Comparative anatomy and phylogeny of the cloacae of salamanders (amphibia: Caudata). III. Amphiumidae.

Histological descriptions of female and male cloacae are presented for the three species of Amphiuma. Little interspecific variation occurs in either females or males. Females possess numerous spermathecae around the walls of the posterior cloacal tube and anterior cloacal chamber as well as rudimentary vent glands, of uncertain homology, in the caudal end of the cloacal chamber. Ventral glands such as those known from female salamanders in other families are absent, likely a secondary loss in the amphiumid clade. In a female A. tridactylum with large ovarian follicles (5- to 8-mm diameter), the lips of the cloacal orifice are highly cornified, and stored sperm in the spermathecae appear in organized bundles, either free in the lumen or with the sperm heads seemingly buried in the spermathecal epithelium. Males possess some gland clusters present in other members of the suborder Salamandroidea (dorsal pelvic glands, vent glands, Kingsbury's glands, anterior ventral glands) but lack others (lateral pelvic glands, posterior ventral glands), resulting in the most plesiomorphic cloacal anatomy in the suborder. However, males possess an autapomorphic gland cluster, called the pit glands, in the posterior wall of the cloacal chamber. The pit glands consist of shallow infoldings of the cloacal epithelium unlike other cloacal glands, which are relatively tubular and elongate. The "pits" contain columnar eosinophilic cells that give positive staining reactions for acid mucopolysaccharides and proteins.