Hauffenia steffeki, Haase & Grego & Erőss & Farkas & Fehér, 2021

Haase, Martin, Grego, Jozef, Erőss, Zoltán Péter, Farkas, Roland & Fehér, Zoltán, 2021, 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, European Journal of Taxonomy 775, pp. 143-184 : 166-174

Hauffenia steffeki sp. nov.

urn:lsid:zoobank.org:act:0637E21D-A9F8-429D-B025-2D3ED9E1E28E

Figs 2 , 5 72a–b, 12D–E, 13C–D, 14B, 15D–F, 16, 17C–E, 19; Tables 3–4

Diagnosis

Hauffenia steffeki sp. nov. is characterized by a valvatiform shell with eventually detached peristome and an only initially sculptured embryonic shell, a thin, orange operculum without peg-like structure in the nucleus and the massive, trapezoidal penis with a rounded lobe on the left side. There were nine diagnostic COI alignment positions of type 1, 13 of type 3, and 85 of type 4 ( Table 3 ). Uncorrected genetic distances to congeners ranged from 0.108 [ H. michleri (Kuščer, 1932) ] to 0.141 ( H. kissdalmae ; see Supp. file 3).

Etymology

Hauffenia steffeki sp. nov. is named after Jozef Šteffek (1 Jan. 1952 – 19 Apr. 2013), ever-helpful friend and unforgettable Slovak malacologist, initiator of the intensive field work dedicated to the genus Hauffenia in Slovakia.

Material examined

Holotype ( Fig. 12D–E ) BOSNIA AND HERCEGOVINA • Bosnian Federation , Bihać district, Donji Kamengrad, 4 km W of Sanski Most; 44°46.7682′ N, 16°33.9859′ E; alt. 228 m; 24 Mar. 2016; J. Grego, G. Jakab and B. Šmída leg.; spring and well in dry canyon; NHMW 113640 . ( Fig. 5 72a, b)

Paratypes ( Fig. 13C–D ) BOSNIA AND HERCEGOVINA – Bosnian Federation • 13 specs; same collection data as for holotype; JG F0679 • 10 specs; same collection data as for holotype; NHMW 113641 • 8 specs; same collection data as for holotype; 4 Apr. 2015; J. Grego, M. Grego, R. Lohaj and B. Šmída leg.; JG F0473 • 10 specs; same collection data as for holotype; 24–25 Mar. 2016; J. Grego, M. Olšavský, G. Jakab and B. Šmída leg.; HNHM 105308 • 33 specs; same collection data as for holotype; JG F0561 • 10 specs; same collection data as for holotype; NMBE 569389 • 10 specs; same collection data as for holotype; SMF 363256 • 10 specs; same collection data as for holotype; SMOPAJ 2/2021 • 10 specs; same collection data as for holotype; ZMH 140885 • 35 specs; Bosanska Krajina , Sanski Most district , Dabarska Pećina , cave in Donji Dabar near Bankovići; 44°42.6′ N, 16°38.31′ E; alt. 173 m; 25 Mar. 2016; J. Grego, M. Olšavský ¸ G. Jakab and B. Šmída leg; sand on cave river bottom; JG0565 • 68 specs; same collection data as for preceding; 15 Mar. 2017; J. Grego, G. Jakab and B. Šmída leg.; JG F0692 • 12 specs; Bihać district , Spring Krivina , Lušči Palanka-Miljevci, 400 m from village toward Sanski Most, at right side of road R405; 44°45.7345′ N, 16°27.0528′ E; alt. 413 m; 15 Mar. 2017; J. Grego, G. Jakab and B. Šmída leg; JG F0675 • 3 specs; Bihać district , Vrelo Zdena 3 km W of Sanski Most; 44°45.7186′ N, 16°37.8371′ E; alt. 176 m; 15 Mar. 2017; J. Grego, G. Jakab and B. Šmída leg.; JG F1888 • 7 specs; Bihać district , Izvor Banovac between Gorice and Naprelie, left side of the road; 44°46.8065′ N, 16°30.0365′ E; alt. 375 m; 15 Mar. 2017; J. Grego, G. Jakab and B. Šmída leg.; JG F0668 .

Other material tentatively allocated to H. steffeki

BOSNIA AND HERCEGOVINA – Bosnian Federation • 1 spec.; Bihać district , Bastaško Vrelo, 5 km W of Drvar; 44°23.584′ N, 16°19.572′ E; alt. 527 m; 15 Mar. 2017; J. Grego, G. Jakab and B. Šmída leg.; JG F 0743 . – Republika Srpska • 7 specs; Donja Pećka , Izvor Sana; 44°19.0302′ N, 16°50.1773′ E; alt. 446 m; 15 Mar. 2017; J. Grego, M. Olšavský, G. Jakab and B. Šmída leg.; sand on bank of river; JG F 0571 /.

Description

SHELL ( Figs 12D–E , 13C–D , 14B ; Table 4 ). Valvatiform, maximum height measured at type locality 1.08 mm, maximum width 1.77 mm, with up to 2.875 whorls, on average twice as wide as high, considerably smaller in Krivina and Dabarska Pećina, there also flatter, final part of last whorl may be detached, fairly variable in shape with a coefficient of variation of 12.87% for the height/width ratio at the type locality, less pronounced at the other two localities where measurements have been taken; embryonic shell with 1.1–1.2 whorls, first 0.75 whorls faintly wrinkled, then smooth; aperture almost round only slightly higher than wide, ortho- or prosocline, lip continuous, not thickened; umbilicus wide.

OPERCULUM ( Fig. 19 ). Thin, chitinous, orange, almost circular with central nucleus, the latter without peg or peg-like modification.

EXTERNAL FEATURES. Epidermis without pigment, no eye spots visible, tentacles without particular ciliation.

MANTLE CAVITY. 9–12 gill filaments (N = 2), osphradium short, oval, behind middle of gill. DIGESTIVE SYSTEM ( Figs 15D–F , 16 ). Radula with formula R 5-6 1 5-6/1 1, L 5 1 5, M1 22-24, M2 17- 19 (Donji Kamengrad, N = 1), R 5 1 5/1-2 1-2, L 4-5 1 5, M1 22-26, M2 19-23 (spring Krivina, N = 2); 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; all live collected specimens were male.

MALE GENITALIA ( Fig. 17C–E ). Testis starting at least one whorl below apex, comprising ca 0.25 whorls, reaching stomach; prostate kidney shaped; penis massive, trapezoidal, with rounded lobe on left side, small, brown, probably chitinous stylet anterior right near penial opening.

Habitat

The type locality is a small well in a meadow in a dry valley west of Donji Kamengrad associated to a nearby karst spring at the north side of the valley. The Dabarska Pećina is a large river cave, one of the two main sources of the Dabar River (left tributary of the Sana River), draining the karst waters from the plateaus above the Jezernica basin through Bobijaško oko Estavella and numerous sinkholes. The other localities are rather smaller permanent karst springs and wells associated to small alluvial stripes with forested surroundings. These localities suggest a habitat preference for the phreatic rhizosphere as discussed for H. kissdalmae . However, the live specimens were also found in sediment on the bottom of the cave which indicates also a wider subterranean distribution including open cave conduits in a similar way as suggested for the species of Hauffenia from Slovenia, Italy and Croatia ( Bole 1970; Bole & Velkovrh 1987; Rysiewska et al. 2016).

Distribution

Known only from the type locality and four nearby localities at Bosanska Krajina west and northwest of Sanski Most.

Conservation

The new species is known from only five locations and it is very probable that the area of occupancy is smaller than 20 km 2. At this stage, we could not identify any plausible threats so that H. steffeki sp. nov. is assessed as Near Threatened (NT; IUCN Standards and Petitions Committee 2019).

Remarks

Hauffenia steffeki sp. nov. extends the distribution of the genus further south and is genetically/ phylogenetically distinct from all other species for which sequence information is available. It is likely that the species has a wider distribution in springs draining to and in the alluvium of the Sana River up to its source. In the type locality, it is syntopic with Lanzaia bosnica Bole, 1970 and in Dabarska Pećina with stygobiotic gastropods including Dabriana bosniaca Radoman, 1974 , Lanzaia bosnica Bole, 1970 and Paladilhiopsis blihensis ( Glöer & Grego, 2015) ( Glöer & Grego 2015) .

Figures

Fig. 2. Localities in Slovakia (A) and Hungary (B). Abbreviation: LT = type locality.

Fig. 5. Type locality of H. lozekiana sp. nov. (71) and Dabarska Pećina, a locality of H. steffeki sp. nov. (72) (see Table 1).

Fig. 12. Holotypes. A–C. H. lozekiana sp. nov. (NHMW 113638). D–E. H. steffeki sp. nov. (NHMW 113640). Scale bar = 500 µm.

Fig. 13. Paratypes from type localities (SEM micrographs). A–B. H. lozekiana sp. nov. (NHMW 113639). C–D. H. steffeki sp. nov. (NHMW 113641). Scale bar = 500 µm.

Fig. 14. Protoconchs (SEM micrographs). A. H. lozekiana sp. nov., paratype (NHMW 113639). B. H. steffeki sp. nov., paratype (NHMW 113641). Scale bars = 100 µm.

Fig. 19. Operculum of H. steffeki sp. nov., from type locality; internal side. Scale bar = 100 µm.

Fig. 15. Radulae (SEM micrographs). A–C. H. lozekiana sp. nov. D–F. H. steffeki sp. nov. D. Type locality. E–F. Spring Krivina (locality 74 from Table 1). Scale bars: A = 10 µm; B–E = 2 µm; F = 3 µm.

Fig. 16. Schematic representations of intestinal loop configurations.

Fig. 17. Penes (SEM micrographs). A–B. H. lozekiana sp. nov. (different specimens). C–E. H. steffeki sp. nov. (one specimen). Arrow points at penial stylet, which is withdrawn in A and B. Scale bars: A–C = 50 µm; D = 10 µm; E = 2 µm.