Ultrastructure of the annual cycle of female sperm storage in spermathecae of the torrent salamander, Rhyacotriton variegatus (Amphibia: Rhyacotritonidae)

This study is the first report on the ultrastructure of the sperm storage glands (spermathecae) in the salamander Rhyacotriton variegatus. The population studied is associated with cold-water, rocky streams of the redwood (Sequoia) zone in northern California. Males possess sperm in their vasa deferentia and undergo spermiation throughout the year, but mating is seasonal. Most females with large, vitellogenic follicles (2.0-3.9 mm mean dia.) collected from February-June contain sperm in their spermathecae, although some females with large follicles lack sperm. Other mature-size females collected during this period have small ovarian follicles (0.9-1.2 mm mean dia.) and lack stored sperm. All females collected from September-November have small follicles (0.6-1.6 mm mean dia.) and lack sperm, except in one instance in which a female collected in November had a small amount of degraded sperm, apparently retained from the previous breeding season. The spermathecae consist of simple tubulo-alveolar glands in which the neck tubules produce a mucoid secretory product, and the distal bulbs, where sperm are stored, contain secretory vacuoles of uniform density that stain positively for glycosaminoglycans. In specimens containing sperm, some bulbs have abundant sperm and others lack sperm, but the ultrastructure is similar in both conditions. The acini contain columnar epithelial cells with wide intercellular canaliculi, and a merocrine process releases the secretion. Spermiophagy occurs. In specimens from spring and summer with small ovarian follicles, the neck tubules are similar to those of breeding females, but the distal bulbs are reduced to cords of cells lacking a discernible lumen. Secretory activity in the distal bulbs is initiated in the fall. Spermathecae of R. variegatus are most similar to those of a stream-dwelling plethodontid, Eurycea cirrigera.