Reproductive traits and sexual selection in the mangrove littorinid snails, Littoraria ardouiniana and L. melanostoma

Abstract

According to Darwin, any inheritable traits that enhance fitness (i.e. survival and reproductive success) would be adaptive in a given environment, and therefore subject to natural selection. Some traits that enhance reproductive success but not necessarily survival may, however, evolve through the process of sexual selection. The importance of sexual selection has been intensively studied in “higher animals” birds, mammals and insects but has largely been neglected in “lower animals” such as gastropods. Using two mangrove snails, Littoraria ardouiniana HEUDE 1885 and L. melanostoma GRAY 1839 (Littorinidae) as model species, this thesis documents their various reproductive traits and the occurrence of sexual selection, in an attempt to understand how these animals optimize their fitness.

Being one of a few gastropod families that live closely associated with mangrove habitats, various adaptive reproductive traits are likely to have evolved in the genus Littoraria. The two Littoraria species exhibited several, contrasting, reproductive traits (e.g. reproductive mode; length of reproductive season; seasonal fecundity and egg- or larvae-releasing rate), which may represent species-specific strategies to optimize reproductive success. Both species adopted mucus trail following as a mate-searching strategy; with males being able to discriminate the trails laid by conspecific females and trail orientation during the mating season. Since these two snails mate in the complex habitat of mangrove tree canopies, incorporating some cue into mucus trails to aid mate recognition would increase potential encounter rates and hence reproductive success. 1D proteomic techniques identified two potential pheromones in the trail mucus of female L. ardouiniana, but further investigations are needed to confirm their possible role as sex pheromones.

Evidence for sexual selection was found in L. ardouiniana through male mate choice and male-male competition. Large male L. ardouiniana showed a preference for mucus trails laid by large (= more fecund) females, but this preference was not seen in small males, suggesting a size-based male mate preference. Males of this species also mated with larger females for a longer duration and they showed physical aggression (i.e. they pushed each other) when they encountered a female. These sexual selection mechanisms may drive size-assortative mating patterns in littorinid species and in other animals.

Conversely, L. melanostoma exhibited no obvious male mate preference or male aggression. Such differences in sexual selection patterns from L. ardouiniana were proposed to be a result of differences in operational sex ratio and variation in female quality. As many gastropods show similar mating behaviour to littorinids (i.e. trail following, shell mounting and copulation), sexual selection may operate in a comparable manner in other gastropods. By revealing the complexity of sex roles of males and females (e.g. the mutual occurrence of male mate choice and male-male competition) and condition-dependent male mate preference in these mangrove snails, this thesis has contributed to a missing link between sexual selection and the “lower animals” and provides new insights into the operation of sexual selection in animals.