CHAPTER XXV

BRANCH CHORDATA (_Continued_). CLASS BATRACHIA: THE BATRACHIANS

The structure, life-history, and habits of the garden-toad (_Bufo lentiginosus_) have already been studied (see Chapter II and Chapter XII).

OTHER BATRACHIANS.

The class Batrachia includes the animals familiarly known as cœcilians, sirens, mud-puppies, salamanders, toads, and frogs. Although differing plainly from fishes in appearance and habits, the batrachians are really closely related to them, resembling them in all but a few essential characters. Among the distinctive characters of batrachians may be noted the absence of fins supported by fin-rays, the presence usually of well-developed legs for walking or leaping, and the absence or reduction of certain bones of the head connected with the gills and lower jaw and which are well developed in the fishes. The batrachians stand in somewhat intermediate position between the fishes and the reptiles, showing some of the characters of both. They are, like fishes and reptiles, cold-blooded. In their adult condition some are terrestrial and some aquatic as to habitat, but all have an aquatic larval life. The water-inhabiting young breathe at first by means of gills, later lungs begin to develop, and for a time both gills and lungs are used in respiration. Finally in the adult condition in almost all of the forms the gills are wholly lost and breathing is done by the lungs and skin solely. Correlated with the change of habits from larval to adult stage there is usually a well-marked metamorphosis in post-embryonic development. This metamorphosis is specially striking among the frogs and toads. None of the aquatic forms is marine, salt water always killing eggs, larvæ or adults. Batrachians are found all over the world, although there are few in the extreme North. They are most abundant in warm and tropical lands.

=Body form and organization.=--The body varies from a long and slender, truly snake-like form as in the tropical cœcilians through the usual salamander (fig. 119) shape, where it is more robust but still elongate and tailed, to the heavy, squat, tailless condition of the toads. Legs, with five digits, are usually present, and are used for swimming, walking, or leaping. The legs are longest and best developed in the short tailless frog and toad forms which are mostly terrestrial, and are short and weak in the tailed salamander forms, many of which are aquatic. The skin is almost always naked, showing a marked difference from the scaled condition of reptiles and most of the fishes, and its cells secrete a slimy, sticky, usually whitish fluid, which in some cases is irritating, or even poisonous. The skin is sometimes thrown up into folds or ridges, and in some species is elevated to form a kind of fin on the tail or back. This unpaired fin differs from the dorsal fin (and other fins) of fishes in not being supported by rayed processes of the skeleton. There are in some batrachians traces of an exoskeleton in the presence of scale-like structures in the skin or in the horny nails on the digits, but these cases are rare. The skin contains pigment-cells and many of the batrachians are brilliantly colored and patterned; some of the pigment is carried by special contractile or expansile cells, the chromatophores (see account of chromatophores of the Cephalopoda, p. 256), so that the animal can change its tint and markings more or less rapidly. All the batrachians possess external gills in their aquatic larval stage, and in a few forms, as the sirens and mud-puppies, gills are retained all through life. These gills are branched folds of the skin abundantly supplied with blood-vessels.

In the organization of the batrachian body the usual vertebrate characters appear, the body-organs being arranged with reference to a supporting and protecting internal bony skeleton. The head is plainly set off from the rest of the body and bears the mouth and the organs of hearing and sight. Certain so-called lateral sense organs, the function of which is not exactly known, occur arranged in three lines on each side of the body of some of the forms. Both pairs of limbs are present and functional in almost all of the species. In the cœcilians the limbs are wholly wanting; in the sirens only the fore legs are present.

=Structure.=--The most obvious skeletal differences among batrachians are those due to variations in external form. While there are as many as 100 vertebræ in some of the elongate long-tailed salamanders (even 250 in the strange snake-like cœcilians), there are but 10 (the last or tenth being the rod-shaped bone called the urostyle) in the short, tailless frogs and toads. To any of the vertebræ except the first (the single cervical vertebra) and the last, ribs may be attached and the cœcilians have about as many pairs of ribs as vertebræ. In the frogs and toads, however, the ribs are lost. In any case they are never fastened by their lower ends to the breast-bone.

The alimentary canal is usually not much longer than the body and is plainly divided into mouth, pharynx, œsophagus, small intestine, large intestine or rectum, and anal opening. The teeth when present occur on both the jaws and the palate. They are small, sharp, point backward and are fused to the bones. They are wholly wanting in the toad and in some other allied forms. The tongue may be wanting, or may be immovably fixed to the floor of the mouth, or as in the frogs, fastened at its front end but free behind, so that the hinder end can be protruded far from the mouth for the purpose of catching insects.

The organs of respiration are gills, external and internal, lungs, trachea or windpipe, and the skin. In the earliest larval stages all batrachians have gills; later, in most cases, the gills become reduced and disappear, while at the same time lungs are developing. In some salamanders the lungs never develop, but the animals, in their adult stage, breathe wholly by means of the skin. In a few cases, as in the siren and mud-puppies, gills are retained through the whole life, although lungs are also present in the adult stage. The lungs are two in number, a right and a left lung, and are simple sacs with the walls more or less folded or thrown into ridges and richly supplied with blood-vessels. The front end of the lungs opens directly into the pharynx or, in the more elongate batrachians, is connected with it by a tubular trachea or windpipe. In the frogs and toads there are vocal cords stretched across the short windpipe; the vibration of these cords produces the croaking.

The heart is always three-chambered, consisting of the right and left auricles and a single ventricle. The circulation of the more generalized salamanders like the mud-puppies is essentially like that of a fish. In the frogs and toads there is a distinct advance beyond this condition. The red corpuscles of the blood are oval in shape and are the largest found among any of the vertebrates.

In the nervous system the small size of the hindbrain or cerebellum is noticeable. The sense organs are fairly well developed. The skin of the whole body is provided with tactile nerve-endings. There are special taste organs on the lining membrane of the tongue and mouth-cavity. The eyes have no lids in some of the lower forms; most of the frogs and toads have an upper lid but no under one, although a thin membrane, called the nictitating membrane, arises from the lower margin of the eye and can be drawn up over it. The ears have no external parts, other than the thin tympanic membranes. The nostrils of frogs and toads can be closed by the contraction of certain special muscles.

=Life-history and habits.=--The sexes are distinct, and in most cases the young hatch from eggs. A few of the salamanders give birth to free young. The eggs are usually in strings or chains enclosed in a clear gelatinous substance; these chains of eggs are either simply dropped into the water or are fastened to water-plants. The young, called tadpoles (fig. 120), in their earlier larval stages are extremely fish-like in character, long-bodied, tailed, swimming freely about by means of the fin-like flattened tail, and breathing by means of external gills. Nor do they show any sign of legs. As the tadpoles grow and develop the legs begin to appear, the hind legs first in the frogs and toads, the fore legs first in the salamanders; lungs develop and the gills disappear (except in the cases of the few forms which retain gills through life). The tail shortens and finally disappears in the frogs and toads; with the salamanders the tail-fin only is lost. At the same time the change from water to land is made. Further growth is very slow; frogs are not really adult, that is, capable of producing young, until they are five years old, and they may continue to increase in size until they are ten years old.

The food of the adult batrachians is almost exclusively small animals, particularly insects and worms. Crustaceans, snails, and young fish are also eaten. The tadpoles also eat vegetable matter. Almost all batrachians are nocturnal in habit, remaining concealed by day. In the zones in which cold winters occur they hibernate or pass the winter in a torpid condition, or state of "suspended animation," or, as it is said, they sleep through the winter. Frogs burrow into the mud at the bottom of ponds at the approach of winter and come forth early in the spring to lay their eggs. Most batrachians are very tenacious of life, being able to withstand long periods of fasting and serious mutilation, and most of them can regenerate certain lost parts, such as the tail or legs.

=Classification.=--The living Batrachia are divided into three orders, viz., the Urodela, including the sirens, mud-puppies, salamanders, and newts, batrachians which retain the tail throughout life, having generally two pairs of limbs of approximately equal size, and sometimes possessing gills or gill-slits in the adult condition; the Anura, or frogs and toads, with no tail in the adult condition, with short and broad trunk, with hind limbs greatly exceeding the fore limbs in size, and never with gills or gill-slits in the adult stage; and the Gymnophiona, or cæcilians, snake-like batrachians having neither limbs nor tail, with a dermal exoskeleton and without gills or gill-slits in the adult.

=Mud-puppies, salamanders, etc. (Urodela).=--TECHNICAL NOTE.--If possible obtain specimens of mud-eels (_Siren_), common in the South, or mud-puppies (_Necturus_), common in the central North, as examples of batrachians with gills persisting in the adult stage. One or more species of _Amblystoma_ may be found in almost any part of the country, and larvæ of large size may be found with the external gills. For an example of the general long-tailed or Urodelous type of batrachian any salamander or newt occurring in the vicinity of the school may be used. The little green triton or eft (_Diemictylus viridiscens_) of the eastern States, or its larger brown-backed congener of the Pacific coast (_D. torosus_) is common in water, while another eft, the little red-backed salamander, (_Plethodon_) is common in the woods under logs and stones. The external characters of the body should be compared with those of the toad. The skeleton should be prepared by macerating away the flesh (for directions, see p. 452), and the presence of the many caudal vertebræ and the ribs, the equality in size of the legs, and other points should be noted. Compare with skeleton of toad. Make drawings. It will be well, also, to dissect out and examine the various internal organs of the salamander, comparing them with the same organs in the toad. The salamander, indeed, is in many ways better than the toad as an example of the class. Its body is less adaptively modified and shows the essentially fish-like character of the batrachian structure.

The batrachians which retain external gills in the adult stage are the members of two families of which the American representatives are known as mud-eels (_Siren_) and mud-puppies or water-dogs (_Necturus_). The mud-eels, which are found "in the ditches in the swamps of the southern States from South Carolina to the Rio Grande of Texas and up the Mississippi as high as Alton, Illinois," are blackish in color, have no hind legs and are long and slender, with the tail shorter than the rest of the body. They reach a length of nearly three feet. The mud-puppies, found in the Great Lakes and in the rivers of the upper Mississippi valley, are brown with colored spots, and are about two feet long when full grown. They have both fore and hind legs.

A few salamanders, while not possessing external gills when adult, have a spiracle or small circular opening in the side of the neck which leads into the throat. The best-known American salamander of this kind is the large heavy-bodied blackish water-dog or "hellbender" (_Cryptobranchus_) of the Ohio River. It is about two feet long, and is "a very unprepossessing but harmless creature." It has a conspicuous longitudinal fold of skin along each side of the body. The largest known batrachian, the giant salamander of Japan (_Megalobatrachus_), reaching a length of three feet, is related to the water-dog.

Of all the salamanders the most interesting are the blunt-nosed salamanders (_Amblystoma_). A dozen or more species of _Amblystoma_ occur in North America, of which _tigrinum_, a dark-brown species with many irregular yellow blotches sometimes arranged in cross-bands, is the most widespread. The larvæ of some _Amblystoma_ retain their gills until they have reached a large size, and in one or two species the usual metamorphosis is very long delayed and the salamanders produce young while in the larval condition, that is, while retaining the gills and a compressed fin-like tail. In the case of a certain Mexican species (_A. maculatum_) it is believed that the final metamorphosis never occurs. The Mexicans call these gilled larval _Amblystoma_ axolotls, and use them for food. For a long time naturalists supposed the _Amblystoma_ larvæ which produce young to be the adults of a species of salamanders which retained their gills through life, like the sirens and mud-puppies, and classified them in a distinct genus.

Of the various common salamanders or newts some are found in streams, ponds, and ditches, and some under logs and stones in the woods. The aquatic forms have the tail compressed (flattened from side to side), while the land forms have the tail cylindrical, tapering to a point. Most of the land-salamanders produce their young alive, while the water forms lay eggs which are usually attached to a submerged plant-stem. The salamanders are, almost without exception, found only in the northern hemisphere.

=Frogs and toads (Anura).=--There are about a dozen species of frogs in the United States. The largest of these, and indeed the largest of all the frogs, is the well-known bullfrog (_Rana catesbiana_), which reaches a length (head to posterior end of body) of eight inches. It is found in ponds and sluggish streams all over eastern United States and in the Mississippi valley. It is greenish in color with the head usually bright pale green. Its croaking is very deep and sonorous. The pickerel-frog (_R. palustris_), which is bright brown on the back with two rows of large oblong square blotches of dark brown on the back, is found in the mountains of eastern United States. The little pale reddish-brown wood-frog (_R. sylvatica_) with arms and legs barred above is common in damp woods and is "an almost silent frog." The peculiar and infrequently seen frogs known as the "spade-foots" (_Scaphiopus_) are subterranean in habit and usually live in dry fields or even on arid plains and deserts. They pass through their development and metamorphosis very rapidly, appearing immediately after a rain and laying their eggs in temporary pools. At this time of egg-laying they utter extraordinarily loud and strange cries. Some frogs in other parts of the world live in trees, and the eggs of one species are deposited on the leaves of trees, leaves which overhang the water being selected so that the issuing young may drop into it.

The true tree-frogs or tree-toads (Hylidæ) constitute a family especially well represented in tropical America. They have little disk- or pad-like swellings on the tips of their toes to enable them to hold firmly to the branches of the trees in which they live. Some, like the swamp tree-frog and the cricket-frog, are not arboreal in habit, remaining almost always on the ground. The common tree-frog of the eastern States (_Hyla versicolor_) is green, gray, or brown above with irregular dark blotches, and yellow below. It croaks or trills, especially at evening and in damp weather. Pickering's tree-frog (_Hyla pickeringii_) makes the "first note of spring" in the eastern States. This tree-frog is the one most frequently heard in the autumn too, but "its voice is less vivacious than in the spring and its lonely pipe in dry woodlands is always associated with goldenrods and asters and falling leaves." The tree-frogs of North America lay their eggs in the water on some fixed object as an aquatic plant, in smaller packets than those of the true frogs, and not in strings as do the toads.

The toads (Bufonidæ) differ from the true frogs in having no teeth and in not having, as the frogs do, a cartilaginous process uniting the shoulder-bones of the two sides of the body. The absence of this uniting process makes the thoracic region capable of great expansion. There are only a few species of toads in North America, but one of these species, the common American toad (_Bufo lentiginosus_), is very abundant and widespread. It appears also in two or three varieties, the common toad of the southern States differing in several particulars from that of the northern. The toad is a familiar inhabitant of gardens, and does much good by feeding on noxious insects. It is most active at twilight. Its eggs are laid in a single line in the centre of a long slender gelatinous string or rope, which is nearly always tangled and wound round some water-plant or stick near the shore on the bottom of a pond. The eggs are jet black and when freshly laid are nearly spherical. At the time of egg-laying the toads croak or call, making a sort of whistling sound and at the same time pronouncing deep in the throat "bu-rr-r-r-r." The toad does not open its mouth when croaking, but expands a large sac or resonator in its throat. The toad-tadpoles are blacker than those of frogs or salamanders, and undergo their metamorphosis while of smaller size than those of frogs. When they leave the water they travel for long distances, hopping along so vigorously that in a few days they may be as far as a mile from the pond where they were hatched. They conceal themselves by day, but will appear after a warm shower; this sudden appearance of many small toads sometimes gives rise to the false notion that they have fallen with the rain.

=Cœcilians (Gymnophiona).=--The third order of batrachians, the cœcilians, includes about twenty species of slender worm- or snake-like limbless forms which are confined to the tropics. Some of them are wholly blind and the others have only rudimentary eyes. In them the skin is folded at regular intervals so that the body appears to be rigid or segmented, and in some species there are small concealed horny scales in the skin.