Site selection in a polymorphic mangrove snail

Three species of littorinid snail live on mangroves of the genus Rhizophora on the north coast of Papua New Guinea. They have recently been placed in the genus Littoraria. The species are L. scabra and L. intermedia, which are monomorphic and cryptic and live on bark, and L. pallescens which is polymorphic and lives in foliage. Littoraria intermedia was found mostly on mangroves facing on to partially fresh water, while L. scabra occurred only at sites facing on to the sea; L. pallescens occurs at both types of site. When kept together in plastic containers, L. pallescens shows a greater propensity to climb than the other two species. The morphs of L. pallescens are principally yellow, orange or dark. Orange is present at a frequency of 0–9% and yellow at 10–50%. There is little evidence of geographical variation in frequency over 80 km of coast, although one central locality has a significantly higher frequency of yellow than sites to the north or south of it. Throughout the tropics where the species occurs, orange is the rarest and yellow usually the next rarest morph. Larger individuals are on average higher in the foliage than smaller ones, and there is a higher frequency of yellows among larger shells than smaller ones. As the air temperature increases, a progressively larger fraction of a sample was found to be on the under-surfaces of leaves rather than on the upper epidermis or on flower or leaf buds. When over 30% of a sample is situated on lower surfaces there is a significant excess of yellows and a deficiency of darks on the upper surfaces. This indicates that individuals selectively take up different positions depending on morph colour, resulting in a degree of optimal background matching. Such a choice of location may be of direct relevance to the maintenance of the polymorphism. A difference in morph frequency between the same sites examined in 1979 and 1984 is interpreted as being a result of site selectivity.