Island Life/XX

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.

[[Image:Wallace_Island_Life_p452.png|center|375px|thumb| 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.

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.

The question at issue can only be finally determined by geological investigations. If Celebes has once formed part of Asia, and participated in its rich mammalian fauna, which has been since destroyed by submergence, then some remains of this fauna must certainly be preserved in caves or late Tertiary deposits, and proofs of the submergence itself will be found when sought for. If, on the other hand, the existing animals fairly represent those which have ever reached the island, then no such remains will be discovered, and there need be no evidence of any great and extensive subsidence in late Tertiary times.

Birds of Celebes.—Having thus clearly placed before us the problem presented by the mammalian fauna of Celebes, we may proceed to see what additional evidence is afforded by the birds and any other groups of which we have sufficient information. About 164 species of true land-birds are now known to inhabit the island of Celebes itself. Considerably more than half of these (ninety-four species) are peculiar to it; twenty-nine are found also in Borneo and the other Malay Islands, to which they specially belong; while sixteen are common to the Moluccas or other islands of the Australian region; the remainder being species of wide range and not characteristic of either division of the Archipelago. We have here a large preponderance of western over eastern species of birds inhabiting Celebes, though not to quite so great an extent as in the mammalia; and the inference to be drawn from this fact is, simply, that more birds have migrated from Borneo than from the Moluccas—which is exactly what we might expect both from the greater extent of the coast of Borneo opposite that of Celebes, and also from the much greater richness in species of the Bornean than the Moluccan bird-fauna.

It is, however, to the relations of the peculiar species of Celebesian birds that we must turn, in order to ascertain the origin of the fauna in past times; and we must look to the source of the generic types which they represent to give us this information. The ninety-four peculiar species above noted belong to about sixty-six genera, of which about twenty-three are common to the whole Archipelago, and have therefore little significance. Of the remainder, twelve are altogether peculiar to Celebes; twenty-one are Malayan, but not Moluccan or Australian; while ten are Moluccan or Australian, but not Malayan. This proportion does not differ much from that afforded by the non-peculiar species; and it teaches us that, for a considerable period, Celebes has been receiving immigrants from all sides, many of which have had time to become modified into distinct representative species. These evidently belong to the period during which Borneo on the one side, and the Moluccas on the other, have occupied very much the same relative position as now. There remain the twelve peculiar Celebesian genera, to which we must look for some further clue as to the origin of the older portion of the fauna; and as these are especially interesting we must examine them somewhat closely.

Bird-types Peculiar to Celebes.—First we have Artamides, one of the Campephaginæ or caterpillar-shrikes—a not very well-marked genus, and which may have been derived, either from the Malayan or the Moluccan side of the Archipelago. Two peculiar genera of kingfishers—Monachalcyon and Cittura—seem allied, the former to the widespread Todiramphus and to the Caridonax of Lombok, the latter to the Australian Melidora. Another kingfisher, Ceycopsis, combines the characters of the Malayan Ceyx and the African Ispidina, and thus forms an example of an ancient generalised form analogous to what occurs among the mammalia. Streptocitta is a peculiar form allied to the magpies; while Basilornis (found also in Ceram), Enodes, and Scissirostrum, are very peculiar starlings, the latter altogether unlike any other bird, and perhaps forming a distinct sub-family. Meropogon is a peculiar bee-eater, allied to the Malayan Nyctiornis; Rhamphococyx is a modification of Phænicophaes, a Malayan genus of cuckoos; Prioniturus (found also in the Philippines) is a genus of parrots distinguished by raquet-formed tail feathers, altogether unique in the order; while Megacephalon is a remarkable and very isolated form of the Australian Megapodiidæ, or mound-builders.

Omitting those whose affinity may be pretty clearly traced to groups still inhabiting the islands of the western or the eastern half of the Archipelago, we find four birds which have no near allies at all, but appear to be either ancestral forms, or extreme modifications, of Asiatic or African birds—Basilornis, Enodes, Scissirostrum, Ceycopsis. These may fairly be associated with the baboon-ape, anoa, and babirusa, as indicating extreme antiquity and some communication with the Asiatic continent at a period when the forms of life and their geographical distribution differed considerably from what they are at the present time.

But here again we meet with exactly the same difficulty as in the mammalia, in the comparative poverty of the types of birds now inhabiting Celebes. Although the preponderance of affinity, especially in the case of its more ancient and peculiar forms, is undoubtedly with Asia rather than with Australia; yet, still more decidedly than in the case of the mammalia, are we forbidden to suppose that it ever formed a part of the old Asiatic continent, on account of the total absence of so many important and extensive groups of Asiatic birds. It is not single species or even genera, but whole families that are thus absent, and among them families which are pre-eminently characteristic of all tropical Asia. Such are the Timaliidæ, or babblers, of which there are twelve genera in Borneo, and nearly thirty genera in the Oriental Region, but of which one species only, hardly distinguishable from a Malayan form, inhabits Celebes; the Phyllornithidæ, or green bulbuls, and the Pycnonotidæ, or bulbuls, both absolutely ubiquitous in tropical Asia and Malaya, but unknown in Celebes; the Eurylæmidæ, or gapers, found everywhere in the great Malay Islands; the Megalæmidæ, or barbets; the Trogonidæ, or trogons; and the Phasianidæ, or pheasants, all pre-eminently Asiatic and Malayan but all absent from Celebes, with the exception of the common jungle-fowl, which, owing to the passion of Malays for cock-fighting, may have been introduced. To these important families may be added Asiatic and Malayan genera by the score; but, confining ourselves to these seven ubiquitous families, we must ask,—Is it possible, that, at the period when the ancestors of the peculiar Celebes mammals entered the island, and when the forms of life, though distinct, could not have been quite unlike those now living, it could have actually formed a part of the continent without possessing representatives of the greater part of these extensive and important families of birds? To get rid altogether of such varied and dominant types of bird-life by any subsequent process of submersion is more difficult than to exterminate mammalia; and we are therefore again driven to our former conclusion—that the present land of Celebes has never (in Tertiary times) been united to the Asiatic continent, but has received its population of Asiatic forms by migration across narrow straits and intervening islands. Taking into consideration the amount of affinity on the one hand, and the isolation on the other, of the Celebesian fauna, we may probably place the period of this earlier migration in the early part of the latter half of the Tertiary period, that is, in middle or late Miocene times.

Celebes not Strictly a Continental Island.—A study of the mammalian and of the bird-fauna of Celebes thus leads us in both cases to the same conclusion, and forbids us to rank it as a strictly continental island on the Asiatic side. But facts of a very similar character are equally opposed to the idea of a former land-connection with Australia or New Guinea, or even with the Moluccas. The numerous marsupials of those countries are all wanting in Celebes, except the phalangers of the genus Cuscus, and these arboreal creatures are very liable to be carried across narrow seas on trees uprooted by earthquakes or floods. The terrestrial cassowaries are equally absent; and thus we can account for the presence of all the Moluccan or Australian types actually found in Celebes without supposing any land-connection on this side during the Tertiary period. The presence of the Celebes ape in the island of Batchian, and of the babirusa in Bouru, can be sufficiently explained by a somewhat closer approximation of the respective lands, or by a few intervening islands which have since disappeared, or it may even be due to human agency.

If the explanation now given of the peculiar features presented by the fauna of Celebes be the correct one, we are fully justified in classing it as an "anomalous island," since it possesses a small but very remarkable mammalian fauna, without ever having been directly united with any continent or extensive land; and, both by what it has and what it wants, occupies such an exactly intermediate position between the Oriental and Australian regions that it will perhaps ever remain a mere matter of opinion with which it should properly be associated. Forming, as it does, the western limit of such typical Australian groups as the Marsupials among mammalia, and the Trichoglossidæ and Meliphagidæ among birds, and being so strikingly deficient in all the more characteristic Oriental families and genera of both classes, I have always placed it in the Australian Region; but it may perhaps with equal propriety be left out of both till a further knowledge of its geology enables us to determine its early history with more precision.

Peculiarities of the Insects of Celebes.—The only other class of animals in Celebes, of which we have a tolerable knowledge, is that of insects, among which we meet with peculiarities of a very remarkable kind, and such as are found in no other island on the globe. Having already given a full account of some of these peculiarities in a paper read before the Linnean Society—republished in my Contributions to the Theory of Natural Selection,—while others have been discussed in my Geographical Distribution of Animals (Vol. I. p. 434)—I will only here briefly refer to them in order to see whether they accord with, or receive any explanation from, the somewhat novel view of the past history of the island here advanced.

The general distribution of the two best known groups of insects—the butterflies and the beetles—agrees very closely with that of the birds and mammalia, inasmuch as Celebes forms the eastern limit of a number of Asiatic and Malayan genera, and at the same time the western limit of several Moluccan and Australian genera, the former perhaps preponderating as in the higher animals.

Himalayan Types of Birds and Butterflies in Celebes.—A curious fact of distribution exhibited both among butterflies and birds, is the occurrence in Celebes of species and genera unknown to the adjacent islands, but only found again when we reach the Himalayan mountains or the Indian Peninsula. Among birds we have a small yellow flycatcher (Myialestes helianthea), a flower-pecker (Pachyglossa aureolimbata), a finch (Munia brunneiceps), and a roller (Coracias temminckii), all closely allied to Indian (not Malayan) species,—all the genera, except Munia, being, in fact, unknown in any Malay island. An exactly parallel case is that of a butterfly of the genus Dichorrhagia, which has a very close ally in the Himalayas, but nothing like it in any intervening country. These facts call to mind the similar case of Formosa, where some of its birds and mammals occurred again, under identical or closely allied forms, in the Himalayas; and in both instances they can only be explained by going back to a period when the distribution of these forms was very different from what it is now.

Peculiarities of Shape and Colour in Celebesian Butterflies.—Even more remarkable are the peculiarities of shape and colour in a number of Celebesian butterflies of different genera. These are found to vary all in the same manner, indicating some general cause of variation able to act upon totally distinct groups, and produce upon them all a common result. Nearly thirty species of butterflies, belonging to three different families, have a common modification in the shape of their wings, by which they can be distinguished at a glance from their allies in any other island or country whatever; and all these are larger than the representative forms inhabiting most of the adjacent islands. No such remarkable local modification as this is known to occur in any other part of the globe; and whatever may have been its cause, that cause must certainly have been long in action, and have been confined to a limited area. We have here, therefore, another argument in favour of the long-continued isolation of Celebes from all the surrounding islands and continents—a hypothesis which we have seen to afford the best, if not the only, explanation of its peculiar vertebrate fauna.

Concluding Remarks.—If the view here given of the origin of the remarkable Celebesian fauna is correct, we have in this island a fragment of the great eastern continent which has preserved to us, perhaps from Miocene times, some remnants of its ancient animal forms. There is no other example on the globe of an island so closely surrounded by other islands on every side, yet preserving such a marked individuality in its forms of life; while, as regards the special features which characterise its insects, it is, so far as yet known, absolutely unique. Unfortunately very little is known of the botany of Celebes, but it seems probable that its plants will to some extent partake of the speciality which so markedly distinguishes its animals; and there is here a rich field for any botanist who is able to penetrate to the forest-clad mountains of its interior.

APPENDIX TO CHAPTER XX

The following list of the Land Birds of Celebes and the adjacent islands which partake of its zoological peculiarities, in which are incorporated all the species discovered up to 1890, has been drawn up from the following sources:—


 * 1. A List of the Birds known to inhabit the Island of Celebes, By Arthur, Viscount Walden, F.R.S. (Trans. Zool. Soc. 1872. Vol. viii. pt. ii.)


 * 2. Intorno al Genere Hermotimia. (Rchb.) Nota di Tommaso Salvadori. (Atti della Reale Accademia delle Scienze di Torino. Vol x. 1874.)


 * 3. Intorno a due Collezioni di Ucelli di Celebes—Note di Tommaso Salvadori. (Annali del Mus. Civ. di St. Nat. di Genova. Vol. vii. 1875.)


 * 4. Beiträge zur Ornithologie von Celebes und Sangir. Von Dr. Friedrich Brüggemann. Bremen, 1876.


 * 5. Intorno a due piccole Collezioni di Ucelli di Isole Sanghir e di Tifore. Nota di Tommaso Salvadori. (Annali del Mus. Civ. di St. Nat. di Genova. Vol. ix. 1876-77.)


 * 6. Intorno alle Specie di Nettarinie delle Molucche e del Gruppo di Celebes. Note di Tommaso Salvadori. (Atti della Reale Accad. delle Scienze di Torino. Vol. xii. 1877.)


 * 7. Descrizione di tre Nuove Specie di Ucelli, e note intorno ad altre poco conosciute delle Isole Sanghir. Per Tommaso Salvadori. (L. c. Vol. xiii. 1878.)


 * 8. Field Notes on the Birds of Celebes. By A. B. Meyer, M.D., &c. (Ibis, 1879.)


 * 9. On the Collection of Birds made by Dr. Meyer during his Expedition to New Guinea and some neighbouring Islands. By R. Boulder Sharpe. (Mitth. d. kgl. Zool. Mus. Dresden, 1878. Heft 3.) New species from the Sula and Sanghir Islands are described.


 * 10. List of Birds from the Sula Islands (East of Celebes) with Descriptions of the New Species. By Alfred Russel Wallace, F.Z.S. (Proc. Zool. Soc. 1862, p. 333.)


 * 11. The Zoological Record, and "The Ibis" to 1890.

LIST OF LAND BIRDS OF CELEBES

N.B.—The Species marked with an * are not included in Viscount Walden's list. For these only, an authority is usually given.

167

168 For outline figures of the chief types of these butterflies, see my Malay Archipelago, Vol. I. p. 441, or p. 216 of the tenth edition.