This Papuan fauna is so interesting and remarkable, that it seems advisable to give lists of these several classes of generic types.
I. Genera occurring in the Papuan Islands which are characteristic of the Australian region (89). Those marked with an asterisk are exclusively Papuan.
The chief points of interest here are the richness and specialization of the parrots, pigeons, and kingfishers; the wonderful paradise-birds; the honeysuckers; and some remarkable flycatchers.The most prominent deficiencies, as compared with Australia, are in Sylviidæ, Timaliidæ, Ploceidæ, Platycercidæ, and Falconidæ.
II. The genera which are characteristic of the whole Malay Archipelago are the following (10):—
III. The curious set of genera apparently of Indo-Malayan origin, but unknown in the Moluccas, are as follows:—
The above six birds are very important as indicating past changes in the Austro-Malay Islands, and we must say a few words about each. (1)Eupetesis very remarkable, since the New Guinea birds resemble in all important characters that which is confined to Malacca and Sumatra. They are probably the survivors of a once wide-spread Malayan group. (2)AlcippeorDrymocataphus(for in which genus the birds should be placed is doubtful) seems another clear case of a typical Indo-Malayan form occurring in New Guinea and Java, but in no intervening island. (3)Pomatorhinusis a most characteristic Himalayan and Indo-Malayan genus, occurring again in New Guinea and also in Australia, but in no intermediate island. The New Guinea bird seems as nearly related to Oriental as Australian species. (4)Arachnotherais exactly parallel toAlcippe, occurring nowhere east of Borneo except in New Guinea. (5)Prionochilus, a small black bird, sometimes classed as a distinct genus, but evidently allied to thePrionochiliof the Indo-Malay Islands. (6)Eulabes, the genus which contains the well known Mynahs of India, extends east of Java as far as Flores, but is not found in Celebes or the Moluccas. The two New Guinea species are sometimes classed in different genera, but they are undoubtedly allied to the Mynahs of India and Malaya.
We find then, that while the ornithology of New Guinea ispreeminently Australian in character and possesses many peculiar developments of Australian types, it has also—as might be expected from its geographical position, its climate, and its vegetation—received an infusion of Malayan forms. But while one group of these is spread over the whole Archipelago, and occasionally beyond it, there is another group which presents the unusual and interesting feature of discontinuous distribution, jumping over a thousand miles of island-studded sea from Java and Borneo to New Guinea itself. It is a parallel case to that of Java in the Oriental region, which we have already discussed, but the suggested explanation in that case is more difficult to apply here. The recent soundings by theChallengershow us, that although the several islands of the Moluccas are surrounded by water from 1,200 to 2,800 fathoms deep, yet these seas form inclosed basins with rims not more than from 400 to 900 fathoms deep, suggesting the idea of great lakes or inland seas which have sunk down bodily with the surrounding land, or that enormous local and restricted elevations and subsidences have here occurred. We have also the numerous small islands and coral banks south of Celebes and eastward towards Timor-Laut and the Aru Islands, indicating great subsidence; and it is possible that there was an extension of Papua to the west, approaching sufficiently near to Java to receive occasional straggling birds of Indo-Malay type, altogether independent of the Moluccas to the north.
Bright Colours and Ornamental Plumage of New Guinea Birds.—One of the most striking features of Papuan ornithology is the large proportion which the handsome and bright-coloured birds bear to the more obscure species. That this is really the case has been ascertained by going over my own collections, made at Aru and New Guinea, and comparing them with my collection made at Malacca—a district remarkable for the number of handsome birds it produces. Using, as nearly as possible, the same standard of beauty, about one-third of the Malacca birds may be classed as handsome,[14]while in Papua the proportion comes out exactly one-half. This is due, in part to the great abundance ofparrots, cockatoos, and lories, almost all of which are beautiful; and of pigeons, more than half of which are very beautiful; as well as to the numerous kingfishers, most of which are excessively brilliant. Then we have the absence of thrushes, and the very small numbers of the warblers, shrikes, and Timaliidæ, which are dull-coloured groups; and, lastly, the presence of numerous gay pittas, flycatchers, and the unequalled family of paradise-birds. A large number of birds adorned with metallic plumage is also a marked feature of this fauna, more than a dozen genera being so distinguished. Among the remarkable forms arePeltops, a flycatcher, long classed as one of the Indo-Malayan Eurylæmidæ, which it resembles both in bill and coloration;Machærirhynchus, curious little boat-billed flycatchers; andTodopsis, a group of terrestrial flycatchers with the brilliant colours ofPittaorMalurus. The paradise-birds present the most wonderful developments of plumage and the most gorgeous varieties of colour, to be found among passerine birds. The great whiskered-swift, the handsomest bird in the entire family, has its head-quarters here. Among kingfishers the elegant long-tailedTanysipteræare preeminent, whether for singularity or beauty. Among parrots, New Guinea possesses the great black cockatoo, one of the largest and most singular birds in the order;Nasiterna, the smallest of known parrots; andCharmosyna, perhaps the most elegant. Lastly, among the pigeons we have the fine crowned-pigeons, the largest and most remarkable group of the order.
Plate X.
SCENE IN NEW GUINEA, WITH CHARACTERISTIC ANIMALS.
SCENE IN NEW GUINEA, WITH CHARACTERISTIC ANIMALS.
Plate X. Illustrating the Ornithology of New Guinea.—The wonderful ornithological fauna we have just sketched, could only be properly represented in a series of elaborate coloured plates. We are obliged here to confine ourselves to representing a few of the more remarkable types of form, as samples of the great number that adorn this teeming bird-land. The large central figure is the fine twelve-wired paradise-bird (Seleucidesalbus), one of the most beautiful and remarkable of the family. Its general plumage appears, at first sight, to be velvety black; but on closer examination, and by holding the bird in various lights, it is found that every part of it glows with the most exquisite metallic tints—rich bronze, intense violet, and, on theedges of the breast-feathers, brilliant green. An immense tuft of dense plumes of a fine orange-buff colour, springs from each side of the body, and six of these on each side terminate in a black curled rachis or shaft, which form a perfectly unique adornment to this lovely bird. To appreciate this wonderful family (of which no good mounted collection exists) the reader should examine the series of plates in Mr. Elliot's great work on the Paradiseidæ, where every species is figured of the size of life, and with a perfection of colouring that leaves little to be desired.
Below theSeleucidesis one of the elegant racquet-tailed king-hunters (Tanysiptera galatea) whose plumage of vivid blue and white, and coral-red bill, combined with the long spatulate tail, renders this bird one of the most attractive of the interesting family of kingfishers. On a high branch is seated the little Papuan parroquet (Charmosyna papuensis), one of the Trichoglossidæ, or brush-tongued parrots,—richly adorned in red and yellow plumage, and with an unusually long and slender tail. On the ground is the well-known crowned pigeon (Goura coronata), a genus which is wholly confined to New Guinea and a few of the adjacent islands. One of the very few Papuan mammals, a tree-kangaroo (Dendrolagus inustus), is seated on a high branch. It is interesting, as an arboreal modification of a family which in Australia is purely terrestrial; and as showing how very little alteration of form or structure is needed to adapt an animal to such a different mode of life.
Reptiles and Amphibia.—Of these classes comparatively little is at present known, but there is evidence that the same intermixture of Oriental and Australian forms that occurs in birds and insects, is also found here. Dr. A. B. Meyer, the translator of this work into German, and well known for his valuable discoveries in New Guinea, has kindly furnished me with a manuscript list of Papuan reptiles, from which most of the information I am able to give is derived.
Of Snakes, 24 genera, are known, belonging to 11 families. Six of the genera are Oriental,—Calamaria,Cerberus,Chrysopelea,Lycodon,Chersydrus, andOphiophagus. Four are Australian,—Morelia,Liasis,Diemenia, andAcanthophis; while four others are more especially Papuan,—Dibamus(Typhlopidæ),Brachyorros—a sub-genus of the wide-spreadRhabdosoma(Calamariidæ), found also in Timor;NardoaandEnygrus(Pythonidæ), ranging from the Moluccas to the Fiji Islands. The rest are either common to the Oriental and Australian regions or of wide range.
Of Lizards also, 24 genera are recorded, belonging to 5 families. Three only are peculiarly Oriental,—Eumeces,Tiaris, andNycteridium; but another,Gonyocephalus, is Malayan, ranging from Java and Borneo to the Pelew Islands. Three are Australian,—Cyclodus,Heteropus, andGehyra; while six are especially Papuan,—Keneuxia(extending to the Philippines),Elania,Carlia(to North Australia),Lipinia(to the Philippine Islands), andTribolonotus,—all belonging to the Scincidæ; andAruabelonging to the Agamidæ. We must addCryptoblepharus, which is confined to the Australian region, except a species in Mauritius. The other genera have a wider distribution.
The preponderant Oriental element in the snakes as compared with the lizards, is suggestive of the dispersal of the former being dependent on floating trees, or even on native canoes, which for an unknown period have traversed these seas, and in which various species of snakes often secrete themselves. This seems the more probable, as snakes are usually more restricted in their range than lizards, and exhibit less numerous examples of widespread genera and species. The other orders of reptiles present no features of interest.
Of Amphibia only 8 genera are known, belonging to 6 families.Rana,Hylarana, andHylaare wide-spread genera, the former being, however, absent from Australia.Hyperolius,Pelodryas,Litoria, andAsterophrysare Australian; whilePlatymantisis Polynesian, with a species in the Philippine Islands. Hence it appears that the amphibia, so far as yet known, exhibit no Oriental affinity; and this is a very suggestive fact. We have seen (p. 29) that salt water is almost a complete barrier to the dispersal of these creatures; so that the wholly Australian character of the Papuan batrachia is what we might expect, if, as here advocated, no actual land connection betweenthe Oriental and Australian regions, has probably occurred during the entire Tertiary and Post-tertiary periods.
Insects.—The general character of the Papuan insects has been sufficiently indicated in our sketch of the Entomology of the region. We will here only add, that the metallic lustre so prevalent among the birds, is also apparent in such insects asSphingnotus mirabilis, a most brilliant metallic Longicorn;Lomaptera wallaceiandAnacamptorhina fulgida, Cetonii of intense lustre;Calodema wallaceiamong the Buprestidæ; and the elegant blueEupholiamong the weevils. Even among moths we haveCocytia durvillii, remarkable for its brilliant metallic colours.
The Moluccas.—The islands of Gilolo, Bouru, and Ceram, with several smaller islands adjacent, together with Sanguir, and perhaps Tulour or Salibaboo to the north-west, and the islands from Ke to Timor-Laut to the south-east, form the group of the Moluccas or Spice-Islands, remarkable for the luxuriance of their vegetation and the extreme beauty of their birds and insects. Their Mammalia are of Papuan character, with some foreign intermixture. Two genera of the New Guinea marsupials,BelideusandCuscus, abound; and we have also the wide-spreadSus. But besides these, we find no less than five genera of placental Mammals quite foreign to the Papuan or Australian faunas. These are 1.Cynopithecus nigrescens, found only in the small island of Batchian, and probably introduced from Celebes, where the same ape occurs. 2.Viverra tangalunga, a common Indo-Malayan species of civet, probably introduced. 3.Cervus hippelaphus, var.Moluccensis, a deer abundant in all the islands, very close to a Javan species and almost certainly introduced by man, perhaps very long ago. 4.Babirusa alfurus, the babirusa, found only in the island of Bouru, and perhaps originally introduced from Celebes. 5.Sorexsp., small shrews. With the exception of the last,allthese species are animals habitually domesticated and kept in confinement by the Malays; and when we consider that none of the smaller Mammalia of Java and Borneo, numbering at least fifty different species, are foundin any of the Moluccas, we can hardly suppose that such large animals as the deer and ape, could have reached them by natural means. There is every reason to believe, therefore, that the indigenous Mammalia of the Moluccas are wholly of Papuan stock, and very limited in number.
The birds are much more varied and interesting. About 200 species of land-birds are now known, belonging to 85 genera. Of the species about 15 are Indo-Malayan, 32 Papuan, and about 140 peculiar. Of the genera only two are peculiar,—Semioptera, a paradise bird, andLycocorax, a singular form of Corvidæ; but there is also a peculiar rail-like wader,Habroptila. One genus,Basilornis, is found only in Ceram and Celebes; another,Scythrops, is Australian, and perhaps a migrant. About 30 genera are characteristic Papuan types, and 37 others, of more or less wide range, are found in New Guinea and were therefore probably derived thence. There remains a group of birds which are not found in New Guinea, and are either Palæarctic or Oriental. These are 13 in number as follows:—
Of these theMonticola, found only in Gilolo, appears to be a straggler or migrant from the Philippine islands.Acrocephalus, of which four species occur, is a wide-spread group; one of the Moluccan birds is an Australian and another a North-Asian species, which perhaps indicates that there has long been some migration southward from island to island, across the Moluccas.Cisticolais a genus of very wide range, extending to Australia.Hypolaisis probably a modified form of a Chinese or Javanese species.Crinigeris a pure Indo-Malay form, represented here by three fine species.Butalisis a Chinese species, no doubt straggling southward.BudytesandCorydallaare widespread Oriental and Palæarctic species or slight modifications of them.Hydrornisis a Malayan form of Pittidæ.Batrachostomusis a distinct representative of a purely Indo-Malay genus.Loriculusis Malayan, and especially Philippine, but it reaches as far as Mysol.Treronis here at its eastern limit, and is represented in Bouru and Ceram by one of the most beautiful species.Neopus, a Malayan eagle, is said to occur in the Moluccas. We find then only three characteristic Indo-Malay types in the Moluccas,—Criniger,Batrachostomus, andTreron. All are represented by distinct and well marked species, indicating a somewhat remote period since their ancestors entered the district, but all are birds of considerable powers of flight, so that a very little extension of the islands in a south-westerly direction would afford the means of transmission, but this could not well have been by way of Celebes, because the two former genera are unknown in that island.
It is evident, therefore, that the Moluccas are wholly Papuan in their zoology; yet they are no less clearly derivative, and must have obtained their original immigrants under conditions that rendered a full representation of the fauna impossible. Such remarkable and dominant types as the eleven genera of Paradiseidæ, withCracticus,Rectes,Todopsis,Machærirhynchus,Gerygone,Dacelo,Podargus,Cyclopsitta,Microglossum,Nasiterna,Chalcopsitta, andGoura,—all characteristic Papuan groups, found in almost all the islands and most of them very abundant, are yet totally absent from the Moluccas. Taking this, in conjunction with the absence of the two genera of Papuan kangaroos and the other smaller groups of marsupials, and we must be convinced that the Moluccas cannot be mere fragments of the old Papuan land, or they would certainly, in some one or other of their large and fertile islands, have preserved a more complete representation of the parent fauna. Most of the Moluccan birds are very distinct from the allied species of New Guinea; and this would imply that the entrance of the original forms took place at a remote period. The two peculiar genera with clearly Papuan affinities, show the same thing. The cassowary, found only in the large island of Ceram and distinct from any Papuan species, would however seem to have required a land connection for its introduction, almost as much as any of the larger mammalia.
Taking all the facts into consideration, I would suggest as the most probable explanation, that if the Moluccas ever formed part of the main Papuan land, they were separated at an early date, and subsequently so greatly submerged as to destroy a large proportion of their fauna. They have since risen, and have probably been larger than at present, and rather more closely approximated to the parent land, whence they received a considerable immigration of such animals as were adapted to cross narrow seas. This gave them several Papuan forms, but still left them without a number of the types more especially confined to the forest depths, or powerful enough to combat the gales which often blow weaker flyers out to sea. Most of the birds whose absence from the Moluccas is so conspicuous belong to one or other of these classes.
Among the most characteristic birds of the Moluccas are the handsome crimson lories of the generaLoriusandEos. These are found in every island (but not in Celebes or the Timor group); and a fine species ofEos, peculiar to the small islands of Siau and Sanguir, just north of Celebes, obliges us to place these with the Moluccas instead of with the former island, to which they seem most naturally to belong. The crimson parrots of the genusEclectusare almost equally characteristic of the Moluccas, and add greatly to the brilliancy of the ornithology of these favoured islands.
Reptiles.—The Reptiles, so far as known, appear to agree in their distribution with the other vertebrates. In some small collections from Ceram there were no less than six of the genera peculiar to the Australian region, and which were before only known from Australia itself. These are, of snakes,LiasisandEnygrus, genera of Pythonidæ; withDiemeniaandAcanthophis(Elapidæ); of lizards,Cyclodus, a genus of Scincidæ; and of Amphibia, a tree-frog of the genusPelodryas.
Insects—Peculiarities of the Moluccan Fauna.—In insects the Moluccas are hardly, if at all, inferior to New Guinea itself. The islands abound in grandPapiliosof the largest size and extreme beauty; and it is a very remarkable fact, that when the closely-allied species of the Moluccas and New Guinea are compared,the former are almost always the largest. As examples may be mentioned,Ornithoptera priamusandO. helenaof the Moluccas, both larger than the varieties (or species) of Papua;Papilio ulyssesanddeiphobusof Amboyna, usually larger than their allies in New Guinea;Hestia idea, the largest species of the genus;Diadema pandarusandCharaxes euryalus, both larger than any other species of the same genera in the whole archipelago. It is to be noted also, that in the Moluccas, the very largest specimens or races seem always to come from the small island of Amboyna; even those of Ceram, the much larger island to which it is a satellite, being almost always of less dimensions. Among Coleoptera, the Moluccas produceEuchirus longimanus, one of the largest and most remarkable of the Lamellicornes;Sphingnotus dunningi, the largest of the Austro-Malayan Tmesisterninæ; aSphenura, the largest and handsomest of an extensive genus; an unusually largeSchizorhina(Cetoniidæ); and some of the most remarkable and longest-horned Anthotribidæ. Even in birds the same law may be seen at work,—in theTanysiptera naisof Ceram, which has a larger tail than any other in the genus; inCentropus goliathof Gilolo, being the largest and longest-tailed species; inHydrornis maximusof Gilolo, the largest and perhaps the most elegantly and conspicuously coloured of all the Pittidæ; inPlatycercus amboinensis, being pre-eminent in its ample blue tail; in the two Moluccan lories andEos rubra, being more conspicuously red than the allied New Guinea species; and inMegapodius wallaceiof Bouru, being the only species of the genus conspicuously marked and banded.
All these examples, of larger size, of longer tails or other appendages, and of more conspicuous colouring, are probably indications of a less severe struggle for existence in these islands than in the larger tract of New Guinea, with a more abundant and more varied fauna; and this may apply even to the smaller islands, as compared with the larger in the immediate vicinity. The limited number of forms in the small islands compared with a similar area in the parent land, implies, perhaps, less competition and less danger; and thus allows, where all other conditions are favourable, an unchecked and continuousdevelopment in size, form, and colour, until they become positively injurious. This law may not improbably apply to the New Guinea fauna itself, as compared with that of Borneo or any other similar country; and some of its peculiarities (such as its wonderful paradise-birds) may be due to long isolation, and consequent freedom from the influence of any competing forms. The difference between the very sober colours of the Coleoptera, and in a less degree of the birds, of Borneo, as compared with their brilliancy in New Guinea, always struck me most forcibly, and was long without any, even conjectural, explanation. It is not the place here to go further into this most curious and interesting subject. The reader who wishes for additional facts to aid him in forming an opinion, should consult Mr. Darwin'sDescent of Man, chapters x. to xv.; and my ownContributions to the Theory of Natural Selection, chapters iii. and iv.
Timor Group.—Mammalia.—In the group of islands between Java and Australia, from Lombok to Timor inclusive, we find a set of mammals similar to those of the Moluccas, but some of them different species. A wide-spread species ofCuscusrepresents the Papuan element. ASorexand a peculiar species of wild pig, we may also accept as indigenous. Three others have almost certainly been introduced. These are, (1.)Macacus cynomolgus, the very commonest Malay monkey, which may have crossed the narrow straits from island to island between Java and Timor, though it seems much more probable that it was introduced by Malays, who constantly capture and rear the young of this species. (2.)Cervus timoriensis, a deer, said to be a distinct species, inhabits Timor, but it is probably only a variety of theCervus hippelaphusof Java. This animal is, however, much more likely to have crossed the sea than the monkey. (3.)Paradoxurus fasciatus, takes the place ofViverra tangalungain the Moluccas, both common and wide-spread civets which are often kept in confinement by the Malays. TheFelis megalotis, long supposed to be a native of Timor, has been ascertained by Mr. Elliot to belong to a different country altogether.
Birds.—The birds are much more interesting, since they aresufficiently numerous to allow us to determine their relations, and trace their origin, with unusual precision. There are 96 genera and 160 species of land-birds known to inhabit this group of islands; and on a careful analysis, they are found to be almost equally related to the Australian and Oriental regions, 30 genera being distinctly traceable to the former, and the same number to the latter. Their connection with the Moluccas is shown by the presence of the generaMimeta,Geoffroyus,Cacatua,Ptilopus, andIanthænas, together withMegapodiusandCerchneisrepresented by Moluccan species.Turacœnashows a connection with Celebes, andScopsis represented by a Celebesian species. The connection with Australia is shown by the generaSphæcothera,Gerygone,Myiagra,Pardalotus,Gliciphila,Amadina, andAprosmictus; whileMilvus,Hypotriorchis,Eudynamis, andEurystomus, are represented by Australian species. Other genera confined to or characteristic of the Australian region, areRhipidura,Monarcha,Artamus,Campephaga,Pachycephala,Philemon,Ptilotis, andMyzomela.
We now come to the Indo-Malay or Javan element represented by the following genera:
Such genera asMeropsandStrix, which are as much Australian as Oriental, are inserted here because they are represented by Javan species. The list is considerably swelled by genera which have reached Lombok across the narrow strait from Baly, but have passed no further. Such areZoothera,Orthotomus,Pycnonotus,PericrocotusandStrix. A much larger number (12) stop short at Flores, leaving only 13, indicated in the list by (T) after their names, which reach Timor. It is evident, therefore, that these islands have been stocked from three chief sources,—theMoluccas (with New Guinea and Celebes,) Australia, and Java. The Moluccan forms may well have arrived as stragglers from island to island, aided by whatever facilities have been afforded by lands now submerged. Most of the remainder have been derived either from Australia or from Java; and as their relations to these islands are very interesting, they must be discussed with some detail.
Origin of the Timorese Fauna.—We must first note, that 80 species, or exactly one-half of the land-birds of the islands, are peculiar and mostly very distinct, intimating that the immigration commenced long enough back to allow of much specific modification. There is also one peculiar genus of kingfishers,Caridonax, found only in Lombok and Flores, and more allied to Australian than to Oriental types. The fine white-banded pigeons (s. g.Leucotreron) are also almost peculiar; one other less typical species only being known, a native of N. Celebes. In order to compare the species with regard to their origin, we must first take away those of wide distribution from which no special indications can be obtained. In this case 49 of the land-birds must be deducted, leaving 111 species which afford good materials for comparison. These, when traced to their origin, show that 62 came from some part of the Australian region, 49 from Java or the Oriental region. But if we divide them into two groups, the one containing the species identical with those of the Australian or Oriental regions, the other containingalliedorrepresentativespecies peculiar to the islands, we have the following result:
This table is very important, as indicating that the connectionwith Australia was probably earlier than that with Java; since the majority of the Australian species have become modified, while the majority of the Oriental species have remained unchanged. This is due, no doubt, in part to the continued immigration of fresh individuals from Java, after that from Australia, the Moluccas and New Guinea had almost wholly ceased. We must also notice the very small proportion of the genera, either of Australia or Java, that have found their way into these islands, many of the largest and most wide-spread groups in both countries being altogether absent. Taking these facts into consideration, it is pretty clear that there has been no close and long-continued approximation of these islands to any part of the Australian region; and it is also probable that they were fairly stocked with such Australian groups as they possess before the immigration from Java commenced, or a larger number of characteristic Oriental forms would have been able to have established themselves.
On looking at our map, we find that a shallow submerged bank extends from Australia to within about twenty miles of the coast of Timor; and this is probably an indication that the two countries were once only so far apart. This would have allowed the purely Australian types to enter, as they are not numerous; there being about 6 Australian species, and 10 or 12 representatives of Australian species, in Timor. All the rest may have been derived from the Moluccas or New Guinea, being mostly wide-spread genera of the Australian region; and the extension of Papua in a south-west direction towards Java (which was suggested as a means of providing New Guinea with peculiar Indo-Malay types not found in any other part of the region) may have probably served to supply Timor and Flores with the mass of their Austro-Malayan genera across a narrow strait or arm of the sea. Lombok, Baly, and Sumbawa were probably not then in existence, or nothing more than small volcanic cones rising out of the sea, thus leaving a distance of 300 miles between Flores and Java. Subsequently they grew into islands, which offered an easy passage for a number of Indo-Malay genera into such scantily stocked territories as Flores and Timor. Thenorth coast of Australia then sank, cutting off the supply from that country; and this left the Timorese group in the position it now occupies.
The reptiles and fishes of this group are too little known to enable us to make any useful comparison.
Insects.—The insects, though not numerous, present many fine species, some quite unlike any others in the Archipelago. Such are—Papilio liris,Pieris læta,Cirrochroa lamarckiiandC. leschenaultiiamong butterflies. The Coleoptera are comparatively little known, but in the insects generally the Indo-Malay element predominates. This may have arisen from the peculiar vegetation and arid climate not being suitable to the Papuan insects. Why Australian forms did not establish themselves we cannot conjecture; but the field appears to have been open to immigrants from Java, the climate and vegetation of which island at its eastern extremity approximates to that of the Timorese group. The insects are, however, so peculiarly modified as to imply a very great antiquity, and this is also indicated by a group of Sylviine birds here classed underOreicola, but some of which probably form distinct genera. There may, perhaps, have been an earlier and a later approximation to Java, which, with the other changes indicated, would account for most of the facts presented by the fauna of these islands. One deduction is, at all events, clear: the extreme paucity of indigenous mammals along with the absence of so many groups of birds, renders it certain that the Timorese islands did not derive their animal life by means of an actual union with any of the large islands either of the Australian or the Oriental regions.
Celebes Group.—We now come to the Island of Celebes, in many respects the most remarkable and interesting in the whole region, or perhaps on the globe, since no other island seems to present so many curious problems for solution. We shall therefore give a somewhat full account of its peculiar fauna, and endeavour to elucidate some of the causes to which its zoological isolation may be attributed.
Mammalia.—The following is the list of the mammalia ofCelebes as far as at present known, though many small species may yet be discovered.
The first—a large black ape—is itself an anomaly, since it is not closely allied to any other form of quadrumana. Its flat projecting muzzle, large superciliary crests and maxillary ridges, with the form and appearance of its teeth, separate it altogether from the genusMacacus, as represented in the Indo-Malay islands, and ally it closely to the baboons of Africa.[15]We have already seen reason to suppose that it has been carried to Batchian, and there is some doubt about the allied species or variety (C. niger) of the Philippines being really indigenous there; in which case this interesting form will remain absolutely confined to Celebes. (2.) The tarsier is a truly Malayan species, but it is said to occur in a small island at the northern extremity of Celebesand on some of the Philippine Islands. It might possibly have been introduced there. (3) and (4)—a civet and a deer—are, almost certainly, as in the Moluccas, introduced species. (5.)Anoa depressicornis.This is one of the peculiar Celebesian types; a small straight-horned wild-bull, anatomically allied to the buffaloes, and somewhat resembling the bovine antelopes of Africa, but having no near allies in the Oriental region. (6.)Sus Celebensis; a peculiar species of wild-pig. (7.)Babirusa alfurus; another remarkable type, having no near allies. It differs in its dentition from the typical Suidæ, and seems to approach the African Phacochœridæ, The manner in which the canines of the upperjaware reversed, and grow directly upwards in a spiral curve over the eyes, is unique among mammalia. (8.) Five squirrels inhabit Celebes, and all are peculiar species. (9.) These are forest rats of the sub-genusGymnomys, allied to Australian species. 10.Cuscus.This typicalAustralian form is represented in Celebes by two peculiar species.
Leaving out the Indo-Malayspecies, which may probably have been introduced by man, and are at all events comparatively recent immigrants, and the wild pig, a genus which ranges over the whole archipelago and which has therefore little significance, we find two genera which have come from the Australian side,—CuscusandMus; and four from the Oriental side,—Cynopithecus,Anoa,Babirusa, andSciurus. ButSciurusalone corresponds toCuscus, as a genus still inhabiting the adjacent islands; the other three being not only peculiar to Celebes, but incapable of being affiliated to any specially Oriental group. We seem, then, to have indications of two distinct periods; one very ancient, when the ancestors of the three peculiar genera roamed over some unknown continent of which Celebes formed, perhaps, an outlying portion;—another more recent, when from one side there enteredSciurus, and from the otherCuscus. But we must remember that the Moluccas to the east, possess scarcely any indigenous mammals exceptCuscus; whereas Borneo and Java on the west, have nearly 50 distinct genera. It is evident then, that the facilities for immigration must have been much less with the Oriental than with the Australian region, and we may be pretty certain that at this later period there was no land connection with the Indo-Malay islands, or some other animals than squirrels would certainly have entered. Let us now see what light is thrown upon the subject by the birds.
Birds.—The total number of birds known to inhabit Celebes is 205, belonging to about 150 genera. We may leave out of consideration the wading and aquatic birds, most of which are wide-ranging species. There remain 123 genera and 152 species of land-birds, of which 9 genera and 66 species are absolutely confined to the island, while 20 more are found also in the Sula or Sanguir Islands, so that we may take 86 to be the number of peculiar Celebes species. Lord Walden, from whose excellent paper on the birds of Celebes (Trans. Zool. Soc.vol. viii. p. 23) most of these figures are obtained, estimates, that of the species which are not peculiar to Celebes, 55 are of Oriental and 22 ofAustralian origin, the remainder being common to both regions. This shows a preponderant recent immigration from the West and North, which is not to be wondered at when we look at the long coast line of Java, Borneo, and the Philippine islands, with an abundant and varied bird population, on the one side, and the small scattered islands of the Moluccas, with a comparatively scanty bird-fauna, on the other.
But, adopting the method here usually followed, let us look at the relations of thegenerafound in Celebes, omitting for the present those which are peculiar to it. I divide these genera into two series:—those which are found in Borneo or Java but not in the Moluccas, and those which inhabit the Moluccas and not Borneo or Java; these being the respective sources from which,primâ facie, the species of these genera must have been derived. Genera which range widely into both these districts are rejected, as teaching us nothing of the origin of the Celebesian fauna. In a few cases, sub-genera which show a decided eastern or western origin, are given.
These tables show a decided preponderance of Oriental over Australian forms. But we must remember that the immediately adjacent lands from whence the supply was derived, isvery much richer in the one case than in the other. The 24 genera derived from Borneo and Java are only aboutone fourthof the characteristic genera of those islands; while the 15 Moluccan and Timorese genera are fullyone thirdof their characteristic types. Theproportionderived from the Australian, is greater than that derived from the Oriental side.
We shall exhibit this perhaps more clearly, by giving a list of the important groups of each set of islands which are absent from Celebes.
If we reckon the absent families to be each represented by only two important genera, we shall find the deficiency on the Oriental side much the greatest; yet those on the side of the Moluccas are sufficiently remarkable. The Meliphagidæ are not indeed absolutely wanting, since aMyzomelahas now been found in Celebes; but all its larger and more powerful forms which range over almost the entire region, are absent. This may be balanced by the absence of the excessively abundant Timaliidæ of the Indo-Malay islands, which are represented byonly a single species; and by the powerful Phasianidæ, represented only by the common Malay jungle fowl, perhaps introduced. The entire absence of Pycnonotidæ is a very anomalous fact, since one of the largest genera,Criniger, is well represented in several islands of the Moluccas, and one has even been found in the Togian islands in the great northern inlet of Celebes; but yet it passes over Celebes itself.Ceyx, a genus of small kingfishers, is a parallel case, since it is found everywhere from India to New Guinea, leaving out only Celebes; but this comes among those curiosities of the Celebesian fauna which we shall notice further on. In the list of genera derived from Borneo or Java, no less than 6 are represented by identical species (indicated by sp. after the name); while in the Moluccan list 5 are thus identical. These must be taken to indicate, either that the genus is a recent introduction, or that stragglers still occasionally enter, crossing the breed, and thus preventing specific modification. In either case they depend on the existing state of things, and throw no light on the different distribution of land and sea which aided or checked migration in former times; and they therefore to some extent diminish the weight of the Indo-Malay affinity, as measured by the relations of the peculiar species of Celebes.
From our examination of the evidence thus far,—that is, taking account firstly, of thespecies, and, secondly, of thegenera, which are common to Celebes and the groups of islands between which it is situated, we must admit that the connexion seems rather with the Oriental than with the Australian region; but when we take into account theproportionof the genera and species present, to those which are absent, and giving some weight to the greater extent of coast line on the Indo-Malay side, we seem justified in stating that the Austro-Malay element is rather the most fully represented. This result applies both to birds and mammals; and it leads us to the belief, that during the epoch of existing species and genera, Celebes has never been united with any extensive tract of land either on the Indo-Malay or Austro-Malay side, but has received immigrants from both during a very long period, the facilities for immigration having been rather thegreatest on the Austro-Malay or Australian side. We have now to consider what further light can be thrown on the subject by the consideration of thepeculiar generaof Celebes, and of those curiosities or anomalies of distribution to which we have referred.
Nine genera of birds are altogether peculiar to Celebes; three more are found only in one other island, and seem to be typically Celebesian; while one is found in the Sula islands (which belongs to the Celebes group) and probably exists in Celebes also. The following is a list of these 13 genera:
Of the above,Artamides,Monachalcyon,Cittura, andMegacephalon, are modifications of types characteristic of the Australian region. All are peculiar to Celebes exceptCittura, found also in the Sanguir islands to the northward, but which seems to belong to the Moluccan group.Streptocitta,Charitornis, andGazzola, are peculiar types of Corvidæ; the two former allied to the magpies, the latter to the jackdaws.Charitornisis known only from the Sula islands east of Celebes, and is closely related toStreptocitta. There is nothing comparable to these three groups in any of the Malay islands, and they seem to have relations rather with the Corvidæ of the old-world northern continent.Basilornis,Enodes, andScissirostrum, are remarkable forms of Sturnidæ.Basilornishas a beautiful compressed crest, which in the allied species found in Ceram is elongated behind.Enodeshas remarkable red superciliary streaks, but seems allied toCalornis.Scissirostrumseems also allied toCalornisin general structure, but has a very peculiarly formed bill and nostrils. We can hardly say whether these three forms show more affinity to Oriental or to Australian types, but they add to the weight of evidence as to the great antiquity and isolation of the Celebesian fauna.Scissirostrumhas been classed withEuryceros, aMadagascar bird, and withBuphaga, an African genus; but the peculiar beak and nostrils approximate more toCracticusand its allies, of the Australian region, which should probably form a distinct family.Ceycopsisis undoubtedly intermediate between the MalayanCeyxand the AfricanIspidina, and is therefore especially interesting.Meropogonis a remarkable form of bee-eater, allied to the Indo-MalayanNyctiornis.Prioniturus(the raquet-tailed parrots) of which two species inhabit Celebes, and one the Philippines, appears to be allied to the Austro-MalayanGeoffroyus.
We must finally notice a few genera found in Celebes, whose nearest allies are not in the surrounding islands, and which thus afford illustrations of discontinuous distribution. The most remarkable, perhaps, isCoracias, of which a fine species inhabits Celebes; while the genus is quite unknown in the Indo-Malay sub-region, and does not appear again till we reach Burmah and India; and the species has no closer affinity for Indian than for African forms.Myialestes, a small yellow flycatcher, is another exmple; its nearest ally (M. cinereocapilla) being a common Indian bird, but unknown in the Malay islands. The Celebesian bird described by me asPrionochilus aureolimbatus, is probably a third case of discontinuous distribution, if (as a more careful examination seems to show) it is not aPrionochilus, but congeneric withPachyglossa, a bird only found in the Himalayas. The fine pigeon,Carpophaga forsteni, belongs to a group found in the Philippines, Australia, and New Zealand; but the Celebes species is very distinct from all the others, and seems, if anything, more allied to that of New Zealand.
The Sula islands (Sula-mangola, Sula-taliabo, and Sula-besi) lie midway between Celebes and the Moluccas, being 80 miles from the nearest part of Celebes, with several intervening islands, and 40 miles from Bouru, all open sea. Their birds show, as might be expected, a blending of the two faunas, but with a decided preponderance of that of Celebes. Out of 43 land birds which have been collected in these islands, we may deduct 6 as of wide range and no significance. Of the 37 remaining, 21 are Celebesian species, and 4 are new species butallied to those of Celebes; while there are 10 Moluccan species and 2 new species allied to those of the Moluccas. It is curious that no less than 3 Moluccan genera, quite unknown in Celebes itself, occur here,—Monarcha,Pachycephala, andCriniger; but all these, as well as several other of the Moluccan birds, are rather weak flyers, and such as are likely to have been carried across by strong winds. Of thegenera, 23 are from Celebes, 10 from the Moluccas. These facts show, that the Sula islands form part of the Celebes group, although they have received an infusion of Moluccan forms, which will perhaps in time spread to the main island, and diminish the remarkable individuality that now characterises its fauna.
Insects.—Of the reptiles and fishes of Celebes we have not sufficient information to draw any satisfactory conclusions. I therefore pass to the insects of which something more is known.
The Butterflies of Celebes are not very numerous, less than 200 species in all having been collected; but a very large proportion of them, probably three-fourths of the whole, are peculiar. There is only one peculiar genus,Amechania, allied toZethera(a group confined to the Philippine Islands), with which it should perhaps be united. Most of the genera are of wide distribution in the archipelago, or are especially Malayan, only two truly Australian genera,ElodinaandAcropthalmia, reaching Celebes. On the other hand, 7 peculiar Oriental genera are found in Celebes, but not further east, viz.,Clerome,Adolias,Euripus,Apatura,Limenitis,Iolaus, andLeptocircus. There are also several indications of a direct affinity with the continent rather than with Malaya, as in the cases already enumerated among birds. A fine butterfly, yet unnamed, almost exactly resemblesDichorragia nesimachus, a Himalayan species.Euripus robustusis closely allied toE. halithersesof N. India; there are no less than 5 species ofLimenitis, all quite unlike those found in other parts of the archipelago. The butterflies of Celebes are remarkably distinguished from all others in the East, by peculiarities of form, size, and colour, which run through groups of species belonging to different genera. Many Papilionidæ and Pieridæ, and someNymphalidæ, have the anterior wings elongated, with the apex often acute, and, what is especially remarkable, an abrupt bend or shoulder near the base of the wing. (SeeMalay Archipelago, 3rd Ed. p. 281, woodcut.) No less than 13 species ofPapilio, 10 Pieridæ, and 4 or 5 Nymphalidæ, are thus distinguished from their nearest allies in the surrounding islands or in India. In size again, a large number of Celebesian butterflies stand preeminent over their allies. The fine Papilios—adamantius,blumei, andgigon—are perfect giants by the side of the closely-allied forms of Java; whileP. androclesis the largest and longest-tailed, of all the true swallow-tailed group of the Old World. Among Nymphalidæ, the species ofRhinopalpaandEuripus, peculiar to Celebes, are immensely larger than their nearest allies; and several of the Pieridæ are also decidedly larger, though in a less marked degree. In colour, many of the Celebesian butterflies differ from the nearest allied species; so that they acquire a singularity of aspect which marks them off from the rest of the group. The most curious case is that of three butterflies, belonging to three distinct genera (Cethosia myrina,Messaras mæonides, andAtella celebensis) all having a delicate violet or lilac gloss in lines or patches, which is wholly wanting in every allied species of the surrounding islands. These numerous peculiarities of Celebesian butterflies are very extraordinary; and imply isolation from surrounding lands, almost as much as do the strange forms of mammals and birds, which more prominently characterise this interesting island.
Of the Coleoptera we know much less, but a few interesting facts may be noted. There are a number of fine species ofCicindela, some of peculiar forms; and oneOdontochila, a South American genus; whileCollyrisreaches Celebes from the Oriental region. In Carabidæ it has one peculiar genus,Dicraspeda; and a species of the fine Australian genusCatadromus. In Lucanidæ it has the Oriental genus,Odontolabris. In Cetoniidæ it has a peculiar genus,Sternoplus, and several fineCetoniæ; but the characteristic Malayan genus,Lomaptera, found in every other island of the archipelago from Sumatra to New Guinea, is absent—an analogous fact to the case ofCeyxamong birds.In Buprestidæ, the principal Austro-Malay genus,Sambus, is found here; whileSponsor, a genus 8 species of which inhabit Mauritius, has one species here and one in New Guinea. In Longicorns there are four peculiar genera,Comusia,Pytholia,Bityle, andOmbrosaga; but the most important features are the occurrence of the otherwise purely Indo-Malayan generaAgelasta,Nyctimene, andAstathes; and of the purely Austro-MalayanArrhenotus,Trysimia,Xenolea,Amblymora,Diallus, andÆgocidnus. The remaining genera range over both portions of the archipelago. In the extensive family of Curculionidæ we can only notice the elegant genus,Celebia, allied toEupholus, which, owing to its abundance and beauty, is a conspicuous feature in the entomology of the island.
Origin of the fauna of Celebes.—We have now to consider, briefly, what past changes of physical geography are indicated by the curious assemblage of facts here adduced. We have evidently, in Celebes, a remnant of an exceedingly ancient land, which has undergone many and varied revolutions; and the stock of ancient forms which it contains must be taken account of, when we speculate on the causes that have so curiously limited more recent immigrations. Going back to the arrival of those genera which are represented in Celebes by peculiar species, and taking first the Austro-Malay genera, we find among them such groups asZonœnas(s.g.),Phlogœnas,Leucotreron(s.g.), andTuracœna, which are not found in the Moluccas at all; andMyzomela, found in Timor and Banda, but not in Ceram or Bouru, which are nearest to Celebes. This, combined with the curious absence of so many of the commonest Moluccan genera, leads to the conclusion that the Austro-Malay immigration took place by way of Timor and the southern part of New Guinea. It will be remembered, that to account for the Indo-Malayan forms in New Guinea, we suggested an extension of that country in a westerly direction just north of Timor. Now this is exactly what we require, to account for the stocking of Celebes with the Australian forms it possesses. At this time Borneo did not approach so near, and it was at a somewhat later period that the last great Indo-Malay migration set in; butfinding the country already fairly stocked, comparatively few groups were able to establish themselves.
Going back a little farther, we come to the entrance of those few birds and insects which belong to India or Indo-China; and this probably occurred at the same time as that continental extension southward, which we found was required to account for a similar phenomenon in Java. Celebes, being more remote, received only a few stragglers. We have now to go much farther back, to the time when the ancestors of the peculiar Celebesian genera entered the country, and here our conjectures must necessarily be less defined.
On the Australian side we have to account forMegacephalon, and the other genera of purely Papuan type. It may perhaps be sufficient to say, that we do not yet know that these genera, or some very close allies, do not still exist in New Guinea; in which case they may well have entered at the same time with the species, already referred to. If, on the other hand, they are really as isolated as they appear to be, they represent an earlier communication, either by an approximation of the two islands over the space now occupied by the Moluccas; or, what is perhaps more probable, through a former extension of the Moluccas, which have since undergone so much subsidence, as to lead to the extinction of a large proportion of their ancient fauna. The wide-spread volcanic action, and especially the prevalence of raised coral-reefs in almost all the islands, render this last supposition very probable.
On the Oriental side the difficulty is greater; for here we find, what seem to be clear indications of a connection with Africa, as well as with Continental Asia, at some immensely remote epoch.Cynopithecus,Babirusa, andAnoa;Ceycopsis,Streptocitta, andGazzola(s. g.), and perhapsScissirostrum, may be well explained as descendants of ancestral types in their respective groups, which also gave rise to the special forms of Africa on the one hand, and of Asia on the other. For this immigration we must suppose, that at a period before the formation of the present Indo-Malay Islands, a great tract of land extended in a north-westerly direction, till it met the old Asiatic continent. This may have been beforethe Himalayas had risen to any great height, and when a large part of what are now the cold plateaus of Central Asia may have teemed with life, some forms of which are preserved in Africa, some in Malaya, and a few in Celebes. Here may have lived the common ancestor ofSus,Babirusa, andPhacochœrus; as well as ofCynopithecus,Cynocephalus, andMacacus; ofAnoaandBubalus; ofScissirostrumandEuryceros; ofCeyx,Ceycopsis, andIspidina. Such an origin accounts, too, for the presence of the North-Indian forms in Celebes; and it offers less difficulties than a direct connection with continental Africa, which once appeared to be the only solution of the problem. If this south-eastward extension of Asia occurred at the same time as the north-eastward extension of South Africa and Madagascar, the two early continents may have approached each other sufficiently to have allowed of some interchange of forms:Tarsiusmay be the descendant of some Lemurine animal that then entered the Malayan area, while the progenitors ofCryptoproctamay then have passed from Asia to Madagascar.
It is true that we here reach the extremest limits of speculation; but when we have before us such singular phenomena as are presented by the fauna of the island of Celebes, we can hardly help endeavouring to picture to our imaginations by what past changes of land and sea (in themselves not improbable) the actual condition of things may have been brought about.
II. Australia and Tasmania, or the Australian Sub-region.
A general sketch of Australian zoology having been given in the earlier part of this chapter, it will not be necessary to occupy much time on this sub-region, which is as remarkably homogeneous as the one we have just left is heterogeneous. Although much of the northern part of Australia is within the tropics, while Victoria and Tasmania are situated from 36° to 43° south latitude, there is no striking change in the character of the fauna throughout the continent; a number of important genera extending over the whole country, and giving a very uniform character to its zoology. The eastern parts, including the colonies of New South Wales and Queensland, are undoubtedly the richest, several peculiar types being found only here. The southern portion is somewhat poorer, and has very few peculiar forms; and Tasmania being isolated is poorer still, yet its zoology has much resemblance to that of Victoria, from which country it has evidently not been very long separated. The north, as far as yet known, is characterised by hardly any peculiar forms, but by the occurrence of a number of Papuan types, which have evidently been derived from New Guinea.
Plate XI.