The History of the European Fauna.

by R. F. Scharff.

PREFACE.

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 comprehensive 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 Geographical Distribution of Species. Collectors of Beetles, b.u.t.terflies, Sh.e.l.ls, 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 revolutions which have moulded our islands and continents. To geographers, a survey of some of the more important changes in the distribution of land and water in past times--based upon the composition 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. lvi., 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 a.s.sisted 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 ill.u.s.trations 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 _Lepus timidus_. After each chapter I have endeavoured to give a short summary of contents, while a bibliography of the princ.i.p.al 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.

Gunther and Dr. Boulenger. The valuable works on Mammalia by Sir W.

Flower, Mr. Lydekker, Mr. Greve, 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 beautiful photographs. The Council of the Royal Irish Academy also kindly gave me permission to reprint the maps used in ill.u.s.tration of my paper.

Professor Haddon first suggested my writing this book, and gave me many useful hints; and great a.s.sistance 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 grat.i.tude.

R. F. SCHARFF.

THE HISTORY OF THE EUROPEAN FAUNA.

CHAPTER I.

INTRODUCTION.

Every student of natural history, whether he be interested in birds, b.u.t.terflies, or sh.e.l.ls, 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 b.u.t.terfly in England 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 b.u.t.terflies, 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 b.u.t.terflies 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 indicated later on.

Mammals and Snails tell us their story more plainly. The bones of the former and the sh.e.l.ls of snails are easily preserved, and thus furnish us with the necessary 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 discovered 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 evidence 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 a.s.sume this, because its remains have never been found in high northern lat.i.tudes; 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 apparently 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, throughout 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, therefore, 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)[1] as occurring in the cave deposit known as the Ichtham fissure, and also from 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 maculosus_).[2] 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.

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, 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 princ.i.p.al areas inhabited at the present time by the species of northern, southern, and eastern origin (Fig. 1).

[Ill.u.s.tration: Fig. 1.--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 correspond 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 "diagrams," and not faithful representations of all the geographical revolutions witnessed by some of our remote forefathers at any particular period.

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 geological 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, Scandinavia, 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 follow.

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 extremes 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 surroundings, whilst mammals are comparatively short-lived. Being driven from one country to another, and exposed to innumerable enemies, new types appear and old ones rapidly vanish; in fact, there are almost constant changes in the mammalian fauna as we pa.s.s 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 investigations 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[3] element, which 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 princ.i.p.al objects of this work is to show how the autochthonous animals of Europe, _i.e._, 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 distribution of living and extinct organisms alone.

A terrestrial mammal like the deer can, under ordinary circ.u.mstances, only reach one part of a country from another by walking or running to it; but a beetle, such as the c.o.c.kchafer, has two different modes of progression. It may walk or fly. In both, however, there is a third mode of transport--an involuntary 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 pa.s.sive means of distribution 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, unsuitability 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 sp.a.w.n, 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 accidental transportal takes place, but the question is whether the accidentally transported animals arrive living and reach a spot where suitable food is procurable, 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 animal.

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 sh.o.r.e, 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 sh.o.r.es we may witness the occasional arrival of such vegetable products 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 possibility of a pa.s.sive 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-sh.e.l.ls 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-sh.e.l.ls, when hibernating and having a membranous diaphragm over the mouth of the sh.e.l.l, might be floated in c.h.i.n.ks 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 sh.e.l.l, the _Helix 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 sh.e.l.l 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-sh.e.l.ls, belonging to ten species, in a box pierced with holes, and immersed it for a fortnight in the sea. Out of the hundred sh.e.l.ls, twenty-seven recovered. The presence of an operculum 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-sh.e.l.ls have often been thus transported; the feet of birds offer a more probable method."

We have here positive evidence that such sh.e.l.ls 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 sh.o.r.e, 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-sh.e.l.l of the type of _Cyclostoma elegans_, it may be asked, with some justification, what chance slugs or the smaller non-operculated species would have to reach an island like Ireland alive from the mainland, and to colonise it successfully.

Both slugs and their eggs are killed by a short immersion in sea-water, as I have proved experimentally. 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."

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 maritime 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 were strongly supported by Wallace, who defines these islands (p. 243) as those which are of volcanic or coralline formation usually far from continents, entirely without indigenous land mammals or amphibians, but with a fair number of birds and insects, and usually with some reptiles.

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