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majority are fluviatile, one group (Neritodryas) actually occurring in
the Philippines on trees of some height, at a distance of a quarter of
a mile from any water. Navicella is a still further modified form of
Neritina, occurring only on wet rocks, branches, etc., in non-tidal
streams (Fig. 13).

Fig. 13.—Illustrating the development of the fresh-water genus

Navicella, through the brackish-water Neritina, from the marine
Nerita, with corresponding changes in the operculum. 1.
Nerita; 2, 3. Neritina; 4. Neritina, intermediate form; 5, 6.
Navicella.
The great family of the Melaniidae, which occurs in the rivers of
warm countries all over the world, and that of the Pleuroceridae,
which is confined to North America, are, in all probability, derived
from some form or forms of Cerithium. The origin of the Paludinidae,
Valvatidae, and Ampullariidae is more doubtful. Their migration from
the sea was probably of an early date, since the first traces of all
three appear in the lower Cretaceous, while Melaniidae are not
known until Tertiary times. Ampullaria, however, shows distinct signs
of relationship to Natica, while the affinities of Paludina and Valvata
cannot as yet be approximately affirmed.
 (2) Pulmonata.—Intermediate between the essentially fresh-water
and the essentially marine species come the group sometimes
known as Gehydrophila, consisting of the two families Auriculidae
and Otinidae. These may be regarded as Mollusca which, though
definitely removed from all marine species by the development of a
true lung or lung cavity in the place of a gill, have yet never become,
in respect of habitat, genuine fresh-water species. Like Potamides,
they haunt salt marshes, mangrove swamps, and the region about
high-water mark. In some cases (Otina, Melampus, Pedipes) they
live on rocks which are moistened, or even bathed by the spray, in
others (Cassidula, Auricula) they are immersed in some depth of
brackish water at high tide, in others again (Scarabus) they are more
definitely terrestrial, and live under dead leaves in woods at some
little distance from water. Indeed one genus of diminutive size
(Carychium) has completely abandoned the neighbourhood of the
sea, and inhabits swampy ground almost all over the world.
Fig. 14.—Examples of the Auriculidae: A,
Auricula Judae Lam., Borneo; B, Scarabus
Lessoni Blainv., E. Indies; C, Cassidula
mustelina Desh., N. Zealand; D, Melampus
castaneus Mühlf., S. Pacific; E, Pedipes
quadridens Pfr., Jamaica.
 Fig. 15.—An example of
Amphibola (avellana
Chem.), the only true
Pulmonate which
possesses an
operculum.
To this same section Gehydrophila have been assigned two
remarkable forms of air-breathing “limpet,” Siphonaria and Gadinia
(see page 151), and the aberrant Amphibola, a unique instance of a
true operculated pulmonate. Siphonaria possesses a pulmonary
cavity as well as a gill, while Gadinia and Amphibola are exclusively
air-breathing. Siphonaria lives on rocks at or above high-water mark,
Gadinia between tide marks, Amphibola (Fig. 15) in brackish water
at the estuaries of rivers, half buried in the sand. There can be little
doubt that all these are marine forms which are gradually becoming
accustomed to a terrestrial existence. In Gadinia and Amphibola the
process is so far complete that they have exchanged gills for a
pulmonary cavity, while in Siphonaria we have an intermediate stage
in which both organs exist together. A curious parallel to this is found
in the case of Ampullaria, which is furnished with two gills and a
pulmonary chamber, and breathes indifferently air and water. It is a
little remarkable that Siphonaria, which lives at a higher tide level
than Gadinia, should retain the gill, while Gadinia has lost it.
The ultimate affinities of the essentially fresh-water groups,
Limnaea, Physa, Chilina, cannot be precisely affirmed. The form of
shell in Latia, Gundlachia, and perhaps Ancylus, may suggest to
some a connexion with the Otinidae, and in Chilina, a similar
connexion with the Auriculidae. But, in a question of derivation,
similarities of shell alone are of little value. It is not a little
remarkable, for instance, that we should find a simple patelliform
shell in genera so completely distinct from one another in all
anatomical essentials as Ancylus, Patella, Siphonaria, Propilidium,
Hipponyx, Cocculina, and Umbrella.
Some recent authors, on grounds of general organisation, regard
the Limnaeidae and their allies as Opisthobranchs adapted to an
aerial life. It is held[24] that the Nudibranchiate Opisthobranchs have
given birth to the Pulmonata Stylommatophora or land snails, and
the Tectibranchiate Opisthobranchs to the Pulmonata
Basommatophora or fresh-water snails. Such a view seems at first
sight open to some objection from other views than those which deal
simply with anatomy. The Opisthobranchiata are not, to any marked
extent, littoral genera, nor do they specially haunt the mouths of
rivers. On the contrary, they inhabit, as a rule, only the very lowest
part of the littoral zone, and are seldom found, except where the
water is purely salt. In other cases, when the derivation of land or
fresh-water genera is fairly well established, intermediate forms
persist, which indicate, with more or less clearness, the lines along
which modification has proceeded. It has, however, recently been
shown that Siphonaria[25] and Gadinia,[26] which have, as has been
already mentioned, hitherto been classified as Pulmonata, are in
reality modified forms of Opisthobranchiata, which are in process of
adaptation to a life partly marine, partly on land. They may therefore
be regarded as supplying the link, hitherto missing, between the land
Pulmonata and the marine groups from one or other of which the
latter must have been derived. The general consensus of recent
opinion inclines towards accepting these views, some writers[27]
being content to regard the Pulmonata, as a whole, as derived from
the Tectibranchiate Opisthobranchs, while others[28] go further and
regard the Stylommatophora as derived directly from the
Basommatophora.

Origin of the Land Fauna

 Gasteropoda.—(1) Operculate. On a priori grounds, one might
predict a double origin for land operculates. Marine species might be
imagined to accustom themselves to a terrestrial existence, after a
period, more or less prolonged, of littoral probation. Or again, fresh-
water species, themselves ultimately derived from the sea, might
submit to a similar transformation, after a preliminary or intermediate
stage of life on mudbanks, wet swamps, branches overhanging the
water, etc. Two great families in this group, and two only, seem to
have undergone these transformations, the Littorinidae and the
Neritidae. The derivation of almost all existing land operculates may
be referred to one or other of these groups.

Fig. 16.—Two rows of the radula of Littorina littorea L., × 72.

The power of the Littorinidae to live for days or even weeks
without being moistened by the sea may be verified by the most
casual observer. In the tropics this power seems even greater than
on our own shores. I have seen, in various parts of Jamaica, Littorina
muricata living at the top of low cliffs among grass and herbage. At
Panama I have taken three large species of Littorina (varia, fasciata,
pulchra), on trees at and above high-water mark. Cases have been
recorded in which a number of L. muricata, collected and put aside,
have lived for three months, and L. irrorata for four months.[29]
These facts are significant, when we know that the land operculates
almost certainly originated in a tropical climate.
The Cyclophoridae, Cyclostomatidae, and Aciculidae, which, as
contrasted with the other land operculates, form one group, have
very close relations, particularly in the length and formation of the
radula, or lingual ribbon, with the Littorinidae.

Fig. 17.—Two rows of the radula of Cyclophorus sp., India, ×

40.
On the other hand, the Helicinidae, Hydrocenidae, and
Proserpinidae are equally closely related to Neritina. The
Proserpinidae (restricted to the Greater Antilles, Central America and
Venezuela) may perhaps be regarded as the ultimate term of the
series. They have lost the characteristic operculum, which in their
case is replaced by a number of folds or lamellae in the interior of
the shell. It has already been noticed how one group of Neritina
(Neritodryas) occurs normally out of the water. This group furnishes
a link between the fresh-water and land forms. It is interesting to
notice that here we have the most perfect sequence of derivatives;
Nerita in the main a purely marine form, with certain species
occurring also in brackish water; Neritina in the main fresh-water, but
some species occurring on the muddy shore, others on dry land;
Helicina the developed land form; and finally Proserpina, an aberrant
derivative which has lost the operculum.[30]
 Fig. 18.—A, Neritina reticularis Sowb., Calcutta (brackish
water); B, Helicina neritella Lam., Jamaica (land); C,
Proserpina (Ceres) eolina Ducl., Central America (land).
Gasteropoda.—(2) Pulmonata. The origin of these, the bulk of
the land fauna, must at present be regarded as a problem not yet
finally solved. Some authorities, as we have seen, regard them as
derived from the Nudibranchiate, others, probably more correctly,
from the Tectibranchiate Opisthobranchs.
The first known members of the land Pulmonata (Pupa [?],
Hyalinia) are from the Carboniferous of North America. Similar but
new forms appear in the Cretaceous, from which time to the present
we have an unbroken series. The characteristically modern forms,
according to Simroth,[31] are Helices with thick shells. According to
the same author, Vitrina and Hyalinia are ancestral types, which give
origin not only to many modern genera with shells, but to many shell-
less genera also, e.g. Testacella is probably derived through
Daudebardia from Hyalinia, while from Vitrina came Limax and
Amalia. A consideration of the radulae of the genera concerned
certainly tends in favour of these views.
Godwin-Austen, speaking generally, considers[32] genera of land
Pulmonata with strongly developed mantle-lobes and rudimentary
shell as more advanced in development than genera in which the
shell is large and covers all or nearly all the animal.
 CHAPTER II
LAND AND FRESH-WATER MOLLUSCA, THEIR HABITS AND GENERAL
ECONOMY

The majority of the Land Mollusca are probably more sensitive

than is usually believed. The humidity of the air must affect the
surface of their skin to a considerable extent. Every one has noticed
how the snails ‘come out’ on a damp evening, especially after rain.
As a rule, they wait till rain is over, probably objecting to the patter of
the drops upon their delicate tentacles. Snails kept in captivity under
a bell-glass are acutely sensitive of a damp atmosphere, and will
bestir themselves after rain just as if they were in the open air.
Certain Helices which are accustomed to live in moist places, will
find their way to water, if removed from their usual haunts. A case is
recorded[33] of a specimen of H. arbustorum, kept in a kitchen, which
used to find its way directly under the cold water tap, and appeared
to enjoy the luxury of a douche. How delicately the conditions of life
are balanced in some of these creatures is seen in the case of
Omalonyx, a genus akin to Succinea, which is found in Brazil and
the northern parts of South America. It lives creeping on plants which
overhang the margin of water, but perishes equally, if placed in the
water itself, or removed to a distance from it for any length of time.
[34]

Endurance of Heat and Cold.—The Mollusca are capable, at

least as far as some species are concerned, of enduring severe
extremes both of cold and heat. The most northern pulmonate yet
observed is a fresh-water species, Physa (Aplecta) hypnorum L. This
hardy mollusc, whose shell is so fragile as to need most careful
handling, has been noticed on the peninsula of Taimyr, North Siberia,
in 73° 30’ N. lat, a region whose mean annual temperature is below
10° F. with a range of from 40° F. in July to -30° F. in January.
It is well known that the Limnaeidae, and probably most fresh-
water Mollusca of sub-temperate regions, can continue to live not
merely under, but enveloped in ice, and themselves frozen hard.
Garnier relates[35] that, during the winter of 1829–30, some large
Limnaea auricularia, which had been placed in a small basin, were
frozen into a solid mass, experiencing a cold of -2° F. He supposed
they were dead, but, to his surprise, when the basin thawed, the
Limnaea gradually revived. Paludina vivipara and Anodonta anatina
have been known to resist a temperature of 23° F., and the former
has produced young shortly after being thawed out of the ice.[36] As
far north as Bodø in Norway (67° 37’ N. lat., well within the Arctic
circle) there are found no less than fourteen species of terrestrial
Mollusca, among them being Balea perversa and Clausilia rugosa.
[37]

Vitrina is one of our most hardy molluscs, and may be observed

crawling on bright mornings over the frost-covered leaves of a wood
or copse. V. glacialis is said by Charpentier to live in the Alps at a
height where the stones are covered with snow from nine to ten
months of the year. Many of the Hyaliniae are very hardy. Arion, in
spite of having no external shell to protect it, is apparently less
affected by the cold than Helix, and does not commence hibernation
till a later period in the autumn. The operculate land Mollusca, in
spite of the protection which their operculum may be supposed to
afford, are exceedingly sensitive to cold, and the whole group is
without doubt a product of tropical or semi-tropical regions (see map
at frontispiece). A species of Helicina which inhabits the southern
States of North America has been known to be almost exterminated
from certain districts by the occurrence of an unusually severe
winter.
One of the highest altitudes at which a land shell is known to live
appears to be the Liti Pass (Himalayas, 14,000 ft.). At this enormous
altitude, two species of Buliminus (arcuatus Hutt. and nivicola Bens.)
live on juniper bushes among patches of snow. An Anadenus is said
to have been found in a similar locality at 15,000 ft., while Limnaea
Hookeri has been taken from over 16,400 ft. in Landour. In the
Andes of Peru and Bolivia, five species of Bulimulus, one of Pupa,
and one of Limax occur at an elevation of 10,500 to 15,000 ft.
Several fresh-water Mollusca inhabit Lake Titicaca, which stands at a
height of 12,550 ft. in the Bolivian table-land.
 In certain parts of the desert of Algeria, where there is not a trace
of vegetation to be seen, and the temperature at mid-day is 110° F.,
the ground is sometimes so covered with Helix lactea as to appear
perfectly white. Dr. F. H. H. Guillemard has told me that he noticed,
in somewhat similar surroundings between Fez and Tangier, H.
pisana in such extraordinary abundance that they hung from the low
scrub in bunches the size of a man’s two fists. It is singular that
Mollusca should live, and not only live, but flourish, in localities
apparently so unpromising. Shells which occur in the Algerian
Sahara are actually larger and altogether finer than the ordinary
European form of the same species. In order to protect themselves
to some extent against the scorching heat and consequent
evaporation, desert species are frequently modified in one of two
ways; the shell becomes either white or a light dusky brown, as in
the familiar Helix desertorum, or else it gains immensely in
thickness. Specimens of H. pomatia, recently procured from Fez, are
of extraordinary thickness as compared with forms from our own
chalk downs of Kent and Surrey.
Fresh-water Mollusca are frequently found inhabiting hot springs.
Thus Neritina fluviatilis lives at Bagnères de Bigorre in water at about
68° F. In another hot spring in the eastern Pyrenees a Bithynia lives
at a temperature of over 73° F.; while Blainville mentions another
case of a Bithynia living in water at 122° F.
Hibernation and Aestivation.—As autumn begins to draw on,
and the first frosts to nip vegetation, terrestrial species retire beneath
stones, into cracks in old walls, holes in tree trunks, deep fissures in
rocks, and nooks and crannies of every kind, or else bury
themselves deeply in the earth or in moss and heaps of leaves. They
thus commence their period of hibernation, which varies in length
according to the duration of winter. Frequently masses of Helices
may be found attached to one another, probably not so much for the
sake of warmth, for their temperature is but low, as to share the
comforts of a cosy retreat in common. Slugs generally hibernate
alone, excavating a sort of nest in the earth, in which they encyst
themselves, contracting their bodies until they are almost round, and
secreting a covering of their own slime. The Helices usually close up
the mouth of their shell by the formation of a membranous or chalky
epiphragm, which will be further described below. Both snails and
slugs take care to be in good condition at the time their winter sleep
begins, and for this reason the former are said to be most esteemed
by foreign epicures if captured just at this period.[38]
During hibernation, the action of the heart in land Pulmonata
ceases almost entirely. This appears to be directly due to the effect
of cold. Mr. C. Ashford has related[39] some interesting experiments
made upon H. hortensis and Hyal. cellaria, with the view of
ascertaining the effect of cold upon their pulsations. His observations
may be tabulated as follows:—
Number of pulsations per minute
Helix hortensis Hyal. cellaria At degrees Fahr.
22 21 52°
14 12 44°
10 11 38°
4 9 30°
At low temperatures the character, as well as the number of the
pulsations changed; they became imperfect and intermittent,
although exceptionally at 31° F. a H. rufescens gave five or six
pulsations a minute, very full and deliberate. The result of taking the
Hyalinia suddenly into the heat of a greenhouse was to bring on
palpitations. Further experiments resulted in evidence of a similar
kind. Hyal. radiatula, placed upon a deal table in a room, showed 52
pulsations per minute at 62° F. Placed upon the palm of the hand,
the action soon rose to 108. Hyal. alliaria, similarly treated, rose from
72 pulsations to 110. Floated upon water, the action of the heart of
the latter suddenly fell to 29.
Fresh-water Pulmonata do not appear to hibernate. Unio and
Anodonta, however, bury themselves more deeply in the mud, and
Dreissensia casts off its byssus and retires under the mud in deeper
water.[40] Limnaea and Planorbis have often been noticed to crawl
about under the lower surface of a thick coating of ice. In periods of
prolonged drought, when the water in the ponds dries up, the
majority of genera bury themselves in the mud. I have known
Limnaea peregra bury itself three inches deep, when surprised by a
sudden fall of the water in the ditch on Coe Fen, behind Peterhouse,
Cambridge. Physa hypnorum frequents by preference ditches which
dry up in summer, as does also Planorbis spirorbis, the latter often
forming a sort of epiphragm against evaporation. Ancylus has been
observed to spend the whole winter out of water, and P. spirorbis has
been noticed alive after four months’ desiccation.[41]
True aestivation, however, occurs mainly in the tropics, where
there is no winter, but only a period when it is not quite so hot as the
rest of the year, or on a coast like the Mediterranean, which is
subject to sudden and severe heat. This period is usually rainless,
and the heat is therefore a dry heat. At this season, which may last
for three or four months, most of the land Mollusca enter upon a
period of inaction, either burying themselves deeply in the ground, or
else permanently attaching themselves to the stalks of grass and
other herbage, or the under sides of rocks. For instance, the large
and beautifully painted Orthalicus, Corona, and Porphyrobaphe,
which inhabit Brazil, Ecuador, and eastern Peru, bury themselves
deeply in the ground during the dry season, while in the rains they
climb to the topmost branches of the great forest trees.[42] Thus it
may well happen that a visitor to a tropical island, Ceylon for
instance, or one of the Greater Antilles, if he times his visit to
coincide with the rainless season, may be grievously disappointed at
what seems its unaccountable poverty in land Mollusca. But as soon
as the weather breaks, and the moisture penetrates their retreats,
every bush and every stone, in favoured localities, will be alive with
interesting species.
The Epiphragm.—A considerable number of the land Pulmonata
(and a very few of the fresh-water) possess the power of closing the
aperture of their shell by means of what is known as an epiphragm
or covering of hardened mucus. This epiphragm is habitually formed
by certain species during hibernation or aestivation, or even during
shorter periods of inactivity and retirement, the object being, either to
check evaporation of the moisture of the body, or to secure the
animal against the cold by retaining a thin layer of slightly warm air
immediately within the aperture of the shell.
 The epiphragm differs widely in character in different species,
sometimes (Clausilia, Pupa, Planorbis) consisting of the merest
pellicle of transparent membrane, while at others (Helix aperta, H.
pomatia) it is a thick chalky substance, with a considerable
admixture of carbonate of lime, with the consistency of a hardened
layer of plaster of Paris. Within these extremes every variety of
thickness, solidity, and transparency occurs. During long hibernation
several epiphragms are not unfrequently formed by the same
individual snail, one within the other, at gradually lessening
distances. The epiphragm thus performs, to a certain extent, the part
of an operculum, but it must be remembered that it differs radically
from an operculum physiologically, in being only a temporary
secretion, while the operculum is actually a living part of the animal.
The actual mode of formation of the epiphragm would seem to
differ in different species. According to Fischer,[43] the mollusc
withdraws into its shell, completely blocking all passage of air into
the interior, and closing the pulmonary orifice. Then, from the middle
part of the foot, which is held exactly at the same plane as the
aperture, is slowly secreted a transparent pellicle, which gradually
thickens, and in certain species becomes calcareous. Dr. Binney,
who kept a large number of Helix hortensis in confinement, had
frequently an opportunity of noticing the manner in which the
epiphragm was formed.[44] The aperture of the shell being upward,
and the collar of the animal having been brought to a level with it, a
quantity of gelatinous matter is thrown out [? where from]. The
pulmonary orifice is then opened, and a portion of the air within
suddenly ejected, with such force as to separate the viscid matter
from the collar, and to project it, like a bubble of air, from the
aperture. The animal then quickly withdraws farther into the shell,
and the pressure of the external air forces back the vesicle to a level
with the aperture, when it hardens and forms the epiphragm. In
some of the European species in which the gelatinous secretion
contains more carbonate of lime, solidification seems to take place at
the moment when the air is expelled, and the epiphragm in these is
in consequence strongly convex.
 Thread-spinning.—A considerable number of fresh-water
Mollusca possess the power of stretching a thread, which is no more
than an exceedingly elongated piece of mucus, to the surface of the
water, and of using it as a means of locomotion. This thread bears
no analogy whatever to the fibrous byssus of certain bivalves, being
formed in an entirely different manner, without the need of a special
gland.
The threads are ‘spun’ by several species of Limnaea, Physa, and
Planorbis, by Bithynia tentaculata, and several of the Cycladidae.
They are anchored to the surface by a minute concavity at the upper
end, which appears to act like a small boat in keeping the thread
steady. The longest threads are those of the Physae, which have
been noticed to attain a length, in confinement, of 14 inches. They
are always spun in the ascent, and as a rule, when the animal
descends, it rolls the thread up and carries it down as it goes. A
single thread is never spun on the descent, but occasionally, when a
thread has become more or less of a permanence, it becomes
stronger by the addition of more mucus each time it is used, whether
for ascending or descending purposes. Cyclas cornea appears to be
an exception to the rule that threads are only spun on the ascent.
This species, which is particularly fond of crawling along the under
surface of the water, has been noticed to spin a thread half an inch in
length while on the surface, and to hang suspended from it for a
considerable time.
What the exact use of the thread may be, must to a certain extent
be matter of conjecture. The Limnaeidae are, in the great majority of
cases, compelled to make periodic visits to the surface in order to
inspire oxygen. It is also a favourite habit with them to float just
under the surface, or crawl about on its under side, perhaps in
pursuit of tiny vegetable organisms. Whatever may be the object of
an excursion to the surface, a taut thread will obviously be a nearer
way up than any other which is likely to present itself; indeed, without
this thread-spinning power, which insures a tolerably rapid arrival at
the surface, the animal might find itself asphyxiated, or at least
seriously inconvenienced, before it could succeed in taking in the
desired supply of oxygen. With the Cycladidae, which do not breathe
air, such an explanation is out of place; in their case the thread
seems to be a convenient means of resting in one position in the
intervals of the periods of active exercise to which several of the
species are so much addicted.
The power of suspension by a thread is also possessed by certain
of the Cyclostomatidae, by some Cerithidea, several Rissoa and
other marine genera, prominent among which is Litiopa bombyx,
whose name expresses its power of anchoring itself to the Sargasso
weed by a silken thread of mucus. Several species of slugs are
known to be able to let themselves down by threads from the
branches of trees. Limax arborum is especially noted for this
property, and has been observed suspended in pairs during the
breeding time. According to Binney, all the American species of
Limax, besides those of Tebennophorus, possess this singular
property. Limax arborum appears to be the only slug which has been
noticed to ascend, as well as descend, its thread. It has also been
observed[45] that when this species is gorged with food, its slime is
thin and watery, and unable to sustain its weight, but that after the
process of digestion has been performed, the mucus again becomes
thick and tenacious. It appears therefore that when the animal is
hungry and most in need of the power of making distant excursions
in search of food, its condition enables it to do so, but that when no
such necessity is pressing, the thread-forming mucus is not
secreted, or is perhaps held in suspense while the glands assist in
lubricating the food before digestion.[46]
Food of Land and Fresh-water Mollusca.—Arion ater, the great
black slug, although normally frugivorous, is unquestionably
carnivorous as well, feeding on all sorts of animal matter, whether
decaying, freshly killed, or even in a living state. It is frequently
noticed feeding on earthworms; kept in captivity, it will eat raw beef;
it does not disdain the carcases of its own dead brethren. An old
man near Berwick-on-Tweed, going out one morning to mow grass,
found a black slug devouring, as he supposed, a dead mouse. Being
of an inquisitive turn, and wishing to ascertain if it were really thus
engaged, he drew the mouse a little back. When he returned in the
evening, the mouse was reduced almost to a skeleton, and the slug
was still there.[47] Indeed it would seem almost difficult to name
anything which Arion ater will not eat. Dr. Gray mentions[48] a case
of a specimen which devoured sand recently taken from the beach,
which contained just enough animal matter to render it luminous
when trodden on in the dark; after a little time the faeces of the slug
were composed of pure sand, united together by a little mucus. A
specimen kept two days in captivity was turned out on a newspaper,
and commenced at once to devour it. The same specimen ate dead
bodies of five other species of slugs, a dead Unio, pupae of
Adimonia tanaceti, part of the abdomen of a dragon-fly, and Pears’
soap, the latter reluctantly.[49]
According to Simroth[50] and Scharff[51] the food of several of our
British slugs, e.g. Limax maximus, L. flavus, Arion subfuscus, A.
intermedius, consists of non-chlorophyllaceous substances only,
while anything containing chlorophyll is as a rule refused. On the
other hand L. agrestis and Amalia carinata feed almost entirely on
green food, and are most destructive in gardens. The latter species
lives several inches under ground during the day, and comes to the
surface only at night. It is largely responsible for the disappearance
of bulbs, to which it is extremely partial. L. marginatus (= arborum
Bouch.) feeds exclusively on lichens, and in captivity absolutely
refuses green leaves and a flesh diet. It follows therefore, if these
observations are correct, that the popular notions about slugs must
be revised, and that while we continue to exterminate from our
gardens those species which have a taste for chlorophyll, we ought
to spare, if not encourage those whose tastes lie in the opposite
direction.
Limax agrestis has been seen devouring the crushed remains of
Arion ater. Five specimens of the same species were once noticed
busily devouring a May-fly each, and this in the middle of a large
meadow, where it may be presumed there was no lack of green
food. The capture and eating of insects by Mollusca seems very
remarkable, but this story does not stand alone. Mr. T. Vernon
Wollaston once enclosed in a bottle at least three dozen specimens
of Coleoptera together with 4 Helix cantiana, 5 H. hispida, and 1 H.
virgata, together with an abundant supply of fresh leaves and grass.
About a fortnight afterwards, on the bottle being opened, it was
found that every single specimen of the Coleoptera had been
devoured by the snails.[52] Amalia marginata in captivity has been
fed upon the larvae of Euchelia jacobaeae, eating three in two hours.
[53]

Limax maximus (Fig. 19) has been seen frequently to make its
way into a dairy and feed on raw beef.[54] Individuals kept in
confinement are guilty of cannibalism. Mr. W. A. Gain kept three
specimens in a box together, and found one of them two-thirds
eaten, “the tail left clean cut off, reminding one of that portion of a
fish on a fishmonger’s stall.” That starvation did not prompt the crime
was proved by the fact that during the preceding night the slug had
been supplied with, and had eaten, a considerable quantity of its
favourite food. On two other occasions the same observer found one
of his slugs deprived of its slime and a portion of its skin, and in a
dying condition.[55] An adult L. maximus, kept for thirty-three days in
captivity with a young Arion ater, attacked it frequently, denuded it of
its slime, and gnawed numerous small pieces of skin off the body
and mantle.[56] The present writer has found no better bait for this
species on a warm summer night than the bodies of its brethren
which were slain on the night preceding; it will also devour dead
Helix aspersa. Mr. Gain considers it a very dainty feeder, preferring
fungi to all other foods, and apparently doing no harm in the garden.
 Fig. 19.—Limax maximus L. PO, pulmonary
orifice: × ⅔.
Limax flavus, which is fond of inhabiting the vicinity of cellars,
makes its presence most disagreeable by attacking articles of food,
and especially by insinuating itself into vessels containing meal and
flour.[57] It is particularly partial to cream.
Slugs will sometimes bite their captor’s hands. Mr. Kew relates
that a Limax agrestis, on being stopped with the finger, while
endeavouring to escape from the attack of a large Arion, attempted
to bite fiercely, the rasping action of its radula being plainly felt.
According to the same authority, probably all the slugs will rasp the
skin of the finger, if it is held out to them, and continue to do so for a
considerable time, without however actually drawing blood.[58] While
Mr. Gain was handling a large Arion ater, it at once seized one of the
folds of skin between the fingers of the hand on which it was placed;
after the action of the radula had been allowed to continue for about
a minute, the skin was seen to be abraded.[59] Another specimen of
Arion ater, carried in the hand for a long time enclosed in a dock leaf,
began to rasp the skin. The operation was permitted until it became
too painful to bear. Examination with a lens showed the skin almost
rasped away, and the place remained tender and sore, like a slight
burn, for several days.[60]
 Helix pisana, if freshly caught, and placed in a box with other
species, will set to work and devour them within twenty-four hours.
The present writer has noticed it, in this position, attack and kill large
specimens of H. ericetorum, cleaning them completely out, and
inserting its elongated body into the top whorls of its unfortunate
victims in a most remarkable manner. Amongst a large number of
species bred in captivity by Miss F. M. Hele,[61] was Hyalinia
Draparnaldi. In the first summer the young offspring were fed on
cabbage, coltsfoot, and broadleafed docks. They would not
hibernate even in the severest frosts, and, no outdoor food being
available, were fed on chopped beef. This, Miss Hele thinks, must
have degenerated their appetites, for in the following spring and
summer they constantly devoured each other.
Zonites algirus feeds on decayed fruit and vegetables, and on
stinking flesh.[62] Achatina panthera has been known to eat meat,
other snails (when dead), vegetables, and paper.[63] The common
Stenogyra decollata of the South of Europe has a very bad character
for flesh-eating habits, when kept in captivity. Mr. Binney[64] kept a
number for a long time as scavengers, to clean the shells of other
snails. As soon as a living Helix was placed in a box with them, one
would attack it, introduce itself into the upper whorls, and completely
remove the animal. One day a number of Succinea ovalis were left
with them for a short time, and disappeared entirely! The Stenogyra
had eaten shell as well as animal. This view of Stenogyra is quite
confirmed by Miss Hele, who has bred them in thousands. “I can
keep,” she writes,[65] “no small Helix or Bulimus with them, for they
at once kill and eat them. They will also eat raw meat.”
Even the common Limnaea stagnalis, which is usually regarded
as strictly herbivorous, will sometimes betake itself, apparently by
preference, to a diet of flesh. Karl Semper frequently observed the
Limnaeae in his aquarium suddenly attack healthy living specimens
of the common large water newt (Triton taeniatus), overcome them,
and devour them, although there was plenty of their favourite
vegetable food growing within easy reach.[66] The same species has
also been noticed to devour its own ova, and the larvae of Dytiscus.