On the origin and diversification of the stygobiotic freshwater snail genus Hauffenia (Caenogastropoda: Hydrobiidae) with special focus on the northern species and the description of two new species Author Haase, Martin 8CE4B347-14C5-464F-99C4-DE2263FC55D5 Vogelwarte, Zoological Institute and Museum, University of Greifswald, Soldmannstrasse 23, 17489 Greifswald, Germany. martin.haase@uni-greifswald.de Author Grego, Jozef 44C83AAB-CF4E-46E8-8146-DFBADA150AB6 Horná Mičiná 219, 97401 Banská Bystrica, Slovakia. jozef.grego@gmail.com Author Erőss, Zoltán Péter 0B2C74A7-0001-4E25-984C-88CB0C705313 Levendula utca 68 / 4, 2119 Pécel, Hungary. erosspeter@hotmail.com Author Farkas, Roland 9BCFB1AA-7605-4B43-937C-A659D020912B Bükk National Park Directorate, Sánc utca 6, 3304 Eger, Hungary. farkasro@yahoo.com Author Fehér, Zoltán E801EC76-8B1E-450B-993E-BBBE57C00EA9 Hungary, Álmos vezér útja 69 / A, 1141 Budapest, Hungary. feher.zoltan.nhmus@gmail.com text European Journal of Taxonomy 2021 2021-10-25 775 143 184 journal article 3885 10.5852/ejt.2021.775.1555 e0c68699-90c8-4b60-8ed4-e0bd6a7d066f 2118-9773 5596674 264FC47B-9D2B-47F2-9461-F3A6CA5EB58B Hauffenia lozekiana sp. nov. urn:lsid:zoobank.org:act: 63FE4F3F-32A5-4F83-869F-C72B6E2D56AE Figs 2 , 5 71a–b, 12A–C, 13A–B, 14A, 15A–C, 17A–B, 18; Tables 3–4 Diagnosis The combination of valvatiform shell with eventually detached peristome and an only initially sculptured embryonic shell, a thin, yellow operculum without peg-like structure in the nucleus and the massive, trapezoidal penis, whose left side is flattened, distinguishes H. lozekiana sp. nov. from all other congeneric species. Only three diagnostic positions of the COI alignment were of type 1, a single of type 3, and 107 of type 4 ( Table 3 ). Uncorrected genetic distances to congeners ranged from 0.077 ( H. kerschneri ) to 0.133 ( H. steffeki sp. nov. ; see Supp. file 3). Etymology Hauffenia lozekiana sp. nov. is dedicated to Vojen Ložek ( 26 Jul. 1925 – 15 Aug. 2020 ), renowned and influential Czech malacologist and paleontologist, who contributed significantly to Quaternary malacostratigraphy, Czechoslovakian zoogeography of mollusks and was the first to discover the genus Hauffenia in Slovakia . Material examined Holotype ( Fig. 12A–C ) SLOVAKIA • Bystrica Highlands , vicinity of Poniky/Oravce , Kalinovský prameň (Kalinovec); 48°42.027′ N , 19°15.952′ E ; alt. 482 m ; 4 Jan. 2014 ; J. Grego leg.; karst spring; NHMW 113638 . ( Fig. 5 71a–b) Paratypes ( Fig. 13A–B ) SLOVAKIA • 5 specs ; same collection data as for holotype; JG F1684 • 4 specs ; same collection data as for holotype; NHMW 113639 • 2 specs ; same collection data as for holotype; ZPE • 2 specs ; same collection data as for holotype; 25 Jan. 2014 ; HNHM 105307 • 2 specs ; same collection data as for holotype; JG F1649 • 2 specs ; same collection data as for holotype; 18 May 2017 ; JG F1650 • 4 specs ; same collection data as for holotype; 2 Jul. 2020 ; JG F1886 • 2 specs ; same collection data as for holotype; NMBE 569388 • 2 specs ; same collection data as for holotype; SMF 363255 • 2 specs ; same collection data as for holotype; SMOPAJ 1/2021 • 2 specs ; same collection data as for holotype; ZMH 140884 • 3 specs ; same collection data as for holotype; 21 Dec. 2020 ; JG F1887 . Description SHELL ( Figs 12A–C , 13A–B , 14A ; Table 4 ). Valvatiform, maximum height measured 1.03 mm , maximum width 1.58 mm , on average twice as wide as high, with up to 2.75 whorls, final part of last whorl may be detached, fairly variable in shape with a coefficient of variation of 13.31% for the height/ width ratio; embryonic shell with 1.1–1.2 whorls, first 0.75 whorls wrinkled, sculpture then fading to become smooth; aperture almost round only slightly higher than wide, prosocline, lip continuous, not thickened; umbilicus wide. OPERCULUM. Thin, chitinous, yellow, almost circular with central nucleus, the latter without peg or peglike modification. EXTERNAL FEATURES. Epidermis without pigment, no eye spots visible, tentacles without particular ciliation. MANTLE CAVITY. 9 gill filaments (N = 1), osphradium short, oval, behind middle of gill. DIGESTIVE SYSTEM ( Figs 15A–C , 16 ). Radula with formula R 4-5 1 4-5/1 1, L 4-6 1 5-6. M1 23-29, M2 15-20 (N = 3); intestine forming a long loop in roof of mantle cavity, the loop is additionally coiled, its appearance seems to vary depending on the presence and number of fecal pellets. FEMALE GENITALIA. Not observed; the only live collected female specimen was subadult, hence, the distal genitalia were not yet developed. Fig. 12. Holotypes. A–C . H. lozekiana sp. nov. (NHMW 113638). D–E. H. steffeki sp. nov. (NHMW 113640). Scale bar = 500 µm. MALE GENITALIA ( Figs 17A–B , 18 ). Not observed except for kidney-shaped prostate and penis; penis massive, trapezoidal, left side flattened, small, brown, probably chitinous stylet anterior right near penial opening. Habitat The type locality is a small permanent karst spring emerging from the small karst massif of Drienok formed by Middle Triassic (Aegean–Ladinian) Wetterstein, Reifling and Gutenstein Limestones of the Silicikum Unit (Silica Nappe) at the contact zone to impermeable Early Triassic shales and sandstones of Szin and Bódvaszilas Members with a small discontinuous stripe of alluvial gravel at the bottom of the spring valley. The character of the type locality indicates the same habitat preference in the phreatic rhizosphere as discussed for H. kissdalmae . Distribution Known only from the type locality so far. Conservation There is only one known location and the area of occupancy is probably considerably smaller than 20 km 2 . The combination of the restricted range and the threat imposed by the apparent trend of logging in the area around the type locality qualify for a threat assessment of Vulnerable (VU, D2) according to the IUCN Standards and Petitions Committee (2019) . Remarks The genetic distances of H. lozekiana sp. nov. from the more widespread H. kissdalmae and the Austrian species suggest that it is a relict probably from the earlier colonization of the Hron River Valley ca 4.5 Ma ago (see Discussion).