Abstract
The phylogeny of the genus Echinolittorina, and phylogeography of some of its members, have previously been examined using molecular data, so that species can now be defined by a combination of phylogenetic, morphological and geographical criteria. The 26 species recognized in the Indo-West Pacific biogeographical region form a monophyletic group, here defined as the subgenus Granulilittorina Habe & Kosuge, 1966. Morphological descriptions are provided for these 26 species, including details of shell, pigmentation of headfoot, reproductive anatomy, spermatozoa, egg capsules and radulae. Diagnoses include reference to mitochondrial gene sequences (COI). A key is based on shells, tentacle pigmentation, penial shape and geographical distribution. Seven new species are described: E. marisrubri, E. omanensis, E. austrotrochoides, E. marquesensis, E. wallaceana, E. tricincta, E. philippinensis. Three name changes are proposed: E. malaccana (Philippi, 1847) and E. cecillei (Philippi, 1851) are valid names for two members of the former ‘E. trochoides’ group; E. biangulata (von Martens, 1897) replaces ‘E. quadricincta’. Full synonymies are given for all taxa, and the taxonomic, evolutionary and ecological literature reviewed. Distribution maps are based on examination of 1701 samples and reliable literature records. The contrast between continental and oceanic distribution patterns is emphasized; one clade of five species and two additional species are shown to have an association with upwelling areas. All species are known (or predicted from protoconch size and oviduct anatomy) to have planktotrophic development, and rare extralimital records suggest a maximum open-water dispersal distance of 1000–2100 km. The most useful morphological characters for identification are the shell, penial shape and copulatory bursa in the pallial oviduct. Sister species can be morphologically similar, but are almost always entirely allopatric, so that distributional information is important for identification. Substantial intraspecific variation is present in the shell shape and sculpture of most species; where there is a pronounced geographical pattern this may have a genetic basis, but ecophenotypic effects are also implicated, e.g. by predictable associations in some species of strongly nodulose sculpture with limestone substrates and with dry habitats where growth rate may be slow. Morphological characters are superimposed on a molecular phylogeny to demonstrate the synapomorphies of clades. This is essentially a morphostatic radiation of largely allopatric species with little morphological differentiation; ecological divergence is limited to specialization to oceanic, continental or upwelling areas and to small differences in zonation level.
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