The History of the European Fauna Part 4

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 _Salmonidae_--are typical northern immigrants. The Stickleback (_Gasterosteus 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 _Cottus_ as far south as California and j.a.pan, 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 _Cottus_, 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 Arctic.

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 succeeding 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 sh.e.l.l, and our freshwater pearl-mussel, _Unio (Margaritana) margaritifer_, belong to this migration. Then among b.u.t.terflies we may cite the Marsh-ringlet (_Coenonympha typhon_), and among beetles, _Pelophila borealis_ and _Blethisa multipunctata_. 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 ill.u.s.tration 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 pensylvanica_, 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 contribution to this subject (_a_, p. 475) I a.s.sumed 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, however, 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 circ.u.mpolar distribution." We have indeed quite similar examples in the Oriental migration, of which part is very ancient, surviving here and there and exhibiting discontinuous distribution. 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.

Gulo luscus.

* Mustela erminea.

* " putorius.

* " vulgaris.

* Sorex vulgaris.

Lagomys pusillus.

* Castor fiber.

Spermophilus Eversmanni.

" erythrogenoides.

* Mus minutus.

* Arvicola agrestis.

* " amphibius.

" arvalis.

* " glareolus.

" gregalis.

" ratticeps.

Equus caballus.

Saiga tartarica.

Ovibos moschatus.

Cricetus songarus.

Myodes lemmus.

Cuniculus torquatus.

Alces latifrons.

" machlis.

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 before, 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 significant 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 circ.u.mstances, 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 percentage to the British fauna. The bulk of that fauna is derived from the various European centres of dispersal, 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 remembered that they need not necessarily have come from what is known among zoologists as the "Oriental Region."

The terms "Oriental animals" and "Oriental 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 s.p.a.ce of time, as in the case of the Siberian invasion; the immeasurable ages which pa.s.sed, 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 enormous 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 flammea_) and the Painted Lady b.u.t.terfly (_Vanessa cardui_).

A great number of our British Mammals, Birds, b.u.t.terflies, 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 sh.o.r.es 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 number 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 _Geomalacus 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 hybrids.

Just as we distinguish in the British Islands the parts inhabited by Englishmen, Scotchmen, and Irishmen, 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 occupied 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 a.s.semblages 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 cla.s.sical essay, and remains one of the most remarkable 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-west } and West of Ireland. } Lusitanian type.

II. South-west of England, and } South-east of Ireland. } Gallican type.

III. South-east of England.

IV. Mountains of Scotland, c.u.mberland, } and Wales. } Scandinavian type.

V. General Flora. Germanic type.

Professor Forbes points out, in connection with the plants of the Germanic type, that the fauna accompanying 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, according 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 molluscs which are almost entirely absent from the first, such as _Geomalacus maculosus_, _Testacella Maugei_, _Helix pisana_, _Helix revelata_, _Helix acuta_, and _Pupa ringens_. These are all of Lusitanian origin, and do not occur in Central Europe. Scotland alone cannot be cla.s.sed 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 cla.s.sification is of particular interest, since the first district represents part of a larger Atlantic province, the second a portion of the Germanic province of the European sub-region. The latter he looks upon as one of the sub-regions of the great Palaearctic 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 provinces belong to Mr. Jordan's greater Germanic Region (p. 302).

In the collection ill.u.s.trating the geographical distribution 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 undetermined origin.