Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae) Author Sawada, Naoto Author Fuke, Yusuke text Contributions to Zoology 2023 2022-08-22 92 1 1 37 http://dx.doi.org/10.1163/18759866-bja10035 journal article 267806 10.1163/18759866-BJA10035 c846d9c5-4d9d-441f-919a-aa40017d9b93 1875-9866 8349612 Semisulcospira nakanoi Sawada sp. nov. [New Japanese name: Chikubu-kawanina] ( table 2 , supplementary table S2 ; figs 10 , 12M, N ) urn:lsid:zoobank.org:act: 9CB53BD3-F776- 4ACB-9C9D-DE0755AADA16 Melania niponica – Annandale, 1916: 45 , pl. 3, fig. 3B (part). Melania biwae – Annandale, 1916: 45 , pl. 3, fig. 4 . Semisulcospira niponica – Kuroda, 1929: 186 , 189, pl. 5, fig. 31 (part); Hirase & Taki, 1951 : pl. 82, fig. 15; Kuroda, 1962: 86 (part; TAU); Kuroda & Habe, 1965: 57 (part); Habe & Kosuge, 1967: 28 , pl. 11, fig. 21 (part); Watanabe, 1984 : fig. 3b, f ; Watanabe, 1989: 11–13 , 47, figs 10 , 27, 28 (part). Semisulcospira niponica var. biwae – Fukuoka , 1933: 114 , 117, fig. 6 . Semisulcospira ( Biwamelania ) niponica – Nishino, 1991: 9–10 , unnumbered figures (part); Kihira et al., 2009: 24–25 (part). Semisulcospira nipponica biwae – Higo & Goto, 1993: 97 ; Goto & Poppe, 1996: 204 . Semisulcospira ( Biwamelania ) niponica ( biwae type ) – Watanabe & Nishino, 1995: 17–18 , fig. 5b (part). Biwamelania niponica – Nishino & Tanida, 2018: 56 , 251 (part). Material examined : Holotype : KUZ Z4122, adult female with embryonic shells treated with sodium hypochlorite, collected from depth of 3.0 m of Lake Biwa at Chikubu-shima , Nagahama City , Shiga Prefecture , Japan , on 9 September 2020 by first author . Paratypes : KUZ Z4123–Z4126, 3 adult females, 1 adult male, collected with holotype . Additional materials: KUZ Z4127, 11 females , KUZ Z4128, 2 males , collected with type specimens; KUZ Z4129, 16 females , KUZ Z4130, 5 males , collected from Onoe Port on 25 July 2021 . Etymology : The specific name is dedicated to Dr Takafumi Nakano, who greatly supported the first author’s study of Japanese Semisulcospira . Diagnosis : Viviparous semisulcospirid. Adult shell medium sized (SH 30.6 ± 2.7, 23.1 ± 3.0 mm; BWL 18.9 ± 1.3, 14.8 ± 1.7 mm ), nearly triangular (SA 21.5 ± 2.6, 21.4 ± 3.5 degrees); color in black background; outer lip of aperture moderately swell (ASL 0.20 ± 0.05, 0.24 ± 0.06 mm ); 2.5 ± 0.5, 2.3 ± 0.6 BCN; longitudinal ribs relatively few, prominently nodulated on penultimate whorl (RN 11.3 ± 1.3, 9.4 ± 1.2); 3.7 ± 0.6, 3.4 ± 0.5 SCN; 1.6 ± 0.1, 1.6 ± 0.1 ASR; 2.9 ± 0.2, 3.2 ± 0.6 WER. Embryonic shell small, with distinct nodes on surface; color in black background or beige background with 3 brown or black bands. Description of holotype : Adult shell ( fig. 10A–C ): AH 11.5 mm , AL 12.5 mm , ASR 1.48, AW 8.5 mm , BCN 2, BWL 19.7 mm , FWL 4.4 mm ; NL 0.18 mm ; PWL 7.3 mm , RN 9, SA 23.1 degrees, SCN 4, SH 34.1 mm , SW 15.7 mm , TWL 5.9 mm , WER 2.61; WN 4.75; shell nearly thick triangular; suture hardly undulating; whorl sides slightly convex; outer lip of aperture moderately swell; longitudinal ribs slightly curved, oblique, prominently nodulated in upper to body whorl; apex of shell eroded; shell colored black background without color band. Operculum ( fig. 10D ): 8.9 mm in long diameter; nearly egg-shaped subcircular, paucispiral, comprising around 3 whorls. Nucleus subcentral. Embryonic shells ( fig. 10E–G ): EN 62, RNE 12, SHE 2.3 mm , SWE 2.2 mm , WNE 2.5; shell globose; suture prominently undulating; remarkable nodes on central part of whorls and weak keels on lower part of whorls; shell colored beige background with 3 brown or black bands on upper and/or lower part of each whorl and/or on basal part of shell. Radula: Taenioglossa consisting of rachidian in single, lateral teeth, interior and exterior marginal teeth in double row. Rachidian roughly triangular with large central denticle and 3 minor pointed triangular cusps on each side. Lateral teeth with large central denticle, 3–4 inner and outer pointed cusps. Central denticle tip of rachidian and lateral teeth mostly flat; central denticle of rachidian approximately regular triangular, about 2.5 times longer than other triangular cusps; central denticle of lateral teeth irregular triangular, about 1.5 times longer than other triangular cusps. Interior and exterior marginal teeth spoon-shaped with 5–6 rounded denticles. Reproductive organ: Female: Long narrow oviduct, entering near seminal receptacle with long protrusions. Ventral edge of spermatophore bursa with curved sperm gutter, extending toward mantle cavity. Brood pouch on dorsal side of spermatophore bursa, inflated dorsally, separated into many cells, including eggs and embryos; eggs and embryos radially developing from base of brood pouch near seminal receptacle and embryos in anterior or dorsal cells more developed. Variation : Adult shells ( fig. 10M, P ): Measurements and counts shown in table 2 and supplementary table S2 . Shell nearly thick to elongated triangular; surface sculptures almost straight to slightly curved, oblique, rarely ribbed on upper whorls; shell colored black to brackish olive background with or without several color bands on upper and lower part of each whorl and basal part of shell; surface of shells covered with thin algae layer before treatment. Operculums ( fig. 10I, L, N, Q ): 7.1–8.9 mm in long diameter. Embryonic shells ( fig. 10J, Q ): Measurements and counts shown in table 2 and supplementary table S2 . Shell with or without keels on lower part of whorls; shell colored blackish brown background or beige background with 3 brown or black bands on upper and/or lower part of each whorl and/or on basal part of shell. Radulae ( fig. 12M, N ): Rachidian with 2–4 minor pointed triangular cusps on each side; lateral teeth with 2–4 inner and outer pointed cusps; central denticle of rachidian about 1.5 to 4 times longer than other triangular cusps; central denticle of lateral teeth about 1.5 to 4.0 times longer than other triangular cusps; 4–6 rounded denticles in interior and exterior marginal teeth. Reproductive organs: Male: Gonad consisting of testes, vas deferens, and prostate without penis. Posterior ventral part of inflated prostate with deep groove, forming U-shape in transverse section. Anterior prostate narrowly opening to mantle cavity. Distribution and ecology : The new species has been found only in Chikubu-shima Island and the coastal Onoe Port in Lake Biwa ( fig. 1 ). Semisulcospira nakanoi sp. nov. inhabits the wharf and rocky bottom of Chikubu-shima Island and concrete blocks around Onoe Port. This species was found with S. morii and S. habei at Chikubu-shima Island. Remarks : The new species has been identified as S. biwae together with S. salebrosa sp. nov. and has been treated as a junior synonym of S. niponica since Kuroda (1929) . Semisulcospira nakanoi sp. nov. resembles S. salebrosa sp. nov. with well-developed nodes on the shell surface. However, the former species is distinguishable from the latter by the smaller adult and embryonic shell size and less undulating outer lip of the aperture. The new species can be discriminated from other congeners by its large, blackish, triangular adult shell, prominent granulated ribs, the moderately undulating outer lip of the aperture, and globose embryonic shells with several brown bands (supplementary fig. S1 ).