We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September-April. Populations within large patches of forest were annuals; all or most individuals died between October-January (spring-midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these "peripheral sources" are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats.

Recent theoretical models have considered how nutritional status and predation risk affect the decision by an individual to forage. We consider the feeding behaviour of a scavenger under risk of predation. The intertidal gastropod Nassarius dorsatus Röding was used to test the following hypotheses: (1) a damaged conspecific should be a less preferred meal than a damaged predator or another species; (2) a scavenger should be able to discriminate between a damaged conspecific in the presence and one in the absence of its predator, preferring the latter; and (3) the decision by a scavenger to feed should reflect a trade-off between nutritional status and the above preferences. Results from the laboratory and field were consistent with predictions. Two hypotheses are suggested from the work: scavengers may generally be less willing to feed on carcases of conspecifics than on those of other species, and intertidal scavengers may commonly face a relative or absolute shortage of food.

Many species of arboreal marine snails in the genus Littoraria are polymorphic for shell colour, with morphs that correspond in both colour and frequency to those predominant in the habitat. Although the combined effects of selection for crypsis and apostasy have been suggested as the most likely reasons for this, they have not been demonstrated directly in the field. We investigated whether two parasitoid flies, Sarcophaga megafilosia and Sarcophaga meiofilosia, select for crypsis in Littoraria filosa. It was possible to compare the proportions that matched and did not match the background between samples of live and dead snails because the shells of individuals killed by these parasitoids, within which the larva and pupa develop, remain attached to the substratum. This comparison was necessary because these snails frequently move among different coloured microhabitats, which will tend to obscure any effect of selection for crypsis if only live individuals are censused. The method appeared reliable since there was no change in background colour and very little loss of shells between larviposition and emergence of flies. S. megafilosia killed a significantly greater than expected proportion of snails that did not match their background. In contrast, there was no evidence of selective attack by S. meiofilosia. These results are discussed in relation to previous work on polymorphisms in Littoraria species. We also suggest that the role of parasitoid insects as selective agents contributing to the maintenance of colour polymorphisms in terrestrial molluscs may have been overlooked. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 367−377.

The marine snail, Littoraria filosa, is polymorphic for shell colour, with yellow, brown, and pink morphs that correspond in both appearance and frequency to the predominant background colours of its habitat. Previous research on this polymorphism has found indirect evidence of apostatic selection and selection for crypsis by unknown agents, probably crabs, and direct evidence of selection for crypsis by the parasitoid fly Sarcophaga megafilosia. In the present study, we report on field experiments to investigate whether S. megafilosia and shell-crushing predators exert apostatic selection on L. filosa. For S. megafilosia, seven experimental treatments containing yellow and brown snails in the proportions of 0.1, 0.2, 0.4, 0.5, 0.6, 0.8, and 0.9 of each colour were established on mangrove trees and used to separately quantify the proportions of each colour attacked on grey/brown trunks and yellow/green leaves. The results obtained confirmed an earlier finding of selection for crypsis, but only showed slight, but significant, anti-apostatic selection by S. megafilosia. For shell-crushing predators, seven experimental treatments containing yellow and brown snails in the proportions of 0.08, 0.17, 0.33, 0.50, 0.66, 0.83, and 0.92 were established on two types of trees that differed in their background proportions of brown and green: (1) trees which had been pruned of approximately 90% of their foliage and (2) unpruned trees. The results obtained showed both selection for crypsis and apostatic selection. Furthermore, a selectively neutral frequency for yellow L. filosa was found for each background, and was less on pruned trees than unpruned ones (and vice versa for brown L. filosa), which can therefore account for the maintenance of a colour polymorphism where the proportions of each morph tend to resemble and correspond in frequency to the colours of the background.

We provide descriptions of two species of sarcophagid flies seen attacking the intertidal snail Littoraria filosa (Sowerby 1832), which occurs on leaves and branches of mangroves in central Queensland, Australia. This is the first description of sarcophagid parasitoids of a marine snail. Sarcophaga megafilosia sp. n. and Sarcophaga meiofilosia sp. n. belong to the subgenus Sarcorohdendorfia Baranov 1938 and both are parasitoids of L. filosa. These two parasitoids cause considerable mortality of L. filosa, which is an annual in large patches of mangroves (where both flies are present), but a perennial in smaller, more isolated patches (where the larger fly is usually absent).