10. Garrulus taivanus.The Formosan jay is a very close ally of the preceding, confined to the island of Formosa.
11. Garrulus japonicus.The Japanese jay is nearly allied to our common British species, being somewhat smaller and less brightly coloured, and with black orbits; yet these are the most widely separated species of the genus. According to Mr. Seebohm this species is equally allied to the Chinese and Siberian jays.
In the accompanying map (see frontispiece) we have laid down the distribution of each species so far as it can be ascertained from the works of Sharpe and Dresser for Europe, Jerdon for India, Swinhoe for China, and Mr. Seebohm's recent work for Japan. There is, however, much uncertainty in many places, and gaps have to be filled up conjecturally, while such a large part of Asia is still very imperfectly explored, that considerable modifications may have to be made when the country becomes more accurately known. But though details may be modified we can hardly suppose that the great features of the several specific areas, or their relations to each other will be much affected; and these are what we have chiefly to consider as bearing on the questions here discussed.
The first thing that strikes us on looking at the map, is, the small amount of overlapping of the several areas, and the isolation of many of the species; while the next most striking feature is the manner in which the Asiatic species almost surround a vast area in which no jays are found. The only species with large areas, are the European G. glandarius and the Asiatic G. Brandti. The former has three species overlapping itin Algeria, in South-eastern and North-eastern Europe respectively. The Syrian jay (No. 4), is not known to occur anywhere with the black-headed jay (No. 3), and perhaps the two areas do not meet. The Persian jay (No. 5), is quite isolated. The Himalayan and Chinese jays (Nos. 7, 8, and 9) form a group which are isolated from the rest of the genus; while the Japanese jay (No. 11), is also completely isolated as regards the European jays to which it is nearly allied. These peculiarities of distribution are no doubt in part dependent on the habits of the jays, which live only in well-wooded districts, among deciduous trees, and are essentially non-migratory in their habits, though sometimes moving southwards in winter. This will explain their absence from the vast desert area of Central Asia, but it will not account for the gap between the North and South Chinese species, nor for the absence of jays from the wooded hills of Turkestan, where Mr. N. A. Severtzoff collected assiduously, obtaining 384 species of birds but no jay. These peculiarities, and the fact that jays are never very abundant anywhere, seem to indicate that the genus is now a decaying one, and that it has at no very distant epoch occupied a larger and more continuous area, such as that of the genus Parus at the present day.
Discontinuous generic Areas.It is not very easy to find good examples of genera whose species occupy two or more quite disconnected areas, for though such cases may not be rare, we are seldom in a position to mark out the limits of the several species with sufficient accuracy. The best and most remarkable case among European birds is that of the blue magpies, forming the genus Cyanopica. One species (C. cooki) is confined (as already stated) to the wooded and mountainous districts of Spain and Portugal, while the only other species of the genus (C. cyanus) is found far away in North-eastern Asia and Japan, so that the two species are separated by about 5,000 miles of continuous land. Another case is that of the curious little water-moles forming the genus Mygale, one species M. muscovitica, being found only on the banks of the Volga and Don in South-eastern Russia, while the other, M. pyrenaica, is confined to streams on the northern side of the Pyrenees. In tropical America there are four different kinds of bell-birds belonging to the genus Chasmorhynchus, each of which appears to inhabit a restricted area completely separated from the others. The most northerly is C. tricarunculatus of Costa Rica and Veragua, a brown bird with a white head and three long caruncles growing upwards at the base of the beak. Next comes C. variegatus, in Venezuela, a white bird with a brown head and numerous caruncles on the throat, perhaps conterminous with the last; in Guiana, extending to near the mouth of the Rio Negro, we have C. niveus, the bell-bird described by Waterton, which is pure white, with a single long fleshy caruncle at the base of the beak; the last species, C. nudicollis, inhabits South-east Brazil, and is also white, but with black stripes over the eyes, and with a naked throat. These birds are about the size of thrushes, and are all remarkable for their loud, ringing notes, like a bell or a blow on an anvil, as well as for their peculiar colours. They are therefore known to the native Indians wherever they exist, and we may be the more sure that they do not spread over the intervening areas where they have never been found, and where the natives know nothing of them.
A good example of isolated species of a group nearer home, is afforded by the snow-partridges of the genus Tetraogallus. One species inhabits the Caucasus range and nowhere else, keeping to the higher slopes from 6,000 to 11,000 feet above the sea, and accompanying the ibex in its wanderings, as both feed on the same plants. Another has a wider range in Asia Minor and Persia, from the Taurus mountains to the South-east corner of the Caspian Sea; a third species inhabits the Western Himalayas, between the forests and perpetual snow, extending eastwards to Nepal; while a fourth is found on the north side of the mountains in Thibet, and the ranges of these two perhaps overlap; the last species inhabit the Altai mountains, and like the two first appears to be completely separated from all its allies.
There are some few still more extraordinary cases in which the species of one genus are separated in remote continents or islands. The most striking of these is that of the tapirs, forming the genus Tapirus, of which there are two or three species in South America, and one very distinct species in Malacca and Borneo, separated by nearly half the circumference of the globe. Another example among quadrupeds is a peculiar genus of moles named Urotrichus, of which one species inhabits Japan and the other British Columbia. The cuckoo-like honey-guides, forming the genus Indicator, are tolerably abundant in tropical Africa, but there are two outlying species, one in the Eastern Himalaya mountains, the other in Borneo, both very rare, and recently an allied species has been found in the Malay peninsula. The beautiful blue and green thrush-tits forming the genus Cochoa, have two species in the Eastern Himalayas and Eastern China, while the third is confined to Java; the curious genus Eupetes, supposed to be allied to the dippers, has one species in Sumatra and Malacca, while four other species are found two thousand miles distant in New Guinea; lastly, the lovely ground-thrushes of the genus Pitta, range from Hindostan to Australia, while a single species, far removed from all its near allies, inhabits West Africa.
Peculiarities of Generic, and Family Distribution.The examples now given sufficiently illustrate the mode in which the several species of a genus are distributed. We have next to consider genera as the component parts of families, and families of orders, from the same point of view.
All the phenomena presented by the species of a genus are reproduced by the genera of a family, and often in a more marked degree. Owing, however, to the extreme restriction of genera by modern naturalists, there are not many among the higher animals that have a world-wide distribution. Among the mammalia there is no such thing as a truly cosmopolitan genus. This is owing to the absence of all the higher orders except the mice from Australia, while the genus Mus, which occurs there, is represented by a distinct group, Hesperomys, in America. If, however, we consider the Australian dingo as a native animal we might class the genus Canis as cosmopolite, but the wild dogs of South America are now formed into separate genera by some naturalists. Many genera, however, range over three or more continents, as Felis (the cat genus) absent only from Australia; Ursus (the bear genus) absent from Australia and tropical Africa; Cervus (the deer genus) with nearly the same range; and Sciurus (the squirrel genus) found in all the continents but Australia. Among birds Turdus, the thrush, and Hirundo, the swallow genus, are the only perching birds which are truly cosmopolites; but there are many genera of hawks, owls, wading and swimming birds, which have a world-wide range.
All the phenomena presented by the species of a genus are reproduced by the genera of a family, and often in a more marked degree. Owing, however, to the extreme restriction of genera by modern naturalists, there are not many among the higher animals that have a world-wide distribution. Among the mammalia there is no such thing as a truly cosmopolitan genus. This is owing to the absence of all the higher orders except the mice from Australia, while the genus Mus, which occurs there, is represented by a distinct group, Hesperomys, in America. If, however, we consider the Australian dingo as a native animal we might class the genus Canis as cosmopolite, but the wild dogs of South America are now formed into separate genera by some naturalists. Many genera, however, range over three or more continents, as Felis (the cat genus) absent only from Australia; Ursus (the bear genus) absent from Australia and tropical Africa; Cervus (the deer genus) with nearly the same range; and Sciurus (the squirrel genus) found in all the continents but Australia. Among birds Turdus, the thrush, and Hirundo, the swallow genus, are the only perching birds which are truly cosmopolites; but there are many genera of hawks, owls, wading and swimming birds, which have a world-wide range.
As a great many genera consist of single species there is no lack of cases of great restriction, such as the curious lemur called the "potto," which is found only at Sierra Leone, and forms the genus Perodicticus; the true chinchillas found only in the Andes of Peru and Chili south of 9° S. lat. and between 8,000 and 12,000 feet elevation; several genera of finches each confined to limited portions of the higher Himalayas, the blood-pheasants (Ithaginis) found only above 10,000 feet from Nepal to East Thibet; the bald-headed starling of the Philippine islands, the lyre-birds of East Australia, and a host of others.
It is among the different genera of the same family that we meet with the most striking examples of discontinuity, although these genera are often as unmistakably allied as are the species of a genus; and it is these cases that furnish the most interesting problems to the student of distribution. We must therefore consider them somewhat more fully.
Among mammalia the most remarkable of these divided families is that of the camels, of which one genus Camelus, the true camels, comprising the camel and dromedary, is confined to Asia, while the other Auchenia, comprising the llamas and alpacas, is found only in the high Andes and in the plains of temperate South America. Not only are these two genera separated by the Atlantic and by the greater part of the land of two continents, but one is confined to the Northern and the other to the Southern hemisphere. The next case, though not so well known, is equally remarkable; it is that of the Centetidæ, a family of small insectivorous animals, which are wholly confined to Madagascar and the large West Indian islands Cuba and Hayti, the former containing five genera and the latter a single genus with a species in each island. Here again we have the whole continent of Africa as well as the Atlantic ocean separating allied genera. Two families (or subfamilies) of rat-like animals, Octodontidæ and Echimyidæ, are also divided by the Atlantic. Both are mainly South American, but the former has two genera in North and East Africa, and the latter also two in South and West Africa. Two other families of mammalia, though confined to the Eastern hemisphere, are yet markedly discontinuous. The Tragulidæ are small deer-like animals, known as chevrotains or mouse-deer, abundant in India and the larger Malay islands and forming the genus Tragulus; while another genus, Hyomoschus, is confined to West Africa. The other family is the Simiidæ or anthropoid apes, in which we have the gorilla and chimpanzee confined to West and Central Africa, while the allied orangs are found only in the islands of Sumatra and Borneo, the two groups being separated by a greater space than the Echimyidæ and other rodents of Africa and South America.
Among birds and reptiles we have several families, which, from being found only within the tropics of Asia, Africa, and America, have been termed tropicopolitan groups. The Megalæmidæ or barbets are gaily coloured fruit-eating birds, almost equally abundant in tropical Asia and Africa, but less plentiful in America, where they probably suffer from the competition of the larger sized toucans. The genera of each country are distinct, but all are closely allied, the family being a very natural one. The trogons form a family of very gorgeously coloured and remarkable insect-eating birds very abundant in tropical America, less so in Asia, and with a single genus of two species in Africa.
Among reptiles we have two families of snakesthe Dendrophidæ or tree-snakes, and the Dryiophidæ or green whip-snakeswhich are also found in the three tropical regions of Asia, Africa, and America, but in these cases even some of the genera are common to Asia and Africa, or to Africa and America. The lizards forming the family Amphisbænidæ are divided between tropical Africa and America, a few species only occurring in the southern portion of the adjacent temperate regions; while even the peculiarly American family of the iguanas is represented by two genera in Madagascar, and one in the Fiji and Friendly Islands. Passing on to the Amphibians the worm-like Cæciliadæ are tropicopolitan, as are also the toads of the family Engystomatidæ. Insects also furnish some analogous cases, three genera of Cicindelidæ, (Pogonostoma, Ctenostoma, and Peridexia) showing a decided connection between this family in South America and Madagascar; while the beautiful family of diurnal moths, Uraniidæ, is confined to the same two countries. A somewhat similar but better known illustration is afforded by the two genera of ostriches, one confined to Africa and Arabia, the other to the plains of temperate South America.
General features of Overlapping and Discontinuous Areas.These numerous examples of discontinuous genera and families form an important section of the facts of animal dispersal which any true theory must satisfactorily account for. In greater or less prominence they are to be found all over the world, and in every group of animals, and they grade imperceptibly into those cases of conterminous and overlapping areas which we have seen to prevail in most extensive groups of species, and which are perhaps even more common in those large families which consist of many closely allied genera. A sufficient proof of the overlapping of generic areas is the occurrence of a number of genera of the same family together. Thus in France or Italy about twenty genera of warblers (Sylviadæ) are found, and as each of the thirty-three genera of this family inhabiting temperate Europe and Asia has a different area, a great number must here overlap. So, in most parts of Africa, at least ten or twelve genera of antelopes may be found, and in South America a large proportion of the genera of monkeys of the family Cebidæ occur in many districts; and still more is this the case with the larger bird families, such as the tanagers, the tyrant shrikes, or the tree-creepers, so that there is in all these extensive families no genus whose area does not overlap that of many others. Then among the moderately extensive families we find a few instances of one or two genera isolated from the rest, as the spectacled bear, Tremarctos, found only in Chili, while the remainder of the family extends from Europe and Asia over North America to the Mountains of Mexico, but no further south; the Bovidæ, or hollow-horned ruminants, which have a few isolated genera in the Rocky Mountains and the islands of Sumatra and Celebes; and from these we pass on to the cases of wide separation already given.
Restricted Areas of Families.As families sometimes consist of single genera and even single species, they often present examples of very restricted range; but what is perhaps more interesting are those cases in which a family contains numerous species and sometimes even several genera, and yet is confined to a narrow area. Such are the golden moles (Chrysochloridæ) consisting of two genera and three species, confined to extratropical South Africa; the hill-tits (Liotrichidæ), a family of numerous genera and species mainly confined to the Himalayas, but with a few straggling species in the Malay countries and the mountains of China; the Pteroptochidæ, large wren-like birds, consisting of eight genera and nineteen species, almost entirely confined to temperate South America and the Andes; and the birds-of-paradise, consisting of nineteen or twenty genera and about thirty-five species, almost all inhabitants of New Guinea and the immediately surrounding islands, while a few, doubtfully belonging to the family, extend to East Australia. Among reptiles the most striking case of restriction is that of the rough-tailed burrowing snakes (Uropeltidæ), the five genera and eighteen species being strictly confined to Ceylon and the southern parts of the Indian Peninsula.