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 In the fixed inequivalves (e.g. Chama) it is sometimes the right,
sometimes the left valve which is undermost, but the fixed valve,
whether right or left, is always deep, and the free valve flat. Ostrea
and Anomia are always fixed by the left valve.
The lunule is a well-marked area in front of and close to the
umbones, usually more or less heart-shaped, and limited by a ridge.
Generally it is shallow, but sometimes, as in Dosinia, Opis, and some
Cardium, deeply impressed. A corresponding area behind the
umbones, enclosing the ligament, is called the escutcheon (Fig.
186), but it seldom occurs.
The ligament is a more or less elastic band, which unites the two
valves along a line adjacent to the umbones. As a rule, the greater
part of the ligament is external to the shell, but it may be entirely
internal. It is placed, normally, behind the umbones, but in a few
cases, when the hinge line is very long (Arca, Pectunculus), it
extends in front of them as well. The edges of the valves, when the
ligament is mainly external, are more or less excavated for its
reception. When internal it is generally contained in a groove or
spoon-shaped pit, known as the fossette (compare Fig. 187).

Fig. 186.—Venus
subrostrata Lam.: es,
escutcheon; li,
ligament; lu, lunule; u,
u, umbones.
 The ligament consists of two distinct parts, which may occur
together or separately, the external, or ligament proper, and the
internal, or cartilage. Only the external portion can be seen when the
valves are closed. As a rule, the two portions are intimately
connected with one another, the ligament folding over the cartilage,
but in some cases, e.g. Mya, Mactra, where the cartilage is lodged
within the hinge, they are completely disconnected (Fig. 187).
In Pecten the external ligament is very thin, and runs along the
dorsal margin, while the internal ligament is large, solid, and situated
in a shallow pit. In Perna, where the hinge is toothless, the ligament
is folded into a number of transverse ridges, which fit into
corresponding grooves in the shell.
The ligament proper is inelastic and insoluble in caustic potash.
The cartilage is very elastic, composed of parallel fibres, slightly
iridescent, and soluble in caustic potash.
The operation of the ligament—using the word as including the
whole ligamental process—is in opposition to that of the adductor
muscles. When the latter close the valves, they compress the
ligament, an action which its elasticity resists: thus its operation
tends in part towards keeping the valves open. But when ligament
and cartilage are both fully developed, they work in opposition to one
another, the ligament, by its resistance to compression, preventing
any straining of the adductor muscles when the valves are open, and
the cartilage, for the same reason, preventing the ventral margins of
the shell from closing too rapidly upon one another when the valves
are being shut.
 Fig. 187.—Hinge of A, right
valve, and B, left valve of
Mulinia edulis King; ca,
cardinals; l.a, anterior
laterals; l.p, posterior
laterals; f, fossette; c,
cartilage; l, ligament.
The Hinge.—The valves of Pelecypoda are generally articulated,
below the umbones, by a hinge which is furnished, in the majority of
cases, with interlocking teeth, small pits or depressions in each valve
corresponding to the teeth in the other. The teeth are distinguished
as cardinal, or those immediately below the umbo, and lateral, or
those to either side of the cardinals, the latter being also
distinguished as anterior and posterior laterals, according as they
are before or behind the umbo (Fig. 184). In shells which are
tolerably equilateral there is no difficulty in distinguishing between
cardinal and lateral teeth. But when they are very inequilateral, the
whole hinge may share in the inequality of growth, and an anterior
lateral may be thrown backward and simulate a cardinal, or a
cardinal may be thrown backward and simulate a posterior lateral
(e.g. Cardita, Unio, Fig. 188). In many Chama the cardinals are
pushed up into the umbo and become a mere ridge, while the strong
anterior lateral becomes nearly central and simulates a cardinal.
 Fig. 188.—Hinges of A, Cardita semiorbiculata Brug., and B,
Unio rectus Lam., showing how, in inequilateral shells, the
lateral teeth tend to shift their position. a.m, anterior
adductor, p.m, posterior adductor muscle; c, c, cardinal
teeth; p.l, posterior lateral teeth; l, ligament.
Some bivalves, e.g. Anodonta, Ostrea, Pedum, many Mytilus,
have no hinge teeth at all, in others the laterals are wanting
(Psammobia, Diplodonta). In the Arcadae the hinge consists of a
number of very similar denticles, which are often serrated like the
teeth of a comb (Fig. 189).
 Fig. 189.—The hinge in
Arcadae: A, Nacula Loringi
Ang. × 4/3; B, Arca
granosa L.; u.a, umbonal
area.

Fig. 190.—A, Tridacna scapha Lam.; B,

Cardium enode Sowb., showing the
interlocking of the ventral margins.
Hinge-teeth are probably, in origin, derived from the crenulations
or ribbings of the surface of the shell, the upper ends of which
impinge upon the dorsal margin and mark it in a way which is quite
recognisable when the shell is thin. Similar crenulations, resulting in
interlocking of the valves, are not uncommon upon the ventral
margin in certain genera (Fig. 190). The mechanical effect of these
continued riblets, when fitted together on the opposing valves, would
be to prevent the valves sliding upon one another while closing, or
after being closed. Thus there would be a probability of their
surviving, even after the ribbing had disappeared from the surface of
the shell, the increased strength given by the hinge compensating
for, and making it possible to do without, the extra strength supplied
by the ribs. It is therefore possible that the teeth of the Nuculidae and
Arcadae, which have no distinction between cardinals and laterals,
represent a very ancient type, from which have been evolved the
various forms of hinge in which cardinals and laterals are
distinguished. Even in some forms of Arcadae (comp. Pectunculus)
we get a hint how the transverse teeth of the typical Arca may have
become transformed into the longitudinal tooth of the normal lateral.
[350]

The developed hinge-teeth, then, ensure the opening of the

valves in one direction; they also secure their accurate closure upon
one another in exactly the same plane. Exposed shells and gaping
siphons matter little to animals which are protected by their
burrowing propensities, but to those which live in material which can
be easily penetrated by their foes, it must be of advantage to be able
to buckle their armour absolutely tight. The edentulous hinge of
Anodonta is a degeneration from a dentate type, which retains its
teeth (in Unio, etc.) when subject to the jar of rapid streams, but
tends to lose them in the stiller waters of canals, lakes, and ponds.
Other processes in the bivalve shell.—In Anatina each umbo is
fissured and strengthened on the inside by a kind of umbonal plate
which carries the ligament. Some forms of Liligna develop a strong
internal umbonal rib, which serves as a buttress to strengthen the
shell. In Pholas, the so-called falciform process serves as a point of
attachment for the muscles of the foot and viscera. There is no
ligament or hinge-teeth, the place of the latter being taken by the
anterior adductor muscle, which is attached to the hinge-plate, the
latter being reflected back into the shell.
 In Septifer the anterior adductor muscle is carried on a sort of
shelf or myophore, and in Cucullaea the posterior adductor is partly
raised on a similar and very prominent formation.
Length and breadth of bivalve shells is variously measured. Most
authorities measure length, or ‘antero-posterior diameter,’ by a
straight line drawn from the extreme anterior to the extreme posterior
margin, and breadth by a similar line, drawn from the umbones to a
point, not very clearly marked, on the opposite ventral margin (see
Figs. 184 and 185). Others, less correctly, reverse these terms.
Thickness is measured by the extreme distance of the opposite
faces of the closed valves. As a rule, the length exceeds, and often
greatly exceeds, the thickness, but in a few cases—e.g. the Cardissa
section of Cardium—this is reversed.
The periostracum.—Nearly all shells are covered, at some period
of their growth, by a periostracum,[351] or surface skin, which serves
the purpose of protecting the shell against the destructive effects of
the chemical action set up by water or air. It also, in some cases (see
p. 258), acts as a kind of base upon which the shell is deposited. In
old shells it is commonly worn away, especially at those parts which
are likely to become abraded.
The form and composition of the periostracum varies greatly.
Sometimes (e.g. Oliva) it is a mere transparent film, at others
(Zonites) it is transparent, but stout and solid. It is corneous in
Solenomya, covered with fine hairs in many Helicidae, in Conus,
Velutina, and Cantharus it is thick, fibrous, and persistent; in
Trichotropis and some Triton it is furnished with long bristles on a
thick ground (Fig. 191). In fresh-water shells it is usually rather thick,
in order to protect the shell from the erosive powers of certain kinds
of water. In some cases (Mya, Anatina) the periostracum is
continued over the siphons, so as to form a protection throughout
their whole length.
 Fig. 191.—Triton olearium L.,
Mediterranean, an example
of a shell with a stout and
hairy periostracum. × ½.
Erosion.—The fresh-water Mollusca generally, and marine
mollusca in a few rare cases (Purpura, Littorina) are subject to
erosion, or decay in the shell substance. In univalves erosion usually
sets in near the apex (Fig. 192), where the life of the shell may be
regarded as weakest, and in bivalves near the umbones. It is
commonest in old shells, and rarely occurs in the very young. So
long as the periostracum is present to protect the shell, erosion
cannot set in, but when once it has been removed the shell is liable
to the chemical changes set up in its substance by water. There is
abundant evidence to show that erosion is caused by pollution of
water. Out of many instances one must suffice. In a certain stream
near Boston, U.S., numbers of Mollusca occurred, the shells of
which were very perfect and free from disease. Some little way down
stream an alkaline manufactory drained its refuse into the water. At
and below this point for some distance every shell was more or less
eroded, most of them seriously. Farther down, when the alkali refuse
became diluted away, the shells retained their normal condition.[352]
 Fig. 192.—Example of an eroded
fresh-water shell (Melania
confusa Dohrn, Ceylon).
A small percentage of lime in the water appears to produce
erosion. The result of some experiments by G. W. Shrubsole, in the
investigation of this point, may be tabulated as follows:[353]—
Water from Lime present Result
per gall.
River Dee, near Chester 3·00 grs. acted strongly on shells
Wrexham 4·00 grs. „ „ „
River Dee, near Llanderyel 0·53 grs. „ „ „
Trent Canal 8·33 grs. no action „
 CHAPTER X
GEOGRAPHICAL DISTRIBUTION OF LAND AND FRESH-WATER MOLLUSCA—
THE PALAEARCTIC, ORIENTAL, AND AUSTRALASIAN REGIONS

The Mollusca afford specially valuable evidence on problems of

geographical distribution. This fact is largely due to their extreme
susceptibility to any change in the conditions of life. Genera which
are accustomed to live in a certain temperature and on certain food,
cannot sustain life if the temperature falls or rises beyond certain
limits, or if the required food be not forthcoming. There is therefore a
marked contrast between the Mollusca of the tropics and of the
temperate zones, while different regions in the same latitude,
whether within or without the tropics, often show great diversity in
their fauna. Every region is thus characterised by its Mollusca. The
Mollusca, for instance, of Australia or of South Africa characterise
those countries quite as much as do the kangaroo and the emu, the
hartebeest and the ostrich; there is nothing like them anywhere else
in the world. In the Greater Antilles the Mollusca stand out beyond all
other forms of life as characteristic of the islands as a whole, and of
each separate island in particular.
The geographical distribution of the land and fresh-water Mollusca
must be considered quite apart from that of the marine Mollusca.
The sea offers no such serious barriers to the spread of the latter as
the land does to the spread of the former. If we were to journey to
the Azores, and turn our attention to the land-snails, we should find
them almost wholly peculiar, while amongst the sea-shells we should
recognise many as occurring also on our southern coasts, and few
that were different from those of the Mediterranean. The marine
Mollusca of the Sandwich Islands, in spite of the enormous
intervening distance, are not very different from those of Natal, but
the land Mollusca of the two countries are as widely different as is
possible to imagine.
Land Mollusca are, as has been remarked, fettered to the soil.
Quadrupeds, birds, fishes, and reptiles are provided with organs of
motion which enable them to overpass barriers of various kinds.
Even plants, although themselves incapable of motion, may be
conveyed in every direction by means of seeds, which are either
wafted by the wind or adhere to the skin of animals. But the Mollusca
have no such regular means of transport, and are, in a large number
of instances, limited to districts of a certain character of soil, or
producing certain kinds of vegetation.
The localisation, both of genera and species, occurs all over the
world. The genus Achatinella, which is peculiar to the Sandwich
Islands, is found there in a profusion of species. It lives in the
mountain valleys which radiate from the central ridge of each island,
and each valley is characterised by its own peculiar set of species.
The great carnivorous Glandina is restricted to Central America and
the adjacent parts of the two continents, with one or two species in
Southern Europe. Bulimus proper is restricted to South America;
Achatina to Africa south of the Sahara; Tornatellina to the Pacific
Islands; Cochlostyla to the Philippines; Cylindrella and Bulimulus are
peculiar to the New World; Buliminus, Nanina, Scarabus, and
Cassidula to the Old.
Extreme cases of this restriction of habitat sometimes occur. Thus
Limnaea involuta is found only in a single small mountain tarn in
Ireland; Clausilia scalaris along a narrow strip of limestone in Malta;
Strophia nana is confined to a few square rods on an island that is
itself a mere dot in the Caribbean Sea; the genus Camptonyx occurs
only in the neighbourhood of Mt. Girnar, in Gujerat; and Lantzia in
moss on the top of a mountain in Bourbon.
Attempts to colonise snails in strange localities have usually
resulted in failure, especially when the attempt has involved serious
changes of environment. The common Cochlicella acuta of our
coasts resists all endeavours to establish it beyond a certain
distance from the sea. Snails brought from the Riviera and placed
under almost similar conditions of climate on our own southern
coasts have lived for a while, but have very rarely taken permanent
root. Mr. H. W. Kew[354] has collected a good many of these
attempts to acclimatise species, the general success of which seems
to depend almost entirely on a restoration of the old conditions of life.
 At the same time there are certain species which exhibit a
curiously opposite tendency, and which seem capable of flourishing
in almost any part of the world, and under the most varied
surroundings. Our own common garden snail (Helix aspersa) is a
striking instance of this adaptability to new conditions. It has been
established, by art or by accident, in Nova Scotia, Maine, South
Carolina, New Orleans, California, Mexico city, Cuba, Hayti,
Cayenne, Brazil, Valparaiso, Cape Town, the Azores, St. Helena,
Mauritius, Loyalty Islands, and Australia. The great Achatina fulica of
East Africa has been established first in Mauritius, and from thence
has been carried to the Seychelles and Calcutta. Helix lactea, a
common Mediterranean species, has been carried to Teneriffe and
Montevideo; Helix similaris, whose fatherland is Eastern Asia, has
been transported to Mauritius, Bourbon, West Africa, West Indies,
Brazil, and Australia; Ennea bicolor (Eastern Asia) to India, Bourbon,
Mauritius, West Indies; Stenogyra decollata (Mediterranean basin) to
South Carolina; S. Goodallii (West Indies) to British pineries; Helix
Hortensis to New Jersey. Seven common English species (Limax
gagates, Hyalinia cellaria, H. alliaria, Helix aspersa, H. pulchella,
Pupa umbilicata) have become naturalised in St. Helena,[355] and as
many as nineteen in Australia.[356]
Cases of artificial transport of this kind are readily detected; they
follow the lines of trade. The snails themselves or their ova have
been accidentally enclosed with plants or mould, or have adhered to
packing-cases, or to hay and grass used in packing. Thus they
constitute no disturbance to the general rule of the persistent
localisation of species and genera, and there is little fear that the
evidence which the geographical distribution of the Mollusca brings
to bear upon the general problems of distribution will be confused by
any intermixture of fauna naturally distinct.
Land Mollusca: Barriers to Dispersal.—The chief natural barriers
to dispersal are extremes of temperature, the sea, mountain ranges,
and deserts. Rivers, however large, seem of little effect in checking
dispersal. There is no appreciable difference between the land
Mollusca north and south of the Ganges, or north and south of the
Amazon. Living snails, or their ova, are no doubt transported from
one bank to another on floating débris of various kinds. The barrier
offered by the sea is obvious, and at first sight appears
insurmountable; but the facts with regard to oceanic groups of
islands like the Azores and Canaries (see p. 297) show that even a
stretch of salt water many hundred miles in breadth may be
ineffectual in preventing the dispersal of Mollusca.
Mountain ranges, provided they are too high to be scaled, and too
long to be turned in flank, offer a far more effective barrier than the
sea. Every thousand feet upward means a fall of so many degrees in
the mean temperature, and a change, more or less marked, in the
character of the vegetation. There is generally, too, a considerable
difference in the nature of the climate on the two sides of a great
mountain range, one side being often arid and cold, the other rainy
and warm. The combined effect of these influences is, as a rule,
decisive against the dispersal of Mollusca. Thus the Helices of
California are almost entirely peculiar; one or two intruders from
states farther east have succeeded in threading their way through
the deep valleys into the Pacific provinces, but not a single genuine
Californian species has been able to scale the heights of the
Cascade Mountains. The land Mollusca of India are numbered by
hundreds; not one penetrates north of the Himalayas. According to
Mr. Nevill,[357] the change from the Indo-Malayan to the so-called
European molluscan fauna at the northern watershed of the Kashmir
valley is most abrupt and distinct; in two days’ march northward,
every species is different. Ranges of inferior altitude, such as the
Pyrenees, the Carpathians, or the Alleghanies, may be turned in
flank as well as scaled, and we find no such marked contrast
between the Mollusca on their opposite sides.
The most effective barrier of all, however, is a desert. Its scorching
heat, combined with the absence of water and of vegetable life,
check dispersal as nothing else can. The distribution of the Mollusca
of the Palaearctic Region is an excellent instance of this. Their
southern limit is the great desert which stretches, with scarcely a
break, from the west coast of Africa to the extreme east coast of
Asia. The Mediterranean offers no effectual barrier; shells of
southern Europe are found in profusion in Morocco, Tunis, and
Egypt, while all through Siberia to the extreme of Kamschatka the
same types, and even the same species, of Mollusca occur.
A detailed examination of the means, other than voluntary, by
which Mollusca are transported from one place to another hardly
comes within the scope of this work. Ocean currents, rivers, floods,
cyclonic storms of wind, birds, and even beetles and frogs, play a
part, more or less considerable, in carrying living Mollusca or their
ova, either separately or in connexion with floating débris of every
kind, to a distance from their native home. Accidental locomotion, of
one or other of these kinds, combined with the well-known tenacity of
life in many species (p. 37), may have contributed to enlarge the
area of distribution in many cases, especially in the tropics, where
the forces of nature are more vigorous than in our latitudes. The
ease with which species are accidentally spread by man increases
the probability of such cases occurring without the intervention of
human agency, and numbers of instances may be collected of their
actual occurrence.[358]
A point, however, which more concerns us here is to remark on
the exceedingly wide distribution of the prevailing forms of fresh-
water Mollusca. It might have been expected that the area of
distribution in the fresh-water forms would be greatly restricted, since
they cannot migrate across the land from one piece of water to
another, and since the barriers between pond and pond, lake and
lake, and one river system and another are, as far as they are
concerned, all but insuperable. We might have expected, therefore,
as Darwin and Wallace have remarked, to find a great multiplicity of
species confined to very restricted areas, since the possibility of
communication with the parent stock appears, in any given case, to
be so exceedingly remote.
As is well known, the exact reverse occurs. The range, not merely
of genera, but even of individual species, is astonishingly wide. This
is especially the case with regard to the Pulmonata and Pelecypoda.
The genera Limnaea, Planorbis, Physa, Ancylus, Unio, and Cyclas
are world-wide. Out of about ten genera of fresh-water Mollusca in
New Zealand, one of the most isolated districts known, only one is
peculiar. In South Africa and the Antilles no genus is peculiar. In the
latter case, this fact is remarkable, when we consider that the same
sub-region has at least ten peculiar genera of operculate land
Mollusca alone.
To give a few instances of the distribution of particular species:—
Limnaea stagnalis L. occurs in the whole of Europe, and northern
Asia to Amoorland, Turkestan, Afghanistan, North Persia, and
Kashmir; Greenland, North America from the Atlantic to the Pacific,
and from North Canada and British Columbia as far south as Texas.
The distribution of L. peregra Müll., L. truncatula Müll., and L.
palustris Müll, is almost equally wide.
Planorbis albus occurs in the whole of Europe, and northern Asia
to Amoorland, Kamschatka, and Japan; Turkestan, the Altai-Baikal
district, Alaska and Greenland, North Canada, and the whole of
eastern North America.
The distribution of Anodonta anatina L., Cyclas cornea L., and
Pisidium pusillum Gmel. is almost equally wide.
It is evident that the accidental means of transport mentioned
above are insufficient to account for the facts as we find them; we
are therefore compelled to seek for further explanation. Anything in
the nature of a current furnishes a ready means of transport for
Mollusca which have obtained a footing in the upper waters of a
river, and there is no difficulty in imagining the gradual spread of
species, through the agency of floods or otherwise, over a whole
river system, when once established at any point upon it. The feeble
clinging power of newly-hatched Limnaea has often been noticed as
contributing to the chances of their range of distribution becoming
extended. Fresh-water Mollusca, too, or their ova, are exceedingly
likely, from their extreme abundance, to be transported by water-
birds, which fly without alighting from one piece of water to another.
Again, the isolation of one river system from another is, in many
instances, by no means well marked or permanent, and a very slight
alteration of level will frequently have the effect of diverting the
supplies of one watershed into another. When we know what
enormous oscillations in level have taken place over practically the
whole surface of the globe, we can recognise the probability that the
whole river system of the earth has been mixed up and
reconstructed again and again, with a very thorough blending of
adjacent fauna.
It is possible that the very uniform conditions under which fresh-
water Mollusca live may have something to do with the uniformity of
their distribution and the comparative sameness in their
development. There can scarcely be any question that the
environments of fresh-water species are in themselves less varied
and less liable to fluctuation than those of species whose home is
the land. Water is very like water, all the world over; it may be
running or motionless, warm or cold, clear or muddy, but the general
tendency is for it to be free from extremes of any kind. Even if the
surface water of a lake or river freezes, or becomes unusually hot,
there is generally plenty of water at a lower stratum which maintains
a less extreme temperature, and to which creatures can retire on the
first symptoms of a change. From this two results will follow. Not only
will the inhabitants of a piece of water not be inclined to vary much
from the type, since their whole surroundings, food, etc., continue
very much the same, but, if transported by any accident or cataclysm
elsewhere, they will be exceedingly likely to arrive at a place which
closely resembles their former home in all essentials. Thus the
tendency for new types to be formed would be constantly checked,
or rather would very seldom arise.
Mr. Belt, while recognising the importance of changes of level as
affecting the distribution of fresh-water species, appears to regard
the operations of such changes from a rather different point of view
to that described above. “I think it probable,” he writes,[359] “that the
variation of fresh-water species of animals and plants has been
constantly checked by the want of continuity of lakes and rivers in
time and space. In the great oscillation of the surface of the earth, of
which geologists find so many proofs, every fresh-water area has
again and again been destroyed.... Thus species of restricted range
were always exposed to destruction, because their habitat was
temporary and their retreat impossible, and only families of wide
distribution could be preserved.”
 The terrestrial surface of the globe has been divided, as indicating
the facts of geographical distribution, into six regions—the
Palaearctic, Oriental, Australasian, Ethiopian, Nearctic, and
Neotropical. To these is sometimes added a seventh, the
Neantarctic, consisting of Chili and Patagonia (and certain islands of
the south Atlantic); but since the Mollusca of Chili unmistakably form
a part of the Neotropical fauna, it seems hardly worth while to
recognise a separate region for those of the extreme south of South
America, which have no peculiar characteristics.
In certain points the exact limits of these regions, as indicated by
the Mollusca, will probably not correspond to those which are
marked out by other zoological classes. Wallace’s line, for instance,
does not exist, as far as the Mollusca are concerned.
These regions may be further subdivided into sub-regions, thus:—
Regions Sub-regions
Septentrional
Palaearctic Mediterranean
Central Asiatic
Indo-Malay
Oriental
Chinese
Papuan
Australasian Australian
Polynesian
Central African
Ethiopian South African
Malagasy
American
Nearctic
Californian
Neotropical Antillean
Central American
Colombian
Brazilian
Chilian
 A. The Palaearctic Region

The southern boundary of this region is the northern limit of the

African Sahara, the Mediterranean forming no break whatever in its
continuity. In Asia this boundary is less well marked, but roughly
corresponds to the southernmost of the vast ranges of mountains
which border the great tablelands of central Asia. Across Africa the
line of desert is well defined; but in the north-east, as the desert
approaches more nearly to the sea, the African extent of the region
is correspondingly narrowed until it becomes little more than a strip
of coast land, scarcely widening even in Lower Egypt. On the
Morocco coast, Palaearctic land forms penetrate as far south as
Cape Nun.[360] At its eastern extremity the line becomes less well
defined, but probably proceeds along the snowy mountains west of
Setchouan, the Pe-ling and Tan-sia-shan ranges, so as to include all
the high ground of Thibet and of the upper waters of the Hoang-ho,
and ultimately reaches its eastern limit at some point on the shores
of the Sea of Japan.
The region thus includes all Europe, Africa north of the Sahara,
with the Atlantic islands (the Azores, Canaries, etc.), North Arabia,
Asiatic Turkey, the greater part of Persia, Afghanistan, Thibet, all
Asiatic Russia, and a very large portion of the Chinese empire.
The principal characteristics of the region as a whole are:—
(1) The rich development of Helix, Arion, Limax, Buliminus, and
Clausilia.
(2) The comparative absence of land operculates (see map,
frontispiece).
 (3) The uniform character of the fresh-water fauna.
It is in the southern portion of the region that Helix (in the sub-
genera Macularia, Iberus, Pomatia, and Xerophila) and Buliminus
(Zebrina, Chondrula, Ena) attain their maximum. In the north,
Fruticicola is the characteristic group; in the mountainous districts of
the south-east, Campylaea, with Clausilia. The Arionidae have their
headquarters in the damp and warm regions of western Europe, but
are rare in the south. They only approach the Mediterranean coast in
Algeria, near Gibraltar, and in the region between the base of the
Pyrenees and the Maritime Alps, and are very poor in species
throughout Italy and Sardinia. They are absent from almost the
whole of northern Africa, the Mediterranean islands (except
Sardinia), the whole Balkan district, the Crimea, Caucasus, and
western Asia.[361]
The uniformity of the fresh-water fauna is disturbed only in the
extreme south. A few species of Melanopsis, with Neritina, occur in
southern Spain and Austria, Galicia, and southern Russia, while a
Melania or two (absent from Spain) penetrate the south-eastern
parts of Europe as far as Germany. Cyrena begins to replace Cyclas
in southern Russia and the Caucasus.
The Palaearctic region falls into three sub-regions:—
(1) The Northern or Septentrional Sub-region, i.e. the district
north of the line formed by the Pyrenees,[362] Alps, Carpathians, and
which, passing to the northward of the Aralo-Caspian district, follows
the great central mountain range of Asia until it reaches the Sea of
Japan, perhaps somewhere in the neighbourhood of Vladivostok.
(2) The Mediterranean Sub-region, i.e. the countries bordering
on the Mediterranean, the Black and Caspian Seas, with the Atlantic
Islands.
(3) The Central Asiatic Sub-region, i.e. Turkestan, Afghanistan,
Thibet, and probably the districts of Mongolia and Manchuria.[363]
(1) The Septentrional Sub-region has been divided by some
writers into two provinces, the European and the Siberian. There
seems, on the whole, but little occasion to separate off northern
Asia, the characteristic of which is, as will be seen below, rather the
gradual disappearance, as we proceed eastward, of European
species and genera, than the development of any new and peculiar
groups. The remarkable fauna of Lake Baikal stands apart, not only
from European, but also from the Siberian types occurring in its
immediate neighbourhood.
On the whole, the Septentrional Sub-region is poor in species
except those which inhabit fresh water. This fact is probably due to
the extreme vicissitudes of temperature which prevail, and it is
interesting to notice that the number of land Mollusca appears to
touch its lowest point in districts where the annual range of
temperature is greatest. On the other hand, in the western portions
of the region, where the climate is moist and temperature more
equable, the Mollusca are considerably more abundant and varied.
The line which separates the Septentrional from the
Mediterranean Sub-region must of necessity be very roughly drawn,
and stragglers from the south will be found to make their way
northward, and vice versâ, under favouring circumstances of
temperature and geological formation. Jordan has noticed[364] that
species which in southern countries are not confined to any
particular quality of soil are in more northern latitudes found only on
limestone, which absorbs more heat than other formations.
Conversely, the higher elevations of the Alps, Pyrenees, and even
Carpathians are like islands in a sea, and support a thoroughly
northern fauna, quite strange to that of the plains below. Thus Helix
harpa Say, a completely boreal shell, which is at home in Canada,
Sweden, Lapland, and the Amoor district, is found on the Riffel Alp,
at a height of 6000 feet.[365] Vertigo arctica Wall., a species
abundant in Lapland, North Siberia, Iceland, and Greenland, occurs
on the high Alps of the Tyrol.
Circumpolar Species.—A certain number of species are common
to the extreme north both of the Palaearctic and Nearctic regions,
and are, in fact, circumpolar. The number of these species, however,
is so small, not exceeding about 40 species (= 16 genera), that it