I. Fins fringed with fulcra, or scales coated with ganoin; notochord usually continuous through the vertebrae.

Vertebral centra not more than rings; fins with fulcra; scales rhombic, united by peg-and-socket joints .......... 1. Pholidophoridae.†

Vertebral centra not more than rings; fins with fulcra; scales cycloid .......... 2. Archaeomaenidae.†

Vertebral centra complete or with minute perforation; fins with fulcra; scales cycloid .......... 3. Oligopleuridae.†

Vertebral centra nearly complete, but with perforation; no fulcra; scales cycloid .......... 4. Leptolepididae.†

II. Fins without fulcra; scales without ganoin; vertebral centra complete.

A. Supraoccipital separated from the frontals by the parietals or upper surface of skull; no adipose fin.

1. Ventral fins with 10 to 16 rays.

An intergular bone; parasphenoid narrow .......... 5. Elopidae.

No intergular bone; parasphenoid very broad .......... 6. Albulidae.

2. Ventrals with not more than 7 rays.

a. Supratemporal very large, plate-like.

Praemaxillary single, its posterior extremity free from the maxillary; symplectic absent; basis cranii simple .......... 7. Mormyridae.

Praemaxillary paired, its posterior extremity firmly attached to the maxillary; symplectic present; basis cranii double .......... 8. Hyodontidae.

b. Supratemporal small; maxillary firmly attached to posterior extremity of praemaxillary.

Praemaxillary paired; a large hole on each side of the skull, between the post-frontal and the squamosal; basis cranii double; suboperculum absent; ribs sessile .......... 9. Notopteridae.

Praemaxillary paired; basis cranii simple; suboperculum reduced; ribs inserted on parapophyses .......... 10. Osteoglossidae.

Praemaxillary single; basis cranii simple; suboperculum and interoperculum absent; ribs inserted on parapophyses .......... 11. Pantodontidae.

c. Supratemporal small; maxillary movable; ribs sessile; ventral fins below the pectorals .......... 12. Ctenothrissidae.†

B. Supraoccipital separating parietals, or adipose fin present.

1. Interoperculum enormous; symplectic absent; basis cranii simple .......... 13. Phractolaemidae.

2. Interoperculum normal; symplectic present; basis cranii double.

a. Teeth in sockets; maxillary firmly attached to praemaxillary. Symplectic exposed .......... 14. Saurodontidae.†

b. Teeth not in sockets.

Symplectic concealed between the quadrate and the hyomandibular .......... 15. Chirocentridae.

Postclavicle on outer side of clavicle; no adipose fin .......... 16. Clupeidae.

Postclavicle on inner side of clavicle; an adipose dorsal fin .......... 17. Salmonidae.

Postclavicle absent; no adipose fin .......... 18. Alepocephalidae.

3. Interoperculum normal; basis cranii simple.

Maxillary large, toothed; praecaudal vertebrae without well-marked parapophyses; scales cycloid or absent; adipose dorsal fin present or absent .......... 19. Stomiatidae.

Mouth small, toothless; vertebrae with strong parapophyses; head and body covered with spiny scales .......... 20. Gonorhynchidae.

Mouth small, toothless; no symplectic; head and body naked .......... 21. Cromeriidae.

† This sign indicates that the group is represented by fossil forms only.

Fam. 1. Pholidophoridae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter large and loosely attached; teeth small and conical. Parietal bones separating the supraoccipital from the frontals; opercular bones well developed. Vertebral centra never advanced beyond the annular stage; ribs delicate; no fused or expanded haemal arches at the base of the tail; no epipleurals or epineurals. Fin-fulcra present, but usually small; dorsal and anal fins small, the former above or behind the ventral fins, which are small or may be absent. Scales ganoid, rhombic, those on the flanks united by peg-and-socket joints.

This family, which appears to me to be related to the Dapediidae, is provisionally placed here by A. S. Woodward on account of its resemblance to the Leptolepididae, but it is not yet quite clear that the mandible was destitute of splenial and coronoid elements, while the bones at the base of the pectoral fin have not hitherto been observed. The principal genera are Pholidophorus, ranging from the Upper Trias to the Purbeck; Thoracopterus, from the Upper Trias; and Pleuropholis, from the Upper Jurassic. The species of Pholidophorus are very numerous in the Jurassic period, and Woodward has observed that the scales of the later species are more elaborately ornamented than those of earlier date.

Fam. 2. Archaeomaenidae.—Distinguished from the preceding by the thin, cycloid scales. Conspicuous obtuse ridge-scales are present along the dorsal and ventral lines. Archaeomenes, from the Jurassic (?) of New South Wales.

Fam. 3. Oligopleuridae.—Characters as in Pholidophoridae, but scales cycloid and vertebrae completely or nearly completely ossified.

Oligopleurus, from the Jurassic of England and France; Oenoscopus, from the Jurassic and Cretaceous of France, Germany, and Italy; and Spathiurus, from the Cretaceous of Mount Lebanon and Dalmatia.

Fam. 4. Leptolepididae.—Praemaxillaries very small; maxillaries large, loosely attached; teeth small and conical. Parietal bones separating the supraoccipital from the frontals; opercular bones well developed. Vertebral centra well ossified, but always pierced by the notochord; ribs delicate; epipleurals present; no fused or expanded haemal arches at the base of the caudal fin. Dorsal and anal fins small, the former above or behind the ventrals. Ventrals with 5 to 10 rays. Scales thin, cycloid and deeply imbricate, usually coated with ganoin in their exposed portion.

fig325

Fig. 325.Leptolepis dubius. (Restoration of skeleton by A. S. Woodward.)

Leptolepis, with numerous species, from the Jurassic and Cretaceous of Europe and New South Wales; Vidalia, Jurassic of France; Aethalion, Jurassic of Bavaria; Thrissops, Jurassic and Cretaceous of Europe; and Lycoptera, Jurassic of Asia.

Fam. 5. Elopidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter the more developed, and movably articulated above the former to the ethmoid. Parietal bones in contact behind the frontals; opercular bones well developed. Basis cranii double. A bony intergular or sublingual plate. Jaws, palatines, pterygoids, vomer, parasphenoid, glossohyal, and pharyngeals toothed. Ribs mostly sessile, inserted very low down, behind parapophyses; epineurals similar to the ribs, but directed upwards. Pectorals low down, folding like the ventrals. Post-temporal forked, the upper branch attached to the epiotic, the lower to the opisthotic; post-clavicle small; scapular foramen in the scapula; pterygials well developed, three in contact with coracoid. Ventrals with 10 to 16 rays. Branchiostegal rays very numerous (over 20). Air-bladder large.

This family is abundantly represented in Cretaceous times by the genera Osmeroides and Elopopsis, and from the Lower Eocene to the present day by Elops and Megalops. Elops saurus is a handsome elongate silvery Fish, found in all the warm and tropical seas; the young are ribbon-shaped like those of Albula. A second species, E. lacerta, is from the West Coast of Africa, entering rivers. Megalops, distinguished by larger scales, the absence of pseudobranchiae, and the curious prolongation of the last ray of the dorsal fin, includes the well-known Tarpon M. atlanticus, and the Indian M. cyprinoides. The Tarpon occurs from the south-eastern coasts of North America and the West Indies to Brazil, and reaches a length of 6 feet and a weight of 110 lbs. It often leaps out of the water, after the manner of Grey Mullets, and its chase when hooked affords good sport, the landing of so active a giant being attended with great difficulties. Its remarkably large scales, over two inches in diameter, are much prized for fancy work in the Florida curiosity shops.

fig326

Fig. 326.—Tarpon, Megalops atlanticus, much reduced. (After Goode.)

Fam. 6. Albulidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter movably articulated above the former to the ethmoid. Parietal bones separating the supraoccipital from the frontals; suboperculum large; interoperculum small and entirely or nearly entirely hidden below the praeoperculum. Basis cranii double. Praemaxillaries, mandible, vomer, palatines, pterygoid, parasphenoid, glossohyal, and pharyngeals toothed. Ribs sessile, inserted behind and somewhat below small parapophyses, which are absent or merely indicated on the anterior vertebrae, and gradually increase in size towards the caudal region; these parapophyses, as well as the neural and haemal arches, are autogenous bones; epineurals, no epipleurals. Pectorals low down, folding like the ventrals. Post-temporal trifid, the upper branch attached to the epiotic, the median to the squamosal, the lower to the opisthotic; post-clavicle large (formed of three pieces); scapular foramen between scapula and clavicle; pterygials well developed, two in contact with coracoid. Ventrals with 10 to 14 rays. Branchiostegal rays 6 to 14. Air-bladder large, not communicating with the ear.

Elongate fusiform Fishes, covered with large silvery scales forming regular longitudinal series; head naked; mouth rather small, with thick lips; gill-openings wide. Dorsal fin originating in front of ventrals; anal short; caudal well developed, forked.

fig327

Fig. 327.Albula conorhynchus. ¼ nat. size.

The type of this family, Albula or Butirinus, is remarkable among Teleosts in possessing a rudiment of a conus arteriosus to the heart, provided with two rows of valvules.[642] Its single species inhabits all the warm and tropical seas. Prof. Gilbert has ascertained that the young pass through a metamorphosis, analogous to that of the Eels. The "Leptocephalid" described as Esunculus by Kaup is probably a larval Albula.

fig328

Fig. 328.—Larva of Albula conorhynchus. (After Gilbert.)

The deep-sea Japanese Pterothrissus (Bathythrissa) must be placed in this family; its skeleton is very similar to that of Albula. It differs in the elongate dorsal fin, in the presence of small teeth on the maxillary bone, and in the small number of branchiostegal rays (6 instead of 12 to 14).

Albula is represented in the Eocene (London Clay and Bruxellian); and the Cretaceous Istieus and Anogmius are believed to be possibly related to Pterothrissus. Four Cretaceous types (Plethodus, Thryptodus, Syntegmodus, and Ancylostylus) are referred with doubt to the Albulidae.

Fam. 7. Mormyridae.—Margin of the upper jaw formed by the single praemaxillary and the maxillaries, the latter articulated above the former to the ethmoid. Parietal bones separating the supraoccipital from the frontals; a large hole on each side of the skull, between the squamosal, the epiotic, and the opisthotic, covered by a large, thin, bony plate (the supratemporal), which may extend over a part of the parietal; symplectic absent; suboperculum small and hidden under the operculum, or absent; interoperculum well developed. Basis cranii simple. No pharyngeal teeth. Opercular bones hidden under the skin; gill-clefts narrow. Anterior ribs sessile; epineurals, no epipleurals. Pectorals directed upwards. Ventrals with 6 or 7 rays. Air-bladder communicating with the ear.

fig329

Fig. 329.Mormyrus caballus. ⅕ nat. size.

Curious-looking Fishes, very variable in the form of the head and body and in the extent of the fins. Mouth often very small; teeth in jaws usually few; teeth usually present on the parasphenoid, working against a similar patch on the glossohyal; eye covered over by skin, sometimes very indistinct; scales small, cycloid; branchiostegal rays 4 to 8. The dorsal and anal fins may be nearly equally developed (Genyomyrus, Gnathonemus); or the former (Mormyrus) or the latter (Hyperopisus) are several times the longer. Gymnarchus, Eel-shaped, apodal, and deprived of the caudal fin, lacks the anal fin, the dorsal extending over the whole extent of the body. Some species of Mormyrops show how a form like Gymnarchus may have been evolved out of a more typically-formed Fish. Nothing is more striking than the variation in shape of the snout within one and the same genus, and the names given to some of the species (ovis, caballus, elephas, tamandua, numenius, ibis) are suggestive of resemblances with the heads of various animals.

fig330

Fig. 330.—Head of Gnathonemus curvirostris.

fig331

Fig. 331.—Head of Gnathonemus numenius.

The Mormyrids are highly remarkable for the enormous development of the brain, the weight of which equals 152 to 182 of the total, a thing unparalleled among lower Vertebrates; and for the problematic organ which surmounts it; also as being among the few Fishes in which an electric organ has been discovered. The organ, situated on each side of the caudal region, is derived from the muscular system and is of feeble power, as ascertained by Babuchin and by Fritsch; it was long considered as "pseudo-electric." The natural affinities of this family appear to be with the Albulidae, and there is nothing to justify the term "Nilhechte" (Nile-pike) which has been bestowed on them by German authors. Ninety-three species are known from the fresh waters of Africa north of the Tropic of Capricorn, and may be referred to two sub-families and ten genera[643]:—

(i.) Mormyrinae, with teeth on the parasphenoid and tongue, with ventral, anal, and caudal fins, and a simple air-bladder; vertebrae 37 to 64; peculiar (Gemmingerian) linear bones, without known homologues, along each side of the tail, above and beneath the electric organ; scapular foramen in the scapula, or between the scapula and the coracoid. Mormyrops, Petrocephalus, Isichthys, Marcusenius, Stomatorhinus, Myomyrus, Gnathonemus, Genyomyrus, Mormyrus.

(ii.) Gymnarchinae, without teeth on the parasphenoid and tongue, without ventral, anal, or caudal fins, and with a cellular air-bladder; vertebrae about 120; Gemmingerian bones absent; scapular foramen in the coracoid. Gymnarchus.

Fossil Mormyrids are unknown.

Venerated by the ancient Egyptians, the Mormyrs of the Nile are frequently represented on hieroglyphics and mural paintings as well as in bronze models. Very little is known of the habits of these Fishes. Prof. G. Fritsch, of Berlin, during his stay in Egypt for the purpose of experimenting on electric Fishes, observed that they perish very rapidly when removed from the river, and he had the greatest difficulty in keeping some alive in an aquarium for two or three days. The species with comparatively large mouths (Mormyrops, Gymnarchus) feed principally on fishes and crustaceans, the others on tiny animals and vegetable and more or less decomposed matter. Delhez, on the Congo, found that many are attracted to the borders of the river in the neighbourhood of human dwellings, where they feed on the refuse thrown into the water. It is probable that the species with a rostrum use it to procure small prey hidden between stones or buried in the mud, and that the fleshy mental appendage with which many are provided is a tactile organ compensating the imperfection of the vision in the search for food. A small Mormyrid from the Congo (Stomatorhinus microps) has the eyes so much reduced and the skin so feebly pigmented as to convey the impression of a cave Fish. Until quite recently, absolutely nothing was known of the breeding habits and development in this important family. To the late J. S. Budgett we owe some very interesting observations made in the Gambia on Gymnarchus niloticus.[644] The Fish makes a floating nest, emerging on three sides, over which the male keeps a fierce watch; the recently-hatched larvae are remarkable for the enormous size of the yolk-sac, which hangs down, acting as a sort of anchor, and for the presence of long external branchial filaments, as in Selachian embryos. The Fish propels itself through the water entirely by the action of its dorsal fin, forwards and backwards with equal facility; when swimming rapidly backwards, it may be seen to use the end of its tail as a feeler to guide the way. Budgett has also identified, with some doubt, the eggs of Hyperopisus bebe, out of which emerged embryos not unlike those of some tailless Batrachians, which hung suspended to rootlets of grass in swamps by means of threads of viscid mucus secreted from glands on the top of the head.

fig332

Fig. 332.Gymnarchus niloticus. ¼ nat. size.

Fam. 8. Hyodontidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter the more developed and firmly united to the end of the former. Parietal bones separating the supraoccipital from the frontals; a large hole on each side of the skull, between the parietal, the squamosal, and the epiotic (paroccipital), closed by a large, thin, bony plate (the supratemporal), which extends over the greater part of the parietal; suboperculum and interoperculum small, the latter partly hidden below the praeoperculum. Basis cranii double. Jaws, palatines, pterygoids, vomer, parasphenoid, and glossohyal toothed; no pharyngeal teeth. Ribs sessile, inserted above and behind well-developed parapophyses; epineurals, no epipleurals. Pectorals low down, folding like the ventrals. Post-temporal forked; the upper branch attached to the epiotic, the lower to the squamosal; no post-clavicle; coracoids forming together a ventral keel; scapular foramen between scapula and clavicle; pterygials well developed, three in contact with coracoid. Ventrals with 7 rays. Branchiostegal rays in moderate number (8 to 10). Air-bladder communicating with the ear. No oviducts, the eggs falling into the abdominal cavity before exclusion.

fig333

Fig. 333.—Upper (A) and posterior (B) views of skull and pectoral arch of Hyodon alosoides (the supratemporal removed on the left side). bo, Basioccipital; cl, clavicle; cor, coracoid; eo, exoccipital; eot, epiotic; eth, ethmoid; f, frontal; m, maxilla; mcor, mesocoracoid; n, nasal; oo, opisthotic; p, parietal; pcl, postclavicle; pm, praemaxilla; por, praeorbital; ptf, postfrontal; ptr, pterygials; ptte, post-temporal; scl, supraclavicle; so, supraoccipital; sor, suborbital; sq, squamosal; ste, supratemporal.

Elongate, compressed, silvery Fishes, covered with moderate-sized cycloid scales; head naked; mouth large, with strong dentition; gill-openings wide; dorsal fin short, posterior to the ventrals; anal rather elongate; caudal well developed, forked.

A single genus (Hyodon) with three species (Moon-Eyes of the Americans), all reaching the length of about a foot, inhabiting the fresh waters of North America east of the Rocky Mountains.

Fam. 9. Notopteridae. The Fishes which form this family may be regarded as an eccentric modification of a type very similar to the preceding, with which they agree in most osteological features as well as in the dentition, in the connexion between the air-bladder and the ear, and in the absence of oviducts. They differ in the absence of the lid-like supratemporal, the hole which it covers in Hyodon being here bordered by the post-frontal and the squamosal (fused with the opisthotic), sometimes also by the epiotic, in the absence of the suboperculum, in the absence or incomplete ossification of the upper branch of the post-temporal (the lower articulating with the opisthotic), and in the presence of accessory bones (named adpleurals) attached to or fused with the distal extremity of the anterior ribs, which they prolong to the mid-ventral line, where they are embraced by dermal ossifications forming a doubly serrated crest.

fig334

Fig. 334.Notopterus afer, skeleton, with outline of soft parts. ⅔ nat. size.

The bones of the head are cavernous, the mouth is large; the anterior nostril is produced into a tentacle. The body is very strongly compressed, with very short precaudal region, attenuate behind; the ventral fins are much reduced or absent; the dorsal is short or absent, and the anal is much elongate and confluent with the caudal, which may be regarded as aborted. The scapular foramen is entirely in the scapula. The air-bladder is very large, with several divisions, forked in front and behind, and prolonged along each side of the caudal region; its extraordinary condition has been described by Bridge.[645]

These Fishes live in marshes and lakes, fresh-water or brackish, and feed on worms and insects. Nothing is known of their breeding habits and development.

Two genera: Notopterus, with a dorsal fin and 6 to 9 branchiostegal rays—three species from India, Burma, and the Malay region, and one from West Africa; Xenomystus, without dorsal fin and with only 3 branchiostegal rays, the unique species inhabiting the White Nile and West Africa. Remains of Notopterus have been found in the marl slates (Tertiary) of Padang, Sumatra. The largest species, the Indian N. chitala, attains 4 feet in length; its flesh is said to be uncommonly rich and well flavoured, but a strong prejudice exists against it, owing to the Fish being supposed to live on human carcases.

Fam. 10. Osteoglossidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter the more developed and firmly attached to the end of the former. Parietal bones separating the supraoccipital from the frontals; suboperculum much reduced, and entirely or partially concealed under the praeoperculum. Basis cranii simple. Teeth in jaws, and on pterygoid and hyoid bones; no pharyngeal teeth. Head scaleless, the thin skin confluent with the bones; body covered with large bony scales, composed of pieces like mosaic. Ribs inserted on the strong parapophyses; epineurals, no epipleurals. Pectoral fins low down. Post-temporal forked, the upper branch attached to the epiotic, the lower to the squamosal; post-clavicle present; scapular foramen in scapula; pterygials well developed, only one in contact with coracoid. Dorsal and anal fins originating behind the ventrals; latter with 5 or 6 rays. No oviducts, the eggs falling into the abdominal cavity before exclusion (at least in Heterotis, as observed by Budgett).

This family is represented at the present day by five species, referred to four genera; thus characterised:—

fig335

Fig. 335.—Principal forms of Osteoglossids. A, Dapedoglossus testis (restoration); B, Scleropages leichardti; C, Osteoglossum bicirrhosum; D, Arapaima gigas; E, Heterotis niloticus. All much reduced.

Scleropages.—Mouth large; vomer, palatines, pterygoids, and glossohyal toothed; mandibular barbels; branchiostegal rays 15 to 17; body compressed, with trenchant abdomen; coracoids forming a ventral keel; dorsal fin short; ventral fins nearly equally distant from end of snout and caudal fin; vertebrae 29 to 31 + 30; air-bladder not cellular. One species from the northern parts of Australia, and one from Sumatra, Banka, and Borneo.

Osteoglossum.—Mouth large; vomer, palatines, pterygoids, and glossohyal toothed; mandibular barbels; branchiostegal rays 10; body compressed, with trenchant abdomen; coracoids forming a ventral keel; dorsal fin long; ventral fins nearly twice as far from the caudal as from the end of the snout; vertebrae 28 + 59; air-bladder not cellular.—South America (Guianas, Brazil).

Arapaima.—Mouth rather large; vomer, palatines, pterygoids, and glossohyal toothed; branchiostegal rays 16; belly rounded; dorsal fin rather long; ventral fins equidistant from head and caudal fin; vertebrae 36 to 38 + 41 to 42; air-bladder cellular.—South America (Guianas, Brazil).

Heterotis.—Mouth moderate; branchiostegal rays 7; belly rounded; dorsal fin rather long; ventral fins nearer end of snout than caudal fin; vertebrae 27 + 42 to 43; air-bladder cellular; fourth branchial arch with an accessory breathing-organ. Africa (Nile, Senegal, Gambia, Niger).

Dapedoglossus, from the Eocene of Wyoming, appears to be nearest to Scleropages, and Brychaetus, from the Eocene (London Clay) of Sheppey, Kent, to Arapaima, so far as the state of preservation of these fossils enables us to form an opinion.

fig336

Fig. 336.—Distribution of the Osteoglossids.

Dr. Günther has directed attention to the remarkable coincidence of the geographical distribution of this family and the Dipneusti, although, however, the latter are not known to be represented in the Malay Archipelago. "Not only," he adds, "are the corresponding species found within the same region, but also in the same river systems; and although such a connexion may and must be partly due to a similarity of habit, yet the identity of this singular distribution is so striking that it can only be accounted for by assuming that the Osteoglossidae are one of the earliest Teleosteous types which have been contemporaries of and have accompanied the present Dipnoi since or even before the beginning of the Tertiary epoch."

The Queensland species of Scleropages (S. leichardti) is known to the settlers by the name of Barramunda, which has also been applied to Neoceratodus. Arapaima gigas is one of the largest fresh-water Fishes known, exceeding a length of 15 feet and a weight of 400 pounds. Its flesh is much valued. Sir R. Schomburgh has observed that the mother protects the young, who, for some time after their birth, always swim in front of her. A similar observation has been made in the Gambia on Heterotis niloticus by the late J. S. Budgett, who states that the Fish builds enormous nests in swamps, in about two feet of water; the walls of the nest are made of the stems of the grasses removed by the Fish from the centre; the floor is the swamp-bottom, and is made perfectly smooth and bare. The nest appears to be used for at most four or five days; the newly-hatched larvae are provided with long external gill-filaments of a blood-red colour.[646]

Fam. 11. Pantodontidae.—The little West African Fish described by Peters as Pantodon buchholzi is the unique representative of a family closely related to the Osteoglossidae, but distinguished by the very small, single praemaxillary and the absence of suboperculum and interoperculum. The pectoral fins are very large and are remarkable for the fleshy process to which the inner ray is adnate; the ventrals, formed of 7 rays, some of which are simple and prolonged into filaments, are placed more forward than in any other type of this sub-order, the Ctenothrissidae excepted, viz. immediately behind the pectorals. Teeth in the jaws and on the vomer, palatines, pterygoids, parasphenoid, glossohyal, and pharyngeal bones. Mesocoracoid arch slender, strongly curved, and meeting its fellow on the median line; coracoids forming a ventral keel. Vertebrae few (16 + 14).

fig337

Fig. 337.Pantodon buchholzi, natural size.

Observed by M. de Brazza to be a freshwater Flying-Fish.

fig338

Fig. 338.Ctenothrissa vexillifer (restored by A. S. Woodward).

Fam. 12. Ctenothrissidae.—A curious type characterised by small praemaxillaries, large maxillaries, with feeble dentition, the parietals in contact on the median line, vertebral centra without transverse processes, a moderately large dorsal with simple anterior rays, and large ventrals advanced far forwards and formed of 8 rays. Its affinities are still obscure, but the condition of the jaws decides its allocation to the suborder Malacopterygii, whilst in the position of the ventrals it is most nearly approached by the Pantodontidae. Small Fishes known only by two genera, of the Cretaceous period (England and Mount Lebanon), one with ctenoid scales (Ctenothrissa), the other with cycloid scales (Aulolepis).

Fam. 13. Phractolaemidae.—Mouth edentulous, projectile, bordered by the very slender praemaxillaries and maxillaries. Supraoccipital in contact with the frontals, widely separating the small parietals; operculum and suboperculum well developed; praeoperculum small; interoperculum enormous, covering the gular region and overlapping its fellow; symplectic absent. Basis cranii single. No pharyngeal teeth. Only 3 slender branchiostegal rays. Ribs stout, sessile, nearly completely encircling the body; slender epineurals; no epipleurals; caudal region very short. Supratemporal small, simple, fixed to the parietal and squamosal; no postclavicle; scapular foramen in the scapula. Pectoral fin inserted low down, folding like the ventrals; latter with 6 rays.

fig339

Fig. 339.Phractolaemus ansorgii. ⅔ nat. size.

The remarkable little Fish, Phractolaemus ansorgii, discovered by Dr. W. J. Ansorge in the Niger Delta in 1900, and which has since also been found in the Congo, stands quite apart among the Malacopterygians, its nearest allies being apparently the Osteoglossidae. The body is elongate and subcylindrical, covered with large striated scales; the head is small, the skull strongly ossified, covered with thin skin; the mouth small, proboscidiform, capable of being thrust forwards, when at rest folded over and received into a depression on the upper surface of the head; the narial orifice is single, and preceded by a barbel; the gill-openings are narrow, restricted to the sides. The ventral fins are inserted far back, the dorsal and anal are short. The air-bladder is very large, and the intestine extremely long and much convoluted. Vertebrae 26 + 8.

Fam. 14. Saurodontidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter the more developed and firmly united to the former; these bones, as well as the mandible, with teeth implanted in deep sockets; palate toothless. Supraoccipital separating the small parietals; opercular bones well developed; symplectic present, exposed. Basis cranii double. Ribs sessile, very low down on the centra; no parapophyses; neural arches not fused with the centra. Pectorals inserted very low down; postclavicle apparently absent. Caudal fin deeply forked, without fused hypurals.

This family, comprising several Cretaceous genera, may be regarded as ancestral to the Chirocentridae, with or near which it has been placed by Cope and various later authors. The normal position of the symplectic, however, entitles its members to rank as a separate family, and the autogenous neural arch, as well as the distinctness of the bones supporting the caudal fin, are also indicative of a greater generalisation. The restoration of Ichthyodectes as given by Loomis, shows a general form similar to an ordinary Herring, but it does not appear to be reliable.

The members of the Saurodontidae have been referred to two groups: (a) with praedentary (praesymphysial) bone, Saurocephalus, Saurodon; (b) without praedentary, Chirocentrites, Portheus, Ichthyodectes, Spathodactylus, Cladocyclus. These Fishes are from the Chalk of Europe and North America, and some among them attain a very large size, perhaps not less than two metres in length.

Fam. 15. Chirocentridae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries, the latter the more developed and firmly united to the former; these bones, as well as the mandible, with large teeth not implanted in true sockets; minute teeth on the palatines, pterygoids, and hyoid bones, Supraoccipital in contact with the frontals, separating the small parietals; opercular bones well developed; symplectic hidden between the inner surface of the quadrate and a descending process of the hyomandibular. Basis cranii double. Ribs very slender, sessile, very low down on the centra; no parapophyses; epipleurals and epineurals. Pectorals inserted very low down. Post-temporal forked; postclavicle absent; a thin bony lamina, similar to the postclavicle, above the pectoral fin, attached to the scapula; scapular foramen in scapula; coracoids in contact with each other, forming a keel. Ventrals very small, with 7 rays. Brachiostegal rays 8. Air-bladder large, not communicating with the ear, incompletely divided into cells. Mucous membrane of the intestine forming a spiral fold.

The body is very elongate and strongly compressed, covered with thin, deciduous scales; the vertebrae number 75. The dorsal fin is short and opposite to the anal, which is long.

fig340

Fig. 340.—Side view of skull and pectoral arch of Chirocentrus dorab.

Chirocentrus dorab, the only representative of this family, inhabits the Indian Ocean and the seas of China and Japan.

Fam. 16. Clupeidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries. Supraoccipital separating the small parietals; opercular bones well developed. Basis cranii double. Ribs mostly sessile, inserted behind parapophyses; intermuscular bones (epineurals, epipleurals, adpleurals) usually numerous. Post-temporal forked, the upper branch attached to the epiotic, the lower to the opisthotic; post-clavicle applied to outer side of clavicle. Ventrals with 6 to 11 rays. Air-bladder large, communicating with the ear.

Four sub-families:—

(i.) Thrissopatrinae.—Mouth large; praemaxillaries very small; maxillaries large, with rather narrow supplemental bone, firmly attached to praemaxillaries; branchiostegals about 30; abdomen compressed to an edge, without serration; no lateral line. Thrissopater, from the Gault of Folkestone.

(ii.) Engraulinae.—Mouth moderate or large; praemaxillaries very small; maxillaries large, with narrow supplemental bones, more or less firmly attached to praemaxillaries; branchiostegals 6 to 19; abdomen rounded or more or less compressed, with or without serration; no lateral line. Recent genera: Dussumieria, Etrumeus, Engraulis, Cetengraulis, Heterothrissa, Coilia. Fossil: Spaniodon, Upper Cretaceous.

(iii.) Clupeinae.—Mouth small or moderate; maxillaries freely movable behind the praemaxillaries, usually with large supplemental bones; branchiostegals 5 to 10; abdomen usually serrated; lateral line usually absent. Recent genera: Clupea, Hyperlophus (Diplomystus), Opisthonema, Brevoortia, Pellonula, Clupeichthys, Odaxothrissa, Pellona, Chirocentrodon, Pristigaster, Raconda, Chatoessus. Fossil: Pseudoberyx, Histiothrissa, Scombroclupea, Leptichthys, Upper Cretaceous.

fig341

Fig. 341.—Showing the wide range of variation, within the family, of the bones (pm, praemaxillary, m, maxillary) forming the upper border of the mouth. A, Dussumieria; B, Coilia; C, Pellona; D, Chatoessus; E, Chanos. In these semi-diagrammatic figures the orbit is represented of the same size in all, as affording the best term of comparison in judging of the relative development of the bones of the upper jaw.

(iv.) Chaninae.—Mouth small, toothless; maxillaries firmly attached to praemaxillaries; branchiostegals 4, very broad; abdomen rounded or flattened; lateral line distinct. Chanos, recent; Chanoides, Upper Eocene; Prochanos, Cretaceous.

Heralded by the genus Thrissopater,[647] which may be regarded as a connecting type between the Elopidae and the Clupeidae, this family is largely represented in Cretaceous times, more abundantly still in the Eocene and Miocene, where Clupea and Engraulis occur in numerous species; Hyperlophus, distinguished from Clupea by the presence of a dorsal serrated ridge similar to the ventral, occurs in the Upper Cretaceous of Syria, Southern Europe, and South America, in the Eocene of North America and Europe, and is represented at the present day on the West Coast of South America and on the coast and in the rivers of New South Wales. About 200 Clupeids are known to live at the present day, mostly marine species, but a few are confined to fresh-waters; none may be termed deep-sea forms; some, like the Allis Shad (Clupea alosa) and Twait Shad (C. finta), are anadromous, ascending rivers to spawn. The range of the family is almost cosmopolitan. Several species are remarkable for the extreme abundance of individuals, as for example the Herring (Clupea harengus), the Pilchard or Sardine (C. pilchardus), and the Anchovy (Engraulis encrasicholus). The Herring inhabits the northern parts of the Atlantic and the seas north of Asia. As Dr. Günther first showed, the so-called "Whitebait" consists chiefly of the fry of Herrings, which, like those of the Sprat (C. sprattus), have a predilection for brackish water. The Anchovy and the Pilchard, on the other hand, seldom if ever enter estuaries. The eggs of the Herring, contrary to those of most British marine food-fishes, are heavy and adhesive, sticking firmly to stones or fixed objects on the sea bottom, whilst those of the Sprat and Pilchard float on the surface. The larvae are long, slender, and transparent. The Sardine, which affords so valuable a fishery on the West Coast of France, is the immature state of the Pilchard, which grows to a length of 10 to 14 inches. Its movements are not yet well understood, and its scarcity during certain years in the waters where it usually swarms has caused periodical crises in an important industry. Ripe Pilchards are mostly found at a considerable distance from the coasts. The Anchovy is especially abundant in the Mediterranean, but it is also regularly fished in Holland, especially in the Zuydersee, where it breeds, as well as in the Mediterranean; it makes only temporary appearances, and has not been observed to spawn, in the English Channel, although eggs have recently been obtained off the coast of North Lancashire.[648]

The imperfectly known Cretaceous Crossognathidae (Crossognathus and Scyllaemus), referred by some authors to the Percesoces, should probably be placed with or near the Clupeidae.

Fam. 17. Salmonidae.—Margin of the upper jaw formed by the praemaxillaries and the maxillaries. Supraoccipital in contact with the frontals, but frequently overlapped by the parietals, which may meet in a sagittal suture; opercular bones all well developed. Basis cranii double. Ribs sessile, parapophyses very short or absent; epineurals, sometimes also epipleurals, present. Post-temporal forked, the upper branch attached to the epiotic, the lower to the opisthotic; postclavicle, as usual, applied to inner side of clavicle. A small adipose dorsal fin. Air-bladder usually present, large. Oviducts rudimentary or absent, the ova falling into the cavity of the abdomen before exclusion.

Marine and fresh-water Fishes, mostly from the temperate and Arctic zones of the northern hemisphere: one genus (Retropinna) on the coasts and in the rivers of New Zealand; a few deep-sea forms (Argentina, Microstoma, Nansenia, Bathylagus) occur in the Arctic Ocean, the North Atlantic Ocean, the Mediterranean, and the Antarctic Ocean, down to 2000 fathoms. Apparently of comparatively recent age, no remains older than Miocene (Osmerus, Thaumaturus, Prothymallus) being certainly referable to this family. The recent genera may be grouped as follows:—