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R. F. t SCHARFF, B.Sc., PH.D., F.Z.S. 

Keeper of the Natural History Collections, Science and Art Museum t 

Dublin ; Member of the Royal Irish Academy ; Corresponding 

Member of the Senckenbergische Naturforschende 








INTRODUCTION .................. 1-36 



THE FAUNA OF BRITAIN ... ......... 89-13! 


THE ARCTIC FAUNA ............. .. 132-188 


THE SIBERIAN MIGRATION ............ 189-244 


THE ORIENTAL MIGRATION ... . ........ 245-286 


THE LUSITANIAN FAUNA ............ 287-308 


THE ALPINE FAUNA ... ............ 

BIBLIOGRAPHY ............... 35^354 

INDEX .................. 355-364 



OUR knowledge of the present and past fauna of 
Europe is as yet insufficient to indicate with precision 
the original homes of its component elements, but I 
hope that the lines of research laid down here, and 
the method of treatment adopted, will aid zoologists 
and geologists in collecting materials for a more com- 
prehensive study of the history of our animals. I 
trust also that a fresh impulse will be given by the 
publication of this book to the study of the Geo- 
graphical Distribution of Species. Collectors of 
Beetles, Butterflies, Shells, and Fossils may derive 
some useful hints by its perusal and thus direct their 
studies, so as to add, by accuracy in observation, 
to our knowledge of the former geographical re- 
volutions which have moulded our islands and con- 
tinents. To geographers, a survey of some of the 
more important changes in the distribution of land 
and water in past times based upon the composi- 
tion of our fauna will be interesting. The subject, 
however, is a complex one. I have ventured to 
indicate a suitable method of treatment, and as such 
this attempt to elucidate the history of the European 
fauna should be received. 


This work was written as the outcome of a paper 
published in the Proceedings of the Royal Irish 
Academy (3rd series, vol. iv., 1897), "On the Origin 
of the European Fauna." A summary of that paper 
appeared in Nature (vol. Ivi., 1897), and fuller extracts 
of more important parts, with some criticisms, in the 
Geological Magazine (N.S., sec. iv., vol. iv., 1897). I 
freely acknowledge the value of these criticisms, 
which have largely assisted me to amplify and to 
improve upon the ideas laid down in the paper. 

I have found that it greatly facilitates comprehension 
of the arguments used, to give a few maps indicating 
in a general way the extent of former seas and 
continents. I may in this way, as Mr. Kendall 
has pointed out, have submerged many square 
miles of land which had never been covered by the 
sea, at least not within recent geological times, 
but the maps were intended as illustrations of my 
views in a broad spirit only. 

Some zoologists may be surprised that, in some 
cases, I have not followed the latest views in revised 
nomenclature. I felt that in a work of this kind 
it was of supreme importance to employ names still 
current in our leading text-books, such as Lepus 
variabilis for the Mountain Hare, instead of Lefius 
timidus. After each chapter I have endeavoured to 
give a short summary of contents, while a biblio- 
graphy of the principal works and papers consulted 
will be found at the end. I should also acknowledge 
the aid which I have received from such excellent 


works of reference as the British Museum Catalogues 
of Birds, by Dr. Bowdler Sharpe, and those of Reptiles, 
Amphibia, and Fishes, by Dr. Giinther and Dr. 
Boulenger. The valuable works en Mammalia by 
Sir W. Flower, Mr. Lydekker, Mr. Grev, and Dr. 
Trouessart, were indispensable to me. 

To Sir William Flower, Mr. Lydekker, Professor 
Sars, and Professor Smitt, I am especially indebted 
for allowing me to reproduce drawings from their 
works, and to my friend Mr. Welch for some beauti- 
ful photographs. The Council of the Royal Irish 
Academy also kindly gave me permission to reprint 
the maps used in illustration of my paper. Professor 
Haddon first suggested my writing this book, and 
gave me many useful hints ; and great assistance was 
rendered me by my colleague, Mr. G. H. Carpenter, 
in revising the proofs. To both of these kind friends 
I desire to acknowledge my deep sense of gratitude. 





/ EVERY student of natural history, whether he be 
' interested in birds, butterflies, or shells, contributes 
his share of facts which help to show how the fauna 
of his country has originated. The capture of a 
Swallow-tail or of a Marbled White Butterfly in Eng- 
land at once furnishes material for reflection as to the 
reason of its absence from Scotland and Ireland. 
Why should the Nightingale allow its beautiful song 
to be heard in England, and never stray across 
the Channel to the sister isle or cross the borders of 
North Britain? Lovers of bird-life and sportsmen, 
who have observed the habits of the Ptarmigan in the 
wild mountain recesses of Scotland, are aware that 
nowhere else in the British Islands do we meet with 
this interesting member of the grouse family, and 
many no doubt have allowed their minds to dwell 
upon the causes of its singularly local distribution. 


All these animals have a wide range in other parts 
of the world. In past times, before man began to 
make observations on the geographical distribution of 
birds and butterflies, or even before the appearance of 
man in Northern Europe, they may have lived all 
over the British Islands. For some reason or other 
they are perhaps dying out or withdrawing towards 
their original home, which may either be northward, 
or to the east or south. If we had some clue as to 
their former history from fossil evidence or, in other 
words, if their remains had been preserved to us in 
geological deposits, we should have less difficulty in 
deciding this problem. But butterflies are scarcely 
ever preserved in a fossil state, and birds very rarely. 
We know little or nothing, therefore, of their past 
history from direct evidence, and are obliged to trust 
to indirect methods of research which will be indi- 
cated later on. 

X^Mammals and Snails tell us their story more plainly. 
* The bones of the former and the shells of snails are 
easily preserved, and thus furnish us with the neces- 
sary data as to their past history, for we find them 
abundantly in most of the recent geological deposits. 
Among the mammals of the British Islands there 
are some instances of distribution which much 
resemble those I have quoted. Thus the Arctic Hare 
(Lepus variabilis] is in the British Islands confined to 
Ireland and to the mountains of Scotland ; and if it 
were not for the fact that its bones have been dis- 
covered in a cave in the south-west of England, we 


should perhaps never have known that, formerly, it 
must have inhabited that country as well. Of other 
mammals we possess fossil and also historical evi- 
dence of their having once lived in these islands. 
Such are the Wolf and the Wild Boar, both of which 
were abundant in Great Britain and Ireland. The 
latter is a distinctly southern species. We assume 
this, because its remains have never been found in 
high northern latitudes ; nor does it now occur in 
Northern Europe or Northern Asia, whilst all its 
nearest relatives live in sub-tropical or tropical 
climates. The Arctic Hare, on the contrary, has 
probably come to us from the north. Its remains 
are unknown even in Southern Europe, and the 
more we approach the Arctic Regions, the more 
abundant it becomes. Thus we have here two 
instances of British mammals, one of which, the 
Wild Boar, has died out as it were in a southerly 
direction ; whilst the other, the Arctic Hare, is ap- 
parently retreating towards the north. 

There are also some British mammals of which we 
have no fossil history, at least of which no remains 
have as yet been found in these islands. Such a 
one is the Harvest Mouse (Mus minutus). It has 
a somewhat restricted range in England, and only 
just crosses the Scottish border in the east. From 
the rest of Scotland and from the whole of Ireland 
it is absent. To judge from this distribution, in 
connection with the fact of its being unknown as a 
British fossil species, it is probably a late immigrant 


to England, and has not had time to spread, through- 
out Scotland at any rate. But it is also absent 
from Scandinavia, from the Spanish peninsula, from 
almost the whole of Italy and the Alps, as also from 
the Mediterranean Islands, whilst the little mouse 
occurs abundantly right across Siberia. We shall 
learn more about centres of dispersion later on ; 
meanwhile I should mention that such a distribution 
indicates that the Harvest Mouse has most likely 
originated in the east, and has spread from there 
westward in recent geological times. 

Conchologists have long ago been acquainted with 
the fact that many molluscs, for example the so-called 
"Stone-cutter" Snail (Helix lapicida) and the "Cheese 
Snail " (Helix obvoluta), have a very restricted range 
in the British Islands. Both are entirely absent 
from Scotland and Ireland, the Cheese Snail being 
confined to South-eastern England. The Stone- 
cutter has rather a wider range, is even known from 
a Welsh locality, and is met with as far north as 
Yorkshire. Their distribution would indicate, there- 
fore, that while both are recent immigrants, the 
Cheese Snail is probably the last comer. This 
supposition is in so far supported by fossil evidence, 
as the latter is unknown in the fossil state, whilst the 
Stone-cutter has been described by Messrs. Kennard 
and Woodward (p. 243) a as occurring in the cave 
deposit known as the Ichtham fissure, and also from 

1 The numbers in brackets throughout this work refer to the page- 
number in the Bibliography at the end. 


several English pleistocene and holocene deposits. 
The Stone-cutter can scarcely be looked upon as a very 
recent immigrant in the light of this evidence, though 
we have no proof of its having ever had a much 
wider range in the British Islands than it has to-day. 

Among the lichens, which so abundantly cover the 
rocks and trees in South-western Ireland, and which 
impart such a characteristic feature to the scenery, 
we find a beautifully spotted slug (Geomalacus macu- 
losus}. 1 It is a stranger to the rest of the British 
Islands, and indeed occurs nowhere else in Northern 
Europe. We have to travel as far as Northern 
Portugal before we again meet with it, and it is 
there also that its nearest relations live. 

Many more similar examples might be quoted, but 
enough, I think, has been said to show that the 
British fauna is made up of several elements whose 
original homes may lie widely apart and in different 
directions. We have fossil evidence that some of 
the northern species, and also a few of the southern 
ones, have become extinct within comparatively 
recent times ; others are apparently on the verge of 
extinction, whilst many not only maintain their 
position in the constant struggle for existence, but 
are even extending their range. 
/X^The problem of tracing the origin of the British 
fauna, or at least that of some of the more 
characteristic members of every. section or element, 

1 A map giving its exact distribution in Ireland will be found on 
p. 300, and a figure of the slug on p. 298. 


appears at first a somewhat difficult task. Indeed, 
the means of dispersal of the various groups of 
animals are so different that it occurred to me it 
might be better to deal with the mammals, the 
birds, the reptiles, and so forth, all separately. This 
idea I have attempted to follow to some extent, with 
most satisfactory results. The British fauna of the 
present day is no doubt complex, but no more so 
than the fauna of the most recent of our geological 
deposits the Pleistocene. However, when we go 
back still further and look at the earlier Tertiary 
remains, we find the fauna becoming less complex. 
Northern species disappear, and the strata are 
entirely filled with the remains of southern animals 
and plants. Geologists indeed are quite unanimous 
in their belief, that the fauna of the British Islands 
during the earlier epochs of the Tertiary Era was a 
southern one ; that it then gradually became more 
temperate, until at last, in more recent times, decidedly 
northern forms invaded the country. These seem 
to have driven out to some extent at least the 
southern species; but more recently again, the 
southerners, reinforced by an eastern contingent, 
appear to have gained territory and are advancing 
into the area held by the northerners. The eastern 
invasion does not seem to have affected Ireland at 
all, and we find the country there divided between the 
southern and northern animals. We can thus roughly 
construct a map as I have done here, showing, by 
means of horizontal and sloping lines, the principal 


areas inhabited at the present time by the species of 
northern, southern, and eastern origin (Fig. i). 

FIG. i. Map of the British Islands, indicating approximately the 
areas inhabited by the northern, southern, and eastern animals. 
The horizontal lines represent the areas of northern species, the 
sloping lines those of southern and eastern ones. 

In the problems which are being discussed in this 
work I have often found it of advantage, in order to 


facilitate the comprehension of the arguments used, to 
give maps. Some of these represent the geographical 
conditions at the particular epoch referred to in the 
text, but they merely claim to give a general idea. 
There was never any intention to make them corre- 
spond with all the data of which we have geological 
evidence. They are what I might call "diagrammatic." 
In comparing them with reconstructions of former 
physical geography such as have been attempted from 
time to time, I hope geologists will therefore deal 
leniently with the faults I may have committed, and 
remember that the maps are " impressions," or " dia- 
grams," and not faithful representations of all the 
geographical revolutions witnessed by some of our 
remote forefathers at any particular period. 
r. The knowledge we gain from a study of the British 
Tertiary deposits enables us to affirm positively that 
both the eastern and the northern species arrived in 
these islands comparatively recently, but that the 
southern forms must have migrated northward from 
the Continent long ages ago. Since the northern 
and the eastern migrations that is to say, those 
coming from the north and east were the last to 
arrive in Northern Europe, the remains of the animals 
contained in the most recent deposits of that portion 
of our continent will furnish us with a clue as to the 
extent of the area inhabited by them. This is not 
all, however. It is also possible to discover from these 
remains the direction which the animals that they 
belonged to came from. As we shall learn later on, 


a migration on a vast scale entered Europe during 
the Pleistocene epoch the most recent of the geo- 
logical epochs, during which great extensions of 
glaciers occurred in the mountainous regions of 
Europe. The latter period is known to us as the Ice 
Age or Glacial period. This will be described more 
fully in Chapter II., meanwhile I may mention that 
we presume that this migration came from the east, 
because no remains of the members of that particular 
fauna are known from Spain, Southern Italy, Scandi- 
navia, Ireland, or from the Balkan peninsula. The 
number of species evidently belonging to this same 
migration, moreover, become fewer as we proceed 
westward, and a large proportion of them still inhabit 
Northern Asia, though most of them are now extinct 
in Europe. After having thoroughly studied such a 
recent geological migration, we learn to understand 
others better, though the more ancient they are, the 
fewer are the traces and the more difficult are they to 

Then again we have to take into consideration the 
fact, that whilst mammals, particularly the larger 
herbivores, are forced to migrate frequently owing 
to scarcity of food or temporary changes of climate, 
many of the invertebrates remain practically unaffected 
by either. Most of our land mollusca, for instance, 
are satisfied with meagre provender, and stand ex- 
tremes of climate well, as long as there is sufficient 
moisture. As a result of their peculiar disposition, 
many of them, no doubt, have survived through 


several geological epochs, and have witnessed vast 
geographical revolutions in their immediate surround- 
ings, whilst mammals are comparatively short-lived. 
Being driven from one country to another, and ex- 
posed to innumerable enemies, new types appear and 
old ones rapidly vanish ; in fact, there are almost 
constant changes in the mammalian fauna as we 
pass from one epoch to another. 

I have until now referred more particularly to the 
British fauna and the North European in general, 
because the history of our own animals interests 
us all more than those of any other European area. 
It is, moreover, preferable to commence our inves- 
tigations into the origin of the European fauna 
by the study of a small district. This should, if 
possible, be an island. If we took a slice of the 
continent like France or Germany, we should find 
the problem more complex. Instead of choosing 
the British Islands, we might, however, take an island 
like Corsica or Sardinia. In either of these we should 
discover peculiarities in the composition of their fauna 
precisely similar to those which I have indicated to 
be present in the British fauna. We should find 
probably a more striking endemic 1 element, which 

1 The term endemic will be employed throughout this work as applied 
to species peculiar to a country and not found elsewhere. Autoch- 
thonous will be used in speaking of a species which has originated in a 
country to which, however, it is not peculiar ; e.g. , the Chamois is an 
autochthonous Alpine species, but occurs also in the Pyrenees and 
Caucasus. An indigenous species is one native to a country, as 
opposed to the term " introduced," and is applicable to all species 
v\hich have reached it by ordinary migration. 


with us is so meagre that it can almost be left 
unnoticed ; the main features, however, remain 
nearly the same. The fauna of both of these 
islands is composed of a strong southern element, 
of an eastern and a northern one, and in addition 
we have here species whose ancestors lived in 
Western Europe. 

Before investigating more minutely the problems 
suggested by the composition of the faunas of these 
insular and also of some continental areas, it is 
necessary that we should thoroughly understand 
all about the migrations of animals. One of the 
principal objects of this work is to show how the 
autochthonous animals of Europe, /.&, those which 
have originated there, may be distinguished from the 
immigrants, and to trace the latter to the home 
of their ancestors. But in doing so, it is necessary 
to refer to the many important geographical changes 
which have occurred in Europe during the latest 
geological epochs. The study of the geographical 
distribution of the European fauna, as expounded in 
this work, will in many instances confirm the theories 
as to geographical changes based upon geological 
foundations. But in every case the views herein 
advocated are founded upon the geographical distri- 
bution of living and extinct organisms alone. 

A terrestrial mammal like the deer can, under 
ordinary circumstances, only reach one part of a 
country from another by walking or running to it; 
but a beetle, such as the cockchafer, has two different 


modes of progression. It may walk or fly. In both, 
however, there is a third mode of transport an in- 
voluntary one. The deer may be suddenly seized by 
a flood whilst crossing a river, and carried far away 
without necessarily coming to grief. The beetle in 
a similar manner could be transported to a distant 
country, or it might be caught in a whirlwind and 
blown hundreds of miles off. 

We may thus distinguish between the natural or 
active and the accidental or passive means of distribu- 
tion of animals. The active mode of dispersal again 
may be only migratory, as in most animals, or periodic 
and migratory, as in some birds and fishes. It is of 
course the tendency of every species to spread in all 
directions from its original home, provided it does not 
encounter obstacles, such as want of food, unsuita- 
bility of climate or soil, or barriers such as mountains, 
rivers, or the sea. Birds might be thought to be little 
interfered with by any of these barriers, but, as Dr. 
Wallace has shown, they are almost as much affected 
by them in their distribution as mammals are. 

This then is the ordinary migratory distribution. 
Periodic distribution obtains with migratory birds 
and fishes. The annual flight of swallows to their 
northern summer residence comes under the heading 
of periodic migration or distribution, but apart from 
this, the swallow must seek to extend its range by 
the ordinary method, like every other animal. 
Similarly, the herring migrates periodically into 
shallow water to spawn, only to return again to its 


deeper home, where, as its numbers increase, there 
must be a tendency to spread. We have in these 
cases, therefore, both a periodic and an ordinary 
movement of migration. 

Now, in studying the composition of a fauna, and 
especially its origin, it is of the utmost importance to 
be able to determine approximately the percentage of 
accidental arrivals and of the ordinary migrants 
that is to say, of those which have reached the 
country owing to accidental distribution, and of 
animals which have adopted the usual course of 
migration. It is of all the more import to review 
this subject of accidental, or, as Darwin called it, " the 
occasional means" of distribution, as both he and 
Dr. Wallace have, I venture to think, somewhat over- 
estimated its significance. No one doubts that acci- 
dental transportal takes place, but the question is 
whether the accidentally transported animals arrive 
living and reach a spot where suitable food is pro- 
curable, and whether they are able to propagate their 
own species in the new locality. For it must be clear 
to anybody that the accidental transportal of a beetle 
or of a snail to a new country cannot affect its fauna 
or add one permanent member to it unless all these 
conditions are fulfilled. As a matter of fact, only 
exceedingly few instances are on record of man 
having witnessed, for example, the accidental 
transportal across the sea to an island of a live 

To mention an example, Colonel Feilden informs 


us (Zoologist, 1888) that, when living on the island 
of Barbadoes, an alligator arrived one day on the 
shore, and at the same time a tree measuring 40 feet 
in length, which was recognised as a Demerara species, 
was likewise stranded. He thinks that there can be 
no doubt that the alligator, which was alive when it 
reached Barbadoes, was transported by the tree, thus 
covering a distance of 250 miles from the nearest 
land. Numerous observations on the accidental 
transportal of seeds and tree-trunks from one island 
to another, and from a continent to an island, have 
been recorded, and even on our own shores we may 
witness the occasional arrival of such vegetable pro- 
ducts from a far distant land. On the west coast of 
Ireland it not unfrequently happens that large West 
Indian beans are stranded, and in this as well as in 
many other similar cases the seeds have often proved 
none the worse for their prolonged immersion in sea- 
water. That locusts are sometimes blown to great 
distances from the land is not so surprising, since 
their power of steering through the air is very limited. 
Darwin mentions (p. 327) having caught one 370 
miles from the coast of Africa, and that swarms of 
them sometimes visited Madeira. Sir Charles Lyell 
relates that green rafts composed of canes and brush- 
wood are occasionally carried down the Parana River 
in South America by inundations, bearing on them 
the tiger, cayman, squirrels, and other quadrupeds. 

But though actual observations of such abnormal 
instances of the dispersal of animals are few, many 


experiments have been made to demonstrate the pos- 
sibility of a passive transportal of species over wide 
distances. It was especially Darwin who gave a great 
stimulus by setting the example to those interested 
in natural history in the conduct of such researches. 
He was struck by the fact that, though land- 
shells and their eggs are easily killed by sea-water, 
almost all oceanic islands, even the smallest and most 
isolated, are inhabited by them, and felt that there 
must be some unknown but occasionally efficient 
means for their transportal (p. 353). To quote his 
words: "It occurred to me that land-shells, when 
hibernating and having a membranous diaphragm 
over the mouth of the shell, might be floated in chinks 
of drifted timber across moderately wide arms of the 
sea. And I find that several species in this state 
withstand uninjured an immersion in sea-water during 
seven days: one shell, the HelLr pomatia, after having 
been thus treated and again hibernating, was put into 
sea-water for twenty days, and perfectly recovered. 
During this length of time the shell might have been 
carried by a marine current of average swiftness to a 
distance of 660 geographical miles. As this Helix 
has a thick calcareous operculum, I removed it, and 
when it had formed a new membranous one, I again 
immersed it for fourteen days in sea-water, and again 
it recovered and crawled away. Baron Aucapitaine 
has since tried similar experiments : he placed one 
hundred land-shells, belonging to ten species, in a 
box pierced with holes, and immersed it for a 


fortnight in the sea. Out of the hundred shells, 
twenty-seven recovered. The presence of an oper- 
culum seems to have been of importance, as out of 
twelve specimens of Cyclostoma elegans which it thus 
furnished, eleven revived. It is remarkable, seeing 
how well the Helix pomatia resisted with me the salt- 
water, that not one of fifty-four specimens belonging 
to four other species of Helix tried by Aucapitaine, 
recovered. It is, however, not at all probable that 
land-shells have often been thus transported ; the feet 
of birds offer a more probable method." 

We have here positive evidence that such shells as 
Helix pomatia and Cyclostoma elegans might easily be 
transported to an island from the mainland. The 
former occurs in France, Holland, and England, and 
the latter all along western continental Europe and 
England. And yet neither of these species inhabits the 
Canary Islands, Madeira, or Ireland, none of which are 
at too great a distance from Europe to be within easy 
reach for a floating object. The fact that Cyclostoma 
elegans does not live in Ireland is of particular interest 
in connection with the floating-theory just quoted, as 
on all sides of Ireland dead specimens have been picked 
up on the shore, showing that marine currents carry 
specimens and have thus transported them for 
countless centuries. Nevertheless the species has not 
established itself in Ireland. If such a fate meets a 
land-shell of the type of Cyclostoma elegans, it may be 
asked, with some justification, what chance slugs or 
the smaller non-opcrculated species would have to 


reach an island like Ireland alive from the main- 
land, and to colonise it successfully. 

Both slugs and their eggs are killed by a short 
immersion in sea-water, as I have proved experi- 
mentally. I have also subjected slugs, in the act of 
crawling on twigs, to an artificial spray of sea-water. 
This seemed to irritate their tender skins to such an 
extent that they curled themselves up, released their 
hold on the twig and let themselves drop to the 
ground. If we supposed, therefore, that a slug had 
successfully reached the sea, transported on a tree- 
trunk, the moisture would tend to lure it forth from 
its hiding-place under the bark, whilst the mere spray 
would prove fatal to its existence. Those species of 
snails and slugs which lead an underground existence, 
rarely venturing above ground, such as Testacella and 
Coecilianella, would have even less chance of being 
accidentally carried to some distant island. 

The suggestion advanced by Darwin (p. 353), that 
young snails just hatched might sometimes adhere to 
the feet of birds roosting on the ground and thus be 
transported, appears to me so extremely improbable 
as to be scarcely worth serious consideration. Indeed, 
as Darwin himself acknowledged later on, it does not 
help us very much to suggest possible modes of 
transport. What we require is direct evidence. How 
far we are, however, from obtaining it, may be inferred 
from Mr. Kew's remark (p. 119), that " we have 
little or no actual evidence of precise modes of 
dispersal even for short distances on land." 


" 'Hv> 

A very curious statement was made by a well- 
known French conchologist, the late M. Bourguignat, 
with regard to introductions of mollusca. Whether 
he had any actual facts collected in support of it, I 
cannot say, but he maintained that species accidentally 
transported, with the exception of those under mari- 
time influence, can only be acclimatised from north to 
south, and not from south to north from east to 
west, but not from west to east (p. 353). 

The whole theory of the accidental or abnormal 
dispersal of mollusca appears to have been originated 
by Darwin, in order to account for their presence on so- 
called Oceanic islands. His views v/ere strongly sup- 
ported by Wallace, who defines these islands (p. 243) 
as those which are of volcanic or coralline formation 
usually far from continents, entirely without indi- 
genous land mammals or amphibians, but with a fair 
number of birds and insects, and usually with some 

I do not wish it to be understood that I am in any 
way undervaluing the great works of these dis- 
tinguished naturalists. Darwin's views have had 
more influence in advancing Zoology than those of any 
man, and his fame is unassailable. Nevertheless, I 
feel that his theories regarding the origin of the faunas 
of oceanic islands require revision. 

The formerly prevalent belief of the permanence of 
ocean basins has been shaken by the utterances of 
some of the greatest geologists of our day, whilst 
many positively assert that what is now deep sea 


of more than 1000 fathoms was dry land within 
comparatively recent geological epochs. Thus the 
Azores are classed by Darwin and Wallace among 
the oceanic islands that is to say, among such 
as have received their fauna and flora by flotsam 
and jetsam. But Professor Neumayr believes, on 
geological grounds, that the Old and New Worlds 
were connected by a land-bridge during Tertiary 
times right across the Atlantic, and that the 
Canary Islands, Madeira, and Azores (p. 547) 
are the last remnants of this continent. This 
meets with the entire approbation of Dr. von 
Ihering, who has recently re-investigated the sub- 
ject from a faunistic point of view (p. 135). Take 
another instance of one of Wallace's most typical 
oceanic islands, the Galapagos Group. Their fauna 
and flora have recently been most thoroughly 
re-explored by an American expedition, the result 
of which, according to Dr Baur, goes to show 
that these islands must have formed part of the 
mainland of South America at no distant date. The 
fauna and flora are therefore to be regarded as having 
reached them in the normal mode, viz., by migration 
on land. According to Mr. Beddard (p. 138), it is 
difficult to see how earthworms could be transported 
across the sea. Floating tree-trunks have been ob- 
served far out at sea, but unless the water remained 
absolutely calm during the long period necessary for 
the drifting by currents so that no splashing occurred, 
the worms would probably be killed. Yet earthworms 


do occur on oceanic islands. It is indeed quite 
possible that our views with regard to the origin of 
the remainder of the Pacific Islands may change very 
materially, and once more revert to what Dr. Gould 
expressed nearly fifty years ago in the following 
words : " From a consideration of the land-shells on 
the Pacific Islands, it seems possible to draw some 
fair inferences as to the relations of the lands which 
once occupied the area of the Pacific Ocean, and 
whose mountain peaks evidently now indicate or 
constitute the islands with which it is now studded." 
Indeed Dr. von Ihering goes so far as to positively 
state that in his opinion the Polynesian Islands are 
not volcanic eruptions of the sea floor, which being 
without life were successively peopled from Australia 
and the neighbouring islands, but the remains of 
a great Pacific continent, which was in early 
mesozoic times connected with other continental land 
masses (a, p. 425). 

Before coming to a decision on the part played by 
flotsam and jetsam in the constitution of an island 
fauna, those who have studied the problem on the 
spot should, however, have a voice in the matter. 
And though, from my experience in northern latitudes, 
I feel sure that island faunas there are but slightly 
affected by occasional dispersal of species, Mr. 
Hedley, who has made the fauna of the Pacific 
Islands his special study, assures me that drift 
migration plays an important rdle in that region. 
I hope we may soon have a more detailed account 


of his particular observation bearing on this interest- 
ing subject. 

On the other hand, Mr. Simpson, who has gained 
considerable experience of oceanic dispersal in the 
West Indian region, though he acknowledges having 
often noticed bamboo rafts, which would be suitable in 
the transportal of invertebrates, nevertheless does not 
attach much importance to this means of distribution. 
" The fact," he remarks, " that the operculates (oper- 
culate land-shells) form so large a proportion of the 
Antillean land-snail fauna, that a majority of the 
genera are found on two or more of the islands and 
the mainland, while nearly every species is absolutely 
restricted to a single island, appears to me to be 
very strong testimony in favour of a former general 
land connection" (p. 428). 

Amphibians are affected in the same manner by 
sea-water as slugs are. The accidental transportal of 
an amphibian from the mainland to an island is there- 
fore almost inconceivable. And the presence of frogs, 
toads, and newts in the British Islands, in Corsica 
and Sardinia, indicates, if nothing else did, that all 
these islands were at no distant date united with the 
continent of Europe. 

As regards the terrestrial reptiles, the case is some- 
what different. Many of them readily take to the 
sea, and, as probably all snakes and some Hzards are 
able to swim, it is possible that sometimes, though 
very rarely, they might reach islands if not too far 
from a continent. Instances of accidental transportal 


of land-reptiles to islands have actually been observed. 
But the fact of the occurrence of such instances by no 
means proves that reptiles thus conveyed are able to 
establish themselves permanently in their new home. 
Sir Charles Lyell records in his Principles of Geology 
that a large boa-constrictor was once seen floating to 
the island of St. Vincent, twisted round the trunk of 
a tree. It appeared so little injured by its long 
voyage from South America, that it captured some 
sheep before it was killed. 

Mammals might be accidentally conveyed to islands 
on such rafts as have been described by Sir Charles 
Lyell, and there are instances on record of their 
having crossed short distances of sea by swimming. 
Elephants and also deer and pigs are good swimmers, 
the former having been known to swim for six hours 
at a stretch. "But," remarks Mr. Lydekker (p. 13), 
"it may be assumed that about twenty miles is the 
utmost limit which mammals are likely to cross by 
swimming, even when favoured by currents. Such 
passages as these must, however, be of very rare 
occurrence, for a terrestrial mammal is not likely 
to take it into its head to swim straight out to sea 
in an unknown direction. Moreover, supposing a 
mammal, near to a particular island, to have arrived 
there by swimming, unless it happen to be a pregnant 
female, or unless another individual of the same 
species but of the opposite sex should arrive soon 
after (a most unlikely event), it would in due course 
die without being able to propagate its kind." 


All zoologists, indeed, are quite in accord with Dr. 
Wallace's view as expressed in Island Life (p. 74). 
" Whenever we find that a considerable number of 
the mammals of two countries exhibit distinct marks 
of relationship, we may be sure that an actual land- 
connection, or at all events an approach to within 
a very few miles of each other, has at one time 
existed." As all the European islands come under 
this category, their mammals exhibiting distinct 
relationship with those on the European continent, 
they all have been connected with it formerly. 

Perhaps the most powerful of all agents in the 
transportal of species by accidental means is man. 
But his actions may be accidental as well as in- 
tentional. We have therefore to distinguish between 
the animals disseminated all over the world by pure 
chance, and those which have been introduced into 
new countries purposely. Invertebrates, such as 
snails, centipedes, woodlice, beetles, and cockroaches, 
are constantly being unintentionally carried with 
vegetables, fruit, trees, and with timber from one 
country to another. Earthworms are sometimes 
transported in the balls of earth in which the roots 
of trees are enveloped. As regards molluscs, Mr. 
Kew believes (p. 178) that during the last three 
centuries at least, human agency has influenced 
their disposal more than all other causes taken 
together. A large number of species of invertebrates 
in America are said to owe their existence in that 
country to accidental introduction by man. In 


most cases, however, no particular reason can be 
assigned why they should have been thus introduced, 
and as a matter of fact there are always individual 
differences of opinion as to the precise number of 
such. Certain it is, that though the number of 
supposed introductions from Europe to America 
is very large, those which have been carried from 
America to Europe is exceedingly small. In fact, 
I remember only two instances of accidental animal 
importations from America which have firmly 
established themselves in Europe, viz., a small 
fresh-water mollusc, Planorbis dilatatus^ and the 
much-dreaded vine-pest, Phylloxera vastatrix. 

As a rule the animals die out very shortly after 
their arrival on foreign soil. Many instances, 
nevertheless, are on record, especially in the case 
of molluscs, where snails thus transported have not 
only survived but are apparently in a flourishing 
condition and spreading. Helix aspersa, for example, 
our large garden snail, has been naturalised in many 
foreign countries by French and Portuguese sailors, 
who had taken them on board their ships as food. 

It certainly cannot be denied that a number of 
species among almost all groups of invertebrates have 
been unintentionally conveyed by man from Europe 
into foreign countries. It has been proposed by Dr. 
von Ihering to apply the term "cenocosmic" to those 
species which have become spread all over the world 
through artificial means, and thus to distinguish 
them from cosmopolitan ones which have attained a 


similar range naturally. The latter he calls " palin- 
cosmic " species (a, p. 422). Many so-called ceno- 
cosrnic ants are believed by Dr. von Ihering to be 
palincosmic. We are altogether too apt to regard 
cosmopolitan as synonymous with introduced, and we 
should hesitate before concluding that because one of 
.our common European species occurs in Australia or 
South America, it must have been transported there 
recently by human agency. Some of our widely- 
distributed forms are probably of very great antiquity, 
and may have spread to distant lands in early Ter- 
tiary times, when a different state of the geographical 
conditions enabled them to do so. 

I cannot quote a more appropriate instance 
than the molluscan fauna of Madeira. No less 
than thirteen of the Madeiran snails are looked 
upon as having been introduced from Europe by 
human agency, on the sole evidence that these 
happen to be common European species. Yet the 
correctness of this supposition must be questioned 
in face of the interesting observation made by 
Darwin (p. 357), "that Madeira and the ad- 
joining islet of Porto Santo possess many distinct 
but representative species of land-shells, some of 
which live in crevices of stone ; and although large 
quantities of stone are annually transported from 
Porto Santo to Madeira, yet this latter island has not 
become colonised by the Porto Santo species. Never- 
theless, both islands have been colonised by European 
land -shells, which no doubt had some advantage 


over the indigenous species." Darwin, therefore, 
meets the evident anomaly by suggesting that the 
European species are supposed to possess some 
advantages as colonisers. But the true explanation 
appears to me to lie in the supposition that the 
European land-shells found in the Madeiran Islands 
are all, or for the greater part, ancient forms which 
survived both there and on the continent, whilst the 
remainder of the forms inhabiting these islands are 
either such as are now extinct in Europe, or have 
become modified since their arrival there from the 
continent at a time when extensive land-connections 
allowed a free migration by land. 

The theory of accidental introductions is an ex- 
tremely popular one. It allows free scope to a host 
of speculations, and once the idea has taken firm 
root that a certain species is introduced, especially 
among the class of naturalists who by way of ex- 
periment are wont to create new centres of dispersion 
in their own neighbourhood, evidence to the contrary 
must be of the most convincing nature to shake the 
popular belief. Thus, it is almost regarded as an 
established fact by conchologists and others, that the 
fresh-water mussel (Dreyssensiapolymorpka) was intro- 
duced into England at the beginning of this century. 
Though it has been proved that this species is quite 
unable to live in pure sea-water, yet the view that it 
has been carried from the Black Sea ports to this 
country attached to the bottom of ships is maintained 
by many, whilst others incline to the theory that the 


shell came with timber. But Dreyssensia polymorpha 
was by no means always confined to the Caspian and 
Black Sea areas; it occurs abundantly in the lower 
continental boulder-clay (see p. 230), and no doubt it 
had at one time a much wider geographical distribution. 
It p appears to me possible, that it was able to maintain 
itself in certain fresh-water lakes and slow-flowing 
rivers in Northern Europe, from which it might have 
spread since the introduction of canals into Europe at 
the beginning of the century. As the larva of this 
fresh-water mussel is free-swimming, its propagation 
is much favoured by canals. Quickly-flowing rivers 
are fatal to its existence, since the delicate larvae are 
swept out to sea and perish. Such an hypothesis as 
this is strengthened by the fact of its recent discovery 
in a sandy layer fifteen feet below the present surface 
under the streets of London in a deposit which prob- 
ably, as Mr. Woodward remarks (p. 8), was accumu- 
lated in the early days of the city's existence. In 
spite of Mr. Woodward's interesting find, and Dr. 
Jeffreys' opinion, who always maintained that this 
shell was indigenous to England, popular belief still 
clings tenaciously to the introduction theory. 

Among man's intentional introductions into a new 
country, no instance is better known than that of the 
rabbit to Australia. Rabbits are entirely confined to 
Europe. In their transplantation to Australia they 
were carried to a country with a different climate and 
among new surroundings. Yet the rabbits flourished, 
and within comparatively few years increased to such 


an extent as to become a burden and pest to the 
country. It may be remembered though, that, owing 
to the complete absence of small carnivores, which 
act with us as a check upon the too rapid increase of 
this rodent, the speed with which it established itself 
in the new surroundings is not so very surprising. 

Many of the English settlers in the New World felt 
that America lacked the presence of our familiar 
birds. The homely sparrow was therefore brought 
over, with the result that the Agricultural Department 
of the United States is now devising means for its 
destruction, so rapid has been its increase. 

Similarly, the inhabitants of Jamaica, annoyed by 
the great profusion of rats in their island, sent over to 
India for a number of mongoose. These have deci- 
mated the rats since their arrival, but they have 
multiplied to such an extent as to be a serious 
menace to the native fauna. 

To give an instance nearer home, the Capercaillie 
(Tetrao urogallus) was successfully introduced into 
Scotland in 1837. From its different centres of 
distribution it is spreading in all directions where 
sufficient cover is obtainable. But this case differs 
from the others very materially, in so far as this bird 
was formerly a native of Scotland, and only became 
extinct during the last century. 

However, although there are many examples of un- 
doubtedly successful introductions by human agency, 
quite as many, or perhaps more, unsuccessful ones 
might be quoted. In fact, it is by no means easy to 


establish a species in any new locality. Frequently 
it happens that the species seems to be on the increase 
at first, but then there is a decline, and after a few 
years the new plantation has entirely vanished. In 
other cases, the species disappears immediately after 
the introduction takes place, or lingers on for many 
years if it receives special and uninterrupted pro- 

It may not be generally known that the English 
Hare {Lepus Europeans) is not found in Ireland, where 
the Mountain Hare (Lepus variabilis) alone occurs. 
Attempts to acclimatise the English species have 
been made in a number of places in Ireland, but 
many of them have been failures, and not one of 
them has been a signal success. 1 Similarly, the 
endeavour to introduce the French or Red-legged 
Partridge (Caccabis rufd) into Ireland has met with a 
like result. According to Dr. Day, it was tried during 
the summer of 1869 to naturalise the Sterlet (Acipenser 
ruthenus) from Russian waters into the Duke of 
Sutherland's River Fleet by importing artificially im- 
pregnated ova. From one hundred and fifty to two 
hundred lively young sterlets are said to have been 
turned out, but nevertheless the experiment met with 
no success. Several fortunately abortive efforts were 
also made in British rivers to establish Silurus glanis, 
a hideous monster of a fish, and quite unpalatable. 

1 I might refer any one more specially interested in these intro- 
ductions to an article on this subject in the Irish Naturalist of March 
1898, by Mr. Barrett- Hamilton. 


The Natterjack Toad (Bufo calamitd} has a very 
local distribution in the British Islands. In Ireland 
it is found only along the coast of Dingle Bay in 
County Kerry, where it is known among the peasantry 
as the Black Frog. There is no doubt about its 
being indigenous there, and though it has not spread 
beyond the very limited area of its habitat, the Irish 
climate cannot be said to be unsuited to its existence. 
Yet it seems to be extremely difficult to acclimatise 
it elsewhere, for though no less than sixty specimens 
were turned out in Phcenix Park, Dublin, about 
forty years ago, none of them were ever seen 
afterwards. They were placed in the vicinity of one 
of the lakes, so as to give them ample scope for 
breeding and developing the young, and in sur- 
roundings which were considered eminently suitable 
at the time. 

It has occasionally happened, too, that animals are 
introduced by kindly-disposed persons with the view 
of adding a species to their fauna, in complete 
ignorance of their previous existence in the country 
where they wished to naturalise them. Thus we are 
told that in the year 1699 one of the Fellows of 
Trinity College, Dublin, procured Frog's spawn from 
England in order to add that amphibian to the Irish 
fauna. It was placed in a ditch in the College Park, 
whence the species is supposed to have gradually 
spread all over the island. This story is quoted by 
many writers as the true history of the Frog in 
Ireland, and is given as an example of the rapidity 


with which animals spread. Unfortunately the would- 
be introducer seemed unaware that, according to 
Stuart's History of Armagh, the first Frog which was 
ever seen in Ireland made its appearance in a pasture 
field near Waterford about the year 1630, that is to 
say, seventy years before its introduction in Dublin. 1 
But even Stuart was mistaken in supposing that no 
Frog had ever been seen in Ireland before, since 
Giraldus Cambrensis, in his Topography of Ireland, 
mentions that a Frog was found in a meadow near 
Waterford in the year 1187. 

Certain British species of vertebrates are generally 
looked upon as introduced species, though we cannot 
trace any record of their first establishment, and it is 
quite possible that, though there was local extinction 
and subsequent local re-introduction, they are truly 
indigenous and may never have become totally 
extinct. Such are, for instance, the Rabbit (Leptts 
cuniculus) and the Pheasant (fhasianus colchicus}. 
The latter certainly had become naturalised in 
England before the Norman invasion. 

But cases of introduction such as those above re- 
ferred to are by no means confined to the vertebrates, 
similar instances among invertebrates being numer- 
ous enough. I am sure every naturalist is personally 
acquainted with a good number, and it is hardly 
necessary that I should quote in any detail after 

1 I should recommend those who are particularly interested in the 
full history of the Irish frog to read the notes on this subject contained 
in vol. ii. of the Irish Naturalist. 


what has been said on the subject generally. The two 
species of snails, Helix pomatia and Cyclostoma elegans^ 
both of which occur in England, and which I had occa- 
sion to mention among those experimented on by 
Darwin, were turned out in several suitable localities in 
Ireland by Thompson, but failed to establish them- 
selves. The former, according to Mr. Kew, was also 
introduced into Scotland and Norway, whilst fifty or 
sixty specimens were brought to Petersfield in Eng- 
land, but none of these trials at acclimatisation 
were successful. As among vertebrates, a large num- 
ber of the so-called successful introductions rest upon 
insufficient evidence. 

When we once more carefully review the evidence 
as to the undoubted difficulty attendant on intentional 
introduction of animals by human agency, placed as 
they often were in most suitable localities, we must 
feel that accidental introduction cannot play an im- 
portant r61e in the making of the fauna of any country. 
Especially is this the case with an island fauna. 
Vertebrates are almost altogether excluded, and in- 
vertebrates must arrive singly as a rule, often stranded 
on an inhospitable and unsuitable shore. Their 
chances of surviving a passage by sea, of finding 
suitable food and shelter and a mate in order to pro- 
create their species, appear to me infinitesimally small. 
Yet there may be some such cases. However, I quite 
agree with Mr. Andrew Murray a high authority 
on geographical distribution that "colonisation or 
'occasional dispersal is insufficient to account for the 


character of the faunas and floras of oceanic islands; 
and I believe that the normal mode in which islands 
have been peopled, has been by direct continuity with 
the land at some former period, or by contiguity so 
close as to be equivalent to junction " (p. 15). " That 
a slight intermixture," he continues, "due to Mr. 
Darwin's colonisation, occurs in many (probably in 
all) I am ready to admit ; and from instances to be 
afterwards noticed, I am disposed to reckon the pro- 
portions of such intermixtures in the flora, in the 
most favourable circumstances, at not more than two 
per cent. In the fauna I think it must be much less." 

Mr. Murray's views, though they relate only to 
oceanic islands, are likewise applicable to continental 
islands such as our own. I think we might take the 
admixture in the British fauna due to occasional, 
including human introduction, as amounting to five 
per cent. It is better to take a high estimate, so as to 
include all the species about whose native land there 
might be some reasonable doubt. Now of what 
importance, after all, is this five per cent? The 
remaining ninety-five per cent, of the species of 
animals belonging to the British fauna undoubtedly 
migrated to these islands in the normal way by land. 

It is of great importance, in dealing with the 
question of the origin of the British fauna, to 
thoroughly grasp this conclusion that ninety-five 
per cent, of the animals have reached us by land. 
We can afford in fact to ignore the five per cent, 
altogether. It is an insignificant factor. As regards 



the botanical aspect of the question, botanists are 
quite in accord with the zoologists, and entirely 
share their views in the belief of a former land- 
continuity between the British Islands and the 
Continent. " It cannot be denied," says Professor 
Blytt (p. 32), "that a plant of one or another 
species may, in an exceptional case, migrate, without 
human assistance, all at once, across large tracts 
of land and sea, and that such migration, if operating 
during geological periods, might introduce a number 
of species even into distant oceanic islands ; but 
when the question is of whole communities of plants, 
such as the above enumerated elements in our flora, 
then such an accidental and sudden transport across 
large tracts can only be conceived to be at all 
probable in the case of Arctic plants carried by 
drifting ice to a bare country without native flora ; 
as to the other species, we must imagine that the 
migration during the gradual change of climate has 
proceeded sloivly and step by step across connected 
tracts of cotmtty. In that manner we may assume 
that our country has in the course of time obtained 
its present covering of plants. Each of the above- 
named elements in our flora has doubtless its 
corresponding element in our fauna. The fauna 
and flora of a region stand in relation of com- 
plicated dependence to each other. The animals 
live on the plants. The fecundation of the plants 
takes place in a great degree by means of insects ; 
their seeds are often scattered by resident birds and 


quadrupeds. Everything indicates that conveyance 
1o small distances is the rule, and that sudden and 
long migration is the exception." 

The conviction which has been gained by zoologists 
and botanists, that the British Islands once formed 
part of the Continent, is based on the present British 
fauna and flora. The remains, however, of animals 
which used formerly to live in these countries, such as 
the Mammoth, the Irish Elk, the Cave Bear, and many 
others, tell us the same tale. They could not have 
peopled England by swimming across the Channel, 
or even by walking across solid ice, as has once been 
suggested. Nothing but a land-connection induced 
them to explore this country more closely, and 
finally to decide on settling there. 

The origin of the British fauna will be discussed 
more in detail in the third chapter. The methods 
of investigation adopted, along with a general scheme 
of this book, will be found in the next. 

The manner in which the origin of the fauna of any 
particular continental area can be traced is very 
similar to that adopted in the case of an island. 
Portions of the continent of Europe can be shown 
to have been islands in former times. Thus the 
Crimea, now a peninsula united to the mainland 
by the narrow isthmus of Perekop, must have been 
an island in comparatively recent times. The absence 
of a number of striking and familiar South Russian 
species of mammals and reptiles proves this to have 
been the case. It was probably long after the appear- 


ance of man, though before historic times, that these 
changes took place. 

We shall learn in the subsequent chapters, that 
by a careful study of the fauna and flora the 
fact can be established, not only of the former 
connection of an island with a continent, but also 
whether such union existed (geologically speaking) 
within recent or more remote times. The better 
the fauna is known, both recent and fossil, the more 
precisely can the period of connection be indicated, 
and its duration determined. 



I INTEND to give in this chapter a general outline 
of the subject which will be discussed in the 
subsequent ones. This will include a brief history 
of the great events, in recent geological times, which 
have modified the evolution of the European fauna 
by the influence which they have exerted on the 
course of the successive streams of migration. 

The composition of the European fauna is the 
first item which will have to be taken into con- 
sideration. But not only must the existing species 
of animals be dealt with : the extinct ones, too, 
at least those which have lived in Europe during 
late Tertiary times, will be useful for our inquiries. 
A knowledge of the past faunas is a most important 
factor in tracing the original home of the European 

Where a species first originated, whether this was 
in one or several places, or, in other words, where 
it first had its home, cannot be determined with 
absolute certainty in the present state of our 
knowledge, but as a rule it can be indicated 
approximately with a fair amount of precision. In 



a few instances, species may possibly have had a 
dual origin. The majority of naturalists doubt 
that there are any such, but it seems to me 
that almost the same forces may have acted in 
different localities on certain forms so as to produce, 
in very exceptional circumstances, similar species. 
The vast majority of animals, however, have no 
doubt originated in one locality; or, we might say, 
almost all species have but one home. 

We may assume that every animal gradually 
extends its range by migration, as the result of the 
natural increase of the species necessitating a search 
for fresh feeding grounds. Every species thus tends 
to slowly take possession of all the habitable parts 
of the globe to which it has access. They would 
all naturally spread from their original homes in 
every direction, unless prevented by an impassable 
barrier. We have already learned that to all land 
animals, the sea acts as such a barrier. Mountains 
and rivers act also in a similar way, but not to the 
same extent. It is not difficult to understand also 
that a forest may be a formidable barrier to a typical 
inhabitant of the open country and vice versa, whilst 
a desert is impassable to almost all terrestrial organ- 
isms. Some species are scarcely affected by climate, 
and flourish equally well in the tropics and in 
temperate or cold countries ; the majority, however, 
are greatly influenced by it. " No more striking 
illustration," remarks Merriam (p. 38), "could be 
desired of the potency of climate compared with 


the inefficiency of physical barriers, than is presented 
by the almost total dissimilarity of the North 
American Tropical and Sonoran Regions, though 
in direct contact, contrasted with the great similarity 
of the Boreal Regions of North America and Eurasia, 
now separated by broad oceans, though formerly 
united, doubtless, in the region of Behring Sea." 

To return to the composition of the European 
fauna, we now know positively that a number of the 
mammals and birds inhabiting Central and Eastern 
Europe are of Siberian origin. How they came, and 
when, will form the subject for discussion in Chapter 
V. At present it will suffice to mention that in the 
superficial deposits belonging to the Pleistocene 
series of the North European plain have been dis- 
covered the remains of many typical members of the 
Siberian Steppe-fauna. Some of these, such as the 
Saiga- Antelope (Saiga tartarica), Fig. 2, still inhabit 
portions of Eastern Europe, whilst others have re- 
treated to their native land. But it might be asked, 
how is it known that these species did not originate 
in Europe, and thence migrate to Siberia? Because 
if they had originated on our continent, they would 
have spread there. They would have invaded 
Northern and Southern Europe, and they would 
probably have left some remains in Spain, Italy, or 
Greece. They would also have left some of their 
relations in Europe ; but all their nearest allies, 
too, are Asiatic. Moreover, and this completes, I 
think, the proof of their Siberian origin, the Pleisto- 




cene remains of these animals in Europe become 
less abundant, and the number of species likewise 
decreases, as we proceed from east to west. With 
these remains of Steppe animals are generally asso- 
ciated those of others, which we must also look upon 
as Siberian emigrants, such as the Pikas or tailless 
Hares belonging to the genus Lagomys, the pouched 
Marmots (Spennophilus), and others. Some of them, 
as I have mentioned, still inhabit Central and Eastern 
Europe, whilst others have a wider distribution on our 

This migration must have been an unusually large 
one. It has been suggested that the Glacial period 
had some connection with it, and there can be little 
doubt, as we shall see later on, that a change of 
climate probably brought about this great Siberian 
invasion of Europe. But other causes might tend 
in the same direction, such as want of sufficient 
food after a few years of great increase of any parti- 
cular species. It is not known to what we owe 
the periodic visits of the Central Asiatic Sandgrouse 
(Syrrhaptes paradoxus\ Fig. 3, but certain it is that 
immense flocks of these birds invade Europe from 
time to time at the present day, just as those 
mammals may have done in past ages. 

The Siberian migrations will be spoken of in the 
subsequent pages, as the Siberian element of the 
European fauna. These migrations, however, are not 
the only ones which reached Europe from Asia. 
The sixth chapter deals with migrations which have 


influenced our fauna far more than the Siberian. The 
latter did not last long, nor did they affect the whole 
of Europe. But what I may call the Oriental migra- 
tions spread to every corner of Europe and certainly 
lasted throughout the whole of the Tertiary Era. The 
Oriental element came probably from Central and 

FIG. 3. Central Asiatic Sandgrouse (Syrrhaftes faradoxus}. 

Southern Asia, and in its march to Northern Europe 
it was joined by local European migrations. For on 
our continent, too, animals originated and spread in 
all directions from their centres of dispersal. A 
separate chapter has been given to the Alpine fauna, 
and another to that of South-western Europe, which 
will be known by the name of the Litsitanian element. 


Finally, animals have also reached us from the north, 
and in the fourth chapter the history of that remark- 
able migration will be fully discussed under the title 
of the Arctic element of the European fauna. 

It is generally believed that Africa played an im- 
portant role in the peopling of our continent, but 
this is quite a mistake. The eminent Swiss palae- 
ontologist Riitimeyer was quite right in saying (p. 
42) that it is much more probable that Morocco, 
Algeria, and Tunis were stocked with animals by way 
of Gibraltar, and perhaps also by Sicily and Malta, 
from Europe, than the South of Europe from Africa. 

I have already referred to what are known as 
"centres of dispersion" of animals, but before con- 
tinuing to explain the general outline of this book, 
it will be necessary to make a few additional remarks 
on the subject. 

Since every animal naturally tends to spread in 
every direction from its original home that is to 
say, from the place of its origin the latter should 
correspond with the centre of its range. And in 
any particular group of animals the maximum 
number of species should be formed in the area or 
zone which is the centre of its distribution. In the 
great majority of instances this is probably the case, 
in the higher animals perhaps less so than in the lower; 
still the rule must hold good that the original home 
of a species is generally indicated by the centre of 
its geographical distribution. 

Take for example our familiar Badger (Meles taxus). 


It inhabits Europe and Northern Asia. It is absent 
apparently from many parts of Central Asia, but it 
appears again farther south in Palestine, Syria, Persia, 
Turkestan, and Tibet. West Central Asia would be 
about the centre of its range. That this corresponds 
to its place of origin is indicated by the fact that the 
only three other Badgers known viz., M. anakuma, 
M. leucuruS) and M. albogularis are confined to Asia. 
If we examine the fossil history of the genus, we find 
that the two most ancient instances of the existence 
of Badgers have been discovered in Persia, where M. 
Polaki and M. maraghanus occur in miocene deposits. 
The latter had migrated as far west as Greece in mio- 
cene times ; no other trace of the Badger, however, is 
known from Europe until we come to the pleistocene 
beds. There are a good many cases known among 
mammals where the centre of dispersion would indicate 
to us a similar origin. On the other hand, there may 
be no fossil evidence of the occurrence of a species, 
or of its ancestors, in Asia, whilst such has been 
discovered in Europe. I think, however, that the 
present range of a species forms a safer criterion for 
the determination of its original home, as the Asiatic 
continent is still practically unworked from a palaeon- 
tological point of view. In a letter which I received 
from Professor Charles Deperet, he advocates the 
view that the wild Boar (Sus scrofa) is probably of 
European, and not, as I maintained (c, p. 455), of 
Asiatic origin ; because there seemed to be a direct 
descent from Hyotherium of the middle miccene of 


Europe, through the upper miocene Pig of the Mount 
Leberon (Sus major) and of Eppelsheim (Sus 
antiquus\ and the pliocene Pigs of Montpellier (Sus 
provincialis) and of the Auvergne (Sus arvernensis}. 
No doubt this appears rather a strong case in favour 
of the European origin of the wild Boar, but although 
the Tertiary strata of Asia, as I remarked, are as yet 
little known, a number of fossil pigs are known from 
India, Persia, and China, the oldest being the upper 
miocene Persian Pig (Sus maraghanus). Pigs are 
therefore as old in Asia as in Europe, and as a direct 
intercourse between the two continents probably never 
ceased since miocene times, it is not surprising that 
this genus should occur in both. Even if the genus had 
its origin in Europe, it is quite possible that in later 
Tertiary times, the active centre of origin was shifted 
to the neighbouring continent, and that henceforth 
many new species issued forth from Asia, some of 
which may subsequently have been modified on 
reaching our continent. The wild Boar (Sus scrofa\ 
however, to judge from its general range, I must look 
upon as merely an immigrant in Europe. I have no 
doubt that it originated somewhere in Asia, probably 
in the south. 

The view I take of the origin of our European 
Boar is also supported by Dr. Forsyth Major's recent 
researches. He was led to a re-investigation of the 
history of the Pig while examining a la'-ge number 
of fossil skulls in the Museum at Florence, and came 
to the conclusion that only three or four species of 


recent wild pigs can be clearly distinguished (b, p. 298). 
One of these, viz., Sus vittatus^ he thinks, is trace- 
able in slight modifications frcm Sardinia to New 
Guinea and from Japan to South Africa. The centre 
of distribution of this species lies in Southern Asia. 
Of the three remaining species, two, viz., Sus ver- 
rucosus and *S. barbarus, are entirely confined to the 
great islands which form part of the Malay Archi- 
pelago. Finally, Sus scrofa, our Central European 
wild Boar, is so closely related to 5. vittatus that the 
Sardinian Boar might be looked upon as a variety of 
either the one or the other. At any rate, Dr. Major 
recognises clearly in Sus vittatiis the representative 
of the ancestral stock of which Sus scrofa is a some- 
what modified offshoot. 

The fauna of Europe consists, as I have mentioned, 
to a large extent of immigrants from the neighbouring 
continents. This is especially noticeable among the 
higher animals. When we come to the lower, such as 
the amphibia, we find a larger percentage, and among 
the land mollusca the great majority, to be of Euro- 
pean origin. The foreigners are, as we learned, called 
Orientals, Siberians, and Arctics. For the sake of 
convenience, only two of the great European centres 
of origin have a chapter devoted to themselves, 
namely, the Alpine and the Lusitanian centres. 
There is another, - however, of almost equal im- 
portance which lies in the east. 

In the British Islands there is only an exceedingly 
small and insignificant group of species which are 


peculiar, and which we may consider to have had 
their origin there. Almost the whole of the British 
fauna is composed of streams of migrants which came 
from the north, south, and east, though many of 
these immigrant species have since their arrival been 
more or less distinctly modified into varieties or local 

The eminent French conchologist Bourguignat (a, p. 
352) was of opinion that, as far as terrestrial mollusca 
were concerned, there are in Europe three principal 
centres of creation or dispersion all situated in 
mountainous countries and not in the plains. He 
distinguished the Spanish, Alpine, and Tauric centres, 
and believed that almost all species known from 
Europe had originated in one of these three, and that 
each of them possessed quite a distinct type of its 
own. This theory seems to agree very well with the 
facts of distribution. Let us take, for instance, the 
genus Clausilid) a pretty turret-shaped snail, which 
abounds on old ruined walls. Only two species, viz., 
CL laminata and CL bidentata, are met with in Ireland. 
In England we find the same species with the addition 
of two others, CL biplicata and Cl. Rolphii. Crossing 
over the Channel to Belgium, these four species occur 
again, and also several others not known in England. 
In Germany the list of Clausilice mounts up to twenty- 
five species, including all those found in the British 
Islands. As we proceed eastward the number of 
species of this genus increases steadily, and when we 
reach the Caucasus or the Balkan Peninsula the con- 


chologist is able to make a collection of several 
hundred different kinds, whilst farther east again they 
diminish. This clearly indicates there is in South- 
Eastern Europe a powerful centre of creation of 
Clausilice, from which the species have spread all 
over Europe. But it is by no means certain that this 
centre was always in our continent, for in South- 
Eastern Asia and the Malay Archipelago Clausilice 
increase once more. It is interesting to note, however, 
that almost all these eastern forms belong to the sub- 
genus Phcedusa (vide Boettger), which had only been 
known as a fossil genus from a few species in the 
Eocene and Oligocene of Southern Europe. The 
first centre of origin, therefore, may possibly have 
been in Southern Asia, and in these early Tertiary 
limes a second centre may have become established 
in Southern Europe from which the sub-genus Gar- 
nieria went eastward, Macroptychia southward, and 
Nenia westward across the Atlantis to South America. 
Only a few remnants of these primitive Clausilicz 
are now left in Europe, such as the interesting Cl. 
(Laminifera] Fault. 

As an example of a genus which has its centre of 
distribution in South-Western Europe we might take 
that to which our common brown garden slug belongs, 
viz., Arion. Dr. Simroth, who was the first to point 
out that the species of Arion had spread over our con- 
tinent from South-Western Europe (p. 5), is inclined 
to the belief that the Arionidce had originated on the 
old land-bridge between Europe and North America. 


which is generally known by the name of " Atlantis." 
From this a branch went westward to the New 
World and another eastward as far as Southern Asia, 
but Arion and a number of other genera are more or 
less confined to South-Western Europe. Only a few 
species of Anon have a wide range in Europe, one of 
them, A. subfuscus, crossing the borders of our con- 
tinent into Siberia. In the British Islands and in 
Western Germany, which are about equi-distant from 
the supposed creative centre of the genus, there are 
found five species. In France six or seven species 
are met with, and in Spain and Portugal about ten. 
Towards the east, Arions diminish in number. This 
genus, therefore, forms part of a migration which I 
have designated as " Lusitanian " from Lusitania, 
the name applied by the Romans to what we 
now call Portugal. Another genus of slugs, Geo- 
malacuS) is interesting from the fact that one species 
occurs in the British Islands, being otherwise con- 
fined to the Lusitanian province. Parmacella, a slug- 
like animal bearing a tiny shell at the extremity of its 
tail, has probably likewise had its origin in this part of 
Europe. All this, however, will be more fully referred 
to in the seventh chapter, which deals with the 
Lusitanian fauna. 

As regards the Alpine centre of origin, Dr. Kobelt 
considers three groups of mollusca as especially 
characteristic of the Alps, viz., the sub -genus Carnpylaea 
of the great and widely-spread genus Helix, and the 
genera Pomatias and Zonites. The latter, which is not 



to be confounded with our British Hyalinia (formerly 
united with Zonites\ does not extend very far south 
or north of the Alps. There may be others too, which 
owe their origin to these mountains, but most of 
the terrestrial mollusca are exceedingly ancient, and 
many genera have existed long before the Alps had 
made their appearance above the surface of the early 
Tertiary seas. It should be remembered that Hya- 
linia and Pupa, both British genera, are known from 
carboniferous deposits in forms which closely approach 
those living at the present day, and in these and a 
great number of other instances, it is quite impossible 
to determine the original home of the genus. 

This little digression on centres of dispersion will 
help us to understand in what manner the indigenous 
element of the European fauna joined in with the 
alien members as they arrived in our continent. The 
species confined to South-Eastern England need not 
necessarily have come to us from Eastern Europe or 
Siberia. Alpine species spread northward probably 
at the same time as the Siberian animals went west- 
ward. An Alpine form may therefore have joined a 
batch of the latter and entered Eng-land with them. 
Even a Lusitanian animal may have mingled with 
these migrants, so that all three elements may occur 
together in one locality. 

But these are exceptions. The migrations have, as 
a rule, not joined to any great extent; indeed, all 
those naturalists who have carefully examined the 
problem of the origin of the European fauna, have felt 


that it was composed of elements which arrived at 
different times. 

The great Russian naturalist, the late Professor 
Brandt, distinguished five phases in the history of the 
Eurasian mammalian fauna (pp. 249-254). During the 
first phase an uncertain period of long duration the 
mammals held intact their position in the northern 
half of Asia. The Mammoth, the Hairy Rhinoceros, 
Bison, Musk Ox, Wild Sheep, Reindeer, and perhaps 
Tigers, Hyaenas, etc., lived then, with numerous 
peculiar Rodents, under such climatic conditions, 
according to Brandt, that they were able to extend 
their range along with tree vegetation to the extreme 
north of the Asiatic continent This, he thinks, seems 
to have been the case especially with the Reindeer, 
Mammoth, Rhinoceros, and Musk Ox. The second 
phase was characterised by the dispersion of the 
Northern Asiatic mammalian fauna towards Central, 
Southern, and Western Europe, and this period lasted 
until the complete extermination of the Mammoth. 
The third phase dates from the time when the Mam- 
moth and the Hairy Rhinoceros had become extinct, 
whilst the fourth commenced with the disappearance 
of the Reindeer in Europe, and terminated when the 
Wild Ox in the feral state had become unknown. 
Finally, the last phase constitutes the present time. 
Lartet held similar views, and also believed that 
Europe was peopled by successive migrations from 

Botanists have worked at the problem of the 


European flora much more systematically, and our 
knowledge of the origin of that flora has been greatly 
increased within the last twenty years, chiefly by 
the researches of Professor Engler. More recently, 
detailed studies have been made in Scandinavia by 
Professor Blytt, in the Alps by Dr. Christ and Mr. 
Ball, in Germany by Professor Drude, Dr. Schulz, 
and many others. Dr. Schulz (p. i) is of opinion 
that the great majority of the European plants have 
either migrated to or have originated in our conti- 
nent since the beginning of the Pliocene epoch, and 
that the original home of the immigrants must be 
looked for in Asia and in Arctic America. From the 
latter an almost uninterrupted migration must have 
taken place during the greater part of Tertiary times 
up to the commencement of the Pliocene epoch, partly 
over a direct land-connection with Europe by way of 
Greenland, Iceland, and the Faroes, and also vid 
Spitsbergen, Franz Josef Land and Novaya Zemlya, 
and partly by an indirect one across the Behring 
Straits between Alaska and Kamtchatka. 

A good deal of work still requires to be done before 
zoologists have acquired the same intimacy with the 
European fauna as botanists have with the flora. 
However, the view that our animals all come from 
Asia, as was long ago believed, has been abandoned 
for some time. The first to bring under the notice 
of naturalists the hypothesis, that there must have 
been two distinct migrations of northern animals to 
Central Europe one from the north, and another 


from the cast was the late Mr. Bogdanov. The 
Arctic species, of which remains have been discovered 
in the Pyrenees namely, the Reindeer, Arctic Hare, 
Willow Grouse, etc., he thought had nothing to do 
with those which invaded Europe from Siberia during 
the Glacial period. He maintained that the former 
had quite a distinct origin, and came from Scandi- 
navia (p. 26). 

As I shall deal with this problem more fully in a 
subsequent chapter, I need only mention that I fully 
agree with the view expressed by Mr. Bogdanov that 
two distinct migrations of northern species to Central 
Europe can be traced. 

No one, I think, has done more in fostering a care- 
ful study of the migrations of animals than our 
distinguished geologist Professor Boyd Dawkins. He 
did not follow Bogdanov in distinguishing two Arctic 
migrations; however, he did more in constructing a 
very ingenious chart (a, p. ill) representing the geo- 
graphy of Europe during the last and most recent 
geological epoch the Pleistocene and indicating on 
it the probable extent, during that time, of an eastern 
and a southern migration of mammals. The map is 
very instructive, and is the first ever published giving 
a clear idea of a southern and an eastern migration 
to Europe. He believed that the migration of the 
southern mammals northward, took place conjunctly 
with the westward movement of the eastern species. 
Having once reached Europe, the southern species 
are supposed to have passed northward in summer 


time, whilst the eastern forms (he calls them northern) 
would swing southwards. The two migrations would 
thus occupy, at different times of the year, the 
same tract of ground (a, p. 1 13). From the mingling 
of the remains of the Hyaena with those of the 
Reindeer and Hippopotamus in the Kirkdale Cavern, 
he infers that the former preyed upon the Reindecr 
at one time of the year, and on the Hippopotamus at 
another. He argues that in such a manner might 
be explained the curious mixture of northern and 
southern types which we find in the British pleistocene 
and in cave deposits. 

Besides mammals, the only European animals 
which have received some attention with a view to a 
study of their origin, are the Butterflies and the Land- 
Snails. The entomologists who have taken up the 
problem have in so far scarcely produced satisfactory 
results, as they all seemed to be bound down to the 
hypothesis that practically all the butterflies had 
been destroyed in Europe during the Glacial period. 
Hofman, in his interesting little work, comes to the 
conclusion (p. 50), that only in Greece and Spain 
could a small remnant of the butterflies have survived 
the extreme rigours of climate. Greece was at that 
time connected with Asia Minor, and Spain with 
North Africa ; and the author supposes that the semi- 
alien fauna inhabiting these tracts was mainly re- 
sponsible for the re-stocking of Southern Europe, but 
that the main mass of our butterflies are post-glacial 
Siberian immigrants. 


The work published by Messrs. Speycr deals only 
with the origin of the Central European Butterflies. 
The period during which our European species 
originated is not specified, but the authors believe 
that they had their home either in Southern Russia 
or Central Asia. The fact that the number of 
butterflies decreases very considerably as we pro- 
ceed north-westward in Europe appears to them 
to substantiate these views. The apparent dislike 
evinced by butterflies to the damp Atlantic Coast 
climate, they think, clearly indicates that they had 
originated in a dry and more continental climate. 
The history of the North European Butterflies 
and Moths has been carefully described by Mr. 
Petersen. He adopts Hofman's theory as to the 
almost total extinction of the Lepidoptera in 
Europe during the Glacial period. The chief immi- 
gration to Europe after that period is, he thinks, 

At first there appeared species which belonged 
to a cold climate, and which now live in ele- 
vated regions ; then came forms suited to a milder 
climate, which established themselves on the north- 
easterly slopes of the Alps. The most recent addition 
which our continent has received from Siberia is, 
according to Mr. Petersen, the present Scandinavian 
fauna. Scandinavia has obtained a larger number of 
species than the European plain, because to this last 
migration were added such as prefer a northern or 
Alpine climate. 


As a contribution to the history and composition of 
the European fauna, by far the most important work 
ever published is that of Dr. Kobelt, the eminent 
German conchologist. Whilst the researches into the 
origin of the Lepidoptera, above described, have been 
marred by the prevalent prejudice as to the dele- 
terious effects of a glacial climate on the butterflies, 
the present author boldly works out the problem on 
independent lines. He shuns theories and specula- 
tions almost altogether. His great work, as yet 
practically unknown, the result of a lifetime of the 
most painstaking labour, ranks among the most im- 
portant contributions to zoogeography. I shall have 
frequent occasion to refer to it throughout these pages. 
Meanwhile some of his more remarkable conclusions 
may be mentioned. " Comparing all classes of 
animals as to their zoogeographical importance, the 
highest rank must undoubtedly be accorded to the 
land-snails " (i., p. 7). " The Pleistocene, and with it 
the land and fresh-water molluscan fauna of the 
present day has been gradually evolved from the 
Tertiary one, and its roots can be traced through the 
Cretaceous to the Jurassic epoch. During the whole 
of that time no sudden appearance of a new fauna 
can be demonstrated. Quite slowly, step by step, the 
Cretaceous is succeeded by the Tertiary fauna, and 
one after the other of the characteristic palaearctic 
genera appear first the fresh-water, then the land 
forms" (p. 141). "The division of the North Alpine 
from the South Alpine fauna must be older than the 


Glacial period ; and the present Central European 
fauna had already become developed from the Plio- 
cene in all its details of form and distribution before 
the commencement of the Ice Age" (p. 162). "We 
must draw the conclusion from the preceding remarks, 
that the present (palaearctic) molluscan fauna in its 
distribution is older than the Glacial period, and that 
the latter produced merely a retreat of the fauna 
from the most inhospitable regions of Europe with 
a subsequent re-immigration, but did not cause its 
destruction" (i., p. 169). 

A few attempts have also been made by naturalists 
to trace the origin of the fauna of some smaller 
European areas. Thus Riitimeyer, in dealing with 
the mammalian fauna of Switzerland, remarks (p. 31) 
"that it seems certain that, in spite of many local 
disturbances, the continuity of generations was never 
interrupted throughout the whole of the Tertiary 
period until the present day." 

An even more interesting memoir is that of Mr. 
Koppen on the origin of the Crimean fauna. It is 
only recently, according to this author, that this pen- 
insula has become connected with Southern Russia. 
And it is for this reason that the Squirrel and a number 
of other animals, and also plants, present in Russia, 
are absent from the Crimea. Originally the latter prob- 
ably formed a westward continuation of the Caucasus, 
and at that time it was surrounded by the sea on 
all other sides. " Much later," he continues, " after 
and in consequence of a local subsidence, the country 


between the Caucasus and the Crimea became 
interrupted. The latter existed for a long time as 
an island, and only much later, in recent geological 
times, did it become united with Southern Russia' 
by means of the isthmus of Perekop." 

There is, on the whole, a great diversity of opinion 
as to how the European fauna has originated ; how- 
ever, except in Dr. Kobelt's work, no attempt has 
hitherto been made to collect together all the available 
information, and to include in the inquiry more than 
one class of animals. The little work which I venture 
to bring before the public will not by any means ex- 
haust the subject, nor is our knowledge of the Euro- 
pean fauna sufficient to give more than a mere sketch 
of many of the animal groups mentioned. As we 
have learned in the introduction, different classes of 
animals are not all of equal importance in indicating 
the changes which have taken place in the distribution 
of land and water. While Dr. Kobelt is of opinion 
that the land-snails are by far the most important 
in such an inquiry, Mr. Lydekker believes that 
mammals afford the safest and truest indications 
of such changes. Mr. Beddard puts in a claim for 
earthworms, as even a narrow strait of sea-water 
forms an insuperable barrier to their dispersion. 
Dr. Wallace agrees with Mr. Lydekker, and goes 
so far as to say (p. 74) that " whenever we find 
that a considerable number of the mammals of two 
countries exhibit distinct marks of relationship, we 
may be sure that an actual land-connection, or at 


all events an approach to within a very few miles 
of each other, has at one time existed." Besides 
the groups referred to, I claim that particular 
attention should be devoted to Amphibia, which, 
contrary to Wallace, I hold do not possess special 
facilities for dispersal ; and also to spiders and to 
all wingless animals leading a subterranean life, such 
as some of the wood-lice, planarian worms and 
apterous beetles. 

A thorough knowledge of the changes in the dis- 
tribution of land and water is desirable in order to 
appreciate the extent and variations of former mi- 
grations. A study of the British fauna, for example, 
teaches us that the British Islands were once con- 
nected with one another and with the continent of 
Europe between England and France. It was 
Professor James Geikie, I believe, who first pointed 
out, many years ago, that the area now covered by 
the Irish Sea was formerly in all probability a fresh- 
water lake. This had its outlet at the southern ex- 
tremity in the form of a stream into which most likely 
flowed the smaller rivers from the south-east of 
Ireland, and which was joined from the east by the 
Severn, and finally debouched into the Atlantic 
(Fig. 4). The range in the British Islands of those 
species which have migrated to them from the south, 
indicates that whilst the Atlantic Ocean had gradually 
crept up and flooded the area between Ireland and 
Wales, and had turned the fresh-water lake into a bay, 
communication between Scotland and Ireland was 



still possible. The occurrence of many Scandinavian 
species in Scotland which are absent on the continent 


FIG. 4. Map of the British Islands and surrounding area at a time 
when the earlier members of the southern migration reached 
England. (Only some of the rivers have been indicated. The 
shaded parts represent water, the light land.) 

of Europe, indicates that these two countries also 
were united formerly. Most geologists hold that such 


a connection, if it existed, must have broken down 
in Pliocene times. Professor Judd, however, has ex- 
pressed his belief (p. 1008) that it still existed until 
after the appearance of man in Northern Europe, and 
that our forefathers might have been able to walk 
dry foot from Scotland to Norway. 

I shall also show on distributional evidence, in the 
fourth chapter, that until recent geological times 
Scandinavia was continued northward, by way of 
Bear Island, with Spitsbergen and probably Franz 
Josef Land, which islands again were joined with 
North Greenland and Arctic North America, and 
that the polar fauna and flora were able to spread on 
this land-connection to both America and Europe. 

That Gibraltar was connected with Morocco, and 
Sicily with Southern Italy and Greece on the one 
hand, and with Tunis on the other, is more generally 
recognised; whilst Professor Suess has shown (vol. i., 
p. 442), on purely geological grounds, that the 
Egean Sea was dry land up till quite recently 
certainly, he thinks, till after the appearance of man. 
This supposition enables us to understand, as will be 
more fully discussed in the sixth chapter, how the 
Oriental fauna entered Europe. Such minor zoo- 
geographical problems as the occurrence of the Wild 
Goat of Asia Minor {Capra cegagtus) on the islands of 
Crete and on some of the Cyclades now almost 
explain themselves. The Sea of Marmora is prob- 
ably a modern formation, so that Asia Minor ex- 
tended not long ago beyond the Turkish capital, but 


Dr. Kobelt believes that an arm of the Black Sea 
communicated up till recent times along the lower 
course of the Maritza with the Gulf of Saros. 
It can be shown also that Sardinia and Corsica 
formed part of the continent of Europe, and that 
their present fauna and flora reached them by migra- 
tion on land. 

The Russian naturalists, Brandt and Koppen, be- 
lieved that at no very distant date a sea extended right 
across Eastern Russia from the Caspian to the Arctic 
Ocean, whilst Professor Boyd Dawkins expressed 
himself in very similar language as follows (c, p. 35) : 
" Before the lowering of the temperature in Central 
Europe the sea had already rolled through the low 
country of Russia, from the Caspian to the White 
Sea and the Baltic, and formed a barrier to western 
migration to the Arctic mammals of Asia." These 
naturalists based their opinions on distributional 
evidence, but additional facts will be brought 
forward in the fifth chapter to substantiate these 

These are some of the more important geographical 
events which will be dealt with in detail in the subse- 
quent chapters in connection with the history of the 
migrations of the European fauna. 

A separate chapter has been devoted to the British 
fauna and its origin, since it plays a very important 
part in the evolution of that of our continent. So 
essential is a thorough knowledge of this fauna, that 
I think it would be difficult to understand, without 


it, the main features of the great migrations ; and I 
have before now expressed the opinion that the 
British fauna forms the key to the solution of the 
problem of the origin of European animals. W T e 
know that our British species came to us by land 
at least the bulk of them. But we want to 
know what direction they came from, and at what 
time they arrived. When Ireland became discon- 
nected from Great Britain, and the latter from 
Scandinavia and France, is another interesting 
problem. Professor Boyd Dawkins has indicated 
to us a method of the special line of research to 
meet such inquiries. " The absence," he says (b y 
p. xxix), " of the beaver and the dormouse from Ire- 
land must be due to the existence of some barrier 
to their westward migration from the adjacent main- 
land, and the fact that the Alpine hare is indi- 
genous, while the common hare is absent, implies 
that, so far as relates to the former animal, the barrier 
did not exist." 

Many members of the great Siberian invasion 
reached England, but Ireland remained entirely 
free from these migrants. The assumption there- 
fore seems not unreasonable, that the latter country 
at the time of their arrival was no longer joined 
to England. The great bulk of the Irish fauna 
is composed of Lusitanian, Alpine, and Oriental 
immigrants, and there is besides a distinctly Arctic or 
North American element. All these, of course, must 
have established themselves in Ireland before the 

^^SJ B *AJT^. 


Siberian fauna set foot in England, since it has been 
shown that a continuous land-surface was necessary 
for their migration. Owing to the perfect preserva- 
tion of the remains of the Siberian migrants in recent 
continental deposits, the history of that migration 
can be clearly followed, and it is possible even to 
determine the date of its arrival in England in 
geological language at any rate. The time of the 
colonisation of Ireland can be thus approximately 
fixed as having taken place at a period prior to the 
arrival of the Siberian migrants in England. 

All those who have seriously studied the problems 
presented by our British fauna notably the late 
Professor Forbes, and more recently Mr. Carpenter 
and myself are agreed that the Lusitanian element 
is the oldest, and that the newest is that which has 
come to us from the east. 

The sequence of events in the British Islands was 
probably as follows : The first comers were the 
members of that fauna which issued from South- 
western Europe; then came the Alpine, and at the 
same time probably the Arctic and the Oriental; 
and finally the Eastern or Siberian. The migrations 
of all but the last continued, uninterruptedly, for very 
long periods. 

The study of these migrations has convinced me 
that, though climate was a powerful factor in the 
evolution or history of the European fauna, the 
geographical changes which took place on our con- 
tinent in later Tertiary times exerted a yet stronger 


influence. The principal climatic disturbance is 
generally supposed to have been the so-called " Ice 
Age." So firmly rooted is the conviction, among 
naturalists of the present day, of the enormous 
destruction which this period produced on our 
European fauna, so that all animal life practically 
disappeared from large areas of our continent, that 
it is desirable that we should now shortly review 
the history of that remarkable period in order to 
ascertain in how far these views are corroborated 
by facts. Frequent reference, moreover, will be 
made throughout this work to the theories con- 
nected with the Glacial period. 

It has been stated by an eminent geologist that 
during part of the Glacial period the climate was such 
that neither plants nor animals could have existed in 
the British Islands. If that had been so, it is evident 
that very few organisms could have even survived in 
France, though a number of Arctic species might 
have dragged on an existence in Southern Europe. 
At any rate, on the return of more genial conditions, 
the Arctic species would undoubtedly have been the 
first to gain admission to the British Islands, to 
re-people the arid wastes. Our supposition that the 
Lusitanian element in the British fauna is the oldest 
would therefore be wrong. From early Tertiary 
times onward, the climate of Europe, which was 
then semi-tropical, gradually became more and more 
temperate; until finally the Ice Age or Glacial 
period arrived, during which, according to Professor 



J. Geikie one of our highest authorities on this 
subject a great part of Northern Europe became 
practically uninhabitable owing to the severity of 
the climate. 

To enable us to judge better of the true value of 
the many hypotheses which have been advanced 
to account for this supposed extraordinary fall of 
temperature during the "Ice Age," we must compare 
the views of other authorities with the one just 
quoted. I do not propose to discuss the causes which 
have led to the production of the Glacial period 
those interested in these questions should consult 
the writings of Dr. Croll, Professor J. Geikie, Pro- 
fessor Bonney, Mr. Falsan, and others but merely to 
give the climatic aspects from a physical, zoological, 
and botanical point of view. 

According to Professor Penck (a, p. 12), the nature 
of the glacial climate can be determined by comparing 
the snow-line of the Glacial period with that of the 
present day. The position of the snow-line is de- 
pendent on two climatic factors viz., precipitation 
and temperature. We know the height at which 
snow must have lain permanently during the Glacial 
period, or during the maximum phase of glaciation. 
If the Ice Age had been produced solely by an 
increase of snowfall, as has been suggested, Professor 
Penck tells us that then it must have snowed three or 
four times as much as it does now. But he does not 
adopt the view that the Ice Age is due to an increase 
of snowfall alone. His calculations, based upon the 


height of the snow-line, tend to show that a general 
decrease of temperature to the extent of from 4-5 
degrees Centigrade (all other atmospheric conditions 
remaining the same as now) would be sufficient to 
give us back the Glacial period. 

Professor Neumayr (p. 619) adopted a similar prin- 
ciple in determining the temperature which prevailed 
in Europe during the Glacial period. Snow now lies 
in the Pyrenees 1000 metres higher 'than it did then, 
1,200 metres higher in the Alps, and 800 metres 
higher in the Tatra mountains. Since the tempera- 
ture in Central Europe decreases by half a degree 
Centigrade for every 100 metres of elevation, it follows 
that if the glacial phenomena had only been brought 
about by a decrease of temperature without an in- 
crease of moisture, we should have had a reduction 
of temperature during the Glacial period of six 
degrees Centigrade in the Pyrenees, of seven degrees 
in the Alps, and of four in the Tatra mountains. 
The general lowering of the temperature of Europe, 
says Professor Neumayr, could not have amounted to 
more than six degrees Centigrade. Moreover, he is 
of opinion that the very low snow-line in the British 
Islands proves that even during the Ice Age a com- 
paratively mild climate prevailed there, and that 
the climatic conditions generally, in the different 
parts of Europe, were relatively about the same as 
they are now. 

Professor J. Geikie does not give us his views as to 
the temperature of the Glacial period, but he main- 


tains that a lowering of the temperature is evinced 
not only by the widespread phenomena of glaciation, 
but by the former presence in our temperate latitudes 
of a northern fauna and flora. 

Mr. Charles Martins, who based his calculations 
on the temperature during the Glacial period on 
the glaciers of Chamounix, concluded that it only 
needed a lowering of the temperature to the extent 
of four degrees Centigrade to bring the glaciers down 
to the plain of Geneva, and in fact give us back the 
Glacial period. It need not surprise us, therefore, 
that the French geologist, Mr. Falsan, the author 
of La periode glaciere, is of opinion (p. 230) that the 
mean annual temperature of France during the 
Glacial period was approximately from 6-9 degrees 
Centigrade, perhaps more. Close to the immense 
glaciers of the Rhone, it might have been about 
six degrees. This is the actual mean annual tempera- 
ture of the South-west of Sweden and Norway, or 
the North of Scotland. 

Although all these investigations tend to show 
that the climate of Europe during the Glacial period 
was by no means so severe as we are often led to 
believe, yet there exists also a school of geologists 
who maintain there was actually a higher temperature 
than at present. The inconsistency of mentioning 
heat in connection with ice and snow is more 
apparent, however, than real, for we must remember 
Tyndall's original remark on this subject. It is the 
snow, he says, which feeds the glaciers. But the 


snow comes from the clouds, and these again 
originate from the vapours which the sun causes 
to be absorbed from the ocean. Without the sun's 
heat, we should have no water vapour in the 
atmosphere ; without vapour, no clouds ; without 
clouds, no snow ; without snow, no glaciers. The 
ice of glaciers, therefore, owes its origin indirectly 
to the sun's heat. It has been supposed that if the 
sun's heat diminished, larger glaciers would form 
than those existing to-day, but the diminution of the 
solar heat would infallibly reduce the amount of 
water vapour in the air, and would thus stop the 
very source of glaciers. 

Mr. Falsan even admits that without a change of 
the mean annual temperature (p. 201) of Europe, 
the central portions of our continent might at this 
period have enjoyed an insular climate. This more 
equable and humid climate could, within certain 
limits, favour the development of the ancient glaciers 
by increasing the snowfall and slackening the summer 
rate of melting. 

It seems evident then, according to these views, 
that with a comparatively slight change of the 
atmospheric conditions in the British Islands, we 
might have glaciers back again on all our highest 
mountain ranges in England, Scotland, and Ireland. 
But a widespread belief seems to prevail that the 
presence of glaciers implies a very low temperature. 
Snow and ice, however, are formed as soon as the 
temperature falls below freezing point; it does not 


matter whether there be I or 20 degrees of cold. 
Winters with a few degrees of frost will be just as 
favourable for the growth of glaciers as winters with 
the most severe cold. 

Let us now see what the fauna and flora, as far 
as we know it, tell us of the climate of the Glacial 
period. At the very outset of our inquiry we are 
confronted with one very serious difficulty in the 
problem, and that is the supposed occurrence of inter- 
glacial mild phases alternating with colder ones during 
the Ice Age. At first, when traces of a temperate flora 
and fauna were discovered intercalated between two 
layers of boulder clay, their presence was explained 
by the supposition of a mild inter-glacial period. 
The famous Forest-bed on the east coast of England 
was also pronounced to be an inter-glacial deposit, 
though not coming precisely under this definition. 
In a few places one such bed was found, in some two 
or more, and in others none at all. Professor James 
Geikie discovered the evidences of no less than five 
of such inter-glacial epochs (p. 612) in Europe. 
Lest a reader of that author's remarkable work on 
the Ice Age might carry with him the idea that his 
hypotheses had met with general acceptance, a few 
quotations from almost equally high authorities on 
glacial matters will be useful. "That the glaciers," 
remarks Professor Bonney (p. 245), " were liable to 
important oscillations seems to be proved, but whether 
the evidence suffices to establish inter-glacial epochs, 
in the usual sense of the words, is more doubtful. 


When the snow-fields, as in the Alps, were much 
more extensive than they are at present, the glaciers 
which radiated from them would be more sensitive 
to minor climatal change. Even now they oscillate 
considerably. But during a Glacial epoch, an inch, 
either more or less, of precipitation might mean a 
considerable advance or retreat of the ice in the low- 
lands." French geologists look with even less favour 
on Professor Geikie's theories. Mr. Falsan (p. 212) 
says that he agrees with Messrs. Favre, de Saporta, 
Lory, de Mortillet, Desor, de Lapparent, Lortet, 
Chantre, Benoit, Fontannes, Deperet, and many 
other geologists, that there was only a single Glacial 
period, which, according to each particular region, 
might be divided into several phases, or into their 
equivalents viz., one or more extensions of the 
ancient glaciers. But, on the whole, the view that 
there was at least one inter-glacial phase in the 
Glacial period meets with more general acceptance 
among geologists, I think, though the other opinion 
agrees much better with the nature of the fauna 
and flora as it has been revealed to us from the 
pleistocene deposits. 

The occurrence of the remains of such arctic species 
of mammals as the Musk-Ox, Arctic Fox, Glutton, 
Lemming, and many others in these deposits, is 
frequently held up to us by geologists as a proof 
of the prevalence of an arctic climate while these 
beds were laid down. And indeed this appears 
at first a most satisfactory explanation of the 


phenomenon. But we must not judge the climate 
of Europe by their presence alone. As I shall 
explain more fully in Chapter V., these species 
invaded Europe owing to two circumstances. Firstly, 
because the climate of Siberia was becoming colder, 
necessitating a southward movement, with a con- 
sequent over-population in a reduced area; secondly, 
because a new short route to Europe had been 
opened up for them about the same time (see 
p. 221). An invasion of Europe therefore took place 
from east to west. Similar invasions occur even at 
the present day, though not caused by a change in 
our climate, for every now and then immense flocks 
of the Siberian Sandgrouse emigrate to our continent. 
The mammalian migrants referred to are not to be 
looked upon as constituting the whole of our fauna 
at that time. Europe had a fauna of its own, and 
these invaders merely mingled with our animals. 
There was, no doubt, a keen struggle for existence, 
as the result of which the weaker in many cases 
succumbed. The hypothesis, however, that these 
Siberian migrants occupied an empty continent, 
forsaken by its pre-glacial inhabitants, is not sup- 
ported by any facts. 

All those who have investigated the pleistocene 
fauna have been struck by the extraordinary mixture 
of northern and southern types of animals. Professor 
Dawkins attempted to explain these facts by the sup- 
position (p. 113) that "in the summer time the southern 
species would pass northwards, and in the winter 


time the northern would sway southwards, and thus 
occupy at different times of the year the same 
tract of ground, as is now the case with the elks 
and reindeer." " In some of the caverns," he 
continues (p. 114), "such as that of Kirkdale, the 
hyaena preyed upon the reindeer at one time of the 
year, and the hippopotamus at another." 

A similar mingling of northern and southern 
faunas has also been observed in France. Mr. 
Falsan tells us (p. 236), that the remains of the 
mammals gathered and determined by Lartet and 
Gaudry belong partly to species which have been 
wrongly regarded as indications of a severe climate, 
and partly to such as are accustomed to a relatively 
mild temperature. In several localities in France, 
viz., at Levallois, St. Acheul, and Arcy, the remains of 
the Hippopotamus have occurred together with those 
of the Reindeer; whilst, according to Sir H. Howorth, 
the Lion has been found together with northern Voles 
at Bicetre, near Paris. It is stated by the same 
authority (p. 115) that much the same conditions 
exist in Germany. "The lion and the spotted hyaena, 
the mammoth and rhinoceros, were found with the 
marmot, the suslik, the lemming, the pica, and the 
reindeer." At another locality near Thiede, remains 
of the Mammoth, woolly Rhinoceros, Horse, Ox, 
Reindeer, Arctic Fox, Lemming, and Pica are met 
with in the same deposit. In quoting the presence 
of these northern animals in Europe as evidence of an 
arctic climate, we commit a fatal mistake. Indeed, 


breeders of animals and those acquainted with zoo- 
logical gardens know perfectly well that it is much 
easier to keep a northern species in a southern 
climate, than a southern species in a northern one. 
If in a Central European deposit occur a mixture of 
northern and southern forms of animals, the presence 
of the latter is more remarkable than that of the 
former. Logically, we should look upon the occurrence 
of southern species in the north, therefore, as support- 
ing the view that a mild climate had induced them 
to travel northward. The only indication, indeed, 
of the presence of a Monkey in the British Isles 
in former times comes to us from the very same 
strata which have also yielded the remains of the 
Siberian mammals. 

Before I conclude the consideration of the pleisto- 
cene fauna, it may be of interest to hear what 
Mr. Lydekker, one of our highest authorities 
on fossil mammals, has to say on this subject. 
" The most remarkable feature connected with 
this fauna is the apparently contradictory evidence 
which it affords as to the nature of the climate 
then prevalent. The Glutton, Reindeer, Arctic 
Fox, and Musk-Ox are strongly indicative of 
a more or less arctic climate; many of the Voles 
{Micr<otus\ Picas (Lagomys), and Susliks {Sper- 
mophilus\ together with the Saiga Antelope, 
appear to point equally strongly to the prevalence 
of a Steppe-like condition ; while the Hippopotamus 
and Spotted Hyaena seem as much in favour of a 


sub-tropical state of things. Many attempts have 
been made to reconcile these apparently contra- 
dictory circumstances ; one of the older views being 
that while the tropical types of animals lived during 
a warm interlude, they migrated southwards with 
the incoming of colder conditions to the arctic type 
of fauna. Since, however, it has now been ascertained 
that the remains of both tropical and arctic forms 
have been found lying side by side in the same bed, 
it is perfectly certain that such an explanation will 
not meet the exigencies of the case " (p. 300). 

In Germany the remains of the Siberian mammals 
occur to a large extent in a pleistocene deposit known 
as " loess," and the theory has of late years gained 
ground that the latter is the fine dust-like sand 
accumulated during an intensely arctic dry climate. 
That many of the mammals discovered in the " loess" 
now inhabit the dry steppes of Eastern Europe and 
North-Western Asia seems to lend support to this 
supposition; but besides the mammals there are also 
land and freshwater shells in this deposit. The mol- 
luscan fauna certainly indicates no steppe-character, 
according to Dr. Kobelt (b, i. p. 166). 

The attempt to utilise the Siberian migrants to 
Europe as indicators of a severe climate there, cer- 
tainly fails to establish conviction. But it may be 
asked, surely the remains of the Alpine and Arctic 
plants which have been found in pleistocene deposits 
must decide this question in favour of one or the 
other hypothesis? Let us test it. 


Plants being more directly affected than animals 
by changes of temperature and rainfall, remarks 
Mr. Clement Reid (p. 185), give evidence of the 
highest value when we inquire into former climatic 
conditions. The severity of the climate during the 
Glacial period is often assumed from the occurrence 
in pleistocene strata of such plants as Dryas octopetala^ 
some species of willow, the dwarf birch, and others, 
which are now found in high latitudes and in the Alps, 
but are, as a rule, absent from the plain of Northern 
Europe. Professor J. Geikie goes so far as to state 
(p. 398) that it was unlikely that southern England 
during the climax of the glacial cold had much if any 
vegetation to boast of, and continues, " It is certain, 
however, that it was clothed and peopled by an Arctic 
flora and fauna when the climatic conditions were 
somewhat less severe, relics of that flora having been 
detected at Bovey Tracey." He believes, therefore, 
that an Arctic flora took possession of England as 
soon as the climate enabled it to live in the country. 
Arctic plants, according to this explanation of the 
sequence of events, were the first immigrants to 
reconquer the dreary, plantless wastes and make 
them habitable for mammals. 

Fortunately these views do not at all agree with 
those of many of our leading European botanists and 
others entitled to have a voice in the matter. Pro- 
fessor Warming is of opinion that the main mass of 
the present flora of Greenland survived the Glacial 
period in that country (p. 403); whilst Professor 


Drude has shown (p. 288) that all plant life could 
not possibly have been destroyed in northern 
countries. He maintains that the greater part of 
the Arctic floral elements which unite Greenland 
and Scandinavia must have survived the Glacial 
period in these countries in sheltered localities. 
Indeed, he justly remarks, where at the present 
moment do we find such plantless wastes ? Green- 
land, Franz- Josef Land, and Grinnell Land, situated 
in high Arctic latitudes, all have a flora composed 
of flowering plants and cryptograms. " I cannot 
understand," he continues (p. 286), " why a flora, 
possibly mixed with northern forms but in the main 
points agreeing with our present floral elements, 
should not have persisted throughout the Ice Age 
even in the heart of Germany." "To my mind," 
says Col. Feilden, the well-known Arctic traveller 
(b, p. 51), "it seems indisputable that several plants 
now confined to the polar area must have originated 
there, and have outlived the period of greatest ice- 
development in that region." The theory in favour of 
a survival of the pre-glacial flora has been especially 
strengthened by the late Mr. Ball (than whom 
probably no botanist possessed a better knowledge 
of Alpine plants), who was strongly in favour 
of this view as far as the Alps are concerned. 
"Is it credible," he says (p. 576), "that in the 
short interval since the close of the Glacial period 
hundreds of very distinct species and several genera 
have been developed on 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 dis- 
continuous mountain chain some 1,500 miles in length, 
from the Pyrenees to the Eastern Carpathians ? " Mr. 
Ball's remarks, indeed, just touch upon a very important 
characteristic of all the so-called Alpine plants. In 
Europe they chiefly occur in Scandinavia and the 
central and southern mountain ranges, whilst they 
are mostly absent from the intervening lowlands. 
Again, we find a large number of species in the 
mountains of Central Asia and in some of the North 
American mountains. Almost all species of Alpine 
plants, in fact, are examples of discontinuous distribu- 
tion; and this, as every naturalist knows, is always, in 
both animals and plants, a proof of antiquity. 

The glacial or Alpine flora is very old, and must 
have originated long before the Ice Age. But it might 
be urged, why should these plants be now almost con- 
fined to the Arctic regions and the higher mountain 
ranges, where the temperature undoubtedly is very 
low, if they had originated during a pre-glacial 
period probably much milder than the present? 
The answer can be given by those who have made 
Alpine plants their special study, and who have 
attempted to grow them by administering to them 
a temperature and such climatic conditions as to 
be most conducive to good health. We should all 
expect these plants to be very robust, and especially 
to be able to stand extremely low temperatures. But, 


strange to say, the very opposite is the case. Pro- 
fessor Blytt tells us (p. 19) that "Arctic and Alpine 
species in the Christiania Botanic Gardens endure the 
strongest summer heat without injury, while they are 
often destroyed when not sufficiently covered during 
winter." The English climate then, one would 
think, ought to suit these plants, since the winters 
are not too cold; but we find that at Kew Gardens 
the large collection of Alpine plants have to be 
wintered in frames under glass in order to keep 
them in good health ; and Professor Dyer, the 
Director of the Gardens, thinks they are mostly 
intolerant of very low temperatures (compare also 
pp. 161-164). 

Such being the constitution of Alpine plants, how 
could they possibly have originated during the Glacial 
period and wandered from the mountains into the 
plains, across numbers of formidable barriers, often 
exposed to icy winds, for thousands of miles? As a 
matter of fact, Alpine plants have survived in the 
high North and in the Alps because they are there 
permanently protected during winter by a covering of 
snow from very low temperatures, and they are at the 
same time prevented from drying up. If they are 
given sufficient moisture and a constant, mild tempera- 
ture they seem to do very well. Such conditions are 
afforded them in many parts of the British Islands, and 
we find indeed the Mountain Avens (Dryas octopetala), 
one of the most typically Arctic plants, growing wild 
in profusion on the coast of Galway, in Ireland, at sea- 


level. The winter temperature of that part of Ireland 
resembles that of southern Europe, being no less than 
12 Fahr. above freezing point This fact appears 
to strengthen the view not only that the Alpine 
flora is of pre-glacial origin, but that the climate of 
Europe during the Glacial period was mild. 

Having now shortly reviewed the state of our 
knowledge with regard to the former presence in 
our temperate latitudes of Arctic animals and plants, 
it still remains for me to give a succinct statement of 
the light thrown by this fauna and flora on the wide- 
spread phenomena of glaciation. It is necessary to 
do so, because, though the greater development of 
glaciers on the mountains of Europe in former times 
does not presuppose the prevalence of an Arctic 
climate, the survival through the Ice Age of a fauna 
and flora could not possibly have taken place in 
northern Europe if the theories of glaciation now so 
much in vogue are really true. Professor Geikie 
reminds us, in speaking of his native country (p. 
67), that "we must believe that all the hills 
and valleys were once swathed in snow and 
ice; that the whole of Scotland was at some 
distant date buried underneath one immense mer 
de glace, through which peered only the higher 
mountain tops." That under such conditions no 
fauna or flora to speak of could have survived in 
Scotland is evident. Then again he argues (p. 426) 
that because in the great plain of Europe we meet 
occasionally with striated rock-surfaces and roches 


moutonnfos very similar to those produced by the 
glaciers of Switzerland, it must have been traversed 
by "inland ice" flowing from Scandinavia and the 
Baltic southward. The boulder clay of Germany is 
supposed to have accumulated underneath this 
vast " mer de glace" as he calls it. There is 
no question here of a simple local development 
of glaciers, such as could have existed under a 
mild and moist climate; practically all the plants 
and animals would have been annihilated in northern 
Europe under such conditions, as there were no areas 
free from ice. A more vivid idea of the state of 
Europe during the epoch of maximum glaciation 
will be obtained by looking at Professor Geikie's 
map (p. 437). The whole of Scandinavia, Iceland, 
Scotland, Ireland, and Switzerland is there repre- 
sented as having been completely enveloped in ice, 
and also the greater part of Russia, Germany, and 
England. In speaking of Scandinavia (p. 424) he 
remarks that "the whole country has been moulded 
and rubbed and polished by one immense sheet 
of ice, which in its deeper portions could hardly 
have been less than 5000 feet or even 6000 
feet thick." The greater portion of the area in- 
dicated as having been underneath a sheet of ice is 
thickly covered with superficial accumulations of 
gravel, sand, and clay. The latter is generally 
spoken of as " boulder clay," and, with the associated 
sand and gravel, it may be observed equally well in 

Russia or Germany, in England or Ireland. As a 



rule these stony clays thicken out as they are traced 
from the high-lying tracts to the low grounds; and 
especially near the mountains the rock-surfaces are 
often polished and striated. " For many years it was 
believed," continues Professor Geikie (p. 432), "that 
all those superficial deposits were of iceberg origin. 
The low grounds of Northern Europe were supposed 
to have been submerged at a time when numerous 
icebergs, detached from glaciers in Scandinavia and 
Finland, sailed across the drowned countries, dropping 
rock-rubbish on the way. Such was thought to have 
been the origin of the erratics, stony clay, and other 
superficial accumulations, and hence they came to be 
known as the 'great northern drift formation."' " But," 
he adds (p. 433), " when the phenomena came to 
be studied in greater detail and over a wider area, 
this explanation did not prove satisfactory. The 
facts described in the preceding paragraphs the 
occurrence of striated surfaces and roches mou- 
tonnees, the disturbed appearances associated with 
the till, and the not infrequent presence of giants' 
kettles convinced geologists that all the vast 
regions over which boulder-clay is distributed were 
formerly occupied by the 'inland ice' of Scan- 

I think Professor Geikie over-estimates the value of 
the evidences which appear to be in favour of his 
theory. His treatise on the Ice Age leaves one 
under the impression that the older view of the 
marine origin of the boulder-clay is not only 


done with for good and all, but that no geologists 
nowadays believe in it. If a more careful study of 
the glacial phenomena has led most geologists to 
abandon what I might call the "marine view" in 
favour of the terrestrial one, a more careful study 
of the fauna and flora will, I venture to think, have 
the opposite effect. However, it appears that even 
from a purely geological point of view more can 
be said in favour of the old theory than Pro- 
fessor Geikie and his school are ready to admit. 
Thus we are told by Professor Bonney (p. 280), 
in referring to the boulder-clay, that "the singular 
mixture and apparent crossing of the paths of 
boulders are less difficult to explain on the hypo- 
thesis of distribution by floating ice than on that of 
transport by land-ice, because, in the former case, 
though the drift of winds and currents would be 
generally in one direction, both might be varied at 
particular seasons. So far as concerns the distribu- 
tion and thickness of the glacial deposits, there is not 
much to choose between either hypothesis; but on 
that of land-ice it is extremely difficult to explain 
the intercalation of perfectly stratified sands and 
gravel and of boulder-clay, as well as the not in- 
frequent signs of bedding in the latter." " Anything," 
writes Professor Cole (p. 239), "that keeps open the 
position maintained by Lyell and others, that 
extensive glaciation is compatible with mild and 
sheltered nooks and corners, and that much of 
the distribution of boulder-clay was performed in 


seas and not on land, may be welcomed by 
rationalists, at any rate until further research has 
been carried on among the Arctic glaciers. At 
present every year brings evidence of modern 
marine boulder-clays in high latitudes, and removes 
us farther and farther from belief in a moraine 
profonde" That foraminifera are occasionally found 
in boulder-clay has been known for a long time, 
but it is only within recent years that these marine 
organisms have been shown to occur in so many 
localities, that Mr. Wright, who examined a large 
number of samples, says (p. 269), " I am forced 
to the conclusion that the Scottish as well as 
the Irish boulder-clay is a true marine sedimentary 
deposit " 

In the fourth and fifth chapters I shall return to 
this subject again, and mention a number of facts 
of distribution which appear to me much easier of 
explanation by means of the marine than by the land- 
ice theory. But I do not propose to go into further 
geological details in this volume, as I think I have 
clearly conveyed my position in this controversy. 

Before concluding this short review of the glacial 
problem, so far as it affects the origin of the European 
fauna, I should like to refer to the opinion of one 
who has devoted years to the study of the glacial 
phenomena in the Arctic Regions, viz., Col. 
Feilden. "To a certain extent," he says (a, p. 57), 
"all boulder clays at home arc fragmentary when 
compared with the boulder-bearing beds of Kolguev, 


which we may safely assume are 50 miles in length 
by 40 in width, with a thickness of not less than 250 
feet, probably far more, all lying in one undisturbed 
mass. It is suggestive that all the glacial deposits 
which I have met with in Arctic and Polar lands, 
with the exception of the terminal moraines now 
forming above sea-level in areas so widely separated 
as Smith's Sound, Grinnell Land, North Greenland, 
Spitsbergen, Novaya Zemlya, and Arctic Norway, 
should be glacio-marine beds. Throughout this 
broad expanse of the Arctic Regions I have come 
across no beds that could be satisfactorily assigned 
to the direct action of land-ice; that is to say, beds 
formed in situ by the grinding force and pressure of 
an ice-sheet. On the contrary, so far as I can judge, 
the glacial beds which I have traced over the exten- 
sive area mentioned above have all been deposited 
subaqueously and re-elevated." 

One of the strongest arguments that can be used 
against the view of the marine origin of the glacial 
phenomena in Northern Europe seems to me the fact 
that we find polished rock-surfaces far removed from 
the source of glaciers, and so exactly resembling those 
produced at the present day by our Alpine glaciers as 
to appear identical to the experienced eye. Most of 
such striated and polished rocks occurring in the 
higher mountain ranges of Scandinavia, and also of 
the British Islands, have no doubt been actually pro- 
duced by glaciers, whilst those in the plain, some- 
times hundreds of miles away from the mountains, 


must have originated in a similar manner; that is to 
say, by a heavy mass of material containing stones 
being slowly dragged over the rock-surfaces. The 
weight which causes the stones to polish the latter is 
generally ice, but it is quite conceivable that any other 
substance, especially if it is in a semi-solid state, must 
act and operate in much the same way. All polished 
rock-surfaces are carved by glaciers, because we can 
see them done by glaciers every day, is the argument 
commonly used nowadays. It was not so formerly. 
But Mr. Mallet and his views are almost forgotten 
now; his name does not even appear in our great 
modern works on the Ice Age. His argument was 
that as the land rose out of the glacial sea, the mud 
which had accumulated round the shore slipped 
downward in a direction determined by the contour 
of the surrounding valleys and mountains. The 
moment the land rose above water-level, the large 
mass of gravel and mud lying upon it slipped down- 
ward. During a steady rising of the land there would 
therefore be produced a continuous sliding down of 
this mud-glacier, which would groove and polish the 
rock underneath it, in the same manner as the ice- 
glaciers do in the Alps (p. 47). Professors Sedgwick 
and Haughton became strong adherents of Mr. 
Mallet's theory at the time, but it seems later on 
to have fallen into disfavour with geologists, who 
may not even be thankful to have it brought to 
light again. 



I have endeavoured to show in this chapter how we can deter- 
mine approximately the original home of an animal. By this 
means we are able to study the component elements of the 
European fauna, which is found to consist to a large extent of 
migrants from the neighbouring continents. There is a Siberian, 
an Oriental, and an Arctic element in it. The remainder of the 
fauna is derived from local centres of dispersal. "What was 
formerly believed to have been one great northern migration 
now resolves itself, on closer study, into two very distinct 
ones the Siberian and the Arctic. The mammals have 
received most attention hitherto, because their remains are so 
frequently met with, thus enabling us more easily to investigate 
their past history; but butterflies and snails have not been 
neglected, and at least one very remarkable work on the latter 
has been published dealing with their origin in Europe and 
in the remainder of the Palsearctic region. 

The former distribution of land and water is intimately con* 
nected with the origin of the European fauna, and the changes 
which have taken place in this respect may be best traced by 
the present distribution of mammals, snails, and earthworms. 
In this manner the British Islands may be shown to have been 
connected with one another and with the Continent; Spain 
with Morocco across the Straits of Gibraltar; Greece with Asia 
Minor, and so forth. 

The British fauna has played such an important part in the 
evolution of the European fauna, that it forms the key to the 
solution of the wider problem. In it five elements are 
recognisable, of which the Lusitanian element is the oldest, 
and the Siberian the most recent. It has been deemed 
advisable to conclude this chapter with a short review of the 


history of the Glacial period in its climatic effects on the 
animals and plants of Europe. A number of writers are 
quoted who have conducted special researches in determining 
the temperature of our continent at the time. The fauna of 
Europe is frequently described as having been of an Arctic 
nature, but as a matter of fact there existed during the Ice 
Age a striking and most remarkable mingling of a northern 
and a southern fauna. The presence of Siberian mammals in 
Europe is said to have been due to the prevalence of a dry 
steppe climate, but this view is not supported by other evidence. 
The Alpine flora in a wide sense is probably pre-glacial in 
origin, and appears to have survived the Ice Age where it 
is now known to exist. A few words on the phenomena of 
glaciation are added before bringing the chapter to a close. 



THE British Islands are, as I have remarked, very 
suitable as a starting-point for our investigations. 
Their fauna and flora are fairly well known, and the 
distribution of the large animals at any rate, which 
are of course of much importance in these researches, 
has been as much studied as that of any other area 
in Europe. We possess in England an abundance of 
the remains of past animal life, and a combination of 
the data furnished by both of these important factors 
will enable us to draw up a history of the origin of 
the present British fauna. 

In the first chapter I indicated that in the fauna of 
the British Islands three divisions or elements are 
recognisable a northern, a southern, and an eastern. 
These elements correspond to migrations which can 
be proved to have arrived in this country at different 
periods in past times. When we investigate these 
migrations more closely, the eastern is found to be 
composed partly of European and partly of Siberian 
species. The southern is made up of European 
and of Central and Southern Asiatic species. To 
make matters still more complex, the southern and 



eastern migrations insensibly merge into one another, 
so that it is often very difficult to determine to which 
of them an animal may belong. The European 
species spread principally from three centres over 
Europe viz., from the Lusitanian, Alpine, and the 
Balkan centres. The southern element of the British 
fauna is therefore composed of animals which have 
originated in these three centres, and in Central and 
Southern Asia. The Balkan species have been 
included with those coming from the latter centre 
under the term "Oriental" migration. The sixth 
chapter is devoted to it, whilst the Lusitanian and 
Alpine migrations have each a chapter to them- 

The Arctic Hare is, as I have already mentioned, 
one of the mammals of the northern element of the 
British fauna. It is now confined to the mountains of 
Scotland and the plain and mountains of Ireland. 
But in former times it had a wider range in the 
British Islands. The Stoat is another distinctly 
northern mammal. It occurs with us, as Messrs. 
Thomas and Barrett-Hamilton have pointed out, 
in two distinct varieties or species, the one being 
confined to Great Britain, the other to Ireland. As 
I shall explain more fully later on (p. 135), I have 
reasons to believe that the Irish Stoat came from the 
Arctic Regions as a northern migrant, but that the 
English Stoat, on the other hand, reached England 
with the Siberian fauna from the east. A third 
northern animal, now extinct in the British Islands, 


is the Reindeer. It is supposed to have died out in 
these countries not very many centuries ago, and 
records have been handed down to us that it still 
inhabited Scotland as late as the thirteenth century. 
Like the Stoat, it occurred in two well-known varie- 
ties, distinguished from one another by the shape 
and form of the antlers. In the English pleistocene 
deposits the remains of both kinds are met with 
mingled together, whilst in Ireland only one of them 
has been found. The explanation of this case is 
similar to that of the two stoats. One of the varieties, 
which we may call the northern one, came to us from 
the Arctic Regions; the second wandered to the 
British Islands at a later period, when Ireland had 
probably become separated from England. It was 
therefore unable to penetrate so far west. 

One of the most familiar examples of a northern 
British bird is the Red Grouse (Lagopus scoticus). By 
most authorities it is looked upon as a species distinct 
from the Scandinavian Willow Grouse (Lagopus 
albus\ but except in colour it is undistinguishabie 
from it, and the eggs are identical. The whole genus 
Lagopus is a distinctly Arctic one, and there can be 
no doubt that the British Grouse belongs to the 
northern migration, just like the Arctic Hare. The 
Ptarmigan (Lagopus mutus) and the Snow Bunting 
are also migrants from the north. Though as resident 
British birds they are quite confined to Scotland, the 
remains of the former have been found in a cave in 
the south of Ireland, showing that its range in the 


British Islands was formerly more extensive. Another 
bird which probably came to our shores with this 
same migration, though it is now unfortunately ex- 
tinct, is the Great Auk (Alca impennis), of which 
some specimens have luckily been preserved in our 
museums. From the occurrence of its remains in 
kitchen-middens and other recent deposits, the Great 
Auk is known to have inhabited the coasts of Scotland, 
Ireland, and Scandinavia, as well as those of New- 
foundland. Mr. Ussher recently found the bones of 
this bird near Waterford, which, I believe, is the most 
southern locality known. The manner of their occur- 
rence leaves no doubt that the bird had been used as 
food by the early races of man. In all probability it 
originated in the Arctic Regions, and subsequently 
spread south on either side of the Atlantic. We 
need not here refer to the many winter visitants, 
northern birds which appear regularly, or at more 
or less long intervals, in these islands, although in 
most of the ornithological works they are included 
under the term "British Birds." 

All the British reptiles and amphibia appear to 
have reached us from the south or east, but among 
the fishes there are a good many northern forms. 
The whole salmon family the Salmonidcs are 
typical northern immigrants. The Stickleback (Gas- 
terosteus aculeatus\ too, has undoubtedly come to us 
from the north. The genus Cottus, like Gasterosteus> 
is certainly Arctic in origin. Originally freshwater 
forms, many species are now found between tide- 


marks, and of these a few have migrated southward 
along the coasts of the great continents. Thus we 
meet with various species of Coitus as far south as 
California and Japan, on the American and Asiatic 
coasts of the Pacific respectively. In Europe, two 
species, viz., C. scorpio and C. bubalis, range as far 
south as the French coast. Our freshwater Cottns, 
the Miller's thumb (Cottus gobio), has migrated to 
us from the north with the Arctic species. All the 
freshwater forms, indeed, of this genus are typically 

A large number of land and freshwater invertebrates 
too have no doubt reached us from the north. Some 
of them may have originated in Scandinavia or within 
the Arctic Circle, but others probably came still 
farther, either from America or even from Asia, and 
used the Arctic land-connection via Greenland in their 
migration to Europe. As I shall give a number of 
additional instances of such migrants in the succeed- 
ing chapters, I need not, perhaps, dwell upon them 
now any longer, except to mention a few of the 
more typical ones. Vertigo alpestris, a minute snail 
with an amber-coloured shell, and our freshwater 
pearl-mussel, Unto (Margaritana) margaritifer y belong 
to this migration. Then among butterflies we may 
cite the Marsh-ringlet (Coenonympha t)phon\ and 
among beetles, Pelophila borealis and BletJiisa multi- 
punctata. There are a number of northern spiders, 
among which a few certainly indicate an Arctic 
origin, or at any rate, that they have wandered to 


Europe across Greenland and the old Arctic land- 
connections. Bathyphantes nigrinus^ Linyphia insignis, 
and Drapetisca socialis, for instance, are three British 
species whose range indicates a northern origin, and 
which also occur, according to Mr. Carpenter, in 
North America. Mr. Carpenter also tells me that the 
Collembolan, Isotoma littoralis, is a typical northern 
migrant He has recently discovered it in the west 
of Ireland, its only station in the British Islands. 

Among the Crustacea, the genus Apus forms an 
exceedingly interesting illustration of the northern 
migration, Apus glacialis having been discovered in 
a Scottish pleistocene freshwater deposit, whilst it is 
now almost confined to the Arctic regions. 

To the same group of animals also belong the three 
remarkable species of freshwater sponges, Ephydatia 
crateriformis, Heteromeyenia Ryderi, and Tubella pen- 
sylvanica y which Dr. Hanitsch has described from some 
lakes in Western Ireland. None of these are known 
from Great Britain or from the continent of Europe. 
A few North American plants grow wild in the same 
district. That any of these should owe their existence 
in Ireland to accidental introduction appears to me 
exceedingly improbable. In a former contribu- 
tion to this subject (a, p. 475) I assumed that 
these American plants and animals had migrated 
to Europe at the same time as the other northern 
forms referred to. My friend Mr. Carpenter, how- 
ever, takes exception to this (p. 383), and I quite 
recognise the force of his argument. "Their very 


restricted and discontinuous ranges," he says, " along 
the extreme western margin of Europe mark them 
as decidedly older than those northern animals and 
plants which have a circumpolar distribution." We 
have indeed quite similar examples in the Oriental 
migration, of which part is very ancient, surviving 
here and there and exhibiting discontinuous distri- 
bution. We may therefore look upon these American 
immigrants as among the oldest members of that 
northern stock which have survived in our islands 
probably a mere remnant of a once luxuriant flora 
and fauna. 

In order to show the importance of the Eastern or 
Siberian element in the English, or, we might say 
with Dr. Sclater, the Anglo-Scotian mammalian 
fauna, I herewith give a list of the species of 
mammals which probably migrated to Great Britain 
from Siberia. I have marked with an asterisk those 
which still exist in this country (not in Ireland), or 
have become extinct within historic times. 

Canis lagopus. * Mus minutus. 

Gulo luscus. * Arvicola agrestis. 

* Mustela erminea. * amphibius. 

* putorius. arvalis. 

* vulgaris. * glareolus. 

* Sorex vulgaris. ,, gregalis. 
Lagomys pusillus. ,, ratticeps. 

* Castor fiber. Equus caballus. 
Spermophilus Eversmanni. Saiga tartarica. 

erythrogenoides. Ovibos moschatus. 


Cricetus songarus. Alces latifrons. 

Myodes lemmus. machlis. 

Cunictilus torquatus. Rangifer tarandus. 

We have evidence that most of these twenty-six 
species of mammals came from Eastern Europe, but 
there is no reason to suppose that they originated there. 
On the contrary, it is highly probable, as I said be- 
fore, that their native home is Siberia, and that they 
entered Europe to the north of the Caspian. Along 
with these, vast numbers of other forms of life, and 
also plants, swarmed into our continent, and as we 
advance eastward from England we meet with them 
in increasing numbers to the present day. But not 
only on the Continent do we find these survivals of 
the great Siberian migration, which has been so ably 
described by Professor Nehring; no less than nine 
species still inhabit Great Britain (if we include the 
recently extinct Beaver). On the other hand, not 
more than three have been found fossil in Ireland, 
and of these only one still survives. This very signifi- 
cant fact will be referred to again more fully on p. 153. 
Meanwhile it should be remembered that these 
three species, viz., Mustela erminea, Equus caballus, 
and Rangifer tarandus, occur in Ireland in varieties 
distinct from those found in Central Europe. It is 
upon this, and many other circumstances, that 
I founded my belief that Ireland was already 
separated from England at the time of the arrival 
of the Siberian emigrants in the latter country. As 


we shall see, the Irish Stoat, Horse, and Reindeer 
probably came by a different route from that taken 
by the English representatives of the same species. 

Very few of the lower animals of Siberian origin 
have reached the British Islands. Most of those 
which were formerly thought to be Siberian are either 
of East European or of Central and South Asiatic 
origin, though they probably joined the Siberian 
migration on their way to England. The Arctic 
migration brought a greater variety of species to 
this country than the Siberian, but neither the one 
nor the other has contributed more than a small per- 
centage to the British fauna. The bulk of that fauna 
is derived from the various European centres of dis- 
persal, and especially from Central and Southern Asia. 

Those animals which have their home in the latter 
area, I have named Orientals, though it must be re- 
membered that they need not necessarily have come 
from what is known among zoologists as the " Oriental 
Region." The terms " Oriental animals " and " Ori- 
ental migration " are used here in a wider sense, and 
include even those species which reached Central and 
Northern Europe from South-Eastern Europe. It is 
astonishing, what a vast number of both vertebrate 
and invertebrate animals can be traced back to this 
Oriental migration. Great tracts of Europe were 
repeatedly submerged beneath the sea during Tertiary 
times, and on their re-appearance were formed into 
green fields and pastures new for the rich Asiatic 
fauna, which was ever ready to flood the neighbouring 



continent. This went on, and not for a comparatively 
short space of time, as in the case of the Siberian 
invasion ; the immeasurable ages which passed, 
whilst several of the Tertiary epochs dawned upon 
Europe, witnessed an almost constant stream of 
Asiatic immigrants pouring in upon us. Europe 
returned her own products in exchange, but they 
must have been scanty in comparison to the enor- 
mous number of species which have undoubtedly 
originated in Central and Southern Asia. Very 
many of the widely distributed forms in the British 
Islands are of Oriental origin. Among these are 
also the cosmopolitan species, such as the Barn 
Owl (Strix flammed] and the Painted Lady Butter- 
fly ( Vanessa cardui}. A great number of our 
British Mammals, Birds, Butterflies, and Beetles have 
come to us with the Oriental migration. But, as 
I shall explain in the special chapter devoted to it, 
the earlier migrants from the south-east found their 
northward progress barred by a great sea which 
stretched through Central Europe from west to east. 
The Mediterranean was then divided into two smaller 
basins. On their arrival in Greece, which was then 
connected with Asia Minor and Southern Italy, 
the Oriental migrants seem to have turned westward, 
skirting the shores of the Mediterranean. When they 
finally reached Spain, many then changed their course 
northward (see Fig. 5, p. 117) and wandered to the 
British Islands with the Lusitanian animals which 
came from South- Western Europe. 


Dr. Wallace makes mention of a fairly large num- 
ber of species and varieties of Lepidoptera, Coleoptera, 
and land and freshwater Mollusca, supposed to be 
peculiar to the British Islands. Even if these were all 
found to be of British origin, most of their nearest 
relatives are continental species. Many, however, must 
be looked upon as mere races or sub-species of 
familiar continental forms. But others, such as Geo- 
jualacus maculosus and Asiminea Grayana, are by no 
means confined to the British Islands. Some of 
the so-called varieties enumerated by Dr. Wallace 
are merely slight individual variations in form and 
colour, which, only by the extraordinary tendency of 
the variety-monger to advertise himself, have received 
a distinct Latin denomination. The number of the 
remaining species, after weeding out the unworthy 
ones, will be found to be insignificant. 

Similarly, the list of seventy -five species and 
varieties of flowering plants included by Dr. Wallace 
among the forms peculiar to the British Islands 
(p. 360) is reduced by Sir Joseph Hooker to twenty. 
The remainder are to be considered as varietal forms 
of a very trifling departure from the type, or as 

Just as we distinguish in the British Islands the 
parts inhabited by Englishmen, Scotchmen, and Irish- 
men, so we can recognise three divisions in the animal 
world, and these roughly correspond to the boundaries 
of England, Scotland, and Ireland. Most of the 
eastern species inhabit England, most of the northern 


ones are confined to Scotland, whilst Ireland is occu- 
pied chiefly by southern animals. This, however, is 
only a very rough-and-ready method of sub-dividing 
the British Islands into their component parts 
according to the origin of their faunas. On closer 
study such a division is found to be unsatisfactory. 
The eastern species do not really stop at the Scottish 
frontier, they range far into Scotland. Nor are the 
northern forms confined to the latter country. Many 
of them range into Ireland, and also into England. I 
have constructed a map of the British Islands showing 
approximately the boundaries of the northern, eastern, 
and southern species (p. 7), but even this may not 
altogether meet with the views of an ornithologist 
or conchologist. For every group of animals the 
boundaries would probably require to be marked 
differently. There is also a good deal of overlapping, 
so that the attempt to define the limits of the various 
elements meets with great difficulties. But the map 
represents, I think, fairly well the general impression 
one receives as to the disposition of its component 
elements, after a careful study of the British fauna 
as a whole. 

The distribution of the British plants has been 
worked out much more thoroughly than that of the 
animals. It need not surprise us, therefore, that the 
first attempt to separate the British Islands into 
natural divisions was made by a botanist the late 
Mr. Watson. As he himself pointed out, in making 
these divisions he did not take into consideration the 


origin of the British species. They represent merely 
groups of assemblages of plants of different types 
of vegetation. Edward Forbes, on the other hand, 
founded his districts on the origin of plants. His 
work is not only the first of the kind, but it is a 
classical essay, and remains one of the most remark- 
able contributions to the literature on the geographical 
distribution of living organisms known to science. 
The vegetation of the British Islands, he informs us 
(p. 4), presents a union of five well-marked floras, 
four of which are restricted to definite provinces, 
whilst the fifth, besides exclusively claiming a great 
part of the area, overspreads and commingles with 
all the others. These are 

I. Mountainous districts of South-) T 

west and West of Ireland . . [Lusitaman type. 

II. South-west of England, and),- ,,. 

South-east of Ireland . , .) Galilean type. 

III. South-east of England. 

IV. Mountains of Scotland, Cumber-) c j- 

land, and Wales . . . .j Scandinavian type. 

V. General Flora Germanic type. 

Professor Forbes points out, in connection with the 
plants of the Germanic type, that the fauna accom- 
panying this flora presents the same peculiarities and 
diminishes westward and to the north. This type 
includes, therefore, almost all the species which can 
be shown to have come to us directly from the east, 
few if any of which have penetrated to Ireland. 

On a previous occasion, the same author had 


divided the British Islands into ten districts, accord- 
ing to the distribution of their molluscan fauna. 
These are 

I. The Channel Isles. 
II. South-east of England (including Cambridgeshire). 

III. South-west of England. 

IV. North-east of England. 

V. North-west of England (including Isle of Man). 
VI. North of Ireland. 
VII. South of Ireland. 
VIII. South of Scotland. 
IX. North of Scotland. 
X. Shetland Isles. 

In a short paper on this subject (b, p. 5), I have 
shown that some of these districts are founded on 
erroneous data, whilst, with the knowledge now at 
our disposal, others can no longer be maintained as 
distinct. I thought then that the molluscan fauna 
warranted a division of the British Islands into the 
following two provinces : 

I. England and Wales (except the South-west). 
II. South-west of England and Wales and the whole of 
Ireland and Scotland. 

The second district contains some species of mol- 
luscs which are almost entirely absent from the first, 
such as Geomalacus maculosus, Testacella Maugei^ 
Helix piscina, Helix revelata, Helix acuta, and Pupa 
ringens. These are all of Lusitanian origin, and do 
not occur in Central Europe. Scotland alone cannot 


be classed as a separate province, since it does not 
contain a single species peculiar to itself. But, along 
with Ireland and the South-west of England and 
Wales, it is distinguished from the remainder of 
these countries by the almost total absence of what 
have been called Germanic types. 

A French conchologist, the late Dr. Fischer, dealt 
with the British molluscan fauna in a somewhat 
similar spirit (p. 57). He divided the British area 
into two districts, but these differ from mine in so 
far as the South-west of England and Wales and the 
West of Ireland form one ; the remainder of England 
and Ireland as well as the whole of Scotland the 
other. His classification is of particular interest, 
since the first district represents part of a larger 
Atlantic province, the second a portion of the Ger- 
manic province of the European sub-region. The 
latter he looks upon as one of the sub-regions of 
the great Palsearctic Region. Attention is thus 
drawn to the intimate relationship existing between 
the western parts of the British Islands and the 
Spanish peninsula on the one hand, and between the 
eastern portions and Central Europe on the other. 

Mr. Jordan's North-Sea-and-Baltic district includes 
Scotland and the North of Ireland, whilst England 
joined with the West and South of Ireland forms part 
of his Celtic province. Both of these districts or pro- 
vinces belong to Mr. Jordan's greater Germanic 
Region (p. 302). 

In the collection illustrating the geographical dis- 


tribution of animals in the Dublin Museum, the 
British species have been grouped into three divisions. 
One contains those with a wide range over the British 
Islands, another the characteristic forms of the south- 
east and lowland districts of Great Britain, and the 
third the Irish and the western and highland Anglo- 
Scotian species. Mr. Carpenter has named the last 
two divisions the "Teutonic" and the "Celtic" More 
recently, he has recognised that this last division 
contains two distinct groups ; one including animals 
of northern, the other those of southern origin. 
He acknowledges indeed, just as I do, three distinct 
faunas in the British Islands, with the addition of 
the group of generally distributed species of un- 
determined origin. 

Many other naturalists have worked in the direc- 
tion I have indicated namely, in grouping the 
British animals into several distinct assemblages, 
without, however, taking their foreign range into 
consideration, or their origin. I have already referred 
to the useful work done by botanists, who have been 
the pioneers in the science of the geographical dis- 
tribution of living organisms. Among the British 
naturalists who have applied the principles of Watson 
to zoology, A. G. More deserves to be specially men- 
tioned. He was the first to make a serious study of 
the British fauna on the lines laid down by that dis- 
tinguished botanist. In conjunction with E. Boyd, 
he published a valuable essay on the " Distribution of 
Butterflies in Great Britain," and later on the birds 


were similarly dealt with. All the more important 
groups of animals are now being studied with a view 
to determining their exact range in these islands. 
Mr. Harvie-Brown, Mr. J. W. Taylor, Mr. Eagle Clarke, 
Mr. Miller Christy, Mr. Ussher, Mr. Harrington, and 
a number of others have considerably advanced our 
knowledge in this direction in recent years. 

Any such contributions are to be welcomed as 
furnishing us with the necessary data to solve the 
problem of the origin of the British fauna. Mean- 
while we know enough to enable us to assert 
positively that the latter has reached us by land- 
connections from various parts of Europe (cf. p. 35). 
This statement of course refers to the bulk jgf the 
Britishfauna^__The small proportion of indigenous 
species, or such as have been introduced accidentally, 
may be left out of consideration when dealing with 
the great mass of animals which have evidently 
migrated to the British Islands on land now sunk 
beneath the sea (see Fig. 4, p. 60). Opinions of 
zoologists, botanists, and geologists are practically 
unanimous on this subject ; yet there are two other 
theories, which have from time to time been advanced 
to arrniini-fnrJJTej^i^itL^if thgJRrifrfch fauna. Only 

the first of these, however, can claim the serious 
attention of those interested in the problem. Its 
chief contention lies in the oft-asserted dictum of 
the "imperfection of geological record" It has been 
suggested, in fact, that the British fauna, instead 
of having migrated to our islands, might have 


orjgimited there, but that, owing to the fragmentary 
nature of our Tertiary deposits, all trace of their 
early history had disappeared. "The origin of 
European species," remarks Professor Cole (p. 238), 
"within the area of the British Isles, and their 
migration outwards when local conditions became 
less favourable for their multiplication, are pos- 
sibilities that seem too often disregarded. Yet 
the geologist must see in the western borderland 
of modern Europe a diminished continent from 
which land-animals must have again and again 
moved eastward." " Hence geologists may fairly be 
unwilling to look on our isles as barren lands waiting 
to be peopled in pliocene or later times. Far rather 
has the breaking up of a broad land-area along the 
present continental edge sent our land-fauna to the 
new steppes that opened eastward, leaving us a 
mere diminished remnant to struggle with the glacial 

There are in Professor Cole's views many points 
with which I readily agree. In the first place, he 
acknowledges that migration has taken place on 
land, so that we have our land-connection between 
Great Britain and the Continent whatever theory we 
accept as to the direction taken by the migrants. 
That the western borderland of Europe has given 
rise to many important assemblages of animals in 
past times, seems to me also exceedingly probable, 
nor do I look upon the British Islands as " barren 
lands waiting to be peopled in pliocene or later 


times." On the contrary, I believe an almost un- 
interrupted stream of migrants poured into the 
British Isles before pliocene times from the south. 
But what I thoroughly disagree with, is the remark 
that our British land-fauna has been sent to the new 
steppes that opened eastward. These are the more 
or less arid portions of Eastern Europe. Professor 
Cole no doubt has in mind those species of mammals 
which I have included in what I called the Siberian 
migration, and of which we have fossil evidence 
in the late Tertiary deposits of Europe. It would 
be impossible here to discuss this subject fully, 
especially as I have done so in the subsequent 
chapters; but, even if we had no geological record 
whatsoever, the present range of the species in 
question and their nearest relatives must convince 
us that they could not have originated in Western 
Europe. However, on the strength of the geological 
evidence, Professor Nehring the only one who has 
made this fauna his special study remarks (p. 228), 
that there seems scarcely any doubt that this steppe- 
fauna just referred to had come to us from the east. 
Professors Boyd Dawkins, Brandt, and Lartet held 
similar views. 

The theory that an ice-sheet stretched across a 
narrow sea might be the means of aiding a fauna 
across from the mainland to an island, is particularly 
inapplicable to the British Islands. Neither Mr. 
Kinahan nor Mr. Lamplugh, the two supporters of 
this view, have, however, taken the trouble to apply 


it to more than one species of the British fauna. An 
ice-bridge, thinks Mr. Kinahan, " could easily have 
connected Scotland and Ireland, thus giving a land 
causeway for migration" (p. 3). Mr. Lamplugh 
throws more light on this interesting speculation by 
giving us the name of an animal which he believes 
crossed a narrow sea on a bridge of ice. This animal 
unfortunately happens to be one whose remains have 
never been found in high northern latitudes, viz., the 
Irish elk (Cervus giganteus). And because he is of 
opinion that this species of extinct deer found its 
way to the Isle of Man from the mainland on a 
waning ice-sheet, he sees no reason why certain 
elements of the Irish fauna should not have been 
similarly introduced. 

It seems of no advantage to begin the discussion 
on the origin of the British fauna by assuming the 
former existence of ice-bridges, and the possibility of 
a migration across them of some of its members. If 
a glacier connected Scotland and Ireland, the climate 
of both countries (since they were highlands and 
acted as the feeders of the ice-sheet) must have 
been uncomfortable to the majority of the British 
species. What were the inducements that could 
have prompted those which had braved the dis- 
comforts of Scotland to emigrate to Ireland at 
such a time? What light does it throw on the 
origin of the Irish fauna as a whole, to advance the 
extremely improbable hypothesis that certain ele- 
ments of it may have reached Ireland by an ice* 


bridge? If any species came to that country in such 
an unusual manner, surely they must have been Arctic 
or northern forms. But what about the southern 
species, which form the bulk of the Irish fauna and 
also the flora? Even the Arctic element of the 
British fauna, which probably includes, besides the 
Reindeer, many hundreds of species, could not, I think, 
have migrated to these islands on an ice-bridge. In- 
deed, I agree with most of the writers who have dealt 
with the subject, in asserting that the northern as well 
as all the other elements of our fauna utilised for their 
migration the old land-bridges which connected these 
islands with one another and with the Continent. 

There is a greater diversity of opinion as to the age 
during which the British fauna arrived in these islands. 
This is naturally a much more complicated problem, 
but it is one which I am convinced will ultimately be 
solved mainly by means of a study of the geographical 
distribution of animals and plants. I ft we can settle 
the relative ages of the various migrations, wejhereby 
supply afTTTTTpoTtant fink in our attempt to reconstruct 
the past geographicallealures of the BritTsrTTslands. 
The range ofthe~7Brttish species will give us ^rr 
idea of the nature of the land-connections and their 
gradual changes in course of time. Geological data 
are exceedingly valuable in these inquiries, but it is 
a fatal mistake to build our geographical theories 
and the origin of the British fauna as a whole 
entirely on the assumptions of a certain school of 
geologists. Unfortunately, Dr. White's very interest- 


ing remarks on the British fauna for this reason lose 
much of the value which they might otherwise 

In his remarkable essay the late Edward Forbes 
affirms that the flora peculiar to the west of 
Ireland, of which the strawberry tree (Arbutus unedo) 
is the most striking example, and which exhibits such 
strong southern affinities, is not only much the most 
ancient of our island floras, but that it is actually of 
miocene age. It migrated to Ireland from Spain at 
a very remote period, during which he supposed that 
a direct land-connection existed between the two 
countries. The destruction of this old land-bridge, 
he thinks, must have taken place before the com- 
mencement of the Glacial period. Climatal changes 
during that time destroyed the mass of the southern 
flora which had thus reached Ireland, the survivors 
being species such as were most hardy (saxifrages, 
heaths, etc.), which he considers to be the only relics 
of this most ancient portion of our flora. 

The northern or Arctic fauna and flora, according 
to the same author, established themselves in the 
British Isles during the Glacial period at a time 
when these were groups of islands in the midst of 
an ice-bound sea. Finally, the great mass of our 
animals and plants migrated from the Continent to 
England after the Glacial period. " The migration of 
the species," he says, " less speedy of diffusion, which 
are now peculiar to England was arrested by the 
breaking up of the land-connection between England 


and Ireland, and thence the famous deficiencies of 
the sister isle, as, for instance, its freedom from 
reptiles" (p. 10). He is also of opinion, that the 
separation between England and the Continent took 
plade at a later date than that between England 
and Ireland. 

According to Dr. A. R.Wallace (p. 338), we possessed 
just before and during the Glacial period "a fauna 
almost or quite identical with that of adjacent parts 
of the Continent, and equally rich in species." But 
the submersion, he thinks, which is supposed to have 
occurred during the latter part of the Glacial period, 
destroyed the greater part of the life of our country. 
When England again became continental, continues 
Dr. Wallace, this fauna was succeeded by an assem- 
blage of animals from Central Europe. " But sufficient 
time does not seem to have elapsed for the migra- 
tion to have been completed before subsidence again 
occurred, cutting off the further influx of purely 
terrestrial animals, and leaving us without the number 
of species which our favourable climate and varied 
surface entitle us to." The comparative zoological 
poverty of Ireland he attributes to the fact that " the 
depth of the Irish Sea being somewhat greater than 
that of the German Ocean, the connecting land 
would there probably be of small extent and of less 
duration, thus offering an additional barrier to 

Dr. Wallace's explanation of the origin of the 
British fauna is disappointing after Forbes's careful 


study and critical inquiry into its component ele- 
ments. So great an authority on geographical 
distribution might have given us more lucid state- 
ments of his views on a variety of topics connected 
with this subject. 

In speaking of the fauna of Ireland, Professor 
Leith Adams, Professor Dawkins, and Mr. Alston are 
evidently only thinking of the mammals, which form 
but a very small proportion of it. The first-men- 
tioned palaeontologist held that there was a land- 
communication between Scotland and Ireland at the 
close of the Glacial period, by which the greater 
portion of the mammals that had found their way to 
the former country crossed to the latter (p. 100). 
And, he continues, the severance between the two 
countries must have taken place before the slow- 
travelling Mole, the Beaver, the forest-haunting Elk 
and the Roebuck had time to arrive. 

Much in the same spirit are Mr. Alston's remarks 
on this subject (p. 5). " The absence from the known 
fossil fauna of Scotland and Ireland of most of the 
characteristic post-pliocene English animals, shows 
that the northward migration of these forms was slow, 
gradually advancing as the glacial conditions of 
the northern parts of our islands decreased in in- 
tensity. Thus it is not difficult to suppose that the 
Hedgehog, Ermine, Badger, Squirrel, and Mountain 
Hare may have found their way through southern 
Scotland into Ireland long before they were able to 
penetrate into the still sub-arctic regions of the High- 

^ n- 


lands. Subsequently, when the improvement of the 
climate had continued, the Shrews and Voles may 
well have found their way northward along the com- 
paratively genial coasts, before the larger beasts of 
prey could find a sufficient stock of game." 

That the Bear, Wolf, Stag, Horse, Mammoth, and 
Reindeer lived in Ireland before the Glacial period 
is considered highly probable by Professor Boyd 
Dawkins (a, p. 152). 

Only the Butterflies are dealt with in Dr. Buchanan 
White's clever little essay on distribution. And, 
as I remarked before, his conclusions are some- 
what marred by the unwarrantable assumption that 
our islands at no distant date were totally destitute 
of all plant-life, and were therefore uninhabitable by 
animals. But his paper differs in so far from most of 
the others, that he has made a thorough study of the 
one group he deals with. In some respects it may serve 
as a model to future students in its general treatment 
of the problem he has set himself to work out He 
adopts the principle, even for butterflies, that though 
it is possible for them to be blown over from the 
Continent, they have probably migrated with the rest 
of our indigenous fauna and flora across the dry bed 
of the German Ocean. His conclusions are that 
Britain derived its butterfly fauna from continental 
Europe in post-glacial times, that the Arctic and 
Alpine species were the first arrivals, and that one 
part of the Irish species reached Ireland by way of 
Scotland, another from the south. He assumes, of 


course, that Great Britain and Ireland were connected 
at that time. 

Within the last few years the spell which has bound 
naturalists to accept the theory of a total destruction 
of life during the Glacial period is happily vanishing, 
and more enlightened views are gaining ground. 
The Lusitanian species of plants in the west of Ire- 
land, which had already furnished Forbes with an 
argument in favour of survival, are also regarded by 
Mr. Bulman as the remnants of a pre-glacial flora 
which was exterminated everywhere else by the 
cold (p. 265). This view of the survival of a pre- 
glacial fauna and flora has since been accepted by 
Mr. Carpenter, whilst I also have endeavoured to 
bring fresh evidence into the field in its favour. 
We both agree with Edward Forbes in considering 
the Lusitanian element as the oldest section of our 
fauna and flora, and that it came long before the 
Glacial period. But we differ somewhat from him, 
in so far as we do not limit that element to Ireland. 
It seems also to be represented in South-western 
England and Wales, though it is there less con- 

This decision as to the relative age of the British 
South-western fauna has not been arrived at from any 
geological considerations. The conviction that it 
must be older than the other sections has been gained 
solely from a study of the geographical distribution 
of the species belonging to that fauna. Many of 
them exhibit what is known as "discontinuous distri- 


bution," which zoologists are agreed to regard as a 
sign of antiquity. Thus Geomalacus maculosus, the 
Kerry Slug, is in the British Islands confined to South- 
western Ireland (see Fig. 19, p. 300), and on the Con- 
tinent it is unknown north of North-western Spain. 
The Millepede, Polydesmus gallicus, has a wider range 
in Ireland, and is also known from France and the 
Azores. Two Earthworms of the Spanish and 
Mediterranean region, viz., Allolobophora veneta and 
Georgii^ have been discovered in Ireland, but are 
apparently unknown in England or France ; whilst 
the Weevil, OtiorrJiyncJius auropunctatus, does not 
occur north of the Auvergne Mountains in France 
except in Ireland. A very large number of instances 
might be mentioned of species found in South- 
western Europe, France, the South-west of England 
and Ireland. Enough, however, has been said to 
show the nature of the fauna, and there is, as 
Forbes has pointed out, a corresponding flora. 

A great number of the species belonging to the 
South-western British element seem to have origin- 
ated in South-western Europe, or at any rate to have 
spread over our continent from that part. Their 
home lay therefore probably in a warm, damp 
climate, and it seems a reasonable inference to 
suppose that they spread north at a time when 
the temperature over the British Islands was much 
higher than what it is now. Any one familiar with 
our Bristle fern, or Killarney fern, as it is called in 
Ireland (Trichomanes radicans}, will readily admit that 


it must have come to us at such an epoch. It at once 
suggests some shady waterfall in a tropical forest, and 
indeed the home of the genus is South America. It 
is one of those plants which have evidently migrated 
to us from South-western Europe, 'a mere remnant 
of a once luxuriant flora. 

Sir Archibald Geikie tells us (p. 837), and in the 
main every one agrees with him, that at the beginning 
of the Tertiary era in which we now live, the climate 
was of a tropical and subtropical character in Europe. 
Gradually it became more temperate, and eventually 
it passed into a phase of extreme cold, but since that 
time the cold has again gradually diminished. It is 
quite evident, therefore, that from a purely geological 
point of view our south-western flora must have 
migrated northward before the cold came on, and 
survived in sheltered localities under the influence 
of the mild coast climate. Some, however, suppose 
that there occurred a phase of extreme mildness im- 
mediately after the Glacial period, and that it was 
during that time that the Lusitanian fauna and flora 
became established in the British Islands. To this 
Professor James Geikie replies (, p. 169), "there are few 
points we can be more sure of than this, that since 
the close of the Glacial epoch since the deposition of 
the clays with Arctic shells and the Saxicava sands 
there have been no great oscillations, but only a 
gradual amelioration of climate. It is quite impossible 
to believe that any warm period could have intervened 
between the last Arctic and the present temperate 


conditions without leaving some notable evidence in 

the superficial deposits of Scotland, Scandinavia, and 
North America." Thus it appears that on the whole 


the assumption that the Lusitanian fauna and flora 
are very ancient and pre-glacial is also supported 
on geological evidence. 

The course of events in the origin of the British 
fauna might have been therefore somewhat as 
follows : In early Tertiary times, when the climate 
all over Western Europe was moist and semi-tropical, 
a migration proceeded northward from the south- 
western corner of Europe. This was strengthened 
by Oriental migrants which had moved westward 
along the Mediterranean basin (Fig. 5, No. i). 
Owing to geographical changes supervening, the 
Alpine fauna (No. 2) was then enabled to colonise 
the British Islands, and subsequently another migra- 
tion had begun to come in from the south-east 
(No. 3). The climate had meanwhile gradually 
become more temperate and drier, About the same 
time, or even earlier, an Arctic migration commenced 
to pass southward (No. 4), and finally the Siberian 
animals (No. 5) poured into our continent. The 
arrows in the map indicate the directions followed by 
the different migrants as they travelled to the British 
Islands. The arrows are not meant to represent the 
whole nor the full extent of the migrations from 
any particular centre, but only in so far as they 
affect our islands. Moreover, it would be im- 
possible to indicate on one map the geographical 
conditions which obtained during the several migra- 
tions. It must be remembered that during the time 
which elapsed while they passed into the British 


Islands, these were joined in the north to Scandinavia 
and in the south to Belgium and France. The various 
phases of geographical evolution of Europe will be 
studied in the subsequent chapters, and maps will 
then be given to show as far as possible in a 
general way the leading characteristics of these 
great changes. 

I have now given some reasons for the belief that 
several different migrations of animals entered the 
British Islands in later Tertiary times. I have also 
shown why some of them must be looked upon as 
being older than others, and in so far we have come 
to a decision as to their relative ages. It still remains 
for us, however, to examine how their geological ages 
can be approximately determined. We require for 
this purpose palasontological aid. 

In the fifth chapter will be found the history of 
the Siberian migration. And since we possess most 
valuable records of it in the numerous fossil remains 
discovered in Central and Western Europe, we are 
able to trace their progress from the east to the 
west in a very complete and satisfactory manner. In 
England their first appearance dates from the Forest- 
Bed, for here we find remains of the Glutton (Gulo 
luscus\ Musk-Ox (Ovibos moschatus\ and others (see 
p. 204). It seems reasonable to suppose, therefore, 
that the first entry of these Siberian mammals into 
Europe took place at or just before the Forest-Bed 
period. But Professor Nehring tells us in his remark- 
able work on the Tundra and Steppes (p. 222), that in 


Germany the remains of the same mammals occur in 
deposits which are certainly more recent than the 
lower continental boulder clay; and he is inclined to 
the belief that they migrated into Europe during the 
inter-glacial phase which is supposed to have separated 
the earlier from the later stage of the Glacial period. 
It is evident that in this case the inter-glacial period 
in Germany would have corresponded to, and be con- 
temporaneous with, our Forest-Bed period. The 
deposits immediately preceding the Forest-Bed would 
also be contemporaneous with the lower continental 
boulder clay. Although this may seem rather a 
startling statement to make, from the evidence 
which will be brought forward in the fourth and 
fifth chapters I am inclined to the belief that such 
is probably the case. 

Having once arrived at a determination of the 
exact geological period during which the Siberian 
mammals invaded our continent, and having also 
previously determined the relative ages of the various 
other migrations, we have advanced another step in 
the direction we are aiming at Let us suppose that 
the Siberian migration actually reached the British 
Islands during the Forest-Bed period. Since the 
Siberian migration is the most recent of those which 
entered the British Islands, the others must have 
commenced their march before the Forest-Bed 
period. Now it was Professor Boyd Dawkins who 
first indicated to us, as I have remarked before, 
the method of research to be adopted in an attempt 


to determine the geological age of the different 
migrations in so far as they affected the British 
Islands. I may be excused, therefore, for again 
quoting the following important passage in one of 
his works. " The absence," he says (, p. xxix), " of 
the beaver and the dormouse from Ireland must be 
due to the existence of some barrier to their westward 
migration from the adjacent mainland, and the fact 
that the Alpine hare is indigenous, while the common 
hare is absent, implies that, so far as relates to the 
former animal, the barrier did not exist." The Beaver, 
Dormouse, and Common Hare are either Siberians or 
later migrants from elsewhere, and there can be no 
doubt that at the Forest-Bed period Ireland was 
already, or was just being, separated from England. 
All the southern species, that is to say all the 
Lusitanian, Alpine, and Oriental forms occurring in 
Ireland, must therefore be older than that period. I 
have advocated similar views in a former essay on 
this subject. Mr. Carpenter recently advanced some 
interesting and valuable criticisms on these views, 
which we may examine a little more closely (p. 385). 
" While, then," he remarks, " I find myself in almost 
complete agreement with Dr. Scharff with regard to 
the older sections of our fauna, I think that those 
widespread species which survived the Glacial period 
must have been confined to the more southern parts 
of our area, and have only subsequently spread 
northwards and westwards to Scotland and Ireland." 
He suggests, in fact, that the widespread British 


species belong to a younger or newer section of our 
fauna than the local ones. In many cases this may 
be quite true, but we possess also a large number of 
common and widely-spread forms which bear the 
impress of antiquity upon them. We have the most 
positive proof of the antiquity of the very common 
small circular Snail (Helix: rotundata}, since it was 
found in miocene freshwater deposits near Bor- 
deaux. Many other examples might be mentioned 
to show that, though discontinuous range is generally 
a proof of antiquity, continuous range is not always a 
sign of the opposite. Some species, in fact, appear to 
be short-lived and disinclined to spread, whilst others 
multiply rapidly even under a change of temperature 
and climate, and are to be found almost everywhere. 
But even if we supposed, with Mr. Carpenter, that 
these widely-ranging species must have been confined 
during the Glacial period to the more southern parts 
of England, the idea that they afterwards made their 
way northwards along the eastern shore of the Irish 
Sea and then passed into Ireland, does not appeal to 
me. Southern England was occupied at that very 
same time by an assemblage of Siberian mammals. 
Mr. Carpenter thinks these might have been kept 
out of Ireland by an arm of the sea until the land- 
connection with North-western England had broken 
down. But if an arm of the sea could keep out the 
Siberian mammals it would also keep out the widely- 
spread British species of the general fauna. On the 
other hand, I quite admit that my view of the survival 


in Ireland of the pre-glacial fauna is somewhat 
difficult to accept, considering that we have such 
undoubted evidence of a very extensive submergence. 
The case of Isle of Man, quoted by Mr. Carpenter, 
can be met, I think, by the supposition that it was 
connected with Cumberland until quite recently, and 
quite independently of any connection between Eng- 
land and Ireland; that the Isle of Man, in fact, 
was always a cape or peninsula of the mainland, and 
only recently became separated by local subsidences 
or by the action of the sea. 

Part of the history of the British fauna will be 
referred to again in the next chapter, which deals 
with the Arctic migration. We need not therefore 
dwell any longer on this subject here. There is one 
matter, however, which is of importance in connection 
with the geographical conditions of the British Islands 
at the time when the greater portion of our fauna 
arrived from abroad. 

On page 60 will be found a map indicating the 
physical geography of that part of the ancient con- 
tinent on which what are now the British Islands were 
situated. Only one large river has been marked on 
that map, namely, that flowing out of a lake which 
occupied part of the Irish Sea. Another probably dis- 
charged its waters into the Atlantic midway between 
France and England, whilst the Thames may have 
been a tributary of the Rhine, as it emptied itself 
into the sea near our south-east coast. I have shown 
in a previous essay that the former presence of a fresh- 


water lake between England and Ireland is indicated 
by the distribution of the Charrs and also by the 
various species of British Coregonus. There are 
three British species of Coregonus, viz., C. clupeoides, 
C. vandesius, and C. pollan. These are confined to 
the lakes of North Wales, North-western England, 
South-western Scotland, and Ireland. All but the 
latter communicate at present directly with the Irish 
Sea. The lakes of the latter country, however, must 
have done so at a time when the west of Ireland 
stood at a higher level than it does now. The 
ancestors of the three Coregonus species, and also 
those of the Charrs, then lived in the large freshwater 
lake indicated on the map (p. 60), and when the sea 
gradually crept up the river valley and finally con- 
verted the lake into a gulf, the freshwater fish took 
refuge^ in the rivers which supplied it with water. 
^NQW as for the continuous sea-shore between the 
coast of Brittany and the south-west of Ireland, 
zoological distribution again aids us in proving that 
such must have actually existed at no very distant 
geological date. Most of our common shore forms 
of life migrate along the coast exactly as land 
animals do step by step. Their eggs are care- 
fully attached to fixed objects, so as not to be 
carried away by the waves, whilst the young often 
remain and grow old in some particular little pool, 
rarely venturing farther than a few yards from the spot 
where they first saw the light of day. A number of 
such shore forms are found on the west coast of France, 


the same species recurring again on the south-west 
coasts of England and Ireland, thus clearly indicating 
a former continuity of coast-line between these points, 
now separated by deep sea. A very familiar example 
to British zoologists is the purple rock-boring Sea- 
urchin {Strongylocentrotus lividus), but there are a 
great many others, such as the semi-marine Beetles 
Octhebiiis Lejolisii and dEpophilus Bonnairei^ the 
Crustaceans Achceus Cranchii, Inachus leptochirus, 
Gonoplax angulata, T/iia assidua, Callianassa sub- 
terranea y the Fishes Blennius galerita and Lepado- 
gaster Decandollii^ and the Molluscs Otina otis, 
Donax politus, and Amphidesma castaneum. 

Before concluding this chapter, a few words as to 
my views on the conditions prevailing during the 
Glacial period will not be out of place. They do not 
differ very much from those held formerly by most 
geologists; and even at present there are, as I have 
mentioned before, a few upholders of those older 

The sea, I think, must have gradually crept across 
England from the east during, or shortly after, the 
Forest-Bed period, so as to separate the south from 
the north, whilst Ireland and Scotland were then still 
connected with one another. At a later stage, the sea 
also partially invaded Ireland, and this condition is 
very roughly represented on the accompanying map. 
Mr. Kendall kindly drew my attention to the fact 
that several notable areas on which shelly drift has 
been observed are here placed upon the land; but 


it must be remembered that one stage only can be 
shown on the map, and that the sea covered more 
ground a little later. Many of the smaller islands in 
the glacial sea, too, are not shown. The map, in fact, 

FlG. 6. Map of the British Islands, showing approximately in what 
manner the sea may have invaded the country from the east during, 
or shortly after, the Forest-Bed period. The darkly shaded 
parts indicate the areas covered by water, and the lightly shaded 
and white portions what was land at that time. 


is merely meant to give a general idea of the manner 
in which the great northern sea moved westward and 
slowly covered a large portion of the British Islands. 
These peculiar geographical conditions explain, I 
think, better than anything, the absence from parts 
of the Midlands and the north of England of 
such a number of terrestrial invertebrates which 
are otherwise widely distributed over the British 
Islands. In spite of the fact that a large portion 
of the British Islands became submerged, we 
possessed at that time an extensive area which 
has since been claimed by the sea, so that there 
was ample room for the present fauna to survive 
the Glacial period. The climate during this period 
was probably much the same as it is at present, 
though moister, with cooler summers and milder 

It may be asked what proof we have of such an 
extensive submergence of England and Ireland. My 
own views are principally based on the general 
distribution of the fauna in the British Islands, and 
the belief that nothing but a mild climate during 
the Glacial period could have brought it about. On 
purely geological grounds, however, some geologists, 
notably Mr. Mellard Reade, have come to a similar 
conclusion. " The whole of Lancashire and Cheshire," 
he remarks (a, p. 542), " from sea-level up to about 
400 feet, and in places 600 feet, is covered by a 
continuous mantle of boulder-clay and sands." 
" These clays, as a rule, contain distributed through 


them, in a greater or less degree, fragments of shells 
and some perfect ones. I myself have recorded 
forty-four species." Again he continues (pp. 545 
and 546) : " A large part of Ayrshire is covered with 
similar shelly boulder-clays from sea-level to 1061 
feet at Dippal. These Ayrshire high-level shells 
have, in the majority of cases, been taken, not from 
sand and gravel beds, but from boulder-clay, and in 
that respect they are most important and unique. 
In Moel Try fan the shells are found in sands and 
gravels at 982 feet; on the range of hills from Miaera 
to Llangollen from 1000-1200 feet; also in sands and 
gravels at Gloppa, near Oswestry, at 1100-1200 feet; 
and near Macclesfield at a level of about 1200 feet. 
In Ireland marine shells can be traced almost from 
sea-level to a height of over 1000 feet." 

"Again," continues the same author, "if we look 
broadly at the distribution of these shelly deposits, 
we find that they occur all round our maritime coasts 
in Lancashire, Cheshire, and Wales, in Cumberland 
and Westmoreland, Wigtonshire and Ayrshire, and 
along the eastern coast of Ireland. The same is 
to be said of the eastern coasts of England and 

That a very considerable change of sea-level has 
taken place in some parts of the British Islands would 
appear to a zoologist the most logical conclusion after 
an examination of these " high-level shelly sands and 
gravels," but the shells contained in them are now 
generally supposed to have been carried there frozen 


in the sole of a glacier or pushed up in front of it 
The older view, however, which agrees so much better 
with the facts of distribution, fortunately has not 
disappeared among geologists. "When we call up," 
says Mr. Mellard Reade (3, p. 435), "before our 
mental vision the simple and well-known facts of 
nature which suffice to explain the marine drifts on 
the theory of submergence, it seems unnecessary 
to resort to the ingenious and artificial system of 
physics elaborated to explain the phenomena of 

" When we have more knowledge of the glaciers 
of the Arctic Regions, and facts, in place of 
ingenious suppositions, to base our reasoning upon, 
we may possibly have to revise all our glacial con- 
ceptions. In the meantime, the submergence theory 
of the origin of high-level shelly gravels and sands 
seems to me by far the simpler of the two theories, 
and the most consistent with the facts and phenomena 
which the labours of a succession of enthusiastic geo- 
logists have made us acquainted with." 

Among those geologists, and they form the majority, 
who hold that Ireland was covered by land-ice, there is 
a great diversity of opinion as to its extent. Messrs. 
Close, Kinahan, J. Geikie, and others believe that 
the ice covered practically everything, whilst others 
who claim to have examined the ground with equal 
care, such as Professor Carvill Lewis, were led to 
believe that the south of Ireland, with the exception 
of a few local glaciers, was free from ice. The glacial 



phenomena of the country can therefore be inter- 
preted in different ways, even by those who are 
convinced that they are due to land-ice and not to 
icebergs or mud-glaciers. 


The history of the British fauna is not only of interest to us 
from a sentimental point of view, it is a convenient starting- 
point in the study of the larger European problem. The fauna, 
broadly speaking, is composed of three foreign elements, viz., the 
northern, eastern, and southern, to which may be added a small 
endemic one. Examples are given of the more noteworthy forms 
belonging to each of these. This leads us to the subject of the 
natural divisions of the British Islands according to their animal 
inhabitants. Zoologists attempted at first to subdivide these 
countries, on the lines laid down by botanists, into a large 
number of provinces. Forbes proposed ten such divisions for 
mollusca, and subsequently five, which were ultimately reduced 
by others to two or three. 

The opinions of biologists are almost unanimous in attributing 
the bulk of the British fauna and flora to migrations by land 
from the Continent, but two other theories, viz., those of Pro- 
fessor Cole and Messrs. Kinahan and Lamplugh, are also 
referred to. The first believes in a possible migration eastward 
from Western Europe, and the latter support the view of the 
former existence of ice-bridges to assist the fauna in their 

An endeavour is next made to determine at what geological 
periods the various migrations entered the British Islands. 
There is considerable difference of opinion on this subject. 
Some believe that the British fauna is altogether post-glacial ; 
a few think that it is partly so and the remainder glacial ; 


others again hold that a portion is pre-glacial and the rest 
glacial and post-glacial. Those who have studied the subject 
most closely feel convinced that the south-western or Lusitanian 
fauna, and also the flora, must have arrived before the Glacial 
period and survived the latter in these Islands. It seems 
reasonable to suppose, therefore, that the climate cannot 
have been very severe during the so-called Ice-Age. This 
Lusitanian fauna must be looked upon as the oldest portion of the 
British fauna. The Alpine and Oriental migrations arrived next. 
After these came the Arctic, and finally the Eastern or Siberian. 
As the fossil evidence is most complete with regard to the last, 
we are able to determine with precision not only the direc- 
tion whence this migration came, but approximately its geo- 
logical age. It arrived in Germany from the east after the 
deposition of the lower boulder-clay. Since the boulder-clay 
is looked upon as a glacial deposit, the Siberian migration 
reached Central Europe after the first portion of the Glacial 
period had passed. In England it makes its first appearance 
in the Forest-Bed, which would therefore correspond to the 
" Loess " formation of Central Europe. All the other migrations 
are older than the Siberian. They must therefore have come 
to Great Britain during the earlier part of the Glacial period 
or before it. 

The chapter concludes with a short statement on the physical 
geography of the British Islands during the time when these 
migrations entered them. That there existed a continuous 
coast-line between France and Ireland is proved by the 
occurrence of a considerable number of identical shore 
species, whilst the former existence of a freshwater lake on 
the site of the present Irish Sea is indicated by the dis- 
tribution of some freshwater fishes. 



THE lands lying within the Polar Circle are inhabited 
by an assemblage of animals and plants, many of 
which are peculiar to those regions. They are mostly 
adapted to the abnormal conditions of life prevailing 
in the high latitudes of our globe the long, dark 
winters, and the short summers of one long day. 
Though the numbers of species and of individuals arc 
few, there is a keen struggle for existence in those 
regions. The prevailing colour of the ground is white, 
and since a resemblance in the colour of an animal 
to the ground it lives on acts as a protection to weak 
ones, and also enables Carnivores to approach their 
prey with greater facility, it is not surprising that we 
should find the majority of polar animals coloured 
white. As I remarked, the polar area contains a very 
distinct set of species; most of them, however, range 
beyond the confines of the Arctic Circle. It is there- 
fore scarcely justifiable to raise this Arctic area into a 
distinct zoological region equivalent to the great zoo- 
geographic regions, which have been established by 
Sclater and Wallace, though we might, with Dr. 
Brauer, look upon it as a sub-region. 



There are six typical Polar Land-mammals, one of 
which, the Polar Bear, is semi-aquatic. The Reindeer 
(Rangifer tarandus) occurs upon almost all the polar 
lands, and it has often been a source of speculation 
in what manner it has reached such remote islands as 
Spitsbergen and Novaya Zemlya the former of the 
two being so remote from a continent. There is no 
doubt that Reindeer are great wanderers, owing to 
the difficulty of rinding sufficient food-supply for the 
large herds in which they are accustomed to travel ; 
and for this reason they can cross, and have been 
known to cross, distances of from ten to twenty miles 
on ice. The Behring Straits, when frozen over in 
winter, is frequently traversed by them. But I quite 
agree with Dr. Brauer (p. 260) that it is impossible to 
account for their presence in Spitsbergen by an im- 
migration from either Novaya Zemlya, Greenland, or 
Scandinavia, under the present geographical con- 
ditions. The seas between the former island and 
the other land-masses referred to are rarely entirely 
frozen over. Even if this should occur, the dis- 
tances between Spitsbergen and Greenland, Novaya 
Zemlya, or Scandinavia are so great, that a migration 
across ice is quite excluded from the range of pos- 
sibilities, since Reindeer could not subsist without 
food during the time it would take to travel from 
one to the other. The manner in which it did reach 
Spitsbergen and Greenland will be discussed more 
fully below, and I will therefore proceed to mention 
the other Arctic mammals. 


One of the most important and most typical species 
is the Polar Bear (Ursus maritimus\ the greater part 
of whose life is spent on the ice and in the sea. The 
fact that its favourite nourishment consists of seals 
proves its excellent and keen faculties of sight and 
hearing, and its facility in swimming. But it is not a 

FIG. 7. The Musk-Ox ( Ovibos moschatus). (From Flower & Lydekker's 
Mammals , p. 358. London: Adam & Chas. Black.) 

dainty feeder, and lives upon almost all animals which 
come within its reach ; birds, land-mammals, or fish 
are not despised in times of scarcity. Its fur through- 
out the year is coloured white, though in old bears it 
assumes a more yellowish hue. 

Another large mammal, perhaps less well known, 
is the Musk-Ox {Ovibos moschatus, Fig. 7), which 


resembles in size the smaller varieties of Oxen, but in 
structure and habits is closely allied to the Sheep. As 
is implied by the specific name, it exhales a musky 
odour ; this does not, however, appear to be due to 
the secretion of a special gland, as is the case in other 
animals with a similar smell. The skin is covered 
with long brown thickly-matted hair, interspersed 
with white. It is confined to the most northerly 
parts of North America and the American Arctic 
islands, and to North Greenland. Though not now 
living in the Old World, it seems formerly to have 
been abundant in Siberia, and, as we shall learn later 
on, it was one of the species which took part in the 
great Siberian invasion of Europe. Its remains have 
been found not only in Germany and France, but also 
in the south of England. 

The Polar Fox (Cants lagopus] occurs throughout 
the Polar Regions, and on islands where even the 
Reindeer and the Musk-Ox are unknown. Beyond 
the Polar Circle, its range extends into Northern 
Asia, to the extreme north of North America, and the 
mountains of Scandinavia. Like its congeners, it 
had in pleistocene times a more southerly extension, 
and fossil remains have been met with in various 
parts of continental Europe and in England. 

The Stoat (Mttstela erminea\ which is known and 
much valued in commerce under the name of Ermine, 
was formerly believed to occur only in Arctic America 
and the northern parts of the Old World, but in more 
recent years it has been discovered in a number of 


the northern islands, such as Saghalien, in the islands 
of the Behring Straits, the Aleutian islands, and also 
in Greenland and Spitsbergen. In Europe, it is found 
as far south as the Arctic Hare, or perhaps even 
farther, and it flourishes in the Alps up to a height 
of 9000 feet. It offers a parallel to the Arctic Hare 
in the fact that in some countries, such as Ireland, it 
only rarely turns white in winter. The Irish form of 
the Stoat differs so much from the English, that 
Messrs. Thomas and Barrett-Hamilton are of opinion 
that it is specifically distinct, as I mentioned in 
speaking of the divisions of the British fauna 
(p. 90). 

The Arctic Hare (Lepus variabilis] is almost the 
only one of the typical Arctic mammals which still 
inhabits the British Islands, and for that reason it is 
to most of us more familiar than any of the preceding 
species. Hares have been described from Green- 
land by the name of Lepus glacialis, from the 
European Alps as Lepus alpinus, and under other 
names from Arctic North America; but though slight 
differences in the fur and even in the skull can be 
pointed out, there is no doubt that all these are only 
varieties or races of what, in the British Islands, is 
known as the Irish or the Scotch Mountain Hare, 
Lepus variabilis. In the Arctic Regions this Hare 
remains white throughout the year, but in Scan- 
dinavia and some other parts its fur becomes brown 
in the summer, and in Ireland it frequently remains 
entirely brown during the whole year, and never, or 



only in very rare cases, becomes entirely white in 
winter. Besides Scandinavia, Scotland, and Ireland, 
it is found in Northern Russia, and also in the 
Pyrenees, the Alps, and the Caucasus. In Asia it 
occurs not only on the mainland of Siberia, but it has 

FIG. 8. Map of the northern hemisphere, to show the geographical 
distribution of the Arctic Hare (Leptis variabilis] indicated in 

been obtained on the Akita Mountains in Japan and 
on the Mioko San Mountain, and also on the island 
of Saghalien. It had in former times a more exten- 
sive range, and its remains have been discovered in 


England and in a number of places on the continent 
of Europe. The peculiarity of its range, which will 
be explained more fully directly, lies in the fact of 
the occurrence of isolated colonies in the mountains 
of Europe, in Ireland and Scotland, and in the 
mountains of Japan (Fig. 8). From a distributional 
point of view, it is one of the most interesting species 
of mammals, and its history throws a flood of light 
on the geographical changes which have occurred in 
former times. 

One more species must be mentioned, and that is 
the Banded Lemming {Cuniculus torquatus}, which 
occurs chiefly in Arctic America, Northern Siberia, 
and Greenland. Though frequently mistaken for the 
Scandinavian Lemming, there is a striking difference 
in the character of the teeth, which has induced 
zoologists to put them into distinct genera. The 
Arctic Lemming, moreover, is distinguished from 
the Scandinavian by the absence of external ears, the 
densely furred feet, and by the great length of the two 
middle claws in the fore-feet. There are two species 
of the true Lemming, namely, the one just referred to, 
Myodus leimnuS) and Myodus obensis. These may be 
looked upon as more or less Arctic species, since they 
occur within the Polar Circle, but they are not so 
exclusively confined to that region as the Banded 
Lemming {Cuniculus torquatus}. The remains of 
both Cuniculus torquatus and of Myodus leinmus have 
been found in British pleistocene deposits. 

Until recently no Lemming remains had been found 


to the south of France, but Mr. Barrett-Hamilton 
announced to us a short time since that Dr. Gadow 
had discovered some skeletons with their skins still 
preserved in a cave in Northern Portugal. These 
were found to belong to the Scandinavian Lemming 
(M. lemmus) y and the author incidentally expressed the 
opinion that there was some possibility of this species 
still inhabiting the mountains of Spain. 

The Lemming multiplies with great rapidity under 
favourable conditions. In speaking of his experiences 
in Siberia Dr. Brehm says (p. 79): "All the young of 
the first litter of the various Lemming females thrive, 
and six weeks later at the most these also multiply. 
Meanwhile the parents have brought forth a second 
and a third litter, and these in their turn bring forth 
young. Within three months the heights and low 
grounds of the tundra teem with lemmings, just as 
our fields do with mice under similar circumstances. 
Whichever way we turn we see the busy little crea- 
tures, dozens at a single glance, thousands in the course 
of an hour. But the countless and still increasing 
numbers prove their own destruction. Soon the lean 
tundra ceases to afford employment enough for their 
greedy teeth. Famine threatens, perhaps actually sets 
in. The anxious animals crowd together and begin 
their march, hundreds join with hundreds, thousands 
with other thousands, the troops become swarms, the 
swarms armies. They travel in a definite direction, 
at first following old tracks, but soon striking out 
new ones; in unending files defying all computation 


they hasten onwards; over the cliffs they plunge 
into the water. Thousands fall victims to want and 
hunger; the army behind streams on over their 
corpses; hundreds of thousands are drowned in the 
water or are shattered at the foot of the cliffs; the 
remainder speed on; other hundreds and thousands 
fall victims to the voracity of Arctic and red foxes, 
wolves and gluttons, rough-legged buzzards and 
ravens, owls and skuas which have followed them; 
the survivors pay no heed. Where these go, how they 
end, none can say; but certain it is, that the tundra 
behind them is as if dead, that a number of years 
pass ere the few who have remained behind and have 
managed to survive slowly multiply and visibly re- 
people their native fields." This eloquent passage 
reminds us of the manner in which migrations of 
all kinds of animals have taken place in former times, 
and are still taking place. It is principally want 
of food which compels them to search for new 

On page 91 I have referred to some birds which 
have come to us from the north. One of these, the 
Snow Bunting (Plectrophenax nivalis], is a typically 
Arctic species. In summer it is widely distributed, 
and is found in Spitsbergen, Novaya Zemlya, Siberia, 
and the Arctic Regions generally. In winter it 
migrates down into North America, into Japan, 
Northern China, Turkestan, Southern Russia, and 
occasionally even across Europe into North Africa. 
Very characteristic Arctic birds are the Eider Ducks 



belonging to the genus Somateria. Three species 
have visited the British Islands. The common Eider 
Duck (S. mollissima}, which is of such high com- 
mercial value, is abundant in Norway and northward, 

FIG. 9. The Great Auk (Aka imfcniris). 

throughout the Polar Regions. The appearance of 
the King Eider (S. spectabilis) on our coasts is an 
extremely rare occurrence, and even in Norway it is 
only known as a visitor, but on Novaya Zemlya and 
along the Arctic shores of Siberia, in Greenland and 


Arctic North America, it is known to breed. The 
third species, Steller's Eider (S. Stelleri), seems to be 
still rarer, and only in the Aleutian islands and in the 
north of Alaska can it be said to be at all abundant. 
It is probable that the famous Great Auk (Alca 
impennis, Fig. 9) also was a typical Arctic species. 
Its range extended to both sides of the Atlantic. In 
Newfoundland and on the coast of Iceland it is known 
to have been met with in considerable numbers 
within historic times; and no doubt, like all Arctic 
species, it extended farther southwards at a more 
remote period. 

The members of the genus Lagopus, including the 
various species of Grouse, are likewise of northern 
origin. The British Red Grouse (L. scoticiis), which 
may be looked upon as a form of the Scandinavian 
Willow Grouse (L. albus) (compare p 91), constitutes 
in some respects a curious case of parallelism with 
the Arctic Hare, since the latter, in its more southern 
station, generally retains the summer fur throughout 
the year. The allied Ptarmigan (L. mutus] inhabits 
Scandinavia, the Ural Mountains, and some of the 
Asiatic mountain ranges. It is also found in the 
European Alps and in the Pyrenees. The North 
European range of the Ptarmigan suggests that 
we are dealing with an ancient species which came 
south from the Arctic Regions at about the same 
time as the Arctic Hare; but it is more probable, 
as I have shown in a subsequent chapter (p. 334), 
that this species has entered Europe more recently 


with the Siberian migrants from Central Asia, where 
indeed the genus had its original home. The Black 
Cock (Tetrao tetrix) and the Capercaillie (Tetrao 
urogallus) have also come to us from the east, 
and have even penetrated into Ireland. They are 
therefore some of the few instances of members 
of the Siberian invasion having become temporarily 
established there. 

Reptiles and amphibia are altogether unknown in 
the Polar Regions, but a large number of fish, chiefly 
marine, have taken their origin there. The Salmon 
family are of Arctic origin, as also are the Stickle- 
backs and the Perches, many of the Cod family, the 
Herrings, and several of the Flat fish. 

It would lead me too far to refer to the invertebrate 
fauna of the Polar Regions, but a few remarks on the 
Arctic plants may not be out of place. 

The principal Arctic genera are Satix, Ranunculus, 
Draba, Pedicularis, Potentilla, Saxifraga, Carex, 
Juncus, Luzula, Eriophorum, and others. 

Among the most characteristic Arctic plants may 
be mentioned Dry as octopetala, to which I have already 
referred as occurring in the west of Ireland; Saxifraga 
oppositifolia, another British species, occurs in the 
higher mountains of Scotland, Ireland, and Wales; 
Braya alpina, Papaver nudicaule. Lychnis apetala, 
Diapensia lapponica, and Lobelia Dortmanna, which is 
found in the lakes of Scotland and Ireland. The 
dwarf birch (Betula nand) also, which still occurs 
in Scotland and the North of England, and which 


had formerly a wider range in the British Islands, 
should be included among these; but there are other 
plants probably of Arctic origin, though not now 
occurring in the Arctic Regions, and to these may 
be classed the so-called American species of plants 
which are found on the northern and western coasts 
of Ireland, in the Hebrides, in Scotland, and in 
North America. These are no doubt the relics of an 
Arctic flora which flourished in high latitudes in past 
times when the climate there was more temperate 
A list of these species will be found on page 166. 

As none of them occur in Siberia, they must 
either have found their way to North America and 
to Europe from the Arctic Regions, or have travelled 
from North America across the latter to Europe. 
In any case a former land-connection between the 
two continents must have existed. This becomes 
the more evident when we examine the remarkable 
results obtained by the late Professor Heer, who first 
described the Tertiary plant-beds in North Greenland. 
No less than 282 species of plants have been described 
by this eminent botanist from these deposits. A large 
number of the plants found were trees belonging to 
the genus Sequoia^ Thujopsis^ and Salisburia, besides 
beeches, oaks, planes, poplars, limes, and magnolias. 
That they grew on the spot is proved by the fruits, 
which have been obtained from these beds in various 
stages of growth. 

From a similar deposit in Spitsbergen a large 
number of fossil plants have also been brought to 


light, many of which are identical with those found 
in Greenland ; and some of the Greenland forms 
(such as Taxodium distichum and Sequoia Langsdorfii) 
have been found too in Alaska, showing that there 
was probably a continuity of land between Spits- 
bergen and North America by way of Greenland. 
Two species of Sequoias, namely, S. sempervirens 
and 5. gigantea, the well-known Californian giant 
trees, are very closely allied to the Greenland forms 
discovered by Professor Heer. 

Heer assigned the Arctic plant-bearing beds to the 
Miocene epoch, but doubts have been recently thrown 
upon this opinion by Mr. Starkie Gardner, who 
brought forward arguments in support of his theory 
of their being of the Eocene age. Professor Heer, 
however, was able to meet these criticisms, and he is 
ably supported in his views by Professor Engler and 
other eminent continental botanists. 

It is evident that under the present conditions of 
temperature none of those plants could have flourished 
in Greenland. The climate must have been much 
milder than it is at present. Professor Heer estimated 
from the general aspect of the fossil flora that the 
mean annual temperature of North Greenland was 
at least nine degrees centigrade, and that the mean 
winter temperature was not below zero. 

It will hardly be necessary for me to review here 
the various theories which have been advanced by 
geologists and botanists to account for this remark- 
ably high temperature in such northern latitudes 



Any one who has read the writings of the late Dr. 
Croll cannot help being struck by the facts he adduces 
to show the importance of ocean currents in relation 
to the distribution of heat over the globe, and it seems 
to me that the view which attributes the mild climate 
prevailing in former times in Greenland to warm 
ocean currents reaching the Polar Circle is the 
one least open to serious objections. If we suppose 
that the North Atlantic Ocean was bridged by a 
land-connection between Scandinavia and Greenland 
by way of Spitsbergen, and between Greenland and 
North America, the Polar Ocean would be practically 
a closed sea. If, then, a wide passage existed some- 
where about Behring Straits to allow a warm current 
to enter and circulate within the Arctic Seas, we 
should have the southern shores of Greenland washed 
by the warm Atlantic current and the northern shores 
by a warm Pacific current, which combination would 
undoubtedly produce the effect of raising the tem- 
perature throughout the Polar Regions very con- 
siderably; and especially would that be the case 
with regard to Greenland and the neighbouring 

It might be urged that the constant darkness 
during winter must have had an injurious action 
upon the flora, but it is found that in countries 
such as Northern Russia, where southern plants are 
housed during winter in greenhouses, the light being 
almost entirely excluded by a covering of straw, no 
serious damage is done thereby to the plants. 


It seems probable that a similar gradual refrigera- 
tion 'of climate in northern latitudes has taken place 
after Miocene times as has been proved to have 
occurred in Europe. 

Some years ago Dr. Haacke propounded the hypo- 
thesis that the centre of creation of all the larger 
groups of animals was situated in the region of the 
North Pole, and that the newly originated groups must 
always push the older ones farther and farther south 
into the most remote corners of the earth. As 
instances of the correctness of his view he quotes 
the fact that the more ancient mammals, such as 
Monotremes, Marsupials, Lemurs, Edentates, and 
Insectivores, all inhabit the more southerly parts 
of the world. The Apteryx, Moa, Rhea, and the 
Ostrich, as well as ^Epyornis, which is only recently 
extinct, are found in the same regions. But we have 
no palseontological evidence in favour of these ex- 
travagant views. Fossil Edentates and Marsupials 
are almost entirely confined to the Southern Hemi- 
sphere, and the supposition that because these 
primitive mammals inhabit the extreme south of our 
great continental land-masses, they therefore came 
from the north, cannot be said to be an argument. 
Nevertheless, I am quite with Dr. Haacke in consider- 
ing that the North Pole, or, we might say, the lands 
within the Arctic Circle, have been the place of 
origin of some of our European mammals, and there 
can be no doubt that certain species in other groups, 
among invertebrates and also plants, have originated 


in the Polar Regions. The facts of geographical 
distribution teach us that in these regions there has 
been a centre of origin within comparatively recent 
geological times. I have on a previous occasion 
drawn attention to the range of the Reindeer: that 
it lives almost throughout the Polar lands, and that it 
spreads into North America, Northern Europe, and 
Northern Asia. We have, again, fossil proof that its 
range extended down to the Pyrenees in Europe in 
pleistocene times. But there is not a scrap of 
evidence that it ever during any time occurred 
farther south, either in Europe, Asia, or North 
America. Its original home must therefore have 
been in the Polar Regions, for if it had originated 
either in Central Europe, Asia, or America, there is no 
reason why it should not, in the natural course of 
events, have extended its range to the south as well 
as to the north. 

The Arctic Hare presents us with a very similar 
case of distribution. Like the Reindeer, it inhabits, as 
we have learned, the Polar Regions and the northerly 
parts of the Old World and the New; but while we 
have only fossil evidence of the former, more southerly, 
extension of the range of the Reindeer, the Arctic Hare 
furnishes us with a still stronger proof of its past 
southward range in the survival of small isolated 
colonies in some of the southern mountain ranges of 
Europe and Asia. It is generally believed that the 
occurrence of the Arctic Hare in these southern moun- 
tains is a standing testimony to the severity of the 


climate at the time when it commenced its southerly 
increase of range, but I have already shown that the 
climate of Europe at that time was not necessarily 
colder than it is at present, but that it may have been 
somewhat milder (p. 80). I think that a vast increase 
of ice in the Polar Regions has taken place only at. a 
comparatively recent date, and that both the Reindeer 
and the Arctic Hare originated there during a much 
more temperate climate than obtains at present. 
A great sensation was produced among European 
zoologists and anthropologists when the discovery 
was first announced that the remains of the Reindeer 
had been found in the Pyrenees, and it naturally gave 
rise to many speculations as to the nature of the 
climate at the time when its range extended so far 
south. 1 The greater number of our best authorities 
are still of opinion that the existence of the Reindeer 
in Southern Europe points to the prevalence of an 
arctic climate in that region. It is generally over- 
looked, however, that the Reindeer-remains occur in 
company with many typically southern animals, which, 

1 A very interesting piece of information has been given us, recently, 
by Mr. Barrett- Hamilton on the Arctic Fox of Spitsbergen. In com- 
paring the skulls of Spitsbergen Foxes with those of Europe, he found 
that the former are much smaller, and represent a distinct race or sub- 
species. This small race he believes to be confined to Greenland, 
Iceland, Spitsbergen, and Novaya Zemlya, whilst the larger one occurs 
in Europe, Asia, and on the Commander Islands. This fact favours 
the view which I have advocated in Chapter V., that the Arctic Fox in 
Europe is a Siberian migrant, and did not come from the north with 
the Reindeer and Arctic Hare. 


if they had been found alone, would have been held 
to be a certain indication of a warm climate. The 
French geologist Professor Lartet, indeed, was of 
opinion that the temperature during the time when 
the Reindeer lived in the Pyrenees must have been 
rather milder than it is at present (compare pp. 71-75). 
Similarly, Mr. Harle argues, that the extremely 
cold climate probably did not extend to South- 
western France, since that area only received occa- 
sional visits from some of the representatives of the 
Arctic fauna. 

Long ago North American zoologists recognised 
the existence in their country of two well-marked 
races of the Reindeer (Caribou) a smaller one with 
rounded antlers (Fig. 10), and a larger one in which 
the antlers are more or less flattened out (Fig. n). 
Two somewhat similar races can also be traced in the 
fossil remains of the Reindeer in Europe. It was, I 
think, Gervais who first pointed out that the Reindeer 
remains from the north of France differed from those 
found in the south; and Lartet referred to the fact 
that the southern remains were more like what,, in 
America, is called the Barren-ground. Caribou, while 
those from Central European deposits all belonged to 
the Siberian variety, which is more like the Wood- 
land Caribou ot North America. In Ireland, Pro- 
fessor Leith Adams also drew attention to the 
curious fact that all the Irish Reindeer remains 
resemble the Norwegian variety rather than the 
Siberian; and Mr. Murray was so much struck by 


FlG. 10 Head of a Barren-ground Reindeer in the Dublin Museum 
(photographed by Mr. McGoogan). 


FIG. II. Head of a Woodland Reindeer in the Dublin Museum 
(photographed by Mr. McGoogan). 


the close resemblance between the Spitsbergen and 
Greenland forms with the Barren-ground Caribou, that 
he based some speculations on a former land-connec- 
tion between these countries on this circumstance. 

We have, therefore, records of the present or the 
former existence of a Reindeer resembling the North 
American Barren-ground form in Greenland, Spits- 
bergen, Scandinavia, Ireland, and the South of 
France. In England the remains of the two forms 
occur mixed, but I do not know in how far either the 
one or the other predominates. The Barren-ground 
Reindeer is in Europe altogether confined to the 
west; the most easterly locality that I am acquainted 
with being Rixdorf, near Berlin. The majority of 
the European remains of the Reindeer seem to 
belong to the Siberian or Woodland variety, and it 
would appear as if some intercrossing between the 
two forms had occurred in Lapland, since it is stated 
that in that country the Reindeer is somewhat inter- 
mediate between the two. All the Asiatic remains 
also resemble the Woodland variety. 

As far as I know, no explanation has been 
attempted to account for this peculiar range in 
Europe of the two forms of Reindeer. But if we look 
more closely into the mode of occurrence of the Rein- 
deer remains, we find that the Barren-ground form, 
seems to have existed in Western Europe long before 
the other variety made its appearance there. It was 
pointed out by Struckmann that the Reindeer in 
Southern Europe occurs in older deposits than in 


the north. In speaking of the northern ones, he had of 
course chiefly the German deposits in view. It is in 
one of the oldest pleistocene deposits in Germany 
that the isolated instance, referred to above, of the 
occurrence of the Barren-ground Reindeer, near Berlin, 
has been noted. 

There is still a further point which illustrates the 
supposition that the Barren-ground Reindeer was a 
more ancient inhabitant of Europe than the Wood- 
land one. The latter in all Central European stations 
(in fact almost wherever it occurs fossil) is asso- 
ciated with the remains of the typical inhabitants 
of Siberia, such as the Glutton, Sousliks, Lemmings, 
and others; but in the deposits in which the Barren- 
ground Reindeer have been found in South-western 
France, no other Arctic mammal finds a place. 
Again, in Irish deposits none of the Siberian 
migrants are found. The only explanation of this 
remarkable fact is that the two varieties of the 
Reindeer have come to Europe by different routes. 
We have learned already from the observations of 
Mr. Murray that there are evidences of the existence 
of a former land-connection between North America, 
Greenland, and Spitsbergen. Professor Petersen 
tells us that, according to recent surveys, a high 
submarine plateau with a sharp fall of 1000 fathoms 
towards the Atlantic Ocean begins from Northern 
Norway and is continued as far as Spitsbergen. 
Several islands, such as Bear Island, King Charles 
Land, and others, arise from this plateau, and these 


must be looked upon as the remains of a sunken 
land (Fig. 12). 

From Arctic America, thinks Professor Schulz 
(p. i), we probably have had an uninterrupted 
migration during the greater part of later Tertiary 
times up to the commencement of the Pliocene epoch 
partly over a direct land-connection between Green- 
land, Iceland, and the Faroes, and also between Arctic 
America, Spitsbergen, Franz Josef Land, etc. There 
was also a connection between Asia and Alaska. 

The distribution of the Barren-ground Reindeer in 
Europe seems to warrant the belief that, at the time 
it began its southward wanderings from the Polar 
area, Northern Norway must have been connected 
with Greenland in the manner just indicated, but, 
as I shall explain later on, Russian Lapland and 
part of Northern Russia, or the land between the 
White Sea and the Baltic, must at that time have 
been submerged by the sea. The greater part of 
Denmark and the lowlands of Sweden were likewise 
submerged, but Scandinavia extended south as far as 
Scotland, while Scotland was connected with Ireland, 
and the latter with England and France. The Rein- 
deer migrating south into Scandinavia could only 
reach the continent of Europe by way of the British 
Islands. It appeared there in the west and gradually 
extended its range east, where, as I mentioned above, 
it has occurred in a few isolated localities. 

The advent of the Woodland form of the Reindeer 
in Europe took place at a much later stage. It came, 


as I indicated, with the hordes of Siberian migrants 
which invaded Europe during what is known as the 
Inter-glacial phase of the Glacial period. Scan- 

FiG. 12.- Map of Europe, indicating the parts which were probably 
submerged (shaded) at the commencement of the Glacial period. 
The light portions represent, approximately, the extent of the 
land at that time. 

dinavia, not being then directly connected with con- 
tinental Europe, was not accessible to it; neither 
was Ireland, which had by that time become dis- 


connected from Great Britain. None of the Siberian 
migrants seem to have been able to cross the River 
Garonne, and \ve therefore find neither the Woodland 
Reindeer nor any of the typical Siberian species 
represented in the Pyrenean deposits. 

The Woodland Reindeer persisted in continental 
Europe until comparatively recent times, and it has 
since made its way into Scandinavia across Northern 
Russia, and probably mingled with the older stock of 
the Barren -ground form. In the same way, it may 
have come about that in the English pleistocene 
deposits the remains of the two races occur. 

In a recent contribution to our knowledge of the 
deer tribe (c, p. 88), Mr. Lydekker suggests that the 
former division of the Reindeer races into the two 
forms of Woodland and Barren-ground Caribou, no 
longer holds good. He now recognises no less than 
six races, as follows : 

1. Rangifer tarandus typicus. 

2. spitzbergensis. 

3. caribou. 

4. terrae-novse. 

5. grcenlandicus. 

6. arcticus. 

I hardly think these can be considered of equal 
value; indeed, though there may be differences 
between R. groenlandicus, typicus, arcticus, and spilz- 
bergensis, the antlers exhibit a certain much closer 
relationship among one another than to R. terrcz- 
novce and caribou. But the whole subject is by no 


means as well known as could be wished, and a very 
careful comparative study of recent and fossil remains 
of the Reindeer from various parts of the Old and 
New Worlds is much needed to put our views on a 
firmer basis. 

The presence of the Arctic Hare in Ireland and 
the absence of the common European Hare (Lepus 
europceus) can be explained in a somewhat similar 
manner. The Arctic Hare is the older of the two 
species corresponding with the Barren-ground Rein- 
deer and the European Hare the newer one, 
associating, like the Woodland Reindeer, in its 
westward migration with Siberian animals, though 
probably of Oriental origin. 

Let us once more refer back again to the map on page 
137 indicating the geographical distribution of the 
Arctic Hare. Its discontinuous range and its isolated 
position in the Alps, Pyrenees, and the Japanese 
mountains, all tend to show that it is an ancient 
species. Moreover, its presence in Ireland in the 
plain as well as in the mountains, clearly points to the 
fact that, in the British Islands at any rate, the Arctic 
Hare was the first comer, and that subsequently the 
European Hare invaded these countries. It probably 
found Ireland then no longer accessible, having since 
become separated from England. Again and again 
do we find the statement repeated, that the presence 
of the Arctic Hare in Europe is a clear proof of the 
former prevalence in our continent of an Arctic climate. 
But if so, why should this Hare at present live and 


thrive in Ireland, which has a particularly mild climate 
in winter, and be absent from so many continental 
stations where the temperature more resembles that 
of its native home? If we suppose that the European 
Hare migrated to Europe from the east, after the 
Arctic Hare had become established in Western 
Europe, and drove the latter into the mountains 
or northward whenever the two came into contact, 
we should have, it seems to me, a better ex- 
planation of the range presented by the two species. 
I was formerly of opinion that the European Hare 
had come with the Siberian animals from Siberia, 
but it appears to me more likely now, that it reached 
our continent with the Oriental migrants, and only 
then joined the Siberians in Eastern Europe. 

The evidence in favour of a former land-connection 
between Scandinavia and Greenland, rests on many 
other facts besides those already brought forward. 
That some form of land-connection formerly existed 
between Europe and Greenland is now indeed almost 
universally accepted. That it was situated more to 
the south between Scotland and Greenland is a sup- 
position which has been actively supported by many 
leading authorities, but it seems to me that if such a 
land-bridge existed, it must have been in very early 
Tertiary times, whilst the northern one, such as I 
have indicated, may have originated later and per- 
sisted until a recent geological date. 

The distribution of few groups of animals is now 
better known than that of the larger butterflies and 


moths (Macro-lepidoptera}\ even those of Siberia have 
been fairly well investigated. The interesting facts ob- 
tainable from their distribution are therefore of special 
value. No less than 243 species of Lepidoptera are 
mentioned by Moschler as being common to North 
America and Europe. It is extremely probable that 
a fair number of these have either migrated direct 
from America to Europe or vice 'versa, though many 
may be of Asiatic origin, and have wandered east 
and west from their original home. The following 
twelve species are mentioned by Petersen (p. 38) 
as occurring in Arctic Europe and also in Arctic 
North America, but not in Asia : Colzas nastes, 
Colias hecla, Syrichthus centaurece, Pachnobia carnea, 
Plusia parilis, Anarta Richardsoni, Anarta Schon- 
herri, Anarta lapponica, Anarta Zetterstedti^ Cidaria 
frigidaria, Cidaria polata, Eupithecia hyperboreata; 
and these, as he remarks, point to the possibility 
of a former direct land-connection between Europe 
and North America. 

Mr. Petersen believes that the chief immigration 
into the Arctic area of Europe is post-glacial and 
took place from Siberia, since the majority of the 
species are still to be found in that country at the 
present day (p. 57). He also draws particular atten- 
tion to a fact, which I shall discuss more fully in the 
next chapter, namely, that the most characteristically 
Arctic forms of Northern Europe, which also partly 
occur in the Alps, are entirely absent from the 


Adopting the glacial views of some of our leading 
geologists, Petersen comes to the logical conclusion 
that Central Europe could not have possessed 
any butterflies during the height of the Glacial 
period, but since all evidences seem to point to the 
chief migration from Siberia having taken place after 
the Glacial period, he concludes that they must have 
survived the severe cold of that time in Central Asia. 
He leaves us, however, to imagine .under what possible 
geographical conditions the climate in Europe could 
be too severe for a lepidopterous fauna, while at the 
same time Central Asia could maintain an abundant 

In a suggestive note on the origin of European 
and North American Ants, Professor Emery states 
(p. 399) that a great number of North American 
ants are specifically identical with European ones; 
whilst Dr. Hamilton tells us (p. 89), as an instance, 
that specimens of the beetle Loricera ccemJescens 
from Lake Superior and from Scotland do not seem 
to vary to the extent of a hair on the antennae. 
He enumerates 487 species of Coleoptera as being 
common to North America, Northern Asia, and 
Europe, many of which no doubt have migrated by 
the Americo-European land-connection. 

Arctic Scandinavia or Lapland, according to Sir 
Joseph Hooker, contains three-fourths of the entire 
number of species of plants known from the whole 
circumpolar area. His view, that the Greenland 
flora is almost exclusively Lapponian, having only 



an extremely slight admixture of American or 
Asiatic types, again points to a former more 
intimate connection between North America and 
Arctic Europe, and indeed he remarks (p. 252), 
" It is inconceivable to me that so many Scan- 
dinavian plants should, under existing conditions 
of sea, land, and temperature, have not only found 
their way to Greenland by migration across the 
Atlantic, but should have stopped short on its 
western coast and not crossed to America." 

Hooker's view, that the Scandinavian flora is of 
great antiquity, that, at the advent of the Glacial 
period, it was everywhere driven southwards, and 
that during the succeeding warm epoch the sur- 
viving species returned north, has been adopted by 
the great majority of naturalists. 

The natural corollary of this 'theory is that there 
must have been, between the beginning of the Glacial 
period and the present time, either two independent 
land-connections between the Polar Regions and 
Northern Europe at different epochs to enable 
animals and plants to travel southwards and once 
more to regain their former northern home, or, 
that during the whole of the Glacial period the 
Polar Regions were uninterruptedly connected with 
Northern Europe, until the fauna and flora had once 
more reached their northern goal, after the Polar 
lands had been desolated by the supposed rigours of 
that period. 

In following the history of the Arctic migration to 


Europe, it is of great importance to determine the 
nature and the time of duration of these land- 
connections. The Greenland flora is a very in- 
structive one in helping us to understand many 
of the problems connected with the origin of the 
European plants and animals. To judge from the 
remarks of Professor James Geikie and Mr. Clement 
Reid, no flowering plants could have existed in 
the British Islands during the height of the Glacial 
period, and one would suppose that the cold in Green- 
land at that time must have been far more intense 
than in England. If no flowering plants could exist 
in the latter country, then very surely none could in 
Greenland, where the climate was of necessity by far 
more rigorous. It will be a surprise, therefore, to 
those who are acquainted only with Professor Geikie's 
views of the nature of the Glacial period, that two of 
the most eminent Swedish botanists, who have made 
a special study of the flora of Greenland, have come 
to the conclusion that a survival of flowering plants 
has taken place in Greenland itself from pre-glacial 
times. According to Professor Nathorst (p. 200), 
only a few plants could have survived the Glacial 
period in Greenland. The species now peculiar to 
that country may perhaps, he thinks, be the remnants 
of those which existed in pre-glacial times. Mr. 
Warming, on the other hand, is of opinion that the 
main mass of Greenland's present flora survived the 
Glacial period there (p. 403), and that the remainder 
was carried from Europe and North America by 


occasional means of distribution of the nature in- 
dicated by Darwin. 

Very similar views on the origin of the present 
Polar flora are expressed by Colonel Feilden, who 
says, " To my mind it seems indisputable that several 
plants now confined to the Polar area must have 
originated there and have outlived the period of 
greatest ice-development in that region" (, p. 50). 
No land-connection at all need be supposed to have 
existed in recent geological times, that is to say, 
during the Glacial period or after, if Mr. Warming's 
and Colonel Feilden's views be adopted. A pre-glacial 
connection would be sufficient to explain the general 
features of distribution. An admission is thus ob- 
tained from these two independent authorities that 
the climate during the Glacial period must have 
been vastly less severe in the Polar Regions than 
is generally conceded. I am of opinion that not 
only the whole of the present flora, but also the 
fauna of Greenland survived the Glacial period in 
that country. 

If we suppose that an extensive centre of origin 
existed in the Polar area, or we may say in Green- 
land, both animals and plants would have been able 
to spread from it into Northern Europe and North 
America by means of the land-connections which are 
generally supposed to have existed in pliocene times, 
that is to say, just before the commencement of the 
Glacial period. There must have been at this time 
a connection too between Scotland and Scandinavia, 


which will be dealt with more fully presently. The 
important point is to consider what light the Green- 
land flora arid fauna will throw upon the problem 
of the continuity of the aforesaid land-connec- 
tion during the Glacial period. We have seen 
that the Barren-ground Reindeer, a typically Polar 
species, penetrated as far south as the Pyrenees, the 
Arctic Hare went as far, while a number of other 
species of Polar animals and also of plants occur 
in the Alps. Of these it remains to be seen how 
many have come direct by way of Northern Europe 
or from the Polar Regions by way of Asia. At any 
rate, as the origin of the Alpine animals and plants 
will be discussed in another chapter, there is no need 
to dwell on this subject at present. 

From the nature of the distribution in Ireland of 
Arctic plants and animals, which occur mostly on the 
north and west coasts, it would seem that a stream of 
migration entered from Scotland, and I have no doubt 
that that same migration came into Scotland directly 
from Scandinavia by a route over which now roll the 
waves of the North Sea. There is, moreover, as I 
already mentioned on p. 94, a very interesting so-called 
American element in the north-western European 
flora, that is to say, plants now found in North-west 
Europe and North America without occurring in 
Greenland or any of the islands which might have 
formed the former highway between the Old World 
and the New. These are probably some of the more 
ancient Polar plants which have become extinct in 


the Arctic Regions and survive in isolated patches in 
favourable localities. We find seven species of these 
American plants in Ireland, almost entirely confined 
to the north and west coasts. These are Spiranthes 
Romanzoviana, Sisyrinchium anceps, Naias flexilis, 
Eriocaulon septangular e^Juncus tennis , and Polygonum 
sagittifoliuui. To them must be added another plant 
recently discovered by the Rev. Mr. Marshall in the 
south of Ireland, namely Sisyrinchium calif or nicum. 
As I have mentioned in former writings, there are 
three species of North American freshwater-sponges 
in Ireland which have not hitherto been discovered 
elsewhere in Europe or in Asia. These, namely 
Ephydatia crater if ormis, Heteromeyenia Ryderi, and 
Tubella pennsylvanica, all occur in some of the lakes 
near the western coast of Ireland. 

There are in all groups of animals instances of species 
which are confined to Europe and North America, while 
unknown from the Asiatic continent, but none, as far 
as is known, have such a very discontinuous range as 
that of the animals and plants just referred to. In 
some cases the species still occur in Greenland, and 
in this way make it still clearer that their migration 
in former times took place from one continent to 
the other by way of that country. As an interesting 
instance of such distribution may be mentioned the 
Common Stickleback (Gasterostens aculeatus], which 
is found in Greenland, North America, and Europe, 
but is quite absent from Asia. Then again, the Nine- 
spined Stickleback (Gasterosteus pungitius] is confined 


to Western Europe and North America, though an 
allied species, Gasterostcus sinensis, lives in China and 
has probably penetrated there from the New World 
across the old Behring Straits land-connection. 

The Coleoptera Diachila arctica, Elaphrus lapponicus, 
and Blethisa multipunctata are good instances of 
species which have come to us from North America 
by way of Greenland. I have already referred to 
the Lepidoptera, but might add that eleven species 
of Anarta occur in Scandinavia, eight of which 
reappear again in Labrador, none of them, however, 
being met with in Siberia. Then again, take the in- 
teresting Crustacean Lepidurus (Apus] glacialis. It 
is found in Greenland, Spitsbergen, Lapland, and 
Norway; and formerly, as we know from fossil 
evidence, it ranged into Scotland. Another Phyllo- 
pod, viz., Branchinecta palndosa, inhabits Greenland, 
Lapland, and Norway. Mr. Kennard suggests that 
the freshwater Snail Planorbis glaber might also 
belong to the same migration. And there are no 
doubt large numbers of others. 

Professor Emery mentions that Northern Europe 
possesses one peculiar genus of Ant, viz., Anergates. 
This is closely allied to Epoccus, another genus con- 
fined to North America. It seems probable, there- 
fore, that both of these have sprung from an Arctic 
genus which sent two branches southward into the 
two continents without there being any migration 
through Asia. 

The general range of the Arctic plants and animals 


gives no reason to suppose that the Greenland fauna 
and flora of the present day were exterminated by the 
Glacial period and then reintroduced into that country. 
Nor have we any evidence that such a fauna and flora 
migrated across the British Islands northward. The 
Greenland animals and plants too are altogether much 
more like the Lapland ones than those of Scotland. 
It will also become evident to the reader of this work 
that no very extensive migrations could have taken 
place during the post-glacial period, and that almost 
everything points to a survival of both fauna and 
flora in northern latitudes throughout the Glacial 

If we take into consideration the palaeontological 
evidence of the two races of Reindeer in Europe, one of 
which came to us from the north, and that the Arctic 
Hare and one of the races of the Stoat entered our 
continent from the same direction when we, more- 
over, carefully review the numerous other instances 
quoted of plants and animals which could only have 
reached us from the north, the irresistible conclusion is 
forced upon us that a land-connection existed at no 
very distant period between Northern Europe and the 
Arctic Regions of North America. This is not a new 
hypothesis. Many geologists are of opinion that a 
land-passage did exist within comparatively recent 
times, uniting Europe, Greenland, and North America. 
But the position of this old land-bridge, as I have 
mentioned, has been generally placed somewhat 
farther south than I should feel inclined to put it. 


The fact that very extensive glaciers formerly 
covered the mountains of Scandinavia on the eastern 
side, whilst they scarcely reached the sea on the west 
(Feilden, a, p. 721), seems to favour the view of a 
warm current having washed the western shores. As 
I shall attempt to show later on (p. 179), the Arctic 
Ocean extended across Northern Russia at that time 
from the White Sea to the Baltic that is to say, to 
the eastern shores of Scandinavia, which country was 
then joined to the north of Scotland. The predis- 
posing agents to a copious snowfall existed in 
Scandinavia, viz., an excessive evaporation of the 
warm Atlantic waters and unusual precipitation in 
the form of snow owing to the cold given off by 
the Arctic waters on the east side of the mountains. 
It is therefore probable that the land -connection 
which united Europe and North America was farther 
north than has been supposed. 

If we sail straight across from Northern Scandinavia 
to Greenland, we traverse an exceedingly deep marine 
basin ; but if we examine the sub-marine bank which 
runs all along the coast of the former country from 
south to north, we find that it does not end when the 
extreme north of the land is reached. The bank 
extends much farther north, and is continued as far 
as Spitsbergen. As I have said before, the latter, 
as well as Bear Island, must be looked upon as the 
remains of a large mass of sunken land the ancient 
Scandinavia stretching far into the Arctic Circle. Pro- 
fessor Nathorst speaks of Spitsbergen as a northern 


continuation of Europe, not only geographically, but 
also botanically and geologically. However, this 
northern land must have stretched even farther not 

FIG. 13 Map of Europe, indicating approximately the distribution of 
land and water during the earlier stages of the Glacial period 
shortly after the period represented in Fig. 12, p. 156. The darkly 
shaded parts indicate the areas covered by water, and the white 
portions what was land at the time. 

perhaps farther north, but farther west. Here lay the 
old land-connection between Scandinavia, Greenland, 
and North America (Fig. 13). One of the highest 


authorities on the geographical distribution of plants, 
Professor Engler, maintains that the arguments in 
favour of this Arctic connection of America with 
Europe are more weighty than those for a land-bridge 
between Greenland, Iceland, the Faroes, and Great 
Britain. Moreover, he is of opinion that a certain 
number of species of plants belonging to the Alpine 
flora of Arctic Siberia have travelled from Scandinavia 
via Greenland and North America to Eastern Asia, 
and not direct from Scandinavia to Siberia (p. 143). 

That this ancient Arctic land-connection existed 
almost throughout the Glacial period appears to me 
probable. It has often been suggested that such a 
land-barrier was one of the principal causes of the 
production of the glacial phenomena in Europe, 
and as such it must have existed intact certainly 
during the earlier stages of the Glacial period. 
The barrier must then have gradually subsided in 
one or two places ; and once a breach was formed, 
the complete union between the Atlantic and 
the Arctic Oceans could not have been long 

The terrestrial fauna and flora, as we have seen, 
lend strong support to the view of the former 
connection between Scandinavia and Greenland, but 
many other facts point in the same direction. It was 
Edward Forbes who first drew attention to the pres- 
ence of a number of species of littoral molluscs on 
the coast of Finmark which also occur on the coast of 
Greenland, and he expressed the firm conviction that 


they indicated by their existence on both sides of the 
Atlantic some ancient continuity of the coast-line. 
He held that the line of migration of these mollusca 
was probably from west to east, and that it must 
have taken place during physical conditions entirely 
different from those prevailing at present If Forbes's 
view is correct, a current must have existed from the 
north coast of North America along the northern 
shore of the ancient land which stretched east as 
far as Europe. We have also some palaeontological 
evidence bearing on the existence of such a current 

(P- 173). 

As we shall learn presently, the early stages of 

the Glacial period were accompanied by a marine 
transgression over Northern Russia and Germany 
an overflow, as it were, of the waters of the Arctic 
Ocean covering a great part of Northern Europe, 
with the exception of Norway. One continuous 
ocean ultimately extended from the east coast 
of England across Holland, Northern Germany, and 
Russia to the White Sea (Fig. 12, p. 156). The 
south of England being at that time joined to 
France, and Scotland to Scandinavia, there was no 
direct communication between this large North 
European Sea and the Atlantic. The glaciers 
which took their origin in the Scandinavian Moun- 
tains discharged icebergs into this sea, and many of 
them no doubt were stranded on the east coast of 
England. The boulders of Scandinavian origin which 
have been discovered in recent geological deposits on 


that coast have generally been traced to the action of 
land-ice, but the supposition that they have been 
carried by icebergs the older theory appears to 
me the more probable one. Such boulders begin 
to make their first appearance in the Red Crag, a 
deposit which is now looked upon as belonging to 
the newer pliocene series. But whether we call it 
pliocene or pleistocene really matters little. The 
important fact is, that glacial phenomena, consisting 
of the appearance of boulders foreign to the country 
together with an invasion of Arctic shells, are now 
ushered in upon a coast which shortly before teemed 
with the southern life of a Mediterranean character. 
Among the new arrivals in these English crags there 
are no less than eighteen species of North American 
marine mollusca. Since the German Ocean had then 
no direct communication with the Atlantic, these 
mollusca could only have come from the White 
Sea, and Forbcs's Arctic current would offer an 
explanation of the manner in which they were 
enabled to migrate there from their original 

It might be urged that we have no grounds 
for the supposition that the German Ocean was 
practically a closed basin; and that these American 
species probably inhabited at that time the whole of 
the North Atlantic Ocean. But if such had been the 
case, we ought to have evidence of the occurrence of 
some of these species in the newer Tertiary deposits 
along the west coasts of the British Islands. Such 


beds exist; there is, however, not a trace in any 
of them of any American mollusca. In examin- 
ing the marine deposits of St. Erth, on the coast of 
Cornwall, which are believed to be of about the same 
age as the newer crags, Messrs. Kendall and Bell 
were much struck by the absence of the species 
characteristic of the latter. The St. Erth fauna led 
them to believe that the Arctic Ocean could not 
then have opened into the Atlantic, but that a land- 
communication had existed between Europe and 
North America, so as to form a barrier of separation 
between the two oceans. This again perfectly har- 
monises with the views I have expressed, and 
supports them. 

Let us now look a little more closely at the 
history and the fauna of the Baltic and the adjoining 
lakes, in order to gain additional information as to the 
geographical changes which have had such lasting 
influence on the peninsula of Scandinavia, The 
Baltic is a shallow sea covering an area of 184,496 
square miles, and its waters are decidedly brackish. 
The fauna is a poor one, being too salt for the purely 
freshwater species and not salt enough for the typical 
marine forms. The absence of some animals which 
we should expect to find there is one of the remark- 
able features about the Baltic, but, on the other hand, 
some species occur which are altogether strangers to 
the fauna. And these, moreover, are confined to the 
extreme northern end of the sea. I need only refer 
to the Arctic Seal (P/wca annelata), which is confined 


to the Gulf of Bothnia, and to the four-horned sting- 
fish (Cottus quadricornis, Fig. 14, p. 178), neither of 
which occur on the west coast of Scandinavia. But 
there are others which point in an equally unmistakable 
manner to the former existence of a marine connection 
between the Baltic and the southward prolongation of 
the Arctic Ocean known as the White Sea. It is 
generally admitted now that such a union between 
these two seas, viz., the Baltic and the White Sea, 
occurred in recent geological times, but opinions 
differ as to the duration of this connection. I 
adhere to the view expressed by Murchison and 
others, that the boulder-clay is a marine deposit. I 
am also convinced that the Arctic Ocean, as I have 
already mentioned, transgressed over the lowlands 
of Northern Russia at about the time when the 
newer crags were being deposited on the east coast 
of England; that the same large sea also covered 
Northern Germany, Denmark, Holland, and the low- 
lands of Sweden, and laid down the lower continental 
boulder-clay which is spread over such vast tracts of 
land in those countries. I shall have occasion to refer 
to this again more fully in the next chapter; mean- 
while, it should be remembered that this stage was 
followed by a partial retreat of the northern sea, 
though Scandinavia did not become joined to the 
Continent. The date of this retreat of the sea, repre- 
sented in Fig. 13, corresponds probably to what is 
know as the inter-glacial phase of the Glacial period, 
and I think it must have been during this time that 


the Forest-Bed on the coast of Norfolk was laid 
down. 1 

None of the Siberian mammals apparently entered 
Scandinavia at the time when they invaded Central 
Europe and penetrated as far west as England and 
Western France. Nor did the great Oriental mammals, 
like the Mammoth and others, reach Scandinavia; and 
Professor Pohlig argued, on the strength of these 
facts, that the latter country was either for a 
very short time only free from ice, or that it had 
defective land-communication with the Continent 
during inter -glacial times. This seems to me 
scarcely to explain the facts of distribution and 
account satisfactorily for the absentees. Nor does it, 
of course, harmonise with the views that I have 
announced above. Professor Engler's remark (p. 131), 
that Scandinavia probably projected above the glacial 
sea as an island, is more in accordance with these 
views, though the term island is scarcely applicable 
to that country, since it was always, as I said, in- 
directly joined to the Continent (vide Fig. 13, p. 170). 
The fauna of Scandinavia, both fossil and recent, points 
to a direct isolation of that country from the continent 
of Europe during a considerable period. 

Another proof that Northern Russia and the low- 
lands of Sweden were covered by the sea comes to us 
from a study of the fauna of the relict lakes the 
" Reliktenseen " of Leuckart. This name was first 
applied by Leuckart to lakes containing marine 

1 I have already expressed this view on p. 120. 


organisms, which are supposed to have been 
flooded by, or to have been in close communication 
with the sea at some former period, like the lakes 
Ladoga and Onega in Russia. His views have been 
worked out subsequently in greater detail by Loven 
and O. Peschel, who gave them their strong adherence. 
Many leading zoologists, such as Professor Sars and 
others, have since adopted them, and though dis- 
credited by Professor Credner, the theory still 
offers the best explanation for the origin of marine 
animals in freshwater lakes. 

Professor Credner's contention, that marine mollusca 
are always absent from these relict lakes, seems at first 
sight a stumbling-block to the theory. But the ex- 
planation is really simple enough. It is to Dr. Sollas 
that we owe a very ingenious explanation of the origin 
of freshwater faunas. He showed that all freshwater 
organisms in their early stages of development are pro- 
vided either with some process enabling them to attach 
themselves to a foreign object, or that they pass this 
period within the body of the parent. This is a 
provision of nature to prevent freshwater organisms 
from being floated out to sea, where they would 
perish, until they reach maturity and can cope with 
floods and currents. Had Professor Credner been 
aware of Dr. Sollas's views, no doubt he would have 
modified his criticisms, for, as most marine mollusca 
have free- swimming larvae, they would have little 
chance of becoming permanent residents of lakes. 
During their larval stage, marine molluscs are quite a 



prey to the currents of the sea. They have practically 
no swimming organs, and only move by lashing to and 
fro the tender cilia with which they are provided. 

This disposes, therefore, of Professor Credner's 
main criticisms. As for the fauna of the relict 
lakes, we are now only concerned with those of 
Northern Russia, Finland, and Sweden. In the 
lakes Wetter and Wener in the latter country 
occurs the four-horned sting-fish (Coitus quadricornis y 

FlG. 14. The Four-horned Sting- fish (Coitus qitadricornis], reduced 
from Professor Smitt's figure in the Fishes of Scandinavia. 

Fig. 14), which, as we have learned, also inhabits the 
northern part of the Baltic, and, as was suggested, 
migrated there at a time when the latter was 
connected with the White Sea. The principal food 
of this little fish consists in a marine Crustacean 
called Idotea entomon, an animal allied to our 
common woodlouse. This is a typical marine 
species, but it occurs also in the relict lakes of 
the countries mentioned above, as well as in the 
Baltic and the Caspian. Perhaps the best known 


form with a similar range is the Schizopod crus- 
tacean Mysis relzcta 1 (Fig. 15), which is clearly a des- 
cendant of the Arctic marine Mysis oculata> of which 
it was formerly considered a mere variety. The two 
Amphipods Gammaracanthus relictus and Pontoporeia 
ajjinis, and the Copepod Limnocalanus macrurus, are 
three additional well-known Arctic crustaceans whose 
range differs but little from those above-mentioned. 2 

FIG. 15. Mysis relicta, a small shrimp-like Crustacean, after Sars 

These facts all go to prove that the sea formerly 
covered the lowlands of Sweden, Finland, and 
Northern Russia. The fauna of Scandinavia, as 
we have seen, indicates that during the greater part 
of the Glacial period the country was not directly 
connected with continental Europe as it is now. It 
seems that the barrier of separation probably con- 

1 The occurrence of this species in Lough Neagh in Ireland, pointing 
to a connection between the Irish Sea and the Baltic, will be referred to 
later on; as also that of two allied forms in the Caspian Sea. 

- For additional species with a similar range, vide Nordquist. 


sisted of a broad expanse of ocean on which floated 
numerous icebergs, which originated from the Scan- 
dinavian glaciers as they reached the sea. This was 
a cold sea, whilst Western Scandinavia was washed 
by the Gulf Stream (vide Fig. 12, p. 156). We 
might look upon the boulder-clay which covers such 
vast tracts of country in Northern Germany, Russia, 
and Holland as deposits formed by this sea rather 
than the ground-moraine of a huge Scandinavian 
glacier. I shall refer to this subject again in the 
next chapter ; meanwhile it may be remembered 
that the boulder-clay of Northern Europe exactly 
resembles in all important particulars the similar 
accumulations met with in the British Islands. 
They resemble one another also in the occasional 
occurrence of sea-shells, the frequent appearance of 
bedded deposits, and the often inexplicable course 
taken by boulders from their source of origin. There 
occurs often a singular mixture and an apparent 
crossing of the paths of boulders in the boulder- 
clay. Professor Bonney remarks (p. 280) that these 
are less difficult to explain on the hypothesis of 
distribution by floating ice than on that of transport 
by land-ice, because, in the former case, though the 
drift of winds and currents would be generally in one 
direction, both might be varied at particular seasons. 
So far as concerns the distribution and thickness of 
the glacial deposits, he says there is not much 
to choose between either hypothesis; but on that 
of land-ice it is extremely difficult to explain the 


intercalation of perfectly stratified sands and gravels 
and of boulder-clay, as well as the not infrequent 
signs of bedding in the latter. Two divisions are 
generally recognisable in the continental boulder-clay 
a lower and an upper. An inter-glacial phase 
characterised by a less severe climate is assumed 
to have intervened between the deposition of the 
two. In Russia no such division can as a rule be 
made out, and sea-shells are either entirely absent or 
extremely scarce. It has been pointed out by Pro- 
fessor J. Geikie that the erratics a name applied to 
boulders in boulder-clay in the upper division have 
travelled in a different direction from those contained 
in the lower. Taking for granted that the boulder- 
clay is a marine deposit, this phenomenon seems 
to indicate that the current which prevahed during 
the early part of the Glacial period in this North 
European ocean was different from the prevailing 
current during the latter part. I have attempted 
to explain this circumstance by the supposition 
that during the early part of the Glacial period 
the Northern Sea had a connection with the Ponto- 
Caspian Sea a sea formed by the junction of the 
Black Sea and the Caspian (Fig. 12, p. 156). There 
is geological evidence, as will be explained in the 
following chapter, that the area of these two seas was 
considerably larger in glacial times than it is now, 
and that they were joined across the valley of the 
Manytch. After the inter-glacial phase of the Glacial 
period, the North European Ocean became connected 


with the Atlantic Ocean across the north of England 
(Fig. 6, p. 126), the junction between the former 
and the Ponto-Caspian having meanwhile become 
dry land (Fig. 13, p. 170). A fresh current, now 
flowing westward, was set up in the North Euro- 
pean Ocean, which accounts for the fact just cited 
that the erratics in the upper continental boulder- 
clay have travelled in a different direction from 
those in the lower. The boulder-clay laid down by 
the sea on the midland and northern counties of 
England, just as was the case with the similar deposit 
on the Continent, is generally accredited to the 
action of land-ice. It is by most geologists looked 
upon as the ground-moraine, partly of the huge 
Scandinavian glacier which is supposed to have 
impinged upon the English coast, partly of local 
British glaciers. 

But renewed geological investigations on this point 
throw doubts upon these theories. Thus Mr. Harmer 
remarks in a recent contribution to glacial literature 
(p. 775), that "it is difficult to see how the Baltic 
glacier could have reached East Anglia, though ice- 
floes with Scandinavian boulders might easily have 
done so, while had the Norwegian ice filled the North 
Sea and overflowed the county of Norfolk, some evi- 
dence of its presence ought to be found in the glacial 
beds of Holland." 

All the phenomena of distribution of the British 
fauna and flora are, as we have seen, much more easily 
explained by the supposition of a damp, temperate 


climate, such as might have been produced by the 
proximity of a cold sea on one side and of a warm 
one at the other, than by invokirg an arctic climate 
with enormous glaciers. Most of the living animals 
and plants would have been exterminated under the 
latter conditions. Palaeontological evidence in Great 
Britain clearly indicates that southern species migrated 
first to these islands, that Arctic species were then 
driven south from their native lands, probably owing 
to insufficient food-supply and climatic changes in 
the north, that finally eastern species invaded the 
country all this without the annual temperature 
of Europe being apparently much affected. For we 
find in the British pleistocene deposits and Mr. 
Lydekker draws particular attention to this remark- 
able fact a curious intermingling of southern and 
northern mammals, which undoubtedly lived side by 
side. Everybody knows that northern and Arctic 
species can live perfectly well in a temperate climate, 
but that it is almost impossible to acclimatise 
southern animals in an Arctic or even temperate 
one. We have in this circumstance almost a proof, 
therefore, that the climate cannot have been very cold. 
Though a cold sea bathed the shores of Eastern 
England, and even eventually invaded a portion of 
Northern England, the warm ocean on the west 
must have effectually prevented any great lowering 
of temperature. 

At the time when the North European Sea flooded 
a portion of England, Scandinavia was still connected 


with Scotland, and the latter with Ireland (Fig. 6, 
p. 126). There is no doubt that the food-supply in the 
Arctic Regions was decreasing with an increase of 
snowfall and with the gradual lowering of the land, 
which reduced also the habitable area. Arctic species 
therefore were driven south in search of fresh pastures. 
But it need not be supposed that anything like a vast 
destruction of the fauna of the Arctic Regions took 
place. Only fewer mammals were able to find food 
in a given space than heretofore. This southward 
migration may have commenced, in the case of 
plants and the invertebrates, at a much earlier time, 
during the Miocene or Pliocene Epochs, but it 
is doubtful whether the mammals and birds which 
we find in our pleistocene and recent deposits 
began to travel south much before the commence- 
ment of the Glacial period. The beginning of the 
Glacial period in England, I think, is indicated by 
the deposition of the Red Crag, though the latter is 
generally regarded as belonging to the pliocene 
series. Much of the northward migration from 
the British Islands of Lusitanian and other forms 
had then ceased, but we have in Scandinavia, just 
as in these islands, a southern relict fauna and flora, 
plants and animals which had wandered across what 
is now the German Ocean from Scotland to Scan- 
dinavia, and have never become extinct in that 
country to the present day. I need only mention 
the Red Deer, the Badger, and Slugs of the genus 
A rion. 


Professor Blytt directs attention to some such 
southern relict species of plants now only found in the 
extreme south-west of Scandinavia, such as Asplenium 
niarinuni) Hymenophyllum Wilsoni, Carex binervis^ 
Scilla verna, Erica cinerea, Conopodimn denudalum, 
Meum athamanticum, and Rosa involuta (p. 28). 

The Arctic fauna and flora in Scandinavia that 
is to say, the descendants of those species which 
migrated direct from Greenland and Spitsbergen, 
as we have seen, are numerous. They of course per- 
sisted throughout the Glacial period in the country, 
and are now in many localities being exterminated 
partly by change of climate, partly by a keen com- 
petition with more vigorous rivals which have come 
to Scandinavia from the east. It is a curious circum- 
stance, as pointed out by Professor Blytt, that the 
Arctic plants in the Botanic Gardens at Christiania 
are able to stand almost any amount of sunshine, 
but are very liable to be injured by the frost, and have 
to be covered in the winter. A similar observation 
has been made in the case of the Alpine plants at 
Kew Gardens, which have to be wintered in frames, 
though their homes are either in the high Alps among 
the everlasting snows or in the intensely cold climate 
of Greenland. Many of the Scandinavian plants ex- 
hibit instances of discontinuous distribution, thus show- 
ing their ancient origin ; and there is altogether nothing 
in the fauna and flora of that country which might 
lead us to believe that these were exterminated 
during the Glacial period and reintroduced subse- 


qucntly. The climate during that period in Scan- 
dinavia was probably more equable and moister, 
with a greater snowfall in winter and with less 
sun to melt the snow during summer, so that the 
development of glaciers took more formidable dimen- 
sions, chiefly on the east side. The lowlands of 
Sweden were covered by the sea, whilst many of 
the valleys were choked with ^ great glaciers, which 
cast off portions of ice as they reached the sea, just 
as the Greenland and other northern glaciers do {vide 
p. 237). A country which at the present day probably 
somewhat resembles the former Scandinavia climati- 
cally is Tierra del Fuego, in the extreme south of 
South America. Though there is an abundant 
snowfall, so that glaciers reach the sea in many 
parts of the country, the flora has been described by 
travellers as luxuriant; and it appears that the fauna 
also is richer than might be expected from the cheer- 
less climate. 

Towards the latter part of the Glacial period the 
land-connection between Scandinavia, Spitsbergen, 
and Greenland broke down, and the waters of the 
Arctic and Atlantic Oceans joined. Whether it was 
at this time or later that the other land-connection 
between Scandinavia and Scotland collapsed is 
difficult to determine; but it is certain, I think, 
that Scotland was still united with Ireland even after 
these two great land-bridges ceased to exist. 




The fauna of the Arctic Regions is much poorer than that of the 
other regions which are dealt with in this work. In some groups, 
such as Reptiles and Amphibia, there are no representatives at all, 
but no doubt a larger number of species existed there in earlier 
Tertiary times. At least we have fossil evidence that during 
the Miocene Epoch plants of many families flourished in Green- 
land of which no vestige is now left in the Polar area. Climatic 
conditions must therefore have changed, as in Europe. A 
gradual refrigeration took place, owing probably to the slow 
withdrawal of the current which supplied the Arctic Sea with 
warmth. Greenland and Europe were then connected, and 
the Arctic Ocean was separated from the Atlantic. This land- 
connection is supposed to have lain far north between Scan- 
dinavia, Spitsbergen, and Greenland, and must have persisted 
until towards the end of the Glacial period. 

As the temperature decreased and the land-area available in 
the north diminished, the surplus population, consisting of 
animals and plants, and possibly also of human beings, moved 
southward. We have traces in Europe, and especially in the 
British Islands, of a very early migration from the north in the 
so-called American plants and in the freshwater sponges. The 
geographical distribution of some of the Arctic species of mam- 
mals is referred to in greater detail, to show how the relative 
age of their entry into Europe can be determined. Two forms 
of Reindeer, resembling the Barren-ground and Woodland 
varieties, have been met with in European deposits, but only 
the former occurs in Ireland and the south of France, whilst 
eastward the other becomes more common, and finally is the 
only one found. It is believed that the Barren-ground is the 
older form as far as Europe is concerned, and that it came to 
us with the Arctic migration, and that the other Reindeer 
reached Europe much later from Siberia, when Ireland had 
already become detached from England. The range of the 


Arctic Hare is equally instructive. It must have been a native 
of Europe since early glacial or pre-glacial times before the 
common English Hare had made its appearance in Central 
Europe. Along with other Arctic forms, it entered Northern 
Europe directly from the Arctic Regions, by means of the former 
land-connection which joined, as I remarked, Lapland with 
Spitsbergen, Greenland, and North America. There need not 
have been a post-glacial connection between Europe and 
Greenland ; the present flora of that country may have survived 
the Glacial period in the Arctic Regions, as has been main- 
tained by some botanists and other authorities. Professor 
Forbes argued from the occurrence of the same species of 
shore mollusca on the coast of Finmark and Greenland that 
these two countries were not long ago joined, so that a slow 
migration from west to east along an ancient coast-line could 
have taken place. That such a migration actually occurred is 
further made probable, judging from the presence of American 
mollusca in the Crag deposits on the east coast of England. 
These came into the North Sea in the first place direct from the 
Arctic Ocean at a time when the two oceans freely communi- 
cated with one another across the lowlands of Northern Russia, 
Northern Germany, and Holland. Arctic shells are also found 
below the boulder-clay on the Baltic coast, and a free com- 
munication such as indicated is generally held to have taken 
place at no very distant date. The so-called "relict species" 
marine animals left in freshwater lakes in districts formerly 
covered by this sea lend some support to this view. But the 
view that the continental boulder-clay is a marine deposit is not 
now held except by a few, though I here bring it forward again, 
as it seems to me to fit in so much better with the known facts 
of distribution. The sea just referred to probably existed 
throughout the greater part of the Glacial period; and icebergs, 
which originated from the Scandinavian glaciers, would have 
brought detritus and boulders to the lowlands. Scandinavia 
was then connected with Scotland, and England with France. 



IN dealing with the British fauna in particular, I 
have drawn attention to the fact that it is chiefly in 
the south of England that we find fossil remains 
of eastern species of mammals in recent geological 
deposits. We can actually trace the remains of 
these species and their course of migration across 
part of the Continent towards Eastern Europe, and 
as none of their bones have been discovered in the 
southern or northern parts of our Continent, it must 
be assumed that their home lay in Siberia, where 
many still exist to the present day, and where 
closely allied forms also are found. Some of these 
Siberian migrants have remained in England and 
on the Continent to the present day. Many have 
become extinct. But the animals forming this 
eastern migration did not all originate in Siberia, 
though I have sometimes spoken of them collectively 
as Siberian migrants. There must have been other 
centres of dispersion of species in Europe. We know 
that a very active centre of development at any rate 
for land-mollusca lay in South-eastern Europe, either 
in the Caucasus or in the Balkan peninsula, or more 


probably in both. The Alps no doubt produced a 
number of species which have spread north and 
south, and may in their wanderings have joined the 
Siberian migrants in their western course, and thus 
have reached the British Islands. Nevertheless, the 
majority of the mammals belonging to the eastern 
clement of the British fauna (vide p. 95) have un- 
doubtedly originated in Siberia. The Polecat (Mus- 
tela putorius} and the Harvest Mouse (Mus minutus], 
for instance, are members of that eastern migration. 
Both occur throughout Central Europe and a large 
portion of Siberia, but are absent from the extreme 
north and south of Europe and also from all the 
Mediterranean Islands. A Siberian species, which 
has never penetrated so far west as the British 
Islands, nor even so far north as Scandinavia or 
south to Italy, is what is known in Germany as the 
"Hamster" (Cricetus frumentarius], a little Rodent 
which spends the winter asleep in its burrows, and 
surrounds itself with a great accumulation of food- 
material carried there during autumn. The common 
English Hare, which I formerly regarded as an 
instance of a Siberian mammal, must now find a 
place among the Oriental migrants. Its history is 
very instructive, and I shall have an opportunity later 
on to refer to it again. Meanwhile, it may be men- 
tioned that though this Hare inhabits Europe in two 
varieties or races, one of which, Lepus mediterra- 
neus, is confined to Southern Europe, the latter owes 
its origin to an earlier migration from Asia. 


When we come to consider the eastern birds, we 
have to distinguish between resident species and 
migratory ones. The Black-throated Thrush (Turdus 
atrigularis], which has been twice obtained in the 
British Islands, is a mere straggler to Europe, and 
is not known to breed there at all. Better known 
birds, perhaps, are the Golden Thrush ( Turdus varius\ 
which has even occurred as far west as Ireland, the 
Rock-Thrush (Monticola saxatilis) and the Scarlet 
Grosbeak {Carpodacus erythrinus], which breed in 
Eastern Europe, but are known only as occasional 
visitors in the west. 

To judge by their distribution, the Bullfinches 
(Pyrrhula) are of Asiatic origin, for seven species out 
of ten are confined to that continent. Our common 
Bullfinch (P. europed] probably came with the Oriental 
migrants, or perhaps its ancestors did. But the larger 
Northern or Russian Bullfinch (P. major] has no doubt 
entered our Continent directly from the east. We have 
in many groups similar instances of closely allied species 
er varieties, one of which, originating at a somewhat 
later stage than the other, took a different route of 
migration from that followed by its near relative. 

The Pine -Grosbeak (Pimcola enudeator) is only 
known to British ornithologists as an exceedingly 
rare visitor. Its real home lies in the northern parts 
of Europe, Asia, and North America, and it is one 
of the most typical of the Siberian migrants. 

But there are a number of other species of birds, 
which, though probably not of Siberian origin, 


only migrated westward recently, and have either 
not yet reached the British Islands, or which lead one 
to suppose, from their British range, that they are 
eastern forms. 

Such, for instance, is the Nightingale (Danlias 
luscinia\ which is probably of Oriental origin, but 
only visits England regularly in spring. There is no 
authenticated record of its ever having migrated 
either to Scotland or Ireland. 

The Bearded Titmouse (Panurus biarmicus) is one 
of the eastern birds still resident in England, though 
unfortunately it seems tcr be on the verge of extinc- 
tion. It is unknown in Scotland and Ireland. 
Another resident eastern species is the Nuthatch 
(Sitta ca>sia\ but neither of these is probably of 
Siberian origin. 

The majority of the European Reptiles are probably 
of eastern origin. Among our British species, the 
Common Viper (Pelias berus\ for example, is a 
typically eastern form. It is almost unknown in 
Southern Europe proper that is to say, in Italy, 
the Balkan peninsula, and the Mediterranean Islands, 
but its range extends in the west as far as Spain, and 
in the east right across the Asiatic continent to 
Japan. It is well known that the Viper occurs in 
Scotland, and that neither it nor any other snake is 
found in Ireland. There is a legend, indeed, that 
snakes did once exist in Ireland and were banished 
from the island by St. Patrick, but unfortunately 
we have no historical evidence that such an 


interesting event actually took place. The Sand- 
Lizard (Lacerta agilis\ another British species, may 
be looked upon as an eastern form. It is quite absent 
from Italy, the Balkan peninsula, and the Medi- 
terranean Islands, but extends throughout Central 
Europe to the east. 

Among the species of eastern Reptiles which have 
a mere local range in Europe might be mentioned 
the two Lizards, Phrynocephalus auritus and Agarna 
sanguinolenta. They belong to the family Iguanidcs^ 
which includes some very large species. Both of 
them are Asiatic forms, which have only just pene- 
trated across the eastern steppes into Europe, where 
they inhabit the arid regions between the Caspian 
and the River Don in Southern Russia. 

The species of Mammals living in Europe at the 
present day have, with few exceptions, migrated 
to our continent from other parts of the world. 
With regard to the Birds, it is possible that a 
somewhat larger number proportionally may be 
of European origin. Still, the great majority are, 
I think, to be regarded as immigrants. The autoch- 
thones are about equal to the immigrant reptiles, 
but many of the European Amphibians and the 
majority of the Fishes have probably originated on 
our continent. Some of the European Amphibia 
especially among the tailless forms appear to be 
immigrants from Asia. Thus the distribution of 
Rana arvalis in Europe is remarkably like that of 
a Siberian migrant. This frog occurs in Siberia, 



ranging southward as far as Persia and parts of 
Asia Minor. Crossing the European border, we 
find it in Russia, Upper Hungary, North and Central 
Germany, being rarer in the south, Denmark, and 
Scandinavia. According to Bedriaga, it crosses the 
Rhine only in Alsace, but occurs no farther west. It 
only just enters Holland. If we suppose the species 
to have originated in Central Europe, we should 
expect to find it in Switzerland, France, and perhaps 
England. If it had its ancestral home in Eastern 
Europe, we might expect it to occur on the Balkan 
peninsula. It seems to me more probable, therefore, 
that Rana arvalis came with the Siberian migration. 
This need not cause surprise, as the genus Rana is 
certainly not European. Out of about no species, 
only four are peculiar to Europe, the rest are scattered 
over all parts of the globe. Moreover, the fact that 
these four species are confined to Southern Europe 
would seem to indicate that the first species entered 
from the south, and there either became modified or 
spread over nearly the whole continent, as did, for 
instance, Rana esculenta and R. temporaries Neither 
of these is by any means confined to Europe. R. 
esculenta ranges right across the Asiatic continent to 
Japan, and also enters North Africa, while the other 
has a wide distribution in northern and temperate 

The various groups of Vertebrates are not dependent 
on each other in their migrations. Mammals and Birds 
extend their range with so much greater facility than 


Reptiles and Amphibians, that the surplus population 
of our neighbouring continents readily poured into 
Europe when owing to changes of climate perhaps 
they forsook their original homes. 

We observe much the same differences of origin in 
the various groups of European Invertebrates. The 
Central European Molluscan fauna, remarks Dr. 
Kobelt, had already developed from the pliocene 
in almost all its details, as regards formation of 
species and distribution when the Ice-Age com- 
menced (3, i. p. 162). Certain very interesting disloca- 
tions, however, in the range of land mollusca can be 
proved to have taken place about that time. Thus, 
as Dr. Kobelt has pointed out, the genus Zonites, 
which is now almost confined to the south-east of 
Europe, occurs in inter-glacial deposits in the valley 
of the Neckar, and even as far west as the Seine. If 
we might judge from this single instance, a molluscan 
migration from the east to the west seems to have 
occurred either in early or pre-glacial times. That 
Helix pomatia has migrated only comparatively 
recently from the East to Western Europe is 
rendered probable by its general range in northern 
and western Europe, but I cannot agree with Dr. 
Kobelt in the belief that Helix aspersa is of an 
equally recent origin in the North. No matter 
whether it has been found fossil or no, its range in 
the British Islands points to its having penetrated to 
Ireland when the latter was still connected with the 
Continent by way of England. Its migration from 


the Mediterranean dates therefore from early pleisto- 
cene or late pliocene times. 

In referring to the sixty-five species of Land and 
Freshwater Mollusca which have been described from 
the continental "Loess," Dr. Kobelt states (p. 166) 
that this fauna has certainly not a steppe-character. 
It does not therefore strengthen Professor Nehring's 
view that Europe during the deposition of the loess 
had a climate comparable to that of the Siberian 
steppes. The Glacial period had hardly any effect 
on the molluscan fauna of Europe. Dr. Kobelt 
believes in a certain movement of that fauna from 
the least favourable areas, with a subsequent 
re-immigration; but even that could not have 
taken place on a large scale. Nothing like a 
destruction of the fauna occurred, as far as we 
know from fossil evidence. 

Not a single species of land or freshwater mollusc 
can be quoted as having migrated to Europe from 
Siberia in recent geological times. The molluscan 
fauna of the latter country is so closely connected 
with that of Europe, that it is quite impossible to 
elevate it to the rank of a sub-region of the Holarctic 
Region. Dr. Kobelt insists that Siberia cannot even 
claim to be placed into a distinct province. Accord- 
ing to the same authority, we find no species in the 
whole Siberian molluscan fauna which we might 
regard as having immigrated since the close of the 
Glacial period. Even to attempt the location of 
the original homes of many of the species which 


Siberia has in common with Europe, seems hopeless. 
Such forms as Arion hortensis, which has been 
obtained in Siberia, and which, as we have seen, must 
have originated in Western Europe, migrated in 
pliocene or miocene times, possibly along the 
shores of the Mediterranean and across Asia Minor. 
We have evidence, therefore, of an eastward migration 
among the land and freshwater mollusca in later 
Tertiary times, but not of a westward one from 

A very different view is presented to us by the 
coleopterous fauna of Europe. Many of our Euro- 
pean Beetles are Siberian migrants. Let us take, 
for instance, the Tiger Beetles (Cicindelidce). There 
are over forty species of the genus Cicindela in 
Europe, five of which reach the British Islands. 
This seems a large number; but there are altogether 
no less than 6co species of the genus scattered over 
the greater part of the world, many of them being 
Asiatic. The genus is certainly not of European 
origin, for not only are most of the European species 
confined to the Caucasus and the south-east generally, 
but no Cicindelidce whatsoever occur, for example, in 
Madeira or the Canaries, where we should expect 
some to have persisted if the genus had originated 
on our continent. Moreover, of the five tribes into 
which the large family of Cicindelidce can be sub- 
divided, only two range to Europe, and one of 
them is represented by only a single species on our 


Some of the Cicindclas may have come with the 
Oriental migration. I think this was the case with 
the only Irish species of the genus, C. campestris. It 
occurs all over continental Europe and Northern Asia, 
and varieties of the species are known from Corsica, 
Sicily, Crete, the Cyclades, Sardinia, Asia Minor, 
Greece, and Spain. Five species of Cicindela^ as I 
said, are known from England, of which C. silvatica 
and C. maritima are certainly Siberian migrants, and 
perhaps C. hybrida too. Neither of the two first 
species is found in Southern Europe or in Spain, 
where we should expect them to occur had they 
originated on our continent. C- silvatica and 
maritima have no doubt entered Europe from 
Siberia in recent geological times, probably soon 
after a way was opened up across the Tchornosjem 
district of Southern Russia that is to say, in 
inter-glacial times. The former spread along the 
Central European plain as far west as the south-east 
of England when Great Britain still formed part of 
France. C. maritima y which preferred the proximity 
of the sea, migrated along the shores of the Caspian 
and then across Russia to the shores of the Baltic 
and North Sea, and has penetrated a little farther 
north and west in England than its near relative. 
C. litterata has a very similar distribution and origin, 
but instead of wandering so far west as the British 
Islands, it seems to have preferred extending its 
range southward, and has just reached Northern 


The closely allied Ground-beetles (Carabida) furnish 
us with equally interesting and instructive proofs of a 
migration from Asia. Over 300 species of Carabus 
are known to science. The number of species 
inhabiting Asia and Europe are about equal. But 
the genus does not extend its range to Southern 
Asia or to South America or Australia. Very 
few species enter Africa, and only nine North 
America, of which three also occur in Siberia. The 
genus is unknown in Madeira, and only represented 
by three species in the Canary Islands. To judge 
from its distribution, it has probably originated in 
Western Asia. Probably some Carabi of European 
origin have spread into Asia, but the Asiatic or 
we might say the Siberian origin and subsequent 
migration westward of a number of well-known forms 
appears to me evident. Such forms as C. clatkratus, 
C. granulatus, and C. cancellatus are no doubt of 
European origin, and have only in recent geological 
times extended their range across Northern Asia, 
whilst C. marginalis t coming fiom Siberia, can hardly 
be said to have invaded Europe, since it has never 
been met with farther west than the eastern provinces 
of Prussia. 

Among the Carabidtz there are altogether very 
many examples pointing to a migration from Asia to 
Europe, but I do not wish here to give a list of all 
such cases, and only refer to a few of the more, 
remarkable ones. One of the European species of 
Demetrias (D. unipunctatus), known to English ento- 


mologists as a south-eastern form, seems to have 
arrived with the Siberian migration, whilst the closely 
allied D. atricapillus, which has been able to reach 
Ireland, has a wider range and came earlier with the 

Messrs. Speyer state (p. 68) that almost all those 
species of Central European Butterflies whose 
northern limit is deflected southward as we approach 
the west coast of Europe, inhabit also the Volga 
country and the adjoining parts of Asia. Many of 
them are much commoner there than in Central 
Europe, and it appears probable to the authors of 
the Geographical Distribution of Butterflies that 
these species came from the east. Asia and Central 
Europe have, according to Messrs. Speyer, no fewer 
than 156 species in common. Mr. Petersen esti- 
mates that no less than 91 per cent, of the Arctic- 
European Butterflies also occur in Siberia. He 
made a special study of the Arctic Macro-lepi- 
doptera, and came to the conclusion that Central 
Asia, not having been glaciated in the Ice-Age, 
offered a possibility of existence to both animals 
and plants. Here, he thinks, was the principal centre 
to which Europe owed its re-population in post-glacial 
times. Mr. Petersen is of opinion (p. 40) that the 
Arctic-European Lepidoptera are composed of two 
elements the pliocene relics which persisted in 
Europe during the Glacial period, and the new 
immigrants from Siberia. 

No doubt Siberia supplied Europe with a number 


of species of Butterflies and Moths in recent geo- 
logical times, but we need not necessarily suppose 
that these arrived only after the Glacial period. Even 
the most extreme glacialists admit that large areas 
on our continent were free from ice at the height of 
the Ice-Age, Siberia had therefore no particular 
advantage over Europe in giving an asylum to 
Butterflies and Moths which were escaping from the 
rigours of a supposed arctic climate. But we have 
already learned (p. 80) that the climate during the 
Glacial period probably differed but little from that 
which we enjoy at the present day, and we may 
assume, therefore, that the Lepidoptera of Siberia 
migrated during that time or even earlier to 

Let us for a moment reconsider some instances of 
mammalian migration from Siberia, with a view to 
studying more closely the nature of these great 
events. I mentioned the fact that some of the 
Siberian migrants have remained in England, that 
more have settled down permanently on our con- 
tinent, but that many others have either become 
entirely extinct or do not live any longer in 

Of the mammals which made their appearance in 
Great Britain in recent geological times, t.e. y during 
and since the deposition of the Forest-Bed for example, 
the following species probably came direct from 
Siberia across the plains of Europe, as already men- 
tioned (p. 95): 


Canis lagopus. * Mus minutus. 

Gulo luscus. * Arvicola agrestis. 

* Mustela erminea. * amphibius. 

* putorius. arvalis. 

* vulgaris. * glareolus. 

* Sorex vulgaris. gregalis. 
Lagomys pusillus. ratticeps. 

* Castor fiber. Equus caballus. 
Spermophilus Eversmanni. Saiga tartarica. 

erythrogenoides. Ovibos moschatus. 

Cricetus songarus. Alces latifrons. 

Myodes lemmus. ,, machlis. 

Cuniculus torquatus. Rangifer tarandus. 

* Those marked with an asterisk still inhabit Great Britain, or did so 
within historic times. 

Of the arrival of many of these in Europe we have 
geological proof, as they have left their bones in 
recent pleistocene deposits, and are unknown from 
older European strata. The remote ancestors of 
others, such as Sorex and Lagomys, no doubt lived in 
Europe, but the recent species probably had their 
original homes in Asia. It is evident that in recent 
geological times there existed no active centre of 
origin for mammals in Europe, and that our continent 
was largely dependent on the neighbouring one for 
the supply of its mammalian fauna. A shifting of the 
centre of development from Europe to Asia appears 
to have taken place occasionally, as already men- 
tioned (p. 45). Mr. Lydekker has drawn attention 
to the fact that though the remote ancestors of the 
Elephantidcz resided in Europe, neither the latter 


continent nor North America was the home of the 
direct ancestor of any of the true Elephants. 
Similarly, though we have had our Sorex in Europe 
from the Upper Eocene and Lagomys from the 
Middle Miocene, the geographical distribution of 
Sorex vidgaris and Lagomys pusillus does not support 
the view that they are of European origin and have 
migrated to Asia. Their absence from most of 
the European islands indicates either an extremely 
recent origin or a recent immigration from Asia, 
and the latter view seems to me much the more 

No less than twenty-six species of the Siberian 
mammals penetrated as far west as the British Islands, 
and nine of these still inhabit Great Britain. Some 
of the remaining seventeen species probably lived only 
for a very short time in England, and the rest 
gradually became extinct one by one. This process 
of extinction of the aliens still continues. The Beaver 
(Castor fiber) has died out within recent historic times. 
We possess legends and uncertain historic records 
pointing to the existence of the Reindeer in Scotland 
as recently as about seven centuries ago. But much 
the same state of things has happened on the Con- 
tinent. The Glutton (Gulo hiscus), which still lived 
in Northern Germany last century, has now entirely 
vanished from that country, as also the Reindeer. 
The Lemmings have found an asylum in Scandinavia. 
The Musk-Ox (Ovibos moschatus) has disappeared 
not only from Europe but also from Asia, and is now 


confined to Arctic America and Greenland. The 
Horse no longer occurs in Europe in the wild state, 
and the Saiga Antelope (Saiga tartaricd) has retreated 
to the Steppes of Eastern Europe and Western 

As we proceed more and more eastward across 
Central Europe, we find that a larger and larger 
percentage of the Siberian migrants have adopted 
the new country as their permanent home, though in 
France and Germany, as well as in Austria, we have 
evidence that a great number of Siberian species, which 
formerly lived there, have either become entirely 
extinct, or have retreated towards the land of their 
origin. There is a prevalent belief that these migrants 
have taken refuge on the higher European mountain 
ranges, but this idea is altogether erroneous, as will 
be shown in the chapter dealing with the origin of the 
Alpine fauna. 

One of the Jerboas (Alactaga jaculus) occurs fossil 
as far west as Western Germany, but it is now con- 
fined to Russia and Western Siberia. The Bobak 
marmot (A rctoniys bobak\ which has a similar range 
now, probably inhabited France in former times. 
A Siberian species which has retreated but little is 
the Hamster (Cricetus vulgaris). Its fossil remains 
have been found in Central France, but it does not 
now occur west of the Vosges Mountains. 

It appears, therefore, as if a wave of migration had 
swept over Central Europe from east to west, that 
those species which were able to adapt themselves to 


the new surroundings had remained, and as if the rest 
had died out or were gradually retreating to the 

Ornithologists are well acquainted with the fact 
that in some years there is an unusually large exodus 
from Eastern Europe and Siberia of birds; and that 
species like the Waxwing (Ampelis garrulus) then 
appear in great numbers. But the appearance of this 
bird in Western Europe is not looked upon as so 
remarkable as that of Pallas's Sandgrouse (Syrrhaptes 
paradoxus, Fig. 3, p. 42), a typical inhabitant and resi- 
dent of the Arctic Steppes. The last great irruption 
took place in 1888, and many birds reached even the 
extreme west of Ireland in May and June of that 
year. A few weeks before, it had been announced to 
the German papers that large flocks of this peculiar 
pigeon-like bird had arrived in the eastern provinces; 
and though the vast majority vanished as quickly as 
they had come, a certain number remained for a year 
or so in the newly visited countries, and some even 
bred in England. 

Twenty-five years before, in 1863, a similar migra- 
tion had occurred, though not perhaps on quite such 
a vast scale, and a few small flocks had made their 
appearance in Western Europe on several occasions 
between these dates. 

It may not be generally known that no other bird 
has been honoured by our Government in a like 
manner, for it is the only animal for whose protec- 
tion a separate Act of Parliament has been passed. 


In spite of this unusual precaution, the species has 
not survived to add another member to the resident 
British fauna. The wave of migration from the east 
has come and vanished again just like so many others 
with which history is familiar. 

These migrations from the east occurring at the 
present day give us some idea of those of which we 
have fossil evidence, and which all had their origin 
in Central and Northern Asia. Almost all the species 
of mammals to which I have referred as being of 
Siberian origin have been found in the fossil state 
in comparatively recent geological deposits within a 
certain very limited area. None of the typical species 
have ever been found in Southern Europe proper, 
including the Mediterranean islands. It must be 
remembered that though the Reindeer is a Siberian 
migrant, the form of the Reindeer which was found 
in the Pyrenees belonged to a distinct variety in 
fact, to a much earlier migration which issued from 
the Arctic European Regions, and to which I have 
referred in detail (pp. 150-158). Curiously enough, 
no deposits of these typical Siberian mammals 
have ever been obtained in Scandinavia only in 
Russia, Austria, Switzerland (the lowlands), Germany, 
Belgium, France, and England. To facilitate a study 
of the extent of these migrations, I have constructed 
a map on which the probable course taken across 
Central Europe is roughly indicated by dots 
(Fig. 16). 

In the migrations of to-day we perceive the same 



tendency as in the older ones of which we have fossil 
evidence, viz., generally a spreading of species on a 
large scale over new territory, and then a gradual 

FIG. 1 6. Map of Europe. The doited portions represent, approxi- 
mately, the course of migration of the Siberian mammals. The 
principal mountain ranges are roughly indicated in black. 

shrinkage towards their original home, with an occa- 
sional survival of small colonies in the invaded part. 
It must not be supposed that this observation applies 


alone to the Siberian migration. In the case of the 
Arctic one, precisely the same thing has happened, 
and we shall see that the Southern (migration from 
the south) agrees in this respect with the others. 

As for the immediate cause of these migrations, it is 
to be looked for either in the scarcity of food dependent 
upon a temporary or permanent change of climate, or 
in an excessive increase in numbers of a particular 
species. I do not propose to trace back migrations 
beyond the Pliocene Epoch, or indeed much beyond 
the beginning of the Glacial period, which is regarded 
as a phase of the most recent geological epoch, viz., 
the Pleistocene. During the period in question, we 
have indirect evidence of one vast migration from 
Siberia into Europe across the lowlands lying to the 
north of the Caspian and to the south of the Ural 
Mountains. There is a general consensus of opinion 
that this migration took place in Pleistocene times. 
Professor Nehring thinks that there can be no doubt 
(p. 222) that the Siberian migrants arrived in 
Northern Germany after the first stage or division 
of the Glacial period, and lived there probably 
during the inter-glacial phase which occurred be- 
tween the first and second stages if indeed we look 
upon this period as being divisible into two distinct 

Judging from the evidence of distribution of 
mammals in pleistocene Europe, Professor Boyd 
Dawkins came to the conclusion (p. 113) that the 
climate of our continent "was severe in the north 


and warm in the south, while in the middle zone, 
comprising France, Germany, and the greater part 
of Britain, the winters were cold and the summers 
warm, as in Middle Asia and North America." " In 
the summer time the southern species would pass 
northwards, and in the winter time the northern 
would swing southwards, and thus occupy at different 
times of the year the same tract of ground, as is now 
the case with the Elks and Reindeer." Very different 
are the views of Professor Nehring on this subject. 
According to him, the climate in Germany must 
have been extremely cold and damp, resembling 
that of Greenland, though perhaps not quite so 
arctic. Professor Nehring does not at all believe 
that southern and northern species of mammals could 
have lived in Central or Northern Europe at the 
same time; though of this we have undoubted geo- 
logical evidence (pp. 72-75). He thinks that the 
supposed commingling of southern and northern 
types, which has actually been shown by Professor 
Dawkins to occur, is either due to careless observa- 
tion or to the fact that some of the species need not 
necessarily have lived where their bones were found 

(P- 133). 

The most reliable conclusions as regards former 

conditions of vegetation and climate can be drawn, 
according to Professor Nehring, from the smaller 
burrowing mammals, such as the marmots, sousliks, 
etc. He is of opinion that a great portion of 
Northern Europe, where their remains have been 


discovered, must have possessed tundras and steppes, 
as we find them nowadays in Siberia, and a climate 
similar to that of Northern Asia. It is presumed 
that the climate, after the maximum cold of the first 
stage of the Ice- Age, ameliorated so far as to permit 
these mammals to exist in Europe. 

The natural question, however, which is forced 
upon us in reading Professor Nehring's interesting 
and suggestive work is, where did all these steppe 
animals live during the earlier part of the Ice- Age? 
No traces of their remains have been discovered in 
Southern Europe, and it can therefore certainly be 
affirmed that they could not have lived there. If 
Central and Northern Europe were uninhabitable 
for mammals, Central and Northern Asia must have 
been even more so, and we have to fall back upon the 
Oriental Region as a possible home of these species 
during the assumed maximum cold of the Glacial 
period. In invading Europe from the Oriental 
Region these Siberian mammals would have taken 
the shorter route by Asia Minor and Greece, which 
was open to them. This they certainly did not do, 
which proves that they came directly from Siberia to 
Europe without retreating first to Southern Asia. 

But it seems to me that there is no necessity for 
assuming such drastic changes of climate to have 
taken place at all (compare pp. 75-80). We really 
have no idea under what precise climatic conditions 
the Siberian mammals lived in their original home. 
The only thing we can be certain of is that the 


smaller burrowing mammals would not have chosen 
a wood to live in, if they could possibly help it. 
Prairies, or sand-dunes with short grass or shrubs, 
such as abound in Europe near the sea-coast, would 
suit these species perfectly. If we suppose Northern 
Germany to have been covered by sea (p. 156) during 
part of the Pleistocene Epoch, forests would probably 
not have grown there for a very considerable time 
afterwards, owing to the excessive salinity of the soil, 
but a tract of sandy country would have been left 
on the retreat of the sea. Possibly a slight change 
of climate in the original home of these steppe-species 
may have reduced their habitable area, and thus 
caused their migration into Europe. 

But this migration problem cannot be solved 
without tracing the mammals to their place of 
origin and investigating their early history. This I 
shall attempt to do presently; meanwhile, it would 
be interesting to note whether other groups of 
animals support Professor Nehring's steppe-theory. 

Among groups other than mammals, the most 
important, for the purpose of drawing conclusions 
as to former physical conditions and climate, are the 
mollusca. Their remains have been well preserved, 
and are easily identified. Though Professor Nehring 
argues that the molluscs found along with the small 
mammals harmonise perfectly with the assumption 
of a steppe-climate (p. 212), I cannot at all agree 
with him. He enumerates the following sixteen 
species as having been discovered by him : 


1. Pupa muscorum. 9. Helix pulchella. 

2. Chondrula tridens. 10. Do. hortensis. 

3. Cionella lubrica. u. Do. obvoluta. 

4. Patula ruderata. 12. Hyalinia radiatula. 

5. Do. rotundata. 13. Succinea oblonga. 

6. Helix striata. 14. Limnaea peregra. 

7. Do. hispidia. 15. Clausilia sp. 

8. Do. tenuilabris. 16. Pisidium pusillum. 

Only two of these can be looked upon as typically 
northern species, viz., Patula ruderata and Helix 
tenuilabris^ though both of them are still found 
living locally in Germany. Some of the others 
are decidedly southern species, like Chondrula tridens, 
Helix obvoluta, H. rottmdata, and H. striata. All the 
rest live and flourish, for example, in Ireland at the 
present day, where, as we all know, anything but a 
dry steppe-climate prevails. 

Dr. Kobelt quite agrees with me in thinking that 
the remains of the mollusca found along with the 
so-called "steppe-mammals" afford no proof of a 
steppe-character of the country at the time when 
they were alive (p. 166). Nor do the mollusca which 
have been found in England in the Forest-Bed and 
the succeeding pleistocene strata support such a 
view. The Forest-Bed, generally regarded as belong- 
ing to the Upper Pliocene, I believe to be an inter- 
glacial pleistocene deposit contemporaneous with the 
loess formation in Germany. Of fifty-nine species 
of land and freshwater mollusca which have been 
discovered in this bed, forty-eight species, according 


to Mr. Clement Reid (p. 186), are at present living 
in Norfolk, six are extinct, two are continental 
forms living in the same latitudes as Norfolk, and the 
other three are all southern forms. Not a single 
species has a particularly northern range. Of the 
land and freshwater mollusca of the South of Eng- 
land in the succeeding pleistocene deposits, six 
species are now no longer living in the British 
Islands, but only one (Helix ruderata) can be looked 
upon as an Arctic or Alpine form. After this short 
digression on the mollusca, I will briefly recapitulate 
what is known about the early history of the Siberian 
mammals, which will assist us in tracing the cause of 
their migration to Europe. 

We have in Siberia problems quite as difficult of 
solution as the European ones. Volumes have been 
written to explain the former presence of Arctic 
mammals like the Reindeer in Southern Europe, and 
the most extraordinary demands on the credulity of 
the public have been made by some geologists in 
their attempts to account for this comparatively simple 
problem. In Northern Asia a somewhat similar 
phenomenon, but much more difficult of explanation, 
has taken place. Mammals have been found fossil in 
recent geological deposits in localities where they do 
not now occur, and apparently the Siberian and the 
European deposits are of about the same age. Now, 
however, comes the extraordinary difference. In 
Europe the Arctic mammals went southward, but 
in Siberia the Southern ones went northward. Not 


only do we find the Saiga-Antelope, Tiger, Wild 
Horse, European Bison, Mammoth, and Rhinoceros 
in the extreme north of the mainland of Siberia; their 
remains have even been obtained in the New Siberian 
Islands. As these islands are situated in the same 
latitude as the northern part of Novaya Zemlya, 
indeed, not far south of the latitude of Spitsbergen, 
the fact of such huge mammals having been able 
to find subsistence there at apparently quite a recent 
geological period seems an astounding fact. It may 
be urged that their bones might have been carried so 
far north by ice, or by some other equally powerful 
agency. But Tcherski and all other palaeontologists 
who have examined these northern deposits are 
unanimous in the belief that these herbivores and 
carnivores lived and died where their remains are 
now found. "It is evident," says Tcherski (p. 451), 
"that these large animals could only have lived in 
those extremely northern latitudes under correspond- 
ingly favourable conditions of the vegetation, viz., 
during the existence of forests, meadows, and 
steppes." He also is of opinion that the moist climate 
which evidently prevailed in Europe during Post- 
tertiary (Pleistocene) times must have modified the 
Siberian climate in so far as to render it milder. 
The existence of the Aralo-Caspian basin (Fig. 12, 
p. 156) must also have tended in the same direction. 
It appears then that, at the time when plants and 
animals are believed to have retired southward in 
Europe before the supposed advancing Scandinavian 


ice-sheet, no agency existed in North Siberia which 
was able to suppress and to annihilate the forest and 
meadow vegetation, and drive away the fauna con- 
nected with it. We know, continues Tcherski, that 
such genera as Bison, Colus (Saiga\ Rhinoceros, 
Elephas, and Equus are met with in all horizons 
of the diluvium of West Siberia. He therefore 
comes to the conclusion (p. 474), that these and 
other facts imply that the retreat of the North 
Asiatic fauna commenced about the end of the 
Tertiary Era (Pliocene), and that it was continued 
very slowly throughout the Post-tertiary (Pleistocene) 
Epoch, without any visible changes in its southward 
direction, even during the time of the most important 
glacial developments in Northern Europe. Only after 
the conditions disappeared which had produced the 
augmentation of an atmospheric moisture, did the 
climate of North Siberia become deadly to a 
temperate fauna and flora. Tundras then spread 
over the meadow-lands and remnants of forests, 
whilst arctic animals replaced the large ungulates 
and carnivores which had wandered far away from 
their native southern home. 

This is Tcherski's explanation of the extraordinary 
events which he has chronicled, after years of the 
most arduous labour and under conditions of peculiar 
hardship. And though his work cannot be over- 
estimated, and his opinions should receive the most 
careful consideration, yet I fear the explanation will 
not be looked upon as entirely satisfactory. Every one 


will agree with him that the climate of Siberia must 
have been greatly moister in pliocene and pleistocene 
times than it is now. The Aralo- Caspian covered 
a vast area of South-western Siberia. Freshwater 
basins existed along the east of the Ural Mountains, 
while Central Asia was studded over with a number of 
large lakes, which have now almost entirely vanished. 
But that the generally assumed refrigeration of 
Europe must have had a chilling effect on the 
Siberian atmosphere seems to me evident. That the 
whole of Northern Europe should have been made 
uninhabitable owing to the advance of the Scandi- 
navian ice-sheet, while North Siberia at the same 
time supported forests, meadows, and a temperate 
fauna, is incredible. At the approach of winter, at 
any rate, the animals would have been driven south- 
ward for thousands of miles to seek shelter from the 
snows and cold and to obtain nourishment, and it 
would scarcely have been possible for them to 
undertake such vast migrations at every season. 
Professor James Geikie's suggestion (p. 706), that 
the Mammoth and Woolly Rhinoceros could have 
survived the Pleistocene Epoch in Southern Siberia, 
does not appear to solve the problem, as that part 
of Asia must have participated in the great cold 
which is said to have prevailed all over Europe. 

Let us now concede, for the sake of argument, 
that the current views regarding the pleistocene 
climate of Europe are correct. We are told by Pro- 
fessor Geikie that the climate of Scotland during 


part of the Pleistocene Epoch was so cold, that the 
whole country was buried underneath one immense 
mer de glace, through which peered only the higher 
mountain-tops (p. 67). If this was the state of climate 
in close proximity to the Atlantic, it must probably 
have been still more severe on the European con- 
tinent. Now at the present time Siberia has the 
reputation of being the coldest country in the world, 
and the mercury of the thermometer is said to remain 
frozen for weeks during winter, even in the south. 

With the prevailing dampness in pleistocene times 
the snowfall throughout Siberia would have been 
much heavier than at present, though it would have 
modified the temperature to some extent. Under 
such circumstances Southern Siberia could not have 
been a desirable place of residence for large mammals. 
It would have been necessary for the Mammoth and 
the other species referred to, to wander farther into 
the extreme south of Asia or Europe to find a 
suitable refuge during the arctic conditions which 
are supposed to have prevailed in Northern Europe. 
To quote Professor J. Geikie's own words (p. 706): 
" They (Mammoth, etc.) would seem to have lived in 
Southern Siberia throughout the whole Pleistocene 
period, from which region doubtless they originally in- 
vaded our Continent. But with the approach of our 
genial forest-epoch (penultimate inter-glacial stage) 
they gradually vanished from Europe, to linger for a 
long time in Siberia before they finally died out." 
It is suggested, therefore, by the author that the 


Mammoth and the other mammals whose remains 
have been discovered on the New Siberian Islands 
found their way there during one of the late inter- 
glacial stages of the Ice-Age. But there is no 
astonishment expressed by Professor Geikie at the 
extraordinary change of climate which must have 
occurred in Siberia to allow of such migrations. I 
can find no very definite statement in this author's 
work as to the nature of the climate in Europe during 
those inter-glacial phases, but he remarks (p. 129) 
" that the evidence of the Scottish inter-glacial beds, 
so far as it went, did not entitle us to infer that 
during their accumulation local glaciers may not 
have existed in the Highland valleys." There is no 
evidence, in other words, of the existence in Europe 
of a milder climate than that prevailing at present. 
Still less can there be any ground for the supposition 
that the climate of the whole of Siberia ameliorated 
to such an extent that forests and meadows could 
develop as far north as the New Siberian Islands ; 
for if the temperature in Europe was then about 
the same as now, that of Siberia could not have 
been vastly higher than it is at present. 

It is highly improbable, therefore, that a sufficiently 
mild climate prevailed in the extreme north of 
Siberia during the so-called later inter-glacial periods 
to induce the mammals to which I have referred to 
seek fresh pastures there, 

The late Professor Brandt, one of the highest 
zoological authorities in Russia, was of opinion that 


at the commencement of the Glacial period the great 
mammals of Northern Siberia either perished or 
migrated southward. From there they gradually 
penetrated into European Russia. He believed that 
before the Glacial period a connection qxisted 
between the Aralo-Caspian Sea and the Arctic 
Ocean, carrying warm water northward. The gradual 
disappearance of this marine channel caused a 
decrease of warmth in Northern Asia, so that large 
accumulations of frozen soil and ice were formed, 
which still more depressed the temperature. This, 
he suggested, probably took place at the time when 
the Glacial period commenced in North-western 

It has been urged against these views of Tcherski 
and Brandt, that the bone beds in the Liakov Islands 
(New Siberian Islands) rest partly upon a solid layer 
of ice of nearly seventy feet thick. This mass of ice, 
it was thought, must have accumulated during the 
Glacial period. As the bones rest upon it, the 
mammals could only have lived in those islands in 
more recent times, after the Ice- Age had passed away. 
Nothing, apparently, can be clearer, and yet in the 
face of this seeming proof one feels, as I have men- 
tioned before, that if such an extraordinary revolution 
of climate as is implied by this admission had taken 
place, we should be able to perceive the traces 
throughout the northern hemisphere. In this di- 
lemma, a suggestion made by Dr. Bunge, who visited 
the New Siberian Islands recently at the instance of 


the Imperial Academy of St. Petersburg, helps 
us out of the difficulty. He found that, as a 
rule, these so-called fossil glaciers contain seams 
of mud and sand, and he argued that the ice had 
formed, and is still forming at the present day, in 
fissures of the earth. I entirely concur with this 
view. Neither palaeontology nor the geographical 
distribution of animals lend any support to the other 
theory, and I think we may conclude that Brandt's 
view in the main is probably the correct explanation 
of the phenomena which we have discussed. Some 
important facts of distribution are more easily explic- 
able on this assumption. Why, for instance, should 
the Siberian fauna of pliocene times have remained 
in Siberia and not have migrated to Europe at that 
time? The pliocene mammals of Siberia are mostly 
of southern origin. Their range increased enormously 
during the epoch throughout Northern Asia. We 
should expect them, therefore, to have crossed the 
Caspian plains, or even the low-lying Ural Mountains, 
to pour into the neighbouring continent. But Pro- 
fessor Brandt explained how they were prevented 
from spreading west. An arm of the sea stretched 
from the Aralo-Caspian to the Arctic Ocean, thus 
raising an effectual barrier between the two con- 
tinents. There is some evidence for the belief, as we 
shall learn presently, that this marine barrier existed 
also during the early part of the pleistocene epoch, 
After having greatly expanded during pliocene times, 
the fauna of Siberia gradually withdrew from the 


northern regions during the earlier portion of the 
succeeding epoch. It was only after the marine 
connection above referred to ceased to exist, or 
became disconnected, that an entry into Europe 
was possible. 

A fauna, to some extent composed of species now 
inhabiting the steppes of Eastern Europe and Siberia, 
poured into the neighbouring continent. On p. 95 I 
have given a list of those which reached as far west 
as the British Islands, but, as I mentioned, many 
other species came from the east about this time. 
With regard to the early history of the Siberian 
mammals, I favour a view somewhat between that 
of Tcherski and that of Brandt. The outpouring of 
the fauna into Europe seems to me to indicate that 
there was a sudden change of climate in Siberia. 
This was produced, perhaps, by the rupture of the 
marine connection between the Arctic Ocean and 
the Aralo-Caspian. Such an event would not only 
have caused the sudden shrinkage of the area avail- 
able for food-supply by lowering the temperature in 
Siberia, it would have acted also as a means in 
assisting the fauna to enter a new continent where 
an inconsiderable number of mammals, already estab- 
lished, were mostly dispossessed of their homes by the 
advancing eastern host. 

Brandt's theory, however, of a marine connection 
between the Arctic Ocean and the Aralo-Caspian is 
by no means generally accepted. That the Caspian 
Sea was at that time greatly larger than it is at 


present, and joined to the Sea of Aral and the Black 
Sea, is acknowledged by everybody. That the 
deposits laid down by this huge inland sea reach 
as far north as the shores of the river Kama, in 
Central Russia, is also well known to geologists. 
But what comes rather as a surprise, is that Professor 
Karpinski, whom we must take as one of the highest 
authorities on the geology of Russia, asserts that this 
Aralo-Caspian Sea was probably joined by a system 
of lakes or channels to the Arctic Ocean (p. 183). He 
was by no means the first, though, to put forward 
such a theory. We have already learned that Professor 
Brandt held a somewhat similar view, though he 
believed in something more than a connection by 
mere channels, and Mr. Koppen, and also the Russian 
traveller Mr. Kessler, agreed with him. So much 
was Professor Boyd Dawkins impressed with their 
arguments at the time, that he wrote (c, p. 148) : 
" Before the lowering of the temperature in Central 
Europe, the sea had already rolled through the low 
country of Russia, from the Caspian to the White 
Sea and the Baltic, and formed a barrier to western 
migration to the Arctic mammals of Asia" 

In one particular Professor Dawkins's views differ 
from those of almost all the previous writers. His 
connection between the Caspian and the Arctic 
Ocean is placed to the west of the Ural Mountains, 
while it had always been assumed by the Russian 
writers to have lain on the eastern or Asiatic side of 
that mountain range. Thus, when Tcherski in recent 


years announced that the tract on this eastern side of 
the mountains was covered by freshwater deposits, 
his discovery seemed once for all to settle the problem 
of the arctic marine connection in the negative. As 
Professor Dawkins's theory has, however, received 
much additional affirmative evidence by current 
faunal researches, a connection between the Caspian 
(or Aralo-Caspian) and the Arctic Ocean (White Sea) 
may have actually existed within recent geological 

What relict lakes are, has already been explained 
(p. 176), and their fauna will again be referred to in 
a subsequent chapter. I might perhaps be allowed 
to repeat that such lakes are supposed to have been 
flooded by, or to have been in close connection with, 
the sea at some former period. Many of the Swedish 
lakes are spoken of as relict lakes (Reliktenseen), 
because they contain a number of marine species of 
animals which have now become adapted to live in 
fresh water, but all of whose nearest relatives inhabit 
the sea. One of these, the schizopod crustacean Mysis 
relicta, a shrimp-like creature, which was formerly 
believed to inhabit also the Caspian, is of particular 
interest. More recently, the occurrence of this Mysis 
in the Caspian was denied, but though this denial has 
been confirmed by Professor Sars in his memoir on 
the crustaceans of the great Russian inland sea, he 
has been enabled to add two new species of Mysis 
to the list of those already known to science. These 
are M. caspia and M. micropthalma, and both are 


closely related to the arctic marine Mysis oculata. 
According to Professor Sars, the genus Mysis as a 
whole may be regarded as arctic in character. The 
occurrence of these two species, therefore, in his 
opinion, points to a recent connection of the Caspian 
with the Glacial Sea. 

A large number of other crustaceans have been 
described by the same author from the Caspian. Of 
the order Cumacea, which is exclusively marine, 
ten species are mentioned, but none of these seems 
to range beyond the Caspian. Among the smaller 
species of crustaceans, a minute pelagic copepod 
(Limnocalanus grimaldii) also inhabits the Baltic 
and the Arctic Ocean. The marine isopod Idctea 
entomon, related to the common wood-louse, has a 
similar distribution. 

Genuine Arctic species of Fishes do not seem to 
occur in the Caspian, though some, viz., Clupea 
caspia y Atherina pontica, Clupionella Grimmi, and 
Syngnathus bucctilentus, are almost certainly the 
descendants of marine forms. 

The Seal of the Caspian (Phoca caspica) is closely 
allied to the Arctic Seal, and its presence alone in 
that sea indicates that at no very distant date at 
any rate since pliocene times a closer connection 
with the Arctic Ocean existed than at present. 

I am sure it will be readily granted that there is 
zoological evidence for the belief of such a connection 
or union between the two great seas. However, it 
may be urged that owing to the presence of an ice- 



sheet in Northern Europe during the Glacial period, 
such a connection must either have been pre-glacial 
or have existed after that period. But the connec- 
tion must have occurred at a time when the Caspian 

FIG. 17. Map of European Russia (after Karpinski). The faintly 
dotted parts indicate the areas covered by boulder-clay, the 
strongly dotted ones those exhibiting Aralo-Caspian and other 
post-pliocene deposits. 

extended far to the north when indeed the so-called 
post-tertiary Caspian deposits were laid down (Fig. 
17). Since the boulder-ciay which covers the plain 
of Northern Russia is assumed to be the ground- 



moraine of the great northern ice-sheet, we might 
expect to find that the Caspian deposits were not 
contemporaneous with it. Curiously enough, it has 
been shown by Mr. Sjogren that all observations 
have pointed to the fact that these two deposits do 
not overlie one another, but occur side by side, 
and are therefore contemporaneous. This seems to 
warrant our belief, that while the boulder-clay was 
being laid down in Northern Europe, the Aralo- 
Caspian Sea had some communication with the 
White Sea. 

The boulder-clay of Northern Continental Europe, 
as already stated, is now generally recognised to 
be the product of a huge ice-sheet which invaded 
the lowlands of Continental Europe from the Scan- 
dinavian mountains. Though Alpine glaciers at the 
present day produce little or no ground moraine, 
these ancient larger ice-sheets, or "mers-de-glace," are 
believed to have deposited immense layers of mud 
containing scratched and polished stones. Many of 
the latter have been carried great distances from 
their source of origin. The Scandinavian ice-sheet 
is supposed to have advanced as far south as the 
line indicated on the map, after which it gradually 
retreated. On this point, however, as in almost every 
detail connected with the Glacial period, geologists 
are at variance. Professor James Geikie maintains, 
that there were no less than four Glacial periods, 
separated from one another by milder inter-glacial 
phases. On the Continent the view of two Glacial 


and one inter-glacial period is, I think, more gene- 
rally adopted. Professor Geikie's four periods seem 
to me to have originated i-n a desire to correlate 
the British pleistocene deposits with the continental 
ones, and at the same time to retain the old view of 
the inter-glacial position of the Forest-Bed. The 
two theories agree in so far as that in both the 
glacial conditions culminate in a maximum glacia- 
tion, followed by a more temperate phase of climate, 
with consequent retreat of the ice-sheets, and finally 
by a renewed advance of the glaciers. 

We are told that there is not the slightest doubt 
about it that a marked but gradual decrease of 
temperature took place all over Europe either 
during the beginning of the Pleistocene or towards 
the end of the Pliocene Epoch. 

We might reasonably suppose, then, that a similar 
climatic effect was produced in Siberia, in con- 
sequence of which the fauna would have been 
obliged to retreat from the extreme northern lati- 
tudes southward. No doubt great efforts would 
have been made by the members of the Siberian 
fauna at any rate by those possessing strong 
power of locomotion to extend their range in 
other directions. But we have no evidence that 
a migration from Siberia came to Eastern Europe 
at that time. It seems, therefore, as if the barrier 
referred to by Brandt, Koppen, Boyd Dawkins, 
and others (p. 222), had existed at this time. This 
would have effectually prevented an overflow of 


the fauna from Siberia. Only in deposits later 
than the lower continental boulder-clay do we find 
traces of a Siberian migration. The time of maxi- 
mum glaciation had then passed away ; the great 
glacier which was believed to have invaded the 
lowlands of Northern Europe had again retreated, 
before the Siberian mammals made their appearance 
in Germany. 

It has been stated above (p. 226) that while the 
Russian boulder-clay was being laid down, the Aralo- 
Caspian probably had some communication with the 
White Sea. 

But how can this view be reconciled with the 
existence of a huge mer de glace in the northern 
plains of Russia? The existence of the ice-sheet 
has been conjured up in order to explain the 
presence of the boulder-clay. But not long ago a 
very different interpretation of the origin of this clay 
was given; and one, I may say, which explains the 
history of the Siberian and the European fauna in a 
more satisfactory manner than is done by the ice- 
sheet hypothesis. It is that the boulder-clay is not 
the product of land-ice, but has been deposited by a 
sea with floating icebergs. Thus the latter hypothesis 
does not deny the existence of glaciers, but allows 
.the mud to be deposited on the floor of a turbid 
sea, instead of beneath an immense mer de glace. I 
need hardly mention that this view, which was 
formerly universally accepted by geologists, is now 
scouted by almost every authority, both British and 


Continental. I should scarcely venture the attempt 
to revive old memories and stir up again long 
forgotten controversies, were it not for the fact that 
many new points have arisen in the course of the 
above inquiries, which appear to me so very difficult 
to explain by the land-ice hypothesis, while they are 
comparatively easy to understand when we adopt 
the old theory of the marine origin of the boulder- 
clay. But a few geologists even at the present day, 
while believing in the land-ice theory, recognise that 
the marine hypothesis should have some considera- 
tion shown to it. I need only remind glacialists of the 
work recently published by Professor Bonney. " The 
singular mixture," he remarks (p. 280), "and apparent 
crossing of the paths of boulders, as already stated, 
are less difficult to explain on the hypothesis of 
distribution by floating ice than on that of transport 
by land-ice, because, in the former case, though the 
drift of winds and currents would be generally in one 
direction, both might be varied at particular seasons. 
So far as concerns the distribution and thickness of 
the glacial deposits, there is not much to choose 
between either hypothesis; but on that of land-ice 
it is extremely difficult to explain the intercalation of 
perfectly stratified sands and gravels and of boulder- 
clay, as well as the not infrequent signs of bedding 
in the latter." 

Now with regard to the land -ice theory, several 
serious difficulties present themselves in connection 
with the origin of the European fauna. In the first 


place, as the climate renders Northern Siberia 
almost uninhabitable for mammals at the present 
day, how much more severe must it have been 
during the time of the maximum glaciation in 
Europe. As the then existing fauna was not 
driven into Europe, where could it possibly have 
survived ? Secondly, how can we reconcile the 
contemporaneous existence of a great inland sea 
(the Aralo-Caspian) containing survivals of mild 
Sarmatic times with an immense glacier almost 
touching it on its northern shores? How did 
one of the most characteristic species of that sea, 
Dreyssensia polymorpha, come to make its appearance 
in the lower boulder-clay of Prussia and then dis- 
appear in the upper? And finally, how are we to 
explain the sudden appearance of a Siberian fauna 
after the deposition of the lower boulder-clay, except 
by the removal of a barrier which had prevented 
their egress from Siberia? 

If we assume that the continental boulder-clay of 
Russia has been formed in the manner so ably 
explained by Murchison, de Verneuil, and von Key- 
serling, viz., by a sea with floating icebergs, the 
temperature of Siberia might have been higher than 
at present, and have supported a fauna in more 
northern latitudes. 

The contemporaneousness of the deposits of this 
sea with those of the Aralo-Caspian is also rendered 
more intelligible. If we suppose, moreover, the con- 
nection between the Aralo-Caspian and the White 


Sea (Fig. 12, p. 156) to have existed at this time, we 
possess an explanation of the method of migration of 
the Arctic marine species into the Southern and of 
the Caspian species (Dreyssensia) into the Northern 

An inter-glacial phase is believed to have super- 
vened after the deposition of the lower boulder-clay, 
and it is during this period that the Siberian species 
first appeared in Central Europe. If we assume then 
that the retreat of the Northern Sea (Fig. 13, p. 170) 
opened up a passage for the Siberian fauna, we have in 
this very fact also an explanation of the extraordinarily 
large exodus of Asiatic "mammals, because the great 
reduction of the marine area in Northern Europe 
would have had an important influence in lowering 
the temperature in Asia. Only a sudden change of 
climate in Siberia could have brought about the 
migration of the vast hordes of Asiatic mammals 
whose remains we find in Central and Western 
Europe in deposits of that period. 

Throughout this work we are made acquainted 
with facts which bear out the view that the climate 
during the greater part of the Glacial period was 
mild rather than intensely arctic in Europe. That 
a huge ice-sheet could have covered Northern Europe 
under such conditions appears to me very doubt- 
ful. No one can deny, however, that glaciers must 
have existed during the Glacial period in all the 
mountainous regions of Central and Northern 
Europe, though their existence is not incompatible 


with a mild climate. Tree-ferns and other tropical 
vegetation grow at the foot of glaciers in New 
Zealand. We need not even go so far afield, for in 
Switzerland grapes ripen and an abundant fauna and 
flora thrive in close proximity to some of the well- 
known glaciers. 

One matter of importance still remains to be con- 
sidered before concluding this chapter, viz., the 
fauna contained in the English geological deposit 
known as the " Forest-Bed." This interesting 
deposit is exposed at the base of a range of cliffs 
on the coast of Norfolk. It is composed of beds 
of estuarine and marine origin. The tree-stumps 
formerly believed to be the remains of trees in situ 
have, after more careful examination, proved to be 
in all cases drifted specimens. A portion of the 
"Forest-Bed" no doubt was laid down in close proxi- 
mity to a large river, and subject to being periodi- 
cally flooded by it. It is not absolutely certain, there- 
fore, that all the mammals whose remains occur in 
this deposit lived in England or whether only on the 
banks of the river farther south. Nevertheless, we 
may take for granted that some of them did. 
England was at the time connected with France 
and Belgium, and for our purpose it matters little 
whether they had crossed the Channel or inhabited 
those parts of the Continent through which the great 
river flowed which sent its alluvial detritus as far as 
the plains of Norfolk. All we have to remember 
is that certain mammals, which appear to have 


originated in Siberia, and of which we have some 
evidence that they crossed Central Europe in their 
westward course, had now reached the great river 
just alluded to, which some geologists believe to have 
been the Rhine. 

I have had occasion to refer to a number of British 
mammals (p. 202) some of which are now extinct 
which I believe to have migrated across the plains of 
continental Europe direct from Siberia. There were 
twenty-six species of these Siberian mammals; and no 
less than ten of these occur in the Forest-Bed. None 
appear in any older British deposit. It is perfectly 
clear, therefore, that the Forest-Bed must have been 
laid down after their immigration into Europe. They 
probably wandered to Western Europe very soon 
after crossing the eastern boundaries of our conti- 
nent ; the deposits in which they are found are there- 
fore contemporaneous. But we have learned above 
(p. 208), that the beds in Eastern Europe in which the 
Siberian mammal-remains are found are more recent 
than the lower boulder-clay. As already stated, the 
Forest-Bed must also be more recent than the lower 
continental boulder-clay, and should be included in 
the pleistocene series. 

That the Forest-Bed is an inter-glacial deposit has 
been urged long ago by various writers. Professor 
Geikie regards it as stratigraphically contempor- 
aneous with the peat and freshwater beds below the 
lower diluvium of Western and Middle Germany, and 
as having been laid down during the first Inter-glacial 


Epoch of the great Ice- Age. The fact that no 
boulder-clay underlies the Forest-Bed seems rather 
a strong argument against the view of its being an 
inter-glacial deposit. It lies directly on what is 
known as the Newer Pliocene Crags. If the Forest- 
Bed is included in the pleistocene series, as I sug- 
gested it should, the crags, or a portion of them, 
would therefore be equivalent as regards time of 
deposition to the lower continental boulder-clay. 
And again, if the lower continental boulder-clay is 
contemporaneous with the Newer Crags, the latter 
should also be classed with the pleistocene strata. I 
can scarcely hope that geologists will be ready to 
admit such a sweeping change of nomenclature 
without a protest. I venture, therefore, to explain 
more fully my reasons for adhering to these un- 
orthodox views. 

Let us look once more at the map which I con- 
structed (Fig. 12, p. 156) to elucidate the migration 
of the Arctic terrestrial species to the British Islands. 
It will be noticed that one continuous ocean extends 
from the east coast of England across Holland, 
Northern Germany, and Russia to the White Sea. 
At the same time Greenland and Northern Scandi- 
navia, Scotland and Southern Scandinavia, are united 
by a narrow strip of land, and so are England and 
France. The waters of the Atlantic and this North 
European Sea do not therefore intermingle at any 
point, the two seas being absolutely independent of 
one another. 


Such I assume to have been the geographical con- 
dition of Northern Europe during the deposition of 
the Red Crag. Arctic mollusca were then brought 
to the east coast of England, and boulders were 
scattered through the beds laid down on that coast 
by icebergs which had been cast off by Scandinavian 
glaciers on reaching the sea. Bedded clays which 
have yielded arctic shells He beneath the lower con- 
tinental boulder-clay on the Baltic coast-lands and on 
the coast of the White Sea. According to Professor 
Geikie, marine clays on the same geological horizon 
reach an elevation of some 230 feet. " It would seem, 
then," he says, "that before the deposition of the 
lower boulder-clay of those regions the Baltic Sea 
had open communication with the German Ocean " 
(p. 442). All these clays are evidently deposits of the 
same sea. But apart from the fact that the Red Crag 
and these Baltic deposits are the oldest of the upper 
Tertiary beds containing arctic shells, there is no 
evidence that they are contemporaneous. 

Overlying the same Baltic deposits comes the lower 
boulder-clay, reaching a thickness of several hundred 
feet in some parts of Germany. It presents, like the 
upper clay, frequent interstratification with well- 
bedded deposits of sand and gravel. The scarcity of 
marine mollusca, the occurrence of striated surfaces, 
and the occasional presence of so-called giants' 
kettles, appear to favour the view, which at present 
is generally adopted by both British and Continental 
geologists, that the boulder-clay owes its origin to 


land-ice. I have stated on several occasions that the 
view of the marine origin of the boulder-clay agrees 
best with the known facts of distribution, and with 
the history of the European fauna (pp. 80-86, and 
p. 129). It may be urged that if the lower boulder- 
clay were contemporaneous with the British Crags 
which succeeded the Red Crag, how can we explain 
the fact that these crags contain plenty of shells, 
while in the lower continental boulder-clay there 
are scarcely any ? 

But as yet our knowledge of the conditions of life 
of the marine mollusca and of their distribution is 
extremely scanty. We are apt to imagine that the 
bottom of the sea is covered by a more or less 
uniform thick layer of shells; but whenever a careful 
survey of the nature of the deposits now forming 
there has been made, such is by no means found to 
be the case. Some of the best results obtained by 
that useful body, the Liverpool Marine Biological 
Committee, have been precisely in this direction. 
A most interesting account has been published by 
Professor Herd man and Mr. Lomas on the floor 
deposits of the Irish Sea, in which the authors state 
(p. 217), that "a place may be swarming with life 
and yet leave no trace of anything capable of being 
preserved in the fossil state, whereas in other places, 
barren of living things, banks of drifted and dead 
shells may be found, and remain as a permanent 
deposit on the ocean floor." 

Owing to the fact of the peculiar geographical 


position of Scandinavia at this time an isthmus of 
land with a high mountain range lying between the 
warm Atlantic and the cold Arctic Sea the snowfall 
must have been excessive, and large glaciers were 
evidently forming. These produced icebergs as soon 
as the lower parts had advanced to the Baltic coast- 
land and deposited their detritus in the sea. Immense 
masses of mud and stones were thus cast to the 
bottom of the sea, and under these circumstances no 
delicate mollusca or other marine life probably could 
have developed within a considerable distance from 
the shore. To judge from the direction pursued by 
the majority of the boulders from their source of 
origin, the prevailing current during the deposition of 
the lower boulder-clay was from north-west to south- 
east. It is possible that little marine life, except 
free-swimming forms, would have been able to live 
within the Russian area of this sea. But the free- 
swimming larvae of molluscs and other surface 
species were not prevented from passing from the 
White Sea south-westward, and in sheltered localities 
where little or no mud deposition was going on, 
these no doubt might have developed into adults 
on the sea-floor. It is quite conceivable, therefore, 
that in one portion of the North European Sea, which 
was fully exposed to the destructive influences of the 
iceberg action, the fauna was scanty or totally absent, 
while in another part there lived a fairly abundant 
one. The unfossiliferous state of the lower continental 
boulder-clay does not, therefore, offer any serious 


difficulty to the supposition that some of the so- 
called Newer Pliocene Crags of the east coast of 
England were laid down at the same time by the 
same sea. 

This would also explain how the Arctic species 
come to inhabit the Caspian, as the old Aralo-Caspian 
Sea could have had some communication (Fig. 12, 
p. 156) with the North European Sea. And this again 
offers an explanation of the otherwise mysterious 
occurrence of the Caspian Dreyssensia polymorpJia 
in the lower continental boulder-clay. 

The climatic reasons for the supposition that the 
boulder-clay is a marine deposit have already been 
given (p. 66). However, it may be asked what about 
the glacial flora which has been proved to have existed 
all over the plains of Northern Europe ? what about 
the relics of this same flora which still linger on in a 
few localities to the great delight of the systematic 
botanist ? They have been spoken of as indications of 
a former Arctic climate in Europe. The presence of 
an Arctic species such as Dryas octopetala in any of 
the pleistocene deposits is often looked upon as an 
absolute proof of a very severe climate having pre- 
vailed at the time they were laid down. Professor 
Geikie tells us that the South of England was 
clothed with an Arctic flora, when the climate be- 
came somewhat less severe than it had been during 
the climax of the glacial cold (p. 398). Relics of 
such a flora have been detected at Bovey Tracey, in 
Devonshire, the Arctic plants found comprising 


Betnla nana and B. alba, Salix cinerea and Arctosta- 
phylos uva-ursi. 

Now three of these four species of plants are still 
natives in the British Islands, and all are forms 
which probably came to us with the Arctic migration 
which I described in Chapter IV. They travelled 
south with the reindeer, or before it, and may have 
covered large tracts of country at the time. With the 
increased struggle for existence on the arrival of the 
Siberian and Oriental migrants, they have probably 
been evicted by these more powerful rivals. A 
discovery of their remains does not necessarily 
indicate that a great change of climate has taken 
place since they lived in the country. And certainly 
these Arctic plants cannot be taken as indicating a 
low temperature, for it has been shown that Alpine 
plants are mostly intolerant of very low temperatures. 
" Arctic and Alpine species in the Botanical Gardens 
at Christiania," says Professor Blytt (p. 19), "endure 
the strongest summer heat without injury, while they 
arc often destroyed when not sufficiently covered 
during the winter." Similar observations have been 
made in other countries. For this reason they have 
to be generally wintered in frames in the Botanic 
Gardens at Kew and Dublin, and are thus exposed to 
higher temperatures than at present obtain in the 
British Islands. This fact suggests that the Alpine 
and Arctic plants really did not originate in countries 
with cold temperatures. They probably made their 
first appearance long before the Glacial period 


perhaps in early Tertiary times chiefly in the Arctic 
Regions, which at that time had a mild climate. 
They have since become adapted to live in cold 
countries where they flourish, provided they receive 
sufficient moisture in the summer, and are protected 
from severe frost in the winter by a covering of snow. 

When we carefully examine the present range 
of Arctic plants in the British Islands, a curious 
fact presents itself which no doubt has frequently 
been noted by botanists, viz., that some of the most 
characteristically Arctic species, and some which 
are often quoted by glacialists in support of their 
theories, flourish at the present moment in very 
mild situations. I have already referred to the fact 
that the Mountain Avens (Dryas octopetala) abounds 
in the west of Ireland (County Galway) down to sea- 
level. Now it is well known that the mean winter 
temperature of that part of Ireland resembles that 
of Southern Europe, being no less than 12 F. 
( = 7 Cent.) above freezing point. The plant, of 
course, is here a native, and not introduced. This 
instance shows clearly, that as long as more vigorous 
competitors are absent, and as long as it is not 
exposed to severe frost or undue dryness, this and 
allied species do just as well in a mild climate as in 
their native Arctic home. 

In his interesting essay on the distribution of the 
Arctic plants in Europe during the Glacial period, 
Professor Nathorst adduces the fact that all the 
localities but one, in which remains of such plants 


have been discovered, lie either within or close to 
the limits of the maximum extension of the supposed 
northern ice-sheet, or within those of the former 
Alpine glaciers. Whether we look upon the boulder- 
clay as a marine or a terrestrial product, it is quite 
conceivable that, in many instances, the remains of 
the Arctic plants may have been carried by ice to 
great distances from where they grew. The prob- 
ability, however, is in favour of most of them 
having lived where their remains are now found. 
Now, it is a remarkable fact, that the single instance 
in Europe of a deposit of Arctic plants having been 
found far removed from the maximum extension 
of the northern ice-sheet is the one quoted above, 
viz., at Bovey Tracey, in Devonshire. Even up 
to recent times Arctic plants may have persisted at 
Bovey Tracey just as they do in Gal way under the 
influence of a mild coast climate. Similar circum- 
stances may have led to their survival along the 
shores of the sea which deposited the North 
European boulder-clay, while they moved north- 
ward from the Alps along with the glaciers, which 
always supplied them with an abundance of moisture. 
Alpine plants probably became exterminated in the 
plain of Central Europe at a much earlier period. 



What has been spoken of in the earlier parts of this book as the 
eastern migration, refers in a general way to the animals which 
have come to England from the east. But these are by no means 
natives of one country alone. We can trace a number of the 
British mammals to a Siberian origin, and also some birds; 
among many of the lower vertebrates and invertebrates, how- 
ever, there are few species which have reached us from Siberia. 
They may have had their original homes in the Alps, in Eastern 
Europe, or in Central and Southern Asia, and have joined in 
their westward course the later, more quickly travelling 
mammals. Many instances are given from all the more im- 
portant groups of animals to show how we may proceed in 
approximately identifying the home of a species. 

The periodical invasion into our continent of Pallas's Sand- 
grouse and other birds, suggests an explanation as to the cause 
of the great westward migration in former times of the Siberian 
mammals. Since a considerable amount of fossil evidence is 
available to show the path of migration pursued by these 
mammals, other important problems, such as the time of their 
arrival in Europe and the geographical conditions surrounding 
them, may perhaps be approximately ascertained, and thus 
throw much light on the general features of the European 
fauna. It has been proved by Professor Nehring that the 
Siberian mammals arrived in Eastern Europe after the deposi- 
tion of the lower continental boulder-clay. He believes 
that the climate of Germany at that time had ameliorated so 
far, after the maximum cold of the Glacial period, that steppes 
with a Siberian fauna could exist. Other groups, such as the 
Mollusca, however, do not support Professor Nehring's theory, 
and in order to arrive at an independent solution of this and the 
other problems referred to, a short history is given of the Siberian 


fauna. Recent geological ages have witnessed the arrival in 
Southern Europe of mammals now almost confined to the arctic 
and subarctic regions. In Siberia, on the other hand, many 
southern species penetrated, apparently about the same time, 
to the extreme northern limits of that country. The greatest 
authority on the Siberian fossil fauna, Tcherski, believes that 
this took place in pliocene times, the gradual retreat occupy- 
ing the whole of the Glacial period. If this were correct, the 
retreat from the Arctic Regions would have occurred at 
the same time when, according to our European authorities, 
Professors Nehring and Geikie, the much more southern 
parts of our continent were already uninhabitable. But 
Siberia could not have supported the large mammals at all 
at a time when Europe was uninhabitable, as it would be 
difficult to conceive under what geographical conditions the 
climate of the latter was arctic and that of the former temperate. 
If the whole fauna was driven into Southern Asia, how is it that 
the Siberian invasion of Europe occurred immediately after the 
deposition of the lower boulder-clay, that is to say, after the 
earlier part of the Glacial period? The difficulty can be met by 
the supposition that both Europe and Siberia had a temperate 
climate at that time. This view is supported by certain 
evidences, fully described, of a connection between the Caspian 
and the White Sea, which would have had the effect of influ- 
encing the climate. The Siberian fauna would thus have been 
prevented from spreading westward in Pliocene and early 
Glacial times. But on the disappearance of the marine con- 
nection, a way would have been opened into our continent, 
which again had an effect on the climate. The latter would 
have become sensibly colder and thus have reduced the 
habitable area of the Siberian fauna. 

Such geographical conditions would have been incompatible 
with a great northern mer de glace, and the boulder-clay in 
Northern Europe could not have represented a ground moraine 
but is a marine deposit. The sea is supposed to have covered 


the Northern Russian and German plains, and into it icebergs 
discharged the detritus which had accumulated on them when 
they were still Scandinavian glaciers. 

As regards the time of the arrival of the Siberian migrants in 
Europe, the English Forest-Bed gives us an additional clue to 
its determination. Since Siberian migrants are unknown from 
earlier deposits than this, it is reasonable to suppose that they 
arrived in England about the time when it was laid down. But 
since they appear in Germany in the inter-glacial beds subse- 
quent to the deposition of the lower boulder-clay, the former are 
probably contemporaneous with the Forest-Bed. Some of the 
deposits generally regarded as upper pliocene by British geolo- 
gists would therefore have to be classed with the lower 
continental boulder-clay as lower pleistocene. In connection 
with this theory some interesting faunistic data are given 
which seem to support it. 

In conclusion, the former presence of Arctic plants in Central 
Europe and their bearing on the climatic problems are 



THE Oriental migration is closely related to the 
Siberian. Both have originated within the Asiatic 
continent, and in many respects a strict line cannot 
be drawn between them. There can be no doubt 
that some of the species which we regard as Siberian 
migrants had their original home in more southern 
latitudes, and thus may have formed part of the 
older Oriental migration. The home of that migra- 
tion I take to be Central and Southern Asia, that is 
to say, everything south of the Altai Mountains and 
the Caucasus. Its members have reached Europe 
across an old land-connection which united Turkey, 
Greece, and Syria, while the Siberian animals in- 
vaded our continent to the north of the Caspian and 

The Siberian immigrants into Europe on the whole 
are not very numerous, but it is different with those 
from the more southern parts of the Asiatic con- 
tinent. The members of the Oriental migration 
form a very large percentage of the European fauna. 
No other migration has affected our continent so 
powerfully, because it continued uninterruptedly for 



a very long time. Hence its results can be traced 
from one corner of Europe to the other. We have 
seen that the Siberian migration only commenced 
after the first portion of the Glacial period had passed 
away. The Oriental, however, persisted throughout, 
or at any rate for the greater part of that period. It 
commenced ages before it, in miocene times, or even 
earlier. And as the ^Egean Sea, which broke up the 
highway of the Oriental migrants, is only of recent 
formation, there was a steady westward march for a 
very considerable time. No doubt the migration was 
also favoured by the fact that scarcely any formidable 
barriers had to be crossed. 

Many instances might be quoted of the same 
species forming part of the Oriental and also of the 
Siberian migration, but as a rule the Siberian mi- 
grant belongs to a distinct variety, or has such well- 
marked racial characters as to be at once detected 
from its more southern relative. Among the ex- 
amples of Oriental migrants which I have occasion 
to bring forward, such instances will be specially 
dealt with. 

In its wild state the Red Deer (Cervus elaphus) is 
almost extinct in the British Islands, though it still 
occurs in the moorlands of Devonshire and Somerset- 
shire in England, in the south-west of Ireland, and 
in some localities in Scotland. Fifty years ago it was 
also found wild in several other of the Irish western 
counties ; and in the seventeenth century it was 
common in most of the mountainous districts of 


Ireland. Its remains have been found fossil in the 
marls and caves of Ireland, and in the Forest-Bed, 
as well as in a large number of caves in England. 
The history of the Red Deer in other countries 
is very similar. In Scandinavia it flourished as 
far north as the sixty-eighth degree of latitude, 
whereas it is now quite extinct on the main- 
land, though still lingering on in some of the 
western islands. Denmark and Switzerland know 
it no more, and it is almost extinct in Belgium. 
Nearly throughout Europe where it occurs, its 
numbers are diminishing, greatly owing, perhaps, to 
the relentless persecution by man, but its gradual 
disappearance must likewise be partly due to other 
causes. Formerly it inhabited every country of 
Europe and all the larger islands. It still exists 
in Corsica and Sardinia, and at an earlier period 
it was also met with on the island of Malta. 
The Red Deer found in Corsica and Sardinia is 
smaller than that inhabiting Central Europe, and is 
by some authorities regarded as a distinct species, 
which has been named Cervus corsicanus. But 
Sir Victor Brooke has pointed out that the antlers 
of some of the Scotch Deer agree in every point 
with those of the Sardinian species. Indeed, the 
West European Red Deer altogether is a small- 
antlered form, compared with the Eastern one. This 
character, however, is only a racial one, and not of 
specific value. In the pleistocene deposits of Eastern 
and Central Europe, a very large-antlered race has 


been discovered, and identified by Professor Nehring 
with Ceruus canadensis the Canadian Red Deer. 
Tcherski, the Siberian traveller, believed that Cervus 
canadensis was identical with, or a variety of, the 
Asiatic species of Deer, Cervus eustephanus, Cervus 
xanthopygus, and Cervus maral. Some authorities 
and to these-belong Mr. Lydekker think that we 
ought perhaps to regard the whole number of Red 
Deer-like forms as local varieties of one widely- 
spread species. Besides the deer already referred to, 
the following belong to this same group: Cervus 
cashmirianuS) Cervus affinis, Cervus Roosvelti, from 
North America, and the North African Cervus bar- 

The question now is, where have these varieties 
originated ? Or, if we go to the root of the matter, 
where is the original home of their ancestors? Con- 
sidering that so many Cervidcz have been found in 
French and English pliocene deposits, and that 
remains of the Red Deer occur not only in the 
English Forest-Bed, but have been found associated 
with those of the Pigmy Hippopotamus in Malta, 
it would only be reasonable to suppose that the 
genus Cervus had originated in Europe. It might 
also be argued with equal force that the Red Deer 
had its birthplace in our continent. But when we 
carefully study its present range this verdict cannot 
be accepted. The view of the Asiatic origin of the 
Red Deer, so ably maintained by Koppen, cor- 
responds far better with its present distribution, 


especially if we look upon the Asiatic, North 
American, and North African forms as varieties of 
the same species. 

If the Red Deer were of European origin, it must 
have come into existence at a time when Malta was 
part of the mainland, when North Africa and the 
British Islands were connected with the continent of 
Europe, and of course before the deposition of the 
Forest-Bed. Such land-connections existed probably 
during the Pliocene Epoch. Migrants would have 
wandered from Europe into Asia. These would 
have developed into larger races, which again 
furnished emigrants for North America. The latter 
crossed by the old land-connection which once joined 
America and Asia at Behring's Straits. During 
pleistocene times the large Siberian race would now 
have re-migrated to the home of its ancestors in 
Europe, for we find the remains only in Central 
and Eastern Europe, indicating that an invasion of 
the Red Deer from Asia must then have taken 

Against this view of the European origin of the 
Red Deer, it may be urged that deer are known from 
Indian as well as from European pliocene deposits, 
and that a migration could have taken place from 
the Oriental Region to Europe just as easily as 
from the latter to Asia. The majority of the species 
of the genus Ceruus (in a wide sense), moreover, 
are Asiatic, ranging to Borneo, Sumatra, and the 
Philippine Islands, all of which islands have been 


separated from the mainland for a considerable time. 
Finally, the original home of a species, as we have 
learned, generally corresponds with the centre of its 
geographical range, and this lies in the case of the 
Red Deer in Central Asia. 

One of the highest authorities on the deer family, 
Sir Victor Brooke, also was of opinion that the 
Cervida originated in Asia, and from there spread 
east and west Of the two divisions into which true 
deer are divided, viz., the Plesiometacarpalia and 
the Telemetacarpalia, the former is almost confined to 
the Old and the latter to the New World. The only 
North American species belonging to the first 
division is the Canadian Red Deer, which fact 
clearly indicates its recent immigration to that 

There were probably two distinct migrations of 
the Red Deer into Europe. An older one coming 
from Asia Minor into Greece, which stocked Sardinia, 
Corsica, Malta, and North Africa in the first place, 
when these were still connected with one another. 
This same migration likewise affected western con- 
tinental Europe, the Irish Red Deer being probably 
the descendant of this very ancient stock. The 
latter entered the island when it was still part of the 
Continent. The later migration of a larger form came 
from Siberia and spread mainly over Eastern and 
Central Europe, but it appears that it also reached 
England, although there is no evidence of any of 
these Siberian deer having ever inhabited Ireland. 


The range of this deer, therefore, to some extent 
corresponds to that of another described on p. 153. 
We found then that two races of Reindeer had 
migrated to the British Islands one from the Arctic 
Regions, and the other from Siberia, but that only 
the former had reached Ireland. 

The so-called Irish Elk (Cervus giganteus) has 
been referred to the Oriental migration, but, as 
stated below, it has some claims to be regarded 
as a European. Unfortunately it is now extinct; 
it seems not unlikely, however, that it inhabited 
Ireland when man had already made his appear- 
ance on the island. Although its remains are found 
in such extraordinary abundance in Ireland, it 
certainly did not originate there. It lived also 
in England and Scotland, and in the Isle of Man, 
in France, Denmark, Germany, Austria, North Italy, 
and Russia. Its remains have been discovered 
even in Siberia. It must either have originated 
in Europe and then migrated to Asia, or have had its 
birthplace in Asia and wandered to Europe. There 
is nothing to lead any one to assert positively that 
either of these two continents was the one in which 
the original home of the Irish Elk was situated, and 
we can only be guided in this case by the history of 
its nearest relatives. These are the Fallow Ueer 
(Cervus dama). There are two very closely allied 
species, the Persian and the European, but several 
others have been discovered in the Forest-Bed and 
the pliocene deposits of the Auvergne. As no 


remains of the Fallow Deer are known from Asia, 
it seems probable that it and also the Irish Elk 
originated in Southern Europe, and only invaded 
Asia in early pleistocene times. 

The Mammoth (Elephas primigenius) is a familiar 
example among a large number of mammals which 
have come to us about the same time from Asia by 
the Asia Minor route. It had a much wider range 
than the' Irish Elk, since its remains have been dis- 
covered in a large number of European localities as 
far west as Ireland, also in Siberia, and even North 
America. Though we have had Proboscidea in Europe 
from the Middle Miocene onwards, Mr. Lydekker 
(d, p. viii.) holds that " our comparatively full know- 
ledge of Lower Miocene and Upper Eocene mam- 
malian faunas of the greater part of Europe and 
North America, renders it almost certain that neither 
of those regions was the home of the direct ancestors 
of the Elephantidce ; and we must therefore look 
forward to the discovery of marnmaliferous Lower 
Miocene or Upper Eocene strata in some other 
region of the (probably old) world which may yield 
these missing forms." 

The genus Elephas makes its first appearance in 
the Upper Miocene of India. Our European E. 
antiquus is, according to Professor Zittel, probably 
identical with E. armeniacus of Asia Minor, while 
E. meridionalis agrees in all essential characters 
with the Indian E. hysudricus. The Indian and 
European species of fossil elephants altogether are 


very closely related, and the supposition that they 
all have had their original home in the Oriental Region 
offers, I think, no serious obstacle. The view of the 
European origin of the mammoth especially is open 
to very serious objections. It does not occur in any 
European pliocene deposits, and could not therefore 
have originated in our Continent until pleistocene 
times. That it should then have commenced its 
travels through Europe and Siberia to the New 
Siberian Islands and North America seems almost 
an impossibility. But if we suppose the mammoth 
to have had its home in India in pliocene times, it 
could then easily have migrated to all the parts of 
the world where its remains have been discovered. 

Of the Asiatic mammals still living, some have 
only just crossed the borders of Europe and then 
died out again. Similar cases have been referred to 
in discussing the Siberian migration. Thus remains 
of the camel have been found in Roumania and in 
Southern Russia in pleistocene deposits. Others have 
lingered on to the present day. Crocidura etrusca, 
for instance, still lives in Southern France, Italy, 
Sicily, and North-western Africa. All its nearest 
relations are typically Oriental species. In spite of 
the fact that a Crocidura is known from French and 
German miocene deposits, the general range of the 
genus suggests an Oriental origin. In early Tertiary 
times a section spread into African territory and 
another eastward as far as the island of Timor. This 
may possibly have happened in miocene times, when 


a few species likewise found their way into Europe. 
Many other mammals have wandered still farther 
west, and now form an important percentage of the 
European fauna. 

Of Birds, too, a large number might be mentioned 
which had their home in Asia and have found their 
way to Europe with the Oriental migrants. A few 
instances have already been alluded to, and somq 
additional ones may be specified at random, without 
attempting to give a complete list. 

Some of the Wagtails (Motacilla), as I men- 
tioned in the last chapter, have certainly come to 
us with the Siberian migration; but others seem 
to be Oriental, such as Motacilla melanope, which is 
resident in Southern Europe and migratory in the 
North. M. campestris the Yellow Wagtail has 
a most peculiar discontinuous range. One colony 
breeds in the British Isles and Western Europe 
generally, where it is known as a summer visitor, 
retiring to W T est Africa during winter; another is 
found from South-east Russia to Turkestan in 
summer, and winters in Southern Africa. This fact 
may possibly be due to two distinct migrations from 
Asia having taken place: an earlier one from the 
South-east that is to say, an Oriental one and 
a Siberian one more recently. In this case the 
members of the two migrations have not become 
sufficiently differentiated to be regarded as distinct 
varieties. Though most of the Wagtails have a 
somewhat northern range, none (except perhaps M. 


borealis) are truly Arctic; and indeed, as almost all of 
them pass the winter in southern latitudes, it may 
be assumed that they are of southern and not of 
northern origin. 

The Dippers (Cinclus) are practically unknown in 
the Central European plain, but they occur in 
Western Europe as far north as Scandinavia, also 
in the Alps, Carpathians, and Southern Europe, in- 
cluding Sicily and Sardinia. Some authorities dis- 
tinguish three species, others only one. As a matter 
of fact, the difference between the three forms is very 
slight, and their nests and eggs are undistinguishable. 
Eight other species have been recognised, and all 
these are either Asiatic or American. As one of the 
American forms is peculiar to Peru and another to 
Ecuador and Columbia, and since the genus as a 
whole is a mountain-genus, it probably is an ancient 
one. Its European range alone, however, implies that 
it has inhabited our continent for a considerable 
time and is no new-comer. We may look upon it as 
of Asiatic origin. The ancestors have spread east 
and west, the European species having arrived with 
the earlier Oriental migrants, and wandered along 
the Mediterranean at a time when the geographical 
conditions of that sea were vastly different from what 
they are to-day. 

Not quite so ancient as the Dippers, but like- 
wise Asiatic in their origin, are the Bullfinches 
(Pyrrhida). The closely allied Pine-Grosbeak {Pint- 
cola enucleator] has already been referred to (p. 191) 


as a member of the Siberian migration. The 
distribution of the European Bullfinch (P. europcea) 
is very interesting, as it occurs in two distinct 
forms, by some authorities regarded as races, by 
others as species. In all probability these two 
races owe their origin to two different migrations 
from the same ancestral stock. We may suppose 
that P. europcza came to Europe along with the 
Oriental migration, spreading chiefly over the south 
and west, while another branch developed in Siberia 
into the larger and more brilliant race (P. major), 
which subsequently entered the neighbouring con- 
tinent with the Siberian fauna. The latter race 
inhabits, according to Mr Saunders, Northern and 
Eastern Europe, and also Siberia. All the other 
species there are eight more except one, are found 
in Asia. This one species, which inhabits the Azores, 
appears to be more closely related to one of the 
Siberian bullfinches than to the European. It stands 
isolated, and is an extraordinary instance of discon- 
tinuous distribution, as no Bullfinch inhabits either 
Madeira or the Canary Islands. We must assume 
that the form connecting it with the Asiatic prob- 
ably lived in Southern Europe, and has become 

One of the most typically Oriental genera of birds 
is Phaslanus, to which our Common Pheasant belongs. 
Out of twenty species, nineteen are found exclusively 
in Asia, most of them being confined to the central 
plateaux of that continent. Only one species passes 


the confines of Asia into Greece, Turkey, and 
Southern Russia. This is Phasianus colchicus. For- 
merly, however, the Pheasant appears to have had a 
wider range in Europe, for three species are known 
fossil from France. Altogether, it is not quite certain 
whether the Pheasant is not really an indigenous bird 
in the British Islands, having survived from pre- 
glacial times. It is believed that the Romans brought 
it to England, but there is no record of an introduc- 
tion at that time. 

Among the older Oriental bird migrants might be 
mentioned the Fire-crested WrQr\(Regutusignicapiltus) y 
which has even occasionally visited England. It be- 
comes commoner as we go south-eastward. In Asia 
Minor it is more abundant than the Gold-crest ; and 
throughout the year it is resident in Southern Europe, 
where it occurs in Turkey, Greece, Italy, Spain, 
Sardinia, and Malta. On the opposite shore, in 
North-west Africa, it again makes its appearance, 
and its range extends westward to the Canaries (R. 
teneriffa) and Madeira (R. maderensis). 

The genus to which our common Goldfinch belongs, 
viz., Carduelis, is also probably of Oriental origin, and 
may be looked upon as one of the earlier migrants. 
That species (C. elegans) breeds throughout Europe, 
except in the extreme north, but it is especially 
abundant in Southern Europe and North-west Africa. 
It is also resident in Madeira and the Canaries. 
Eastward its range extends to Persia. A larger 
race (C. major) inhabits Western Siberia and crosses 



the European border into Russia. It interbreeds 
in Siberia with C. caniceps^ an East Siberian 

A few instances of Reptiles and Amphibia with a 
similar range will show that the Oriental migration 
was not confined to the higher vertebrates. 

Two species of the genus Eremias (fodarcis] occur 
in South-eastern Europe. This is a genus of Lizards 
with rather a wide distribution, ranging from Central 
Asia to South Africa southward and China eastward. 
Altogether there are twenty-four species, two of which 
just enter Europe; and of the rest half are Asiatic 
and half African. Even if the genus were of African 
origin, it is extremely unlikely that the Asiatic 
species came by way of Europe. We may assume, 
therefore, with a fair degree of probability that the 
two European species wandered westward along with 
the Oriental migrants. 

The genus Ablepharns belongs to a family of 
Lizards in which the legs are either very fully 
developed, or quite absent as in the Slow-worm 
(Anguis fragilis). It is an ancient genus, having a 
wide range from Central Asia to Australia on the one 
hand, and to South Africa on the other. One species 
of this Scink-like Lizard, viz., Ablepharus pannonicus, 
enters Europe in the south-east, inhabiting Greece as 
far north as Southern Hungary. In Asia it is found 
in Syria and North Arabia. This clearly signifies 
that the Lizard is an Oriental migrant. 

Among the Snakes which participated in the 


Oriental migration might be mentioned Eryx jaculus> 
whose home is probably in Western Asia. It is 
known in Europe from the Greek islands of Tinos 
and Naxos, from Turkey and Southern Russia. 
Another, a peculiar worm-like form, lives under- 
ground in damp earth and under stones Typhlops 
lumbricalis. This species inhabits the mainland of 
Greece as well as the Greek islands, and Asia Minor 
as far as the Caucasus. 

A most interesting case of distribution is that of 
the pretty little Toad so well known on the Continent 
under the name of " fire-toad " (Bombinator igneus). 
Though some authorities, such as Boulenger, recog- 
nise only one form of Bombinator^ others are of 
opinion that two well-marked varieties exist in 
Europe. These are looked upon by Dr. von Bedriaga 
as good species, but he acknowledges that they are 
rather critical and difficult to identify. No other 
species of Bombinator occur in Europe. Bombinator 
pachypus, the western race, or if we choose to call it 
species, occurs in France, Germany, Switzerland, 
Austria, Sicily, and Greece. B. igneus the eastern 
race is found in Southern Sweden, Denmark, Ger- 
many, Austria, and Russia. The latter has therefore 
a more northerly and easterly range. The species is 
not known from Siberia, but makes its appearance 
again in China in a form which, according to Dr. von 

1 Since writing the above account, Mr. Boulenger, in his new work 
on the Batrachia of Europe, has accepted the specific distinctions 
between the two fire-toads. 


Bedriaga, docs not quite agree with either of the two 
European races. 

Now if we supposed Bombinator to have originated 
in Europe, its absence from the British Islands, most 
of the Mediterranean islands, and the greater part 
of Scandinavia would not be easy of explanation, 
while as an Asiatic migrant the European range is 
more readily understood. Its apparent absence from 
Western Asia might quite likely be due to the fact 
that the zoology of that part of the Continent is only 
now being investigated. The latter has, moreover, 
undergone great physical changes in recent geological 
times. The supposition that one migration of Bom- 
binator from the south-east has taken place, and 
then another from the east, seems to explain this 
case of distribution, as other similar ones, in a most 
satisfactory manner. 

The Tree-Frog (Hyla arborea) must be an ancient 
species, but it is not of European origin. Few genera 
of Amphibia have a wider distribution than Hyla. 
There are only three species in Asia, Europe, and 
Africa, the remaining 129 being confined to America 
and Australia. Two of the three Old World Tree- 
frogs are so closely allied that until recently they 
were regarded as mere varieties of one another. 
These are Hyla arborea and //. chinensis. The 
former is found in Asia Minor, Persia, China and 
Japan, and in most of the Mediterranean islands and 
Southern Europe generally. It does not occur in the 
British Islands, Norway, or North Russia, but in 


South Sweden, Germany, France, and Spain. It is 
also known from North Africa and from Madeira, the 
Canaries, and the Salvages. The occurrence of the 
Tree-Frog an so many of the Mediterranean islands 
is of particular interest, especially as four well-marked 
varieties have been distinguished by our leading her- 
petologists, so that the more minute features of the 
various forms can be traced from island to island, 
adding one more proof if proof were needed of 
their former continuity. Of course, that Hyla arborea 
must be considered an Oriental migrant seems so 
evident that it scarcely needs further comment. 

A number of mollusca might be mentioned whose 
range indicates that they have migrated to Europe 
from Asia Minor. Buliminus pupa is one of these. 
It is known from Asia Minor, Greece, South Italy, 
Sicily, and Algeria. Buliminus detritus is perhaps 
better known, being common in some parts of 
Germany. From there its range spreads east as 
far as Asia Minor. Many closely allied species 
inhabit Western Asia, to which they are confined, 
while others enter on European territory in some 
of the Greek islands. B. fasciolatus occurs on the 
islands of Crete, Rhodes, Cyprus, and in Greece 
and Syria. Most of the species of Buliminus have 
a very restricted range, but Buliminus obscurus is 
found almost all over Europe, from Ireland in the 
west to the Crimea and Transcaucasia in the east. 

Whether the sub-genus Pomatia of the genus 
Helix to which the so-called Roman Snail belongs 


is of Asiatic origin, or whether some of the species 
have migrated from Europe to Asia, I am not pre- 
pared to say; but there can be no doubt that Helix 
pomatia has reached Western Europe from the east. 

On the whole, the number of mollusca which we 
might point to as having migrated to Europe is not 
large, the great majority being indigenous to our 
continent. However, some of the other groups of 
invertebrates differ very materially in that respect 
from the mollusca. I cannot leave the consideration 
of the mollusca without referring to the fact that 
there appears to be a very important centre of 
distribution in South-eastern Europe. It is from 
this centre that many species have spread north 
and south, east and west Take, for example, the 
genus Clausilia, a small land-shell shaped like a 
pointed round tower, and abundant on old walls 
and tree trunks. In England we have four species 
of Clausilia> in Ireland only two. In the greater 
part of Spain only our common Cl. bidentata occurs. 
As we go east the number of species rapidly 
increases. A maximum is reached in South-eastern 
Europe, where hundreds of different kinds are found. 
Towards Northern Europe a similar decrease of 
species takes place. So far the history of the 
Clausilice seems perfectly simple. An active centre 
of origin appears to exist in South-eastern Europe, 
from which the species radiate out in all directions. 
But when we come to look more closely into the 
extra-European distribution of the genus, and 


especially when we examine its past history, we 
find that its origin is extremely complex, and dates 
back to a much more remote period than would 
have been imagined, had we merely taken into 
account its present range in our own continent. 
Professor Boettger, who is the highest authority on 
Clausilia, tells us that the genus is known from 
the earliest deposits of the Tertiary Era. About 
700 species are now known, and these have 
been sub-divided by Professor Boettger and others 
into a number of sub-genera. Some of these are 
extinct, but the great majority are still living. 
The sub-genus Phcedusa occurs in the eocene 
and oligocene of Southern Europe, but it is extinct 
as far as our continent is concerned. Close upon 
a hundred species, however, still inhabit India, 
the Malayan Islands, China, Ceylon, and Japan. 
Then again, the sub-genus Laminifera occurs in the 
oligocene and miocene of Central Europe, and 
survives in a single species, CL Pauli> in South- 
western France. The groups Garnieria of China, 
Macroptyckia of East Africa, Boettgeria of Madeira, 
and Nenia of South America, have no fossil repre- 
sentatives. We have here some very remarkable 
cases of discontinuous distribution which testify to 
the antiquity of the genus, and this is certainly 
confirmed by the fossil evidence. However, it is 
hardly likely that the headquarters, as it were, of 
Clausilia have always been in South-eastern Europe. 
Most of that part of the Continent has been sub- 


merged since eocene times more than once. The 
peculiar distribution of the genus might be explained, 
I think, if we supposed the original home of Clausilia 
to have been in Southern Asia, that from this centre 
Southern Europe was colonised, where a new centre 
developed in oligocene and miocene times, sending 
colonies off to Madeira and across the old land- 
connection which united Northern Africa and South 
America about that time. The most active centre 
of development then gradually shifted eastward 
again, while the older centres were perhaps sub- 
merged during the physical changes in the distri- 
bution of land and water. 

I should have mentioned that the species 
wandering westward and northward from this 
South-European centre of distribution, would 
naturally have joined the migrants which came 
from beyond the borders of our continent. They 
might thus appear to be true Oriental migrants, 
and on a previous occasion I grouped all these 
together under the term of "Southern Fauna," as 
I assumed the observer to be stationed in the 
British Islands. All new-comers from the south- 
east, south, or south-west of Europe would be to 
him southerners quite irrespective of their original 
home, which might be in Southern Europe, Asia, 
or Africa. 

The Swallow-tail is well known to all collectors 
of Butterflies in England, though it has of late 
years become very rare and is now confined to a 


few localities in the east of England. The members 
of the family Papilionidce, to which it belongs, are 
mostly large and striking species, and their distri- 
bution is therefore more accurately known than 
that of the smaller and less conspicuous butterflies. 
Only four different kinds of Swallow-tail Butterflies 
inhabit Europe, but in Southern Asia and the Malay 
peninsula they attain their maximum as regards 
numbers; and there we find a great many species of 
this genus Papilio. Of the four European species only 
one, viz., Papilio hospiton, is peculiar to Europe ; all 
the others range into Asia. It would seem, therefore, 
as if this genus was an Asiatic one and had migrated 
to Europe, and that the route taken was the one from 
Asia Minor across to Greece. We have a similar 
case in the closely allied genus Tliais two of the 
three European species living also in Asia Minor. 
Thais cerisyi inhabits some of the Greek islands, as 
well as the mainland of Turkey and Greece. 

Another genus of the great family Papilionidce 
with which most lepidopterists are well acquainted 
is Parnassius. What butterfly- hunter has been in 
Switzerland without hearing of, or seeing, the famous 
Parnassius Apollo ? We have four European species of 
Parnassius t only one of which is peculiar to our con- 
tinent, but the locality where it occurs, the Caucasus, 
is on the borders of Asia. Almost all the other 
species are Asiatic, none however range to the south. 
Its headquarters, and I think its original home, are 
the mountains of Central Asia. From there it 


has spread some species to the Himalayas, and 
a few to Europe and North America. But these 
migrations are not of very recent date. Parnassius 
no doubt arrived accompanied by a large number 
of other Central Asiatic mountain insects and 
plants. I shall refer to the latter again when 
dealing with the origin of the Alpine fauna, but 
meanwhile it might be mentioned that the famous 
Swiss "Edelweiss" (Leontopodium alpinuni), which 
we are accustomed to regard as a typical Alpine 
plant, is certainly of Asiatic origin. In some parts 
of Southern Siberia it is one of the common meadow- 
flowers, and ranges from there south into Kashmere, 
but not northward. Like the Apollo, it does not 
occur in Scandinavia or Northern Siberia. Both 
plant and insect evidently migrated from Central 
Asia, directly westward along the southern border of 
the sea, which extended from that region as far as 
the European Alps in early Tertiary times. At that 
time the Caucasus was possibly still connected with 
the Balkan Mountains, across what is now the Black 
Sea, and that may have been the highway on which 
they travelled west. 

Some of the Clouded-Yellows butterflies apper- 
taining to the genus Colias formed part of the 
Oriental migration. The genus is undoubtedly of 
Asiatic origin, and while many of the species have 
turned northward, ranging across Siberia and North 
America, others have taken a southern and westward 
turn and thus reached Europe. We have two 


Clouded-Yellows in Western Europe, and both of 
them must have come with this migration. 

A very good example of an Oriental migrant is 
Danais clirysippus, a magnificent butterfly found in 
Greece and Southern Italy. In Asia it is known from 
Syria, Persia, and from the whole of the southern 
portion of the Continent. The genus Danais (in 
its wide sense) is a large one, and principally 
occurs in the warmer regions of Asia. Three 
species are found in North America and only one in 

Among the beetles belonging to this migration, 
there is one of very considerable interest from a dis- 
tributional point of view, for all the species of the 
genus even the whole family to which the genus 
belongs are what is known by zoologists as " Com- 
mensalists." These are animals habitually associating 
and living in close connection with others with which 
they are not tied by any family relations or kinship. 
Such a state of close and permanent friendship is 
called " commensalism." Now it appears as if the 
members of this family of beetles (Clavigeridce) had of 
their own free will formed such a close connection 
with colonies of ants sometimes with one species, 
sometimes another. They are the permanent guests 
of the ants, and in return they secrete a fluid which is 
apparently highly prized by them. All of the Clavi- 
gers are provided with peculiar club-shaped antennae, 
with which they ungraciously beat their hosts, when 
they are in want of food. According to some authori- 


ties, they even occasionally gnaw at the pupae and 
larvae of the ant with which they live. 

Such beetles naturally can only have extremely 
limited means of distribution, and they are com- 
parable in that respect with the woodlice of the genus 
Platyarthrus, to which I have already had occasion 
to refer. All the species of Claviger are confined to 
Europe, chiefly to the south, but one species, CL 
testaceus, has wandered farther north and occurs in 
the nest of the ant Lasius flavus in the south of 
England, Ireland, and Scotland. Though none of the 
Clavigers can be claimed as Oriental migrants, the 
centre of distribution of the genera belonging to the 
Clavigerida is in Southern Asia, and it is probable 
that the ancestors of the European Clavigers have 
spread westward from that region to Europe, eastward 
to Australia and Japan, and southward to Madagascar 
and South Africa. The genus Hopatroides, belonging 
to the same family as the so-called Spanish-fly (Tene- 
brionidcz}) has twelve species in Western Asia and 
Greece. One only, H. thoracicus an instance of dis- 
continuous distribution occurs in Andalusia. Amphi- 
coma is represented in Western Asia and the Balkan 
peninsula by fifteen species, while three others are 
met with in North-west Africa and Southern Spain. 

A genus of Dragon - fly, Onychogomphus y has in 
Europe a somewhat similar distribution to Claviger, 
but it has besides a very extensive foreign range. 
There are altogether thirty-five species ; of these ten 
are Holarctic, twelve Oriental, five Mascarene, and 


eight Ethiopian. The centre of distribution is there- 
fore in the Oriental region, and we may assume that 
in all probability the genus has originated there, 
the European species having travelled west with 
the Oriental migration at an early date of the 
Tertiary Era. 

Ryothemis, another genus of Dragon - flies, has 
originated perhaps somewhat farther east than the 
last,, for no less than thirteen species are found 
in Australia, a like number in India, five in Mada- 
gascar and Africa, and five in the Holarctic region. 
Both of these genera are entirely absent from 
America, and they have possibly travelled to Europe 

Among the European OrtJwptera the group to 
which our Earwigs and Grasshoppers belong there 
are also a good many instances of Oriental migrants. 
One of the most striking of these is the curious 
"praying insect" (Mantis religiosd]. It occurs all 
over Southern Europe, and ranges as far north as the 
north of France. It is also found in Southern Ger- 
many and in Austria, and has a vast extra-European 
range. There are even records of its occurrence from 
all parts of Southern Asia and Java and a great part 
of Africa. That it belongs to an extreme'y ancient 
genus is testified by the fact of its presence in 
Mauritius, Japan, Australia, New Zealand, South 
America, and Madagascar. The genus Bacilhis to 
which the typical Stick-insects belong has a some- 
what similar geographical distribution. But no less 


than four species of Bacillus are known from Europe, 
according to our great authority Mr. Brunner von 
Wattenwyl all from the south ; and some of these 
also range into North Africa. There are thirty-two 
other species distributed over Southern Asia, Africa, 
Australia, New Zealand, and the Sandwich Islands. 

Volumes, indeed, might be filled with lists of 
species and genera of terrestrial invertebrates of 
Oriental origin, but I will not weary the reader 
with further enumeration of such instances. Just 
two more, however, before concluding, as I have 
not alluded to the large group of the Arachnida. 

Two peculiar spider-like genera, viz., Galeodes and 
Rhax> are found in Southern Europe. Both occur 
also in North Africa, and in Western and a portion of 
Southern Asia. As the whole family altogether has 
an Asiatic character, I cannot agree with Mr. Pocock, 
who considers them of European origin and believes 
that they are migrating eastward. 

But not only terrestrial forms migrated to Europe 
from Western and Southern Asia. Freshwater 
species also took part in this great Oriental migra- 
tion. 1 need only refer to the freshwater Crab 
(Thelphusa fluviatilis), with which Southern Euro- 
peans are familiar. It is the sole representative of a 
large genus which ranges east as far as Australia and 
southward to Madagascar and the Cape of Good 
Hope. The European species is found in Turkey, 
Cyprus, Greece, Southern Italy, Sicily, North Africa, 
Southern Spain, Syria, and Persia. 


There is a corresponding flora with a range exactly 
similar to that of some of the animals quoted. Thus 
the Balkan Rhododendron (Rhododendron ponticmri) 
is again met with in the western Mediterranean 
region in Southern Spain. The Cedar occurs in 
local varieties in the Himalayan Mountains, in the 
Lebanon, and the Atlas Mountains. Both of these 
are instances of discontinuous distribution, a proof of 
their antiquity; but a large number of plants have a 
continuous range between Asia Minor and Spain. 

On looking through these few instances of what have 
been called Oriental migrants, one cannot help being 
struck by the fact that the species after their entry 
into Europe evidently did not all follow the same path 
during their westward advance. We have seen that 
a good many seem to have travelled either due west 
or north-west on entering our continent from Asia 
Minor. They may now perhaps be found in Greece, 
Southern Italy, Algiers, and Spain, also probably on 
some of the intervening islands in the Greek Archipe- 
lago, in Sicily, Sardinia, and Corsica, or they may have 
travelled north-east and occur in the Alps. This distri- 
bution indicates undoubtedly, as I have already set forth 
in another memoir (c, p. 459), that land extended from 
Asia Minor across Greece to Southern Italy, that the 
latter again was disconnected with Central Italy, but 
united with Sicily, Sardinia, and Tunis, and that the 
Straits of Gibraltar did not exist at the time when 
these species migrated westward. Some species 
are only to be found as far west as Southern Italy, 


while others occur in Central and Northern Europe, 
scarcely in the South, and not at all in the 
larger Mediterranean islands or in North Africa. 
This appears to me to indicate that the late comers 
from the east found that geographical changes had 
taken place in Southern Europe which prevented 
them from following the same track as the older 
immigrants. They were now obliged to turn directly 
northward and then westward. It may be asked, 
why should not the earlier migrants have taken the 
same route? This question will be answered imme- 
diately. Meanwhile it should be clearly understood 
that there probably was an older and a newer migra- 
tion from the east. The Oriental genera from 
whose general range we know that they must be 
very ancient indeed, such as Mantis and Bacillus 
are almost invariably confined to Southern Europe. 
There they are frequently found on some of the 
Mediterranean islands. The earlier migrants there- 
fore went westward and the later ones north- 

Let us now inquire a little into the reasons why 
such different courses were pursued by the migrants 
why the Oriental migration divided into two 
streams, an older and a newer. 

During early Tertiary times, and probably through- 
out the Miocene and Pliocene Epochs, the ^Egean 
Sea did not exist. From the island of Crete to the 
Peloponnesus, and from Asia Minor to Thessaly 
and Macedonia, stretched a vast and fertile plain 


dotted over with numerous freshwater lakes. Grad- 
ually the sea encroached upon this land from the 
south, owing chiefly to extensive subsidences having 
taken place. Only very recently, says Professor 
Suess, did the whole of the ^Egean continent subside 
(i., p. 437). Huge cliffs of levantine freshwater 
deposits now mark the new coast-line, and the Medi-. 
terranean advances steadily towards the Black Sea 
and the Sea of Asov. A new order of things is now 
established, continues the famous author of Das 
Antlitz der Erde; where there were high mountains 
we now behold a deep sea, in some places many 
thousand feet deep. All this took place quite 
recently, geologically speaking, certainly in post- 
glacial times; and man may even have witnessed 
these imposing events. Most geologists admit the 
correctness of these views. They are, moreover, 
built upon such solid geological evidence, that even 
if the science of zoogeography had not yet taught us 
anything, naturalists would not hesitate in accepting 

Animals and plants were free to migrate from 
Central and Southern Asia to Greece by land for 
untold ages. The vast accumulation of mammalian 
bones which have been discovered at Pikermi, and so 
ably described by Gaudry, are probably to a large 
extent the remains of Asiatic immigrants to Europe. 
Many of these resemble forms still living in South 
Africa, which implies that a highway existed also at 
that time between Asia and Africa. Among these is 



a giraffe and antelopes closely allied to African 
species, and other most interesting mammals. 

In still earlier European deposits the Miocene 
we find the ancestors of modem Elephants, which 
are probably of Asiatic origin. The remains of 
several kinds of monkeys occur, whose nearest 
relations are now confined to Southern Asia. 
Altogether the fauna bears a strong Asiatic facies. 
Many of our European terrestrial invertebrates 
probably arrived about this time from Asia. The 
struggle for existence being keener and the facility 
for migration much greater in the higher vertebrates, 
they or at any rate the mammalian faunas were 
subjected to more rapid changes than the inverte- 
brates. I have repeatedly expressed my belief that 
a great number of our familiar insects and mollusca 
inhabited Europe long before our present mammals 
came into existence. 1 

Let us now follow one of the miocene Oriental 
migrants starting from Central Asia on its way to 
Europe. Very soon after leaving its home, it must 
have encountered a sea which extended at that time 
from the Eastern Mediterranean to the borders of 
Afghanistan. In following a westward course, the 
emigrant was compelled to keep along the northern 
shore of it. We do not know the state of the 
physical geography of the region between the Black 

1 In some cases the accuracy of this view is proved by fossil evidence, 
Helix rottindata, a common and widely spread British species, having 
been found in miocene strata near Bordeaux. 


Sea and the Tianshan Mountains, but it seems 
certain that a considerable extent of dry land 
enabled a wanderer from Central or Southern Asia 
to reach the Balkan peninsula by skirting the 
northern shore of that large miocene sea. No 
miocene deposits occur north of Teheran or of the 
Upper Euphrates, nor are they known from the 
islands of the ^Egean Sea or the lands surrounding it. 
From the Balkan peninsula it was possible for our 
migrant to reach the European Alps, which were 
then slowly rising as a peninsula out of the western 
portion of the great miocene sea. What are now 
the Alps was then hilly ground, which was being 
raised from the bottom of the sea. It was no doubt 
connected with the Balkan peninsula, so that an 
intercourse of species could take place between this 
newly-formed peninsula and Central Asia. I say 
peninsula, because the miocene sea almost completely 
surrounded it. From the Western Mediterranean a 
wide gulf extended up the Rhone valley into that of 
the Rhine as far north as Maintz. Then skirting along 
the northern outliers of the Tyrol, the gulf can be 
followed as far east as Transylvania. It is quite 
probable that it extended much farther east still, but 
there is as yet no geological evidence forthcoming. 
At any rate, our Asiatic migrant turning northward 
from the Balkan peninsula found its farther progress 
barred once more by an arm of the same sea which in 
its earlier peregrinations had stopped it from going 
south (cf. Suess, i., p. 406). 


In later miocene times the sea does not seem to 
have surrounded the Alps to the same extent as it 
did before, but it certainly extended from the Eastern 
Alps to the shores of the Sea of Asov, so that the 
direct northward passage was still more or less barred 
to the Oriental immigrants. At the same time Alpine 
species were now able to emigrate to the North 
European provinces. During the last stages of this 
epoch, the same sea increased its area very consider- 
ably in an eastward direction. One continuous 
expanse of water now stretched from the Alps as 
far as the Sea of Aral in Central Asia, perhaps even 

During pliocene times especially, the northern parts 
of the Balkan peninsula were occupied by a series of 
freshwater lakes, while Greece was joined to Southern 
Italy, Sicily, and Tunis. Central and Northern Italy 
were represented by a long narrow peninsula con- 
nected in the north with the Alps. Corsica and 
Sardinia were joined to Sicily, and the Straits of 
Gibraltar did not exist. When I first published my 
views regarding these geographical conditions of the 
Mediterranean area, Professor Deperet was good 
enough to send me his criticisms from a purely 
geological standpoint. He is of opinion that though 
Sicily and Sardinia might at this time have still been 
connected with Tunis, the Straits of Messina must 
already have been formed in other words, Southern 
Italy and Sicily could no longer have been connected 
with one another. This opinion is based upon the 


fact that in the upper strata of the enormously thick 
Sicilian pliocene deposits are found a number of 
arctic or subarctic species of mollusca which are 
entirely foreign to the Mediterranean fauna. It is 
generally supposed that these reached the Mediter- 
ranean area by the newly opened Straits of Gibraltar 
in later pliocene times, and that the lower Sicilian 
deposits must therefore have been laid down earlier. 
So far the deductions are perfectly correct, if we assume 
the northern mollusca to have arrived in the Atlantic 
at the time stated. However, they must have reached 
the Atlantic much later not till pleistocene times 
if we adopt the above-stated suggestions as to the 
age of the Forest-Bed (cf. p. 125). Moreover, the 
great similarity between the faunas of Southern 
Spain and North-western Africa indicate that the 
formation of the Straits of Gibraltar is of very 
recent date. The northern mollusca, of course, 
could not have reached Sicily till later. To suppose 
that the Sicilian deposits have been uplifted 7000 feet 
since then is no doubt contrary to all our geological 
teaching, but we must remember that this is altogether 
an exceptional case. The area in question has prob- 
ably ever since been in the immediate neighbourhood 
of an active volcano, and the rate of the uplift has 
therefore been immeasurably greater than at other 
localities with which this one might be compared. The 
disconnection between Tunis, Sicily, and Southern 
Italy was evidently produced by a subsidence of the 
tract of land uniting these countries. If we suppose 


that this happened in early pliocene times, we have 
either to take for granted that the terrestrial fauna 
and flora of these countries are of miocene origin, or 
that they were joined again during the Pleistocene 
Epoch. The range of a very large number of animals 
and plants is such as can only be explained by 
assuming that Tunis, Sicily, Sardinia, Corsica, and 
Southern Italy were connected with one another. Of 
such extensive land-connections subsequent to the 
arrival of the northern marine mollusca we possess, 
however, no geological evidence whatsoever; and it is 
extremely improbable that the land-areas which had 
sunk were once more raised before again subsiding. 
The many animals whose presence in the Mediter- 
ranean Region bears witness to these ancient land- 
connections could not have arrived there in miocene 
times in fact, they could hardly have lived there 
before the end of the Pliocene Epoch. On the other 
hand, it seems difficult to believe, once the Straits of 
Gibraltar were open and the waters of the Atlantic 
able to enter the Mediterranean, that the sunken 
parts between Sicily, Italy, and Tunis could have 
been raised without affecting the entire area of that 
sea. Nor is it likely that the junction between these 
countries could have then been brought about by a 
general lowering of the Mediterranean waters. As it 
may be asked what evidences we possess at all for 
the supposition of such land-connections as I have 
indicated, also that Southern Italy and Greece 
were connected, a few of the more salient instances 


of distribution bearing on this problem may be of 

I have already referred to the occurrence of the 
remains of a small race of Red Deer in the caves 
of Malta, similar to those still living in North- 
west Africa, Corsica, and Sardinia. The Black- 
mouthed Weasel (Mustela boccamela) inhabits Persia, 
Asia Minor, Greece, South Italy, Sicily, and Sardinia, 
while Mustela africana is found in Malta and Algiers. 
The European Porcupine inhabits Asia Minor, the 
island of Rhodos, Greece, Southern Italy, Sicily, 
North Africa, and Spain. Then we have the Wild 
Sheep of Asia Minor, Cyprus, Sardinia, and Corsica, 
all of which are closely allied. The small shrew-like 
Crocidura etrusca occurs in South France, Italy, 
Sicily, and North Africa. Many other mammalia 
might be quoted, but these are sufficient for our 

There are a good many reptiles and amphibians 
with a similar distribution. The European Chamaeleon 
(Chamczleon vulgaris) has been found in South Spain, 
North Africa, and Sicily. The Snake Periops hippo- 
crepis is confined to Spain, Sardinia, and Greece. 
The worm-like Lizard Blanus cinereus inhabits some 
of the Greek islands, North Africa, and Spain. 
Another Lizard belonging to the Scincid& has also 
been found in some of the Greek islands, Sicily, 
Sardinia, Southern Spain, and the Canary Islands. 
Discoglosstis pictus a toad" occurs in Spain, North- 
west Africa, Malta, Sicily, Sardinia, and Corsica. 


A variety of the Tree Frog (Hyla arborea Savignyi} 
is found in Europe only in Corsica, Sardinia, and the 
Greek Archipelago. 

Eight species of Reptiles and Amphibia some of 
which I have just referred to are enumerated by Dr. 
Forsyth Major as occurring eastward and westward 
of the Italian peninsula (and almost all also in North 
Africa) without being known on the mainland of Italy. 
And in order to show that Sardinia and Corsica are 
more closely related to North Africa than to Italy, 
he indicates the general range of the Reptiles and 
Amphibians found in these islands. Of the twenty- 
one species, only twelve inhabit Italy, but at least 
sixteen North Africa and seventeen Spain. Indeed, 
he shows that Corsica, Sardinia, Sicily, and North- 
west Africa form a zoogeographical province, from 
which Italy, with the exception of a few localities 
on its west coast, is excluded. It is a remarkable 
fact that there are a few localities on the west 
coast of Italy which in their fauna and flora 
exhibit closer relationship with Corsica and Sar- 
dinia than with the mainland. Thus Dr. Major 
pointed out that the Catena Mettalifera, the Monte 
Argentario, and Monte Circeo all belong to what 
we may call the former Tyrrhenian continent. 
They are to be regarded as its eastern limits, which 
remained standing, while the central portion now 
occupied by the Tyrrhenian Sea subsided, and is 
at present covered by deep sea. Subsequently these 
remnants- of the old continent became joined with 


the newly-formed Italian peninsula, but the plants 
and animals belonging to the older flora and fauna 
were mostly destroyed by newer and more vigorous 
immigrants. A few of the more hardy ones survived, 
and are a standing testimony of the geographical 
revolutions of that part of Southern Europe. 

That the Mediterranean area has undergone such 
profound geographical changes as I have endeavoured 
to indicate is no new theory. Many zoologists who 
have investigated the fauna of that region, and have 
attempted to explain the faunistic relations, had to 
acknowledge that the migrations must have taken 
place under geographical conditions entirely different 
from those obtaining at present. Riitimeyer long 
ago remarked that it seemed to him much more 
probable that Morocco, Algeria, and Tunis were 
peopled by way of Gibraltar, and perhaps also by 
Sicily and Malta from Europe, than Southern 
Europe from Africa. After careful conchological 
researches in the Western Mediterranean region, 
Dr. Kobelt came to the conclusion that formerly 
Southern Spain and Morocco must have been 
united by a broad land-connection. Sicily and 
Algeria do not apparently show any very intimate 
relationship conchologically, but farther west in the 
mountains of Tetuan Dr. Kobelt discovered a colony 
of Sicilian forms. 1 

1 There are a great many instances of discontinuous distribution 
among Oriental Invertebrates. Thus the Freshwater Crab ( Telphitsa 
faiviatilis] occurs in Southern Italy, Greece, Turkey, Cyprus, and 


" The close relationship," remarks Dr. Major (a, 
p. 106), "shown in the fauna of Corsica and Sardinia 
to Africa, permits the supposition that the connec- 
tion with these islands had persisted to a much more 
recent date than that with Europe." 

Many other authors have pointed out the close 
similarity existing between the faunas of Southern 
Europe and North Africa. We need only refer to 
the writings of Professor Suess, Milne-Edwards, and 
Boyd Dawkins. Mr. Blanchard went even so far as 
to say, " a comparer les plantes et les animaux de la 
Sicile et de la Tunesie, on se croirait sur le meme 
terrain" (p. 1047). 

No less than 113 species of phanerogamic plants 
are enumerated by Professor Engler (p. 53) as occur- 
ring in the Mediterranean coast region east and west 
of Italy without being found in that peninsula, or at 
least only in the extreme south of it. But he tells us 
that these species represent only a portion of such 
plants, which are extremely numerous. 

In taking a general survey of these plants, Pro- 
fessor Engler is of opinion that their range implies 
that a large number of the Mediterranean species 
have migrated along a line which can be drawn 
between North Africa, Sicily, Greece, Crete, and Asia 
Minor, and that from this line the distribution started 
northward again. 

Asia Minor. Another crustacean a Freshwater Crayfish (Hemicari- 
dina Desmaresti] inhabits Spain, Corsica, Sardinia, Sicily, and Asia 


Many of these plants then, and also some of the 
animals I have referred to, formed part of the older 
stream of migration which entered Europe from Asia 
Minor (vide Fig. 5, p. 117). There were only two 
courses open to them as they arrived on our continent 
during earlier Tertiary times. They could either go 
straight west towards Greece, or in a more northward 
direction to the newly-formed Alps. As the latter 
were raised, some of the immigrants were modified 
so as to adapt themselves to the new surroundings. 
Others became extinct ; but a great many have per- 
sisted in the Alps to the present day and exhibit 
discontinuous distribution, having meanwhile dis- 
appeared in the intermediate tract between the latter 
and their original home in Asia. The lowlands of 
Eastern and Central Europe were either occupied by 
the sea or by large freshwater lakes, so as effectually 
to prevent a direct migration northward. 

When the newer migrants arrived from Asia not 
only had the Alps risen to a lofty, mountain chain 
acting as an effectual barrier, but Southern Italy and 
Greece had become disconnected. Some time after, 
Sicily and Southern Italy also became separated. 
Meanwhile the stream of migrants which consisted 
less and less of typically southern forms, emigrants 
from Central Asia and even Southern Siberia, 
mingled with the southern forms on their way to 
Europe, and these now poured across the newly 
opened plain of Central and Northern Europe. But 
it was not until some time after this that the Mediter- 


ranean Sea broke across the ^Egean region, and that 
the Northern Sea retired from the plains of Eastern 
Russia to admit the typical Siberian fauna and flora 
into our continent (vide pp. 189-241). 

I cannot close this chapter without referring to the 
active distributional centre or I might say, centre of 
origin of species situated in South-eastern Europe. 
No group of animals is more instructive in eluci- 
dating the paths of migration from this centre than 
the terrestrial mollusca. Wherever the original home 
of the genus Clausilia may have been in early Tertiary 
times, it is certain that the most active centre of 
origin is now, and has been for a considerable 
time past, in South-eastern Europe. One of the 
earliest migrants from that modern centre of this 
interesting genus is Clausilia bidentata, which is the 
only species found in Southern Spain, and one of 
the two met with in Ireland, and which has been 
observed in high altitudes in the Alps and in 
Scandinavia. As we go eastward from Western 
Europe the number of species of Clausilia^ as we 
have seen, increases until we reach a maximum in 
the Balkan peninsula and the region of the Caucasus. 
Limax, Agriolimax, and Amalia, three genera of 
slugs, likewise appear to have originated in the same 
region and spread over Europe from there. Some 
species like Limax maximus and L. marginatus are 
very ancient, and probably commenced their wander- 
ings in early Tertiary times. In this manner many 
animals of European origin have joined the Oriental 


migrants in their westward and also in their later 
northward travels. In a similar way species of plants 
and animals of Alpine origin might have joined these 
migrants in their northward course, and it is only 
when we come to carefully analyse the constituent 
parts of all these members which have come to us in 
England from the south, that we realise the com- 
plexity of their origin. Finally, even the Siberian 
migrants mingled with the later Oriental ones, and 
in some cases the decision as to whether a certain 
species belongs to the former or to the latter migration 
becomes a matter of great difficulty. 


LIKE the last chapter, this deals with the Asiatic migrants. 
But while the former described the history of the northern 
invasion, those animals which entered Europe from the south- 
east are here more particularly referred to. They originated in 
Central, Southern, and Western Asia. It is not easy to dis- 
criminate in all cases between this Oriental migration and the 
Siberian. To a certain extent, even an entry of Northern 
Asiatic species has taken place by the southern route, and 
vice versa. On the other hand, southern species might have 
come to Europe by the southern route that is to say, to 
the south of the Caspian and also by the northern, which 
lay to the north of that great inland sea. The Red Deer is a 
good example. It arrived on our continent by both routes. 
However, there is a racial difference in the members of the two 
migrations. The small race now found in Corsica, Sardinia, 
North-west Africa, and Western Europe, is probably the older 


of the two, while the larger one resembling the American 
Wapiti Deer arrived very much later from Siberia. 

The Mammoth, Wild Boar, Badger, the Dippers and Phea- 
sants, are all Oriental species which have come to us from the 
south-east ; but there are also Reptiles and Amphibians, and a 
host of Invertebrates. Not all the animals, for instance, which 
have reached us in England from the south-east are of Asiatic 
origin. There is an active centre of distribution in South- 
eastern Europe itself, from which species radiate out in all 
directions. This fact is well illustrated by the genus Clausilia. 
Species from this centre, and also from the Alps, joined the 
Oriental stream in their northward course. 

In reviewing a number of instances of Oriental species in 
Europe, one is struck by the peculiarity of their having ap- 
parently followed two distinct routes. All entered from Asia 
Minor, which is proved to have been connected with Greece 
until recent geological times. From here some seem to have 
proceeded straight west, others northward. Further study 
reveals the fact that the first route was followed by a much 
older set of migrants at a time when the Mediterranean area 
was greatly different from what it is at the present day. 
Greece was then joined to Southern Italy, Sicily, and Tunis. 
The latter was also connected with Sardinia and Corsica, and 
the Straits of Gibraltar did not exist. Under such geographical 
conditions a direct migration on land from Southern Greece to 
Spain was not only possible, but was actually undertaken by 
a very large number of Oriental species. 



UNDER the Roman Emperor Augustus, the Spanish 
peninsula was divided into three provinces, one of 
which namely Lusitania occupied a large portion of 
the present area of Portugal. The term " Lusitanian " 
is therefore almost synonymous with Portuguese, but 
it has frequently been applied by zoologists and 
botanists in a much wider sense, so as to vaguely 
include the extreme south-west of Europe without 
any definite limits. Neither do I propose to restrict 
the term to everything found within the borders of 
Portugal For the sake of convenience, we may 
designate as Lusitanian forms those animals and 
plants which have migrated to Central, Southern, or 
Northern Europe from South-western Europe. They 
may really be North-west African species, or they 
may have originated on land which lay to the west of 
Portugal, and which is now mostly buried beneath a 
deep sea. Nevertheless, we have received them from 
the extreme south-western portion of our continent 
they have come to greater Europe from that 

In discussing the component elements of the British 


fauna and flora in the third chapter, I have already 
referred to the distinguishing characters of the Lusi- 
tanian migrants and to their distribution. I need 
only repeat, therefore, that these are now principally 
confined to the south-western portions of the British 
Islands. The late Edward Forbes was the first to 
trace the Lusitanian flora to its native home. In his 
classical memoir on the geological relations of the 
existing fauna and flora of the British Isles, he laid 
the foundations of a new method of research. We are 
as yet only beginning to realise the far-reaching 
conclusions obtainable by a careful study of the 
geographical distribution of animals and plants, 
though the lines of investigation were indicated 
by him more than fifty years ago. Forbes was 
of opinion that the Lusitanian element in the British 
flora was of miocene age, and that it survived the 
Glacial period on a now sunken land to the south- 
west of Ireland. Mr. Carpenter and myself agree 
in so far that we are both inclined to look upon 
this Lusitanian flora and the accompanying fauna 
in Ireland as of pre-glacial origin. But I am 
not quite satisfied that the Lusitanian migration 
ceased to come north then. It may have received 
a temporary check ; but the presence, for instance, 
of the Dartford Warbler {Melizophilus undatus) in 
the south-east of England would seem to indicate 
that its northward migration took place in very 
recent times. It is possible also that the very 
restricted occurrence of the Dartford Warbler may 


imply that it is gradually withdrawing towards its 
centre of origin from a former wider range. Such 
an eventuality, as we have seen, has actually taken 
place in a great number of instances. 

It is not only in the British Islands that we 
perceive the influence of the Lusitanian element. 
Scandinavia, Russia indeed almost every part of 
Europe can boast of some migrants which have 
originated in South-western Europe or on the 
mysterious lands which lay beyond it. As a rule, 
however, we notice a marked decrease of Lusitanian 
species as we travel eastward from Western Europe. 
Nevertheless, certain forms have travelled far beyond 
the confines of our continent, and we certainly meet 
with them in Asia and Northern Africa. 

It is remarkable that we are apt to mistake some- 
times for Lusitanian migrants species which are of 
Oriental origin. In a previous paper I classed such 
animals which had apparently originated in South- 
western Europe, but had really come from Asia by a 
circuitous southern route, with the Lusitanians. How- 
ever, there is really no reason why the two should 
not be kept apart, provided we can discriminate 
between the pseudo-Lusitanians and the true ones. 
I have already indicated in the last chapter how 
these pseudo-Lusitanian migrants originated. 

Supposing an Oriental species had left Asia for 
Europe in miocene times, it would on its arrival in 
Greece have had to decide between two courses. It 
could either advance into the newly-formed Alpine 



peninsula and there remain, or at once push on west- 
ward into Southern Italy, Sicily, and Tunis, by means 
of the old land-connections, and thence into Southern 
Spain. The Atlantic communicated at that time with 
the Mediterranean across the valley of the Guadal- 
quivir ; but that connection ceased to exist towards 
the end of the Miocene Epoch, when the Oriental 
migrants were free to ramble through Spain and the 
whole of the North European plain. I have indicated 
on a previous occasion (a, p. 484) that the earliest 
members of the Red Deer migration, which have left 
their traces in the caves of Malta, and whose descend- 
ants still live in Corsica, Sardinia, and North Africa, 
may have found their way to Northern Europe in 
this manner. Many other Asiatic mammals probably 
reached the British Islands in a similar way. 

I cannot call to mind any large species of 
mammal which we might reasonably suppose to 
have originated in South-western Europe. Even 
among the smaller ones, few give us any definite 
clue in this respect. For instance, the present range 
of the genus Myogale a small Insectivore belonging 
to the Mole family (Talpidce) teaches us nothing. 
The two living species show discontinuous distribu- 
tion, and are almost confined to Europe. Myogale 
occurs fossil in French miocene deposits, but is 
unknown beyond the confines of our continent. It is 
therefore probably of West European origin. The 
gap between the South Russian M. moschata and the 
Spanish M.pyrenaica is bridged over in so far as we 


know from fossil evidence that the former had a 
much wider range in pleistocene times, being then 
found in England, Belgium, and Germany. Ta!pa> 
too, to which genus our common Mole belongs, 
seems to be a West European genus, since it occurs 
in French miocene deposits. However, it would be 
difficult to name many more recent genera which could 
be included in the area which I propose to investigate 
in this chapter. The genus Lepus is probably not of 
Lusitanian origin, but the sub-genus Oryctolagus to 
which our common Rabbit belongs has no doubt had 
its original home in that region. Only two species of 
Lepus {Oryctolagus) are known, one of which Lepus 
lacostei has been met with in French pliocene 
deposits. The other is the Rabbit (L. cuniculus). 
Though generally considered to have been intro- 
duced into the British Islands, no reason can be 
brought forward in favour of such a supposition, 
especially as it is known to have spread into 
Germany in pleistocene times from South-western 
Europe. It occurs in France, the Spanish peninsula, 
North-western Africa, and on some of the Mediter- 
ranean islands. Its nearest living relatives, as we 
should almost expect, are found in South America. 

Of the Lusitanian Birds I have already mentioned 
the so-called Dartford Warbler {Melizophilus undatus\ 
which ranges from the south of England to the 
extreme south-west of Europe. A second species 
occurs on the Balearic Islands and on Corsica, 
Sardinia, and Sicily. The Andalusian Bush-quail 


(Turnix sylvatica) is probably of North African 
origin, and has subsequently spread into Southern 
Spain and Portugal, and eastward as far as Sicily. 
It is an instance of a migrant utilising the old 
Mediterranean land-connections in the opposite direc- 
tion from that described in the last chapter. 

Two of our British Wagtails are very closely 
related, so much so that it requires a very critical eye 
to distinguish them even at close range. They also 
frequently interbreed. In their distribution, however, 
there is a considerable difference between the White 
Wagtail (Motacilla alba} and the Pied Wagtail 
(M. lugubris). While the former ranges almost 
all over Europe and Asia, the latter is a local 
form resident in the British Islands, Southern 
Scandinavia, and France, and a winter visitor to 
Spain and North-west Africa. The genus Motacilla 
is probably Oriental in its origin, but it seems as if 
the Pied Wagtail was a Lusitanian species which 
had gradually spread northward, only to return to 
South-western Europe in severe weather for shelter. 

The Bearded Titmouse (Panurus biarmicus] the 
only representative of the family Panuridce may 
possibly be a Lusitanian bird. The fact of its 
being absent from Scandinavia and Northern Russia 
is suggestive of a southern origin. It is doubtful 
whether the bird occurs on the south side of the 
Mediterranean, but it is common in the south of 
France and Spain, and has also been observed in 
Sicily, Greece, and Asia Minor. In Central Europe 


it is found sparingly, and eastward its range extends 
as far as Turkestan. 

The genus FringilUi> which belongs to the great 
family of the Finches, appears to be not only of 
European origin, but, if the range of the species 
counts for anything, I should feel inclined to locate 
their home in the south-west. Altogether, five 
species are known. One of them, viz., Fringilla 
leydea, is confined to the Island of Teneriffe ; another, 
F. madeirensis, is found in Madeira, the Canaries, 
and the Azores ; a third, F. spodiogenys, inhabits 
North-west Africa. The two remaining species 
have a much wider range. F. ccclebs the common 
Chaffinch occurs in Europe, while its range extends 
eastward to Western Siberia, Persia, and Turkestan. 
The other F. montifringilla, known as the Bramb- 
ling is more common in Northern Europe, and 
generally frequents the more northern latitudes of 
Asia as far as Japan. 

It might be urged that the peculiar little blue 
Magpie of Spain Cyanopolius Cooki should find a 
place among the Lusitanian species, since there is 
no bird like it anywhere else in Europe. But in 
Eastern Siberia there lives a bird so closely allied 
as to be barely distinguishable from it. Neverthe- 
less, since there are some distinguishing characters, it 
has received a distinct name C. cyanus. This is a 
most interesting and remarkable case of discontinuous 
distribution, which may perhaps be explained by the 
supposition that the genus is of Oriental origin, and 


has died out at its former headquarters in Southern 
Asia and all along the line of migration, except at 
the extreme limits of the range in both directions 
east and west. 

As we go down in the scale of life among the lower 
vertebrates and invertebrates we meet with a greater 
number of prominent members of the Lusitanian 
migration. The Bullfinch, Dipper, and Chough, 
which might be thought to be of Lusitanian origin, 
are, as I have shown in the last chapter, Asiatic. 

The European snakes seem to be all of eastern 
origin, unless TropidonotiLS viperinus might be claimed 
as a Lusitanian form. Of very great interest from 
a zoogeographical point of view is our only European 
member of the South American and African family 
Amphisb&nida. This species Blanus cinereus is 
of the size and shape of an ordinary earth-worm, 
from which, however, it may be distinguished by its 
snake-like wriggling motions. It lives under stones 
in Spain and Portugal, North-west Africa, and 
Greece. It has, therefore, a somewhat similar dis- 
tribution to that of many of the animals and 
plants referred to in the last chapter. But here 
we have an animal which has evidently utilised the 
old Mediterranean route described on p. 271, from 
west to east. Two other species of Blanus inhabit 
Asia Minor and Syria, but most of its nearest relations 
either live in South America or tropical Africa. In 
migrating to North and West Africa, its ancestors 
probably made use of the land-bridge which spanned 


the Atlantic in early Tertiary times. Another 
Lusitanian Lizard belonging not to an aberrant 
group, but to the typical Lacertidae is Psammo- 
dromus hispanicus. It is rather variable in colour 
generally of a brown or green and grows to a length 
of about four or five inches. It occurs throughout 
the Spanish peninsula and also in Southern France. 
One of the handsomest European Lizards, which 
reaches almost a foot in length, of an olive colour 
with greenish or mother-of-pearl reflection, and with 
two yellow stripes along each side of the body, is 
an allied species (P. algirus). From the Spanish 
peninsula it passes into Southern France and North 
Africa. Two other species of the genus are confined 
to North-west Africa. 

It is quite possible that the genus Pelobates is of 
south-western origin. Of the two known species of 
this genus of Toads, one is found in the Central 
European plain and the other on the Spanish penin- 
sula and in France. The closely allied Pelodytes 
punctatus, too, is confined to this south-western 
district, and their nearest relations are found in 
Mexico. Similarly, the genus to which the Midwife 
Toad (Alytes obstetricans) belongs may have its 
original home in that part of Europe. Of the two 
species, one is confined to France, Switzerland, 
Belgium, and Western Germany, and the other, viz., 
Alytes cisternasii, to Spain. Discoglossus pictus a 
well-known and conspicuous Toad in Southern 
Europe inhabits Spain, Algiers, and Tunis, the 


islands of Malta, Sicily, Sardinia, and Corsica. From 
the general range of the family Discoglossidtv, as 
given in Mr. Boulenger's excellent catalogue, it 
appears that nowhere in the vast space between 
China and New Zealand has any member of the 
family been discovered. The peculiar genus of 
Salamander CJiioglossa is quite confined to the 
Spanish peninsula. 

The Butterflies Nemeobius htcina and Charaxes 
jasius may also have had their home in that south- 
western district. To this migration also seems to 
belong the genus Gonepteryx, which has so peculiar a 
range in the British Islands. The only British species, 
known as the Brimstone Butterfly (Goneptcryxrhanmi\ 
occurs in the south of England and in the south and 
west of Ireland. It is met with over the greater part 
of Europe, and its range extends into Asia Minor and 
Northern India, and then it reappears again in dis- 
tinct varieties in Japan and the Amur district. Three 
other species of Gonepteryx are known from Tibet 
and India, and one (G. cleopatrd] from Southern 
Europe and Northern Africa. All the remaining 
species inhabit the west, viz., Brazil, Mexico, and 
Venezuela. That the genus has migrated from 
America eastward to Europe appears to be more 
probable than a migration in the opposite direction. 
At any rate, that an exchange of species between 
the south-western portion of the Holafctic Region 
and the Neotropical area took place is indicated by 
the fact, not only that a variety of G. cleopatra has 


been found in Madeira, but also that the Canary 
Islands possess a distinct form of Goneptetyx, viz., 
G. cleobule. 

Dr. Kobelt has given us such an exhaustive memoir 
on the characteristic Mollusca of the different 
zoogeographical provinces of Europe, that we are 
particularly well informed as regards that group of 
Invertebrates. He tells us that the group Torquilla 
of the genus Pupa which is a small chrysalis-like 
snail is especially characteristic of the Pyrenees, 
Spain, and Portugal. In a certain measure they 
replace there the Clausilice which, as we have seen in 
the last chapter, have come from the east and are 
almost entirely absent in the south-west of Europe. 
Of about seventy species of Torquilla, the larger 
number are confined to this district, and some, which 
like Pupa (Torquilla} granum, range eastward, have 
travelled along the old Mediterranean highway, via 
Algiers, Sicily and Greece, to Asia Minor. They 
are still found along the whole of this route. 

Sifnilarly, we are told by the same author, that 
Gonostoma a group of the large genus Helix has a 
number of species in the same south-western district, 
while only one, viz., Helix obvoluta, occurs in England 
and Germany, and two in the Alps. Southward we 
again find many representatives crossing over to 
North Africa, among which Helix lenticula has a 
similar range to Piipa granum, which I have just 
referred to. The Alpine sub-genus Cainpylcea is 
quite absent in the Lusitanian district. 


Among our own British testaceous Land Mollusca, 
several Helices, viz., Helix pisana, ericetorum, virgata, 
acuta, fusca, rotundata, aculeata, and probably many 
others, have come to us from the south-west. The 
species of Hyalinia are undoubtedly of very remote 
origin, and it would be futile at the present state of 
our knowledge to speculate as to their home. Some 
of our species may possibly be of British origin. 
Balea perversa is probably a south-western species, 
and certainly Pupa anglica, which is quite confined to 
Western Europe. 


FIG. 1 8. The Spotted Slug (Geomalacus maculosus}. 

Much more characteristic of South-western Europe, 
however, than these land-shells are some of the slugs. 
The peculiar genus Geomalacus is almost entirely 
confined to Portugal. One species, which I have 
had several occasions to refer to in illustration of the 
term "discontinuous distribution," ranges far beyond 
the confines of that country. This is Geomalacus 
maculosus (Fig. 18), first discovered in the south- 
west of Ireland, and more recently also in Portugal. 
Although careful search has been made for it in 


other parts of the British Islands, this slug has only 
been found in the portion of Ireland just indicated. 
Within the last few years I have taken it, up to a 
height of over a thousand feet, on the promontory 
north of the Kenmare River, also from sea-level up 
to a considerable height near Glengarirf, and more 
recently Messrs. Praeger and Welch discovered it in 
abundance near the town of Kenmare. But beyond 
this rather circumscribed area in the counties of Cork 
and Kerry it does not occur (vide Fig. 19). Several 
Portuguese species of this interesting genus have 
since been added to science by Dr. Simroth and 
others. Dr. Simroth, too, has promulgated the view 
that the genus Arion to which our common brown 
garden slug belongs is of Lusitanian origin. Indeed, 
the number of species of Arion diminishes as we 
leave that province, though one extends beyond the 
borders of Europe into Siberia. The same number of 
species, viz. five, occur in Germany and in England. 
Testacella a slug-like mollusc which lives under- 
ground on earthworms, and of which genus three 
species, viz. T. mangei, T. haliotidea, 7\ scutulum, 
are known to inhabit the British Islands, is another 
Lusitanian animal. All the species are confined to 
\Vestern Europe and North Africa ; they do not even 
reach Germany or Switzerland. 

I have had occasion to mention once before an ex- 
tremely interesting genus of blind Woocllouse, viz., 
Platyarthnis. Like Testacella, it lives underground, 
and also resembles it in its general range. Its distribu- 


tion is therefore of particular interest. It is difficult to 
conceive that Platyarthrus, from its peculiar mode of 
life, could have crossed any formidable barrier, such 

FIG. 19. Map of the British Islands on which the geographical distri- 
bution of Geomalaats maculosus is indicated in black. 

as even a narrow straits of sea. Its occurrence in 
Spain and North Africa indicates, therefore, that the 
Straits of Gibraltar did not exist at the time when 


it undertook the migration southward, just as the 
English Channel and the Irish Sea could not have 
been there when it wandered to England and Ireland. 
The species which occurs in the south of England 
has a wide range in Ireland, and reaches in Scotland 
its most northern European limit of distribution. 
Platyarthrus is only one of the Lusitanian genera of 
woodlice. In Ireland chiefly on the west coast we 
also find a brilliantly coloured Woodlouse, which is 
absent from Great Britain, viz. Metoponorthuscingendus. 
It reappears again on the Continent in the south of 
France. Its range is therefore suggestive of a Lusi- 
tanian origin ; and indeed, when we examine the 
general distribution of the genus Metoponorthus, we 
find that out of the forty-four known species, fully 
one-half arc confined to Western Europe and North 

My friend and colleague, Mr. Carpenter, informs 
me that among the Irish Spiders he is acquainted 
with, the following are to be looked upon as Lusi- 
tanian species : 

Dysdera crocota. Agroeca celans. 
Oonops pulcher. do. gracilipes. 

Tegenaria hibernica. Teutana grossa. 

Theridion aulicum. Cnephalocotes curtus. 

Lasseola inornata. Porrhomma myops. 

Of the Coleoptera, the genera Trichis, Glycia, and 
Singilis, all belonging to the Running Beetles (Cara- 
bidce], are almost confined to the Spanish penin- 


The beetles Rhopalomesites Tardyi, Eurynebria 
complanata, and Otiorrhynchus auropimctatus also 
belong to this fauna, as also the Earthworms Allolo- 
bophora veneta and A. Georgii, and the Millipede 
Polydesmus gallicus. 

It will be evident to every one from these few 
instances of Lusitanian species, that somewhere in 
South-western Europe and. North-western Africa, 
and also, perhaps, in a larger now submerged western 
land-area, there existed an active centre of develop- 
ment, from which animals spread in all directions. 

If the presence of Platyarthrus in North-west Africa 
proves that the Straits of Gibraltar had come into 
existence after its southward migration, it also 
suggests that the ancestral home of this woodlouse 
was in the Spanish peninsula. Whether this sup- 
position is correct or not, does not affect the Straits 
of Gibraltar problem, for in a migration northward 
into Spain from Morocco a land-connection would be 
equally necessary. Almost every group of vertebrates 
and invertebrates furnishes instances of species which 
must have crossed the Straits on dry land. Many 
naturalists have come to this conclusion, and have 
clearly expressed their views on the subject At 
the commencement of the present period, says Mr. 
Bourguignat (p. 354), the north of Africa was a 
peninsula of Spain, the Straits of Gibraltar did not 
exist, and the Mediterranean communicated by the 
Sahara with the Atlantic. 

The faunas of North-west Africa and the south- 


western portion of our continent are so closely 
related, that an uninterrupted intercourse by land 
must have existed for a very long period. The 
Mediterranean, however, throughout the Tertiary 
period at any rate since miocene times must have 
had almost constant communication with the Atlantic. 
According to Professor Suess, this was the case. The 
Atlantic was joined with the Mediterranean across 
the valley of the Guadalquivir during the Miocene 
Epoch, so that Andalusia must have belonged to 
North Africa in those days. The Straits of Gibraltar 
are supposed to have been formed in the next epoch. 
I have already expressed my disagreement with that 
theory from a zoogeographical point of view. The 
old Guadalquivir connection probably persisted much 
longer, though interrupted by temporary periods of 
a partial retreat so as to uncover sufficient land to 
allow of an interchange during miocene as well as 
pliocene times between the European and North 
African faunas. It is in this way, perhaps, that some 
of the members of the Alpine fauna have reached 
Spain by way of Corsica, Sardinia, and North-western 
Africa, and vice versd. The Balearic Islands were 
then connected with Spain; and we find there many 
curious survivals which have long ago become extinct 
on the mainland 

That the Straits of Gibraltar are only of recent 
formation has been suggested on zoogeographical 
evidence by Bourguignat, Simroth, Kobelt, and many 
others. Dr. Kobelt believes that the former land- 


connection between the south of Spain and Morocco 
was much wider than is generally assumed, and that 
the coast-line stretched from Oran in Algeria straight 
across to Cartagena in Spain (^, ii., p. 228). 

My allusions to the lands lying beyond the Lusi- 
tanian province, refer chiefly to the Canary Islands 
and Madeira. Whatever doubts Dr. Wallace had 
on the subject of their former connection with 
Morocco, it cannot be denied that they used to be 
of much larger extent, especially in miocene and 
pliocene times. It seems extremely probable that 
these islands formed part of the mainland of North 
Africa until comparatively recently, and that they 
are the last traces of a sunken continent which united 
Africa and South America. A discussion of this 
problem, however, must be deferred, as it is a com- 
plicated one, and one which would lead me alto- 
gether outside the scope of this little volume. I 
hope I shall have an opportunity to publish my 
views on this subject before long, meanwhile the 
reader must content himself with this mere state- 

During the greater portion of the Miocene, and I 
think for part of the Pliocene Epoch too, the advance 
of the Lusitanian species eastward was barred on 
the continent of Europe by an arm of the sea which 
stretched northward along the Rhone valley from 
the Mediterranean. The Lusitanian forms which 
originated in Southern Spain were able to travel east 
during these times by way of North-west Africa, 


Sicily, Southern Italy, and Greece; and it is possible 

FIG. 20. The Strawberry-tree (Arbutus unedo) in its native habitat in 
the south-west of Ireland. (From a photograph by Robert Welch. ) 

that some may have reached the Alps in this manner, 

and Eastern Europe generally. That the Lusi- 




tanian centre was never a very active one compared 
with, for instance, the Oriental is indicated by many 
distributional facts. It is difficult to understand, 
however, why the Oriental species, on the whole, 
have migrated so far west, while few Lusitanians have 
gone very far east. This seems to have been noted 

FlG. 21. The Irish Spurge {Euphorbia hiberna] in its native habitat 
in the south of Ireland. (From a photograph by Robert Welch.) 

particularly in the case of the flora. Mr. Bonnet drew 
attention to the fact that in Tunis there are none of the 
absolutely characteristic plants of Morocco and Spain, 
while the Oriental flora is represented by a good 
many species. Lusitanian species have spread chiefly 
southward into North Africa, and northward into 
France, the British Islands, and even Scandinavia. 


As I have mentioned in the third chapter, there are a 
good many species of Lusitanian origin in the British 
Islands. However, we have only a mere remnant 
of what we ought to have, had the climate been less 
trying. It is probable, too, that the submergence 
destroyed a good many plants and the insects depen- 
dent on them. That the Lusitanian fauna is very 
ancient in the British Islands is proved by the fact 
of the discontinuous distribution of so many species. 
A greater number survived in Ireland than in 

Altogether and this was strongly urged by 
Edward Forbes the Lusitanian element is the 
oldest of the components of our fauna, and it must 
have poured into the British Islands for many geo- 
logical periods almost without cessation. The same 
author, in his classic essay, refers especially to the 
Lusitanian flora, two prominent members of which 
are the British plants, Arbutus unedo (Fig. 20, p. 305) 
and EupJiorbia hiberna (Fig. 21, p. 306). The former 
has a wide range in the Mediterranean region, and 
occurs in the British Islands only in the south-west 
of Ireland. The Spurge, on the other hand, is also 
found in the south-west of England, besides Ireland 
and Southern Europe, 



The term " Lusitanian " is in this chapter employed in the 
wide sense, as indicating the South-west of Europe and North- 
western Africa. From this centre, and probably also from a 
now sunken land which lay to the west of it, issued a fauna and 
flora of which we have abundant evidence in our own islands, 
especially in Ireland. Edward Forbes held that the Lusitanian 
element of the British flora was of miocene age, and that it 
survived the Glacial period in this country. 

At the time when the Straits of Gibraltar did not exist, and 
when there was free land communication between Asia Minor, 
Greece, and Tunis, many Oriental species migrated westward 
by this ancient Mediterranean route as far as Spain. They 
would then have invaded the more central parts of Europe from 
the south-west, without however being of Lusitanian origin. Of 
the true Lusitanian mammals a typical example is the Rabbit. 
Then we have a few birds and several interesting reptiles and 
amphibians. The genus to which the Brimstone Butterfly 
belongs is also of south-western origin. A number of Mollusca 
are mentioned which from their range likewise indicate a Lusi- 
tanian origin. Most of our British Slugs and many of our 
larger Snails belong to this group. 

All these are merely a small remnant of what we received 
from South-western Europe during the Miocene and Pliocene 
Epochs. But they spread into many parts of Europe, and a 
few even crossed into Asia. The antiquity of the Lusitanian 
element in our fauna is especially indicated by the frequent 
recurrence of "discontinuous distribution" among the species 
belonging to that section. 



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 abun- 
dance 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 any- 
where along the ^Egean 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 main- 
land 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 doubt- 
ful whether many of these species ever reached the 

The fauna of the Alps, and also the flora, is 
therefore made up of a number of component ele- 
ments. 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 sur- 
roundings 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. cegagrus) occurs in 
Asia Minor, Persia, the island of Crete, and some of 
the Cyclades. This exemplifies what I remarked in the 
last chapter about ihe 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 Nemorhcedus 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 mar motto) 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 Moun- 
tains 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. Sqrexalpinus 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 5. minutus (pygmceus) 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 
(vulgar is}> subsequently migrated towards the old 
home of its forefathers, since we find it more or less 
confined to Central and Northern Asia and Northern 

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 advan- 
tages 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. Europceus) 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 sup- 
posed, a "standing testimony of a former arctic 
climate" in Europe, but merely the necessary con- 
sequence of the weaker species being thrust into 
less accessible regions by a stronger rival. 

Muscardinus avellanarhis, the common Dormouse, 
though by no means confined to the Alps, has prob- 
ably 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, Scot- 
land, Norway, and Northern Russia, it seems as 
if it had only diffused northward in more recent 

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 ^gean 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 (Petronza stultd) 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 (Monti- 
fringilla 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 (Sala- 
mandra mac^llosa} 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. caucasicd) 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 Sala- 
mandridcz are certainly Asiatic. Our common Newt 
(Molge vnlgaris] 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 ^Egean 
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. Hagenmillleri, while the 
Moroccan M. Waltlii passes into the south of Spain. 
Here Molge boscce, M. aspera, and M. mannorata 
originated, the latter passing into France. 

Another branch of the Molge tribe turned north- 
ward 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 
temper aria> 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 

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 t 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 Apen- 
nines 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 Heidel- 
berg. 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 differ- 
ing but little from our present forms, while undoubted 
Campylaece are not met with till we reach the upper 

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 discon- 
tinuous 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 dees 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 

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 geo- 
graphical 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 de- 
light, and he generally brings home with him a rich 
harvest of specimens. The more experienced Butter- 
fly 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. mnemo- 
syne is known from the Pyrenees, Sweden, Hungary, 
Sicily, Russia, and Western Asia. One other Par- 
nassius inhabits Europe, viz., P. Nordmanni of the 
Caucasus, but all the remaining species of the genus 
and there arc 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 

Melitcea, 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. mattirna, 
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 Melitcea 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. czthiops are taken at first sight 
for true Arctic migrants. As neither of them, how- 
ever, 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 


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 high- 
way, at a still more recent period. These are Erebia 
cethiops, medusa, ligea, and ambla. 

Only one species of the well-known Polar genus 
(Eneis, viz. (E. 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 CEneis came 
to Europe by the Oriental route. But the Lapland 
species at any rate CE. jutta and (E. 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 

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 Euro- 
pean 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 practi- 
cally 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 impres- 
sions 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. ScJimidti 
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, Den- 
mark, 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 

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 Grass- 
hopper, 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 dis- 
tribution, 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 migra- 
tion. 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 Rein- 
deer 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 con- 
genial 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 con- 
ditions of the climate at that time and the size of 
the Scandinavian glaciers have been greatly ex- 
aggerated, 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 

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 con- 
nected 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 North- 
ern Scandinavian and the Alpine faunas in a pre- 
vious 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 with- 
out 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 Tetraonida. 

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. rnpestris 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 dis- 
tinguishable, 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 specifi- 
cally 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 dis- 
appeared 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 con- 
fined 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 them- 
selves in the Alps. 

The Capercaillie (Telrao urogaUus) 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 pro- 
tected, 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 be- 
longing 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 dis- 
tribution 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 

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 Scandi- 
navia 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 modifica- 
tion or change of species seems to have been pro- 
duced 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 moun- 


tains 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 alpinuvi} 
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 observa- 
tion 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. 53 2 )- 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 main- 
tain 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 fresh- 
water 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 migra- 
tion 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 discon- 
tinuous chain some 1500 miles in length, from the 
Pyrenees to the Eastern Carpathians? Nor would 
the difficulties cease there. You would have left un- 
explained 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 com- 
pletely 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 clim.ate 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 im- 
pression 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. 


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 migra- 
tion. 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 Heli.\\ 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 Lcpidoptera^ 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 versd. 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 ex- 
plained 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. 


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Brehm, A. E. "From North Pole 
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Carpenter, G. II. Problems of the 



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Christ, H. "Das Pflanzenleben 
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Emery, C. On the Origin of Eu- 
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Engler, A. "Versuch einer Ent- 
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Falsan, A." La Periode Glaci- 
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Fischer, P. Faune malacologique 
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Forbes, E. The Geological Rela- 
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the British Isles, etc., "Memoiis 
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Harle, E. Sur la succession de 
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Abkpharus pannonicu s, 258 
Accentor collaHs, 316 

modidaris, 317 

Accidental distribution, 12, 26 
Acipenser ruthenus, 29 
Acme, 323 

Adams, Leith, 112, 150 
/Egean continent, subsidence of, 


Agriolimax, 284 
AlactagajacuhiS) 204 
Aha impennis, 92, 142 
Alligator, accidental dispersal of, 

AHolobophora Georgit, 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- 


Hare, as a test of climate, 316 
Snow-finch. 318 
Alps, component elements of fauna, 


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, 1 66 
sponges in Ireland, 166 
Anipelis garruhts t 205 

Amphibia of Europe, 193-194 

dispersal of, 21 
Aniphicoma, 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, 

Arbutus unedo, IIO, 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 
Arctotnys bobak, 204, 313 

mannoi 7a, 313 
Arion, 48, 299 

snbf^tsc^ls, 49 

Arionidie, 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 
B latins cinereus, 279, 294 
Blelhisa multipunctata, 93 
Blytt, A., 34, 52, 79, 185, 239 
Boar, distribution of, 44 
Bobak Marmot, 204 
Boettger, O. , 48, 263 
Bogdanov, M. M., 53 
Bombinatof, 259-260 
igneus, 259 
pachypus, 259 
Bonnet, 306 
Bonney, T. G., 66, 70, 83, 180, 


Boulenger, G. A., 259 
Boulder-clay, foraminifera in, 84 
origin of, 81, 175, 180-181, 229- 

of Continental Europe, 180, 226- 


Bourguignat, J. R., 18, 47, 302 
Bovey Tracy, arctic plant beds at, 


Boyd, E., 104 
Branchinecta palludosa, 167 
Brandt, J. F., 51, 62, 107, 218, 


Brauer, A., 132-133 

Brehm, A. E., 139 

Bristle-fern, 115 

British flora during Glacial period, 


British Islands, Lusitanian flora in, 
288, 306-307 

submergence of, 127 
Brooke, Sir V., 247, 250 
Brunner von Wattenwyl, 270 
Btifo 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, 

origin of North European, 55 

Caccabis rufa, 29 

Campylaea, sub-genus of Helix, 49, 


Canary Islands, origin of, 19 
Cam's lagopus, 135 
Capercaillie, 28, 143, 336-337 
Capra ibex, 312 

tzgagrus, 61, 312 

pyrenaica, 312 

sibirica, 312 

sinaitica, 312 
Carabus, 199 
Carduelis, 257 

caniceps, 258 

elegans, 257 

major t 257 
Carpenter, G. H., 64, 94-95, 104, 

114, 121-123, 288 > 3 01 
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 
Cervida, origin of, 248 



Cervus dama, 251 

elaphus, 246-250 

giganiciis, 108, 251 
Chaincelcon 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 

biplicala, 47 

faminafa, 47 

Pauli, 48, 263 

Rolphii, 47 
Claviger, 267-268 

tes/acetts, 268 

Climate in Glacial period, 65 8 1, 
127, 149, 182-183 

in Pleistocene Europe, 208-209 
Close, Maxwell, 129 
Cod-fish, origin of, 143 
Coecilianella, 17 
Coenonyjiipha typhon, 93 
Cole, G. J., 83, 106-107 
Coleoptera of Europe, origin of, 


common to Europe and North 

America, 161 
Colias, 266 
Coregonus, 124 

clubeoides, 124 

pollan, 124 

vandesiiiS) 124 
Cosmopolitan species, 24 
Coitus, 92-93 

bubalis, 93 

gobio, 93 

quadricornis, 175 

scorpio, 93 

Credner, R., 177-178 
Cricetus frunifntariuS) 190 

vttlgaris, 204 

Crimea formerly an island, 35 
Crimean fauna, 57 
Croll, J., 66 

Crocidura, 253 

etnisca, 253, 279 
Ctitiiciilus tonjuatus^ 138 
Current, Arctic, 172-173 
Cyanopolius Coo/a, 293 

cyamts, 293 
Cyclostoma elegans, 16 

Danais chrysipftts, 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, II2-II3, 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- 


Drift, a marine deposit, 129 
Drude, O., 52, 77 
Dryas octopetela, 79, 238 
Dual origin, possibility of, 38 
Dyer, Th., 79 

Earth-worms, distribution of, 19, 

23,. 58 

Edelweiss, 266, 342 
Egean Continent, subsidence of, 


Eider-duck, range of, 141 
ElepkttmtidaSi origin of, 202, 252- 


Elephas primigemus, 214, 252 
Emery, C., 161, 167 




Endemic species, 10 

Engler, A., 52, 145, 171, 176, 282, 


English Hare, 29 
Ephydatia craierifonnis, 94 
Epoccus, 167 
Erebia, 325-326 

athiops, 325-326 

epiphron, 325 

glacialis, 326 

lappona, 326 
Eremias, 258 
Erratics, 181-182 
Eryx jacu/us, 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 
Etirynebria coniplanata, 302 
Evotomys Nageri, 314 
Extension of range, mode of, 38 

Fallow Deer, 251 

Falsan, A., 66, 68-69, 7* 73 

Feilden, H. W., 13, 77, 84-85, 

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, no. 114-115, 

171, 288, 339 

Forest-Bed an inter-glacial deposit, 

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 

ccelebs, 293 

madeirensis, 293 

niontifringilla, 293 

spodiogenys, 293 

teydea, 293 

Frog, introduction of, into Ireland, 

Gadow, H., 139 
Galapagos Islands, 19 
Gale odes, 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, 181, 216- 
217, 226-227, 233, 235, 238 
Geographical changes, importance 

of, 64 
Geomalacus maculostis, 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 

ckobule, 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, 1 7 1 

Tertiary plants, 144-145 
Grouse, 91, 334 

black, 336 
Gu/o hiscus, 119, 203 

Ilaacke, 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, 


Harle, E., 150 
Harmer, F. W., 182 
Harvest- mouse, 3, 190 
Harvie-Brown, J. A , 105 
Plaughton, W./86 
Hazel-grouse, 337 
Hedge Accentor, 317 
Hedley, C, 20 
Heer, O., 144-145 
Helix, 4 

actifa, 102, 298 

aculeata, 298 

aspersa, 24, 195 

er ice tor urn, 298 

fusca, 298 

lapicida.) 4 

obvolula> 4, 297 

pi sana, 102, 298 

pomatia, 15, 32, 195, 262 

revelata, 102 

rotundata, 122, 274, 298 

ruder at a, 212-213 
Herdman, W. A., and Lomas, J., 


Herring, origin of, 143 
Heteroineyenia Ryderi, 94 
Hofman, E., 54 
Hooker, Sir J., 99, 161 
Hopatroides thoracictis, 268 
Howorth, Sir H., 73 
Hyalinia, antiquity of, 50 
Hyla, 260 

arborea, 260-261, 280 

chinensisy 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 
hotoma lit tor a Us, 94 

Jeffreys,;. 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, II. 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, 


Koppen, F. T., 51, 62, 222, 248 
Krasan, F., 347 

Lagomys, origin of, 41, 203 
Lagopus, 334 

albus, 91, 142, 334 
hyperboreus, 334 
leucttnts, 335 
mutus, 91, 142, 334 
1-upestris, 334 
scoticus, 334 
LaDiinifera,) sub-genus of Clatisilia^ 


Lamplugh, G. W., 107-108 
Land-connection between America 

and Northern Europe, 61 
British Islands and France, 59- 

British Islands and Scandinavia, 


Europe and North Africa, 61 
Europe and North America, 6 1, 


Greece and Asia Minor, 61 
Greenland and Europe, 155, 159 
Ireland and Spain, no 
Land Mollusca, migrations of, 9 



Land shells, West Indian, 21 
Lartet, E., 51, 73, 107, 150 
Lemming, range of, 138-140 
Leon'.opodium alpinu/n, 266, 342 
Lepus, 27, 29, 31 
) 136 

iS) 27, 31, 291 
us, 29, 316 

glacialis, 136 

lacostei, 291 

variabilts, 2, 29, 136, 315 
Lepidoptera, range of, 159-160 

Arctic, 200 

surviving Glacial period, 54 
Leuckart, R., 176 
Lewis, Carvill, 129 
Liwax, 284 

marginal-its ; 284 

maximus, 284 

Limnocalanus macriirus y ] 79 
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 
Lyrtirtis tetrix^ 336 

Macro ptyckia, sub-genus of Clatt- 
silia, 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 

Marine connection between Caspian 
Sea and Arctic Ocean, 62, 219- 

mollusca, distribution of, 236 
origin of boulder-clay, 82-86, 228- 

Marine shells above sea-level, 127- 

transgression in Northern Russia, 

Marmot, Alpine, 313 

Bobak, 204 
Marsh-ringlet, 93 
Marshall, Rev., id6 
Martins, C, 68 
Mediterranean land- connections, 

Meles, 43 

albogularis, 44 

anakunia, 44 

t'eucurus, 44 

maraghanus, 44 

polaki, 44 

taxus, 43 
Melitcea aster ia, 325 

cynthia, 325 

mattirna, 325 
Melizoph : lus undatus, 288 
Merriam, C. H., 38 
Metoponorthus cingendus, 301 
Microtus, 313 

brecciensis, 314 

leucurus, 314 

nivalis, 313 
Mid- wife Toad, 295 
Migrations, 8-9 

cause of, 208 

of British shore-forms, 124 
Migration from Asia to Europe by 

North America, 327 
Migration on ice-bridge, 108 

waves of, 204-206 
Miller's thumb, 93 
Milne-Edwards, A., 282 
Mingling of Southern and Northern 

Mammals, 72-75, 209 
Miocene geography, 274-276 

temperature in Greenland, 146 
Mi o gale mo s chat a ^ 290 

pyrenaica, 290 
Molge alpestris, 320 

cristata, 320 

montana, 320 

palmata^ 320 

rusconii, 320 

vulgaris, 319 
Mollusca in Loess, 196 



Mollusca, distribution of marine, 236 
Molluscan fauna, divisions of British, 

Madeira, 25 

Porto Santo, 25 
Molluscs, dispersal of, 17, 24 
Mongoose, 28 
Montifringilla nivalis, 318 
More, A. G., 104 
Moschler, E., 160 
Motacillct) 254, 292 

alba, 292 

boreal is, 255 

campestris, 254 

lugubris, 292 

melanope, 254 
Mountain Avens, 79 
Mountain-ringlet, 325 
Mouse, distribution of Harvest, 3 
Mud-glaciers, 86 
Murchison, R., 175, 230 
Murray, Andrew, 32-33, 150, 154 
Muscardinus avellanarius, 316 
Musk-Ox, range of, 119, 134-135, 


Mus minutusy 3, 95, 190 
Miistela africana, 279 

boccamela, 279 

erminea, 135- 136 

putorius, 190 
Myodus lemmuS) 138 

obensis, 138 
Myoxus, 316 
Mysis caspica, 223 

necropthalma, 223 

relicta, 179, 223 

Nathorst, A. G., 163, 169, 240- 


Natterjack Toad, 30 
Nebria atrata, 328 

livida, 328 
Nehring, A., 96, 107, 119, 196, 

208-211, 340 

Nenia, sub-genus of Claiisilia, 48 
Neumayr, M., 19, 67 
Newt, 319 
New Siberian Islands, origin of 

bone-beds, 219 
extinct fauna of, 218 
Nightingale, 192 

Nordquist, 179 

North American marine mollusca 

in crag deposits, 173 
North European Sea, 172 

Ocean basins, permanence of, 18 
Oceanic Islands, 18 
Oeneis, 326-327 

aello, 326 

Onychogomphus, 268 
Oriental migration, old and new, 


Oriental plants, 282-283 
Origin, centre of, shifting, 202 
Otiorrhynchus auropunctatus, 115, 

Ovibos moschatus, 119, 134-135, 203 

Pacific Continent, 20 
Painted-lady Butterfly, 98 
Palincosmic species, 25 
Pallas's Sandgrouse, 205 
Panurus biarmicus, 292 
Fapilio hospifon, 265 
Parmacella, 49 
Parnassitis, 265-266, 324 

apollo, 265, 324 

delitts, 324 

mnemosyne, 324 

Nordmanni, 324 
Partridge, 29 
Pelias beruS) 192 
Pelobates, 295 
Pelodytes punclattts, 295 
Pelophila boreatis, 93 
Penck, A., 66 
Perches, origin of, 143 
Periops hippocrepis, 279 
Peschel, O., 177 
Petersen, VV., 55, 154, 160, 200 
Petronia stulta, 318 
Pezotettix, 329-330 

alpinus, 329 

pedestris, 329 
Phasianus, 256-257 

cokhicus, 31, 257 
Pheasant, origin of, 256-257 
Phoca annelata, 174 

caspica, 224 

Phcedusa, sub-genus of Clausili 



Phylloxera vastalrix, 24 
Pigs, origin of, 44-46 
Pika, 41 

Pine-Grosbeak, 191 
Pinicola enucleator, 191 
Planorbis dilatetus, 24 

glaber, 167 
Plants, American, in Ireland, 166 

as tests of climate, 75-80 

migration of, 34 
Platyarthrus, 299-302 
Plectrophenax nivalis, 140 
Pleistocene climate of Asia, 210, 

Europe, climate in, 208-209 

fauna, northern and southern 
animals in, 72-75 

mollusca, 211-213 
Pliocene deposits of Sicily, 277 

geography, 276-277 
Pohlig, H., 176 
Polar Bear, 134 

Fox, range of, 135, 149 

origin of animals, 147 
Pole-cat, 190 

Polydesnius gallicus, 115, 302 
Polynesian Islands, 20 
Pomalias, 49, 321-322 
Pontoporeia affiu/s, 179 
Porto Santo, molluscan fauna of, 25 
Pouched Marmot, 41 
Praeger, R. L , 298 
Praying insect, 269 
Proboscidea, origin of, 252 
Psanirnodromus algirus, 295 

hispanicuS) 295 
Ptarmigan, 91, 142 
Ptipa, antiquity of, 50 
Pupa, 297-298 

anglica, 298 
granum, 297 

ringens, 102 

Pyrrhocorax alpimis, 317 
Pyrrhula, 191, 255 

e^^rop<za, 191, 256 

major, 191, 256 

Rabbit, introduction of, 27, 31, 291 
Rana temporaria, 194, 320 
Range, extension of, 38 
Rangifer tarandus, 133, 150-158 

Reade, Mellard, 127-129 

Red Crag, Arctic mollusca in, 235- 


Red Deer, 246-250 
Red Grouse, 91, 142 
Red-legged Partridge, 29 
Reguhis ignicapilhis, 257 
maderensis, 257 
teneriffiz, 257 
Reid, Clement, 76, 163 
Reindeer, range and varieties of, 91, 

133, 149-158 

Relict lakes, 176-179, 223 
Reptiles, dispersal of, 21 
Reptiles of Europe, 192-193 
Rhax, 270 

Rhinoceros, distribution of, 214-216 
Rhododendron ponticum^ 271 
Rhopalomesites Tardyi, 302 
Rock Sparrow, 318 
Route of migration of Siberian 

mammals, 210 
Rupicapra tragus, 312 
Russia, marine transgression in, 


submergence of North, 155 
RiUimeyer, L., 43, 57, 281 
Ryothemis, 269 

Saiga Antelope, 39, 204 
Saiga tartarica, 39, 204 
Salamandra atra, 319 
caitcasica, 319 
maculosa, 319 

Salnionidce, origin of, 92, 143 
Sandgrouse, migration of Pallas's, 

41, 205 

Sars, O., 177, 223-224 
Saunders, H., 256 
Seal, Arctic, 174 
Sea-urchin, 125 
Sedgwick, A., 86 
Scandinavia, absence of Oriental 

and Siberian mammals from, 

176, 206 

during Glacial period, 176 
Scandinavian boulders, origin of, 


butterflies in America, 167 
flora, antiquity of, 162, 185 
glaciers, 172 



Scandinavian lepidopterous fauna, 55 

Schulz, A., 52, 155 

Sclater, P. L., 95 

Scotch Argus, 325 

Scotch and Irish Hare, 136 

Siberia, climate of, during Glacial 

period, 217 

Siberian birds in British Islands, 

mammals absent from Scandi- 
navia, 176, 206 

mammals absent from Southern 
Europe, 206 

mammals in Great Britain, 95-96, 
190, 202-204 

migration, date of, 208 

steppe-fauna, 39 
Sicily, Pliocene deposits in, 277 
Silurus glanis, 29 
Simpson, C. T., 21 
Simroth, H. , 48, 299 
Sjogren, H., 226 
Slugs, dispersal of, 16-17 
Snails, dispersal of, 17 
Snow-Bunting, range of, 91, 140 
Snow-Finch, 318 
Sollas, W., 177 
Somateria mollissima, 141 
Sorex^ 202, 3 14-3 1 5 

alpinus, 3 14-3 1 5 

araneus, 315 

minulus, 315 
Southern mammals in Arctic Siberia, 


Sparrow, introduction of, 28 
Spcmtop1tilus t 41 
Speyer, A. and A., 55, 200 
Spiders, Lusitanian, 301 
Sponges, American, in Ireland, 166 
St. Erth fauna, 174 
Steinbock, 312 
Steppe-fauna, Siberian, 39 
Sterlet, introduction of, 29 
Stick insect, 269 
Stickleback, origin of, 92, 143, 166- 


Stoat, range of, 90-91, 135-136 
Strails of Gibraltar, age of, 277- 

278, 281 

Strix Jlammea> 98 
Stronglyocentrotus lividus, 125 

Struckmann, C. , 153 
Studer, Th., 340 

Sub-marine plateau between Nor- 
way and Spitsbergen, 154 
Suess, E., 61, 273, 282, 303 
Stts scrofa, 44-46 
Swallow-tail Butterfly, 264-265 
Syrrhaptes paradoxus, 41, 205 

Tailless Hare, 41 
Talpa, 291 
Taylor, J. W., 105 
Tcherski, J. D., 214-215, 222 
Tertiary plants in Greenland, 144- 

Test ace I la, 17, 299 

haliotidea, 299 

maugeii 102, 299 

scutulum.) 299 
Tetrao, 28 

urogallus, 28, 143, 336-337 
Tetrastes bonasia, 337 
Tettix, 330 

bipunctatus, 330 
Thais cerisyi) 265 
Thelphusa fluviatilis, 270, 281 
Thomas, O., 90 
Thompson, W., 32 
Time of disconnection between 
Great Britain and Ireland, 63 
Titmouse, bearded, 292 
Toad, Natter-jack, 30 
Tree-frog, 260, 280 
Trichomanes radicans, 115 
Tropidonotus viperimts, 294 
Tubclfa pensylvam'ca, 94 
Tundras in Northern Europe, 209- 


Turnix sylvatica, 291-292 
Tyndall, J., 68 
Typhlops lumbricalis, 259 
Tyrrhenian Continent, 280 

Unio(Maigaritana} margaritifer^ 
Ursus maritinwS) 134 
Ussher, R. J., 92, 105 

Vanessa cardui^ 98 

Vertigo alpestris, 93 
Viper, range of, 192 
Vole, range of, 313-314 



Wag-tails, origin and range of, 254, 

Wallace, A. R., 12-13, 18-19, 23, 

58, 99, in, 304 
Warming, E., 76, 163 
Watson, H. C., 100 
Wax-bill, migrations of, 205 
Welch, R., 298 
West-Indian land-shells, 21 

White, Buchanan, 109, 113 
Willow-grouse, 91, 142 
Woodward, B. B. (vide Kennard 

and Woodward) 
Wright, J., 84 

Zittel, 252 

Zonites, 49, 195, 322-323 



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