The epidermis of the young larvae of Amphibia is furnished with cilia, which later on are suppressed by the development of a thin hyaline layer or cuticula, but clusters of such cilia remain, at least during the larval life and during the periodical aquatic life of the adult, in the epidermal sense-organs. In the frog, currents are set up by the ciliary action at an earlier stage, and are maintained to a later stage than in the newt. In the latter the tail loses its ciliation, whereas in the frog it remains active almost up to the time of the metamorphosis. In tadpoles of 3-10 mm. nearly the whole surface is ciliated (Assheton).[[14]] The cilia work from head to tail, causing the little animal, when perfectly quiet, to move forwards slowly in the water. Beneath the cuticula, in the Perennibranchiata and the larvae of the other Urodela, lies a somewhat thicker layer of vertically striated cells, the so-called pseudo-cuticula, which disappears with the transformation of the upper layers of the Malpighian cells into the stratum corneum. The latter is very thin, consists of one or two layers of flattened cells, and is shed periodically by all Amphibia in one piece. In the Urodela it generally breaks loose around the mouth, and the animal slips out of the delicate, transparent, colourless "shirt," which during this process of ecdysis or moulting becomes inverted. In the Anura it mostly breaks along the middle line of the back, the creature struggles out of it, pokes it into its mouth, and swallows it. Urodela also eat this skin. As a rule the first ecdysis takes place towards the end of the metamorphosis, preparatory to terrestrial life. So long as the animal grows rapidly, the skin has to be shed frequently, since this corneous layer is practically dead and unyielding. Adult terrestrial Urodela do not seem to moult often, mostly only when they take to the water in the breeding season. Anura, on the other hand, moult often on land, at least every few months. The surface of the new skin is then quite moist and slimy, but it soon dries and hardens.
The Malpighian stratum consists of several layers, thickest in the Perennibranchiata; in them it contains mucous cells throughout life, in others such slime-cells are restricted to larval life. Later, regular slime-glands are developed, which open on the surface. They are very numerous, and more evenly distributed, over most parts of the body, than the specific or poison-glands, which are restricted to certain parts, often forming large clusters, especially on the sides of the body. They reach their greatest development in the "parotoid glands" of the Anura. Both kinds of glands are furnished with smooth muscle-fibres, which are said to arise from the basal membrane underlying and forming part of the Malpighian layer; these muscle-cells extend later downwards into the corium. For the action of the poison, see p. [37].
The stratum corneum is mostly thin, but on many parts of the body, especially in Anura, the epidermal cells proliferate and form hard spikes or other rugosities, generally stained dark brown. With these may be grouped the nuptial excrescences so frequent in the Anura, especially on the rudiment of the thumb, and on the under surface of the joints of the fingers and toes. In many Anura, less frequently in the Urodela, the tips of the fingers and toes are encased in thicker horny sheaths, producing claws or nails. They are best developed among newts in Onychodactylus, among the Anura in Xenopus and Hymenochirus. The horny covering of the metatarsal tubercles reaches its greatest size in the digging spur or spade of Pelobates. In most of these cases the cutis is elevated into more or less wart-like papillae, covered, of course, by the proliferated and cornified epidermis. In the female of Rana temporaria nearly the whole surface of the body becomes covered with rosy papillae during the breeding season. Similar nuptial excrescences are common, and are most noteworthy in the male of the Indian Rana liebigi.
The epidermis also contains sense-organs. They attain their highest development in the larvae; later on they undergo a retrogressive change. Each of these sense-organs is a little cup-shaped papilla, visible to the naked eye. It is composed of elongated cells which form a mantle around some central cells, each of which ends in a stiff cilium perforating a thin, hyaline membrane which lines the bottom of the cup, and is perhaps the representation of the cuticula. These ciliated cells are connected with sensory fibres, the nerve entering at the bottom of the whole organ. The cilia are in direct contact with the water, but the outer rim of the whole apparatus is protected by a short tube of hyaline cuticula-like secretion. These sense-organs are, in the larvae, scattered over the head, especially near the mouth and around the eyes, whence they extend backwards on to the tail, mostly in three pairs of longitudinal rows, one near the vertebral column, the others lateral. They are supplied by the lateral branch of the vagus nerve. They disappear during the metamorphosis, at least in the Anura, with the exception of Xenopus, in which they form conspicuous white objects. The white colour is caused by the tubes becoming choked with the débris of cells or coagulating mucous matter, so that it is doubtful if these organs, which moreover have sunk deeper into the skin, are still functional. In the terrestrial Urodela these organs undergo a periodical process of retrogression and rejuvenescence. During the life on land they shrink and withdraw from the surface, and their nerves likewise diminish, but in the breeding season, when the newts take again to aquatic life, they revive, are rebuilt and become prominent on the surface. They are an inheritance from the fishes, in which such lateral line organs are universally present.
The cutis of most Amphibia is very rich in lymph-spaces, which, especially in the Anura, assume enormous proportions, since the so-called subcutaneous connective tissue forms comparatively few vertical septa by which the upper and denser layers, the corium proper, are connected with the underlying muscles. The spaces are filled with lymph, and into some of them the abnormally expanded vocal sacs extend, notably in Paludicola, Leptodactylus, and other Cystignathidae, and in Rhinoderma.
The cutis frequently forms papillae and prominent folds, sometimes regular longitudinal keels on the sides of the back; but dermal, more or less calcified or ossified scales are restricted to the Stegocephali and to the Apoda, q.v., pp. [79], [87]. We conclude that the Urodela and Anura have entirely lost these organs. Dermal ossifications, besides those which now form an integral part of the skeleton, like many of the cranial membrane-bones, are rare, and are restricted to the Anura. They are least infrequent on the head, where the skin is more or less involved in the ossification of the underlying membrane-bones, for instance in Triprion, Calyptocephalus, Hemiphractus and Pelobates. The thick ossifications in the skin of the back of several species of Ceratophrys are very exceptional. In Brachycephalus ephippium these dermal bones enter into connection with the vertebrae; small plates fuse with the dorsal processes of the first to third vertebrae, while one large and thick plate fuses with the rest of the dorsal vertebrae. Simple calcareous deposits in the cutis are less uncommon, for instance, in old specimens of Bufo vulgaris. We are scarcely justified in looking upon these various calcifications and even ossifications as reminiscences of Stegocephalous conditions.
The skin contains pigment. This is either diffuse or granular. Diffuse pigment, mostly dark brown or yellow, occurs frequently in the epidermis, even in the stratum corneum. The granular pigment is stored up in cells, the chromatophores, which send out amœboid processes, and are restricted to the cutis, mostly to its upper stratum, where they make their first appearance. Contraction of the chromatophores withdraws the pigment from the surface, expansion distributes it more or less equally. The usual colours of the pigment are black, brown, yellow, and red. Green and blue are merely subjective colours, due to interference. A peculiar kind of colouring matter is the white pigment, which probably consists of guanine, and is likewise deposited within cells; cf. the description of the white spots in the skin of Hyla coerulea.
Most Amphibia are capable of changing colour, the Urodela, however, far less than the Anura, some of which exhibit an extraordinary range and adaptability in their changes.
The mechanism by which the change of colour is produced in frogs has been recently studied by Biedermann.[[15]] If we examine the green skin of the common Tree-frog, Hyla arborea, under a low power and direct light, we see a mosaic of green, polygonal areas, separated by dark lines and interrupted by the openings of the skin-glands. Seen from below the skin appears black. Under a stronger power the black layer is seen to be composed of anastomosing and ramified black pigment-cells. Where the light shines through, the skin appears yellow. The epidermis itself is quite colourless. The mosaic layer is composed of polygonal interference-cells, each of which consists of a basal half which is granular and colourless, while the upper half is made up of yellow drops. Sometimes the tree-frog appears blackish, and if then the black pigment-cells are induced to contract, for instance, by warming the frog, it appears silver-grey; in this case the pigment in the yellow drops is no longer diffuse, but is concentrated into a round lump lodged between the interstices of the granular portions; the black pigment-cells are likewise balled together. These black chromatophores send out numerous fine branches, which occasionally stretch between and round the polygonal cells. When each of these is quite surrounded and covered by the black processes, the frog appears black. On the other hand, when the black pigment-cells withdraw their processes, shrink up, and, so to speak, retire, then the light which passes through the yellow drops is, by interference, broken into green.
Stoppage of the circulation of the blood in the skin causes the black chromatophores to contract. Carbon dioxide paralyses them and causes them to dilate. This is direct influence without the action of nerves. But stimulation of the central nerve-centres makes the skin turn pale. Low temperature causes expansion, high temperature contraction, of the chromatophores. Hence hibernating frogs are much darker than they are in the summer. Frogs kept in dry moss, or such as have escaped into the room and dry up, turn pale, regardless of light or darkness, probably owing to a central, reflex, nerve-stimulus.