /q.a./, generative
apertures; /r.e./, renal openings; /e.o./, excretory opening; /f/,
posterior portion of foot; /o.e.m./, edge of shell.
2, ventral view: /h/, head; /t/, tentacle; /m/, mouth; /b/, outline
of body; /e.m./, edge of mantle; /s.f./, lower surface of foot;
/c.t./, ctenidia, or gills; /f/, posterior portion of foot.
3, horizontal section, showing heart, nephridia, and gonads:
/m/, mouth; /o/, esophagus; /g/, gonad; /p/, pericardium; /n/,
nephridium, or renal organ; /v/, ventricle; /a/, auricles; /g´/,
gonad aperture; /n´/, renal opening; /e/, excretory opening; /f/,
posterior portion of foot.
4, vertical median section: /m/, mouth; /b.m./, buccal mass; /l/,
liver; /d.a./, digestive tract; /q/, gonad; /f/, foot; /q.o./,
gonad aperture, or opening of oviduct; /r.o./, renal opening;
/e.o./, end of digestive tract, excretory opening; /n/, mantle;
/r/, nephridium, or renal organ; /o.f./, recurved portion of
mantle; /sh./, shell; /v/, ventricle; /p/, pericardium.]
The digestive tract, beginning at the mouth, opens immediately into a
muscular throat, which is called the /buccal mass/. Within the buccal
mass there is a long, fleshy ribbon that bears upon its upper surface
numerous rows of sharp teeth; these rows of teeth run crossways on
the ribbon, each row containing more or less prominent median teeth,
with several laterals upon each side, and with marginal teeth on each
side of the laterals. This is called the /radula/, and is used for
tearing or rasping the food. When [pg319] not in use it lies coiled
up within the buccal mass. Beyond the buccal mass, food passes into
the esophagus, which widens out into a muscular pouch or stomach;
this may also be provided with chitinous teeth or hard knobs or
plates for the purpose of further triturating the food. Above and
partially surrounding the stomach is a large reddish or brownish
liver, which furnishes to the alimentary canal the digestive juices.
From the stomach the intestine leads to the terminal anal opening.
There are two organs, serving the function of kidneys, placed
dorsally one upon each side of the body (/nephridia/). A heart lies
within the median line of the body, and consists of a ventricle and
two lateral auricles. The vascular system is almost entirely closed;
only in so far as it is partially open does it essentially differ
from that of higher groups; the blood is whitish or light bluish,
and receives its oxygenation in the gills or along the free mantle
surface, through which it courses in exceedingly thin capillaries. In
this manner respiration is carried on to some extent throughout the
whole outer surface of the body. There are two /branchiæ/ (gills).
These have the appearance of small feathers and are placed upon
each side of the animal posteriorly; they project slightly. Spread
over the top of this animal, fitting it perfectly, is a thin fleshy
covering attached to its dorsal portion like a skin, but hanging
free, like a flap, over the sides, the head, and the posterior end
of the animal. Upon the sides it may just cover the tips of the
branchiæ, but in front the head is seen to project from under it.
The loose-hanging portion of this sac-like covering is somewhat
thickened, and perhaps recurved upward upon itself. Upon the edge of
this flap all the way around may occur a row of cilia, or tentacular
processes. This cloak-like covering is called the mantle, and, as
already noted, is the most peculiar and persistent characteristic
of the /Mollusca/. The "cavity" formed between the loose-hanging
portion of the mantle and the sides of the foot is called the /mantle
cavity/, and contains posteriorly the gills, or branchiæ. From the
outer surface of this mantle a calcareous secretion takes place,
forming a shell, which, in our ideal mollusk, will be shaped in
conformity with the surface outlines of the mantle, oblong and convex
above. Over the lower portion of the shell the edge of [pg320] the
mantle recurves. When the creature is disturbed or is at complete
rest it may withdraw that portion of the mantle which protrudes from
below the shell entirely within the edge of the shell, so that the
"soft parts" may be wholly covered, and thus protected from harm. In
such a state of rest nothing may be seen from above but a shell. The
entire upper surface of the mantle is firmly attached to the under or
inner side of the shell, except the mantle margin, this being free.
The nervous system consists of a circumesophageal ring of
nerve-cells, with two cords passing along the digestive tract to the
posterior end of the animal. In the head and above the buccal mass
are several series of nerve-ganglia. These also communicate with the
posterior portion of the animal through two parallel cords which pass
along its ventral side. From these four principal branches, which
traverse the entire length of the creature, minor nerve-cords are
thrown off to the tentacles, the foot, the branchiæ, and the various
organs of the viscera. The sexes are separate.
Reviewing our "typical mollusk," we find that it is bilaterally
symmetrical--that is, uniform upon both sides of a median line from
mouth to excretory opening; is unsegmented; has the ventral surface
thickened into a creeping foot or disk; and is possessed of a head
with tentacular processes, a radula, lateral branchiæ, a heart with a
ventricle and two auricles, a well-developed mantle, and a shell.
If the reader can learn these few details of organization he knows
the structure of a mollusk. In turning to the actual living forms he
will find merely modifications (though sometimes confusing ones) of
these various parts.
CLASSIFICATION
The /Mollusca/ are divided into five great classes: /Amphineura/,
/Gasteropoda/, /Scaphopoda/, /Pelecypoda/, and /Cephalopoda/. These
names, it will be noticed, with the exception of the first, all refer
to the /foot/, the modifications of the foot being selected as the
basis of the first great divisions of the phylum. The /Gasteropoda/
crawl upon the thickened ventral surface of their bodies (already
[pg321] referred to as the foot), as the common garden-snail, the
periwinkles, and in general all those mollusks which have a spirally
coiled shell. The /Scaphopoda/ have a long, worm-like foot, with
which they burrow in the sand or mud. Their shells are like miniature
elephant-tusks, but are open at both ends. The /Pelecypoda/ have a
more or less club-shaped foot, utilized, in the many families, for
a great variety of purposes. They are always inclosed in a bivalve
shell. Familiar examples are the oysters, the clams, mussels, etc.
The /Cephalopoda/ have the /foot/ modified into a number of /arms/,
which encircle the head or the mouth. They are the cuttlefishes, the
octopi, squids, etc.
CLASS =AMPHINEURA=
ORDER =POLYPLACOPHORA=
The /Amphineura/, as already observed, approach most closely to the
ideal mollusk just described. They are bilaterally symmetrical.
This fact is so important that it constitutes them a class,
notwithstanding the fact that in respect to the foot (the basis of
division into classes) they would be included with the /Gasteropoda/,
for (barring some exceptional instances) they creep along upon a foot
quite as our ideal mollusk would, and as the /Gasteropoda/ do. The
head carries no tentacles, thus essentially differing in this respect
from the ideal form. The mantle is extended down in front, completely
covering the head. The branchiæ are confined to a few pairs of
/ctenidia/, or plume-like gills, within the mantle groove or cavity,
and are arranged upon each side of the excretory opening like small
feathers.
There are two orders of the /Amphineura/, the /Polyplacophora/ and
the /Aplacophora/ (or /Solenogastres/), the one name meaning "bearing
many plates," and the other "without plates," the word "plate" in
this sense being synonymous with "shell." The shell of the first
order consists of eight calcareous disks arranged in a longitudinal
row along the back or dorsal side of the animal, which overlap like
shingles on a roof and admit of great variation of form in the
various families. [pg322]
It will be observed from the examples given that the shell plates do
not entirely cover the mantle, but that a considerable portion of the
upper mantle surface is left exposed along the side of the animal.
This exposed portion of the mantle is called the /girdle/. It is
always very much thickened and is often exceedingly tough. In some
species it is covered by a great number of calcareous spicules or
granules which are embedded in the leathery mantle tissue. Strangely
enough, the /Chitonidæ/ (a large family of the /Polyplacophora/)
possess a great number of eyes scattered over the shell plates like
minute dots.
[Illustration: /Chiton spinosus/, dorsal view.]
The members of this order are to be found in all parts of the world,
but their greatest development is upon the west coast of America.
In California the student of marine life has a splendid opportunity
to examine the chitons. They are generally to be found adhering
to the under surface of stones about low-tide mark. They are very
sluggish in their movements and may readily be removed from their
resting-places by suddenly thrusting a knife-blade under the foot
and prying them off. When thus detached they have a habit of rolling
up into a ball and so remaining indefinitely; to prevent this they
should be placed at once in a jar of sea-water, where they will
resume their natural position. For preservation as cabinet specimens,
chitons should be tied down between two boards or flat substances,
to hold them firmly extended until dried; otherwise they are apt to
contract out of shape.
[Illustration:/Chiton/, ventral view: /an/, anus; /cten/, ctenidia;
/ft/, foot; /mant/, mantle-edge; /mo/, mouth.]
Six or eight species are exceedingly common upon the west [pg323]
coast. The collector who has successfully taken a number of these
will not fail to note the varying degrees of width of the girdle in
the different species. In some cases the girdle partially covers the
overlapping shell plates. The largest of all the chitons inhabits
the California shores and is known as ^Cryptochiton stelleri^.
It measures fully six inches in length and three in width. In
/Cryptochiton/ the girdle is continued over the shell plates as a
tough, gritty, brownish covering. The shell plates are white, and,
as detached pieces, are frequently to be found cast up upon the
California beaches, where, from their curious shape, they are often
referred to as "butterfly-shells."
[Illustration: /Chiton ruber./ /Chiton apiculatus./]
On the east coast of America there are comparatively few species, and
these are all small and inconspicuous. The three commonest ones are
^Chiton ruber^, ^C. albus^, and ^C. apiculatus^. They may readily be
found on stones or dead shells fished up from below low-tide mark, or
in rock-pools left by the receding tide.
ORDER =APLACOPHORA=
This second order of the /Amphineura/ is entirely destitute of the
shell plates characteristic of the first. The outer mantle surface,
however, develops a number of calcareous granules which correspond to
a shell. The /Aplacophora/ are symmetrical, having the mouth and the
excretory opening at the anterior and posterior ends respectively.
The foot is greatly modified, being narrowed at times to a mere slit
along the ventral surface. Some of them have long, worm-like bodies.
This is no doubt a very primitive form of mollusk, or, if not a
primitive form, it has greatly degenerated from some higher type.
They are supposed to be commensal in habit and live in the deeper
water offshore.
The /Aplacophora/ are not littoral animals, and it is extremely
unlikely that one will ever encounter them upon the beach, unless,
possibly, after the severest of storms. We may therefore leave them
with this mere mention and proceed to the second class of mollusks,
the /Gasteropoda/. [pg324]
TABLE SHOWING THE CLASSIFICATION OF THE GASTEROPODS AND SCAPHOPODS
DESCRIBED IN THIS CHAPTER
Phylum =MOLLUSCA=
Class =GASTEROPODA=
Order =OPISTHOBRANCHIATA=
Suborder =TECTIBRANCHIATA=
Families Genera Species
BULLIDÆ ^Bulla^ ^B. (Haminea) solitaria^
^B. occidentalis^
^B. nebulosa^
APLYSIIDÆ ^Aplysia^ ^A. Wilcoxii^
Suborder =NUDIBRANCHIATA=
DENDRONOTIDÆ ^Dendronotus^ ^D. arborescens^
ÆOLIDIDÆ ^Æolis^ ^Æ. papillosa^
DORIDIDÆ ^Doris^ ^D. bilamellata^
Order =PROSOBRANCHIATA=
Suborder =DIATOCARDIA=
Families Genera Species
ACMÆIDÆ ^Acmæa^ ^A. testudinalis^
^A. testudinalis,^
Var. ^alveus^
^A. mitra^
^A. patina^
^Lottia^ ^L. gigantea^
FISSURELLIDÆ ^Fissurella^ ^F. alternata^
^F. barbadensis^
^F. (Glyphis) aspersa^
^Lucapina^ ^L. crenulata^
HALIOTIDÆ ^Haliotis^ ^H. splendens^
^H. rufescens^
^H. cracherodii^
TROCHIDÆ ^Margarita^ ^M. cinerea^
^M. helicina^
^M. undulata^
^Calliostoma^ ^C. occidentale^
^C. jujubinum^
^C. annulatum^
^C. canaliculatum^
^C. costatum^
^Chlorostoma^ ^C. funebrale^
^C. brunneum^
^Trochiscus^ ^T. norrisi^
^Livona^ ^L. pica^
TURBINIDÆ ^Turbo^ ^T. castaneus^
Var. ^crenulatus^
NERITIDÆ ^Nerita^ ^N. peleronta^
^N. tessellata^
^N. versicolor^
^Neritina^ ^N. reclivata^
^N. viridis^
Suborder =MONOTOCARDIA=
JANTHINIDÆ ^Janthina^ ^J. fragilis^
SCALIDÆ ^Scala^ ^S. lineata^
^S. multistriata^
^S. groenlandica^
^S. angulata^
NATICIDÆ ^Polynices^ ^P. (Lunatia) heros^
^P. (Lunatia) triseriata^
^P. (Lunatia) lewisii^
^P. (Neverita) duplicata^
^P. (Neverita) recluziana^
^Natica^ ^N. clausa^
^N. canrena^
^Sigaretus^ ^S. perspectivus^
CAPULIDÆ ^Crepidula^ ^C. fornicata^
^C. plana^
^C. aculeata^
^Crucibulum^ ^C. striatum^
^C. spinosum^
LITTORINIDÆ ^Littorina^ ^L. litorea^
^L. rudis^
^L. palliata^
^L. irrorata^
^L. angulifera^
^L. scutulata^
^L. planaxis^
^Lacuna^ ^L. vincta^
^Tectarius^ ^T. muricatus^
^T. nodulosus^
CERITHIIDÆ ^Cerithium^ ^C. floridanum^
^C. muscarum^
^C. minimum^
^Bittium^ ^B. nigrum^
^Cerithidea^ ^C. scalariformis^
^C. sacrata^
VERMETIDÆ ^Vermicularia^ ^V. spirata^
STROMBIDÆ ^Strombus^ ^S. pugilis^
^S. gigas^
CYPRÆIDÆ ^Cypræa^ ^C. exanthema^
^C. spurca^
^C. spadacea^
^Trivia^ ^T. pediculus^
^T. quadripunctata^
^T. californica^
^T. solandri^
DOLIIDÆ ^Dolium^ ^D. galea^
^D. perdix^
^Pyrula^ ^P. papyratia^
CASSIDIDÆ ^Cassis^ ^C. tuberosa^
^C. cameo^
^C. testiculus^
^C. inflata^
Families Genera Species
MURICIDÆ Subfamily Muricinæ
^Murex^ ^M. rufus^
^M. pomum^
^Trophon^ ^T. clathratus^
^Urosalpinx^ ^U. cinerea^
^Eupleura^ ^E. caudata^
^Pteronotus^ ^P. festivus^
^Ocinebra^ ^O. poulsoni^
^O. lurida^
^O. interfossa^
^Cerostoma^ ^C. nuttallii^
^C. foliatum^
MURICIDÆ Subfamily Purpurinæ
^Purpura^ ^P. lapillus^
^P. patula^
^P. hæmastoma^
^P. crispata^
^P. lima^
^P. saxicola^
^Monoceras^ ^M. lapilloides^
^M. engonatum^
^Chorus^ ^C. belcheri^
COLUMBELLIDÆ ^Columbella^ ^C. mercatoria^
^C. (Anachis) avara^
^C. similis^
^C. (Astyris) lunata^
^C. (Amphissa) corrugata^
^C. (Astyris) gausapata^
NASSIDÆ ^Nassa^ ^N. trivittata^
^N. obsoleta^
^N. vibex^
^N. fossata^
^N. perpinguis^
^N. tegula^
^N. mendica^
BUCCINIDÆ ^Buccinum^ ^B. undatum^
^Chrysodomus^ ^C. decemcostatus^
^Sipho^ ^S. Stimpsoni^
^S. pygmæus^
^Siphonalia^ ^S. kellettii^
^Tritonidea^ ^T. tincta^
TURBINELLIDÆ ^Fulgur^ ^F. carica^
^F. canaliculata^
^F. perversa^
^F. pyrum^
^Melongena^ ^M. corona^
FASCIOLARIIDÆ ^Fasciolaria^ ^F. gigantea^
^F. tulipa^
^F. distans^
VOLUTIDÆ ^Voluta^ ^V. junonia^
MARGINELLIDÆ ^Marginella^ ^M. apicina^
OLIVIDÆ ^Oliva^ ^O. literata^
^O. mutica^
^Olivella^ ^O. biplicata^
^O. boetica^
CONIDÆ ^Conus^ ^C. floridanus^
^C. pealii^
CANCELLARIIDÆ ^Cancellaria^ ^C. reticulata^
Class =SCAPHOPODA=
DENTALIDÆ ^Dentalium^ ^D. dentale^
^D. pretiosum^
^D. hexagonum^
^Entalis^ ^E. striolata^
[pg328]
CLASS =GASTEROPODA=
The class /Gasteropoda/ is the largest and most comprehensive
subdivision of the /Mollusca/, and within its wide range of families
many differing details of organization are to be found. As a class it
possesses all the most characteristic features of the phylum, though
it is frequently modified in a high degree. This class includes
all the univalve mollusks (except /Nautilus/ and /Dentalia/), such
as the snails, the whelks, and the host of spirally coiled land,
fresh-water, and marine shells.
The gasteropod foot is, as has been remarked, primarily the same
as in our schematic mollusk--a flat, muscular disk caused by
the thickening of the ventral body-surface. The neck, head, and
tentacles are also quite the same, but considerable modification
of these organs will be found when we come to examine some
specimens. The mantle is always present, except in the nudibranch
or non-shell-bearing forms; but in few of the /Gasteropoda/ is the
mantle so regularly simple as in our ideal mollusk.
Perhaps the most striking feature of the /Gasteropoda/, and
one that will at first surprise him who has in mind the simple
structure of the ideal mollusk, is the fact that they are always
asymmetrical--that is to say, a median line drawn longitudinally
through a gasteropod will not divide it into halves of similar
anatomical structure.
The quality of symmetry is an important one throughout the lower
orders of animal life. In nearly all phyletic or class descriptions
the word "symmetry" occurs, and its exact meaning must be understood.
Take, for example, a human being; a median line drawn vertically
would divide him into two similar halves--upon each side would be
an eye, an arm, a leg, etc., of similar shape and construction. So
far at least as the external features [pg329] go, man is then a
symmetrical creature; likewise the vertebrates in general. Cases
of actual symmetry are found in the lobsters and crabs and in the
segmented worms. It is usual in describing mollusks broadly and in
general terms to call them symmetrical animals, yet here is the
largest class within the phylum, whose representatives are none of
them symmetrical. In the /Gasteropoda/ the mouth is anteriorly placed
in the head, just as in the ideal form, but the digestive tract,
after traversing the visceral cavity in the usual fashion, suddenly
turns forward and terminates in an excretory opening either on the
right or on the left side of the animal, just back of the head. The
various other orifices for the genital and renal glands are also
placed in this unexpected anterior position. The gills, or branchiæ,
are to be found upon one side only, forward of the heart, and
differ somewhat in appearance from the plume-like processes of the
/Amphineura/.
What, then, has become of the corresponding gill we should expect to
find on the other side? The symmetrical shape of the foot is not in
the least altered, but what has happened to cause this distortion of
the visceral portion of the animal? And, finally, will this torsion
of the body of the snail perhaps account for the spiral nature of the
shell?
An evolutionary process is supposed to have taken place in mollusks,
which, if real, would no doubt account for these curious conditions
of torsion in the /Gasteropoda/; but the theory upon which this
process is based is altogether speculative, and is not fully
sustained by the facts in the case. Originally all mollusks are
presumed to have been symmetrical, and are assumed to have resembled
very much in form the schematic creature we have already described.
Now, for some reason, certain of the primitive mollusks, but not all
of them, began to develop a larger visceral mass, which, continuing
to enlarge throughout many generations, began finally to protrude
above and form a hump on the dorsal side of the animal. This hump,
containing the liver, a portion of the intestines, and the generative
glands, as it increased in bulk became so much elevated that it could
no longer maintain itself in an erect position over the body, but,
impelled by its own [pg330] weight, began to sag over to one side.
Such a process would of course tend to twist the digestive tract and
bring the excretory opening constantly farther forward on one side.
At the same time, this process would necessarily interfere, by the
weight of the overhanging viscera and the shell covering the mass,
with the development of the branchiæ lying upon that side and now
crowded under the visceral mass. Thus, according to this theory, this
process has continued until, in the modern gasteropod, the digestive
tract has been bent upon itself and twisted from a straight course
into a curve of almost 180°, bringing the excretory opening near or
just over the head, where it empties into the mantle cavity; while
the original right gill, by the same movement, has been brought to a
position on the /left/ side of the head, forward of the heart, the
original left gill having been crowded out, eventually to atrophy and
disappear. The same torsion is found in the nerve-cords; the heart,
situated at about the pivotal position of this twisting process,
has turned about upon itself, leaving an auricle in front of the
ventricle, one auricle, like one of the gills, having been lost. The
mantle portion covering the visceral hump naturally continued to
secrete its shell, though always in conformity with the change, the
result being the familiar spiral form of the usual gasteropod shell.
This theory may not be satisfactory, but the asymmetry of gasteropods
is a problem to be solved, and a more interesting line of biological
investigation could not be found.
Let us now take a good example of a gasteropod and locate its various
organs; at the same time we may use the occasion to refer to more
important modifications of these organs which will be encountered
later in the various genera. The most available gasteropod on the
east coast of the United States for this purpose, is /Fulgur/, both
on account of its large size and its abundance. /Buccinum/ may be
used if the student is north of Cape Cod and therefore unable to
secure a good living specimen of /Fulgur/; the anatomical differences
between the two are slight.
Note the siphon protruding forward from a notch in the shell. This
consists merely in an elongation of a fold of the mantle, which is
held in a manner to constitute a tube, through which [pg331] the
animal draws in water to supply the gills, lying just back of the
siphon and concealed under the mantle in the cavity already spoken
of as the mantle cavity. With the exception of that portion of the
mantle which constitutes the siphon, no part of its margin, it will
be observed, can be seen, the edge of the shell completely hiding it
from view.
In some genera the siphon is exceedingly long--in fact, longer than
the body of the animal. In these cases the mollusk is generally fond
of burying itself in the sand and maintaining communication with
the world above only by means of this long, fleshy tube. In some
genera the siphon is protected by an elongation of the shell, as in
/Fasciolaria/ (Plate LXXVI). An extreme example of this is given in
the figure on page 343, which represents a species of /Murex/, where
the siphon is not only protected by the shell, but the shell itself
is further protected by the double row of sharp spines with which it
is beset.
[Illustration: /Buccinum undatum/: /si/, siphon.]
There are some genera which possess no siphons at all, in all of
which cases there is no notch in the shell opening, the aperture
being round, as in /Natica/ and in /Littorina/ (pages 368, 371). The
mollusks of this type make a fold in the forward part of the mantle
which serves as a siphon. Such a muscular folding in the mantle would
be technically called a /functional siphon/.
On the posterior dorsal side of the foot is attached a horny plate,
of concentric structure about a central or subcentral point or
nucleus. This is called the /operculum/, and serves to close the
entrance of the shell when the animal is withdrawn, the object no
doubt being protective. The opercula vary to a great extent in the
different genera; in some cases they are calcareous, in others
cartilaginous. When the aperture of the shell is very large it often
happens that the operculum does not entirely close the opening, as
in /Fulgur/, but in /Buccinum/ it is a close fit. The operculum is
entirely wanting in the /Pulmonata/, one of the [pg332] large orders
of the /Gasteropoda/. The figures represent some of the commoner
types of opercula.
[Illustration: Various forms of opercula. /Turbo/ (/Sarmaticus/).
/Turbo/ (/Callopoma/). /Livona./ /Ampullaria./ /Natica./ ]
The opercula have been utilized a great deal in the classification
of the /Gasteropoda/, and especially in some groups that have become
terrestrial in habit. Often, when shell-characters utterly fail to
give a clue to the proper generic position of a new or rare mollusk,
the operculum may be relied upon to indicate it.
[Illustration: Various forms of opercula. /Pyrula./ /Purpura./
/Littorina./ /Aulopoma./ /Torinia./ /Neritopsis./ /Strombus./
/Conus./]
TENTACLES
Two tentacles project like fleshy horns from the sides of the head.
These are not retractile, as in the land-snails, and their office
appears to be that of sensory organs. In both /Fulgur/ and /Buccinum/
the tentacles are comparatively short and stout, and have expanded
bases; in many other genera they are long and slender. Almost without
exception the /Gasteropoda/ possess tentacles. [pg333]
EYES
In /Fulgur/ the eyes are situated near the bases and upon the outer
sides of the tentacles; in /Buccinum/ they are extremely small and
are mounted about midway between the bases and the tips of the
tentacles. This is the usual position for the eyes in the marine
/Gasteropoda/. Only in the pulmonates, or land-snails, are the eyes
placed at the summit of the tentacles. Sight does not appear to be
a very important sense in the gasteropods, although the eye in some
instances becomes a highly developed organ. A number of forms that
have the habit of burrowing deep into the sand or mud are quite
destitute of visual organs, and certain cave-dwelling land-shells
have entirely lost the sense of vision.
MOUTH AND PROBOSCIS
The mouth will easily be seen in both /Fulgur/ and /Buccinum/. It is
a small slit with slightly thickened, fleshy lips, and is placed on
the lower forward portion of the head. In the case of both of these
genera the mouth is situated at the end of a proboscis. It will be
noted that /Fulgur/ has a comparatively long and stout proboscis, or
snout, which is not retractile. /Buccinum/ has a smaller one. It is
fairly safe to assume that all gasteropods that possess a proboscis
are carnivorous. This organ is greatly developed in some genera,
where it actually exceeds in length the rest of the animal. In such
instances, no doubt, the creature is enabled to reach the interior
portions of the shells of its victims and to devour every vestige
of their flesh, for the mouth is most conveniently placed just at
the end of this long, trunk-like organ. The genera which have no
proboscis are, almost without exception, vegetable feeders.
We must now remove the animal from its shell. It is well to boil
it first, for this does away with the annoying mucous secretion
that is freely exuded by glands in the foot when the creature is
roughly handled. Alcoholic specimens are apt to be much contracted
and hardened, and are therefore less satisfactory as subjects for
anatomical study. If it is not convenient to boil the [pg334]
specimen, break away the shell with a hammer, using care not to
lacerate the soft, fleshy portion within; when this is accomplished,
wash the animal carefully to remove the slimy exudation. Note the
"visceral hump," which is spiral, and which formerly occupied the
upper-whorl portion of the shell. The thin skin covering it is the
mantle, which below is greatly thickened and free, lying about the
foot like a heavy fleshy flap.
MANTLE
The mantle-edge in both of these examples is simple; that is to say,
it possesses no fringe of tentacles, nor is it supplemented by extra
processes, characters which mark many genera of marine /Gasteropoda/.
In /Fulgur/ and /Buccinum/ the mantle-edge does not protrude below
the edge of the shell; but in many genera, especially those which
possess smooth, glossy shells, like the cowries (/Cypræa/) and the
graceful /Oliva/, the mantle is proportionately very much larger. In
these two genera, when the animal is extended, as in crawling about
the sand, the mantle curves upward and incloses a large portion of
the shell itself. Indeed, in some genera the shell is almost entirely
concealed by this extension of the mantle (/Sigaretus/, /Natica/,
etc.).
When the shell is removed, the folding of the mantle which
constitutes the siphon can be plainly seen. The office of the siphon
has already been referred to, also the fact that the presence of a
siphon in the gasteropod mollusk may always be determined by merely
glancing at the shell alone, for a notch at the base of the aperture
indicates the place through which the siphon passed. In /Buccinum/
this is merely a notch, but in /Fulgur/ the siphonal canal of the
shell is much longer. Just why the long siphons of some mollusks
should be naked and exposed to danger, while others are so carefully
protected by elongated, portions of the shell, is a mystery, but
nature is full of such contradictions.
THE GASTEROPOD FOOT
The foot is long, broad, and flat on the under side, like a disk.
The variations in the gasteropod foot are almost infinite. [pg335]
In some of the species it is amazingly large and powerful, as
in /Polynices/ (/Natica/), and in most of the forms which live
along sandy beaches. These are sometimes provided in front with a
wedge-shaped process called the /propodium/, which serves admirably
as a plow to push aside the heavy wet sand through which the animal
forces its way. In /Nassa/, which is so common all along our coasts,
the foot has two terminal appendages or points behind.
[Illustration: /Sigaretus l[oe]vigatus/, showing excessive
development of the propodium (/pr./) and metapodium (/met./) in a
mollusk living in sand (the shell, which covers only the liver and
adjacent parts, has been removed): /l/, liver; /s. ap./, aperture of
proboscis, here deflected from the median line; /t/, /t/, tentacles;
/f/, foot.]
[Illustration: /Oliva textilina/ showing how the front part of the
foot (/f/) is developed into a sort of fender, the propodium (/pr./):
/e/, /e/, eyes; /m. ap./, front appendage of mantle; /m. ap.´/,
hinder appendage of mantle, folded into the suture when the animal is
at rest; /si./, siphon; /t/, /t/, tentacles.]
There is no single feature of mollusks more important than the foot,
for upon the many modifications of this organ the various molluscan
classes are founded. The name /Gasteropoda/ means "stomach-foot,"
the latter organ being merely a thickening of [pg336] the ventral
surface of the animal, giving to the creature, when active, the
appearance of crawling upon its stomach.
If a living gasteropod is placed in a jar of sea-water and left
undisturbed it will soon crawl up the side of the glass and thus
afford an excellent view of the extended under surface of its foot.
The majority of the gasteropods are lethargic and slow in movement,
but others will be found to be exceedingly lively and able to creep
up the sides of the jar and make their escape in a surprisingly short
time. Further modifications of the foot will be mentioned later.
THE MANTLE CAVITY
Now let us place the animal with the foot below and the head facing
us. The spiral visceral hump will point back and away from the
observer. The mantle covering the visceral portion is thin and
translucent like an ordinary skin, but at the base of the visceral
mass the mantle suddenly thickens very greatly, and spreads over
the dorsal portion of the foot or the body like a cloak. The space
between the body and the loose-lying mantle is called the /mantle
cavity/, and within this area are to be found several important
external organs. To find these it is well to cut the mantle flap in
a straight line, beginning just back of the head and ending at the
point where the mantle is attached to the body. This will most easily
be done with scissors. Having thus divided the mantle in front, throw
back the two flaps. On the right-hand side (left of the animal), and
attached to the inner side of the mantle thus exposed, will be seen
the branchiæ, or gills.
THE BRANCHIÆ, OR GILLS
They consist of a row of flattened filaments bound together at the
base like the leaves of a book. The blood is conveyed to these
gills by a large vein, and is then forced through the thinly walled
filaments, being thus brought into close contact with the water, to
which it releases its carbon dioxide, and from which it receives the
life-giving oxygen. In certain gasteropods there are two sets of
gills, one placed upon each side of the body [pg338] (as in the
ideal mollusk); but the presence of two sets of gills only indicates,
in these gasteropods, that the twisting process of the viscera,
already alluded to as so remarkable a feature of this class, has not
proceeded far enough to destroy the functional activity of one of the
gills.
[Illustration: Anatomy of the internal organs of a female gasteropod,
viewed from the dorsal side. The roof of the mantle cavity has been
divided by a longitudinal incision and the flaps laid out, that on
the left bearing the ctenidium and osphradium, and that on the right
the rectum and terminal part of the oviduct. The muscular dorsal wall
of the body and the introvert have been divided so as to bring into
view the anterior part of the alimentary canal and a portion of the
nervous system. The buccal cavity has been tilted up and opened so
as to show the odontophore, and the esophagus has been cut through
near the anterior end. A portion of the ventral wall of the crop
has been removed so as to bring the internal folds into view, and
the interior of the nephridium with the contained portion of the
intestine has been exposed. The stomach is not seen, being hidden by
the nephridium, and the ovary is not represented. /an./, anus; /ant.
aort./, anterior aorta; /aur./, auricle; /buc./, buccal cavity; /cer.
buc. con./, cerebro-buccal connective; /cer. g./, cerebral ganglia;
/crop/, crop; /cten./, ctenidium; /int./, intestine; /jaw/, jaw;
/l. buc. g./, left buccal ganglion; /l. sal. gl./, left salivary
gland; /neph./, nephridium; /neph. ap./, nephridial aperture; /od./,
odontophore; /oes./, esophagus; /oes'./, anterior end of same, cut
and turned aside; /osph./, osphradium; /ovid./, oviduct; /ovid'./,
terminal thick-walled portion of oviduct; /pleur. g./, pleural
ganglion; /post. aort./, posterior aorta; /post. oes./, posterior
esophagus; /rad. s./, radula sac; /r. sal. gl./, right salivary
gland; /rect./, rectum; /sal. du./, salivary duct; /siph./, siphon;
/supra. g./, supra-esophageal visceral ganglion; /tent./, tentacle;
/vent./, ventricle.]
In general, it may be said that the breathing process of mollusks
does not markedly differ from the respiratory methods of higher
forms of animals. Whether the breathing is effected by gills or by
a pulmonary sac, the essential features of the system are the same
as in fishes or mammals. There are, however, some rather peculiar
developments in the breathing system of gasteropods that are worthy
of notice. The gills themselves differ throughout the various orders
of this class both as to their structure and their relative position;
in one of the orders (the /Pulmonata/) the gills have entirely
disappeared, and are replaced by a pulmonary sac which fulfils the
purpose of a lung.
It is a generally accepted theory that all land or air-breathing
mollusks were originally marine, but by a gradual change in their
habits have developed into terrestrial forms, losing their gills
and acquiring in their place organs which correspond to lungs. The
theory is supported by the fact that many species of marine mollusks
live between tide-marks, and for several hours daily are obliged to
maintain life outside of their native element. The /Littorinidæ/,
which may always be found in abundance on both the east and the west
American coasts, usually live high above low-tide line, and some
tropical species of this genus actually live in the trees above
the reach of high tide. They have gills just as in /Buccinum/ or
/Fulgur/, but the /Littorina/ shell is strongly suggestive of certain
land forms, and there is reason to believe that the genus is in
course of evolution into a pulmonate. On the other hand, there are to
be found in fresh water numerous genera which are true pulmonates,
and which are obliged to come to the surface every few minutes to
obtain air. One notable genus of marine gasteropods possesses both
lungs and gills, and one marine form has entirely lost its gills
and breathes only by means of lungs. It is quite likely that all
mollusks are able to breathe more or less all along the exposed
mantle surface, which is usually [pg339] crowded with capillaries
with exceedingly thin walls. This very remarkable method of surface
respiration is proved to exist in the nudibranch or shell-less marine
/Gasteropoda/, for they possess neither gills nor lungs.
OSPHRADIUM
Lying just under the gills, and side by side with them, is a smaller,
plume-like organ having the appearance of a smaller gill; it is
called the /osphradium/. The office of this organ is not definitely
known, but the supposition is that it is the seat of the olfactory
nerves. The osphradia are not always present, but whenever found they
are placed in close conjunction with the gills, in order, probably,
that the animal may best determine the quality of the water brought
by the siphon to the breathing-organs. In /Buccinum/ the osphradium
is especially prominent. In all mollusks the sense of smell is highly
developed. A dead animal anchored to the bottom will very soon
attract the carnivorous /Mollusca/ from every direction. This is not
a bad way to trap certain species that live upon rocky stations and
are otherwise difficult to secure by the dredge.
On the left-hand side (right of the animal), and adhering to the
under side of the mantle flap, which has been thrown back, a somewhat
inflated and convoluted tube, the rectum, will be seen. It terminates
in an opening just within the mantle-edge. By the side of this are
the renal and genital openings.
* * * * *
Having now examined the external organs, we may investigate the
internal anatomy of our subject. To do this thoroughly requires
considerable skill, but for the purpose of merely discovering the
relative positions of the principal internal organs a little care
only is necessary. The work will be greatly facilitated by boiling
the animal for a few minutes, for this serves to harden the tissues
and to separate the various organs in a most satisfactory manner.
To begin, open the proboscis from the mouth, cutting back so as to
expose the esophagus. It will be seen that the throat is long and
very muscular. [pg340]
RADULA
Hidden among the red stripes of muscular fiber which line the inner
surface of the throat will be found a fleshy strip covered upon
one side with many transverse rows of chitinous teeth. This is the
/lingual ribbon/, or /radula/. Remove as much of it as possible, and
examine it with a hand-glass. The radula is a most important organ,
and is a unique possession of /Mollusca/, except the pelecypods, or
bivalves. Usually it lies coiled up like a spiral when not in use;
its purpose is to tear and rasp food.
[Illustration: Three rows of teeth from the radula of /Fasciolaria
trapezium/.]
[Illustration: Portion of the radula of /Imbricaria marmorata/.]
The radula has been the subject of much study by conchologists, and
has furnished the basis for an arrangement of the numerous sections
of gasteropods into somewhat well-defined groups. If the observer has
a strong glass he will notice that each transverse row of teeth on
the ribbon consists of a median or central tooth with several points
or "cusps." /Fulgur/ has a median tooth with three cusps; /Buccinum/
one with six. On each side of the median tooth is a lateral tooth,
which in /Fulgur/ will be seen to have six cusps, and in /Buccinum/
five. In many patterns of radulæ there is, besides the median and
lateral teeth, still another paired row, called the marginals. In
the accompanying figures are given some examples which show small
sections of lingual ribbons. While all the radulæ of mollusks are
capable of being grouped by their general characteristics, yet in no
two species of mollusks are the radulæ quite identical. [pg341]
VITAL ORGANS, STOMACH, LIVER, RENAL GLANDS, ETC.
The digestive tract will be found to widen out a short distance back
into a crop or stomach, and, continuing still further, to lose itself
in a soft, brownish mass within the coiled spire, which is the liver.
The very large size of the liver leads one to suspect that /Fulgur/
and /Buccinum/ must be voracious creatures to need so large an organ
for the secretion of bile. In many species of mollusks the stomach
and intestines are filled quite solidly, at times, with a gelatinous
transparent substance called the /crystalline stylet/. Just why the
digestive tract should be clogged with this substance no one has
yet been able to explain, so here again is a chance for original
investigation. The intestine curves about after reaching the liver,
and comes forward again to appear once more as the rectum, clinging
to the inner surface of the free portion of the mantle.
Closely associated with the liver, but differing slightly in color,
is the gonad, or organ in which the genital products are formed.
Situated dorsally and forward of the liver and gonad is a large renal
gland, which may readily be detected by its peculiar structure. In
some forms the kidney is closely associated with the gonad, and seems
to coöperate with the latter in the generative functions.
HEART AND VASCULAR SYSTEM
To find the heart, make an incision into the body just at the
posterior end of the gills. The heart is white and round, and
is inclosed within a cavity known as the /pericardium/; it has
a ventricle and one or two auricles, although in /Buccinum/ and
/Fulgur/ there is but one auricle.
There is nothing remarkable about the vascular system to distinguish
it from that of many higher forms of animals. It is, however, not
completely closed--by which is meant that the blood is not always
contained within arteries or veins, and that it does sometimes flow
into other organs and floods certain other body-cavities, although
the vascular system of mollusks is by no means so completely open as
is that of insects and crustaceans. [pg342]
Having examined the animal which inhabits the shell and gained an
idea of its anatomy, we may now turn to an inspection of the house
which it has built for itself, and here we shall find a delightful
subject for study.
THE GASTEROPOD SHELL
An industrious hour of collecting upon almost any beach will provide
the student with an abundance of interesting shells. Putting aside
the bivalves, the univalve or gasteropod shells remaining should be
closely examined for the purpose of becoming acquainted with their
various conchological features. The collector will see at once that
he has a number of species that differ more or less widely in their
general form, size, texture, and sculpture. Some shells will be long
in proportion to their width, with many turns of the spire which
probably terminates in a sharply pointed apex; others will be almost
round, with comparatively few spiral turns which end in a blunt apex,
giving to the specimen outlines suggestive of a dome. A wide range of
characters will be found in the mouths or apertures of the shells,
some being almost if not quite round, others oblong and with a notch
cut into the lower portion of the opening, and others possibly with
this notch extended into a sort of semi-inclosed channel. A first
lesson in classification of the /Mollusca/ may be taken by dividing
the results of the first day's collecting of univalve shells into
groups according to these prominent shell-characters.
[Illustration: Parts of a gasteropod shell: /a/, anterior canal; /b/,
body-whorl; /c/, columella; /l/, outer lip; /m/, mouth or aperture;
/p/, posterior canal; /s/, sutures; /sp/, spire; /u/, umbilicus.]
A good knowledge of the parts of a gasteropod shell is essential to
the student, and it can readily be acquired. Four specimens are given
in the accompanying figures, representing highly divergent [pg343]
types of marine gasteropods, and their respective characteristics may
be noted by comparing them one with another: /Fasciolaria tulipa/
(Plate LXXVI.), a Floridian shell; /Polynices heros/, a common
species found on all the sandy shores of the American Atlantic
coast; /Murex tenuispina/, an Indo-Pacific species much prized by
collectors; and /Acmæa testudinalis/, commonly found in Maine and
Massachusetts at low tide clinging to rocks.
[Illustration: /Polynices heros/; /Acmæa testudinalis/, from below;
/Acmæa testudinalis/, profile; /Murex tenuispina/.]
The extreme top of the shell is called the /apex/, and it may be
either sharply pointed or obtuse. Each turn is called a /whorl/; the
last one, and always the largest, is called the /body-whorl/, the
others collectively forming the /spire/ of the shell. When the animal
first emerges from the egg it is always provided with one or two
whorls, which generally may be distinguished at the apex as smoother
than the rest of the spire, and a faint line may be discovered where
the shell began its growth after birth. These [pg344] first whorls
of embryonic growth are usually referred to as the /nepionic whorls/.
The interstices between the successive whorls are called /sutures/,
and these lines of juncture may be shallow or deep according to the
convexity of the whorls. The sculpturing of the whorls is often
of great complexity, and exhibits /nodules/, /varices/, /ribs/,
/revolving striæ/, /spinous processes/, and /lines of growth/.
It frequently happens that there is a complete absence of any
sculpturing whatever, the whole surface of the shell being perfectly
smooth, save for the faint lines of growth to be detected only under
a magnifying-glass. The outer edge of the mouth or aperture of the
shell is called the /outer lip/. This may be more or less thickened,
expanded, or simple. The inner lip is called the /columella/, and may
be greatly extended anteriorly. When the mouth of the shell is round,
and there is no notch or indentation either above or below, it is
said to be /continuous/.
The /anterior canal/ is for the accommodation of the siphon of the
animal, which, as noted above, may be very long or almost entirely
absent; therefore the character of the shell generally indicates
whether or not the animal itself possesses a long siphon. There is in
some forms a notch at the upper extremity of the mouth, which is for
the accommodation of the anal discharges. When present it is called
the /posterior canal/.
It will be noted that all the specimens figured as examples, save
one, are spiral, the coils revolving about an imaginary axis. When
the whorls are not too closely coiled there is left open to the view
a hollow space throughout the entire length of the shell, which is
called the /umbilicus/. When the coils of the shell are very compact
there is no umbilicus to be seen. The umbilical opening may be very
large, giving a clear view of the inner surfaces of all the whorls
right through to the apex. The usual gasteropod shell is in reality a
hollow tube wound about an imaginary axis. Since the tube increases
in size as it progresses, the base of the shell is necessarily
wider and larger than the apex. Thus all gasteropod shells may be
reduced in shape to the single type of a cone or top, modified by
the excessive length or shortness of the spire, or otherwise altered
by special peculiarities. In /Polynices duplicata/ (page 368), it
will be observed that while the [pg345] umbilicus is fairly large,
its external opening, in the adult shell, has been almost completely
closed by a callous process thrown off from the inner or columellar
lip.
Gasteropod shells may be /sinistral/ or /dextral/, according as
the whorls turn to the left or right. The great majority of marine
species are dextral, having the opening on the right, although a
few species exhibit the curious property of being constructed upon
either the one or the other plan without apparent reason. Sinistral
specimens of many normally dextral species have been discovered, but
they are so rare as to warrant the belief that such specimens are
merely deformities or monstrosities resulting from some accident of
birth.
The collector will soon learn to distinguish between young and adult
specimens, for those finishing touches of wonder and beauty which
the mollusk gives to his protective covering are made only by the
adult. The outer lip of young gasteropod shells is usually thin and
fragile; even if slightly thickened, it has an unformed or unfinished
appearance.
Shells differ greatly in their /structure/, many species being
porcelanous like china, others glassy, and many more are of a softer
chalky composition. In the latter case the shell is usually covered
with a thick membranous skin, which, when removed, leaves a dull,
lusterless, white body beneath. Shells of this texture, when cast
upon the beach, soon yield to the weathering action of sun and air.
The porcelanous shells are composed of successive layers of carbonate
of lime, throughout which is a filmy membranous framework of a
substance similar to that of which the covering of the /Crustacea/ is
formed. The mode of deposition of the various layers of crystalline
calcic carbonate and the peculiar lineation of their outer surfaces
give rise to the iridescent or nacreous appearance of some shells.
With the exception of those species in which the mantle of the
animal is extended over the edge of the aperture and more or less
envelops the entire test, there is an outer skin of membranous or
organic matter which overlies the surface of the shell. This skin
is sometimes quite thick, often hairy, and is usually of a dull
greenish- or brownish-yellow color. In many genera this [pg346]
outer covering of the shell is excessively thin. In old specimens
the epidermis is usually worn away from the apical whorls. It often
happens that the shells of the most exquisite colors and markings are
thus rendered somber and unattractive by their tenants during life;
when the animal dies, or when such specimens are kept in a cabinet
for a period of time, the epidermis dries, cracks, and falls off,
revealing the wealth of color and design beneath.
The growth of the gasteropod shell is accomplished by the exudation
from the margin of the animal's mantle of a liquid containing the
shelly matter in solution. The mantle-edge is provided with a
complicated system of glands and pores, from which is secreted this
"stony liquor." This more or less viscous liquid, containing the
carbonate of lime and the other inorganic materials of which the
shell is composed, hardens upon exposure, and the shelly matter is
then deposited in crystalline form around the edges or lip of the
shell aperture. The gasteropod shell therefore grows by the continual
building out of its aperture through successive depositions of shelly
matter at the extreme edge of the lip. At the extreme edge of the
mantle margin are situated those glands which secrete the materials
for the epidermis of the shell, and as one would therefore expect,
this outermost layer of epidermis is first produced in the advancing
growth of the shell. There also are situated the pigment-glands,
which produce the color-secretions. The various layers of the shelly
substance are successively deposited inside the mouth of the shell by
glands situated just back of the extreme edge of the mantle margin.
Thus in the growth stage, if one could examine closely the aperture
of a gasteropod shell, one would observe at the extreme tip of the
lip this projecting epidermis, just beneath it and just inside the
aperture a thin deposit of shelly matter, just beneath this, and
farther in, another layer, and still farther in a third layer.
The growth of nearly all gasteropod shells is marked by periods of
rest. During the inactive seasons the creature may thicken the edge
of the aperture to a greater or less extent by an extra deposit of
shelly matter, for otherwise the thin lip might soon be [pg347]
injured or broken. These places where the lip was thickened, as a
temporary protection until the young animal decided to continue
building, appear on the surface of adult shells, and persist as
/varices/, "raised lines of growth," "longitudinal ribs," etc. Such
forms of sculptural markings generally occur at equidistant points
and preserve a scheme of regularity which would indicate that the
periods of rest were of seasonal occurrence. All irregularities and
all surface features of the external shell simply reflect certain
peculiarities of the mantle margin of the animal; all spinous
processes, for instance, as in the marvelous /Murex tenuispina/,
indicate the existence of finger-like processes extending from the
mantle margin, which once occupied the hollow thorns projecting
from the shell. When the outer surface of a shell is devoid of
any sculpturing and is perfectly smooth, one may assume that the
mantle margin of the animal was simple. If the edge of the lip has
a crenulated appearance and the surface of the whorls is decorated
with revolving ribs, the mantle margin was probably folded or wavy.
There can be no doubt but that the entire mantle surface is provided
to some extent with glands for the secretion of shelly matter, for
if any portion of the shell is accidentally injured the animal soon
repairs the break with a calcareous deposit. But such repairs are
never homogeneous in texture with the other parts of the shell of
normal growth; the patches are never covered externally with an
epidermis, and they are always devoid of color. The function of the
epidermis is to protect the calcareous shell from the corroding
agents contained in sea-water. Hence it is that, when the epidermis
is removed, "dead shells" exposed to the influence of the water
or the weather soon lose their brilliancy and luster, and become
undesirable for specimens either for the cabinet or for study. It
often happens that living shells, inhabiting a region where, for one
cause or another, the water is highly charged with impurities of an
acid nature, are discovered to be badly corroded about the apex,
where the epidermis is thin and likely to be rubbed off. For this
reason specimens collected in harbors near large cities or near the
mouths of rivers are apt to be poor and defective.
The inner surface of all shells, if not actually nacreous, is
[pg348] always perfectly smooth. When foreign substances, such as
grains of sand, enter the shell and cannot be removed by the animal,
the irritation caused thereby to the soft, fleshy creature induces
discharges of a liquid from the glandular surface of the mantle,
which hardens about the offending substance and glazes it over with a
smooth, pearly deposit.
/Acmæa testudinalis/ (page 343) presents a special type of gasteropod
shell which is found in several families. Here the spire seems to
be wholly absent, and the entire shell consists of but one large
body-whorl. There are very many of these patelliform species,
inhabiting many seas and belonging to many different genera, and
in nearly all cases their embryonic shells display a spiral form.
After birth the animal does not build his house upon the spiral
plan, but expands the shell into one large shield-like covering.
The student, however, must not presume that /Acmæa/ is an ancestral
type just because the simple character of the shell is suggestive
of the model chosen to represent a schematic mollusk; anatomically
/Acmæa/ presents the complications of body-torsion which show a very
considerable evolutionary change, and indicate that its simple shell
is probably a degenerate form of a once more highly developed and
convoluted one.
The forms, the architecture, and the painting of gasteropod shells
are so infinite in variety that it would be unwise to attempt a
description of their marvels. A close observer of nature's works
soon becomes prepared for every surprise, but he never ceases to be
charmed and fascinated by his new discoveries. The careful student
alone can learn really to see and appreciate the wonders of nature,
and this is especially true in the study of the /Mollusca/.
CLASSIFICATION OF GASTEROPODS
The /Gasteropoda/ far exceed all the other divisions of the
/Mollusca/ in the number of their genera and species. Apparently
this has not always been the case. There is evidence tending to show
that in past geological epochs the pelecypods (the bivalve shells)
outnumbered the gasteropods, but that in the course of time the
increase in the genera of gasteropods has been more [pg349] rapid.
To judge also from the very large number of extinct families among
the cephalopods, which once flourished in astonishing abundance, it
would appear that they are a dying race.
The classification of the /Gasteropoda/ is primarily a division into
three great orders: the /Opisthobranchiata/, the /Prosobranchiata/,
and the /Pulmonata/. The /Pulmonata/, which include the land and
fresh-water gasteropod shells, are characterized by the presence of
a breathing-sac, or lung, instead of gills. They are, therefore,
essentially air-breathers, and would perish if kept too long under
water. The /Opisthobranchiata/ and /Prosobranchiata/ are marine,
and, like all animals which breathe by means of gills, soon die
when removed from the water. We have already noted, however, how
some genera of marine gasteropods that find their station about
high-tide mark are able to live for considerable periods out of the
water, notwithstanding the fact that they are fully equipped with
gills and not possessed of lungs. Indeed, there is one large family
of prosobranchs (the /Cyclostomatidæ/) that has become entirely
terrestrial in habit, its members having lost their gills and
acquired lungs, but their organization otherwise is so essentially
that of the prosobranchs that they have never been considered as
pulmonates.
The main difference between these two orders of marine gasteropods
is that in the prosobranchs the breathing-organs (the gills) are
placed in a position forward of the heart, and the degree of torsion
(page 330) characteristic of this molluscan class has been continued
until the auricle of the heart is in front of the ventricle. There
is always a shell, usually spiral, and, with few exceptions, an
operculum. In the opisthobranchs, on the other hand, the relative
position of the heart and gills is reversed, and they further differ
from the prosobranchs in that the sexes are always united in each
individual. The opisthobranchs are not always provided with a shell.
One division of the order, known as the "nudibranchs," are entirely
naked. These are commonly called the "sea-slugs," and are to be
found crawling about the marine vegetation in shallow water, in
tide-pools, and on the piling of old docks. Curiously enough, the
sea-slugs (see page 354) have no gills at all, but, having lost these
apparently essential organs, are [pg350] enabled to "breathe water"
through their skin. They are wonderfully beautiful creatures, and the
collector must not fail to secure some specimens, put them into a jar
of sea-water, and watch them expand.
There are other more superficial differences between the prosobranchs
and the opisthobranchs, which will enable even a beginner to
distinguish them at a glance. When the latter are possessed of an
external shell, it is bulbous, generally glassy, and with a simple
lip, the aperture extending the entire length of the shell. Again,
the mantle or the propodium of the foot is greatly extended and
usually covers the shell almost wholly.
With the exception of the nudibranchs, which are common all along
the Atlantic shore, especially north of Cape Cod, there are very few
opisthobranchs to be found in American waters. Their shells are not
very abundantly found anywhere, though in Florida there are two or
three species which an untrained collector might discover.
Practically all the gasteropod or univalve shells that will be taken
by the ordinarily expert collector along the shore are prosobranchs.
This order includes the great majority of marine gasteropods, and
is entitled to the distinction of claiming, in the great number of
its genera and species, the most startling eccentricities of form
and color to be found among the /Mollusca/, if not among all marine
invertebrates.
Only those species which are actually abundant on the Atlantic and
Pacific shores of the United States, and which are likely to be found
by the untrained collector, are selected from the long list of forms
which belong to the three faunal regions involved.
ORDER =OPISTHOBRANCHIATA=
SUBORDER ~TECTIBRANCHIATA~
To avoid confusion, it is well to keep the systematic table in view:
Class Orders Suborders
=GASTEROPODA= =OPISTHOBRANCHIATA= =TECTIBRANCHIATA=
=NUDIBRANCHIATA=
=PROSOBRANCHIATA=
=PULMONATA=
[Illustration: PLATE LXV. Bulla occidentalis, enlarged. Bulla
nebulosa. Lottia gigantea, inside view. Fissurella alternata.]
As has already been remarked, shell-bearing opisthobranchs are not
abundantly represented in number of species in North American waters.
South of Cape Cod, in muddy bays and well-sheltered places, the
little shell, /Haminea solitaria/, may, however, often be found in
considerable numbers.
GENUS ^Haminea^
^H. solitaria.^ Like all external opisthobranch tests, the shell of
this species is thin and fragile, and would appear to be of small
importance to the creature it only partially protects. It is devoid
of spire, is shining bluish-white, sometimes brownish in color,
and is marked with revolving grooves, which, being cut across by
irregular growth-lines, give its surface a faintly decussated
appearance. The aperture is as long as the shell; the columella is
incurved. This little species is especially abundant in shallow
water about Vineyard Sound and in Peconic Bay.
[Illustration: /Haminea solitaria./]
GENUS ^Bulla^
^B. occidentalis.^ This species, which is found in the waters
of Florida on all sandy beaches, is a relative of the Northern
species just described. The shell is larger than that of /Haminea
solitaria/, and has a color-pattern of mottled or clouded
reddish-brown on a white foundation. There is no spire; there
is a pit in place of an apex; and the aperture extends the full
length of the shell. To the naked eye the surface appears smooth
and shining. It is an exceedingly variable species, occurring all
through the West Indies, but not north of Florida. The shores of
the keys along the west coast of Florida are often strewn with
these shells after storms. The animal, like all the shell-bearing
opisthobranchs, is very large as compared with the shell, and the
large mantle folds recurve upward, almost completely hiding the
shell from view. (Plate LXV.)
^B. nebulosa.^ This is one of the largest and finest species of
the family /Bullidæ/. It is found on the coast of California south
of San Francisco. The shell is much larger than that of either of
the Atlantic species mentioned above, but very closely resembles
them in outline, although it is built upon a more generous plan
and is more richly painted. It is brownish, mottled with white and
yellow patches, and is very suggestive of certain kinds of birds'
eggs. The animals have a greatly extended mantle which almost
completely envelops the shell. The foot is extremely large, with
great wing-like developments upon each side, called parapodia. Some
species of /Bulla/ have been seen to swim by means of the lazy
flapping of the parapodia. (Plate LXV.)
SEA-HARES
Closely allied to these outer-shell-bearing opisthobranchs is a large
and important group of tectibranchs, known as [pg351] "sea-hares."
They have a mantle so greatly developed that it actually covers the
shell, and its edges unite and fuse over the top. The shell, in
consequence, having almost ceased to be of use as a protection, has
degenerated into a mere horny plate, and has lost all resemblance to
the ordinary gasteropod shell. Having practically lost its protective
office and become a mere internal plate, it is quite probable that
it will in time wholly disappear. The gills of the sea-hares are
concealed under a flap of the mantle, their position being posterior
to the heart. The most conspicuous representative of this type of
tectibranchs in the United States is the following:
GENUS ^Aplysia^
^A. Wilcoxii.^ This species appears at times in vast numbers in
the waters of Florida, until the sea may truly be said to be
fairly alive with them. They swim lazily with a waving motion of
the parapodia. They disappear as mysteriously as they come, and
for months not a specimen will be seen. There is a variety of this
Floridian /Aplysia/ which occurs at Cape May, but no sea-hares are
to be found north of that point. The tropical Pacific furnishes an
astonishing wealth of these creatures, belonging to many genera
and species, and among them are some of the most beautiful of the
invertebrate animals.
SUBORDER ~NUDIBRANCHIATA~
The opisthobranchs are divided into two suborders, the second of
which is called /Nudibranchiata/. The name is well chosen and
very suggestive, for it means "naked or exposed gills"; but this
anatomical feature is only one of the peculiarities of this suborder.
The nudibranchs are commonly known as "sea-slugs"; for, like the
land-slugs, which are also true mollusks, they possess no shell at
all. That they at one time carried a shell is evident from the fact
that they are born with a rudimentary testaceous covering, which soon
afterward disappears.
A striking peculiarity of the nudibranchs is that the conventional
molluscan mantle is not usually apparent. Instead of seeing the usual
flaps or folds of the mantle which more or less encircle mollusks,
and which one seems to have a right to expect, [pg352] they are
entirely absent, and the body of the sea-slug assumes in consequence
a worm-like appearance. Other notable features of the nudibranchs
are the great number of tentacular processes that usually project
from the dorsal region of the animal, and, in many of the genera, an
entire absence of gills. When the gills are present, as is the case
in several of the nudibranch families, they are not placed along the
side of the animal, where one would naturally look for them, but are
arranged in the form of a rosette of plume-like processes situated in
the posterior dorsal region, or, in other words, on the animal's back.
Nudibranchs are commonly to be found all along the Atlantic and
Pacific coasts of North America, and more especially in the colder
waters north of Cape Cod. They are essentially littoral, and live
upon algæ in shallow water, upon eel-grass in sheltered places,
and in tide-pools, where there is more or less vegetation. They
crawl about the fronds of algæ, or swim, foot upward, with a gentle
and undulating motion, or, when caught between tides, may be seen
clinging to the under surface of rocks. Protective resemblances
have been so remarkably developed in the nudibranchs that they are
not always easy to discover; indeed, one may actually be looking at
one for some time without suspecting it to be other than some torn
fragment of seaweed.
* * * * *
There are many species belonging to several genera which frequent
the Maine and Massachusetts coasts, but those most likely to be
encountered are the following:
GENUS ^Dendronotus^
^D. arborescens.^ This species is about one inch long and variable
in color, but is usually reddish-brown or rose. There are no
tentacles, but in their place are two antler-like appendages
pointing forward and branched like a tree. All along the back are
two rows of these curiously branched processes, which give to
the animal the appearance of a plant. These /cerata/, or dorsal
papillæ, are delicately transparent, contractile, and richly
colored. The function of these papillæ is not fully known. As
the animal has no specialized breathing-organs, it is reasonable
to suppose that respiration is carried on through the outer skin
and perhaps all over the surface of these branched papillæ. The
liver, which [pg354] in most nudibranchs is extremely large and
completely surrounds the stomach, in /Dendronotus/ also extends
into these dorsal cerata, so that they may have some sort of
digestive function.
[Illustration: /Dendronotus arborescens./]
GENUS ^Æolis^
^Æ. papillosa.^ This is probably the commonest nudibranch upon
the North Atlantic coast, and it occurs as frequently in European
waters. It may readily be found clinging to stones, algæ, the
piling of wharves, eel-grass, etc., in bays or in rocky tide-pools.
The color is yellowish-gray to orange, with purplish or olive
spots. The cerata are very numerous, and cover the dorsal portion
of the animal save for an open space along the middle of the back.
There are two sets of tentacles, the two dorsal and the two simple
labial tentacles. The foot tapers behind to a sharp point, and is
truncate in front. As in /Dendronotus/, the liver is diffused into
the cerata, which in /Æolis/ are simple tubular processes without
the branching character of those of the former. Both species lay
their eggs in a gelatinous, bobbin-like cord, which is hung in
festoons over rocks or upon zoöphytes, or at times they wind the
cord in a coil upon the surfaces of stones. When the young first
hatch out, they are provided with a glassy nautiloid shell, a fact
which would indicate that at some remote period they were, like
most mollusks, provided with a shell. /Æolis/ swims in an inverted
position, and is at times exceedingly active. It is very variable
in coloring and in the number of cerata.
[Illustration: /Æolis papillosa./]
GENUS ^Doris^
^D. bilamellata.^ In /Doris/ we find a quite different type of
nudibranch from that presented by the two genera /Dendronotus/
and /Æolis/. In one there were numerous branched cerata or dorsal
papillæ, in the other many plain cerata, and in neither case were
there any specialized [pg355] breathing-organs. In /Doris/,
however, there are no dorsal papillæ at all, the back being covered
with calcareous spicules, which form a sort of secondary shell.
There is a circlet, or rosette, of retractile plume-like gills, or
branchiæ, placed in the middle of the back posteriorly. There are
various species of /Doris/, and of closely allied genera belonging
to the family /Dorididæ/, which are to be found along the New
England coast in tide-pools and among rocks. Those seen by the
writer never appeared to be other than very sluggish creatures, and
proved to be rather unsatisfactory tenants of the aquarium.
[Illustration: /Doris bilamellata./]
When discovered at low tide upon moist seaweed or stones, or when
disturbed in the water, nudibranchs often appear like small lumps of
jelly-like tissue, without a single attractive feature. It is only
when they are placed in a jar of sea-water and left unmolested for
a few moments that they unfold their beauties to the view. On the
Pacific coast there are numerous species of nudibranchs, occupying
corresponding shore stations and exhibiting the same degree of high
coloration. There are over a thousand species of sea-slugs scattered
about the various seas. They find their greatest development in
tropical waters, and for brilliancy of tint and variation in form
are unequaled by any other class of animal life, save perhaps the
butterflies.
ORDER =PROSOBRANCHIATA=
To this exceedingly large and comprehensive order of gasteropods
belong the great majority of univalve mollusks. In point of number of
genera and species and in abundance of individuals, the prosobranchs
probably exceed all the other orders of mollusks combined. They may
be called the typical gasteropods, the shell being univalvular and
generally spiral, and the animal exhibiting the singular torsion
of the visceral mass which has placed the breathing-organs forward
of the heart. They are provided with an operculum (placed upon the
posterior dorsal portion of the foot), which is used to close the
aperture of the shell against [pg356] enemies when the animal
retires within its fortress. They are all marine except a very
few families, which, supposedly of marine derivation, have become
terrestrial in habit.
The prosobranchs are further subdivided into suborders according to
certain peculiarities of the heart and breathing-organs. There is
a group of these prosobranchs which gives evidence of an inferior
degree of that visceral torsion which is always found in the
gasteropods. In this group, or suborder, the heart has two auricles,
and there is a pair of gills instead of only a single one. Other
internal organs are paired just as they were represented to be in
the schematic mollusk. This group also seems to show its primitive
character in the want of a proboscis and a siphon, or, in some
families, by having the ventricle of the heart traversed by the
intestinal canal, just as in the lower class of mollusks, which
includes the clams and oysters. For the most part the shells of this
group are not typically spiral, but are patelliform, shield-like
coverings, with only a suggestion of a spiral form at the very tip
of the apex. This group of primitive prosobranchs is included in the
following suborder:
SUBORDER ~DIATOCARDIA~
This suborder is named from the presence of two auricles in the heart.
FAMILY ~ACMÆIDÆ~
The first family to be noted is the /Acmæidæ/. Its principal genus,
/Acmæa/, is well represented on both the east and the west coast of
the United States.
GENUS ^Acmæa^
^A. testudinalis.^ This species is found in vast numbers all along
the New England shore, clinging to the rocks between tides. They
are usually called limpets. The shell is solid, conical, with an
oval outline, and with no trace of a spiral form in the adult. When
the shell is thoroughly cleaned, it generally presents a mottled
coloration of pale green, brown, and white. Inside it is white
and nacreous, with a large brown [pg357] area under the apex.
The animal has a powerful foot, by means of which it is enabled
to cling to a rock with great strength. In order to dislodge it,
it is necessary to approach carefully, and, taking the creature
unawares, quickly slip a broad knife-blade under the foot,
otherwise it is quite impossible to tear it from its resting-place.
/A. testudinalis/ lives entirely between tides, and ranges from the
northernmost waters to New York. It feeds upon algæ and is a very
sluggish animal. It has been said to leave its resting-place and
wander about in search of food, returning to its original and usual
spot when the tide begins to ebb. (Page 343.)
^A. testudinalis,^ variety ^alveus.^ This variety is smaller, more
fragile, and oblong. It lives upon eel-grass, its oblong shell
being adapted to the narrow leaves of the grass. The coloration is
brighter--reddish-brown spots on a white surface. It is exceedingly
common on the New England coast.
[Illustration: /Acmæa testudinalis/, var. /alveus./]
^A. mitra.^ On the Pacific coast there are a number of acmæas. /A.
mitra/ is often found dead upon the beaches. It is conical in form
and pure creamy-white in color. It looks very much like a clown's
pointed cap.
^A. patina.^ This is also very abundant in California. Outside it
is dark in color and is often incrusted with mineral deposits.
Within there is a dark ring around the edge, then a zone of
bluish-white, and a patch of brown just beneath the apex.
GENUS ^Lottia^
^L. gigantea.^ This is the largest of the California limpets.
Specimens three inches long have been found. The outer surface
of the shell is rough and brownish in color. The apex is near
one end. Within it is almost black, shining, lustrous, with a
horseshoe-shaped muscle-scar under the apex. The color is bluish
and brown. (Plate LXV.)
FAMILY ~FISSURELLIDÆ~
This is an extensive family, including several genera and a number
of species, commonly known as keyhole-limpets. In the general form
of the shell they closely resemble the true limpets, the /Acmæidæ/,
except that they have a hole, or rather a slit, in the shell just
back of the apex. Often this slit is so long that it has entirely
removed the apex of the conical shell.
GENUS ^Fissurella^
^F. alternata^, ^F. barbadensis^. These species occur upon the
southern shores of the United States, the latter, however, being
confined to [pg358] extreme southern Florida. /F. barbadensis/
has heavy longitudinal ribs and is light green within, with a rosy
circle about the apical perforation. (Plate LXV.)
GENUS ^Glyphis^
^G. aspersa.^ One of the numerous species belonging to this family
which are found on the west coast of the United States. It is about
one and a half inches long, grayish-white, with sharply raised
longitudinal ridges, slightly rayed, crossed by revolving ribs,
which give to the outer surface a decussated appearance. The apex
is forward of the center, and is entirely replaced by a round
perforation. The edge is wrinkled, and within smooth and white.
GENUS ^Lucapina^
^L. crenulata.^ The largest of the keyhole-limpets; the shell
is often four inches in length, while the animal, with its huge
yellowish foot and dark-colored mantle, which is thrown back,
almost concealing the shell upon its back, is much larger. The
apical perforation is very large. Shell flattened, with radiating,
rounded, crowded ribs; brownish-white in color; edge crenulated;
within pure white. It is found at Monterey, but live specimens are
not very frequently seen near the beach. (Plate LXVI.)
FAMILY ~HALIOTIDÆ~
GENUS ^Haliotis^
This family is closely allied to the /Fissurellidæ/. The species are
known on the Pacific coast by the name of abalone shells, and in
England and the Channel Islands as ormers or sea-ears. There are no
species of /Haliotis/ on the east coast of the United States, but
one has recently been dredged at a considerable depth in West Indian
waters. This family, with the last, possesses the striking anatomical
feature of having the heart traversed by the digestive tract. It
also has two gills, the degree of torsion in the visceral mass not
being sufficient to have crowded out and destroyed the original
right gill. The shell is spiral, but is so greatly flattened, and
the body-whorl is so greatly extended, that the shell quite loses
the spiral appearance. Along the dorsal side of the shell is a row
of holes, through which project numerous tentacular processes from
the mantle. The outer surface of the shell, before it is polished,
is usually rough and unattractive, but within, when the [pg359]
animal is removed, it displays a most beautiful and highly colored
nacreous surface. The spot near the center of the inner shell
surface where the muscles of the foot were attached is usually
most brilliantly colored. The shell of the abalone is susceptible
of taking a very high degree of polish, and is extensively used in
commerce for colored mother-of-pearl and for inlaid work. /Haliotis/
is a vegetable-feeding genus. They cling with great tenacity to rocks
about low tide, and it requires skill to remove them without breaking
the shell. There are several species in California. The Chinese use
the abalone for food, and have waged a persistent war upon the family
along the Pacific coast until the specimens are not nearly as common
as formerly. They are also eaten in France and in Japan.
^H. splendens.^ The largest and perhaps the most attractive in
appearance of the Californian species. Speaking of this beautiful
shell, Professor Keep says: "Within, a whole rainbow is condensed
in one of the magnificent shells, though the shades of green are
most conspicuous. The coloring in the center is particularly fine,
resembling a peacock's tail. There are about six open holes near
one side of the shell, and its length is about the same number of
inches." (Plate LXVI.)
^H. rufescens.^ A large abalone, which sometimes attains a length
of eight or nine inches. It is red in color, with three open holes
in the body-whorl. The outer portion of the shell is usually
incrusted with mineral deposit and overgrown with vegetation.
^H. cracherodii.^ Very dark green without, with five to nine holes;
length from one to six inches; spire exceedingly short. Common on
the Californian coast in crevices of rocks at low tide.
FAMILY ~TROCHIDÆ~
This is one of the largest and most interesting families of the
/Mollusca/. It contains many apparently widely separated genera
and a host of species, which for the most part are littoral, the
majority actually living between tide-marks. The typical trochid
shell is top-shaped or pyramidal, having a broad base and many
closely wound flat whorls terminating in a sharp apex. All the
trochids are nacreous within the aperture--a character which is
constant throughout the family. The animal has but one gill (the
left), a short snout, and often frontal lobes on the head. The
edge of the mantle or the epipodial line of the foot is usually
ornamented with from three to five cirri. The tentacles are [pg360]
long and slender, with short peduncles for the eyes; the operculum
is corneous, with a central nucleus. The animals are herbivorous,
feeding upon algæ. The trochids are essentially tropical shells, and
the most of the genera which are comprised in the family are only to
be encountered in the warm waters of the Pacific and Indian oceans.
Some forms are very beautiful, and frequently are used as mantel
ornaments, and the shells of one little species, which is opalescent
in its coloring, are still extensively gathered in the East Indies,
to be polished and strung like pearls in necklaces.
GENUS ^Margarita^
On the Atlantic coast north of Cape Cod the trochids are represented
by the genus /Margarita/, with five or six species. The shells are
small, thin, and globosely depressed, with smooth or transversely
striated whorls. The aperture is nearly circular, with a simple lip.
[Illustration: /Margarita cinerea./]
^M. cinerea.^ This species has several prominent revolving ridges
upon the upper side of the whorls, with finer ones on the base.
Very fine growth-lines cover the entire shell. It ranges all along
the coast north of Cape Cod, but is not usually found between
tides. The writer has dredged many specimens in shallow water at
Eastport and Bar Harbor.
[Illustration: /Margarita helicina./]
^M. helicina.^ A thinner and more globose species than the last,
with a translucent, shining, smooth surface of a yellowish or olive
color. /M. helicina/ is very fond of the leaves of /Laminaria/, and
is often found clinging to them when storms have torn these great
algæ from the bottom and cast them upon the shore. At Bar Harbor
they are common upon the eel-grass in Rodicks Weir. This species
can generally be distinguished by its iridescent, metallic luster.
[Illustration: /Margarita undulata./]
^M. undulata.^ A commoner species, perhaps, than either of the
preceding, sometimes found on the rocks of sheltered coves at
exceptionally low tides. Judging from the number often to be found
in the stomachs of fishes, they must be considered excellent food
by the cod and its allied species which thrive along the Maine
coast. No doubt millions of /M. undulata/ are yearly destroyed
in this way. The shell is depressed, with four rounded whorls, a
flattish base, and a large umbilical opening. In color it varies
from rose-red to brown. The surface is decorated with numerous
revolving raised lines placed at uniform distances. Just below
the suture the body-whorl is somewhat undulated with short folds.
Height three tenths of an inch, base four tenths of an inch.
[pg361]
GENUS ^Calliostoma^
This genus is better represented on the Pacific coast of the United
States than upon the Atlantic. It comprises a series of marvelously
beautiful shells, often exquisitely colored. They are regularly
conical or pyramidal in shape, with flattened bases. One never tires
in the search for calliostomas. They are not common enough to cause
one to lose interest, and whenever a good specimen is captured it
seems as though one had found some gem cut and polished by nature's
skilful hand and prepared for a place of honor in the cabinet.
^C. occidentale.^ The only species of this genus found on the
northeast coast. It is larger than /Margarita cinerea/, but
somewhat resembles it. It is shining nacreous within and without,
and has strong revolving ribs, the upper one on each whorl often
being broken into a circular row of white dots. The lip is
crenulated. This very pretty species is not likely to be met with
upon the shore, but may be dredged in shallow water on gravelly
bottoms along the Maine coast. Height about one half of an inch.
[Illustration: /Calliostoma occidentale./]
^C. jujubinum.^ A species which occurs in the waters of Florida.
Its form is almost that of a true pyramid. The sutures can scarcely
be distinguished. The shell is marked by brown and purplish-red
spots on a white background, and has numerous revolving ribs broken
into rows of white dots like little glazed beads. The umbilicus
is funnel-shaped and wide. In Tampa Bay this shell is frequently
found on sponges, and may sometimes be gathered on the beach after
storms, even as far north as Hatteras. There are as many as twenty
species of this genus in American Atlantic waters, but they are
either rare or have deep-water stations.
^C. annulatum.^ A remarkable species of /Calliostoma/, found in
California. The whorls are adorned with revolving rows of raised
dots, and the sutures are frescoed in exquisite purple. This
beautiful species lives upon seaweed, and on pleasant days comes
to the surface of the water. It can then be collected from a boat
by drawing in quantities of seaweed. It is very unlikely that the
collector will ever find a specimen upon the beach, for the shell
is too fragile to withstand the rough handling of the waves. Length
one inch.
^C. canaliculatum.^ The largest member of this genus to be found
in American waters. It resembles the last species in form, but
lacks the purple sutures. The revolving ribs are very numerous and
prominent. There is no umbilicus. Length one to one and a half
inches. Found on the Pacific coast.
^C. costatum.^ A heavier shell than the preceding, with somewhat
more rounded whorls. Reddish-brown in color; numerous revolving
ridges; no umbilicus; about three quarters of an inch in length. It
is found in rocky places, and sometimes in considerable numbers,
just at low-tide mark. Found in California. [pg362]
GENUS ^Chlorostoma^
This genus is represented in California by several species, the
commonest of which is probably /C. funebrale/--a doleful name, no
doubt given on account of the jet-black color.
^C. funebrale.^ The shell is thick and strong, like most
between-tide species, which are constantly exposed to the buffeting
of the waves. It is to be found in countless thousands upon rocks
exposed at low tide, and may be gathered at any time except full
flood-tide, like the littorinas and purpuras of the east coast.
Within the aperture the surface is nacreous and greenish in tint;
the last whorl is drawn in, like gathers, at the suture. The
umbilicus is closed. There is a white nodule at the base of the lip
of the columella. Length one half of an inch to one inch. (Plate
LXVII.)
^C. brunneum.^ This species is brown, as its name would indicate,
and there is greenish nacre within the aperture. It lives upon
kelp, or upon rocks at very low tide. Length about an inch. (Plate
LXVI.)
GENUS ^Trochiscus^
^T. norrisi.^ A flattened shell with rounded, dome-like spire and
obtuse apex; a fairly common species. It has a wide, deep umbilicus
and a sharp lip. It is of a rich brown color, with dark chestnut
about the umbilical region and greenish-white within the umbilicus;
there is a band of dark olive-green about the inner margin of the
lip. The shell is about two inches in diameter and of a somewhat
greater height. The operculum is multispiral, with a central
nucleus, and is shaggy and rough. Like all other trochids, it feeds
upon algæ. It is often found upon the beach after violent storms.
Found in California, south of San Francisco. (Plate LXVI.)
GENUS ^Livona^
^L. pica.^ This large and interesting trochid is a West Indian
species which sometimes occurs in Florida. It lives about coral
reefs and rocky shores, attaching itself in vast numbers to the
rocks. This shell is a favorite refuge for large hermit-crabs. It
is a curious sight in certain of the West India islands to see a
/Livona pica/ shell scrambling up a tree, looking very much out of
place upon the back of some terrestrially inclined crustacean. When
well cleaned it is a beautiful shell, with black wavy lines over a
greenish-white nacreous foundation. The animal has a row of waving
cirri upon each side of the mantle, and long, slender tentacles.
It is largely used as an article of diet in the West Indies and
Central America. (Plate LXVII.)
FAMILY ~TURBINIDÆ~
This family is very closely allied to the trochids, the most striking
difference being in the operculum, which in the /Turbinidæ/ is
calcareous and usually smooth and very convex on the outside.
[pg363] The shells are nacreous within, and the animal, with its rows
of waving cirri upon each side, is very suggestive of /Trochus/. They
are, for the most part, shallow-water or littoral forms.
[Illustration: PLATE LXVI. 1, Lucapina crenulata. 3, Chlorostoma
brunneum. 2, Haliotis splendens. 4, Neritina reclivata, enlarged. 5,
Trochiscus norrisi.]
[Illustration: PLATE LXVII. Livona pica, reduced. Turbo castaneus,
enlarged. Nerita peleronta. Chlorostoma funebrale. Nerita tessellata.]
GENUS ^Turbo^
^T. castaneus.^ This species has a range as far north as Cape
Hatteras, and is especially abundant at Tampa, on the west coast of
Florida. The peculiar operculum is sufficient to identify the genus
at once. One variety of this species has a crenulated shoulder upon
the body-whorl and is referred to as ^Turbo crenulatus^, A series
of intermediate forms establishes the specific identity of the two
varieties. (Plate LXVII.)
FAMILY ~NERITIDÆ~
The /Neritidæ/ are strictly littoral forms, almost entirely confined
to the warmer waters of tropical seas. The animal preserves the usual
diatocardian features--has a short snout and long tentacles. Unlike
the trochids and /Turbo/, it has no cirri along the epipodial margin.
The shells are peculiarly shaped, the spire being greatly flattened
and scarcely noticeable on account of the unduly large development
of the body-whorl; thus the shell takes on a decidedly patelliform
appearance. It is without an umbilicus. In the principal genus
/Nerita/ the outer lip is sharp on the edge, but greatly thickened
just within.
GENUS ^Nerita^
^N. peleronta.^ This shell has two teeth on the wide, flat
columellar lip, and about them is a blotch of red, suggesting
blood. The common name of this shell, "bleeding-tooth," is very
appropriate. Found in southern Florida. (Plate LXVII.)
^N. tessellata^ is a smaller species, with heavy revolving ribs,
and is further decorated with transverse oblique black lines. The
operculum is calcareous. Both of these species have the habit of
absorbing the entire inner portion of their shells. They belong
to the West Indian fauna, and occur in great numbers on rocky or
coral shore stations. They also may be found on the east Florida
coast, well to the south. A third species, ^N. versicolor^, often
occurs, associated with the other two. It is somewhat smaller than
the others and can easily be distinguished by the four teeth on
the columellar lip, the edge of the columella being convex. (Plate
LXVII.)
GENUS ^Neritina^
The genus /Neritina/ is very closely allied to the last, having quite
the same form of shell; but it is usually more globular and [pg364]
variously ornamented by bright spots or zigzag lines of coloration.
The neritinas have acquired the habit of ascending rivers, until
they have become almost wholly a brackish- or fresh-water genus.
The metropolis of this genus is in the South Sea Islands, where it
attains a wonderful development in the clear running streams of the
volcanic islands.
^N. reclivata.^ A very pretty olive-green species, with very
fine, wavy, longitudinal lines of coloration, found in almost
all Floridian streams above the action of the tide. The nacre of
the columella and within the aperture is bluish-green, and the
operculum is rich olive. (Plate LXVI.)
^N. viridis.^ A small, intensely green species, which, unlike most
neritinas, is strictly marine. It belongs to the great West Indian
faunal province, but is occasionally found upon the shores of
Florida and of Texas.
The prosobranch gasteropods thus far considered (belonging to the
suborder /Diatocardia/) all show by their anatomical structure that
the process of visceral torsion, though carried very far within them,
has not been complete enough to crowd out and finally destroy one
of each of the paired organs. Both auricles of the heart (with some
exceptions) were left intact.
SUBORDER ~MONOTOCARDIA~
In this the second suborder of the prosobranch gasteropods, the
twisting visceral process has been carried to the extreme. The
heart has but one auricle. There is but one gill (on the left
side), and this is attached to the inner side of the mantle flap.
In other respects the presence of certain specialized organs would
probably indicate that gasteropods belonging to this suborder are
a step higher in the scale of life than those which belong to the
/Diatocardia/.
FAMILY ~JANTHINIDÆ~
GENUS ^Janthina^
The genus /Janthina/ has an exceedingly thin and semi-transparent
shell, deep violet in color on the base and lighter blue on the
spire. An interesting feature of this genus is the mode of depositing
the ova. The female exudes from a gland in the foot a [pg365]
glutinous secretion which hardens in water, and, being filled with
air-bubbles, constitutes a float. On the under side of this are
deposited the eggs in rows of little capsules. While attached to this
float it is impossible for a /Janthina/ to sink, and hence it is that
so many of these creatures are sacrificed in onshore gales of wind.
^J. fragilis.^ The shell of this species is so brittle and fragile
that it is very clearly not adapted to a life near shore. It is in
reality a pelagic species which is occasionally blown ashore during
easterly gales along the Atlantic coast of the United States. Vast
numbers of these pretty creatures are sometimes encountered far out
at sea, floating quietly on the surface. When storms drive them
upon the beach, they become utterly helpless; since their foot
is not adapted for crawling upon the sand, they soon perish, and
their brittle shells are demolished by the surf. In Florida the
beaches are sometimes fairly lined with /Janthina/ shells, which
make a band of purple along the high-tide mark as far as the eye
can reach; then it may be years before they again appear. (Plate
LXVIII.)
[Illustration: /Janthina fragilis/: /FL/, float; /O/, ova; /Pr/,
proboscis; /Br/, branchiæ; /F/, foot.]
FAMILY ~SCALIDÆ~
GENUS ^Scala^
The shells of /Scala/ have such a peculiar scheme of decoration
that once seen they can never be mistaken. They are generally
pure white, with well-rounded whorls, all of which are crossed
at even distances by greatly elevated and smooth ribs. Each rib
represents a rest-period, when the creature thickened the rim
of the shell-aperture. The aperture is generally round, with a
continuous lip. The animal has a retractile proboscis and long,
slender tentacles with eyes at the outer bases, and is a predaceous,
[pg366] carnivorous creature. Some Asiatic species of this genus,
remarkable for their beauty and rarity, have been greatly prized by
collectors. A single specimen of the now well-known ^S. pretiosa^ of
China has been sold for two hundred dollars--a fancy price, indeed,
for a shell which can now be bought for a dollar! There are over
fifty species of /Scala/ on the Atlantic coast, but most of them are
either rare or belong to a zone of deeper water; there are, however,
four or five species which are exceedingly common.
^S. lineata.^ A species which ranges from Hatteras to New England.
It has about eight whorls, and is slightly brownish in color. The
ribs are robust and not greatly elevated; there are from seventeen
to nineteen on the body-whorl. The shell is sometimes painted with
a few revolving brownish lines.
[Illustration: /Scala lineata./]
^S. multistriata.^ The transverse ribs are much smaller but very
numerous; the small spaces between them are marked with many fine
revolving lines. Found from Cape Cod southward.
[Illustration: /Scala multistriata./]
^S. groenlandica.^ Essentially an arctic species, which has found
its way down to the New England coast. It is readily distinguished
by the flattened, coarsely rounded, revolving ribs, which follow
the volutions of the spire. Over them are the usual transverse
heavy ribs peculiar to this genus.
[Illustration: /Scala groenlandica./]
^S. angulata.^ The whorls touch one another only by the ribs,
of which there are nine to each volution. This species has a
remarkably wide range, occurring from Cape Cod to southern Florida.
(Plate LXVIII.)
These four species vary from one half of an inch to one inch in
length. All of them are found on the beach after storms or may be
dredged in shallow water near the shore.
FAMILY ~NATICIDÆ~
This large and interesting family is well represented in the Atlantic
waters of the United States, but its more beautiful members live
in the tropics. The New England and New Jersey species are dull in
color, but offer much of interest to the collector and student. The
foot is enormously large, and carries in front a great shield-like
fleshy process, which curves back over the head of the animal and
serves as a plow in pushing its way through the heavy wet sand of
the beach. When the creature is thus seen extended in the act of
crawling, one wonders how it is possible for it to withdraw so great
an amount of body into its shell; but if it is suddenly seized or
irritated, it will quickly [pg367] demonstrate its ability
to hide itself completely within its house and to close the door
very effectively by means of its operculum. The eyes seem to be
wanting, or they are concealed under the skin of the head. The shell
is usually quite large, with a depressed spire and well-rounded
whorls--especially the body-whorl, which appears to be greatly
swollen. The umbilicus is usually open and moderately large, the lip
simple.
[Illustration: PLATE LXVIII. Janthina fragilis. Sigaretus
perspectivus, upper side. Scala angulata, enlarged. Sigaretus
perspectivus, under side. Natica canrena. Crepidula aculeata.]
GENUS ^Polynices^ (^Lunatia^, ^Natica^).
^P. heros^ (generally referred to as ^Lunatia^ or ^Natica
heros^). One of the commonest large shells and one of the most
characteristic species of the New England and New Jersey littoral
fauna. It is exceedingly common along the Long Island shore, where
it may be found on the open beach, in pools with a sandy bottom
left by the receding tide. It is usually partially and frequently
wholly buried in the sand. The umbilicus is open and large, the
operculum corneous, and the shell heavy and ashy-white to brownish,
with (when young) a yellowish epidermis. Its length is from two to
four inches. It has no ornamentation whatever. /P. heros/ is a most
voracious creature and spends its time in hunting for flesh--either
alive or dead--to devour. It feeds upon dead fish, or upon other
mollusks whose shell it is able to pierce by means of its radula,
making a little round hole through which it sucks out the flesh
from within. The curious egg-cases of this species have already
been referred to. (See Plate I.) It glues together particles of
sand into the form of a basin with the bottom knocked out and
broken on one side. In the gelatinous substance of this basin it
deposits its eggs in a regular order. These hatch out in midsummer.
Egg-cases of this kind can always be found wherever /Polynices/
lives. For a long time naturalists were greatly puzzled by these
curious things, and their blunders are recorded in earlier works,
where these egg-cases have been elaborately described as living
animals belonging to various invertebrate orders. The largest and
best specimens of /P. heros/ are to be found south of Cape Cod.
(Page 343.)
^P. triseriata.^ A small shell of exactly the same shape as /P.
heros/, but decorated with three revolving series of bluish or
chestnut spots. It is pretty well determined that this so-called
species is only the young of /P. heros/. It is very abundant all
along the coast.
[Illustration: /Polynices triseriata/, young; /Polynices
triseriata/, older specimen.]
^P. duplicata.^ This is even more abundant than /P. heros/. It
has a flatter shell, with an obtuse apex and dome-like spire.
The umbilicus is partly or wholly closed by a thick, callous,
shelly process thrown off from the columellar lip, and is chestnut
in color. The surface of the shell is smooth, often polished,
ashy-white below and light chestnut above. The operculum is
corneous. The length of the shell varies in different localities
from one half of an inch to about two [pg368] inches; the breadth
slightly exceeds the length. /P. duplicata/ has a very extensive
range, from the Gulf of Mexico to Newfoundland. The largest and
finest specimens are found in the vicinity of New York and at
Hatteras. In Florida they are usually smaller. This species cannot
be confounded with any other upon our coast on account of the heavy
callous deposit over the umbilicus. Its habits are similar to those
of /P. heros/.
[Illustration: /Polynices duplicata./]
[Illustration: /Natica clausa./]
^Natica clausa.^ A Northern species which is fairly abundant on the
Maine coast. It has a shelly operculum, and the umbilicus is neatly
closed by a pure white, shining callosity. Its length is about one
half an inch, its color livid white to light brown, white within.
The calcareous operculum at once determines this shell. This
species (as well as the following one) has retained the generic
name of /Natica/ because of the shelly operculum, as explained
below.
^N. canrena.^ One of the handsomer species of /Natica/, which
occurs in Florida. Sometimes it is seen as far north as Hatteras.
The shell is white, with bars of light chestnut circling the
whorls, and with zigzag lines of darker purplish hue crossing them.
The base of the shell is white. The aperture is large and flaring,
and is purplish within. The umbilicus is partially closed by an
entering callous plug. The operculum is calcareous, with eccentric,
deeply cut grooves. Found in sandy stations just about low-water
mark. (Plate LXVIII.)
^P. lewisii.^ A species which occurs on the west coast of the
United States. It resembles its New England relative /P. heros/
very closely. /P. lewisii/ is a Northern species, and does not
range south of Oregon. Professor Keep mentions one specimen of the
size of a six-inch globe, but such dimensions are very unusual. A
good-sized specimen need not exceed four inches in height.
^P. recluziana.^ A species well known on the southern coast of
California. The umbilicus is closed by a thick, highly polished
white callosity, and the general shape of the shell is strongly
suggestive of the common east-coast species /P. duplicata/. A large
specimen is about two inches in diameter.
[There has been much confusion in the generic nomenclature of
this family. The old name /Natica/ once did service for all the
species; then the names /Lunatia/, /Neverita/, and /Polynices/ were
applied to certain special forms; but it has been wisely proposed
by conchologists to use the name /Natica/ for those forms having
a shelly operculum, and to adopt the name /Polynices/ for those
having a corneous operculum. /Lunatia/ and /Neverita/ have become
subgenera of /Polynices/.] [pg369]
GENUS ^Sigaretus^
A very interesting species of this genus occurs on the east coast,
sparingly in New Jersey, but very abundantly south of Hatteras.
/Sigaretus/ is a modified /Natica/. The spire is flattened and
minute. The body-whorl, being greatly expanded, gives a wide,
flaring, oblique aperture. The shell is ear-shaped and white, with
fine revolving lines. There is no umbilicus. The operculum is small
and rudimentary. The animal is large, with an enormous foot and
greatly developed propodium, and is a sand-dweller. The common
east-coast form is ^S. perspectivus^. (Plate LXVIII.)
FAMILY ~CAPULIDÆ~
GENUS ^Crepidula^
[Illustration: /Crepidula fornicata./]
^C. fornicata.^ All collectors of mollusks on the Atlantic coast
sooner or later encounter this exceedingly common species adhering
to oyster-shells or scallops, and often to other large live shells.
It is cast upon the beach along the entire length of the Atlantic
coast. The shell is obliquely oval, dull whitish in color, and
either smooth or rough or even ribbed, according to the nature
of the surface to which it clings. The spire is almost entirely
suppressed, the little inconspicuous apex being turned to one
side and closely pressed down against the body of the shell. When
looking into the interior of the shell one is reminded of a boat,
for the upper portion of the aperture is covered by a horizontal
shelly partition, called the diaphragm, a space being left below
which would correspond to the forecastle of a ship. The "stern" is
round, and the "bow" is suggestively pointed.
[Illustration: /Crepidula plana./]
^C. plana.^ In this species the shell is white and flat, or
slightly concave. Although it is pointed at the "bow" end and
square at the "stern," the nautical resemblance stops there,
because, being flat, there are no swelling sides and bow. The
diaphragm is about one half the length of the shell, convex,
shining, white, and translucent. /C. plana/ lives generally within
the aperture of large dead shells. It has a wide range, from Maine
to Florida.
^C. aculeata.^ A common Floridian species which has a remarkable
range, being found in nearly all the tropical and semi-tropical
waters of the world. It is smaller than /C. fornicata/, is
reddish-brown with a white diaphragm, and has several radiating
ribs on the back, which are inclined to be nodulous. (Plate
LXVIII.) [pg370]
On the Pacific coast the following species are found: ^C. adunca^,
with a strongly recurved apex, and less than one inch long; ^C.
rugosa^, rough, brown, with the apex lying on the edge of the
shell, and one inch in length; ^C. navicelloides^, almost identical
with the east-coast /C. plana/. /C. aculeata/ also appears.
GENUS ^Crucibulum^
/Crucibulum/ has a peculiar rounded shield-like form, with a very
small apex on one side. Within there is a cup-shaped appendage
attached by one side to the inner margin of the shell. This latter
feature at once determines the genus.
[Illustration: /Crucibulum striatum/, from above.]
[Illustration: /Crucibulum striatum/, from below.]
^C. striatum.^ This species has radiating riblets, cut by circular
lines of growth. No dimension would quite reach an inch. It is a
common shell on the Atlantic coast, and will be found adhering
to stones and other shells, but it is not, strictly speaking, a
littoral species.
^C. spinosum.^ The shell exhibits a strong tendency to spinous
processes on its back. Found along the southern part of the
California shore.
FAMILY ~LITTORINIDÆ~
GENUS ^Littorina^
/Littorina/ is probably the most characteristic genus of Northern
littoral regions. Together with some of its allied genera it is also,
probably, equally characteristic of various tropical littoral faunas
all over the world. The family comprises strictly between-the-tides
genera and species. Indeed, it is suspected that some species of
/Littorina/ are making very fair progress toward a terrestrial
condition, for they actually live above high-tide mark,--even in
the branches of overhanging trees,--and must certainly pass days at
a time out of their natural element. That such a transformation is
possible need not for a moment be doubted, for there are many land
mollusks to-day that give abundant evidence of having been at some
past time aquatic or marine species. These changes in nature are
constantly going on, and the gradual substitution of a lung for a
gill is no very startling metamorphosis. [pg371]
The littorinas fairly swarm in favorable localities upon all shore
stations. In Maine and Massachusetts the bold, rocky coast furnishes
a home for several species. Often the rocks at low tide are black
with them; the algæ that cling in wet masses to the exposed rocks
are alive with them. One cannot walk about in such localities
without crushing hundreds of specimens. Sometimes they will be found
clinging in clusters upon the piling of old wharves, or crawling
about the bottom at or about the low-tide mark. The best specimens
of /Littorina/ are found in stations where they are bathed twice a
day by pure, uncontaminated sea-water; those living near the mouths
of streams, or where the water is brackish or impure, are usually
small and degenerate. They are vegetable feeders, and have received
the common name of "periwinkles." In Great Britain they are used
among the poorer classes for food. The animal has a short, broad
muzzle, and eyes at the outer bases of the tentacles. The foot is
longitudinally grooved, and there is a rudimentary siphonal fold in
the mantle. The shells are turbinated, usually heavy, few-whorled,
and with a round aperture.
[Illustration: /Littorina litorea./]
^L. litorea.^ This is supposed to be an importation from the Old
World--to have come over by way of Iceland and Greenland, and
then to have migrated down the Labrador coast. For many years
Cape Cod formed a barrier to its advance, but now the species is
abundant at Newport, and is reported at New York. It occurs on
the Maine coast in astonishing numbers, living in vast colonies
on the rocks exposed at low tide. The shell is thick, imperforate
(no umbilicus), and usually has flat, spiral ribs. The columella
is broad and white; the lip thin and black. The general color
varies from black to olive or to dingy gray--sometimes reddish.
The operculum is corneous, with the nucleus near the outer edge.
Despite the variableness of this very common shell (the variations
being chiefly in the height of the spire), it has certain
unmistakable characteristics which, once seen, will enable the
collector to determine it at once.
[Illustration: /Littorina rudis./]
^L. rudis.^ A smaller species than the last. It is strong
and coarse, with revolving grooves and ribs, or smooth, with
interrupted whitish bands and spots. A very common variety of
this species is much smaller than the typical form, being about
one sixth to one fourth of an inch long, smooth, with white and
yellowish spots on olive. It clings to the rocks near high-tide
mark, and is usually found attached to its resting-place by a
bit of hardened mucus. While the typical /L. rudis/ is heavy and
banded, with a moderately high spire and no color, this [pg372]
variety is thinner, with a higher spire and smoother surface,
and has a color-scheme of decoration. It would be impossible
to disconnect the two, for a large series of specimens will
demonstrate beyond doubt, through every minute degree of variation,
that the two forms belong to one and the same species. The station
of /L. rudis/ is much the same as that of /L. litorea/, but the
smaller colored variety of /rudis/ prefers quieter and more
sheltered spots. It is occasionally found on reeds and grasses,
on the piling of wharves, on large boulders above the line of
algæ, and on algæ. It has been the writer's experience not to find
/rudis/ and /litorea/ associated together.
[Illustration: /Littorina palliata./]
^L. palliata.^ A low-spired, globular shell with a large, tumid,
smooth body-whorl. The columella is flattened, curved below, and
imperforate. The color is exceedingly variable, but is usually
bright, shining olive, and this is especially the case with those
individuals that find their station in the dense masses of algæ
that form so conspicuous a feature of a low-tide scene on a rocky
New England coast. From pure olive-green to yellow or bright red,
with revolving black bands, seems a long chromatic leap, but it
is not too great for the Designer of these pretty little globular
shells. As a rule, the color of the shell simulates pretty closely
that of the seaweed upon which it lives, and inexperienced eyes
may easily overlook hundreds of specimens, all within close reach.
The banded varieties are less common. The head of the animal is
somewhat orange in color, the foot slate. The distinguishing
features of this species are the smooth, globular shell, the low
spire, the broadly flattened columella, and the orange-colored head
of the animal. It is found on /Fucus/ between tides, and often
associated with /L. rudis/, on the whole coast north of New Jersey.
The three preceding species are distinctly boreal in their range,
but their place is taken in southern Atlantic waters of the United
States by the following exceedingly common species:
[Illustration: /Littorina irrorata./]
^L. irrorata.^ A solid, robust shell, which attains a length of
about one inch. It is in many respects suggestive of /L. litorea/,
its Northern relative, but the spire is higher, with straighter
outlines, and the apex is acute. The surface is ornamented with
closely set, revolving ribs; the sutures are indistinct; the lip
is thin, though thickened just within; and the color is whitish to
pale cinereous greenish, sometimes spotted with broken brownish
lines. Within it is white, with a reddish tinge on the columella,
and brownish spots on the edge of the lip. This species ranges from
Maryland to Texas; its station is between tides.
^L. angulifera.^ As the last species resembles /L. litorea/, so
this other common Southern form resembles a greatly developed and
large /L. rudis/ of the smoother variety. It has a high-spired
shell, with an acute black tip. It has about six whorls, which are
variously decorated by wavy, oblique black lines and revolving
black lines broken into series of dots and larger spots near the
suture. The ground-color varies from yellow to purple. None of
the colors are very vivid. Within it is yellowish-white. This
species has a much thinner and more delicate shell than any of
the littorinas thus far considered. It is common on the piling of
wharves and in sheltered nooks everywhere [pg373] between tides.
It is essentially a tropical species of very wide and extended
range, but it does not occur north of St. Augustine. At Tampa it
has been found in vast numbers. (Plate LXIX.)
The California coast furnishes two more species of /Littorina/.
Their station is among the rocks between tides, and their habits
are quite the same as those of the east-coast species.
^L. scutulata.^ In this species the color is of various shades,
generally dark gray, mottled with white spots. A good-sized
specimen measures one half of an inch in length. The spire is high;
the columella flattened, broad; the shell is purplish-white within
the aperture. (Plate LXIX.)