A depth cline in deep-sea octopods (Cephalopoda: Graneledone) in the northeast Pacific Ocean

Bathyal octopods of the genus Graneledone Joubin, 1918 in the northeast Pacific differ dramatically in skin texture. To test the hypothesis that these differences are associated with geography, we quantified the skin warts and tubercles of 50 specimens collected between 36 degrees N and 46 degrees N from 1116 to 2850 m depth. Using Principal Component Analysis (PCA), we analyzed the numbers of mantle and head warts, mantle and web tubercles, arm suckers, and inner and outer gill lamellae, adding head width as a size proxy. We used Canonical Correlation Analysis to explicitly test the relationship of morphology to depth and latitude. Sequences of 12S rRNA, 16S rRNA, and COI genes from 12 individuals quantified genetic divergence. PCA revealed correlated variation in head width, arm suckers, and gill lamella counts opposed by skin texture on PC1; PC2 showed correlated variation in skin texture opposed by gill lamella counts. Both components correlate with depth and latitude but, as most shallow specimens came from southern localities, the correlation with latitude may be artefactual. This relationship is corroborated by the canonical correlations. Octopods from greater depths are smaller than those from shallower depths, have rougher skin texture, and fewer suckers and gill lamellae. Clades of shallow and deep octopods show 0.8% genetic divergence in COI (Kimura two-parameter distance), not inconsistent with intraspecific differentiation. We conclude that the variation is likely clinal with depth. The observed morphological differences may result from differences in food availability and oxygen saturation with depth.