Mating behaviour and spermatophore transfer in the shrimpHeptacarpus pictus(Stimpson) (Decapoda: Caridea: Hippolytidae)

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Abstract
Heptacarpus pictus, a small caridean shrimp inhabiting the low intertidal of southern and Baja California, breeds during the winter, spring, and summer months. Fall is a period of growth. Life span of an individual does not exceed 18 months, with fish predation as the most likely source of mortality. Females are multiple brooders, carrying developing embryos concomitant with increase in ovary size. Hatching of larvae is followed by a moult, after which the female is attractive to males and receptive to copulation. A distance pheromone does not appear to be involved in attraction of males to females. Males apparently respond to a non-diffusible substance on the exoskeleton of newly moulted females. Precopulatory behaviour is absent. Copulation can be divided into a series of relatively stereotyped events. Female rejection of the male or his spermatophore is the chief cause of unsuccessful matings. Males deposit the spermatophore on the underside of the female's first abdominal segment. Sperm packets are formed upon extrusion from the male's genital openings, and are composed of a mucoid material in which sperm are mixed. The glutinous spermatophores adhere to the female's smooth abdominal sternite. The endopods of pleopods 1 and 2 of the male are different in shape, size and setation from homologous rami of females and juveniles. Endopods of pleopod 1 possess a distally located appendix interna, absent in juveniles and females. An anteriorly projecting process, the appendix masculina, is located on the endopod of pleopod 2 in males. Experiments were performed which showed that these modifications insure proper deposition of spermatophores. Males which had the copulatory rami removed did not transfer spermatophores as successfully as normal males. Transfer of the spermatophore from the male to the female is a result of the interplay of male pleopods 1 and 2 during copulation. The large expanded abdominal pleurae of females prevent the male's genital opening from contacting her abdominal sternite. Thus, the male's anterior pleopods have become modified to lift the emitted spermatophore from his genital orifices to the first abdominal sternite of the female.