Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

The freshwater snail genus Semisulcospira has adaptively radiated in the ancient Lake Biwa, Japan. Although significant geographical variations have been known in the shell morphology of Semisulcospira niponica, their systematics remain uncertain. We evaluated the systematic status of the geographical variations in S. niponica and its congenerics S. biwae and S. fuscata through morphological and genetic investigations. Genome-wide snp s were used to estimate their genetic relationships. Random Forest algorithms explored the morphological variation of the shells of the name-bearing types and newly collected specimens. Our results detected five genetic clusters and revealed the consistency of adult shell morphology with genetic relationships, reinforcing the robustness of shell morphology-based species delimitation of the genus. The present systematics synonymized S. biwae under S. niponica. The five genetic clusters corresponded to the two described species, S. niponica and S. fuscata, and three new species, Semisulcospira watanabei sp. nov., Semisulcospira nakanoi sp. nov., and Semisulcospira salebrosa sp. nov. discovered among the geographical variations of S. niponica. The observed similarities in substrate preference and differences in distribution patterns among species suggest that the diversification of the genus was accelerated by geographical isolation after the initial ecological niche differentiation. We suggest that the distributions of Semisulcospira species may have been influenced by competitive exclusion among them, and that multiple speciation events have occurred in similar patterns.