Chapter VIII: Introduction: General Characters—anatomy of Amphioxus—embryology and (3)

Pelagic Fishes, like the Herring, Mackerel, Flying-Fish (_Exocoetus_), and many others, often have the belly and sides silvery or white, and the back dark green, black, or steely blue. Seen from below against the light sky, or viewed from above against the background of the dark water, these Fishes would seem to be practically invisible to their predatory foes, whether Fishes or Birds, or at all events not easily detected.

Coloration may not only be protective, but also aggressive, by helping to conceal the proximity of an animal from its prey; add to this some device for deceiving and attracting the prey, and we have an example of "alluring" coloration.[143]

As an example of coloration which is both aggressive and alluring, the Angler-Fish or Fishing-Frog (_Lophius piscatorius_) of our own coasts may be quoted. Naturally sluggish and inactive in its habits, and often using its muscular pectoral fins for crawling about the sea-bottom, the Angler-Fish usually hides itself in the sand or amongst sea-weeds, which it closely resembles in general colour. Curious branched tag-like processes of soft skin fringe the sides of the head and body, and in appearance and colour resemble the smaller fronds of the surrounding sea-weed. So far the coloration is simply aggressive, and helps to conceal the Fish from its prey, but in addition the animal is provided with a special device for luring its prey within the reach of its capacious and Frog-like mouth. The first three spines {174}of the dorsal fin are detached from one another and greatly elongated, and moreover extend along the middle of the dorsal surface of the head. The first, which is the longest, terminates in lobes or lappets of skin, and can be freely moved in every direction by the muscles inserted into its base. By the agitation of this lure or bait smaller Fishes, probably mistaking the disturbance for the presence of a wriggling worm, are tempted to their fate, and soon find themselves engulfed in the enormous mouth of the artful angler.[144] In some allied forms (e.g. _Ceratias bispinosus_ and _Oneirodes eschrichtii_)[145] which live in the abyssal darkness of the deep sea, use is made of the attraction which light has to aquatic animals, and the fishing-rod spine terminates in a phosphorescent organ, which is probably used for enticing smaller Fishes within the reach of the jaws of these singularly modified Angler-Fishes.[146]

It is by no means improbable that examples of "warning" coloration occur amongst Fishes. The brilliant colours of some of the Trigger-Fishes (_Balistes_, _Monacanthus_), Coffer-Fishes (_Ostracion_), and Globe-Fishes (_Tetrodon_) are perhaps of this nature. They are often associated with the presence of strong spines, defensive and often erectile, either in connexion with the dorsal fin or on the general surface of the body, and may therefore serve the purpose of a danger signal to such predatory foes of these Fishes as might otherwise be tempted to attack them—to the mutual advantage of the Fishes themselves and their would-be enemies. The British Weever-Fish (_Trachinus_) may perhaps offer another example of warning coloration.[147] The Fish is armed with poisonous spines on its opercula, and, in addition, has a conspicuous black dorsal fin. When the body of the Fish is buried in the sand, only its projecting dorsal fin remains to indicate its whereabouts to predatory Gurnards, which might otherwise mistake the Weever for harmless Fishes of similar size and habits. The existence of "recognition" colours or markings peculiar to the species, to enable individuals of the same species to recognise one another and to keep together in shoals, has not yet been proved. It is probable that the relatively {175}limited range of vision, even in the clearest water, would render coloration unsuitable for this purpose. Recognition sounds are likely to be far more effective, and there is evidence of their production by a special vocal mechanism in many Fishes.[148]

The examples given above show how natural selection may lead to the evolution of distinctive forms of coloration which are advantageous to the Fish either for concealment, aggression, or protection, and in conclusion it may be pointed out that by the same cause colour may be eliminated or its development checked if in any way harmful to the animal; and further, that if a particular coloration becomes useless to the Fish by reason of a change in its habits or environment, natural selection ceasing to act where its intervention is no longer necessary to maintain the coloration, the latter will sooner or later tend to disappear.

The absence of pigment is sometimes protective. The surface-swimming larvae of many Teleosts have no chromatophores, and therefore no obvious pigmentary colours. Their bodies are so translucent that they can be seen through, and hence are visible only with difficulty. The transparency of the body may even be increased by the absence of the red haemoglobin of the blood, as is the case with the pelagic _Leptocephalus_-larvae of the Eel.[149] The iridocytes of the reflecting tissue may also disappear under the influence of changed surroundings. The larvae of various species of _Onus_ (Gadidae) are silvery in hue during their pelagic career, owing to the presence of iridocytes in the skin, but on becoming mature they change to a dull dark colour, and live under stones or in holes and crevices in the rocks. During the change of habit the reflecting tissue (argenteum) is lost, and the needful chromatophores are acquired.[150]

Instances of the loss of pigmentary colours, owing to the cessation of the controlling influence of natural selection, are to be found in the absence of chromatophores on the white under surface of the Flat-Fishes, where such colours are useless but not necessarily harmful, and in the colourless, cave-inhabiting Fishes, of which the Blind-Fish (_Amblyopsis_) of North America may be taken as an example.

{176}POISON GLANDS OF FISHES.

A few Teleosts are provided with weapons of offence or defence in the shape of poison-glands, probably derived from the epidermis, and associated with spines on the gill-covers, or in connexion with the dorsal fin, or with both.

The two British species of "Weever" (_Trachinus draco_ and _T. vipera_) are both provided with poison-organs in connexion with a spine on the operculum and with the five or six spiny rays of the anterior dorsal fin.[151] The first of these spines is a structure projecting backwards from the hinder margin of the opercular bone of the gill-cover, and is traversed along both its upper and lower margins, from base to point, by a deep groove. Except at its protruding naked point the spine is ensheathed in an extension of the external skin. Along each of the grooves there extends a solid pear-shaped mass of gland-cells, the broad base of which coincides with the base of the spine, while the gradually tapering, narrower portion is continued as far as the sharp point. The glands enclose no cavity, and there is no duct, so that whatever poisonous fluid their cells secrete is probably set free by the rupture of the cells and discharged into the grooves, along which it passes to the point of the spine, somewhat after the fashion of a hypodermic syringe. The origin of the gland-cells from an in-pushing of the epidermis is indicated by the continuity of the two structures near the point of the spine. Both in structure and in their relation to poison-glands each of the spines of the {177}dorsal fin is almost precisely similar to the opercular spine. There is no evidence as to how the poison is ejected into a wound, and it can only be conjectured that it may be caused by the pressure exerted on the gland when the spine is forcibly thrust for some distance into the flesh. Certain it is that these structures are capable of inflicting painful and troublesome wounds when the Fish is incautiously handled and the skin accidentally punctured, and no doubt they can be used with great effect as offensive organs.

A similar poison apparatus exists in certain species of Batrachidae, such as _Thalassophryne reticulata_,[152] which is by no means uncommon at Panama. This apparatus is formed by a spinous outgrowth from the opercular bone and by the first two dorsal spines. Instead, however, of having two grooves, the opercular spine resembles the fang of a venomous snake, and is perforated by a complete canal which is only open at the base and point of the spine. A poison-sac at the base of the spine discharges its contents into the canal. The nature of the glands which secrete the poison has yet to be discovered, but it is probable either that there are glands in connexion with the poison-sac, or that the latter is lined by a glandular epithelium. The structure of the dorsal spines is similar. In some species of the Scorpaenoid genus _Synancia_[153] (e.g. _S. verrucosa_, from the Indian Ocean), the terminal portions of the dorsal spines are deeply grooved on each side, and at the origin of each groove there is a pear-shaped bag containing a milky poison. The bag is prolonged into a duct which, after traversing the groove, opens at the extremity of the spine.

Many Siluridae are armed with powerful and often serrated dorsal and pectoral spines which are certainly capable of inflicting dangerous wounds, and not a few of them possess a sac-like organ with an external opening in the axilla of the pectoral fin. It is possible that the sac secretes a poison for anointing the spine, but at present there is no evidence that such is the case, or that the sac produces any poisonous secretion at all.[154]

Among the Elasmobranchs the Eagle-Rays (_Aëtobatis_),[155] and the Sting-Rays (_Trygon_), have barbed or serrated spines on the tail, which inflict wounds far more severe than those caused by {178}mere mechanical laceration; but, except the mucus secreted by the gland cells of the skin, which may possess venomous properties, no special poison-forming glands in connexion with the spines are at present known.

PHOSPHORESCENT ORGANS.[156]

In common with many other animals of similar habitat, phosphorescent organs (photophores) are highly characteristic structures in many deep-sea Teleosts belonging to widely different families (e.g. _Stomiatidae_, _Scopelidae_, _Halosauridae_, and _Anomalopidae_). These organs probably had their origin in local aggregations of the gland cells of the epidermis, which had acquired the power of secreting a luminous slime. Luminous organs vary greatly in number and in their mode of distribution in the skin. Usually they are found on the sides and ventral surface of the body and head, very rarely on the dorsal surface, and they often present the appearance of brightly glistening jewels set in the skin. A very frequent method of arrangement is in one or two longitudinal lines along the lateral and ventral surfaces, sometimes extending continuously from the head to the end of the tail (Fig. 371, A, and Fig. 379), but occasionally interrupted and limited to portions of the body and tail; and in a few a distinctly metameric disposition is obvious. On the other hand, the very numerous and simple organs of _Opostomias_ are disposed in many transverse bands along the sides of the Fish. In addition to these organs, which are usually numerous, and whose arrangement is linear, specially large and often structurally complex luminous organs are present on different parts of the head and body. In _Opostomias micripnus_ there is a phosphorescent organ on a median barbel depending from the chin. _Sternoptyx diaphana_ has one on the lower jaw. The presence of one or two organs beneath the eyes (Fig. 96) is characteristic of several species (e.g. _Opostomias micripnus_, _Astronesthes niger_, _Pachystomias microdon_, _Scopelus benoitii_, _Malacosteus indicus_). _Opostomias micripnus_ has a luminous organ on the isolated and elongated first fin ray of the pectoral fin, while in certain deep-sea Angler-Fishes (e.g. _Ceratias_) there is one on the anterior cephalic fin-ray of the dorsal fin. The Scopelid _Ipnops {179}murrayi_[157] (Fig. 371, C) has a singular organ, probably luminous, beneath the transparent superficial bones of each side of the roof of the skull. Another member of the same family (_Scopelus benoitii_) is interesting in having a phosphorescent organ in the middle of the back, which is directed backwards. An American genus of Batrachidae (_Porichthys_) has about 350 photophores in relation with the lateral sense-organs of each side of the head and body.[158] The existence of luminous organs has also been noticed in the Haddock (_Gadidae_).[159] A primitive form of photophore, distributed in considerable numbers on the head and trunk, either in lines or diffused over the surface, exists in eleven species of Selachii (_Spinacidae_), of which some are known to be luminous.[160]

Diversity of structure is equally marked. The essential part of each luminous organ is always a collection of gland cells, usually disposed so as to form the lining of a series of radially arranged gland-tubules in the deeper part of the organ, which also contains ganglion cells, and is supplied with nerves from contiguous spinal or cranial nerves. The simplest form of phosphorescent organ consists of little more than these essential elements. In the more complex organs an investing pigment-sheath, reflecting and lens-like structures, and an iris diaphragm, either singly or in combination, may be added. Fig. 97 represents one of the simplest types of phosphorescent organ, which, in groups of {180}50 to 100, are arranged in transverse bands on the sides of _Opostomias micripnus_, and appear as small white spots on the otherwise black skin of this Fish.

Each organ has the shape of a biconvex lens, sunk to about half its thickness in the skin. The inner half is formed of radially-arranged gland tubes filled with small granular cells, and converging towards the centre of the organ. Into the connective-tissue walls of the tubes extend blood-vessels and nerves. External to the gland tubes there is a layer of long slender cells arranged perpendicularly to the surface, and more externally still a layer of ganglion cells. There is evidence that these organs multiply by division. Such simple phosphorescent organs as these differ little from the groups of epidermic gland cells, which probably formed the evolutionary starting-point in the development of these singular structures.

A much more complex type of luminous organ is to be found in the suborbital organs of _Pachystomias microdon_, of which there are two on each side, appearing as conspicuous white masses, one in front of the other, and situated just below the eye. The more anterior of the two organs is somewhat pouch-shaped in section, its walls consisting of several concentric layers (Fig. 98). Externally there is a layer of black pigment, within which is a stratum of irregular gland tubes. More internally still there is a thick layer of light-reflecting spicules, probably derived from an inverted and modified dermal scale. The axial part of the organ is occupied by a number of radial-disposed structures, probably similar to the gland tubes of the simple organs of _Opostomias_, and continuous with a lens-like structure which, as it were, closes the expanded mouth of the pouch. The superficial skin which forms the margin of the aperture partially projects over the outer surface {181}of the lens-like body, somewhat after the fashion of an iris-diaphragm. The organ is supplied by a branch of the fifth cranial nerve. Between such simple and complex organs as those above described there are various other types which are more or less intermediate in character.

A particular type of phosphorescent organ is not necessarily restricted to the same species; both the simplest and one or more of the more complex types may be represented in the same Fish. Thus, _Opostomias micripnus_, which frequents depths of over 2000 fathoms, has not only the simple organs described above, but also others differing from the former in having an external pigmentary sheath, which are scattered all over the body at intervals of 1 to 3 mm. There are also larger and still more complex organs which are disposed in two parallel rows along each side of the body; and finally, the same species has special luminous organs on a median chin-barbel, and also on an elongated fin-ray pertaining to the pectoral fin.

The light emitted by phosphorescent organs is probably of use to deep-sea Fishes in enabling them to seek and detect their prey in the sunless depths which they frequent. The position of the organs on the sides and ventral surface of the body, and the frequent presence of special luminous organs in the vicinity of the mouth, render them admirably adapted to light up the water in front of and beneath the Fish, while the existence of optical accessories for intensifying the luminous beams, and for regulating their distribution, combined with an abundant nervous supply, suggests that the emission of light is under the control of the Fish, and may be varied as the occasion requires. That these organs may also be defensive, in some instances at all events, seems not improbable. A flash-light from the dorsal luminous organ or "stern-chaser" of _Scopelus benoitii_ would probably dazzle and frighten an enemy in hot pursuit of the _Scopelus_. The use of phosphorescent organs as baits or lures for enticing prey has already been alluded to. There is some evidence that the colour of the emitted light differs in different Fishes; and as there is considerable variety in the precise disposition of the organs, it seems probable that in deep-sea Fishes recognition lights may take the place of the recognition colours and sounds of those whose lot is cast in a sunnier habitat.

{182}CHAPTER VII

THE SKIN AND SCALES

The skin of the Cyclostomata and Fishes consists (1) of the epidermis, formed of several layers of epidermic cells, which are constantly being recruited by the division of the cells of the basal layer; and (2) of a stratum of connective tissue with intermingled unstriped muscle-fibres, blood-vessels and nerves, which constitutes the deeper layer or dermis. From the epidermis are formed the various unicellular or multicellular glands with which the skin is provided; and from one or both of the skin layers originate the different calcareous structures which constitute the hard exoskeleton.

In the Cyclostomata the epidermis is particularly rich in goblet-shaped, mucus-secreting, gland-cells. The Myxinoids also possess numerous pockets of so-called "thread-cells." In each of these cells the protoplasm secretes a long spirally-coiled thread, and under the influence of appropriate stimuli the thread is shot out and unwound to a great length. The threads and the mucus are so abundant that one of these animals will convert a bucket of water into a thick mass of jelly. No scales or other hard exoskeletal structures are present in any of the Cyclostomata.

In Fishes mucus-glands are also abundant in the epidermis, and to their activity is due the slimy mucus which lubricates the surface of the body. They are specially numerous in the Dipnoi (e.g. _Protopterus_), where, in addition, there are many simple multicellular glands which secrete the "cocoon" or capsule in which the Fish is enclosed during the dry season. From the epidermis are derived the poison-glands of some Teleosts, and also the "glandula pterygopodia" in relation with the claspers of the male Elasmobranchs. The glandular structures in connexion with {183}the phosphorescent organs of the deep-sea Fishes will no doubt be traced to the same source.

In the great majority of Fishes the skin becomes the seat of calcareous deposit, and gives rise to such diverse exoskeletal structures as the varied forms of spines and scales with which the surface of a Fish is invested.[161] These structures, probably the most ancient form of Vertebrate skeleton owing its existence to the presence of lime salts in the tissues of the body, present highly characteristic modifications in the different groups.

Exoskeletal structures are of two kinds: (1) those which owe their formation to the secretory activity of cells belonging both to the epidermis and the dermis, and (2) those which are derived solely from the dermis. To the first belong the dermal denticles or so-called placoid scales of most Elasmobranchs, and to the second the scales which form the skin-skeleton of living and extinct Teleostomi and Dipnoi. With the exception of enamel, which is always formed by the cells of the epidermis, the hard exoskeletal tissues owe their existence to the secretion of certain cells of the dermis (scleroblasts),[162] the inclusion of which in a growing calcifying tissue is the cause of whatever cellular structure the tissue may present. It will shortly be apparent that the dermic scleroblasts are by no means uniform in their products, and that in different Fishes they give rise to widely different hard tissues.

The dermal denticles or "shagreen" of the ordinary Sharks and Dog-Fishes (Elasmobranchii) probably represent the most primitive form of exoskeleton. In the development of a dermal denticle a papilla of the dermis grows up into the overlying epidermis, pushing before it the basal layer of epidermic cells, which forms an investment to the papilla and constitutes the so-called "enamel organ" (Fig. 100). The papilla itself subsequently becomes converted into dentine, leaving, however, a central pulp-cavity, while the apex of the papilla is invested by a cap of enamel formed by the enamel organ. Ultimately the base of the papilla widens out into a more or less rhomboidal basal plate formed of bone. In this way there is formed a {184}pointed, enamel-tipped spine of dentine which protrudes through the epidermis, and projects backwards on the surface of the body, but is firmly fixed in the skin by the basal plate with which it is continuous. The centre of the under surface of the basal plate is perforated for the entrance of the blood-vessels which pass to the cellular pulp in the axis of the spine. In the adult Fish the denticles form a fairly close-set covering to the whole body, including the head and even the surfaces of the fins, and are larger on the dorsal than on the ventral surface (Fig. 99). In the Rays (_Raia_) they are more sparsely scattered, and in different parts of the body may form spines of considerable size for offensive or defensive purposes. The spines vary greatly in shape in different members of the group, sometimes being acutely pointed, and sometimes flattened or depressed, and often they are furnished with smaller accessory spines developed at their bases or from the surface of the basal plate. An arrangement of the denticles in oblique transverse rows is observable in some genera (e.g. _Scyllium_). In the Saw-Fishes (e.g. _Pristis_) the denticles which fringe the lateral margins of the long flattened rostrum are not only enormously enlarged, but are implanted in sockets and form the teeth of the saw (Fig. 262). In the Holocephali the smooth skin is almost entirely devoid of exoskeletal structures, but dermal denticles are present on the frontal and anterior claspers, and in the young there may be a double row of small denticles along the back.

{185}

In the remaining groups of Fishes, the Teleostomi and the Dipnoi, the spine of the primitive dermal denticle is either evanescent or entirely wanting, while the equivalent of the basal plate remains to form the unit of a scaly armature. Evidence of this may be found in the presence of transitory evanescent spines, provided with an enamel-cap, secreted by the basal epidermis, on the developing rhomboidal scales, as in the young _Lepidosteus_[163] (Fig. 101); while the entrance of blood-vessels into the scales through perforations on their inner surfaces, as in _Polypterus_ and _Lepidosteus_, obviously recalls the perforated base of a dermal denticle (Fig. 99). The epidermis now ceases to take any part in the formation of the scales, and hence enamel no longer enters into their structure. A more regular and definite arrangement of the scales is noticeable, and whether distinct, or articulating with one another, or overlapping like the slates on the roof of a house, they are usually disposed in a series of successive oblique transverse rows. In some of these Fishes the embryonic epidermic covering of the scales becomes lost, and their outer surfaces are naked. More frequently, as in the generality of Teleosts, and in the Dipnoi, the reverse is the case, and the scales are more or less completely invested both by the dermis and the epidermis. As regards their shape, size, and minute structure there is much variation. In some Teleostomi {186}the primitive rhomboidal shape of the dermal denticle is retained; in others a rounded or cycloid scale supplants the earlier rhombic type. Within the limits of the same group (_e.g._ Crossopterygii) there are examples of the independent evolution of a cycloid from a pre-existing rhombic squamation; and with the introduction of the cycloid type an overlapping or imbricated disposition of the scales always takes the place of the marginal articulation of the rhombic type.

As to the causes which may have determined the shape and mutual relations of scales interesting suggestions have been made.[164] Scales bear a segmental relation to the subjacent muscle-segments or myotomes, sometimes being disposed in oblique transverse rows coinciding with the latter, or the rows may be so far increased as to be multiples of the myotomes. From mechanical considerations depending on the sigmoid shape and interdigitating relations of the myotomes and their separating fibrous septa or myocommata, and the attachment of the myocommata to the dermis, the contraction of the myotomes during the lateral flexions of the trunk in swimming has a tendency to wrinkle the skin into definitely circumscribed rhombic areas, thus determining the shape, limits, and disposition of the scales which are developed in those areas. The rhombic was probably the primitive shape of scales, and is certainly characteristic of the palaeontologically older types of scaly Fishes. Generally the rhombic condition is associated with a peg-and-socket articulation between the upper and lower margins of adjacent scales. But a rhombic squamation is not without {187}disadvantages, and would certainly impose some restriction on the lateral flexures of the body in swimming, and hence in the different groups of Fishes it may happen that, in the more specialised forms, an imbricated cycloid squamation supersedes a rhombic condition, and with the change the Fish acquires greater lateral mobility. Even in the same Fish the gradual substitution of the cycloid for the rhombic type may be observed. In the Australian _Aetheolepis_,[165] a fossil genus related to the European Liassic _Dapedius_, there is a gradual transition along the sides of the body between the articulated rhombic scales of the relatively immobile trunk and the cycloid overlapping scales of the flexible tail; and it may be mentioned that, even where a typical rhombic squamation exists, the peg-and-socket articulation may be wanting in the caudal region, so as to ensure greater freedom of movement. Mechanical considerations may also explain the overlapping of cycloid scales. From the mode of attachment of the myocommata to the dermis, the contractions of the myotomes, through the pull which they exert on the former, tend to deflect or depress the scale-areas, particularly at their anterior margins.

In the surviving Crossopterygii, as in _Polypterus_, the scales are rhomboidal and thick, and they only slightly overlap. They articulate with one another by means of marginal peg-and-socket articulations (Fig. 106, B). A thick layer of hard, glistening, enamel-like substance or "ganoin" forms the outer layer of the scale; the inner layer consisting of bone in which dentinal tubules as well as bone-cells are present. In the numerous fossil members of the group the scales are either rhomboidal or cycloid.

{188}The oldest representatives of the Chondrostei, the Palaeoniscidae (Fig. 283) possessed a complete armature of rhombic scales, but in all the surviving members of the group the scales have undergone considerable modification in some respects, and in others are degenerate. In the Sturgeon (_Acipenser_)[166] the primitive rhombic squamation is retained only on the sides of the terminal part of the tail, and there they are in close apposition in oblique rows. The rest of the body is traversed by five widely-separated longitudinal rows of large bony scutes, which, like the rhombic scales, are furnished with ridges and projecting spines (Fig. 102). Between the rows of large scales there are numerous denticle-like structures arranged in oblique rows. Each of these consists of a basal plate imbedded in the dermis, and of one or more projecting spines which perforate the epidermis. All the scales have the same minute structure, consisting mainly of bone; but the surface layer and the spines seem to be composed of a hard laminated substance from which bone-cells are absent (ganoin). In _Polyodon_ the scutes are wanting, but vestigial denticles are retained.

Among the Holostei the scales are very different in the two surviving members of the group. In _Lepidosteus_ (Fig. 103) the {189}scales are rhombic, and both in arrangement and structure, as well as in their method of articulating with one another, they closely resemble those of _Polypterus_. In _Amia_,[167] on the contrary, the relatively thin scales are cycloid in shape, and in their imbricated arrangement, in their enclosure in pouches of the dermis, and in the absence of any superficial covering of ganoin, they are very similar to the scales of the more typical Teleosts (Fig. 295). The resemblance extends even to histological structure, for each scale consists of an outer layer of bone, which gradually passes into an inner fibrous stratum.

In Teleosts the usually thin and flexible scales are primarily developed from dermic papillae, but subsequently they come to lie in pockets or pouches in the dermis. As a rule no spines are developed, and so far no trace of an enamel organ has been detected during their development. The scales in their dermal pouches are disposed obliquely to the surface of the body, so that the hinder part of one scale overlaps the anterior portion of the scale next behind it (Fig. 104). Only the free hinder part of each scale has an epidermic investment (Fig. 105). In minute structure each scale consists of an outer layer of bone, which, like the bone of the endoskeleton, may either be homogeneous except for a feeble lamination, or it may contain bone-cells arranged in successive layers, parallel to the surface of the scale. In addition, there is an inner fibrous stratum in which the fibrous bundles in any one plane cross those in planes above or below them. The scales are either "cycloid," that is, they have smooth, unbroken margins (Fig. 105), or the free margin of each scale is produced into a series of tooth-like spines, and the scale is said to be "ctenoid" (Fig. 106, A).

Some Teleosts, however, have scales which are neither cycloid nor ctenoid, and in certain features seem to be intermediate between ordinary Teleostean scales and dermal denticles. Thus, on certain parts of the body of _Centriscus_,[168] each scale consists {190}of a rhombic basal plate, produced into a curved, backwardly-inclined spine, the axis of which contains a pulp-cavity opening on the inner surface of the basal plate (Fig. 107). Some Malthidae (e.g. _Malthe_[169]) have similar scales, but with round basal plates and solid spines (Fig. 108, B). Similar scales (Fig. 109), sometimes rhombic in shape, with one or more spines, which may be simple or branched, are also found in the Sclerodermi (e.g. _Balistes_, _Monacanthus_, _Triacanthus_).[170]

FIG. 106.—A, Ctenoid scale; B, "Ganoid" scale. (After Günther; from Parker and Haswell.)]

Amongst some of the usually scaleless Siluroid Fishes the scales assume a very peculiar structure. In _Hypostoma_[171] (_Plecostomus_) the sides and back of the Fish are covered by large bony plates, but on the under surface and on the fins these are replaced by much smaller ones. Both kinds, however, carry numerous small movable spines implanted in sockets (Fig. 110), a fact which suggests comparison with a stage in the development of the scales of _Lepidosteus_, when the independently formed and evanescent spines have not yet fused with the basal plates.

In other Teleosts, as in the Agonidae and some Triglidae, the body is completely cuirassed with large keeled bony plates. The singular appearance of many of the Plectognathi is largely {191}the result of the curious modifications which their scales undergo. In some of the Coffer-Fishes (_Ostracion_) these structures assume the form of polygonal bony plates which suturally articulate with one another and enclose the trunk in a rigid cuirass, from which the scaleless tail protrudes behind (Fig. 438); while in some Globe-Fishes and Porcupine-Fishes (e.g. _Tetrodon_, _Diodon_) the prolongation of the scales into strong erectile spines equally well serves the purpose of protection (Fig. 439).

Most Teleostomi have the scales along the "lateral line" perforated by single or multiple apertures, through which the sensory canal communicates with the exterior.

In a few Teleosts scales are entirely absent, as in most Siluridae; or they exist only as microscopic vestiges hidden in the skin, as in Eels; or, as in such naked forms as _Antennarius marmoratus_ and _Lepadogaster_, and in some Siluridae, they become reduced to mere papillae of the dermis.

The concentric rings observable (Fig. 105) on the surface of many Teleostean scales are an index to the age of the {192}Fish.[172] The formation of these rings depends on the fact that the lines of growth on the surface of the scale are more widely separated from one another on that portion of the scale formed during summer, and relatively closer together on that part which is formed during the winter; the more rapid growth in the warmer season probably being due to favourable conditions as to food and temperature, and the retarded growth of the colder season to the reverse. Hence, by counting the alternating zones of close-set winter lines and less closely approximated summer lines of growth, a reliable clue may be gained as to the age of the Fish.

In the Dipnoi,[173] as in Teleosts, the scales are enclosed in dermal pockets, and exhibit a regular, imbricated disposition in oblique rows (Fig. 304, A). In shape they are nearly cycloid, or slightly oval, with the long axis coinciding with that of the body. Structurally, also, they bear some resemblance to Teleostean scales, although differing in details. On the outer surface of the scales there are numerous small conical spines. No significance, other than as an example of evolutionary convergence, can be attached to the resemblance between the scales of Fishes so widely separated as the Dipnoi and the Teleosts.

All known fossil Dipnoi had scales of a similar character, although differing greatly in size in different genera. In some (e.g. _Dipterus_) a layer of enamel-like substance invests the exposed portions of the scales.

{193}CHAPTER VIII

THE SKELETON

All Fishes possess an internal skeleton which, in order that it may be distinguished from the more superficial scaly exoskeleton described in