Zospeum spelaeum (ROSSMAESSLER, 1838)

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 : 141-150

Zospeum spelaeum clade

Distribution (Supplementary Fig. S8 ) This clade occurs in the northern Dinarides of northwestern Croatia and Slovenia as well as in the region of Ljubljana, the Julian Alps and the Italian Prealps east of the Brenta River.

Both genetic trees have the same topology ( Fig. 3 , red part). We follow here the delimitations C-E ( Table 1) and accept two species: Z. spelaeum with genetic data from four caves and two specimens from Jama 2 pri Jabljah, which are classified here with Z. costatum due to morphological similarities and geographical proximity to the type locality. The topology within Z. spelaeum is not well-supported in either tree.

Morphology Shell conical, translucent when fresh, in most cases comparatively taller than other clades; whorl often shouldered; ribs in most cases present, though very variable; aperture taller than wide; parietal shield well-differentiated from the lip, not broad, its margin generally straight, sometimes slightly convex or concave; upper part of the palatal margin of the aperture expanded; parietalis and columellaris present in aperture and well-developed; angularis and palatalis present. Differing from all other clades through the presence of an angularis and a palatalis.

Differing in the LDA from the Z. obesum clade by the ratios aw/D1 ( Z. spelaeum clade: 1.389 –2.225, Z. obesum clade: (1.718–) 2.2–2.703) and sh/sw ( Z. spelaeum clade: 1.338 – 1.934, Z. obesum clade: 1.208 –1.455); from the Z. pretneri clade by the ratios cjl/D1 ( Z. spelaeum clade: 1.456 –2.075, Z. pretneri clade: 0.837–1.42) and lh/lw ( Z. spelaeum clade: 1.01–1.339, Z. pretneri clade: 0.884 –1.045).

None of the species could be clearly separated via the measurement-based PCA (Supplementary Fig. S2 , top).

The geometric PCA (Supplementary Fig. S2 , middle) separated Z. costatum , Z. lamellatum and Z. trebicianum from each other through a combination of the PC1- and PC2-axes.

The CVA (Supplementary Fig. S2 , bottom) separated Z. lamellatum from all the other species on the CV1-axis (Shape changes from negative to positive: apex getting more blunt (landmark 10 approach landmarks 9 and 11); parietalis getting smaller (landmark 14 extending more to the upper left)). The CV2-axis separated Z. lautum from the rest (Shape changes from negative to positive: palatal side of the aperture ceasing to stand out (landmark 2 extending more to the left); last whorl decreasing in size (landmarks 7, 8 extending to the right)).

Zospeum spelaeum (ROSSMAESSLER, 1838)

Figures 4a–n, S 14a–b, S20a–f, S21a–c

1838 Carychium spelaeum ROSSMAESSLER, Icon. , (1) 2 (1/2): 36–37, pl. 49 fig. 661 (shell) [in der Aldelsberger Höhle in Krain an Stalaktiten (Postojnska jama, near Postojna)].

1854 Carychium schmidtii FRAUENFELD, Verh. zool-bot. Ges. Wien, Abh., 4 (1): 32, 33, 34, pl. [4] fig. 5 (shell) [Aus der PasizaGrotte (F.J. SCHMIDT leg.; Velika Pasica, near Gornji Ig)].

1855 Carychium pulchellum FREYER, Sitzungsber. mathem.-naturw. Cl. kais. Akad. Wiss. Wien, 15 (1): 20, pl. [1] fig. 4a–d (shell) [In der Grotte am Krimberge bei Laibach zuerst aufgefunden von Anton KUKEK (Krimska jama, near Gorenja Brezovica)] .

1856 Zospeum aglenum BOURGUIGNAT, Rev. Mag. Zool. , (2) 8 (11): 507–508 [la caverne de Pasiza, en Carniole (Velika Pasica, near Gornji Ig)].

1856 Zospeum nycteum BOURGUIGNAT, Rev. Mag. Zool. , (2) 8 (11): 509–510 [la caverne de Pasiza, en Carniole (Velika Pasica, near Gornji Ig)].

1856 Carychium reticulatum HAUFFEN, Verh. zool.-bot. Ges. Wien, Abh., 6: 623–624, pl. 7 fig. 4 (shell) [In der Grotte Bidou sturm und in einer hinter Laak (HAUFFEN leg.; Dobruška jama, near Vodice)] (see Hauffen 1856a).

1858 Carychium carinatum HAUFFEN, Jahresh. Ver. krain. Landes-Mus., 2: 96 [Sie kommt in der Grotte Bidov sturm und in einer hinter Laak vor (Dobruška jama, near Vodice; Communicated at a meeting on 11 June 1856, but published in 1858; clearly reticulatum is meant)].

1889 Zospeum tellinii POLLONERA, Bull. Soc. malac. ital., 14 (2): 49–50, pl. 2 fig. 6 (shell) [nelle posature del Natisone nel Friuli ... a nord di Cividale (A. TELLINI leg., 1 ex.; sieved from the Natisone river north of Cividale )] .

1899 Zospeum auritum STOSSICH, Boll. Soc. adriat. sci. natur. Trieste, 19: 41 + 3 fig. (shell) [Scoperta dal Dott. MOSER nella grotta di Corgnale (? Near Lokev, Slovenia)].

1899 Zospeum istrianum STOSSICH, Boll. Soc. adriat. sci. natur. Trieste, 19: 42 + 3 fig. (shell) [Grotta di San Servolo; rinvenuta dal Dott. MOSER (Sveta jama, near Socerb)].

1905 Zospeum lyratum POLLONERA, Proteus , 3 (1): 4, fig. 1 (shell) [nella posatura del Natisone, nel Friuli, alquanto a nord di Cividale (A. TELLINI leg., 1 ex.; sieved from the Natisone river north of Cividale )] .

1905 Zospeum venetum POLLONERA, Proteus , 3 (1): 3–4, fig. 2 (shell) [nella posatura del Natisone, nel Friuli, alquanto a nord di Cividale (A. TELLINI leg.; sieved from the Natisone river north of Cividale)].

1912 Zospeum alpestre rossmaessleri A.J. WAGNER, Verh. kais. königl. zool.-bot. Ges. Wien, 62 (8/9): 257–258 [Johannesgrotte in den Adelsberger Höhlen (Postojnska jama, near Postojna)] .

Type specimens spelaeum : Lectotype (by subsequent designation of ZILCH, 1959, Handb. Paläozool., 6 (2, 1): 65, fig. 206) SMF 158543, ROSSMAESSLER leg., October 1835, ± 20 exx. schmidtii : syntype NHMW 71844/1 (imaged by K. Jaksch-Mason 2013). A shell ( NHMUK 1991027), mentioned by Jochum et al. (2015b fig. 6G) as a syntype, can not be unambiguously verified as such. NHMW /MO/13032: the label record contains a label stating B carniolica^, which is a manuscript name forwarded to Frauenfeld by Schmidt. Frauenfeld Von (1854: 34) explains that he does not accept this name and names the species B schmidtii^ instead. Thus, this specimen is considered a syntype of schmidtii ; pulchellum : collection Freyer in Trieste contains no Zospeum specimens (pers. comm. De Mattia 2017), type specimens probably lost; aglenum : replacement name for Zospeum lautum sensu Freyer 1855 non lautum Frauenfeld, 1854 ; no Zospeum specimens in coll. Freyer in Trieste, type specimens probably lost; nycteum : based on Carychium obesum sensu FREYER (1855: 21 , pl. 1 fig. 6a–c B Aus der Pasica-Höhle bei Sonneg in Krain; gesammelt von Herrn ERJAVEC^) (non FRAUENFELD), type specimen lost. reticulatum : no type specimens found in NHMW nor in Ljubljana; carinatum no type specimens found in NHMW nor in Ljubljana; tellinii : MZUT Turin; specimen inaccessible, no information available (pers. comm. Elena Gavetti); auritum : Stossich collection is lost ( De Mattia 2004), no type specimens available; istrianum : Stossich collection is lost ( De Mattia 2004), no type specimens available; lyratum : MZUT Turin; specimen inaccessible, no information available (pers. comm. Elena Gavetti); venetum : MZUT Turin; specimen inaccessible, no information available (pers. comm. Elena Gavetti); rossmaessleri: syntype NHMW 69394.

Specimens examined Italy: NMBE 553314 /1, Grotta d’ Ercole, 14.6.2011, leg. Prodan ; NMBE 553315 /1, Grotta d’ Ercole, 14.6.2011, leg. Prodan ; NMBE 553316 /1, Grotte Bac , 22.9.2011, leg. Prodan ; NMBE 553317 /1, Grotte Bac , 22.9.2011, leg. Prodan ; NMBE 554704 /4, Grotta Valentina , leg. De Mattia ; NMBE 554705 /2, Grotta dell’ Alce, leg. De Mattia ; NMBE 554706 /2, Grotta delle Gallerie , leg. De

sh: 1.791 mm. — j NMBE 553303, Jama pod križ, sh: 1.782 mm. — k NMBE 553301, Vrlovka, sh: 2.17 mm. — l NMBE 553302, špilja Pećina pod Stržen, sh: 2.03 mm. — m NMBE 553316, Grotte Bac, sh: 2 mm. — n NMBE 554711, Grotta di Boriano. sh: 2.62 mm. — All phot. × 20; a S. Hof, SMF; (b6) M. Kampschulte, G. Martels, University of Giessen

Mattia; NMBE 554707 /4, Grotta dell’ Orso, leg. De Mattia ; NMBE 554708 /6, Caverna presso Basovizza, leg. De Mattia ; NMBE 554710 /2, Grotta Bac , leg. De Mattia ; NMBE 554711 /2, Grotta di Boriano , leg. De Mattia ; NMBE 554712 /2, Grotta di Padriciano , leg. De Mattia ; NMBE 554713 /3, Grotta Romana , leg. De Mattia ; NMBE 554714 / 3, Grotta delle Torri di Slivia , leg. De Mattia ; NMBE 554715 /3, Grotta di Crogole , leg. De Mattia ; NMBE 554716 /2, Grotta ad E di S. Martino del Carso, leg. De Mattia ; NMBE 554717 /3, Abisso di Bonetti , leg. De Mattia ; NMBE 554718 /4, Antro della Biscia Morta , leg. De Mattia ; NMBE 554719 /1, Grotta delle Radici , leg. De Mattia ; NMBE 554721 /1, Grotta Rosica , leg. De Mattia ; NMBE 554722 /3, Grotta Azzurra , leg. De Mattia ; NMBE 554723 /2, Grotta d’ Ercole, leg. De Mattia ; NMBE 554724 /3, Sorgenti di Vedronza , leg. De Mattia. — Slovenia : NMBE 553306 /23, Betalov Spodmol , 4.10.2007, leg. Jochum ; NMBE 553307 / 8, Betalov Spodmol , 4.10.2007, leg. Jochum ; NMBE 553308 / 19, Betalov Spodmol , 4.10.2007, leg. Jochum ; NMBE 553309 /14, Betalov Spodmol , 4.10.2007, leg. Jochum ; NMBE 553311 /22, Velika Pasica , 2.9.2010, leg. Jochum ; NMBE 553312 /3, Velika Pasica , 2.9.2010, leg. Jochum ; NMBE 553313 /1, Velika Pasica , 2.9.2010, leg. Jochum ; NMBE 553310 /1, Mačkovica , 26.6.2009, leg. Jochum ; NMBE 554396 , Hotiške Ponikve , 30.10.2012, leg. Polak ; NHMW 71838 /2, no further data available ; NMBE 554709 / 4, Jama v Bukovniku, leg. De Mattia ; NMBE 554720 /3, Miškotova jama v Lokah, leg. De Mattia ; NMBE 554725 /1, Miškotova jama v Lokah, leg. De Mattia. — Croatia: NMBE 553301 /5, Vrlovka , 17.3.2009, leg. Bedek ; NMBE 553304 /1, Čampari jama, 29.11.2008, leg. Raguz ; NMBE 553303 /4, Jama pod križ, 27.6.2009, leg. Lukic ; NMBE 553302 /1, Pećina pod Stržen, 27.6.2009, leg. Bregovic ; NMBE 553305 /5, Pećina pod Stržen, 27.6.2009, leg. Bilandzija.

Diagnosis Shell ca. 1.96 mm, transparent, elongate or elongate-conical with an auriform and more or less thickened peristome, bearing two to three apertural barriers; columella with a single lamella.

Measurements (n = 148) sh: 1.49–2.62 mm (mean: 1.962 ± 0.223 mm); sw: 0.88–1.5 mm (mean: 1.198 ± 0.135 mm); ah: 0.68–1.25 mm (mean: 0.988 ± 0.131 mm); aw: 0.58– 1.04 mm (mean: 0.796 ± 0.097 mm); number of whorls: 4.5–6.25 (mean: 5.18 ± 0.306).

Description Very variable species. Shell conical, often more slender than other species, translucent when fresh; whorl nev- er well rounded (as, for example, in Zospeum frauenfeldii ), usually ovate-conic in form; shell surface usually ribbed, but variable, which has been used to differentiate subspecies in the past (present on the whole whorl as in B Z. s. costatum^, present on the upper part of the whorl as in B Z. s. schmidti^ ( Fig. 4f–n ) or completely absent as in B Z. s. spelaeum^ ( Fig. 4a–e )), but the spectrum of variation within populations is too broad to establish congruence here; aperture taller than wide, the palatal side usually shouldered; the parietal shield well-differentiated from the lip, its shape either convex or straight; parietalis well-developed in the aperture, extending one whorl into the shell; columellaris visible in the aperture, in some cases ( Fig. 4b ) disappearing within half a whorl into the shell or continuously weakening ( Fig. 4e ); a more or less oblique lamella is present above the middle of the short columella; a basal bulge is present just above the umbilicus ( Fig. 4f 2 ); angularis in most cases present as a tooth of variable prominence; palatalis usually present. Protoconch covered with pits (Supplementary Figs. S14a–b); interconnected pits on top of the protoconch densely aggregated into rows; pits on the side of the whorl distributed more randomly. Radula: The radula of topotypic Zospeum spelaeum (Postojna; Supplementary Figs. S20a–d) is very different from that of Zospeum spelaeum from Velika Pasica (Supplementary Figs. S20e–f; S21a–c). In topotypic Z. spelaeum (collected by Bole in 1968), the right laterals mostly bear two pointed cusps of uneven length, whereby the left cusp is ¼ longer than the right (Supplementary Fig. S20b). An occasional third, stubby cusp may be present. On the other hand, the equally pointed left laterals are either triscuspid or bicuspid, whereby the median cusp is 1/2 cusp shorter than the longer flanking two cusps of the tricuspid crowns. The crowns of the transitional teeth consist of 5–6 very pointed subequal cusps, which do not extend beyond the edge of the individual, concave basal plates (Supplementary Fig. S20c). The broad marginal teeth are highly and unevenly serrated with very pointed cusps (Supplementary Fig. S20d). The massive, irregular basal plates of the marginals meld into eachother with occasional spaces in between the individual plates. The claw-like crown bears up to 10 pointed cusps of different widths and lengths (Supplementary Fig. S20d). The radular ribbon is long (Supplementary Fig. S20a). Since it folded and ripped upon positioning on the stub, the ends are undecipherable. On the other hand, the radula of Z. spelaeum from Velika Pasica shows multiple rows of obtuse, tricuspid crowns of which the third cusp is a mere nub of an ectocone very close to the top of the basal plate (Supplementary Fig. S21a). The two major cusps are medially grooved and show a wrinkled, marbled-like surface texture. The cusps are melded together and are only partially detached from the middle of the crown. The basal plates are concave and show wing-like prominences (Supplementary Fig. S20f). The obtuse transitional teeth are quadricuspid and sometimes quincuspid of almost equal lengths except the pointed, outermost endocones and ectocones, which are about 1/3–1/2 the length of the preceding, equal-lengthed cusps (Supplementary Fig. S21b). Many cusps are grooved. Each cusp of the transitional teeth is partially detached midway through the crown. The marginal teeth bear 6–7 cusps on each crown of which the furthest endocones and ectocones are sometimes mere stubs. The marginal teeth also show cusp detachment midway from the middle of the crown to the tip of each cusp (Supplementary Fig. S21)c. A median groove is visible on some of the marginal cusps. Laterally, the crowns of the marginal teeth extend beyond the edge of each basal plate by two cusps. The radular ribbon is long and tapered at both ends (Supplementary Fig. S20e).

Differing from Z. costatum by a less expanded upper part of the palatal side of the aperture; differing from Z. lamellatum by its greatly reduced external ribbing and number and size of lamellae and apertural barriers; usually differing from Z. lautum by a slenderer shell, presence of ribs, and its bearing a simple inclined lamella versus the more elaborate 3-tiered lamellar configuration in Z. lautum ; differing from Z. trebicianum by its larger size. This species cannot be separated through PCA and LDA from its closest relatives ( Z. costatum , Z. lamellatum , Z. lautum , Z. trebicianum ).

Distribution (Supplementary Fig. S8 ) This species was report- ed from the region of Ljubljana westwards to the Brenta River in Italy ( Bole 1974, Giusti and Pezzoli 1982, Pezzoli 1992, Figs. 4a–h ( Slovenia only)), southwards in the region around Trieste ( De Mattia 2003, Figs. 4m, n ), in Istria ( Slapnik and Ozimec 2004, Figs. 4j +l) and on the island of Cres (this study, Fig. 4i ). Eastwards, this species was reported from along and just north of the Sava River and to about 14° 00′ E where the Sava River enters the Julian Alps. An isolated record was made from the Vrlovka cave in the northeastern part of the Karlovac region, Croatia ( Bole 1974, Fig. 4k ).

Zospeum costatum ( FREYER, 1855)

Figure 5a–f

1855 Carychium costatum FREYER, Sitzungsber. mathem.- naturw. Cl. kais. Akad. Wiss. Wien, 15 (1): 20, pl. [1] fig. 5a–c (shell) [Von Herrn Ferd. SCHMIDT in der Grotte bei Goričane unweit des fürstbischöflichen Schlosses Görtschach entdeckt (Babja Luknia, near Goričane)].

1856 Carychium bidentatum HAUFFEN, Verh. zool.-bot. Ges. Wien, Abh., 6: 701, pl. 7 fig. 5 (shell) [in der Grotte

Grotte von Klince/Utik (mentioned in Frauenfeld 1856), sh: 1.75 mm. — m NHMW 71834, Mala Bukovije (glatt, No 5 in Frauenfeld 1856), sh: 1.599 mm. — n–r Zospeum trebicianum , n NMBE 554702, Grotta di Trebicianum, sh: 1.525 mm; o ditto, sh: 1.466 mm. — p NMBE 554701, Grotta Lazzaro Jerko, sh: 1.639 mm. — q NHMW MO75000- E-46317, Cave at Trebic, sh: 1.63 mm; r sh: 1.66 mm. — All phot. × 20; a, b, q, r P. Albano, NHMW; c1–3, g2–3 B. Ruthensteiner, ZSM; l1, m E. Bochud, NMBE; l 2 M. Kampschulte, G. Martels, University of Giessen am Glaven vrh B (HAUFFEN leg., 17 exx.; Ručigajev izvir (Jama pod Ručigajem), near Dobeno)] (see Hauffen 1856b).

Type specimens costatum : collection Freyer in Trieste contains no Zospeum specimens (pers. comm. De Mattia 2017). bidentatum : no type specimens found neither in NHMW nor in Ljubljana;

Specimens examined Slovenia: NHMW 71848 /2, Babja Luknja , leg. Schmidt & Erjavec; NHMW 71847 /1, Babja Luknja , leg. Schmidt & Erjavec; NMBE 553383 /8, Jama 2 pri Jabljah, 25.6.2009, leg. Jochum; NHMUK 1939-6 - 14.504 -506, 3 shells, Babja Luknja B loc. class.!^; NHMUK 1935-1 - 1.280 -289, 10 shells, Jama pri Račigajevem mlinu (near Dobeno) (= Hauffen’ s orig. Loc. Cerote im B Glaven vrh^), ex Kuščer. According to the Cave registry of the Speleological Assoc. of Slovenia, this cave is most probably #1712 Ručigajev izvir (= Jama pod Račigajem) 46.142393°N 14.537118°E, 374 m.

Diagnosis Shell ca. 2 mm, transparent, ribbed, with weak spiral lines, elongate-conical with an auriform and more or less thickened peristome, bearing four apertural barriers, two of which continue on the columella as lamellae.

Measurements (n = 10) sh: 1.86–2.22 mm (mean: 2.002 ± 0.112 mm); sw: 1.2–1.44 mm (mean: 1.298 ± 0.088 mm); ah: 0.96–1.13 mm (mean: 1.026 ± 0.06 mm); aw: 0.83– 0.98 mm (mean: 0.896 ± 0.058 mm); number of whorls: 4.75–5.25 (mean: 5.075 ± 0.206).

Description Shell conical, translucent when fresh; whorls somewhat shouldered, usually stronger than in Z. spelaeum ; pronounced equidistantly-spaced axial ribs present on each whorl, though not always covering an entire whorl; weak spiral striae are usually visible (such as in Fig. 5b and e ); aperture taller than wide, the upper half of the palatal rim shouldered and expanded; the parietal shield well differentiated from lip, its margin more curved than straight; parietalis well developed in the aperture, extending one whorl into the shell; columellaris visible in aperture, extending ¾ of a whorl into the shell ( Fig. 5c 2 ); palatalis always present; angularis sometimes present.

Differing from Z. spelaeum by the generally more expand- ed upper palatal rim and the thickened basal columellar configuration; differing from Z. lamellatum by its considerably smaller lamellae and dentition; differing from Z. lautum by the marked presence of ribs and presence of a single lamella; differing from Z. trebicianum by its enhanced degree of ribbing and larger size.

Differing in the LDA from Z. lamellatum by the ratios ah/ aw ( Z. costatum : 1.06–1.19; Z. lamellatum : 1.25–1.35) and sh/ lh ( Z. costatum : 1.44–1.54; Z. lamellatum : 1.56–1.7); from Z. lautum by the ratio sw/ah ( Z. costatum : 1.224 –1.306, Z. lautum : 1.351 –1.445).

Distribution (Supplementary Fig. S8 ) Known from two caves north of Ljubljana: Babja Luknja near Goričane ( Fig. 5a–c ) and Jama 2 pri Jabljah near Loka pri Mengsu ( Fig. 5d–f ).

Remarks Considered a separate species here because of the distinct columellar configuration corroborated by genetic evidence ( Weigand et al. 2013).

Zospeum lamellatum BOLE, 1974

Figure 5g –k

1974 Zospeum spelaeum lamellatum BOLE, Razpr. Slov. Akad. Znan. Umetn., Cl. IV , 17 (5): 276–278, 288, fig. 17d– e (shell), 18, 19 (distribution) [Höhle Krasnica oberhalb Slap na Idrijci, 10 km SO von Tolmin].

Type specimens A holotype was not selected. We hereby, select MCSMNH 35995/1 (labeled B holotyp^), Krasnica, as lectotype.

Specimens examined Slovenia: MCSMNH 7057/5, Ukovnik , 21.10.1972 .

Diagnosis Shell ca. 1.8 mm, densely ribbed, transparent, elongate with prominent apertural barriers; columella bears a dense 3-tiered lamella configuration starting at the lower surface of the penultimate whorl and ending with the columellaris as a prominent basal lamella above the umbilicus (in dorsal cutout/CT view).

Measurements (n = 5) sh: 1.73–2.06 mm (mean: 1.81 ± 0.14 mm); sw: 1–1.2 mm (mean: 1.082 ± 0.075 mm); ah: 0.86–0.95 mm (mean: 0.903 ± 0.033 mm); aw: 0.66– 0.72 mm (mean: 0.692 ± 0.022 mm); number of whorls: 5.25–6 (mean: 5.5 ± 0.306).

Description Shell elongate-conical, translucent when fresh; whorls slightly shouldered; strongly ribbed, though rib density as well as rib prominence can vary considerably within a population ( Fig. 5h–k ); aperture is taller than wide, the palatal side not or only slightly shouldered; parietal shield well differentiated from the lip, its margin either straight or convex; parietalis and angularis very prominent, usually dominating the aperture; columellaris and palatalis both prominent as well, though to a lesser degree; palatalis and angularis do not extend into the whorl; the columellaris continuing with the same size as in the aperture for at least ½–¾ whorls; parietalis extending somewhat over one whorl into the shell, decreasing in size, though still more pronounced than in any other species ( Fig. 5g 3 ).

Differing from all other species through its remarkably large lamellae and teeth.

Differing in the LDA from Z. costatum by the ratios ah/aw ( Z. costatum : 1.06–1.19; Z. lamellatum : 1.25–1.35) and sh/lh ( Z. costatum : 1.44–1.54; Z. lamellatum : 1.56–1.7); from Z. trebicianum by the ratio sh/lh ( Z. lamellatum : 1.563 – 1.694, Z. trebicianum : 1.368–1.47).

Distribution (Supplementary Fig. S8 ) Bole (1974) reported the taxon from four caves in northwestern Slovenia.

Remarks Due to its morphological distinctness, this taxon is considered an independent species though not yet confirmed via genetic analysis.

Zospeum lautum ( FRAUENFELD, 1854)

Figure 5l–m

1854 Carychium lautum FRAUENFELD, Verh. zool-bot. Ges. Wien, Abh., 4 (1): 32, 33–34, pl. [4] fig. 4 (shell) [Aus der Grotte am Krimberg (F. J. SCHMIDT leg.; Krimska jama, near Gorenja Brezovica)].

Figures

Fig. 8 (a–e) Zospeum frauenfeldii, (a1–2) NMBE 553388, Podpeška Jama, sh: 1.982 mm. — (b1–2) NHMW 71882, Podpeška Jama, sh: 1.986 mm. — (c1–2) Paratype of Z. frauenfeldii osolei, MCSMNH 6835, Osoletova jama, sh: 2.01 mm; (d1–2) Paratype of Z. frauenfeldii osolei, ditto, sh: 1.805 mm. — (e1–2) NMBE 548795, Hrustovača špilja, sh: 1.494 mm. —(f–j) Zospeum subobesum, (f1–2) Syntype, MCSMNH 2083, Tounjčica, sh: 1.718 mm. — (g1–2) NMBE 553326, Tounjčica, sh: 1.655 mm. — (h) NMBE 553323, Sustav Đula Medvedica, sh: 1.637 mm. — (i) NMBE 553322, Jama ispod heliodroma, sh: 1.35 mm.

Fig. 3 The major clades recovered within the Dinaride and Alpine Zospeum radiation. Top: Bayesian inference tree; bottom: maximum likelihood consensus tree. * refers to individuals sequenced in this study

Fig. 2 Top: The nine measurements as illustrated on the frontal (left) and apical (right) view of a Zospeum manitaense shell (NMBE 549731). Bottom right: Position of the 14 landmarks on the shell. Bottom left: Nomenclature to dentition indicated on shell of Zospeum lamellatum (MCSMNH 35995)

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,

Fig. 5 a–f Zospeum costatum, (a) NHMW 71848, Babja luknja, Goričane, sh: 2.214 mm; (b) ditto, sh: 2.043 mm. — (c1–3) NHMW 71847, Babja luknja, Goričane, sh: 2.08 mm. — (d) NMBE 553383, Jama 2 pri Jabljah, sh: 1.86 mm; (e) ditto, sh: 1.92 mm; (f) ditto, sh: 2.03 mm. —(g–k) Zospeum lamellatum, (g1–3) Lectotype, MCSMNH 35995, Krasnica, sh: 2.059 mm. — (h) MCSMNH 7057, Ukovnik pri sp. Idriji, sh: 1.751 mm; (i) ditto, sh: 1.739 mm; (j) ditto, sh: 1.733 mm; (k) ditto, sh: 1.766 mm. —(l–m) Zospeum lautum, (l1–2) NHMW 71828,