Unparalleled disjunction or unexpected relationships? Molecular phylogeny and biogeography of Melanopsidae (Caenogastropoda: Cerithioidea), with the description of a new family and a new genus from the ancient continent Zealandia.

Melanopsidae are an important component of the freshwater fauna of the subtropical to temperate regions of the Western Palaearctic and also are reported from Zealandia, representing an unparalleled disjunction among a group of freshwater animals. We sequenced markers for species of all constituent genera covering the entire range of the group. Our phylogenetic analyses indicate that Melanopsidae are only monophyletic when excluding Holandriana from the Balkans, which was found to be more closely related to Pleuroceridae (eastern North America) and Semisuclospiridae (western North America and eastern Asia) than to the remaining melanopsids. It is, therefore, placed in the previously proposed family Amphimelaniidae. Zemelanopsis from New Zealand and Caledomelanella gen.n. from New Caledonia were recovered as a deeply divergent clade, here suggested to represent Zemelanopsidae fam.n. as the sister group of Melanopsidae s.s. The genus Microcolpia, including M. daudebartii and the extinct M. parreyssii, was recovered as a moderately well-supported sister group of Melanopsis s.s., which comprises three distinct lineages: one from the western Mediterranean region, a second from the eastern Mediterranean region and the Near to Middle East, and a third from thermal springs in Tuscany. Although the Italian populations show little variability, both the eastern and western Melanopsis clades exhibit a high degree of variation in shell form and/or sculpture. Whether this is indicative of more than a species pair each (M. praemorsa/cariosa and M. buccinoidea/costata) within the eastern and western clade, respectively, or the result of parallel evolution currently cannot be answered with certainty. Distinct populations were not recovered as monophyletic clades in some cases, which may indicate incomplete reproductive isolation. We compare the timeframe for the evolution of major "melanopsid" lineages with geological events, and elucidate scenarios that may have shaped the distribution patterns observed today.