# The Geographical Distribution of Animals/Chapter 23

CHAPTER XXIII.
SUMMARY OF THE DISTRIBUTION, AND LINES OF MIGRATION, OF THE SEVERAL CLASSES OF ANIMALS.
Having already given summaries of the distribution of the several orders, and of some of the classes of land animals, we propose here to make a few general remarks on the special phenomena presented by the more important groups, and to indicate where possible, the general lines of migration by which they have become dispersed over wide areas.
Mammalia.
This class is very important, and its past history is much better known than that of most others. We shall therefore briefly summarise the results we have arrived at from our examination of the distribution of extinct and living forms of each order.
Primates.—This order, being pre-eminently a tropical one, became separated into two portions, inhabiting the Eastern and Western Hemispheres respectively, at a very early epoch. In consequence of this separation it has diverged more radically than most other orders, so that the two American families, Cebidæ and Hapalidæ, are widely differentiated from the Apes, Monkeys, and Lemurs of the Old World. The Lemurs were probably still more ancient, but being much lower in organisation, they became extinct in most of the areas where the higher forms of Primates became developed. Remains found in the Eocene formation indicate, that the North American and European ​Primates had, even at that early epoch, diverged into distinct series, so that we must probably look back to the secondary period for the ancestral form from which the entire order was developed.
Chiroptera.—These are also undoubtedly very ancient. The most generalised forms—the Vespertilionidæ and Noctilionidæ—are the most widely distributed; while special types have arisen in America, and in the Eastern Hemisphere. Remains found in the Upper Eocene formation of Europe differ little from species still living in the same countries; so that we can form no conjecture as to the origin or migration, of the group. Their power of flight would, however, enable them rapidly to spread over all the great continents of the globe.
Insectivora.—This very ancient group, now probably verging towards extinction, appears to have originated in the Northern continent, and never to have reached Australia or South America. It may, however, have become extinct in the latter country owing to the competition of the numerous Edentata. The Insectivora now often maintain themselves amidst more highly developed forms, by means of some special protection. Some burrow in the earth,—like the moles; others have a spiny covering,—as the hedgehog's and several of the Centetidæ; others are aquatic,—-as the Potamogale and the desman; others have a nauseous odour,—as the shrews; while there are several which seem to be preserved by their resemblance to higher forms,—as the elephant-shrews to jerboas, and the tupaias to squirrels. The same need of protection is shown by the numerous Insectivora inhabiting Madagascar, where the competing forms are few; and by one lingering in the Antilles, where there are hardly any other mammalia.
Carnivora.—Although perhaps less ancient than the preceding, this form of mammal is far more highly organised, and from its earliest appearance appears to have become dominant in the world. It would therefore soon spread widely, and diverge into the various specialised types represented by existing families. Most of these appear to have originated in the Eastern Hemisphere, the only Carnivora occurring in North ​American Miocene deposits being ancestral forms of Canidæ and Felidæ. It seems probable, therefore, that the order had attained a considerable development before it reached the Western Hemisphere. The Procyonidæ, now confined to America, are not very ancient; and the occurrence of a few allied forms in the Himalayas (Ælurus and Æluropus) render it probable that their common ancestors entered North America from the Palæarctic region during the Miocene period, but being a rather low type they have succumbed under the competition of higher forms in most parts of the Eastern Hemisphere. Bears and Weasels are probably still more recent emigrants to America. The aquatic carnivora (Seals, &c.) are, as might be expected, more widely and uniformly distributed, but there is little evidence to show at what period the type was first developed.
Ungulata.—These are the dominant vegetable-feeders of the great continents, and they have steadily increased in numbers and in specialisation from the oldest Tertiary times to the present day. Being generally of larger size and less active than the Carnivora, they have somewhat more restricted powers of dispersal. We have good evidence that their wide range over the globe is a comparatively recent phenomenon. Tapirs and Llamas have probably not long inhabited South America, while Rhinoceroses and Antelopes were once, perhaps, unknown in Africa, although abounding in Europe and Asia. Swine are one of the most ancient types in both hemispheres; and their great hardiness, their omnivorous diet, and their powers of swimming, have led to their wide distribution. The sheep and goats, on the other hand, are perhaps the most recent development of the Ungulata, and they seem to have arisen in the Palæarctic region at a time when its climate already approximated to that which now prevails. Hence they are pre-eminently a Temperate group, never found within the Tropics except upon a few mountain ranges.
Proboscidea.—These huge animals (the Elephants and Mastodons) appear to have originated in the warmer parts of the Palæarctic region, but they soon spread over all the great ​continents, even reaching the southern extremity of America. Their extinction has probably depended more on physical than on organic changes, and we can clearly trace their almost total disappearance to the effects of the Glacial epoch.
Rodentia.—Rodents are a very dominant group, and a very ancient one. Owing to their small size and rapid powers of increase, they soon spread over almost every part of the globe, whence has resulted a great specialisation of family types in the South American continent which remained so long isolated. They are capable of living wherever there is any kind of vegetable food, hence their range will be determined rather by organic than by physical conditions; and the occupation of a country by enemies or by competing forms, is probably the chief cause which has prevented many of the families from acquiring a wide range. The occurrence of isolated species of the South American families, Octodontidæ and Echimyidæ in the Ethiopian and Palæarctic regions, is an indication that the range of many of the families has recently become less extensive.
Edentata.—These singular and lowly-organised animals appear to have become almost restricted to the two great Southern lands—South Africa and South America—at an early period; and, being there free from the competition of higher forms, developed a number of remarkable types often of huge size, of which the Megatherium is one of the best known. The incursion of the highly-organised Ungulates and Carnivora into Africa during the Miocene epoch, probably exterminated most of them in that continent; but in America they continued in full force down to the Post-Pliocene period; and even now, the comparatively diminutive Sloths, Ant-eaters, and Armadillos, form a large and important portion of the fauna.
Marsupialia and Monotremata.—These are probably the representatives of the most ancient and lowly-organised types of mammal. They once existed in the northern continents, whence they spread into Australia; and being isolated, and preserved from the competition of the higher forms which soon arose in other parts of the world, they have developed into a variety of types, which, however, still preserve a general ​uniformity of organisation. One family, which continued to exist in Europe till the latter part of the Miocene period, reached America, and has there been preserved to our day.
Lines of Migration of the Mammalia.—The whole series of phenomena presented by the distribution of the Mammalia, looked at broadly, are in harmony with the view that the great continents and oceans of our own epoch have been in existence, with comparatively small changes, during all Tertiary times. Each one of them has, no doubt, undergone considerable modifications in its area, its altitude, and in its connection with other lands. Yet some considerable portion of each continent has, probably, long existed in its present position, while the great oceans seem to have occupied the same depressions of the earth's crust (varied, perhaps, by local elevations and subsidences) during all this vast period of time. Hence, allowing for the changes of which we have more or less satisfactory evidence, the migrations of the chief mammalian types can be pretty clearly traced. Some, owing to their small size and great vitality, have spread to almost all the chief land masses; but the majority of the orders have a more restricted range. All the evidence at our command points to the Northern Hemisphere as the birth-place of the class, and probably of all the orders. At a very early period the land communication with Australia was cut off, and has never been renewed; so that we have here preserved for us a sample of one or more of the most ancient forms of mammal. Somewhat later the union with South America and South Africa was severed; and in both these countries we have samples of a somewhat more advanced stage of mammalian development. Later still, the union by a northern route between the Eastern and Western Hemispheres appears to have been broken, partly by a physical separation, but almost as effectually by a lowering of temperature. About the same period the separation of the Palæarctic region from the Oriental was effected, by the rise of the Himalayas and the increasing contrast of climate; while the formation of the great desert-belts of the Sahara, Arabia, Persia, and Central Asia, helped to complete the separation of ​the Temperate and Tropical zones, and to render further intermigration almost impossible.
In a few cases—of which the Rodents in Australia and the pigs in Austro-Malaya are perhaps the most striking examples—the distribution of land-mammals has been effected by a sea-passage either by swimming or on floating vegetation; but, as a rule, we may be sure that the migrations of mammalia have taken place over the land; and their presence on islands is, therefore, a clear indication that these have been once connected with a continent. The present class of animals thus affords the best evidence of the past history of the land surface of our globe; and we have chiefly relied upon it in sketching out (in Part III.) the probable changes which each of our great regions has undergone.
Birds.
Although birds are, of all land-vertebrates, the best able to cross seas and oceans, it is remarkable how closely the main features of their distribution correspond with those of the Mammalia. South America possesses the low Formicaroid type of Passeres,—which, compared with the more highly developed forms of the Eastern Hemisphere, is analogous to the Cebidæ and Hapalidæ as compared with the Old World Apes and Monkeys; while its Cracidæ as compared with the Pheasants and Grouse, may be considered parallel to the Edentata as compared with the Ungulates of the Old World. The Marsupials of America and Australia, are paralleled among birds, in the Struthionidæ and Megapodiidæ; the Lemurs and Insectivora preserved in Madagascar are represented by the Mascarene Dididæ; the absence of Deer and Bears from Africa is analogous to the absence of Wrens, Creepers, and Pheasants; while the African Hyracidæ and Chrysochloridæ among mammals, may well be compared with the equally peculiar Coliidæ and Musophagidæ among birds.
From these and many other similarities of distribution, it is clear that birds have, as a rule, followed the same great lines of migration as mammalia; and that oceans, seas, and deserts, have ​always to a great extent limited their range. Yet these barriers have not been absolute; and in the course of ages birds have been able to reach almost every habitable land upon the globe. Hence have arisen some of the most curious and interesting phenomena of distribution; and many islands, which are entirely destitute of mammalia, or possess a very few species, abound in birds, often of peculiar types and remarkable for some unusual character or habit. Striking examples of such interesting bird-faunas are those of New Zealand, the Sandwich Islands, the Galapagos, the Mascarene Islands, the Moluccas, and the Antilles; while even small and remote islets,—such as Juan Fernandez and Norfolk Island, have more light thrown upon their past history by means of their birds, than by any other portion of their scanty fauna.
Another peculiar feature in the distribution of this class is the extraordinary manner in which certain groups and certain external characteristics, have become developed in islands, where the smaller and less powerful birds have been protected from the incursions of mammalian enemies, and where rapacious birds—which seem to some degree dependent on the abundance of mammalia—are also scarce. Thus, we have the Pigeons and the Parrots most wonderfully developed in the Australian region, which is pre-eminently insular; and both these groups here acquire conspicuous colours very unusual, or altogether absent, elsewhere. Similar colours (black and red) appear, in the same two groups, in the distant Mascarene islands; while in the Antilles the parrots have often white heads, a character not found in the allied species on the South American continent. Crests, too, are largely developed, in both these groups, in the Australian region only; and a crested parrot formerly lived in Mauritius,—a coincidence too much like that of the colours as above noted, to be considered accidental.
Again, birds exhibit to us a remarkable contrast as regards the oceanic islands of tropical and temperate latitudes; for while most of the former present hardly any cases of specific identity with the birds of adjacent continents, the latter often show hardly any differences. The Galapagos and Madagascar ​are examples of the first-named peculiarity; the Azores and the Bermudas of the last; and the difference can be clearly traced to the frequency and violence of storms in the one case and to the calms or steady breezes in the other.
It appears then, that although birds do not afford us the same convincing proof of the former union of now disjoined lands as we obtain from mammals, yet they give us much curious and suggestive information as to the various and complex modes in which the existing peculiarities of the distribution of animals have been brought about. They also throw much light on the relation between distribution and the external characters of animals; and, as they are often found where mammalia are quite absent, we must rank them as of equal value for the purposes of our present study.
Reptiles.
These hold a somewhat intermediate place, as regards their distribution, between mammals and birds, having on the whole rather a wider range than the former, and a more restricted one than the latter.
Snakes appear to have hardly more facilities for crossing the ocean than mammals; hence they are generally absent from oceanic islands. They are more especially a tropical group, and have thus never been able to pass from one continent to another by those high northern and southern routes, which we have seen reason to believe were very effectual in the case of mammalia and some other animals. Hence we find no resemblance between the Australian and Neotropical regions, or between the Palæarctic and Nearctic; while the Western Hemisphere is comparatively poor as regards variety of types, although rich in genera and species. Deserts and high mountains are also very effectual barriers for this group, and their lines of migration have probably been along river valleys, and occasionally across narrow seas by means of floating vegetation.
Lizards, being somewhat less tropical than snakes, may have passed by the northern route during warm epochs. They are also more suited to traverse deserts, and they possess some unknown ​means of crossing the ocean, as they are not unfrequently found in remote oceanic islands. These various causes have modified their distribution. The Western Hemisphere is much richer in lizards than it is in snakes; and it is also very distinct from the Eastern Hemisphere. The lines of migration of lizards appear to have been along the mountains and deserts of tropical countries, and, under special conditions, across tropical seas from island to island.
Crocodiles are a declining group. They were once more generally distributed, all the three families being found in British Eocene deposits. Being aquatic and capable of living in the sea, they can readily pass along all the coasts and islands of the warmer parts of the globe. Tortoises are equally ancient, and the restriction of certain groups to definite areas seems to be also a recent phenomenon.
Amphibia.
The Amphibia differ widely from Reptiles in their power of enduring cold; one of their chief divisions, the Urodela or Tailed-Batrachia, being confined to the temperate parts of the Northern Hemisphere. To this class of animals the northern and southern routes of migration were open; and we accordingly find a considerable amount of resemblance between South America and Australia, and a still stronger affinity between North America and the Palæarctic continent. The other tropical regions are more distinct from each other; clearly indicating that, in this group, it is tropical deserts and tropical oceans which are the barriers to migration. The class however is very fragmentary, and probably very ancient; so that descendants of once widespread types are now found isolated in various parts of the globe, between which we may feel sure there has been no direct transmission of Batrachia. Remembering that their chief lines of migration have been by northern and southern land-routes, by floating ice, by fresh-water channels, and perhaps at rare intervals by ova being carried by aquatic birds or by violent storms,—we shall be able to comprehend most of the features of their actual distribution.
Fresh-water Fishes.
Although it would appear, at first sight, that the means of dispersal of these animals are very limited, yet they share to some extent the wide range of other fresh-water organisms. They are found in all climates; but the tropical regions are by far the most productive, and of these South America is perhaps the richest and most peculiar. There is a certain amount of identity between the two northern continents, and also between those of the South Temperate zone; yet all are radically distinct, even North America and Europe having but a small proportion of their forms in common. The occurrence of allied fresh-water species in remote lands—as the Aphritis of Tasmania and Patagonia, and the Comephorus of Lake Baikal, distantly allied to the mackerels of Northern seas—would imply that marine fishes are often modified for a life in fresh waters; while other facts no less plainly show that permanent fresh-water species are sometimes dispersed in various ways across the oceans, more especially by the northern and southern routes.
The families of fresh-water fishes are often of restricted range, although cases of very wide and scattered distribution also occur. The great zoological regions are, on the whole, very well characterized; showing that the same barriers are effectual here, as with most other vertebrates. We conclude, therefore, that the chief lines of migration of fresh-water fishes have been across the Arctic and Antarctic seas, probably by means of floating ice as well as by the help of the vast flocks of migratory aquatic birds that frequent those regions. On continents they are, usually, widely dispersed; but tropical seas, even when of small extent, appear to have offered an effectual barrier to their dispersal. The cases of affinity between Tropical America, Africa, Asia, and Australia, must therefore be imputed either to the survival of once widespread groups, or to analogous adaptation to a fresh-water life of wide-spread marine types; and these cases cannot be taken as evidence of any former land connection between such remote continents.
Insects.
It has already been shown (Vol. I. pp. 209-213 and Vol. II. pp. 44-48) that the peculiarities of distribution of the various groups of insects depend very much on their habits and general economy. Their antiquity is so vast, and their more important modifications of structure have probably occurred so slowly, that modes of dispersal depending on such a combination of favourable conditions as to be of excessive rarity, may yet have had time to produce large cumulative effects. Their small specific gravity and their habits of flight render them liable to dispersal by winds to an extent unknown in other classes of animals; and thus, what are usually very effectual barriers have been overstepped, and sometimes almost obliterated, in the case of insects. A careful examination will, however, almost always show traces of an ancient fauna, agreeing in character with other classes of animals, intermixed with the more prominent and often more numerous forms whose presence is due to this unusual facility of dispersal.
The effectual migration of insects is, perhaps more than in any other class of animals, limited by organic and physical conditions. The vegetation, the soil, the temperature, and the supply of moisture, must all be suited to their habits and economy; while they require an immunity from enemies of various kinds, which immigrants to a new country seldom obtain. Few organisms have, in so many complex ways, become adapted to their special environment, as have insects. They are in each country more or less adapted to the plants which belong to it; while their colours, their habits, and the very nature of the juices of their system, are all modified so as to protect them from the special dangers which surround them in their native land. It follows, that while no animals are so well adapted to show us the various modes by which dispersal may be effected, none can so effectually teach us the true nature and vast influence of the organic barrier in limiting dispersal.
It is probable that insects have at one time or another taken advantage of every line of migration by which any terrestrial ​organisms have spread over the earth, but owing to their small size and rapid multiplication, they have made use of some which are exclusively their own. Such are the passage along mountain ranges from the Arctic to the Antarctic regions, and the dispersal of certain types over all temperate lands. It will perhaps be found that insects have spread over the land surface in directions dependent on our surface zones—forests, pastures, and deserts;—and a study of these, with a due consideration of the fact that narrow seas are scarcely a barrier to most of the groups, may assist us to understand many of the details of insect-distribution.
Terrestrial Mollusca.
The distribution of land-shells agrees, in some features, with that of insects, while in others the two are strongly contrasted. In both we see the effects of great antiquity, with some special means of dispersal; but while in insects the general powers of motion, both voluntary and involuntary, are at a maximum, in land-molluscs they are almost at a minimum. Although to some extent dependent on vegetation and climate, the latter are more dependent on inorganic conditions, and also to a large extent on the general organic environment. The result of these various causes, acting through countless ages, has been to spread the main types of structure with considerable uniformity over the globe; while generic and sub-generic forms are often wonderfully localized.
Land-shells, even more than insects, seem, at first sight, to require regions of their own; but we have already pointed out the disadvantages of such a method of study. It will be far more instructive to refer them to those regions and sub-regions which are found to accord best with the distribution of the higher animals, and to consider the various anomalies they present as so many problems, to be solved by a careful study of their habits and economy, and especially by a search after the hidden causes which have enabled them to spread so widely over land and ocean.
The lines of migration which land-shells have followed, can ​hardly be determined with any definiteness. On continents they seem to spread steadily, but slowly, in every direction, checked probably by organic and physical conditions rather than by the barriers which limit the higher groups. Over the ocean they are also slowly dispersed, by some means which act perhaps at very long intervals, but which, within the period of the duration of genera and families, are tolerably effective. It thus happens that, although the powers of dispersal of land-shells and insects are so very unequal, the resulting geographical distribution is almost the opposite of what might have been expected,—the former being, on the whole, less distinctly localized than the latter.
CONCLUSION.
The preceding remarks are all I now venture to offer, on the distinguishing features of the various groups of land-animals as regards their distribution and migrations. They are at best but indications of the various lines of research opened up to us by the study of animals from the geographical point of view, and by looking upon their range in space and time as an important portion of the earth's history. Much work has yet to be done before the materials will exist for a complete treatment of the subject in all its branches; and it is the author's hope that his volumes may lead to a more systematic collection and arrangement of the necessary facts. At present all public museums and private collections are arranged zoologically. All treatises, monographs, and catalogues, also follow, more or less completely, the zoological arrangement; and the greatest difficulty the student of geographical distribution has to contend against, is the total absence of geographical collections, and the almost total want of complete and comparable local catalogues. Till every well-marked district,—every archipelago, and every important island, has all its known species of the more important groups of animals catalogued on a uniform plan, and with a uniform nomenclature, a thoroughly satisfactory account of the Geographical Distribution of Animals will not be possible. But more than this is wanted. Many of the most curious relations between animal ​forms and their habitats, are entirely unnoticed, owing to the productions of the same locality never being associated in our museums and collections. A few such relations have been brought to light by modern scientific travellers, but many more remain to be discovered; and there is probably no fresher and more productive field still unexplored in Natural History. Most of these curious and suggestive relations are to be found in the productions of islands, as compared with each other, or with the continents of which they form appendages; but these can never be properly studied, or even discovered, unless they are visibly grouped together. When the birds, the more conspicuous families of insects, and the land-shells of islands, are kept together so as to be readily compared with similar associations from the adjacent continents or other islands, it is believed that in almost every case there will be found to be peculiarities of form or colour running through widely different groups, and strictly indicative of local or geographical influences. Some of these coincident variations have been alluded to in various parts of this work, but they have never been systematically investigated. They constitute an unworked mine of wealth for the enterprising explorer; and they may not improbably lead to the discovery of some of the hidden laws (supplementary to Natural Selection), which seem to be required, in order to account for many of the external characteristics of animals.
In concluding his task, the author ventures to suggest, that naturalists who are disposed to turn aside from the beaten track of research, may find in the line of study here suggested a new and interesting pursuit, not inferior in attractions to the lofty heights of transcendental anatomy, or the bewildering mazes of modern classification. And it is a study which will surely lead them to an increased appreciation of the beauty and the harmony of nature, and to a fuller comprehension of the complex relations and mutual interdependence, which link together every animal and vegetable form, with the ever-changing earth which supports them, into one grand organic whole.