Kitabı oku: «Island Life; Or, The Phenomena and Causes of Insular Faunas and Floras», sayfa 29

Of the flora of the smaller Madagascarian islands we possess a fuller account, owing to the recent publication of Mr. Baker's Flora of the Mauritius and the Seychelles, including also Rodriguez. The total number of species in this flora is 1,058, more than half of which (536) are exclusively Mascarene—that is, found only in some of the islands of the Madagascar group, while nearly a third (304) are endemic or confined to single islands. Of the widespread plants sixty-six are found in Africa but not in Asia, and eighty-six in Asia but not in Africa, showing a similar Asiatic preponderance to what is said to occur in Madagascar. With the genera, however, the proportions are different, for I find by going through the whole of the generic distributions as given by Mr. Baker, that out of the 440 genera of wild plants fifty are endemic, twenty-two are Asiatic but not African, while twenty-eight are African but not Asiatic. This implies that the more ancient connection has been on the side of Africa, while a more recent immigration, shown by identity of species, has come from the side of Asia; and it is already certain that when the flora of Madagascar is more thoroughly worked out, a still greater African preponderance will be found in that island.

A few Mascarene genera are found elsewhere only in South America, Australia, or Polynesia; and there are also a considerable number of genera whose metropolis is South America, but which are represented by one or more species in Madagascar, and by a single often widely distributed species in Africa. This fact throws light upon the problem offered by those mammals, reptiles, and insects of Madagascar which now have their only allies in South America, since the two cases would be exactly parallel were the African plants to become extinct. Plants, however, are undoubtedly more long-lived specifically than animals—especially the more highly organised groups, and are less liable to complete extinction through the attacks of enemies or through changes of climate or of physical geography; hence we find comparatively few cases in which groups of Madagascar plants have their only allies in such distant regions as America and Australia, while such cases are numerous among animals, owing to the extinction of the allied forms in intervening areas, for which extinction, as we have already shown, ample cause can be assigned.

Curious Relations of Mascarene Plants.—Among the curious affinities of Mascarene plants we have culled the following from Mr. Baker's volume. Trochetia, a genus of Sterculiaceæ, has four species in Mauritius, one in Madagascar, and one in the remote island of St. Helena. Mathurina, a genus of Turneraceæ, consisting of a single species peculiar to Rodriguez, has its nearest ally in another monotypic genus, Erblichia, confined to Central America. Siegesbeckia, one of the Compositæ, consists of two species, one inhabiting the Mascarene islands, the other Peru. Labourdonasia, a genus of Sapotaceæ, has two species in Mauritius, one in Natal, and one in Cuba. Nesogenes, belonging to the verbena family, has one species in Rodriguez and one in Polynesia. Mespilodaphne, an extensive genus of Lauraceæ, has six species in the Mascarene islands, and all the rest (about fifty species) in South America. Nepenthes, the well-known pitcher plants, are found chiefly in the Malay Islands, South China, and Ceylon, with species in the Seychelles Islands, and in Madagascar. Milla, a large genus of Liliaceæ, is exclusively American, except one species found in Mauritius and Bourbon. Agauria, a genus of Ericaceæ, is found in Madagascar, the Mascarene islands, the plateau of Central Africa, and the Camaroon Mountains in West Africa. An acacia, found in Mauritius and Bourbon (A. heterophylla), can hardly be separated specifically from Acacia koa of the Sandwich Islands. The genus Pandanus, or screw-pine, has sixteen species in the three islands—Mauritius, Rodriguez, and the Seychelles—all being peculiar, and none ranging beyond a single island. Of palms there are fifteen species belonging to ten genera, and all these genera are peculiar to the islands. We have here ample evidence that plants exhibit the same anomalies of distribution in these islands as do the animals, though in a smaller proportion; while they also exhibit some of the transitional stages by which these anomalies have, in all probability, been brought about, rendering quite unnecessary any other changes in the distribution of sea and land than physical and geological evidence warrants.¹⁶⁶

Fragmentary Character of the Mascarene Flora.—Although the peculiar character and affinities of the vegetation of these islands is sufficiently apparent, there can be little doubt that we only possess a fragment of the rich flora which once adorned them. The cultivation of sugar, and other tropical products, has led to the clearing away of the virgin forests from all the lowlands, plateaus, and accessible slopes of the mountains, so that remains of the aboriginal woodlands only linger in the recesses of the hills, and numbers of forest-haunting plants must inevitably have been exterminated. The result is, that nearly three hundred species of foreign plants have run wild in Mauritius, and have in their turn helped to extinguish the native species. In the Seychelles, too, the indigenous flora has been almost entirely destroyed in most of the islands, although the peculiar palms, from their longevity and comparative hardiness, have survived. Mr. Geoffrey Nevill tells us, that at Mahé, and most of the other islands visited by him, it was only in a few spots near the summits of the hills that he could perceive any remains of the ancient flora. Pine-apples, cinnamon, bamboos, and other plants have obtained a firm footing, covering large tracts of country and killing the more delicate native flowers and ferns. The pine-apple, especially, grows almost to the tops of the mountains. Where the timber and shrubs have been destroyed, the water falling on the surface immediately cuts channels, runs off rapidly, and causes the land to become dry and arid; and the same effect is largely seen both in Mauritius and Bourbon, where, originally, dense forest covered the entire surface, and perennial moisture, with its ever-accompanying luxuriance of vegetation, prevailed.

Flora of Madagascar Allied to that of South Africa.—In my Geographical Distribution of Animals I have remarked on the relation between the insects of Madagascar and those of south temperate Africa, and have speculated on a great southern extension of the continent at the time when Madagascar was united with it. As supporting this view I now quote Mr. Bentham's remarks on the Compositæ. He says: "The connections of the Mascarene endemic Compositæ, especially those of Madagascar itself, are eminently with the southern and sub-tropical African races; the more tropical races, Plucheineæ, &c., may be rather more of an Asiatic type." He further says that the Composite flora is almost as strictly endemic as that of the Sandwich Islands, and that it is much diversified, with evidences of great antiquity, while it shows insular characteristics in the tendency to tall shrubby or arborescent forms in several of the endemic or prevailing genera.

Preponderance of Ferns in the Mascarene Flora.—A striking character of the flora of these smaller Mascarene islands is the great preponderance of ferns, and next to them of orchideæ. The following figures are taken from Mr. Baker's Flora for Mauritius and the Seychelles, and from an estimate by M. Frappier of the flora of Bourbon given in Maillard's volume already quoted:—

The cause of the great preponderance of ferns in oceanic islands has already been discussed in my book on Tropical Nature; and we have seen that Mauritius, Bourbon, and Rodriguez must be classed as such, though from their proximity to Madagascar they have to be considered as satellites to that great island. The abundance of orchids, the reverse of what occurs in remoter oceanic islands, may be in part due to analogous causes. Their usually minute and abundant seeds would be as easily carried by the wind as the spores of ferns, and their frequent epiphytic habit affords them an endless variety of stations on which to vegetate, and at the same time removes them in a great measure from the competition of other plants. When, therefore, the climate is sufficiently moist and equable, and there is a luxuriant forest vegetation, we may expect to find orchids plentiful on such tropical islands as possess an abundance of insects adapted to fertilise them, and which are not too far removed from other lands or continents from which their seeds might be conveyed.

Concluding Remarks on Madagascar and the Mascarene Islands.—There is probably no portion of the globe that contains within itself so many and such varied features of interest connected with geographical distribution, or which so well illustrates the mode of solving the problems it presents, as the comparatively small insular region which comprises the great island of Madagascar and the smaller islands and island-groups which immediately surround it. In Madagascar we have a continental island of the first rank, and undoubtedly of immense antiquity; we have detached fragments of this island in the Comoros and Aldabra; in the Seychelles we have the fragments of another very ancient island, which may perhaps never have been continental; in Mauritius, Bourbon, and Rodriguez we have three undoubtedly oceanic islands; while in the extensive banks and coral reefs of Cargados, Saya de Malha, the Chagos, and the Maldive Isles, we have indications of the submergence of many large islands which may have aided in the transmission of organisms from the Indian Peninsula. But between and around all these islands we have depths of 2,500 fathoms and upwards, which renders it very improbable that there has ever been here a continuous land surface, at all events during the Tertiary or Secondary periods of geology.

It is most interesting and satisfactory to find that this conclusion, arrived at solely by a study of the form of the sea-bottom and the general principle of oceanic permanence, is fully supported by the evidence of the organic productions of the several islands; because it gives us confidence in those principles, and helps to supply us with a practical demonstration of them. We find that the entire group contains just that amount of Indian forms which could well have passed from island to island; that many of these forms are slightly modified species, indicating that the migration occurred during late Tertiary times, while others are distinct genera, indicating a more ancient connection; but in no one case do we find animals which necessitate an actual land-connection, while the numerous Indian types of mammalia, reptiles, birds, and insects, which must certainly have passed over had there been such an actual land-connection, are totally wanting. The one fact which has been supposed to require such a connection—the distribution of the lemurs—can be far more naturally explained by a general dispersion of the group from Europe, where we know it existed in Eocene times; and such an explanation applies equally to the affinity of the Insectivora of Madagascar and Cuba; the snakes (Herpetodryas, &c.) of Madagascar and America; and the lizards (Cryptoblepharus) of Mauritius and Australia. To suppose, in all these cases, and in many others, a direct land-connection, is really absurd, because we have the evidence afforded by geology of wide differences of distribution directly we pass beyond the most recent deposits; and when we go back to Mesozoic—and still more to Palæozoic—times, the majority of the groups of animals and plants appear to have had a world-wide range. A large number of our European Miocene genera of vertebrates were also Indian or African, or even American; the South American Tertiary fauna contained many European types; while many Mesozoic reptiles and mollusca ranged from Europe and North America to Australia and New Zealand.

By very good evidence (the occurrence of wide areas of marine deposits of Eocene age), geologists have established the fact that Africa was cut off from Europe and Asia by an arm of the sea in early Tertiary times, forming a large island-continent. By the evidence of abundant organic remains we know that all the types of large mammalia now found in Africa (but which are absent from Madagascar) inhabited Europe and Asia, and many of them also North America, in the Miocene period. At a still earlier epoch Africa may have received its lower types of mammals—lemurs, insectivora, and small carnivora, together with its ancestral struthious birds, and its reptiles and insects of American or Australian affinity; and at this period it was joined to Madagascar. Before the later continental period of Africa, Madagascar had become an island; and thus, when the large mammalia from the northern continent overran Africa, they were prevented from reaching Madagascar, which thenceforth was enabled to develop its singular forms of low-type mammalia, its gigantic ostrich-like Æpyornis, its isolated birds, its remarkable insects, and its rich and peculiar flora. From it the adjacent islands received such organisms as could cross the sea; while they transmitted to Madagascar some of the Indian birds and insects which had reached them.

The method we have followed in these investigations is to accept the results of geological and palæontological science, and the ascertained facts as to the powers of dispersal of the various animal groups; to take full account of the laws of evolution as affecting distribution, and of the various ocean depths as implying recent or remote union of islands with their adjacent continents; and the result is, that wherever we possess a sufficient knowledge of these various classes of evidence, we find it possible to give a connected and intelligible explanation of all the most striking peculiarities of the organic world. In Madagascar we have undoubtedly one of the most difficult of these problems; but we have, I think, fairly met and conquered most of its difficulties. The complexity of the organic relations of this island is due, partly to its having derived its animal forms from two distinct sources—from one continent through a direct land-connection, and from another by means of intervening islands now submerged; but, mainly to the fact of its having been separated from a continent which is now, zoologically, in a very different condition from that which prevailed at the time of the separation; and to its having been thus able to preserve a number of types which may date back to the Eocene, or even to the Cretaceous, period. Some of these types have become altogether extinct elsewhere; others have spread far and wide over the globe, and have survived only in a few remote countries—and especially in those which have been more or less secured by their isolated position from the incursions of the more highly-developed forms of later times. This explains why it is that the nearest allies of the Madagascar fauna and flora are now so often to be found in South America or Australia—countries in which low forms of mammalia and birds still largely prevail;—it being on account of the long-continued isolation of all these countries that similar forms (descendants of ancient types) are preserved in them. Had the numerous suggested continental extensions connecting these remote continents at various geological periods been realities, the result would have been that all these interesting archaic forms, all these defenceless insular types, would long ago have been exterminated, and one comparatively monotonous fauna have reigned over the whole earth. So far from explaining the anomalous facts, the alleged continental extensions, had they existed, would have left no such facts to be explained.

CHAPTER XX
ANOMALOUS ISLANDS: CELEBES

Anomalous Relations of Celebes—Physical Features of the Island—Zoological Character of the Islands Around Celebes—The Malayan and Australian Banks—Zoology of Celebes: Mammalia—Probable Derivation of the Mammals of Celebes—Birds of Celebes—Bird-types Peculiar to Celebes—Celebes not Strictly a Continental Island—Peculiarities of the Insects of Celebes—Himalayan Types of Birds and Butterflies in Celebes—Peculiarities of Shape and Colour of Celebesian Butterflies—Concluding Remarks—Appendix on the Birds of Celebes.

The only other islands of the globe which can be classed as "ancient continental" are the larger Antilles (Cuba, Haiti, Jamaica, and Porto Rico), Iceland, and perhaps Celebes. The Antilles have been so fully discussed and illustrated in my former work, and there is so little fresh information about them, that I do not propose to treat of them here, especially as they fall short of Madagascar in all points of biological interest, and offer no problems of a different character from such as have already been sufficiently explained.

Iceland, also, must apparently be classed as belonging to the "Ancient Continental Islands," for though usually described as wholly volcanic, it is, more probably, an island of varied geological structure buried under the lavas of its numerous volcanoes. But of late years extensive Tertiary deposits of Miocene age have been discovered, showing that it is not a mere congeries of volcanoes; it is connected with the British Islands and with Greenland by seas less than 500 fathoms deep; and it possesses a few mammalia, one of which is peculiar, and at least three peculiar species of birds. It was therefore almost certainly united with Greenland, and probably with Europe by way of Britain, in the early part of the Tertiary period, and thus afforded one of the routes by which that intermigration of American and European animals and plants was effected which we know occurred during some portion of the Eocene and Miocene periods, and probably also in the Pliocene. The fauna and flora of this island are, however, so poor, and offer so few peculiarities, that it is unnecessary to devote more time to their consideration.

There remains the great Malay island—Celebes, which, owing to its possession of several large and very peculiar mammalia, must be classed, zoologically, as "ancient continental"; but whose central position and relations both to Asia and to Australia render it very difficult to decide in which of the primary zoological regions it ought to be placed, or whether it has ever been united with either of the great continents. Although I have pretty fully discussed its zoological peculiarities and past history in my Geographical Distribution of Animals, it seems advisable to review the facts on the present occasion, more especially as the systematic investigation of the characteristics of continental islands we have now made will place us in a better position for determining its true zoo-geographical relations.

Physical Features of Celebes.—This large and still comparatively unexplored island is interesting to the geographer on account of its remarkable outline, but much more so to the zoologist for its curious assemblage of animal forms. The geological structure of Celebes is almost unknown. The extremity of the northern peninsula is volcanic; while in the southern peninsula there are extensive deposits of a crystalline limestone, in some places overlying basalt. Gold is found in the northern peninsula and in the central mass, as well as iron, tin, and copper in small quantities; so that there can be little doubt that the mountain ranges of the interior consist of ancient stratified rocks.

MAP OF CELEBES AND THE SURROUNDING ISLANDS.

The depth of sea is shown by three tints: the lightest indicating less than 100 fathoms, the medium tint less than 1,000 fathoms, and the dark tint more than 1,000 fathoms. The figures show depths in fathoms.

It is not yet known whether Celebes is completely separated from the surrounding islands by a deep sea, but the facts at our command render it probable that it is so. The northern and eastern portions of the Celebes Sea have been ascertained to be from 2,000 to 2,600 fathoms deep, and such depths may extend over a considerable portion of it, or even be much exceeded in the centre. In the Molucca passage a single sounding on the Gilolo side gave 1,200 fathoms, and a large part of the Molucca and Banda Seas probably exceed 2,000 fathoms. The southern portion of the Straits of Macassar is full of coral reefs, and a shallow sea of less than 100 fathoms extends from Borneo to within about forty miles of the western promontory of Celebes; but farther north there is deep water close to the shore, and it seems probable that a deep channel extends quite through the straits, which have no doubt been much shallowed by the deposits from the great Bornean rivers as well as by those of Celebes itself. Southward again, the chain of volcanic islands from Bali to Timor appears to rise out of a deep ocean, the few soundings we possess showing depths of from 670 to 1,300 fathoms almost close to their northern shores. We seem justified, therefore, in concluding that Celebes is entirely surrounded by a deep sea, which has, however, become partially filled up by river deposits, by volcanic upheaval, or by coral reefs. Such shallows, where they exist, may therefore be due to antiquity and isolation, instead of being indications of a former union with any of the surrounding islands.

Zoological Character of the Islands around Celebes.—In order to have a clear conception of the peculiar character of the Celebesian fauna, we must take into account that of the surrounding countries from which we may suppose it to have received immigrants. These we may divide broadly into two groups, those on the west belonging to the Oriental region of our zoological geography, and those on the east belonging to the Australian region. Of the first group Borneo is a typical representative; and from its proximity and the extent of its opposing coasts it is the island which we should expect to show most resemblance to Celebes. We have already seen that the fauna of Borneo is essentially the same as that of Southern Asia, and that it is excessively rich in all the Malayan types of mammalia and birds. Java and Bali closely resemble Borneo in general character, though somewhat less rich and with several peculiar forms; while the Philippine Islands, though very much poorer, and with a greater amount of speciality, yet exhibit essentially the same character. These islands, taken as a whole, may be described as having a fauna almost identical with that of Southern Asia; for no family of mammalia is found in the one which is absent from the other, and the same may be said, with very few and unimportant exceptions, of the birds; while hundreds of genera and of species are common to both.

In the islands east and south of Celebes—the Moluccas, New Guinea, and the Timor group from Lombok eastward—we find, on the other hand, the most wonderful contrast in the forms of life. Of twenty-seven families of terrestrial mammals found in the great Malay islands, all have disappeared but four, and of these it is doubtful whether two have not been introduced by man. We also find here four families of Marsupials, all totally unknown in the western islands. Even birds, though usually more widely spread, show a corresponding difference, about eleven Malayan families being quite unknown east of Celebes, where six new families make their appearance which are equally unknown to the westward.¹⁶⁷

We have here a radical difference between two sets of islands not very far removed from each other, the one set belonging zoologically to Asia, the other to Australia. The Asiatic or Malayan group is found to be bounded strictly by the eastward limits of the great bank (for the most part less than fifty fathoms below the surface) which stretches out from the Siamese and Malayan peninsula as far as Java, Sumatra, Borneo, and the Philippines. To the east another bank unites New Guinea and the Papuan Islands as far as Aru, Mysol, and Waigiou, with Australia; while the Moluccas and Timor groups are surrounded by much deeper water, which forms, in the Banda and Celebes Seas and perhaps in other parts of this area, great basins of enormous depths (2,000 to 3,000 fathoms or even more) enclosed by tracts under a thousand fathoms, which separate the basins from each other and from the adjacent Pacific and Indian Oceans (see map). This peculiar formation of the sea-bottom probably indicates that this area has been the seat of great local upheavals and subsidences; and it is quite in accordance with this view that we find the Moluccas, while closely agreeing with New Guinea in their forms of life, yet strikingly deficient in many important groups, and exhibiting an altogether poverty-stricken appearance as regards the higher animals. It is a suggestive fact that the Philippine Islands bear an exactly parallel relation to Borneo, being equally deficient in many of the higher groups; and here too, in the Sooloo Sea, we find a similar enclosed basin of great depth. Hence we may in both cases connect, on the one hand, the extensive area of land-surface and of adjacent shallow sea with a long period of stability and a consequent rich development of the forms of life; and, on the other hand, a highly broken land-surface with the adjacent seas of great but very unequal depths, with a period of disturbance, probably involving extensive submersions of the land, resulting in a scanty and fragmentary vertebrate fauna.

Zoology of Celebes.—The zoology of Celebes differs so remarkably from that of both the great divisions of the Archipelago above indicated, that it is very difficult to decide in which to place it. It possesses only about sixteen species of terrestrial mammalia, so that it is at once distinguished from Borneo and Java by its extreme poverty in this class. Of this small number four belong to the Moluccan and Australian fauna—there being two marsupials of the genus Cuscus, and two forest rats said to be allied to Australian types.

The remaining twelve species are, generally speaking, of Malayan or Asiatic types, but some of them are so peculiar that they have no near allies in any part of the world; while the rest are of the ordinary Malay type or even identical with Malayan species, and some of these may be recent introductions through human agency. These twelve species of Asiatic type will be now enumerated. They consist of five peculiar squirrels—a group unknown farther east; a peculiar species of wild pig; a deer so closely allied to the Cervus hippelaphus of Borneo that it may well have been introduced by man both here and in the Moluccas; a civet, Viverra tangalunga, common in all the Malay Islands, and also perhaps introduced; the curious Malayan tarsier (Tarsius spectrum) said to be only found in a small island off the coast;—and besides these, three remarkable animals, all of large size and all quite unlike anything found in the Malay Islands or even in Asia. These are a black and almost tailless baboon-like ape (Cynopithecus nigrescens); an antelopean buffalo (Anoa depressicornis), and the strange babirusa (Babirusa alfurus).

None of these three animals last mentioned has any close allies elsewhere, and their presence in Celebes may be considered the crucial fact which must give us the clue to the past history of the island. Let us then see what they teach us. The ape is apparently somewhat intermediate between the great baboons of Africa and the short-tailed macaques of Asia, but its cranium shows a nearer approach to the former group, in its flat projecting muzzle, large superciliary crests, and maxillary ridges. The anoa, though anatomically allied to the buffaloes, externally more resembles the bovine antelopes of Africa; while the babirusa is altogether unlike any other living member of the swine family, the canines of the upper jaws growing directly upwards like horns, forming a spiral curve over the eyes, instead of downwards, as in all other mammalia. An approach to this peculiarity is made by the African wart-hogs, in which the upper tusk grows out laterally and then curves up; but these animals are not otherwise closely allied to the babirusa.

Probable Derivation of the Mammals of Celebes.—It is clear that we have here a group of extremely peculiar, and, in all probability, very ancient forms, which have been preserved to us by isolation in Celebes, just as the monotremes and marsupials have been preserved in Australia, and so many of the lemurs and Insectivora in Madagascar. And this compels us to look upon the existing island as a fragment of some ancient land, once perhaps forming part of the great northern continent, but separated from it far earlier than Borneo, Sumatra, and Java. The exceeding scantiness of the mammalian fauna, however, remains to be accounted for. We have seen that Formosa, a much smaller island, contains more than twice as many species; and we may be sure that at the time when such animals as apes and buffaloes existed, the Asiatic continent swarmed with varied forms of mammals to quite as great an extent as Borneo does now. If the portion of separated land had been anything like as large as Celebes now is, it would certainly have preserved a far more abundant and varied fauna. To explain the facts we have the choice of two theories:—either that the original island has since its separation been greatly reduced by submersion, so as to lead to the extinction of most of the higher land animals; or, that it originally formed part of an independent land stretching eastward, and was only united with the Asiatic continent for a short period, or perhaps even never united at all, but so connected by intervening islands separated by narrow straits that a few mammals might find their way across. The latter supposition appears best to explain the facts. The three animals in question are such as might readily pass over narrow straits from island to island; and we are thus better enabled to understand the complete absence of the arboreal monkeys, of the Insectivora, and of the very numerous and varied Carnivora and Rodents of Borneo, all of which except the squirrels are entirely unrepresented in Celebes by any peculiar and ancient forms.