Supratidal existence drives phenotypic divergence, but not speciation, in tropical rocky-shore snails

Abstract

Abstract The mechanisms underlying the evolutionary adaptation of animals that transcend the ecological barrier separating the intertidal and supratidal zones of rocky shores are poorly understood. Different wetting frequencies in these zones in tropical regions (daily vs. seasonally, respectively) impose different physical stressors, which should drive phenotypic variation and ultimately speciation in the animals that inhabit them. We studied morphological, physiological and genetic variation in a tropical high-shore gastropod that transcends these zones [Echinolittorina malaccana (Philippi, 1847)]. Variation in melanization, shell features and evaporative water loss was linked to regular seawater wetting, frequent activity and feeding, and solar exposure in intertidal snails, and to inactivity and prolonged aestivation in the shade during continuous air exposure in supratidal snails. Despite selective pressure for phenotypic divergence, and reproductive isolation of the populations in either zone, their mitochondrial COI gene sequences confirmed that they represent a single species. Speciation in our study case is probably constrained by the limitation on activity, mating and reproduction of supratidal snails, such that their populations can only be sustained through intertidal pelagic larval recruitment. Comparisons with other studies suggest that supratidal speciation and specialization for life in this zone probably require moderation of the abiotic (desiccative) conditions, to facilitate greater activity and interaction of animals during air exposure.