Habitat and zonation patterns of Littoraria species (Gastropoda: Littorinidae) in Indo-Pacific mangrove forests

The zonation of 15 species of Littoraria in the scabra group is described, using a transect method, at 13 mangrove localities in Australia, Thailand, Malaysia and Hawaii. At these localities the littorinids occurred only on trees, and showed both vertical and horizontal patterns of zonation, with broad overlaps between co-occurring species. It is suggested that vertical distribution is determined mainly by behavioural responses, while the landward limits of horizontal distribution may be set by physiological tolerance. Several species occurred predominantly on either leaf or bark substrates, but no consistent associations with tree species were found. On the basis of local distribution patterns, Littoraria species can be classified as continental or oceanic in character. Oceanic species may be less tolerant of sediment in the water or on the substrate.