Comparative studies of the reproductive strategies of New Zealand grapsid crabs (Brachyura: Grapsidae) and the effects of parasites on their reproductive success.

The reproductive strategies of four intertidal grapsid crabs, Hemigrapsus sexdentatus, H. crenulatus, Cyclograpsus lavauxi, and Helice crassa, were studied in the field and laboratory, with emphasis on mating behaviour, duration of female receptivity, and sperm competition. Mating occurred in all species during the intermoult on the days prior to oviposition, when the gonopore opercula of females became temporarily mobile. Female Helice crassa mated up to three weeks after oviposition, but in all other species mating typically ceased at egg-laying. Male Hemigrapsus spp. used a female-centered competition strategy in which they searched for and defended receptive females until they laid eggs. In contrast, male C. lavauxi searched for and intercepted receptive females only for the duration of copulation and then pursued other receptive females (a mating system termed encounter rate competition with pure search and interception). Male Helice crassa searched for receptive females in their immediate neighbourhood and mated with them briefly on the substrate or in the burrow after which the female left (a mating system termed encounter rate competition with neighbourhoods of dominance). The mating season was short and highly synchronous for Hemigrapsus sexdentatus and Cyclograpsus lavauxi and asynchronous for Hemigrapsus crenulatus and Helice crassa. In the laboratory, the mean duration of receptivity for females housed with three males varied between 4.1 and 12.4 days, and the copulation frequency of females varied before oviposition between 2.1 and 24.3 times (mean) depending on the species. Female Hemigrapsus spp. Isolated from males stayed receptive significantly longer than females held continuously with males. This suggests that females are able to control the duration of their receptivity, and therefore the time available for mating, according to the absence or presence of males. The operational sex ratio (OSR) had no effect on the duration of female receptivity, but female Hemigrapsus crenulatus mated more often when several males were competing for access. Therefore, male-male competition increased the number of matings per female and hence sperm competition within the female spermathecae. Larger males mated significantly more often than smaller males in all species. However, male size did not affect ejaculate size, meaning that small and large males transferred similar-sized ejaculates, e.g., in Hemigrapsus spp. Males of the two Hemigrapsus species followed a different strategy of sperm allocation. Male H. crenulatus, which are typically confronted with a high mating frequency of the female and a long, asynchronous mating season, distributed similar-sized ejaculates, irrespective of female size. By contrast, male H. sexdentatus, which experience a comparatively lower risk of sperm competition during a short, synchronised mating season, invested larger ejaculates for larger females than for smaller females. In addition, the size of the first and second ejaculates transferred to a female by a male H. crenulatus were not significantly different, whereas the first was larger than the second for H. sexdentatus. A parasitological survey was undertaken of the four grapsid crabs and the presence, seasonal variation and relationship with host gender and size of parasites determined. Four internal parasites were discovered: Nectonema zealandica n. sp. (Nematomorpha: Nectonematoidea), Portunion sp. (Isopoda: Entoniscidae), Profilicollis novaezelandensis n. sp. and Profilicollis antarcticus (Acanthocephala: Polymorphidae). Portunion sp. castrated its female hosts, but not the males thereby creating a more male-biased sex ratio. Males parasitized with Portunion sp. were equally successful during male-male competition and the number of matings they achieved. The above findings are important for our current understanding of mating strategies in Grapsidae, which are more diverse than previously thought. Females with a restricted duration of sexual receptivity have some control over their receptive period and can therefore influence the OSR and the extent of male-male competition. As females mated multiple times during their receptive period, sperm competition is a common feature in Grapsidae. However, males employed different tactics in regards to sperm competition such as longer mating duration (e.g., C. lavauxi), high number of matings (Helice crassa), or post-copulatory mate guarding until oviposition (Hemigrapsus spp.)