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Otoconcha is peculiar. The fresh-water Mollusca, besides the Isidora
characteristic of the sub-region, are partly related to New Caledonia
through the occurrence of Melanopsis, partly to Tasmania through
Potamopyrgus, while the peculiar Latia is possibly akin to
Gundlachia (Tasmania). The land operculates number only 5 genera
and 14 species in all, excluding a doubtful Diplommatina.[374]
Land and Fresh-water Mollusca of the Neozealanian Province
Schizoglossa 1
Paryphanta 5
Rhytida 6
Rhenea 2
Helicarion 1
Otoconcha 1
Microcystis 1
Trochonanina 1
Phacussa 3
Thalassohelix 5
Gerontia 2
Allodiscus 10
Pyrrha 1
Therasia 7
Phenacohelix 3
Suteria 1
Flammulina 13
Laoma 23
Endodonta 10
Charopa 28
Placostylus 1
Carthaea 1
Tornatellina 1
Janella 3
Latia 2
Ancylus 2
Limnaea 5
Amphipeplea 2
Planorbis 1
Isidora 7
Melanopsis 2
Potamopyrgus 4
Paxillus 1
 Lagochilus 7
Omphalotropis 1
Realia 4
Hydrocena 1
Unio 9
Sphaerium 1
Pisidium 2
Lord Howe’s I. is remarkable as containing a Placostylus, which
thus links the island with this province. The remainder of the fauna is
Polynesian, with the exception of a species (common to the Fijis) of
Parmella, a slug akin to Helicarion, Parmacochlea, and Cystopelta.
(3) The Polynesian Sub-region includes all the island groups of
the central and southern Pacific (except those classified in the
Papuan and Australian sub-regions), from the Pelews and Carolines
in the west to the Marquesas and Paumotus in the east, and from
the Tonga group in the south to the Sandwich Is. in the north. It may
be subdivided into (a) the Polynesian province proper, and (b) the
Hawaiian province, which includes the Sandwich Is. only.

Fig. 216.—Characteristic
Polynesian Mollusca: A,
Achatinella vulpina Fér.,
Sandwich Is.; B, Partula
planilabrum Pease, Society
Is.
(a) The general features of the Polynesian province are very
similar throughout, although the Mollusca of each island group are in
the main peculiar. The species are mostly small and obscure. Helix
scarcely occurs, its place being taken by small Zonitidae
(Microcystis, Charopa, Trochomorpha, etc.), and by groups of so-
called Patula (Endodonta, Pitys, etc.), the exact position of which is
not yet settled. Libera, remarkable for its method of ovipositing (p.
128), is peculiar to the Society and Hervey Is.; Partula is almost
universal, attaining its maximum (40 sp.) in the Society Is.;
Tornatellina, Pupa, and Vertigo occur throughout.
The land operculates consist chiefly of Omphalotropis, Pupina,
Realia, and Helicina. Diplommatina and Palaina are abundant on the
Pelews, and a Moussonia occurs in the Samoa Is. Ostodes, a small
form of Cyclophorus, is found in some of the southern groups. The
fresh-water operculates are Melania, Neritina (including Clithon, a
sub-genus furnished with spines), and Navicella; there are no
Unionidae, while fresh-water Pulmonata are very scarce.
(b) The land Mollusca of the Hawaiian province are distinguished
by the possession of four entirely peculiar genera—Achatinella,
Leptachatina, Carelia, and Auriculella. More than 300 of the two
former genera have been described, every mountain valley of some
of the islands having its own peculiar species. The destruction of the
indigenous herbage by goats is rapidly extinguishing many forms.
Partula, and the small land operculates, so characteristic of the other
groups, are, with the exception of Helicina, entirely wanting. The
occurrence of one of the Merope group of Helix (Solomon Is.) is
remarkable, and there is a rich development of Succinea. “Patula,”
Microcystis, Tornatellina, and the other small Polynesian land
Pulmonata are well represented. The presence of Isidora, absent
from the central Pacific groups, is remarkable, and Erinna is a
peculiar genus belonging to the Limnaeidae.
 CHAPTER XI
GEOGRAPHICAL DISTRIBUTION OF LAND MOLLUSCA (continued)—THE
ETHIOPIAN, NEARCTIC, AND NEOTROPICAL REGIONS

D. The Ethiopian Region

The Ethiopian region includes the whole of Africa south of the
Great Desert, and Southern Arabia, together with the outlying
islands, excepting those of the Atlantidean province (p. 297).
Regarded as a whole, the Ethiopian is poorest in land Mollusca of
all the tropical regions. And yet its characteristics are very
remarkable. The entire Achatina group is peculiar, and takes,
especially in W. Africa, some curious forms (Columna, Perideris,
Pseudachatina). Carnivorous Mollusca (Ennea, Gibbus, etc.) are
highly developed, especially in the south and east, the largest known
helicoid form (Aerope) being from Natal. In the possession of these
types of the Agnatha, Africa is more closely related to the
Australasian than to the Oriental region. The true Cyclostoma are
entirely peculiar to the region, but are absent from West Africa.
Fresh-water Mollusca are abundant and characteristic, especially
in and near the Great Lakes. Lanistes, Cleopatra, and Meladomus,
among the operculates, together with Mutela and Aetheria
(Unionidae), Galatea and Fischeria (Cyrenidae), are peculiar.
In its negative, as well as its positive features, the Ethiopian
region is markedly isolated. Helicidae and Naninidae are equally
deficient, the former, indeed, attaining some numerical
predominance in the extreme south, but the species are nearly all
insignificant in size and colouring. It is only in Madagascar that Helix
asserts itself. Arion, Limax, Hyalinia, Clausilia, and a number of other
genera abundant along the Mediterranean, are either altogether
absent, or are very scantily represented. Land operculates, so
characteristic of other tropical countries, are almost entirely wanting.
If we disregard the Malagasy sub-region, there are scarcely forty
species of land operculates on the whole African continent.
 The Ethiopian region may be divided into three sub-regions: (1)
the Central African; (2) the South African; (3) the Malagasy.
(1) The Central African Sub-region is bounded on the north by
the Great Desert, on the east and west by the ocean, and on the
south by a line roughly drawn between the mouth of the Orange
River and Delagoa Bay; it also includes S. Arabia. No natural
features exist which tend to break up this vast district into areas of
independent zoological development. The absence of long and lofty
mountain ranges, the enormous size of the great river basins, and
the general uniformity of climate, equalise the conditions of life
throughout. It will be convenient to break the sub-region up into
provinces, but in most cases no precise line of demarcation can be
laid down.
(a) The Senegambian Province may be regarded as extending
from the mouth of the Senegal River to Cape Palmas. Only 8 genera
of land Mollusca are known, including 4 Limicolaria and 3 Thapsia,
with 1 small Cyclophorus. Fresh-water genera are abundant, and
include most of the characteristic Ethiopian forms.
(b) The West African Province extends from Cape Palmas to the
mouth of the Congo, and is rich in Mollusca. The great Achatina,
largest of land snails, whose shell sometimes attains a length of 6½
in., Limicolaria, Perideris, and Pseudachatina are the characteristic
forms. The Agnatha are represented by Ennea, Streptaxis, and
Streptostele. Rachis and Pachnodus, sub-genera of Buliminus, occur
also on the east coast. A special feature is the development of
several peculiar slug-like genera, e.g. Oopelta, perhaps a form of
Arion; Estria, a slug with an external shell, akin to Parmacella; and
Aspidelus, a form intermediate between Helicarion and Limax.
Claviger, a handsome group akin to Cerithium, is peculiar to the
estuaries of West African rivers.
About sixteen species are known from the Cameroons District, but
no peculiar genera occur. The French Congo District has not yet
been well explored. Tomostele, a genus allied to Streptostele, is
peculiar, and Pseudachatina attains its maximum.
 Fig. 217.—Columna
flammea Müll., Princes I.
St. Thomas and Princes Is., in the Gulf of Guinea, are well known.
Princes I. has 22 species, 14 peculiar, and 2 common to St. Thomas
only, one of the latter being the great sinistral Achatina bicarinata
Chem. The remarkable genus Columna (Fig. 217) is peculiar, and
Streptostele (4 sp.) attains its maximum. Peculiar to St. Thomas are
Pyrgina, a turreted form of Stenogyra; Thyrophorella, a sinistral form
of Zonites; and Atopocochlis, a large bulimoid shell, whose true
relationships are not yet known. Homorus, a group of Achatina with
an elongated spire, occurring also in the Angola District and on the
east coast, has 4 species. No fresh-water species have as yet been
discovered in either of the islands.
The Angola and Benguela District, extending from the Congo to
the Cunene R., probably belongs to the West African Sub-region, but
until its fauna is better known it is advisable to consider it apart.
Achatina continues abundant, but the other characteristic West
African forms (Pseudachatina, Streptostele, Perideris) diminish or
are absent altogether. No Helix and only 1 Cyclophorus occur.
Ovampo, Damara, and Great Namaqualand, lying between the
Cumene and Orange rivers, seem to form a transition district
between the West and South African faunas. Helix reappears, while
the characteristic West African genera are almost entirely wanting.
 (c) The East African Province extends from about Delagoa Bay to
the Abyssinian shores of the Red Sea. In general out-line the
province consists of a flat marshy district, extending inland for many
miles from the sea; this is succeeded by rising ground, which
eventually becomes a high table-land, often desolate and arid,
whose line of slope lies parallel to the trend of the coast. The
Mollusca are little known, and have only been studied in isolated
districts, usually from the discoveries of exploring expeditions.
The Mozambique District, from Delagoa Bay to Cape Delgado,
includes no genus which does not occur on the west coast, except
Cyclostoma (2 sp.). Trochonanina (4 sp.), Urocyclus, a characteristic
African slug (2 sp.), Rachis (6 sp.), Pachnodus (2 sp.), and Achatina
(5 sp.), are the principal groups.

Fig. 218.—Urocyclus comorensis Fisch., Comoro Is.:

G, Generative orifice; M, mucous gland; O,
orifice leading to internal shell; P, pulmonary
orifice; T, tentacles. (After Fischer.)
The Zanzibar District, from Cape Delgado to the Somali country,
has the same general features. Meladomus, a large sinistral
Ampullaria, is characteristic, while Cyclostoma (5 sp.) becomes more
abundant. Helix is still absent, but the carnivorous forms (Streptaxis
2 sp., Ennea 7 sp.) are rather numerous.
The Somali District is characterised by operculate groups of the
Otopoma type (Georgia, Rochebrunia, Revoilia) whose generic value
is rather doubtful. Petraeus, in an Arabian type, supplants Rachis
and Pachnodus. Achatina is nearly wanting, but Limicolaria has 9
species. A few Helix, said to be of the Pisana group, occur.
 The District between the Great Lakes and the coast region is fairly
well known through recent explorations, especially those associated
with Emin Pasha. Streptaxis (6 sp.) and Ennea (24 sp.) are
numerous, Helix is wanting, and the Naninidae are represented by
Trochonanina (7 sp.), and other forms at present grouped under
Nanina or Hyalinia. On the high ground Buliminus, Cerastus, and
Hapalus replace, to some extent, the Achatina and Limicolaria of the
marshy plains. Land operculates (Cyclophorus 1, Cyclostoma 8) are
more numerous; among fresh-water genera we have Lanistes (5
sp.), Cleopatra (3 sp.), Meladomus (1 sp.), and Leroya, a sinistral
form with the facies of a Littorina. The characteristic African bivalves
(Mutela, Spatha, etc.) are few in number.
(d) Province of the Great Lakes.—The Mollusca of the four great
lakes of Eastern Central Africa—Lakes Albert Nyanza (Luta Nzige,
2720 ft.), Victoria Nyanza (Oukéréwé, 3700 ft.), Nyassa (1520 ft.),
and Tanganyika (2800 ft.)—are well known, and supply an
interesting problem in distribution. Those of the three first mentioned
lakes differ in no way from the rest of tropical Africa, but the Mollusca
of Tanganyika include, in addition to the ordinary African element, a
number of peculiar operculate genera, belonging principally to the
Melaniidae and Hydrobiidae. Several of these possess a solidity of
form and compactness of structure which is unusual in fresh-water
genera, and has led to the belief, among some authorities, that they
are the direct descendants of marine species, and that Tanganyika
represents an ancient marine area. This view appears untenable.
The Victoria Nyanza and Nyassa are part of the same system as
Tanganyika, and it is not easy to see how, if Tanganyika were once
an arm of the sea, they were not equally so, especially as they are
several hundred miles nearer the Indian Ocean as at present
defined. Nor, as will be seen from the figures given above, is there
anything in the altitudes which would make us expect anything
exceptional in Tanganyika. The similar case of L. Baikal must be
compared (p. 290), where again a number of specialised forms of
Hydrobia occur.
Of the genera concerned, Paramelania and Nassopsis are forms
of Melaniidae; Tiphobia (Fig. 219), which is allied to Paludomus, is a
compact shell with angulated spinose whorls; Lacunopsis,
Ponsonbya, Limnotrochus, and Tanganyicia are probably forms of
Lithoglyphus, some, as their names denote, being of decidedly
marine facies; Syrnolopsis and Turbonilla (?) look like
Pyramidellidae, Horea and Reymondia like Rissoina; Bourguignatia
appears to belong to Vivipara, with which has now been merged the
genus Neothauma. Recently discovered forms from the adjacent L.
Mweru are evidently of kindred origin.
(e) The Afro-Arabian Province includes Abyssinia, with S. Arabia,
the African shores of the Gulf of Aden, and Socotra. The province
contains a singular mixture of types. The high ground of Abyssinia
stands like a lofty European island in the midst of a tropical plain,
with Palaearctic genera flourishing like hardy northern plants on a
mountain in low latitudes. Helix, Vitrina, and Pupa abound, with a
few Clausilia and even a Limax. On the lower levels occur
Limicolaria (3 sp.), Subulina (7 sp.), Helicarion, and Homorus, but
land operculates are entirely wanting. Characteristic of the province
as a whole are various forms of Buliminus, which in Socotra are
represented by two peculiar sub-genera, Achatinelloides and
Passamaiella. In S. Arabia the mixture of types produces curious
results: the Helix, Clausilia, and Vitrina being Palaearctic, the
Limicolaria and all the operculates Ethiopian, while the single
Trochomorpha is Indian. Indian influence, indeed, comes out
unmistakably throughout the province. Thus in Socotra there are two
Cyclotopsis, in Abyssinia two Africarion (closely related to the Indian
Girasia), two Microcystis, and a Glessula, and in the Scioa district
there is a Sitala. The fresh-water Mollusca of Socotra are Indian
forms.
 Fig. 219.—Tiphobia Horei E.
A. Smith, L. Tanganyika.

Fig. 220.—Mollusca
characteristic of L.
Tanganyika: A, Nassopsis
nassa Woodw.; B, Spekia
zonata Woodw.; C,
Syrnolopsis lacustris E. A.
Smith.
 Fig. 221.—Achatina zebra
Lam., S. Africa. × ½.
(2) The South African Sub-region.—The principal characteristic
of the Mollusca of S. Africa is the occurrence of numerous small
species of Helicidae, belonging chiefly to the groups Pella, Phasis,
Dorcasia, and Sculptaria, all of which are practically peculiar.
Carnivorous genera are also prominent, Ennea here attaining its
maximum. Rhytida (to which several species still regarded as Pella
belong) is common only to the S. Pacific and Australasia, and forms,
with Isidora among the fresh-water pulmonates, a remarkable link of
connexion. Aerope, the largest of all helicoid carnivorous genera,
and Chlamydephorus, a carnivorous slug with an internal shell, are
peculiar. Achatina is still abundant, but Limicolaria is wanting.
Livinhacea, a form with a continuous peristome, perhaps akin to
Bulimus; Apera, a form of slug; and Coeliaxis, a genus perhaps akin
to the Papuan and Queensland Perrieria, are all peculiar. The land
operculates, which are not numerous, are of the East African type.
Land Mollusca of the S. African Sub-region
Chlamydephorus 1
Ennea 31
Aerope 5
Rhytida 3
Helicarion 3
Trochonanina 1
Trochozonites 1
Limax 1
Apera 1
 Vitrina 7
Nanina 6
Conulus 2
Patula 2
Pella 44
Dorcasia 8
Phasis 1
Sculptaria 2
Helix (inc. sed.) 4
Rachis 1
Pachnodus 3
Buliminus (?) 4
Pupa 20
Vertigo 2
Achatina 18
Livinhacea 1
Stenogyra 4
Coeliaxis 1
Succinea 3
Vaginula 2
Cyclophorus 1
Cyclostoma 7
Cyclotus (?) 1
Blanfordia 1
St. Helena.—The Molluscan fauna of St. Helena is perhaps the
most puzzling, as regards its geographical affinities, of any in the
world. It consists of 29 peculiar species of land Mollusca (fresh-water
species being unknown), 19 of which are recently extinct, partly
owing to the destruction of the forest, but are found in considerable
abundance in a state of good preservation.[375] The genera are—
Hyalinia 1
Patula 4 (3 extinct)
Endodonta 10 (7 extinct)
Bulimulus 7 (5 extinct)
Pachyotus 1 (extinct)
Tomigerus (?) 1 (extinct)
Pupa 2 (extinct)
Succinea 3
The 5 genera which concentrate our attention are Patula,
Endodonta, Pachyotus (Fig. 222), Tomigerus, and Bulimulus, all of
which appear utterly strange to an oceanic island in the middle of the
S. Atlantic. Patula and Endodonta are essentially Polynesian forms,
occurring abundantly on all the island groups in the Central Pacific.
Pachyotus, Tomigerus (assuming its correct identification), and
Bulimulus are all S. American forms, the two former being especially
characteristic of Brazil. How this mixture of genera now confined to
regions so widely distant, not only from St. Helena itself, but from
one another, became associated here, is a problem obviously not
easy of solution. The fauna is probably a remnant of a very ancient
type, possibly at one time much more widely distributed. Endodonta
(an essentially insular form, like Omphalotropis) actually occurs on
Fernando Noronha, off the Brazil coast, and we shall see how an
Indian and even a Polynesian element is present off the eastern
coasts of Africa.
Ascension I.—One indigenous species, a so-called Limax, is all
that has ever been discovered.
(3) The Malagasy Sub-region includes Madagascar with its
attendant satellites Bourbon, Mauritius, and Rodriguez, and the
Seychelles and Comoro groups. No land Mollusca are known from
the Amirantes, the Chagos, or from Aldabra. The special
characteristics of the sub-region are the great development of the
carnivorous land Mollusca (Ennea, Gibbus), the occurrence of a
considerable number of true Helicidae of great size and beauty, and
the prominence of the genus Cyclostoma.
(a) The Madagascan Province.—The land Mollusca of
Madagascar, although as yet imperfectly known, possess a striking
individuality. Two of the chief characteristics of the Ethiopian region
are the paucity of its land operculate and of its Helix fauna;
Madagascar is especially distinguished by the rich development of
both these groups. For size, colouring, and beauty of shape, the
Helicidae of the two sub-genera Ampelita and Helicophanta rival, if
they do not surpass, any in the world. They are quite peculiar to this
sub-region, not a trace of them occurring on the Mascarenes,
Seychelles, or even on the Comoros. Helicophanta is distinguished
by the enormous size of its embryonic shell, which persists in the
adult (Fig. 223), and in this respect the group appears to be related
to Acavus (Ceylon, Fig. 204) and Panda (N.E. Australia). As is usual
when Helix is well developed, Nanina (about 12 sp.) is
proportionately scanty.
The African Bulimini (Pachnodus and Rachis) are represented by
two species, but Achatina, so abundant on the mainland, is scarce.
Two other groups of Buliminus, Leucotaenia and Clavator, are
peculiar. The presence of a single Kaliella, specifically identical with
a common Indian form, is very remarkable.
Cyclostoma proper, of which Madagascar is the metropolis, is
richly developed (54 sp.). Many of the species are of great size and
of striking beauty of ornamentation. Unlike its Helicidae, this genus is
not restricted to Madagascar; several species occur on the mainland,
6 on the Comoros, one on the Seychelles, and 16 in Mauritius. The
sub-genera Acroptychia and Hainesia are peculiar.

Fig. 222.—Pachyotus auris

vulpina Desh., St. Helena (sub-
fossil).
 Fig. 223.—Helix (Helicophanta)
Souverbiana Fisch.,
Madagascar, showing
embryonic shell. × ⅔.

Fig. 224.—Cyclostoma
campanulatum Pfr., Madagascar.
The fresh-water Mollusca of Madagascar contain further traces of
Indian relationship. Thus we find two species of Paludomus, a genus
whose metropolis is Ceylon, India, and Further India, and which is
barely represented on the Seychelles and in the Somali district.
Melanatria, which is peculiar to Madagascar, has its nearest affinities
in the Cingalese and East Indian faunas. Several of the Melania and
the two Bithynia are of a type entirely wanting in Africa, but common
in the Indo-Malay sub-region. Not a single one of the characteristic
African fresh-water bivalves (Mutela, Spatha, Aetheria, Galatea, etc.)
has been found in Madagascar. On the other hand, certain African
Gasteropoda, such as Cleopatra and Isidora, occur, indicating, in
common with the land Mollusca, that an ultimate land connexion with
Africa must have taken place, but at an immeasurably remote period.
Land and Fresh-water Mollusca of Madagascar
Ennea 9
Urocyclus 2
Helicarion (?) 1
Macrocyclis (?) 1
Kaliella 1
Nanina (inc. sed.) 9
Ampelita 35
Helicophanta 17
Pachnodus 2
Rachis 2
Leucotaenia 2
Clavator 2
Achatina 3
Opeas 2
Subulina 3
Vaginula 4
Limnea 2
Planorbis 3
Isidora 3
Melania 7
Melanatria 4
Paludomus 2
Vivipara 1
Bithynia 2
Cleopatra 2
Ampullaria 6
Cyclophorus 2
Cyclotus (?) 1
Cyclostoma 54
Otopoma 5
Lithidion 1
Acroptychia 3
Hainesia 3
Unio 1
Corbicula 2
Sphaerium 1
Pisidium 1
 The Comoro Islands.—This isolated group possesses about 100
species, almost all of which are peculiar. The principal feature is the
rich development of Ennea (30 sp.). On the whole the group shows
more relationship to Madagascar than to the mainland. Thus we
have six species of true Cyclostoma, and only one Achatina, while
among the fresh-water genera is Septaria, which is characteristic of
the whole Malagasy Sub-region, but is absent from the mainland.
The Helicidae are all of insignificant size. Peculiar to the group is the
remarkable genus Cyclosurus (Fig. 152, p. 247).
(b) The Mascarene Province (Mauritius, Bourbon, Rodriguez, and
the Seychelles).—The percentage of peculiar species, which is very
high, can only be paralleled in the case of some of the West Indian
islands, and sufficiently attests the extreme isolation of the group
from Madagascar. We have—
Total sp. Land sp. Fresh-water Peculiar Peculiar to
sp. group
Mauritius 113 104 9 78 102 (90 p.c.)
Bourbon 45 40 5 19 38 (84 p.c.)
Rodriguez 23 19 4 15 21 (95 p.c.)
Seychelles 34 27 7 24 30 (90 p.c.)
The Mollusca of the group exhibit three distinct elements, the
Indigenous, the Madagascan, and the Indian and Australasian.
The genus Pachystyla (Naninidae) is quite peculiar, forming the
main portion of the land snails proper. It attains its maximum in
Mauritius (17 sp.), with 5 sp. in Bourbon and one sub-fossil sp. in
Rodriguez, while in the Seychelles it is absent. But the principal
feature of the Mascarene group is the extraordinary development of
the carnivorous genus Gibbus, which has 27 sp. in Mauritius, 8 in
Bourbon, 4 in Rodriguez; in the Seychelles, it is replaced by
Edentulina and Streptostele. The principal link with Madagascar is
found in a part of the operculate land fauna. Cyclostoma is present
(with Otopoma) in several fine living forms, and the number of sub-
fossil species is a clear indication that this group was, not long ago,
much more abundant, for of the 16 Cyclostoma known from
Mauritius 10 are sub-fossil. The operculates form a decided feature
of the land fauna; thus in Mauritius there are 32 species, or more
than 28 per cent of the whole.

Fig. 225.—Characteristic Mauritian land

shells: A, Gibbus palanga Fér.; A´,
young of same; B, Gibbus
lyonetianus Pall.
Indian and Australasian affinities are unmistakably present. Thus
Omphalotropis, a genus characteristic of small islands, is profusely
represented, but it does not occur in Madagascar or Africa. Two
Helicina (Mauritius and Seychelles) and a single Leptopoma
(possibly a Leptopomoides) are also of eastern relationship.
Cyclotopsis, Cyathopoma, and Geostilbia are markedly Indian
genera. Microcystis, Patula, and Tornatellina are Polynesian.
Hyalimax—and this is a very striking fact—occurs nowhere else but
in the Andamans and Nicobars, and on the Aracan coast. The
nearest relation to the Seychelles Mariaella appears to be the
Cingalese Tennentia. Not a single representative of these eleven
genera has been found even in Madagascar.
The fresh-water Mollusca (omitting the Neritidae) are: Mauritius 9
species, Bourbon 5, Rodriguez 4, Seychelles 6, with only 15 species
in all. The one Planorbis and the Vivipara, the Paludomus and two of
the Melania are of Indian types. The Lantzia (peculiar to Bourbon) is
probably allied to the Indian Camptonyx. Owing to the paucity of
permanent streams, no fresh-water bivalves occur. Among the
Neritidae is a single Septaria, a genus which, though occurring in
Madagascar, is entirely strange to Africa, and is abundant in the
Oriental and Australasian regions.
It would seem probable that when the closer connexion which at
one time undoubtedly existed between India and Eastern Africa
began to be less continuous,[376] the Mascarene group was first
severed from what ultimately became Madagascar, while the
Seychelles, and perhaps the Comoros, still continued united to it.
The Comoros, which lack the great Helices, separated off from
Madagascar first, while the Seychelles continued in more or less
direct union with that island sufficiently long to receive the
progenitors of Stylodonta (a peculiar group of Helix), but became
disunited at an exceedingly remote period.

E. The Nearctic Region

The southern boundary of this region may be regarded as roughly

corresponding to that of the United States, i.e. Lower California and
Mexico are excluded. The southern portion of Florida belongs to the
Antillean sub-region.
The principal characteristic of the Nearctic Region is the
remarkable poverty of its land Mollusca. No district in the world of
equal extent is so poor in genera, while those which occur are
generally of small size, with scarcely anything remarkable either in
colouring or form. The elongated land shells (Clausilia, Buliminus),
so characteristic of Europe, are entirely wanting, but a few
Bulimulus, of Neotropical origin, penetrate Texas, and from the same
sources come a few species of Glandina (as far north as S.
Carolina), Holospira (Texas), and Helicina.
The region falls into two well-marked sub-regions, the N.
American and the Californian, with the Rocky Mountain district as a
sort of debatable ground between them. The Californian sub-region
consists of the narrow strip of country between the Sierra Nevada,
the Cascade Mountains and the coast-line, from San Diego to
Alaska; the N. American sub-region consists of the remainder of the
region.
(1) The N. American Sub-region.—The Carnivorous genera are
represented solely by the few Glandina mentioned above, and by the
indigenous genus Selenites, a form midway between Testacella and
Limax, whose metropolis is on the Pacific slope, but which spreads
eastward into the Antilles. Among the Limacidae, Limax is common
to both sub-regions, but Tebennophorus (4 sp., 3 of which belong to
the genus Pallifera), a genus found also in China and Siam, and
Vitrinozonites do not occur in the Californian. Hyalinia (Zonites) is
fairly abundant, especially in the groups Mesomphix and
Gastrodonta (peculiar to this sub-region), and Hyalinia proper. Patula
is well represented. The Helicidae belong principally to the groups
Mesodon, Stenotrema, Triodopsis, Polygyra, and Strobila, only 6 of
which, out of a total of 84, reach the Pacific slope. Land operculates
are conspicuous for their almost complete absence (see map,
frontispiece).

Fig. 226.—Characteristic North American Mollusca. A,

Helix (Mesodon) palliata Say, Ohio. B, Helix
(Polygyra) cereolus Mühlf., Texas. C, Patula
alternata Say, Tennessee.
The poverty of the land fauna is atoned for by the extraordinary
abundance and variety of the fresh-water genera. A family of
operculates, the Pleuroceridae, with 10 genera and about 450
species, is quite peculiar, a few stragglers only reaching Central
America and the Antilles. The nucleus of their distribution is the
Upper Tennessee River with its branches, and the Coosa River.
They appear to dislike the neighbourhood of the sea, and are never