Chapter VIII: The Distribution of Animals as Indicating Geographical Changes.63 (2)

Further south, in Bolivia, the Pampas, and Patagonia, we also find abundant fossil remains, probably a little older than the cave fauna of Brazil, and usually referred to the newer part of the Pliocene period. The same families of rodents and Edentata are here abundant, many of the genera being the same but several new ones also appearing. There are also horses, peccaries, a mastodon, llamas, and deer; but besides these there are a number of altogether peculiar forms, such as the Macrauchenia, allied to the Tapir and Palæotherium; the Homalodontotherium, allied to the Miocene Hyracodon of North America; and the Toxodontidæ, a group of very large animals having affinities to Ungulates, rodents, Edentata, and Sirenia, and therefore probably the representative of a very ancient type.

Here then we meet with a mixture of highly developed and recent, with low and ancient types, but the latter largely predominate; and the most probable explanation seems to be that the same concurrence of favourable conditions which allowed the megatherium and megalonyx to enter North America also led to an immigration of horses, deer, mastodons, and many of the Felidæ into South America. These inter-migrations appear to have taken place at several remote intervals, the northern and southern continents being for the most part quite separated, and each developing its own peculiar forms of life. This view is supported by the curious fact of a large number of the marine fishes of the two sides of Central America being absolutely identical--implying a recent union of the two oceans and separation of the continents--while the mollusca of the Pacific coast of America bear so close a relation to those of the Caribbean Sea and the Atlantic coasts, as to indicate a somewhat more remote but longer continued sea-passage. The straits connecting the two oceans were probably situated in Nicaragua and to the south of Panama, leaving the highlands of Mexico and Guatemala united to North America.

Around the Gulf of Mexico and the Caribbean Sea there is a wide belt of rather shallow water, and during the alternate elevations and subsidences to which this region has been subjected, the newly raised land would afford a route for the passage of immigrants between North and South America. The great depression of the ocean, believed to have occurred during the Glacial period (caused by the locking-up of the water in the two polar masses of ice), may perhaps have afforded the opportunity for those latest immigrations which gave so striking a character to the North American fauna in Post-Pliocene times.

Among the changes which South America itself has undergone, perhaps the most important has been its separation into a group of large islands. Such a change is clearly indicated by the immense area and low elevation of the great alluvial plains of the Orinoko, Amazon, and La Plata, as well as by certain features in the distribution of the existing Neotropical fauna. A subsidence of less than 2,000 feet would convert the highlands of Guiana and Brazil into islands separated by a shallow strait from the chain of the Andes. When this occurred the balance of the land was probably restored by an elevation of the extensive submerged banks on the east coast of South America, which in South Brazil and Patagonia are several hundred miles wide, embracing the Falkland Islands, and reaching far to the south of Cape Horn.

Looking, then, at the whole of the evidence at our command, we seem justified in concluding that the past histories of North and South America have been different, and in some respects strongly contrasted. North America was evidently in very early times so far connected with Europe and Asia as to interchange with those continents the higher types of animal life as they were successively developed in either hemisphere. These more perfectly organised beings rapidly gained the ascendency, and led to the extinction of most of the lower forms which had preceded them. The Nearctic has thus run a course parallel to that of the Palæarctic region, although its fauna is, and perhaps always has been, less diversified and more subject to incursions of lower types from adjacent lands in the southern hemisphere.

South America, on the other hand, has had a history in many respects parallel to that of Africa. Both have long existed either as continents or groups of large islands in the southern hemisphere, and for the most part completely separated from the northern continents; and each accordingly developed its peculiar types from those ancestral and lowly-organised forms which first entered it. South America, however, seems to have had a larger area and more favourable conditions, and it remained almost completely isolated till a later period. It was therefore able to develop a more-varied and extensive fauna of its own peculiar types, and its union with the northern continent has been so recent, and is even now maintained by so narrow an isthmus, that it has never been overrun with the more perfect mammalia to anything like the extent that has occurred in Africa. South America, therefore, almost as completely as Australia, has preserved for us examples of a number of low and early types of mammalian life, which, had not the entire country been isolated from the northern continent during middle and late Tertiary times, would long since have become extinct.

_The Australian Region._--There only remains for us now to consider the relation of the island-continent of Australia to Asia and South America, with both which countries it has a certain amount of zoological connection.

Australia, including New Guinea (which has in recent times been united with it), differs from all the other continents by the extreme uniformity and lowly organisation of its mammalia which almost all belong to one of the lowest orders--the marsupials. Monkeys, carnivora, insectivora, and the great and almost ubiquitous class of hoofed-animals, are all alike wanting; the only mammals besides marsupials being a few species of a still lower type--the monotremes, and a few of the very smallest forms of rodents--the mice. The marsupials, however, are very numerous and varied, constituting 6 families and 33 genera, of which there are about 120 known species. None of these families is represented in any other continent; and this fact alone is sufficient to prove that Australia must have remained almost or quite isolated during the whole of the Tertiary period.

In birds there is, as we might expect, less complete isolation; yet there are a number of very peculiar types. About 15 families are confined to the Australian region, among which are the paradise-birds, the honey-suckers, the lyre-birds, the brush-tongued lories, the mound-makers, and the cassowaries.

Our knowledge of the former mammalian inhabitants of Australia is imperfect, as all yet discovered are from Post-Tertiary or very late Tertiary deposits. It is interesting to find, however, that all belong to the marsupial type, although several are quite unlike any living animals, and some are of enormous size, almost rivalling the mastodons and megatheriums of the northern continents. In the earliest Tertiary formation of Europe remains of marsupials have been found, but they all belong to the opossum type, which is unknown in Australia; and this supports the view that no communication existed between the Palæarctic and Australian regions even at this early period. Much farther back, however, in the Oolite and Trias formations, remains of a number of small mammalia have been found which are almost certainly marsupial, and bear a very close resemblance to the Myrmecobius, a small and very rare mammal still living in Australia. An animal of somewhat similar type has been discovered in rocks of the same age in North America; and we have, therefore, every reason to believe, that it was at or near this remote epoch when Australia, or some land which has been since in connection with it, received a stock of mammalian immigrants from the great northern continent; since which time it has almost certainly remained completely isolated.

The occurrence of the marsupial opossums in America has been thought by some writers to imply an early connection between that continent and Australia; but the fact that opossums existed in Europe in Eocene and Miocene times, and that no trace of them has been found in North or South America before the Post-Pliocene period, renders it almost certain that they entered America from Europe or North Asia in middle or late Tertiary times, and have flourished there in consequence of a less severe competition with highly-developed forms of life.

The birds of Australia and South America only exhibit a few cases of very remote and general affinity, which are best explained by the preservation in each country of once wide-spread types, but is quite inconsistent with the theory of a direct union between the two countries during Tertiary times.

Reptiles are even more destitute of proofs of any such connection than even mammalia or birds; but in amphibia, fresh-water fishes, and insects the case is different, all these classes furnishing examples of the same families or genera inhabiting the temperate parts of both continents. But the fact that such cases are confined to these three groups and to plants, is the strongest possible proof that they are not due to land-connection; for all these organisms may be transmitted across the ocean in various ways. Violent storms of wind, floating ice, drift-wood, and aquatic birds, are all known to be effective means for the distribution of these animals or their ova, and the seeds of plants. All of them too, it must be noted, are to a considerable degree patient of cold; the reverse being the case with true reptiles and land-birds, which are essentially heat-loving; so that the whole body of facts seems to point rather to an extension of the Antarctic lands and islands reducing the width of open sea, than to any former union, or even close approximation of the Australian and South American continents.

_Summary and Conclusion._

Let us now briefly review the conclusions at which we have arrived. If we look back to remote Tertiary times, we shall probably find that all our great continents and oceans were then in existence, and even bore a general resemblance to the forms and outlines now so familiar to us. But in many details, and especially in their amount of communication with each other, we should observe important changes. The first thing we should notice would be a more complete separation of the northern and the southern continents. Now, there is only one completely detached southern land--Australia; but at that period Africa and South America were also vast islands or archipelagos, completely separated from their sister continents. Examining them more closely, we should observe that the great Euro-Asiatic continent had a considerable extension to the south-east, over what are now the shallow seas of Japan, China, and Java. In the south-west it would include Northern Africa, the Mediterranean then forming two inland seas; while to the west and north-west it would include the British Isles, and perhaps extend even to Iceland and Greenland. As a balance to these extensions, much of Northern Siberia and North-Western Asia may have been under water; the peninsula of India would be an island with a considerable south-west extension over what are now the Laccadive and Maldive coral-reefs. The Himalayas would be a moderate range of hills; the great desert plateau of Central Asia a fertile plain; the greater part of the continent would enjoy a tropical or subtropical climate, while even the extreme north would support a luxuriant vegetation. This great continent would abound in animal life, and would be especially remarkable for its mammalia, which would comprise ancestral forms of all our existing higher types, along with a number of those lower grades of organisation (such as lemurs and opossums) now found chiefly in the southern hemisphere.

Connected with this continent by what is now Behring Straits and the Sea of Kamschatka, we should find North America, perhaps somewhat diminished in the east, but more extensive in the south and north, and abounding as now with great inland lakes which were situated to the west of the present lake district. This continent seems to have had a less tropical climate and vegetation than prevailed in the eastern hemisphere, but it supported an almost equally varied though very distinct fauna. Ancestral horses no larger than dogs; huge tapir-like and pig-like animals; strange forms allied to rhinoceroses; the Dinocerata--huge horned animals allied to elephants and to generalised Ungulata; and the Tillodontia, still more unlike anything now living, since they combined characters now found separated in the carnivora, the Ungulata, and the rodents. Ancestral Primates, allied to both the lemurs and the South American monkeys, also inhabited this continent.

The great land masses of the northern hemisphere thus appear to have possessed between them all the higher types of animal life; and these seem to have been developed for a time in one continent and then to have been in part transferred by migration to the other, where alone they have sometimes maintained themselves. Thus, the elephants and the camels appear to have descended from what were once exclusively American types, while the opossums were as certainly European. Many groups, however, never passed out of the continent in which they originated--the civets, hyænas, and the giraffes being wholly eastern, while the Oreodontidæ and Brontotheridæ were no less exclusively western.

South America seems to have been united to the northern continent once at least in Secondary or early Tertiary times, since it was inhabited in the Eocene period by many forms of mammalia, such as rodents, felines, and some ancient forms of Ungulata. It must also have possessed the ancestors of the Edentata (though they have not yet been discovered), or we should not find such a variety of strange and gigantic forms of this order in later Tertiary deposits in this part of the world only. During the greater part of the Tertiary period, therefore, South America must have been separated from the North and protected from incursions of the higher forms of mammalia which were there so abundant. Thus only does it seem possible to understand the unchecked development of so many large but comparatively helpless animals as the Edentata of the Pampas and the Brazilian caves--a development only comparable with that of the Australian marsupials, still more completely shut off from all competition with higher forms of life.

In Africa the evidence of a long period of insulation is somewhat more complex and less easily apparent, but, it seems to me, equally conclusive. We have first, the remarkable fauna of Madagascar, in which lemurs and insectivora predominate, with a few low forms of carnivora; but none of the higher animals, such as apes, antelopes, buffaloes, rhinoceroses, elephants, lions, leopards, and hyænas, which swarm on the continent. The separation of Madagascar from Africa must therefore have occurred before these important groups existed there. Now, we know that all these large animals lived in Europe and Asia during late Miocene times, while lemurs are only known there during the Eocene period, and were probably more abundant in late Mesozoic times. It is almost certain, therefore, that Southern Africa must have been cut off from Europe and Asia during the whole intervening period, or the same development of high forms and extinction of low would have gone on in the one country as in the other. The persistence of a number of low and isolated types in South and West Africa, which are probably a remnant of the ancient fauna of the country, is also favourable to this view. At the time we are considering, therefore, we look upon tropical and South Africa, with Madagascar, as forming a completely isolated land or archipelago; while the Seychelles and Chagos banks, with Bourbon and Mauritius, perhaps, formed another island or group permanently separated from the larger masses. The extra-tropical portion of South Africa was also probably more extensive, affording an area in which its remarkable flora was being developed.

Turning to Australia, we should probably find it, at this remote period, more extensive than it is now, including in its area New Guinea and some of the adjacent islands, as well as Tasmania; while another extensive land probably occupied the site of the New Zealand group. It may be considered certain that, whatever elevations and subsidences these countries may have undergone, they have not been connected either with Asia, Africa, or South America during the whole Tertiary period.

In conclusion, I would especially remark that the various changes in the outlines and mutual relations of our continents, which I have now endeavoured to establish, must not be supposed to have been all strictly contemporaneous. Some may have been a little earlier or a little later than others; some changes may have been slower, others more rapid; some may have had but a short duration, while others may have persisted through considerable geological periods. But, notwithstanding this uncertainty as to details, the great features of the geographical revolutions which I have indicated, appear to be established by a mass of concurring evidence; and the lesson they teach us is, that although almost the whole of what is now dry land has undoubtedly once lain deep beneath the waters of the ocean, yet such changes on a great scale are excessively slow and gradual; so that, when compared with the highest estimates of the antiquity of the human race, or even with that of most of the higher animals, our existing continents and oceans may be looked upon as permanent features of the earth’s surface.

ERRATUM.

At page 59 I have said that there are only three or four species of Mimosa which are sensitive. This is a mistake, as the greater portion of the species in the extensive genus Mimosa, as well as some species of several other genera of Leguminosæ, and also of Oxalidaceæ, possess this curious property. I cannot find, however, that any one has suggested in what way the sensitiveness may have been useful to the species which first acquired it. My guess at an explanation may therefore induce botanists who are acquainted with the various species in a state of nature, to suggest some better solution of the problem.

INDEX.

A.

_Abrus precatoria_, perhaps a case of mimicry, 226

Absorption-colours or pigments, 183

Acræidæ, warning colours of, 174

Adaptive characters, 150, 155

Affinities, how to determine doubtful, 148

African large mammalia, recent immigrants, 323

Allen, Mr. Grant, on protective colours of fruits, 225

Alpine flowers, why so beautiful, 232

Amboyna, large-sized butterflies of, 258

American monkeys, 118

American Continents, past history of, 332

Ancient races of North and South America, 298

Andaman Islands, pale butterflies of, 260
white-marked birds of, 263

Anderson, Mr. W. Marshall, on cranium from N. American mound, 296

Andes, very rich in humming-birds, 139

Animal colours, how produced, 184
life in tropical forests, 70

Anthribidæ, 95

Ants, wasps, and bees, 80
numbers of, in India and Malaya, 81-88
destructive to insect-specimens, 85
and vegetation, special relation between, 89

Apatura and Heterochroa, resemblance of species of, 257

Apes, 116

Aqueous vapour of atmosphere, its influence on temperature, 9
quantity at Batavia and Clifton, 10

Arctic plants, large leaves of, 236
flowers and fruits brightly coloured, 237

Areca palm, 45

_Arenga saccharifera_, 43

Argus-pheasant, wonderful plumage of, 205

Arums, 48

Assai of the Amazon, 43

Auckland Isles, handsome flowers of, 238

Audubon, on the ruby humming-birds, 130, 137

Australian Region, mammalia of, 340
birds of, 340
extinct fauna of, 341
its supposed union with S. America, 341

Azara, on food of humming-birds, 135

B.

Bamboos, 52
uses of, 53-58

Bananas, wild, 47

Banana, 48

Barber, Mrs. on colour changes of pupa of _Papilio nireus_, 168

Barbets, 105

Bark, varieties of in tropical forests, 33

Barometer, range of, at Batavia, 24

Batavia, Meteorology of, 4
and London, diagram of mean temperatures, 5
greatest rainfall at, 24
range of barometer at, 24

Bates, Mr., on climate at the Equator, 24
on scarcity of forest-flowers on Amazon, 61
on animal life in Amazon valley, 70
on abundance of butterflies at Ega, 75
on importance of study of butterflies, 78
on leaf-cutting ants, 86
on blind ants, 88
on bird-catching spider, 97
on use of toucan’s bill, 106
on large serpents, 115
on the habits of humming-birds, 132

Bats, 118

Beetles, 94
abundance of, in New Forest-clearings, 96
probable use of horns of, 202

Belt, Mr., on virgin forests of Nicaragua, 62
on aspects of tropical vegetation, 67
on leaf-cutting ants, 86
on an Acacia inhabited by ants, 89
on uses of ants to the trees they live on, 90
on a leaf-like locust, 93
on tree-frogs, 116
on the habits of humming-birds, 133, 134
on uneatable bright-coloured frog, 175
on use of light of glow-worm, 205

Betel-nut, 45

Bill of humming-birds, 129

Biology, by-paths of, illustrated, 251

Birds, 99
how many known, 124
cases of local variation of colour among, 262
influence of locality on colours of, 255
which fertilize flowers, 273, 274
and insects blown to oceanic islands, 308
of Palæarctic Region, 316
of Ethiopian Region, 318
of Oriental Region, 320

Bonelli, Mr., on the Sappho comet humming-bird, 132

Bullock on food of humming-birds, 153

Buprestidæ, 94

Burchell, Dr., on the “stone mesembryanthemum,” 223

Butterflies, abundance of, in tropical forests, 72
conspicuousness of in tropical forests, 73
colours and form of, 74
peculiar habits of tropical, 76
tropical and temperate compared as to colour, 164
females do not choose their partners, 200
with gaily-coloured females, 204
numbers and variety of, 255
influence of locality on colours of, 255

Buttressed trees, 31