Trans-Japan Sea land-bridge disjunction: A case of vicariance in the subterranean genus Nipponasellus (Crustacea, Isopoda, Asellidae) in a large-scale biogeographical context

Abstract

This study examines nineteen phreatobiological hand pump samples collected in 2009–2010 in three separate areas of salmon river basins in the southern part of Primorye, in the Far East of Russia. For the first time, faunal groundwater patterns were assessed for the rivers of Eastern Manchuria, the Ussury River Basin and the rivers of the south-western slope of Sikhote Alin. A total of 164 species (including 32 stygobionts) belonging to the phyla Annelida, Mollusca and Arthropoda are first records of groundwater animals, and two of the stygobionts are described below as new species for Science. Nipponasellus sudzukhensis spec. nov. and N. matsumotoi spec. nov. are described and illustrated, and their taxonomic affinities with congeners are discussed. The morphology of male pleopod 2 of the genus Nipponasellus is re-examined using scanning electron microscopy (SEM), which revealed fine details of the pleopodal structures. The two new ‘continental’ species of Nipponasellus were characterised by a distinct ‘cannula’ completely immersed in ctenoid cuticular scales and the absence of a ‘labial spur’. Informal macrogroups were proposed based on the structure of the ‘appendix masculina’ and an analysis of family distribution. The results of cladistic analyses revealed relationships between the proposed groups, but phylogenetic relationships within the Caecidotea-Proasellus group remain challenging. Common diagnostic characters of Nipponasellus showed that the group occupies an independent position close to the Caecidotea-Proasellus group rather than the ‘Asellus-pattern’ sensu Magniez. Revised diagnosis for the genus Nipponasellus is given, including the two new species and the five previously described ones. The species diversity of East Asian stygobiotic asellid isopods is briefly reviewed to infer a biogeographic distribution pattern that emphasises the strict endemicity of members of the genus Nipponasellus. The intracontinental separation between continental Far East Asia and the Japanese archipelago highlights the biogeographic importance of the land-bridge in the Oligocene and raises questions about hypotheses explaining the (disjunct) distribution in East Asia, together with the circumstances preceding an ancient colonisation of the region.