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Island Life/IV

CHAPTER IV

EVOLUTION THE KEY TO DISTRIBUTION

 * Importance of the Doctrine of Evolution—The Origin of New Species—Variation in Animals—The Amount of Variation in North American Birds—How New Species arise from a Variable Species—Definition and Origin of Genera—Cause of the Extinction of Species—The Rise and Decay of Species and Genera—Discontinuous Specific Areas, why Rare—Discontinuity of the Area of Parus Palustris—Discontinuity of Emberiza Schœniclus—The European and Japanese Jays—Supposed Examples of Discontinuity among North American Birds—Distribution and Antiquity of Families—Discontinuity a proof of Antiquity—Concluding Remarks.

In the preceding chapters we have explained the general nature of the phenomena presented by the distribution of animals, and have illustrated and defined the new geographical division of the earth which is found best to agree with them. Before we go further into the details of our subject, and especially before we attempt to trace the causes which have brought about the existing biological relations of the islands of the globe, it is absolutely necessary to have a clear comprehension of the collateral facts and general principles to which we shall most frequently have occasion to refer. These may be briefly defined as, the powers of dispersal of animals and plants under different conditions, such as geological and climatal changes, and the origin and development of species and groups by natural selection. This last is of the most fundamental importance, and its bearing on the dispersal of animals has been much neglected. We therefore devote the present chapter to its consideration.

As we have already shown in our first chapter that the distribution of species, of genera, and of families, present almost exactly the same general phenomena in varying degrees of complexity, and that almost all the interesting problems we have to deal with depend upon the mode of dispersal of one or other of these; and as, further, our knowledge of most of these groups, in the higher animals at least, is confined to the tertiary period of geology, it is therefore unnecessary for us to enter into any questions involving the origin of more comprehensive groups, such as classes or orders. This enables us to avoid most of the disputed questions as to the development of animals, and to confine ourselves to those general principles regulating the origin and development of species and genera which were first laid down by Mr. Darwin thirty years ago, and have now come to be adopted by naturalists as established propositions in the theory of evolution.

The Origin of New Species.—How, then, do new species arise, supposing the world to have been, physically, much as we now see it; and what becomes of them after they have arisen? In the first place we must remember that new species can only be formed when and where there is room for them. If a continent is fully stocked with animals, each species being so well adapted for its mode of life that it can overcome all the dangers to which it is exposed, and maintain on the average a tolerably uniform population, then, so long as no change takes place, no new species will arise. For every place or station is supposed to be filled by creatures in all respects adapted to surrounding conditions, able to defend themselves from all enemies, and to obtain food notwithstanding the rivalry of many competitors. But such a perfect balance of organisms nowhere exists upon the earth, and probably never has existed. The well-known fact that some species are very common, while others are very rare, is an almost certain proof that the one is better adapted to its position than the other; and this belief is strengthened when we find the individuals of one species ranging into different climates, subsisting on different food, and competing with different sets of animals, while the individuals of another species will be limited to a small area beyond which they seem unable to extend. When a change occurs, either of climate or geography, some of the small and ill-adapted species will probably die out altogether, and thus leave room for others to increase, or for new forms to occupy their places.

But the change will most likely affect even flourishing species in different ways, some beneficially, others injuriously. Or, again, it may affect a great many injuriously, to such an extent as to require some change in their structure or habits to enable them to get on as well as before. Now "variation" and the "struggle for existence" come into play. All the weaker and less perfectly organised individuals die out, while those which vary in such a way as to bring them into more harmony with the new conditions constantly survive. If the change of conditions has been considerable, then, after a few centuries, or perhaps even a few generations, one or more new species will be almost sure to be formed.

Variation in Animals.—To make this more intelligible to those who have not considered the subject, and to obviate the difficulty many feel about "favourable variations occurring at the right time," it will be well to discuss this matter a little more fully. Few persons consider how largely and universally all animals are varying. We know, however, that in every generation, if we could examine all the individuals of any common species, we should find considerable differences, not only in size and colour, but in the form and proportions of all the parts and organs of the body. In our domesticated animals we know this to be the case, and it is by means of the continual selection of such slight varieties to breed from that all our extremely different domestic breeds have been produced. Think of the difference in every limb, and every bone and muscle, and probably in every part, internal and external of the whole body, between a greyhound and a bull-dog! Yet, if we had the complete series of ancestors of these two breeds before us, we should probably find that in no one generation was there a greater difference than now occurs in the same breed, or sometimes even the same litter. It is often thought, however, that wild species do not vary sufficiently to bring about any such change as this in the same time; and though naturalists are well aware that this is a mistake, it is only recently that they have been able to adduce positive proof of their opinion.

The Amount of Variation in North American Birds.—An American naturalist, Mr. J. A. Allen, has made elaborate observations and measurements of the birds of the United States, and he finds a wonderful and altogether unsuspected amount of variation between individuals of the same species. They differ in the general tint, and in the markings and distribution of the colours; in size and proportions; in the length of the wings, tail, bill, and feet; in the length of particular feathers, altering the shape of the wing or tail; in the length of the tarsi and of the separate toes, and in the length, width, thickness, and curvature of the bill. These variations are very considerable, often reaching to one-sixth or one-seventh of the average dimensions, and sometimes more. Thus Turdus fuscescens (Wilson's thrush) varied in length of wing from 3·58 to 4·16 inches, and in the tail from 3·55 to 4·00 inches; and in twelve specimens, all taken in the same locality, the wing varied in length from 14·5 to 21 per cent., and the tail from 14 to 22·5 per cent. In Sialia sialis (the blue bird) the middle toe varied from ·77 to ·91 inch, and the hind toe from ·58 to ·72 inch, or more than 21·5 per cent. on the mean, while the bill varied from ·45 to ·56 inch in length, and from ·30 to ·38 inch in width, or about 20 per cent. in both cases. In Dendrœca coronata (the yellow-crowned warbler) the quills vary in proportionate length, so that the 1st, the 2nd, the 3rd, or the 4th, is sometimes longest; and a similar variation of the wing involving a change of proportion between two or more of the feathers is recorded in eleven species of birds. Colour and marking vary to an equal extent; the dark streaks on the under surface of Melospiza melodia (the American song-sparrow) being sometimes reduced to narrow lines. while in other specimens they are so enlarged as to cover the greater part of the breast and sides of the body, sometimes uniting on the middle of the breast into a nearly continuous patch. In one of the small spotted wood-thrushes, Turdus fuscescens, the colours are sometimes very pale, and the markings on the breast reduced to indistinct narrow lines, while in other specimens the general colour is much darker, and the breast markings dark, broad, and triangular. All the variations here mentioned occur between adult males, so that there is no question of differences of age or sex, and the pair last referred to were taken at the same place and on the same day.

These interesting facts entirely support the belief in the variability of all animals in all their parts and organs, to an extent amply sufficient for natural selection to work with. We may, indeed, admit that these are extreme cases, and that the majority of species do not vary half or a quarter so much as shown in the examples quoted, and we shall still have ample variation for all purposes of specific modification. Instead of an extreme variation in the dimensions and proportions of the various organs of from 10 to 25 per cent. as is here proved to occur, we may assume from 3 to 6 per cent. as generally occurring in the majority of species; and if we further remember that the above excessive variations were found by comparing a number of specimens of each species, varying from 50 to 150 only, we may be sure that the smaller variations we require must occur in considerable numbers among the thousands or millions of individuals of which all but the very rare species consist. If, therefore, we were to divide the population of any species into three groups of equal extent, with regard to any particular character—as length of wing, or of toes, or thickness or curvature of bill, or strength of markings—we should have one group in which the mean or average character prevailed with little variation, one in which the character was greatly, and one in which it was little, developed. If we formed our groups, not by equal numbers, but by equal amount of variation, we should probably find, in accordance with the law of averages, that the central group in which the mean characteristics prevailed was much more numerous than the extremes, perhaps twice, or even three times, as great as either of them, and forming such a series as the following—10 maximum, 30 mean, 10 minimum development. In ordinary cases we have no reason to believe that the mean characters or the amount of variation of a species changes materially from year to year or from century to century, and we may therefore look upon the central group as the type of the species which is best adapted to the conditions in which it has actually to exist. This type will therefore always form the majority, because the struggle for existence will lead to the continual suppression of the less perfectly adapted extremes. But sometimes a species has a wide range into countries which differ in physical conditions, and then it often happens that one or other of the extremes will predominate in a portion of its range. These form local varieties, but as they occur mixed with the other forms, they are not considered to be distinct species, although they may differ from the other extreme form quite as much as species often do from each other.

How New Species arise from a Variable Species.—It is now very easy to understand how, from such a variable species, one or more new species may arise. The peculiar physical or organic conditions that render one part of the area better adapted to an extreme form may become intensified, and the most extreme variations thus having the advantage, they will multiply at the expense of the rest. If the change of conditions spreads over the whole area occupied by the species, this one extreme form will replace the others; while if the area should be cut in two by subsidence or elevation, the conditions of the two parts may be modified in opposite directions, so as to be each adapted to one extreme form; in which case the original type will become extinct, being replaced by two species, each formed by a combination of certain extreme characters which had before existed in some of its varieties.

The changes of conditions which lead to such selection of varieties are very diverse in nature, and new species may thus be formed, diverging in many ways from the original stock. The climate may change from moist to dry, or the reverse, or the temperature may increase or diminish for long periods, in either case requiring a corresponding change of constitution, of covering, of vegetable or of insect food, to be met by the selection of variations of colour or of swiftness, of length of bill or of strength of claws. Again, competitors or enemies may arrive from other regions, giving the advantage to such varieties as can change their food, or by swifter flight or greater wariness can escape their new foes. We may thus easily understand how a series of changes may occur at distant intervals, each leading to the selection and preservation of a special set of variations, and thus what was a single species may become transformed into a group of allied species differing from each other in a variety of ways, just as we find them in nature.

Among these species, however, there will be some which will have become adapted to very local or special conditions, and will therefore be comparatively few in number and confined to a limited area; while others, retaining the more general characters of the parent form, but with some important change of structure, will be better adapted to succeed in the struggle for existence with other animals, will spread over a wider area, and increase so as to become common species. Sometimes these will acquire such a perfection of organisation by successive favourable modifications that they will be able to spread greatly beyond the range of the parent form. They then become what are termed dominant species, maintaining themselves in vigour and abundance over very wide areas, displacing other species with which they come into competition, and, under still further changes of conditions, becoming the parents of a new set of diverging species.

Definition and Origin of Genera.—As some of the most important and interesting phenomena of distribution relate to genera rather than to single species, it will be well here to explain what is meant by a genus, and how genera are supposed to arise.

A genus is a group of allied species which differs from all other groups in some well marked characters, usually of a structural rather than a superficial nature. Species of one genus usually differ from each other in size, in colour or marking, in the proportions of the limbs or other organs, and in the form and size of such superficial appendages as horns, crests, manes, &c.; but they generally agree in the form and structure of important organs, as the teeth, the bill, the feet, and the wings. When two groups of species differ from each other constantly in one or more of these latter particulars they are said to belong to different genera. We have already seen that species vary in these more important as well as in the more superficial characters. If, then, in any part of the area occupied by a species some change of habits becomes useful to it, all such structural variations as facilitate the change will be accumulated by natural selection, and when they have become fixed in the proportions most beneficial to the animal, we shall have the first species of a new genus.

A creature which has been thus modified in important characters will be a new type, specially adapted to fill a new place in the economy of nature. It will almost certainly have arisen from an extensive or dominant species, because only such are sufficiently rich in individuals to afford an ample supply of the necessary variations, and it will inherit the vigour of constitution and adaptability to a wide range of conditions which gave success to its ancestors. It will therefore have every chance in its favour in the struggle for existence; it may spread widely and displace many of its nearest allies, and in doing so will itself become modified superficially and become the parent of a number of subordinate species. It will now have become a dominant genus, occupying an entire continent, or perhaps even two or more continents, spreading in every direction till it comes in contact with competing forms better adapted to the different environments. Such a genus may continue to exist during long geological epochs; but the time will generally come when either physical changes, or competing forms, or new enemies are too much for it, and it begins to lose its supremacy. First one then another of its component species will dwindle away and become extinct, till at last only a few species remain. Sometimes these soon follow the others and the whole genus dies out, as thousands of genera have died out during the long course of the earth's life-history; but it will also sometimes happen that a few species will continue to maintain themselves in areas where they are removed from the influences that exterminated their fellows.

Cause of the Extinction of Species.—There is good reason to believe that the most effective agent in the extinction of species is the pressure of other species, whether as enemies or merely as competitors. If therefore any portion of the earth is cut off from the influx of new or more highly organised animals, we may there expect to find the remains of groups which have elsewhere become extinct. In islands which have been long separated from their parent continents these conditions are exactly fulfilled, and it is in such places that we find the most striking examples of the preservation of fragments of primeval groups of animals, often widely separated from each other, owing to their having been preserved at remote portions of the area of the once widespread parental group. There are many other ways in which portions of dying out groups may be saved. Nocturnal or subterranean modes of life may save a species from enemies or competitors, and many of the ancient types still existing have such habits. The dense gloom of equatorial forests also affords means of concealment and protection, and we sometimes find in such localities a few remnants of low types in the midst of a general assemblage of higher forms. Some of the most ancient types now living inhabit caves like the Proteus, or bury themselves in mud like the Lepidosiren, or in sand like the Amphioxus, the last being the most primitive of all vertebrates; while the Galeopithecus and Tarsius of the Malay islands and the potto of West Africa, survive amid the higher mammalia of the Asiatic and African continents owing to their nocturnal habits and concealment in the densest forests.

The Rise and Decay of Species and Genera.—The preceding sketch of the mode in which species and genera have arisen, have come to maturity, and then decay, leads us to some very important conclusions as to the mode of distribution of animals. When a species or a genus is increasing and spreading, it necessarily occupies a continuous area which gets larger and larger till it reaches a maximum; and we accordingly find that almost all extensive groups are thus continuous. When decay commences, and the group, ceasing to be in harmony with its environment, is encroached upon by other forms, the continuity may frequently be broken. Sometimes the outlying species may be the first to become extinct, and the group may simply diminish in area while keeping a compact central mass; but more often the process of extinction will be very irregular, and may even divide the group into two or more disconnected portions. This is the more likely to be the case because the most recently formed species, probably adapted to local conditions and therefore most removed from the general type of the group, will have the best chance of surviving, and these may exist at several isolated points of the area once occupied by the whole group. We may thus understand how the phenomenon of discontinuous areas has come about, and we may be sure that when allied species or varieties of the same species are found widely separated from each other, they were once connected by intervening forms or by each extending till it overlapped the other's area.

Discontinuous Specific Areas, why Rare.—But although discontinuous generic areas, or the separation from each other of species whose ancestors must once have occupied conterminous or overlapping areas, is of frequent occurrence, yet undoubted cases of discontinuous specific areas are very rare, except, as already stated, when one portion of a species inhabits an island. A few examples among mammalia have been referred to in our first chapter, but it may be said that these are examples of the very common phenomenon of a species being only found in the station for which its organisation adapts it; so that forest or marsh or mountain animals are of course only found where there are forests, marshes, or mountains. This may be true, and when the separate forests or mountains inhabited by the same species are not far apart there is little that needs explanation; but in one of the cases referred to there was a gap of a thousand miles between two of the areas occupied by the species, and this being too far for the animal to traverse through an uncongenial territory, we are forced to the conclusion that it must at some former period and under different conditions have occupied a considerable portion of the intervening area.

Among birds such cases of specific discontinuity are very rare and hardly ever quite satisfactory. This may be owing to birds being more rapidly influenced by changed conditions, so that when a species is divided the two portions almost always become modified into varieties or distinct species; while another reason may be that their powers of flight cause them to occupy on the average wider and less precisely defined areas than do the species of mammalia. It will be interesting therefore to examine the few cases on record, as we shall thereby obtain additional knowledge of the steps and processes by which the distribution of varieties and species has been brought about.

Discontinuity of the Area of Parus palustris.—Mr. Seebohm, who has travelled and collected in Europe, Siberia, and India, and possesses extensive and accurate knowledge of Palæarctic birds, has recently called attention to the varieties and sub-species of the marsh tit (Parus palustris), of which he has examined numerous specimens ranging from England to Japan. The curious point is that those of Southern Europe and of China are exactly alike, while all over Siberia a very distinct form occurs, forming the sub-species P. borealis. In Japan and Kamschatka other varieties are found, which have been named respectively P. japonicus and P. camschatkensis and another P. songarus in Turkestan and Mongolia. Now it all depends upon these forms being classed as sub-species or as true species whether this is or is not a case of discontinuous specific distribution. If Parus borealis is a distinct species from Parus palustris, as it is reckoned in Gray's Hand List of Birds, and also in Sharpe and Dresser's Birds of Europe, then Parus palustris has a most remarkable discontinuous distribution, as shown in the accompanying map, one portion of its area comprising Central and South Europe and Asia Minor, the other an undefined tract in Northern China, the two portions being thus situated in about the same latitude and having a very similar climate, but with a distance of about 4,000 miles between them. If, however, these two forms are reckoned as sub-species only, then the area of the species becomes continuous, while only one of its varieties or sub-species has a discontinuous area. It is a curious fact that P. palustris and P. borealis are found together in Southern Scandinavia and in some parts of Central Europe, and are said to differ somewhat in their note and their habits, as well as in colouration.

Discontinuity of Emberiza schœniclus.—The other case is that of our reed bunting (Emberiza schœniclus), which ranges over almost all Europe and Western Asia as far as the Yenesai valley and North-west India. It is then replaced by another smaller species, E. passerina, which ranges eastwards to the Lena river, and in winter as far south as Amoy in China; but in Japan the original species appears again, receiving a new name (E. pyrrhulina), but Mr. Seebohm assures us that it is quite indistinguishable from the European bird. Although the distance between these two portions of the species is not so great as in the last example, being about 2,000 miles, in other respects the case is an interesting one, because the forms which occupy the intervening space are recognised by Mr. Seebohm himself as undoubted species.

The European and Japanese Jays.—Another case somewhat resembling that of the marsh tit is afforded by the European and Japanese jays (Garrulus glandarius and G. japonicus). Our common jay inhabits the whole of Europe except the extreme north, but is not known to extend anywhere into Asia, where it is represented by several quite distinct species. (See Map, Frontispiece.) But the great central island of Japan is inhabited by a jay (G. japonicus) which is very like ours, and was formerly classed as a sub-species only, in which case our jay would be considered to have a discontinuous distribution. But the specific distinctness of the Japanese bird is now universally admitted, and it is certainly a very remarkable fact that among the twelve species of jays which together range over all temperate Europe and Asia, one which is so closely allied to our English bird should be found at the remotest possible point from it. Looking at the map exhibiting the distribution of the several species, we can hardly avoid the conclusion that a bird very like our jay once occupied the whole area of the genus, that in various parts of Asia it became gradually modified into a variety of distinct species in the manner already explained, a remnant of the original type being preserved almost unchanged in Japan, owing probably to favourable conditions of climate and protection from competing forms.

Supposed Examples of Discontinuity among North American Birds.—In North America, the eastern and western provinces are so different in climate and vegetation, and are besides separated by such remarkable physical barriers—the arid central plains and the vast ranges of the Rocky Mountains and Sierra Nevada, that we can hardly expect to find species whose areas may be divided maintaining their identity. Towards the north however the above-named barriers disappear, the forests being almost continuous from east to west, while the mountain range is broken up by passes and valleys. It thus happens that most species of birds which inhabit both the eastern and western coasts of the North American continent have maintained their continuity towards the north, while even when differentiated into two or more allied species their areas are often conterminous or overlapping.

Almost the only bird that seems to have a really discontinuous range is the species of wren, Thryothorus bewickii, of which the type form ranges from the east coast to Kansas and Minnesota, while a longer-billed variety, T. bewickii spilurus, is found in the wooded parts of California and as far north as Puget Sound. If this really represents the range of the species there remains a gap of about 1,000 miles between its two disconnected areas. Other cases are those of Vireo bellii of the middle United States and the sub-species pusillus of California; and of the purple red-finch, Carpodacus purpureus, with its variety C. californicus; but unfortunately the exact limits of these varieties are in neither case known, and though each one is characteristic of its own province, it is possible that they may somewhere become conterminous, though in the case of the red-finches this does not seem likely to be the fact.

In a later chapter we shall have to point out some remarkable cases of this kind where one portion of the species inhabits an island; but the facts now given are sufficient to prove that the discontinuity of the area occupied by a single homogeneous species, by two varieties of a species, by two well-marked sub-species, and by two closely allied but distinct species, are all different phases of one phenomenon—the decay of ill-adapted, and their replacement by better-adapted forms, under the pressure of a change of conditions either physical or organic. We may now proceed with our sketch of the mode of distribution of higher groups.

Distribution and Antiquity of Families.—Just as genera are groups of allied species distinguished from all other groups by some well-marked structural characters, so families are groups of allied genera distinguished by more marked and more important characters, which are generally accompanied by a peculiar outward form and style of colouration, and by distinctive habits and mode of life. As a genus is usually more ancient than any of the species of which it is composed, because during its growth and development the original rudimentary species becomes supplanted by more and more perfectly adapted forms, so a family is usually older than its component genera, and during the long period of its life-history may have survived many and great terrestrial and organic changes. Many families of the higher animals have now an almost worldwide extension, or at least range over several continents; and it seems probable that all families which have survived long enough to develop a considerable variety of generic and specific forms have also at one time or other occupied an extensive area.

Discontinuity a Proof of Antiquity.—Discontinuity will therefore be an indication of antiquity, and the more widely the fragments are scattered the more ancient we may usually presume the parent group to be. A striking example is furnished by the strange reptilian fishes forming the order or sub-order Dipnoi, which includes the Lepidosiren and its allies. Only three or four living species are known, and these inhabit tropical rivers situated in the remotest continents. The Lepidosiren paradoxa is only known from the Amazon and some other South American rivers. An allied species, Lepidosiren annectens, sometimes placed in a distinct genus, inhabits the Gambia in West Africa, while the recent discovery in Eastern Australia of the Ceratodus or mud-fish of Queensland, adds another form to the same isolated group. Numerous fossil teeth, long known from the Triassic beds of this country, and also found in Germany and India in beds of the same age, agree so closely with those of the living Ceratodus that both are referred to the same genus. No more recent traces of any such animal have been discovered, but the Carboniferous Ctenodus and the Devonian Dipterus evidently belong to the same group, while in North America the Devonian rocks have yielded a gigantic allied form which has been named Heliodus by Professor 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.

9 These facts are taken from a memoir on The Mammals and Winter Birds of Florida, by J. A. Allen; forming Vol. II., No. 3, of the Bulletin of the Museum of Comparative Zoology at Harvard College, Cambridge, Massachusetts.

10 The great variation in wild animals is more fully discussed and illustrated in the author's Darwinism (Chapter III.).

11 See Ibis, 1879, p. 32.

12 In Mr. Seebohm's latest work, Birds of the Japanese Empire (1890), he says, "Examples from North China are indistinguishable from those obtained in Greece" (p. 82).

13 Ibis, 1879, p. 40. In his Birds of the Japanese Empire (1890), Mr. Seebohm classes the Japanese and European forms as E. schœniclus, and thinks that their range is probably continuous across the two continents.

title: Island Life/IV


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Island Life/V

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 with unsuitability of climate.

The Dispersal of Birds.—Wherever mammals can migrate other vertebrates can generally follow with even greater facility. Birds, having the power of flight, can pass over wide arms of the sea, or even over extensive oceans, when these are, as in the Pacific, studded with islands to serve as resting places. Even the smaller land-birds are often carried by violent gales of wind from Europe to the Azores, a distance of nearly a thousand miles, so that it becomes comparatively easy to explain the exceptional distribution of certain species of birds. Yet on the whole it is remarkable how closely the majority of birds follow the same laws of distribution as mammals, showing that they generally require either continuous land or an island-strewn sea as a means of dispersal to new homes.

The Dispersal of Reptiles.—Reptiles appear at first sight to be as much dependent on land for their dispersal as mammalia, but they possess two peculiarities which favour their occasional transmission across the sea—the one being their greater tenacity of life, the other their oviparous mode of reproduction. A large boa-constrictor was once floated to the island of St. Vincent, twisted round the trunk of a cedar tree, and was so little injured by its voyage that it captured some sheep before it was killed. The island is nearly two hundred miles from Trinidad and the coast of South America, whence the reptile almost certainly came. Snakes are, however, comparatively scarce on islands far from continents, but lizards are often abundant, and though these might also travel on floating trees, it seems more probable that there is some as yet unknown mode by which their eggs are safely, though perhaps very rarely, conveyed from island to island. Examples of their peculiar distribution will be given when we treat of the fauna of some islands in which they abound.

The Dispersal of Amphibia and Fresh-water Fishes.—The two lower groups of vertebrates, Amphibia and fresh-water fishes, possess special facilities for dispersal, in the fact of their eggs being deposited in water, and in their aquatic or semi-aquatic habits. They have another advantage over reptiles in being capable of flourishing in arctic regions, and in the power possessed by their eggs of being frozen without injury. They have thus, no doubt, been assisted in their dispersal by floating ice, and by that approximation of all the continents in high northern latitudes which has been the chief agent in producing the general uniformity in the animal productions of the globe. Some genera of Batrachia have almost a world-wide distribution; while the tailed Batrachia, such as the newts and salamanders, are almost entirely confined to the northern hemisphere, some of the genera spreading over the whole of the north temperate zone. Fresh-water fishes have often a very wide range, the same species being sometimes found in all the rivers of a continent. This is no doubt chiefly due to the want of permanence in river basins, especially in their lower portions, where streams belonging to distinct systems often approach each other and may be made to change their course from one to the other basin by very slight elevations or depressions of the land. Hurricanes and water-spouts also often carry considerable quantities of water from ponds and rivers, and thus disperse eggs and even small fishes. As a rule, however, the same species are not often found in countries separated by a considerable extent of sea, and in the tropics rarely the same genera. The exceptions are in the colder regions of the earth, where the transporting power of ice may have come into play. High ranges of mountains, if continuous for long distances, rarely have the same species of fish in the rivers on their two sides. Where exceptions occur, it is often due to the great antiquity of the group, which has survived so many changes in physical geography that it has been able, step by step, to reach countries which are separated by barriers impassable to more recent types. Yet another and more efficient explanation of the distribution of this group of animals is the fact that many families and genera inhabit both fresh and salt water; and there is reason to believe that many of the fishes now inhabiting the tropical rivers of both hemispheres have arisen from allied marine forms becoming gradually modified for a life in fresh water. By some of these various causes, or a combination of them, most of the facts in the distribution of fishes can be explained without much difficulty.

The Dispersal of Insects.—In the enormous group of insects the means of dispersal among land animals reach their maximum. Many of them have great powers of flight, and from their extreme lightness they can be carried immense distances by gales of wind. Others can survive exposure to salt water for many days, and may thus be floated long distances by marine currents. The eggs and larvæ often inhabit solid timber, or lurk under bark or in crevices of logs, and may thus reach any countries to which such logs are floated. Another important factor in the problem is the immense antiquity of insects, and the long persistence of many of the best marked types. The rich insect fauna of the Miocene period in Switzerland consisted largely of genera still inhabiting Europe, and even of a considerable number identical, or almost so, with living species. Out of 156 genera of Swiss fossil beetles no less than 114 are still living; and the general character of the species is exactly like that of the existing fauna of the northern hemisphere in a somewhat more southern latitude. There is, therefore, evidently no difficulty in accounting for any amount of dispersal among insects; and it is all the more surprising that with such powers of migration they should yet be often as restricted in their range as the reptiles or even the mammalia. The cause of this wonderful restriction to limited areas is, undoubtedly, the extreme specialisation of most insects. They have become so exactly adapted to one set of conditions, that when carried into a new country they cannot live. Many can only feed in the larva state on one species of plant; others are bound up with certain groups of animals on whom they are more or less parasitic. Climatal influences have a great effect on their delicate bodies; while, however well a species may be adapted to cope with its enemies in one locality, it may be quite unable to guard itself against those which elsewhere attack it. From this peculiar combination of characters it happens, that among insects are to be found examples of the widest and most erratic dispersal and also of the extremest restriction to limited areas; and it is only by bearing these considerations in mind that we can find a satisfactory explanation of the many anomalies we meet with in studying their distribution.

The Dispersal of Land Mollusca.—The only other group of animals we need now refer to is that of the air-breathing mollusca, commonly called land-shells. These are almost as ubiquitous as insects, though far less numerous; and their wide distribution is by no means so easy to explain. The genera have usually a very wide, and often a cosmopolitan range, while the species are rather restricted, and sometimes wonderfully so. Not only do single islands, however small, often possess peculiar species of land-shells, but sometimes single mountains or valleys, or even a particular mountain side, possess species or varieties found nowhere else upon the globe. It is pretty certain that they have no means of passing over the sea but such as are very rare and exceptional. Some which possess an operculum, or which close the mouth of the shell with a diaphragm of secreted mucus, may float across narrow arms of the sea, especially when protected in the crevices of logs of timber; while in the young state when attached to leaves or twigs they may be carried long distances by hurricanes. Owing to their exceedingly slow motion, their powers of voluntary dispersal, even on land, are very limited, and this will explain the extreme restriction of their range in many cases.

Great Antiquity of Land-Shells.—The clue to the almost universal distribution of the several families and of many genera, is to be found, however, in their immense antiquity. In the Pliocene and Miocene formations most of the land-shells are either identical with living species or closely allied to them, while even in the Eocene almost all are of living genera, and one British Eocene fossil still lives in Texas. Strange to say, no true land-shells have been discovered in the Secondary formations, but they must certainly have abounded, for in the far more ancient Palæozoic coal measures of Nova Scotia two species belonging to the living genera Pupa and Zonites have been found in considerable abundance.

Land-shells have therefore survived all the revolutions the earth has undergone since Palæozoic times. They have been able to spread slowly but surely into every land that has ever been connected with a continent, while the rare chances of transfer across the ocean, to which we have referred as possible, have again and again occurred during the almost unimaginable ages of their existence. The remotest and most solitary of the islands of the mid-ocean have thus become stocked with them, though the variety of species and genera bears a direct relation to the facilities of transfer, and the shell fauna is never very rich and varied, except in countries which have at one time or other been united to some continental land.

Causes Favouring the Abundance of Land-Shells.—The abundance and variety of land-shells is also, more than that of any other class of animals, dependent on the nature of the surface and the absence of enemies, and where these conditions are favourable their forms are wonderfully luxuriant. The first condition is the presence of lime in the soil, and a broken surface of country with much rugged rock offering crevices for concealment and hibernation. The second is a limited bird and mammalian fauna, in which such species as are especially shell-eaters shall be rare or absent. Both these conditions are found in certain large islands, and pre-eminently in the Antilles, which possess more species of land-shells than any single continent. If we take the whole globe, more species of land-shells are found on the islands than on the continents—a state of things to which no approach is made in any other group of animals whatever, but which is perhaps explained by the considerations now suggested.

The Dispersal of Plants.—The ways in which plants are dispersed over the earth, and the special facilities they often possess for migration have been pointed out by eminent botanists, and a considerable space might be occupied in giving a summary of what has been written on the subject. In the present work, however, it is only in two or three chapters that I discuss the origin of insular floras in any detail; and it will therefore be advisable to adduce any special facts when they are required to support the argument in particular cases. A few general remarks only will therefore be made here.

Special Adaptability of Seeds for Dispersal.—Plants possess many great advantages over animals as regards the power of dispersal, since they are all propagated by seeds or spores, which are hardier than the eggs of even insects, and retain their vitality for a much longer time. Seeds may lie dormant for many years and then vegetate, while they endure extremes of heat, of cold, of drought, or of moisture which would almost always be fatal to animal germs. Among the causes of the dispersal of seeds De Candolle enumerates the wind, rivers, ocean currents, icebergs, birds and other animals, and human agency. Great numbers of seeds are specially adapted for transport by one or other of these agencies. Many are very light, and have winged appendages, pappus, or down, which enable them to be carried enormous distances. It is true, as De Candolle remarks, that we have no actual proofs of their being so carried; but this is not surprising when we consider how small and inconspicuous most seeds are. Supposing every year a million seeds were brought by the wind to the British Isles from the Continent, this would be only ten to a square mile, and the observation of a life-time might never detect one; yet a hundredth part of this number would serve in a few centuries to stock an island like Britain with a great variety of continental plants.

When, however, we consider the enormous quantity of seeds produced by plants, that great numbers of these are more or less adapted to be carried by the wind, and that winds of great violence and long duration occur in most parts of the world, we are as sure that seeds must be carried to great distances as if we had seen them so carried. Such storms carry leaves, hay, dust, and many small objects to a great height in the air, while many insects have been conveyed by them for hundreds of miles out to sea and far beyond what their unaided powers of flight could have effected.

Birds as Agents in the Dispersal of Plants.—Birds are undoubtedly important agents in the dispersal of plants over wide spaces of ocean, either by swallowing fruits and rejecting the seeds in a state fit for germination, or by the seeds becoming attached to the plumage of ground-nesting birds, or to the feet of aquatic birds embedded in small quantities of mud or earth. Illustrations of these various modes of transport will be found in Chapter XII. when discussing the origin of the flora of the Azores and Bermuda.

Ocean-currents as Agents in Plant-dispersal.—Ocean-currents are undoubtedly more important agents in conveying seeds of plants than they are in the case of any other organisms, and a considerable body of facts and experiments have been collected proving that seeds may sometimes be carried in this way many thousand miles and afterwards germinate. Mr. Darwin made a series of interesting experiments on this subject, some of which will be given in the chapter above referred to.

Dispersal along Mountain Chains.—These various modes of transport are, as will be shown when discussing special cases, amply sufficient to account for the vegetation found on oceanic islands, which almost always bears a close relation to that of the nearest continent; but there are other phenomena presented by the dispersal of species and genera of plants over very wide areas, especially when they occur in widely separated portions of the northern and southern hemispheres, that are not easily explained by such causes alone. It is here that transmission along mountain chains has probably been effective; and the exact mode in which this has occurred is discussed in Chapter XXIII., where a considerable body of facts is given, showing that extensive migrations may be effected by a succession of moderate steps, owing to the frequent exposure of fresh surfaces of soil or débris on mountain sides and summits, offering stations on which foreign plants can temporarily establish themselves.

Antiquity of Plants as affecting their Distribution.—We have already referred to the importance of great antiquity in enabling us to account for the wide dispersal of some genera and species of insects and land-shells, and recent discoveries in fossil botany show that this cause has also had great influence in the case of plants. Rich floras have been discovered in the Miocene, the Eocene, and the Upper Cretaceous formations, and these consist almost wholly of living genera, and many of them of species very closely allied to existing forms. We have therefore every reason to believe that a large number of our plant-species have survived great geological, geographical, and climatal changes; and this fact, combined with the varied and wonderful powers of dispersal many of them possess, renders it far less difficult to understand the examples of wide distribution of the genera and species of plants than in the case of similar instances among animals. This subject will be further alluded to when discussing the origin of the New Zealand flora, in Chapter XXII.

14 Lyell's Principles of Geology, ii., p. 369.

15 Mr. Darwin found that the large Helix pomatia lived after immersion in sea-water for twenty days. It is hardly likely that this is the extreme limit of their powers of endurance, but even this would allow of their being floated many hundred miles at a stretch, and if we suppose the shell to be partially protected in the crevice of a log of wood, and to be thus out of water in calm weather, the distance might extend to a thousand miles or more. The eggs of fresh-water mollusca, as well as the young animals, are known to attach themselves to the feet of aquatic birds, and this is probably the most efficient cause of their very wide diffusion.

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