many being close allies of the present inhabitants of the deep sea. At this period the spiny-rayed Teleosteans, dominant in the seas of the present day, made their first appearance.
With the Eocene, the fish-fauna has assumed the essential character which it now bears. A few Pycnodonts survive as the last representatives of typically Mesozoic ganoids, whilst in the marine deposits of Monte Bolca (Upper Eocene) the principal families of living marine fishes are represented by genera identical with or more or less closely allied to those still existing; it is highly remarkable that forms so highly specialized as the sucking-fish or remoras, the flat-fish (Pleuronectidae), the Pediculati, the Plectognaths, &c., were in existence, whilst in the freshwater deposits of North America Osteoglossidae and Cichlidae were already represented. Very little is known of the freshwater fishes of the early Tertiaries. What has been preserved of them from the Oligocene and Miocene shows that they differed very slightly from their modern representatives. We may conclude that from early Tertiary times fishes were practically as they are at present. The great hiatus in our knowledge lies in the period between the Cretaceous and the Eocene.
At the present day the Teleosteans are in immense preponderance, Selachians are still well represented, the Chondrostean ganoids are confined to the rivers and lakes of the temperate zone of the northern hemisphere (Acipenseridae, Polyodontidae), the Holostean ganoids are reduced to a few species (Lepidosteus, Amia) dwelling in the fresh waters of North America, Mexico and Cuba, the Crossopterygians are represented by the isolated group Polypteridae, widely different from any of the known fossil forms, with about ten species inhabiting the rivers and lakes of Africa, whilst the Dipneusti linger in Australia (Neoceratodus), in South America (Lepidosiren), and in tropical Africa (Protopterus). The imperfections of the geological record preclude any attempt to deal with the distribution in space as regards extinct forms, but several types, at present very restricted in their habitat, once had a very wide distribution. The Ceratodontidae, for instance, of which only one species is now living, confined to the rivers of Queensland, has left remains in Triassic, Rhaetic, Jurassic and Cretaceous rocks of Europe, North America, Patagonia, North and South Africa, India and Australia; the Amiidae and Lepidosteidae were abundant in Europe in Eocene and Miocene times; the Osteoglossidae, now living in Africa, S.E. Asia and South America, occurred in North America and Europe in the Eocene.
In treating of the geographical distribution of modern fishes, it is necessary to distinguish between fresh-water and marine forms. It is, however, not easy to draw a line between these categories, as a large number of forms are able to accommodate themselves to either fresh or salt water, whilst some periodically migrate from the one into the other. On the whole, fishes may be roughly divided into the following categories:—
I. Marine fishes. A. shore-fishes; B. pelagic fishes; C. deep-sea fishes.
II. Brackish-water fishes.
III. Fresh-water fishes.
IV. Migratory fishes. A. anadromous (ascending fresh waters to spawn); B. catadromous (descending to the sea to spawn).
About two-thirds of the known recent fishes are marine. Such are nearly all the Selachians, and, among the Teleosteans, all the Heteromi, Pediculati and the great majority of Apodes, Thoracostei, Percesoces, Anacanthini, Acanthopterygii and Plectognathi. All the Crossopterygii, Dipneusti, Opisthomi, Symbranchii, and nearly all the Ganoidei and Ostariophysi are confined to fresh-water.
The three categories of marine fishes have thus been defined by Günther:—
“1. Shore Fishes—that is, fishes which chiefly inhabit parts of the sea in the immediate neighbourhood of land either actually raised above, or at least but little submerged below, the surface of the water. They do not descend to any great depth,—very few to 300 fathoms, and the majority live close to the surface. The distribution of these fishes is determined, not only by the temperature of the surface water, but also by the nature of the adjacent land and its animal and vegetable products,—some being confined to flat coasts with soft or sandy bottoms, others to rocky and fissured coasts, others to living coral formations. If it were not for the frequent mechanical and involuntary removals to which these fishes are exposed, their distribution within certain limits, as it no doubt originally existed, would resemble still more that of freshwater fishes than we find it actually does at the present period.
2. Pelagic Fishes—that is, fishes which inhabit the surface and uppermost strata of the open ocean, and approach the shores only accidentally or occasionally (in search of prey), or periodically (for the purpose of spawning). The majority spawn in the open sea, their ova and young being always found at a great distance from the shore. With regard to their distribution, they are still subject to the influences of light and the temperature of the surface water; but they are independent of the variable local conditions which tie the shore fish to its original home, and therefore roam freely over a space which would take a freshwater or shore fish thousands of years to cover in its gradual dispersal. Such as are devoid of rapidity of motion are dispersed over similarly large areas by the oceanic currents, more slowly than the strong swimmers, but not less surely. An accurate definition, therefore, of their distribution within certain areas equivalent to the terrestrial regions is much less feasible than in the case of shore fishes.
3. Deep-Sea Fishes—that is, fishes which inhabit such depths of the ocean that they are but little or not at all influenced by light or the surface temperature, and which, by their organization, are prevented from reaching the surface stratum in a healthy condition. Living almost under identical tellurian conditions, the same type, the same species, may inhabit an abyssal depth under the equator as well as one near the arctic or antarctic circle; and all that we know of these fishes points to the conclusion that no separate horizontal regions can be distinguished in the abyssal fauna, and that no division into bathymetrical strata can be attempted on the base of generic much less of family characters.”
A division of the world into regions according to the distribution of the shore-fishes is a much more difficult task than that of tracing continental areas. It is possible perhaps to distinguish four great divisions: the Arctic region, the Atlantic region, the Indo-Pacific region and the Antarctic region. The second and third may be again subdivided into three zones: Northern, Tropical and Southern. This appears to be a more satisfactory arrangement than that which has been proposed into three zones primarily, each again subdivided according to the different oceans. Perhaps a better division is that adopted by D. S. Jordan, who arranges the littoral fishes according to coast lines; we then have an East Atlantic area, a West Atlantic, an East Pacific and a West Pacific, the latter including the coasts of the Indian Ocean. The tropical zone, whatever be the ocean, is that in which fishes flourish in greatest abundance and where, especially about coral-reefs, they show the greatest variety of bizarre forms and the most gorgeous coloration. The fish-fauna of the Indo-Pacific is much richer than that of the Atlantic, both as regards genera and species.
As regards the Arctic and Antarctic regions, the continuity or circumpolar distribution of the shore fishes is well established. The former is chiefly characterized by its Cottids, Cyclopterids, Zoarcids and Gadids, the latter by its Nototheniids. The theory of bipolarity receives no support from the study of the fishes.
Pelagic fishes, among which we find the largest Selachians and Teleosteans, are far less limited in their distribution, which, for many species, is nearly world-wide. Some are dependent upon currents, but the great majority being rapid swimmers able to continue their course for weeks, apparently without the necessity of rest (many sharks, scombrids, sword-fishes), pass from one ocean into the other. Most numerous between the tropics, many of these fishes occasionally wander far north and south of their habitual range, and there are few genera that are at all limited in their distribution.
Deep-sea fishes, of which between seven hundred and eight hundred species are known, belong to the most diverse groups and quite a number of families are exclusively bathybial (Chlamydoselachidae, Stomiatidae, Alepocephalidae, Nemichthyidae, Synaphobranchidae, Saccopharyngidae, Cetomimidae, Halosauridae, Lipogenyidae, Notacanthidae, Chiasmodontidae, Icosteidae, Muraenolepididae, Macruridae, Anomalopidae, Podatelidae, Trachypteridae, Lophotidae, Ceratiidae, Gigantactinidae). But they are all comparatively slight modifications of the forms living on the surface of the sea or in the shallow parts, from
