Derived sperm morphology in the interstitial sea cucumber Rhabdomolgus ruber, with observations on oogenesis and spawning behavior


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Some life history features of the interstitial sea cucumber Rhabdomolgus ruber are described from intertidal specimens collected from the northern coast of Maine. Histological studies suggest that the population consists of hermaphrodites with gametogenesis being initiated in April and reproduction beginning in May and continuing through the summer months. Sexually mature adults possess a single, blind-ended gonadal tubule that functions as an ovotestis by producing both eggs and sperm. The ovotestis wall consists of an outer peritoneum composed of flagellated epithelial cells and muscles; an inner germinal epithelium of germ and somatic cells; and a middle connective tissue (hemal) compartment bounded by the basal laminas of the peritoneum and germinal epithelium. During the reproductive season, the gonadal tubule contains all stages of oocyte development. Vitellogenesis appears to involve the biosynthetic activities of the Golgi complex and rough endoplasmic reticulum. A few specimens had transitional ovotestes with mature sperm in the gonad lumen and asynchronously developing oocytes and a small number of spermatocytes within the germinal epithelium. The mature spermatozoon is an ent-aquasperm with ultrastructural features significantly different from those described from other echinoderm classes including a highly elongated acrosome, a large periacrosomal region between the acrosome and nucleus, numerous unfused mitochondria in the midpiece, and a cytoplasmic sleeve or collar extending posteriorly along the proximal portion of the flagellum. The sperm head reaches 11.5 μm in length (combined midpiece, nucleus, periacrosomal region, acrosome), making it the longest yet reported from the Holothuroidea and among the longest in the Echinodermata. Some elements of this derived morphology could be attributed to fertilization biology, but others may have phylogenetic significance. Spawning behavior was observed in which two individuals appeared to pseudocopulate by intertwining their oral tentacles for several minutes before one of them abruptly secreted an egg mass containing three eggs.