CHAPTER VIII.

THE ALPINE FAUNA.

We are told by Sir Archibald Geikie (p. 851) that "from the Pyrenees

eastwards, through the Alps and Apennines into Greece, and the southern

side of the Mediterranean basin, through the Carpathian Mountains and

the Balkan into Asia Minor, and thence through Persia and the heart of

Asia to the shores of China and Japan, a series of massive limestones

has been traced, which, from the abundance of their characteristic

foraminifera, have been called the Nummulitic Limestone. Unlike the

thin, soft, modern-looking, undisturbed beds of the Anglo-Parisian area,

these limestones attain a depth of sometimes several thousand feet of

hard, compact, sometimes crystalline rock, passing even into marble, and

they have been folded and fractured on such a colossal scale that their

strata have been heaved up into lofty mountain crests sometimes 10,000,

and in the Himalaya range more than 16,000 feet above the sea." "Nowhere

in Europe," continues the same author (p. 860), "do oligocene strata

play so important a part in the scenery of the land, or present on the

whole so interesting and full a picture of the state of Europe when

they were deposited, as in Switzerland. Rising into massive mountains,

as in the well-known Rigi and Rossberg, they attain a thickness of more

than 6000 feet." "By far the larger portion of these strata is of

lacustrine origin. They must have been formed in a large lake, the area

of which probably underwent gradual subsidence during the period of

deposition, until in Miocene times the sea once more overflowed the

area."

From these remarks by our most eminent British geologist, we gather that

in early Tertiary times much of the present area of Switzerland was

either a sea or a large freshwater lake. The Alps were then appearing in

this sea, probably as a chain of islands, and in the beginning of the

Miocene Epoch one large elongated island had made its appearance--the

future European Alps. I have already mentioned that the Miocene Sea

skirted the Alps from the Mediterranean up the valley of the Rhone and

along its northern and eastern margin. Miocene marine deposits are also

known from the Southern Alps and the east side of the Apennines, from

Corsica, Sardinia, and Malta. No trace, however, of them has been

noticed anywhere along the AEgean Sea or on the Balkan peninsula. The

Alps were therefore connected to the east with the outliers of the

Balkan Mountains, and in this way with Asia, from which they received so

large a proportion of their fauna and flora. In pliocene times the sea

still washed the southern shore of the Alps, but to the north dry land

gradually supplemented the sea, and the Alpine fauna and flora were

able to pour into the plain. It was then that the Arctic species--which

we have learned had migrated into Northern Europe from the north--found

their way to the Alps. In a similar way Lusitanian forms--in fact,

species from almost all parts of Europe--were now free to wander to the

newly opened up peninsula which had become part of the mainland of

Europe. The typical Siberian species had not entered our continent at

that time, it was not till much later--not until the middle of the

Pleistocene Epoch--that they made their appearance at the foot of the

Alps, but, as we shall see later on, it is doubtful whether many of

these species ever reached the mountains.

The fauna of the Alps, and also the flora, is therefore made up of a

number of component elements. In the first place we have the Oriental

element--the migrants from Central and Southern Asia. When the nature

and origin of the Oriental fauna in Europe was discussed, reference was

made to the fact (p. 272) that we can distinguish an older from a newer

Oriental migration. Both of these have entered the Alps. As we might

anticipate, many of the older Oriental migrants have developed into new

species, laying the foundation of an indigenous Alpine element. From the

fact that they set foot on the Alpine peninsula, it might be expected

that there could have existed no mountains to speak of. The climate was

mild and damp. Now as the country rose, and a formidable mountain range

took the place of a hilly island, the whole fauna was lifted up and

transferred to entirely different conditions. A modification of their

structure to suit the new surroundings was therefore to be anticipated,

and that is exactly what occurred, though not in all cases.

Take, for example, the goats which are of Asiatic origin. Every one has

heard of the "Steinbock,"--the Alpine mountain goat (_Capra

ibex_)--though very few have seen it in its native haunts, where it is

now on the verge of extinction. A closely allied species (_Capra

sibirica_) inhabits the Altai and Himalayan Mountains; a third species

(_Capra sinaitica_) lives in Palestine, and has entered Egypt by way of

the Sinaitic peninsula. Another (_C. aegagrus_) occurs in Asia Minor,

Persia, the island of Crete, and some of the Cyclades. This exemplifies

what I remarked in the last chapter about the former land-connection

between Greece and the Asiatic continent. Finally, we have the Pyrenean

Goat (_Capra pyrenaica_), which is found in the Pyrenees, the higher

ranges of Central Spain, in Andalusia, and Portugal, thus indicating

that it probably reached the Spanish peninsula from the south by means

of the old Sicilo-Algerian highway, especially as remains of the species

occur in the cave deposits of Gibraltar. The ancestors of the goat-like

Antelope--known as the Chamois (_Rupicapra tragus_)--no doubt also came

from Asia. The genus is not represented there, but _Nemorhoedus_ and

_Budorcas_ are allied Asiatic genera, while the Rocky Mountain Goat

(_Haploceros montanus_) also has certain affinities with the Chamois.

Besides the Alps, the latter occurs in the Caucasus and the Pyrenees.

The Alpine Marmot (_Arctomys marmotta_) is sometimes quoted as owing its

origin to the Siberian pleistocene migration, but it does not occur in

Siberia now, nor is there any palaeontological evidence that it was ever

found there. The genus _Arctomys_ is an ancient Asiatic genus, to judge

from its general range. Only two species occur in Europe, one of which,

the true Siberian Marmot (_A. bobac_), just enters our continent in the

east--or rather, it is one of those species which came to us in

pleistocene times and are now gradually retreating towards their native

land. The genus, however, is probably not of Siberian origin. No less

than seven other species occur in Asia, six of which are confined to

Central Asia and the Himalayan Mountains, while four have wandered to

North America. The sequence of events, therefore, was that the ancestor

of _Arctomys marmotta_ probably came to the Alps direct from Central

Asia by way of Asia Minor in miocene or pliocene times. It has since

become modified into a distinct species, and has spread to the European

plain, where it occurs fossil in pleistocene strata, and to the

Carpathian Mountains and the Pyrenees.

The great majority of species of the large genus _Microtus_ (_Arvicola_)

are Asiatic, and there can be little doubt that it has originated in

that continent. There is one species of Vole (_Microtus nivalis_) which

occurs in the high Alps, and which has been supposed to be a typical

Alpine form. It is known, however, to occur also in North Italy and in

Bohemia, while _Microtus leucurus_ of the Pyrenees is identical with

this species. But its range is by no means confined to Europe, for it

has also been discovered in Syria and Palestine, while a closely allied

form exists in the Himalayan Mountains. This shows clearly that the

species has migrated to the Alps from Asia Minor. That this migration

may have taken place at an early period--at a time when Sardinia and

Corsica were still connected with Southern Europe--is indicated by the

occurrence of an extinct Vole (_Microtus brecciensis_) in Sardinian and

Corsican pleistocene (?) deposits.

All the Alpine species mentioned except the Chamois can be easily traced

to their former Asiatic home. But even it has its nearest relations in

Asia. I might also refer to another Vole (_Evotomys Nageri_) which is

practically confined to the Alps and Northern Italy, and which has

probably originated there, though most of its nearest relations are

either Asiatic or North American species.

But besides these Asiatic immigrants and their modified descendants we

have a small truly native Alpine mammalian fauna. _Sorex alpinus_--the

Alpine Shrew--occurs only in the Alps, the Harz Mountains, Pyrenees, and

Carpathians. The genus has been found in European eocene strata,--in

vastly older deposits in our own continent than elsewhere,--so that it

is extremely probable that it has originated there. It may then have

developed a new centre of distribution in the newly-formed Alps where

both _Sorex alpinus_ and _S. minutus_ (_pygmaeus_) have their home. From

there they again spread--perhaps already in miocene times--to Asia and

North America, where a large number of new species originated. It seems

to me even probable that one of these Asiatic species of _Sorex_, viz.

_S. araneus_ (_vulgaris_), subsequently migrated towards the old home of

its forefathers, since we find it more or less confined to Central and

Northern Asia and Northern Europe.

Though the origin of the Alpine Hare has already been referred to and

fully discussed in a previous chapter (p. 148), the conclusions arrived

at may be once more repeated. The Alpine Hare (_Lepus variabilis_) is of

Arctic origin. It spread southward into Europe, North America, and Asia

in early glacial times, and reached our continent from Spitsbergen by

means of a direct land-connection with Lapland. The Scandinavian

peninsula was then separated from Russia, but connected with Scotland

and Ireland (Fig. 13, p. 170). Since England was then united to France,

the Alpine Hare was able to invade western continental Europe and all

the mountain ranges. Its range is very discontinuous, small colonies

being scattered all over the mountainous parts of the Northern

Hemisphere, while the European Hare--a closely allied species--occurs in

the plain, and now occupies to some extent the former haunts of the

Alpine Hare (cf. Fig. 8, p. 137). Might not the European Hare, as

suggested, possess some advantages which enabled it to drive the other

into more inaccessible parts, thus producing the peculiarity of range?

The present distribution of the Alpine and the European Hare (_L.

Europaeus_) appears to me to strongly support such an assumption. It is

not the cold which has driven the Alpine Hare to the Alps; and its

presence there is not, as is often supposed, a "_standing testimony of a

former arctic climate_" in Europe, but merely the necessary consequence

of the weaker species being thrust into less accessible regions by a

stronger rival.

_Muscardinus avellanarius_,--the common Dormouse,--though by no means

confined to the Alps, has probably originated there. It is found up to a

height of nearly 5000 feet in these mountains, and is spread over Europe

at nearly equal distances from the Alps in all directions. Being absent

from Ireland, Scotland, Norway, and Northern Russia, it seems as if it

had only diffused northward in more recent times.

The closely allied genus _Myoxus_ is likewise of European extraction,

some species being known from French eocene deposits.

There are only a few typically Alpine Birds. One of these is the Alpine

Accentor (_Accentor collaris_), which on rare occasions visits England,

and Northern Europe generally. It is, however, by no means peculiar to

the European Alps; a variety of this species occurs in Central Asia,

Eastern Siberia, and Japan. The only other Accentor inhabiting our

continent is the Hedge Accentor (_A. modularis_), which is resident over

the greater part of it, and also in North Africa and the Mediterranean

Islands. It also extends its range across the AEgean Sea to Asia Minor,

so that really not a single Accentor is peculiar to Europe.

Both the European species are evidently old forms, and the genus, as

might be expected, is certainly Asiatic. No less than ten other species

of Accentor are known, all of which are confined to Central Asia and the

Himalayan Mountains, and are therefore all Holarctic. I may mention that

much difference of opinion still exists as to the true zoological

position of this anomalous genus. It has been located in several

different families by various ornithologists, but has not yet found a

permanent resting-place. Another bird generally considered to be

peculiar to Switzerland is the Alpine Chough (_Pyrrhocorax alpinus_),

but its range extends across Asia Minor to the Himalayas. Whether the

European Chough should not form a distinct genus is a matter of opinion.

Some of our leading ornithologists, like Dr. B. Sharpe, are inclined to

separate it from _Pyrrhocorax_; however, there is no doubt that it is

closely related to the Alpine Chough, whatever view we may take of the

generic distinctness. It inhabits principally Western and Southern

Europe, also North Africa; and its range extends eastward to the

Himalayas, China, and Eastern Siberia. If any doubt still existed as to

the Asiatic origin of the Choughs, it may be noted that the only two

other closely allied genera, viz., _Corcorax_ and _Podoces_, live in

Australia and Central Asia respectively.

There are two other birds to which I should like to refer. These are the

Rock Sparrow and the Alpine Snow Finch. The first of these (_Petronia

stulta_) is by no means peculiar to the Alps. It is the only species of

the genus inhabiting Europe; and besides the Alps it occurs in Southern

Europe generally, and ranges as far west as the Canaries and Madeira.

Eastward it is not found beyond Central Asia. Of the remaining five

species of _Petronia_, two occur in Asia (including India) and three in

Africa. Whether the genus is African or Asiatic is immaterial for our

purpose, since, in any case, the only European species came to us from

the east with the Oriental migration. The distribution of the Alpine

Snow Finch (_Montifringilla nivalis_) is very similar to that of the

birds we have just been considering. It inhabits the Alps up to a great

height, but occurs also on the Pyrenees and other South European

mountain ranges as far east as Palestine, where again it is found in the

Lebanon. The genus _Montifringilla_ has seventeen other species. Twelve

of these live in Central Asia and Japan, extending as far north as

Kamtchatka, while five inhabit Western North America right down to

Mexico. There is every probability that in this case also we have to

deal with an Asiatic genus which spread eastward to America, and

westward to Europe.

As regards the Reptiles, there are _no_ peculiar Alpine forms, but among

the Amphibia some species deserve to be mentioned. Up to an elevation of

10,000 feet we find in the Alps the Black Salamander (_Salamandra

atra_); and it is apparently quite peculiar to them, never having been

observed in the plains. The handsome black and yellow Salamander

(_Salamandra maculosa_)--so well known as a terrarium specimen--likewise

occurs in the Alps, and it has besides a fairly wide distribution in

Europe. It is known from Southern Germany, the Pyrenees, Spain,

Portugal, Sardinia, Corsica, Greece, Syria, and Algiers. A third species

(_S. caucasica_) inhabits the Caucasus. The evidence of distribution

here points emphatically to an Alpine origin of the genus _Salamandra_.

We cannot tell where the ancestors of _Salamandra_ may have come from,

but several other genera of _Salamandridae_ are certainly Asiatic. Our

common Newt (_Molge vulgaris_) belongs to a genus with nineteen species,

several of which are peculiar to Europe. The general range of the genus,

however, extends to North America, and it is more probable therefore

that it originated in Asia. If so, it certainly must have passed into

Europe at a very early date. Let us assume the first _Molges_ to have

traversed the AEgean Sea on _terra firma_ to Greece in miocene times,

they might thus have been able to travel straight on to the old

Tyrrhenian continent of which Corsica and Sardinia now form the

remains, and also on to North-west Africa. Indeed, we find high up in

the Corsican mountains an interesting large brownish-grey Newt (_Molge

montana_), and another in Sardinia (_Molge Rusconii_). Again, in Algiers

there are two species, viz., _Molge Poireti_ and _M. Hagenmuelleri_,

while the Moroccan _M. Waltlii_ passes into the south of Spain. Here

_Molge boscae_, _M. aspera_, and _M. marmorata_ originated, the latter

passing into France.

Another branch of the _Molge_ tribe turned northward from Greece towards

the newly forming Alps; and there originated _Molge alpestris_ and _M.

palmata_, which more recently have spread into England (one at least),

Germany, France, Austria, and Southern Italy. _Molge vulgaris_ is an

Asiatic species which wandered northward after entering Europe, covering

a large area, but never reached the extreme south or south-west. _M.

cristata_--the large Water Newt--has a similar but not quite so extended

a range, while _M. vittata_ never managed to cross the borders of Asia

Minor. Some of the other species occur in China, Japan, and North

America.

None of the tailless Batrachians--the Frogs and Toads--are peculiar to

the Alps, but one, viz. _Rana temporaria_, ascends to the height of no

less than 10,000 feet. It is our common British Frog. No other Frog

probably ranges so far north or to such heights.

Let us now inquire what the invertebrate fauna of the Alps teaches us.

We are told by Dr. Kobelt, the great authority on European land shells,

that a uniformity of character marks the Alpine Molluscan fauna (_b_,

i., p. 251). One of the characteristic genera _Campylaea_--often looked

upon as a sub-genus of _Helix_--is a group containing somewhat flattened

conspicuous snails of large size. These are found everywhere in the

Alps, and wherever they occur beyond the confines of these mountains,

remarks Dr. Kobelt, their origin from the main stock is easily traced.

They have been gathered in the Apennines in Sicily, and even beyond the

Mediterranean in Algeria. On the Balkan peninsula they occur right down

to the most southern point of Greece, but are not met with either in

Crete or Asia Minor. One species has been found sub-fossil in Thuringia

in Northern Germany.

Another truly Alpine genus, says Dr. Kobelt, is the operculate

_Pomatias_, which in its geographical distribution offers some

interesting modifications from that of _Campylaea_. Less limited to high

elevations, it has spread over a greater part of the plains. This has

happened especially in France, while in Germany one species advances

almost as far north as Heidelberg. In other directions also the genus

has travelled beyond the limits of range of _Campylaea_. _Pomatias_

occurs in the Pyrenees and Northern Spain, in Sardinia and Crete, and

may, according to the same author, be expected in Asia Minor, although

no species has as yet been met with there. In Greece, again, it has been

observed, and numerous species inhabit Tunis and Algeria. Dr. Kobelt

connects the wider range of _Pomatias_ with the geological history of

the genus (_b_, i., p. 253). He tells us that species of Pomatias have

been found in eocene deposits differing but little from our present

forms, while undoubted _Campylaeae_ are not met with till we reach the

upper Miocene.

_Zonites_ is, according to Dr. Kobelt, a third Alpine genus, whose range

scarcely differs from the other two (_b_, i., p. 254). The centre of

distribution lies at present in one of the branches of the most southern

Alpine chain which help to form a large portion of the Balkan peninsula.

The bulk of the species inhabit that peninsula, the Greek Islands

(except Crete) and Asia Minor. Neither in the Tyrol nor in Switzerland

do we find any _Zonites_, and the few species that do occur in the

south-eastern Alps only just cross the outliers of these mountains.

Between the south-western Alps and the Rhone we again find a

_Zonites_--a remarkable case of discontinuous distribution, since the

nearest other habitat of the genus is Monte Gargano in South-eastern

Italy, which is known to harbour a good many interesting geographical

puzzles.

We still have a good deal to learn as regards the molluscan fauna of

Sicily, Sardinia, and Corsica. These islands have scarcely been more

than skimmed by conchologists, and _Zonites_ may inhabit one or all of

these, which might indicate to us the manner in which this genus

travelled from Southern Italy to Provence in the south of France. The

distribution of _Zonites_ certainly does not seem to imply an Alpine

origin, because it is almost completely absent from the Alps proper.

But I do not think my views differ materially from those of Dr. Kobelt,

since the Alps, in the wide sense, include the mountains of the Balkan

peninsula, where I should feel inclined to locate the ancestral home of

the genus.

The small operculate genus _Acme_ is a similar case. Dr. Kobelt places

the centre of distribution on the southern slope of the Alps, but

scarcely any of the species inhabit the Alps proper. Some occur in

France, others in North Africa, Sicily, Southern Italy, and the

Caucasus. It is evidently a very ancient genus. The species live in moss

or underground, and are not likely to be transported across the sea by

accidental or occasional means of distribution.

Still another genus, which resembles _Acme_ in its geographical

distribution, is _Daudebardia_--a small slug-like mollusc with a tiny

shell. It does not, however, range nearly so far north or west as

_Acme_, for it occurs neither in the British Islands nor in Spain or the

Pyrenees.

I shall not be able to refer to more than a few of the most typical

Alpine species of Lepidoptera, but they may be taken as fair examples of

the geographical distribution of the rest of the group.

Even those visitors to Switzerland who do not claim to be naturalists

have heard of the remarkably handsome and stately Butterfly known as

Apollo. To the ardent entomologist, the first sight of this typical

Alpine species is a never-to-be-forgotten delight, and he generally

brings home with him a rich harvest of specimens. The more experienced

Butterfly hunter knows that there are no less than three different kinds

of Apollo--or, as we should say more correctly, of Parnassius--in

Switzerland. There is first the common Apollo (_Parnassius Apollo_),

then the rarer and more local _P. delius_, which inhabits more elevated

regions, and finally the still scarcer _P. mnemosyne_, which is only

known from the highest mountain ranges. It may be a surprise to those

who have accustomed themselves to connect Apollo with the Alps, and who

think the two belong together and cannot do without one another, to hear

that it is by no means confined to them. It is also found in

Scandinavia, France, Spain, Russia, and in Siberia. _Parnassius delius_

is confined to the European Alps and the mountains of Central Asia,

while _P. mnemosyne_ is known from the Pyrenees, Sweden, Hungary,

Sicily, Russia, and Western Asia. One other Parnassius inhabits Europe,

viz., _P. Nordmanni_ of the Caucasus, but all the remaining species of

the genus--and there are nearly thirty more--are confined to Central

Asia. A few, as we have seen, have reached Europe, some have travelled

to the Himalayan Mountains, and others to Western North America. The

centre of distribution is certainly in Central Asia, and we have no

reason to suppose that the original home in this case does not agree

with that centre.

_Melitaea_, a genus to which some of our British Fritillaries belong, has

also some typically Alpine members. Two of these, viz. _M. cynthia_ and

_M. asteria_, are peculiar to the Alps, the latter being only found at

considerable elevations. Most of the remaining fourteen European species

are also found in Central Asia. Thus the isolated _M. maturna_, which in

Europe is confined to Lapland, is also known from the Altai Mountains,

which again are near the centre of distribution, since some species of

_Melitaea_ range across the Northern Pacific to Western North America.

The small British Mountain Ringlet, and also the Scotch Argus, belong to

a genus of butterflies which is very characteristic of the European

Alps. But owing to its enormous geographical distribution, its probable

home is somewhat difficult to ascertain. Nevertheless it is a noteworthy

genus, especially so from the fact that the two British species _Erebia

epiphron_ and _E. aethiops_ are taken at first sight for true Arctic

migrants. As neither of them, however, occurs in Scandinavia, Greenland,

or Arctic America, this supposition must be abandoned. They must be

looked upon as species which once had a wider range in the southern

parts of the British Islands, and which have survived in a few isolated

localities, where they are apparently on the verge of extinction.

About sixty species of _Erebia_ are known to science, half of which are

found in Europe, the remainder in Siberia, the Himalayas, Arctic

America, Chili, Patagonia, South Africa, and Madagascar. Though a few do

range into these outlying regions of the earth, Central Asia seems to

lie near the centre of distribution of the genus, and the probability is

that it also was its original home. Most of the European species are

high Alpine forms--_E. glacialis_ being met with at a height of 10,000

feet--and these are generally quite peculiar to the Alps, showing that

their ancestors came from Asia at an early date, probably by way of Asia

Minor and Greece. A few, as for instance _E. lappona_, range right

across to the Altai Mountains from the Alps, and at least one--_E.

melas_--is found in Greece. _Erebia_ migrations seem therefore to have

taken place by the Southern or Oriental route at different geological

periods. But some of the European species which are more or less

confined to the plain, and are either absent from Switzerland or do not

reach the higher elevations, appear to me to have come by the more

direct northern or Siberian highway, at a still more recent period.

These are _Erebia aethiops_, _medusa_, _ligea_, and _ambla_.

Only one species of the well-known Polar genus _Oeneis_, viz. _Oe.

aello_ occurs in the Alps. It has always been taken at very high

elevations near the verge of the snow-line on the most lofty parts of

the Simplon Pass, and other similar situations. Altogether about a

dozen species of this genus of butterfly are known, most of which are

confined to the polar regions of the Old World and the New, though some

have found their way to the extreme south end of South America, in what

manner is still a mystery. Like the preceding genera, this also appears

to have emerged from Central Asia. The genus, too, is closely allied to

the last, and though its range is not quite so extensive, it resembles

it in many respects. The Alpine species of _Oeneis_ came to Europe by

the Oriental route. But the Lapland species--at any rate _Oe. jutta_

and _Oe. bore_--have taken a somewhat circuitous route to reach our

continent. They first migrated from Asia to North America, and then by

the old land-connections by way of Greenland to Lapland. It is

noteworthy that Professor Engler felt convinced (cf. p. 171) that the

occurrence of many of the Arctic plants in North Scandinavia and Siberia

could be best explained by the assumption of such a migration from Asia

_via_ North America to Europe rather than by the shorter route.

There are far more Alpine beetles than butterflies, but their

geographical distribution is less well known, and it is therefore not at

all safe to base important conclusions as to the origin of a fauna on

that group alone; however, as far as my limited knowledge of the

_Coleoptera_ of the Alps goes, their general range seems to agree

perfectly with other orders of insects. Many can also be traced to an

Asiatic home, and the route they came by is the Oriental and not what I

have called the Siberian.

Take, for instance, the genus _Nebria_, of which we have one species in

England--a black insect with a bright reddish-yellow border and long

light legs--known as _N. livida_. There are about eighty European

species, most of which are confined to the Alps, the Caucasus, the

Pyrenees, Spain, and Greece. The genus, however, ranges all over the

Holarctic Region, that is to say roughly, over Europe, Central and

Northern Asia, and North America. The centre of distribution lies in

Central Asia. If the genus had poured into Europe by the northern or

Siberian route, we should probably now find many species in Northern

Russia, Germany, and France; but this is not the case, and we may

therefore assume with some justification that the Southern or Oriental

route was the only one available at the time when the bulk of the

species of _Nebria_ wandered to Europe. Many of the _Nebrias_ occur in

Switzerland and in the Alps, generally on the margins of the snow-fields

and glaciers, like _N. Germari_ and _Brunii_. Others, for example, _N.

atrata_, ascend to the highest limit of animal life, having been

observed at a height of over 10,000 feet.

Of the remaining orders of insects we know as yet very little. Central

Asia and even Siberia are only beginning to be explored, and their

invertebrate fauna--except _Lepidoptera_ and _Coleoptera_--is

practically unknown. However, I cannot conclude this short summary of

some of the more characteristic Alpine animals without referring to the

Grasshoppers which are so conspicuous in the mountains. The mountain air

simply rings during a bright summer's day with the loud and cheerful

song of millions of these insects. It is one of the most vivid

impressions a tourist brings back from Switzerland--this constant shrill

sound issuing from an apparently invisible source.

Among these Grasshoppers there are some highly characteristic Alpine

genera. _Pezotettix_--formerly known as _Podisma_--is one of these. _P.

alpinus_ is almost confined to the high Alps; with _P. mendax_ it occurs

in lower levels chiefly towards the south-east, that is to say, in the

direction of Hungary, Servia, and Dalmatia. _P. frigidus_ occurs not

only in the high Alps, but also in Lapland. _P. Schmidti_ and _P.

salamandra_ are found in Carinthia, Servia, and Transylvania; and one

species also inhabits the Pyrenees and another the Italian Mountains.

Finally, the only English species of _Pezotettix_, viz. _P. pedestris_,

has been taken in Sweden, Denmark, and then again in Austria, Hungary,

Servia, etc., as far east as the Volga, and also on the high Alps, in

Sardinia and the Abruzzi Mountains in Italy.

Very little, as I remarked, is known of the Asiatic range of this genus,

but either the same or a closely allied one has many representatives in

North and South America. Whether _Pezotettix_ is therefore of Asiatic

origin we cannot positively affirm, but whatever view we take, the

general range of the European species indicates that the migration took

place from the Alps in a south-easterly direction, or to them in a

north-westerly one. That is to say the Oriental route, and not the

Siberian, was utilised by the migrants.

Fortunately, we know a little more about another Grasshopper genus,

called _Chrysochraon_. There are only two species, one of which, _Chr.

dispar_, has been found from Northern France to the mountains of Servia,

but not in the Alps. The other, _Chr. brachypterus_, has a somewhat

similar range in the plain; but, moreover, it inhabits the Alps up to a

considerable height. It is interesting to note that both these

Grasshoppers again turn up on the Amur in Eastern Siberia.

In conclusion, I might mention one more Grasshopper, viz. _Tettix_,

because it includes a species--_T. bipunctatus_--which, though well

known in the plain of Middle and North Europe, ascends the Alps to a

height of nearly 10,000 feet. It is one of the few instances I know of

an animal occurring in the same form in such an enormous range of

altitude--from sea-level to the highest regions where animal life is

known to exist. It is also known from Asia Minor and Siberia. _T.

subulatus_ has a similar distribution, but is more common in Southern

Europe than the other. _T. fuliginosus_ occurs in Lapland and Siberia,

_T. meridionalis_ and _T. depressus_ all along the shores of the

Mediterranean. There can be no doubt that here also we can trace

migration to or from Siberia, and again, as on previous occasions, by

the Oriental route.

We now possess a fair general idea of the fauna of the Alps. We have

learned that a good many of the animals are indigenous, and that others

have migrated to the Alps by various routes. The majority of these have

come from Central and Southern Asia with what has been described as the

Oriental migration. A much smaller number have reached the Alps from the

north and the west, but none of the latter are among the high Alpine

forms. What will be the most surprising revelation is that the eastern

species, which arrived in Europe with the Siberian migration, are

practically absent from the Alps proper. No doubt some of them still

survive in the lowlands of Switzerland and the Tyrol, but none of the

true Alpine fauna owes its origin to the Siberian migration. If we

compare the Alpine mammals with the Siberian forms which reached England

(_vide_ p. 202), we at once perceive the difference. We should expect to

find in the Alps--if not the Reindeer and the Glutton--the Arctic Fox,

the little Pica, the Lemmings, and the pouched Marmots. It might be

urged that some of the smaller Siberian carnivores and rodents do

inhabit the Alps. So they do. The Stoat and Weasel have found such a

congenial home in Europe, both in the plain and mountains, that they

have spread rapidly to the latter, and no doubt reached within a

comparatively short time the great heights at which they now occur in

the Alps. But the Voles (_Arvicola_) have scarcely spread beyond the

region of fields and cultivated ground. A height of 5000 feet at the

most marks their maximum altitude in the Alps.

The fauna which reached the Alps in miocene and pliocene times, as well

as the indigenous element, must have survived the Glacial period in

their mountain home. Though I think that the conditions of the climate

at that time and the size of the Scandinavian glaciers have been greatly

exaggerated, there can be no doubt at all about the enormous size of

many of the Alpine glaciers at this period. The climate was probably

much moister but not colder than what it is now, possibly warmer. The

snowfall was therefore greater, so that glaciers filled many of the

lower valleys of Switzerland which are now quite free from ice, and even

invaded the plain. But there is no reason whatsoever why the Alps should

not even then have supported a luxuriant fauna and flora as they do now.

Possibly many of the miocene plants and animals became extinct then, but

extinction of species occurs at the present day. We hear complaints that

the Chamois and the Steinbock have nearly vanished; we know that the

Marmot is now much scarcer than it used to be, and that the Edelweiss

and many other plants are more and more difficult to find, and seem

rapidly to disappear. No doubt all this is in a great measure due to

the influence of man, but not altogether. There is a constant struggle

for existence going on among the animals and plants themselves--the

stronger and fitter species driving the less fit and weaker into a

corner, where they finally succumb. This happens now just as it did in

pliocene and pleistocene times, and need not imply change of climate.

As soon as the Miocene sea to the north of the mountains had retreated,

a portion of the Alpine fauna poured into the plain, and many species

have found their way to the British Islands, a few to Scandinavia and

Russia. Westward too, the sea soon after retired and opened a way for

those Alpine species which were vigorous enough to extend their range in

that direction. South-eastward, of course, a highway had long ago been

open, and Alpine forms which were able to migrate towards the incoming

Oriental stream, had no difficulty in doing so. When they arrived in

Greece, some turned westward again and populated Sicily, Southern Italy,

Sardinia, Corsica, and Northern Africa, while others crossed over to

Asia Minor, which was then connected with Greece, and wandered towards

the Central Asiatic or the Himalayan Mountains.

But, as I remarked, few of the typical Alpine species reached

Scandinavia and Lapland. I have already referred to the similarity

between the Northern Scandinavian and the Alpine faunas in a previous

chapter, and I have shown that this resemblance cannot altogether be

explained by the supposition of an interchange in the faunas of the two

regions. That this has taken place to some extent is probable, but the

resemblance appears more especially due to the fact that the Alps and

Scandinavia have been peopled from the same centres of distribution.

In order to make this matter quite clear, I will give a familiar example

as an instance of the manner in which the present distribution can be

explained without taking recourse to direct migration from the Alps to

Scandinavia or _vice versa_. The example I will take is that of a family

of birds, not only of extreme interest from the fact of its northern

range, but also from the pleasure it gives to those addicted to sport.

This is the grouse family, the _Tetraonidae_.

Let us commence with our British Grouse (_Lagopus scoticus_), which is

peculiar to the British Islands. In Norway we find a Grouse (_L. albus_)

which differs in habit, and in the fact of its turning white in winter;

otherwise it is so closely allied to our Grouse that many ornithologists

do not separate them specifically. No doubt the British Grouse is a

descendant of this Scandinavian Willow-grouse. The latter is known also

to inhabit Greenland and Arctic North America, and it is even found

beyond Behring Straits in Northern Siberia. _En route_ between

Scandinavia and Asia, travelling in a westward direction, we meet with

two other very local species of Grouse, which may be looked upon as

offshoots of _L. rupestris_--viz., _L. hyperboreus_ of Spitsbergen, and

_leucurus_ of Western North America. In Asia we then again find two

kinds of Grouse, very closely related, and by some indeed regarded as

belonging to the same species. These are _L. rupestris_ and _L. mutus_.

Mr. Ogilvie-Grant tells us of the former (p. 49), that it is merely a

more northern rufous form of _L. mutus_, and that it goes through

similar changes of plumage. In summer the males are readily

distinguishable, but in winter it is impossible to tell one from the

other. "_L. rupestris_ taken as a whole," says Mr. Ogilvie-Grant,

"appears to us barely specifically distinct from _L. mutus_." _L.

rupestris_ occurs not only in Northern Asia, but crosses the Behring

Straits to Arctic America, being still found on the Aleutian Islands,

which represent the last remains of the former land-bridge between Asia

and North America, then eastward to Greenland and Iceland. However,

while this form does not cross the confines of Asia in a westerly

direction, its near relative _L. mutus_--better known as the

Ptarmigan--does; and may perhaps have entered Europe as a Siberian and

also as an Arctic migrant. It is still found in the Ural Mountains, in

Finland, and the highlands of Scandinavia. It is gradually being driven

out of the Alpine lowlands, while it has long ago disappeared from

Germany, France, and Austria--in fact, from all the lowlands of Europe.

It has also been met with in the Pyrenees and in some of the Spanish

mountains. Similarly, the bird has become extinct in England and

Ireland, while it is becoming more and more scarce in Scotland. The

centre of distribution of the genus lies in Arctic America, and from

there the genus has spread to Europe and Asia. _L. albus_ and _L. mutus_

appear in our continent chiefly as Arctic migrants.

The Black Grouse (_Lyrurus tetrix_) belongs to a closely allied genus,

which has only two species. One of these is very local in distribution,

being confined to the Caucasus, but the smallness of range is to some

extent compensated for by the peculiarity of its name, which is _L.

mlokosiewiczi_. The Black Grouse, on the contrary, is widely

distributed. It inhabits Northern Asia from the Pacific to the Ural

Mountains, and extends as far south as Pekin and the Tian Shan range. In

Europe it is found from the extreme east to the Pyrenees, the Apennines

on the south, and to Great Britain and Scandinavia in the north. It is

important to note its absence from Spain, the Mediterranean islands, and

Ireland; and we have learned that it is one of those Siberian migrants

which have succeeded in establishing themselves in the Alps.

The Capercaillie (_Tetrao urogallus_)--another great favourite with

sportsmen--is now generally separated generically from the Black Grouse,

though they are of course near relations. Its range greatly resembles

that of the Black Grouse, except that it does not go quite as far east

in Siberia, not having been met with beyond Lake Baikal. From there it

is found westward as far as the Pyrenees. It occurs also in the

Carpathians and the Alps. In England, where it used to be known by the

name Cock of the Wood, it became extinct at some remote period in

history, while it lingered on in Scotland and Ireland until the end of

the last century. In Scotland it has been reintroduced into several

counties, and being protected, it appears to spread from these

artificial centres of distribution.

Like the Black Cock, the Capercaillie is a Siberian migrant, and it is

one of the few Siberian species which have reached Ireland, as I have

had occasion to mention in dealing with the origin of the British fauna.

Two other species of Capercaillie and an allied genus (_Falcipennis_)

are met with in the extreme north-east of Siberia, and six other genera,

all belonging to the grouse family, are confined to North America. We

have therefore a very intimate relationship between the grouse of Asia

and those of North America, some species even ranging right across the

two continents.

The last genus of this very interesting family is _Tetrastes_. This

grouse is not familiar to British ornithologists, since it is entirely

absent from the British Islands. But sportsmen who have tramped over

Scandinavia know it well by the name of Hazel Grouse. It is ashy grey in

colour, barred and vermiculated with black. The Common Hazel Grouse

(_Tetrastes bonasia_) is found from Northern Spain in the west right

through the mountainous parts of Central and Northern Europe and

Northern Asia to Kamtchatka and the Russian convict island of Saghalien

in the Pacific. Besides the Common Hazel Grouse, two other species are

known, one from Eastern Russia and the other from China.

Having now shortly reviewed the whole grouse family, we have seen that,

although some species live within the Polar Circle, the majority are

more or less confined to the more temperate or rather the less arctic

parts of the Northern Hemisphere. They are quite absent from Southern

Asia and even the southern parts of North America, and almost so from

the Mediterranean basin. The whole range of the family is therefore

suggestive of a northern origin, and this view agrees perfectly with all

the details of distribution. The centre of distribution lies in Northern

Asia, or in Arctic North America. From there the great genus _Lagopus_

spread east and west, reaching Europe by these vastly divergent routes

at a time when the physical geography was very different from what it is

to-day. Several of the species common to the Alps and Scandinavia have

migrated from Siberia direct to Eastern Europe. But we can now imagine

how from a similar centre in Asia--perhaps at a rather more remote

time--a species spread eastward across North America and Greenland to

Scandinavia, and westward along the mountain ranges of the Tian Shan and

the mountains of Asia Minor to Greece, and finally to the Alps. We

should then have the same species in the Alps and in Scandinavia, not

far removed from one another; but how different were their paths of

migration! This, however, is not an imaginary instance. Such a migration

must have actually taken place in a good number of instances among the

terrestrial invertebrates and also among plants.

The view still current among many zoologists and botanists, that animals

and plants were driven down into the plain from the mountains of Europe

during the height of the Glacial period and there lived together till

the return of a more genial temperature, when they retreated to their

mountain homes, is a very plausible one. During their sojourn in the

plain, the plants and animals--say from Scandinavia--intermingled with

those from the Alps; and when the time of separation came, many Alpine

forms retired northward with the Scandinavians, while many Scandinavians

would go with the Alpines to their home. In this way the similarity

between the Alpine and Scandinavian faunas and floras is assumed to have

been brought about. These theories, first promulgated by Edward Forbes,

were hailed with general satisfaction by the scientific world. Even

Darwin says of them (p. 331), that grounded as they are on the perfectly

well-ascertained occurrence of a former Glacial period, they seemed to

him to explain in a satisfactory manner the present distribution of the

Alpine and Arctic productions of Europe. To the present day this view

meets with much favour among naturalists. It is somewhat similar to one

which has recently been strongly supported by Professor Nehring and

accepted by Professor Th. Studer and many others. They have never made

it quite clear whether the pre-glacial fauna and flora are supposed to

have been absolutely destroyed by the glacial climate, or whether part

of them have been able to take refuge somewhere in the south; but the

great mass of our Alpine plants and animals are believed to have been

derived from the Siberian invasion, which I have fully described in the

fifth chapter. This invasion spread over the European plain, and when

the climate ameliorated, both animals and plants migrated north and

south to the mountains. This view agrees with the earlier theory, except

that the adaptation to Alpine conditions would, according to the former,

have taken place since the close of the Glacial period, during which

time no such modification or change of species seems to have been

produced in other parts of the world. The characteristic fauna of the

Alps, as has been gathered from the preceding pages, is mainly of

Central Asiatic rather than of Siberian origin. Migration to the Alps

took place by the Oriental route long before the Siberian invasion. Some

of the species of the latter have penetrated to the Alps, but these

Siberian species have not given to the fauna of the highest European

mountain range the striking character with which we all associate it.

Before concluding this chapter, a few remarks on the botanical aspect

of the Alpine problem might not be out of place. It will enable us to

judge which of the views indicated is the more probable, and will add to

the interest which may have been aroused by the perusal of this sketch

of the fauna of the Alps. Very much the same train of argument was

applied as to the course of events in the formation of the Alpine flora

as in the case of the fauna. The plants were all supposed to have been

killed or driven away by the arctic temperature of the Glacial period,

and their place taken by new migrants from the north or east when the

climate ameliorated.

Professor Engler, one of the highest living authorities on the

geographical distribution of plants, is of opinion (p. 102) that a large

number of the indigenous Alpine species did not originate till after the

close of the Glacial period, because so many of them are absent from the

Sierra Nevada in Spain, where the condition for their well-being exists,

while many have evidently spread from the Alps to the Carpathian

Mountains and to the Pyrenees. He does not believe that a glacial flora

could have existed in the plain between the Sierra Nevada and the

Pyrenees during the Glacial period (p. 109). In speaking of the

Caucasus, Professor Engler informs us (p. 117) that a good many species

which do not occur in the Alps reached these mountains from Siberia.

Apart from the northern glacial plants, the Caucasus has only few

species in common with the Alps, more with the Balkan mountains and

Northern Persia. Turning to Afghanistan, our author mentions (p. 121) a

few Alpine plants as occurring in that country, and likewise in the

Caucasus and the Himalayas. He considers it probable that the route of

migration of some glacial plants from the east to the west, and _vice

versa_, lay across the Afghan mountains. Many of our Alpine plants occur

in the Siberian mountains, but in the Altai and Eastern Siberia

generally a considerable number of these are by no means confined to the

mountains (p. 125). They are also met with in the lower regions, and the

rare Alpine Edelweiss (_Leontopodium alpinum_) frequently covers wide

tracts in the plain, and is passed by almost unnoticed by the Siberian

botanist.

Special attention is drawn by Professor Engler to the fact (p. 130) that

several of the Siberian plants inhabit the Alps and the Caucasus, but

are not found in Scandinavia. And from this he deduces the conclusion

that part of the Siberian flora migrated in a south-westerly direction

towards the Caucasus and the mountains of the Mediterranean area,

exactly in the manner indicated in respect to the fauna of the Alps. We

learned that the migration to the Alps from Central and perhaps also

parts of Northern Asia took a south-westerly course first, and was then

followed by one in an easterly direction. I called the former the

Oriental migration and the latter the Siberian. Later on Professor

Engler states (p. 142) that the main mass of the Siberian forms of

plants certainly wandered westward to the south of the Ural. This is

proved by the numerous glacial plants found in the Caucasus, while the

glacial flora of the Ural Mountains is poor. Finally, he expresses the

opinion that the probability of most of the Alpine plants occurring in

Arctic Siberia, having wandered from the Alps, by way of Scandinavia,

Greenland, and North America, to North-eastern Siberia, is greater than

the direct migration from Europe to Siberia (p. 143).

Another continental writer on the Alpine flora who deserves special

mention is Dr. Christ. His observation that Alpine plants by no means

suffer from a high temperature (p. 309), but solely from a drying up of

the soil, seems to me to point to the correctness of the view I have

expressed on several occasions, that these plants have originated long

before the Glacial period at a time when the climate was warmer and

moister than it is now. It seems quite natural to Dr. Christ that the

Arcto-Alpine flora should have originated in Asia, but he excepts thirty

species which are absent from Northern Asia, though occurring in America

(p. 327). These he thinks have penetrated direct from America to the

Alps by way of Scandinavia, since no less than twenty-three still occur

in the latter country. In the human population of the Alps, he continues

(p. 336), one can distinguish an indigenous Celtic race, a Germanic

colder and more apathetic race, and a more lively Roman one. The flora

is composed of quite a similar mixture. We find also an indigenous

element--an Arctic and a Mediterranean one. The last element is a

survival of the Tertiary flora of the Central European plateau (p. 532).

The plants were driven down to the shores of the Mediterranean, and it

is only after the retreat of the glaciers that a few of them have been

able to regain their ancient territory. The incoming Asiatic and North

American flora likewise retired at the end of the Glacial period to the

Alps and the Arctic countries, and left isolated traces of its former

abundance on the North European plain. The bulk of the Arctic or Alpine

flora is held to be of Asiatic origin. Since Siberia shows little trace

of having been glaciated, owing to the dryness of the climate, a rich

flora was able to develop there, which spread into Europe as soon as the

vanishing glaciers made room for it.

These are the views of Professor Engler and Dr. Christ. They agree in so

far as both of them maintain that the bulk of the Alpine flora is

post-glacial--that is to say, that it has developed quite recently, or

migrated to the Alps after the glaciers had retreated from the plain to

the mountain recesses. It is assumed by Dr. Christ that while Europe was

practically uninhabitable, a rich flora survived in Northern Asia,

because the climate there was too dry for the development of glaciers.

Due consideration in this interesting speculation, however, is not given

to the fact which he himself emphasised, that Alpine plants are

particularly prone to suffer from a dry climate. Even a moderately dry

cold kills most of them. How can we then reconcile this fact with the

theory of their origin in a dry and intensely cold climate? I quite

agree with the views as to the Asiatic origin of the bulk of the Alpine

flora, while the dry state of the Siberian climate is certainly

indicated by the extremely feeble development of the glaciers during a

large part of the Glacial period. We know, however, that in Pliocene and

even in early Glacial times the atmospheric conditions must have been

very different in Siberia. A great slice of Central Asia was under

water, and numerous freshwater lakes covered the lowlands in the north,

so that the climate must have been damp though not cold enough for the

formation of extensive glaciers. Everything, in fact, seems to indicate

that the migration of the Asiatic Alpine flora took place at a very

early date--probably long before the Glacial period--either by the

Oriental or by the Arctic route _via_ North America, Greenland, and

Scandinavia. But would this not necessitate a survival of the Alpine

plants in the Alps themselves? That is the view which has already been

expressed with regard to the fauna, and the flora probably followed a

very similar course. This is by no means a novel theory, however, and

though unfortunately an untimely death has removed one of our very best

authorities on the Alpine flora before he had completed his life's work,

we have some indications in the earlier writings of John Ball that his

opinions on the origin of that flora did not coincide with those held by

the leading continental authors. To quote the words of this

distinguished botanist (p. 576): "Is it credible that in the short

interval since the close of the Glacial period hundreds of very distinct

species and several genera have been developed in the Alps, and--what is

no less hard to conceive--that several of these non-Arctic species and

genera should still more recently have been distributed at wide

intervals throughout a discontinuous chain some 1500 miles in length,

from the Pyrenees to the Eastern Carpathians? Nor would the difficulties

cease there. You would have left unexplained the fact that many of the

non-Arctic types which are present in the Alps are represented in the

mountains of distant regions, not by the same, but by allied species,

which must have descended from a common ancestor; that one species of

_Wulfenia_, for example, inhabits one small corner of the Alps, that

another is found in Northern Syria, while a third allied species has its

home in the Himalaya." Mr. Ball is of opinion (p. 584) that the effects

of the Glacial period have been greatly overrated. "Even during the

period of maximum cold the highest ridges of the Alps were not

completely covered with snow and ice; for we still see by the appearance

of the surface, the limit above which the ancient ice did not reach, and

in the middle zone the slopes that rose above the ancient glaciers had a

summer climate not very different from that which now prevails. In my

opinion the effect of the Glacial period on the growth of plants in the

Alps was to lower the vertical height of the zones of vegetation by

from one to two thousand feet." He acknowledges that there was probably

a moderate diminution of the mean temperature of Europe with an

increased snowfall, so as to cause a great extension of glaciers on all

the mountains of Northern Europe. "But that the climate of Middle Europe

was such that the plants of the high Alps could spread across the plains

seems to me an improbable supposition" (p. 584).

On the Continent, also, some botanists seem to feel that Forbes's

theories of the origin of the Alpine flora, which were at first hailed

with such delight, and accepted by almost every naturalist as the final

verdict, must be modified in the light of recent researches. Professor

Krasan believes that many plants which now live in the high Alps

flourished in pliocene times at sea-level (p. 37). "Especially the

evergreen species exhibit the impression of an originally mild

climate--of a climate without winter frosts--for otherwise the plants

would have developed into species with deciduous leaves." To the

favourable conditions, consisting in periodic snowfalls and high summer

temperature, must be attributed the fact that in the highlands so many

more species from Tertiary times have survived than in the plains. The

temperature was probably much higher during the Glacial period than is

generally believed. The climate was more moist, thus contributing to an

abundant snowfall, while the survivors of ancient Tertiary times were

able to repeople the parts which were temporarily devastated by the

advancing glaciers.

In so short a chapter it is impossible to deal with the Alpine fauna in

a manner more deserving of this theme. I have merely sought to give a

sketch of the general outlines of the subject and to suggest another

possible mode of origin of Alpine animals than that currently believed

in by naturalists. It is to be hoped these suggestions will be useful to

those intending to reinvestigate the problems raised in this chapter.

When our knowledge of the fauna of Asia is more complete, it will be

possible to give a more thorough and in many respects a more

satisfactory history of the European fauna than at present.

SUMMARY OF CHAPTER VIII.

In early Tertiary times the area now covered by the European Alps was

covered by the sea. Islands slowly rose above the surface of the waters,

which finally coalesced to form a peninsula connected with the mainland

in the east. Animals now began to invade the new territory which

continued to rise, while the sea retired farther and farther to the

north and south. During the Pliocene Epoch the sea ceased to wash the

northern shores of the Alps, and both emigration and immigration became

possible in that direction, and also from and to the west.

The Alpine fauna and also the flora are made up of a number of elements,

the eastern one being the oldest. The latter is represented in the Alps

by the older and newer Oriental migration. The general range of the

Alpine Steinbock, Chamois, Marmot, Vole, Shrew, and Hare are specially

referred to. The Alpine birds are few in number, and all of them are

readily traceable to an Asiatic ancestry. Among the Amphibia, the

Salamanders are considered of Alpine origin.

Dr. Kobelt tells us that a uniformity of character marks the Alpine

molluscan fauna. _Campylaea_,--often considered a sub-genus of

_Helix_,--_Pomatias_, _Zonites_, are looked upon as truly Alpine genera.

For very long periods the Alps seem to have received no addition to

their molluscan fauna from other areas. The case is very different with

the _Lepidoptera_, some of the most striking species being evidently

Asiatic immigrants. Some examples of _Coleoptera_ and _Orthoptera_ are

mentioned, and their origin discussed.

We find as the result of these considerations that the majority of the

Alpine species are either indigenous or have come from Asia with the

Oriental migration. None of the northern or western immigrants appear to

be among the characteristic Alpine species, and it seems that the

Siberian migrants have not retired to the Alps, as some naturalists have

been led to suppose. It is evident that the fauna must have survived the

Glacial period on the Alps, though according to geological evidence

glaciers of enormous size originated on these mountains.

The identity of many Alpine species with Scandinavian ones appears at

first sight due to a direct migration from the Alps to Scandinavia or

_vice versa_. Perhaps such a migration has taken place to some extent,

but it is probable that from a Central Asiatic centre some species

spread across Arctic America into Northern Europe, and also westward to

the Alps. The Grouse family forms an interesting example.

There are two older theories which explain the similarity between the

Scandinavian and Alpine faunas. Forbes's view, which gained most

adherents among naturalists, was that the Scandinavian and Alpine

animals were driven into the plain by the cold during the Glacial

period, and when they ultimately regained their homes, some individuals

of the northern species moved southward, and a few of the southern ones

northward. By the more recent theory of Nehring, the Siberian animals

which invaded our continent from the east, and then spread northward to

Scandinavia and southward to the Alps, formed the nucleus of the faunas

of these two areas. The objections to both of these views are fully set

forth in this chapter.

A few remarks on the botanical aspect of the Alpine problem conclude the

chapter. The origin of the flora has been explained in a very similar

manner to that of the fauna. But already Ball and Krasan have raised

their voices against the current theories, as the facts of distribution

appear to them more satisfactorily explained on lines more consonant

with those which I have used in discussing the origin of the Alpine

fauna. One of the most important conclusions obtained by this study of

the flora in conjunction with the fauna, is that I have emphasised in

most of the preceding chapters--viz., that the Glacial period in Europe

was not a time of extreme cold, and that its destructive effect on the

animals and plants was by no means such as is currently believed.

BIBLIOGRAPHY.

(_Titles of Works and Papers referred to in the Text._)

Adams, A. Leith.--Report on the History of Irish Fossil Mammals, "Proc.

Royal Irish Acad." (2nd series), vol. iii., 1878.

Alston, E. R.--"The Fauna of Scotland," 1880.

Ball, J.--On the Origin of the Flora of the European Alps, "Proc. Royal

Geograph. Soc.," vol. i., 1879.

Barrett-Hamilton, G. E. H. (_vide_ also Thomas and

Barrett-Hamilton).--Notes on the Introduction of the Brown Hare in

Ireland, "Irish Naturalist," vol. vii., 1898.

Beddard, F. E.--"A Text-book of Zoogeography," 1895.

Bell (_vide_ Kendall and Bell).

Blanchard, E.--Preuves de la Formation Recente de la Mediterranee,

"Comptes Rendus," vol. xciii., 1881.

Blanford, W. T.--Address delivered at the Anniversary Meeting of the

Geological Society of London, 1890.

Blytt, A.--"Essay on the Immigration of the Norwegian Flora," 1876.

Boettger, O.--=a.= System. Verzeich. d. lebenden Arten d.

Landschneckengattung Clausilia, "Ber. Offenbach Ver. f. Naturkunde,"

1878. =b.= Entwicklung d. Pupa Arten d. Mittelrheingebiets, "Jahrbuch d.

nass. Ver. f. Naturw.," vol. xlii.

Bogdanov, M. M.--Quelques mots sur l'histoire de la faune de la Russie

d'Europe, "Archives des Sciences physiques et naturelles" (N.S.), vol.

lvi., 1876.

Bonney, T. G.--"Ice-work, Present and Past," 1896.

Boulenger, G. A.--British Museum Catalogue (Amphibia), 1882.

Bourguignat, J. R.--Distribution geographique des Mollusques terrestres

et fluviatiles en Algerie, "Annales des Sciences naturelles" (5th ser.),

vol. v., 1864.

Brandt, J. F.--Zoogeographische und Palaeontologische Beitraege,

"Verhandl. der Kais. russ. Mineral. Gesell." (2nd ser.), vol. ii., 1867.

Brauer, A.--Die Arktische Subregion, "Zool. Jahrbuecher" (Abth. System.),

1888.

Brehm, A. E.--"From North Pole to Equator," 1896.

Brooke, Sir V.--On the Classification of the Cervidae, "Proc. Zool. Soc.

London," 1878.

Bulman, G. W.--The Effect of the Glacial Period on the Fauna and Flora

of the British Islands, "Natural Science," vol. iii., 1893.

Bunge, D.--La faune eteinte des bouches de la Lena et des iles de la

Nouvelle Siberie, "Congres Intern. de Zoologie," vol. ii., Moscow, 1892.

Carpenter, G. H.--Problems of the British Fauna, "Natural Science,"

vol. ii., 1897.

Christ, H.--"Das Pflanzenleben der Schweiz," 1879.

Cole, G. J.--Borderland, "Irish Naturalist," vol. vi., 1897.

Credner, R.--Die Reliktenseen, "Petermann's Geog. Mitth.," 1887.

Croll, J.--"Climate and Time," 1875.

Darwin, C.--"Origin of Species," 6th ed., 1878.

Dawkins, W. Boyd.--=a.= "Early Man in Britain," 1880. =b.= "The British

Pleistocene Mammalia," part A, Preliminary Treatise, 1878. =c.= The

Former Range of the Reindeer in Europe, "Popular Science Review," vol.

vii., 1868.

Drude, O.--Betrachtungen ii. d. hypothetischen vegetationslosen Einoeden

im temperirten Klima der Noerdlichen Hemisphaere zur Eiszeit, "Petermann's

Geog. Mitth.," vol. xxxv., 1889.

Emery, C.--On the Origin of European and North American Ants, "Nature,"

vol. lii., 1895.

Engler, A.--"Versuch einer Entwicklungsgeschichte d. extratrop.

Florengebiete d. Noerdlichen Hemisphaere," 1879.

Falsan, A.--"La Periode Glaciaire," 1889.

Feilden, H. W.--=a.= Notes on the Glacial Geology of Arctic Europe and

its Islands, part 2, "Quart. Journ. Geol. Soc.," vol. lii., 1896. =b.=

Notes on a small Collection of Spitsbergen Plants, "Trans. Norfolk and

Norwich Nat. Hist. Soc.," vol. vi., 1894.

Fischer, P.--Faune malacologique de Cauterets, suivie d'une etude sur la

repartition des Mollusques, "Journal de Conchyliologie" (3rd series),

vol. xvi., 1876.

Forbes, E.--The Geological Relations of the Fauna and Flora of the

British Isles, etc., "Memoirs Geol. Survey," vol. i., 1846.

Geikie, Sir A.--"Text-book of Geology," 1882.

Geikie, J.--=a.= "The Great Ice Age," 1894. =b.= On Changes of Climate

during the Glacial Period, "Geological Magazine," vol. ix., 1872.

Haacke, W.--Der Nordpol als Schoepfungs-zentrum der Landfauna,

"Biologisches Centralblatt," vol. vi., 1887.

Harle, E.--Sur la succession de diverses faunes a la fin du quaternaire,

dans le sud-ouest de la France, "Societe d'Histoire Naturelle,"

Toulouse, 1893.

Harmer, F. W.--On the Pliocene Deposits of Holland, and their relation

to the English and Belgian Crags, "Quarterly Journal Geological Soc.,"

vol. lii., 1896.

Herdman, W. A., and Lomas, J.--On the Floor Deposits of the Irish Sea,

"Proc. Liverpool Geological Society," vol. viii., 1898.

Hofman, E.--Die Isoporien der Europaeischen Tagfalter, "Inaugural Diss.

d. Phil. Fakultaet in Jena," 1873.

Hooker, J. D.--Outlines of the Distribution of Arctic Plants, "Trans.

Linnean Soc.," vol. xxiii., 1862.

Howorth, Sir H.--"The Mammoth and the Flood," 1887.

Ihering, H. von.--=a.= Die Ameisen von Rio Grande d. Sul, "Berliner

Entomolog. Zeitschrift," vol. iii., 1894. =b.= Najaden von Sao Paulo und

die geographische Verbreitung d. Suesswasserfaunen von Sued Amerika,

"Archiv. f. Naturgesch.," 1893.

Jordan, H.--Die Binnenmollusken der noerdl. gemaess. Laender, "Nova Acta

Acad. Carol.-Leop.," vol. xlv., 1883.

Judd, J. W.--Address to the Geological Section, "British Association

Report," Aberdeen, 1885.

Karpinski, A.--Uebersicht d. physiko-geograph. Verhaeltnisse d. Europ.

Russlands, etc., "Beitraege z. Kenntniss d. Russ. Reichs" (3 Folge), vol.

iv., 1888.

Kendall, P. F., and Bell, A.--On the Pliocene Beds of St. Erth, "Quart.

Jour. Geol. Soc.," vol. xlii, 1886.

Kennard, A. S., and Woodward, B. B.--The Mollusca of the English

Cave-Deposits, "Proc. Malacological Soc.," vol. ii., 1897.

Kew, H. W.--"The Dispersal of Shells," 1893.

Kinahan, G. H.--On possible land-connections in recent geological times

between Ireland and Great Britain, "Trans. Manchester Geol. Soc.," vol.

xxiv., 1896.

Kobelt, W.--=a.= Das Verhaeltniss d. Europ. Landmoll. z. Westindien und

Central Amerika, "Nachrichtsblatt d. deutschen Malak. Gesell.," 1886.

=b.= "Studien zur Zoogeographie," vols. i., ii., 1897-98.

Koeppen, F. T.--Das Fehlen d. Eichhoernchens, etc., in der Krim, "Beitraege

zur Kenntniss d. Russ. Reichs" (2 Folge), vol. vi., 1883.

Krasan, F.--Zur Abstammungsgesch. d. autochthonen Pflanzenarten, "Mitth.

d. Naturw. Ver. Steiermark," 1896.

Lamplugh, G. W.--The Glacial Period and the Irish Fauna, "Nature," vol.

lvii., 1898.

Lomas (_vide_ Herdman and Lomas).

Lydekker, R.--=a.= "A Handbook of the British Mammalia," 1895. =b.= "A

Geographical History of Mammals," 1896. =c.= "The Deer of all Lands,"

1898. =d.= Catalogue of the Fossil Mammalia of the British Museum, pt.

4, 1886.

Major, C. J. Forsyth.--=a.= Die Tyrrhenis, "Kosmos," 7 Jahrg., 1883.

=b.= Studien zur Geschichte der Wildschweine, "Zool. Anzeiger," vol.

vi., 1883.

Mallet, R.--Some Remarks upon the Movements of Post-tertiary and other

Discontinuous Masses, "Journal Geol. Soc. Dublin," vol. v., 1850-53.

Merriam, C. H.--The Geographic Distribution of Life in North America,

"Proc. Biological Soc. Washington," vol. vii., 1892.

Murray, Andrew.--"The Geographical Distribution of Mammals," 1866.

Nathorst, A. G.--Kritische Bemerkungen ueber die Geschichte d. Vegetation

Groenlands, "Botanische Jahrbuecher," vol. xiv., 1891.

Nehring, A.--"Tundren und Steppen der Jetzt und Vorzeit," 1890.

Neumayr, M.--"Erdgeschichte," vol. ii., 1887.

Ogilvie, W. R. Grant.--"Game Birds," 1895.

Penck, A.--Die grosse Eiszeit, "Himmel und Erde."

Petersen, W.--Die Lepidopterenfauna d. arktisch. Gebiete von Europa,

"Beitraege z. Kenntniss d. Russ. Reichs," vol. iv., 1888.

Pettersen, K.--Arktis, "Arch. f. Math. og Naturvid.," 1882.

Pohlig, H.--Dentition und Kraniologie des Elephas antiquus (part 2),

"Nova Acta Acad. Carol.-Leop.," vol. liii., 1889.

Reade, Mellard--=a.= The Present Aspects of Glacial Geology, "Geological

Magazine," vol. iii., 1896. =b.= High Level Shelly Sands and Gravels,

"Natural Science," vol. iii., 1893.

Reid, C.--The Pliocene Deposits of Britain, "Mem. Geol. Survey of the

United Kingdom," 1890.

Ruetimeyer, L.--"Die Herkunft unserer Thierwelt," 1867.

Sars, G. O.--Crustacea Caspica, "Bull. Acad. Imp. d. Sciences St.

Petersburg," vol. xiii., 1893-94.

Scharff, R. F.--=a.= On the Origin of the European Fauna, "Proc. Royal

Irish Acad." (3rd. ser.), vol. iv., 1897. =b.= Some Remarks on the

Distribution of British Land and Freshwater Mollusca, "Conchologist,"

vol. ii., 1893. =c.= Etude sur les mammiferes de la Region Holarctique,

"Mem. de la Soc. Zool. de France," vol. viii., 1894.

Schulz, A.--"Grundzuege einer Entwicklungsgeschichte d. Pflanzenwelt

Mittel-europas," 1894.

Sharpe, R. Bowdler.--British Museum Catalogue (Birds).

Simroth, H.--Eine Bearbeitung d. Russischen Nacktschnecken fauna, "Ann.

Musee Zool. St. Petersburg," 1896.

Sjoegren, H.--Ueber das diluviale Aralo-Kaspische Meer und die

nord-europaeische Vereisung, "Jahrb. d. Kais. K. Geol. Reichsanstalt,"

vol. xl., 1890.

Sollas, W. J.--On the Origin of Freshwater Faunas, "Trans. Royal Dublin

Soc." (2nd ser.), vol. iii., 1884.

Speyer, A. and A.--"Die Geographische Verbreitung der Schmetterlinge

Deutschlands und der Schweiz," vol. i., 1858.

Struckmann, C.--Ueber die Verbreitung d. Rennthiers, in d. Gegenwart,

etc., "Zeitschr. d. deutsch. Geol. Gesell.," vol. iii., 1883.

Suess, E.--"Das Antlitz der Erde," vol. i., 1892.

Tcherski, J. D.--Das Janaland und die Neusibirischen Inseln, "Mem. Acad.

Imp. St. Petersburg," vol. xl., 1892.

Thomas, O., and Barrett-Hamilton, G. E. H.--The Irish Stoat distinct

from the British species, "Zoologist," 1895.

Wallace, A. R.--"Island Life," 2nd edit., 1892.

Warming, E.--Ueber Groenlands Vegetation, "Botanische Jahrbuecher," vol.

x., 1888-89.

White, F. B.--Some Thoughts on the Distribution of the British

Butterflies, "The Entomologist," vol. xiv., 1881.

Woodward, B. B.--On the Pleistocene (non-marine) Mollusca of the London

District, "Proc. Geologists' Association," vol. ii. (_vide_ also Kennard

and Woodward).

Wright, J.--Boulder-clay a marine deposit, "Trans. Geol. Soc. Glasgow,"

vol. x., 1896.

INDEX.

_Ablepharus pannonicus_, 258

_Accentor collaris_, 316

_modularis_, 317

Accidental distribution, 12, 26

_Acipenser ruthenus_, 29

Acme, 323

Adams, Leith, 112, 150

AEgean continent, subsidence of, 272-273

Agriolimax, 284

_Alactaga jaculus_, 204

_Alca impennis_, 92, 142

Alligator, accidental dispersal of, 14

_Allolobophora Georgii_, 115, 302

_veneta_, 115, 302

Alpine Accentor, 316

Chough, 317

fauna, survival of, during Glacial period, 332

flora, 341-348

flora, age of, 78, 79

flora, suitable conditions for, 78-79

Hare, as a test of climate, 316

Snow-finch, 318

Alps, component elements of fauna, 311-312

Alston, E. R., 312

_Alytes cisternasii_, 295

_obstetricans_, 295

_Amalia_, 284

America, introductions from, 24

American beetles in the British Islands, 167

plants, origin of, in the British Islands, 144

plants in Ireland, 166

sponges in Ireland, 166

_Ampelis garrulus_, 205

Amphibia of Europe, 193-194

dispersal of, 21

_Amphicoma_, 268

Andalusian Bush-quail, 291-292

_Anergates_, 167

Anglo-Scotian fauna, 95

Animals as tests of climate, 71-75

Antelope, 39

Ants, European origin of, 161

_Apollo_, 325

_Apus glacialis_, 94, 167

Aralo-Caspian Sea and Arctic Ocean, connection between, 219-231

_Arbutus unedo_, 110, 307

Arctic animals in Caspian Sea, 238

flora in Europe, 238-241

Hare, origin of, 148-149, 158-159

Hare, range of, 136-138

Lepidoptera, 200

plant-beds at Bovey Tracy, 238

plants, 143-144

plants, delicacy of, 185

mollusca in Red Crag, 235-236

Seal, 174

sub-region, 132

_Arctomys bobak_, 204, 313

_marmotta_, 313

_Arion_, 48, 299

_subfuscus_, 49

_Arionidae_, origin of, 48, 299

_Asiminea Grayana_, 99

Aucapitaine, Baron, experiments by, 15

Autochthonous species, 10

Azores, remains of a continent, 19

_Bacillus_, 269-270

Badger, distribution of, 43

_Balea perversa_, 298

Ball, J., 77, 345-347

Baltic Sea, fauna of, 174

Baltic and White Seas, connection between, 175

Banded Lemming, range of, 138

Barn Owl, 98

Barrett-Hamilton, G. E. H., 29, 90, 139, 149

Barriers to animal migration, 38

Barrington, R. M., 105

_Bathyphantes nigrinus_, 94

Baur, G., 19

Bearded Titmouse, 292

Beaver, 203

Beaver absent from Ireland, 121

Beddard, F. E., 19, 58

Bedriaga, J. von, 259-260

Beetles common to North America and Europe, 161

American, in British Islands, 167

Bell (_vide_ Kendall and Bell)

Black Cock, 143

Black Grouse, 336

Blanchard, E., 282

_Blanus cinereus_, 279, 294

_Blethisa multipunctata_, 93

Blytt, A., 34, 52, 79, 185, 239

Boar, distribution of, 44

Bobak Marmot, 204

Boettger, O., 48, 263

Bogdanov, M. M., 53

_Bombinator_, 259-260

_igneus_, 259

_pachypus_, 259

Bonnet, 306

Bonney, T. G., 66, 70, 83, 180, 229

Boulenger, G. A., 259

Boulder-clay, foraminifera in, 84

origin of, 81, 175, 180-181, 229-230

of Continental Europe, 180, 226-231

Bourguignat, J. R., 18, 47, 302

Bovey Tracy, arctic plant beds at, 238

Boyd, E., 104

_Branchinecta palludosa_, 167

Brandt, J. F., 51, 62, 107, 218, 222

Brauer, A., 132-133

Brehm, A. E., 139

Bristle-fern, 115

British flora during Glacial period, 163

British Islands, Lusitanian flora in, 288, 306-307

submergence of, 127

Brooke, Sir V., 247, 250

Brunner von Wattenwyl, 270

_Bufo calamita_, 30

_Buliminus detritus_, 261

_fasciolatus_, 261

_obscurus_, 261

_pupa_, 261

Bullfinch, origin of, 191, 255-256

Bulman, G. W., 114

Bunge, D., 219

Bush-quail, Andalusian, 291-292

Butterflies, Arctic distribution of, 159-161

origin of North European, 55

_Caccabis rufa_, 29

_Campylaea_, sub-genus of _Helix_, 49, 321

Canary Islands, origin of, 19

_Canis lagopus_, 135

Capercaillie, 28, 143, 336-337

_Capra ibex_, 312

_aegagrus_, 61, 312

_pyrenaica_, 312

_sibirica_, 312

_sinaitica_, 312

_Carabus_, 199

_Carduelis_, 257

_caniceps_, 258

_elegans_, 257

_major_, 257

Carpenter, G. H., 64, 94-95, 104, 114, 121-123, 288, 301

Caspian Fishes, 224

post-pliocene deposits, 222, 226

Sea, Arctic animals in, 238

seal, 224

_Castor fiber_, 203

Cave deposits, mixture of Northern and Southern animals in, 54

Cenocosmic species, 24

Centre of distribution, 12, 43, 47

origin, shifting of, 202

_Cervidae_, origin of, 248

_Cervus dama_, 251

_elaphus_, 246-250

_giganteus_, 108, 251

_Chamaeleon vulgaris_, 279

Chamois, 312

Charrs, origin of, 124

_Chioglossa_, 296

Chough, Alpine, 317

Christ, H., 52, 343-344

Christy, Miller, 105

_Chrysochraon_, 330

_Cicindela_, 197-198

_Cinclus_, 255

Clarke, Eagle, 105

_Clausilia_, 47, 262-264, 284

_bidentata_, 47, 262, 284

_biplicata_, 47

_laminata_, 47

_Pauli_, 48, 263

_Rolphii_, 47

_Claviger_, 267-268

_testaceus_, 268

Climate in Glacial period, 65, 81, 127, 149, 182-183

in Pleistocene Europe, 208-209

Close, Maxwell, 129

Cod-fish, origin of, 143

_Coecilianella_, 17

_Coenonympha typhon_, 93

Cole, G. J., 83, 106-107

Coleoptera of Europe, origin of, 197

common to Europe and North America, 161

_Colias_, 266

_Coregonus_, 124

_clupeoides_, 124

_pollan_, 124

_vandesius_, 124

Cosmopolitan species, 24

_Cottus_, 92-93

_bubalis_, 93

_gobio_, 93

_quadricornis_, 175

_scorpio_, 93

Credner, R., 177-178

_Cricetus frumentarius_, 190

_vulgaris_, 204

Crimea formerly an island, 35

Crimean fauna, 57

Croll, J., 66

_Crocidura_, 253

_etrusca_, 253, 279

_Cuniculus torquatus_, 138

Current, Arctic, 172-173

_Cyanopolius Cooki_, 293

_cyanus_, 293

_Cyclostoma elegans_, 16

_Danais chrysippus_, 267

Dartford Warbler, 288

Darwin, C., 13-15, 17-19, 25, 32-33, 339

_Daudebardia_, 323

_Daulias luscinea_, 192

Dawkins, Boyd, 53, 62-63, 72-73, 107, 112-113, 120, 208, 222-223, 282

Day, F., 29

Deperet, C., 44, 276

Dippers, origin of, 255

_Discoglossus pictus_, 279, 295-296

Dispersal of Amphibia, 59

of beetles (_apterous_), 59

of British butterflies, 113

of earthworms, 58

of planarian worms, 59

of spiders, 59

of wood-lice, 59

Distribution, centres of, 12, 43, 47

discontinuous, 114

Dormouse, 316

absent from Ireland, 121

_Drapetisca socialis_, 94

_Dreyssensia polymorpha_, 26, 230-231

Drift, a marine deposit, 129

Drude, O., 52, 77

_Dryas octopetala_, 79, 238

Dual origin, possibility of, 38

Dyer, Th., 79

Earth-worms, distribution of, 19, 23, 58

Edelweiss, 266, 342

Egean Continent, subsidence of, 272-273

Eider-duck, range of, 141

_Elephantidae_, origin of, 202, 252-253

_Elephas primigenius_, 214, 252

Emery, C., 161, 167

Endemic species, 10

Engler, A., 52, 145, 171, 176, 282, 341-342

English Hare, 29

_Ephydatia crateriformis_, 94

_Epoccus_, 167

_Erebia_, 325-326

_aethiops_, 325-326

_epiphron_, 325

_glacialis_, 326

_lappona_, 326

_Eremias_, 258

Erratics, 181-182

_Eryx jaculus_, 259

_Euphorbia hiberna_, 307

European beetles, origin of, 197

butterflies, origin of, 200

land and fresh-water mollusca, origin of, 196

mammals, origin of, 106, 193

_Eurynebria complanata_, 302

_Evotomys Nageri_, 314

Extension of range, mode of, 38

Fallow Deer, 251

Falsan, A., 66, 68-69, 71, 73

Feilden, H. W., 13, 77, 84-85, 164

Finmark, Greenland mollusca on coast of, 171

Fire-toads, origin of, 259

Fischer, P., 103

Fishes, Caspian, 224

"Flotsam and jetsam" theory, 20

Foraminifera in boulder-clay, 84

Forbes, E., 64, 101, 110, 114-115, 171, 288, 339

Forest-Bed an inter-glacial deposit, 70

corresponding to continental inter-glacial deposits, 120

fauna of, 232-234

mollusca, 212-213

Fossil glaciers, 220

Fresh-water faunas, origin of, 177

_Fringilla_, 293

_coelebs_, 293

_madeirensis_, 293

_montifringilla_, 293

_spodiogenys_, 293

_teydea_, 293

Frog, introduction of, into Ireland, 30-31

Gadow, H., 139

Galapagos Islands, 19

_Galeodes_, 270

_Gammaracanthus relictus_, 179

Gardner, J. S., 145

_Garnieria_, sub-genus of _Clausilia_, 48

_Gasterosteus aculeatus_, 92

Gaudry, A., 73, 273

Geikie, Sir A., 116, 309

Geikie, J., 59, 66-67, 70, 75-76, 80-83, 116, 129, 163, 182, 216-217,

226-227, 233, 235, 238

Geographical changes, importance of, 64

_Geomalacus maculosus_, 5, 49, 99, 102, 115, 298-299

Gervais, E., 150

Glacial climate in France, 150

period, climate of, 65-81, 127, 149, 182-183

period in Scandinavia, 176

period, survival of animals and plants during, 65

Glaciation of Ireland, 129

Glutton, 119, 203

Goldfinch, origin of, 257

_Gonepteryx_, 296-297

_cleobule_, 297

_cleopatra_, 296

_rhamni_, 296

Gould, J. E., 20

Great Auk, range of, 92, 142

Greenland flora, 161-162

flora, survival of, 163-164

miocene temperature in, 146

mollusca on coast of Finmark, 171

Tertiary plants, 144-145

Grouse, 91, 334

black, 336

_Gulo luscus_, 119, 203

Haacke, W., 147

Hamilton, John, 161

Hamster, 190, 204

Hanitsch, R., 94

Hare, 2, 29, 90

Arctic, 2, 90-91

English, 29

Hare, Alpine, as a test of climate, 316

Harle, E., 150

Harmer, F. W., 182

Harvest-mouse, 3, 190

Harvie-Brown, J. A., 105

Haughton, W., 86

Hazel-grouse, 337

Hedge Accentor, 317

Hedley, C., 20

Heer, O., 144-145

_Helix_, 4

_acuta_, 102, 298

_aculeata_, 298

_aspersa_, 24, 195

_ericetorum_, 298

_fusca_, 298

_lapicida_, 4

_obvoluta_, 4, 297

_pisana_, 102, 298

_pomatia_, 15, 32, 195, 262

_revelata_, 102

_rotundata_, 122, 274, 298

_ruderata_, 212-213

Herdman, W. A., and Lomas, J., 236

Herring, origin of, 143

_Heteromeyenia Ryderi_, 94

Hofman, E., 54

Hooker, Sir J., 99, 161

_Hopatroides thoracicus_, 268

Howorth, Sir H., 73

Hyalinia, antiquity of, 50

_Hyla_, 260

_arborea_, 260-261, 280

_chinensis_, 260

Ice-Age, climate of, 65, 81

Ice-bridge, migration on, 108

_Idotea entomon_, 178, 224

Ihering, H. von, 19, 20, 24-25

Indigenous species, 10

Inter-glacial deposit, the Forest-Bed an, 70

periods, climate of, 181, 218

Introduced species, 10, 23, 27-29

Introduction by man, 32-33

Introductions from America, 24

Ireland, glaciation of, 129

Irish and Scotch Hare, 136

Elk, 108, 251-252

Irish fauna, composition of, 63

Stoat, 136

Isle of Man, fauna of, 123

_Isotoma littoralis_, 94

Jeffreys, J. G., 27

Jerboa, 204

Jordan, H., 103

Judd, J. W., 61

Karpinski, H., 222

Kendall, P. J., 125

Kendall, P. J., and Bell, A., 174

Kennard, A. S., 167

Kennard, A. S., and Woodward, B. B., 4

Kessler, H., 222

Kew, H. W., 17, 23, 32

Killarney fern, 115

Kinahan, G. H., 107-108, 129

Kirkdale Cavern, remains in, 54

Kobelt, W., 49, 56, 58, 62, 75, 195-196, 212, 281, 297, 303, 321-323

Koeppen, F. T., 51, 62, 222, 248

Krasan, F., 347

_Lagomys_, origin of, 41, 203

_Lagopus_, 334

_albus_, 91, 142, 334

_hyperboreus_, 334

_leucurus_, 335

_mutus_, 91, 142, 334

_rupestris_, 334

_scoticus_, 334

_Laminifera_, sub-genus of _Clausilia_, 48

Lamplugh, G. W., 107-108

Land-connection between America and Northern Europe, 61

British Islands and France, 59-60

British Islands and Scandinavia, 61

Europe and North Africa, 61

Europe and North America, 61, 168-172

Greece and Asia Minor, 61

Greenland and Europe, 155, 159

Ireland and Spain, 110

Land Mollusca, migrations of, 9

Land shells, West Indian, 21

Lartet, E., 51, 73, 107, 150

Lemming, range of, 138-140

_Leontopodium alpinum_, 266, 342

_Lepus_, 27, 29, 31

_alpinus_, 136

_cuniculus_, 27, 31, 291

_europaeus_, 29, 316

_glacialis_, 136

_lacostei_, 291

_variabilis_, 2, 29, 136, 315

Lepidoptera, range of, 159-160

Arctic, 200

surviving Glacial period, 54

Leuckart, R., 176

Lewis, Carvill, 129

_Limax_, 284

_marginatus_, 284

_maximus_, 284

_Limnocalanus macrurus_, 179

_Linyphia insignis_, 94

"Loess" fauna, 75, 196

Lomas (_vide_ Herdman and Lomas)

Loven, S., 177

Lusitanian flora in British Islands, 288, 306-307

spiders, 301

Lydekker, R., 14, 22, 32, 58, 157, 183, 202, 248, 252

Lyell, Sir C., 14, 22

_Lyrurus tetrix_, 336

_Macroptychia_, sub-genus of _Clausilia_, 48

Madeira, 19

molluscan fauna of, 25

remains of a continent, 19

Major, Forsyth, 45-46, 280, 282

Mallet, R., 86

Mammoth in Siberia, 214-217

range of, 252-253

Mammals, dispersal of, 9, 21

_Mantis religiosa_, 269

Maps, general plan of, 8

Margaritana, 93