Comparative anatomy and phylogeny of the cloacae of salamanders (Amphibia: Caudata). II. Cryptobranchidae, Hynobiidae, and Sirenidae.

Cloacae were examined from salamanders representing the three families in which fertilization of eggs is known or inferred to occur externally. The cloacae of male and female sirenids are aglandular and lack cilia. Sexual dimorphism in sirenid cloacae occurs only in the extent of epithelial stratification in the cloacal chamber in females (entire chamber) versus males (posterior angle of the vent). Both male and female Cryptobranchus alleganiensis possess ventral glands that secrete an acid mucopolysaccharide and have ciliated cloacal linings. The ventral glands are more numerous and hypertrophied in breeding male than female C. alleganiensis, but in males, ventral glands secrete only onto the surface of the cloacal lips along the anterior three-fifths of the cloacal orifice, whereas in females, the glands secrete onto the border of the entire cloacal orifice. Except for male Onychodactylus japonicus, male and female hynobiids also possess only ventral glands and have ciliated cloacal linings. Hynobiid ventral glands secrete a glycoprotein. Much variation occurs, however, among these hynobiids in cloacal conformation, extent of epidermis into the cloaca, and anatomy of the ventral gland. Male O. japonicus possess an unciliated cloaca in which three types of cloacal glands occur, each giving unique reactions to tests for carbohydrates and proteins. The glands in male O. japonicus do not seem to be homologous to those found in spermatophore producing salamanders in the Salamandroidea, but this does not negate the possibility that O. japonicus makes spermatophores. Examination of cloacal characters in additional species of hynobiids may be useful in resolving intrafamilial phylogenetic relationships.