Increased copulation duration does not necessarily reflect a proportional increase in the number of transferred spermatozoa

Post-copulatory sexual selection research tends to focus on the numerous adaptations that have evolved to increase the chances of donated spermatozoa fertilizing oocytes. Even though fertilization obviously directly depends on the presence of sufficient, viable spermatozoa, the quantification of the sperm transfer process itself has not received the attention it deserves. Here, we present experimental work on a simultaneously hermaphroditic snail in combination with a review of the literature focussing on the relationship between the duration of copulation and the number of sperm that are transferred. Based on classical work, this relationship is often assumed to be linear, but as we show here this need not be the case. Both our experimental data and the reviewed literature indicate that there are clear instances where the process of sperm transfer is not a linear process, i.e., longer copulation duration does not necessarily imply more transfer of sperm. As we point out, there seems to be a bias in the literature towards investigating this in insects, but other animal groups in which this has been investigated do show similar relationships. To conclude, we discuss how the specific patterns of sperm transfer that have been reported can be biologically interpreted and we caution that simply using copulation duration as a proxy for the number of sperm transferred can be misleading.