Zospeum alpestre (FREYER, 1855)
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https://doi.org/ 10.1007/s13127-019-00400-8 |
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https://treatment.plazi.org/id/300F1A79-B426-B74E-E780-FE46FD9EFB13 |
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Felipe |
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scientific name |
Zospeum alpestre |
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Zospeum alpestre View in CoL clade
Distribution This clade is known from the Kamnik-Savinja Alps, the Julian Alps and the Karawanks of Slovenia, northeastern Italy and southeastern Austria (Supplementary Fig. S9 View Fig ).
The BI tree and the ML consensus tree are mostly identical ( Fig. 3 View Fig , blue part), excepting the lack of resolution of the position of Z. isselianum in the Bayesian tree. Here, we retain the classification of Jochum et al. (2015c; delimitation C (see Table 2)) since little additional material (MCSMNH 40651a; RS 2037) was included here and since the phylogenetic trees essentially group the same specimens together. We choose to separate Z. isselianum from Z. amoenum (despite delimitation E) due to the geographical separation and the lack of support in both trees.
Morphology Shell conical, translucent when fresh; shell size small to medium; shell surface smooth; aperture somewhat taller than wide; parietalis and columellaris usually present. Morphological overlap with the Z. frauenfeldii clade (tends to be larger) and the Z. obesum clade (usually has an expanded last whorl and spiral lines). Differing from the Z. spelaeum clade by the absence of an angularis and a palatalis. Differing from the Z. pretneri clade by the comparatively larger aperture.
Differing in the LDA from the Z. obesum clade by the ratios aw/D1 ( Z. alpestre : 1.437–2(–2.202), Z. obesum : (1.718–) 2.2 – 2.703) and sh/lh ( Z. alpestre : 1.387 –1.739, Z. obesum : 1.215 – 1.448).
Neither the PCAs nor the CVA were able to separate Z. amoenum from other species. The measurement-based PCA (Supplementary Fig. S3 View Fig , top) is able to separate Z. alpestre from Z. kupitzense and Z. isselianum on the PC1- axis, but not the latter two from each other. The geometric PCA separated Z. isselianum and Z. alpestre on the PC1- axis (Supplementary Fig. S3 View Fig , middle) and these two from Z. kupitzense on the PC2-axis. The CVA (Supplementary Fig. S3 View Fig , bottom) is able to separate the three aforementioned species on the CV1-axis.
Zospeum alpestre ( FREYER, 1855)
Figure 6a–e View Fig
1855 Carychium alpestre FREYER, Sitzungsber. mathem.- naturw. Cl. kais. Akad. Wiss. Wien, 15 (1): 19, pl. [1] fig. 2a–d (shell) [Im Jahre 1854 von Herrn Franz ERJAVEC in der Höhle Dioja griča, nächst der Veternica-Höhle auf der Velika planina der Steineralpen in Oberkrain aufgefunden (Dovja griča, Kamnik)].
1856 Zospeum nyctozoilum BOURGUIGNAT, Rev. Mag. Zool. , (2) 8 (11): 513–514 [les cavernes de Dioja-Grica et [sic!] de Veternica, en Carniole^ (Dovja griča, Kamnik). [reprint: 15–16, pl. 8 fig. 4–6 (shell; copy from FREYER)]].
1991 Zospeum alpestre bolei SLAPNIK, Razpr. Slov. Akad. Znan. Umetn., Cl. IV , 32 (1): 54–56, 61–63, fig. 5 + map 1 (distribution), pl. 1 fig. 3a–b, pl. 3 fig. 3 (shell) [Tomaževčeva zijalka, Podvolovljek (J. BOLE leg., 18-VI-1964)].
Type specimens Z. alpestre : collection Freyer in Trieste contains no Zospeum specimens (pers. comm. De Mattia 2017). nyctozoilum : partially based on Carychium alpestre FREYER (partim: pl. 1 fig. 2b, d; Im Jahre 1854 von Herrn Franz ERJAVEC in der Höhle Dioja griča, nächst der Veternica-Höhle auf der Velika planina der Steineralpen in Oberkrain aufgefunden); bolei: Holotype + 32 paratypes ( ZRC SAZU 3169) [synonymised by Jochum et al. 2015c].
Specimens examined: Slovenia NHMW 71817 View Materials / 2, B Juchantza^ (Ihanšica), leg. Tusek ; NHMW 71815 View Materials /5 ; NHMW 75000 View Materials /E12483/3, Dovja griča, leg. Kuščer. NMBE 553369 View Materials /1, Jama pod Mokrico, 5.5.2001, leg. Slapnik ; NMBE 553370 View Materials /1, Jama pod Mokrico, 29.8.2008, leg. Slapnik ; NMBE 553371 View Materials /2, Jama pod Mokrico, 18.4.2013, leg. Slapnik ; NMBE 553372 View Materials /3, Jama pod Mokrico, 18.4.2013, leg. Slapnik ; NMBE 553373 View Materials /3, Jama pod Mokrico, 2.10.2007, leg. Jochum ; MCSMNH 40651a, Jelenska zijalka, 23.7.2012, leg. Slapnik.
Diagnosis Shell ca. 1.6 mm, transparent, ovate-conical with a lunate aperture and a singular, obliquely extended lamella. The lamella is frequently turned upwards in its maximal extension within the shell cavity.
Measurements (n = 8) sh: 1.36–1.71 mm (mean: 1.578 ± 0.107 mm); sw: 1.05–1.2 mm (mean: 1.105 ± 0.051 mm); ah: 0.65–0.84 mm (mean: 0.756 ± 0.058 mm); aw: 0.56– 0.68 mm (mean: 0.657 ± 0.048 mm); number of whorls: 4.75–5.5 (mean: 5.346 ± 0.347).
Description Shell ovate-conical, translucent when fresh; shell surface smooth; whorls convex; aperture lunate; peristome somewhat taller than wide; parietal shield well differentiated from the lip, comparatively thin (see Supplementary Fig. S3 View Fig , PC2-axis in the geometric PCA and the CV1-axis in the CVA), its margin usually straight; parietalis present in the aperture, though not particularly pronounced; parietalis extends one whorl into the shell, most pronounced half a whorl in, sometimes after increasing considerably ( Fig. 6a–c View Fig ); columellaris starting behind the aperture, sometimes completely absent, extending up to one whorl into the shell, most pronounced half a whorl in. The singular, inclinate, extensive lamella shows a wide variation in the degree of horizontal extension in mature shells, directed upward, almost wing-like, away from the columella.
Differing from Z. kupitzense by the starting point of the columellaris behind the aperture, by its shorter parietal shield and internally, by its oblique and extensive wing-like lamella projecting away from the columella; from Z. isselianum , by its wing-like extension of the inclinate lamella; from Z. amoenum , by the discrete parietalis increasing in size until half a whorl into the shell and the frequent lack of this structure in edentate morphs known within Z. amoenum ( Jochum et al. 2015c) ; differing in the LDA from Z. isselianum by the ratio sh/ah ( Z. alpestre : 1.983 –2.208, Z. isselianum : 1.737 –1.794).
Distribution (Supplementary Fig. S9 View Fig ) Known from a small area in the Kamnik-Savinja Alps, encompassing the municipalities of Kamnik and Luče.
Zospeum kupitzense A. STUMMER, 1984
Figures 6f–i, S View Fig 14g –h
1984 Zospeum alpestre kupitzense A. STUMMER, Heldia , 1 (1): 13–14, pl. 1b fig. 1, 2a-c, 3a–b (shell) [Kupitzklamm südöstlich Eisenkappel, Nordostkarawanken, Kärnten].
2015 Zospeum kupitzense , − Jochum et al., Subterranean Biology 16: 123–165.
Type specimens Holotype SMF 256354 About SMF , paratypes SMF 256355 About SMF /3, NMBE 549730 View Materials (ex coll. Subai); NMW, Landesmuseum für Kärnten, coll. STUMMER 8682 + 8682a, coll. FALKNER 5492/5, coll. SEIDL, coll. STOJASPAL.
S p e c i m e n s e x a m i n e d Sloveni a: NMBE 553391/2, Ložekarjeva zijalka, 2.10.2007, leg. Jochum; NMBE 553392/5, Ložekarjeva zijalka, 2.7.1999, leg. Slapnik; NMBE 553393/4, Ložekarjeva zijalka, 12.6.2012, leg. Jochum; NMBE 553394/1, Ložekarjeva zijalka, 12.6.2012, leg. Weigand.
Diagnosis Shell ca. 1.6 mm, transparent, conical, a long angular parietal shield and a low-lying, robust parietalis extending from the outermost parieto-columellar corner into the aperture, columellaris present in the peristome.
Measurements (n = 17) sh: 1.44–1.66 mm (mean: 1.569 ± 0.072 mm); sw: 1.02–1.15 mm (mean: 1.104 ± 0.042 mm); ah: 0.72–0.94 mm (mean: 0.836 ± 0.062 mm); aw: 0.62– 0.77 mm (mean: 0.701 ± 0.0308 mm); number of whorls: 5– 5.75 (mean: 5.365 ± 0.242).
Description Shell conical, translucent when fresh; convex whorls; shell surface smooth; aperture somewhat taller than wide; angular parietal shield; parietalis present in aperture and slightly more pronounced than in Z. alpestre , extending one whorl into the shell at a more or less constant size keeping on the parietal side of the whorl inside the shell, instead of wandering onto the columella as in most species; columellaris often present in aperture, extending up to one whorl into the shell, being most pronounced half a whorl in; protoconch covered with concentric rows of interconnecting pits on the first three whorls (Supplementary Fig. S 14g –h), most densely on the top (sometimes organized into striae) with density decreasing abapically; teleoconch with striae of densely interconnected pits.
Differing from Z. alpestre by the columellaris starting in the aperture and by its parietalis staying on the parietal side of the whorl within the shell; from Z. isselianum , by its much smaller parietal shield; from Z. amoenum , by the parietalis increasing in size until half a whorl into the shell. Differing in the LDA from Z. isselianum by the ratio sw/lh ( Z. kupitzense : 0.998 –1.145, Z. isselianum : 1.189 –1.253).
Distribution (Supplementary Fig. S9 View Fig ) Known from the Kupitzklamm near Bad Eisenkappel (Carinthia, Austria) and adjacent regions in Slovenia.
Zospeum isselianum POLLONERA, 1887
Figures 6j–k, S View Fig 14e–f, S18a–c
1887 Zospeum isselianum POLLONERA, Bull. Soc. malac. ital., 12 (3): 205, pl. 6 fig. 13 (shell) [raccolte nelle posature del Natisone nel Friuli .... a nord di Cividale (A. TELLINI leg., 2 exx.; sieved from the Natisone river north of Cividale)].
Type specimens Syntype MZUT M3232 View Materials - heavily deteriorated by Byne’ s degradation and unrecognizable . Neotype: MCSMNH 37013 (from Turjeva jama, near Robič) designat- ed by Jochum et al. 2015c.
Specimens examined Slovenia: NMBE 553389 View Materials , Turjeva jama, 19.10.2007, leg. Slapnik ; NMBE 553390 View Materials , Turjeva jama, 4.6.2013, leg. Slapnik.
Diagnosis Shell ca. 1.45 mm, transparent with ovate-conic form, peristome with a strongly developed parietal shield, a parietalis extending into the shell on the columella, a weak columellaris inside the shell.
Measurements (n = 5) sh: 1.36–1.5 mm (mean: 1.441 ± 0.05 mm); sw: 1.07–1.14 mm (mean: 1.105 ± 0.025 mm); ah: 0.76–0.84 mm (mean: 0.815 ± 0.03 mm); aw: 0.67– 0.76 mm (mean: 0.706 ± 0.03 mm); number of whorls: 5.25–5.5 (mean: 5.4 ± 0.137).
Description Shell conical, translucent when fresh, usually broader than in its relatives; shell surface smooth; aperture somewhat taller than wide; parietal shield well differentiated from the lip, usually broader than in related species; parietalis present in aperture, though not very pronounced, extending less than a whorl into the shell, being most pronounced half a whorl in; columellaris sometimes present in the aperture, sometimes starting just behind, extending up to ¾ a whorl into the shell, being most pronounced ½ a whorl in. Protoconch covered with spiral lines of pits with smooth bands lacking pits in between (Supplementary Fig. S14e); teleoconch with striae of densely interconnected pits (Supplementary Fig. S14f). Radula: Adjacent to the sharply pointed tricuspid rachidian tooth, the lateral teeth of the Z. isselianum (Turjeva jama) radula have long pointed cusps bearing an endocone and an ectocone on both the left and right sides respectively (Supplementary Fig. S18c). These cusps are 1/3 to 1/2 the length of the long mesocone. Laterals to the right of the rachidian tooth have a short, nub-like endocone to the left of the mesocone and a longer ectocone on the right, which is about ½ the length of the mesocone. Grooves run down the length of the mesocones. Some rows have a consistent pattern of pointed tricuspid teeth with both endocones and ectocones measuring about ¼–½ the length of the mesocone (Supplementary Fig. S18a). The marginal teeth of Z. isselianum have 3–4 pointed cusps of uneven length whereby the longest cusps in a given row are about the same length (Supplementary Fig. S18b). The basal plates have wing- or arm-like appendages with conspicuous wart-like prominences on the inner surfaces of these wings (Supplementary Fig. S18a) (lowest row). Martins (1996) reported similar basal plate structures in the radula of Melampus monile ( Bruguière, 1789) . Despite the folding during preparation, the radular ribbon of Z. isselianum appears shorter and broader, attenuating to a wide triangular base (Fig. not shown).
Differing from most species by its broad parietal shield; from Z. amoenum by the parietalis increasing in size until half a whorl into the shell. Differing in the LDA from Z. alpestre by the ratio sh/ah ( Z. isselianum : 1.737 –1.794, Z. alpestre : 1.983 –2.208); from Z. kupitzense by the ratio sw/lh ( Z. isselianum : 1.189 –1.253, Z. kupitzense : 0.998 –1.145); from Z. amoenum barely by a combination of the ratios ah/lh ( Z. isselianum : 0.88–0.926, Z. amoenum : 0.712 –0.891) and sw/ D1 ( Z. isselianum : 2.836–3.09, Z. amoenum : 2.194 –3.292).
Distribution (Supplementary Fig. S9 View Fig ) Originally described from debris collected from the Natisone River in Italy, this species is known from Turjeva jama near Kobarid ( Slovenia, neotype locality) ( Jochum et al. 2015c) and from the Grotta Nuova di Villanova (Villanova, Italy; Pezzoli 1992). Other Italian reports ( Pezzoli 1992) from Grotta di S. Giovanni d’ Antro (San Pietro al Natisone) and from Sorgente B Potcouch^ (Vedronza) are probably this species as well.
Zospeum amoenum ( FRAUENFELD, 1856)
Figures 6l–p, S View Fig 14c–d, S18d–e
1856 Carychium amoenum FRAUENFELD, Sitzungsber. mathem.-naturw. Cl. kais. Akad. Wiss. Wien, 19 (1): 82–83, pl. fig. 1, 1a–b (shell) [Ich habe sie selbst in der Pasizagrotte gesammelt, und besitze ausserdem ein Exemplar durch Herrn SCHMID aus der Grotte Juhanča (Velika Pasica, near Gornji Ig and Ihanšica, near Ihan respectively)].
Type specimens There is no type material of amoenum present in NHMW coming from Velika Pasica. The shell NHMW 71976 from the Frauenfeld collection originates from Juhanča (today known as Ihanščica) and thus has to be considered a syntype of C. amoenum . Recent exploration in the Velika Pasica (visited several times) yielded no Z. amoenum at all.
Specimens examined Slovenia: RS 2037/2, Ihanšica , 29.7.2016, leg. Slapnik; MZRS 0059 / 1, Potočka zijalka, 8.6.2012, leg. Slapnik; NMBE 553374 View Materials / 1, Konečka zijalka, 2.10.2007, leg. Jochum; NMBE 553375 View Materials / 20, Konečka zijalka, 2.10.2007, leg. Jochum; NMBE 553376 View Materials / 1, Konečka zijalka, 27.6.2008, leg. Slapnik; NMBE 553377 View Materials / 1, Konečka zijalka, 16.8.2008, leg. Slapnik; NMBE 553378 View Materials / 1, Konečka zijalka, 16.8.2008, leg. Slapnik; NMBE 553379 View Materials / 7, Konečka zijalka, 2.10.2007, leg. Jochum; NMBE 553380 View Materials / 1, Konečka zijalka, 27.6.2008, leg. Slapnik; NMBE 553381 View Materials / 2, Jama 2 pri Jabljah, 25.6.2009, leg. Jochum.
Diagnosis Shell ca. 1.5 mm, transparent with ovate-conic form, sometimes lacking obvious apertural barriers, but often with an obsolete lamella (denticle) in the parietal-columellar region.
Measurements (n = 47) sh: 1.24–1.78 mm (mean: 1.531 ± 0.132 mm); sw: 0.95–1.24 mm (mean: 1.103 ± 0.073 mm); ah: 0.66–0.89 mm (mean: 0.778 ± 0.05 mm); aw: 0.58– 0.76 mm (mean: 0.678 ± 0.033 mm); number of whorls: 4.75–5.75 (mean: 5.245 ± 0.224).
Description Shell conical, translucent when fresh; shell surface smooth; aperture somewhat taller than wide. Parietal shield well-differentiated from lip, moderately broad to narrow, its margin being straight; parietalis may or may not be present; if present, extending one whorl into the shell and maintaining a constant size; a very weak columellaris sometimes present; protoconch with pits aligned into spiral lines interspersed with irregular zones of smooth bands lacking pits (Konečka zijalka; Supplementary Fig. S14c), though this is not always easily visible (Ihanščica; Jochum et al. 2015c, Fig. 9M–O View Fig ); teleoconch with striae of densely interconnected pits (Supplementary Fig. S14d). Radula: The obtuse tricuspid teeth of the Z. amoenum radula from Konečka zijalka (L 2 in Jochum et al. 2015c) are structurally dense and show a deep median groove down the mesocone (Supplementary Fig. S18d). The transitional teeth bear four cusps of unequal length and width. The endocones are generally semi-detached. One transitional tooth (positioned sideways here) (Supplementary Fig. S18e) shows four wide subequal cusps with the broadest on the right flanked by the second middle cusp bearing furrows on their leftmost edges. The rightmost cusp also has a discrete furrow at its rightmost edge covered in part by crust. The crown of this tooth is very skewed to the right. The radular ribbon completely folded upon positioning onto the SEM stub and thus, we have no complete perspective.
Differing from all other species by the consistent absence of the columellaris and the either very weak or absent parietalis. Barely differing in the LDA from Z. isselianum through a combination of the ratios ah/lh ( Z. amoenum : 0.712 –0.891, Z. isselianum : 0.88–0.926) and sw/D1 ( Z. amoenum : 2.194 –3.292, Z. isselianum : 2.836–3.09).
Distribution (Supplementary Fig. S9 View Fig ) Endemic to Slovenia. Known from four caves east and south from the distribution areas of Z. alpestre and Z. kupitzense : Ihanščica (Ihan, Domžale), Jama 2 pri Jabljah (Loka pri Mengšu, Mengeš), Konečka zijalka (Šmihel nad Mozirjem, Mozirje), and Potočka zijalka (Olševa mountain, Solčava).
Remarks Likely the locality data to the shells Frauenfeld collected in the Pasica cave got mixed by him or his companion Schmidt. Only Zospeum spelaeum is known from Pasica cave, a relatively small and easily navigable cave.
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
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