The Holocephali and Ganoidei show numerous deviations from the Chondropterygian type, all leading in the direction towards the Teleosteans. As a whole they take an intermediate position between the preceding types and the Teleosteans, but they show a great variation among themselves, and have in common only the imperfect separation of the branchial sacs and the presence of a single outer branchial aperture.
In Chimæra the septum separating the branchial sacs is confluent with the wall of the gill-cavity in a part of its extent only, and still more imperfect is the separation of those branchial divisions in Ceratodus (Fig. [60]). The other Ganoids show no such division whatever. In Chimæra the first gill is incomplete (uniserial), and belongs to the hyoid; then follow three complete gills; the last, belonging to the fourth branchial arch, being again incomplete. Acipenser, Scaphirhynchus, Lepidosiren, Protopterus, and Lepidosteus, possess likewise an anterior incomplete gill (opercular gill), followed by four complete gills in the Sturgeons and Lepidosteus, whilst in Lepidosiren and Protopterus a part of the branchial arches is gill-less. In Polyodon, Ceratodus, and Polypterus, an opercular gill is absent, the two former having four complete gills, the latter three and a half only. Spiracles are still in some Ganoids present, viz. in the Sturgeons and Polypterus. In all the Ganoids an osseous gill-cover is now developed.
In the Teleostei the gills with their supporting branchial arches lie in one undivided cavity; more or less wide clefts between the arches lead from the pharynx to the gills, and a more or less wide opening gives exit to the water after it has washed the gills. The interbranchial clefts have sometimes nearly the same extent as the branchial arches; sometimes they are reduced to small openings, the integuments stretching from one arch to the other. Sometimes there is no cleft behind the fourth arch, in which case this arch has only an uniserial gill developed. The gill-opening likewise varies much in its extent, and when reduced to a foramen may be situated at any place of the posterior boundary of the head. In the Symbranchidæ the gill-openings coalesce into a single narrow slit in the median line of the isthmus. In the majority of Teleosteans the integument of the concave side of the branchial arches develops a series of horny protuberances of various form, the so-called gill-rakers. They are destined to catch any solid corpuscles or substances which would be carried into the gill-cavity with the water. In some fishes they are setiform, and form a complete sieve, whilst in others they are merely rough tubercles, the action of which must be very incomplete if they have any function at all.
Most Teleosteans possess four complete gills, but frequently the fourth arch is provided with an uniserial gill only, as mentioned above, or even entirely gill-less. The most imperfect gills are found in Malthe, which has two and a half gills only, and in Amphipnous cuchia, in which one small gill is fixed to the second arch.
The gills of the Teleosteans as well as of the Ganoids are supported by a series of solid cartilaginous or horny pointed rods, arranged along the convex edges of the branchial arches. Arches bearing a complete gill have two series of those rods, one along each edge; those with uniserial gills bear one row of rods only. The rods are not part of the arch, but fixed in its integument, the several rods of one row corresponding to those of the other, forming pairs (feuillet, Cuvier) (Fig. [59]). Each rod is covered by a loose mucous membrane passing from one rod to its fellow opposite, which again is finely transversely plaited, the general surface being greatly increased by these plaits. In most Teleostei the branchial lamellæ are compressed, and taper towards their free end, but in the Lophobranchs their base is attenuated and the end enlarged. The mucous membrane contains the finest terminations of the vessels, which, being very superficial, impart the blood-red colour to living gills. The Arteria branchialis, the course of which lies in the open canal in the convexity of the branchial arch, emits a branch (a) for every pair of lamellæ which ascends (b) along the inner edge of the lamella, and supplies every one of the transverse plaits with a branchlet. The latter break up into a fine net of capillaries, from which the oxygenised blood is collected into venous branchlets, returning by the venous branch (d), which occupies the outer edge of the lamella.
Fig. 59.—A pair of branchial lamellæ (magnified) of the Perch.
a, Branch of Arteria branchialis; b, Ascending branch of the same; c, Branch of Vena branchialis; d, Descending branch of the same; e, Transverse section through the branchial arch.
The so-called Pseudobranchiæ (Fig. [60]) are the remains of an anterior gill which had respiratory functions during the embryonic life of the individuals. By a change in the circulatory system these organs have lost those functions, and appear in the adult fish as retia mirabilia, as they receive oxygenised blood, which, after having passed through their capillary system, is carried to other parts of the head. In Palæichthyes the pseudobranchia is a rete mirabile caroticum for the brain and eye; in Teleosteans a rete mirabile ophthalmicum only. Pseudobranchiæ are as frequently absent as present in Chondropterygians as well as Teleosteans. As to the Ganoids, they occur in Ceratodus, Acipenser, Polyodon, and Lepidosteus, and are absent in Lepidosiren, Protopterus, Scaphirhynchus, Polypterus, and Amia.
In Chondropterygians and Sturgeons the pseudobranchiæ are situated within the spiracles; in those, in which spiracles have become obliterated, the pseudobranchiæ lie on the suspensorium, hidden below cellular tissue; but pseudobranchiæ are not necessarily co-existent with spiracles. In the other Ganoids and Teleosteans the pseudobranchiæ (Fig. [60], h) are within the gill-cavity, near the base of the gill-cover; in Ceratodus even rudiments of the gill-rakers (x’, x”) belonging to this embryonic gill are preserved, part of them (x”) being attached to the hyoid arch. Pseudobranchiæ are frequently hidden below the integuments of the gill-cavity, and have the appearance of a glandular body rather than of a gill.