CAPITONIDae (Barbets).
93. Cyanops pulcherrimus.
94. ,, monticulus.
95. *Megalaema chrysopsis.
BUBONIDae (Owls).
96. Heteroscops luciae.
97. *Syrnium leptogrammic.u.m.
FALCONIDae (Hawks, &c.).
98. Spilornis pallidus.
99. *Accipiter nigrotibialis.
100. Microhierax latifrons.
PHASIANIDae (Pheasants).
101. Polyplectron schliermacheri.
102. Lobiophasis bulweri.
103. *Argusia.n.u.s grayi.
104. *Euplocamus pyrronotus.
{380}
TETRAONIDae (Grouse, &c.).
105. Bambusicola hyperythra.
106. ,, erythrophrys.
107. Haematortyx sanguiniceps.
RALLIDae (Rails).
108. Rallina rufigenys.
Representative forms of the same character as those noted above are found in all extensive continental areas, but they are rarely so numerous. Thus, in Mr. Elwes" paper on the "Distribution of Asiatic Birds," he states that 12.5 per cent. of the land birds of Burmah and Tena.s.serim are peculiar species, whereas we find that in Borneo they are about 25 per cent., and the difference may fairly be imputed to the greater proportion of slightly modified representative species due to a period of complete isolation. Of peculiar genera, the Indo-Chinese Peninsula has one--Ampeliceps, a remarkable yellow-crowned starling, with bare pink-coloured orbits; while two others, Temnurus and Crypsirhina--singular birds allied to the jays--are found in no other part of the Asiatic continent though they occur in some of the Malay Islands. Borneo has seven peculiar genera of pa.s.seres,[89] as well as Haematortyx, a crested partridge; and Lobiophasis, a pheasant hardly distinct from Euplocamus; while two others, Pityriasis, an extraordinary bare-headed bird between a jay and a shrike, and Carpococcyx, a pheasant-like ground cuckoo formerly thought to be peculiar, are said to have been discovered also in Sumatra.
The insects and land-sh.e.l.ls of Borneo and of the surrounding countries are too imperfectly known to enable us to arrive at any accurate results with regard to their distribution. They agree, however, with the birds and mammals in their general approximation to Malayan forms, but the number of peculiar species is perhaps larger.
The proportion here shown of less than one-fourth peculiar species of mammalia and fully one-fourth peculiar species of land-birds, teaches us that the possession of the power of flight affects but little the distribution of {381} land-animals, and gives us confidence in the results we may arrive at in those cases where we have, from whatever cause, to depend on a knowledge of the birds alone. And if we consider the wide range of certain groups of powerful flight--as the birds of prey, the swallows and swifts, the king-crows, and some others, we shall be forced to conclude that the majority of forest-birds are restricted by even narrow watery barriers, to an even greater extent than mammalia.
_The Affinities of the Bornean Fauna._--The animals of Borneo exhibit an almost perfect ident.i.ty in general character, and a close similarity in species, with those of Sumatra and the Malay Peninsula. So great is this resemblance that it is a question whether it might not be quite as great were the whole united; for the extreme points of Borneo and Sumatra are 1,500 miles apart--as far as from Madrid to Constantinople, or from the Missouri valley to California. In such an extent of country we always meet with some local species, and representative forms, so that we hardly require any great lapse of time as an element in the production of the peculiarities we actually find. So far as the forms of life are concerned, Borneo, as an island, may be no older than Great Britain; for the time that has elapsed since the glacial epoch would be amply sufficient to produce such a redistribution of the species, consequent on their mutual relations being disturbed, as would bring the islands into their present zoological condition. There are, however, other facts to be considered, which seem to imply much greater and more complex revolutions than the recent separation of Borneo from Sumatra and the Malay Peninsula, and that these changes must have been spread over a considerable lapse of time. In order to understand what these changes probably were, we must give a brief sketch of the fauna of Java, the peculiarities of which introduce a new element into the question we have to discuss. {382}
JAVA.
The rich and beautiful island of Java, interesting alike to the politician, the geographer, and the naturalist, is more especially attractive to the student of geographical distribution, because it furnishes him with some of the most curious anomalies and difficult problems in a place where such would be least expected. As Java forms with Sumatra one almost unbroken line of volcanoes and volcanic mountains, interrupted only by the narrow Straits of Sunda, we should naturally expect a close resemblance between the productions of the two islands. But in point of fact there is a much greater difference between them than between Sumatra and Borneo, so much further apart, and so very unlike in physical features.[90] Java differs from the three great land ma.s.ses--Borneo, Sumatra, and the Malay Peninsula, far more than either of these do from each other; and this is the first anomaly we encounter. But a more serious difficulty than this remains to be stated. Java has certain close resemblances to the Siamese Peninsula, and also to the Himalayas, which Borneo and Sumatra do not exhibit to so great a proportionate extent; and looking at the relative position of these lands respectively, this seems most incomprehensible. In order fully to appreciate the singularity and difficulty of the problem, it will be necessary to point out the exact nature and amount of these peculiarities in the fauna of Java.
_General Character of the Fauna of Java._--If we were only to take account of the number of peculiar species in Java, and the relations of its fauna generally to that of the surrounding lands, we might pa.s.s it over as a less interesting island than Borneo or Sumatra. Its mammalia (ninety species) are nearly as numerous as those of Borneo, but are apparently less peculiar, none of the genera and only five or six of the species being confined to the island. In land-birds it is decidedly less rich, having only 300 species, of which about forty-five are peculiar, and only one {383} or two belong to peculiar genera; so that here again the amount of speciality is considerably less than in Borneo. It is only when we proceed to a.n.a.lyse the species of the Javan fauna, and trace their distribution and affinities, that we discover its interesting nature.
_Difference Between the Fauna of Java and that of the other great Malay Islands._--Comparing the fauna of Java with that which may be called the typical Malayan fauna as exhibited in Borneo, Sumatra, and the Malay Peninsula, we find the following differences. No less than thirteen genera of mammalia, each of which is known to inhabit at least two, and generally all three, of the above-named Malayan countries, are totally absent from Java; and they include such important forms as the elephant, the tapir, and the Malay bear. It cannot be said that this difference depends on imperfect knowledge, for Java is one of the oldest European settlements in the East, and has been explored by a long succession of Dutch and English naturalists. Every part of it is thoroughly well known, and it would be almost as difficult to find a new mammal of any size in Europe as in Java.
Of birds there are twenty-five genera, all typically Malayan and occurring at least in two, and for the most part in all three of the Malay countries, which are yet absent from Java. Most of these are large and conspicuous forms, such as jays, gapers, bee-eaters, woodp.e.c.k.e.rs, hornbills, cuckoos, parrots, pheasants, and partridges, as impossible to have remained undiscovered in Java as the large mammalia above referred to.
Besides these absent _genera_ there are some curious ill.u.s.trations of Javan isolation in the _species_; there being several cases in which the same species occurs in all three of the typical Malay countries, while in Java it is represented by an allied species. These occur chiefly among birds, there being no less than seven species which are common to the three great Malay countries but are represented in Java by distinct though closely allied species.
From these facts it is impossible to doubt that Java has had a history of its own, quite distinct from that of the other portions of the Malayan area. {384}
_Special Relations of the Javan Fauna to that of the Asiatic Continent._--These relations are indicated by comparatively few examples, but they are very clear and of great importance. Among mammalia, the genus Helictis is found in Java but in no other Malay country, though it inhabits also North India; while two species, _Rhinoceros javanicus_ and _Lepus kurgosa_, are natives of Indo-Chinese countries and Java, but not of typical Malaya. In birds there are five genera or sub-genera--Zoothera, Notodela, Crypsirhina, Allotrius, and Cochoa, which inhabit Java, the Himalayas, and Indo-China, all but the last extending south to Tena.s.serim, but none of them occurring in Malacca, Sumatra, or Borneo. There are also two species of birds--a trogon (_Harpactes oreskios_), and the Javanese peac.o.c.k (_Pavo muticus_), which inhabit only Java and the Indo-Chinese countries, the former reaching Tena.s.serim and the latter Perak in the Malay Peninsula.
Here, then, we find a series of remarkable similarities between Java and the Asiatic continent, quite independent of the typical Malay countries--Borneo, Sumatra, and the Malay Peninsula, which latter have evidently formed one connected land, and thus appear to preclude any independent union of Java and Siam.
The great difficulty in explaining these facts is, that all the required changes of sea and land must have occurred within the period of existing species of mammalia. Sumatra, Borneo, and Malacca have, as we have seen, a great similarity as regards their species of mammals and birds, while Java, though it differs from them in so curious a manner, has no greater degree of speciality, since its species, when not Malayan, are almost all North Indian or Siamese.
There is, however, one consideration which may help us over this difficulty. It seems highly probable that in the equatorial regions species have changed less rapidly than in the north temperate zone, on account of the equality and stability of the equatorial climate. We have seen, in Chapter X., how important an agent in producing extinction and modification of species must have been the repeated changes from cold to warm, and from warm to cold {385} conditions, with the migrations and crowding together that must have been their necessary consequence. But in the lowlands, near the equator, these changes would be very little if at all felt, and thus one great cause of specific modification would be wanting. Let us now see whether we can sketch out a series of not improbable changes which may have brought about the existing relations of Java and Borneo to the continent.
_Past Geographical Changes of Java and Borneo._--Although Java and Sumatra are mainly volcanic, they are by no means wholly so. Sumatra possesses in its great mountain ma.s.ses ancient crystalline rocks with much granite, while there are extensive Tertiary deposits of Eocene age, overlying which are numerous beds of coal now raised up many thousand feet above the sea.[91] The volcanoes appear to have burst through these older mountains, and to have partly covered them as well as great areas of the lowlands with the products of their eruptions. In Java either the fundamental strata were less extensive and less raised above the sea, or the period of volcanic action has been of longer duration; for here no crystalline rocks have been found except a few boulders of granite in the western part of the island, perhaps the relics of a formation destroyed by denudation or covered up by volcanic deposits. In the southern part of Java, however, there is an extensive range of low mountains, about 3,000 feet high, consisting of basalt with limestone, apparently of Miocene age.
During this last named period, then, Java would have been at least 3,000 feet lower than it is now, and such a depression would probably extend to considerable parts of Sumatra and Borneo, so as to reduce them all to a few small islands. At some later period a gradual elevation occurred, which ultimately united the whole of the islands with the continent. This may have continued till the glacial period of the northern hemisphere, during the severest part of which a few Himalayan species of birds and mammals may have been driven southward, and {386} have ranged over suitable portions of the whole area. Java then became separated by subsidence, and these species were imprisoned in the island; while those in the remaining part of the Malayan area again migrated northward when the cold had pa.s.sed away from their former home, the equatorial forests of Borneo, Sumatra, and the Malay Peninsula being more especially adapted to the typical Malayan fauna which is there developed in rich profusion. A little later the subsidence may have extended farther north, isolating Borneo and Sumatra, in which a few other Indian or Indo-Chinese forms have been retained, but probably leaving the Malay Peninsula as a ridge between them as far as the islands of Banca and Biliton. Other slight changes of climate followed, when a further subsidence separated these last-named islands from the Malay Peninsula, and left them with two or three species which have since become slightly modified. We may thus explain how it is that a species is sometimes common to Sumatra and Borneo, while the intervening island (Banca) possesses a distinct form.[92]
In my _Geographical Distribution of Animals_, Vol. I., p. 357, I have given a somewhat different hypothetical explanation of the relations of Java and Borneo to the continent, in which I took account of changes of land and sea only; but a fuller consideration of the influence of changes of climate on the migration of animals, has led me to the much simpler, and, I think, more probable, explanation above given. The amount of the relationship between Java and Siam, as well as of that between Java and the Himalayas, is too small to be well accounted for by an independent geographical connection in which Borneo and Sumatra did not take part. It is, at the same time, too distinct and indisputable to be ignored; and a change of climate which should drive a portion of the Himalayan fauna southward, leaving a few species in Java and Borneo from which they could not return owing to the subsequent isolation of those islands by subsidence, seems {387} to be a cause exactly adapted to produce the kind and amount of affinity between these distant countries that actually exists.
THE PHILIPPINE ISLANDS.
A general account of the fauna of these islands, and of their biological relations to the countries which form the subject of this chapter, has been given in my _Geographical Distribution of Animals_, Vol. I. pp. 345-349; but since the publication of that work considerable additions have been made to their fauna, having the effect of somewhat diminishing their isolation from the other islands. Four genera have been added to the terrestrial mammalia--Crocidura, Felis, Pteromys, and Mus, as well as two additional squirrels; while the black ape (_Cynopithecus niger_) has been struck out as not inhabiting the Philippines. This brings the true land mammalia to twenty-one species, of which fourteen are peculiar to the islands; but to these we must add no less than thirty-three species of bats of which only ten are peculiar.[93] In these estimates the Palawan {388} group has been omitted as these islands contain so many Bornean species that if included they obscure the special features of the fauna.
_Birds._--The late Marquis of Tweeddale made a special study of Philippine birds, and in 1873 published a catalogue in the _Transactions of the Zoological Society_ (Vol. IX. Pt. 2, pp. 125-247). But since that date large collections have been made by Everett, Steere, and other travellers, the result of which has been to more than double the known species, and to render the ornithological fauna an exceedingly rich one. Many of the Malayan genera which were thought to be absent when the first edition of this work was published have since been discovered, among which are Phyllornis, Criniger, Diceum, Prionochilus, and Batrachostomus. But there still remain a large number of highly characteristic Malayan genera whose absence gives a distinctive feature to the Philippine bird fauna. Among these are Tiga and Meiglyptes, genera of woodp.e.c.k.e.rs; Phaenicophaes and Centropus, remarkable cuckoos; the long-tailed paroquets, Palaeornis; all the genera of Barbets except Xantholaema; the small but beautiful family Eurylaemidae; many genera allied to Timalia and Ixos; the mynahs, Gracula; the long-tailed flycatchers, Tchitrea; the fire-backed pheasants, Euplocamus; the argus pheasants, the jungle-fowl, and many others.
The following tabular statement will ill.u.s.trate the rapid growth of our knowledge of the birds of the Philippines:--
Land-birds.Water-birds.Total.
+-----------+------------+------ Lord Tweeddale"s Catalogue (1873)15860218 Mr. Wardlaw Ramsay"s List (1881)26575340 Mr. Everett"s MSS. List of Additions (1891)370102472
The number of peculiar species is very large, there being about 300 land and forty-two water birds, which are not {389} known to occur beyond the group. We have here, still more p.r.o.nounced than in the case of Borneo, the remarkable fact of the true land birds presenting a larger amount of speciality than the land mammals; for while more than four-fifths of the birds are peculiar, only a little more than half the mammals are so, and if we exclude the bats only two-thirds.
The general character of the fauna of this group of islands is evidently the result of their physical conditions and geological history. The Philippines are almost surrounded by deep sea, but are connected with Borneo by means of two narrow submarine banks, on the northern of which is situated Palawan, and on the southern the Sulu Islands. Two small groups of islands, the Bashees and Babuyanes, have also afforded a partial connection with the continent by way of Formosa. It is evident that the Philippines once formed part of the great Malayan extension of Asia, but that they were separated considerably earlier than Java; and having been since greatly isolated and much broken up by volcanic disturbances, their species have for the most part become modified into distinct local forms, representative species often occurring in the different islands of the group. They have also received a few Chinese types by the route already indicated, and a few Australian forms owing to their proximity to the Moluccas. Their comparative poverty in genera and species of the mammalia is perhaps due to the fact that they have been subjected to a great amount of submersion in recent times, greatly reducing their area and causing the extinction of a considerable portion of their fauna. This is not a mere hypothesis, but is supported by direct evidence; for I am informed by Mr. Everett, who has made extensive explorations in the islands, that almost everywhere are found large tracts of elevated coral-reefs, containing sh.e.l.ls similar to those living in the adjacent seas, an indisputable proof of recent elevation.
_Concluding Remarks on the Malay Islands._--This completes our sketch of the great Malay islands, the seat of the typical Malayan fauna. It has been shown that the peculiarities presented by the individual islands may be all {390} sufficiently well explained by a very simple and comparatively unimportant series of geographical changes, combined with a limited amount of change of climate towards the northern tropic. Beginning in late Miocene times when the deposits on the south coast of Java were upraised, we suppose a general elevation of the whole of the extremely shallow seas uniting what are now Sumatra, Java, Borneo, and the Philippines with the Asiatic continent, and forming that extended equatorial area in which the typical Malayan fauna was developed. After a long period of stability, giving ample time for the specialisation of so many peculiar types, the Philippines were first separated; then at a considerably later period Java; a little later Sumatra and Borneo; and finally the islands south of Singapore to Banca and Biliton. This one simple series of elevations and subsidences, combined with the changes of climate already referred to, and such local elevations and depressions as must undoubtedly have occurred, appears sufficient to have brought about the curious, and at first sight puzzling, relations, of the faunas of Java and the Philippines, as compared with those of the larger islands.
We will now pa.s.s on to the consideration of two other groups which offer features of special interest, and which will complete our ill.u.s.trative survey of recent continental islands.
{391}
CHAPTER XVIII
j.a.pAN AND FORMOSA
j.a.pan, its Position and Physical Features--Zoological Features of j.a.pan--Mammalia--Birds--Birds Common to Great Britain and j.a.pan--Birds Peculiar to j.a.pan--j.a.pan Birds Recurring in Distant Areas--Formosa--Physical Features of Formosa--Animal Life of Formosa--Mammalia--Land-birds Peculiar to Formosa--Formosan Birds Recurring in India or Malaya--Comparison of Faunas of Hainan, Formosa, and j.a.pan--General Remarks on Recent Continental Islands.