III.—Anatomical Works.
The number of authors who worked on the anatomy of fishes is almost as great as that of faunists; and we should go beyond the limits of the present work if we mentioned more than the most prominent and successful. M. H. Rathke, J. Müller, J. Hyrtl, and H. Stannius left scarcely any organ unexamined, and their researches had a direct bearing either on the relation of the class of fishes to the other vertebrates, or on the systematic arrangement of the fishes themselves. E. E. von Baer, F. de Filippi, C. Vogt, W. His, W. K. Parker, and F. M. Balfour worked at their embryology; A. Kölliker and G. Pouchet at their histology. The osteology was specially treated by G. Bakker, F. C. Rosenthal, L. Agassiz, and C. Gegenbaur; the nervous system by Gottsche, Philipeaux, Stannius, L. de Sanctis, L. Stieda, Baudelot and Miclucho-Maclay; the organ of hearing by E. H. Weber, C. Hasse, and G. Retzius. The electric fishes were examined by E. Geoffroy, C. Matteuci, P. Pacini, T. Bilharz, and Max Schultze. The development and metamorphosis of the Lamperns was made the subject of research by H. Müller, M. Schultze, and P. Owsjannikow; Müller’s examination of Branchiostoma was continued by J. Marcusen, A. Kovalevsky, L. Stieda, W. Müller, C. Hasse, T. Huxley, and F. M. Balfour. The most comprehensive accounts of the anatomy of fishes are contained in the following works:—
1. H. Stannius, “Zootomic der Fische,” 2d edit. (Berl. 1854, 8vo.)
2. R. Owen, “Anatomy of Vertebrates,” vol. i. (Lond. 1866, 8vo.)
3. R. Owen, “Lectures on the Comparative Anatomy and Physiology of the Vertebrate Animals.” Part I. Fishes. (Lond. 1846, 8vo.)
4. T. Huxley, “A Manual of the Anatomy of Vertebrated Animals.” (Lond. 1871, 16mo.)
Latest Systematic Works.
It has been mentioned above that the great work of Cuvier and Valenciennes had been left incomplete. Several authors, therefore, supplied detailed accounts of the orders omitted in that work. Müller and Henle published an account of the Plagiostomes, and Kaup of the Murænidæ and Lophobranchii. A. Duméril, finally, commenced an “Histoire naturelle des Poissons ou Ichthyologie générale,” of which, however, two volumes only appeared, containing a complete account of the “Plagiostomes” (Paris, 1865, 8vo.), and of the “Ganoids and Lophobranchs.” (Paris, 1870, 8vo.)
So great an activity had prevailed in Ichthyology since the publication of the “Histoire naturelle” by Cuvier and Valenciennes, and the results of the manifold enquiries were scattered over such a multitude of publications, that it became imperative to collect again all these materials in one comprehensive work. This was done in the “Catalogue of Fishes,” published by the Trustees of the British Museum, in eight volumes (Lond. 1859-70). Beside the species previously described many new forms were added, the number total of species referred to in those volumes amounting to 8525. As regards the systematic arrangement—Müller’s system was adopted in the main, but the definition of the families is much modified. This, however, need not be further entered into here, and will become sufficiently apparent in the subsequent parts of the present work.
Fig. 1.—Lower aspect of head of Raia lemprieri.
CHAPTER II.
TOPOGRAPHICAL DESCRIPTION OF THE EXTERNAL PARTS OF FISHES.
Form of the body.
In the body of a fish four parts are distinguished: the head, trunk, tail, and the fins; the boundary between the first and second being generally indicated by the gill-opening, and that between the second and third by the vent. The form of the body and the relative proportions of those principal parts are subject to much variation, such as is not found in any other class of Vertebrates. In fishes which are endowed with the power of steady and more or less rapid locomotion, a deviation from that form of body, which we observe in a perch, carp, or mackerel, is never excessive. The body forms a simple, equally-formed wedge, compressed or slightly rounded, well fitted for cleaving the water. In fishes which are in the habit of moving on the bottom, the whole body, or at least the head, is vertically depressed and flattened; the head may be so enormously enlarged that the trunk and tail appear merely as an appendage. In one family of fishes, the Pleuronectidæ or Flat-fishes, the body is compressed into a thin disk; they swim and move on one side only, which remains constantly directed towards the bottom, a peculiarity by which the symmetry of all parts of the body has been affected. A lateral compression of the body, in conjunction with a lengthening of the vertical and a shortening of the longitudinal axis, we find in fishes moving comparatively slowly through the water, and able to remain (as it were) suspended in it. This deviation from the typical form may proceed so far that the vertical axis greatly exceeds the longitudinal in length; generally all the parts of the body participate in this form, but in one kind of fish (the Sun-fish or Orthagoriscus) it is chiefly the tail which has been shortened, and reduced so much as to present the appearance of being cut off. An excessive lengthening of the longitudinal axis, with a shortening of the vertical, occurs in Eels and eel-like fishes, and in the so-called Band-fishes. They are bottom-fish, capable of insinuating themselves into narrow crevices and holes. The form of the body of these long fish is either cylindrical, snake-like, as in the Eels and many Cod-fishes, or strongly compressed as in the Band-fishes (Trichiurus, Regalecus, etc.) It is chiefly the tail which is lengthened, but frequently the head and trunk participate more or less in this form. Every possible variation occurs between these and other principal types of form. The old ichthyologists, even down to Linnæus, depended in great measure on them for classification; but although often the same form of body obtains in the same group of fishes, similarity of form by no means indicates natural affinity; it only indicates similitude of habits and mode of life.
Eye.
The external parts of the Head.—The Eye divides the head into the ante-orbital and post-orbital portion. In most fishes, especially in those with a compressed head, it is situated on the side and in the anterior half of the length of the head; in many, chiefly those with a depressed head, it is directed upwards, and sometimes situated quite at the upper side; in very few, the eyes look obliquely downwards. In the Flat-fishes both eyes are on the same side of the head, either the right or the left, always on that which is directed towards the light, and coloured.
Fishes in general, compared with other Vertebrata, have large eyes. Sometimes these organs are enormously enlarged, their great size indicating that the fish is either nocturnal, or lives at a depth to which only a part of the sun’s rays penetrate. On the other hand, small eyes occur in fishes inhabiting muddy places, or great depths to which scarcely any light descends, or in fishes in which the want of an organ of sight is compensated by the development of other organs of sense. In a few fishes, more particularly in those inhabiting caves or the greatest depths of the ocean, the eyes have become quite rudimentary and hidden under the skin.
Snout.
In the ante-orbital portion of the head, or the Snout, are situated the mouth and the nostrils.
Mouth.
The Mouth is formed by the intermaxillary and maxillary bones, or by the intermaxillary only in the upper jaw, and by the mandibulary bone in the lower. These bones are either bare or covered by integument, to which frequently labial folds or lips are added. As regards form, the mouth offers as many variations as the body itself, in accordance with the nature of the food, and the mode of feeding. It may be narrow, or extremely wide and cleft to nearly the hind margin of the head; it may be semi-elliptical, semicircular, or straight in a transverse line; it may be quite in front of the snout (anterior), or at its upper surface (superior), or at its lower (inferior), or extending along each side (lateral); sometimes it is subcircular, organised for sucking. The jaws of some fishes are modified into a special weapon of attack (Sword-fish, Saw-fish); in fact, throughout the whole class of fishes the jaws are the only organ specialised for the purpose of attacking; weapons on other parts of the body are purely defensive.
Both jaws may be provided with skinny appendages, barbels, which, if developed and movable, are sensitive organs of touch.
Nostrils.
In the majority of fishes the Nostrils are a double opening on each side of the upper surface of the snout; the openings of each side being more or less close together. They lead into a shallow groove; and only in one family (the Myxinoids) perforate the palate. In this family, as well as in the Lampreys, the nasal aperture is single. In many Eels the openings are lateral, the lower perforating the upper lip. In the Sharks and Rays (Fig. [1], p. 34) they are at the lower surface of the snout, and more or less confluent; and, finally, in the Dipnoi and other Ganoids, one at least is within the labial boundary of the mouth.
The space across the forehead, between the orbits, is called the interorbital space; that below the orbit, the infraorbital or sub-orbital region.
Gill-cover.
In the post-orbital part of the head there are distinguished, at least in most Teleosteous Fishes and many Ganoids, (Fig. [24]) the præoperculum, a sub-semicircular bone, generally with a free and often serrated or variously-armed margin; the operculum, forming the posterior margin of the gill-opening, and the sub-operculum and interoperculum along its inferior margin. All these bones, collectively called opercles, form the gill-cover, a thin bony lamella covering the cavity containing the gills. Sometimes they are covered with so thin a membrane that the single bones may be readily distinguished; sometimes they are hidden under a thick integument. In some cases the interoperculum is rudimentary or entirely absent (Siluroids).
Gill-opening.
The Gill-opening is a foramen, or a slit behind or below the head, by which the water which has been taken up through the mouth for the purpose of breathing is again expelled. This slit may extend from the upper end of the operculum all round the side of the head to the symphysis of the lower jaw; or it may be shortened and finally reduced to a small opening on any part of the margin of the gill-cover. Sometimes (Symbranchus) the two openings, thus reduced, coalesce, and form what externally appears as a single opening only. The margin of the gill-cover is provided with a cutaneous fringe, in order to more effectually close the gill-opening; and this fringe is supported by one or several or many bony rays, the branchiostegals. The space on the chest between the two rami of the lower jaw and between the gill-openings is called the isthmus.
Fig. 2.—Head of Mordacia mordax, showing the single nostril, and seven branchial openings.
The Sharks and Rays differ from the Teleosteous and Ganoid fishes in having five branchial slits (six or seven in Hexanchus and Heptanchus), which are lateral in the Sharks, and at the lower surface of the head in the Rays (Fig. [1], p. 34). In Myxine only the gill-opening is at a great distance from the head; it is either single in this family (Cyclostomi), or there are six and more on each side (Fig. [2]).
Tail.
In the Trunk are distinguished the back, the sides, and the abdomen. It gradually passes in all fishes into the Tail; the termination of the abdominal cavity and the commencement of the tail being generally indicated by the position of the vent. The exceptions are numerous: not only certain abdominal organs, like the sexual, may extend to between the muscles of the tail, but the intestinal tract itself may pass far backwards, or, singularly, it may be reflected forwards, so that the position of the vent may be either close to the extremity of the tail or to the foremost part of the trunk.
In many fishes the greater part of the tail is surrounded by the fins, leaving only a small portion (between dorsal, caudal, and anal fins) finless; this part is called the free portion or the peduncle of the tail.
Fins.
The Fins are divided into vertical or unpaired, and into horizontal or paired fins. Any of them may be present or absent; and their position, number, and form are most important guides in determining the affinities of fishes.
The vertical fins are situated in the median dorsal line, from the head to the extremity of the tail, and in the ventral line of the tail. In fishes in which they are least developed or most embryonic, the vertical fin appears as a simple fold of the skin surrounding the extremity of the tail In its further progress of development in the series of fishes, it gradually extends more forwards, and may reach even the head and vent. Even in this embryonic condition the fin is generally supported by fine rays, which are the continuations of, or articulated to, other stronger rays supported by the processes or apophyses of the vertebral column. This form of the vertical fin is very common, for instance in the Eels, many Gadoid, Blennioid and Ganoid fishes in which, besides, the rays have ceased to be simple rods, showing more or less numerous joints (simple articulated rays; Fig. [3]). Branched rays are dichotomically split, the joints increasing in number towards the extremity.
The continuity of the vertical fin, however, is interrupted in the majority of fishes; and three fins then are distinguished: one in the dorsal line—the dorsal fin; one in the ventral line behind the anus—the anal fin; and one confined to the extremity of the tail—the caudal fin.
Fig. 3.
- 1. Simple ray.
- 2. Spine.
- 3. Simple articulated ray (soft).
- 4. Branched ray (soft).]
The caudal fin is rarely symmetrical, so that its upper half would be equal to its lower; the greatest degree of asymmetry obtains in fishes with heterocercal termination of the vertebral column (see subsequently, Figs. [31], [41]). In fishes in which it is nearly symmetrical it is frequently prolonged into an upper and lower lobe, its hind margin being concave or more or less deeply excised; in others the hind margin is rounded, and when the middle rays greatly exceed in length the outer ones the fin assumes a pointed form.
Fig. 4.—Labrax lupus (Bass), an Acanthopterygian with anterior spinous, and posterior soft dorsal fin.
Many and systematically important differences are observed in the dorsal fin, which is either spiny-rayed (spinous) (Acanthopterygian), or soft-rayed (Malacopterygian). In the former, a smaller or greater number of the rays are simple and without transverse joints; they may be flexible, or so much osseous matter is deposited in them that they appear hard and truly spinous (Fig. [3]); these spines form always the anterior portion of the fin, which is detached from, or continuous with, the remaining jointed rays. The spines can be erected or depressed at the will of the fish; if in the depressed position the spines cover one another completely, their points lying in the same line, the fish is called homacanth; but if the spines are asymmetrical, alternately broader on one side than on the other, the fish is called heteracanth. The spinous division, as well as the one consisting of jointed rays, may again be subdivided. In the Malacopterygian type all the rays remain jointed; indeed, sometimes the foremost ray, with its preceding short supports, is likewise ossified, and a hard spine, but the articulations can nearly always be distinctly traced. Sometimes the dorsal fin of Malacopterygian fishes is very long, extending from the head to the end of the tail, sometimes it is reduced to a few rays only, and in a few cases it is entirely absent. In addition to the rayed dorsal fin, many Malacopterygian fishes (as the Salmonoids, many Siluroids, Scopeloids, etc.) have another of greater or lesser extent, without any rays; and as always fat is deposited within this fold, it is called a fatty fin (pinna adiposa).
Fig. 5.—Saurus undosquamis, a Malacopterygian with anterior soft dorsal, and additional adipose fin.
The anal fin is built on the same plan as the dorsal, and may be single or plural, long or short, or entirely absent; in Acanthopterygians its foremost rays are frequently simple and spinous.
The horizontal or paired fins consist of two pairs: the pectorals and ventrals.
The pectoral fins (with their osseous supports) are the homologues of the anterior limbs of the higher Vertebrata. They are always inserted immediately behind the gill-opening; either symmetrical with a rounded posterior margin, or asymmetrical, with the upper rays longest and strongest; in Malacopterygians with a dorsal spine the upper pectoral ray is frequently developed into a similar defensive weapon.
The ventral fins are the homologues of the hind-limbs, and inserted on the abdominal surface, either behind the pectorals (Pisces s. Pinnæ abdominales), or below them (Pisces s. Pinnæ thoracicæ), or in advance of them (Pisces s. Pinnæ jugulares). They are generally narrow, composed of a small number of rays, the outer of which is frequently osseous. In some small groups of fishes, like the Gobies, the fins coalesce and form a suctorial disk.
Fig. 6.—Salmo salar (Salmon), with abdominal ventral fins.
Fig. 7.—Mullus barbatus (Red Mullet), with thoracic ventral fins.
Fig. 8.—Burbot (Lota vulgaris), with jugular ventral fins.
For the definition of the smaller systematic groups, and the determination of species, the numbers of the spines and rays are generally of the greatest importance. This holds good, especially for the ventral rays, by the number of which the Acanthopterygian affinities of a fish can nearly always be determined. The numbers of the dorsal and anal rays generally correspond to the number of vertebræ in a certain portion of the spine, and are therefore constant specific, generic, or even family characters; but when their number is very great, a proportionally wide margin must be allowed for variation, and the taxinomic value of this character becomes uncertain. The numbers of the pectoral and caudal rays are rarely of any account.
Function of the Fins.
The fins are organs of motion; but it is chiefly the tail and the caudal fin by which the fish impels itself forward. To execute energetic locomotion the tail and caudal fin are strongly bent, with rapidity, alternately towards the right and left; whilst a gentle motion forwards is effected by a simple undulating action of the caudal fin, the lobes of which act like the blades of a screw. Retrograde motions can be made by fish in an imperfect manner only, by forward-strokes of the pectoral fins. When the fish wants to turn towards the left, he gives a stroke of the tail towards the right, the right pectoral acting simultaneously, whilst the left remains ad-pressed to the body. Thus the pectoral fins assist in the progressive motions of the fish, but rather directing its course than acting as powerful propellers. The chief function of the paired fins is to maintain the balance of the fish in the water, which is always the most unsteady where there is no weight to sink it: when the pectoral of one side, or the pectoral and ventral of the same side are removed, the fish loses its balance and falls on the side opposite; when both pectorals are removed, the fish’s head sinks; on removal of the dorsal and anal fins the motion of the fish assumes a zig-zag course. A fish deprived of all fins, as well as a dead fish, floats with the belly upwards, the back being the heavier part of the body.
In numerous groups of fishes which live in mud, or are enabled to pass a longer or shorter time in soil periodically dried and hardened during the hot season, forms occur entirely devoid of, or with only rudimentary, ventral fins (Cyprinodon, Ophiocephalidæ, Galaxiidæ, Siluridæ). The chief function of these fins being to balance the body of the fish whilst swimming, it is evident that in fishes moving during a great part of their life over swampy ground, or through more or less consistent mud, this function of the ventral fins ceases, and that nature can readily dispense with these organs altogether.
Fig. 9.—Ventrals of Gobius.
In certain fishes the shape and function of the fins are considerably modified: thus, in the Rays, locomotion is almost entirely effected and regulated by the broad and expanded pectoral fins acting with an undulatory motion of their margins, similar to the undulations of the long vertical fins of the Flat-fishes; in many Blennies the ventral fins are adapted for walking on the sea-bottom; in some Gobioids (Periophthalmus), Trigloids, Scorpænioids, and Pediculati, the pectoral fins are perfect organs of walking; in the Gobies, Cyclopteri, and Discoboli the ventral fins are transformed into an adhesive disk, and finally in the Flying-fish, in which the pectorals act as a parachute. In the Eels and other snake-like fishes, the swimming as well as the gliding motions are effected by several curvatures of the body, alternate towards the right and left, resembling the locomotion of Snakes. In the Syngnathi (Pipe-fishes) and Hippocampi, whose body admits of but a slight degree of lateral curvature, and whose caudal fin is generally small, if present at all, locomotion is very limited, and almost wholly dependent on the action of the dorsal fin, which consists of a rapid undulating movement.
Fig. 10.—Cycloid scale of Gadopsis marmoratus (magn.)
Fig. 11.—Cycloid scale of Scopelus resplendens (magn.)
Skin and Scales.
The skin of fishes is either covered with scales, or naked, or provided with more or less numerous scutes of various forms and sizes. Some parts, like the head and fins, are more frequently naked than scaly. All fishes provided with electric organs, the majority of Eels, and the Lampreys, are naked. Scales of fishes are very different from those of Reptiles; the latter being merely folds of the cutis, whilst the scales of fishes are distinct horny elements, developed in grooves or pockets of the skin, like hairs, nails, or feathers. Very small or rudimentary scales are extremely thin, homogeneous in structure, and more or less imbedded in the skin, and do not cover each other. When more developed, they are imbricated (arranged in the manner of tiles), with the posterior part extruded and free, the surface of the anterior portion being usually covered by the skin to a greater or less extent. On their surface (Figs. [10] and [11]) may be observed a very fine striation concentric and parallel to the margin, and coarser striæ radiating from a central point towards the hind margin. Scales without a covering of enamel, with an entire (not denticulated) posterior margin, and with a concentric striation, are called Cycloid scales. Ctenoid scales (Figs. [12–15]) are generally thicker, and provided with spinous teeth on the posterior edges of the layers of which the scale consists. In some species only the layer nearest to the margin is provided with denticulations (Fig. [14]). Scales, the free surface of which is spiny, and which have no denticulation on the margin, have been termed Sparoid scales; but their distinction from ctenoid scales is by no means sharp, and there are even intermediate forms between the cycloid and ctenoid types. Both kinds of scales may occur not only in species of the same genus of fishes, but in the same fish.
Fig. 12.—Ctenoid scale of Scatophagus multifasciatus (magn.)
Fig. 13.—Ctenoid scale of Platycephalus cirrhonasus (magn.)
Fig. 14.—Ctenoid scale of Gobius ommaturus (magn.)
Fig. 15.—Ctenoid scale of Lethrinus (magn.)
Fig. 16. Ganoid Scales.
Ganoid scales are hard and bony, covered with a layer of enamel; they are generally rhombic or quadrangular, rarely rounded and imbricate; and arranged in oblique rows, those of one row being linked together by an articulary process. This type of scales, common in fossil Ganoid fishes, occurs among recent fishes in Lepidosteus and Polypterus only.
Finally, in Sharks, the Balistidæ, and others, true scales are absent and replaced by the ossified papillæ of the cutis, which give the surface the appearance of fine-grained chagreen. These generally small bodies, as well as the large osseous scutes of the Rays, Sturgeons, etc., have been comprised under the common name Placoid scales; a term which deservedly is being abandoned.
Fig. 17.—Dermal papillæ of Monacanthus trossulus.
Fig. 18.—Dermal papillæ of Monacanthus hippocrepis (magn.)
Fig. 19.—Cycloid scale from the lateral line of Odax lineatus (magn.)
Along the side of the body of osseous fishes runs a series of perforated scales, which is called the lateral line (Fig. [21]). The perforating duct is simple at its base, and may be also simple at its outer opening (Fig. [19]), or (and this is frequently the case) the portion on the free surface of the scale is ramified (Fig. [20]). The lateral line runs from the head to the tail, sometimes reaching the caudal fin, sometimes stopping in front of it, sometimes advancing over its rays. It is nearer to the dorsal profile in some fishes than in others. Some species have several lateral lines, the upper one coasting the dorsal, the lower the abdominal outline, one running along the middle as usual. The scales of the lateral line are sometimes larger than the others, sometimes smaller, sometimes modified into scutes, sometimes there are no other scales beside them, the rest of the body being naked. The foramina of the lateral line are the outlets of a muciferous duct which is continued on to the head, running along the infraorbital bones, and sending off a branch into the præopercular margin and mandible. In many fishes, as in many Sciænoids, Gadoids, and in numerous deep-sea fishes, the ducts of this muciferous system are extraordinarily wide, and generally filled with mucus, which is congealed or contracted in specimens preserved in spirits, but swells again when the specimens are immersed in water. This system is abundantly provided with nerves, and, therefore, has been considered to be the seat of a sense peculiar to fishes, but there cannot be any doubt that its function is the excretion of mucus, although probably mucus is excreted also from the entire surface of the fish.
Fig. 20.—Cycloid scale from the lateral line of Labrichthys laticlavius (magn.)
The scales, their structure, number and arrangement, are an important character for the determination of fishes; in most scaly fishes they are arranged in oblique transverse series; and as the number of scales in the lateral line generally corresponds to the number of transverse series, it is usual to count the scales in that line. To ascertain the number of longitudinal series of scales, the scales are counted in one of the transverse series, generally in that running from the commencement of the dorsal fin, or the middle of the back to the lateral line, and from the lateral line down to the vent or ventral fin, or middle of the abdomen.[4]
Fig. 21.—Arrangement of scales in the Roach (Leuciscus ratilus): Ll = Lateral line; tr = Transverse line. a, Transverse line from lateral line to ventral fin.
The scales of many fishes are modified for special purposes, especially to form weapons of defence or a protective armour, but the details of such modifications are better mentioned under the several families in which they occur. All scales are continually growing and wasting away on the surface, and it seems that some fish, at least,—for instance, Salmonoids—“shed” them periodically; during the progress of this shedding the outlines of the scales are singularly irregular.
CHAPTER III.
TERMINOLOGY AND TOPOGRAPHY OF THE SKELETON.
In order to readily comprehend the subsequent account of the modifications of the skeleton in the various sub-classes and groups of Fishes, the student has to acquaint himself with the terms used for the numerous bones of the fish skeleton, as well as with their relative position. The skeleton of any of the more common kinds of osseous fish may serve for this purpose; that of the Perch is chosen here.
The series of bones constituting the axis of the body, and destined to protect the spinal chord and some large longitudinal blood-vessels, is called the vertebral or spinal column; the single bones are the vertebræ. The skull consists of the bones surrounding the brain and organs of sense, and of a number of arches suspended from it, to support the commencement of the alimentary canal and the respiratory organs.
The vertebra (Fig. [22]) consists of a body or centrum (c), with a concave anterior and posterior surface, and generally of several processes or apophyses, as—1. Two neurapophyses (na), which, on the dorsal side, rising upwards, form the neural arch over the canal, in which the spinal chord is lodged. 2. Two parapophyses (pa) usually projecting from the lower part of the sides of the body, or two hæmapophyses (ha) which actually coalesce to form on the ventral side the hæmal canal for a large trunk of the vascular system. 3. A neural spine (ns), which crowns the neurapophyses, or is interposed between their tips. 4. A hæmal spine (hs), having the same relation to the hæmapophyses. 5. Two pleurapophyses or floating ribs, suspended from, or from the base of, the parapophyses. 6. In most fishes the neural arches are connected together by articular or oblique processes, zygapophyses (za), which are developed from the base of each neurapophysis.
Fig. 22.—Side and Front view of Fish-vertebra.
The vertebræ are either abdominal or caudal vertebræ, the coalescence of the parapophyses into a complete hæmal ring, and the suspension of the anal fin generally forming a sufficiently well-marked boundary between abdominal and caudal regions (Fig. [23]). In the Perch there are twenty-one abdominal and as many caudal vertebræ. The centrum of the first vertebra or atlas is very short, with the apophyses scarcely indicated, and lacking ribs like the succeeding vertebra. All the other abdominal vertebræ, with the exception of the last or two last, are provided with ribs, many of which are bifid (72). A series of flat spines (74), called interneurals, to which the spines and rays of the dorsal fins are articulated, are supported by the neural spines, the strength of the neurals and interneurals corresponding to that of the dermal spines (75). The caudal vertebræ differ from the abdominal in having the hæmapophyseal elements converted into spines similar to the neurals, the anterior being likewise destined to support a series of interhæmals (79), to which the anal rays are articulated. The last and smallest caudal vertebra articulates with the hypural (70), a fan-like bone, which, together with the dilated hindmost neural and hæmal elements, supports the caudal rays.
Looking at a perch’s skull from the side (Fig. [24]), the most superficial bones will be found to be those of the jaws, a chain of thin bones round the lower half of the eye, and the opercles.
The anterior margin of the upper jaw is formed by the intermaxillary or premaxillary (17) which bears teeth, terminates in a pedicle above, to allow of a forward sliding motion of the jaw, and is dilated into a flat triangular process behind, on which leans the second bone of the upper jaw, the maxillary (18). This bone is toothless, articulates with the vomer and palatine bone, and is greatly dilated towards its distal extremity. Both the maxillary and intermaxillary lie and move parallel to each other, being connected by a narrow membrane; in many other fishes their relative position is very different.
The mandible or lower jaw consists of a right and left ramus; their union by a ligament in front is called symphysis. Each ramus is formed of several pieces; that which, by a sigmoid concavity articulates with the quadrate, is the articulary bone (35); it sends upwards a coronoid process, to which a ligament from the maxillary and the masticatory muscles are attached; and forwards a long-pointed process, to be sheathed in the deep notch of the anterior piece. A small separate piece (36) at the lower posterior angle of the mandible is termed angular. The largest piece (34) is tooth-bearing, and hence termed dentary; at its inner surface it is always deeply excavated, to receive a cylindrical cartilage, called Meckel’s cartilage, the remains of an embryonic condition of the jaw, the articulary and angular being but ossified parts of it. In other Teleostei this number is still more increased by a splenial and other bones.
The infraorbital ring of bones (Fig. [23], 19) consists of several (four) pieces, of which the anterior is the largest, and distinguished as præorbital.
The so-called præoperculum (30) belongs rather to the bones of the suspensorium of the mandible, presently to be described, than to the opercles proper. It is narrow, strong, angularly bent, so as to consist of a vertical and horizontal limb, with an incompletely closed canal running along both limbs. As it is quite a superficial bone, and frequently armed with various spines, its form and configuration form an important item in the descriptive details of many fishes.
The principal piece of the gill-cover is the operculum (28), triangular in shape, situated behind, and movably united with, the vertical limb of the præoperculum. There is an articulary cavity at its upper anterior angle for its junction with the hyomandibular. The oblong lamella below the operculum is the sub-operculum (32), and the one in front of this latter, below the horizontal limb of the præoperculum, is the interoperculum (33), which is connected by ligament with the angular piece of the lower jaw, and is also attached to the outer face of the hyoid, so that the gill-covers cannot open or shut without the hyoid apparatus executing a corresponding movement.
The chain of flat bones which, after the removal of the temporal muscles, appear arranged within the inner concavity of the præoperculum (Fig. [24]), are comprised with the latter under the common name of mandibulary suspensorium. They connect the mandible with the cranium. The uppermost, the epitympanic or hyomandibular (23), is articulated by a double articulary head with the mastoid and posterior frontal. Another articulary head is destined for the opercular joint. The mesotympanic or symplectic (31) appears as a styliform prolongation of the lower part of the hyomandibular; is entirely cartilaginous in the young, but nearly entirely ossified in the adult. The position of this bone is noteworthy, because, directly inwards of its cartilaginous junction with the hyomandibular, there is situated the uppermost piece of the hyoid arch, the stylohyal. The next bone of the series is the pretympanic or metapterygoid (27), a flat bone forming a bridge towards the pterygoid, and not rarely absent in the teleosteous sub-class. Finally, the large triangular hypo-tympanic or quadrate (26) has a large condyle for the mandibulary joint.
The palatine arch (Fig. [26]) connects the suspensorium with the anterior extremity of the skull, and is formed by three bones: the entopterygoid (25), an oblong and thin bone attached to the inner border of the palatine and pterygoid, and increasing the surface of the bony roof of the mouth towards the median line; it constitutes also the floor of the orbit. The pterygoid (24) (or os transversum) starts from the quadrate, and is joined by suture to the palatine, which is toothed, and reaches to the vomer and anterior frontal.
In the occipital region there are distinguished the basi-occipital (5), readily recognised by the conical excavation corresponding and similar to that of the atlas, with which it is articulated through the intervention of a capsule filled with a gelatinous substance (the remains of the notochord); the exoccipitals (10), articulated, one on each side, to the basi-occipital, and expanding on the upper surface of that bone, so as to meet and support the spinal column; a superficial thin lamella (13), suturally connected with the exoccipitals, not constant in fishes, and erroneously believed by Cuvier to be the petrosal (os petrosum) of higher animals; further, the paroccipitals (9), which are wedged in between the exoccipitals and supraoccipital. This last bone (8) forms the key of the arch over the occipital foramen, and raises a strong high crest from the whole length of its mesial line; a transverse supraoccipital ridge, coming from each side of the base of this spine runs outwards laterally to the external angles of the bone. The supraoccipital separates the parietals, and forms a suture with the frontals.
In front of the basi-occipital the base of the skull is formed by the basisphenoid (parasphenoid of Huxley) (6). This very long and narrow bone extends from the basi-occipital beyond the brain-capsule to between the orbits, where it forms the support of the fibro-membranous interorbital septum. Anteriorly it is connate with another long hammer-shaped bone (16), the vomer, the head of which marks the anterior end of the palate, and is beset with teeth. The alisphenoids (11) are short broad bones, rising from the basisphenoid; their posterior margins are suturally connected with the anterior of the basi- and exoccipitals.
Fig. 23.—Skeleton of the Perch.
Fig. 24.—Skeleton of a Perch’s Skull.
Fig. 25.—Hyoid arch, branchial apparatus, and scapulary arch of the Perch.
Fig. 26.—Lower view of Skull of Perch.
Fig. 27.—Hyoid bone of the Perch.
The formation of the posterior part of the side of the skull is completed by the mastoid and parietal bones. The former (12) projects outwards and backwards farther than the paroccipital, forming the outer strong process of the side of the cranium. This process lodges on its upper surface one of the main ducts of the muciferous system, and affords the base of articulation to a part of the hyomandibular. Its extremity gives attachment to the strong tendon of the dorso-lateral muscles of the trunk. The parietals (7) are flat bones, of comparatively much smaller extent than in higher Vertebrates, and separated from each other by the anterior prolongation of the supraoccipital.
The anterior wall of the brain-capsule (or the posterior of the orbit) is formed by the orbitosphenoids (14), between which, superiorly, the olfactory nerves, and inferiorly, the optic, pass out of the cranium. In addition to this paired bone, the Perch and many other fishes possess another single bone (15),—the os sphenoideum anterius of Cuvier, ethmoid of Owen, and basisphenoid of Huxley; it is Y-shaped, each lateral branch being connected with an orbito-sphenoid, whilst the lower branch rests upon the long basal bone.
A cartilage, the substance of which is thickest above the vomer, and which extends as a narrow stripe along the interorbital septum, represents the ethmoid of higher Vertebrata; the olfactory nerves run along, and finally perforate it.
There remain, finally, the bones distinguishable on the upper surface of the skull; the largest, extending from the nasal cavities to the occipital, are the frontal bones (1), which also form the upper margin of the orbit. The postfrontals (4) are small bones placed on the supero-posterior angle of the orbit, and serving as the point from which the infraorbital ring is suspended. The pre-frontals (2), also small, occupy the anterior margin of the orbit. A pair of small tubiform bones (20), the turbinals, occupy the foremost part of the snout, in front of the frontals, and are separated from each other by intervening cartilage.
After removal of the gill-cover and mandibulary suspensorium, the hyoid arch, which encloses the branchial apparatus, and farther behind, the humeral arch are laid open to view (Fig. [25]). These parts can be readily separated from the cranium proper.
The hyoid arch is suspended by a slender styliform bone, the stylohyal (29), from the hyomandibulars; it consists of three segments, the epihyal (37), ceratohyal (38), which is the longest and strongest piece, and the basihyal, which is formed by two juxtaposed pieces (39, 40). Between the latter there is a median styliform ossicle (41), extending forwards into the substance of the tongue, called glossohyal or os linguale; and below the junction of the two hyoid branches there is a vertical single bone (42), expanded along its lower edge, which, connected by ligament with the anterior extremity of the humeral arch, forms the isthmus separating the two gill-openings. This bone is called the urohyal. Articulated or attached by ligaments to the epihyal and ceratohyal are a number of sword-shaped bones or rays (43), the branchiostegals, between which the branchiostegal membrane is extended.
The branchial arches (Figs. [25] and [27]) are enclosed within the hyoid arch, with which they are closely connected at the base. They are five in number, of which four bear gills, whilst the fifth (56) remains dwarfed, is beset with teeth, and called the lower pharyngeal bone. The arches adhere by their lower extremities to a chain of ossicles (53, 54, 55), basibranchials, and, curving as they ascend, nearly meet at the base of the cranium, to which they are attached by a layer of ligamentous and cellular tissue. Each of the first three branchial arches consists of four pieces movably connected with one another. The lowest is the hypobranchial (57), the next much longer one (58) the cerato-branchial, and, above this, a slender and a short irregularly-shaped epibranchial (61). In the fourth arch the hypobranchial is absent. The uppermost of these segments (62), especially of the fourth arch, are dilated, and more or less confluent; they are beset with fine teeth, and generally distinguished as the upper pharyngeal bones. Only the cerato-branchial is represented in the fifth arch or lower pharyngeal. On their outer convex side the branchial segments are grooved for the reception of large blood-vessels and nerves; on the inner side they support horny processes (63), called the gill-rakers, which do not form part of the skeleton.
The scapular or humeral arch is suspended from the skull by the (suprascapula) post-temporal (46), which, in the Perch, is attached by a triple prong to the occipital and mastoid bones. Then follows the (scapula) supraclavicula (47), and the arch is completed below by the union of the large (coracoid) clavicula (48) with its fellow. Two flat bones (51, 52), each with a vacuity, attached to the clavicle have been determined as the (radius and ulna) coracoid and scapula of higher vertebrates, and the two series of small bones (53) intervening between the forearm and the fin as carpals and metacarpals. A two-jointed appendage the (epicoracoid) postclavicula, is attached to the clavicle: its upper piece (49) is broad and lamelliform, its lower (50) styliform and pointed.
The ventral fins are articulated to a pair of flat triangular bones, the pubic bones (80).
The bones of the skull of the fish have received so many different interpretations that no two accounts agree in their nomenclature, so that their study is a matter of considerable difficulty to the beginner. The following synonymic table will tend to overcome difficulties arising from this cause; it contains the terms used by Cuvier, those introduced by Owen, and finally the nomenclature of Stannius, Huxley, and Parker. Those adopted in the present work are printed in italics. The numbers refer to the figures in the accompanying woodcuts (Figs. [23–27]).
| Cuvier. | Owen. | Stannius. | Huxley, Parker, etc. | ||||
| 1. | Frontal principal | Frontal | Os frontale | ||||
| 2. | Frontal antérieur | Prefrontal | Os frontale anterius | Lateral ethmoid (Parker) | |||
| 3. | Ethmoid | Nasal | Os ethmoideum | ||||
| 4. | Frontal postérieur | Postfrontal | Os frontale posterius | Sphenotic (Parker) | |||
| 5. | Basilaire | Basioccipital | Os basilare | ||||
| 6. | Sphénoide | Basisphenoid | Os sphenoideum basilare | Sometimes referred to as “Basal” | |||
| 7. | Pariétal | Parietal | Os parietale | ||||
| 8. | Interpariétal or occipital supérieure | Supraoccipital | Os occipitale superius | ||||
| 9. | Occipital externe | Paroccipital | Os occipitale externum | Epioticum (Huxley) | |||
| 10. | Occipital lateral | Exoccipital | Os occipitale laterale | ||||
| 11. | Grande aile du sphénoide | Alisphenoid | Ala temporalis | Prooticum (Huxley) | |||
| 12. | Mastoidien | Mastoid | Os mastoideum + os extrascapulare |  | Opisthoticum[5] +Squamosal (Huxley) | ||
| 13. | Rocher | Petrosal and Otosteal | Oberflächliche Knochen-lamelle | ||||
| 14. | Aile orbitaire | Orbitosphenoid | Ala orbitalis | Alisphenoid (Huxley) | |||
| 15. | Sphenoide antérieur | Ethmoid and Ethmoturbinal | Os sphenoideum anterius | Basisphenoid (Huxley) | |||
| 16. | Vomer | Vomer | Vomer | ||||
| 17. | Intermaxillaire | Inter- or Pre-maxillary | Os intermaxillare | ||||
| 18. | Maxillaire supérieur | Maxillary | Os maxillare | ||||
| 19. | Sousorbitaires | Infraorbital ring | Ossa infraorbitalia | ||||
| 20. | Nasal | Turbinal | Os terminale | ||||
| 22. | Palatine | Palatin | Os palatinum | ||||
| 23. | Temporal | Epitympanic | Os temporale | Hyomandibular (Huxley) | |||
| 24. | Transverse | Pterygoid | Os transversum s. pterygoideum externum | ||||
| 25. | Ptérygoidien interne | Entopterygoid | Os pterygoideum | Mesopterygoid (Parker) | |||
| 26. | Jugal | Hypotympanic | Os quadratojugale | Quadrate (Huxley) | |||
| 27. | Tympanal | Pretympanic | Os tympanicum | Metapterygoid (Huxley) | |||
| 28. | Operculaire | Operculum | Operculum | ||||
| 29. | Styloide | Stylohyal | Os styloideum | ||||
| 30. | Préopercule | Præoperculum | Præoperculum | ||||
| 31. | Symplectique | Mesotympanic | Os symplecticum | ||||
| 32. | Sousopercule | Suboperculum | Suboperculum | ||||
| 33. | Interopercule | Interoperculum | Interoperculum | ||||
| 34. | Dentaire | Dentary | Os dentale | ||||
| 35. | Articulaire | Articulary | Os articulare | ||||
| 36. | Angulaire | Angular | Os angulare | ||||
| 37. |  | Grandes pièces latérales | Epihyal | | Segmente der Zungenbein-Schenkel | ||
| 38. | Ceratohyal | ||||||
| 39. | | Petites pièces laterales | Basihyal | ||||
| 40. | |||||||
| 41. | Os lingual | Glossohyal | Os linguale s. entoglossum | ||||
| 42. | Queue de l’os hyoide | Urohyal | Basibranchiostegal (Parker) | ||||
| 43. | Rayon branchiostège | Branchiostegal | Radii branchiostegi | ||||
| 46. | Surscapulaire | Suprascapula | Omolita | Post-temporal (Parker) | |||
| 47. | Scapulaire | Scapula | Scapula | Supraclavicula (Parker) | |||
| 48. | Humeral | Coracoid | Clavicula | Clavicula (Parker) | |||
| 49. | | Coracoid | Epicoracoid | Postclavicula (Parker) | |||
| 50. | |||||||
| 51. | Cubital | Radius | | Ossa carpi | Coracoid (Parker) | ||
| 52. | Radial | Ulna | Scapula (Parker) | ||||
| 53. | Os du carpe | Carpals | Ossa metacarpi | | Basalia (Huxley), | ||
| Brachials (Parker) | |||||||
| 53 bis. | | Chaine intermédiaire | Basibranchials | Copula | |||
| 55. | |||||||
| 54. | |||||||
| 56. | Pharyngiens inférieurs | Lower Pharyngeals | Ossa pharyngea inferiora | ||||
| 57. | Pièce interne de partie inférieure de l’arceau branchiale | Hypobranchial | | Segmente der Kiemenbogen-Schenkel | |||
| 58. | Pièce externe „ | Ceratobranchial | |||||
| 59. | Stylet de prémière arceau branchiale | Upper epibranchial of first branchial arch | |||||
| 61. | Partie supérieure de l’arceau branchiale | Epibranchials | |||||
| 62. | Os pharyngian supérieur | Pharyngobranchial | Os pharyngeum superius | Upper pharyngeals | |||
| 63. | Gill-rakers | ||||||
| 65. | Rayons de la pectorale | Pectoral rays | Brustflossen-Strablen | ||||
| 67, 68. | Vertèbres abdominales | Abdominal vertebræ | Bauchwirbel | ||||
| 69. | Vertèbres caudales | Caudal vertebræ | Schwanzwirbel | ||||
| 70. | Plaque triangulaire et verticale | [Aggregated interhæmals] | Verticale Platte | Hypural (Huxley) | |||
| 71. | Caudal rays | Schwanzflossen Strahlen | |||||
| 72. | Côte | Rib | Rippen | ||||
| 73. | Appendices or stylets | Epipleural spines | Muskel-Gräthen | ||||
| 74. | Interépineux | Interneural spines | Ossa interspinalia s. obere Flossentræger | ||||
| 75. | Épines et rayons dorsales | Dorsal rays and spines | Rückenflossen-Strablen u. Stacheln | ||||
| 76. | First interneural | ||||||
| 78. | Rudimentary caudal rays | ||||||
| 79. | Apophyses épineuses inférieures | Interhæmal spines | Untere Flossentræger | ||||
| 80. | Pubic | Becken | |||||
| 81. | Ventral spine | Bauchflossen-Stachel |
CHAPTER IV.
MODIFICATIONS OF THE SKELETON.
The lowermost sub-class of fishes, which comprises one form only, the Lancelet (Branchiostoma [s. Amphioxus] lanceolatum), possesses the skeleton of the most primitive type.
Fig. 28.—Branchiostoma lanceolatum. a, Mouth; b, Vent; c, abdominal porus.
Fig. 29.—Anterior end of body of Branchiostoma (magn.) d, Chorda dorsalis; e, Spinal chord; f, Cartilaginous rods; g, Eye; h, Branchial rods; i, Labial cartilage; k, Oral cirrhi.
The vertebral column is represented by a simple chorda dorsalis or notochord only, which extends from one extremity of the fish to the other, and, so far from being expanded into a cranial cavity, it is pointed at its anterior end as well as at its posterior. It is enveloped in a simple membrane like the spinal chord and the abdominal organs, and there is no trace of vertebral segments or ribs; however, a series of short cartilaginous rods above the spine evidently represent apophyses. A maxillary or hyoid apparatus, or elements representing limbs, are entirely absent.
[J. Müller, Ueber den Bau und die Lebenserscheinungen des Branchiostoma lubricum, in Abhandl. Ak. Wiss. Berlin, 1844.]
The skeleton of the Cyclostomata (or Marsipobranchii) (Lampreys and Sea-hags) shows a considerable advance of development. It consists of a notochord, the anterior pointed end of which is wedged into the base of a cranial capsule, partly membranous partly cartilaginous. This skull, therefore, is not movable upon the spinal column. No vertebral segmentation can be observed in the notochord, but neural arches are represented by a series of cartilages on each side of the spinal chord. In Petromyzon (Fig. [30]) the basis cranii emits two prolongations on each side: an inferior, extending for some distance along the lower side of the spinal column, and a lateral, which is ramified into a skeleton supporting the branchial apparatus. A stylohyal process and a subocular arch with a palato-pterygoid portion may be distinguished. The roof of the cranial capsule is membranous in Myxine and in the larvæ of Petromyzon, but more or less cartilaginous in the adult Petromyzon and in Bdellostoma. A cartilaginous capsule on each side of the hinder part of the skull contains the auditory organ, whilst the olfactory capsule occupies the anterior upper part of the roof. A broad cartilaginous lamina, starting from the cranium and overlying part of the snout, has been determined as representing the ethmo-vomerine elements, whilst the oral organs are supported by large, very peculiar cartilages (labials), greatly differing in general configuration and arrangement in the various Cyclostomes. There are three in the Sea-lamprey, of which the middle one is joined to the palate by an intermediate smaller one; the foremost is ring-like, tooth-bearing, emitting on each side a styliform process. The lingual cartilage is large in all Cyclostomes.
There is no trace of ribs or limbs.
[J. Müller, Vergleichende Anatomie der Myxinoiden. Erster Theil. Osteologie und Myologie, in Abhandl. Ak. Wiss. Berlin, 1835.]
Fig. 30.—Upper (A) and side (B) views, and vertical section (C) of the skull of Petromyzon marinus.
a, Notochord; b, Basis cranii; c, Inferior, and d, Lateral process of basis; e, Auditory capsule; f, Subocular arch; g, Stylohyal process; h, Olfactory capsule; i, Ethmo-vomerine plate; k, Palato-pterygoid portion of subocular arch; l-n, Accessory labial or rostral cartilages; with o, appendage; p, lingual cartilage; q, neural arches; r, Branchial skeleton; s, Blind termination of the nasal duct between the notochord and œsophagus.
Fig. 31.—Heterocercal Tail of Centrina salviani.
a, Vertebræ; b, Neurapophyses; c, Hæmapophyses.
The Chondropterygians exhibit a most extraordinary diversity in the development of their vertebral column; almost every degree of ossification, from a notochord without a trace of annular structure to a series of completely ossified vertebræ being found in this order. Sharks, in which the notochord is persistent, are the Holocephali (if they be reckoned to this order, and the genera Notidanus and Echinorhinus). Among the first, Chimæra monstrosa begins to show traces of segmentation; but they are limited to the outer sheath of the notochord, in which slender subossified rings appear. In Notidanus membranous septa, with a central vacuity, cross the substance of the gelatinous notochord. In the other Sharks the segmentation is complete, each vertebra having a deep conical excavation in front and behind, with a central canal through which the notochord is continued; but the degree in which the primitive cartilage is replaced by concentric or radiating lamellæ of bone varies greatly in the various genera, and according to the age of the individuals. In the Rays all the vertebræ are completely ossified, and the anterior ones confluent into one continuous mass.
In the majority of Chondropterygians the extremity of the vertebral column shows a decidedly heterocercal condition (Fig. [31]), and only a few, like Squatina and some Rays, possess a diphycercal tail
The advance in the development of the skeleton of the Chondropterygians beyond the primitive condition of the previous sub-classes, manifests itself further by the presence of neural and hæmal elements, which extend to the foremost part of the axial column, but of which the hæmal form a closed arch in the caudal region only, whilst on the trunk they appear merely as a lateral longitudinal ridge.
Fig. 32.—Lateral view.
Fig. 33.—Longitudinal section.
Fig. 34.—Transverse section of Caudal vertebra of Basking Shark (Selache maxima). (After Hasse.) a, Centrum; b, Neurapophysis; c, Intercrural cartilage; d, Hæmapophysis; e, Spinal canal; f, Intervertebral cavity; g, Central canal for persistent portion of notochord; h, Hæmal canals for blood-vessels.
The neural and hæmal apophyses are either merely attached to the axis, as in Chondropterygians with persistent notochord, the Rays and some Sharks; or their basal portions penetrate like wedges into the substance of the centrum, so that, in a transverse section, in consequence of the difference in their texture, they appear in the form of an X.[6] The interspaces between the neurapophyses of the vertebræ are not filled by fibrous membrane, as in other fishes, but by separate cartilages, laminæ or cartilagines intercrurales, to which frequently a series of terminal pieces is superadded, which must be regarded as the first appearance of the interneural spines of the Teleostei and many Ganoids. Similar terminal pieces are sometimes observed on the hæmal arches. Ribs are either absent or but imperfectly represented (Carcharias).
The substance of the skull of the Chondropterygians is cartilage, interrupted especially on its upper surface by more or less extensive fibro-membranous fontanelles. Superficially it is covered by a more or less thick chagreen-like osseous deposit. The articulation with the vertebral column is effected by a pair of lateral condyles. In the Sharks, besides, a central conical excavation corresponds to that of the centrum of the foremost vertebral segment, whilst in the Rays this central excavation of the skull receives a condyle of the axis of the spinous column.
The cranium itself is a continuous undivided cartilage, in which the limits of the orbit are well marked by an anterior and posterior protuberance. The ethmoidal region sends horizontal plates over the nasal sacs, the apertures of which retain their embryonic situation upon the under surface of the skull. In the majority of Chondropterygians these plates are conically produced, forming the base of the soft projecting snout; and in some forms, especially in the long-snouted Rays and the Saw-fishes (Pristis) this prolongation appears in the form of three or more tubiform rods.
As separate cartilages there are appended to the skull a suspensorium, a palatine, mandible, hyoid, and rudimentary maxillary elements.
The suspensorium is movably attached to the side of the skull. It generally consists of one piece only, but in some Rays of two. In the Rays it is articulated with the mandible only, their hyoid possessing a distinct point of attachment to the skull. In the Sharks the hyoid is suspended from the lower end of the suspensorium together with the mandible.
What is generally called the upper jaw of a Shark is, as Cuvier has already stated, not the maxillary, but palatine. It consists of two simple lateral halves, each of which articulates with the corresponding half of the lower jaw, which is formed by the simple representative of Meckel’s cartilage.
Some cartilages of various sizes are generally developed on each side of the palatine, and one on each side of the mandible. They are called labial cartilages, and seem to represent maxillary elements.
The hyoid consists generally of a pair of long and strong lateral pieces, and a single mesial piece. From the former cartilaginous filaments (representing branchiostegals) pass directly outwards. Branchial arches, varying in number, and similar to the hyoid, succeed it. They are suspended from the side of the foremost part of the spinous column, and, like the hyoid, bear a number of filaments.
The vertical fins are supported by interneural and interhæmal cartilages, each of which consists of two and more pieces, and to which the fin-rays are attached without articulation.
The scapular arch of the Sharks is formed by a single coracoid cartilage bent from the dorsal region downwards and forwards. In some genera (Scyllium, Squatina) a small separate scapular cartilage is attached to the dorsal extremities of the coracoid; but in none of the Elasmobranchs is the scapular arch suspended from the skull or vertebral column; it is merely sunk, and fixed in the substance of the muscles. Behind, at the point of its greatest curvature, three carpal cartilages are joined to the coracoid, which Gegenbaur has distinguished as propterygium, mesopterygium, and metapterygium, the former occupying the front, the latter the hind margin of the fin. Several more or less regular transverse series of styliform cartilages follow. They represent the phalanges, to which the horny filaments which are imbedded in the skin of the fin are attached.
In the Rays, with the exception of Torpedo, the scapular arch is intimately connected with the confluent anterior portion of the vertebral column. The anterior and posterior carpal cartilages are followed by a series of similar pieces, which extend like an arch forwards to the rostral portion of the skull, and backwards to the pubic region. Extremely numerous phalangeal elements, longest in the middle, are supported by the carpals, and form the skeleton of the lateral expansion of the so-called disk of the Ray’s body, which thus, in fact, is nothing but the enormously enlarged pectoral fin.
The pubic is represented by a single median transverse cartilage, with which a tarsal cartilage articulates. The latter supports the fin-rays. To the end of this cartilage is also attached, in the male Chondropterygians, a peculiar accessory generative organ or clasper.
The Holocephali differ from the other Chondropterygians in several important points of the structure of their skeleton, and approach unmistakably certain Ganoids. That their spinal column is persistently notochordal has been mentioned already. Their palatal apparatus, with the suspensorium, coalesces with the skull, the mandible articulating with a short apophysis of the cranial cartilage. The mandible is simple, without anterior symphysis. The spine with which the dorsal fin is armed articulates with a neural apophysis, and is not immovably attached to it, as in the Sharks. The pubic consists of two lateral halves, with a short, rounded, tarsal cartilage.
The skeleton of the Ganoid Fishes offers extreme variations with regard to the degree in which ossifications replace the primordial cartilage. Whilst some exhibit scarcely any advance beyond the Plagiostomes with persistent cartilage, others approach, as regards the development and specialisation of the several parts of their osseous framework, the Teleosteans so closely that their Ganoid nature can be demonstrated by, or inferred from, other considerations only. All Ganoids possess a separate gill-cover.[7]
The diversity in the development of the Ganoid skeleton is well exemplified by the few representatives of the order in the existing Fish-fauna. Lowest in the scale (in this respect) are those with a persistent notochord, and an autostylic skull, that is, a skull without separate suspensorium—the fishes constituting the suborder Dipnoi, of which the existing representatives are Lepidosiren, Protopterus, and Ceratodus, and the extinct (as far as demonstrated at present) Dipterus, Chirodus (and Phaneropleuron?). In these fishes the notochord is persistent, passing uninterruptedly into the cartilaginous base of the skull. Only now and then a distinct vertical segmentation occurs in the caudal portion of the column, but it does not extend to the notochord itself, but indicates only the limits between the superadded apophyseal elements, each neural being confluent with the opposite hæmal. Some Dipnoi are diphy-, others heterocercal.
Neural and hæmal elements and ribs are well developed. In Ceratodus each neurapophysis consists of a basal cartilaginous portion, forming an arch over the myelon, and of a superadded second portion. The latter is separated from the former by a distinct line of demarcation, and its two branches are more styliform, cartilaginous at the ends and in the centre, but with an osseous sheath, and coalesced at the top, forming a gable over an elastic fibrous band which runs along and parallel to the longitudinal axis of the column (Ligamentum longitudinale superius). To the top of this gable is joined a single long cylindrical neural spine. From the eleventh apophyseal segment a distinct interneural spine, of the same structure as the neural, begins to be developed, and farther on a second interneural is superadded. Towards the extremity of the column these various pieces are gradually reduced in size and number, finally only a low cartilaginous band (the rudiments of the neurapophysis) remaining. The hæmapophyses are in form, size, and structure, very similar to the neurapophyses; and all these long bones, including the ribs, have that in common, that they consist of a solid rod of cartilage enclosed in a bony sheath, which, after the disappearance or decomposition of the cartilage, appears as a hollow tube. Such bones are extremely common throughout the order of Ganoids, and their remains have led to the designation of a family as Cœlacanthi (κοιλος, hollow; and ἀκανθος, spine).
The primordial cranium of the Dipnoi is cartilaginous, but with more or less extensive ossifications in its occipital, basal, or lateral portions, and with large tegumentary bones, the arrangement of which varies in the different genera. There is no separate suspensorium for the lower jaw. A strong process descends from the cranial cartilage, and offers by means of a double condyle (Fig. [35] s) attachment to corresponding articulary surfaces of the lower jaw. Maxillary and intermaxillary elements are not developed, but, perhaps, represented in Ceratodus by some inconstant rudimentary labial cartilages situated behind the posterior nasal opening. Facial cartilages and an infraorbital ring are developed at least in Ceratodus. The presence of a pair of small teeth in front indicates the vomerine portion (v) which remains cartilage, whilst the posterior pair of teeth are implanted in a pterygo-palatine ossification (l), which sometimes is paired, sometimes continuous. The base of the skull is constantly covered by a large basal ossification (o).
Fig. 35.—Palatal view of Skull of Ceratodus.
The hyoid is well developed, sometimes reduced to a pair of ceratohyals, sometimes with a basihyal and glossyhyal. The skeleton of the branchial apparatus approaches the Teleosteous type, less so in Lepidosiren than in Ceratodus, in which five branchial arches are developed, but with the lateral and mesial pieces reduced in number.
A large operculum, and a smaller sub- or interoperculum are present.
The scapular arch consists of a single median transverse cartilage, and a pair of lateral cartilages which bear the articular condyle for the pectoral limb. The latter cartilages form the base of a large membrane-bone, and the whole arch is suspended from the skull by means of an osseous supraclavicle.
The fore-limb of the Dipnoi (Fig. [36]) differs externally greatly from the pectoral fin of other Ganoid fishes. It is covered with small scales along the middle, from the root to its extremity, and surrounded by a rayed fringe similar to the vertical fin. A muscle split into numerous fascicles extends all the length of the fin, which is flexible in every part and in every direction. The cartilaginous framework supporting it is joined to the scapular arch by an oblong cartilage, followed by a broad basal cartilage (a), generally single, sometimes showing traces of a triple division. Along the middle of the fin runs a jointed axis (b), the joints gradually becoming smaller and thinner towards the extremity; each joint bears on each side a three, two, or one-jointed branch (c, d). This axial arrangement of the pectoral skeleton, which evidently represents one of its first and lowest conditions, has been termed Archipterygium by Gegenbaur. It is found in Ceratodus and other genera, but in Lepidosiren the jointed axis only has been preserved, with the addition of rudimentary rays in Protopterus.
Fig. 36.—Fore-limb of Ceratodus.
The pubic consists of a single flattened subquadrangular cartilage, produced into a long single anterior process. Posteriorly it terminates on each side in a condyle, to which the basal cartilage of the ventral paddle is joined. The endoskeleton of the paddle is almost identical with that of the pectoral.
The Ganoid fishes with persistent notochord, but with a hyostylic skull (that is, a skull with a separate suspensorium) consist of the suborder Chondrostei, of which the existing representatives are the Sturgeons (Acipenser, Scaphirhynchus, Polyodon), and the extinct the Chondrosteidæ, Palæoniscidæ, and (according to Traquair) Platysomidæ.
Their spinal column does not differ essentially from that of the Dipnoi. Segmentation is represented only as far as the neural and hæmal elements are concerned. All are eminently heterocercal. Ribs are present in most, but replaced by ligaments in Polyodon.
Fig. 37.—Skull of Polyodon (after Traquair).
n, Nasal cavity; sq, squamosal; mh, hyomandibular; sy, symplectic; pa, palato-pterygoid; m, Meckelian cartilage; mx, maxillary; d, dentary; h, hyoid; op, opercle; br, branchiostegal; s.cl, supra-clavicular; p.cl, post-clavicular; cl, clavicle; i.cl, infra-clavicular.
The primordial cranium of the Sturgeons consists of persistent cartilage without ossifications in its substance, but superficial bones are still more developed and specialised than in the Dipnoi; so it is, at least, in the true Sturgeons, but less so in Polyodon (Fig. [37]). The upper and lateral parts of the skull are covered by well-developed membrane bones, which, from this suborder, upwards in the series, will be found to exist throughout the remaining forms of fishes. They are bones, the origin of which is not in cartilage but in membranous connective tissue. The lower surface of the skull is covered by an extremely large basal bone, which extends from the vomerine region on to the anterior part of the spinal column. The nasal excavation in the skull is rather lateral than inferior. The ethmoidal region is generally much produced, forming the base of the long projecting snout. The suspensorium is movably attached to the side of the skull, and consists of two pieces, a hyomandibular and a symplectic, which now appears for the first time as a separate piece, and to which the hyoid is attached. The palato-maxillary apparatus is more complex than in the Sharks and Dipnoi; a palato-pterygoid consists of two mesially-connected rami in Polyodon, and of a complex cartilaginous disk in Acipenser, being articulated in both to the Meckelian cartilage. In addition, the Sturgeons possess one or two pairs of osseous rods, which, in Polyodon at least, represent the maxillary, and therefore must be the representatives of the labial cartilages of the Sharks. The Meckelian cartilage is more or less covered by tegumentary bones.
In the gill-cover, besides the operculum, a sub- and interoperculum may be distinguished in Acipenser.
The hyoid consists of three pieces, of which the posterior bears a broad branchiostegal in Polyodon.
In the scapulary arch the primordial cartilaginous elements scarcely differ from those of the Dipnoi. The membrane-bones are much expanded, and offer a continuous series suspended from the skull. Their division in the median ventral line is complete.
Fig. 38.—Fore-limb of Acipenser.
The pectoral is supported by a cartilaginous framework (Fig. [38]) similar to that of Ceratodus, but much more shortened and reduced in its periphery, the branches being absent altogether on one side of the axis. This modification of the fin is analogous to the heterocercal condition of the end of the spinous column. To the inner corner of a basal cartilage (a) a short axis (b) is joined, which on its outer side bears a few branches (d) only, the remaining branches (c) being fixed to the basal cartilage. The dermal fin-rays are opposed to the extremities of the branches, as in the Dipnoi.
The pubic consists of a paired cartilage, to which tarsal pieces supporting the fin-rays are attached.
The other living Ganoid fishes have the spinous column entirely or nearly entirely ossified, and have been comprised under the common name Holostei. However, they form three very distinct types; several attempts have been made to coordinate with them the fossil forms, but this task is beset with extreme difficulties, and its solution hitherto has not proved to be satisfactory.
The Polypteroidei have their spinous column formed by distinct osseous amphicœlous vertebræ, that is, vertebræ with concave anterior and posterior surfaces. It is nearly diphycercal; a slight degree of heterocercy obtains, inasmuch as the last vertebra is succeeded by a very thin cartilaginous filament which penetrates between the halves of one of the middle rays of the terminal fin. The rays above this cartilaginous filament are articulated to interneurals, those below lack interhæmals, and are attached either to the hæmals or vertebral centres. The neural arches, though ossified, do not coalesce with the centrum, and form one canal only, for the myelon. There are no intermediate elements between the neural spines. Interneurals developed, but simple, articulating with the dermoneurals. The abdominal vertebræ have parapophyses developed with epipleural spines. Only the caudal vertebræ have hæmal spines, which, like the interhæmals, agree in every essential respect with the opposite neurals. Ribs are inserted, not on the parapophyses, but on the centre, immediately below the parapophyses.
The skull of Polypterus (Fig. [39]) shows a great advance towards the Teleosteous type, the number of separable bones being greatly increased. They are arranged much in the same fashion as in Teleostei. But a great portion of the primordial cranium remains cartilaginous. The membrane-bones which cover the upper and lower surfaces of the brain-case are so much developed as to cause the underlying cartilage to disappear, so that a large vacuity or fontanelle exists in the substance of the upper as well as lower cartilaginous wall. Of ossifications belonging to the primordial skull must be noticed the single occipital with a mastoid on each side. They are separated by persistent cartilage from the sphenoids and postfrontals; the former, which are the largest ossification of the primordial cranium, enclose the anterior half of the brain cavity. Finally, the nasal portion contains a median ethmoid and a pair of præfrontal bones.
Fig. 39.—Skull of Polypterus. (After Traquair.)
Fig. I. Upper aspect of the Primordial Cranium, with the membrane-bones removed. Fig. II. Lower aspect of the same. Fig. III. Side view, with the membrane-bones. Fig. IV. Lower aspect of the Skull, part of the bones being removed on one side. The parts shaded with oblique lines are cartilage of the primordial skull.
An, Angular; ao, ante-orbital; Ar, articulary; B, basal; D, dentary; E, ethmoid; F, frontal; Ma, mastoid; Mp, metapterygoid; Mx, maxillary; N, nasal; O, operculum; Oc, occipital; Pa, parietal; Pl, palatine; Pmx, præmaxillary; po, post-orbital; Prf, prefrontal; Pt, post-temporals; Ptf, postfrontal; Ptr, pterygoid; Q, quadrate; S, suspensorium; So, sub-operculum; Sp, sphenoid; Spl, splenial; St, supratemporals; T, tympanic lamina; Tu, turbinal; v, vomer; x x, small ossicles; x’ x’, spiraculars.
Only a very small portion of the bones described are visible externally, nearly the whole of the primordial cranium being covered by the membrane-bones. Of these are seen on the upper surface a pair of parietals, frontals, “nasals,” and turbinals; on the lower surface a large cross-shaped basal, anteriorly bordered on each side by a pterygoid, parallel to a palatine which forms a suture with the double vomer. The suspensorium has in front a metapterygoid and quadrate bone, and an operculum and sub-operculum are attached to it behind.
Præmaxillaries and maxillaries are now fully developed, but immovably attached to the skull. The lower jaw is ossified, and consists of an articulary, angular, dentary, and splenial. Of labial cartilages a rudiment at the angle of the mouth has remained persistent.
The side of the skull, in front of the operculum, is covered by a large irregularly-shaped bone (T) (corresponding to the “tympanic lamina” of Ceratodus, Fig. [35], q), held by some to be the præoperculum; along its upper circumference lies a series of small ossicles, of which two may be distinguished as spiraculars, as they form a valve for the protection of the spiracular orifice of these fishes. An infraorbital ring is represented by a præ- and post-orbital only.
Each hyoid consists of three pieces, none of which bear branchiostegals, the single median piece being osseous in front and cartilaginous behind. Four branchial arches are developed, the foremost consisting of three, the second and third of two, and the last of a single piece. There is no lower pharyngeal. Between the rami of the lower jaw the throat is protected by a pair of large osseous laminæ (gular plates), which have been considered to represent the urohyal of osseous fishes.
The scapulary arch is almost entirely formed by the well-developed membrane-bones, which in the ventral line are suturally united. The pectoral fin is supported by three bones, pro-, meso-, and metapterygium, of which the dilated middle one alone bears rays, and is excluded from the articulation with the shoulder-girdle.
The pubic consists of paired bone, to which tarsal bones supporting the fin-rays are attached.
In the Lepidosteoidei the vertebræ are completely ossified, and opisthocœlous, having a convexity in front and a concavity behind, as in some Amphibians. Though the end of the body externally appears nearly diphycercal, the termination of the vertebral column is, in fact, distinctly heterocercal (Fig. [40]). Its extremity remains cartilaginous, is turned upwards, and lies immediately below the scutes which cover the upper margin of the caudal fin. It is preceded by a few rudimentary vertebræ which gradually pass into the fully developed normal vertebræ. The caudal fin is suspended from hæmapophyses only, and does not extend to the neural side of the vertebral column. The neural arches coalesce with the centrum; interneurals simple. The abdominal vertebræ have parapophyses, to which the ribs are attached. Only the caudal vertebræ have hæmal spines.
Fig. 40.—Heterocercal Tail of Lepidosteus.
n, Vertebral column; h, hæmal spines; dn, fulcra; dh, lower fulcra.
In the skull of Lepidosteus the cartilage of the endocranium is still more replaced by ossifications than in Polypterus; those ossifications, moreover, being represented by a greater number of discrete bones; especially the membrane-bones are greatly multiplied: the occipital, for instance, consists of three pieces; the vomer is double as in Polypterus; the maxillary consists of a series of pieces firmly united by suture. The symplectic reaches the lower jaw, so that the articulary is provided with a double joint, viz. for the symplectic and quadrate; the component parts of the lower jaw are as numerous as in reptiles, a dentary, splenial, articulary, angular, supra-angular, and coronary being distinct. The sides of the head are covered with numerous bones, and a præoperculum is developed in front of the gill-cover which, again, consists of an operculum and sub-operculum.
Each hyoid consists of three pieces, of which the middle is the longest, the upper bearing the largest of the three branchiostegals which Lepidosteus possesses; a long and large glossohyal is intercalated between the lower ends of the hyoids. There are five branchial arches, the hindmost of which is modified into a lower pharyngeal; upper pharyngeals are likewise present as in the majority of Teleosteous fishes. No gular plate.
Of the scapulary arch the two halves are separated by a suture in the median line; the membrane-bones are well developed, only a remnant of the primordial cartilage remaining; the supraclavicle is very similar to that of Teleosteous fishes, less so the post-temporal. The base to which the limb is attached is a single osseous plate, supporting on its posterior margin semi-ossified rods in small number, which bear the pectoral rays.
The pubic consists of paired bone, the anterior ends of which overlap each other, the extremity of the right pubis being dorsad to that of the left. The elements representing a tarsus are quite rudimentary and reduced in number (two or three).
The vertebral column of the Amioidei shows unmistakable characters of the Palæichthyic type. The arrangement of its component parts is extremely simple. The centra of the amphicœlous vertebræ are well ossified, but the neural and hæmal arches do not coalesce with the centra, from which they are separated by a thin layer of cartilage. Singularly, not every vertebra has apophyses: in the caudal portion of Amia the vertebræ are alternately provided with them and lack them. The heterocercal condition of the spinous column is well marked: as in the other Holostei the hindmost vertebræ are turned upwards, become smaller and smaller in size, and lose their neural arches, the hæmals remaining developed to the end. Finally, the column terminates in a thin cartilaginous band, which is received between the lateral halves of the fifth or sixth upper caudal ray. Interneurals and interhæmals simple. Only the abdominal vertebræ have parapophyses, with which the ribs are articulated.
The configuration of the skull, and the development and arrangement of its component parts, approaches so much the Teleosteous type that, perhaps, there are greater differences in skulls of truly Teleosteous fishes than between the skulls of Amia and many Physostomi. Externally the cranium is entirely ossified; and the remains of the cartilaginous primordial cranium (which, however, has no vacuity in its roof) can only be seen in a section, and are of much less extent than in many Physostomous fishes. The immovable intermaxillary, the double vomer, the plurality of ossifications representing the articulary, the double articulary cavity of the mandible for junction with the quadrate and symplectic bones, remind us still of similar conditions in the skull of Lepidosteus, but the mobility and formation of the maxillary, the arrangement of the gill-covers, the development of the opercles, the suspensorium, the palate, the insertion of a number of branchiostegals on the long middle hyoid piece, the composition of the branchial framework (with upper and lower pharyngeals), are as in the Teleosteous type. A gular plate replaces the urohyal.
The scapular arch is composed entirely of the membrane-bones found in the Teleostei, and the two sides are loosely united by ligament. The base to which the limb is attached is cartilaginous; short semi-ossified rods are arranged along its hinder margin and bear the pectoral rays.
The skeleton of the hind-limb agrees entirely with that of Lepidosteus.
[T. W. Bridge, The Cranial Osteology of Amia calva; in Journ. Anat. and Physiol. vol. xi.]
In the Teleosteous fishes the spinous column consists of completely ossified amphicœlous vertebræ; its termination is homocercal—that is, the caudal fin appears to be more or less symmetrical, the last vertebra occupying a central position in the base of the fin, and being coalesced with a flat osseous lamella, the hypural (Fig. [23], [70]), on the hind margin of which the fin-rays are fixed. The hypural is but a union of modified hæmapophyses which are directed backwards, and the actual termination of the notochord is bent upwards, and lies along the upper edge of the hypural, hidden below the last rudimentary neural elements. In some Teleosteans, as the Salmonidæ, the last vertebræ are conspicuously bent upwards: in fact, strictly speaking, this homocercal condition is but one of the various degrees of heterocercy, different from that of many Ganoids in this respect only, that the caudal fin itself has assumed a higher degree of symmetry.
The neural and hæmal arches generally coalesce with the centrum, but there are many exceptions, inasmuch as some portion of the arches of a species, or all of them, show the original division.
The vertebræ are generally united with one another by zygapophyses, and frequently similar additional articulations exist at the lower parts of the centra. Parapophyses and ribs are very general, but the latter are inserted on the centra and the base of the processes, and never on their extremities. The point of insertion of the rib, more especially on the anterior vertebræ, may be still higher—viz. at the base of the neural arch, as in Cotylis and allied genera, and even on the top of the neurapophysis, as in Batrachus.
There is a great amount of variation as regards the degree in which the primordial cranium persists; it is always more or less replaced by bone; frequently it disappears entirely, but in some fishes, like the Salmonidæ or Esocidæ, the cartilage persists to the same or even to a greater extent than in the Ganoidei holostei. Added to the bones preformed in cartilage are a great number of membrane-bones. The different kinds of these membrane-bones occur with greater or less constancy throughout this sub-class; they often coalesce with, and are no more separable from, the neighbouring or underlying cartilage-bones. All these bones have been topographically enumerated in Chapter IV.
Many attempts have been made to classify the bones of the Teleosteous skull, according to their supposed relation to each other, or with the view to demonstrate the unity of plan on which the skull has been built; but in all either the one or the other of the following two principles has been followed:—
A. The “vertebral doctrine” starts from the undeniable fact that the skull is originally composed of several segments, each of which is merely the modification of a vertebra. The component parts of such a cranial segment are considered to be homologous to those of a vertebra. Three, four, or five cranial vertebræ have been distinguished, all the various bones of the fully-developed and ossified skull being referred, without distinction as to their origin, to one or the other of those vertebral segments. The idea of the typical unity of the osseous framework of Vertebrates has been worked out with the greatest originality and knowledge of detail, by Owen, who demonstrates that the fish-skull is composed of four vertebræ.
The bones of the fish-skull are, according to him, primarily divisible into those of the neuroskeleton, splanchnoskeleton, and dermoskeleton.
The bones of the neuro- or proper endoskeleton are arranged in a series of four horizontally succeeding segments: the occipital, parietal, frontal, and nasal vertebræ; each segment consisting of an upper (neural) and a lower (hæmal) arch, with a common centre, and with diverging appendages.
The neural arches of the four vertebræ, in their succession from the occiput towards the snout, are:—
1. Epencephalic arch, composed of the occipitals.
2. Mesencephalic arch, composed of basisphenoid, alisphenoid, parietal, and mastoid.
3. Prosencephalic arch, composed of presphenoid, orbito-sphenoid, frontal, and postfrontal.
4. Rhinencephalic arch, composed of vomer, prefrontal, and nasal.
The hæmal arches in the same order of succession are:—
1. Scapular or scapulo-coracoid arch, composed of suprascapula, scapula, and coracoid; its appendage consists of the ulna, radius and carpal.
2. Hyoid or stylo-hyoid arch, composed of stylohyal, epihyal, ceratohyal, basihyal, glossohyal, and urohyal; its appendage is the branchiostegals.
3. Mandibular or tympano-mandibular arch, composed of epi-, meso-, pre-, and hypo-tympanic, and the bones of the lower jaw; its appendage consists of the præoperculum and the other opercles.
4. Maxillary or palato-maxillary arch, composed of palatine, maxillary, and premaxillary; its appendage consists of the pterygoid and entopterygoid.
Parts of the splanchnoskeleton are held to be the ear-capsule or petrosal and the otolite, the eye-capsule or sclerotic, the nose-capsule or “ethmoid” and turbinal; the branchial arches.
The bones of the dermoskeleton are the supratemporals, supraorbitals, suborbitals, and labials.
B. In the second method of classifying the bones of the skull prominence is given to the facts of their different origin as ascertained by a study of their development. The parts developed from the primordial skull, or the cartilaginous case protecting the nervous centre are distinguished from those which enclose and support the commencement of the alimentary canal and the respiratory apparatus, and which, consisting of several arches, are comprised under the common name of visceral skeleton of the skull. Further, a distinction is made between the bones preformed in cartilage and those originating in tegumentary or membranous tissue. It is admitted that the primordial cranium is a coalition of several segments, the number of which is determined by that of the visceral arches, these representing the hæmal arches of the vertebral column; but the membrane-bones are excluded from a consideration of the vertebral division of the primordial skull, as elements originally independent of it, although these additions have entered into special relations to the cartilage-bones.
With these views the bones of the Teleosteous skull are classified thus:—
1. Cartilage-bones of the primordial skull.—The basi-occipital (5 in Figs. [23–26]) has retained the form of a vertebral centrum; it is generally concave behind, the concavity containing remains of the notochord; rarely a rounded articulary head of the first vertebra fits into it, as in Symbranchus, and still more rarely it is provided with such an articulary head (Fistularia); frequently it shows two excavations on its inner surface for the reception of the saccus vestibuli. The exoccipitals (10) are situated on the side of the basi-occipital, and contribute the greater portion of the periphery of the foramen magnum; frequently they articulate with the first vertebra, or meet in the upper median line, so as to exclude the supraoccipital from the foramen magnum. The supraoccipital (8) is intercalated between the exoccipitals, and forms a most prominent part by the median crest, which sometimes extends far forwards on the upper side of the skull, and offers attachment to the dorsal portion of the large lateral muscle of the trunk. When the interior portions of this bone remain cartilaginous, some part of the semicircular canals may be lodged in it.
The region of the skull which succeeds the bones described encloses at least the greater portion of the labyrinth, and its component parts have been named with reference to it by some anatomists.[8] The alisphenoids (11) (Prooticum) form sutures posteriorly with the basi- and exoccipitals, and meet each other in the median line at the bottom of the cerebral cavity; they contribute to the formation of a hollow in which the hypophysis cerebri and the saccus vasculosus are received; in conjunction with the exoccipital it forms another hollow for the reception of the vestibulum; generally it is perforated by the Trigeminal and Facial nerves. The paroccipitals (9) (Epioticum) lodge a portion of the posterior vertical semicircular canal, and form a projection of the skull on each side of the occipital crest, to which a terminal branch of the scapular arch is attached. The Mastoid (12 + 13) (Opisthoticum) occupies the postero-external projection of the head; it encloses a part of the external semicircular canal; is generally coalesced with a membrane-bone, the superficial squamosal, which emits a process for the suspension of the scapular arch, and is frequently, as in the Perch, divided into two separate bones.
The anterior portion of the skull varies greatly as regards form, which is chiefly dependent on the extent of the cerebral cavity; if the latter is advanced far forwards, the lateral walls of the primordial cranium are protected by more developed ossifications than if the cerebral cavity is shortened by the presence of a wide and deep orbit. In the latter case parts which normally form the side of the skull are situated in front of the brain-case, between it and the orbit, and generally reduced in extent, often replaced by membranes; especially the interorbital septum may be reduced to membrane. The most constant ossifications of this part of the skull are the orbitosphenoids (14), which join the upper anterior margin of the alisphenoids. They vary much with regard to their development—they are small in Gadoids; larger in the Perch, Pike, Salmonoids, Macrodon, and the Clupeoids; and very large in Cyprinoids and Siluroids, in which they contribute to the formation of the side of the brain-case. The single Y-shaped Sphenoideum anterius (15) is as frequently absent as present; it forms the anterior margin of the fossa for the hypophysis. Finally, the postfrontal (4) belongs also to this group of cartilage-bones.
The centre of the foremost part of the skull is occupied by the ethmoid (3), which shows great variations as regards its extent and the degree of ossification; it may extend backwards into the interorbital septum, and reach the orbitosphenoids, or may be confined to the extremity of the skull; it may remain entirely cartilaginous, or ossify into a lamina which separates the two orbits and encloses an anterior prolongation of the brain-case, along which the olfactory nerves pass: modifications occurring again in higher vertebrates. A paired ossification attached to the fore-part of the ethmoid is the pre-frontals (2), which form the base of the nasal fossa.
2. Membrane-bones attached to the primordial skull.—To this group belong the parietals (7) and frontals (1). The squamosal (12) has been mentioned above in connection with the mastoid. The supraorbital is always small, and frequently absent. The lower surface of the skull is protected by the basisphenoid (parasphenoid) (6) and the vomer (16), both of which, especially the latter, may be armed with teeth.[9]
3. Cartilage bones of the alimentary portion of the visceral skeleton of the skull.—The suspensorium consists of three cartilage-bones, and affords a base for the opercular apparatus as well as a point of attachment to the hyoid, whilst in front it is connected with the palato-pterygo-palatine arch. They are the hyomandibular (23), symplectic (31), and quadrate (26), connected by means of the metapterygoid (27) with the ecto- (24) and ento-pterygoid (25), the foremost bone of the arch being the palatine (22). All these bones have been sufficiently described above (p. 55), and it remains only to be mentioned that the bones of the palatine arch are but rarely absent, as for instance in Murænophis; and that the symplectic does not extend to the articulary of the mandible, as in Amia and Lepidosteus, though its suspensory relation to the Meckelian cartilage is still indicated by a ligament which connects the two pieces. Of the mandibulary bones the articulary (35) is distinctly part of Meckel’s cartilage. Frequently another portion of cartilage below the articulary remains persistent, or is replaced by a separate membrane-bone, the angular.
4. Membrane-bones of the alimentary portion of the visceral skeleton of the skull.—The suspensorium has one tegumentary bone attached to it, viz. the præoperculum (30); it is but rarely absent, for instance in Murænophis. The premaxillary (17) and maxillary (18) of the Teleostei appear to be also membrane-bones, although they are clearly analogous to the upper labial cartilages of the Sharks. The premaxillaries sometimes coalesce into a single piece (as in Diodon, Mormyrus), or they are firmly united with the maxillaries (as in all Gymnodonts, Serrasalmo, etc.) The relative position and connection of these two bones differs much, and is a valuable character in the discrimination of the various families. In some, the front margin of the jaw is formed by the premaxillary only, the two bones having a parallel position, as it has been described in the Perch (p. 53); in others, the premaxillary is shortened, allowing the maxillary to enter, and to complete, the margin of the upper jaw; and finally, in many no part of the maxillary is situated behind the premaxillary, but the entire bone is attached to the end of the premaxillary, forming its continuation. In the last case the maxillary may be quite abortive. The mobility of the upper jaw is greatest in those fishes in which the premaxillary alone forms its margin. The form of the premaxillary is subject to great variation: the beak of Belone, Xiphias is formed by the prolonged and coalesced premaxillaries. The maxillary consists sometimes of one piece, sometimes of two or three. The principal membrane-bone of the mandible is the dentary (34), to which is added the angular (36) and rarely a smaller one, the splenial or os operculare, which is situated at the inside of the articulary.
5. Cartilage-bones of the respiratory portion of the visceral skeleton of the skull.—With few exceptions all the ossifications of the hyoid and branchial arches, as described above (p. 58), belong to this group.
6. Membrane-bones of the respiratory portion of the visceral skeleton of the skull.—They are the following: the opercular pieces, viz. operculum (28), sub-operculum (32), and interoperculum (33). The last of these is the least constant; it may be entirely absent, and represented by a ligament extending from the mandible to the hyoid. The urohyal (42) which separates the musculi sternohyoidei, and serves for an increased surface of their insertion; and finally the branchiostegals (43), which vary greatly in number, but are always fixed to the cerato- and epi-hyals.
7. Dermal bones of the skull.—To this category are referred some bones which are ossifications of, and belong to, the cutis. They are the turbinals (20), the suborbitals (19), and the supratemporals. They vary much with regard to the degree in which they are developed, and are rarely entirely absent. Nearly always they are wholly or partly transformed into tubes or hollows, in which the muciferous canals with their numerous nerves are lodged. Those in the temporal and scapulary regions are not always developed; on the other hand, the series of those ossicles may be continued on to the trunk, accompanying the lateral line. In many fishes those of the infraorbital ring are much dilated, protecting the entire space between the orbit and the rim of the præoperculum; in others, especially those which have the angle of the præoperculum armed with a powerful spine, the infraorbital ring emits a process towards the spine, which thus serves as a stay or support of this weapon (Scorpænidæ, Cottidæ).
The pectoral arch of the Teleosteous fishes exhibits but a remnant of a primordial cartilage, which is replaced by two ossifications,[10] the coracoid (51) and scapula (52); they offer posteriorly attachment to two series of short rods, of which the proximal are nearly always ossified, whilst the distal frequently remain small cartilaginous nodules hidden in the base of the pectoral rays. The bones, by which this portion is connected with the skull, are membrane-bones, viz. the clavicle (49), with the postclavicle (49 + 50), the supraclavicle (47), and post-temporal (46). The order of their arrangement in the Perch has been described above (p. 59). However, many Teleosteous fish lack pectoral fins, and in them the pectoral arch is frequently more or less reduced or rudimentary, as in many species of Murænidæ. In others the membrane-bones are exceedingly strong, contributing to the outer protective armour of the fish, and then the clavicles are generally suturally connected in the median line. The postclavicula and the supraclavicula may be absent. Only exceptionally the shoulder-girdle is not suspended from the skull, but from the anterior portion of the spinous column (Symbranchidæ, Murænidæ, Notacanthidæ). The number of basal elements of each of the two series never exceeds five, but may be less; and the distal series is absent in Siluroids.
The pubic bones of the Teleosteous fishes undergo many modifications of form in the various families, but they are essentially of the same simple type as in the Perch.
CHAPTER V.
MYOLOGY.
In the lowest vertebrate, Branchiostoma, the whole of the muscular mass is arranged in a longitudinal band running along each side of the body; it is vertically divided into a number of flakes or segments (myocommas) by aponeurotic septa, which serve as the surfaces of insertion to the muscular fibres. But this muscular band has no connection with the notochord except in its foremost portion, where some relation has been formed to the visceral skeleton. A very thin muscular layer covers the abdomen.
Also in the Cyclostomes the greatest portion of the muscular system is without direct relation to the skeleton, and, again, it is only on the skull and visceral skeleton where distinct muscles have been differentiated for special functions.
To the development of the skeleton in the more highly organised fishes corresponds a similar development of the muscles; and the maxillary and branchial apparatus, the pectoral and ventral fins, the vertical fins, and especially the caudal, possess a separate system of muscles. But the most noteworthy is the muscle covering the sides of the trunk and tail (already noticed in Branchiostoma), which Cuvier described as the “great lateral muscle,” and which, in the higher fishes, is a compound of many smaller segments, corresponding in number with the vertebræ. Each lateral muscle is divided by a median longitudinal groove into a dorsal and ventral half; the depression in its middle is filled by an embryonal muscular substance which contains a large quantity of fat and blood-vessels, and therefore differs from ordinary muscle by its softer consistency, and by its colour which is reddish or grayish. Superficially the lateral muscle appears crossed by a number of white parallel tendinous zig-zag stripes, forming generally three angles, of which the upper and lower point backwards, the middle one forwards. These are the outer edges of the aponeurotic septa between the myocommas. Each septum is attached to the middle and the apophyses of a vertebra, and, in the abdominal region, to its rib; frequently the septa receive additional support by the existence of epipleural spines. The fibres of each myocomma run straight and nearly horizontally from one septum to the next; they are grouped so as to form semiconical masses, of which the upper and lower have their apices turned backwards, whilst the middle cone, formed by the contiguous parts of the preceding, has its apex directed forward; this fits into the interspace between the antecedent upper and lower cones, the apices of which reciprocally enter the depressions in the succeeding segment, whereby all the segments are firmly locked together (Owen).
In connection with the muscles reference has to be made to the Electric organs with which certain fishes are provided, as it is more than probable, not only from the examination of peculiar muscular organs occurring in the Rays, Mormyrus, and Gymnarchus (the function of which is still conjectural), but especially from the researches into the development of the electric organ of Torpedo, that the electric organs have been developed out of muscular substance. The fishes possessing fully developed electric organs, with the power of accumulating electric force and communicating it in the form of shocks to other animals, are the electric Rays (Torpedinidæ), the electric Sheath-fish of tropical Africa (Malapterurus), and the electric Eel of tropical America (Gymnotus). The structure and arrangement of the electric organ is very different in these fishes, and will be subsequently described in the special account of the several species.
The phenomena attending the exercise of this extraordinary faculty also closely resemble muscular action. The time and strength of the discharge are entirely under the control of the fish. The power is exhausted after some time, and it needs repose and nourishment to restore it. If the electric nerves are cut and divided from the brain the cerebral action is interrupted, and no irritant to the body has any effect to excite electric discharge; but if their ends be irritated the discharge takes place, just as a muscle is excited to contraction under similar circumstances. And, singularly enough, the application of strychnine causes simultaneously a tetanic state of the muscles and a rapid succession of involuntary electric discharges. The strength of the discharges depends entirely on the size, health, and energy of the fish: an observation entirely agreeing with that made on the efficacy of snake-poison. Like this latter, the property of the electric force serves two ends in the economy of the animals which are endowed with it; it is essential and necessary to them for overpowering, stunning, or killing the creatures on which they feed, whilst incidentally they use it as the means of defending themselves from their enemies.
CHAPTER VI.
NEUROLOGY.
The most simple condition of the nervous central organ known in Vertebrates is found in Branchiostoma. In this fish the spinal chord tapers at both ends, an anterior cerebral swelling, or anything approaching a brain, being absent. It is band-like along its middle third, and groups of darker cells mark the origins of the fifty or sixty pairs of nerves which accompany the intermuscular septa, and divide into a dorsal and ventral branch, as in other fishes. The two anterior pairs pass to the membranous parts above the mouth, and supply with nerve filaments a ciliated depression near the extremity of the fish, which is considered to be an olfactory organ, and two pigment spots, the rudiments of eyes. An auditory organ is absent.
The spinal chord of the Cyclostomes is flattened in its whole extent, band-like, and elastic; also in Chimæra it is elastic, but flattened in its posterior portion only. In all other fishes it is cylindrical, non-ductile, and generally extending along the whole length of the spinal canal. The Plectognaths offer a singular exception in this respect that the spinal chord is much shortened, the posterior portion of the canal being occupied by a long cauda equina; this shortening of the spinal chord has become extreme in the Sun-fish (Orthagoriscus), in which it has shrunk into a short and conical appendage of the brain. Also in the Devil-fish (Lophius) a long cauda equina partly conceals the chord which terminates on the level of about the twelfth vertebra.
The brain of fishes is relatively small; in the Burbot (Lota) it has been estimated to be 1/720th part of the weight of the entire fish, in the Pike the 1/1305th part, and in the large Sharks it is relatively still smaller. It never fills the entire cavity of the cranium; between the dura mater which adheres to the inner surface of the cranial cavity, and the arachnoidea which envelops the brain, a more or less considerable space remains, which is filled with a soft gelatinous mass generally containing a large quantity of fat. It has been observed that this space is much less in young specimens than in adult, which proves that the brain of fishes does not grow in the same proportion as the rest of the body; and, indeed, its size is nearly the same in individuals of which one is double the bulk of the other.
Fig. 41.—Brain of Perch.
I. Upper aspect. II. Lower aspect.
a, cerebellum; b, optic lobes; c, hemispheres; e, lobi inferiores; f, hypophysis; g, lobi posteriores; i, Olfactory lobes; n, N. opticus; o, N. olfactorius; p, N. oculo-motorius; q, N. trochlearis; r, N. trigeminus; s, N. acusticus; t, N. vagus; u, N. abducens; v, Fourth ventricle.
The brain of Osseous fishes (Fig. [41]) viewed from above shows three protuberances, respectively termed prosencephalon, mesencephalon, and metencephalon, the two anterior of which are paired, the hindmost being single. The foremost pair are the hemispheres, which are solid in their interior, and provided with two swellings in front, the olfactory lobes. The second pair are the optic lobes, which generally are larger than the hemispheres, and succeeded by the third single portion, the cerebellum. In the fresh state the hemispheres are of a grayish colour, and often show some shallow depressions on their surface; a narrow commissure of white colour connects them with each other. The optic lobes possess a cavity (ventriculus lobioptici), at the bottom of which some protuberances of variable development represent the corpora quadrigemina of higher animals. On the lower surface of the base of the optic lobes, behind the crura cerebri, two swellings are observed, the lobi inferiores, which slightly diverge in front for the passage of the infundibulum, from which a generally large hypophysis or pituitary gland is suspended. The relative size of the cerebellum varies greatly in the different osseous fishes: in the Tunny and Silurus it is so large as nearly to cover the optic lobes; sometimes distinct transverse grooves and a median longitudinal groove are visible. The cerebellum possesses in its interior a cavity which communicates with the anterior part of the fourth ventricle. The medulla oblongata is broader than the spinal chord, and contains the fourth ventricle, which forms the continuation of the central canal of the spinal chord. In most fishes a perfect roof is formed over the fourth ventricle by two longitudinal pads, which meet each other in the median line (lobi posteriores), and but rarely it remains open along its upper surface.
The brain of Ganoid fishes shows great similarity to that of the Teleostei; however, there is considerable diversity of the arrangement of its various portions in the different types. In the Sturgeons and Polypterus (Fig. [42]) the hemispheres are more or less remote from the mesencephalon, so that in an upper view the crura cerebri, with the intermediate entrance into the third ventricle (fissura cerebri magna), may be seen. A vascular membranous sac, containing lymphatic fluid (epiphysis), takes its origin from the third ventricle, its base being expanded over the anterior interspace of the optic lobes, and the apex being fixed to the cartilaginous roof of the cranium. This structure is not peculiar to the Ganoids, but found in various stages of development in Teleosteans, marking, when present, the boundary between prosencephalon and mesencephalon. The lobi optici are essentially as in Teleosteans. The cerebellum penetrates into the ventriculus lobi optici, and extends thence into the open sinus rhomboidalis. At its upper surface it is crossed by a commissure formed by the corpora restiformia of the medulla.
Fig. 42.—Brain of Polypterus. (After Müller.)
I., Upper; II., Lateral; III., Lower aspect.
a, Medulla; b, corpora restiformia; c, cerebellum; d, lobi optici; e, hypophysis; f, fissura cerebri magna; g, nervus opticus; g’, chiasma; h, hemispheres; i, lobus olfactorius; k, sinus rhomboidalis (fourth ventricle).
As regards external configuration, the brain of Lepidosteus and Amia approach still more the Teleosteous type. The prosencephalon, mesencephalon, and metencephalon are contiguous, and the cerebellum lacks the prominent transverse commissure at its upper surface. The sinus rhomboidalis is open.
The brain of the Dipnoi shows characters reminding us of that of the Ganoids as well as the Chondropterygians, Ceratodus agreeing with Protopterus in this respect, as in most other points of its organisation. The hemispheres form the largest part of the brain; they are coalescent, as in Sharks, but possess two lateral ventricles, the separation being externally indicated by a shallow median groove on the upper surface. The olfactory lobes take their origin from the upper anterior end of the hemispheres. Epiphysis and hypophysis well developed. The lobi optici are very small, and remote from the prosencephalon, their division into the lateral halves being indicated by a median groove only. The cerebellum is very small, overlying the front part of the sinus rhomboidalis.
Fig. 43.—Brain of Carcharias. (After Owen.)
ac, Nerv. acusticus; b, corpus restiforme; c, cerebellum; d, lobus opticus; e, hypophysis; g, nervus opticus; h, hemisphere; i, lobus olfactorius; i’, olfactory pedicle; k, nerv. olfactorius; l, epiphysis; m, nerv. oculo-motorius; tr, nerv. trigeminus; v, nerv. vagus.
The brain of Chondropterygians (Fig. [43]) is more developed than that of all other fishes, and distinguished by well-marked characters. These are, first, the prolongation of the olfactory lobes into more or less long pedicles, which dilate into great ganglionic masses, where they come into contact with the olfactory sacs; secondly, the space which generally intervenes between prosencephalon and mesencephalon, as in some Ganoids; thirdly, the large development of the metencephalon.
The hemispheres are generally large, coalescent, but with a median, longitudinal, dividing groove. Frequently their surface shows traces of gyrations, and when they are provided with lateral ventricles, tubercles representing corpora striata may be observed. The olfactory pedicles take their origin from the side of the hemispheres, and are frequently hollow, and if so, their cavity communicates with the ventricle of the hemisphere. The optic lobes are generally smaller than the hemispheres, coalescent, and provided with an upper median groove like the prosencephalon. At their base a pair of lobi inferiores are constant, with the hypophysis and sacsus vasculosus (a conglomeration of vascular loops without medullary substance) between them.
The cerebellum is very large, overlying a portion of the optic lobes and of the sinus rhomboidalis, and is frequently transversely grooved. The side-walls of the fourth ventricle, which are formed by the corpora restiformia, are singularly folded, and appear as two pads, one on each side of the cerebellum (lobi posteriores s. lobi nervi trigemini).
Fig. 44.—Brain of Bdellostoma. (Enlarged, after Müller.)
I., Upper; II., Lower aspect. Letters as in Fig. [45].
Fig. 45.—Brain of Petromyzon. (Enlarged, after Müller.)
I., Upper; II., Lower aspect.
a, Medulla oblongata; ac, nerv. acusticus; b, corpus restiforme or rudimentary cerebellum; d, lobus ventriculi tertii; d’, entrance into the third ventricle; c, hypophysis; fa, nerv. facialis; g, nerv. opticus; h, hemisphere; hy, nerv. hypoglossus (so named by Müller); i, lobus olfactorius; k, sinus rhomboidalis; l, epiphysis; m, nerv. oculo-motorius; q, corpora quadrigemina; tr, nerv. trigeminus; tro, nerv. trochlearis; v, nerv. vagus.
The brain of the Cyclostomes (Figs. [44], [45]) represents a type different from that of other fishes, showing at its upper surface three pairs of protuberances in front of the cerebellum; they are all solid. Their homologies are not yet satisfactorily determined, parts of the Myxinoid brain having received by the same observers determinations very different from those given to the corresponding parts of the brain of the Lampreys. The foremost pair are the large olfactory tubercles, which are exceedingly large in Petromyzon. They are followed by the hemispheres, with a single body wedged in between their posterior half; in Petromyzon, at least, the vascular tissue leading to an epiphysis seems to be connected with this body. Then follows the lobus ventriculi tertii, distinctly paired in Myxinoids, less so in Petromyzon. The last pair are the corpora quadrigemina. According to this interpretation the cerebellum would be absent in Myxinoids, and represented in Petromyzon by a narrow commissure only (Fig. [45], b), stretching over the foremost part of the sinus rhomboidalis. In the Myxinoids the medulla oblongata ends in two divergent swellings, free and obtuse at their extremity, from which most of the cerebral nerves take their origin.
The Nerves which supply the organs of the head are either merely continuations or diverticula of the brain-substance, or proper nerves taking their origin from the brain, or receiving their constituent parts from the foremost part of the spinal chord. The number of these spino-cerebral nerves is always less than in the higher vertebrates, and their arrangement varies considerably.
A. Nerves which are diverticula of the brain (Figs. [41–45]).
The olfactory nerves (first pair) always retain their intimate relation to the hemispheres, the ventricles of which are not rarely continued into the tubercle or even pedicle of the nerves. The different position of the olfactory tubercle has been already described as characteristic of some of the orders of fishes. In those fishes in which the tubercle is remote from the brain, the nerve which has entered the tubercle as a single stem leaves it split up into several or numerous branches, which are distributed in the nasal organ. In the other fishes it breaks up into branchlets spread into a fan-like expansion at the point, where it enters the nasal cavity. The nerve always passes out of the skull through the ethmoid.
The optic nerves (second pair) vary in size, their strength corresponding to the size of the eye; they take their origin from the lobi optici, the development of which again is proportionate to that of the nerves. The mutual relation of the two nerves immediately after their origin is very characteristic of the sub-classes of fishes. In the Cyclostomes they have no further connection with each other, each going to the eye of its own side.[11] In the Teleostei they simply cross each other (decussate), so that the one starting from the right half of the brain goes to the left eye and vice versa. Finally, in Palæichthyes the two nerves are fused together, immediately after their origin, into a chiasma. The nerve is cylindrical for some portion of its course, but in most fishes gradually changes this form into that of a plaited band, which is capable of separation and expansion. It enters the bulbus generally behind and above its axis. The foramen through which it leaves the skull of Teleostei is generally in a membranous portion of its anterior wall, or, where ossification has taken place, in the orbito-sphenoid.
B. Nerves proper taking their origin from the brain
(Figs. [41–45]).
The Nervus oculorum motorius (third pair) takes its origin from the Pedunculus cerebri, close behind the lobi inferiores; it escapes through the orbito-sphenoid, or the membrane replacing it, and is distributed to the musculi rectus superior, rectus internus, obliquus inferior, and rectus inferior. Its size corresponds to the development of the muscles of the eye. Consequently it is absent in the blind Amblyopsis, and the Myxinoids. In Lepidosiren the nerves supplying the muscles of the eye have no independent origin, but are part of the ophthalmic division of the Trigeminus. In Petromyzon these muscles are supplied partly from the Trigeminus, partly by a nerve representing the Oculo-motor and Trochlearis, which are fused into a common trunk.
The Nervus trochlearis (fourth pair), if present with an independent origin, is always thin, taking its origin on the upper surface of the brain from the groove between lobus opticus and cerebellum; it goes to the Musculus obliquus superior of the eye.
C. Nerves taking their origin from the Medulla oblongata (Figs. [41–45]).
The Nervus abducens (sixth pair) issues on the lower surface of the brain, taking its origin from the anterior pyramids of the Medulla oblongata, and supplies the Musculus rectus externus of the eye, and the muscle of the nictitating membrane of Sharks.
The Nervus trigeminus (fifth pair) and the Nervus facialis (seventh pair) have their origins close together, and enter into intimate connection with each other. In the Chondropterygians and most Teleostei the number of their roots is four, in the Sturgeons five, and in a few Teleostei three. When there are four, the first issues immediately below the cerebellum from the side of the Medulla oblongata; it contains motory and sensory elements for the maxillary and suspensorial muscles, and belongs exclusively to the trigeminal nerve. The second root, which generally becomes free a little above the first, supplies especially the elements for the Ramus palatinus, which sometimes unites with parts of the Trigeminal, sometimes with the Facial nerve. The third root, if present, is very small, and issues immediately in front of the acustic nerve, and supplies part of the motor elements of the facial nerve. The fourth root is much stronger, sometimes double, and its elements pass again partly into the Trigeminal, partly into the Facial nerve. On the passage of these stems through the skull (through a foramen or foramina in the alisphenoid) they form a ganglionic plexus, in which the palatine ramus and the first stem of the Trigeminus generally possess discrete ganglia. The branches which issue from the plexus and belong exclusively to the Trigeminus, supply the organs and integuments of the frontal, ophthalmic, and nasal regions, and the upper and lower jaws with their soft parts. The Facial nerve supplies the muscles of the gill-cover and suspensorium, and emits a strong branch accompanying the Meckelian cartilage to the symphysis, and another for the hyoid apparatus.
The Nervus acusticus (eighth pair) is strong, and takes its origin immediately behind, and in contact with, the last root of the seventh pair.
The Nervus glossopharyngeus (ninth pair)[12] takes its origin between the roots of the eighth and tenth nerves, and issues in Teleostei from the cranial cavity by a foramen of the exoccipital. In the Cyclostomes and Lepidosiren it is part of the Nervus vagus. It is distributed in the pharyngeal and lingual regions, one branch supplying the first branchial arch. After having left the cranial cavity it swells into a ganglion, which in Teleostei is always in communication with the sympathic nerve.
The Nervus vagus or pneumogastricus (tenth pair) rises in all Teleostei and Palæichthyes with two discrete strong roots: the first constantly from the swellings of the corpora restiformia, be they thinner or thicker and overlying the sinus rhomboidalis, or be they developed into lateral plaited pads, as in Acipenser and Chondropterygians. The second much thicker root rises from the lower tracts of the medulla oblongata. Both stems leave the cranial cavity by a common foramen, situated in Teleosteous fishes in the exoccipital; and form ganglionic swellings, of which those of the lower stem are the more conspicuous. The lower stem has mixed elements, motory as well as sensory, and is distributed to the muscles of the branchial arches and pharynx, the œsophagus and stomach; it sends filaments to the heart and to the air-bladder where it exists. The first (upper) stem forms the Nervus lateralis. This nerve, which accompanies the lateral mucous system of the trunk and tail, is either a single longitudinal stem, gradually becoming thinner behind, running superficially below the skin (Salmonidæ, Cyclopterus), or deeply between the muscles (Sharks, Chimæra), or divided into two parallel branches (most Teleostei): thus in the Perch there are two branches on each side, the superficial of which supplies the lateral line, whilst the deep-seated branch communicates with the spinal nerves and supplies the septa between the myocommas and the skin. In fishes which lack the lateral muciferous system and possess hard integuments, as the Ostracions, the lateral nerve is more or less rudimentary. It is entirely absent in Myxinoids, but the gastric branches of the Vagus are continued, united as a single nerve, along the intestine to the anus.
No fish possesses a Nervus accessorius. Also a separate Nervus hypoglossus (twelfth pair)[13] is absent, but elements from the first spinal nerve are distributed in the area normally supplied by this nerve in higher vertebrates.
The number of Spinal nerves corresponds to that of the vertebræ, through or between which they pass out. Each nerve has two roots, an anterior and posterior, the former of which has no ganglion, and exclusively contains motor elements. The posterior or dorsal has a ganglionic enlargement, and contains sensory elements only. After leaving the vertebral canal each spinal nerve usually divides into a dorsal and ventral branch. The Gadoids show that peculiarity that each of the posterior roots of some or many of the spinal nerves possesses two separate threads, each of which has a ganglion of its own; the one of these threads joins the dorsal and the other the ventral branch. In fishes in which the spinal chord is very short, as in Plectognaths, Lophius, the roots of the nerves are extremely long, forming a thick Cauda equina. The additional function which the (five) anterior spinal nerves of Trigla have to perform in supplying the sensitive pectoral appendages and their muscles has caused the development of a paired series of globular swellings of the corresponding portion of the spinal chord. A similar structure is found in Polynemus.
Fig. 46.
Brain and anterior portion of the spinal chord of Trigla (Gurnard), showing the globular swellings at the base of the anterior spinal nerves.
A Sympathic nervous system appears to be absent in Branchiostoma, and has not yet been clearly made out in Cyclostomes. It is well developed in the Palæichthyes, but without cephalic portion. This latter is present in all Osseous fishes, in which communication of the Sympathic has been found to exist with all cerebral nerves, except the olfactory, optic, and acustic. The sympathic trunks run along each side of the aorta and the back of the abdomen into the hæmal canal; communicate in their course with the ventral branches of each of the spinal nerves; and, finally, often blend together into a common trunk beneath the tail. At the points of communication with the cerebral and spinal nerves frequently ganglia are developed, from which nerves emerge which are distributed to the various viscera.
CHAPTER VII.
THE ORGANS OF SENSE.
Characteristic of the Organ of Smell in Fishes is that it has no relation whatever to the respiratory function, with the exception of the Dipnoi, in which possibly part of the water received for respiration passes through the nasal sac.
The olfactory organ is single in Branchiostoma and the Cyclostomes. In the former a small depression on the front end of the body, clothed with a ciliated epithelium, is regarded as a rudimentary organ of smell. In the adult Petromyzon a membranous tube leads from the single opening on the top of the head into the cartilaginous olfactory capsule, the inside of which is clothed by membranes prolonged into a posterior blind tube (Fig. [30], s), which penetrates the cartilaginous roof of the palate, but not the mucous membrane of the buccal cavity. In the Myxinoids the outer tube is strengthened by cartilaginous rings like a trachea; the capsule is lined by a longitudinally folded pituitary membrane, and the posterior tube opens backwards on the roof of the mouth; the opening is provided with a valve.
In all other Fishes the organ of smell is double, one being on each side; it consists of a sac lined with a pituitary membrane, and without, or with one or two, openings. The position of these openings is very different in the various orders or suborders of Fishes.
In the Dipnoi the nasal sac opens downwards by two wide openings which are within the boundaries of the cavity of the mouth. The pituitary membrane is transversely folded, the transverse folds being divided by one longitudinal fold. The walls of the sac are strengthened by sundry small cartilages.
Also in Chondropterygians the openings, of which there is one to each sac, are on the lower part of the snout, and in the Rays, Holocephali, and some Sharks, each extends into the cleft of the mouth. The openings are protected by valvular flaps, supported by small cartilages, and moved by muscles, whence it may be concluded that these fishes are able to scent (actively) as well as to smell (passively).
Fig. 47.—Nostrils of Raia lemprieri, with nasal flaps reverted.
In the majority of Teleostei the olfactory capsules are lateral or superior on the snout, covered externally by the skin, each usually pierced by two openings, which are either close together, or more or less remote from each other; the posterior is generally open, the anterior provided with a valve or tube. In the Chromides and Labroidei ctenoidei a single opening only exists for each sac. In the Murænidæ the two openings of each side are either superior, or lateral, or labial, that is, they are continued downwards and pierce the margin of the upper lip. In many Tetrodonts nasal openings are absent, and replaced by a conical papilla, in which the olfactory nerve terminates.
It is certain that fishes possess the faculty of perceiving odours, and that various scents attract or repel them. A mangled carcase or fresh blood attracts Sharks as well as the voracious Serrasal monoids of the South American rivers. There is no reason to doubt that the seat of that perception is in the olfactory sac; and it may be reasonably conjectured that its strength depends mainly on the degree of development indicated by the number and extent of the interior folds of the pituitary membrane.
Organ of Sight.—The position, direction, and dimensions of the eyes of fishes vary greatly. In some they have an upward aspect, and are often very close together; in others they are lateral, and in a few they are even directed downwards. The Flat-fishes represent the extraordinary anomaly that both eyes are on the same side of the head, and rarely on the same level, one being generally placed more forward than the other. In certain species of marine fishes the eyes are of an extraordinary size, a peculiarity indicating that the fish either lives at a great depth, to which only a small proportion of the rays of light penetrate, or that it is of nocturnal habits. In fishes which have descended to such great depths that no rays whatever can reach them, or in freshwater fishes living in caves, or in species which grovel and live constantly in mud, the eyes are more or less aborted, sometimes quite rudimentary, and covered by the skin. In very few this organ appears to be entirely absent. In some Gobioids and Trachinoids (Periophthalmus, Boleophthalmus, Uronoscopus, etc.) the eyes, which are on the upper side of the head, can be elevated and depressed at the will of the fish. In the range of their vision and acuteness of sight, Fishes are very inferior to the higher classes of Vertebrates, yet at the same time it is evident that they perceive their prey or approaching danger from a considerable distance; and it would appear that the visual powers of a Periophthalmus, when hunting insects on mud-flats of the tropical coasts, are quite equal to that of a frog. Again, the discrimination with which fishes sometimes prefer one colour or kind of artificial fly to another affords sufficient evidence that the vision, at least of certain species is by no means devoid of clearness and precision.
The eye of Branchiostoma is of the most rudimentary condition. It is simply a minute speck coated by dark pigment, and receiving the end of a short nerve. In Myxinoids the minute rudiment of the eye is covered by the skin and muscles. This is also the case in many of the blind Teleosteous fishes; however, whilst in the former fishes the organ of sight has not attained to any degree of development, the rudimentary eye of blind Teleostei is a retrogressive formation, in which often a lens and other portions of the eye can be recognised. In fishes with a well-developed eye it is imbedded in a layer of gelatinous and adipose substance, which covers the cavity of the orbit. A lacrymal gland is absent. In the orbit of one fish only, Chorismodentex, an organ has been found which can be compared to a saccus lacrymalis. It is a round, blind, wide sac, of the size of a pea, situated below the anterior corner of the orbit, between the maxillary bone and the muscles of the cheek, communicating by a rather wide foramen with the orbital cavity. The membrane by which it is formed is continuous with that coating the orbita. In the Chondropterygians the eyeball is supported by and moves on a cartilaginous peduncle of the orbital wall. In the majority of Teleosteans, and in Acipenser, a fibrous ligament attaches the sclerotic to the wall of the orbit. The proper muscles of the eyeball exist in all fishes, and consist of the four Musculi recti and the two M. obliqui. In many Teleostei the former rise from a subcranial canal, the origin of the M. rectus externus being prolonged farthest backwards. The Recti muscles are extraordinarily long in the Hammerheaded Sharks, in which they extend from the basis cranii along the lateral prolongations of the head to the eyes, which are situated at the extremities of the hammer.
In all fishes the general integument of the head passes over the eye, and becomes transparent where it enters the orbit; sometimes it simply passes over the orbit, sometimes it forms a circular fold. The anterior and posterior portions may be especially broad and the seat of an adipose deposit (adipose eyelids), as in Scomber, Caranx, Mugil, etc. In many of these fishes the extent of these eyelids varies with the seasons; during the spawning season they are so much loaded with fat as nearly to hide the whole eye. Many Sharks possess a nictitating membrane, developed from the lower part of the palpebral fold, and moved by a proper set of muscles.
Fig. 48.
Vertical section through eye of Xiphias. (After Owen.)
co, Cornea; sc, sclerotica; o, nervus opticus; c, sclerotic capsule; a, membrana argentea; v, membrana vasculosa; u, membrana uvea; ch, choroid gland; r, retina; f, processus falciformis; h, humor vitreus; l, lens; i, iris.
The form of the bulbus (Fig. [48]) is subhemispherical, the cornea (co) being flat. If it were convex, as in higher Vertebrates, it would be more liable to injury; but being level with the side of the head the chances of injury by friction are diminished. The sclerotica (sc) is cartilaginous in Chondropterygians and Acipensers, fibrous and of varying thickness in Teleosteans, in the majority of which it is supported by a pair of cartilaginous or ossified hemispheroid cups (c). In a few fishes, as in Ceratodus, Xiphias, the cups are confluent into one cup, which possesses a foramen behind to allow the passage of the optic nerve (o). The cornea of Anableps shows an unique peculiarity. It is crossed by a dark horizontal stripe of the conjunctiva, dividing it into an upper and lower portion; also the iris is perforated by two pupils. This fish is observed to swim frequently with half of its head out of the water, and it is a fact that it can see out of the water as well as in it.
The membranes situated between the sclerotica and retina are collectively called choroidea, and three in number. The one in immediate contact with the sclerotic, and continued upon the iris, is by no means constantly present; it is the membrana argentea (a), and composed of microscopical crystals reflecting a silvery or sometimes golden lustre. The middle layer is the membrana vasculosa s. halleri (v), the chief seat of the ramifications of the choroid vessels; the innermost layer is the membrana ruyscheana or uvea (u), which is composed of hexagonal pigment-cells, usually of a deep brown or black colour.
In many Teleostei a rete mirabile surrounds the entry of the optic nerve; it is situated between the membrana argentea and vasculosa, and called the choroid gland (ch). It receives its arterial blood from the artery issuing from the pseudobranchia; the presence of a choroid gland always being combined with that of a pseudobranchia. Teleosteans without pseudobranchia lack a choroid gland. In the Palæichthyes, on the other hand, the pseudobranchia is present and a choroid gland absent.
The iris (i) is merely the continuation of the choroid membrane; its capability of contracting and expanding is much more limited than in higher Vertebrates. The pupil is generally round, sometimes horizontally or vertically elliptical, sometimes fringed. In the Rays and Pleuronectidæ a lobe descends from the upper margin of the pupil, and the outer integument overlying this lobe is coloured and non-transparent; a structure evidently preventing light from entering the eye from above.
In most Teleostei a fold of the Choroidea, called the Processus falciformis (f), extends from the vicinity of the entrance of the optic nerve to the lens. It seems to be constantly absent in Ganoids.
The retina (r) is the membrane into which the optic nerve penetrates, and in which its terminal filaments are distributed. It consists of several layers (Fig. [49]). The outermost is an extremely delicate membrane (a), followed by a layer of nerve-cells (b), from which the terminal filaments issue, passing through several granular strata (c, d, e), on which the innermost stratum rests. This stratum is composed of cylindrical rods (f) vertically arranged, between which twin fusiform corpuscles (g) are intercalated. This last layer is thickly covered with a dark pigment. The retina extends over a portion of the iris, and a well-defined raised rim runs along its anterior margin.
Fig. 49.—Vertical section of the Retina of the Perch, magn. X 350.
The vitreous humour (Fig. [48], h) which fills the posterior cavity of the eyeball, is of a firmer consistency than in the higher Vertebrates. The lens is spherical, or nearly so; firm, denser towards the centre, and lies in a hollow of the vitreous humour. When a falciform process is present, it is with one end attached to the lens, which is thus steadied in its position. It consists of concentric layers consisting of fibres, which in the nucleus of the body have marginal teeth, by which they are interlocked together. In Petromyzon this serrature is absent, or but faintly indicated.
Fig. 50.—Interlocking fibres of lens, highly magnified.
The anterior cavity of the eye is very small in Fishes, in consequence of the small degree of convexity of the cornea; the quantity of the aqueous humour, therefore, is very small, just sufficient to float the free border of the iris; and the lessened refractive power of the aqueous humour is compensated by the greater convexity of the lens.
Organ of Hearing.—No trace of an organ of hearing has been found in Branchiostoma. In the Cyclostomes the labyrinth is enclosed in externally visible cartilaginous capsules laterally attached to the skull; it consists of a single semicircular canal in the Myxinoids, whilst the Petromyzontes possess two semicircular canals with a vestibulum.
In all other fishes the labyrinth consists of a vestibule and three semicircular canals, the vestibule dilating into one or more sacs which contain the otoliths. A tympanum, tympanic cavity, and external parts, are entirely absent in the class of fishes.
In the Chondropterygians and Dipnoi, the labyrinth is enclosed in the cartilaginous substance of the skull. In the former the excavation in the cartilage is larger than the membranous labyrinth, but nearly corresponds to it in form; the part which receives the membranous vestibulum is called Vestibulum cartilagineum, from which a canal issues and penetrates to the surface of the skull, where it is closed by the skin in Sharks, but opens by a minute foramen in Rays. The otolithic contents are soft and chalklike.
In the Holocephali part of the labyrinth is enclosed in the cartilage of the skull, another part being in the cranial cavity, as in Ganoids and Teleosteans. The membranous vestibulum is continued by a canal to a single opening in the roof of the skull, from which two smaller canals are continued to two small foramina in the skin covering the occipital region.
In the Teleosteans the sac which contains the otoliths lies on each side of the base of the cranial cavity and is often divided by a septum into two compartments of unequal size, each containing a firm and solid otolith; these bodies (Fig. [51]), possess indented margins, frequently other impressions and grooves, in which nerves from the N. acusticus are lodged; they vary much in size and form, but in both respects show a remarkable constancy in the same kind of fishes. The vestibule is outwards in contact with the osseous side wall of the skull, inwards with the metencephalon and medulla oblongata; it contains another firm concretion, and opens by five foramina into the three semicircular canals. The terminations of the acustic nerve are distributed over the vestibular concretion and the ampulliform ends (Fig. [52] p) of the semicircular canals, without being continued into the latter, which are filled with fluid. The semicircular canals (Fig. [52] g), are sometimes lodged in the cranial bones, sometimes partly free in the cranial cavity. Many Teleostei have fontanelles in the roof of the skull, closed by skin or very thin bone only at the place where the auditory organ approaches the surface, by which means sonorous undulations must be conducted with greater ease to the ear.
Fig. 51.—Otolith of Haddock (Gadus æglefinus). I. Outer, II. Inner aspect.
In many Teleostei a most remarkable relation obtains between the organ of hearing and the air-bladder. In the most simple form this connection is established in Percoids and the allied families, in which the two anterior horns of the air-bladder are attached to fontanelles of the occipital region of the skull, the vestibulum occupying the opposite side of the membrane by which the fontanelle is closed. The condition is similar, but more complicated in many Clupeoids. The anterior narrow end of the air-bladder is produced into a canal at the base of the skull, and divided into two very narrow branches, which again bifurcate and terminate in a globular swelling. An appendage of the vestibulum meets the anterior of these swellings, and comes into close contact with it. Besides, the two vestibules communicate with each other by a transverse canal, crossing the cranial cavity below the brain.
Fig. 52.—Communication between auditory organ and air-bladder in the Carp. (After E. H. Weber.)
a, Basisphenoid; b, Occipital; c, Supraoccipital; d, Exoccipital; e, Paroccipital; f, Alisphenoid; g, Neural arch of first vertebra; h, i, k, second, third, and fourth vertebra; h’, i’, Parapophyses of second and third vertebra; i", process of the third vertebra for the attachment of the air-bladder; k, l, m, Chain of ossicles; n, Air-bladder; o, vestibulum; p, p, Ampullæ; q, q, Canales semicirculares; r, Sinus impar.
The connection is effected by means of a chain of ossicles in Siluridæ, Characinidæ, Cyprinidæ and Gymnotidæ. A canal issues from the communication between vestibule and its sac, and meeting that from the other side forms with it a common sinus impar (Fig. [52], r), lodged in the substance of the basi-occipital; this communicates on each side by a small orifice with two subspherical atria, on the body of the atlas, close to the foramen magnum. Each atrium is supported externally by a small bone (m); a third larger bone (k) completes the communication with the anterior part of the air-bladder. From the sinus impar a bifid canal penetrates into the alisphenoids, in which it terminates. In Cobitis and several Loach-like Siluroids the small air-bladder consists of two globular portions placed side by side, and wholly included within two bullæ, formed by the modified parapophyses of the second and third vertebræ. The three ossicles on each side are present, but concealed by the fore part of the osseous bulla.
Organ of Taste.—Some fishes, especially vegetable feeders, or those provided with broad molar-like teeth, masticate their food; and it may be observed in Carps and other Cyprinoid fish, that this process of mastication frequently takes some time. But the majority of fish swallow their food rapidly, and without mastication, and therefore we may conclude that the sense of taste cannot be acute. The tongue is often entirely absent, and even when it exists in its most distinct state, it consists merely of ligamentous or cellular substance, and is never furnished with muscles capable of producing the movements of extension or retraction as in most higher Vertebrates. A peculiar organ on the roof of the palate of Cyprinoids, is perhaps an organ adapted for perception of this sense; in these fishes the palate between and below the upper pharyngeal bones is cushioned with a thick, soft contractile substance, richly supplied with nerves from the Nervi vagus and glossopharyngeus.
Organs of Touch.—The faculty of touch is more developed than that of taste, and there are numerous fishes which possess special organs of touch. Most fishes are very sensitive to external touch, although their body may be protected by hard horny scales. They perceive impressions even on those parts which are covered by osseous scutes, in the same manner as a tortoise perceives the slightest touch of its carapace. The seat of the greatest sensitiveness, however, appears to be the snout and the labial folds surrounding the mouth. Many species possess soft and delicate appendages, called barbels, which are almost constantly in action, and clearly used as organs of touch. Among the Triglidæ and allied families, there are many species which have one or more rays of the pectoral fin detached from the membrane, and supplied with strong nerves. Such detached rays (also found in the Polynemidæ, Bathypterois) are used partly for locomotion, partly for the purpose of exploring the ground over which the fish moves.
Some fish appear to be much less sensitive than others, or at least lose their sensitiveness under peculiar circumstances. It is well known that a Pike, whose mouth has been lacerated and torn by the hook, continues to yield to the temptation of a bait immediately afterwards. The Greenland Shark when feeding on the carcass of a whale allows itself to be repeatedly stabbed in the head without abandoning its prey. A pair of Congers are so dead to external impression at the time of copulation, and so automatically, as it were, engaged, that they have been taken by the hand together out of the water.
CHAPTER VIII.
THE ORGANS OF NUTRITION AND DIGESTION.
Fishes are either exclusively carnivorous or herbivorous, but not a few feed on vegetable substances as well as animal, or on mud containing alimentary substance in a living or decomposing state. Generally they are very voracious, especially the carnivorous kinds, and the rule of “eat or be eaten” applies to them with unusual force. They are almost constantly engaged in the pursuit and capture of their prey, the degree of their power in these respects depending on the dimensions of the mouth and gullet and the strength of the teeth and jaws. If the teeth are sharp and hooked, they are capable of securing the most slender and agile animals; if this kind of teeth is combined with a wide gullet and distensible stomach, they are able to overpower and swallow other fish larger than themselves; if the teeth are broad, strong molars, they are able to crush the hardest aliments; if they are feeble, they are only serviceable in procuring some small or inert and unresisting prey. Teeth may be wanting altogether. Whatever the prey, in the majority of cases it is swallowed whole; but some of the most voracious fishes, like some Sharks and Characinidæ, are provided with cutting teeth, which enable them to tear their prey to pieces if too large to be swallowed whole. Auxiliary organs for the purpose of overpowering their prey, which afterwards is seized or torn by the teeth, like the claws of some carnivorous mammals and birds, are not found in this class; but in a few fishes the jaws themselves are modified for that purpose. In the Sword-fishes the bones of the upper jaw form a long dagger-shaped weapon, with which they not only attack large animals, but also frequently kill fishes on which they feed. The Saw-fishes are armed with a similar but still more complicated weapon, the saw, which is armed on each side with large teeth implanted in deep sockets, specially adapted for killing and tearing the prey before it is seized and masticated by the small teeth within the mouth. Fishes show but little choice in the selection of their food, and some devour their own offspring indiscriminately with other fishes. Their digestive powers are strong and rapid, but subject in some degree to the temperature, which, when sinking below a certain point, lowers the vital powers of these cold-blooded animals. On the whole, marine fishes are more voracious than those inhabiting fresh waters; and whilst the latter may survive total abstinence from food for weeks or months, the marine species succumb to hunger within a few days. The growth of fishes depends greatly on the nature and supply of food, and different individuals of the same species may exhibit a great disparity in their respective dimensions. They grow less rapidly and to smaller dimensions in small ponds or shallow streams than in large lakes and deep rivers. The young of coast fishes, when driven out to sea, where they find a much smaller supply of food, remain in an undeveloped condition, assuming an hydropic appearance. The growth itself seems to continue in most fishes for a great length of time, and we can scarcely set bounds to—certainly we know not with precision—the utmost range of the specific size of fishes. Even among species in no way remarkable for their dimensions we sometimes meet with old individuals, favourably situated, which more or less exceed the ordinary weight and measurement of their kind. However, there are certain evidently short-lived species of fishes which attain a remarkably uniform size within a very short time; for instance, the Stickleback, many species of Gobius and Clupea.
The organs of nutrition, manducation, and deglutition, are lodged in two large cavities—an anterior (the mouth or buccal cavity), and a posterior (the abdominal cavity). In the former the alimentary organs are associated with those fulfilling the respiratory functions, the transmission of food to the stomach and of water to the gills being performed by similar acts of deglutition. The abdominal cavity commences immediately behind the head, so, however, that an extremely short thoracic cavity for the heart is partitioned off in front. Beside the alimentary organs it contains also those of the urogenital system and the air-bladder. The abdominal cavity is generally situated in the trunk only, but in numerous fishes it extends into the tail, being continued for some distance along each side of the hæmal apophyses.
In numerous fishes the abdominal cavity opens outwards by one or two openings. A single porus abdominalis in front of the vent is found in Lepidosiren and some Sturgeons; a paired one, one on each side of the vent, in Ceratodus, some species of Sturgeon, Lepidosteus, Polypterus, Amia, and all Chondropterygians. As in these fishes semen and ova are discharged by proper ducts, the abdominal openings may serve for the expulsion of semen, and those ova only which, having lost their way to the abdominal aperture of the oviduct, would be retained in the abdominal cavity. In those Teleosteans which lack an oviduct a single porus genitalis opens behind the vent.
The mouth of fishes shows extreme variation with regard to form, extent, and position. Generally opening in front, it may be turned upwards, or may lie at the lower side of the snout, as in most Chondropterygians, Sturgeons, and some Teleosteans. Vogt regards this position as a persistent fœtal condition. In most fishes the jaws are covered by the skin, which, before passing over the jaws, is often folded, forming more or less fleshy lips. In the Sharks the skin retains its external character even within the teeth, but in other fishes it changes into a mucous membrane. A tongue may exist as a more or less free and short projection, formed by the glosso*-hyal and a soft covering, or may be entirely absent. Salivary glands and a velum palati are absent in fishes.
With regard to the dentition, the class of Fishes offers an amount of variation such as is not found in any of the other classes of Vertebrates. As the teeth form one of the most important elements in the classification of fishes, their special arrangement and form will be referred to in the account of the various families and genera. Whilst not a few fishes are entirely edentulous, in others most of the bones of the buccal cavity, or some of them, may be toothed, as the bones of the jaws, the palatines, pterygoids, vomers, basisphenoid, glossohyal, branchial arches, upper and lower pharyngeals. In others teeth may be found fixed in some portion of the buccal membrane without being supported by underlying bone or cartilage; or the teeth have been developed in membrane overlying one of the dentigerous bones mentioned, without having become anchylosed to the bone. When the tooth is fixed to the bone the attachment has generally been effected by the ossification of the bone of the tooth, but in some fishes a process of the bone projects into the cavity of the tooth; in others the teeth are implanted in alveoli. In these, again, frequently a process of bone rises from the bottom, on which the tooth rests.
Many fishes, especially predatory fishes with long, lancet-shaped teeth, have all or some of the teeth capable of being bent inwards towards the mouth. Such “hinged” teeth resume at once the upright position when pressure is removed from them. They are, however, depressible in one direction only, thus offering no obstacle to the ingress, but opposing the egress of prey. Mr. C. S. Tomes has shown that the means by which this mechanism is worked are different in different fishes; for whilst, in the Pediculati and Gadoids (Hake) the elasticity resides solely in the tissue of the hinge (the tooth being as resilient as ever after everything else is severed), in the Pike the hinge is not in the least endowed with elasticity, but the bundles of fibres proceeding from the interior of the dentine cap are exceedingly elastic.
As regards texture the teeth of fishes show considerable variation. The conical teeth of the Cyclostomes and the setiform teeth of many Teleosteans consist of a horny albuminous substance. The principal substance of the teeth of other fishes consists of dentine, with numerous dividing and anastomosing tubercles, sometimes covered by a stratum of unvascular dentine. An enamel-like substance has been observed on the crown of the teeth of Sargus and Balistes, and an ossification of the capsule of their matrix covers the enamel with a thin coating of cement. The teeth either possess a cavity in which the matrix is received, or, more frequently, they are solid, in which case vascular canals of the underlying bone are continued into the substance of the tooth. In the teeth of some fishes numerous sets of canals and tubes are so arranged that they do not anastomose with one another, each set being surrounded by a layer of dentine and cement. These apparently simple teeth are evidently composed of numerous small teeth, and called compound teeth.
The teeth may be, and generally are, very different as regards size or form in the different parts of the mouth; they may be also different according to the age or sex of the fish (Raja). The teeth may be few in number and isolated, or placed in a single, double, or triple series, distant from one another or closely set; they may form narrow or broad bands, or patches of various forms. As regards form, they may be cylindrical or conical, pointed, straight, or curved, with or without an angular bent near their base; some are compressed laterally or from the front backwards; the latter may be triangular in shape, or truncated at the top like incisors of mammals; they may have one apex (cusp) only, or be bi- or tri-lobate (bi- or tri-cuspid); or have the margins denticulated or serrated. Compressed teeth may be confluent, and form a cutting edge in both jaws, which assume the shape of a parrot’s beak (Fig. [53]). In some the apex is hooked or provided with barbs. Again, some teeth are broad, with flat or convex surface, like molar teeth. With regard to size, the finest teeth are like fine flexible bristles, ciliiform or setiform; or, if very short and anchylosed to the bone, they appear only as inconspicuous asperities of the bone. Very fine conical teeth arranged in a band are termed villiform teeth; when they are coarser, or mixed with coarser teeth, they are card-like (dents en rape or en cardes) (Fig. [54]); molar-like teeth of very small size are termed granular.
Fig. 53.—Jaws of Calliodon.
In all fishes the teeth are constantly shed or renewed during the whole course of their life. In fishes which have compound teeth, as the Dipnoi, Chimæroids, Scari,[14] Gymnodonts, as well as in those which have apparently permanent teeth, as in the saw of Pristis, the detrition of the surface is made up by a constant growth of the tooth from its base. When the teeth are implanted in alveoli, they are generally succeeded by others in the vertical direction, but in others they succeed one another, side by side. In the majority of fishes the new tooth is not developed (as in reptiles and mammals) in a diverticulum of the sack of its predecessor, but like this from the free surface of the buccal membrane. Generally there are more than one tooth growing, which are in various stages of development, and destined to replace the one in function. This is very conspicuous in Sharks, in which the whole phalanx of their numerous teeth is ever marching slowly forwards (or in some backwards), in rotatory progress, over the alveolar border of the jaw, the teeth being successively cast off after having reached the outer margin, and fulfilled for a longer and shorter period their special function.
[The richest materials for our knowledge of the teeth of fishes are contained in Owen’s “Odontography.” Lond. 1840. 8vo.]
Fig. 54.—Cardlike teeth of Plectropoma dentex, with canines.
The intestinal tract is divided into four portions: œsophagus, stomach, small and large intestine; two or more of these divisions may coalesce in fishes and become indistinguishable. But it is characteristic of the class that the urinary apertures are constantly situated behind the termination of the intestinal tract.
In Branchiostoma the whole intestinal tract is straight, and coated with a ciliated mucous membrane. The wide pharynx passes into a narrow œsophagus, this into a gastric cavity, the remainder being again narrower and terminating in the anal aperture, which lies somewhat to the left of the median line. The liver is represented by a green coloured cœcal diverticulum of the stomachic dilatation. A mesenterium is absent.
In the Cyclostomi the intestinal tract is likewise straight, and without clearly defined divisions; however, in Petromyzon the œsophagus shows numerous longitudinal folds, and the intestine proper is provided with a single longitudinal fold. A mesentery, which is present in the Myxinoids, is represented by a short median fold only, by means of which the hindmost part of the intestine is fixed.
The Palæichthyes show differences in the structure of their intestinal tract as considerable as are found among the Teleostei, but they have that in common that the absorbent surface of their intestine is enlarged by the development of a spiral valve, evidence of the presence of which in extinct Palæichthyes is still preserved in the fossilised fæces or coproliths, so abundant in some of the older strata.
In Chondropterygians (Fig. [55]) the stomach is divided into a cardiac and pyloric portion, the former frequently terminating in a blind sac, and the latter varying in length. The pyloric portion is bent at its origin and end, and separated from the short duodenum (called Bursa entiana in these fishes) by a valve; the ductus hepaticus and pancreaticus enter the duodenum. This is succeeded by the straight intestine provided with the spiral valve, the coils of which may be either longitudinal and wound vertically about the axis of the intestine, as in Carcharias, Galeocerdo, Thalassorhinus, and Zygœna, or they may be transverse to that axis, as in the other genera. The number of gyrations in the latter case varies: there may be as many as forty. The short rectum passes into a cloaca, which contains also the orifices of the urogenital ducts. Only the commencement and end of the intestinal tract are fixed by mesenterial folds.
In the Holocephali and Dipnoi, the intestinal tract is short, straight, and wide, without stomachic dilatation, a pyloric valve, close to which the ductus choledochus enters, indicating the boundary of the intestine proper (Fig. [57], p). The spiral valve is perfect, and makes from three (Chimæra) to nine (Ceratodus) gyrations. A cloaca is present, as in Chondropterygians. A mesentery fixing the dorsal side of the intestine is absent.
Fig. 55.—Siphonal stomach and spiral valve of Basking-Shark (Selache). (After Home and Owen.)
a, Œsophagus; b, Cardiac portion of stomach; c, pyloric portion; d, pouch intermediate between stomach and duodenum, with circular valves at both ends; e, Duodenum; f, Valve of intestine; g, Ductus hepaticus; h, Spleen.
The other Ganoids resemble again more the Chondropterygians in the structure of their intestinal tract. The stomach has always a distinct pyloric portion, and has a still more complicated structure in Acipenser. The duodenal portion receives the contents of Appendices pyloricæ, which are confluent into a gland-like mass in Acipenser, but separate in Polyodon, and numerous and short in Lepidosteus, whilst Polypterus possesses one such appendage only. A spiral valve is developed in the Sturgeons and Polypterus, but in Amia, in which the intestine performs several convolutions, the four gyrations of the valve are situated far back towards the end of the intestine. In Lepidosteus the valve is rudimentary, and indicated only by three raised lines crossing the terminal portion of the intestine. In all these Ganoids the rectum has a separate opening, without cloaca.
The structure of the intestinal tract of Teleosteous fishes is subject to so numerous modifications that we should go beyond the limits of the present work if we would attempt to enter into details. Great differences in this respect may be found even in groups of the same natural families. Frequently the intestinal tract remains of nearly the same width throughout its course, and only the entrance of the various ducts serves as a guide for the distinction of its divisions. An intestine of such uniform width may be straight and short, as in Scombresocidæ, Symbranchidæ, or it may be more or less convoluted and long, as in many Cyprinidæ, Doradina, etc. On the whole, carnivorous fishes have a much shorter and simpler intestinal tract than herbivorous.
In the majority of Teleosteans, however, œsophagus, stomach, duodenum, small intestine and rectum, can be more or less distinctly, even externally distinguished.
There are two predominant forms of the stomach, intermediate forms being, however, numerous. In the first, the siphonal, it presents the form of a bent tube or canal, one-half of the horse-shoe being the cardiac, the other the pyloric portion. In the second, the cæcal, the cardiac division is prolonged into a long descending blind sac, the cardiac and pyloric openings of the stomach lying close together (Clupea, Scomber, Thynnus, etc.)
Fig. 56.—Siphonal Stomach and Pyloric Appendages of a Female Salmon, 3⅓ feet long. a a a, Pyloric appendages; ch, ductus choledochus; oe, œsophagus; st, lower end of stomach; p, pyloric region; i, ascending; and í’, descending portion of intestine.
The duodenum receives always the hepatic and pancreatic secretions, and, besides, those of the appendices pyloricæ, which, in varying numbers (from 1 to 200), are of very common occurrence in Teleosteans (Fig. [56]). They vary also in length and width, and whilst the narrowest serve only as secretory organs, the widest are frequently found filled with the same contents as the intestine. When few in number, each opens by a separate duct into the duodenum; when their number is greater two or more coalesce into a common duct; in the latter case the appendages cease to be free, and are connected with one another by a more or less firm tissue.
Cœcal appendages at the end of the intestinal tract are of exceedingly rare occurrence in fishes (Box). There is no cloaca in Teleosteans.
In the majority of Teleosteous fishes the vent is situated on the boundary between trunk and tail, behind the ventral fins. In a few it lies farther backwards, not far from the caudal fin; more frequently it is advanced forwards, under the middle of the abdomen or to the scapular arch. In two fishes, Aphredoderus and Amblyopsis, it lies before the pectoral fins.
A peritoneum envelops all the divisions of the intestinal tract within the abdominal cavity. A broad, well-developed omentum has hitherto been found in Gobiesox cephalus only.
Liver.—The existence of a liver in Branchiostoma as a long diverticulum of the intestine has been mentioned above. In the Myxinoids the liver is divided into two glandular bodies, an anterior rounded smaller one, and a posterior larger one of an elongate shape. The gall-bladder lies between both, and receives a cystic duct from each of them. In the other fishes the proportionally large liver is a single large gland, from which only now and then small portions are found to be detached. It is either simple, or with a right and left lobe, or with a third lobe in the middle; each lobe may have incisions or subdivisions, which, however, are very inconstant. The liver of fishes is distinguished by the great quantity of fluid fat (oil) which it contains. The gall-bladder is but rarely absent, and attached to the right lobe, or towards the centre; however, in some fishes it is detached from the liver and connected with it by the cystic duct only. The bile may be conveyed by one or more hepatic ducts into a common duct which is continued towards the gall-bladder as ductus cysticus, and towards the duodenum as ductus choledochus; or some of the hepatic ducts enter directly the gall-bladder, or directly the duodenum, without communicating with the common duct. Individual variations in this respect are of common occurrence.
A pancreas has been found hitherto in all Chondropterygians, Acipenser, and many Teleosteans. In the first it is a glandular mass of considerable size behind the stomach, close to the spleen; its duct leads into the duodenum. In the Sturgeons the pancreas is attached to the duodenum, and opens close to the ductus choledochus. In Silurus glanis it is very large, and the ductus choledochus passes through its substance; it is smaller in Belone and Pleuronectes, and situated in the mesentery; its duct accompanies the terminal portion of the ductus choledochus. In the Salmon, which possesses a large lobed pancreas, the duct is so intimately connected with the ductus choledochus that both appear externally as a single duct only.
The spleen, which is substantially a lymphatic gland, may be mentioned here, as it is constantly situated in the immediate vicinity of the stomach, generally near its cardiac portion. With the exception of Branchiostoma, it is found in all fishes, and appears as a rounded or oblong organ of dark-red colour. In the Sharks frequently one or more smaller pieces are detached from the principal body. In the Dipnoi a thin layer of a very soft substance of brownish-black colour below the mucous membrane of the stomach and upper part of the intestine has been regarded as the homologue of the spleen (Fig. [57], m). In most Teleostei the spleen is undivided, and appended by its vessels and a fold of the peritoneum to the pyloric bend of the stomach or the beginning of the intestine.
Fig. 57.—Upper part of Intestine of Ceratodus. The anterior wall of the intestine is opened, the liver (c) and gall-bladder (e) being drawn forward. A slit is made at n, through which part of the next compartment of the spirally wound intestine may be seen.
é, Mouth of ductus choledochus; f, stomach; i, adipose agglomeration; l, first compartment of intestinal spire; m, spleen; oe, lower part of œsophagus, opened; p, double pyloric fold; q q, glandular patches.
CHAPTER IX.
ORGANS OF RESPIRATION.
Fishes breathe the air dissolved in water by means of gills or branchiæ. The oxygen consumed by them is not that which forms the chemical constituent of the water, but that contained in the air which is dissolved in water. Hence fishes transferred into water from which the air has been driven out by a high temperature, or in which the air absorbed by them is not replaced, are speedily suffocated. The absorption of oxygen by fishes is comparatively small, and it has been calculated that a man consumes 50,000 times more than is required by a Tench. However, some fishes evidently require a much larger supply of oxygen than others: Eels and Carps, and other fishes of similar low vitality, can survive the removal out of their elements for days, the small quantity of moisture retained in their gill-cavity being sufficient to sustain life, whilst other fishes, especially such as have very wide gill-openings, are immediately suffocated after being taken out of the water. In some fishes noted for their muscular activity, like the Scombridæ, the respiratory process is so energetic as to raise the temperature of their blood far beyond that of the medium in which they live. A few fishes, especially such as are periodically compelled to live in water thickened into mud by desiccation and vitiated by decomposing substances, breathe atmospheric air, and have generally special contrivances for this purpose. These are so much habituated to breathing air that many of them, even when brought into pure water of normal condition, are obliged to rise to the surface at frequent intervals to take in a quantity of air, and if they be kept beneath the surface by means of a gauze net, they perish from suffocation. The special contrivances consist of additional respiratory organs, lodged in cavities either adjoining the gill-cavity or communicating with the ventral side of the œsophagus, or of the air-bladder which enters upon respiratory functions (Dipnoi, Lepidosteus, Amia).
The water used by fishes for respiration is received by the mouth, and by an action similar to that of swallowing driven to the gills, and expelled by the gill-openings, of which there may be one or several on each side behind the head; rarely one only in the median line of the ventral surface.
Fig. 58.—Fore-part of the body of an embryon of Carcharias, showing the branchial filaments (natural size).
The gills or branchiæ consist essentially of folds of the mucous membrane of the gill-cavity (laminæ branchiales), in which the capillary vessels are distributed. In all fishes the gills are lodged in a cavity, but during the embryonic stage the Chondropterygians have the gill-laminæ prolonged into long filaments projecting beyond the gill-cavity (Fig. [58]), and in a few young Ganoids external gills are superadded to the internal.
In Branchiostoma the dilated pharynx is perforated by numerous clefts, supported by cartilaginous rods (Fig. [29], h). The water passes between these clefts into the peritoneal cavity, and makes its exit by the porus abdominalis situated considerably in advance of the vent. The water is propelled by cilia.
In the Cyclostomes the gills of each side are lodged in a series of six or more antero-posteriorly compressed sacs, separated from each other by intervening septa. Each sac communicates by an inner duct with the œsophagus, the water being expelled by an outer duct. In Bdellostoma each outer duct has a separate opening, but in Myxine all the outer ducts pass outwards by one common gill-opening on each side. In the Lampreys the ducts are short, the outer ones having separate openings (Fig. [2], p. 39). The inner ducts lead into a single diverticulum or bronchus, blind behind, situated below the œsophagus, and communicating in front with the pharynx, where it is provided with two valves by which the regurgitation of the water into the buccal cavity is prevented.
The same type of branchial organs persists in Chondropterygians, which possess five, rarely six or seven, flattened pouches with transversely plaited walls. The septa between them are supported by cartilaginous filaments rising from the hyoidean and branchial arches. Each pouch opens by a cleft outwards, and by an aperture into the pharynx, without intervening ducts. The anterior wall of the first pouch is supported by the hyoidean arch. Between the posterior wall of the first and the anterior of the second sac, and between the adjacent walls of the succeeding, a branchial arch with its two series of radiating cartilaginous filaments is interposed. Consequently the first and last pouch have one set of gill-laminæ only, viz. the first on its posterior and the last on its anterior wall. The so-called spiracles on the upper surface of the head of Chondropterygians are to be referred to in connection with the respiratory organs. They are the external openings of a canal leading on each side into the pharynx, and situated generally close to and behind the orbit. They frequently possess valves or an irregularly indented margin, and are found in all species during the embryonic stage, but remaining persistent in a part only. The spiracles are the remains of the first visceral cleft of the embryo, and in the fœtal state long branchial filaments have been observed to protrude, as from the other branchial clefts.
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.
[See Müller, “Vergleichende Anatomie des Gefäss-systems der Myxinoiden;” and “Ueber den Bau und die Grenzen der Ganoiden.”]
Fig. 60.—Gills of Ceratodus.
x, Arcus aortæ; gl, Glossohyal; ch, Ceratohyal; u, Attachment of the first gill to the walls of the gill-cavity; h, Pseudobranchia; x’, x”, two series of gill-rakers belonging to the Pseudobranchia.
Accessory respiratory organs for retaining water or breathing air, such as are found in the Labyrinthici, Ophiocephalidæ, certain Siluridæ, and Lutodira, are structures so specialised that they are better described in the accounts of the Fishes in which they have been observed.
Air-Bladder.—The air-bladder, one of the most characteristic organs of fishes, is a hollow sac, formed of several tunics, containing gas, situated in the abdominal cavity, but without the peritoneal sac, entirely closed or communicating by a duct with the intestinal tract. Being compressible, its special functions consist in altering the specific gravity of the fish or in changing the centre of gravity. In a few fishes it assumes the function of the organ of higher Vertebrates, of which it is the homologue—viz. of a lung.
The gas contained in the air-bladder is secreted from its inner surface. In most freshwater fishes it consists of nitrogen, with a very small quantity of oxygen and a trace of carbonic acid; in sea-fishes, especially those living at some depth, oxygen predominates, as much as 87 per cent having been found. Davy found in the air-bladder of a fresh-run Salmon a trace of carbonic acid and 10 per cent of oxygen, the remainder of the gas being nitrogen.
An air-bladder is absent in Leptocardii, Cyclostomi, Chondropterygii, and Holocephali; but occurs in all Ganoids, in which, besides, its respiratory functions more or less clearly manifest themselves. Its occurrence in Teleosteans is most irregular, closely allied species sometimes differing from each other in this respect; it shows in this sub-class the most extraordinary modifications, but has no respiratory function whatever.
Constantly situated within the abdominal cavity, below the vertebral column, but without the sac of the peritoneum which covers only its ventral portion, the air-bladder is frequently prolonged into the tail, the prolongation being either simple and lodged between the non-united parapophyses, or double and penetrating between the muscles and hæmapophyses of each side. In the opposite direction processes of the air-bladder may penetrate into the skull, as has been mentioned above (p. 117). In some fishes the air-bladder is almost loose in the abdominal cavity, whilst in others it adheres most intimately by firm and short tissue to the vertebral column, the walls of the abdomen, and the intestines. In the Cobitina and many Siluroids it is more or less completely enclosed in osseous capsules formed by the vertebræ.
The tunics of the majority of air-bladders are an extremely fine internal one, frequently shining silvery, containing crystalline corpuscles, sometimes covered with a pavement-epithelium; and a thicker outer one of a fibrous texture, which sometimes attains to considerable thickness and yields isinglass. This wall is strengthened in many fishes by muscular layers for the compression of the whole organ or of some portion of it.
A distinction has been made between air-bladders which communicate by a duct with the intestinal tract and those which are entirely closed. However, it is to be remembered that at an early stage of development all air-bladders are provided with such a duct, which in a part of the fishes more or less completely obliterates, being then represented by a fine ligament only. In young Lucioperca of six to eight inches in length the duct may be found still open for a considerable distance; and, on the other hand, in adult Physostomi, that is Teleosteous fishes with a ductus pneumaticus, not rarely the whole duct is found very narrow, or, for some part of its length, even entirely closed.
Fig. 61.—Air-bladder of Otolithus sp.
Fig. 62.—Vertical section through abdominal cavity of Collichthys lucida. b, air-bladder; l, liver; s, stomach; epp and ipp, external and internal laminæ of peritoneum parietale; epv and ipv, external and internal laminæ of peritoneum viscerale; dv, dorsal air-vessels; vv, ventral air-vessels.
Air-bladders without duct are found in Acanthopterygians, Pharyngognaths, Anacanths, and Lophobranchs. They may consist of a single cavity or divided by constrictions into two or three partitions situated behind one another; they may consist of two lateral partitions, assuming a horseshoe-like form, or they may be a single sac with a pair of simple or bifid processes in front or behind (Fig. [61]). The families of Sciænidæ and Polynemidæ possess air-bladders with a most extraordinary development of appendages rising from each side of the air-bladder. In the Sciænoid (Fig. [63]) fifty-two branches issue from each side, each branch being bifurcate and bearing smaller appendages. In Pogonias chromis (Fig. [64]) the sides of the anterior half is provided with irregular broad-fringed appendages, the hindmost of which communicates by a narrow duct with the posterior extremity of the air-bladder. In Collichthys lucida (Fig. [62]) twenty-five appendages issue from each side; the anterior ones are directed towards the front, but the lateral assume a more posterior direction, the nearer they are to the posterior extremity of the air-bladder, where they form an assemblage giving the appearance of a cauda equina. All these appendages soon bifurcate in a dorsal and ventral stem; these stems bifurcate again and again, and either terminate after the first or second bifurcation or are so far prolonged as to reach the median line of the ventral and dorsal sides, anastomosing with the branches of the other side. The branches being enveloped in laminæ of the peritonæum, form a dorsal and ventral sac of beautiful appearance, caused by the regular arrangement of the air-vessels. The dorsal sac is situated between the air-bladder and the roof of the abdominal cavity without being attached to the latter. The ventral sac receives within its cavity the intestine, liver, and ovaries.—A peculiar mechanism has been observed in the air-bladder of the Ophidiidæ, the anterior portion of which can be prolonged by the contraction of two muscles attached to its anterior extremity, with or without the addition of a small bone.
Fig. 63.—Air-bladder of a Sciænoid.
I. Visceral surface opened at b, to show openings of the lateral branches.
II. Isolated lateral branch; a, its opening into the cavity of the air-bladder.
Air-bladders with a pneumatic duct are found in Ganoids and Physostomes, the duct entering the dorsal side of the intestinal tract, with the exception of Polypterus and the Dipnoi, in which it enters on the ventral side of the œsophagus. In the majority the orifice is in the œsophagus, but in some, as in Acipenser, in the cardiac portion of the stomach, or in its blind sac, as in many Clupeoids. The air-bladder may be single, or consist of two divisions situated one behind the other (Fig. [52]); its inner surface may be perfectly smooth, or form manifold pouches and cells. If two divisions are present the anterior possesses a middle elastic membrane which is absent in the posterior; each division has a muscular layer, by which it can be separately compressed, so that part of the contents of the posterior may be driven into the elastic anterior division, and vice versa. The posterior division being provided with the ductus pneumaticus does not require the elasticity of the anterior.
Fig. 64.—Air-bladder of Pogonias chromis.
Some Siluroids possess a peculiar apparatus for voluntarily exercising a pressure upon the air-bladder. From the first vertebra a process takes its origin on each side, expanding at its end into a large round plate; this is applied to the side of the air-bladder, and by pressing upon it expels the air through the duct; the small muscle moving the plate rises from the skull.
The connection of the air-bladder with the organ of hearing in some Physostomes has been described above, p. 117.
In the modifications of the air-bladder, hitherto mentioned, the chief and most general function is a mechanical one; this organ serves to regulate the specific gravity of the fish, to aid it in maintaining a particular level in the water, in rising or sinking, in raising or depressing the front part of its body as occasion may serve. Yet a secretion of gas from the blood into its cavity must take place; and if this be so, it is not at all impossible that also an exchange of gases between the two kinds of blood is effected by means of the extraordinary development of retia mirabilia in many air-bladders.
In all fishes the arteries of the air-bladder take their origin from the aorta or the system of the aorta, and its veins return either to the portal, or vertebral, or hepatic veins; like the other organs of the abdominal cavity it receives arterial blood and returns venous blood. However, in many fishes the arteries as well as veins break up below the inner membrane into retia mirabilia in various ways. The terminal ramifications of the arteries may dissolve into fan-like tufts of capillaries over almost every part of the inner surface, as in Cyprinoids. Or these tufts of radiating capillaries are more localised at various places, as in Esocidæ; or the tufts are so aggregated as to form gland-like, red bodies, the capillaries reuniting into larger vessels, which again ramify freely round the border of the red body; the red bodies are formed not only by minute arteries but also by minute veins, both freely anastomosing with its kind, and being inextricably interwoven. The rest of the inner surface of the air-bladder receives its blood, not from the red bodies, but from normally ramifying vessels. This kind of rete mirabile or “vaso-ganglion” is found in the Perch and Gadoids; it is generally distributed in closed air-bladders, but also sometimes observed in air-bladders’ with pneumatic duct. In Anguilla and Conger two similar vaso-ganglia are situated at the sides of the opening of the pneumatic duct.
Fig. 65.—Lung of Ceratodus opened in its lower half to show its cellular pouches. a, Right half; b, Left half; c, Cellular pouches; e, Vena pulmonalis; f, Arterial blood-vessel; oe, Œsophagus opened, to show glottis (gl.)
Whilst the air-bladders of some Ganoids, anatomically as well as functionally, closely adhere to the Teleosteous type, that of Amia is more cellular and lung-like in its interior than the Teleosteous air-bladder, and Polypterus approaches the Dipnoi not only in having a laterally divided air-bladder but also in its pneumatic duct entering the ventral side of the œsophagus. The air-bladder of the Dipnoi possesses still more the anatomical characteristics of a lung and assumes its functions, though, as it co-exists with gills, only periodically or in an auxiliary manner. The ductus pneumaticus is a membranous bronchus, entering the ventral side of the œsophagus, and provided at its entrance with a glottis. In Ceratodus (Fig. [65]) the lung is still a single cavity, but with a symmetrical arrangement of its internal pouches; it has no pulmonal artery, but receives branches from the arteria cœliaca. Finally, in Lepidosiren and Protopterus the lung is completely divided into lateral halves, and by its cellular structure approaches most nearly that of a reptile; it is supplied with venous blood by a true pulmonary artery.
Fig. 66.—Heart of Lepidosteus osseus.
I. External aspect. II. Conus arteriosus opened.
a, Atrium; b, Conus arteriosus; v, Ventricle; h, Branchial artery for 3d and 4th gill; k, for the second; l, for the first; m, branch for the opercular gill; d, Single valve at the base of the conus; e-g, Transverse rows of Ganoid valves.
CHAPTER X.
ORGANS OF CIRCULATION.
The Blood-corpuscles of fishes are, with one exception, of an elliptic shape; this exception is Petromyzon, which possesses circular, flat, or slightly biconvex blood-corpuscles. They vary much in size; they are smallest in Teleosteans and Cyclostomes, those of Acerina cernua measuring 1/2461 of an inch in their longitudinal, and 1/3000 in their transverse diameter. As far as it is known at present the Salmonidæ have the largest blood-corpuscles among Teleosteans, those of the salmon measuring 1/1524 by 1/2460 in., approaching those of the Sturgeon. Those of the Chondropterygians are still larger; and finally, Lepidosiren has blood-corpuscles not much smaller than those of Perennibranchiates, viz.—1/570 by 1/941 in. Branchiostoma is the only fish which does not possess red blood-corpuscles.
[See G. Gulliver, “Proc. Zool. Soc.,” 1862, p. 91; and 1870, p. 844; and 1872, p. 833.]
Fishes, in common with the other Vertebrates, are provided with a complete circulation for the body, with another equally complete for the organs of respiration, and with a particular abdominal circulation, terminating at the liver by means of the vena portæ; but their peculiar character consists in this, that the branchial circulation alone is provided at its base with a muscular apparatus or heart, corresponding to the right half of the heart of Mammalia and Birds.
The Heart is situated between the branchial and abdominal cavities, between the two halves of the scapulary arch, rarely farther behind, as in Symbranchidæ. It is enclosed in a pericardium, generally entirely separated from the abdominal cavity by a diaphragma, which is, in fact, the anterior portion of the peritoneum, strengthened by aponeurotic fibres. However, in some fishes there is a communication between the pericardial and peritoneal sacs, viz. in the Chondropterygians and Acipenser, whilst in the Myxinoids the pericardial sac is merely a continuation of the peritoneum.
Fig. 67.—Heart of Ceratodus.
a, Atrium; b, Conus arteriosus; d, Papillary valve within the conus; e-g, Transverse rows of Ganoid valves; h, i, Anterior arcus aortæ; k, l, Posterior arcus aortæ; v, Ventricle.
The heart is, relatively to the size of the body, very small, and consists of three divisions: the atrium, with a large sinus venosus into which the veins enter; the ventricle; and a conical hollow swelling at the beginning of the arterial system, the structure of which forms one of the most important characters used in the classification of fishes. In all Palæichthyes (Figs. [66] and [67]) this swelling is still a division of the pulsating heart, being provided with a thick muscular stratum; it is not separated from the ventricle by two valves opposite to each other, but its interior is fitted with a plurality of valves, arranged in transverse series more or less numerous in the various groups of Palæichthyes. Lepidosiren and Protopterus offer an example of a modification of this valvular arrangement, their valves being longitudinal, each valve in fact being formed by the confluence of several smaller ones situated behind one another. This Palæichthyan type is called conus arteriosus.
In Cyclostomes and Teleosteans (Fig. [68]) the enlargement is a swelling of the artery, without muscular stratum and without contractility; with the exception of the Myxinoids its walls are thick, fibrous, with many trabeculæ and pouches, but it has no valves in its interior, and is separated from the ventricle by two valves opposite to each other. This Teleostean type is called bulbus aortæ.
Fig. 68.—Bulbus aortæ of Xiphias gladius, opened.
a, Section through part of the wall of ventricle; b, Section through the bulbus; c, Teleosteous valves of the ostium arteriosum; d, Accessory valves, of rudimentary nature and inconstant; e, Trabeculæ carneæ of the bulbus.
The sinus venosus sends the whole of the venous blood by a single orifice of its anterior convexity into the atrium; two thin membranous valvules turned towards the atrium, prevent the blood from re-entering the sinus. A pair of other valves between atrium and ventricle have the same function. The walls of the ventricle are robust, and, internally, it is furnished with powerful fleshy trabeculæ.
The bulbus or conus arteriosus is prolonged into the branchial artery which soon divides, sending off a branch to each branchial arch. On returning from the respiratory organ the branchial veins assume the structure and functions of arteries. Several branches are sent off to different portions of the head and to the heart, but the main trunks unite to form the great artery which carries the blood to the viscera and all the parts of the trunk and tail, and which, therefore, represents the aorta of higher animals.
In the majority of Teleosteans the aorta has proper walls formed by its own membranes, but in the Sturgeons it is independent at its commencement only, and replaced by a canal formed by hæmal elements of the vertebral column, and clothed inside with a perichondrium. In many Chondropterygians and some Teleosteans (Esox, Clupea, Silurus), the aorta possesses its own firm membranes along its ventral side, dorsally being protected by a very thin membrane only, attached to the concavity of the centra of the vertebræ.
The circulatory system of Branchiostoma and of the Dipnoi shows essential differences from that of other fishes.
Branchiostoma is the only fish which does not possess a muscular heart, several cardinal portions of its vascular system being contractile. A great vein extends forwards along the caudal region below the notochord, and exhibits contractility in a forward direction; it is bent anteriorly, passing into another tube-like pulsatile trunk, the branchial heart, which runs along the middle of the base of the pharynx, sending off branches on each side to the branchiæ; each of these branches has a small contractile dilatation (bulbillus) at its base. The two anterior branches pass directly into the aorta, the others are branchial arteries, the blood of which returns by branchial veins emptying into the aorta. The blood of the intestinal veins is collected in a contractile tube, the portal vein, situated below the intestine, and distributed over the rudimentary liver. Of all other fishes, only in Myxinoids the portal vein is contractile. All the blood-corpuscles of Branchiostoma are colourless and without nucleus.
In Dipnoi a rudimentary division of the heart into a right and left partition has been observed; this is limited to the ventricle in Ceratodus, but in Lepidosiren and Protopterus an incomplete septum has been observed in the atrium also. All Dipnoi have a pulmonal vein, which enters the atrium by a separate opening, provided with a valve. The pulmonal artery rises in Lepidosiren and Protopterus from an arch of the aorta, but in Ceratodus it is merely a subordinate branch, rising from the Arteria cœliaca.
CHAPTER XI.
URINARY ORGANS.
In Branchiostoma no urinary organs have been found.
In Myxinoids these organs are of a very primitive structure: they consist of a pair of ducts, extending from the urogenital porus through the abdominal cavity. Each duct sends off at regular intervals from its outer side a short wide branch (the uriniferous tube), which communicates by a narrow opening with a blind sac. At the bottom of this sac there is a small vaso-ganglion (Malpighian corpuscle), by which the urine is secreted.
In the Lampreys the kidneys form a continuous gland-like body, with irregular detached small portions. The ureters coalesce before they terminate in the urogenital papilla.
In Chondropterygians the kidneys occupy the posterior half or two-thirds of the back of the abdominal cavity, without the sac of the peritoneum (as in all fishes) which forms a firm tendinous horizontal septum. The kidneys of the two sides are never confluent, and generally show a convoluted or lobulated surface. The ureters are short; each is dilated into a pouch, and communicating with its fellow terminates by a single urethra (which also receives the vasa deferentia) behind the end of the rectum in the large common cloaca.
In Ganoids the kidneys occupy a similar position as in Chondropterygians, but these fishes differ considerably with regard to the termination and the arrangement of the ends of the urogenital ducts. The Dipnoi possess a cloaca. In Ceratodus the ureters open into it by a common opening, separate from the genital opening; and no closed urinary bladder has been developed. Lepidosiren has a small urinary bladder; the ureters do not communicate directly with it, but terminate separately on small papillæ in the dorsal compartment of the cloaca. The other Ganoids lack a cloaca, and the urogenital opening is behind the vent as in Teleosteans. In all the genital and urinary ducts coalesce towards their end. The Sturgeons have no urinary bladder, whilst it is present in Amia, the ureters opening separately into it.
The kidneys of Teleosteans are situated likewise without the peritoneal cavity, immediately below some part of the vertebral column, and vary exceedingly with regard to form and extent. Sometimes they reach from the skull to between the muscles of the tail, sometimes they are limited to the foremost part of the abdominal cavity (in advance of the diaphragm), but generally their extent corresponds to that of the abdominal portion of the vertebral column. Frequently they are irregular on their dorsal surface, filling every available recess, flat, attenuated on the sides, more or less coalescent towards the middle; in other fishes they are more compact bodies. The ureters terminate, either separate or united, in a urinary bladder, varying in shape, which opens by a short urethra behind the vent. The urinary opening may be separate or confluent with that of the genital ducts, and is frequently placed on a more or less prominent papilla (papilla urogenitalis). If separate, the urinary opening is behind the genital; and if a papilla is developed, its extremity is perforated by the urethra, the genital opening being situated nearer the base. A few Teleosteans show an arrangement similar to that of Chondropterygians and Dipnoi, the urogenital openings being in the posterior wall of the rectum (Symbranchidæ, Pediculati, and some Plectognathi).
CHAPTER XII.
ORGANS OF REPRODUCTION.
All fishes are dioecious, or of distinct sex. Instances of so-called hermaphroditism are, with the exception of Serranus, abnormal individual peculiarities, and have been observed in the Cod-fish, some Pleuronectidæ, and in the Herring. Either the generative organ of one side was found to be male, that of the other female; or the organ of one or both sides was observed to have been developed partly into an ovary partly into a testicle. In the European species of Serranus a testicle-like body is attached to the lower part of the ovary; but many specimens of this genus are undoubtedly males, having normally developed testicles only.
The majority of fishes are oviparous, comparatively few viviparous; the embryos being developed either in the ovarium or in some dilated portion of the oviduct. In viviparous fishes actual copulation takes place, and the males of most of them are provided with copulatory or intromittent organs. In oviparous fishes the generative products are, during sexual excitement, discharged into the water, a very small quantity of semen being sufficient for effectual impregnation of a number of ova dispersed in a considerable quantity of water; circumstances which render artificial impregnation more practicable than in any other class of animals.
In Branchiostoma the generative organs occupy the ventral side of the abdominal cavity, into which they discharge their contents. No ducts are developed in either sex.
In the Cyclostomes the generative organ is single, and fixed to or suspended from the median line of the back of the visceral cavity by a duplicature of the peritoneum (mesoarium); the testicle and ovary being distinguishable by their contents only. These escape by dehiscence of the cells or capsules and rupture of the peritoneal covering into the abdominal cavity, and are expelled by reciprocal pressure of the intertwined sexes through the porus genitalis, which is sunk between two labia of the skin in Myxine, and produced into a long papilla in Petromyzon.
69.—Ovum of Myxine glutinosa, enlarged.
The ova of the Lampreys are small, globular, like those of Teleosteans. Those of Myxine have a very peculiar shape when mature; they are of an oval form, about 15 millimetres long and 8 millimetres broad, enveloped in a horny case, which at each end is provided with a bundle of short threads, each thread ending in a triple hook. Whilst in the mesoarial fold the eggs are attached to one another by means of these hooks, and after being expelled they probably fix themselves by the same means to other objects. As in all fishes producing ova of large size, the number of ova matured in one season is but small.
In Teleosteans the generative organs are comparatively large. In some families the ovaries are without closed covering and without oviducts, as in Salmonidæ, Galaxiidæ, Notopteridæ, Murænidæ, and others. The surface of such an open ovary—as, for instance, that of the Salmon—is transversely plaited, the ova being developed in capsules in the stroma of the laminæ; after rupture of the capsules the mature ova drop into the abdominal cavity, and are expelled by the porus genitalis. The ovaries of the other Teleosteans are closed sacs, continued into oviducts. Frequently such ovaries coalesce into a single body, or one in which the division is effected internally only by a more or less complete septum. Fixed by a mesoarium, the ovaries occupy generally a position outwards of the intestine or air-bladder; their form varies as well as the thickness and firmness of their covering, which frequently is an extremely thin transparent membrane. The inner surface of the ovarian sac is transversely or longitudinally plaited or covered with fringes, on which the ova are developed, as in the open ovaries. In the viviparous Teleosteans the embryons are likewise developed within the ovary, notably in the Embiotocidæ, many Blenniidæ, and Cyprinodontidæ, Sebastes viviparus, etc. Among the Cyprinodonts the end of the oviduct is attached to the anterior anal rays, which are modified into supports of its termination. In Rhodeus the oviduct is periodically prolonged into a long oviferous tube, by means of which the female deposits her ova into the shells of living Bivalves.
Fig. 70.—Ditrema argenteum, with fully developed young, ready for expulsion by the genital orifice, o; a, folds of the ovarian sac; v, vent.
The ova of Teleosteous Fishes are extremely variable in size, quite independently of the size of the parent species. The ova of large and small individuals of the same species, of course, do not differ in size; but, on the whole larger individuals produce a greater number of ova than smaller ones of the same species. The larger the size of the ova is in a species, the smaller is the number produced during one season. The ova of the Eel are almost microscopic. The small sized roe in the Herring, Lump-fish, Halibut, and Cod-fish, have been estimated at respectively 25,000, 155,000, 3,500,000, and 9,344,000. Larger in size and fewer in number are those of Antennarius, Salmo, Aspredo, Lophobranchs, etc. Comparatively largest are those of Gastrosteus; and the Siluroid genus Arius, the males of which take care of their progeny, produces ova from 5 to 10 millimeters in diameter. The ova of all Teleosteans are perfectly globular and soft-shelled. Teleosteans without oviduct, deposit them separated from one another; whilst in many Teleosteans with an oviduct the ova are enveloped in a glutinous substance, secreted by its glands, swelling in the water and forming lumps or cords, in which the ova are aggregated.
Fig. 71. Ovum of Arius boakii (Ceylon), showing embryo. Nat. size.
Fig. 72.—Abdomen of Aspredo batrachus, with the ova attached; at a, the ova are removed, to show the spongy structure of the skin, and the processes filling the interspaces between the ova. (Natural size.)
Instances of the female taking care of her progeny are extremely scarce in fishes. At present only two examples are known, that of the Siluroid genus Aspredo, and of Solenostoma. In the former, during the time of propagation, the integuments of the lower side of the flat trunk of the female assume a soft and spongy texture. After having deposited the eggs, the female attaches them to, and presses them into, the spongy integument, by merely lying over them. She carries them on her belly, as the Surinam Toad (Pipa) carries her ova on the back. When the eggs are hatched the excrescence on the skin disappears, and the abdomen becomes as smooth as before. In Solenostoma the inner side of the long and broad ventral fins coalesces with the integuments of the body, a large pouch being formed for the reception of the eggs. There is a peculiar provision for the retention of the eggs in the sac, and probably for the attachment of the embryo. The inner walls of the sac are lined with long filaments, arranged in series along the ventral rays, and more numerous and longer at the base of the rays than in the middle of their length, behind which they disappear entirely. They are also more developed in examples in which eggs are deposited in the sac than in those which have the sac empty. The filaments most developed have a length of half an inch, and are beset with mamilliform appendages. A slightly undulated canal runs along the interior of the filament.
Fig. 73.—Solenostoma cyanopterum ♂ (Indian Ocean).
Fig. 74.—Pouch with ova, formed by the ventral fins of Solenostoma. Lower aspect; the edges of the fins have been pushed aside to allow of a view of the inside of the pouch. (Natural size.)
The Testicles of the Teleosteans are always paired, and occupy the same position as the ovaries. Their size varies extraordinarily at the different seasons of the year. Vasa deferentia are constant. In the males of viviparous Teleosteans the urogenital papilla is frequently enlarged, and clearly serves as an intromittent organ. In Clinus despicillatus the vas deferens widens within the abdomen into a cavity occupied by a complex network of loose fasciculi, rising from the mucous membrane. The cavity can be compressed by a special powerful muscle, the accumulated semen being thus expelled with considerable force through the narrow aperture of the penis. In many Cyprinodonts the vas deferens runs along the anterior anal rays, which may be thickened, and prolonged into a long slender organ.
Many Teleostei take care of their progeny, but with the exception of Aspredo and Solenostoma, mentioned above (p. 160), it is the male on which this duty devolves. In some, as in Cottus, Gastrosteus, Cyclopterus, Antennarius, Ophiocephalus, Callichthys, the male constructs with more or less skill a nest, and jealously guards the ova deposited in it by the female. The male of some species of Arius carries the ova (Fig. [71]) about with him in his capacious pharynx. The species of Chromis, inhabiting the sea of Galilee, are said to take care of their ova in the same manner. And, finally, in the Lophobranchs, nature has aided this instinct by the development of a pouch on the abdomen or lower side of the tail. In the Syngnathidæ this pouch is formed by a fold of the skin developed from each side of the trunk and tail, the free margins of the fold being firmly united in the median line, whilst the eggs are being hatched in the inside of the pouch. In Hippocampus the pouch is completely closed, with a narrow anterior opening.
Fig. 75.—Syngnathus acus ♂, with sub-caudal pouch.
Fig. 76.—Sub-caudal pouch of Syngnathus acus, with the young, ready to leave the pouch. One side of the membrane of the pouch is pushed aside to admit of a view of its interior. (Natural size.)
The genital organs of Ganoids show similar diversity of structure as those of Teleosteans, but on the whole they approach the Batrachian type. The ovaries are not closed, except in Lepidosiren; all Ganoids possess oviducts. In the Sturgeons the oviduct as well as the vas deferens is represented by a funnel-shaped prolongation of the peritoneum, which communicates with the wide ureter. The inner aperture of the funnel is on a level of the middle of the testicle or ovary, the outer within the ureter; and it is a noteworthy fact that only at certain periods of the life of the fish this outer aperture is found to be open,—at other times the peritoneal funnel appears as a closed blind sac within the ureter. The mode of passage of the semen into the funnel is not known.
In Polypterus and Amia, proper oviducts, with abdominal apertures in about the middle of the abdominal cavity, are developed; they coalesce with the ureters close to the common urogenital aperture.
In Ceratodus (Fig. [77]), a long convoluted oviduct extends to the foremost limit of the abdominal cavity, where it opens by a slit at a considerable distance from the front end of the long ovary; this aperture is closed in sexually immature specimens. The oviducts unite close to their common opening in the cloaca. During their passage through the oviduct the ova receive a gelatinous covering secreted by its mucous membrane. This is probably also the case in Lepidosiren, which possesses a convoluted oviduct with secretory glands in the middle of its length. The oviduct begins with a funnel-shaped dilatation, and terminates in a wide pouch, which posteriorly communicates with that of the other side, both opening by a common aperture behind the urinary bladder.
The ova of Ganoids, as far as they are known at present, are small, but enveloped in a gelatinous substance. In the Sturgeon have been counted as many as 7,635,200. Those of Lepidosteus seem to be the largest, measuring 5 millimetres in diameter with their envelope, and 3 millimetres without it. They are deposited singly, like those of Newts.
Fig. 77.—Ovaries of Ceratodus.
a, Right ovary shown from the inner surface, which is covered by the peritoneum; a’, Left ovary, showing its outer surface; l, Portion of liver; o, Oviduct; p, the lower part of the oviduct is opened to show the folds of its inner membrane; q, Opening of the left oviduct into the right; r, Abdominal orifice of the oviduct.
In Chondropterygians (and Holocephali) the organs of reproduction assume a more compact form, and are more free from a lengthened attachment to the back of the abdominal cavity. The ovaries of the majority are paired, single in the Carchariidæ and Scylliidæ, one remaining undeveloped. But the oviducts are always paired, beginning immediately behind the diaphragma with a common aperture. They consist of two divisions, separated by a circular valve; the upper is narrow, and provided within its coats with a gland which secretes the leathery envelope in which most of the Chondropterygian ova are enclosed; the lower forms the uterine dilatation, in which the embryoes of the viviparous species are developed. Generally the vitelline sac of the embryoes is free, and without connection with the uterus, which in these cases has merely the function of a protecting pouch; but in Carcharias and Mustelus lævis a placenta uterina is formed, the vascular walls of the vitelline sac forming plaits fitting into those of the membrane of the uterus. The ends of the uteri open by a common aperture behind the ureter into the cloaca.
Fig. 78.—Ventral fins and claspers of Chiloscyllium trispeculare.
The testicles are always paired, rounded, and situated in the anterior part of the abdominal cavity, covered by the liver. Vasa efferentia pass the semen into a much-convoluted epididymis, which is continued into the vas deferens; this, at the commencement of its course, is spirally wound, but becomes straight behind, and has its end dilated into a seminal reservoir. It opens with the urethra in a papilla within the cloaca.
The so-called claspers of Chondropterygians (Fig. [78]) are characteristic of all male individuals. They are semi-ossified appendages of the pubic, with which they are movably joined, and special muscles serve to regulate their movements. Sometimes they are armed with hook-like osseous excrescences (Selache). They are irregularly longitudinally convoluted, and, when closely ad-pressed to each other, form a canal open at their extremity. A gland, abundantly discharging a secretion during the season of propagation, is situated at, and opens into, the base of the canal. It is still doubtful whether the generally-adopted opinion that their function consists in holding the female during copulation is correct, or whether they are not rather an intromittent organ, the canal of which not only conducts the secretion of their proper gland but also the impregnating fluid.
Fig. 79.—Egg of a Scyllium from Magelhan’s Straits (? Sc. chilense). Natural size.
Fig. 80.—Egg-shell of Cestracion philippi, half natural size, linear.
I. External view. II. Vertical section.
a, One spiral ridge; b, The other spiral ridge; c, Cavity for the ovum.
The ova of the oviparous Chondropterygians are large and few in number; they are successively impregnated, and the impregnation must take place before they are invested with a tough leathery envelope which would be impenetrable to the semen, that is, before they enter the uterus; therefore, copulation must take place in all these fishes. The form of the egg-shell differs in the various genera; generally (Fig. [79]) they are flattened, quadrangular, with each of the four corners produced, and frequently prolonged into long filaments which serve for the attachment of the ova to other fixed objects. In Notidanus the surfaces are crossed by numerous ridges. In Cestracion (Fig. [80]) the egg is pyriform, with two broad ridges or plates, wound edgewise round it, the two ridges forming five spires. The eggs of Callorhynchus (Fig. [81]) have received a protective resemblance to a broad-leaved fucus, forming a long depressed ellipse, with a plicated and fringed margin.
Fig. 81.—Egg of Callorhynchus antarcticus. a, Cavity for the embryo.
CHAPTER XIII.
GROWTH AND VARIATION OF FISHES.
Changes of form normally accompanying growth (after absorption of the vitelline sac) are observed in all fishes; but in the majority they affect only the proportional size of the various parts of the body. In young fishes the eyes are constantly larger than in adult relatively to the size of the head; and again, the head is larger relatively to that of the body. Changes amounting to metamorphosis have been hitherto observed in Petromyzon only. In the larval condition (Ammocætes) the head is very small, and the toothless buccal cavity is surrounded by a semicircular upper lip. The eyes are extremely small, hidden in a shallow groove; and the vertical fins form a continuous fringe. In the course of three or four years the teeth are developed, and the mouth changes into a perfect suctorial organ; the eyes grow; and the dorsal fin is divided into two divisions. In Malacopterygians and Anacanths the embryonal fringe from which the vertical fins are developed, is much longer persistent than in Acanthopterygians. A metamorphosis relating to the respiratory organs, as in Batrachians, is indicated in the class of Fishes by the external gills with which fœtal Plagiostomes (Fig. [58], p. 136) and the young of some Ganoids, viz. the Protopterus and Polypterus, are provided.
Fig. 82.—Mouth of Larva of Petromyzon branchialis.
Fig. 83.—Mouth of Petromyzon fluviatilis.
mx, Maxillary tooth; md, Mandibulary tooth; l, Lingual tooth; s, Suctorial teeth.
Fig. 84.—Armature of præoperculum of young Caranx ferdau. (Magnified.)
I. Of an individual, 1¼ inch long. II. Of an individual, 2 inches long.
Fig. 85.—Tholichthys osseus. Six times the natural size.
Fig. 86.—Tholichthys-stage of Heniochus (?).
Fig. 87.—Tholichthys-stage of Pomacanthus (magn.) Atlantic.
Fig. 88.—Young Chætodon citrinellus (30 mill. long).
One of the most extraordinary changes by which, during growth, the form and position of several important organs are affected, occurs in Flat-fishes (Pleuronectidæ); their young are symmetrically formed, with a symmetrical mouth, and with one eye on each side, and, therefore, keep their body in a vertical position when swimming. As they grow they live more on the bottom, and their body, during rest, assumes a horizontal position; in consequence, the eye of the lower side moves towards the upper, which alone is coloured; and in many genera the mouth is twisted in the opposite direction, so that the bones, muscles, and teeth are much more developed on the blind side than on the coloured. In a great number of other Teleostei certain bones of the head show a very different form in the young state. Ossification proceeds in those bones in the direction of lines or radii which project in the form of spines or processes; as the interspaces between these processes are filled with bone, the processes disappear entirely, or at least project much less in the older than in the younger individuals (Fig. [84]). The young of some fishes may be armed with a long powerful præopercular or scapular spine, or show a serrature of which nothing remains in the adult fish except some ridges or radiating lines. These processes seem to serve as weapons of defence during a period in the life of the fish in which it needs them most. In not a few instances a portion of this armature is so much developed that the disappearance of its most projecting parts with the growth of the fish is not only due to its being surrounded by other bone, but, partially at least, caused by absorption. The Carangidæ, Cyttidæ, Squamipinnes, Xiphiidæ, offer instances of such remarkable changes. A fish, described as Tholichthys osseus (Fig. [85]), is probably the young of a Cyttoid, the suprascapula, humerus, and præoperculum forming enormously enlarged plates. In the fish Fig. [86] those bones appear still enlarged, and the frontals develop a remarkably long and curved horn above the orbit. In the Tholichthys-stage of Pomacanthus (specimens 10 millimetres long, Fig. [87]), the frontal bone is prolonged into a straight lancet-shaped process, nearly half as long as the body; the suprascapular and præopercular processes cover and hide the dorsal and ventral fins. The plates attached to the shoulder-girdle remain persistent until the young fish has assumed the form of the adult; thus they are still visible in young Chætodon citrinellus, 30 millimetres long, in which the specific characters are already fully developed.—The Sword-fishes with ventral fins (Histiophorus) belong to the Teleosteans of the largest size; in young individuals, 9 millimetres long (Fig. [89]), both jaws are produced, and armed with pointed teeth; the supraorbital margin is ciliated; the parietal and præoperculum are prolonged into long spines; the dorsal and anal fins are a low fringe, and the ventrals make their appearance as a pair of short buds. When 14 millimetres long (Fig. [90]) the young fish has still the same armature of the head, but the dorsal fin has become much higher, and the ventral filaments have grown to a great length. At a third stage, when the fish has attained to a length of 60 millimetres, the upper jaw is considerably prolonged beyond the lower, losing its teeth; the spines of the head are shortened, and the fins assume nearly the shape which they retain in mature individuals. Young Sword-fishes without ventral fins (Xiphias) undergo similar changes; and, besides, their skin is covered with small rough excrescences longitudinally arranged, which continue to be visible after the young fish has assumed the form of the mature in other respects (Fig. [92]).
Fig. 89.—Young Sword-fish (Histiophorus), 9 mill. long. Atlantic. (Magn.)
Fig. 90.—Young Sword-fish (Histiophorus), 14 mill. long. South Atlantic. (Magn.)
Fig. 91.—Young Sword-fish (Histiophorus), 60 mill. long. Mid-Atlantic.
Fig. 92.—Xiphias gladius, young, about 8 inches long.
The Plectognaths show no less extraordinary changes: an extraordinary form taken in the South Atlantic, and named Ostracion boops, is considered by Lütken to be the young of a Sun-fish (Orthagoriscus). In very young more advanced Sun-fishes (18 to 32 millimetres) the vertical diameter of the body exceeds, or is not much less than, the longitudinal; and small conical spines are scattered over its various parts. The caudal fin is developed long after the other vertical fins.
Fig. 93.—“Ostracion boops” (much magnified).
Fig. 94.—Young of Orthagoriscus, 18 and 32 mill. long. (Natural size.)
Similar changes take place in a number of other fishes, and in many cases the young are so different that they were described as distinct genera: thus Priacantichthys has proved to be the young of Serranus, Rhynchichthys that of Holocentrum, Cephalacanthus of Dactylopterus, Dicrotus of Thyrsites, Nauclerus of Naucrates, Porthmeus of Chorinemus, Lampugus of Coryphæna, Acronurus of Acanthurus, Keris of Naseus, Porobronchus of Fierasfer, Couchia of Motella, Stomiasunculus of Stomias, etc.
The fins are most frequently subject to changes; but, whilst in some fishes parts of them are prolonged into filaments with age, in others the filaments exist during the early life-periods only; whilst in some a part of the dorsal or the ventral fins is normally developed in the young only, in others those very parts are peculiar to the mature age. The integuments are similarly altered: in some species the young only has asperities on the skin, in others the young are smooth and the old have a tubercular skin; in some the young only have a hard bony head; in others (some Siluroids) the osseous carapace of the head and neck, as it appears in the adult, is more or less covered with soft skin whilst the fish is young.
In not a few fishes the external changes are in relation to the sexual development (Callionymus, many Labyrinthici, Cyprinodonts). These secondary sexual differences show themselves in the male individual, only when it commences to enter upon his sexual functions, and it may require two or more seasons before its external characteristics are fully developed. Immature males do not differ externally from the old female. The male secondary sexual characters consist principally in the prolongation of some of the fin-rays, or of entire fins; and in Salmonidæ in the greater development of the jaw-bones. The coloration of the male is in many fishes much brighter and more variegated than that of the female, but in comparatively few permanent (as in some Callionymus, Labrus mixtus); generally it is acquired immediately before and during the season of propagation only, and lost afterwards. Another periodical change in the integuments, also due to sexual influence and peculiar to the male, is the excrescence of wart-like tubercles on the skin of many Cyprinoids; they are developed chiefly on the head, but sometimes extend over the whole body and all the fins.
With regard to size, it appears that in all Teleosteous fishes the female is larger than the male; in many Cyprinodonts the male may be only one-sixth or even less of the bulk of the female. The observations on the relative size of the sexes are few in Palæichthyes, but such as have been made tend to show that, if a difference exists at all, the male is generally the larger (Lepidosteus). In the Rays (Raja) the sexes, after they have attained maturity, differ in the development of dermal spines and the form of the teeth, the female being frequently much rougher than the male. There is much variation in this respect in the different species; but the males are constantly distinguished by an oblong patch of erectile claw-like spines on each pectoral fin, and by having the teeth (all, or only a portion) pointed, and not obtuse, like those of the females. In Sharks no secondary sexual differences have been observed; the male Chimæridæ (see Fig. [96], p. 184), possess a singular comb-like cartilaginous appendage on the top of the head, which can be erected or depressed into a groove, both the appendage and the anterior part of the groove being armed with hooklets. The use of this singular organ is not known.
The majority of Teleostei are mixogamous—that is, the males and females congregate on the spawning-beds, and the number of the former being in excess, several males attend to the same female, frequently changing from one female to another. The same habit has been observed in Lepidosteus. Gastrosteus is truly polygamous, several females depositing their ova into the same nest, guarded by one male only. Some Teleostei (Ophiocephalus), and probably all Chondropterygians, are monogamous; and it is asserted that the connection between the pair is not merely temporary, but lasts until they are separated by accident. Monogamous are probably also all those Teleosteans which bring forth living young, and those, the males of which, for the attraction of the female, are provided with appendages, or ornamented with a bright coloration.
Hybridism is another source of changes and variations within the limits of a species, and is by no means so scarce as has been believed hitherto; it is only apparently of exceptional occurrence, because the life of fishes is more withdrawn from our direct observation than that of terrestrial animals. It has been observed among species of Serranus, Pleuronectidæ, Cyprinidæ, Clupeidæ, and especially Salmonidæ. As in other animals, the more certain kinds of fishes are brought under domestication, the more readily do they interbreed with other allied species. It is characteristic of hybrids that their characters are very variable, the degrees of affinity to one or the other of the parents being inconstant; and as these hybrids are known readily to breed with either of the parent race, the variations of form, structure, and colour are infinite. Of internal organs the dentition, gill-rakers, pyloric appendages, are those particularly affected by such mixture of species.
Some fishes are known to grow rapidly (in the course of from one to three years) and regularly to a certain size, growth being definitely arrested after the standard has been attained. Such fishes may be called “full-grown,” in the sense in which the term is applied to warm-blooded Vertebrates—the Sticklebacks, most Cyprinodonts, and many Clupeoids (Herring, Sprat, Pilchard) are examples of this regular kind of growth.[15] But in the majority of fishes the rate of growth is extremely irregular, and it is hardly possible to know when growth is actually and definitely arrested. All seems to depend on the amount of food and the more or less favourable circumstances under which the individual grows up. Fishes which rapidly grow to a definite size are short-lived, whilst those which steadily and slowly increase in size attain to a great age, Teleosteans as well as Chondropterygians. Carp and Pike have been ascertained to live beyond a hundred years.
It is evident that such diversity and irregularity of growth in the same species is accompanied by considerable differences in the appearance and general development of the fish. No instance is more remarkable than that of the so-called Leptocephali, which for a long time have been regarded either as a distinct group of Fishes, or as the larval stages of various genera of fishes.
Fig. 95.—Leptocephalus.
The Leptocephali proper are small, narrow, elongate, more or less band-shaped fishes, pellucid in a fresh state, but assuming a white colour when preserved in spirits, resembling a tapeworm, being quite as soft and flexible. The skeleton is entirely cartilaginous, or slight ossifications are only now and then visible, especially towards the end of the vertebral column. The latter is replaced by a chorda dorsalis which, in many specimens, is found to be divided into numerous segments. Neural arches are sometimes present in their rudimentary condition. The anterior end of the chorda passes into the cartilaginous base of the skull, the connection not being by means of joint and ligaments. Hæmal arches are found on the caudal portion. Ribs none. The skull, like the vertebral column, is nearly entirely cartilaginous. The basisphenoid, frontal, and jaw-bones are the first which may be distinguished, and the mandible has generally ossifications.
The muscles are generally not attached to the chorda, which is surrounded by a thick gelatinous mass, separating the lateral sets of muscles from each other. These muscles are attached to the external integument, each forming a thin flat angular band, the angle being directed forwards. However, specimens are frequently found in which the muscles are more developed, evidently at the expense of the gelatinous matter, which is diminished in quantity. They are attached to the chorda, and the entire fish has a more cylindrical form of the body (Helmichthys).
The nervous, circulatory, and respiratory organs are well developed. In those with a sub-cylindrical body the blood is red, in those with a flat body the blood-corpuscles show but rarely a faint coloration. There are four branchial arches, and in some (Tilurus) pseudobrauchiæ have been found. The gill-openings are more or less narrow. The nostrils are double on each side, and the posterior is close to the eye.
The stomach has a large blind sac, and in Leptocephalus two lateral cæca. The intestine is straight, running close to the abdominal profile, with a small appendix directed forward and a larger one directed backwards. The vent is nearly always very small, and, in preserved examples at least, cannot always be discovered. Its position is variable, even in examples entirely similar in other points. Air-bladder none. No trace of generative organs.
The vertical fins, when present, are confluent, with more or less conspicuous traces of rays; sometimes they are merely a fold of the skin, without any rays. Pectoral fins sometimes present, sometimes rudimentary, sometimes entirely absent. Ventrals none.
Most examples have series of round black dots along each side of the abdominal profile, along the lateral line, and sometimes along the dorsal fin. They remind us of the luminous organs of many Scopelidæ, Stomiatidæ, and other pelagic fishes, but are composed entirely of pigmentary cells.
These fishes are found floating in the sea, frequently at a great distance from land. Their movements are slow and languid. The largest specimen of Leptocephalus observed was 10 inches, but specimens of that size are very rare.
[See Kölliker, Zeitschr. wiss. Zool. iv. 1852, p. 360; and Carus, Ueber die Leptocephaliden. Leipz. 1861. 4to.]
Taking into account all the various facts mentioned, we must come to the conclusion that the Leptocephalids are the offspring of various kinds of marine fishes, representing, not a normal stage of development (larvæ), but an arrest of development at a very early period of their life; they continue to grow to a certain size without corresponding development of their internal organs, and perish without having attained the characters of the perfect animal. The cause by which this abnormal condition is brought about is not known; but it is quite within the limits of probability that fishes usually spawning in the vicinity of land sometimes spawn in the open ocean, or that floating spawn is carried by currents to a great distance from land; and that such embryoes, which for their normal growth require the conditions afforded by the vicinity of the shore, if hatched in mid-ocean, grow into undeveloped hydropic creatures, such as the Leptocephales seem to be.
Abundance or scarcity of food, and other circumstances connected with the localities inhabited by fishes, affect considerably the colour of their muscles and integuments; the periodical changes of colour in connection with their sexual functions have been referred to above (p. 176). The flesh of many Teleostei is colourless, or but slightly tinged by the blood; that of Scombridæ, most Ganoids and Chondropterygians, is more or less red; but in badly-fed fishes, as well as in very young ones, the flesh is invariably white (anæmic). Many fishes, like the Salmonidæ, feed at times exclusively on Crustaceans, and the colouring substance of these Invertebrates, which by boiling and by the stomachic secretion turns red, seems to pass into the flesh of the fishes, imparting to it the well-known “salmon” colour. Further, the coloration of the integuments of many marine fish is dependent on the nature of their surroundings. In those which habitually hide themselves on the bottom, in sand, between stones or seaweeds, the colours of the body readily assimilate to those of the vicinity, and are thus an important element in the economy of their life. The changes from one set or tinge of colours to another may be rapid and temporary, or more or less permanent; in some fishes—as in the Pediculati, of which the Sea-Devil, or Lophius, and Antennarius are members—scarcely two individuals are found exactly alike in coloration, and only too frequently such differences in coloration are mistaken for specific characters. The changes of colours are produced in two ways: either by an increase or decrease of the black, red, yellow, etc., pigment-cells, or chromatophors, in the skin of the fish; or by the rapid contraction or expansion of the chromatophors which happen to be developed. The former change is gradual, like every kind of growth or development; the latter rapid, owing to the great sensitiveness of the cells, but certainly involuntary. In many bright-shining fishes—as Mackerels, Mullets—the colours appear to be brightest in the time intervening between the capture of the fish and its death: a phenomenon clearly due to the pressure of the convulsively-contracted muscles on the chromatophors. External irritation readily excites the chromatophors to expand—a fact unconsciously utilised by fishermen, who, by scaling the Red Mullet immediately before its death, produce the desired intensity of the red colour of the skin, without which the fish would not be saleable. However, it does not require such strong measures to prove the sensitiveness of the chromatophors to external irritation, the mere change of darkness into light is sufficient to induce them to contract, the fish appearing paler, and vice versa. In Trout which are kept or live in dark places, the black chromatophors are expanded, and, consequently, such specimens are very dark-coloured; when removed to the light they become paler almost instantaneously.
Total absence of chromatophors in the skin, or Albinism, is very rare among fishes; much more common is incipient Albinism, in which the dark chromatophors are changed into cells with a more or less intense yellow pigment. Fishes in a state of domestication, like the Crucian Carp of China, the Carp, Tench, and the Ide, are particularly subject to this abnormal coloration, and are known as the common Gold-fish, the Gold-Tench, and the Gold-Orfe. But it occurs also not rarely in fishes living in a wild state, and has been observed in the Haddock, Flounder, Plaice, Carp, Roach, and Eel.
It will be evident, from the foregoing remarks, that the amount of variation within the limits of the same species—either due to the natural growth and development, or to external physical conditions, or to abnormal accidental circumstances—is greater in fishes than in any of the higher classes of Vertebrates. The amount of variation is greater in certain genera or families than in others, and it is much greater in Teleosteans and Ganoids than in Chondropterygians. Naturally, it is greatest in the few species which have been domesticated, and which we shall mention in the succeeding chapter.
Fig. 96.—Chimæra colliei ♂, west coast of North America. A. Front view of head. B. Palate. a, Peritoneal aperture; b, Nostrils; c, Vomerine teeth; d, Mandibular teeth; e, Palatine teeth; f, Claspers.
CHAPTER XIV.
DOMESTICATED AND ACCLIMATISED FISHES; ARTIFICIAL IMPREGNATION OF OVA—TENACITY OF LIFE AND REPRODUCTION OF LOST PARTS—HYBERNATION—USEFUL AND POISONOUS FISHES.
A few fishes only are thoroughly domesticated—that is, bred in captivity, and capable of transportation within certain climatic limits—viz. the Carp, Crucian Carp (European and Chinese varieties), Tench, Orfe or Ide, and the Goramy. The two former have accompanied civilised man almost to every place of the globe where he has effected a permanent settlement.
Attempts to acclimatise particularly useful species in countries in which they were not indigenous have been made from time to time, but were permanently successful in a few instances only; the failures being due partly to the choice of a species which did not yield the profitable return expected, partly to the utter disregard of the difference of the climatic and other physical conditions between the original and new homes of the fish. The first successful attempts of acclimatisation were made with domestic species, viz. the Carp and Gold-fish, which were transferred from Eastern Asia to Europe. Then, in the first third of the present century, the Javanese Goramy was acclimatised in Mauritius and Guiana, but no care seems to have been taken to insure permanent advantages from the successful execution of the experiment. In these cases fully developed individuals were transported to the country in which they were to be acclimatised. The most successful attempt of recent years is the acclimatisation of the Trout and Sea-Trout, and probably also of the Salmon, in Tasmania and New Zealand, and of the Californian Salmon (Salmo quinnat?), in Victoria, by means of artificially-impregnated ova. The ova were transported on ice, in order to retard their development generally, and thus to preserve them from destruction during the passage of the tropical zone.
Artificial impregnation of fish-ova was first practised by J. L. Jacobi, a native of Westphalia, in the years 1757–63, who employed exactly the same method which is followed now; and there is no doubt that this able observer of nature conceived and carried out his idea with the distinct object of advantageously restocking water-courses which had become unproductive, and increasing production by fecundating and preserving all ova, of which a great proportion, in the ordinary course of propagation, would be left unfecundated or accidentally perish. Physiology soon turned to account Jacobi’s discovery, and artificial impregnation has proved to be one of the greatest helps to the student of embryology.
Fishes differ in an extraordinary degree with regard to tenacity of life. Some will bear suspension of respiration—caused by removal from water, or by exposure to cold or heat—for a long time, whilst others succumb at once. Nearly all marine fishes are very sensitive to changes in the temperature of the water, and will not bear transportation from one climate to another. This seems to be much less the case with some freshwater fishes of the temperate zones: the Carp may survive after being frozen in a solid block of ice, and will thrive in the southern parts of the temperate zone. On the other hand, some freshwater fishes are so sensitive to a change in the water that they perish when transplanted from their native river into another apparently offering the same physical conditions (Grayling, Salmo hucho). Some marine fishes may be abruptly transferred from salt into fresh water, like Sticklebacks, some Blennies, and Cottus, etc.; others survive the change when gradually effected, as many migratory fishes; whilst again, others cannot bear the least alteration in the composition of the salt water (all pelagic fishes). On the whole, instances of marine fishes voluntarily entering brackish or fresh water are very numerous, whilst freshwater fishes proper but rarely descend into salt water.
Abstinence from food affects different fishes in a similarly different degree. Marine fishes can endure hunger less than freshwater fishes, at least in the temperate zones, no observations having been made in this respect on tropical fishes. Goldfishes, Carps, Eels, are known to be able to subsist without food for months, without showing a visible decrease of bulk; whilst the Trigloids, Sparoids, and other marine fishes, survive abstinence from food for a few days only. In freshwater fishes the temperature of the water is of great influence on their vital functions generally, and consequently on their appetite,—many cease to feed altogether in the course of the winter; a few, like the Pike, are less inclined to feed during the heat of the summer than when the temperature is lowered.
Captivity is easily borne by most fishes, and the appliances introduced in our modern aquaria have rendered it possible to keep in confinement, and even to induce to propagate, fishes which formerly were considered to be intolerant of captivity.
Wounds affect fishes generally much less than higher Vertebrates. A Greenland Shark continues to feed whilst his head is pierced by a harpoon or by the knife, as long as the nervous centre is not touched; a Sea-perch or a Pike (Fig. [97]) will survive the loss of a portion of its tail; a Carp that of half of its snout. However, some fishes are much more sensitive, and perish even from the superficial abrasion caused by the meshes of the net during capture (Mullsn.)
The power of reproduction of lost parts in Teleosteous fishes is limited to the delicate terminations of their fin-rays and the various tegumentary filaments with which some are provided. These filaments are sometimes developed in an extraordinary degree, mimicking the waving fronds of the seaweed in which the fish hides. Both the ends of the fin-rays and the filaments are frequently lost, not only by accident, but merely by wear and tear; and as these organs are essential for the preservation of the fish, their reproduction is necessary.
Fig. 97.—Pike caught in the Thames, which, when young, had lost part of the tail with the caudal fin.
In Dipnoi, Ceratodus, and Protopterus, the terminal portion of the tail has been found to have been reproduced, but without the notochord.
Hybernation has been observed in many Cyprinoids and Murænoids of the temperate zones. They do not fall into a condition of complete torpidity, as Reptiles and Mammals, but their vital functions are simply lowered, and they hide in sheltered holes, and cease to go abroad in search of their food. Between the tropics a great number of fishes (especially Siluroids, Labyrinthici, Ophiocephaloids, the Dipnoi), are known to survive long-continued droughts by passing the dry season in a perfectly torpid state, imbedded in the hardened mud. Protopterus, and probably many of the other fishes mentioned, prepare for themselves a cavity large enough to hold them, and coated on the inside with a layer of hardened mucus, which preserves them from complete desiccation. It has been stated that in India fishes may survive in this condition for more than one season, and that ponds known to have been dry for several years, and to the depth of many feet, have swarmed with fishes as soon as the accumulation of water released them from their hardened bed.
The principal use derived by man from the class of Fishes consists in the abundance of wholesome and nourishing food which they yield. In the Polar regions especially, whole tribes are entirely dependent on this class for subsistence; and in almost all nations fishes form a more or less essential part of food, many being, in a preserved condition, most important articles of trade. The use derived by man from them in other respects is of but secondary importance. Cod-liver oil is prepared from the liver of some of the Gadoids of the Northern Hemisphere, and of Sharks; isinglass from the swim-bladder of Sturgeons, Sciænoids, and Polynemoids; shagreen from the skin of Sharks and Rays.
The flesh of some fishes is at times, or constantly, poisonous. When eaten, it causes symptoms of more or less intense irritation of the stomach and intestines, inflammation of the mucous membranes, and not rarely death. The fishes, the flesh of which appears always to have poisonous properties, are Clupea thrissa, Clupea venenosa, and some species of Scarus, Tetrodon, and Diodon. There are many others which have occasionally or frequently caused symptoms of poisoning. Poey enumerates not less than seventy-two different kinds from Cuba; and various species of Sphyræna, Balistes, Ostracion, Caranx, Lachnolæmus, Tetragonurus, Thynnus, have been found to be poisonous in all seas between the tropics. All or nearly all these fishes acquire their poisonous properties from their food which consists of poisonous Medusæ, Corals, or decomposing substances. Frequently the fishes are found to be eatable if the head and intestines be removed immediately after capture. In the West Indies it has been ascertained that all the fishes living and feeding on certain coral banks are poisonous. In other fishes the poisonous properties are developed at certain seasons of the year only, especially the season of propagation: as the Barbel, Pike, and Burbot, whose roe causes violent diarrhœas when eaten during the season of spawning.
Fig. 98.—Portion of tail, with spines, of Aëtobatis narinari, a Sting-ray from the Indian Ocean. a, nat. size.
Poison-organs are more common in the class of Fishes than was formerly believed, but they seem to have exclusively the function of defence, and are not auxiliary in procuring food, as in venomous Snakes. Such organs are found in the Sting-rays, the tail of which is armed with one or more powerful barbed spines. Although they lack a special organ secreting poison, or a canal in or on the spine by which the venomous fluid is conducted, the symptoms caused by a wound from the spine of a Sting-ray are such as cannot be accounted for merely by the mechanical laceration, the pain being intense, and the subsequent inflammation and swelling of the wounded part terminating not rarely in gangrene. The mucus secreted from the surface of the fish and inoculated by the jagged spine evidently possesses venomous properties. This is also the case in many Scorpænoids, and in the Weaver (Trachinis), in which the dorsal and opercular spines have the same function as the caudal spines of the Sting-rays; however, in the Weavers the spines are deeply grooved, the groove being charged with a fluid mucus. In Synanceia the poison-organ (Fig. [99],) is still more developed: each dorsal spine is in its terminal half provided with a deep groove on each side, at the lower end of which lies a pear-shaped bag containing the milky poison; it is prolonged into a membranous duct, lying in the groove of the spine, and open at its point. The native fishermen, well acquainted with the dangerous nature of these fishes, carefully avoid handling them; but it often happens that persons wading with naked feet in the sea, step upon the fish, which generally lies hidden in the sand. One or more of the erected spines penetrate the skin, and the poison is injected into the wound by the pressure of the foot on the poison-bags. Death has not rarely been the result.
Fig. 99.—A dorsal spine, with poison-bags, of Synanceia verrucosa. Indian Ocean.
Fig. 100.—Opercular part of the Poison-apparatus of Thalassophryne (Panama).
1. Hinder half of the head, with the venom-sac* in situ. a, Lateral line and its branches; b, Gill-opening; c, Ventral fin; d, Base of Pectoral fin; e, Base of dorsal.
2. Operculum with the perforated spine.
The most perfect poison-organs hitherto discovered in fishes are those of Thalassophryne, a Batrachoid genus of fishes from the coasts of Central America. In these fishes the operculum again and the two dorsal spines are the weapons. The former (Fig. [100], 2) is very narrow, vertically styliform and very mobile; it is armed behind with a spine, eight lines long, and of the same form as the hollow venom-fang of a snake, being perforated at its base and at its extremity. A sac covering the base of the spine discharges its contents through the apertures and the canal in the interior of the spine. The structure of the dorsal spines is similar. There are no secretory glands imbedded in the membranes of the sacs; and the fluid must be secreted by their mucous membrane. The sacs are without an external muscular layer, and situated immediately below the thick loose skin which envelops the spines to their extremity; the ejection of the poison into a living animal, therefore, can only be effected, as in Synanceia, by the pressure to which the sac is subjected the moment the spine enters another body.
Finally, a singular apparatus found in many Siluroids may be mentioned in connection with the poison-organs, although its function is still problematical. Some of these fishes are armed with powerful pectoral spines and justly feared on account of the dangerous wounds they inflict; not a few of them possess, in addition to the pectoral spines, a sac with a more or less wide opening in the axil of the pectoral fin; and it does not seem improbable that it contains a fluid which may be introduced into a wound by means of the pectoral spine, which would be covered with it, like the barbed arrow-head of an Indian. However, whether this secretion is equally poisonous in all the species provided with that axillary sac, or whether it has poisonous qualities at all, is a question which can be decided by experiments only made with the living fishes.
CHAPTER XV.
DISTRIBUTION OF FISHES IN TIME.
Of what kind the fishes were which were the first to make their appearance on the globe; whether or not they were identical with, or similar to, any of the principal types existing at present; are questions which probably will for ever remain hidden in mystery and uncertainty. The supposition that the Leptocardii and Cyclostomes, the lowest of the vertebrate series, must have preceded the other sub-classes, is an idea which has been held by many Zoologists: and as the horny teeth of the Cyclostomes are the only parts of their body which under favourable circumstances might have been preserved, Palæontologists have ever been searching for this evidence.
Fig. 101. Right dental plate of Myxine affinis.
Indeed, in deposits belonging to the Lower Silurian and Devonian, in Russia, England, and North America, minute, slender, pointed horny bodies, bent like a hook, with sharp opposite margins, have been found and described under the name of Conodonts. More frequently they possess an elongated basal portion, in which there is generally a larger tooth with rows of similar but smaller denticles on one or both sides of the larger tooth, according as this is central or at one end of the base. In other examples there is no prominent central tooth, but a series of more or less similar teeth is implanted on a straight or curved base. Modifications of these arrangements are very numerous, and many Palæontologists entertain still doubts whether the origin of these remains is not rather from Annelids and Mollusks than from Fishes.
[See G. J. Hinde, in “Quarterly Journal of the Geological Society,” 1879.]
The first undeniable evidence of a fish, or, indeed, of a vertebrate animal, occurs in the Upper Silurian Rocks, in a bone-bed of the Downton sandstone, near Ludlow. It consists of compressed, slightly curved, ribbed spines, of less than two inches in length (Onchus); of small shagreen-scales (Thelodus); the fragment of a jaw-like bar with pluricuspid teeth (Plectrodus); the cephalic bucklers of what seems to be a species of Pteraspis; and, finally, the coprolitic bodies of phosphate and carbonate of lime, including recognisable remains of the Mollusks and Crinoids inhabiting the same waters. But no vertebra or other part of the skeleton has been found. The spines and scales seem to have belonged to the same kind of fish, which probably was a Plagiostome. It is quite uncertain whether or not the jaw (if it be the jaw of a fish[16]) belonged to the buckler-bearing Pteraspis, the position of which among Ganoids, with which it is generally associated, is open to doubt.
No detached undoubted tooth of a Plagiostome or Ganoid scale has been discovered in the Ludlow deposits: but so much is certain that those earliest remains in Palæozoic rocks belonged to fishes closely allied to forms occurring in greater abundance in the succeeding formation, the Devonian, where they are associated with undoubted Palæichthyes, Plagiostomes as well as Ganoids.
These fish-remains of the Devonian or Old Red Sandstone, can be determined with greater certainty. They consist of spines or the so-called Ichthyodorulites, which show sufficiently distinctive characters to be referred to several genera, one of them, Onchus, still surviving from the Silurian epoch. All these spines are believed to be those of Chondropterygians, to which order some pluricuspid teeth (Cladodus) from the Old Red Sandstone in the vicinity of St. Petersburg have been referred likewise.
The remains of the Ganoid fishes are in a much more perfect state of preservation, so that it is even possible to obtain a tolerably certain idea of the general appearance and habits of some of them, especially of such as were provided with hard carapaces, solid scales, and ordinary or bony fin-rays. A certain proportion of them, as might have been expected, remind us, with regard to external form, of Teleosteous fishes rather than of any of the few still existing Ganoid types; but it is contrary to all analogy and to all palæontological evidence to suppose that those fishes were, with regard to their internal structure, more nearly allied to Teleosteans than to Ganoids. If they were not true Ganoids, they may be justly supposed to have had the essential characters of Palæichthyes. Other forms exhibit even at that remote geological epoch so unmistakably the characteristics of existing Ganoids, that no one can entertain any doubt with regard to their place in the system. In none of these fishes is there any trace of vertebral segmentation.
The Palæichthyes of the Old Red Sandstone, the systematic position of which is still obscure, are the Cephalaspidæ from the Lower Old Red Sandstone of Great Britain and Eastern Canada; Pterichthys, Coccosteus, and Dinichthys: genera which have been combined in one group—Placodermi; and Acanthodes and allied genera, which combined numerous branchiostegals with chondropterygian spines and a shagreen-like dermal covering.
Among the other Devonian fishes (and they formed the majority) two types may be recognised, both of which are unmistakably Ganoids. The first approaches the still living Polypterus, with which some of the genera like Diplopterus singularly agree in the form and armature of the head, the lepidosis of the body, the lobate pectoral fins, and the termination of the vertebral column. Other genera, as Holoptychius, have cycloid scales; many have two dorsal fins (Holoptychius), and, instead of branchiostegals, jugular scutes; others one long dorsal confluent with the caudal (Phaneropleuron).
In the second type the principal characters of the Dipnoi are manifest, and some of them, for example Dipterus, Palædaphus, Holodus, approach so closely the Dipnoi which still survive, that the differences existing between them warrant a separation into families only.
Devonian fishes are frequently found under peculiar circumstances, enclosed in the so-called nodules. These bodies are elliptical flattened pebbles, which have resisted the action of water in consequence of their greater hardness, whilst the surrounding rock has been reduced to detritus by that agency. Their greater density is due to the dispersion in their substance of the fat of the animal which decomposed in them. Frequently, on cleaving one of these nodules with the stroke of the hammer, a fish is found embedded in the centre. At certain localities of the Devonian, fossil fishes are so abundant that the whole of the stratum is affected by the decomposing remains emitting a peculiar smell when newly opened, and acquiring a density and durability not possessed by strata without fishes. The flagstones of Caithness are a remarkable instance of this.
The fish-remains of the Carboniferous formation show a great similarity to those of the preceding. They occur throughout the series, but are very irregularly distributed, being extremely scarce in some countries, whilst in others entire beds (the so-called bone-beds) are composed of ichthyolites. In the ironstones they frequently form the nuclei of nodules, as in the Devonian.
Of Chondropterygians the spines of Onchus and others still occur, with the addition of teeth indicative of the existence of fishes allied to the Cestracion-type (Cochliodus, Psammodus): a type which henceforth plays an important part in the composition of the extinct marine fish faunæ. Another extinct Selachian family, that of Hybodontes, makes its appearance, but is known from the teeth only.
Of the Ganoid fishes, the family Palæoniscidæ (Traquair) is numerously represented; others are Cœlacanths (Cœlocanthus, Rhizodus), and Saurodipteridæ (Megalichthys). None of these fishes have an ossified vertebral column, but in some (Megalichthys) the outer surface of the vertebræ is ossified into a ring; the termination of their tail is heterocercal. The carboniferous Uronemus and the Devonian Phaneropleuron are probably generically the same; and the Devonian Dipnoi are continued as, and well represented by, Ctenodus.
The fishes of the Permian group are very similar to those of the Carboniferous. A type which in the latter was but very scantily represented, namely the Platysomidæ, is much developed. They were deep-bodied fish, covered with hard rhomboid scales possessing a strong anterior rib, and provided with a heterocercal caudal, long dorsal and anal, short non-lobate paired fins (when present), and branchiostegals. The Palæoniscidæ are represented by many species of Palæoniscus, Pygopterus and Acrolepis, and Cestracionts by Janassa and Strophodus.
The passage from the Palæozoic into the Mesozoic era is not indicated by any marked change as far as fishes are concerned. The more remarkable forms of the Trias are Shark-like fishes represented by ichthyodorulithes like Nemacanthus, Liacanthus, and Hybodus; and Cestracionts represented by species of Acrodus and Strophodus. Of the Ganoid genera Cœlacanthus, Amblypterus (Palæoniscidæ), Saurichthys persist from the Carboniferous epoch. Ceratodus appears for the first time (Muschel-Kalk of Germany).
Thanks to the researches of Agassiz, and especially Sir P. Egerton, the ichthyological fauna of the Lias is, perhaps, the best known of the Mesozoic era, 152 species having been described. Of the various localities, Lyme Regis has yielded more than any other, nearly all the Liassic genera being represented there by not less than seventy-nine species. The Hybodonts and Cestracionts continue in their fullest development. Holocephales (Ischyodus), true Sharks (Palæoscyllium), Rays (Squaloraja, Arthropterus), and Sturgeons (Chondrosteus) make their first appearance; but they are sufficiently distinct from living types to be classed in separate genera, or even families. The Ganoids, especially Lepidosteoids, predominate over all the other fishes: Lepidotus, Semionotus, Pholidophorus, Pachycormus, Eugnathus, Tetragonolepis, are represented by numerous species; other remarkable genera are Aspidorhynchus, Belonostomus, Saurostomus, Sauropsis, Thrissonotus, Conodus, Ptycholepis, Endactis, Centrolepis, Legnonotus, Oxygnathus, Heterolepidotus, Isocolum, Osteorhachis, Mesodon. These genera offer evidence of a great change since the preceding period, the majority not being represented in older strata, whilst, on the other hand, many are continued into the succeeding oolithic formations. The homocercal termination of the vertebral column commences to supersede the heterocercal, and many of the genera have well ossified and distinctly segmented spinal columns. Also the cycloid form of scales becomes more common: one genus (Leptolepis) being, with regard to the preserved hard portions of its organisation, so similar to the Teleosteous type that some Palæontologists refer it (with much reason) to that sub-class.
[See E. Sauvage, Essai sur la Faune Ichthyologique de la période Liasique. In “Bibl. de l’école des hautes études,” xiii. art. 5. Paris 1875. 8o.]
As already mentioned, the Oolithic formations show a great similarity of their fish-fauna to that of the Lias; but still more apparent is its approach to the existing fauna. Teeth have been found which cannot even generically be distinguished from Notidanus. The Rays are represented by genera like Spathobatis, Belemnobatis, Thaumas; the Holocephali are more numerous than in the Lias (Ischyodus, Ganodus). The most common Ganoid genera are Caturus, Pycnodus, Pholidophorus, Lepidotus, Leptolepis, all of which had been more or less fully represented in the Lias. Also Ceratodus is continued into it.
The Cretaceous group offers clear evidence of the further advance towards the existing fauna. Teeth of Sharks of existing genera Carcharias (Corax), Scyllium, Notidanus, and Galeocerdo, are common in some of the marine strata, whilst Hybodonts and Cestracionts are represented by a small number of species only; of the latter one new genus, Ptychodus, appears and disappears. A very characteristic Ganoid genus, Macropoma, comprises homocercal fishes with rounded ganoid scales sculptured externally and pierced by prominent mucous tubes. Caturus becomes extinct. Teeth and scales of Lepidotus (with Sphærodus as subgenus), clearly a freshwater fish, are widely distributed in the Wealden, and finally disappear in the chalk; its body was covered with large rhomboidal ganoid scales. Gyrodus and Aspidorhynchus occur in the beds of Voirons, Coelodus and Amiopsis (allied to Amra), in those of Comen, in Istria. But the Palæichthyes are now in the minority; undoubted Teleosteans have appeared, for the first time, on the stage of life in numerous genera, many of which are identical with still existing fishes. The majority are Acanthopterygians, but Physostomes and Plectognaths are likewise well represented, most of them being marine. Of Acanthopterygian families the first to appear are the Berycidæ, represented by several very distinct genera: Beryx; Pseudoberyx with abdominal ventral fins; Berycopsis with cycloid scales; Homonotus, Stenostoma, Sphenocephalus, Acanus, Hoplopteryx, Platycornus with granular scales; Podocys with a dorsal extending to the neck; Acrogaster, Macrolepis, Rhacolepis from the chalk of Brazil. The position of Pycnosterynx is uncertain, it approaches certain Pharyngognaths. True Percidæ are absent, whilst the Carangidæ, Sphyrænidæ, Cataphracti, Gobiidæ, Cottidæ, and Sparidæ are represented by one or more genera. Somewhat less diversified are the Physostomes, which belong principally to the Clupeidæ and Dercetidæ, most of the genera being extinct; Clupea is abundant in some localities. Scopelidæ (Hemisaurida and Saurocephalus) occur in the chalk of Comen in Istria, and of Mæstricht. Of all cretaceous deposits none surpass those of the Lebanon with regard to the number of genera, species, and individuals; the forms are exclusively marine, and the remains in the most perfect condition.
In the Tertiary epoch the Teleosteans have almost entirely replaced the Ganoids; a few species only of the latter make their appearance, and they belong to existing genera, or, at least, very closely allied forms (Lepidosteus, Amia, Hypamia, Acipenser). The Chondropterygians merge more and more into recent forms; Holocephali continue, and still are better represented than in the present fauna. The Teleosteans show even in the Eocene a large proportion of existing genera, and the fauna of some localities of the Miocene (Oeningen) is almost wholly composed of them. On the whole, hitherto more than one-half have been found to belong to existing genera, and there is no doubt that the number of seemingly distinct extinct genera will be lessened as the fossils will be examined with a better knowledge of the living forms. The distribution of the fishes differed widely from that of our period, many of our tropical genera occurring in localities which are now included within our temperate zone, and being mixed with others, which nowadays are restricted to a colder climate: a mixture which continues throughout the Pliocene.
A few families of fishes, like the freshwater Salmonidæ, seem to have put in their appearance in Post-pliocene times; however, not much attention has been paid to fish-remains of these deposits; and such as have been incidentally examined offer evidence of the fact that the distribution of fishes has not undergone any further essential change down to the present period.
[See E. Sauvage, Mémoire sur la Faune Ichthyologique de la période Tertiaire. Paris 1873. 8°.]
Fig. 102.—Pycnodus rhombus, a Ganoid from the Upper Oolite.
CHAPTER XVI.
THE DISTRIBUTION OF EXISTING FISHES OVER THE EARTH’S SURFACE—GENERAL REMARKS.
In an account of the geographical distribution of fishes the Freshwater forms are to be kept separate from the Marine. However, when we attempt to draw a line between these two kinds of fishes, we meet with a great number of species and of facts which would seem to render that distinction very vague. There are not only species which can gradually accommodate themselves to a sojourn in either salt or fresh water, but there are also such as seem to be quite indifferent to a rapid change from one into the other: so that individuals of one and the same species (Gastrosteus, Gobius, Blennius, Osmerus, Retropinna, Clupea, Syngnathus, etc.), may be found at some distance out at sea, whilst others live in rivers far beyond the influence of the tide, or even in inland fresh waters without outlet to the sea. The majority of these fishes belong to forms of the fauna of the brackish water, and as they are not an insignificant portion of the fauna of almost every coast, we shall have to treat of them in a separate chapter.
Almost every large river offers instances of truly marine fishes (such as Serranus, Sciænidæ, Pleuronectes, Clupeidæ, Tetrodon, Carcharias, Trygonidæ), ascending for hundreds of miles of their course; and not periodically, or from any apparent physiological necessity, but sporadically throughout the year, just like the various kinds of marine Porpoises which are found all along the lower course of the Ganges, Yang-tseKiang, the Amazons, the Congo, etc. This is evidently the commencement of a change in a fish’s habits, and, indeed, not a few of such fishes have actually taken up their permanent residence in fresh waters (as species of Ambassis, Apogon Dules, Therapon, Sciæna, Blennius, Gobius, Atherina, Mugil, Myxus, Hemirhamphus, Clupea, Anguilla, Tetrodon, Trygon): all forms originally marine.
On the other hand, we find fishes belonging to freshwater genera descending rivers and sojourning in the sea for a more or less limited period; but these instances are much less in number than those in which the reverse obtains. We may mention species of Salmo (the Common Trout, the Northern Charr), and Siluroids (as Arius, Plotosus). Coregonus, a genus so characteristic of the inland lakes of Europe, Northern Asia, and North America, nevertheless offers some instances of species wandering by the effluents into the sea, and taking up their residence in salt water, apparently by preference, as Coregonus oxyrhynchus. But of all the Freshwater families none exhibit so great a capability of surviving the change from fresh into salt water, as the Gastrosteidæ (Sticklebacks), of the northern Hemisphere, and the equally diminutive Cyprinodontidæ of the tropics; not only do they enter into, and live freely in, the sea, but many species of the latter family inhabit inland waters, which, not having an outlet, have become briny, or impregnated with a larger proportion of salts than pure sea water. During the voyage of the “Challenger” a species of Fundulus, F. nigrofasciatus, which inhabits the fresh and brackish waters of the Atlantic States of North America, was obtained, with Scopelids and other pelagic forms, in the tow-net, midway between St. Thomas and Teneriffe.
Some fishes annually or periodically ascend rivers for the purpose of spawning, passing the rest of the year in the sea, as Sturgeons, many Salmonoids, some Clupeoids, Lampreys, etc. The two former evidently belonged originally to the freshwater series, and it was only in the course of their existence that they acquired the habit of descending to the sea, perhaps because their freshwater home did not offer a sufficient supply of food. These migrations of freshwater fishes have been compared with the migrations of birds; but they are much more limited in extent, and do not impart an additional element to the fauna of the place to which they migrate, as is the case with the distant countries to which birds migrate.
The distinction between freshwater and marine fishes is further obscured by geological changes, in consequence of which the salt water is gradually being changed into fresh, or vice versa. These changes are so gradual and spread over so long a time, that many of the fishes inhabiting such localities accommodate themselves to the new conditions. One of the most remarkable and best studied instances of such an alteration is the Baltic, which, during the second half of the Glacial period, was in open and wide communication with the Arctic Ocean, and evidently had the same marine fauna as the White Sea. Since then, by the rising of the land of Northern Scandinavia and Finland, this great gulf of the Arctic Ocean has become an inland sea, with a narrow outlet into the North Sea, and its water, in consequence of the excess of the fresh water pouring into it over the loss by evaporation, has been so much diluted as to be nearly fresh at its northern extremities: and yet nine species, the origin of which from the Arctic Ocean can be proved, have survived the changes, propagating their species, agreeing with their brethren in the Arctic Ocean in every point, but remaining comparatively smaller. On the other hand, fishes which we must regard as true freshwater fishes, like the Rudd, Roach, Pike, Perch, enter freely the brackish water of the Baltic.[17] Instances of marine fishes being permanently retained in fresh water in consequence of geological changes are well known: thus Cottus quadricornis in the large lakes of Scandinavia; species of Gobius, Blennius, and Atherina in the lakes of Northern Italy; Comephorus, of the depths of the Lake of Baikal, which seems to be a dwarfed Gadoid. Carcharias gangeticus in inland lakes of the Fiji Islands, is another instance of a marine fish which has permanently established itself in fresh water.
In the miocene formation of Licata in Sicily, in which fish remains abound, numerous Cyprinoids are mixed with littoral and pelagic forms. Sauvage found in 450 specimens from that locality, not less than 266, which were Leucisci, Alburni, or Rhodei. Now, although it is quite possible that in consequence of a sudden catastrophe the bodies of those Cyprinoids were carried by a freshwater current into, and deposited on the bottom of, the sea, the surmise that they lived together with the littoral fishes in the brackish water of a large estuary, which was not rarely entered by pelagic forms, is equally admissible. And, if confirmed by other similar observations, this instance of a mixture of forms which are now strictly freshwater or marine, may have an important bearing on the question to what extent fishes have in time changed their original habitat.
Thus there is a constant exchange of species in progress between the freshwater and marine faunæ, and in not a few cases it would seem almost arbitrary to refer a genus or even larger group of fishes to one or the other; yet there are certain groups of fishes which entirely, or with but few exceptions are, and, apparently, during the whole period of their existence have been, inhabitants either of the sea or of fresh water; and as the agencies operating upon the distribution of marine fishes differ greatly from those influencing the dispersal of freshwater fishes, the two series must be treated separately. The most obvious fact that dry land, which intervenes between river systems, offers to the rapid spreading of a freshwater fish an obstacle which can be surmounted only exceptionally or by a most circuitous route, whilst marine fishes may readily and voluntarily extend their original limits, could be illustrated by a great number of instances. Without entering into details, it may suffice to state as the general result, that no species or genus of freshwater fishes has anything like the immense range of the corresponding categories of marine fishes; and that, with the exception of the Siluroids, no other freshwater family is so widely spread as the families of marine fishes. Surface temperature or climate which is, if not the most, one of the most important physical factors in the limitation of freshwater fishes, similarly affects the distribution of marine fishes, but in a less degree, and only those which live near to the shore or the surface of the ocean; whilst it ceases to exercise its influence in proportion to the depth, the true deep-sea forms being entirely exempt from its operation. Light, which is pretty equally distributed over the localities inhabited by freshwater fishes, cannot be considered as an important factor in their distribution, but it contributes towards constituting the impassable barrier between the surface and abyssal forms of marine fishes. Altitude has stamped the fishes of the various Alpine provinces of the globe with a certain character, and limited their distribution; but the number of these Alpine forms is comparatively small, ichthyic life being extinguished at great elevations even before the mean temperature equals that of the high latitudes of the Arctic region, in which some freshwater fishes flourish. On the other hand, the depths of the ocean, far exceeding the altitude of the highest mountains, still swarm with forms specially adapted for abyssal life. That other physical conditions of minor and local importance, under which fresh water fishes live, and by which their dispersal is regulated, are more complicated than similar ones of the ocean, is probable, though perhaps less so than is generally supposed: for the fact is that the former are more accessible to observation than the latter, and are, therefore, more generally and more readily comprehended and acknowledged. Thus, not only because many of the most characteristic forms of the marine and freshwater series are found, on taking a broader view of the subject, to be sufficiently distinct, but also because their distribution depends on causes different in their nature as well as the degree of their action, it will be necessary to treat of the two series separately. Whether the oceanic areas correspond in any way to the terrestrial will be seen in the sequel.
Fig. 103.—Ganoid scales of Tetragonolepis.
CHAPTER XVII.
THE DISTRIBUTION OF FRESHWATER FISHES.
Having shown above that numerous marine fishes enter fresh waters, and that some of them have permanently established themselves therein, we have to eliminate from the category of freshwater fishes all such adventitious elements. They are derived from forms, the distribution of which is regulated by other agencies, and which, therefore, would obscure the relations of the faunæ of terrestrial regions if they were included in them. They will be mentioned with greater propriety along with the fishes constituting the fauna of the brackish water.
True freshwater fishes are the following families and groups only:—
| Dipnoi | with | 4 | species. |
| Acipenseridæ and Polyodontidæ | „ | 26 | „ |
| Amiidæ | „ | 1 | „ |
| Polypteridæ. | „ | 2 | „ |
| Lepidosteidæ. | „ | 3 | „ |
| Percina | „ | 46 | „ |
| Grystina | „ | 11 | „ |
| Aphredoderidæ | „ | 1 | „ |
| Centrarchina | „ | 26 | „ |
| Dules | „ | 10 | „ |
| Nandidæ | „ | 7 | „ |
| Polycentridæ | „ | 3 | „ |
| Labyrinthici | „ | 30 | „ |
| Luciocephalidæ | „ | 1 | „ |
| Gastrosteus | „ | 10 | „ |
| Ophiocephalidæ | „ | 31 | „ |
| Mastacembelidaæ | „ | 13 | „ |
| Chromides | „ | 105 | „ |
| Comephoridæ | „ | 1 | „ |
| Gadopsidæ | „ | 1 | „ |
| Siluridæ | „ | 572 | „ |
| Characinidæ | „ | 261 | „ |
| Haplochitonidæ | „ | 3 | „ |
| Salmonidæ (3 genera excepted) | „ | 135 | „ |
| Percopsidæ | „ | 1 | „ |
| Galaxiidæ | „ | 15 | „ |
| Mormyridæ (and Gymnarchidæ) | „ | 52 | „ |
| Esocidæ | „ | 8 | „ |
| Umbridæ | „ | 2 | „ |
| Cyprinodontidæ | „ | 112 | „ |
| Heteropygii | „ | 2 | „ |
| Cyprinidæ | „ | 724 | „ |
| Kneriidæ | „ | 2 | „ |
| Hyodontidæ | „ | 1 | „ |
| Osteoglossidæ | „ | 5 | „ |
| Notopteridæ | „ | 5 | „ |
| Gymnotidæ | „ | 20 | „ |
| Symbranchidæ | „ | 5 | „ |
| Petromyzontidæ | „ | 12 | „ |
| Total | 2269 | species. |
As in every other class of animals, these freshwater genera and families vary greatly with regard to the extent of their geographical range; some extend over the greater half of the continental areas, whilst others are limited to one continent only, or even to a very small portion of it. As a general rule, a genus or family of freshwater fishes is regularly dispersed and most developed within a certain district, the species and individuals becoming scarcer towards the periphery as the type recedes more from its central home, some outposts being frequently pushed far beyond the outskirts of the area occupied by it. But there are not wanting those remarkable instances of closely allied forms occurring, almost isolated, at most distant points, without being connected by allied species in the intervening space; or of members of the same family, genus, or species inhabiting the opposite shores of an ocean, and separated by many degrees of abyssal depths. We mention of a multitude of such instances the following only:—
A. Species identical in distant continents—
1. A number of species inhabiting Europe and the temperate parts of eastern North America, as Perca fluviatilis, Gastrosteus pungitius, Lota vulgaris, Salmo solar, Esox lucius, Acipenser sturio, Acipenser maculosus, and several Petromyzonts.
2. Lates calcarifer is common in India as well as in Queensland.
3. Galaxias attenuatus inhabits Tasmania, New Zealand, the Falkland Islands, and the southernmost part of the South American continent.
4. Several Petromyzonts enter the fresh waters of Tasmania, South Australia, New Zealand, and Chili.
B. Genera identical in distant continents—
1. The genus Umbra, so peculiar a form as to be the type of a distinct family consisting of two most closely allied species only, one of which is found in the Atlantic States of North America, the other in the system of the Danube.
2. A very distinct genus of Sturgeons, Scaphirhynchus, consisting of two species only, one inhabiting fresh waters of Central Asia, the other the system of the Mississippi.
3. A second most peculiar genus of Sturgeons, Polyodon, consists likewise of two species only, one inhabiting the Mississippi, the other the Yang-tse-kiang.
4. Amiurus, a Siluroid, and Catostomus, a Cyprinoid genus, both well represented in North America, occur in a single species in temperate China.
5. Lepidosiren is represented by one species in tropical America, and by the second in tropical Africa (Protopterus).
6. Notopterus consists of three Indian and two West African species.
7. Mastacembelus and Ophiocephalus, genera characteristic of the Indian region, emerge severally by a single species in West and Central Africa.
8. Symbranchus has two Indian and one South American species.
9. Prototroctes, the singular antarctic analogue of Coregonus, consists of two species, one in the south of Australia the other in New Zealand.
10. Galaxias is equally represented in Southern Australia, New Zealand, and the southern parts of South America.
C. Families identical in distant continents—
1. The Labyrinthici, represented in Africa by 5, and in India by 25 species.
2. The Chromides, represented in Africa by 25, and in South America by 80 species.
3. The Characinidæ, represented in Africa by 35, and in South America by 226 species.
4. The Haplochitonidæ, represented in Southern Australia by one, in New Zealand by one, and in Patagonia by a third species.
This list could be much increased from the families of Siluridæ and Cyprinidæ, but as these have a greater range than the other Freshwater fishes, they do not illustrate with equal force the object for which the list has been composed.
The ways in which the dispersal of Freshwater fishes has been effected were various; they are probably all still in operation, but most work so slowly and imperceptibly as to escape direct observation; perhaps, they will be more conspicuous, after science and scientific inquiry shall have reached to a somewhat greater age. From the great number of freshwater forms which we see at this present day acclimatised in, gradually acclimatising themselves in, or periodically or sporadically migrating into, the sea, we must conclude that, under certain circumstances, salt water may cease to be an impassable barrier at some period of the existence of freshwater species, and that many of them have passed from one river through salt water into another. Secondly, the headwaters of some of the grandest rivers, the mouths of which are at opposite ends of the continents which they drain, are sometimes distant from each other a few miles only; the intervening space may have been easily bridged over for the passage of fishes by a slight geological change affecting the level of the watershed, or even by temporary floods; and a communication of this kind, if existing for a limited period only, would afford the ready means of an exchange of a number of species previously peculiar to one or the other of those river or lake systems. Some fishes, provided with gill-openings so narrow that the water moistening the gills cannot readily evaporate; and endowed, besides, with an extraordinary degree of vitality, like many Siluroids (Clarias, Callichthys), Eels, etc., are enabled to wander for some distance over land, and may thus reach a water-course leading them thousands of miles from their original home. Finally, fishes or their ova may be accidentally carried by waterspouts, by aquatic birds or insects, to considerable distances.
Freshwater fishes of the present fauna were already in existence when the great changes of the distribution of land and water took place in the tertiary epoch; and having stated that salt water is not an absolute barrier to the spreading of Freshwater fishes, we can now more easily account for those instances of singular disconnection of certain families or genera. It is not necessary to assume that there was a continuity of land stretching from the present coast of Africa to South America, or from South America to New Zealand and Australia, to explain the presence of identical forms at so distant localities; it suffices to assume that the distances were lessened by intervening archipelagoes, or that an oscillation has taken place in the level of the land area.
Dispersal of a type over several distant continental areas may be evidence of its great antiquity, but it does not prove that it is of greater antiquity than another limited to one region only. Geological evidence is the only proof of the antiquity of a type. Thus, although the Dipnoi occur on the continents of Africa, South America, and Australia, and their present distribution is evidently the consequence of their wide range in palæozoic and secondary epochs; the proof of their high antiquity can be found in their fossil remains only. For, though the Siluroids have a still greater range, their wide distribution is of comparatively recent date, as the few fossil remains that have been found belong to the tertiary epoch. The rapidity of dispersal of a type depends entirely on its facility to accommodate itself to a variety of physical conditions, and on the degree of vitality by which it is enabled to survive more or less sudden changes under unfavourable conditions; proof of this is afforded by the family of Siluroids, many of which can suspend for some time the energy of their respiratory functions, and readily survive a change of water.
To trace the geological sequence of the distribution of an ichthyic type, and to recognise the various laws which have governed, and are still governing its dispersal, is one of the ultimate tasks of Ichthyology. But the endeavour to establish by means of our present fragmentary geological knowledge the divisions of the fauna of the globe, leads us into a maze of conflicting evidence; or, as Mr. Wallace truly observes, “any attempt to exhibit the regions of former geological ages in combination with those of our own period must lead to confusion.” Nevertheless, as the different types of animals found at the present day within a particular area have made their appearance therein at distant periods, we should endeavour to decide as far as we can, in an account of the several zoo-geographical divisions, the following questions:—
1. Which of the fishes of an area should be considered to be the remnants of ancient types, probably spread over much larger areas in preceding epochs?
2. Which of them are to be considered to be autochthont species, that is, forms which came in the tertiary epoch or later into existence within the area to which they are still limited, or from which they have since spread?
3. Which are the forms which must be considered to be immigrants from some other region?
The mode of division of the earth’s surface into zoological regions or areas now generally adopted, is that proposed by Mr. Sclater, which recommends itself as most nearly agreeing with the geographical divisions. These regions are as follows:—
I. Palæogæa.
1. The Palæarctic region; including Europe, temperate Asia, and North Africa.
2. The Ethiopian region; including Africa, south of the Sahara, Madagascar, and the Mascarene Islands; also Southern Arabia.
3. The Indian region; including India south of the Himalayas, to Southern China, Borneo, and Java.
4. The Australian region; including Australia, the Pacific Islands, Celebes, and Lombock.
II. Neogæa.
5. The Nearctic region; including North America to Northern Mexico.
6. The Neotropical region; including South America, the West Indies, and Southern Mexico.
Comparatively few classes and orders of animals have been carefully studied with regard to their geographical distribution, but the majority of those which have been examined show that the difference of latitude is accompanied by a greater dissimilarity of indigenous species than that of longitude, and that a main division into an old world and new world fauna is untenable. More especially the Freshwater fishes, with which we are here solely concerned, have been spread in circumpolar zones, and in a but limited degree from north to south. No family, much less a genus, ranges from the north to the south, whilst a number of families and genera make the entire circuit, and some species more than half of the circuit round the globe within the zone to which they belong. Not even the Cyprinoids and Siluroids, which are most characteristic of the freshwater fauna of our period, are an exception to this. Temperature and climate, indeed, are the principal factors by which the character of the freshwater fauna is determined; they form the barriers which interfere with the unlimited dispersal of an ichthyic type, much more than mountain ranges, deserts, or oceans. Hence the tropical zone is an impassable barrier to the northern Freshwater fish in its progress towards the south; where a similarly temperate climate obtains in the southern hemisphere, fish-forms appear analogous to those of the north, but genetically and structurally distinct.
The similarity which obtains in fishes at somewhat distant points of the same degree of longitude, rarely extends far, and is due to the natural tendency of every animal to spread as far as physical conditions will permit. Between two regions situated north and south of each other there is always a debateable border ground, in parts of which sometimes the fishes of the one, sometimes those of the other, predominate, and which is, in fact, a band of demarcation. Within this band the regions overlap each other; therefore, their border lines are rarely identical, and should be determined by the northern and southernmost extent of the most characteristic types of each region. Thus, for instance, in China, a broad band intervenes between temperate and tropical Asia, in which these two faunæ mix, and the actual northern border line of the tropical fauna is north of the southern border line of temperate Asia.
It is the aim of every philosophical classification to indicate the degree of affinity which obtains between the various divisions; but the mode of division into six equivalent regions, as given above, does not fulfil this aim with regard to Freshwater fishes, the distribution of which allows of further generalisation and subdivision. The two families, Cyprinidæ and Siluridæ, of which the former yields a contingent of one-third, and the latter of one-fourth of all the freshwater species known of our period, afford most valuable guidance for the valuation of the degrees of affinity between the various divisions. The Cyprinoids may be assumed to have taken their origin in the Alpine region, dividing the temperate and tropical parts of Asia; endowed with a greater capability of acclimatising themselves in a temperate as well as tropical climate than any other family of freshwater fishes, they spread north and south as well as east and west; in the preglacial epoch they reached North America, but they have not had time to penetrate into South America, Australia, or the islands of the Pacific. The Siluroids, principally fishes of the sluggish waters of the plains, and well adapted for surviving changes of the water in which they live, for living in mud or sea-water, flourish most in the tropical climate, in which this type evidently had its origin. They came into existence after the Cyprinoids, fossil remains being known only from tertiary deposits in India, none from Europe. They rapidly spread over the areas of land within the tropical zone, reaching northern Australia from India, and one species even immigrated into the Sandwich Islands, probably from South America. The Coral Islands of the Pacific still remain untenanted by them. Their progress into temperate regions was evidently slow, only very few species penetrating into the temperate parts of Asia and Europe; and the North American species, although more numerous, showing no great variety of structure, all belonging to the same group (Amiurina). Towards the south their progress was still slower, Tasmania, New Zealand, and Patagonia being without representatives, whilst the streams of the Andes of Chili are inhabited by a few dwarfed forms identical with such as are characteristic of similar localities in the more northern and warmer parts of the South American continent.
After these preliminary remarks we propose the following division of the fauna of Freshwater fishes:—
I. The Northern Zone.—Characterised by Acipenseridæ. Few Siluridæ. Numerous Cyprinidæ. Salmonidæ, Esocidæ.
1. Europo-Asiatic or Palæarctic Region.—Characterised by absence of osseous Ganoidei; Cobitidæ and Barbus numerous.
2. North American Region.—Characterised by osseous Ganoidei, Amiurina, and Catostomina; but no Cobitidæ or Barbus.
II. The Equatorial Zone.—Characterised by the development of Siluridæ.
A. Cyprinoid Division.—Characterised by presence of Cyprinidæ and Labyrinthici.
1. Indian Region.—Characterised by [absence of Dipnoi[18]] Ophiocephalidæ, Mastacembelidæ. Cobitidæ numerous.
2. African Region.—Characterised by presence of Dipnoi and Polypteridæ. Chromides and Characinidæ numerous. Mormyridæ. Cobitidæ absent.
B. Acyprinoid Division.—Characterised by absence of Cyprinidæ and Labyrinthici.
1. Tropical American Region.—Characterised by presence of Dipnoi. Chromides and Characinidæ numerous. Gymnotidæ.
2. Tropical Pacific Region.—Characterised by presence of Dipnoi. Chromides and Characinidæ absent.
III. The Southern Zone.—Characterised by absence of Cyprinidæ, and scarcity of Siluridæ. Haplochitonidæ and Galaxiidæ represent the Salmonoids and Esoces of the Northern zone. One region only.
1. Antarctic Region.—Characterised by the small number of species; the fishes of—
a. The Tasmanian sub-region; b. The New Zealand sub-region; c. The Patagonian sub-region;
being almost identical.[19]
In the following detailed account we begin with a description of the equatorial zone, this being the one from which the two principal families of freshwater fishes seem to have spread.