SYSTEMATIC REVISION OF PATELLOIDA PYGMAEA (DUNKER, 1860) (GASTROPODA: LOTTIIDAE), WITH A DESCRIPTION OF A NEW SPECIES

Patelloida pygmaea (Dunker) and its closely allied species,P. heroldi (Dunker) and P. conulus (Dunker) have caused nomenclaturalconfusion because of their variable shell morphology and distinctivehabitats. According to current nomenclature, these species ofPatelloida have been synonymized and treated as one specieswith two ecological forms. Patelloida pygmaea lives on the shellof Crassostrea gigas (Ostreidae), P. pygmaea form heroldi occurson intertidal rocks on sheltered shores and P. pygmaea formconulus is found on the shell of Batillaria multiformis (Batillariidae).Their taxonomic relationships and possible species status are,however, unclear. Using two mitochondrial genes (fragments ofCOI and 16S ribosomal RNA; total 1192 sites), we analysed 88specimens of P. pygmaea, P. pygmaea form heroldi and P. pygmaeaform conulus from 37 localities in East Asia. In the resultingmolecular phylogenetic trees, the specimens of Patelloida fallinto four clades with high bootstrap probabilities; these cladescorrespond taxonomically to P. pygmaea, P. conulus, P. heroldiand a fourth previously unrecognized taxon, Patelloida ryukyuensisn. sp., described here. A minimum-spanning network for 29 uniquemitochondrial COI haplotypes obtained from 45 specimens in thesame bay in central Japan form three distinct clusters, consistingof P. pygmaea, P. conulus and P. heroldi, respectively. Thissuggests that reproductive isolation has been established betweeneach group. A detailed examination of radular and shell morphologiesof the four taxa clarifies the morphological distinction betweenthese species. The four species form a rather young clade inthe genus Patelloida that diverged during the Pliocene. Theyprovide an example of habitat segregation in a restricted marineenvironment. (Received 21 December 2004; accepted 11 March 2005)     

Evolutionary models of lineage-zones

Among the various kinds of biostratigraphic units, lineage-zones are especially interesting and meaningful, not only in stratigraphy but also in evolutionary biology. The chronological shift of morphology in evolution is fundamentally uninterrupted, but sometimes it seems discontinuous or even saltational. Such breaks have been commonly attributed to gaps in sedimentation, but could be explained, at least in some cases, by more essential mechanisms of evolution. The difference in the patterns of morphological change may be primarily related to the nature of non-sex-associated genetic variation. continuous or discontinuous, as well as the strength of selection pressure. Some theoretical models for phenotypic substitution are presented here in terms of population genetics, though these are not the only possible explanation for breaks in fossil records.