Fig. 28.–Map showing distribution of Aglossa. Hymenochirus to be added in Equatorial Africa.
The pharyngeal opening and the tubes themselves are wide, the tympanic cavities are present, but the tympanic discs are not distinct from the rest of the skin. The fronto-parietal bones are fused into one mass, a rare feature in the Anura. The nasals are large. Pipa and Hymenochirus have no teeth, Xenopus has teeth on the upper jaw. The vertebrae are opisthocoelous and typically epichordal in their development; the second, third, and fourth carry long ribs, which in old specimens fuse with the supporting diapophyses. The sacral diapophyses are enormously dilated, and the sacrum is fused with the os coccygeum. The serial number of the sacral vertebrae exhibits a most interesting gradation. In Xenopus the ilium is carried by the diapophyses of the 9th, in Pipa the 9th and 8th, in Hymenochirus the 7th and 6th. In these cases the two diapophyses of each side are fused together into a single broad blade, and their original duplicity is indicated only by the holes for the spinal nerves. Hymenochirus has consequently only 5 presacral vertebrae, the vertebral column being shortened to the greatest extent known amongst Vertebrata. For further information see p. [22]. The ilia are much broadened vertically, and are firmly attached to the sacrum. The shoulder-girdle is sometimes described as of the arciferous type, but this is quite unjustifiable. The epicoracoid cartilages do not overlap each other, but meet, and partly fuse in the middle line. The three genera exhibit some differences. In Pipa and Hymenochirus the bony portions of the coracoids are much expanded dorsally, and there is a considerable amount of epicoracoid cartilage, that of the precoracoid bars extending backwards as a broad-based and blunt omosternum. Xenopus is devoid of an omosternum, and the configuration of the whole apparatus is more slender. The metasternum of Xenopus and Hymenochirus broadens out laterally. Hymenochirus greatly resembles Breviceps, a genus of Engystomatinae, in the relative position and size of the various parts of the shoulder-girdle and sternum.
The tibio-fibula of Hymenochirus has a wing-like expansion of thin bone on each side, forming a deep groove on the outer aspect. The astragalus and calcaneum are united by a similar bony expansion with wing-like projections.
The lungs are remarkable for the prominent development of trabecular projections and niches, so that their free lumen is much restricted; they have thereby reached a much higher stage than in any other Amphibia or even many Autosauri. The persistence of an arteria sacralis s. caudalis, a vessel absolutely absent in the adult Rana, is a primitive feature, and the same applies to the presence of a true first spinal or suboccipital nerve.
The skin of the back and belly is supplied by two great branches from the arteria anonyma, one arising proximally, the other distally from the subclavian; herewith is correlated the almost complete absence of the arteria cutanea magna, which as a branch of the ductus pulmo-cutaneus plays such a prominent rôle in the other Anura. Only in Pipa, but not in Xenopus, is the great cutaneous vein represented by a very small branch. Both these genera possess a much more complicated "diaphragm" than the other Anura, chiefly owing to a special muscle which arises from the anterior end of the ilia and spreads out fan-like to the oesophagus and to the bases of the lungs.[[71]] This diaphragmatic arrangement is correlated with the great development of the lungs, and is not a primitive but an advanced feature. It is reasonable to suppose that this has caused the reduction of the usual arteria pulmo-cutanea, and that the other two cutaneous arteries have been developed secondarily. The Aglossa are generally considered as the lowest Anura, and only Cope looked upon Pipa and Xenopus as two convergent terminal branches. Beddard came to the conclusion that both are closely related to each other, chiefly on account of their peculiar diaphragmatic arrangement. The whole question has entered upon a new stage since the recent discovery of Hymenochirus, which is in many ways intermediate between the two other genera. Moreover, the mid-Tertiary Palaeobatrachus of Europe is undoubtedly related to them, and we conclude now that all these four genera belong to one group with a distribution formerly much wider than Africa and part of South America. But this does not necessarily mean that the Aglossa are in all respects the most primitive group of living Anura. On the contrary, they possess few decidedly primitive characters, namely, the long typical ribs, the presence of the first spinal nerve, the unimportant persistence of the arteria sacralis, and lastly, the possession in the tadpoles of a right and left opercular "spiracle." The absence of the tongue cannot possibly be an archaic feature, considering its universal presence in all the other Amphibia, including the Apoda, and the suggestive circumstance that this organ is least developed in the entirely aquatic members of the Urodela. In fact, thoroughly aquatic creatures, which seize and swallow their prey under water, require no elaborate tongue; and since we know that the Anura must owe their typical formation to terrestrial life, it follows that those which have again taken to the water and are tongueless, have lost this organ. As I have shown elsewhere,[[72]] the epichordal development of the vertebrae is likewise a secondary feature, far from primitive; and the tendency of the shortening of the vertebral column, which has reached its extreme in Hymenochirus, points to the same conclusion. The apparatus of the shoulder-girdle and sternum is in the last transitional stage from the former arciferous to the typically consolidated firmisternal type. In fact there is little left which is primitive, but much that is very specialised and highly developed in the Aglossa, mostly in adaptation to their absolutely aquatic life, to which they must however have taken very early. They are in a position somewhat analogous to the Ratitae among Birds, which are likewise an old group, although many of their most striking features have been acquired secondarily.
Xenopus s. Dactylethra. The upper jaw is furnished with teeth. The ilia are attached to the ninth vertebra. The pupil is round. The terminal phalanges are pointed. The fingers are free, the toes broadly webbed, and the first three are covered with sharply pointed, horny, black-brown nails, a feature which is alluded to by the alternative generic names. A cutaneous tentacle projects from below the eye and naturally invites comparison with the tentacle of the Apoda and of Urodela. The skin is smooth, rich in mucous glands, besides certain tube-like apparatuses, possibly sensory, which are scattered over the body, especially on the head, and form a conspicuous series of white dots along the dorso-lateral line, from the eye to the vent. The general colour of the upper parts is olive brown, mottled darker, while the under parts are whitish. The female has three cutaneous flaps closing the vent. The male develops black nuptial brushes along the inner side of the fingers. There are several species, all African (Ethiopian).
X. laevis, ranging from the Cape to Abyssinia, is distinguished by the absence of a metatarsal spur. The tentacle is very short. Size about 3 inches. X. muelleri of Zanzibar and Benguella, is smaller. The tentacle is conspicuous, as long as the diameter of the eye. The inner metatarsal tubercle carries a sharp claw. X. calcaratus of tropical West Africa is only 2 inches long, and has strong metatarsal claws, short tentacles and very minute eyes.
The habits and oviposition of the "Clawed Toad" have been described by Leslie.[[73]] The Boers call it "Plathander," i.e. flat hand. Entirely aquatic, it rests floating in the water, with the nostrils exposed, and leaves the water only if it has to change the locality on account of drought or scarcity of food. The pairing takes place, at least at Port Elizabeth, in the early spring, i.e. in the month of August. The only sound which is emitted is heard during this time, a very slight and dull tick-tick, audible at only a few feet distance. The male grasps the female by the loins; the eggs are extruded singly, measuring only 1.5 mm. in diameter, but swell to double that size. They are attached singly to stones or water-plants.
Fig. 29.–Xenopus laevis. Clawed Toad, adult and larvae, × ⅔.