Zospeum robustum Inäbnit, Jochum & Neubert, 2019

Inäbnit, Thomas, Jochum, Adrienne, Kampschulte, Marian, Martels, Gunhild, Ruthensteiner, Bernhard, Slapnik, Rajko, Nesselhauf, Claudia & Neubert, Eike, 2019, An integrative taxonomic study reveals carychiid microsnails of the troglobitic genus Zospeum in the Eastern and Dinaric Alps (Gastropoda, Ellobioidea, Carychiinae), Organisms Diversity & Evolution 19 (2), pp. 135-177 : 165-166

Zospeum robustum Inäbnit, Jochum & Neubert n. sp.

Figure 9f–n

Type specimens Holotype NMBE 548791 , Markov ponor, 20.8.2011, leg. Raguz; paratypes: NMBE 548777 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548778 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548779 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548780 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548781 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548782 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548783 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548784 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548785 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548786 /1, Markov ponor, 20.8.2011, leg. Raguz; NMBE 548787 /1, Markov ponor, 20.8.2011, leg. Raguz.

Specimens examined Croatia: NMBE 548797 /1, Budina špilja, 29.1.2011, leg. Bregovic ; NMBE 553352 /4, Budina špilja, 11.6.2011, leg. Raguz ; NMBE 553353 /3, Budina špilja, 29.1.2011, leg. Bilandzija ; NMBE 553355 /1, Budina špilja, 24.7.2007, leg. Pavlec ; NMBE 548790 /1, Ostrvička špilja, 12.6.2011, leg. Cukovic ; NMBE 553354 /1, Ostrvička špilja, 12.6.2011, leg. Bilandzija ; NMBE 553356 /1, 29.7.2010, leg. Bedek ; NMBE 548773 /1, Budina špilja, 29.1.2011, leg. Bregovic ; NMBE 548809 /1, Ostrvička špilja, 22.8.2009, leg. Lukic ; NMBE 553534 , Vrlovka , 7.6.2009 ,

Mačkovom dragom, sh: 1.562 mm. — (i) Holotype, NMBE 548791 , Markov ponor, sh: 2.15 mm. — (j1–2) NMBE 553352 , Budina špilja, sh: 1.826 mm. — (k1–2) NMBE 553534 , Vrlovka, sh: 1.727 mm. — (l) NMBE 553367 , Vrlovka, sh: 1.595 mm. — (m) NMBE 554399 , Židovske kuće, sh: 1.51 mm. — (n) NMBE 554400 , Pušina jama, sh: 1.61 mm. — All phot. × 20

leg. Lukic; NMBE 553367 /16, Vrlovka , 7.6.2009 , leg. Lukic; NMBE 548799 /7, Vrlovka , 19.3.2009 , leg. Bedek; NMBE 553364 /1, Vrlovka , 19.3.2009 , leg. Bedek; NMBE 553365 / 10, Vrlovka , 6.6.2009 , leg. Bedek; NMBE 553366 /1, Vrlovka , 17.3.2009 , leg. Bedek; NMBE 548776 /1, Vrlovka , 17.9.2009 , leg. Cvitanovic; RS 2210a/1, Vrlovka , 2.12.2015 , leg. Ozimec; NMBE 554399 /1, Židovske kuće, 12.4.2008 , leg. Kljaković; NMBE 554400 /1, Pušina jama, 20.4.2008 , Pavlek; NMBE 553536 , Tonkovića špilja, 24.7.2011 , leg. Bedek; NMBE 554397 /1, Tonkovića špilja, 24.7.2011 , leg. Bedek; NMBE 554398 /1, Tonkovića špilja, 24.7.2011 , leg. Bedek; NMBE 553368 /1, Špilja pod Mačkovom dragom, 11.9.2009 , leg. Lukic.

Diagnosis Shell ca. 1.6 mm, ovate-conic, transparent, last whorl sometimes expanded, peristome roundish, sometimes quite massive, with a straight parietal shield, columella with a parietalis and a weak columellaris.

Measurements (n = 51) sh: 1.45–2.15 mm (mean: 1.649 ± 0.124 mm); sw: 0.99–1.54 mm (mean: 1.108 ± 0.0911 mm); ah: 0.76–1.32 mm (mean: 0.863 ± 0.083 mm); aw: 0.62– 1.13 mm (0.738 ± 0.073 mm); number of whorls: 4.75–6 (mean: 5.025 ± 0.231).

Description Shell ovate-conical, translucent when fresh; shell either smooth or ribbed, either at the uppermost part of the whorl or the entire whorl; shell shape variable: specimens from Markov ponor resembling Z. obesum , specimens from Tonkovića špilja resembling Z. frauenfeldii , others resemble more Z. pagodulum and Z. bucculentum ; aperture taller than wide; parietal shield differentiated from the lip, its margin either straight or, more rarely, concave; parietalis in most cases present in the aperture, though often fairly small, increasing in size for half a whorl; a weak columellaris usually starting behind the aperture; columellaris completely absent in ribbed populations.

Smaller, smooth forms (e.g., Budina špilja, Vlovka) not really separable from other smooth-shelled species within the Z. frauenfeldii clade (such as Z. frauenfeldii , Z. bucculentum , Z. pagodulum ); tentatively differing from Z. bucculentum by its smaller size and its less shouldered whorls; the form from Markov ponor differs from Z. obesum by the presence of ribs on the upper part of the whorl; the form from Tonkovića špilja and Špilja pod Mačkovom dragom differ from Z. frauenfeldii and Z. pagodulum by their lack of a columellaris.

Etymology Named for the solid, robust shell of the Holotype (lat.: robustum: robust)

Distribution (Supplementary Fig. S12) Occurs in Croatia in three disjunct areas: 1) The Lika-Senj region, for which we have genetic records for two caves (Budina špilja ( Fig. 9j ) and Markov ponor ( Fig. 9i )). Two additional populations are tentatively assigned to this species, including Ostrvička špilja and the upper part of the Lukina jama-Trojama cave system (see Fig. 4C in Weigand 2013)); 2) Two caves north of Ogulin, one with a genetic record (Tonkovića špilja ( Fig. 9f, g )) and one without (Špilja pod Mačkovom dragom ( Fig. 9h )); 3) Three caves in and around the Žumberak Mountains (Vrlovka ( Fig. 9k, l ), Pušina jama ( Fig. 9n ) and Židovske kuće ( Fig. 9m )).

Remarks The three disjunct areas are reflected in the phylogenetic trees, albeit with low node support. Although future studies might find them to be independent species, our sample size does not allow definite conclusions.

Figures

Fig. 9 (a–e) Z. pagodulum, (a1–2) Holotype, NMBE 548815, Kučka jama, sh: 1.982 mm. — (b) NMBE 553363, Jama ispod Tominićevog brega, sh: 1.759 mm. — (c) NMBE 553361, Pećina kod sela Puhari, sh: 1.781 mm. — (d) NMBE 553360, Lučićka špilja, sh: 1.613 mm. — (e1– 2) NMBE 548789, Grnjača špilja, sh: 1.765 mm. —(f–n) Z. robustum, (f1–2) NMBE 553536, Tonkovića špilja, sh: 1.826 mm. — (g) NMBE 554397, Tonkovića špilja, sh: 1.52 mm. —(h) NMBE 553368, Špilja pod

Fig. 4 Zospeum spelaeum. a lectotype SMF 158543, Postojnska jama, sh: 1.634 mm. — b1–6 NMBE 553308, Betalov Spodmol, sh: 1.8 mm; c ditto, sh: 1.495 mm. — d NMBE 553306, Betalov Spodmol, sh: 1.8 mm. — e NMBE 553310, Mačkovica jama, sh: 2.39 mm. — f1–2 NMBE 553312, Velika Pasica, sh: 2.055 mm. —(g) NMBE 553311, sh: 1.957 mm; h ditto, sh: 2.07 mm. — i NMBE 553304, Čampari jama,