II. Northern Zone.
The boundaries of the Northern Zone coincide in the main with the northern limit of the Equatorial Zone; but at three different points they overlap the latter, as has been already indicated. This happens in, and east of, Syria, where the mixed faunæ of the Jordan and the rivers of Mesopotamia demand the inclusion of this territory into the Northern Zone as well as the Equatorial; in the island of Formosa, where a Salmonoid and several Japanese Cyprinoids flourish; and in Central America, where a Lepidosteus, a Cyprinoid (Sclerognathus meridionalis), and an Amiurus (A. meridionalis) represent the North American fauna in the midst of a host of tropical forms.
A separate Arctic Zone does not exist for Freshwater fishes; ichthyic life becomes extinct towards the pole as soon as the fresh water remains frozen throughout the year, or thaws for a few weeks only; and the few fishes which extend into high latitudes, in which lakes are open for two or three months in the year, belong to types in no wise differing from those of the more temperate south. The highest latitude at which fishes have been obtained is 82° lat. N., whence the late Arctic Expedition brought back specimens of Charr (Salmo arcturus and Salmo naresii).
The ichthyological features of this zone are well marked: the Chondrosteous Ganoids or Sturgeons, and the families of Salmonidæ and Esocidæ are limited to, and characteristic of, it; Cyprinoids flourish with the Salmonoids, both families preponderating in numbers over the others, whilst the Siluroids are few in number and in variety.
The two regions in which this zone is divided are very closely related to one another, and their affinity is not unlike that which obtains between the sub-regions of the Southern Zone. The subjoined list will show their close agreement with regard to families as well as species. Several of the latter are common to both, viz.—Acipenser sturio, A. maculatus, Perca fluviatilis, Gastrosteus pungitius, Salmo salar, Esox lucius, Lota vulgaris, Petromyzon marinus, P. fluviatilis, and P. branchialis; and all recent investigations have resulted in giving additional evidence of the affinity, and not of the diversity of the two regions.
In Europe and temperate Asia, as well as in North America, mountain ranges elevated beyond the line of perpetual snow would seem to offer physical conditions favourable for the development of a distinct alpine fauna. But this is not the case, because the difference of climate between the mountain districts and the lowlands is much less in this zone than in the Equatorial. Consequently the alpine freshwater fishes do not essentially differ from those of the plains; they are principally Salmonoids; and in Asia, besides, mountain-barbels and Loaches. Salmo orientalis was found by Griffith to abound in the tributaries of the Bamean river at an altitude of about 11,000 feet.
| Europo-Asiatic. | N. American. | |||
| Acipenseridæ— | ||||
| Acipenser | 9 | species. | 12 | species. |
| Scaphirhynchus | 2 | „ | 1 | „ |
| Polyodon | 1 | „ | 1 | „ |
| Lepidosteidæ | 0 | „ | 3 | „ |
| Amiidæ | 0 | „ | 1 | „ |
| Percina [Cosmopol.] | 10 | „ | 30 | „ |
| Grystina [Australia, New Zealand] | 0 | „ | 2 | „ |
| Centrarchina | 0 | „ | 26 | „ |
| Aphredoderidæ | 0 | „ | 1 | „ |
| Cottidæ [partly marine]— | ||||
| Cottus | 3 | „ | 8 | „ |
| Ptyonotus | 0 | „ | 1 | „ |
| Gastrosteidæ | 5 | „ | 5 | „ |
| Comephoridæ | 1 | „ | 0 | „ |
| Gadidæ [marine]— | ||||
| Lota | 1 | „ | 1 | „ |
| Siluridæ— | ||||
| Silurina [India, Africa] | 5 | „ | 0 | „ |
| Bagrina | 2 | „ | 0 | „ |
| Amiurina | 1 | „ | 17 | „ |
| Salmonidæ | 90 | „ | 45 | „ |
| Percopsidæ | 0 | „ | 1 | „ |
| Esocidæ | 1 | „ | 7 | „ |
| Umbridæ | 1 | „ | 1 | „ |
| Cyprinodontidæ Carnivoræ [India, Africa, Neotrop.] | 9 | „ | 30 | „ |
| Heteropygii | 0 | „ | 2 | „ |
| Cyprinidæ— | ||||
| Catostomina | 1 | „ | 25 | „ |
| Cyprinina [India, Africa] | 80 | „ | 30 | „ |
| Leuciscina | 60 | „ | 70 | „ |
| Rhodeina | 10 | „ | 0 | „ |
| Abramidina [India, Africa] | 44 | „ | 10 | „ |
| Cobitidina [India] | 20 | „ | 0 | „ |
| Hyodontidæ | 0 | „ | 1 | „ |
| Petromyzontidæ [Southern Zone] | 4 | „ | 8 | „ |
| 360 | species. | 339 | species. | |
A. The Europo-Asiatic (Palæarctic) Region.—Its western and southern boundaries coincide with those of the Northern Zone, so that only those which divide it from North America have to be indicated. Behring’s Strait and the Kamtschatka Sea have been conventionally taken as the boundary, but this is shown to be artificial by the fact that the animals of both coasts, as far as they are known at present, are not sufficiently distinct to be referred to two distinct regions. As to the freshwater fishes those of North-western America and of Kamtschatka are but imperfectly known, but there can be little doubt that the same agreement exists between them as is the case with other classes of animals. The Japanese islands exhibit a decided Palæarctic fish-fauna, which includes Barbus and Cobitioids, forms strange to the North American fauna. A slight influx of tropical forms is perceived in the south of Japan, where two Bagrina (Pseudobagrus aurantiacus and Liocassis longirostris) have established themselves for a considerable period, for both are peculiar to the island, and have not been found elsewhere.
In the east, as well as in the west, the distinction between the Europo-Asiatic and North American regions disappears almost entirely the farther we advance towards the north. Of four species of the genus Salmo known from Iceland, one (S. salar) is common to both regions, two are European (S. fario and S. alpinus), and one is a peculiarly Icelandic race (S. nivalis). As far as we know the Salmonoids of Greenland and Baffin’s Land they are all most closely allied to European species, though they may be distinguished as local races.
Finally, as we have seen above, the Europo-Asiatic fauna mingles with African and Indian forms in Syria, Persia, and Afghanistan. Capoëta, a Cyprinoid genus, is characteristic of this district, and well represented in the Jordan and rivers of Mesopotamia.
Assuming that the distribution of Cyprinoids has taken its origin from the alpine tract of country dividing the Indian and Palæarctic regions, we find that this type has found in the temperate region as equally favourable conditions for its development as in the tropical. Out of the 360 species known to exist in the Palæarctic regions, no less than 215 are Cyprinoids. In the countries and on the plateaus immediately joining the Himalayan ranges those mountain forms which we mentioned as peculiar to the Indian Alps abound and extend for a considerable distance towards the west and east, mixed with other Cyprinina and Cobitidina. The representatives of these two groups are more numerous in Central and Eastern Asia than in Europe and the northern parts of Asia, where the Leuciscina predominate. Abramidina or Breams are more numerous in the south and east of Asia, but they spread to the extreme north-western and northern limits, to which the Cyprinoid type reaches. The Rhodeina are a small family especially characteristic of the East, but with one or two off-shoots in Central Europe. Very significant is the appearance in China of a species of the Catostomina, a group otherwise limited to North America.
The Cyprinoids, in their dispersal from the south northwards, are met from the opposite direction by the Salmonoids. These fishes are, without doubt, one of the youngest families of Teleostei, for they did not appear before the Pliocene era; they flourished at any rate during the glacial period, and, as is testified by the remnants which we find in isolated elevated positions, like the Trout of the Atlas, of the mountains of Asia-Minor, and of the Hindu Kush, they spread to the extreme south of this region. At the present day they are most numerously represented in its northern temperate parts; towards the south they become scarcer, but increase again in numbers and species, wherever a great elevation offers them the snow-fed waters which they affect. In the rivers of the Mediterranean Salmonoids are by no means scarce, but they prefer the upper courses of those rivers, and do not migrate to the sea.
The Pike, Umbra, several species of Perch and Stickleback, are also clearly autochthont species of this region. Others belong to marine types, and seem to have been retained in fresh water at various epochs: thus the freshwater Cottus (Miller’s Thumb); Cottus quadricornis, which inhabits lakes of Scandinavia, whilst other individuals of the same species are strictly marine; the Burbot (Lota vulgaris); and the singular Comephorus, a dwarfed and much-changed Gadoid which inhabits the greatest depths of Lake Baikal.
Remnants of the Palæichthyic fauna are the Sturgeons and Lampreys. The former inhabit in abundance the great rivers of Eastern Europe and Asia, periodically ascending them from the sea; their southernmost limits are the Yangtse-kiang in the east, and rivers flowing into the Adriatic, Black and Caspian Seas, and Lake Aral, towards the centre of this region. None are known to have gone beyond the boundaries of the Northern Zone. If the Lampreys are justly reckoned among Freshwater fishes, their distribution is unique and exceptional. In the Palæarctic region some of the species descend periodically to the sea, whilst others remain stationary in the rivers; the same has been observed in the Lampreys of North America. They are entirely absent in the Equatorial Zone, but reappear in the Temperate Zone of the Southern Hemisphere. Many points of the organisation of the Cyclostomes indicate that they are a type of great antiquity.
The remaining Palæarctic fishes are clearly immigrants from neighbouring regions: thus Silurus, Macrones, and Pseudobagrus from the Indian region; Amiurus (and, as mentioned above, Catostomus) from North America. The Cyprinodonts are restricted to the southern and warmer parts; all belong to the carnivorous division. The facility with which these fishes accommodate themselves to a sojourn in fresh, brackish, or salt water, and even in thermal springs, renders their general distribution easily comprehensible, but it is impossible to decide to which region they originally belonged; their remains in tertiary deposits round the Mediterranean are not rare.
B. The boundaries of the North American or Nearctic Region have been sufficiently indicated. The main features and the distribution of this fauna are identical with those of the preceding region. The proportion of Cyprinoid species to the total number of North-American fishes (135:339) appears to be considerably less than in the Palæarctic region, but we cannot admit that these figures approach the truth, as the Cyprinoids of North America have been much less studied than those of Europe; of many scarcely more than the name is known. This also applies in a great measure to the Salmonoids, of which only half as many as are found in the Palæarctic region have been sufficiently described to be worthy of consideration. North America will, without doubt, in the end show as many distinct races as Europe and Asia.
Cyprinoids, belonging to genera living as well as extinct, existed in North America in the tertiary period. At present, Cyprinina, Leuciscina, and Abramidina are well represented, but there is no representative of the Old World genus Barbus, or of the Cobitidina[26]; Rhodeina are also absent. On the other hand, a well-marked Cyprinoid type is developed—the Catostomina, of which one species has, as it were, returned into Asia. Very characteristic is the group of Centrarchina, allied to the Perch, of which there are some thirty species; two Grystina. Of the Sticklebacks there are as many species as in Europe, and of Pike not less than seven species have been distinguished. Umbra appears to be as local as in Europe. Some very remarkable forms, types of distinct families, though represented by one or two species only, complete the number of North American autochthont fishes—viz., Aphredoderus, Percopsis, Hyodon, and the Heteropygii (Amblyopsis and Chologaster). The last are allied to the Cyprinodonts, differing from them in some points of the structure of their intestines. The two genera are extremely similar, but Chologaster, which is found in ditches of the rice-fields of South Carolina, is provided with eyes, and lacks the ventral fins. Amblyopsis is the celebrated Blind Fish of the Mammoth Cave of Kentucky: colourless, eyeless, with rudimentary ventral fins, which may be occasionally entirely absent.
A peculiar feature of the North American Fish Fauna is that it has retained, besides the Sturgeons and Lampreys, representatives of two Ganoid families, Lepidosteus and Amia. Both these genera existed in tertiary times: the former occurs in tertiary deposits of Europe as well as North America, whilst fossil remains of Amia have been found in the Western Hemisphere only.
It is difficult to account for the presence of the Amiurina in North America. They form a well-marked division of the Bagrina, which are well represented in Africa and the East Indies, but absent in South America; it is evident, therefore, they should not be regarded as immigrants from the south, as is the case with the Palæarctic Siluroids. Nor, again, has the connection between South and North America been established sufficiently long to admit of the supposition that these Siluroids could have spread in the interval from the south to the northern parts of the continent, for some of the species are found as far north as Pine Islands Lake (54° lat. N.)[27]