The History of the European Fauna

by R. F. Scharff

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 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 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 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 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 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 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-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-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 unassailable. Nevertheless, I feel that his theories regarding the origin of the faunas of oceanic islands require revision.