While our knowledge of the osteology and embryology of most of the families of fishes is very incomplete, it is evident that the relationships of the groups cannot be shown in any linear series or by any conceivable arrangement of orders and suborders. The living teleost fishes have sprung from many lines of descent, their relationships are extremely diverse, and their differences are of every possible degree of value. The ordinary schemes have magnified the value of a few common characters, at the same time neglecting other differences of equal value. No system of arrangement which throws these fishes into large groups can ever be definite or permanent.
Relationships of Isospondyli.—For our purposes we may divide the physostomous fishes as understood by Müller into several orders, the most primitive, the most generalized, and economically the most important being the order of Isospondyli. This order contains those bony fishes which have the anterior vertebræ unaltered (as distinguished from the Ostariophysi), the skull relatively complete, or at least not eel-like, the mesocoracoid typically developed, but atrophied in deep-sea forms and finally lost, the orbitosphenoid present. In all the species the ventral fins are abdominal and normally composed of more than six rays; the air-duct is developed. The scales are chiefly cycloid and the fins are without true spines. In many ways the order is more primitive than Nematognathi, Plectospondyli, or Apodes. It is certain that it began earlier in geological time than any of these. On the other hand, the Isospondyli are closely connected through the Berycoidei with the highly specialized fishes. The continuity of the natural series is therefore interrupted by the interposition of the side branches of Ostariophysans and eels before considering the Haplomi and the other transitional forms. The forms called Iniomi, which lack the mesocoracoid and the orbitosphenoid, have been lately transferred to the Haplomi by Boulenger. This arrangement is probably a step in advance.
Ganoid traits are present in certain families of Isospondyli. Among these are the gular plate (found in Amia and the Elopidæ), doubtless derived from the similar structure in earlier Ganoids; additional valves in the arterial bulb in the cellular air-bladder of Notopterus and Osteoglossum, the spiral intestinal valve in Chirocentridæ, and the ganoid scales of the extinct Leptolepidæ.
The Clupeoidea.—The Isospondyli are divisible into numerous families, which may be grouped roughly under three subdivisions, Clupeoidea, the herring-like forms; the Salmonoidea, the trout-like forms; and the Iniomi, or lantern-fishes, and their allies. The last-named group should probably be removed from the order of Isospondyli. In the Clupeoidea, the allies of the great family of the herring, the shoulder-girdle is normally developed, retaining the mesocoracoid arch on its inner edge, and through the post-temporal is articulated above with the cranium. The fishes in this group lack the adipose fin which is characteristic of most of the higher or salmon-like families.
Fig. 27.—Leptolepis dubius Blainville, Lithographic Stone. (After Woodward.)
The Leptolepidæ.—Most primitive of the Isospondyli is the extinct family of Leptolepidæ, closely allied to the Ganoid families of Pholidophoridæ and Oligopleuridæ. It is composed of graceful, herring-like fishes, with the bones of the head thin but covered with enamel, and the scales thin but firm and enameled on their free portion. There are no fulcra and there is no lateral line. The vertebræ are well developed, but always pierced by the notochord. The genera are Lycoptera, Leptolepis, Æthalion, and Thrissops. In Lycoptera of the Jurassic of China the vertebral centra are feebly developed, and the dorsal fin short and posterior. In Leptolepis the anal is short and placed behind the dorsal. There are many species, mostly from the Triassic and lithographic shales of Europe, one being found in the Cretaceous. Leptolepis coryphænoides and Leptolepis dubius are among the more common species. Æthalion (knorri) differs in the form of the jaws. In Thrissops the anal fin is long and opposite the dorsal. Thrissops salmonea is found in the lithographic stone; Thrissops exigua in the Cretaceous. In all these early forms there is a hard casque over the brain-cavity, as in the living types, Amia and Osteoglossum.
Fig. 28.—Ten-pounder, Elops saurus L. An ally of the earliest bony fishes. Virginia.