Island Life/XVIII

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CHAPTER XVIII

JAPAN AND FORMOSA

Japan, its Position and Physical Features—Zoological Features of Japan—Mammalia—Birds—Birds Common to Great Britain and Japan—Birds Peculiar to Japan—Japan 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 Japan—General Remarks on Recent Continental Islands.

Japan.

The Japanese Islands occupy a very similar position on the eastern shore of the great Euro-Asiatic continent to that of the British Islands on the western, except that they are about sixteen degrees further south, and having a greater extension in latitude enjoy a more varied as well as a more temperate climate. Their outline is also much more irregular and their mountains loftier, the volcanic peak of Fusiyama being 14,177 feet high; while their geological structure is very complex, their soil extremely fertile, and their vegetation in the highest degree varied and beautiful. Like our own islands, too, they are connected with the continent by a marine bank less than a hundred fathoms below the surface—at all events towards the north and south; but in the intervening space the Sea of Japan opens out to a width of six hundred miles, and in its central portion is very deep, and this may be an indication that the connection between the islands and the continent is of rather ancient date. At the Straits of Corea the distance from the main land is about 120 miles, while at the northern extremity of Yesso it is about 200. The island of Saghalien, however, separated from Yesso by a strait only twenty-five miles wide, forms a connection with Amoorland in about 52° N. Lat. A southern warm current flowing a little to the eastward of the islands, ameliorates their climate much in the same way as the Gulf Stream does ours, and added to their insular position enables them to support a more tropical vegetation and more varied forms of life than are found at corresponding latitudes in China.

MAP OF JAPAN AND FORMOSA (with depths in fathoms).
Light tint, sea under 100 fathoms. Medium tint, under 1,000 fathoms. Dark tint, over 1,000 fathoms. The figures show the depth in fathoms.

Zoological Features of Japan.—As we might expect from the conditions here sketched out, Japan exhibits in all its forms of animal life a close general resemblance to the adjacent continent, but with a considerable element of specific individuality; while it also possesses some remarkable isolated groups. Its fauna presents indications of there having been two or more lines of migration at different epochs. The majority of its animals are related to those of the temperate or cold regions of the continent, either as identical or allied species; but a smaller number have a tropical character, and these have in several instances no allies in China but occur again only in Northern India or the Malay Archipelago. There is also a slight American element in the fauna of Japan, a relic probably of the period when a land communication existed between the two continents over what are now the shallow seas of Japan, Ochotsk, and Kamschatka. We will now proceed to examine the peculiarities and relations of the fauna.

Mammalia.—The mammalia of Japan at present known are forty in number; not very many when compared with the rich fauna of China and Manchuria, but containing monkeys, bears, deer, wild goats and wild boars, as well as foxes, badgers, moles, squirrels, and hares, so that there can be no doubt whatever that they imply a land connection with the continent. No complete account of Japan mammals has been given by any competent zoologist since the publication of Von Siebold's Fauna Japonica in 1844, but by collecting together most of the scattered observations since that period the following list has been drawn up, and will, it is hoped, be of use to naturalists. The species believed to be peculiar to Japan are printed in italics. These are very numerous, but it must be remembered that Corea and Manchuria (the portions of the continent opposite Japan) are comparatively little known, while in very few cases have the species of Japan and of the continent been critically compared. Where this has been done, however, the peculiar species established by the older naturalists have been in many cases found to be correct.

List of the Mammalia of the Japanese Islands.

1. Macacus speciosus. A monkey with rudimentary tail and red face, allied to the Barbary ape. It inhabits the island of Niphon up to 41° N. Lat., and has thus the most northern range of any living monkey.
2. Pteropus dasymallus. A peculiar fruit-bat, found in Kiusiu Island only (Lat. 33° N.), and thus ranging further north of the equator than any other species of the genus.
3. Rhinolophus ferrum-equinum. The great horse-shoe bat, ranges from Britain across Europe and temperate Asia to Japan. It is the R. nippon of the Fauna Japonica according to Mr. Dobson's Monograph of Asiatic Bats.
4. R. minor. Found also in Burma, Yunan, Java, Borneo, &c.
5. Vesperugo pipistrellus. From Britain across Europe and Asia.
6. V. abramus. Also in India and China.
7. V. noctula. From Britain across Europe and Asia.
8. V. molossus. Also in China.
9. Vespertilio capaccinii. Philippine Islands, and Italy! This is V. macrodactylus of the Fauna Japonica according to Mr. Dobson.
10. Miniopterus schreibersii. Philippines, Burma, Malay Islands. This is Vespertilio blepotis of the Fauna Japonica.
11. Talpa wogura. Closely resembles the common mole of Europe, but has six incisors instead of eight in the lower jaw.
12. Talpa mizura. Günth. Allied to T. wogura.
13. Urotrichus talpoides. A peculiar genus of moles confined to Japan. An American species has been named Urotrichus gibsii, and Mr. Lord after comparing the two says that he "can find no difference whatever, either generic or specific. In shape, size, and colour, they are exactly alike." But Dr. Günther (P. Z. S. 1880, p. 441) states that U. gibsii differs so much in dentition from the Japanese species that it should be placed in a distinct genus, which he calls Neurotrichus.
14. Sorex myosurus. A shrew, found also in India and Malaya.
15. Sorex dzi-nezumi.
16. S. umbrinus.
17. S. platycephalus.
18. Ursus arctos. var. A peculiar variety of the European brown bear which inhabits also Amoorland and Kamschatka. It is the Ursus ferox of the Fauna Japonica.
19. Ursus japonicus. A peculiar species allied to the Himalayan and Formosan species. Named U. tibetanus in the Fauna Japonica.
20. Meles anakuma. Differs from the European and Siberian badgers in the form of the skull.
21. Mustela brachyura. A peculiar martin found also in the Kurile Islands.
22. Mustela melanopus. The Japanese sable.
23. M. Japonica. A peculiar martin (See Proc. Zool. Soc. 1865, p. 104).
24. M. Sibericus. Also Siberia and China. This is the M. italsi of the Fauna Japonica according to Dr. Gray.
25. Lutronectes whiteleyi. A new genus and species of otter (P. Z. S. 1867, p. 180). In the Fauna Japonica named Lutra vulgaris.
26. Enhydris marina. The sea-otter of California and Kamschatka.
27. Canis hodophylax. According to Dr. Gray allied to Cuon sumatranus of the Malay Islands, and C. alpinus of Siberia, if not identical with one of them (P. Z. S. 1868, p. 500).
28. Vulpes japonica. A peculiar fox. Canis vulpes of Fauna Japonica.
29. Nyctereutes procyonoides. The racoon-dog of N. China and Amoorland.
30. Lepus brachyurus. A peculiar hare.
31. Sciurus lis. A peculiar squirrel.
32. Pteromys leucogenys. The white-cheeked flying squirrel.
33. P. momoga. Perhaps identical with a Cambojan species (P. Z. S. 1861, p. 137).
34. Myoxus japonicus. A peculiar dormouse. M. elegans of the Fauna Japonica; M. javanicus, Schinz (Synopsis Mammalium, ii. p. 530).
35. Mus argenteus. China.
36. Mus molossinus.
37. M. nezumi.
38. M. speciosus.
39. Cervus sika. A peculiar deer allied to C. pseudaxis of Formosa and C. mantchuricus of Northern China.
40. Nemorhedus crispa. A goat-like antelope allied to N. sumatranus of Sumatra, and N. Swinhoei of Formosa.
41. Sus leucomystax. A wild boar allied to S. taeranus of Formosa.

We thus find that no less than twenty-six out of the forty-one Japanese mammals are peculiar, and if we omit the aërial bats (nine in number), as well as the marine sea-otter, we shall have remaining only thirty strictly land mammalia, of which twenty-five are peculiar, or five-sixths of the whole. Nor does this represent all their speciality; for we have a mole differing in its dentition from the European mole; another superficially resembling but quite distinct from an American species; a peculiar genus of otters; and an antelope whose nearest allies are in Formosa and Sumatra. The importance of these facts will be best understood when we have examined the corresponding affinities of the birds of Japan.

Birds.—Owing to the recent researches of some English residents we have probably a fuller knowledge of the birds than of the mammalia; yet the number of true land-birds ascertained to inhabit the islands either as residents or migrants is only 200, which is less than might be expected considering the highly favourable conditions of mild climate, luxuriant vegetation, and abundance of insect-life, and the extreme riches of the adjacent continent,—Mr. Swinhoe's list of the birds of China containing more than 400 land species, after deducting all which are peculiar to the adjacent islands. Only seventeen species, or about one-twelfth of the whole, are now considered to be peculiar to Japan proper; while seventeen more are peculiar to the various outlying small islands constituting the Bonin and Loo Choo groups. Even of these, six or seven are classed by Mr. Seebohm as probably sub-species or slightly modified forms of continental birds, so that ten only are well-marked species, undoubtedly distinct from those of any other country.

The great majority of the birds are decidedly temperate forms identical with those of Northern Asia and Europe; while no less than forty of the species of land-birds are also found in Britain, or are such slight modifications of British species that the difference is only perceptible to a trained ornithologist. The following list of the land-birds common to Britain and Japan is very interesting, when we consider that these countries are separated by the whole extent of the European and Asiatic continents, or by almost exactly one-fourth of the circumference of the globe:—

Land Birds Common to Great Britain and Japan.[146]

(Either Identical Species or Representative sub-species.)

1. Goldcrest Regulus cristatus sub-sp. orientalis.
2. Marsh tit Parus palustris sub-sp. japonicus.
3. Coal tit Parus ater sub-sp. pekinensis.
4. Long-tailed tit Acredula caudata (the sub-sp. rosea, is British).

5.
Common creeper Certhia familiaris.
6. Nuthatch Sitta europæa sub-sp. amurensis.
7. Carrion crow Corvus corone.
8. Nutcracker Nucifraga caryocatactes.
9. Magpie Pica caudata.
10. Pallass' grey shrike Lanius excubitor sub-sp. major.
11. Waxwing Ampelis garrulus.
12. Grey wagtail Motacilla boarula sub-sp. melanope.
13. Alpine Pipit Anthus spinoletta sub-sp. japonicus.
14. Skylark Alauda arvensis sub-sp. japonica.
15. Common hawfinch Coccothraustes vulgaris.
16. Common Crossbill Loxia curvirostra.
17. Siskin Fringilla spinus.
18. Mealy redpole       ,,      linaria.
19. Brambling       ,,      montifringilla.
20. Tree sparrow Passer montanus.
21. Reed bunting Emberiza schœniculus sub-sp. palustris.
22. Rustic bunting        ,,      rustica.
23. Snow bunting        ,,      nivalis.
24. Chimney swallow Hirundo rustica sub-sp. gutturalis.
25. Sand martin Cotyle riparia.
26. Great spotted woodpecker Picus major sub-sp. japonicus.
27. Lesser spotted woodpecker     ,,   minor.
28. Wryneck Jynx torquilla.
29. Hoopoe Upupa epops.
30. Blue rock pigeon Columba livia.
31. Cuckoo Cuculus canorus.
32. Kingfisher Alcedo ispida sub-sp. bengalensis.
33. Eagle owl Bubo maximus.
34. Snowy owl Surnia nyctea.
35. Long-eared owl Strix otus.
36. Short-eared owl    ,,   brachyotus.
37. Scops owl Scops scops.
38. Jer falcon Falco gyrfalco.
39. Peregrine falcon     ,,   peregrinus.
40. Hobby     ,,   subbuteo.
41. Merlin Falco æsalon.
42. Kestrel Tinnunculus alaudarius sub-sp. japonicus.
43. Osprey Pandion haliäctus.
44. Honey-buzzard Pernis apivorus.
45. White-tailed eagle Haliäetus albicilla.
46. Golden eagle Aquila chrysäetus.
47. Common buzzard Buteo vulgaris sub-sp. plumipes.
48. Hen-harrier Circus cyaneus.
49. Marsh-harrier     ,,    æruginosus.
50. Gos-hawk Astur palumbarius.
51. Sparrow-hawk Accipiter nisus.
52. Ptarmigan Tetrao mutus.
53. Common quail Coturnix communis.

But even these fifty-three species by no means fairly represent the amount of resemblance between Britain and Japan as regards birds; for there are also thrushes, robins, stonechats, wrens, hedge-sparrows, sedge-warblers, jays, starlings, swifts, goatsuckers, and some others, which, though distinct species from our own, have the same general appearance, and give a familiar aspect to the ornithology. There remains, however, a considerable body of Chinese and Siberian species, which link the islands to the neighbouring parts of the continent; and there are also a few which are Malayan or Himalayan rather than Chinese, and thus afford us an interesting problem in distribution.

The seventeen species and sub-species which are altogether peculiar to Japan proper, are for the most part allied to birds of North China and Siberia, but three are decidedly tropical, and one of them—a fruit pigeon (Treron sieboldi)—has no close ally nearer than Burmah and the Himalayas. In the following list the affinities of the species are indicated wherever they have been ascertained:—

List of the Species of Land Birds Peculiar To Japan.

1. Accentor rubidus. Nearly allied to our hedge-sparrow, and less closely to the Central Asian A. immaculatus.
(1a. Hypsipetes amaurotis. Migrates to the Corea, otherwise peculiar.)
2. Zosterops japonica. Allied to two Chinese species.
3. Lusciniola pryeri.
4. Garrulus japonicus. Allied to the Siberian and British Jays.
5. Fringilla kawarahiba. Allied to the Chinese greenfinch.
6. Emberiza ciopsis. Allied to the E. Siberian bunting E. cioides, of which it may be considered a sub-species.
7.        ,,      yessoensis. A distinct species.
8.        ,,      personata. A sub-species of E. spodocephala.
9. Gecinus awokera. A distinct species of green woodpecker.
10. Picus namiyei. Allied to a Formosan species.
11. Treron sieboldi. Allied to T. sphenura of the Himalayas, and to a Formosan species.
12. Carpophaga ianthina. A distinct species of fruit-pigeon.
13. Bubo blakistoni. Allied to a Philippine eagle-owl.
14. Scops semitorgues. A distinct species.
15. Phasianus versicolor. A distinct species.
16.        ,,       sœmmeringi. A distinct species.
17.        ,,       scintillaus. A sub-species of the last.

The large number of seventeen peculiar species in the outlying Bonin and Loo Choo Islands is an interesting feature of Japanese ornithology. The comparative remoteness of these islands, their mild sub-tropical climate and luxuriant vegetation, and perhaps the absence of violent storms and their being situated out of the line of continental migration, seem to be the conditions that have favoured the specialisation of modified types adapted to the new environment.

Japan Birds Recurring in Distant Areas.—The most interesting feature in the ornithology of Japan is, undoubtedly, the presence of several species which indicate an alliance with such remote districts as the Himalayas, the Malay Islands, and Europe. Among the peculiar species, the most remarkable of this class are,—the fruit-pigeon of the genus Treron, entirely unknown in China, but reappearing in Formosa and Japan; the Hypsipetes, whose nearest ally is in South China at a distance of nearly 500 miles; and the jay (Garrulus japonicus), whose near ally (G. glandarius) inhabits Europe only, at a distance of 3,700 miles. But even more extraordinary are the following non-peculiar species:—Spizaetus orientalis, a crested eagle, inhabiting the Himalayas, Formosa, and Japan, but unknown in Southern or Eastern China; Ceryle guttata, a spotted kingfisher, almost confined to the Himalayas and Japan, though occurring rarely in Central China; and Halcyon coromanda, a brilliant red kingfisher inhabiting Northern India, the Malay Islands to Celebes, Formosa, and Japan. We have here an excellent illustration of the favourable conditions which islands afford both for species which elsewhere live further south (Halcyon coromanda), and for the preservation in isolated colonies of species which are verging towards extinction; for such we must consider the above-named eagle and kingfisher, both confined to a very limited area on the continent, but surviving in remote islands. Referring to our account of the birth, growth, and death of a species (in Chapter IV.) it can hardly be doubted that the Ceryle guttata formerly ranged from the Himalayas to Japan, and has now almost died out in the intervening area owing to geographical and physical changes, a subject which will be better discussed when we have examined the interesting fauna of the island of Formosa.

The other orders of animals are not yet sufficiently known to enable us to found any accurate conclusions upon them. The main facts of their distribution have already been given in my Geographical Distribution of Animals (Vol I., pp. 227-231), and they sufficiently agree with the birds and mammalia in showing a mixture of temperate and tropical forms with a considerable proportion of peculiar species. Owing to the comparatively easy passage from the northern extremity of Japan through the island of Saghalien to the mainland of Asia, a large number of temperate forms of insects and birds are still able to enter the country, and thus diminish the proportionate number of peculiar species. In the case of mammals this is more difficult; and the large proportion of specific difference in their case is a good indication of the comparatively remote epoch at which Japan was finally separated from the continent. How long ago this separation took place we cannot of course tell, but we may be sure it was much longer than in the case of our own islands, and therefore probably in the earlier portion of the Pliocene period.

Formosa.

Among recent continental islands there is probably none that surpasses in interest and instructiveness the Chinese island named by the Portuguese, Formosa, or "The Beautiful." Till quite recently it was a terra incognita to naturalists, and we owe almost all our present knowledge of it to a single man, the late Mr. Robert Swinhoe, who, in his official capacity as one of our consuls in China, visited it several times between 1856 and 1866, besides residing on it for more than a year. During this period he devoted all his spare time and energy to the study of natural history, more especially of the two important groups, birds and mammals; and by employing a large staff of native collectors and hunters, he obtained a very complete knowledge of its fauna. In this case, too, we have the great advantage of a very thorough knowledge of the adjacent parts of the continent, in great part due to Mr. Swinhoe's own exertions during the twenty years of his service in that country. We possess, too, the further advantage of having the whole of the available materials in these two classes collected together by Mr. Swinhoe himself after full examination and comparison of specimens; so that there is probably no part of the world (if we except Europe, North America, and British India) of whose warm-blooded vertebrates we possess fuller or more accurate knowledge than we do of those of the coast districts of China and its islands.[147]

Physical Features of Formosa.—The island of Formosa is nearly half the size of Ireland, being 220 miles long, and from twenty to eighty miles wide. It is traversed down its centre by a fine mountain range, which reaches an altitude of about 8,000 feet in the south and 12,000 feet in the northern half of the island, and whose higher slopes and valleys are everywhere clothed with magnificent forests. It is crossed by the line of the Tropic of Cancer a little south of its centre; and this position, combined with its lofty mountains, gives it an unusual variety of tropical and temperate climates. These circumstances are all highly favourable to the preservation and development of animal life, and from what we already know of its productions, it seems probable that few, if any islands of approximately the same size and equally removed from a continent will be found to equal it in the number and variety of their higher animals. The outline map (at page 392) shows that Formosa is connected with the mainland by a submerged bank, the hundred-fathom line including it along with Hainan to the south-west and Japan on the north-east; while the line of two-hundred fathoms includes also the Madjico-Sima and Loo-Choo Islands, and may, perhaps, mark out approximately the last great extension of the Asiatic continent, the submergence of which isolated these islands from the mainland.

Animal Life of Formosa.—We are at present acquainted with 35 species of mammalia, and 128 species of land-birds from Formosa, fourteen of the former and forty-three of the latter being peculiar, while the remainder inhabit also some part of the continent or adjacent islands. This proportion of peculiar species is perhaps (as regards the birds) the highest to be met with in any island which can be classed as both continental and recent, and this, in all probability, implies that the epoch of separation is somewhat remote. It was not, however, remote enough to reach back to a time when the continental fauna was very different from what it is now, for we find all the chief types of living Asiatic mammalia represented in this small island. Thus we have monkeys; insectivora; numerous carnivora; pigs, deer, antelopes, and cattle among ungulata; numerous rodents, and the edentate Manis,—a very fair representation of Asiatic mammals, all being of known genera, and of species either absolutely identical with some still living elsewhere or very closely allied to them. The birds exhibit analogous phenomena, with the exception that we have here two peculiar and very interesting genera.

But besides the amount of specific and generic modification that has occurred, we have another indication of the lapse of time in the peculiar relations of a large proportion of the Formosan animals, which show that a great change in the distribution of Asiatic species must have taken place since the separation of the island from the continent. Before pointing these out it will be advantageous to give lists of the mammalia and peculiar birds of the island, as we shall have frequent occasion to refer to them.

List of the Mammalia of Formosa. (The peculiar species are printed in italics.)

  1. Macacus cyclopis. A rock-monkey more allied to M. rhesus of India than to M. sancti-johannis of South China.
  2. Pteropus formosus. A fruit-bat closely allied to the Japanese species. None of the genus are found in China.
  3. Vesperugo abramus. China.
  4. Vespertilio formosus. Black and orange Bat. China.
  5. Nyctinomus cestonii. Large-eared Bat. China, S. Europe.
  6. Talpa insularis. A blind mole of a peculiar species.
  7. Sorex murinus. Musk Rat. China.
  8. Sorex sp. A shrew, undescribed.
  9. Erinaceus sp. A Hedgehog, undescribed.
10. Ursus tibetanus. The Tibetan Bear. Himalayas and North China.
11. Helictis subaurantiaca. The orange-tinted Tree Civet. Allied to H. nipalensis of the Himalayas more than to H. moschata of China.
12. Martes flavigula, var. The yellow-necked Marten. India, China.
13. Felis macroscelis. The clouded Tiger of Siam and Malaya.
14. Felis viverrina. The Asiatic wild Cat. Himalayas and Malacca.
15. Felis chinensis. The Chinese Tiger Cat. China.
16. Viverricula malaccensis. Spotted Civet. China, India.
17. Paguma larvata. Gem-faced Civet. China.
18. Sus taivanus. Allied to the wild Pig of Japan.
19. Cervulus reevesii. Reeve's Muntjac. China.
20. Cervus pseudaxis. Formosan Spotted Deer. Allied to C. sika of Japan.
21. Cervus swinhoii. Swinhoe's Rusa Deer. Allied to Indian and Malayan species.
22. Nemorhedus swinhoii. Swinhoe's Goat-antelope. Allied to the species of Sumatra and Japan.
23. Bos chinensis. South China wild Cow.
24. Mus bandicota. The Bandicoot Rat. Perhaps introduced from India.
25. Mus indicus. Indian Rat.
26. Mus coxinga. Spinous Country-rat.
27. Mus canna. Silken Country-rat.
28. Mus losca. Brown Country-rat.
29. Sciurus castaneoventris. Chestnut-bellied Squirrel. China and Hainan.
30. Sciurus m'clellandi. M‘Clelland's Squirrel. Himalayas, China.
31. Sciuropterus kaleensis. Small Formosan Flying Squirrel. Allied to S. alboniger of Nepal.
32. Pteromys grandis. Large Red Flying Squirrel. Allied to Himalayan and Bornean species. From North Formosa.
33. Pteromys pectoralis. White-breasted Flying Squirrel. From South Formosa.
34. Lepus sinensis. Chinese Hare. Inhabits South China.
35. Manis dalmanni. Scaly Ant-eater. China and the
Himalayas.

The most interesting and suggestive feature connected with these Formosan mammals is the identity or affinity of several of them, with Indian or Malayan rather than with Chinese species. We have the rock-monkey of Formosa allied to the rhesus monkeys of India and Burma, not to those of South China and Hainan. The tree civet (Helictis subaurantiaca), and the small flying squirrel (Sciuropterus kaleensis), are both allied to Himalayan species. Swinhoe's deer and goat-antelope are nearest to Malayan species, as are the red and white-breasted flying squirrels; while the fruit-bat, the wild pig, and the spotted deer are all allied to peculiar Japanese species. The clouded tiger is a Malay species unknown in China, while the Asiatic wild cat is a native of the Himalayas and Malacca. It is clear, therefore, that before Formosa was separated from the mainland the above named animals or their ancestral types must have ranged over the intervening country as far as the Himalayas on the west, Japan on the north, and Borneo or the Philippines on the south; and that after that event occurred, the conditions were so materially changed as to lead to the extinction of these species in what are now the coast provinces of China, while they or their modified descendants continued to exist in the dense forests of the Himalayas and the Malay Islands, and in such detached islands as Formosa and Japan. We will now see what additional light is thrown upon this subject by an examination of the birds.

List of the Land Birds peculiar to Formosa.

Turdidæ (Thrushes).

1. Turdus albiceps. Allied to Chinese species.

Sylvidiæ (Warblers).

2. Cisticola volitans. Allied to C. schœnicola of India and China.
3. Herbivox cantans. Sub-species of H. cantillaus of N. China and Japan.
4. Notodela montium. Allied to N. leucura of the Himalayas; no ally in
China.

Timaliidæ (Babblers).

5. Pomatorhinus musicus. Allies in S. China and the Himalayas.
6. P. erythroenemis.              Do.                  do.
7. Garrulax ruficeps. Allied to G. albogularis of N. India and East Thibet, not to the species of S. China (G. sannio).
8. Janthocincla pœcilorhyncha. Allied to J. cœrulata of the Himalayas. None of the genus in China.
9. Trochalopteron taivanus. Allied to a Chinese species.
10. Alcippe morrisoniana. Near the Himalayan A. nipalensis. None of the genus in China.
11. A. brunnea.                                        Do.                                        do.
12. Sibia auricularis. Allied to the Himalayan S. capistrata. The genus not known in
China.

Panuridæ (Bearded Tits, &c.).

13. Suthora bulomachus. Allied to the Chinese S. suffusa.

Cinclidæ (Dippers and Whistling Thrushes).

14. Myiophoneus insularis. Allied to M. horsfieldi of South India.

Paridæ (Tits).

15. Parus insperatus. Sub-species of P. monticola of the Himalayas and East Thibet.
16. P. castaneiventris. Allied to P. varius of
Japan.

Liotrichidæ (Hill Tits).

17. Liocichla steerii. A peculiar genus of a specially Himalayan family, quite unknown in China.

Pycnonotidæ (Bulbuls).

18. Pycnonotus (Spizixos) cinereicapillus. Very near P. semitorques of China.
19. Hypsipetes nigerrimus. Allied to H. concolor of Assam, not to H. macclellandi of
China.

Oriolidæ (Orioles).

20. Analcipus ardens. Allied to A. traillii of the Himalayas and Tenasserim.

Campephagidæ (Caterpillar Shrikes).

21. Graucalus rex-pineti. Closely allied to the Indian G. macei. No ally in China.

Dicruridæ (King Crows).

22. Chaptia brauniana. Closely allied to C. ænea of Assam. No ally in China.

Muscicapidæ (Flycatchers).

23. Cyornis vivida. Allied to C. rubeculoides of India.

Corvidæ (Jays and Crows).

24. Garrulus taivanus. Allied to G. sinensis of S. China.
25. Urocissa cœrulea. A very distinct species from its Indian and Chinese allies.
26. Dendrocitta formosæ. A sub-species of the Chinese D. sinensis.

Ploceidæ (Weaver Finches).

27. Munia formosana. Allied to M. rubronigra of India and Burmah.

Alaudidæ (Larks).

28. Alauda sala. Allies in South China.
29. A. wattersi.             
 Do.

Pittidæ (Pittas).

30. Pitta oreas. Allied to P. cyanoptera of Malaya and S. China.

Picidæ (Woodpeckers).

31. Picus insularis. Allied to P. leuconotus of Japan and Siberia.

Megalæmidæ.

32. Megalæma nuchalis. Allied to M. oortii of Sumatra and M. faber of Hainan. No allies in China.

Caprimulgidæ (Goatsuckers).

33. Caprimulgus stictomus. A sub-species of C. monticolus of India and China.

Columbidæ (Pigeons).

34. Treron formosæ. Allied to Malayan species.
35. Sphenocercus sororius. Allied to Malay species and to S. sieboldi of Japan. No allies of these two birds inhabit China.
36. Chalcophaps formosana. Allied to the Indian species which extends to Tenasserim and
Hainan.

Tetraonidæ (Grouse and Partridges).

37. Orcoperdix crudigularis. A peculiar genus of partridges.
38. Bambusicola sonorivox. Allied to the Chinese B. thoracica.
39. Arcoturnix rostrata. Allied to the Chinese A. blakistonii.

Phasianidæ (Pheasants).

40. Phasianus formosanus. Allied to P. torquatus of China.
41. Euplocamus swinhoii. A very peculiar and beautiful species allied to the tropical fire-backed pheasants, and to the silver pheasant of North
China.

Strigidæ (Owls).

42. Athene pardalota. Closely allied to a Chinese species.
43. Lempigius hambroekii. Allied to a Chinese
species.

This list exhibits to us the marvellous fact that more than half the peculiar species of Formosan birds have their nearest allies in such remote regions as the Himalayas, South India, the Malay Islands, or Japan, rather than in the adjacent parts of the Asiatic continent. Fourteen species have Himalayan allies, and six of these belong to genera which are unknown in China. One has its nearest ally in the Nilgherries, and five in the Malay Islands; and of these six, four belong to genera which are not Chinese. Two have their only near allies in Japan. Perhaps more curious still are those cases in which, though the genus is Chinese, the nearest allied species is to be sought for in some remote region. Thus we have the Formosan babbler (Garrulax ruficeps) not allied to the species found in South China, but to one inhabiting North India and East Thibet; while the black bulbul (Hypsipetes nigerrimus), is not allied to the Chinese species but to an Assamese form.

In the same category as the above we must place eight species not peculiar to Formosa, but which are Indian or Malayan rather than Chinese, so that they offer examples of discontinuous distribution somewhat analogous to what we found to occur in Japan. These are enumerated in the following list.

Species of Birds common to Formosa and India or Malaya, but not found in China.

1. Siphia superciliaris. The Rufous-breasted Flycatcher of the S. E. Himalayas.

2. Halcyon coromanda. The Great Red Kingfisher of India, Malaya, and Japan.

3. Palumbus pulchricollis. The Darjeeling Wood-pigeon of the S. E. Himalayas.

4. Turnix dussumieri. The larger Button-quail of India.

5. Spizaetus nipalensis. The Spotted Hawk-eagle of Nepal and Assam.

6. Lophospiza trivirgata. The Crested Gos-hawk of the Malay Islands.

7. Bulaca newarensis. The Brown Wood-owl of the Himalayas.

8. Strix candida. The Grass-owl of India and Malaya.

The most interesting of the above are the pigeon and the flycatcher, both of which are, so far as yet known, strictly confined to the Himalayan mountains and Formosa. They thus afford examples of discontinuous specific distribution exactly parallel to that of the great spotted kingfisher, already referred to as found only in the Himalayas and Japan.

Comparison of the Faunas of Hainan, Formosa, and Japan.—The island of Hainan on the extreme south of China, and only separated from the mainland by a strait fifteen miles wide, appears to have considerable similarity to Formosa, inasmuch as it possesses seventeen peculiar land-birds (out of 130 obtained by Mr. Swinhoe), two of which are close allies of Formosan species, while two others are identical. We also find four species whose nearest allies are in the Himalayas. Our knowledge of this island and of the adjacent coast of China is not yet sufficient to enable us to form an accurate judgment of its relations, but it seems probable that it was separated from the continent at, approximately, the same epoch as Formosa and Japan, and that the special features of each of these islands are mainly due to their geographical position. Formosa, being more completely isolated than either of the others, possesses a larger proportion of peculiar species of birds, while its tropical situation and lofty mountain ranges have enabled it to preserve an unusual number of Himalayan and Malayan forms. Japan, almost equally isolated towards the south, and having a much greater variety of climate as well as a much larger area, possesses about an equal number of mammalia with Formosa, and an even larger proportion of peculiar species. Its birds, however, though more numerous are less peculiar; and this is probably due to the large number of species which migrate northwards in summer, and find it easy to enter Japan through the Kurile Isles or Saghalien.[148] Japan too, is largely peopled by those northern types which have an unusually wide range, and which, being almost all migratory, are accustomed to cross over seas of moderate extent. The regular or occasional influx of these species prevents the formation of special insular races, such as are almost always produced when a portion of the population of a species remains for a considerable time completely isolated. We thus have explained the curious fact, that while the mammalia of the two islands are almost equally peculiar, (those of Japan being most so in the present state of our knowledge), the birds of Formosa show a far greater number of peculiar species than those of Japan.

General Remarks on Recent Continental Islands.—We have now briefly sketched the zoological peculiarities of an illustrative series of recent continental islands, commencing with one of the most recent—Great Britain—in which the process of formation of peculiar species has only just commenced, and terminating with Formosa, probably one of the most ancient of the series, and which accordingly presents us with a very large proportion of peculiar species, not only in its mammalia, which have no means of crossing the wide strait which separates it from the mainland, but also in its birds, many of which are quite able to cross over.

Here, too, we obtain a glimpse of the way in which species die out and are replaced by others, which quite agrees with what the theory of evolution assures us must have occurred. On a continent, the process of extinction will generally take effect on the circumference of the area of distribution, because it is there that the species comes into contact with such adverse conditions or competing forms as prevent it from advancing further. A very slight change will evidently turn the scale and cause the species to contract its range, and this usually goes on till it is reduced to a very restricted area, and finally becomes extinct. It may conceivably happen (and almost certainly has sometimes happened) that the process of restriction of range by adverse conditions may act in one direction only, and over a limited district, so as ultimately to divide the specific area into two separated parts, in each of which a portion of the species will continue to maintain itself. We have seen that there is reason to believe that this has occurred in a very few cases both in North America and in Northern Asia. (See pp. 65-68.) But the same thing has certainly occurred in a considerable number of cases, only it has resulted in the divided areas being occupied by representative forms instead of by the very same species. The cause of this is very easy to understand. We have already shown that there is a large amount of local variation in a considerable number of species, and we may be sure that were it not for the constant intermingling and intercrossing of the individuals inhabiting adjacent localities this tendency to local variation in adaptation to slightly different conditions, would soon form distinct races. But as soon as the area is divided into two portions the intercrossing is stopped, and the usual result is that two closely allied races, classed as representative species, become formed. Such pairs of allied species on the two sides of a continent, or in two detached areas, are very numerous; and their existence is only explicable on the supposition that they are descendants of a parent form which once occupied an area comprising that of both of them,—that this area then became discontinuous,—and, lastly, that, as a consequence of the discontinuity, the two sections of the parent species became segregated into distinct races or new species.

Now, when the division of the area leaves one portion of the species in an island, a similar modification of the species, either in the island or in the continent, occurs, resulting in closely-allied but distinct forms; and such forms are, as we have seen, highly characteristic of island-faunas. But islands also favour the occasional preservation of the unchanged species—a phenomenon which very rarely occurs in continents. This is probably due to the absence of competition in islands, so that the parent species there maintains itself unchanged, while the continental portion, by the force of that competition, is driven back to some remote mountain area, where it also obtains a comparative freedom from competition. Thus may be explained the curious fact, that the species common to Formosa and India are generally confined to limited areas in the Himalayas, or in other cases are found only in remote islands, as Japan or Hainan.

The distribution and affinities of the animals of continental islands thus throws much light on that obscure subject—the decay and extinction of species; while the numerous and delicate gradations in the modification of the continental species, from perfect identity, through slight varieties, local forms, and insular races, to well-defined species and even distinct genera, afford an overwhelming mass of evidence in favour of the theory of "descent with modification."

We shall now pass on to another class of islands, which, though originally forming parts of continents, were separated from them at very remote epochs. This antiquity is clearly manifested in their existing faunas, which present many peculiarities, and offer some most curious problems to the student of distribution.


146   Extracted from Messrs. Blakiston and Pryer's Catalogue of Birds of Japan (Ibis, 1878, p. 209), with Mr. Seebohm's additions and corrections in his Birds of the Japanese Empire 1890. Accidental stragglers are not reckoned as British birds.

147   Mr. Swinhoe died in October, 1877, at the early age of forty-two. His writings on natural history are chiefly scattered through the volumes of the Proceedings of the Zoological Society and The Ibis; the whole being summarised in his Catalogue of the Mammals of South China and Formosa (P. Z. S., 1870, p. 615), and his Catalogue of the Birds of China and its Islands (P. Z. S., 1871, p. 337).

148   Captain Blakiston has shown that the northern island—Yezo—is much more temperate and less peculiar in its zoology than the central and southern islands. This is no doubt dependent chiefly on the considerable change of climate that occurs on passing the Tsu-garu strait.