A review of sperm storage methods and post-copulatory sexual selection in the Cephalopoda

In many animal phyla, females have a unique sperm storage organ (SSO). Post-copulatory sexual selection is a powerful driving force of SSO evolution. SSOs are generally considered to have evolved through sexual antagonistic coevolution between male genitalia and the SSO and/or cryptic female choice (CFC). In cephalopods, sperm transfer and fertilization are conducted through complex processes, and sperm storage methods show inter-species variation. In some species, males implant spermatangia superficially under the female skin, and then sperm released from the spermatangia are transferred into a seminal receptacle (SR). Deep-sea cephalopods, which lack a SR, have instead evolved a deep-implanting method by which the spermatangium is embedded deep in the musculature of the mantle wall of the female. In some species, the female stores whole spermatangia within a spermatangium pocket. Because the males of most species do not insert an intromittent organ into the female when transferring sperm, SSO evolution may have been influenced by CFC alone. This review summarizes the sperm storage methods and the mechanisms of post-copulatory sexual selection in cephalopods and it is proposed that these diverse methods evolved as adaptive mechanisms through post-copulatory sexual selection.