Morphometric analysis of male reproductive features of octopodids (Mollusca: Cephalopoda).

Taxonomic accounts of octopodids frequently describe the spermatophore, the penis that releases the spermatophore from the internal organs, and the ligula and calamus that transfer it to a female. To explore relationships among these male features and body size, this study applies principal components analysis to data from 43 species of the family Octopodidae, or benthic octopuses. Covariation in penis and mantle length opposed by covariation in ligula and calamus lengths forms primary shape variation. Secondary shape variation is due to opposing variation between ligula and calamus lengths. Primary shape variation is greatest among shallow-water species. The calami and ligulae of diurnal and crepuscular shallow-water species are short compared to those of nocturnal shallow-water species. Because these structures contain heterogeneous collagen arrays and lack camouflaging chromatophore organs, they are white. Diurnal and crepuscular octopus species may minimize their lengths due to selection imposed by visual predators. Secondary shape variation is greater in deep-sea and high-latitude octopuses. Members of Voss's Eledoninae (except Eledone) and Graneledoninae and two species of Benthoctopus have exceptionally long calami and comparatively short ligulae; these lengths vary among members of the Bathypolypodinae. Variation in spermatophore length is independent of the structures considered.