: The Geographical Distribution of Animals Volume 2 With a study of the relations of living and extinct faunas as elucidating the past changes of the Earth's surface by Wallace Alfred Russel - Paleontology; Zoogeography

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The excessive and altogether unexampled productiveness of the West Indian islands in land-shells, may be traced to two main sets of causes. The first and least known, consist of the peculiar influences and conditions which render islands always more productive than continents. Whatever these conditions are, they will be more effective where the islands have been long separated from the mainland, as is here undoubtedly the case. It seems most probable that the great development of land-shells in islands, is due to the absence or deficiency of the vertebrata, which on continents supply a variety of species adapted to prey upon these molluscs. This view is supported by the fact, that in such islands as have been united to a continent at no very distant epoch, and still maintain a continental variety of vertebrata, no such special development of land-shells has taken place. If we compare the Philippine islands with the Sunda group, we find the development of vertebrata and land-molluscs in inverse ratio to each other. The same thing occurs if we compare New Zealand and Tasmania; and we have a still more striking example in the Antillean group itself, continental Trinidad having only 20 genera and 38 species, while the highly insular Jamaica has about 30 genera and more than 500 species.

The other causes favourable to the increase and development of land-shells are of a physical nature. A great extent of limestone-rock is one; and in the larger West Indian islands we have a considerable proportion of the surface consisting of this rock. But perhaps equally or more important, is the character of the land surface, and the texture of the exposed rock itself. A much broken surface, with numerous deep ravines, cutting up the whole country into isolated valleys and ridges, seems very favourable to the specialization of forms in this very sedentary class of animals. Equally favourable is a honeycombed and highly-fissured rock-surface, affording everywhere cracks and crannies for concealment. Now, taking Jamaica as an example of the archipelago, we find all these conditions in a wonderful degree. Over a large part of this island, a yard of level ground can hardly be found; but ridges, precipices, ravines, and rock-bound valleys, succeed each other over the whole country. At least five-sixths of the entire surface is limestone, and under the influence of tropical rains this rock is worn, fissured, and honeycombed, so as to afford ample shelter and concealment for land-shells.

Although these islands are so wonderfully rich even now, there is good reason to believe that many species have become extinct since the European occupation of them. When small islands are much cultivated, many of these molluscs which can only live under the shade of forests, are soon extirpated. In St. Croix many species have become extinct at a comparatively recent period, from the burning of forests; and as we know that in all the islands many of the species are excessively local, being often confined to single valleys or ridges, we may be sure that wherever the native forests have disappeared before the hand of man, numbers of land shells have disappeared with them. As some of the smaller islands have been almost denuded of their wood, and in the larger ones extensive tracts have been cleared for sugar cultivation, a very considerable number of species have almost certainly been exterminated.

The only large island of whose geology we have any detailed account, is Jamaica; and taking this as a type of what will probably be found in Cuba and Hayti, we must place the continental period as having occurred after the close of the Miocene, or during some part of the Pliocene epoch, since a large portion of the surface of the former island consists of beds of marine limestone from 2,000 to 3,000 thick, believed to be of Pliocene age. After some time, the land between Hayti and South America subsided, and still later that between Central America and Cuba with Jamaica; but a large tract of land remained insulated, and no doubt supported a very much richer and more varied fauna than now. We have evidence of this in extinct Mammalia of large size, belonging to the peculiar South American family of the chinchillas, which have been found in caves in the small islands of Anguilla, and which, from the character of the land-shells associated with them, are believed to be of Pliocene or Post-pliocene age. This discovery is most interesting, and gives promise of very valuable results from the exploration of the numerous caverns that undoubtedly exist in the abundant limestone strata of the larger islands. This extensive Antillean land, after long continuing undivided, was at length broken up by subsidence into several islands; but as this alone would not account for the almost complete annihilation of the mammalian fauna, it seems probable that the subsidence was continued much farther, so as greatly to reduce the size and increase the number of the islands. This is indicated, by the extensive alluvial plains in Cuba and Hayti, and to a less extent in Jamaica; and by elevated beds of Post-pliocene marls in the latter island.

We have already discussed this subject, both in our account of extinct animals, and in various parts of the present chapter. It is therefore only necessary here, briefly to review and summarise the conclusions we have arrived at.

The whole character of Neotropical zoology, whether as regards its deficiencies or its specialities, points to a long continuance of isolation from the rest of the world, with a few very distant periods of union with the northern continent. The latest important separation took place by the submergence of parts of Nicaragua and Honduras, and this separation probably continued throughout much of the Miocene and Pliocene periods; but some time previous to the coming on of the glacial epoch, the union between the two continents took place which has continued to our day. Earlier submergences of the isthmus of Panama probably occurred, isolating Costa Rica and Veragua, which then may have had a greater extension, and have thus been able to develope their rich and peculiar fauna.

The isthmus of Tehuantepec, at the south of Mexico, may, probably, also have been submerged; thus isolating Guatemala and Yucatan, and leading to the specialization of some of the peculiar forms that now characterise those countries and Mexico.

The West Indian Islands have been long isolated and have varied much in extent. Originally, they probably formed part of Central America, and may have been united with Yucatan and Honduras in one extensive tropical land. But their separation from the continent took place at a remote period, and they have since been broken up into numerous islands, which have probably undergone much submergence in recent times. This has led to that poverty of the higher forms of life, combined with the remarkable speciality, which now characterises them; while their fauna still preserves a sufficient resemblance to that of Central America to indicate its origin.

The great continent of South America, as far as we can judge from the remarkable characteristics of its fauna and the vast depths of the oceans east and west of it, has not during Tertiary, and probably not even during Secondary times, been united with any other continent, except through the intervention of North America. During some part of the Secondary epoch it probably received the ancestral forms of its Edentates and Rodents, at a time when these were among the highest types of Mammalia on the globe. It appears to have remained long isolated, and to have already greatly developed these groups of animals, before it received, in early Tertiary times, the ancestors of its marmosets and monkeys, and, perhaps also, some of its peculiar forms of Carnivora. Later, it received its Camelidae, peccaries, mastodons, and large Carnivora; and later still, just before the Glacial epoch, its deer, tapir, opossums, antelopes, and horses, the two latter having since become extinct. All this time its surface was undergoing important physical changes. What its earlier condition was we cannot conjecture, but there are clear indications that it has been broken up into at least three large masses, and probably a number of smaller ones; and these have no doubt undergone successive elevations and subsidences, so as at one time to reduce their area and separate them still more widely from each other, and at another period to unite them into continental masses. The richness and varied development of the old fauna of South America, as still existing, proves, however, that the country has always maintained an extensive area; and there is reason to believe that the last great change has been a long continued and steady increase of its surface, resulting in the formation of the vast alluvial plains of the Amazon, Orinoko, and La Plata, and thus greatly favouring the production of that wealth of specific forms, which distinguishes South America above all other parts of our globe.

The southern temperate portion of the continent, has probably had a considerable southward extension in late Tertiary times; and this, as well as the comparatively recent elevation of the Andes, has given rise to some degree of intermixture of two distinct faunas, with that proper to South Temperate America itself. The most important of these, is the considerable Australian element that appears in the insects, and even in the reptiles and fresh-water fishes, of South Temperate America. These may be traced to several causes. Icebergs and icefloes, and even solid fields of ice, may, during the Glacial epoch, have afforded many opportunities for the passage of the more cold-enduring groups; while the greater extension of southern lands and islands during the warm periods--which there is reason to believe prevailed in the southern as well as in the northern regions in Miocene times--would afford facilities for the passage of the reptiles and insects of more temperate zones. That no actual land-connection occurred, is proved by the total absence of interchange of the mammals or land-birds of the two countries, no less than by the very fragmentary nature of the resemblances that do exist. The northern element consists almost wholly of insects; and is evidently due to the migration of arctic and north temperate forms along the ridges and plateaus of the Andes; and most likely occurred when these organisms were driven southward at successive cold or Glacial periods.

A curious parallel exists between the past history and actual zoological condition of South America and Africa. In both we see a very ancient land-area extending into the South Temperate zone, isolated at a very early period, and developing only a low grade of Mammalian life; chiefly Edentates and Rodents on the one, Lemurs and Insectivora in the other. Later we find an irruption into both of higher forms, including Quadrumana, which soon acquired a large and special development in the tropical portions of each country. Still later we have an irruption into both of northern forms, which spread widely over the two regions, and having become extinct in the land from whence they came, have been long held to be the original denizens of their adopted country. Such are the various forms of antelopes, the giraffe, the elephant, rhinoceros, and lion in Africa; while in America we have deer and peccaries, the tapir, opossums, and the puma.

On the whole, we cannot but consider that the broad outlines of the zoological history of the Neotropical region can be traced with some degree of certainty; but, owing to the absence of information as to the most important of the geological periods--the Miocene and Eocene--we have no clue to the character of its early fauna, or to the land connections with other countries, which may possibly have occurred in early Tertiary times.

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