Chapter 8 of 16 · 15320 words · ~77 min read

CHAPTER V

LISSAMPHIBIA (_CONTINUED_)–URODELA

ORDER II. URODELA OR TAILED AMPHIBIA.

The recent tailed Amphibia, Salamanders and Newts in the wider sense, have been grouped into four families which can be conveniently diagnosed by the following characters:–

Both the upper and lower jaws are furnished with teeth. Fore- and hind-limbs are always present.

Maxillary bones present.

Eyes free and devoid of lids .......... AMPHIUMIDAE, p. 97.

Eyes with movable lids[45] .......... SALAMANDRIDAE, p. 102.

Maxillary bones absent.

Eyes without lids. Perennibranchiate .......... PROTEIDAE, p. 132.

Both jaws are toothless. The hind-limbs, the maxillary bones and eyelids are absent. Perennibranchiate .......... SIRENIDAE, p. 136.

These four families are closely allied to each other, especially the Amphiumidae and the Salamandridae.

The geographical distribution of the Urodela is essentially Periarctic, except that about one dozen species each of _Amblystoma_ and of _Spelerpes_ extend southwards into Central America, and in the case of the latter genus even into the Andesian parts of South America. _Plethodon platense_ inhabits Argentina.

The Urodela afford good reasons for dividing the Periarctic region into three co-ordinate sub-regions, namely, Nearctic, Eastern and Western Palaearctic. The difference between the European and the Eastern Asiatic fauna is well marked; the two are–at least with our present knowledge–separated by a wide stretch of country very poor in Urodele forms; while, lastly {95}there are not a few resemblances between this Eastern Asiatic and the American fauna. The Urodela thus lend no support to the usual division of the Periarctic into a Palaearctic and a Nearctic sub-region. Nor is it possible to divide the Palaearctic into a Eurasian and a Mediterranean province. We have in this case to distinguish between an American, an Asiatic, and a European fauna. The Asiatic or Eastern Palaearctic sub-region assumes the central position, at least from a merely geographical point of view. It would be unjustifiable to assume a spreading from this centre into Europe, and, on the other hand, into America. The centre existed more probably in the Arctic circle, now devoid of Urodela.

[Illustration: FIG. 16.–Map showing the distribution of the Urodela. "Ichthyodea" = Amphiumidae + Proteidae + Sirenidae.]

So far as mere numbers of species are concerned the huge Asiatic or Eastern Palaearctic region is the poorest, but it is also the least explored, and China will probably yield a good many new forms. We know at present only 15 species, nearly all from the eastern half. These 15 species represent no less than 11 genera, 8 of which (= 73 per cent) are peculiar to the sub-region. Next comes the Western Palaearctic or European sub-region with about 21 recent species of 5 genera, 4 of which are peculiar. America is by far the richest, with no less than 66 species (36 eastern, about 16 western, and the rest Central American, etc.), belonging to 19 genera, 17 of which (= 90 per cent) are peculiar to the New World. But this richness in species is due mainly to the abundance of the two genera _Amblystoma_ and _Spelerpes_, just as Europe is characterised by its many Tritons.

One of the most striking features of the Asiatic sub-region is {96}its difference from the European. They have very little in common. _Pachytriton_, _Tylototriton_, and two species of _Triton_ (_T. pyrrhogaster_ and _T. sinensis_) are the only Salamandrinae, while all the rest are Lechriodont (see p. 102), like the American Urodela, excepting the two American Tritons, _T. torosus_ and _T. viridescens_.

GEOGRAPHICAL DISTRIBUTION OF THE URODELA

The occurrence of an _Amblystoma_, _A. persimile_, in the mountains of Siam and Burmah, is most suggestive, and others will in all probability be found. It must also be borne in {97}mind that the differences between the genera of Amblystomatinae are in reality very slight; and the same applies to the sub-families themselves. The presence or absence of teeth on the parasphenoid, the possession of amphi- or opistho-coelous vertebrae, do not mean much, and certainly do not forbid the notion that all the recent Urodela are the offspring of one common generalised stock which inhabited the northern portion of the globe. Nothing is gained by hiding the solitary European species of the essentially American genus _Spelerpes_ under the name of _Geotriton_. It is a _Spelerpes_ in all characteristic points. Speaking broadly, each of the three principal sub-families of Salamandridae is characteristic of a sub-region; the Salamandrinae of the Western Palaearctic, the Plethodontinae of the American, while the Amblystomatinae are chiefly Asiatic, at least so far as diversity of genera is concerned.

FAM. 1. AMPHIUMIDAE.–Without gills in the perfect state. The gill-clefts are in a vanishing stage, being either reduced to one pair of small holes or being altogether absent. The maxillary bones are present. Teeth occur in both jaws; those of the vomers form transverse rows. The vertebrae are amphicoelous. The fore-limbs and hind-limbs are present, but small. The small eyes are devoid of lids.

This family is now represented by two genera, with only three species, found in the United States and in Eastern Asia.

_Cryptobranchus._–The limbs are functional, with four fingers and five toes. The outer digits and the sides of the limbs are bordered with folds of skin. The head and body are stout and depressed; the tail is short, laterally compressed, and provided with a fin. The skin is very glandular and slimy, and forms a thick, irregularly-shaped fold along the side of the body.

_C. (Menopoma) alleghaniensis._–The gill-clefts are normally reduced to one pair, individually to the left cleft, the right closing up. There are, however, four branchial arches and vessels. The general colour is brown or grey above, sometimes with darker patches, lighter below. The "Hellbender" reaches a length of nearly 18 inches (about 46 cm.), is entirely aquatic, and is apparently restricted to the rivers and streams of the mountainous districts of the Eastern United States. It is very voracious, living on worms and on fish, being much disliked by the fishermen, as it takes the angler's bait, and destroys great quantities of the valuable food-fish _Coregonus albus_. Although rather common and easily kept, its larvae still remain unknown.

{98}[Illustration: FIG. 17.–_Cryptobranchus japonicus._ Japanese Giant Salamander. × ⅙.]

{99}_C. japonicus_ s. _maximus_.–The Giant Salamander of Japan differs from its American relation in one essential point only, namely, by the absence of gill-openings and of the modifications of the branchial apparatus connected therewith. It has but three branchial vessels, and the skeletal arches are reduced to two. It lives in Japan and in China, from 600 to 4500 feet above the level of the sea, in small streams of mountain-meadows. It feeds upon fishes, Amphibia, worms, and insects. It is easily fished with the hook and is eaten by the Japanese.

The first living specimen was brought to Europe in 1829 by Th. von Siebold, its discoverer. It grew within a few years from 1 foot to 3 feet in length, and died in 1881, at least fifty-two years old. Another specimen lived in the Hamburg aquarium for fourteen years, during which time it is said to have grown 36 cm. (more than 14 inches), having attained a length of nearly 4½ feet, or 134 cm. The largest specimen known measures 159 cm = 5 feet 3 inches.

The life-history of this species is still imperfectly known. Japanese picture-books contain drawings of the adult and of larvae, the latter showing three pairs of fringed external gills. Young specimens of 16 cm. length have already lost the gills, but still retain a cleft on either side of the neck, in the shape of a horizontal slit, and this is soon after closed up by the skin.

The best account has recently been given by Sasaki.[46] According to him the Giant Salamander leads a solitary life, concealed in dark places, under rocks in swift-flowing, thickly shaded small brooks of clear and cold water.

The animal may be easily captured with a fish-hook, baited with a fish, frog, or several earth-worms, and tied to a string a few feet in length. This is thrust by the aid of a small bamboo-stick into the salamander's retreat. The string is not tied to the stick, but the point of the loaded hook is forced into one end of it, far enough to keep it in place while this end of the rod is pushed under the rock. When the bait has been thus brought near the salamander, any bite will be instantly felt through the {100}rod. The latter is then withdrawn as quietly as possible, the hook and bait being left. As soon as a jerk of the string is noticed, a pull is made, which generally ends in the capture of the unfortunate animal. If the first pull should fail, the bait is replaced as before, and a second opportunity is offered, which the unwary creature accepts as readily as the first. The fisherman, having obtained one bite, is sure of ultimate success, as the salamander does not learn by experience to refuse the proffered morsel. When captured, it emits a peculiar slimy secretion, having an odour much like that of the leaves of the Japan pepper (_Xanthoxylon peperitum_). This secretion hardens into a gelatinous mass after a short exposure to the air.

Temminck and Schlegel state that the act of inspiration is ordinarily performed once every 6-10 minutes. This is true for specimens kept in tubs; but Sasaki is inclined to think that they perform this act less frequently in their native brooks. The eyes are so small that they are obviously of little importance; the salamanders capture their prey not by pursuing, but by waiting for its near approach, whereupon they seize it with their teeth by a swift lateral movement of the head. The eggs are said to be laid in August and September, and they form a string resembling a rosary. Each egg floats in a clear fluid, inclosed in a bead-shaped gelatinous envelope, and this is connected with the next by means of a comparatively small string. The egg measures about 6 mm. by 4 mm., and is yellow everywhere except at the upper pole, where it is whitish. All attempts to make _Cryptobranchus_ breed in captivity have failed hitherto, owing no doubt to the difficulty of obtaining the cool temperature of its mountain streams. Sasaki's smallest specimens measured 19 to 20 cm. These had three pairs of very short branchial processes, from 3 to 5 mm. in length, attached just inside the branchial orifice. Each process was somewhat flattened and tapering, most of them still with branchlets. In another specimen, 20.5 cm. in length, the gills had almost wholly disappeared, but the branchial slits were still visible. One of 24.5 cm. length showed no trace of gills, and the branchial orifice was completely closed, but still marked by a light streak.

{101}[Illustration: FIG. 18.–_Amphiuma means._ × ⅕.]

_Amphiuma means_ s. _tridactyla_.–The limbs are very much reduced, and end in two or three little fingers or toes. Just in front of the fore-limbs lies the pair of small gill-clefts, each guarded by two flaps of the skin. There are four branchial arches. The general colour of this eel-shaped creature is black, lighter below. The head is covered with numerous pores, arranged in several rows, which unite in the region of the neck, so that only two rows extend along the sides of the body. It reaches a length of three feet, and lives in swamps or muddy waters, for instance in the ditches of rice-fields, burrowing occasionally in the mud, feeding on crayfishes, molluscs, small fishes, etc. It is confined to the south-eastern States of North America, from Carolina to Mississippi. According to Davison,[47] copulation takes place in May. The rather hard-shelled eggs are deposited in the following August or September, and are connected by a twisted cord. The female lies about them in a coil. The embryos, which are hatched in the month of November or December, have well-developed external gills. By the following February they have {102}reached a length of from 68 to 90 mm. (about 3 inches), living in damp localities under rocks or rooted stumps, and have already lost their gills. The legs are said to be relatively longer than they are in the adult.

FAM. 2. SALAMANDRIDAE (Salamanders and Newts).–Without gills in the perfect state. Maxillaries are present. Both jaws are furnished with teeth. The eyes are protected by movable lids, except in _Typhlotriton_. Fore- and hind-limbs present, although sometimes very much reduced.

To this family belong by far the greater number of tailed Amphibia. They have been, for the sake of convenience, grouped into four sub-families, the determining characters of which are all internal and of comparatively slight importance. Little better is the division into _Mecodonta_, with the teeth of the palate in two longitudinal rows diverging behind and inserted upon the inner margins of the two palatine processes, which are much prolonged posteriorly, and _Lechriodonta_, in which the series of palatal teeth are restricted to the posterior portion of the vomers and form either transverse or posteriorly converging rows.

I. Series of palatal teeth transverse, restricted to the posterior portion of the vomers. Parasphenoid beset with dentigerous plates.

Vertebrae opisthocoelous: _Desmognathinae_, p. 102.

" amphicoelous: _Plethodontinae_, p. 103.

II. Series of palatal teeth transverse or posteriorly converging, restricted to the posterior portion of the vomers. Parasphenoid toothless. Vertebrae amphicoelous: _Amblystomatinae_, p. 109.

III. Series of palatal teeth in two longitudinal series, diverging behind, inserted on the inner margin of the long palatine processes. Parasphenoid toothless. Vertebrae amphicoelous: _Salamandrinae_, p. 115.

SUB-FAM. 1. DESMOGNATHINAE.–Comprising only three genera, with five species, in North America. Five toes.

_Desmognathus._–The tongue is attached along the median line, free behind, oval in shape. Three species in the eastern half of the United States. _D. fuscus_ is one of the lungless Urodela, for which condition see p. 46. The skin is nearly smooth; parotoids prominent, gular fold strongly marked. General colour above, brown suffused with pink and grey, sometimes with a dark lateral band; under parts mottled brown. The vomerine teeth are frequently absent. Total length, about 4 to 5 inches. They live, carefully concealed in the daytime, under {103}stones in or on the edge of the banks of little mountain streams. The eggs are laid in two long strings, and are wrapped round the body of the female like a rosary, the female having resorted to a hollow in the mud, below a stone or other suitable place. The outer envelope of each egg tapers out into a short stalk, and the several stalks all converge, or are glued together into one common knot, "much like a bunch of toy balloons held in the hand of a street vendor." The egg is said to be meroblastic. The larvae seem to remain in the egg until they are nearly adult, and they emerge at midsummer, with the gills already much reduced. The complete metamorphosis takes place in the autumn of the same year. These little newts can, according to Wilder,[48] be collected all the year round, in Massachusetts from March to December, except during the time of deep snow. They are nocturnal and are easily kept.

[Illustration: FIG. 19.–_Desmognathus fuscus_; female with eggs in a hole underground. × 1. (After Wilder.)]

_Thorius pennatulus_, from Orizaba, Mexico, the only species, is noteworthy for its extremely large nostrils, and for the tongue, which is supported by a central pedicle, free all round, and ending in a thick knob, which can probably be protruded. The limbs are weak, and the digits are also much reduced. Total length, under 2 inches, or 50 mm.

_Typhlotriton spelaeus_, of the Rock House Cave in Missouri, is blind, the eyes becoming concealed by the skin during metamorphosis, when the gills are lost.

SUB-FAM. 2. PLETHODONTINAE.–The five genera of this almost entirely American sub-family (only one species of which, _Spelerpes fuscus_, occurs in Europe) can be distinguished as follows:–

I. The tongue is attached by its central pedicle only, is free all round, ends in a soft knob and can be shot out to a considerable distance.

With 5 toes: _Spelerpes_, p. 104.

With 4 toes: _Manculus_, p. 106.

{104}II. The tongue is attached along the middle line and cannot be protruded out of the mouth.

Jaws with numerous small teeth.

With 5 toes: _Plethodon_, p. 106.

With 4 toes: _Batrachoseps_.

Maxillary and mandibular teeth few in number but very large.

With 5 toes: _Autodax_, p. 107.

_Spelerpes._–Except in a few species the limbs are well developed and possess 4 fingers and 5 toes, which are either free or webbed. But in the Colombian _S. parvipes_, still more in _S. lineolus_ of Orizaba and _S. uniformis_ of Costa Rica the limbs and digits are reduced to mere vestiges, and are practically without function, the body, with the extremely long tail, having assumed a wormlike shape. The young of many, if not all, species have a pair of short balancers below each nostril; in the adult these organs are reduced to little swellings or lost completely. Several species are lungless, see p. 46.

The geographical distribution of this genus, of which some twenty species are known, is very remarkable. The majority live in Mexico and in the United States, a few are found in Colombia and Northern Peru (_S. altamazonicus_ and _Plethodon platense_ being the only Urodeles hitherto recorded from south of the equator), one in Hayti (_S. infuscatus_), two (_S. subpalmatus_ and _S. uniformis_) in Costa Rica, and _S. fuscus_ in Europe.

_S. bilineatus_ is a little newt under 4 inches in length–60-95 mm.–found in the Atlantic States. It is brownish-yellow above, with a black lateral line extending from the eye to nearly the end of the tail. The under parts are bright yellow. It lives on land, in damp places, concealed during the daytime under stones or old trees, whence it emerges after a rain or in the dusk of evening.

According to H. H. Wilder,[49] "the eggs are deposited in May and June in a single layer upon the lower side of submerged stones, each batch containing 30 to 50 eggs. The stones which are suitable for this purpose must be in the form of an arch, allowing the water to flow beneath. They are generally in the more rapidly flowing portions of the brook, but the depth of water must be such that the eggs are at all times entirely submerged. They are attached to the stone by gelatinous threads, proceeding from the outer envelope, and although they are generally contiguous, they {105}are each attached separately." The eggs are holoblastic. The larvae hatch early and continue for a long time in the larval state, probably two or three years.

_S. porphyriticus_ s. _salmoneus._–Yellowish-brown or purplish-grey above with tiny darker dots and markings. The sides of the body are salmon-coloured, with a tinge of yellow. The under parts are whitish, turning into salmon-pink on the tail. This beautiful newt reaches about 6 inches in length and has a very moist, slimy skin, which, combined with the lively motions of the creature, make it as slippery as an eel. It is found in the Alleghany range, from New York to Alabama.

Specimens which I am keeping prefer the wettest part of the cage, where they lie concealed in the moss and mud, leaving their hiding-places at night in search of insects. One of them escaped into the greenhouse and was discovered after nine months, having established its permanent home in a cleft between mossy stones: when the sweepings of a butterfly-net are emptied near its hiding-place it peeps out and with a flash of its long, forked, white-coloured tongue it secures its prey. Occasionally it goes into a tank, when it swims with rapid, undulating motions, the limbs being laid back and remaining inactive; it sometimes rises to the surface to emit and to take in air, but, although mostly resting half in the water, upon a rotten stump, it often lies for hours at the bottom without stirring. When kept in dry surroundings, the skin soon dries and wrinkles, and the animals show every sign of suffocation and general discomfort. The respiration of this lungless species by means of rapid movements of the throat is very limited, most of the necessary oxidisation of the blood being effected through the skin.

_S. fuscus._–This, the only European species, is thoroughly terrestrial. It is found in the mountains bordering the Gulf of Genoa, and in Sardinia. Its total length remains under four inches. The smooth, very delicate and easily broken skin is brown above, light below, and speckled with lighter and darker markings. Below each nostril is a slight swelling, the remnant of the cirri or balancers common to the young of many species. It lives in shady surroundings, under stones, in old trees and in limestone-caves, glued to the walls with spread-out toes, belly and tail, quietly waiting for insects and spiders which it catches by flashing out the long tongue.

{106}According to J. Berg,[50] it keeps well in cool, moist and well-ventilated places. It lives on flies, small beetles, and maggots; ants are also taken at once, probably owing to their lively movements, but a few minutes later the newts roll about in spasms and soon die. Towards the end of March one of Berg's specimens gave birth to four young, which were 36 mm., or nearly 1½ inches long, and differed from the adult only by their exceptionally large nostrils, thereby resembling the Mexican _Thorius_. The little ones shot out their tongues about 10 mm., feeding on Aphides.

[Illustration: FIG. 20.–_Spelerpes fuscus_, showing the position and shape of the partly and fully protruded tongue. The figure on the right side shows the tongue and the skeleton of the hyoid apparatus. _B_, the threadlike, elongated, first branchial arch; _H_, hyoid, in reality attached by its outer end to the vicinity of the quadrate; _T_, tongue. About × 2. (After Berg and Wiedersheim.)]

_Manculus._–The two species of this genus live in Carolina and Florida. _M. quadridigitatus_ is a very slender, graceful little animal, about 3 inches in length, the long and thin tail being considerably larger than the rest of the body. Yellowish, minutely speckled with brown above and on the sides, greyish-white below. Life entirely terrestrial.

_Plethodon._–About seven species in North America. This genus has given its name to that of the subfamily, which might with more reason be called Spelerpinae.

_P. glutinosus_ is slaty or bluish-black, with small whitish specks, especially on the sides of the trunk, where they are large and often confluent. The skin is smooth and shiny. Total length about 5 inches, half of which belong to the tail. Holbrook considered this as one of the commonest of the North American newts, and mostly widely distributed, from Ohio to the Gulf of Mexico. It usually lives concealed under stones, but prefers fallen trees, probably on account of the insects upon which it {107}preys. When taken in the hand it gives off a great quantity of slime.

_P. erythronotus_ extends into Canada and is much smaller. Brown or grey above, mostly with a broad, reddish-brown band over the head, back, and tail. The under parts are white, with grey and brown specks.

_Autodax_ s. _Anaides_.–The large tongue is attached along the median line. The jaws are furnished with few, but surprisingly large, knife-shaped teeth, about ten in the upper and fewer in the lower jaw. The small teeth of the vomers form a chevron-shaped series behind the choanae, those of the parasphenoid stand in one elongated patch. The tail is round; number of toes, five. Three species in Western North America, from California to Oregon.

_A. lugubris._–The eyes are very large and prominent. The upper jaw shows a peculiar recess on either side for the reception of the large lower teeth. The skin is smooth, devoid of parotoid glands, but has a strong gular fold. The upper parts are dark brown or lead-coloured, with whitish dots on the sides; under parts white. Total length some 6 inches, about half of which belongs to the tail. The fingers and toes are very rich in subcutaneous venous sinuses.

The habits of these creatures are in many respects peculiar. Van Denburgh[51] says of _A. iecanus_ "that it usually moves quite slowly, moving one foot at a time, but is capable of motion surprisingly rapid for a salamander. When moving rapidly, it aids the action of its legs by a sinuous movement of its whole body and tail. The latter is prehensile. Several individuals, when held with their heads down, coiled their tails around my finger, and, when the original hold was released, sustained themselves for some time by this means alone. One even raised itself high enough to secure a foothold. This animal's tail is also of use in another way. When caught, it will often remain motionless, but if touched, will either run a short distance with great speed, or quickly raising its tail and striking it forcibly against the surface on which it rests, and accompanying this with a quick motion of its hind-limbs, will jump from four to six inches, rising as high as two or three."

Ritter and Miller[52] have made extensive observations on the life-history of _A. lugubris_. When wishing to pass from the hand to {108}the table, the creature will frequently execute a well co-ordinated spring and alight on its feet some distance away, instead of falling over the edge in the typical salamander-fashion. This species is nocturnal and entirely terrestrial, and seems to be indifferent even to proximity to water. Rotten stumps and logs are the habitations preferred, and wherever these occur in the region about San Francisco Bay, even though at the places remotest from water, specimens are sure to be found.

The eggs are laid in a hollow under ground, and the female seems to remain curled around them until they are hatched, which takes place in two or three weeks. The specimen observed by Ritter and Miller laid 19 eggs. Each was contained in a gelatinous capsule 6 mm. in diameter, and was firmly anchored to a clump of earth by a narrow peduncle about 8 mm. long. The embryos developed very large gills, each being composed of three broad membranous lobes, the latter being thin and delicate, much expanded, highly vascular and widely confluent at their bases, so that the gills of each side really form one three-lobed mass. Their dorsal surfaces are applied to the inner surface of the egg-capsule. The amount of food-yolk is considerable. The whole larval life is passed through within the egg. Before the young is hatched the gills wither and cease to be functional, and the gill-slits close up. The tail is round, and shows no indication of a fin at any time during the larval period. Newly hatched individuals appeared much distressed when put into water, and were quite unable to swim. They immediately sank to the bottom and remained there until they were removed. The integumentary sense-organs, so well developed in the aquatic larvae of Urodeles, are entirely wanting. When hatched the young creature is about 32 mm. long; its general colour is blackish-grey, finely sprinkled with bluish-silver. During the second year this garb is changed to the dusky brown of the adult, and the fine silver speckling is replaced by much larger and less numerous yellow spots.

Although one of the most terrestrial of Urodeles, this species is lungless, but the skin remains delicately smooth and moist throughout life. According to the observers quoted, the pharynx plays an important part in respiration. From 120 to 180 or even more vibrations are made by the throat in a minute, and in some cases these movements are grouped into series of about {109}20 to 25 extremely rapid vibrations, with periods between each two series.

SUBFAM. 3. AMBLYSTOMATINAE.–Composed of seven closely allied genera, the distinguishing characters of which are the grouping of the palatal teeth and the number of the toes, which varies between 4 and 5. The geographical range of the subfamily extends over the whole of North America and Mexico and over the whole of Northern Asia, from Kamtchatka and Japan westwards to the Ural, and southwards into China. The occurrence of one species, _Amblystoma persimile_, in the mountains of Siam, makes it highly probable that other species and genera exist in the hitherto unexplored intervening countries.

Boulenger gives the following synopsis:–

I. The series of palatal teeth converge backwards, forming a V-shaped figure.

With 5 toes: _Hynobius_, 3 species in Japan.

With 4 toes: _Salamandrella_, 2 species Lake Baikal, Ussuri and Schilka rivers, and Kamtchatka, p. 109.

II. The series of palatal teeth form an uninterrupted, doubly arched V-shaped figure.

The 4 fingers and 5 toes are furnished with black, horny claws: _Onychodactylus japonicus_.

III. The series of palatal teeth form two arches, convex forwards, separated by a wide interspace.

The two series are short, confined to the space between the choanae.

With 5 toes: _Ranidens sibiricus_, Eastern Siberia and N.E. China.

With 4 toes: _Batrachyperus sinensis_, Moupin in China.

The series are long and converge backwards, 5 toes: _Dicamptodon ensatus_, California.

IV. The palatal teeth are arranged in a nearly straight, transverse line, or they form an angle which points slightly forwards; they are not separated by a wide median space. With 5 toes: _Amblystoma_. Some 16 species in North and Central America, one in Siam, p. 110.

_Salamandrella keyserlingi._–The mode of propagation of this newt-like species has been observed by Shitkow near Jekaterinburg in the Ural mountains. The eggs were laid at the end of April and were deposited in bags, which were attached to a plant, with one end about an inch below the surface of the water. The bag measured 15 cm. in length and 2 cm. in width and contained 50 to 60 eggs. The larvae were hatched in 14 days in a sunny aquarium; in another with a northern {110}aspect the hatching took 23 days. The larvae were 10 mm. long, and remarkable for the length (1 mm.) of their balancers.

_Amblystoma opacum._–The general shape is very much like that of the European Spotted Salamander. The head is short and broad, the snout is rounded. The eyes are very prominent, with a black pupil and a dark-grey iris. The neck has a well-marked gular fold. The tail is thick and almost round. The hind-limbs are considerably larger than the fore-limbs. The general colour of the shiny, moist skin is a purplish-black with light grey, transverse, partly confluent bars, giving the creature a pretty appearance; the under parts are paler, bluish-grey. Total length between 3 and 4 inches, or 9 cm.

This beautiful species inhabits many of the United States east of the Rocky Mountains, from New Jersey to Florida and Texas. In the perfect state it is thoroughly terrestrial and easily kept. My specimens prefer the holes of rotten and moist, moss-covered stumps, or holes beneath stones, which they leave, at night only, in search of earthworms and insects.

_A. talpoideum_ is closely allied, somewhat stouter and almost uniform brownish-back. According to Holbrook, "it chooses light soil in which it will bury itself in a few seconds like a mole, and there continue its course concealed from view; but its track can often be followed by the elevation produced on the surface of the soil, similar to that seen in fields infested by moles."

[Illustration: FIG. 21.–Egg-sac of _Salamandrella schrenki_. × ½. (After Shitkow.)]

_A. punctatum_ is bluish-black, with a row of roundish yellow spots on each side of the body and tail and upon the limbs.

E. A. Andrews[53] has made observations upon the breeding of this species. Near Baltimore the eggs are very abundant in March and even in February, in small pools in the woods, but the adults are then rarely seen. Even when small pools, but 4 feet wide and 9 inches deep, were thoroughly raked out {111}before and after the eggs appeared, no adults were found, so that it is to be inferred that the laying takes place in the night and that the adults leave the water every day to conceal themselves under stones. One female was found moving away from a bunch of eggs early in the morning. This specimen was kept isolated, and laid many eggs, and as these developed into normal larvae, the existence of internal fertilisation was proved. Previously to the laying of the eggs white spermatophores were found in the small pools, on the dead twigs and leaves covering the bottom.

_A. jeffersonianum._–This very slender and slippery species, reaching a length of 6 inches, is remarkable for its long fingers and toes, and its rather compressed tail. The general colour is brown above, dirty whitish below, generally with numerous, small, light blue and pale brown spots on the sides of the neck, body, limbs, and tail. There are several colour-varieties, one of them with white specks. It is a very active and surprisingly good climber, easily escaping out of high-walled bell-glasses, hiding in the daytime in dark and moist localities. Its range extends from Indiana and Virginia to Quebec.

_A. persimile._–This species is remarkable on account of its geographical distribution. It is the only non-American species, inhabiting the higher mountains of Siam and Upper Burmah. There is no doubt about its belonging to the genus _Amblystoma_, although it had originally been described as a _Plethodon_. It closely resembles _A. jeffersonianum_ in most of its characters, notably in the arrangement of the palatal teeth, general proportions, slender toes, and even in the presence of whitish spots, which are scattered over the sides of its blackish, smooth skin.

_A. tigrinum._–This, the commonest species, is conspicuous for its large, depressed head, which is as broad as it is long, its width being enhanced by the unusually large parotoid glands. The mouth is very wide. The large, prominent eyes are golden, and reticulated with brown. The gular fold is strong. The limbs are stout, the fingers and toes short. The trunk is strongly constricted by twelve intercostal grooves. The tail, which is as long as the rest of the body, is somewhat compressed laterally, but bears no trace of a fin. The general colour is more or less dark brown or bluish black, marked with numerous yellow spots and large blotches; the under surface inclines to {112}grey. The length of the adult male is about half a foot; the females, as usual being larger, sometimes reach the length of 9 inches. The range is from New York to California and to Central Mexico.

The larva of this species is the famous Axolotl. It is provided with three pairs of delicate and much-branched external gills, a flat, long tail with a broad ventral and dorsal fin, the latter extending along the back almost to the neck. The limbs, although comparatively slender, are fully developed, and the head is much more pointed than it is in the perfect form. The larvae usually reach 8 or 9 inches in length; exceptional specimens have been recorded of one foot in length, and have been described as _Triton ingens_.

[Illustration: FIG. 22.–Axolotls or larvae of _Amblystoma tigrinum_. × ½.]

These larvae were found by the Spanish conquerors to occur in great numbers in the lakes near Mexico City, and were called Axolotl by the natives, a word signifying "play in the water." They were, and are still, eaten, either roasted or boiled, with vinegar or cayenne pepper.

For many years these creatures were looked upon as a species of the Perennibranchiata, under the generic name of _Siredon_ (_S. axolotl_, _s. pisciformis_, _s. mexicanus_, etc.), although Cuvier suspected that they were but the larvae of an otherwise unknown terrestrial Urodele. The mystery was not cleared up until the year 1865, when some Axolotls which had been kept for a year in the Jardin des Plantes at Paris, suddenly began {113}to pair, and laid eggs which within six months developed into full-sized Axolotls. This certainly looked as if these creatures were not larvae, but a true Perennibranchiate species. But to the general surprise several of these young Axolotls gradually lost their gills, the clefts closed up, the fins of the back and tail disappeared, the head became broader, the creatures left the water permanently, and in fact turned into the already well-known terrestrial _Amblystoma tigrinum_. The other brothers and sisters of the same brood remained aquatic Axolotls, which thereby revealed themselves after all as the larval and not as the perfect stage of this remarkable species.

At the suggestion of Kölliker and Weismann, Frl. Marie von Chauvin[54] undertook, at the University of Freiburg, long and carefully conducted experiments, showing (1) that little Axolotls can comparatively easily be caused to develop further into the perfect _Amblystoma_ if they are induced to breathe air more frequently than usual; shallow vessels, perhaps also insufficiently aerated water, will produce the desired result; (2) that the commencing metamorphosis can again be checked, the shrinking gills then undergoing fresh development; (3) that they can be forced to remain Axolotls; (4) that the cutting off of the gills has no influence upon their possible metamorphosis, the gills being easily and quickly renewed. The same lady found also that _Amblystoma_, the perfect form, lives in the water during the pairing time and behaves in the same way as the Axolotls.

The latest observations have been made by Metzdorff.[55] Axolotls, at least those which are kept in captivity in Europe, are ready for propagation several times in the year, either in the spring, from April to June, or in December. The male deposits spermatophores, which in the following night are taken up by the female into the cloaca. On the following day, preferably in the afternoon, she grasps a suitable leaf, for instance that of _Vallisneria_, with the hind-limbs, and presses it against the vent. The eggs are expelled by strong wriggling movements of the body, and are formed into three or four packets of six to ten eggs each, so that about thirty eggs are laid at one sitting. {114}Then she takes a rest before proceeding again; the whole process, in which the male takes no further interest, lasting about two days. The most suitable temperature is one of 18-20° C., or about 68° F. The water must be well aerated. Sterile eggs turn white on the second day. The little larvae are hatched in about a fortnight. Eggs which are kept in a higher temperature, from 22-24° C., develop more quickly, but the resulting young are smaller; they show already on the fifth day head, tail, and the beginning of the gills. According to Bedriaga, they live at first upon Infusoria and _Daphnia_; when they are 20-25 mm. long they eat _Tubifex rivulorum_; later on they take scraped meat and are liable, when hungry, to nibble off each other's gills, but these are easily reproduced. When 20-25 cm. long, at the age of about six months, they are able to breed. The chief point of interest is the fact that this species of _Amblystoma_ frequently remains throughout life in the larval state, except that it develops generative organs. The natural causes of this retention are not completely known. According to Shufeldt, who observed them under natural conditions near Fort Wingate in New Mexico, plenty of food, the drying up of the swamps, and the increasing temperature of the diminishing water, hurries on the metamorphosis, while deeper water retards it. Weismann[56] suggested that the specimens in the Mexican lakes which remained Axolotls were prevented from becoming perfect Amblystomas on account of these lakes, after the disappearance of the surrounding forests, having receded from their former boundaries, which are now covered with a saline, uninhabitable crust. This may be an explanation, although Axolotls do not live in brackish water. But Weismann went farther, and with his well-known dialectic powers has succeeded in spreading the belief not only that the Axolotl is a case of reversion to an ancestral stage, but that the present _Amblystoma_, instead of being the progressive, perfect form, is likewise a case of reversion. A reversion from a reversion! The whole line of evolution would then be as follows: _Amblystoma_; its young, owing to adverse circumstances, revert to the stage of the Perennibranchiate ancestors of all Urodela; if some of these Axolotls lose their gills and fins, they revert thereby into the original _Amblystoma_. {115}Surely a roundabout way of explaining a curious but after all rather simple process of Neoteny; cf. p. 63.

Observations on the metamorphosis of _Siredon lichenoides_ into _Amblystoma mavortium_ have been made by Marsh, who also gives figures of the larval and adult forms.[57]

SUB-FAM. 4. SALAMANDRINAE.–The six genera of this subfamily fall into two natural groups: I, True Salamanders, with the palatal teeth arranged in a pair of S-shaped figures, and without a fronto-squamosal arch. II, Tritons, with the palatal teeth in the shape of a Λ, _i.e._ the right and left series meet at an angle; the fronto-squamosal arch is present, either bony, or at least ligamentous. _Triton cristatus_ is, however, exceptional, in that the two palatal series often do not meet and that the arch is absent. The number of fingers is universally four, that of the toes is five except in _Salamandrina_, which has only four.

The geographical distribution of the sub-family, entirely Periarctic, may be said to be the reverse of that of the Amblystomatinae. Of the twenty-five species namely, only two are American, four are Eastern Asiatic, and of the remaining nineteen, two are Algerian, while the rest live in Europe or in Asia Minor. It is in fact an essentially Palaearctic group.

The six genera can be distinguished as follows:–

I. The palatal teeth are arranged in two S-shaped curves. True Salamanders.

Tongue short and thick. _Salamandra_, p. 115.

Tongue long and projectile. _Chioglossa_, p. 121.

II. The palatal teeth are arranged in a Λ shape. True Tritons.

With only four toes. _Salamandrina_, p. 122.

With five toes.

Pterygoids separated from the maxillary and quadrate bones: _Triton_, p. 122.

Pterygoids touching the maxillae and quadrates. Himalo-Chinese: _Tylototriton_, p. 132.

Pterygoids united broadly with the maxillae. Chinese: _Pachytriton_, p. 132.

_Salamandra._–Without fronto-squamosal arch. Five toes. Tail round. Three species in Europe and Western Asia.

_S. maculosa._–The Spotted or Fire Salamander. General habit stout. Usual length about 5 to 6 inches; the females are mostly larger than the males; specimens of more than 8 inches in {116}length are giants. Head as broad as it is long, snout rounded. Limbs and digits stout and short. The skin is smooth, shiny and full of pores, with a strong gular fold. The parotoid glands are large and covered with large pores. A series of distinct swellings, or cutaneous glands, each with a distinct opening, extends along either side of the back, and a shorter series along the flanks. The general colour of the Spotted or Fire-salamander is black, with irregular, large yellow patches on the back and limbs. These markings vary extremely, so much so that scarcely two specimens, collected at random, are alike. In some the yellow patches form two more or less regular bands, in others they are partly confluent; again the yellow may be preponderant on the back or much restricted. Occasionally the chrome-yellow is replaced by orange. The under surface is as a rule bluish grey-black. This combination of shiny yellow and black is a good instance of warning colours. The creature is poisonous, cf. p. 38. When left in peace, or handled gently, it is perfectly harmless, but when treated with violence, or submitted to severe pain, a milky white fluid exudes from the glands and is, under violent contractions of the muscular skin and body, sometimes squirted out in fine jets to the distance of a foot. Burning pain and subsequent inflammation result if this poison gets into the eye. The same applies to the mucous lining of the mouth and throat. A few drops of this poison introduced into the blood or into the stomach of a small animal are sufficient to cause its death. Cold-blooded animals are as susceptible as warm-blooded creatures.

I once put two American bull-frogs into the same outdoor enclosure with a large number of salamanders. Next morning the huge frogs were found dead, each having swallowed a salamander, which they were not acquainted with and had taken without suspicion.

The Fire-salamander has a wide range, namely the whole of Central, Southern, and Western Europe with the exception of the British Isles. It extends southwards into Corsica and Algeria, eastwards through Asia Minor into Syria. Where it does occur it is rather common, provided the terrain is mountainous or hilly and covered with vegetation. There it lives under moss or rotten leaves, in the roots of old trees, in the cracks and clefts of the ground, of rocks or of ruins of buildings; {117}in default of anything better under heaps of stones, or in the holes dug by mice or moles. One chief necessity for its happiness is moisture.

The salamander does not occur everywhere, but is rather local. On certain kinds of limestone it is rare or absent; granitic terrain and red sandstone seem to suit it best, for instance the Hartz Mountains, Thuringia, and Heidelberg are favourite localities. But even there we may spend days and weeks and never come across a single specimen. We may turn stones, rake up the moss and leaves, pry into cracks, and we unearth perhaps a few sorry-looking, listless, dull and dry, half-emaciated creatures. The same place after a thunderstorm will be literally swarming with sleek, lively salamanders, in search of earthworms and all kinds of insects, especially at dusk or during the night. They disappear in the autumn, in October, to hibernate in the ground, out of the reach of frost, and they reappear again in April. Later on they congregate at little springs, always at running water, to reach which they have often to make long migrations. This is the only time when these thoroughly terrestrial creatures approach water, in which they easily get drowned.

Although this species is so common its mode of reproduction has been satisfactorily discovered only quite recently. There are some puzzling facts which it took a long time to observe correctly and to interpret. The larvae are born in April, May, or June, while there are no eggs in the oviducts, but in July these are full of fertilised eggs before copulation takes place. This seems contradictory. The explanation is as follows. In July there is an amplexus of the sexes, short, and often on land–a sort of preliminary exciting performance. Both sexes then descend into the water, but generally remain on land with the fore part of the body. The male deposits a spermatophore and the female takes part of this into its cloaca. In the case of a virgin female the eggs are fertilised in the oviduct and ripen until the autumn, but the larvae nearly ready for birth remain within the uterus until the following May, _i.e._ about ten months. The mother then crawls half into the water, mostly at night, and gives birth to from a few to fifty young, fifteen being perhaps the average. The young are surrounded by the egg-membrane, which either bursts before or shortly after expulsion. This species is consequently viviparous in the proper sense. If she produces a few young only, say from {118}two to five, these are much larger and stronger than those of a large litter. Occasionally a few addled or only partly developed eggs are also expelled.

In the case of old females which have produced offspring before, the whole process is more complicated. The sperma, taken up in July, remains in the receptaculum of the cloaca until the May or June following, _i.e._ until the previous larvae have passed out of the uterus and are born. Then the spermatozoa ascend to the upper ends of the oviducts, where they meet and fertilise the new eggs. After these have descended into and filled the uterus, and are already developing into embryos, copulation takes place again in July, preparatory for next year's eggs.

The new-born salamanders have three pairs of long external gills, a long tail furnished with a broad dorsal and ventral fin, and four limbs, although these are small. The total length is about 25 mm. or 1 inch. The general colour is blackish with a pretty metallic golden and greenish lustre. The little creatures are very active, and at once eat living or dead animal matter. In captivity they are liable to nibble each other's gills and tails. During the first six or eight weeks they assume a row of dark spots on the sides; these spots enlarge, and the whole skin becomes darker. Yellow spots appear next, first above the eyes and on the thighs, later upon the back; the ground-colour at the same time becomes black, until at the beginning of the fourth month they look like the parents.

The metamorphosis is very gradual. The tail-fin diminishes first, but the gills grow until shortly before the little creatures leave the water. Darkness, cold, and insufficient food retard the metamorphosis, sometimes until October. It is easy to rear them artificially provided they are well fed, kept in a light place, and in clean, well aerated water. If prevented from leaving the latter, for instance when kept in a glass vessel with vertical walls, or if hindered by a piece of gauze from rising to the surface and taking in air, they can be kept as larvae well into the winter.

Very young, perfect little salamanders, of from 1 to 2 inches in length, are excessively rare; even specimens of 3 inches are far from common. They probably spend the first two or three years of their life in careful seclusion.

A few adults can be easily kept for many years in shady {119}places provided with moss, rotten stumps and stones, to afford them suitable moist and cool hiding-places, and they readily take earthworms, larvae of beetles, snails, woodlice, etc. But any attempt to keep them in large numbers ends in failure. They congregate together in clumps, all making for the same cavity or recess, as if that were the only one in existence (very likely they are right in so far as that place is probably the best), and they get rapidly enlarging sores, chiefly on the elbows and knees. These are soon infested with fungoid growths, and this disease spreads like an epidemic and soon carries them off.

_S. atra._–The Alpine Salamander differs from the Spotted Salamander by its uniform black colour and smaller size, which averages between 7 and 5 inches. It is restricted to the Alps of Europe, from Savoy to Carinthia, at from 2000 to as much as 9000 feet elevation, living with predilection near waterfalls, the spray of which keeps the neighbourhood moist, or in mossy walls, in the shade of forests near brooks, or under flat stones on northern slopes. The most interesting feature of this species is that it produces only two young at a time. These are nourished at the expense of the partially developed eggs in the uterus, and they undergo their whole metamorphosis before they are born. By far the best and most complete account of this mode of propagation has been given by G. Schwalbe.[58] The length of the ripe embryos is about 45 mm.; they lie mostly bent up, with their heads and tails turned towards the head of the mother. The gills are beautiful, delicate red organs, the first pair being generally directed forwards and ventralwards, the second upwards, the third backwards; they are longest when the creature is about 32 mm. long, while there is still much yolk present. At this stage the gills are so long as to envelop nearly the whole embryo. There is rarely a second embryo in the same uterus, and an extra foetus is generally smaller, frequently a monstrosity not fit to live; it is probable that it is not used as food, but that it is expelled at parturition. The embryo passes through three stages, (1) still enclosed within its follicle and living on its own yolk, (2) free within the vitelline mass which is the product of the other eggs, (3) there is no more vitelline mass, but the embryo is possessed of gills 10-12 mm. in length, and is still growing. During the {120}second stage the yolk is directly swallowed by the mouth. The walls of the maternal uterus are rather red. The exchange of nutritive fluid takes place through the long external gills, which thereby function in the same way as the chorionic villi of the Mammalian egg. Each gill contains a ventral artery and a dorsal vein, each of which looks like the midrib of a pinnate leaf; there is also a fine nerve and a weak bundle of striped muscular fibres. Each gill-filament receives a capillary artery which extends to the epithelium of the tip, where it turns into a capillary vein. The epithelium of these filaments, which are full of blood, is ciliated, the resulting current being directed from the base towards the tip. In older larvae this ciliation becomes restricted to the tips. The body of the gills is furnished with flat epithelium, these non-ciliated portions alone are closely appressed to the uterine wall, and it is here that the exchange of gas takes place between mother and larva. The nutrition takes place through the gills, as they are bathed by the yolk-mass.

Schwalbe also explains the whole question of the reduction of the number of embryos. He says rightly that in _S. maculosa_, which gives birth to many young, there are in the oviduct many eggs which have only partly developed into embryos, and these, perhaps from want of room and nourishment, degenerate into the irregularly shaped whitish-yellow bodies which are occasionally found packed in between the developing embryos. Consequently all those eggs had been fertilised near the ovaries. _S. atra_ exhibits a further stage in so far as most of the eggs, fertilised above in the oviduct, degenerate, and only two or three become fully developed. These few embryos live on the degenerating eggs, which together produce the vitelline material spoken of above. The two full-grown and metamorphosed embryos, each measuring about 50 mm. in length, are equivalent to the numerous new-born larvae of _S. maculosa_, especially if the smaller size of the adult Alpine Salamander is taken into consideration.

Mlle. von Chauvin[59] has experimented with the unborn larvae of this Salamander. She cut out 23 larvae and put them into water. One of them, already 43 mm. long, took earthworms on the next day, and the beautiful long, red gills became pale and shrunk, and on the third day were cast off close to the {121}body. New gills sprouted out on the same day, first in the shape of three tiny knobs on either side. After three weeks they had become round globes, which gradually sprouted out into several branches, far shorter and more clumsy than the original gills. During the whole time the larva was lying quietly at the bottom, in the darkest corner, but showed a good appetite. The fin of the tail disappeared and was supplanted by a stronger one. In the sixth week the skin was shed in flakes, and this process took fifteen days. This larva lived in the water for fourteen weeks and grew to 6 cm. in length! When the new gills gradually shrank, the compressed and finny tail assumed a round shape, the skin became darker and shinier, and after the larva had again shed its skin, there appeared the dark rugose skin of the typical _S. atra_. The gills were reduced to useless appendages–not cast off–and the creature crawled out of the water. A fortnight later the gill-clefts were closed. A second larva behaved similarly, first casting off the feathery gills, substituting a new and stronger set, which, however, fourteen days after excision from the uterus, shrank again, and on the nineteenth day the gill-clefts were closed. The lady also observed that nearly ripe larvae, when cut out, rushed about in the water and ate, just like the new-born larvae of the Spotted Salamander.

A third species, _S. caucasica_, is found in the Caucasus. It rather resembles the Spotted Salamander in coloration, but has a larger tail and lacks the lateral warts. The male is remarkable for the possession of a soft permanent knob or hook at the top of the root of the tail. This pommel possibly prevents the slipping off during the amorous amplexus, provided the sexes then entwine like certain Tritons.

_Chioglossa lusitanica._–The only species of this genus is restricted to the north-western third of the Iberian peninsula. This graceful, slenderly-proportioned and beautiful Salamander is apparently very rare and local, having hitherto been found at a few places, namely, near Coimbra, Oporto and Coruña. It lives under moss, and runs and climbs with an agility surprising in a Urodele. The tongue is long, ending in a fork, and is supported by a median pedicle so that the tip can be quickly protruded to the distance of more than an inch. The whole length of the animal is about 5 to 6 inches, two-thirds of which belong to {122}the long tail, which is compressed at the end. The skin is smooth and shiny, with a gular fold and large parotoids. The general colour is a rich dark brown, with a pair of broad reddish-golden bands along the back and tail, the bands being separated by an almost black vertebral line.

The few specimens which I have been lucky enough to observe made little holes or passages in the moist moss of their cage, peeping out with their heads in wait for little insects, which they caught with flash-like quickness. They seem to be crepuscular.

_Salamandrina perspicillata._–This genus, represented by one species, a native of Liguria and Northern Italy, possibly extending into Dalmatia, is the only Salamander which has but four toes. The skin is not shiny and smooth, but is finely granular and dry, forms no gular fold, and is devoid of parotoid glands. The tail is more than half the length of the animal, which measures from 3 to 4 inches. The general colour is black-brown with a broad V-shaped orange-yellow mark extending from eye to eye over the occiput. A faint irregular yellowish line extends along the middle of the back and tail. The throat is black, with a diffused white patch in the middle; the belly is white, with black dots; the anal region, the inner sides of the legs and the under side of the tail are carmine-red.

This slender and pretty Salamander is diurnal, and feigns death when discovered. Only the female goes into the water, in March, to glue the eggs on to submerged rocks or water-plants. The young finish their metamorphosis by the month of June, and reach full size during the winter, the climate of their home being sufficiently genial to make hibernation scarcely necessary.

_Triton_ s. _Molge_.–The tail is strongly compressed and frequently has a permanent fin. The fronto-squamosal arch is variable, it being either bony as in the South European, Eastern and American species, or reduced to a ligament, or lastly absent as in _T. cristatus_. The males of all the English Newts, of _T. vittatus_ and of _T. marmoratus_, develop a high cutaneous crest on the back and tail during the breeding season, and this crest acts not only as a swimming organ and ornament, but also as a sensory organ.

The whole genus comprises some eighteen species, twelve of which are European, although some of these extend into Western {123}Asia; _T. pyrrhogaster_ and _T. sinensis_ are found in N.E. China, the former also in Japan; _T. poireti_ and _T. hagenmuelleri_ live in Algeria, and only two, _T. torosus_ and _T. viridescens_, are North American. Some of the species have a limited range; thus _T. montanus_ is confined to Corsica, _T. rusconii_ to Sardinia, _T. boscai_ to the north-west of the Iberian peninsula and _T. asper_ to the Pyrenees.

Newts all prefer moisture without heat. During the pairing season they take to the water, mostly to stagnant pools, which sometimes implies long migrations. During this period, which is in some cases rather prolonged, they become thoroughly aquatic and undergo some important changes. The tail-fins are much enlarged; in the males of some species a high cutaneous fold grows out on the back, devoid of muscles, but rich in sense-organs. The whole skin, instead of being dry, possesses numerous mucous glands and, what is of more importance, specialised sensory apparatuses which are arranged chiefly along the lateral lines of the body and part of the tail.

After the breeding season Newts become terrestrial, hiding in cracks, trees, or in the sandy soil. Some species aestivate during the hot and dry season. They hibernate either in the ground, or occasionally in ponds. _T. vulgaris_ is difficult to keep in the water beyond the pairing season, while this is easily done with _T. alpestris_ and _T. cristatus_; _T. waltli_ can live in the water for years. The food consists of all kinds of insects, centipedes, worms, snails, etc., which are searched for chiefly at night. It is astonishing to see a little Triton getting hold of and gradually swallowing a wriggling earthworm almost as thick and as long as itself. When two newts seize the same worm, as these voracious and jealous creatures often do, each gets hold of one end, and swallowing as much as it can, twists and rolls round in a direction opposite to that of its rival, until the worm breaks, or until the jaws of the two newts meet and the stronger of the two draws it out of the weaker one and swallows the whole worm. They do not drink, but soak themselves in the water.

The skin is shed periodically, and rather often by the rapidly growing young; by the adult, during the life in the water, rarely during the sojourn on dry land. The skin breaks round the mouth; assisted by the fingers and by contortions of the {124}body, it is then slipped backwards over the trunk and tail, whereupon the newt seizes the skin with the mouth, draws the shirt off entirely, and–swallows it. Such freshly shed skins are very delicate and pretty objects when suspended in water or some preserving fluid. The shed skin, consisting only of the outermost layer of the epidermis, is entire, but turned inside out, with fingers and toes complete, the only holes being those for the mouth, eyes, and vent.

None of the Tritons are viviparous. The eggs, which are glued singly or in small numbers on to stones or water-plants, are hatched in about a fortnight, sooner or later according to the species and the prevailing temperature. The larvae are always provided with three pairs of branched external gills; the fore-limbs appear much earlier than the hind-limbs. Most, perhaps all, larvae develop two pairs of thread-like protuberances on the sides of the upper jaw, by means of which they attach or anchor themselves on to water-plants shortly after they are hatched. Thus moored they remain motionless in a slanting position, now and then wriggling their tails and shifting their place, or sinking to the bottom. The metamorphosis is finished during the first summer, and the little newts, often partially transparent, leave the water to hide under stones. Not unfrequently the metamorphosis is retarded and not finished by the autumn. The larvae of _T. cristatus_, especially when reared in ponds with abrupt or overhanging banks, so that they cannot leave the water, retain considerable remnants of the gills, still more frequently the clefts, although breathing chiefly by the lungs. Such individuals reach a length of 3 inches, and are larvae so far as the finny tail and the gills are concerned. They hibernate in this condition, and in exceptional cases reach sexual maturity;–at least the females, which develop ripe eggs; the males are not known to produce spermatozoa.

Much has been written on the amorous games of newts, but it is only recently that the mode of fecundation has been actually observed. Gasco[60] placed the newts in glass vessels suspended from the ceiling of his laboratory. The antics of the enamoured male around the female, rubbing the latter with its head, or lashing it gently with the tail, and playing around it in its often beautiful nuptial dress, are meant to excite the {125}female. The male then at intervals emits spermatophores, which sink to the bottom, and the female takes them up into its cloaca. For further information see p. 54.

_Triton cristatus._–The Crested Newt has a slightly tubercular skin with distinct pores on the head, on the parotoid region and on a line along the side of the trunk. There is a strong gular fold. The general colour above is dark or black-brown with an olive tinge, interspersed with darker spots; the sides of the body bear irregular white spots. The under parts are yellow, almost always with large black spots. The iris is golden yellow.–The nuptial dress of the male is very striking. A high, serrated crest occurs on the head and body; the upper surface of the head is marbled with black and white; the under parts are orange-yellow with black spots, and the sides of the tail are adorned with a bluish-white band.–The female, always devoid of a crest, generally exhibits a yellow line along the middle of the back.–The average length of fully adult specimens is about 5-6 inches or 13-15 cm.; the females are as usual larger than the males; 144 and 162 mm. for an English male and female respectively are exceptional records.

[Illustration: FIG. 23.–_Triton cristatus._ 1, Female; 2, male in nuptial dress. × ⅔.]

Propagation takes place in April. The newly hatched larvae are yellowish-green, with two black dorsal bands, and with a whitish edge to the tail-fin. By the middle of July they are about 5 cm. long, and the white-margined tail now ends in a {126}thread 1 cm. in length. The general colour above is light olive-brown, dotted with black; the flanks and belly have a golden shimmer.

The Crested Newt has a wide distribution, extending from England and Scotland through Central Europe into Transcaucasia; the northern limits are Scotland and Southern Sweden. Although found in Greece and Lombardy, it does not occur in the Iberian peninsula nor in the South of France, where it is represented by the next following species.

_Triton marmoratus._–The Marbled Newt is of the same size as the Crested Newt. Its ground colour is grass-green above, brown below, with numerous large and small irregularly shaped marbling patches, spots and dots of black. The crest of the neck and trunk is entire, not serrated, adorned with dark vertical bands, and separated from the high dorsal fin of the tail by a deep indenture or gap. The female has an orange line, slightly sunk in, instead of the crest. This newt is confined to France and the Iberian peninsula. In the North of Portugal and in Galicia it is frequently seen in little streams and ponds during the months of March and April. The rest of the year it spends on land. In France occur hybrids of this species and _T. cristatus_. They have been described as _T. blasii_.

_T. alpestris._–The Alpine Newt is easily distinguished by the rich orange colour of its under parts, which are unspotted, excepting a few dark specks across the throat, below the gular fold. Specimens with many ventro-lateral black spots are exceedingly rare. All the upper parts are dark, but vary individually. The prettiest specimens are dark purplish grey, with black marblings; others incline more towards brown ground-tones, the blackish markings then appearing more prominent. The sides are often stippled with tiny whitish dots. The iris is golden yellow.–The nuptial male has a low, not serrated crest, which extends uninterruptedly from the nape into the dorsal fin of the tail. The crest is pale yellow, with black vertical bands and spots. The ground-colour of the upper parts inclines to blue, especially on the sides. The lower fin of the tail assumes an irregular band of bluish-white confluent patches.

This newt is rather small, females rarely exceeding 100 mm. or 4 inches in length. Its home is chiefly the hilly and mountainous parts of Central Europe, from Holland to Lombardy, {127}Austria-Hungary, and Greece. Although it ascends the Alps to between 6000 and 7000 feet, it is also found in the Netherlands, but not in the North German plain.

_T. vulgaris_ (s. _taeniatus_, s. _punctatus_).–The Common or Spotted Newt usually reaches 3 inches (7-8 cm.) in length. Boulenger's record-specimen measured 104 mm. It is characterised by the yellow, partly orange under surface, which is always spotted with black. The upper parts are olive-green or brown, inclining to white on the flanks; the black spots of the back, sides, and especially of the tail, are arranged in more or less distinct lines, giving a somewhat banded appearance to some females.–The breeding dress of the male shows a non-serrated, but "festooned" high and very wavy crest, which extends from the neck without interruption into the likewise wavy tail-fin. The tail is adorned with a lateral, glittering blue stripe, interrupted by vertical dark spots. The larvae are marked by a series of yellow dots, which extend over the lateral line and the tail, which latter temporarily possesses a terminal filament like that of the larvae of _T. cristatus_.

The distribution of the Spotted Newt is the same as that of _T. cristatus_, namely Europe with the exception of the Iberian Peninsula, and Western Asia.

_T. palmatus_ s. _helveticus_.–This is the smallest of all the European newts, rarely reaching more than 3 inches in length. It is distinguished by several specific characters. The tail ends in a thread which is in some males 10 mm. in length, but is only just indicated in the female. The breeding male develops a cutaneous fold along each side of the back, and a low, entire, vertebral crest; the toes are fully webbed. The under parts are pale yellow, inclining to orange towards the middle of the belly, and with a few blackish dots. The lower caudal crest has its edge blue in the male, orange in the female. The general colour of the smooth skin is olive-brown above, with numerous dark spots, which are arranged in more longitudinal streaks on the head.

The Webbed Newt is a native of Western middle Europe, ranging from Great Britain and Northern Spain to Switzerland and Western Germany.

Closely allied to the last species are _T. boscai_ of Spain and Portugal, _T. italicus_, _T. montadoni_ of Moldavia, and the {128}beautiful _T. vittatus_ of Asia Minor. From China and Japan are known _T. pyrrhogaster_ and _T. sinensis_.

The North American species are _T. torosus_ and _T. viridescens_. The former, of Western North America, is one of the largest newts, reaching a length of more than six inches. The head is much depressed and broad, and has very prominent parotoid and other glands. The limbs are strong, especially in the male. The skin of the upper parts is very granular, uniform dark brown, without a crest. The tail, which is larger than the head and body, is strongly compressed, with a low dorsal and ventral fin. The under parts and the lower edge of the tail are uniform yellow or orange red. The iris is green. A specimen in my keeping spends most of its time in the cracks of rotten stumps or on the top of moss in the darkest shade. It lives on earthworms but despises insects. Like most of the other newts it becomes lively at dusk.

[Illustration: FIG. 24.–_Triton viridescens._ 1, Egg just after deposition, with the outer membrane opened, × 6; 2, a spermatophore just discharged showing its gelatinous base with a projecting spike which bears a tuft of spermatozoa, × 2. (After Jordan.)]

_T. viridescens_ is common throughout the Northern and Eastern parts of the United States. Large females are about 11 cm. long, the males 1 cm. less. The general colour above is brown, with a tinge of green; on each side of the trunk, with a row of bright vermilion spots; the under parts are orange, studded with small black dots. Half-grown specimens are brownish red, with the same lateral red spots as the adult. According to Jordan,[61] this voracious species lives chiefly on the larvae of insects, on small molluscs such as _Cyclas_ and _Planorbis_, on earthworms and on small Crustacea. It is eminently aquatic in the adult stage. The eggs are laid from April to June, the period lasting for one individual four to six weeks, or even longer. One female laid 108 eggs in all from 20th April to 30th May. After having selected a suitable plant, for instance an _Anacharis_ or a bunch of _Fontinalis_ leaflets, she bestrides the plant and gathers in the surrounding shoots with her hind-limbs, {129}pressing the leaves closely around the cloaca. She next turns on her side, or occasionally on her back; with fore-limbs outstretched and rigid, with hind-limbs and leaves completely hiding the cloaca, she remains perfectly motionless for six to eight minutes. Then she slowly leaves the "nest," which now holds an egg well protected by a tangle of shoots glued together by the gelatinous secretion poured out of the cloaca. Jordan concludes, from the fact that he never found spermatozoa in the oviducts, that the eggs are fertilised just before they are expelled, when passing the receptaculum seminis.

The metamorphosed young pass their life on land under stones and logs as the so-called red variety, which is merely a stage in the life-history of the species. It seems to take them several years to reach maturity, and to become again typically aquatic. Young, red individuals which I have myself kept, have behaved for more than a year like the young of other newts, spending their time under moss and bark without going into the water.

The change from the red-spotted stage has been exhaustively studied by Gage.[62] He remarks that this species is very common near Ithaca, in an upland forest and along the head-waters of the Susquehannah. The transformation takes place either in the autumn or in the spring, either while the newt is still on land, or after entering the water.

Of two which were kept in a jar with moist wood, one was especially brilliant, but within two weeks it assumed, in the middle of September, the characteristic coloration of the viridescent form. The two specimens were in the jar until the following July, when they were placed where they could enter the water. This they did with great readiness, and they remained submerged for a considerable time at first. The time under water increased in length, until within two or three days the pharyngeal respiration under water was fully established. On the other hand, viridescent specimens never reassume the red garb when kept out of the water.

Red specimens entering the water in the spring, changed into the greenish form within a few weeks, and established the pharyngeal respiration, losing the ciliated oral epithelium. Branchiate larvae and the adult aquatic forms have non-ciliated {130}epithelium, and the cilia are re-established when a green specimen is forced again to live on land. Ciliation always exists in the red stage, and in the green stage before the newt has taken to the water. The cilia sweep towards the stomach.

The three following South European species belong to the _Euproctus_ group, so called on account of the mostly conical, backward directed, and vividly coloured vent.

_T. asper_ s. _pyrenaeus_.–The Pyrenean newt has hitherto been found only in the Pyrenees, for instance in Lac Bleu and Lac d'Oncet, which latter lies about 7000 feet above the level of the sea. According to Bedriaga,[63] it prefers lakes which are supplied during the whole summer with water from glaciers. It is very sluggish, only moving to breathe and when in search of food, which consists of worms and insects. The general colour is greenish brown, dark above; the under side of the head and body are bright orange red in the female, yellow in the male; dark spots separate this bright colour from the flanks. The tail has a narrow ventral stripe of bright red and yellow. The cloaca of the female is bright red, that of the male dull grey. The total length amounts to about 4 inches or 10 cm.

The pairing time is the end of June, or later in cold seasons. The male gets hold of the female by forming a noose with its tail round her; it lies underneath, the cloacae being pressed together so that the spermatozoa can be taken in directly. The larvae have large yellow-green spots on the back and sides, and a bright red ventral tail-fin; when metamorphosed the greenish spots become more confluent on the back, producing a broad spinal band. Larvae which live in deep water are dark, while those in sunny places are light-coloured and spotted with yellow.

_T. montanus_ in Corsica and _T. rusconii_ in Sardinia are allied forms, but the males are distinguished by a spur-like process or dilatation at the end of the fibula.

_T. waltli_, the Iberian Newt, is olive-brown above, yellowish with blackish markings below. The tail has a yellow or orange ventral line. There is no crest. A remarkable peculiarity of this species (which it shares only with _Tylototriton andersoni_ of the Loo-Choo Islands) is its ribs, which are very long, sharply pointed, and frequently perforate the skin. Before {131}perforation the point of the rib lies in a lymphatic space. This surprising feature has by many authorities been considered as abnormal or pathological. Certainly young, and even many adult, individuals are found in which the skin is not perforated, but when these are handled the wriggling motions of this strong newt force the points of the ribs through the skin, and they remain sticking out to the extent of several millimetres. The wounds heal up, the skin forming a neatly finished-off hole through which the spike projects, not as a formidable, but as a sufficiently awkward, protective weapon.

[Illustration: FIG. 25.–_Triton waltli._ Spanish Newt, adult and larvae. × ⅔.]

Large females reach a length of 10 inches. The larvae metamorphose, as a rule, when they are between 2 and 3 inches long, but those which have been bred in tanks often reach double this length. These newts are frequent inhabitants of the rain-water cisterns common in the South of Portugal and Spain, into which they tumble without ever being able to get out again. This species spends most of its time in the water, {132}preferring ponds, among the vegetation of which they can be watched lying motionless, with their limbs hanging down and with the head close to the surface; but they are lively during the night. When their ponds dry up they leave them, crawling into the most unexpected places, to aestivate under rocks, or even in the walls of old buildings, where they are found by accident only. The range extends from Central Spain and Portugal into Morocco.

_Tylototriton verrucosus_ lives in the Eastern Himalayas and in the mountains of Yunnan. The skin is tubercular, with large parotoids; above uniform black-brown, pale below; the tail has a ventral yellow or orange line. Total length about 6 inches. _T. andersoni_ of the Loo-Choo Islands is remarkable for the pointed ribs which perforate the skin.

_Pachytriton brevipes_, discovered in Kiansi, Southern China, has a smooth skin, olive-brown above, with many black dots; the under parts are yellowish, dotted with black. Total length about 7 inches.

FAM. 3. PROTEIDAE.–The three pairs of fringed external gills persist throughout life. Both fore- and hind-limbs are present. The eyes are devoid of lids. The maxillaries are absent. Teeth are present on the premaxillaries, on the vomers, and on the mandible. The vertebrae are amphicoelous.

This family consists of only three genera, with one species in each.

_Necturus maculatus_ s. _Menobranchus lateralis_.–The eyes are functional, being covered by the thin transparent skin. The limbs, although short, are well developed, and have four fingers and four toes. The whole animal, which reaches the length of one foot, is quite smooth and slimy, brown with irregular dark, blackish spots and patches, which frequently form a dark lateral band extending from the mouth to the tail. The latter, which measures about one-third of the whole length, is strongly compressed, carries a thick dorsal and ventral fin, and is rounded off at the end. The skin of the throat forms a strongly-marked transverse fold. The thick stalks of the gills are brown, while the numerous and delicate fringes are dark red in life; beneath and behind them are two gill-clefts. _N. maculatus_ is found in the eastern half of the United States, chiefly the eastern part of the basin of the Mississippi and the Canadian lakes.

{133}These creatures are rather dull; they remain mostly at the bottom of the water, more or less concealed in the weeds or between rocks during the daytime. Mine, which are kept in a roomy, light-coloured tank, lie motionless, with their gills spread out transversely. Every now and then the gills contract suddenly and become pale, whereupon they are filled again with blood. Very rarely they rise to the surface, but tiny air-bubbles are let out more frequently, especially when the animals are disturbed. Then the gills collapse, are laid flat against the neck, and the creature darts about with quick, eel-like motions. At night they leave their hiding-places, swim about or creep along the ground with slow, undulating movements, the limbs being scarcely used, in search of food, which in their wild state consists of rather large Crustacea, small fishes, worms, insects and frogs. They are most voracious, and absolutely indifferent to cold. The spawning takes place in the months of April and May.

_Proteus anguinus._–The fore- and hind-limbs are fully developed, but possess only three fingers and two toes. The eyes are completely hidden beneath the opaque skin. This peculiar creature is restricted to the subterranean waters of Carniola, Carinthia, and Dalmatia. The vast caves of Adelsberg not far from Trieste are especially celebrated for the occurrence of the "Olm," the German name of this animal. The river Poik, a moderate mountain-stream, but a large, fierce torrent during the rainy season, disappears into the limestone-hills, and rushes through enormous stalactite-grottoes, most of which have been only partially explored, until several miles farther on it reappears on the surface. There, deep down below the surface, in absolute darkness, in an almost constant temperature of about 50° F. is the home of _Proteus_.

Their total length is scarcely one foot. The whole body is white, occasionally suffused with a slight fleshy, rosy tinge, while the three pairs of gill-bunches are carmine-red. They are easily kept in captivity, and live for many years, provided three conditions are strictly adhered to, viz. fresh and clean water, an equable low temperature of about 50° F. = 10° C. and darkness. The question of food is not so very important, since specimens are known to have existed for years, although they refused to take any nourishment. How far darkness is an {134}absolute necessity is not known. Anyhow, the white skin is almost as susceptible to light as is a photographic plate. If light is not absolutely excluded the white skin becomes in time cloudy, with grey patches, and if kept exposed to stronger light, the whole animal turns ultimately jet-black. Mr. Bles has succeeded in producing several totally black specimens, having kept them for several months in a white basin under ordinary conditions of light. No experiments have yet been made to find out if the black pigment deposited is lost again in darkness. Those which are kept in a tank in an absolutely dark cellar of the Cambridge Museum, with permanent water-supply, are doing very well. When approached with a candle they become restless or remain partly hidden in all sorts of seemingly most uncomfortable attitudes, squeezed in between the sharp-edged tiles and drain-pipes with which their lodgings are furnished. But the introduction of a wriggling worm, a little crustacean or other live bait draws them from their hiding-places, and, guided by the motions of the prey in the water, possibly also by the sense of smell, they snap it up and devour it.

[Illustration: FIG. 26.–_Proteus anguinus._ × ⅔. Front view of the mouth in the left upper corner.]

If the water is not sufficiently well aerated, they rise to the surface, emit a bubble of air, and take a new supply into their lungs. As a rule they remain motionless under water, but the gills contract spasmodically and become paler, whereupon they fill again with blood and darken; the contrast between the pure white body and the carmine-red feathery gills is very beautiful.

Until recently the mode of propagation was quite unknown. Several _Proteus_, kept by E. Zeller, laid, in the middle of April, {135}a number of eggs which were then fastened singly on to the under side of projecting stones in the water. The pale yellow yolk measured 4 mm. in diameter and was surrounded by a cover of 1 mm. in thickness, besides an outer gelatinous mantle, so that the whole egg measured about 11 mm. The larvae were hatched after 90 days; they were 22 mm. long, and already much like the adult, except that the fin was not restricted to the tail, but extended over the last quarter of the trunk, and that their eyes were still visible. The fore-limbs were already typical in shape, but the hind-limbs were still toe-less little stumps.[64]

_Typhlomolge rathbuni._–It is of the greatest interest that a subterranean Perennibranchiate newt, in many respects closely resembling _Proteus_, has recently been discovered in Texas. There can be no doubt that similar conditions of life have produced these two forms from _Necturus_- and _Spelerpes_-like ancestors,[65] one in Europe, the other in North America, absolutely independently of each other. The limbs of _Typhlomolge_ are long and very slender, the four fingers and five toes are thin, free and pointed. The head is large, the mouth square. The eyes are completely hidden and the whole animal is colourless and white. The tail is furnished with a dorsal and a ventral fin. The very deep gular fold is nothing but the pair of united but large opercular flaps. The three pairs of gills are remarkable for their blade-like stalks, while the gill-lamellae proper are short and restricted to the tapering ends. Total length about 75 mm., of which the head measures 15, the tail 32 mm.

This peculiar creature inhabits subterranean caves in Texas, to judge from the fact that all the specimens hitherto known have come up with the water of an artesian well 188 feet deep, near San Marcos. According to Blackford,[66] "the legs are used for locomotion and the animals creep along the bottom of the aquarium with a peculiar movement, swinging the legs in irregular circles at each step. They climb easily over the rocks piled in the aquarium, and hide in the crevices between them. All efforts to induce them to eat have been futile, as has also been the case with blind cave-fish in captivity, and they are {136}either capable of long fasts or live on infusoria in the water." It seems more reasonable to suppose that these newts live upon Crustacea, four kinds of which, all new to science, also came up with the water.

FAM. 4. SIRENIDAE.–The three pairs of fringed external gills persist throughout life. The body is eel-like. Hind-limbs are altogether absent, while the fore-limbs are short and have three or four fingers. The maxillary bones are absent. With the exception of small teeth on the vomer the mouth is toothless, but the jaws are furnished with horny sheaths. The eyes are devoid of lids, but shine through the skin.

[Illustration: FIG. 27.–_Siren lacertina._ × ½.]

The Sirenidae are the most degraded members of the Urodela and are represented by two closely-allied genera, each with one species, in the south-eastern parts of the United States. Their most interesting feature, which bears upon the question of neoteny, is their retrograde metamorphosis as described by Cope.[67] The gills atrophy in the young and are subsequently redeveloped. Cope therefrom concludes rightly that the ultimate or persistent gills of _Siren_ are signs of maturity and not a larval character. In young specimens of _Siren_ of 5 to 6 inches in length the gills are functionless; in one of 3 inches they were found to be entirely vestigial and "subepidermal," _i.e._ covered by a common dermal investment. Unfortunately really young larvae are still unknown. Old Sirens can live without gills, as has been shown by aquarium-specimens. In the adult _Pseudobranchus_ all the gills are normally covered up by an investment of the skin so as to be quite without function and movability.

_Siren lacertina_, the "mud-eel," is distinguished by the {137}possession of three pairs of gill-clefts and by its four fingers. It reaches a length of 70 cm., or about 2½ feet, of which about one-third is taken up by the tail, which is strongly compressed and finned. The skin is smooth, mostly blackish, lighter below, sometimes with whitish specks all over the body. This creature is frequently found in ditches and ponds, where it burrows in the mud. When swimming the limbs are folded back. They are said sometimes to leave the water and to crawl about on the moist ground.

_Pseudobranchus striatus_ has only one pair of gill-clefts and only three fingers. The slightly granular skin is dusky brown above, with a broad yellow band on either side and with a paler, narrower stripe below. Total length about 7 inches.

{138}