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Nemastoma pluridentatum (Hadži, 1973) stat. nov. Novak Figs 2, 4D, 5D, 6D, 9D, 10D, 12D; Table 7 Nemastoma (Lugubrostoma) triste pluridentatum Hadži, 1973a: 43, fig 33a (original description) [partim; the description also includes N. bidentatum schmidti ssp. nov., N. bidentatum gruberi ssp. nov. × N. bidentatum schmidti ssp. nov. and N. bidentatum martensi ssp. nov. × N. bidentatum schmidti ssp. nov.]. Nemastoma bidentatum bidentatum – Martens 1978: 107. Nemastoma ? bidentatum pluridentatum – Novak 2005a: 314, fig. 34. Nemastoma bidentatum pluridentatum – Schönhofer 2013: 36. Diagnosis (partly according to Hadži 1973a) Robust, short-legged member of the Nemastoma b. complex, with extremely stout Ch and Pa. Ch-Apo small, slightly trapezoid button-like, wider than high, with secretion field in frontal third. Pa-Tr nearly round, Pa-Fe and Pa-Pt very stout, Pa-Fe club-shaped, with 4 large, wide humps. Material examined Holotype BOSNIA AND HERZEGOVINA – YK10 • 1 ♂; Mt. Paljenik, Vlašić planina Mts; 1933 m a.s.l.; 18 Jun. 1962; E. Pretner leg. (4/2005, JH’s microscopic preparation); PMSL. (Note: UTM code in Novak (2005a) was erroneously cited as YJ19.) Partial redescription Note: male body and penis missing; female unknown. Male (holotype) CHELICERAE. Ch basal article length (without Apo) 0.57, stout, 1.8 times as long as wide at Ch max width at dorsal hump, distally rounded. Ch-Apo rounded low trapezoid, with longest margin distally, secretion field frontally. Distal article length 0.57, max width 0.26, movable finger length 0.28 (Figs 4D, 6D, 9D). PEDIPALPS. Pa stout, Pa-Fe extremely stout, with 4 large, bulky humps (instead of spines), club-shaped (Pa-Fe min:max width ~ 1:3.6), Pa-Pt extremely stout (length:max width ~1:2.8) (Figs 5D, 6D). Pa article lengths in Table 7. LEGS. Pseudoarticle leg-Fe formula I−IV: 0−2−1−2. Tarsomere formula I−IV: 8−13−5−8. Leg article lengths in Table 7. Remarks Under the name Nemastoma (Lugubrostoma) triste pluridentatum, Hadži (1973a) described four taxa: a) N. pluridentatum (Hadži, 1973) stat. nov. (Hadži 1973a: fig. 33a) from Mt. Paljenik (1943 m a.s.l.) in the Vlašić Mts, Bosnia, b) a hybrid N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. (Hadži 1973a: fig. 33b, č) from the surroundings of Novo mesto, Slovenia, c) a hybrid N. b. martensi ssp. nov. × N. b. schmidti ssp. nov. (Hadži 1973a: fig. 33c) from the surroundings of Novo mesto, which Hadži (ibid.) erroneously cited for Mt. Vlašić, and d) Nemastoma b. schmidti ssp. nov. from Mt. Šmarna gora, Slovenia (Hadži 1973a: fig. 34a), from the surroundings of Ljubljana (Hadži 1973a: fig. 34b), and from the cave Jama Sv. Janeza pri Prestranku near Postojna, Slovenia (Hadži 1973a: fig. 34c). Figs 33 and 34 were drawn freehand; consequently, shapes and dimensions only partly correspond to the real measurements. Hadži’s original microscopic preparation labeled “ Nemastoma bidentatum Roewer pulli, A-1065 (Novo Mesto), legit Karaman” includes two males and a female, but no juveniles. Hadži (ibid.) drew some details of the males, including damaged legs; see remarks on Nemastoma (Stridulostoma) seliskari Hadži, 1973 under N. b. martensi ssp. nov. × N. b. schmidti ssp. nov. Given that some important parts, e.g., Pa of one male, are missing, Hadži probably made this incomplete preparation after observation of the specimens under a microscope. In his original paper (Hadži 1973a), the type specimen and the type locality of N. pluridentatum stat. nov. were not designated, but he did that in the catalogue (Hadži 1973b: 13) under a mistyped name: “ N. (L.) triste pleuridentatum Hadži, 1973 ”. There he wrote: “Loc. typ.: Novo mesto ”. Since the specimens from this locality belong to the above-mentioned hybrids, Novo mesto cannot be the type locality of N. pluridentatum stat. nov.; that is the first-mentioned location in Bosnia, as fig. 33a (Hadži 1973a) is the only drawing that refers to this taxon. Unfortunately, only the preparation of one Ch, one Pa and four legs of the male holotype are preserved, while the body, as well as a further male and female mentioned by Hadži (1973a: 44) are missing. Because of the lack of any information on the Pe, the saw on Fe IV in preserved legs is the only available character allowing us to place this taxon into the N. b. complex, interpreted as N. ? bidentatum pluridentatum (Novak 2005a: fig. 34) or N. b. pluridentatum (Schönhofer 2013). In 2019, we visited the type locality Mt. Paljenik, but we did not find further specimens of N. pluridentatum. Instead we found a new, much different N. kozari sp. nov. there, which unlikely hybridize with N. pluridentatum stat. nov. Consequently, we ascribed the species status to N. pluridentatum. See remarks under N. relictum stat. nov., N. kozari sp. nov. and hybrids N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. and N. b. martensi ssp. nov. × N. b. schmidti ssp. nov. Distribution Bosnia and Herzegovina, endemic to central Bosnia, probably endemic to the Vlašić Mts. Type locality: Mt. Paljenik in the Vlašić planina Mts, Bosnia, (44.29° N, 17.64° E, 1933 m a.s.l.), Bosnia and Herzegovina. Ecology Nemastoma pluridentatum stat. nov. inhabit the highest peak areas with the Mastgrass Gentianello crispatae-Nardetum strictae Redžić 1990 communities. Phenology: adults probably eurychronous.

Nemastoma relictum Gruber & Martens, 1968 stat. nov. Komposch Figs 1B, 2, 4C, 5C, 6C, 7D, 8D, 9C, 10C, 11D, 12C, 13C; Tables 5–6 Nemastoma bidentatum relictum Gruber & Martens, 1968: 147–149, figs 14−16, 24 (original description). Nemastoma bidentatum relictum – Martens 1978: 112–113, figs 148−150. — Komposch & Gruber 2004: 485. — Komposch 2009: 476; 2011: 65. — Schönhofer 2013: 36. Diagnosis Species within the N. b. complex with button-shaped Ch-Apo, and secretion field oriented frontally. Pa- Fe without spines, slightly club-shaped (Pa-Fe min:max width ~1:2.3). Rec sem with 5 lobate, tubularsaccular vesicles. Material examined AUSTRIA • 2 ♂♂, 3 ♀♀; Grosses Kar, the Koralpe Mts; 25 Oct. 2005; C. Komposch leg. (CK det.); PMSL. Description (according to Gruber & Martens 1968, slightly modified) Male (from Grosses Kar, Salzburg) BODY. Length 1.55−1.60. CHELICERAE. Ch with flat triangular dorsal margin in front of dorsal indentation; ventral margin straight, with minute triangular hump. Ch-Apo button-like, inferior and frontal margins straight, embracing a 90° angle, Ch-Apo dorsally round, secretion field with dense setae (brush) frontally (Figs 4C, 6C, 9C). Proximal article length 0.54, distal article length 0.53, max width 0.18, movable finger length 0.19. PEDIPALPS. Pa-Tr dorsal margin high, oval, leaned distally; Pa-Fe distended distally (max:min width ~ 1.7:1), without spines. Pa-Pt with conspicuously convex ventral margin and finger-like medial Apo. Pa-Ti and Pa-Ta straight (Figs 5C, 6C). Pa article lengths in Table 5. PENIS. Pe length 1.6, Pe basis ¼ Pe length, glans isosceles triangular (Figs 7C, 8C). LEGS. Leg article lengths in Tables 5–6. Female (from Grosses Kar, Salzburg) BODY. Length 1.70. CHELICERAE. Ch basal article with low-arched dorsal margin in front of indentation, and ventral lowarched concavity, Ch basal article length:width ~ 2.3:1 (Fig. 13C). Proximal article length 0.56, distal article length 0.55, max width 0.19, movable finger length 0.20. PEDIPALPS. Pa short, ~ 2 mm long, Pa-Tr with high, isosceles triangular dorsal margin arched on the top, Pa-Fe distended distally (max:min width ~1.6:1), Pa-Ta short, Pa-Ta length:Pa-Ti length ~1:1.6 (Fig. 13C). Pa article lengths in Table 5. OVIPOSITOR. Rec sem with 5 lobate, tubular-saccular vesicles (Fig. 11D). LEGS. Leg article lengths in Tables 5–6. Remarks We did not examine the original type material; for the current description, we used other specimens from the type location. Rec sem were found much different from the other taxa within the N. b. complex. Distribution Austria, endemite of eastern Central Alps (Gruber & Martens 1968; Martens 1978; Komposch 1999, 2009). Vertical distribution: 1325−2240 m a.s.l. (Komposch 1999, 2009). Type locality: Grosses Kar, near the Tappenkarsee Lake, Kleinartal, Salzburg (47.19° N, 13.32° E, 1760 m a.s.l.), Radstädter Tauern Mts, Austria. Ecology Nemastoma relictum stat. nov. is a montane to alpine species within the N. b. complex inhabiting forests below the krummholz zone, alpine meadows, spring-areas, and gravel and block heaps in the eastern Central Alps (Gruber & Martens 1968; Martens 1978; Komposch 1999, 2009) in 1325–2240 m a.s.l. (Komposch 1999). Phenology: evidence of adults from the end of May till the end of October, but probably eurychronous., Published as part of Novak, Tone, Novak, Ljuba Slana, Kozel, Peter, Schaider, Miriam Gudrun, Komposch, Christian, Lipovšek, Saška, Podlesnik, Jan, Paušič, Igor & Raspotnig, Günther, 2021, Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence, pp. 1-67 in European Journal of Taxonomy 777 on pages 16-18, DOI: 10.5852/ejt.2021.777.1561, http://zenodo.org/record/5637112, {"references":["Gruber J. & Martens J. 1968. Morphologie, Systematik und Okologie der Gattung Nemastoma C. L. Koch (s. str.) (Opiliones, Nemastomatidae). Senckenbergiana Biologica 49 (2): 137 - 172.","Martens J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. In: Senglaub F., Hannemann H. & Schumann H. (eds) Die Tierwelt Deutschlands 64: 1 - 464. Fischer Verlag, Jena.","Komposch C. & Gruber J. 2004. Die Weberknechte Osterreichs (Arachnida, Opiliones). Denisia 12, zugleich Kataloge der Osterreichishen Landesmuseen Neue Serie 14: 485 - 534.","Komposch C. 2009. Opiliones (Weberknechte). In: Rabitsch W. & Essl F. (eds) Endemiten - Kostbarkeiten in Osterreichs Pflanzen- und Tierwelt: 476 - 500. Naturwissenschaftlicher Verein fur Karnten & Umweltbundesamt.","Komposch C. 2011. Endemic harvestmen and spiders of Austria (Arachnida: Opiliones, Araneae). Arachnologische Mitteilungen 40: 65 - 79. https: // doi. org / 10.5431 / aramit 4008","Schonhofer A. L. 2013. A taxonomic catalogue of the Dyspnoi Hansen and Sorensen, 1904 (Arachnida: Opiliones). Zootaxa 3679 (1): 1 - 68. https: // doi. org / 10.11646 / zootaxa. 3679.1.1","Komposch C. 1999. Rote Liste der Weberknechte Karntens (Arachnida: Opiliones). In: Rottenburg T., Wieser Ch., Mildner P. & Holzinger W. E. (eds): Rote Listen gefahrdeter Tiere Karntens. Naturschutz in Karnten 15: 547 - 565. Amt der Karntner Landesregierung, Abt. 20, Landesplanung, Klagenfurt."]}

Nemastoma bidentatum martensi ssp. nov. × N. bidentatum schmidti ssp. nov. Figs 3, 9O Nemastoma (Stridulostoma) seliskari Hadži, 1973a: 48, figs 33c, 37a−e. Nemastoma bidentatum bidentatum – Martens 1978: 107 [partim: Nemastoma (Stridulostoma) seliskari Hadži, 1973]. Nemastoma bidentatum bidentatum × N. bidentatum sparsum – Novak & Gruber 2000: 286 [partim: Novo mesto, Mt. Mirna gora]. Nemastoma bidentatum pluridentatum – Schönhofer 2013: 36 [partim: Nemastoma (Stridulostoma) seliskari Hadži, 1973, and Nemastoma seliskari Hadži, 1973]. Diagnosis Typical male hybrids with Ch basal article with large hump in front of dorsal indentation, Ch-Apo with knee-like frontal flexion and narrowly drawn out upper portion, Pa-Fe club-shaped, but thinner than in N. b. schmidti ssp. nov., with four simple, equidistant ventral spines in distal Pa-Fe half, and Pa-Ti with minute hump. Material examined SLOVENIA – VL15 • 1 ♂; Prelože pri Lokvi; 27 Sep. 2011; L. Slana Novak and T. Novak leg. (310/2011); PMSL. – VL35 • 3 ♂♂, 5 ♀♀; Ribnica – Ostrožno Brdo; 27 Sep. 2011; L. Slana Novak and T. Novak leg. (324/2011, rev. 2015); PMSL. – VL53 • 1 ♂; Zabiče; 27 Sep. 2011; L. Slana Novak and T. Novak leg. (301/2011, rev. 2015); PMSL. – WL05 • 1 ♂; Mt. Mirna gora; 28 Jul. 1948; A. Seliškar leg; PMSL. – WL17 • 1 ♂; Novo Mesto; Karaman leg. (107/1983 microscopic preparation, JH det., TN rev. 2017); PMSL. Remarks In Hadži’s original preparation, labelled “ Nemastoma bidentatum Roewer pulli”, with two cover glasses on the same microscope slide, there are two males and a female. One male corresponds to the type specimen of Nemastoma (Stridulostoma) seliskari Hadži, 1973 (Hadži 1973a). For this species, Hadži (ibid.) founded a new subgenus Stridulostoma based on the existence of an alleged “stridulatory organ” on the Fe III (Hadži 1973a: figs 37č–d). Gruber (1976: 797) explained that the stridulatory organ alone does not justify establishing of a new subgenus, especially because this new species differs from N. (Lugubrostoma) triste pluridentatum Hadži, 1973 exclusively by possession of this organ, and accordingly, synonymized Stridulostoma with Nemastoma. Following these remarks, Schönhofer (2013: 36) synonymized N. seliskari with N. b. pluridentatum. We found that the portion of the Fe III, identified by Hadži (1973a) as a stridulatory organ in the type specimen, is in fact a depression-damage, caused by desiccation. Moreover, such damage is present in nearly all (sic!) the other femora of this freshly moulted specimen, but nothing resembles the fine lamellated structure shown in Hadži’s drawing (1973a: figs 37č–d). No similar structure was found in the specimens of Nemastoma collected later on Mt. Mirna gora. One might imagine that a careless inspection under a microscope at low magnification could result in the illusion of a ʻstridulatory organʼ. However, the drawing of its lamellae cannot be considered anything but a pure fabrication. Moreover, Pa-Fe in Hadži (1973a: fig. 33c), being under the same cover glass in the preparation, belongs to the same specimen, although Hadži (ibid.) ascribed it to N. (Lugubrostoma) triste pluridentatum. See remark under N. pluridentatum stat. nov.

Nemastoma bidentatum sparsum Gruber & Martens, 1968 Figs 2–3, 4B, 5B, 6B, 7C, 8C, 9B, 10B, 11A, C, 12B, 13B; Tables 3–4 Nemastoma bidentatum sparsum Gruber & Martens, 1968: 146–147, figs 1–2, 7−11 (original description) [partim; the type series also includes N. bidentatum gruberi ssp. nov.]. Nemastoma bidentatum – Roewer 1914: 141 [partim: type series also includes N. bidentatum bidentatum and N. triste]. — Šilhavý 1956: 131, figs 243–246. — Loksa 1962: 267, figs 3a–f. — Gruber & Martens 1968: 146–147, figs 1–2, 7−11. — Staręga 1976: 313–315, figs 25–27. — Martens 1978: 111–112, figs 143−145, 147, 150; 2006: 182–184, figs 20, 21c–d, 22a–h. — Novak & Gruber 2000: 286. — Novak et al. 2002: 136 [partim: Radenci]; 2006: 269. — Komposch & Gruber 2004: 485. — Novak 2004b: 245 [partim: Breznica, Slavonski brod, Spačva near Županja]; 2005a: 315 [partim: Bosanski Brod]. — Schönhofer & Holle 2007: 25, figs 2a–i. — Schönhofer 2013: 36. Diagnosis (according to Gruber & Martens 1968, slightly modified) Male Ch basal article with nearly straight dorsal margin in front of dorsal indentation, and short, roundly oblong Ch-Apo, with small apical pinnacle. Pa-Fe moderately club-shaped (Pa-Fe min:max width ~1:2.5), with 2 distal, pointed spines, spine 2 the largest. Pa-Pt Apo conical. Glans equilaterally triangular. Rec sem with 14 balloon-like vesicles. Material examined CROATIA – XM05 • 1 ♀; Jurovčak; 3 Apr. 1983; T. Novak and L. Slana Novak leg. (LSN 7/1983, TN rev. 2008); PMSL. SLOVENIA – WM06 • 1 ♂; Cvitrško sedlo, Otiški vrh; 28 May 2008; T. Novak leg. (160/2011); PMSL. – WM42 • 1 ♂, 2 ♀♀, 1 juv.; Mestinje; 4 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 43/1983); PMSL • 1 ♂, 1 ♀; ibid.; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1138/2002); PMSL • 1 ♀; Šmarje pri Jelšah; 6 Apr. 1986 (1169/1997, rev. 2015); PMSL • 1 ♀; Zgornje Poljčane; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1160/2002); PMSL. – WM45 • 1 ♂; Ledine; 25 Jul. 1987; F. Janžekovič leg. (7/1999); PMSL • 1 ♂; Maribor; 24 Aug. 1986; Malavašić and Koren leg. (265/2002); PMSL. – WM47 • 1 ♀; Kresnica; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (162/2011); PMSL. – WM52 • 1 ♂; Maršečka vas; 11 Oct. 1997; L. Slana Novak and T. Novak leg. (1893/1998); PMSL. – WM53 • 2 ♂♂, 1 ♀; Pečke; 24 Jun. 2007; T. Novak leg. (45/2007); PMSL • 1 ♂, 2 ♀♀; Šega, Makole; 9 Apr. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 70/1983); PMSL. – WM56 • 1 ♀; Kaniža; 20 May 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 80/1983); PMSL • 1 ♀; Mt. Piramida, Maribor, 1−30 Nov. 1978; (909/1997, rev. 2015); PMSL • 1 ♀; Spodnje Dobrenje; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (165/2011); PMSL • Ranca, Pesnica. – WM57 • 1 ♂; Ceršak; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (113/2011); PMSL • 1 ♂; Kozjak pri Ceršaku; 20 Apr. 2008; L. Slana Novak and T. Novak leg. (116/2011); PMSL. – WM66 • 2 ♀♀; Črnetina; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (163/2011); PMSL • 1 ♂; Kremberg; 4 Jun. 1984; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 343/1985); PMSL • 2 ♂♂; ibid.; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (114/2011); PMSL • 1 ♀; Spodnji Žerjavci; 23 Apr. 2008; L. Slana Novak and T. Novak leg. (115/2011); PMSL. – WM67 • 1 ♂; Sv. Ana; 3 Jul. 2002; T. Novak leg. (799/2002); PMSL • 1 ♀; Nasova; 3 Jun. 1984; L. Slana Novak and T. Novak leg. (314/2002, rev. 2015); PMSL • 1 ♂; Stogovci; 13 Apr. 1984; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 323/1985); PMSL • 1 ♂, 1 ♀; Zgornja Velka; 20 Mar. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 8/1983); PMSL. – WM73 • 1 ♀; Repišče; 24 Jun. 2007; T. Novak leg. (31/2007, rev. 2015); PMSL. – WM74 • 1 ♀; Podvinci; 26 Oct. 1994; M. Varžič leg. (21/1999); PMSL • 1 ♂; Žamenci; 22 Oct. 1994; M. Varžič leg. (18/1999); PMSL • 1 ♂; ibid.; (20/1999); PMSL • 1 ♂; ibid.; 23 Oct. 1994; M. Varžič leg. (19/1999, rev. 2015); PMSL. – WM75 • 1 ♂; Trnovce; 2 Jun. 2002; L. Slana Novak and T. Novak leg. (760/2002); PMSL • 7 ♂♂, 4 ♀♀; Velika Slavšina; 13 Oct. 2005; L. Slana Novak and T. Novak leg. (158/2005, rev. 2018); PMSL • 5 ♂♂, 3 ♀♀; Vitomarci; 30 Jun. 2002; T. Novak leg. (787/2002); PMSL • 1 ♀; ibid.; 3 Jul. 2002; T. Novak leg. (792/2002); PMSL • 1 ♂; ibid.; (797/2002); PMSL • 1 ♀; ibid.; 3 Jul. 2002; T. Novak leg. (794/2002, rev. 2008); PMSL • 1 ♀; ibid.; (796/2002); PMSL • 1 ♂; ibid.; (797/2002); PMSL • 1 ♂; ibid.; (800/2002); PMSL • 1 ♀; ibid.; 2 Oct. 2002; T. Novak leg. (1280/2002); PMSL • 1 ♂; ibid.; (1279/2002); PMSL. – WM77 • 1 ♀; Črešnjevci; 24 Apr. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 79/1983, TN rev. 2008); PMSL • 1 ♂, 1 ♀; Korovci; 7 Apr. 1984; L. Slana Novak and T. Novak leg. (LSN 191/1985); PMSL • 1 ♂, 1 ♀; Lomanoše; 1 Oct. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 110/1983); PMSL. – WM78 • 1 ♂; Gerlinci; 11 May 1986; T. Novak and L. Slana Novak leg. (LSN 139/1986); PMSL. – WM86 • 2 ♂♂, 6 ♀♀; Hrašenski vrh; 9 Sep. 1983; T. Novak and L. Slana Novak leg. (LSN 115/1983); PMSL • 4 ♂♂, 3 ♀♀; Kapelski vrh; 17 Sep. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 112/1983); PMSL • 1 ♂; Murski vrh; 19 May 1984; L. Slana Novak and T. Novak leg. (LSN 240/1984); PMSL • 1 ♀; Radenci; 25 Jul. 1983; M. Slana Novak and L. Slana Novak leg. (LSN 111/1983, TN rev. 2007); PMSL • 2 ♂♂, 2 ♀♀; ibid.; 20 Sep. 1983; L. Slana Novak leg. (LSN 119/1983); PMSL • 1 ♂; Rihtarovci; 26 Feb. 1997; B. Drovenik leg. (94/2001); PMSL. – WM87 • 3 ♂♂, 3 ♀♀; Cankova; 15 Sep. 1983; T. Novak and L. Slana Novak leg. (LSN 114/1983); PMSL • 3 ♂♂, 3 ♀♀; Skakovci; 27 Mar. 1983; T. Novak and L. Slana Novak leg. (LSN 4/1983); PMSL • 1 ♂; Sodišinci, Radenci; 19 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 68/1983); PMSL. – WM88 • 1 ♂; Gerlinci; 11 Aug. 1983; L. Slana Novak, M. Slana Novak and T. Novak leg. (1322/1998); PMSL. – WM89 • 2 ♂♂, 4 ♀♀, 1 juv.; Trdkova; 30 Jul. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 49/1983); PMSL. – WM94 • 1 ♀; Pavlovci; 15 Apr. 2000; L. Slana Novak and T. Novak leg. (744/2002); PMSL. – WM95 • 1 ♀; Podgradje, Ljutomer; 27 May 2001; S. Brelih leg. (125/2001); PMSL. – WM96 • 1 juv.; Bratonci; 9 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 335/1985, TN rev. 2008); PMSL. – WM97 • 1 ♂; Andrejci; 28 Jul. 1981; T. Novak leg. (1188/1981, JG det.); PMSL • 1 ♂; Moravci; 16 Mar. 1984; T. Novak and L. Slana Novak leg. (LSN 243/1984); PMSL • 4 ♂♂; Murska Sobota; 24 Nov. 1982; T. Novak leg. (603/1982); PMSL • 3 ♀♀; Rakičan – Noršinci; 23 Sep. 1984; L. Slana Novak leg. (322/1985); PMSL • 1 ♂, 2 ♀♀; ibid.; 13 Sep. 1981; T. Novak leg. (371/1982, JG det.); PMSL. – WM98 • 2 ♂♂, 3 ♀♀; Košarovci; 24 Jul. 1982; L. Slana Novak, T. Novak and M. Slana Novak leg. (529/1982, JG det.); PMSL • 1 ♀; ibid.; 1 Sep. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 318/1985); PMSL • 1 ♂, 3 ♀♀; Mačkovci; 31 Jul. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 67/1983); PMSL. – WM99 • 2 ♂♂; Lucova; 2 Jun. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 239/1984); PMSL. – XM05 • 1 ♀; Bobri; 24 Aug. 1988; F. Velkovrh leg. (13/1995); PMSL • 2 ♂♂, 1 juv.; Dolnja Bistrica; 17 Aug. 1996; I. Kovač leg. (SL 2/1997); PMSL • 2 ♀♀; ibid.; (SL 3/1997); PMSL. – XM06 • 2 ♂♂, 3 ♀♀; Banuta /Bánuta; 10 Aug. 1996; I. Kovač leg. (SL 5/1997); PMSL • 2 ♂♂, 1 ♀; ibid.; (SL 22/1997); PMSL • 2 ♂♂, 2 ♀♀; ibid.; (SL 21/1997); PMSL • 13 ♂♂, 15 ♀♀; ibid.; 13 Aug. 2012; T. Novak leg. (GR 3837−3849, 3792−3797, TN det.); PMSL • 4 ♂♂, 3 ♀♀; ibid.; (GR 3770−3776); PMSL • several specs; Polanski log, Mala Polana; 17 Jul. 2000; T. Novak leg.; PMSL • 1 ♂, 1 ♀; Radmožanci / Radamos; 17 Mar. 1984; T. Novak and L. Slana Novak leg. (LSN 109/1984); PMSL. – XM07 • 4 ♂♂, 3 ♀♀; Dobrovnik /Dobronak; 8 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 57/1983); PMSL • 1 ♀, 1 juv.; ibid.; 18 Aug. 1996; I. Kovač leg. (SL 1/1997); PMSL. – XM08 • 3 ♂♂, 1 ♀; Prosenjakovci / Pártosfalva; 8 Aug. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 50/1983); PMSL • 2 ♀♀; Središče / Szerdahely; 12 Aug. 1996; I. Kovač leg. (SL 6/1997); PMSL • 4 ♂♂; ibid.; 24 Aug. 1996; I. Kovač leg. (SL 7/1997); PMSL • 2 ♀♀; ibid.; (SL 8/1997); PMSL • 1 ♂, 3 ♀♀; ibid.; (SL 20/1997); PMSL • 4 ♂♂; ibid.; (SL 23/1997); PMSL • 4 ♂♂; ibid.; (SL 61/1997); PMSL • 3 ♂♂; ibid.; (SL 108/1997); PMSL • 12 ♀♀; ibid.; (SL 105/1997); PMSL. – XM15 • 3 ♀♀; Benica / Benice; 1Aug. 1996; I. Kovač leg. (SL 11/1997); PMSL • 3 ♀♀; ibid.; (SL 12/1997); PMSL • 2 ♂♂; ibid.; 19 Aug. 1996; I. Kovač leg. (SL 25/1997); PMSL • 2 ♀♀; ibid.; (SL 26/1997); PMSL • 5 ♀♀; ibid.; (SL 28/1997); PMSL • 3 ♂♂; ibid.; (SL 29/1997); PMSL • 2 ♂♂; ibid.; (SL 39/1997); PMSL • 2 ♀♀; ibid.; (SL 64/1997); PMSL • 1 ♂, 1 ♀; ibid.; (SL 65/1997); PMSL • 1 ♂, 3 ♀♀; Gaberje / Gyetyános; 22 Aug. 1996; I. Kovač leg. (SL 9/1997); PMSL • 1 ♀; ibid.; (SL 10/1997); PMSL • 1 ♂; ibid.; (SL 15/1997); PMSL • 3 ♀♀; ibid.; (SL 16/1997); PMSL • 1 ♀; ibid.; 12 Oct. 1996; I. Kovač leg. (SL 30/1997); PMSL • 2 ♀♀; ibid.; (SL 55/1997); PMSL • 5 ♂♂; ibid.; 19 Oct. 1996 (SL 17/1997); PMSL • 4 ♂♂, 4 ♀♀; ibid.; (SL 24/1997); PMSL • 4 ♀♀; ibid.; (SL 103/1997); PMSL • 8 ♀♀; ibid.; (SL 104/1997); PMSL • 2 ♀♀; ibid.; (SL 106/1997); PMSL • 7 ♂♂; ibid.; (SL 107/1997); PMSL • 2 ♂♂; ibid.; (SL 109/1997); PMSL • 14 ♀♀; ibid.; 22 Aug. 1997; I. Kovač leg. (SL 27/1997); PMSL • 1 ♂, 1 juv.; ibid.; (SL 32/1997); PMSL • 1 ♂; ibid.; (SL 68/1997); PMSL • 4 ♂♂; ibid.; 24 Aug. 1997; I. Kovač leg. (SL 19/1997); PMSL • 2 ♀♀; ibid.; 28 Aug. 1997; I. Kovač leg. (SL 13/1997); PMSL • 2 ♂♂; ibid.; (SL 14/1997); PMSL • 1 ♂; ibid.; 12 Oct. 1997; I. Kovač leg. (SL 4/1997); PMSL • 4 ♂♂; ibid.; (SL 20/1997); PMSL • 1 ♀; ibid.; (SL 56/1997); PMSL • 1 ♂; ibid.; (SL 100/1997); PMSL • 4 ♀♀; ibid.; (SL 101/1997); PMSL. – XM16 • 2 ♂♂; Murska šuma; 29 Aug. 1998; T. Novak leg. (1538/1998); PMSL • several specs; ibid.; 18 Jun. 2001; T. Novak leg.; PMSL • 1 ♂, 1 ♀; Dolgovaške Gorice / Hossúfaluhegy; 31 Mar. 1984; T. Novak, D. Kuhar and L. Slana Novak leg. (LSN 141/1984); PMSL • 4 ♂♂, 1 ♀; Lendavske gorice / Lendvahegy; 17 Aug. 1996; I. Kovač leg. (SL 31/1997); PMSL. – XM25 • 1 ♂; Muriša, Murska šuma; B. Drovenik and A. Pirnat leg. (26/2005); PMSL. Description (according to Gruber & Martens 1968, slightly modified) Male (from Prosenjakovci) BODY. Length 1.5−2.0. Males from Slovenia larger than those from the type locality. Length 1.96, width 1.44. CHELICERAE. Ch basal article with nearly straight dorsal margin in front of dorsal indentation and small ventral hump, Ch-Apo short, roundly oblong, with convex anterior margin and small apical pinnacle (Figs 4B, 6B, 9B). Secretion field medially. Proximal article length 0.53, distal article length 0.59, max width 0.18, movable finger length 0.25. PEDIPALPS. Pa-Tr with highly arched dorsal hump. Pa-Fe moderately club-shaped (Pa-Fe min:max width ~ 1:2.5), slightly ventrally bent, with 2 stout, pointed spines in distal Pa-Fe quarter; spine 2 the largest. Pa-Pt Apo triangular-conical. Pa-Ti simple, straight. Pa-Ta conically tapering (Figs 5B, 6B). Pa article lengths in Table 3. PENIS. Pe length 1.6, Pe basis ¼ Pe length, glans equilaterally triangular (Figs 7C, 8C). LEGS. Leg article lengths in Tables 3–4. Female (from Prosenjakovci) BODY. Body length 1.6−2.4. Females from Slovenia larger than those from the type locality. Body length 2.40, body width 1.97. CHELICERAE. Ch basal article with low-arched dorsal margin in front of dorsal indentation, and straight ventral margin with triangular hump, Ch basal article length:width>2.6:1. Proximal article length 0.57, distal article length 0.63, max width 0.21, movable finger length 0.32 (Fig. 13B). PEDIPALPS. Pa-Tr with moderately high, arched dorsal margin; Pa-Fe with distal bonce, Pa-Fe max:min width ~ 1.7:1; Pa-Ti elongated-oval (13 B). Pa article lengths in Table 3. OVIPOSITOR. Ovipositor length 1.0, Rec sem with 14 balloon-like vesicles (Fig. 11A, C). LEGS. Leg article lengths in Tables 3–4. Remarks In Slovenia, N. b. sparsum lives in the northeast, at the margins of the Pannonian basin (Martens & Gruber 1968; Martens 1978; Kovač 1997). The type series deposited in Naturhistorisches Museum Wien (Gruber & Martens 1968) also includes N. b. gruberi ssp. nov. distributed south of the Sava River in Slovenia and Croatia. All prior identifications of N. b. sparsum south of the Sava River (Novak et al. 1995, 2002; Novak 2005b) refer to N. b. gruberi ssp. nov. Martens (1978: 112) reported a sparsum -like population living in the mountain pine association (Hyperico grisebachii-Pinetum mugo Horvat 1938) on Mt. Veliki Snežnik (the peak at 1796 m) at least up to the altitude of 1650 m a.s.l. These individuals may refer to N. b. gruberi ssp. nov., N. b. martensi ssp. nov. or N. b. sneznikensis ssp. nov. Individuals from Bosnia and Herzegovina (Mt. Plasa, Jajce) and Dalmatia, Croatia (Split) (Gruber & Martens 1968) should be revised for identification. See remarks under N. b. gruberi ssp. nov. and N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov., N. b. martensi ssp. nov. and N. b. sneznikensis ssp. nov. Distribution Central and eastern Europe: Austria, Bulgaria, Bosnia and Herzegovina, Czech Republic, Denmark, Hungary, Croatia, Montenegro, Romania, Russia, Slovakia, Slovenia, Serbia and Turkey (Gruber & Martens 1968; Staręga 1976; Martens 1978, 2006; Mučalica 1988; Klimeš 2000; Komposch 2002; Komposch & Gruber 2004; Schönhofer & Holle 2007). Vertical distribution: 120−1650 m a.s.l. (Komposch 1999; Komposch & Gruber 2004), in Slovenia 155–644 m a.s.l. Type locality: Sonnenberg (47.88° N, 16.31° E, 480 m a.s.l.), Leitha Mts, Burgenland, Austria. Ecology Nemastoma b. sparsum is a lowland and hilly land subspecies inhabiting various types of acidophilous and neutrophilous forests of Fagus sylvatica, Carpinus betulus L. and Quercus petraea (Matt.) Liebl., riverine forests, and thermophilous scrubs in north-eastern Slovenia, mostly on non-calcareous soils. Phenology: adults eurychronous.

Nemastoma bidentatum gruberi ssp. nov. × N. bidentatum schmidti ssp. nov. Figs 3, 9M, 10M Nemastoma bidentatum bidentatum – Gruber & Martens 1968: 141 [partim: Dugo selo]. — Martens 1978: 107 [partim: Županja, Mt. Snežnik]. Nemastoma bidentatum bidentatum × Nemastoma bidentatum sparsum – Novak & Gruber 2000: 285 [partim: Mt. Snežnik]. — Novak 2004b: 246 [partim: Dugo selo]. Diagnosis In typical male hybrids, Ch-Apo is boomerang-shaped, Pa-Fe club-shaped, but relatively thin, with four simple, equidistant ventral Pa-Fe spines in the frontal half, Pa-Ti without proximal dorso-medial hump. Material examined CROATIA – WL97 • 1 ♂, 1 ♀; Zagreb − Dugo selo; J. Martens et al. leg. (JM 837); PMSL • 16 ♂♂, 13 ♀♀; ibid.; 9 Sep. 2009; A. Schönhofer leg. (ASc 342, 4/2012, ASc det., TN rev. 2018); PMSL. – XL06 • 3 ♂♂, 2 ♀♀; Prevlaka; (826/2002, rev. 2018); PMSL • 4 ♂♂, 6 ♀♀; ibid.; Oct.1984; (236/2002, rev. 2018); PMSL • 2 ♂♂, 2 ♀♀; ibid.; (278/2002, rev. 2018); PMSL • 1 ♂, 1 ♀; ibid.; (280/2002, rev. 2018) • 1 ♂, 1 ♀; ibid.; 15 Aug. 1985; (LSN 28 /1996, TN rev. 2018); PMSL • 1 ♀; ibid.; (891/2002, rev. 2018); PMSL • 1 ♂; ibid.; 30 Aug. 1985; (609/2002, rev. 2019); PMSL • 2 ♂♂; ibid.; (861/2002, rev. 2018); PMSL • 2 ♂♂, 1 ♀; ibid.; Oct.1985; (896/2002); PMSL • 1 ♀; ibid.; 7 Apr. 1986; I. Kos leg. (328/2002, rev. 2018); PMSL • 1 ♂; ibid.; (843/2002, rev. 2018); PMSL. SLOVENIA – VL13 • 3 ♂♂, 3 ♀♀; Rakitovec; 21 Sep. 2011; L. Slana Novak and T. Novak leg. (263/2011, rev. 2017); PMSL. – VL26 • 5 ♂♂, 2 ♀♀; Fameljska loza, Senožeče; 1 Oct. 2011; L. Slana Novak and T. Novak leg. (350/2011, rev. 2017); PMSL • 1 ♂; Senožeče – Vremščica; 1 Oct. 2011; L. Slana Novak and T. Novak leg. (355/2011, rev. 2017); PMSL. – VL27 • 3 ♂♂, 1 ♀; Mt. Nanos; 1 Sep. 2011; T. Novak leg. (198/2011, rev. 2018); PMSL. – VL34 • 1 ♂, 3 ♀♀; Harije; 19 Sep. 2011; L. Slana Novak and T. Novak leg. (224/2011, rev. 2015); PMSL • 1 ♀; Petelinje; 27 Apr. 1994; S. Brelih leg. (815/1998, rev. 2015); PMSL. – VL58 • 2 ♂♂; Brezovica pri Borovnici; 12 May 1997; S. Brelih leg. (83/2001, rev. 2015); PMSL. – VL67 • 1 ♂, 2 ♀♀; Lužarji; 25 Aug. 2011; T. Novak leg. (168/2011, rev. 2018); PMSL. – VL78 • 1 ♀; Mt. Limberk; Jun.1998; I. Bertoncelj leg. (1544/1998, rev. 2009); PMSL • 1 ♂; Veliko Mlačevo; 13 Mar. 1992; S. Brelih leg. (813/1998, rev. 2018); PMSL. – VL79 • 1 ♂; Veliki Lipoglav; 8 Jul. 1986; L. Slana Novak and T. Novak leg. (472/2002, rev. 2018); PMSL. – VL84 • 1 ♂, 2 ♀♀; Pleš, Borovec pri Kočevski Reki; 19 Sep. 2013; P. Kozel and T. Novak leg. (27/2013, rev. 2015); PMSL. – VL88 • 1 ♂, 1 ♀; Krka, Krška vas pri Muljavi; 7 Jul. 1986; L. Slana Novak and T. Novak leg. (474/2002, rev. 2018); PMSL • 1 ♀; ibid.; Jun. 1998; V. Kovačič leg. (1444/1998, rev. 2017); PMSL. – VL93 • 1 ♂; Slavski Laz; 29 Apr. 2001; S. Brelih leg. (79/2001, rev. 2018); PMSL. – VL99 • 5 ♂♂, 1 ♀; Cirnik; 27 May 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 249 /1984, TN rev. 2015); PMSL • 1 ♀; Temenica; 4 Mar. 1994; S. Brelih leg. (680a/1998, rev. 2018); PMSL. – WL04 • 5 ♂♂; Brezovica pri Predgradu; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (147/1997, rev. 2018); PMSL. – WL05 • 5 ♂♂; Črmošnjice; 18 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (152/1997, rev. 2018); PMSL. – WL06 • 1 ♂, 1 ♀; Baza 20, Kočevske Poljane; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 76 /1986, TN rev. 2018); PMSL • 2 ♂♂; Podstene, Kočevski Rog; 19 Jun. 1986 (672/2002, rev. 2017); PMSL. – WL08 • 1 ♀; Velika jama nad Trebnjem cave, Cad. No. 104, Mokronog; 28 Jul. 2009; M. Zagmajster leg. (PK and TN 6 /2020); PMSL. – WL09 • 1 ♂; Šentrupert; 28 Feb. 1994; S. Brelih leg. (825/1998, rev. 2018); PMSL. – WL19 • 3 ♂♂; Puščava, Mokronog; 28 Feb. 1994; S. Brelih leg. (801/1998, rev. 2018); PMSL. – VL28 • 1 ♂; Mt. Homski hrib, Sela pri Zburah; Jun. 1997; D. Vrček leg. (1709/1998); PMSL. – WL29 • 1 ♂, 2 ♀♀; Dolenji Boštanj; 15 Aug. 2007; L. Slana Novak and T. Novak leg. (73/2007, rev. 2015); PMSL. – WM01 • 1 ♂, 6 ♀♀; Mt. Javor, Znojile; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1194/2002, rev. 2018); PMSL. – WM40 • 1 ♂; Lesično; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1130/2002, rev. 2018); PMSL • 5 ♂♂; ibid.; (1132/2002, rev. 2018); PMSL. – WM42 • 1 ♂; Mestinje; 4 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 44 /1983, TN rev. 2015); PMSL.

Nemastoma bidentatum sneznikensis Novak, Komposch, Slana Novak & Raspotnig ssp. nov. urn:lsid:zoobank.org:act: 7690FA07-5D60-4545-8660-282ADED3F576 Figs 2���3, 4I, 5I, 6I, 7I, 8I, 9I, 10I, 11I, 12I, 13H; Table 11 Etymology The subspecies name �� sneznikensis �� references Mt. Snežnik, southern Slovenia. Diagnosis Short-legged (leg II N. b. complex, with slightly elongated-trapezoidal Ch-Apo, with row of low tubercles on Pa-Fe and with equilateral triangular glans. Rec sem of 18 balloon-like vesicles. Material examined Holotype SLOVENIA ��� VL54 ��� 1 ♂; Mt. Snežnik; 45.59�� N, 14.44�� E; 1553 m a.s.l.; 21 Aug. 1998; Ljuba Slana Novak and Tone Novak leg.; deep-humus in Dinaric Pinus mugo scrub in contact with the Dinaric Fagus sylvatica ��� Abies alba treeline litter sift; PMSL-Opiliones-TN 1494/1998. Paratypes SLOVENIA ��� VL54 ��� 2 ♂♂, 1 ♀; same collection data as for holotype; PMSL-Opiliones-TN 1494/1998. Other material SLOVENIA ��� VL54 ��� 3 ♂♂, 4 ♀♀; Mt. Snežnik; 20 Jul. 1999; L. Slana Novak and T. Novak leg. (185/1999, rev. 2009); PMSL. Description Male (holotype) BODY. Length 1.81, width 1.18. CHELICERAE. Ch basal article (without Apo) length 0.53, 2.7 times as long as wide at Ch max width at dorsal hump. Ch-Apo oval-trapezoid, wider distally, without apical pinnacle. Ch-Apo height 0.13, max width 0.13, min width 0.08 (at the base). Secretion field frontally. Distal article length 0.51, max width 0.17, movable finger length 0.22 (Figs 4I, 6I, 9I). PEDIPALPS. Pa-Tr with dorsal margin slightly inclined distally. Pa-Fe slightly club-shaped (Pa-Fe min:max width ~ 1:2.6) with terminal bonce, and 7 irregular granula in distal third. Pa-Pt length:max width ~ 1:4.3. Pa-Ti slender (Figs 5I, 6I, 10I). Pa article lengths in Table 12. PENIS. Pe length 1.22, base 0.46, glans 0.09, stylus 0.04 (Figs 7I, 8I). LEGS. Pseudoarticle leg-Fe formula I���IV: 0���1���1���2. Tarsomere leg formula I���IV: 9���14���6���8. Leg article lengths in Table 12. Female BODY. Length 2.11, width 1.43. CHELICERAE. Ch basal article length 0.59, 2.7 times as long as wide at dorsal hump, with low-arched dorsal margin in front of indentation and straight ventral margin (Fig. 13H). Distal article length 0.66, max width 0.20, movable finger length 0.30. PEDIPALPS. Pa-Tr low, with roof-like broken dorsal margin, highest in the first two fifths. Pa-Fe distally with low ventral hump, Pa-Pt, Pa-Ti and Pa-Ta elongated (Fig. 13H). Pa article lengths in Table 12. OVIPOSITOR. Ovipositor length 0.62, Rec sem of 18 balloon-like vesicles (Fig. 11I). LEGS. Leg article lengths in Table 12. Remarks Regarding external morphology, N. b. sneznikensis ssp. nov. is very similar to N. relictum, including the habitus, the similarly shaped Pa and relatively short legs. No individuals belonging to the subspecies were found in the last decade. See remarks on taxa of Nemastoma on Mt. Snežnik under N. b. gruberi ssp. nov. Distribution Despite over 30 collections, this taxon has been recorded only in a limited area on Mt. Snežnik. Endemic to Mt. Snežnik, southern Slovenia. Type locality: Mt. Snežnik (45.59�� N, 14.44�� E, 1553 m a.s.l.), Slovenia. Ecology Nemastoma b. sneznikensis ssp. nov. is a psychrophilous subspecies. To date it has been recorded only in a cold, well-drained, deep mull humus Dinaric Pinus mugo scrub in contact with the Dinaric Fagus sylvatica ��� Abies alba treeline on calcareous soils. Likely, it inhabits other cold habitats, like scree and stone heap edges in contact with humus accumulations, as is known for N. relictum (Komposch 1999; Komposch & Gruber 2004), and perhaps a humus-rich shallow subterranean habitat (SSH), i.e., Milieu Souterrain Superficiel (MSS) (Mammola et al. 2016; Halse 2018). Since this habitat is restricted to an area of less than 1 km 2 on Mt. Snežnik, this subspecies is considered highly endangered. Phenology: probably eurychronous. Hybrids among the subspecies of Nemastoma bidentatum Remarks In Austria, a few localities are known where N. b. bidentatum lives in syntopy with N. relictum (Gruber & Martens 1968; Martens 1978; in both papers sub N. b. relictum), and with N. b. sparsum (Komposch & Gruber 2004; Komposch 2009). However, individuals of these three taxa do not interbreed and have been considered as potential species (Gruber & Martens 1968; Martens 1978; Komposch & Gruber 2004; Komposch 2009). In Slovenia, N. b. schmidti ssp. nov., with its central geographic position, interbreeds with all other adjacent subspecies: N. b. bidentatum in the North, and N. b. martensi ssp. nov., N. b. sneznikensis ssp. nov., N. b. gruberi ssp. nov. and N. b. sparsum from the Southwest to the Northeast of the country. Besides, N. b. gruberi ssp. nov. interbreed with N. b. martensi ssp. nov., and with N. b. sparsum. In Bosnia, N. pluridentatum stat. nov. has not been found; consequently, the genitalia have not been examined. Despite this, major differences from all the other known taxa within the N. b. complex allow it to be considered a separate species. This is because interbreeding of N. pluridentatum stat. nov., which might be a subalpine species, with the montane N. kozari sp. nov. from the same mountain is quite unlikely, given their great dissimilarity. However, data on taxa of Nemastoma in Bosnia and Herzegovina are nearly completely missing, allowing no further conclusions. Although being morphologically variable, hybrids possess larger Ch-Apo than pure subspecies and are well distinguishable from these. In interbreeding subspecies, hybrids between the two subspecies frequently occur syntopically with individuals belonging to pure subspecies. Consequently, such a collection consists of either one or two pure subspecies and a series of hybrids with varying Ch-Apo shapes. An overview of interbreeding relations among the taxa of the N. b. complex is presented in Table 13. In Slovenia, N. b. bidentatum and N. b. sparsum do not interbreed, since the areas are several tens of kilometres distant from each other. In contrast, we have found hybrids between all the adjacent subspecies (Fig. 3), except N. b. gruberi ssp. nov. �� N. b. schmidti ssp. nov. �� N. b. sparsum, which is expected in the meeting area of the three taxa. Besides, data on hybridization of N. b. sneznikensis ssp. nov. are too scarce to be properly evaluated in full. The zone of hybridization between N. b. schmidti ssp. nov. and N. b. sparsum is the widest, measuring up to 100 km in north-eastern Slovenia, while these zones are narrow to virtually non-identifiable among the others, and hybrids are accordingly scarce. Hybrids N. b. gruberi ssp. nov. �� N. b. schmidti ssp. nov. were found in a zone up to 10 km wide, while N. b. martensi ssp. nov. �� N. b. schmidti ssp. nov. and N. b. gruberi ssp. nov. �� N. b. martensi ssp. nov. are limited to zones a few kilometers wide. In Slovenia, there are no hybrids between allopatric N. b. martensi ssp. nov. and N. b. sparsum, while these are expected in Croatia. Hybridization patterns were not simple clines in the transition zones between two subspecies. Instead, in some cases, morphologically various hybrids between the neighboring subspecies, sharing various ratios of the hybridizing subspecies characters (cf. Gruber & Martens 1968; Martens 1978), appeared scattered across the whole width of the transition zone. In some places in the hybridization zone, individuals of both subspecies, together with hybrids of various intermittent stages, occurred syntopically., Published as part of Novak, Tone, Novak, Ljuba Slana, Kozel, Peter, Schaider, Miriam Gudrun, Komposch, Christian, Lipov��ek, Sa��ka, Podlesnik, Jan, Pau��i��, Igor & Raspotnig, G��nther, 2021, Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence, pp. 1-67 in European Journal of Taxonomy 777 on pages 40-42, DOI: 10.5852/ejt.2021.777.1561, http://zenodo.org/record/5637112, {"references":["Komposch C. 1999. Rote Liste der Weberknechte Karntens (Arachnida: Opiliones). In: Rottenburg T., Wieser Ch., Mildner P. & Holzinger W. E. (eds): Rote Listen gefahrdeter Tiere Karntens. Naturschutz in Karnten 15: 547 - 565. Amt der Karntner Landesregierung, Abt. 20, Landesplanung, Klagenfurt.","Komposch C. & Gruber J. 2004. Die Weberknechte Osterreichs (Arachnida, Opiliones). Denisia 12, zugleich Kataloge der Osterreichishen Landesmuseen Neue Serie 14: 485 - 534.","Mammola S., Giachino P. M., Piano E., Jones A., Barberis M., Badino G. & Isaia M. 2016. Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS). The Science of Nature 103: e 88. https: // doi. org / 10.1007 / s 00114 - 016 - 1413 - 9","Halse S. 2018. Research in calcretes and other deep subterranean habitats outside caves. In: Moldovan O. T., Kovac L. & Halse S. (eds) Cave Ecology: 415 - 434. Springer, Cham. https: // doi. org / 10.1007 / 978 - 3 - 319 - 98852 - 8","Gruber J. & Martens J. 1968. Morphologie, Systematik und Okologie der Gattung Nemastoma C. L. Koch (s. str.) (Opiliones, Nemastomatidae). Senckenbergiana Biologica 49 (2): 137 - 172.","Martens J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. In: Senglaub F., Hannemann H. & Schumann H. (eds) Die Tierwelt Deutschlands 64: 1 - 464. Fischer Verlag, Jena.","Komposch C. 2009. Opiliones (Weberknechte). In: Rabitsch W. & Essl F. (eds) Endemiten - Kostbarkeiten in Osterreichs Pflanzen- und Tierwelt: 476 - 500. Naturwissenschaftlicher Verein fur Karnten & Umweltbundesamt."]}

Nemastoma bidentatum schmidti Novak, Raspotnig & Slana Novak ssp. nov. urn:lsid:zoobank.org:act: 389E4C27-24B9-44F9-A724-11475CB9E12B Figs 2���3, 4H, 5H, 6H, 7A, H, 8A, H, 9H, 10H, 11H, 12H, 13G; Table 11 Nemastoma triste ��� Roewer 1917: 149 [partim: Maribor / Marburg], 151 [partim: Postojna /Adelsberg, Divača / Divaca, Ljubljana / Laibach, Zagreb / Agram]. Nemastoma bidentatum ��� Roewer 1923: 658 [partim: Postojna /Adelsberg, Ljubljana / Laibach, Zagreb / Agram; the other localities refer to N. bidentatum sparsum]. ��� Hadži 1927: 11���14, figs 81���93; 1928: 9; 1931: 108���109 [partim: Podkoren, Peričnik, Bled]. Nemastoma bidentatum bidentatum ��� Gruber & Martens 1968: 141, figs 3, 23 [partim: Leutscach]. ��� Marcellino 1973: 194. ��� Martens 1978: 107, fig 142 [partim: HR: Dugo selo, Zagreb; I: Laghi di Fusine; SLO: Bled, Cerknica, Ljubljana, Mt. Nanos, Postojna]. ��� Novak 2004a: 218. Nemastoma quadripunctatum humerale ��� Hadži 1931: 41, figs 94���98 (by lapsus). Nemastoma (Lugubrostoma) triste pluridentatum ��� Hadži 1973a: 43 figs 34a���c [partim: Mt. ��marna gora, Prestranek].? Nemastoma bidentatum bidentatum �� N. bidentatum sparsum ��� Novak & Gruber 2000: 285. Diagnosis Subspecies of Nemastoma bidentatum with elongated oval-trapezoid Ch-Apo, sagittally shallowly bilobe apically, wider distally, with a prominent apical pinnacle posteriorly, and conspicuously club-shaped Pa- Fe, with dorsal margin bent in the middle and nearly straight anterior and posterior portions, and evenly arched ventral margin, with 3 stout spines in distal fourth, and low, inconspicuous Pa-Ti hump. Rec sem of one tubular and 12 slightly elongated vesicles. Etymology The subspecies name �� schmidti �� is dedicated to Ferdinand Joseph Schmidt (1791���1878; Ljubljana), who collected the first specimens of this taxon preserved in his arthropod collection in the Slovenian Museum of Natural History, Ljubljana. Material examined Holotype SLOVENIA ��� VM31 ��� 1 ♂; Dolenja Sevnica, Poljane nad ��kofjo Loko; 46.12�� N, 14.15�� E; 517 m a.s.l.; 27 Mar. 2018; T. Novak leg.; mixed forest litter; PMSL-Opiliones-TN 1145/2018. Paratypes SLOVENIA ��� VM31 ��� 2 ♀♀; same collection data as for holotype; PMSL-Opiliones-GR 3809. Other material AUSTRIA ��� VM74 ��� 1 ♂; Eisenkappel / Železna Kapla ��� Ebriach / Obirsko, Carinthia; 7 May 2018; T. Novak leg. (38/2018); PMSL ��� 1 ♀; Seilach / Sele ��� Ebriach / Obirsko, Carinthia; 7 May 2018; T. Novak leg. (41/2018); PMSL. CROATIA ��� WL67 ��� 1 ♂, 1 ♀; Mt. Medvednica; A. Sch��nhofer leg. (6/2012, rev. 2019); PMSL. ITALY ��� UM52 ��� 7 ♂♂, 7 ♀♀; Artegna; 4 Mar.���11 Aug. 1992; F. Gasparo leg. (64/2002, rev. 2017); PMSL. ��� UM62 ��� 1 ♂; Alta Val Torre; 5 Aug. 1992; G. Governatori leg. (1062/2003, rev. 2017); PMSL ��� 1 ♂, 1 ♀; Villanova, Lusevera; 23 Oct. 1997; G. Governatori leg. (244/2003); PMSL ��� 3 ♂♂, 4 ♀♀; Tarcento / Čenta; 19 Aug. 2001; L. Slana Novak and T. Novak leg. (259/2001, rev. 2017); PMSL ��� 1 ♂, 5 ♀♀; ibid.; (257/2001, rev. 2018); PMSL ��� 1 ♂; Valle del Torre Vedronca valley; 5 Aug. 1992; G. Governatori leg. (1062/2003, rev. 2017); PMSL. ��� UM65 ��� 2 ♂♂, 1 ♀; Pontebba; 19 Aug. 2001; L. Slana Novak and T. Novak leg. (240/2001, rev. 2017); PMSL. ��� UM80 ��� 4 ♂♂; Altana / Utana; 26 Aug. 2001; L. Slana Novak and T. Novak leg. (283/2001, rev. 2017); PMSL. SLOVENIA ��� 1 ♂, 1 ♀ (Schmidt���s collection); (84/1985 microscopic preparation, JH det. sub N. triste, TN rev.); PMSL ��� 1 ♂, 1 ♀ (Schmidt���s collection); (89/1985 microscopic preparation, JH det. sub N. triste, TN rev.); PMSL. ��� UL99 ��� 1 ♂, 2 ♀♀; Panovec; 17 Aug. 2000; T. Novak leg. (668/2000, rev. 2017); PMSL ��� 1 ♀; ibid.; 18 Mar. 2001; S. Brelih leg. (63/2001, TN rev. 2019); PMSL ��� 9 ♂♂, 5 ♀♀; ibid.; 11 Nov. 2006; L. Slana Novak and T. Novak leg. (152/2006, rev. 2017); PMSL ��� 1 ♂; Replje; 8 Apr. 1992; S. Brelih leg. (649/1998, rev. 2017); PMSL. ��� UM72 ��� 1 ♂; Mt. Kobari��ki Stol; 24 May 2003; L. Slana Novak and T. Novak leg. (720a/2003, TN and SL rev. 2017); PMSL. ��� UM73 ��� 1 ♂; Mt. Kobari��ki Stol; 8 Jun. 2017; T. Novak leg. (6/2017); PMSL. ��� UL82 ��� 1 ♀; Tavenica, Bela; 10 Oct. 1995; S. Brelih leg. (789/1998, rev. 2018); PMSL ��� 1 ♂; Sedlo ��� Mt. Kobari��ki Stol; 1 Aug. 1998; L. Slana Novak and T. Novak leg. (52/1999, rev. 2018); PMSL. ��� UM90 ��� 2 ♀♀; Avče; 2 Aug. 2000; T. Novak leg. (298/2000, rev. 2019); PMSL ��� 2 ♂♂; Doblar; 2 Aug. 2000; leg. T. Novak (285/2000, rev. 2017); PMSL ��� 1 ♀; ibid.; 17 Jun. 2001; L. Slana Novak and T. Novak leg. (151/2001, rev. 2017); PMSL ��� 1 ♂; Lepenka valley, Doblar; 25 Sep. 1998; L. Slana Novak and T. Novak leg. (1795/1998, rev. 2017); PMSL ��� 1 ♂; Levpa ��� Seni��ki breg; 2 Aug. 2000; T. Novak leg. (319/2000, rev. 2017); PMSL ��� 1 ♀; Marija Snežna, Avče; 2 Aug. 2000; T. Novak leg. (293/2000; rev. 2019); PMSL. ��� UL91 ��� 1 ♂; Kamno; 31 Jul. 1998; L. Slana Novak and T. Novak leg. (1337/1998, rev. 2018); PMSL ��� 1 ♂, 1 ♀; ibid.; 25 Sep. 1998; L. Slana Novak and T. Novak leg. (1809/1998, rev. 2017); PMSL ��� 2 ♂♂, 2 ♀♀; Solarji, Liv��ke Ravne; 8 Aug. 2000; T. Novak leg. (562/2000, TN and SL rev. 2017); PMSL. ��� UM92 ��� 6 ♂♂, 1 ♀; Ladrski vrh; 14 May���11 Jun. 2002; B. Drovenik and A. Gergeli leg. (482/2018); PMSL ��� 1 ♀; ibid.; 11 Jun.���7 Aug. 2002; B. Drovenik and A. Pirnat leg. (792/2018); PMSL ��� 2 ♂♂, 3 ♀♀; ibid.; 8 Jul.���7 Aug. 2002; B. Drovenik and A. Pirnat leg. (PK 9/2013, TN rev. 2019); PMSL ��� 1 ♀; ibid.; (PK 10/2013, TN rev. 2019); PMSL ��� 1 ♀; ibid.; (585/2018); PMSL ��� 1 ♀; ibid.; 7 Aug.���4 Sep. 2002; B. Drovenik and A. Pirnat leg. (780/2018); PMSL ��� 1 ♂; ibid.; (787/2018); PMSL ��� 2 ♂♂, 6 ♀♀; ibid.; 4 Sep.���1 Oct. 2002; B. Drovenik and A. Pirnat leg. (827/2018); PMSL ��� 2 ♂♂; ibid.; (910/2018); PMSL ��� 4 ♀♀; ibid.; (915/2018); PMSL ��� 3 ♀♀; ibid.; (923/2018); PMSL ��� 2 ♀♀; ibid.; (928/2018); PMSL ��� 1 ♂; ibid.; (933/2018); PMSL ��� 3 ♂♂, 5 ♀♀; ibid.; 1���29 Oct. 2002; B. Drovenik and A. Pirnat leg. (959/2018); PMSL ��� 1 ♂, 3 ♀♀; ibid.; (961/2018); PMSL ��� 10 ♀♀; ibid.; (965/2018); PMSL ��� 4 ♀♀, 1 juv.; ibid.; (971/2018); PMSL ��� 1 ♂, 1 ♀; ibid.; (974/2018); PMSL ��� 3 ♂♂, 6 ♀♀; ibid.; (981/2018); PMSL ��� 8 ♀♀; ibid.; (1003/2018); PMSL ��� 1 ♂; ibid.; 29 Oct.���7 Nov. 2002; B. Drovenik and A. Gergeli leg. (PK 36/2013, TN rev. 2019); PMSL ��� 3 ♀♀; ibid.; (PK 37/2013, TN rev. 2019); PMSL ��� 2 ♂♂, 13 ♀♀; ibid.; (871/2018); PMSL ��� 2 ♀♀; ibid.; (877/2018); PMSL ��� 1 ♂, 3 ♀♀; ibid.; (880/2018); PMSL ��� 2 ♀♀; ibid.; (881/2018); PMSL ��� 3 ♂♂, 4 ♀♀; ibid.; (897/2018); PMSL ��� 4 ♂♂, 6 ♀♀; ibid.; (900/2018); PMSL ��� 3 ♂♂, 1 ♀; ibid.; (939/2018); PMSL ��� 2 ♀♀; ibid.; 27 Nov.���21 Dec. 2002; B. Drovenik and A. Gergeli leg. (948/2018); PMSL ��� 3 ♂♂, 2 ♀♀; ibid.; (950/2018); PMSL ��� 1 ♂; ibid.; (1010/2018); PMSL ��� 3 ♀♀; ibid.; (1014/2018); PMSL ��� 2 ♀♀; ibid.; (1019/2018); PMSL ��� 1 ♂; ibid.; (1026/2018); PMSL ��� 1 ♂, 2 ♀♀; ibid.; 21 Dec. 2002 ��� 28 Jan. 2003; B. Drovenik and A. Pirnat leg. (718/2018); PMSL ��� 1 ♂, 2 ♀♀; ibid.; (742/2018); PMSL ��� 2 ♂♂, 1 ♀; ibid.; (772/2018); PMSL ��� 3 ♀♀; ibid.; (778/2018); PMSL ��� 3 ♂♂, 2 ♀♀; ibid.; (813/2018); PMSL ��� 2 ♂♂, 3 ♀♀; ibid.; (1007/2018); PMSL ��� 1 ♂; ibid.; (1013/2018); PMSL. ��� UL92 ��� 1 ♂, 1 ♀; near Kamno; Sep.1915; Andreini leg.; PMSL ��� 1 ♂, 2 ♀♀; Selce; 10 Oct. 1995; S. Brelih leg. (786/1998, rev. 2017); PMSL ��� 1 ♂; ��turmi; 8 Aug. 2000; T. Novak leg. (545/2000, rev. 2017); PMSL. ��� UL93 ��� 1 ♀; Bav��ica; 9 Aug. 1996; S. Brelih leg. (744/1998, rev. 2018); PMSL ��� 3 ♀♀; Brvce; 11 Oct. 1995; S. Brelih leg. (745/1998, rev. 2018); PMSL. ��� VL07 ��� 1 ♀; Velike Žablje; 17 Aug. 2000; T. Novak leg. (677/2000, rev. 2017); PMSL. ��� VL09 ��� 1 ♂, 2 ♀♀; Čepovanska reber; 2 Aug. 2000; T. Novak leg. (344/2000, rev. 2017); PMSL ��� 1 ♂; Lijak spring, Ozeljan; 17 Apr. 2002; B. Drovenik and A. Gergeli leg. (219/2011, rev. 2017); PMSL ��� 1 ♂; ibid.; 17 Apr.���14 May 2002; B. Drovenik and A. Pirnat leg. (540/2018); PMSL ��� 2 ♂♂; ibid.; (553/2018); PMSL ��� 2 ♂♂, 3 ♀♀; ibid.; 14 May���11 Jun. 2002; B. Drovenik and A. Gergeli leg. (503/2018); PMSL ��� 1 ♀; ibid.; (518/2018); PMSL ��� 1 ♂; ibid.; 11 Jun. 2002; B. Drovenik and A. Pirnat leg. (212/2011, rev. 2017); PMSL ��� 1 ♂; ibid.; 8 Jul. 2002; B. Drovenik and A. Pirnat leg. (1093/2018); PMSL ��� 1 ♀; ibid.; (1097/2018); PMSL ��� 1 ♀; ibid.; 7 Aug.���4 Sep. 2002; B. Drovenik and A. Gergeli leg. (567/2018); PMSL ��� 1 ♂; ibid.; 1 Oct. 2002; B. Drovenik and A. Pirnat leg. (313/2018); PMSL ��� 1 ♂, 1 ♀; ibid.; 18 Feb. 2003; B. Drovenik and A. Gergeli leg. (200/2011, TN rev. 2017); PMSL ��� 1 ♂; Lokve; 2 Aug. 2000; T. Novak leg. (350/2000, rev. 2017); PMSL ��� 1 ♀; ibid.; (364/2000, rev. 2017); PMSL ��� 1 ♀; Smrečje, Trnovski gozd; 18 Aug. 1973; I. Sivec leg. (519/1981, JG det. 1982, TN rev. 2018); PMSL. ��� VL16 ��� Divača ��� VL19 ��� 1 ♂; Gorenja Kanomlja; 1 Aug. 2000; T. Novak leg. (401/2000, rev. 2017); PMSL ��� 1 ♀; Predmeja; 21 Aug. 2006; L. Slana Novak and T. Novak leg. (4/2006, rev. 2017); PMSL ��� 2 ♀♀; Srednja Kanomlja; 1 Aug. 2000; T. Novak leg. (391/2000, rev. 2017); PMSL. ��� VL26 ��� 3 ♂♂, 3 ♀♀; Senožeče; 20 Sep. 2011; L. Slana Novak and T. Novak leg. (262/2011, rev. 2017); PMSL ��� 1 ♀; Mt. Vrem��čica; 10 Mar. 1994; S. Brelih leg. (680/1998, rev. 2017); PMSL. ��� VL28 ��� 2 ♂♂, 1 ♀; Črni vrh ��� Col; L. Slana Novak and T. Novak leg. (239/2011, rev. 2017); PMSL. ��� VL29 ��� 3 ♂♂, 3 ♀♀; Godovič ��� Črni vrh; 11 Sep. 2011; L. Slana Novak and T. Novak leg. (241/2011, rev. 2017); PMSL ��� 2 ♂♂, 1 ♀; Gore; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (155/2000, rev. 2017); PMSL ��� 1 ♀; Idrija; Jun. 1997; M. Vončina leg. (1702/1998, rev. 2017); PMSL ��� 1 ♂; ibid.; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (137/2000, rev. 2017); PMSL ��� 8 ♂♂, 1 ♀; ibid.; (229/2000, TN and SL rev. 2017); PMSL ��� 1 ♂; ibid.; 1 Aug. 2000; T. Novak leg. (382/2000, rev. 2017); PMSL ��� 2 ♂♂; Razpotje; 1 Aug. 2000; T. Novak leg. (408/2000, rev. 2017); PMSL ��� 1 ♂; Spodnja Idrija; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (143/2000, rev. 2017); PMSL ��� 2 ♂♂; Srednja Kanomlja; 1 Aug. 2000; T. Novak leg. (389/2000, rev. 2017); PMSL ��� 1 ♀; ibid.; (397/2000, rev. 2017); PMSL ��� 1 ♂; Urbanovec; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (149/2000, rev. 2017); PMSL. ��� VL36 ��� 1 ♂; Jama Sv. Janeza cave, Cad. No. 897, Prestranek; 11 Aug. 1952; E. Pretner leg. (56/1983, JH det., TN rev. 2017); PMSL ��� 1 ♀; ibid.; 29 Aug. 1954; E. Pretner leg. (174/1984, JH det., TN rev. 2018); PMSL ��� 1 ♂; ibid.; 11 Aug. 1957; E. Pretner leg. (221/1984, TN rev. 2017); PMSL ��� 3 ♀♀; Jernejeva jama cave, Cad. No. 929, Koče; 22 Apr. 2001; S. Vidmar, S. Polak, T. Novak leg. (157/2001, rev. 2008); PMSL ��� Postojna ��� VL37 ��� 1 ♂; Belsko; 14 Apr. 2011; L. Slana Novak and T. Novak leg. (86/2011); PMSL ��� 1 ♂; Landol; 9 May 1994; S. Brelih leg. (888/1998); PMSL. ��� VL38 ��� 2 ♂♂, 3 ♀♀; Dolenji Novi Svet; L. Slana Novak and T. Novak leg. (230/2011); PMSL ��� 1 ♀; Novi svet, Hoteder��ica; 22 Mar. 1993; S. Brelih leg. (855/1998, rev. 2019); PMSL. ��� VL39 ��� 3 ♂♂, 3 ♀♀; Kugu, Rovte; 11 Sep. 2011; L. Slana Novak and T. Novak leg. (244/2011); PMSL. ��� VL46 ��� 1 ♂, 1 ♀; Mt. Javorniki; 9 May 1994; S. Brelih leg. (836/1998); PMSL ��� 2 ♂♂, 8 ♀♀; Mali Kožljek, Mt. Javorniki; 2 Oct. 2011; L. Slana Novak and T. Novak leg. (336/2011); PMSL. ��� VL47 ��� 1 ♂, 2 ♀♀; Planina pri Rakeku; 24 Mar. 1992; F. Velkovrh leg. (221/1995, rev. 2017); PMSL ��� 1 ♂; Rakov ��kocjan; Jul. 1994; K. Musar leg. (79/1995, rev. 2017); PMSL ��� 1 ♂; ibid.; 12.���15 May 1994; S. Polak leg. (640/1998, rev. 2018); NM ��� 7 ♂♂, 17 ♀♀; ibid.; 10 Sep. 2009; A. Sch��nhofer leg. (3/2012, rev. 2017); PMSL. ��� VL48 ��� 1 ♀; Bistra; 29 Mar. 1993; S. Brelih leg. (641/1998, rev. 2017); PMSL ��� 1 ♂; Laze; 16 Oct.���21 Nov. 1994; M. Kuntner leg. (176/1995, rev. 2017); PMSL ��� 1 ♀; ibid.; Jul.��� Aug.1994; M. Kuntner leg. (20/1997, rev. 2017); PMSL ��� 1 ♂, 1 ♀; ibid.; Aug. 1994; M. Kuntner leg. (183/1995, rev. 2017); PMSL ��� 1 ♂, 1 ♀; ibid.; Dec. 1994 ��� Mar. 1995; M. Kuntner leg. (170/1995, rev. 2017); PMSL ��� 1 ♀; Vranja jama cave, Cad. No. 88, Laze pri Planini; 7 Jun. 1994; stud. biol. leg. (101/2019); PMSL. ��� VL49 ��� 1 ♀; Retovje, Vrhnika; Jul. 1994, A. Valenčak leg. (17/1997, rev. 2017); PMSL ��� 1 ♂; ibid.; Jun. 1998; M. Zagmajster leg. (1493/1998, rev. 2017); PMSL ��� 1 ♀; Vrhnika; Jul. 1994; A. Valenčak leg. (17/1997, rev. 2017); PMSL. ��� VL54 ��� 1 ♂; Mt. Snežnik; 21 Jul. 2001; B. Bertoncelj, U. Bertoncelj and T. Novak leg. (150/2001, rev. 2017); PMSL ��� 1 ♀; ibid.; 24 Nov. 2003; S. Polak leg. (120/2006, rev. 2018); NM; PMSL. ��� VL56 ��� 1 ♂, 3 ♀♀; Cerkni��ko jezero; 19 Oct. 1994; S. Polak leg. (721/1998, rev. 2017); NM ��� 1 ♀; ibid.; 20 Nov. 1994; S. Polak leg. (687/1998, rev. 2017); NM ��� 2 ♂♂, 1 ♀; ibid.; 24 Apr. 2001; S. Brelih leg. (72/2001, rev. 2017); PMSL. ��� VL57 ��� 2 ♂♂, 4 ♀♀; Dolenje Otave ��� Sel��ček; 14 Aug. 2011; L. Slana Novak and T. Novak leg. (36/2011, rev. 2017); PMSL. ��� VL58 ��� 1 ♂; Mt. Krim; 14 Aug. 2011; L. Slana Novak and T. Novak leg. (42/2011, rev. 2017); PMSL ��� 2 ♀♀; Rakitna; 14 Aug. 2011; L. Slana Novak and T. Novak leg. (40/2011, rev. 2017); PMSL. ��� VL59 ��� 2 ♂♂, 1 ♀; Bevke; 20 Oct. 2018; L. Slana Novak and T. Novak leg. (136/2018); PMSL ��� 1 ♂; Brezovica pri Ljubljani; 25 May 1997; L. Slana Novak and T. Novak leg. (14/1997, rev. 2018); PMSL ��� 1 ♀; Draga pri Igu; 13 Apr. 1995; S. Brelih leg. (732/1998, rev. 2018); PMSL ��� 3 ♂♂, 10 ♀♀; Žabnica; 14 Aug. 2011; L. Slana Novak and T. Novak leg. (33/2011, rev. 2017); PMSL. ��� VL68 ��� 1 ♂, 3 ♀♀; I��ka vas ��� Mt. Krim; 14 Aug. 2011; L. Slana Novak and T. Novak leg. (48/2011, rev. 2017); PMSL ��� 1 ♂; ibid.; 31 Mar. 1997; S. Brelih leg. (53/2001, rev. 2017); PMSL ��� 1 ♀; Kremenica, Ig; 29 Jun. 1980 (1155/1981, JG det., TN rev. 2018); PMSL ��� 1 ♀; ibid.; 2 Mar. 1996; S. Brelih leg. (45/2001, rev. 2017); PMSL ��� 1 ♂; ibid.; 1 May 1997; S. Brelih leg. (101/2001, rev. 2017); PMSL ��� 1 ♂; ibid.; 7 Jun. 1997; S. Brelih leg. (117/2001, rev. 2017); PMSL ��� 1 ♂; ibid.; 10 Jul. 1997; S. Brelih leg. (139/2001, rev. 2017); PMSL ��� 1 ♂, 1 ♀; ibid.; 25 Aug. 1997; S. Brelih leg. (113/2001, rev. 2017); PMSL ��� 1 ♀; Mt. Mokrec; 18 Jul. 1980 (LSN 151/1984, TN rev. 2017); PMSL ��� 1 ♀; ��čurki, Turjak; 3 May 2008; L. Slana Novak and T. Novak leg. (155/2011, rev. 2017); PMSL ��� 1 ♀; ibid.; 20 Jun. 1980; (1149/1981, JG det., TN rev. 2018); PMSL ��� 1 ♀; ��krilje; 20 Jun. 1980; (1159/1981, JG det., TN rev. 2018); PMSL. ��� VL69 ��� 2 ♀♀; ��marje Sap; 10 May 2008; L. Slana Novak and T. Novak leg. (119a/2011, rev. 2017); PMSL ��� 1 ♂, 1 ♀; ibid.; (147/2011, rev. 2018); PMSL. ��� VL79 ��� 1 ♂, 4 ♀♀; Brinjski hrib; L. Slana Novak and T. Novak leg. (99/2012, TN and SL rev. 2017); PMSL ��� 1 ♂; Stara vas; 16 Mar. 1993; S. Brelih leg. (665/1998, rev. 2017); PMSL. ��� VL85 ��� 1 ♀; Mt. Stojna; 21 Oct. 1996; S. Brelih leg. (90/2001, rev. 2017); PMSL. ��� VL89 ��� 1 ♂, 4 ♀♀; Vintarjevec; T. Novak leg. (173/2011, rev. 2017); PMSL ��� 2 ♂♂, 4 ♀♀; ibid.; (178/2011, rev. 2017); PMSL. ��� VL93 ��� 1 ♀; Kostel, Dolenja Žaga; 16 Apr. 1996; S. Brelih leg. (737/1998, rev. 2018); PMSL. ��� VL99 ��� 1 ♀; Cerovec, Gabrovka; L. Slana Novak and T. Novak leg. (74/2007, rev. 2017); PMSL. ��� VM00 ��� 1 ♂, 2 ♀♀; Cvetrež, Voglarji; 2 Aug. 2000; T. Novak leg. (324/2000, TN and SL rev. 2017); PMSL ��� 5 ♂♂, 4 ♀♀; Čepovan; 1 Aug. 2000; T. Novak leg. (361/2000, rev. 2017); PMSL ��� 1 ♂; Čepovan ��� Most na Soči; 10 Jun. 1997; S. Brelih leg. (110/2001, rev. 2017); PMSL ��� 1 ♂; Preval, Lokovec, Čepovan; 2 Aug. 2000; T. Novak leg. (329/2000, rev. 2017); PMSL. ��� VM05 ��� several specs; Podkoren; J. Hadži leg.; PMSL. ��� VM09 ��� 1 ♀; Smrečje; 18 Aug. 1973; I. Sivec leg. (519/1981, JG det., TN rev. 2018); PMSL. ��� VM10 ��� 1 ♂; Bukovo; 8Aug. 2000; T. Novak leg. (501/2000, rev. 2017); PMSL ��� 3 ♂♂, 5 ♀♀; Oblakov vrh; 1 A, Published as part of Novak, Tone, Novak, Ljuba Slana, Kozel, Peter, Schaider, Miriam Gudrun, Komposch, Christian, Lipov��ek, Sa��ka, Podlesnik, Jan, Pau��i��, Igor & Raspotnig, G��nther, 2021, Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence, pp. 1-67 in European Journal of Taxonomy 777 on pages 30-40, DOI: 10.5852/ejt.2021.777.1561, http://zenodo.org/record/5637112, {"references":["Roewer C. F. 1917. Uber Nemastomatiden und ihre Verbreitung. Archives fur Naturgeschichte 83 A (2): 140 - 160.","Roewer C. F. 1923. Die Weberknechte der Erde. Gustav Fischer, Jena.","Hadzi J. 1927. Opilioni Schmidtove zbirke. Glasnik Muzejskega Drustva za Slovenijo 7 - 8: 1 - 41 + VIII.","Gruber J. & Martens J. 1968. Morphologie, Systematik und Okologie der Gattung Nemastoma C. L. Koch (s. str.) (Opiliones, Nemastomatidae). Senckenbergiana Biologica 49 (2): 137 - 172.","Marcellino I. 1973. Notizie su Opilioni (Arachnida) italiani e dell'Alto Isonzo. Annali del Museo Civico di Storia Naturale di Genova 79: 192 - 205.","Martens J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. In: Senglaub F., Hannemann H. & Schumann H. (eds) Die Tierwelt Deutschlands 64: 1 - 464. Fischer Verlag, Jena.","Novak T. 2004 a. Harvestmen of the Museo Friulano di Storia Naturale in Udine (Arachnida: Opiliones). Part 1. Gortania 26: 211 - 241.","Hadzi J. 1931. Opilioni Triglavskoga masiva. Prirodoslovne razprave 1: 107 - 154.","Hadzi J. 1973 a. Novi taksoni suhih juzin (Opilionidea) v Jugoslaviji. Razprave SAZU, IV. razred 16 (1): 1 - 120.","Novak T. & Gruber J. 2000. Remarks on published data on harvestmen (Arachnida: Opiliones) from Slovenia. Annales, Series historiae naturalis 10: 281 - 308.","Hadzi J. 1928. Beitrag zur Kenntnis der Opilioniden-Fauna von Slowenien (Jugoslawien). Zoologischer Anzeiger 77 (1 - 2): 5 - 19.","Novak T., Gruber J. & Slana L. 1984. Remarks on Opiliones from cavities in Slovenia (Yugoslavia). Memoires de Biospeologie 11: 185 - 197.","Novak T., Gruber J. & Slana L. 1995. A contribution to the knowledge of the harvestmen (Opiliones) from the submediterranean region of Slovenia. Annales, Series historiae naturalis 5: 181 - 192.","Novak T., Gruber J. & Slana L. 1996. Weberknechte (Opiliones) des Zentral-Europaischen zoogeographischen Gebietes Sloweniens. Znanstvena revija, Maribor 7: 47 - 60.","Novak T., Slana L., Cervek N., Mlakar M., Zmaher N. & Gruber J. 2002. Harvestmen (Opiliones) in human settlements of Slovenia. Acta Entomologica Slovenica 10: 131 - 154.","Novak T., Janzekovic F., Rak M., Ivajnsic D., Kozel P., Slana Novak L. & Skornik S. 2017. Can highland habitat type distributions replace mapping of harvestman species? Journal of Insect Conservation 21 (2): 331 - 343. https: // doi. org / 10.1007 / s 10841 - 017 - 9979 - x","Komposch C. & Gruber J. 2004. Die Weberknechte Osterreichs (Arachnida, Opiliones). Denisia 12, zugleich Kataloge der Osterreichishen Landesmuseen Neue Serie 14: 485 - 534.","Novak T. 2005 b. The harvestmen fauna (Arachnida: Opiliones) from the Sub Mediterranean region of Slovenia - II. Annales, Series historiae naturalis 15: 103 - 114.","Novak T. 2005 c. Terrestrial fauna from cavities in Northern and Central Slovenia, and a review of systematically ecologically investigated cavities. Acta Carsologica 34: 169 - 210. https: // doi. org / 10.3986 / ac. v 34 i 1.285","Roewer C. F. 1951. Uber Nemastomatiden. Weitere Weberknechte XVI. Senckenbergiana 32: 95 - 153."]}

Nemastoma bidentatum gruberi Novak, Slana Novak, Kozel & Raspotnig ssp. nov. urn:lsid:zoobank.org:act: 17EA28A1-5F36-4DFC-9191-9C8680496FAC Figs 1A, C, 2– 3, 4E, 5E, 6E, 7E, 8E, 9E, 10E, 11E, 12E, 13D; Table 8 Nemastoma bidentatum sparsum – Novak et al. 2002: 136 [partim: Brežice, Črnomelj, Kočevje, Krško, Litija, Metlika, Novo mesto, Ribnica, Sevnica,Trebnje]. — Novak 2004b: 245 [partim: Buje, Karojba, Vižinada]; 2005b: 105. Diagnosis Relatively long-legged subspecies of Nemastoma bidentatum, with strait Ch ventral margin, and Pa-Fe with either 4 subequidistantly positioned spines or pointed tubercles or long setae on low protrusions, or their combination in the distal Pa-Fe half, or alternatively, with 2–3 distal stout thorns. Rec sem with one tubular and 12 elongated balloon-like vesicles. Etymology The subspecies name ʻ gruberi ʼ is dedicated to Jürgen Gruber (Vienna), our teacher, colleague and friend, who generously introduced us to the taxonomy of harvestmen and helped in identifying problematic specimens. Material examined Holotype SLOVENIA – WL18 • 1 ♂; Dolenje Laknice, Mokronog; 45.93° N, 15.21° E; 237 m a.s.l.; 26 Oct. 2013; L. Slana Novak and T. Novak leg.; mixed forest litter sift; PMSL-Opiliones-GR 4661. Paratypes SLOVENIA – WL18 • 1 ♀; same collection data as for holotype; PMSL-Opiliones-GR 4664 • 5 ♂♂, 4 ♀♀; same collection data as for holotype; PMSL-Opiliones-GR4662, GR4666, GR4667, GR4669, GR4673, GR4676 to GR4679. Other material ITALY – VL05 • 1 ♀; Chiusa/Ricmanje; 29 Sep. 1999; L. Slana Novak and T. Novak leg. (275/1999, rev. 2015); PMSL. – VL15 • 1 ♀; Basovizza/Bazovica; 29 Sep. 1999; L. Slana Novak and T. Novak leg. (271/1999, rev. 2015); PMSL. CROATIA – UL92 • 3 ♂♂, 2 ♀♀ (paratypes of N. b. sparsum); vicinity of Buje, and VL02, Vižinada; Sep. 1963; R. Sturany leg.; NHMW 4700. – VL01 • 1 ♂; Karojba; 28 Feb. 1990; F. Potočnik leg. (1307/1998, rev. 2018); PMSL. – VL13 • 1 ♀; Mlini; 30 Sep. 1990; L. Slana Novak and T. Novak leg. (32/1990, rev. 2015); PMSL. – XL90 • 1 ♂, 2 ♀♀; Ponor Sušik, Drežnik; 8 Sep. 2009; A. Schönhofer leg. (Coll. ASc 334, ASc det., TN rev. 5/2012); PMSL. SLOVENIA – UL93 • 4 ♀♀, 1 juv.; Abrami; 19 Jul. 1999; L. Slana Novak and T. Novak leg. (148/1999, rev. 2018); PMSL • 2 ♀♀; ibid.; 20 Jul. 1999; L. Slana Novak and T. Novak leg. (188/1999, rev. 2015); PMSL • 1 ♀; Dragonja; 14 Sep. 2000; L. Slana Novak and T. Novak leg. (154/2001, rev. 2015); PMSL • 4 ♂♂, 4 ♀♀; Dragonja valley; 26 Jun. 2015; T. Novak leg. (78/2015 = GR 5123−5130); PMSL • 2 ♂♂, 3 ♀♀; Ivankovec; 23 Sep. 1998; L. Slana Novak and T. Novak leg. (1791/1998, rev. 2018); PMSL • 1 ♀; Lucija–Seča – Debernardo; 22 Sep. 1998; L. Slana Novak and T. Novak leg. (1817/1998, rev. 2019); PMSL • 1 ♂; Korte; 23 Sep. 1998; L. Slana Novak and T. Novak leg. (1821/1998, rev. 2018); PMSL • 33 ♂♂, 30 ♀♀; Medljan, Korte; 21 Aug. 1999; T. Novak leg. (239/1999, rev. 2018); PMSL • 1 juv.; ibid.; 22 Jun. 2000; B. Bertoncelj, B. Ogrizek and T. Novak leg. (217/2000, rev. 2018); PMSL • 1 ♀; ibid.; (218/2000, TN rev. 2019); PMSL • 1 ♂; Padna; 19 Jul. 1999; L. Slana Novak and T. Novak leg. (168/1999, rev. 2017); PMSL • 2 ♀♀; Parecag; 23 Sep. 1998; L. Slana Novak and T. Novak leg. (1788/1998, rev. 2018); PMSL • 1 ♀; Seča; 10 Oct. 1998; L. Slana Novak and T. Novak leg. (1844/1998, rev. 2008); PMSL • 1 ♂, 1 ♀; ibid.; 24 Oct. 1998, L. Slana Novak and T. Novak leg. (1881/1998, rev. 2007); PMSL • 3 ♀♀; ibid.; 21 Aug. 1999; M. Štangelj and T. Novak leg. (236/1999, rev. 2008); PMSL • 15 ♂♂, 21 ♀♀; ibid.; 21 Aug. 1999; M. Štangelj and T. Novak leg. (231/1999, rev. 2015); PMSL • 1 ♂, 6 ♀♀; Sečovlje; 1 Mar. 2009; L. Slana Novak and T. Novak leg. (1/2015); PMSL • 1 ♂; Sv. Peter; 11 Oct. 1998; L. Slana Novak and T. Novak leg. (1850/1998, rev. 2015); PMSL • 4 ♂♂, 2 ♀♀; Sv. Sab, Padna; 19 Jul. 1999; L. Slana Novak and T. Novak leg. (151/1999, rev. 2009); PMSL • 1 ♀; Škrline, Dragonja; 20 Jul. 1999; L. Slana Novak and T. Novak leg. (194/1999, rev. 2015); PMSL. – VL03 • 2 ♂♂, 1 ♀; Boršt; 14 Apr. 2011; T. Novak leg. (76/2011, rev. 2015); PMSL • 2 ♀♀; ibid.; (81/2011, rev. 2015); PMSL. – VL04 • 1 ♀; Ankaran; 29 Sep. 1999; L. Slana Novak and T. Novak leg. (276/1999, rev. 2015); PMSL • 1 ♂, 1 ♀; Kolombini, Spodnje Škofije; 26 Sep. 2007; L. Slana Novak and T. Novak leg. (94/2007, rev. 2018); PMSL • 5 ♂♂; Mlinarji – Osp; 26 Sep. 2007; L. Slana Novak and T. Novak leg. (97/2007, rev. 2017); PMSL • 1 ♂, 1 ♀; Rižana; 24 Sep. 2016; L. Slana Novak and T. Novak leg. (18/2018); PMSL • 1 ♀; Škocjanski zatok, Koper; 15 Oct. 2009; S. Polak leg. (PK 1/2015); NM • 4 ♂♂, 3 ♀♀; Zgornje Škofije – Tinjan; 26 Sep. 2007; L. Slana Novak and T. Novak leg. (88/2007); PMSL. – VL07 • 4 ♀♀; Hrastje, Šmarje; 17 Aug. 2000 (674/2000, rev. 2015); PMSL. – VL13 • 1 ♂; Poletiči, Kubed; 4 Sep. 2016; T. Marinko leg. (27/2016); PMSL • 1 ♂, 1 ♀; Sočerga; 28 Sep. 1990; L. Slana Novak and T. Novak leg. (31/1990, rev. 2015); PMSL. – VL14 • 4 ♂♂, 4 ♀♀; Bezovica; 29 Sep. 1990; L. Slana Novak and T. Novak leg. (LSN 1/1990, TN rev. 2018); PMSL • 1 ♂; Hrpelje, Kozina; 2 Apr. 1993; S. Brelih leg. (651/1998, rev. 2018); PMSL • 2 ♂♂; Predloka; 29 Sep. 1990; L. Slana Novak and T. Novak leg. (35/1990, rev. 2015); PMSL • 2 ♂♂, 1 ♀; Rižana; 29 Sep. 1990; L. Slana Novak and T. Novak leg. (33/1990, rev. 2015); PMSL. – VL15 • 1 ♀; Botač; 22 Mar. 1990; M. Černila, B. Horvat and I. Sivec leg. (1246/1997); PMSL. – VL16 • 1 ♂; Divača; 2 Apr. 1993; S. Brelih leg. (174/2001, rev. 2015); PMSL. – VL17 • 1 ♂; Vrabče, Podnanos; 10 Mar. 1994; S. Brelih leg. (632/1998, rev. 2017); PMSL. – VL18 • 1 ♂; Uhanje; 3 Sep. 2001; T. Novak leg. (329/2001, rev. 2018); PMSL. – VL23 • 1 ♂, 1 ♀; Poljane pri Podgradu; 18 Aug. 2011; L. Slana Novak and T. Novak leg. (166/2012, rev. 2019); PMSL. – VL25 • 3 ♀♀; Buje; 10 Mar. 1994; S. Brelih leg. (163/2001, rev. 2018); PMSL • 1 ♂, 2 ♀♀; Goriče pri Framljah; L. Slana Novak and T. Novak leg. (343/2011); PMSL • 1 ♀; ibid.; (344/2011); PMSL. – VL28 • 1 ♂, 1 ♀; Podvrh, Raka; 18 Jul. 2012; P. Kozel, T. Kozel and T. Novak leg. (55/2012); PMSL • 9 ♂♂, 7 ♀♀; ibid.; (GR 3734–3746); PMSL • 3 ♀♀; ibid.; (GR 3791); PMSL. – VL29 • 6 ♂♂, 12 ♀♀; Dolnje Brezovo, Sevnica; 30 Jun. 2012; (GR 3670–3683); PMSL. – VL36 • 1 ♂, 1 ♀; Palčje – Mt. Javorniki; 1 Oct. 2011; L. Slana Novak and T. Novak leg. (361/2011, rev. 2019); PMSL. – VL45 • 1 ♀; Koritnice − Mašun; 2 Oct. 2011; L. Slana Novak and T. Novak leg. (329/2011, rev. 2014); PMSL. – VL47 • 1 ♀; Suha reber, Koritnice; 21 Jan. 2007; S. Polak leg. (107/2007); NM. – VL54 • 1 ♀; Sviščaki; 20 May 1989; T. Trilar leg. (271/2000, rev. 2017); PMSL • 1 ♀; ibid.; 25 May 1989; T. Trilar leg. (275/2000, rev. 2017); PMSL • 1 ♀; ibid.; 21 Jul. 2001; B. Bertoncelj, U. Bertoncelj and T. Novak leg. (29/2001, rev. 2018); PMSL • 1 ♀; ibid.; (34/2001, rev. 2011); PMSL • 1 ♂, 3 ♀♀; Travni Dolci, Mt. Snežnik; 12 Aug. 2001; L. Slana Novak and T. Novak leg. (195a/2001, rev. 2020); PMSL • 2 ♂♂, 4 ♀♀; Mt. Veliki Snežnik; 20 Jul. 1999; L. Slana Novak and T. Novak leg. (185a/1999, rev. 2020); PMSL • 1 ♂; ibid.; 12 Sep. 2018; L. Slana Novak and T. Novak leg. (116/2018); PMSL. – VL36 • 1 ♂, 1 ♀; Palčje − Javorniki; L. Slana Novak and T. Novak leg. (361/2011, rev. 2015); PMSL. – VL66 • 1 ♀; Benete, Runarsko; 31 May 2004; L. Slana Novak and T. Novak leg. (33/2004, rev. 2015); PMSL. – VL67 • 5 ♂♂, 3 ♀♀; Nova vas – Volčje; 26 Sep. 2006; L. Slana Novak and T. Novak leg. (25/2006, rev. 2019); PMSL • 1 ♂; Runarsko; 26 Sep. 2006; L. Slana Novak and T. Novak leg. (38/2006, rev. 2015); PMSL • 1 ♂, 1 ♀; Veliki vrh, Runarsko; 26 Sep. 2006; L. Slana Novak and T. Novak leg. (36/2006, rev. 2015); PMSL • 7 ♂♂, 5 ♀♀; Volčje; 26 Sep. 2006; L. Slana Novak and T. Novak leg. (28/2006, rev. 2015); PMSL. – VL69 • 1 ♂; Veliki vrh pri Šmarju; 10 May 2008; L. Slana Novak and T. Novak leg. (156/2011); PMSL. – VL74 • 1 ♂, 1 ♀; Mt. Goteniški Snežnik; 28 May 2000; L. Slana Novak and T. Novak leg. (247/2000, rev. 2007); PMSL • 1 ♀; Strma reber, Zgornji Čačiči; 27 Sep. 1996; L. Slana Novak and T. Novak leg. (LSN 126/1996, TN rev. 2008); PMSL. – VL75 • 2 ♂♂; Draga; 17 Aug. 2011; L. Slana Novak and T. Novak leg. (94/2011); PMSL • 1 ♂, 2 ♀♀; Mt. Goteniški Snežnik; 28 May 2000; L. Slana Novak and T. Novak leg. (253/2000, rev. 2007); PMSL. – VL76 • 1 ♂; Bukovica, Ribnica; Jul.1994; B. Oven leg. (LSN 25/1996, TN rev. 2009); PMSL • 2 ♂♂, 1 ♀; Lipovšica, Sodražica; 22 Aug. 2011; L. Slana Novak and T. Novak leg. (100/2011, rev. 2018); PMSL • 1 ♂; Nova Štifta; 22 Aug. 2011; L. Slana Novak and T. Novak leg. (105/2011); PMSL • 2 ♂♂, 3 ♀♀; Ribnica; 6 Sep. 2000; M. Mlakar leg. (828/2000, rev. 2008); PMSL • 1 ♂, 2 ♀♀; ibid.; (831/2000, rev. 2019); PMSL • 1 ♀; Mt. Turn; 27 Jun. 1980; B. Drovenik leg. (383/2002, rev. 2018); PMSL • 1 ♂; ibid.; 28 Aug. 1980; B. Drovenik leg. (394/2002, rev. 2009); PMSL. – VL77 • 1 ♂; Dolnje Retje; 3 May 2008; L. Slana Novak and T. Novak leg. (161/2011); PMSL • 1 ♀; Rašica, Velike Lašče; 16 Jun. 1986 (261/2002, rev. 2015); PMSL • 1 ♂, 1 ♀; Velike Poljane, Ortnek; 3 May 2008; L. Slana Novak and T. Novak leg. (142/2011, rev. 2019); PMSL. – VL78 • 1 ♂; Laporje; 3 May 2008; L. Slana Novak and T. Novak leg. (151/2011); PMSL • 1 ♀; Zagradec pri Grosupljem; 7 Jul. 1986; L. Slana Novak and T. Novak leg. (500/2002, rev. 2015); PMSL. – VL79 • 1 ♀; Veliki Lipoglav − Dobrinje; 8 Jul. 1986; L. Slana Novak and T. Novak leg. (467/2002, rev. 2015); PMSL. – VL83 • 2 ♂♂; Vršič, Banjaloka; 24 Apr. 2001; S. Brelih leg. (121/2001, rev. 2019); PMSL. – VL84 • 1 ♂; Jelenja vas; 3 May 2008; L. Slana Novak and T. Novak leg. (137a/2011); PMSL • 1 ♂; ibid.; (153/2011); PMSL • 1 ♂; Pleš, Borovec pri Kočevski Reki; 19 Sep. 2013; P. Kozel and T. Novak leg. (71/2013, rev. 2018); PMSL. – VL85 • 1 ♀; Grčarice; 18 Oct. 2013; T. Novak leg. (63/2013, rev. 2015); PMSL • 1 ♂, 2 ♀♀; Grčarske Ravne; 18 Oct. 2013; T. Novak leg. (66/2013, TN rev. 2019); PMSL • 3 ♀♀; Kočevje; 6 Sep. 2000; M. Mlakar leg. (822/2000, rev. 2008); PMSL • 1 ♂, 1 ♀; ibid.; (825/2000, rev. 2007); PMSL • 2 ♂♂, 1 ♀; Mt. Stojna, Kočevje; 25 Oct. 1996; S. Brelih leg. (92/2001, rev. 2008); PMSL. – VL86 • 1 ♀; Dolenja vas; 3 May 2008; L. Slana Novak and T. Novak leg. (110/2011); PMSL • 1 ♂, 1 ♀; ibid.; (154/2012, rev. 2018); PMSL. – VL87 • 1 ♂; Breg pri Zagradcu; 4 Nov. 1993; S. Brelih leg. (805/1998, rev. 2007); PMSL. – VL89 • 1 ♀; Potok, Stična; 4 Mar. 1994; S. Brelih leg. (658/1998, rev. 2015); PMSL. – VL93 • 1 ♂; Brsnik, Fara; 3 May 2008; L. Slana Novak and T. Novak leg. (157/2011, rev. 2018); PMSL. – VL94 • 1 ♂, 1 ♀; Dolenja Žaga; 29 Apr. 2001; S. Brelih leg. (131/2001, rev. 2015); PMSL • 1 ♀; Dolenja Žaga – Podstene; 26 Jun. 1999; L. Slana Novak and T. Novak leg. (90/1999, rev. 2008); PMSL • 1 ♂, 1 ♀; Gorenja Žaga; 3 May 2008; L. Slana Novak and T. Novak leg. (111a/2011); PMSL • 1 ♂, 1 ♀; ibid.; (159/2011); PMSL • 1 ♀; Ograja; 3 May 2008; L. Slana Novak and T. Novak leg. (109a/2011); PMSL • 1 ♀; ibid.; (121a/2011); PMSL • 1 ♀; ibid.; (144/2011); PMSL • 1 ♀; ibid.; (148/2011); PMSL • 1 ♂, 1 ♀; Ograja – Suhor − Morava; 3 May 2008; L. Slana Novak and T. Novak leg. (120a/2011); PMSL • 1 ♂, 1 ♀; ibid.; (138a/2011); PMSL • 1 ♂, 1 ♀; ibid.; (152/2011); PMSL • 1 ♂; Ograja – Zdihovo; 26 Jun. 1999; L. Slana Novak and T. Novak leg. (76/1999, rev. 2008); PMSL • 3 juvs; ibid.; 12 Jun. 1999; L. Slana Novak and T. Novak leg. (32/1999, rev. 2008); PMSL • 2 juvs; Podstene pri Kostelu; 12 Jun. 1999; L. Slana Novak and T. Novak leg. (24/1999, rev. 2008); PMSL • 1 ♂; Rajndol; 21 Mar. 1994; S. Brelih leg. (821/1998, rev. 2008); PMSL • 5 ♂♂, 3 ♀♀; Srednji Potok, Banja Loka; 8 Oct. 2012; P. Kozel and T. Novak leg. (68/2015); PMSL • 1 ♂, 1 ♀, 2 juvs; Štalcerski preval; 12 Jun. 1999; L. Slana Novak and T. Novak leg. (33/1999, rev. 2008); PMSL. – VL95 • 1 ♂, 2 ♀♀; Dolga vas; 17 Aug. 1985; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 97/1986, TN rev. 2007); PMSL • 1 ♂, 1 ♀; Livold; 3 May 2008; L. Slana Novak and T. Novak leg. (123a/2011); PMSL • 1 ♂, 1 ♀; ibid.; (158/2011); PMSL. – VL97 • 1 ♀; Dvor, Lašče; 10 May 2008; L. Slana Novak and T. Novak leg. (112/2011); PMSL • 1 ♂, 2 ♀♀; Železno, Dobrnič; 4 Nov. 1993; S. Brelih leg. (812/1998, rev. 2007); PMSL. – VL99 • 3 ♀♀; Nova gora, Čatež; 15 Aug. 2007; L. Slana Novak and T. Novak leg. (61/2007, rev. 2015); PMSL. – VM80 • 1 ♂; Litija; 10 Sep. 2000; M. Mlakar leg. (809/2000, rev. 2008); PMSL • 1 ♂, 1 ♀; ibid.; (834/2000, rev. 2015); PMSL. – WL03 • 2 ♂♂; Dolenje Kozice; 18 Apr.−13 May 2002; B. Drovenik and A. Gergeli leg. (509/2018); PMSL • 1 ♂; ibid.; 18 Apr.−13 May 2002; B. Drovenik and A. Gergeli leg. (550/2018); PMSL • 1 ♂; ibid.; 5 Jul. 2002; B. Drovenik and A. Pirnat leg. (1135/2018); PMSL • 1 ♀; ibid.; 6 Aug. 2002; B. Drovenik and A. Pirnat leg. (1090/2018); PMSL • 1 ♀; ibid.; 30 Sep. –28 Oct. 2002; B. Drovenik and A. Gergeli leg. (PK 64/2012); PMSL • 1 ♀; ibid.; 25 Nov. 2002; B. Drovenik and A. Pirnat leg. (270/2018); PMSL • 2 ♀♀; ibid.; 25 Nov. −23 Dec. 2002; B. Drovenik and A. Gergeli leg. (1027/2018); PMSL • 4 ♂♂, 5 ♀♀; Srednji Radenci; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 51/1986, rev. 2008); PMSL • 3 ♂♂, 4 ♀♀; Stari trg ob Kolpi; 21 Mar. 1994; S. Brelih leg. (851/1998, rev. 2008); PMSL • 5 ♂♂, 5 ♀♀; Dolenji Radenci – Breg pri Sinjem vrhu; 3 May 2008; L. Slana Novak and T. Novak leg. (135a/2011); PMSL • 5 ♂♂, 2 ♀♀; ibid.; (149/2011); PMSL. – WL05 • 1 ♂, 3 ♀♀, 1 juv.; Črmošnjice; 16 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (160/1997); PMSL • 1 ♀; Koprivnik, Kočevski Rog; 21 Mar. 1994; S. Brelih leg. (867/1998, rev. 2008); PMSL • 1 ♂, 2 ♀♀; Mt. Mirna gora; 8 Oct. 2012; P. Kozel and T. Novak leg. (59/2015); PMSL • 4 ♂♂, 4 ♀♀; ibid.; 8 Oct. 2012; P. Kozel and T. Novak leg. (66/2015); PMSL. – WL06 • 1 ♂; Cerovec − Selišče, Podturn pri Dolenjskih toplicah; 10 May 2008; L. Slana Novak and T. Novak leg. (145/2011, rev. 2017); PMSL. – WL07 • 1 ♂; Boršt pri Dvoru; 10 May 2008; L. Slana Novak and T. Novak leg. (113/2012); PMSL • 1 ♀; Prečna, Mirna; Jun. 2000; V. Jaćimović leg. (PK and TN 100/2020); PMSL. – WL08 • 1 ♂; Rdeči kal, Dobrnič; 4 Nov. 1993; S. Brelih leg. (872/1998, rev. 2007); PMSL • 1 ♂, 1 ♀; Trebnje; 29 Aug. 2000; M. Mlakar leg. (787/2000, rev. 2018); PMSL • 1 ♂; ibid.; (787a/2000, rev. 2018); PMSL • 1 ♀; ibid.; (788/2000, rev. 2008); PMSL. – WL13 • 4 ♀♀; Balkovci, Vinica; 15 Aug. 1999; M. Štangelj and T. Novak leg. (212/1999, rev. 2008); PMSL • 2 ♂♂, 2 ♀♀; Dolenji Radenci – Breg pri Sinjem vrhu; 3 May 2008; L. Slana Novak and T. Novak leg. (146/2011); PMSL • 4 ♂♂, 1 ♀; Kot pri Dramlju; 16 Aug. 1999; M. Štangelj and T. Novak leg. (207/1999, rev. 2015); PMSL • 3 ♂♂, 2 ♀♀; Špeharji, Vinica; 16 Aug. 1999; M. Štangelj and T. Novak leg. (201/1999, rev. 2006); PMSL • 6 ♂♂, 5 ♀♀; Vinica – Sinji vrh; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 59/1986, TN rev. 2007); PMSL • 2 ♂♂; Vukovci; 18 Apr. 1996; S. Brelih leg. (764/1998, rev. 2007); PMSL. – WL14 • 1 ♂, 1 ♀; Bistrica, Črnomelj; 26 Jun. 1999; L. Slana Novak and T. Novak leg. (65/1999, rev. 2017); PMSL • 1 ♀; Črnomelj; 15 Aug. 1999; M. Štangelj and T. Novak leg. (221/1999, rev. 2008); PMSL • 3 ♂♂, 3 ♀♀, 2 juvs; Desinec – Črnomelj; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (385/1998, rev. 2007); PMSL • 2 ♂♂, 2 ♀♀; Tanča gora; 21 Mar. 1994; S. Brelih leg. (861/1998, rev. 2015); PMSL • 5 ♂♂, 2 ♀♀; Sinji vrh, Vinica; 16 Aug. 1999; M. Štangelj and T. Novak leg. (205/1999, rev. 2007); PMSL. – WL15 • 1 ♀; Krupa spring; 10 Mar. 1

Nemastoma bidentatum schmidti ssp. nov. × N. bidentatum sparsum Gruber & Martens, 1968 Figs 3, 9P, 10O Nemastoma triste – Roewer 1917: 149: [partim: Celje / Cilli, Maribor / Marburg]. Nemastoma bidentatum bidentatum × N. bidentatum sparsum – Gruber & Martens 1968: 142, 147, figs 12–13 [partim: Novo mesto; Županja]. — Martens 1978: 107, fig 146 [partim: Županja; Novo mesto, Mt. Snežnik]. — Novak & Gruber 2000: 286 [partim: Novo mesto]. — Novak et al. 2002: 136 [partim: Ptuj]. — Novak 2004b: 246 [partim: Županja]. Nemastoma bidentatum ssp. – Novak & Gruber 2000: 286 [partim: Celje, Maribor]. Diagnosis Hybrids exhibiting varying Ch-Apo morphology, but with knee-like flexion of anterior margin, and either being relatively tiny with more or less elongated superior portion or more massive with superior portion drawn out in tiny apical portion, and club-shaped Pa-Fe with intermediate form with respect to both subspecies. Material examined SLOVENIA – VM91 • 2 ♂♂, 4 ♀♀; Čemšemik; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1183/2002, rev. 2018); PMSL • 1 ♂, 1 ♀; Jesenovo – Znojile; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1187/2002, TN rev. 2018); PMSL • 6 ♂♂, 5 ♀♀; Mt. Čemšeniška planina; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1191/2002, rev. 2018); PMSL • 2 ♀♀; Zaplanina, Ločica pri Vranskem; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1210/2002, rev. 2018); PMSL • 1 ♂, 5 ♀♀; ibid.; (1179/2002, rev. 2018); PMSL. – VM92 • 4 ♂♂, 9 ♀♀; Ločica pri Vranskem; 10 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 101 /1986, TN rev. 2018); PMSL. – WM01 • 7 ♂♂, 5 ♀♀; Marija Reka; 21 Jul. 2002; L. Slana Novak and T. Novak leg. (1186/2002, rev. 2018); PMSL. – WM05 • 4 ♂♂, 7 ♀♀; Slovenj Gradec; 9 Oct. 1983; T. Novak and L. Slana Novak leg. (LSN 113 /1983, TN rev. 2018); 3 ♂♂, 4 ♀♀; ibid.; 6 Jul. 2003; L. Slana Novak and T. Novak leg. (190/2003, rev. 2018); PMSL. – WM06 • 1 ♂, 2 ♀♀; Dravograd; 25 Oct. 1995; S. Hudrap and M. Pavlin leg. (11/1996, rev. 2018); PMSL. – WM10 • 4 ♂♂, 1 ♀; Lože – Globoko; 14 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 45 /1983, TN rev. 2018); PMSL. – WM11 • 1 ♂, 2 ♀♀; Brdce; 22 Apr. 1995; S. Brelih leg. (758/1998, rev. 2018); PMSL • 1 ♂, 3 ♀♀; Udmat; 15 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 47 /1983, TN rev. 2018); PMSL. – WM12 • 7 ♂♂, 3 ♀♀; Arja vas; 10 Jul. 1986; L. Slana Novak and T. Novak leg. (513/2002, rev. 2018); PMSL • 1 ♂, 1 ♀; Velika Pirešica; 10 Jul. 1986; L. Slana Novak and T. Novak leg. (452/2002, rev. 2018); PMSL • 1 ♂, 4 juvs; ibid.; (470/2002, rev. 2018); PMSL. – WM13 • 4 ♂♂, 3 ♀♀; Vodosteč, Črnova; 10 Jul. 1986; L. Slana Novak and T. Novak leg. (453/2002, rev. 2018); PMSL • 2 ♂♂; ibid.; 22 Apr. 1995; S. Brelih leg. (772/1998, rev. 2018); PMSL • 1 ♂, 1 ♀; ibid.; 21 Apr. 2002; L. Slana Novak and T. Novak leg. (737/2002, rev. 2019); PMSL. – WM16 • 2 ♂♂, 3 ♀♀; Sv. Trije Kralji, Muta; 20 Aug. 2005; L. Slana Novak and T. Novak leg. (79/2005, rev. 20018); PMSL. – WM20 • 2 ♂♂, 3 ♀♀; Jurklošter; 15 Aug. 1983; L. Slana Novak and T. Novak leg. (LSN 53 /1983, TN rev. 2018); PMSL. – WM21 • 1 ♂, 2 ♀♀; Lahovna; 22 Apr. 1995; S. Brelih leg. (740/1998, TN rev. 2018); PMSL • 2 ♂♂, 3 ♀♀, 1 juv.; Trobni dol; 16 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 48 /1983, TN rev. 2018); PMSL. – WM22 • Celje • 1 ♂; ibid.; 30 Oct. 2000; N. Žmaher leg. (881/2000, rev. 2018); PMSL • 7 ♂♂, 2 ♀♀; Šmartinsko jezero; 25 Mar. 1995; S. Brelih leg. (704/1998, rev. 2018); PMSL • 1 ♂, 4 ♀♀; Vojnik; 17 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 58 /1983, TN rev. 2018); PMSL. – WM23 • 1 ♂, 1 ♀; Črešnjice; 25 Mar. 1995; S. Brelih leg. (709/1998, rev. 2018); PMSL • 3 ♂♂, 3 ♀♀, l juv.; Frankolovo; 17 Aug. 1983; L. Slana Novak and T. Novak leg. (LSN 76 /1983, TN rev. 2018); PMSL • 1 ♂; Mt. Konjiška gora; 14 Jun. 1981; V. Kuštor, T. Novak leg. (1120/1981, rev. 2018); PMSL • 1 ♂, 2 ♀♀; Mt. Stenica; 30 May 1981; T. Novak, V. Kuštor leg. (228/1981, rev. 2018); PMSL • 1 ♀; Stenica; 24 Jul. 2002; B. Drovenik and A. Gergeli leg. (1138/2018); PMSL • 1 ♀; ibid.; 24 Jul.–21 Aug. 2002; B. Drovenik and A. Gergeli leg. (PK 73 /2012, TN rev. 2018); PMSL • 1 ♀; ibid.; (PK 12 /2013, TN rev. 2018); PMSL • 1 ♂; ibid.; 21 Aug. 2002; B. Drovenik and A. Gergeli leg. (1071/2018); PMSL • 1 ♀; ibid.; 21 Aug.−16 Sep. 2002; B. Drovenik and A. Gergeli leg. (782/2018); PMSL • 1 ♂, 1 ♀; ibid.; (819/2018); PMSL • 1 ♀; ibid.; (856/2018); PMSL • 1 ♂, 1 ♀; ibid.; (804/2018); PMSL • 1 ♀; ibid.; 16 Sep.−14 Oct. 2002; B. Drovenik and A. Gergeli leg. (817/2018); PMSL • 1 ♀; ibid.; 14 Oct. 2002; B. Drovenik and A. Gergeli leg. (361/ 2018); PMSL • 1 ♀; ibid.; (783/2018); PMSL • 1 ♂, 1 ♀; ibid.; (794/2018); PMSL • 1 ♂, 1 ♀; ibid.; (814/2018); PMSL • 1 ♀; ibid.; (824/2018); PMSL • 1 ♀; ibid.; (831/2018); PMSL • 1 ♀; ibid.; (846/2018); PMSL • 1 ♂; ibid.; 11 Nov. 2002; B. Drovenik and A. Gergeli leg. (208/2018); PMSL • 1 juv.; ibid.; (266/2018); PMSL • 1 ♂; ibid.; 11 Nov. 2002 − 4 Mar. 2003; B. Drovenik and A. Gergeli leg. (694/2018); PMSL • 1 ♀; ibid.; (698/2018); PMSL • 2 ♀♀; ibid.; (701/2018); PMSL • 1 ♂, 1 ♀; ibid.; (710/2018); PMSL • 1 ♀; ibid.; (716/2018); PMSL • 1 ♀; ibid.; (721/2018); PMSL • 1 ♂; ibid.; (724/2018); PMSL • 3 ♂♂, 2 ♀♀; ibid.; (729/2018); PMSL • 1 ♂; ibid.; 8 Apr. 2003; B. Drovenik and A. Gergeli leg. (267/ 2018); PMSL • 3 juvs; ibid.; (271/2018); PMSL • 1 ♂; ibid.; 8 Apr. 2003; B. Drovenik and A. Gergeli leg. (189/ 2018); PMSL • 1 juv.; ibid.; (266/2018); PMSL. – WM24 • 1 ♂, 1 ♀; Jurgovo; 24 Jul.–3 Aug. 2005; A. Kapla leg. (178/2005, rev. 2018); PMSL • 3 ♂♂; Resnik; 12 Aug. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 252 /1984, TN rev. 2018); PMSL. – WM25 • 3 ♂♂; Glažuta, Mislinjski jarek; 9 Sep. 2017; M. Delakorda, S. Lipovšek, L. Slana Novak and T. Novak leg. (107/2018); PMSL • 1 ♀; ibid.; 19 Sep. 2018; S. Lipovšek, P. Kozel and T. Novak leg. (132/2018); PMSL • 1 ♂; Ribnica na Pohorju; 4 Jul. 1939; J. Hadži leg. (145/1984, rev. 2018); PMSL. – WM30 • 1 ♂, 1 ♀; Loke – Lažiše; 11 Jul. 2003; T. Novak leg. (154/2003, rev. 2018); PMSL. – WM31 • 3 ♂♂, 2 ♀♀; Bezovje pri Šentjurju; 16 Aug. 1983; L. Slana Novak and T. Novak leg. (LSN 56 /1983, TN rev. 2018); PMSL • 1 ♂; Slivniško jezero; 7 Apr. 1995; S. Brelih leg. (701/1998, rev. 2018); PMSL. – WM33 • 3 ♂♂, 2 ♀♀, 1 juv.; Tepanje; 17 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 62 /1983, TN rev. 2018); PMSL • 2 ♂♂; Žiče monastery; 25 Mar. 1995; S. Brelih leg. (706/1998, rev. 2018); PMSL. – WM40 • 1 ♂; Lesično; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1130/2002, rev. 2018); PMSL. – WM41 • 1 ♂, 5 ♀♀; Podčetrtek; 13 Aug. 1983; L. Slana Novak and T. Novak leg. (LSN 51 /1983, TN rev. 2018); PMSL • 2 ♂♂, 3 ♀♀; ibid.; 7 Jul. 2012; T. Novak leg. (51/2012, rev. 2018); PMSL • 2 ♂♂, 3 ♀♀; Virštajn; 7 Apr. 1995; S. Brelih leg. (699/1998, rev. 2018); PMSL. – WM42 • 1 ♀; Mt. Boč; 17 Apr. 1982; T. Novak and G. Kolar leg. (125/1982, rev. 2018); PMSL • 2 ♂♂, 1 ♀; ibid.; 16 Oct. 1983; L. Slana Novak and T. Novak leg. (LSN 118 /1983, TN rev. 2018); PMSL • 2 ♂♂; Mestinje; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1165/2002, rev. 2018); PMSL • 1 ♂, 1 ♀; ibid.; (1168/2002, rev. 2018); PMSL • 1 ♂, 1 ♀; Zgornji Gabrnik; 11 Oct. 1997; L. Slana Novak and T. Novak leg. (412/1998, rev. 2018); PMSL. – WM43 • 4 ♂♂, 1 ♀; Ložnica; 17 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 75 /1983, TN rev. 2018); PMSL. – WM44 • 2 ♂♂, 4 ♀♀; Kočno pri Polskavi; 8 Aug. 2015; L. Slana Novak and T. Novak leg. (110/2018); PMSL. – WM45 • 1 ♂; Limbuš; 11 Oct. 2001; D. Kos leg. (226/2004, rev. 2018); PMSL. – WM52 • 1 ♀; Sv. Donat, Donačka gora; 16 Sep. 2002; B. Drovenik and A. Pirnat leg. (247/2018); PMSL • 1 ♀; ibid.; (248/2018); PMSL • 1 ♀; ibid.; 14 Oct. 2002; B. Drovenik and A. Pirnat leg. (1032/2018); PMSL • 6 ♂♂, 2 ♀♀, 2 juvs; Tuncovec; 4 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 66 /1983, TN rev. 20018); PMSL. – WM53 • 4 ♂♂, 1 ♀; Majšperk; 24 Jun. 2007; T. Novak leg. (43/2018); PMSL • 4 ♂♂, 1 ♀; Pečke; 10 Apr. 1995; S. Brelih leg. (713/1998, rev. 2018); PMSL • 1 ♂; Stogovci; 24 Jun. 2007; T. Novak leg. (50/2007, rev. 2018); PMSL. – WM62 • 1 ♂, 2 ♀♀; Dobovec pri Rogatcu; 7 Oct. 2001; T. Novak leg. (605/2002, rev. 2007); PMSL • 2 ♂♂; ibid.; 16 Aug. 2003; J. Ravljen and T. Novak leg. (807/2003, rev. 2018); PMSL • 4 ♂♂; ibid.; 16 Aug. 2003; J. Ravljen, L. Slana Novak and T. Novak leg. (812/2003, rev. 2007); PMSL • 1 ♂, 2 ♀♀; ibid.; 30 Sep. 2012; T. Novak leg. (63/2015); PMSL • 7 ♂♂, 4 ♀♀, 1 juv.; Kozminci; 6 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 61 /1983, TN rev. 2018); PMSL • 1 ♀; Mt. Macelj; Jun. 1980; V. Kuštor and T. Novak leg. (LSN 5 /1983, TN rev. 2018); PMSL • 2 ♀♀; Žale, Dobrina; 24 Apr. 1998; S. Brelih leg. (85/2001, rev. 2018); PMSL. – WM63 • 4 ♂♂, 1 ♀; Majski vrh − Tržec; 6 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 65 /1983, TN rev. 2018); PMSL. – WM64 • 3 ♂♂, 3 ♀♀; Ptuj; 6 Aug. 1983, T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 77 /1983, TN rev. 2018); PMSL. – WM65 • 2 ♂♂; Selca; 14 Jun. 1984; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 238 /1984, TN rev. 2018); PMSL • 1 ♂, 2 ♀♀; Zgornja Voličina; 23 Apr. 1998; S. Brelih leg. (124/2001, rev. 2018); PMSL. – WM73 • 1 ♂, 4 ♀♀; Berinjak; 24 Jun. 2007; T. Novak leg. (36/2007, rev. 2018); PMSL • 1 ♂, 8 ♀♀; Borl castle, Dolane; 7 Aug. 1983; L. Slana Novak, M. Štangelj and T. Novak leg. (LSN 59 /1983, TN rev. 2018); PMSL • 1 ♂, 2 ♀♀; Ljubslava, Podlehnik; 24 Jun. 2007; T. Novak leg. (49/2007, TN rev. 2018); PMSL • 5 ♂♂; Spodje Gruškovje, Podlehnik; 24 Jun. 2007; T. Novak leg. (13/2007, rev. 2018); PMSL • 4 ♂♂, 1 ♀; Vareja; 6 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 78 /1983, TN rev. 2018); PMSL. – WM76 • 4 ♂♂, 3 ♀♀; Brengova; 26 Jul. 1983; T. Novak and L. Slana Novak leg. (LSN 63 /1983, TN rev. 2018); PMSL • 2 ♂♂, 1 ♀; Kočki vrh; 30 Jun. 2002; T. Novak leg. (783/2002, rev. 2018); PMSL • 1 ♂, 2 ♀♀; ibid.; (789/2002, rev. 2018); PMSL. – WM83 • 3 ♂♂, 5 ♀♀; Korenjak – Turški vrh; 7 Aug. 1983; T. Novak, M. Štangelj and L. Slana Novak leg. (LSN 55 /1983, TN rev. 2018); PMSL. – WM84 • 4 ♂♂, 3 ♀♀; Vudina, Velika Nedelja; 29 May 2001; P. Igerc and T. Novak leg. (22/2001, rev. 2018); PMSL. – WM85 • 2 ♀♀; Radoslavci; 10 Sep. 1983; T. Novak, M. Slana Novak and L. Slana Novak leg. (LSN 117 /1983, TN rev. 2018); PMSL. – WM94 • 2 ♂♂, 3 ♀♀; Pavlovci; 2 Apr. 1983; T. Novak and L. Slana Novak leg. (LSN 6 /1983, TN rev. 2018); PMSL. – WM95 • Noršinci pri Ljutomeru; 27 Aug. 1983; T. Novak and L. Slana Novak leg. 2 ♂♂, 2 ♀♀ (LSN 109 /1983, TN rev. 2018); PMSL. – WM95 • 3 ♂♂, 3 ♀♀; Gresovščak; L. Slana Novak and T. Novak leg. (163/2012, rev. 2018); PMSL • 2 ♂♂, 2 ♀♀; Slamnjak, Ljutomer; 10 Jun. 1984; L. Slana Novak and T. Novak leg. (LSN 241 /1984, rev. 2018); PMSL. Remarks To date, N. b. schmidti ssp. nov. has been reported under N. b. bidentatum, and N. b. gruberi ssp. nov. under N. b. sparsum. Since hybrids N. b. bidentatum × N. b. sparsum have not been identified in any locality, all these published data likely refer either to N. b. schmidti ssp. nov. × N. b. sparsum or N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. Nemastoma b. bidentatum referred from Germany (Schönhofer & Holle 2007) is a hybrid, probably N. b. schmidti ssp. nov. × N. b. sparsum. Since both N. b. sparsum and N. b. gruberi ssp. nov. inhabit the lowland in Croatia, hybrids south of the Sava River or nearby probably belong to N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov.

Nemastoma bidentatum bidentatum Roewer, 1914 �� N. bidentatum schmidti ssp. nov. Figs 3, 9J, 10J. Diagnosis Typically, males with narrow quarter moon-like, apically bifid Ch-Apo, ~2.4 times as high as wide, with wide, inconspicuous apical pinnacle posteriorly, and Pa-Fe ~ intermediate between both subspecies. Material examined SLOVENIA ��� VM44 ��� 1 ♂, 1 ♀; Gromov rob, Jelendol, Tržič; 23 Aug. 2017; T. Novak leg. (GR 5339, GR 5338); PMSL. ��� WM06 ��� 1 ♂, 1 ♀; Mt. Ko��enjak; 27 Sep. 1981; T. Novak leg. (520/1981, rev. 2018); PMSL ��� 4 ♂♂, 2 ♀♀; ibid.; 14 Aug. 2012; T. Novak leg. (GR 3784, GR 3799, GR 3800, GR 3826, GR 3830, GR 3832); PMSL ��� 2 ♀♀; ibid.; 24 Aug. 2012; T. Novak leg. (153/2012, rev. 2018); PMSL ��� 1 ♂; Sv. Simon, Pernice; 18 Aug. 2005; T. Novak leg. (66b/2005, rev. 2018); PMSL ��� 7 ♂♂, 5 ♀♀; Sv. Urban, Kozji vrh nad Dravogradom; 24 Sep. 2005; L. Slana Novak and T. Novak leg. (127/2005, rev. 2019); PMSL ��� 3 ♂♂, 7 ♀♀; ibid.; 9 Aug. 2017; S. Lipov��ek and T. Novak leg. (GR M1021, GR M1024, GR M1025, GR M1027, GR M1033, GR M1035 ��� GR M1039); PMSL., Published as part of Novak, Tone, Novak, Ljuba Slana, Kozel, Peter, Schaider, Miriam Gudrun, Komposch, Christian, Lipov��ek, Sa��ka, Podlesnik, Jan, Pau��i��, Igor & Raspotnig, G��nther, 2021, Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence, pp. 1-67 in European Journal of Taxonomy 777 on page 42, DOI: 10.5852/ejt.2021.777.1561, http://zenodo.org/record/5637112, {"references":["Roewer C. F. 1914. Die Familien der Ischyropsalidae und Nemastomatidae der Opiliones-Palpatores. Archives fur Naturgeschichte 80 A (3): 99 - 169."]}

While the chemistries of scent gland secretions from a few selected species of three families of Cyphophthalmi, namely Sironidae, Pettalidae, and Stylocellidae, have already been reported and found to consist of complex blends of naphthoquinones and methyl ketones, nothing is known about the other families. We here report on the secretions of Metasiro savannahensis Clouse and Wheeler (Zootaxa 3814:177–201, 2014), a first representative of the family Neogoveidae. The secretions from males, females and one juvenile were extracted and analyzed by gas chromatography–mass spectrometry. Twenty-five compounds were identified, all of which belong to the chemical classes of naphthoquinones and methyl ketones, confirming a hypothesized chemical uniformity of cyphophthalmid exudates. One major naphthoquinone compound, however, was new for cyphophthalmids and for arthropod exocrine secretions in general: a methyljuglone isomer, 6-methyljuglone (= 6-MJ; iupac name: 5-hydroxy-6-methyl-1,4-naphthoquinone), amounted for about 20% of the secretion and was eventually identified by synthesis. Hydroxy-naphthoquinones and their derivatives are known to possess a variety of antibiotic effects, probably enhancing the antimicrobial/antifungal potential of the Metasiro-secretion. Currently, without further data on neogoveids, the compound represents a chemical autapomorphy of M. savannahensis, and—just as the strange chloro-naphthoquinones of Sironidae and Pettalidae—adds to the repertoire of unusual naphthoquinone compounds across the Cyphophthalmi. Electronic supplementary material The online version of this article (10.1007/s00049-019-00288-y) contains supplementary material, which is available to authorized users.

Interactive 3D content of Fig. 2. (PDF 2100 kb)

Additional information for sections Morphology and Chemistry as well as Tables S1 - S4. (PDF 239 kb)

By the possession of unique exocrine scent glands, Opiliones (harvestmen) arise as a perfect model for studies on the evolutionary history of secretion chemistry. Among gland compounds of harvestmen, it is the quinones that represent recurring elements across the secretions of all suborders. Reliable data on quinone-distribution, however, is only known for Laniatores (benzoquinones) and Cyphophthalmi (naphthoquinones). We here unraveled the quinone-distribution across scent gland secretions of the third large harvestman suborder, the Palpatores (= Eu- and Dyspnoi): Naphthoquinones were found in phalangiid Eupnoi across all subfamilies as well as in nemastomatid (and at least one ischyropsalid) Dyspnoi. Benzoquinones (1,4-benzoquinone) were restricted to a small entity within Eupnoi, namely platybunine Phalangiidae, probably misplaced Gyantinae (currently Sclerosomatidae) and Amilenus (incertae sedis). Our findings, combined with data from Laniatores and Cyphophthalmi, allow evaluation of a comprehensive chemosystematic model for Opiliones for the first time. Evolutionary scenarios imply naphthoquinones as scent gland compounds of common ancestry, having evolved in an early harvestman ancestor and present in cyphophthalmids and palpatoreans, but lost in laniatoreans. Benzoquinones evolved later and independently at least twice: once in the secretions of gonyleptoid Laniatores (alkylated benzoquinones), and a second time in a lineage of phalangiid Eupnoi (1,4-benzoquinone).

Summary. This review focuses on advances in opilionid scent gland research between the years 2005 and 2012, along with an evaluation of all currently available chemical information from different groups of Opiliones from a phylogenetic perspective. To date, examples of scent gland secretions from all four suborders of Opiliones have been published, including: 4 species of Cyphophthalmi, 12 species of Eupnoi, 1 species of Dyspnoi, 4 species of insidiatorean Laniatores, and 42 species of grassatorean Laniatores. In addition, preliminary data are given for another 20 species (mainly Insidiatores and Eupnoi). From all species, more than 70 scent gland-derived components from different chemical classes have so far been identified. Secretion profiles appear to be highly specific, and well-suited for both the characterization of phyletic groups as well as for the distinction of closely related species. Secretions may contain more than 20 components (e.g. Cyphopththalmi), or consist of only a few, or even a single compound (e.g. some Insidiatores and early-derivative Grassatores). With respect to gross systematics, the suborders of Opiliones can be chemically characterized as follows: naphthoquinones and acyclic compounds (mainly ketones) appear to be widespread in the secretions of a proposed clade of Cyphophthalmi + Palpatores (Eupnoi, Dyspnoi). For this clade, naphthoquinones are considered synapomorphic, since they have not been detected in the laniatores. Although the phyletic status of acyclic compounds in cyphophthalmid and palpatorean secretions remains unclear, these acycles may also have a common origin. A sub-class of these acycles, the so-called “sclerosomatid compounds”, may represent a chemical synapomorphy of the monophyletic Palpatores. Deep chemical divergence is evident in secretions from the Laniatores, along with a split of laniatorean secretions into at least two chemically distinct groups: travunioid Insidiatores mainly produce an array of nitrogen-containing compounds, whereas phenol- and benzoquinone-rich secretions are characteristic of grassatorean Laniatores. However, a link between the chemistry of laniatoreans and the secretions of Cyphophthalmi-Palpatores is still missing.

Der bodenbewohnende Milbenkanker Cyphophthalmus duricorius Joseph, 1868 (Fam. Sironidae) galt mehr als 60 Jahre lang als die einzige in Österreich vorkommende Art der Weberknecht-Unterordnung Cyphophthalmi. Neuere Aufsammlungen zeigen jedoch, dass mindestens zwei weitere Cyphophthalmi-Arten in Österreich existieren: 1) Siro cf. crassus Novak & Giribet, 2006 wurde an einer Lokalität in der SW Steiermark nahe der slowenischen Grenze gefunden und repräsentiert eine zweite Gattung von Sironiden in Österreich. 2) Ein weiterer, morphologisch distinkter Sironide („Sironidae gen. et sp. nov.?“) − derzeit noch unbeschrieben und taxonomisch nicht zugeordnet − stammt aus Böden im Grenzgebiet Steiermark-Kärnten. Obwohl bislang kein syntopes Vorkommen belegt ist, können alle drei Arten in einem kleinen Areal von wenigen Quadratkilometern gefunden werden. For the last 60 years, the mite-harvestman Cyphophthalmus duricorius Joseph, 1868, a soil-dwelling sironid, has been considered to be the only representative of the opilionid suborder Cyphophthalmi in Austria. However, novel data from recent collections confirm the presence of at least two further Austrian cyphophthalmid species. (1) Siro cf. crassus Novak & Giribet, 2006 occurs in at least one location in SW Styria near the Slovenian border and hence represents a member of a second genus of Austrian sironids. (2) A further morphologically distinct sironid (“Sironidae gen. et sp. nov.?”) – so far undescribed and systematically not placed in detail – was collected in the borderland between Styria and Carinthia. All three species can be found in a small area of a few square-kilometers; although no syntopic occurrence was recorded.

Comparison of Trhypochthonius tectorum populations The biometrical data numbers of genital setae, notogastral setal lengths (%) and notogaster lengths (µm) were measured from each specimen of six Trhypochthonius tectorum populations (3: n=5), (6: n=4), (7: n=5), (8: n=5), (9: n=5) and (10: n=3), as listed in the section “Material examined”. The number of genital setae pairs varies within the populations and obviously also among the populations of T. tectorum (Fig. 10b). The total observed range of setae pairs is 7–11; the median value in populations (9) and (10) is nine, in (3) is ten, in (6), (7) and (8) is eleven. There were no significant differences of notogaster lengths among the tested six populations (Fig. 10a). Only in three of fifteen notogastral setae sets were significant differences found: regarding setae d 2, population (3) is significantly different from all others which are otherwise not different from each other. The significant differences regarding d 1 and p 1 are indicated in figures 11c and 11d: the relative lengths of seta d 1 are partly different between populations and show moderate variability within populations. Population (10) is represented only by three specimens that probably are not fully representative. As two examples, figures 11a and 11b show the non-significant variability between the populations for setae c 2 and c 3; the means are partly different, but the ranges overlap. Remarks. The observed but limited biometrical differences among populations of Trhypochthonius tectorum are not surprising, considering that the species is strictly parthenogenetic; populations consist of clone lines without genetic interchange, which would balance variability within populations as occurs in bisexual species. Similar differences between populations are observed in other biometrically analysed parthenogenetic species, e.g. in Tectocepheus and Trhypochthoniellus species (Weigmann 1999, 2002)., Published as part of Weigmann, Gerd & Raspotnig, Günther, 2009, Comparative morphological and biometrical studies on Trhypochthonius species of the tectorum species group (Acari: Oribatida: Trhypochthoniidae), pp. 1-31 in Zootaxa 2269 (1) on page 20, DOI: 10.11646/zootaxa.2269.1.1, http://zenodo.org/record/5306773, {"references":["Weigmann, G. (1999) Morphological variability in populations of a thelytokous mite, Trhypochthoniellus longisetus (Oribatida), with notes on synonymy. In: Bruin, J., van der Geest, L. P. S. & Sabelis, M. W. (eds), Ecology and evolution of the Acari. Kluwer Academic Publisher, Dordrecht, pp. 581 - 586.","Weigmann, G. (2002) Morphological variability between and within populations of Tectocepheus (Acari, Oribatida, Tectocepheidae) from the velatus-complex in central Europe. In: Bernini, F., Nannelli, R., Nuzzaci, G. & de Lillo, E. (eds), Acarid phylogeny and evolution: Adaptation in mites and ticks. Proceedings of the IV Symposium of the European Association of Acarologists. Kluwer Academic Publisher, Dordrecht, pp. 141 - 152."]}

Trhypochthonius japonicus n. forma occidentalis (Figs 8, 4e, 5e, 9b) Differential diagnosis. Body length about 680–750 µm. Shape of notogastral setae as in nominate form; setae c 1 and c 2 (range 16-21 µm) distinctly shorter than c 3 and d 3 (52-70 µm), d 1 and smaller d 2 (18-26 µm), e 1 of medium length (41-54 µm); posterior notogastral setae longer, p 1 the longest (92-108 µm), spinose and acuminate; p 2 remarkably long (about 82 µm), acuminate and sparsely spinose; h 3 and p 3 smooth setiform (17- 38 µm), less than half the length of p 2. 12–14 pairs of genital setae, mostly 13. General characters. Mean body length 717 µm, measured range 682–746 µm; mean length of notogaster 507 µm, mean width 417 µm (n=6). Prodorsum. Sensillus about 85 µm long with slender head; rostral seta about 83 µm long, acuminate, with setulae (seta type 3: cf. Fig. 1a); lamellar seta about 88 µ m long, bacilliform with setulae (type 4); interlamellar seta about 125 µm long, acuminate, with setulae (type 3). Notogaster. Detailed length values of all setae in table 3; shapes of setae in Figs 4e and 9b. With distinct posterior boss. Ventral region. As typical for the genus. The genital setation (Fig. 5e) is slightly variable within the range from 12–14 pairs, median number is 13, often asymmetrically expressed. Material examined. (1) Austria, Carinthia, Waidischbach near Ferlach; leg. G. Raspotnig, 22.2.200 7 and 24.9.2008; from litter and moss in a Pinus stand. Five specimens. (2) Austria, Tyrol, Kranebitter Au near Innsbruck; leg. Heinrich Schatz, March 2000; moist litter in a deciduous forest. Four specimens. (3) Germany, Bärenklau Forest northern Berlin; leg. G. Weigmann, 1991; from litter and moss in a Pinus stand. One specimen. Remarks. The most obvious difference between the two forms of Trhypochthonius japonicus is in the number of genital setae: 6–10 (median 8) in the Japanese form, 12–14 (median 13) in the European form B (= forma occidentalis). The notogastral setation profile is very similar, but some setae appear a little stronger (cf. the form types indicated in table 3) in forma occidentalis from Austria, compared to those of the nominate form from Shigoku Island, Japan (Aoki 1970). Except seta h 2 and p -setae, all notogastral setae are consistently longer in forma occidentalis, both absolutely and in relative proportions, yet the differences mostly are not significant (see table 4 in the statistical section below). Also there is a tendency for larger bodies in the European forma occidentalis, though we have few data (n=3 for the Japanese form, n=6 for the European form from Carinthia; the Innsbruck-population has the same range) and ranges overlap (tab. 3). Small differences occur also among populations of Trhypochthonius tectorum from different origins (see the statistical section below). With such limited sampling of the Palaearctic region, we do not overestimate the taxonomical relevance of the differences between the studied populations in Japan and Europe, so we propose only a regional form for the European populations. This proposal is supported by the fact that the profile of the oil gland secretion of the European population is identical to that of the Japanese T. japonicus as described by Sakata et al. (2003) (G. R., unpublished). Trhypochthonius misumaiensis Fujikawa, 2000 has the same range of genital setation (11–12 pairs) and a similar notogastral setation as does T. japonicus forma occidentalis. Yet the body is smaller (629–643 µm length) and there are some notogastral setae with stronger setulation: c 3, f 2, h 2, p 2. Most notogastral setae are distinctly shorter, especially c 1, c 2, d 1, d 2, which are less than 15 µm. We regard T. misumaiensis to be a subspecies of T. silvestris (see above). Distribution and ecology. The nominate form of Trhypochthonius japonicus has been found only in Japan (Shigoku Island); the European forma occidentalis has been found only in Austria and northern Germany. In Europe T. japonicus seems to prefer moderately dry to moist litter in forests. The European records in Carinthia and North Germany are syntopic with T. silvestris europaeus n. subsp., Published as part of Weigmann, Gerd & Raspotnig, Günther, 2009, Comparative morphological and biometrical studies on Trhypochthonius species of the tectorum species group (Acari: Oribatida: Trhypochthoniidae), pp. 1-31 in Zootaxa 2269 (1) on pages 16-18, DOI: 10.11646/zootaxa.2269.1.1, http://zenodo.org/record/5306773, {"references":["Aoki, J. - i. (1970) The Oribatid mites of the Islands of Tsushima. Bulletin of the National Science Museum Tokyo, 13, 395 - 442.","Sakata, T., Shimano, S. & Kuwahara, Y. (2003) Chemical ecology of oribatid mites. III. Chemical cmposition of oil gland exudates from two oribatid mites, Trhypochthoniellus sp. and Trhypochthonius japonicus (Acari: Trhypochthoniidae). Experimental and applied Acarology, 29, 279 - 291.","Fujikawa, T. (2000) Five new species of the genera Trhypochthoniellus and Trhypochthonius. Edaphologia, 65, 35 - 53."]}

While considerable knowledge on the chemistry of the scent gland secretions from the opilionid suborders Laniatores and Cyphophthalmi has been compiled, it is the Palpatores (Eupnoi and Dyspnoi) where chemical data are scarce. In particular, the Dyspnoi have remained nearly unstudied, mainly due to their reported general reluctance to release secretions as well as to the phenomenon of production of insoluble—and inaccessible—solid secretion. We here show that at least certain nemastomatid Dyspnoi, namely all three species of genus Carinostoma, indeed produce a volatile secretion, comprising octan-3-one, 6-methyl-5-hepten-2-one and acetophenone in species-specific combinations. In all Carinostoma spp., these volatiles are embedded in a semi-volatile, naphthoquinone matrix (mainly 1,4-naphthoquinone and 6-methyl-1,4-naphthoquinone). In detail, acetophenone and traces of naphthoquinones characterize the secretions of Carinostoma carinatum. A mixture of octan-3-one, 6-methyl-5-hepten-2-one and large amounts of naphthoquinones were found in C. elegans, and 6-methyl-5-hepten-2-one together with small amounts of naphthoquinones in the secretions of C. ornatum. So far, exclusively naphthoquinones had been reported from a single dyspnoan hitherto studied, Paranemastoma quadripunctatum.