We shall perhaps better exhibit the scope and complexity of the subject, and show that any intelligent study of it was almost impossible till quite recently, if we concisely enumerate the great mass of facts and the number of scientific theories or principles which are necessary for its elucidation.
We require then in the first place an adequate knowledge of the fauna and flora of the whole world, and even a detailed knowledge of many parts of it, including the islands of more special interest and their adjacent continents. This kind of knowledge is of very slow growth, and is still very imperfect;2 and in many cases it can never now be obtained owing to the reckless destruction of forests and with them of countless species of plants and animals. In the next place we require a true and natural classification of animals and plants, so that we may know their real affinities; and it is only now that this is being generally arrived at. We further have to make use of the theory of "descent with modification" as the only possible key to the interpretation of the facts of distribution, and this theory has only been generally accepted within the last twenty years. It is evident that, so long as the belief in "special creations" of each species prevailed, no explanation of the complex facts of distribution could be arrived at or even conceived; for if each species was created where it is now found no further inquiry can take us beyond that fact, and there is an end of the whole matter. Another important factor in our interpretation of the phenomena of distribution, is a knowledge of the extinct forms that have inhabited each country during the tertiary and secondary periods of geology. New facts of this kind are daily coming to light, but except as regards Europe, North America, and parts of India, they are extremely scanty; and even in the best-known countries the record itself is often very defective and fragmentary. Yet we have already obtained remarkable evidence of the migrations of many animals and plants in past ages, throwing an often unexpected light on the actual distribution of many groups.3 By this means alone can we obtain positive evidence of the past migrations of organisms; and when, as too frequently is the case, this is altogether wanting, we have to trust to collateral evidence and more or less probable hypothetical explanations. Hardly less valuable is the evidence of stratigraphical geology; for this often shows us what parts of a country have been submerged at certain epochs, and thus enables us to prove that certain areas have been long isolated and the fauna and flora allowed time for special development. Here, too, our knowledge is exceedingly imperfect, though the blanks upon the geological map of the world are yearly diminishing in extent. Lastly, as a most valuable supplement to geology, we require to know approximately, the depth and contour of the ocean-bed, since this affords an important clue to the former existence of now-submerged lands, uniting islands to continents, or affording intermediate stations which have aided the migrations of many organisms. This kind of information has only been partially obtained during the last few years; and it will be seen in the latter part of this volume, that some of the most recent deep-sea soundings have afforded a basis for an explanation of one of the most difficult and interesting questions in geographical biologythe origin of the fauna and flora of New Zealand.
Such are the various classes of evidence that bear directly on the question of the distribution of organisms; but there are others of even a more fundamental character, and the importance of which is only now beginning to be recognised by students of nature. These are, firstly, the wonderful alterations of climate which have occurred in the temperate and polar zones, as proved by the evidences of glaciation in the one and of luxuriant vegetation in the other; and, secondly, the theory of the permanence of existing continents and oceans. If glacial epochs in temperate lands and mild climates near the poles have, as now believed by men of eminence, occurred several times over in the past history of the earth, the effects of such great and repeated changes, both on the migration, modification, and extinction, of species, must have been of overwhelming importanceof more importance perhaps than even the geological changes of sea and land. It is therefore necessary to consider the evidence for these climatal changes; and then, by a critical examination of their possible causes, to ascertain whether they were isolated phenomena, were due to recurrent cosmical actions, or were the result of a great system of terrestrial development. The latter is the conclusion we arrive at; and this conclusion brings with it the conviction, that in the theory which accounts for both glacial epochs and warm polar climates, we have the key to explain and harmonize many of the most anomalous biological and geological phenomena, and one which is especially valuable for the light it throws on the dispersal and existing distribution of organisms. The other important theory, or rather corollary from the preceding theorythat of the permanence of oceans and the general stability of continents throughout all geological time, is as yet very imperfectly understood, and seems, in fact, to many persons in the nature of a paradox. The evidence for it, however, appears to me to be conclusive; and it is certainly the most fundamental question in regard to the subject we have to deal with: since, if we once admit that continents and oceans may have changed places over and over again (as many writers maintain), we lose all power of reasoning on the migrations of ancestral forms of life, and are at the mercy of every wild theorist who chooses to imagine the former existence of a now-submerged continent to explain the existing distribution of a group of frogs or a genus of beetles.
As already shown by the illustrative examples adduced in this chapter, some of the most remarkable and interesting facts in the distribution and affinities of organic forms are presented by islands in relation to each other and to the surrounding continents. The study of the productions of the Galapagosso peculiar, and yet so decidedly related to the American continentappears to have had a powerful influence in determining the direction of Mr. Darwin's researches into the origin of species; and every naturalist who studies them has always been struck by the unexpected relations or singular anomalies which are so often found to characterize the fauna and flora of islands. Yet their full importance in connection with the history of the earth and its inhabitants has hardly yet been recognised; and it is in order to direct the attention of naturalists to this most promising field of research, that I restrict myself in this volume to an elucidation of some of the problems they present to us. By far the larger part of the islands of the globe are but portions of continents undergoing some of the various changes to which they are ever subject; and the correlative proposition, that every portion of our continents has again and again passed through insular conditions, has not been sufficiently considered, but is, I believe, the statement of a great and most suggestive truth, and one which lies at the foundation of all accurate conception of the physical and organic changes which have resulted in the present state of the earth.
The indications now given of the scope and purpose of the present volume renders it evident that, before we can proceed to the discussion of the remarkable phenomena presented by insular faunas and floras, and the complex causes which have produced them, we must go through a series of preliminary studies, adapted to give us a command of the more important facts and principles on which the solution of such problems depends. The succeeding eight chapters will therefore be devoted to the explanation of the mode of distribution, variation, modification, and dispersal, of species and groups, illustrated by facts and examples; of the true nature of geological change as affecting continents and islands; of changes of climate, their nature, causes, and effects; of the duration of geological time and the rate of organic development.
CHAPTER II
THE ELEMENTARY FACTS OF DISTRIBUTION
Importance of Locality as an essential character of SpeciesAreas of DistributionExtent and Limitations of Specific AreasSpecific range of BirdsGeneric AreasSeparate and overlapping areasThe species of Tits as illustrating Areas of DistributionThe distribution of the species of JaysDiscontinuous generic areasPeculiarities of generic and family distributionGeneral features of overlapping and discontinuous areasRestricted areas of FamiliesThe distribution of Orders.
So long as it was believed that the several species of animals and plants were "special creations," and had been formed expressly to inhabit the countries in which they are now found, their habitat was an ultimate fact which required no explanation. It was assumed that every animal was exactly adapted to the climate and surroundings amid which it lived, and that the only, or, at all events, the chief reason why it did not inhabit another country was, that the climate or general conditions of that country were not suitable to it, but in what the unsuitability consisted we could rarely hope to discover. Hence the exact locality of any species was not thought of much importance from a scientific point of view, and the idea that anything could be learnt by a comparative study of different floras and faunas never entered the minds of the older naturalists.
But so soon as the theory of evolution came to be generally adopted, and it was seen that each animal could only have come into existence in some area where ancestral forms closely allied to it already lived, a real and important relation was established between an animal and its native country, and a new set of problems at once sprang into existence. From the old point of view the diversities of animal life in the separate continents, even where physical conditions were almost identical, was the fact that excited astonishment; but seen by the light of the evolution theory, it is the resemblances rather than the diversities in these distant continents and islands that are most difficult to explain. It thus comes to be admitted that a knowledge of the exact area occupied by a species or a group is a real portion of its natural history, of as much importance as its habits, its structure, or its affinities; and that we can never arrive at any trustworthy conclusions as to how the present state of the organic world was brought about, until we have ascertained with some accuracy the general laws of the distribution of living things over the earth's surface.
Areas of Distribution.Every species of animal has a certain area of distribution to which, as a rule, it is permanently confined, although, no doubt, the limits of its range fluctuate somewhat from year to year, and in some exceptional cases may be considerably altered in a few years or centuries. Each species is moreover usually limited to one continuous area, over the whole of which it is more or less frequently to be met with, but there are many apparent and some real exceptions to this rule. Some animals are so adapted to certain kinds of countryas to forests or marshes, mountains or desertsthat they cannot, permanently, live elsewhere. These may be found scattered over a wide area in suitable spots only, but can hardly on that account be said to have several distinct areas of distribution. As an example we may name the chamois, which lives only on high mountains, but is found in the Pyrenees, the Alps, the Carpathians, in some of the Greek mountains and the Caucasus. The variable hare is another and more remarkable case, being found all over Northern Europe and Asia beyond lat. 55°, and also in Scotland and Ireland. In central Europe it is unknown till we come to the Alps, the Pyrenees, and the Caucasus, where it again appears. This is one of the best cases known of the discontinuous distribution of a species, there being a gap of about a thousand miles between its southern limits in Russia, and its reappearance in the Alps. There are of course numerous instances in which species occur in two or more islands, or in an island and continent, and are thus rendered discontinuous by the sea, but these involve questions of changes in sea and land which we shall have to consider further on. Other cases are believed to exist of still wider separation of a species, as with the marsh titmice and the reed buntings of Europe and Japan, where similar forms are found in the extreme localities, while distinct varieties or sub-species, inhabit the intervening districts.
Extent and Limitations of Specific Areas.Leaving for the present these cases of want of continuity in a species, we find the most wide difference between the extent of country occupied, varying in fact from a few square miles to almost the entire land surface of the globe. Among the mammalia, however, the same species seldom inhabits both the old and new worlds, unless they are strictly arctic animals, as the reindeer, the elk, the arctic fox, the glutton, the ermine, and some others. The common wolf of Europe and Northern Asia is thought by many naturalists to be identical with the variously coloured wolves of North America extending from the Arctic Ocean to Mexico, in which case this will have perhaps the widest range of any species of mammal. Little doubt exists as to the identity of the brown bears and the beavers of Europe and North America; but all these species range up to the arctic circle, and there is no example of a mammal universally admitted to be identical yet confined to the temperate zones of the two hemispheres. Among the undisputed species of mammalia the leopard has an enormous range, extending all over Africa and South Asia to Borneo and the east of China, and thus having probably the widest range of any known mammal. The winged mammalia have not usually very wide ranges, there being only one bat common to the Old and New Worlds. This is a British species, Vesperugo serotinus, which is found over the larger part of North America, Europe and Asia, as far as Pekin, and even extends into tropical Africa, thus rivalling the leopard and the wolf in the extent of country it occupies.
Of very restricted ranges there are many examples, but some of these are subject to doubts as to the distinctness of the species or as to its geographical limits being really known. In Europe we have a distinct species of ibex (Capra Pyrenaica) confined to the Pyrenean mountains, while the true marmot is restricted to the Alpine range. More remarkable is the Pyrenean water-mole (Mygale Pyrenaica), a curious small insectivorous animal found only in a few places in the northern valleys of the Pyrenees. In islands there are many cases of undoubted restriction of species to a small area, but these involve a different question from the range of species on continents where there is no apparent obstacle to their wider extension.
Specific range of Birds.Among birds we find instances of much wider range of species, which is only what might be expected considering their powers of flight; but, what is very curious, we also find more striking (though perhaps not more frequent) examples of extreme limitation of range among birds than among mammals. Of the former phenomenon perhaps the most remarkable case is that afforded by the osprey or fishing-hawk, which ranges over the greater portion of all the continents, as far as Brazil, South Africa, the Malay Islands, and Tasmania. The barn owl (Strix flammea) has nearly as wide a range, but in this case there is more diversity of opinion as to the specific difference of many of the forms inhabiting remote countries, some of which seem undoubtedly to be distinct. Among passerine birds the raven has probably the widest range, extending from the arctic regions to Texas and New Mexico in America, and to North India and Lake Baikal in Asia; while the little northern willow-wren (Phylloscopus borealis) ranges from arctic Norway across Asia to Alaska, and southward to Ceylon, China, Borneo, and Timor.