Island Life; Or, The Phenomena and Causes of Insular Faunas and Floras. Alfred Russel Wallace
Newberry. Thus an enormous range in time is accompanied by a very wide and scattered distribution of the existing species.
Whenever, therefore, we find two or more living genera belonging to the same family or order but not very closely allied to each other, we may be sure that they are the remnants of a once extensive group of genera; and if we find them now isolated in remote parts of the globe, the natural inference is that the family of which they are fragments once had an area embracing the countries in which they are found. Yet this simple and very obvious explanation has rarely been adopted by naturalists, who have instead imagined changes of land and sea to afford a direct passage from the one fragment to the other. If there were no cosmopolitan or very wide-spread families still existing, or even if such cases were rare, there would be some justification for such a proceeding; but as about one-fourth of the existing families of land mammalia have a range extending to at least three or four continents, while many which are now represented by disconnected genera are known to have occupied intervening lands or to have had an almost continuous distribution in tertiary times, all the presumptions are in favour of the former continuity of the group. We have also in many cases direct evidence that this former continuity was effected by means of existing continents, while in no single case has it been shown that such a continuity was impossible, and that it either was or must have been effected by means of continents now sunk beneath the ocean.
Concluding Remarks.—When writing on the subject of distribution it usually seems to have been forgotten that the theory of evolution absolutely necessitates the former existence of a whole series of extinct genera filling up the gap between the isolated genera which in many cases now alone exist; while it is almost an axiom of "natural selection" that such numerous forms of one type could only have been developed in a wide area and under varied conditions, implying a great lapse of time. In our succeeding chapters we shall show that the known and probable changes of sea and land, the known changes of climate, and the actual powers of dispersal of the different groups of animals, were such as would have enabled all the now disconnected groups to have once formed parts of a continuous series. Proofs of such former continuity are continually being obtained by the discovery of allied extinct forms in intervening lands, but the extreme imperfection of the geological record as regards land animals renders it unlikely that this proof will be forthcoming in the majority of cases. The notion that if such animals ever existed their remains would certainly be found, is a superstition which, notwithstanding the efforts of Lyell and Darwin, still largely prevails among naturalists; but until it is got rid of no true notions of the former distribution of life upon the earth can be attained.
CHAPTER V
THE POWERS OF DISPERSAL OF ANIMALS AND PLANTS
Statement of the general question of Dispersal—The Ocean as a Barrier to the Dispersal of Mammals—The Dispersal of Birds—The Dispersal of Reptiles—The Dispersal of Insects—The Dispersal of Land Mollusca—Great Antiquity of Land-shells—Causes favouring the Abundance of Land-shells—The Dispersal of Plants—Special adaptability of Seeds for Dispersal—Birds as agents in the Dispersal of Seeds—Ocean Currents as agents in Plant Dispersal—Dispersal along Mountain-chains—Antiquity of Plants as affecting their Distribution.
In order to understand the many curious anomalies we meet with in studying the distribution of animals and plants, and to be able to explain how it is that some species and genera have been able to spread widely over the globe, while others are confined to one hemisphere, to one continent, or even to a single mountain or a single island, we must make some inquiry into the different powers of dispersal of animals and plants, into the nature of the barriers that limit their migrations, and into the character of the geological or climatal changes which have favoured or checked such migrations.
The first portion of the subject—that which relates to the various modes by which organisms can pass over wide areas of sea and land—has been fully treated by Sir Charles Lyell, by Mr. Darwin, and many other writers, and it will only be necessary here to give a very brief notice of the best known facts on the subject, which will be further referred to when we come to discuss the particular cases that arise in regard to the faunas and floras of remote islands. But the other side of the question of dispersal—that which depends on geological and climatal changes—is in a far less satisfactory condition, for, though much has been written upon it, the most contradictory opinions still prevail, and at almost every step we find ourselves on the battle-field of opposing schools in geological or physical science. As, however, these questions lie at the very root of any general solution of the problems of distribution, I have given much time to a careful examination of the various theories that have been advanced, and the discussions to which they have given rise; and have arrived at some definite conclusions which I venture to hope may serve as the foundation for a better comprehension of these intricate problems. The four chapters which follow this are devoted to a full examination of these profoundly interesting and important questions, after which we shall enter upon our special inquiry—the nature and origin of insular faunas and floras.
The Ocean as a Barrier to the Dispersal of Mammals.—A wide extent of ocean forms an almost absolute barrier to the dispersal of all land animals, and of most of those which are aerial, since even birds cannot fly for thousands of miles without rest and without food, unless they are aquatic birds which can find both rest and food on the surface of the ocean. We may be sure, therefore, that without artificial help neither mammalia nor land birds can pass over very wide oceans. The exact width they can pass over is not determined, but we have a few facts to guide us. Contrary to the common notion, pigs can swim very well, and have been known to swim over five or six miles of sea, and the wide distribution of pigs in the islands of the Eastern Hemisphere may be due to this power. It is almost certain, however, that they would never voluntarily swim away from their native land, and if carried out to sea by a flood they would certainly endeavour to return to the shore. We cannot therefore believe that they would ever swim over fifty or a hundred miles of sea, and the same may be said of all the larger mammalia. Deer also swim well, but there is no reason to believe that they would venture out of sight of land. With the smaller, and especially with the arboreal mammalia, there is a much more effectual way of passing over the sea, by means of floating trees, or those floating islands which are often formed at the mouths of great rivers. Sir Charles Lyell describes such floating islands which were encountered among the Moluccas, on which trees and shrubs were growing on a stratum of soil which even formed a white beach round the margin of each raft. Among the Philippine Islands similar rafts with trees growing on them have been seen after hurricanes; and it is easy to understand how, if the sea were tolerably calm, such a raft might be carried along by a current, aided by the wind acting on the trees, till after a passage of several weeks it might arrive safely on the shores of some land hundreds of miles away from its starting-point. Such small animals as squirrels and field-mice might have been carried away on the trees which formed part of such a raft, and might thus colonise a new island; though, as it would require a pair of the same species to be thus conveyed at the same time, such accidents would no doubt be rare. Insects, however, and land-shells would almost certainly be abundant on such a raft or island, and in this way we may account for the wide dispersal of many species of both these groups.
Notwithstanding the occasional action of such causes, we cannot suppose that they have been effective in the dispersal of mammalia as a whole; and 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. But a considerable number of identical mammalian families and even genera are actually found in all the great continents, and the present distribution of land upon the globe renders it easy to see how they have been able to disperse themselves so widely. All the great land masses radiate from the arctic regions as a common centre, the only break being at Behrings Strait, which is so shallow that a rise of less than a thousand feet would form a broad isthmus connecting Asia and America as far south as the parallel of 60° N. Continuity of land therefore may be said to exist already for all parts of the world (except Australia and a number of large islands, which will be considered separately), and we have thus no difficulty in the way of that former wide diffusion of many groups, which we maintain to be the only explanation of most anomalies of distribution other than such as may be connected