Nemastoma bidentatum gruberi Novak, Slana Novak, Kozel & Raspotnig, 2021

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, European Journal of Taxonomy 777, pp. 1-67 : 20-26

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. 1994; S. Brelih leg. (808/1998, rev. 2007); PMSL • 6 ♂♂, 3 ♀♀, 3 juvs; Lokve , Črnomelj; 2 Jul. 1995; L. Slana Novak and T. Novak leg. (105/1997, rev. 2009); PMSL • 3 ♂♂, 2 ♀♀; Mihelja vas; 18 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 55/1986, TN rev. 2017); PMSL • 1 ♀; Praprot ; 6 Aug. 1997; T. Novak leg. (418/1998, rev. 2008); PMSL • 1 ♀; ibid .; (432/1998, rev. 2008); PMSL • 1 ♀; ibid .; 22 Jul. 1999; L. Slana Novak and T. Novak leg. (37/1999, rev. 2008); PMSL • 1 ♀; ibid .; (110/1999, rev. 2008); PMSL • 2 ♂♂, 2 ♀♀; Stranska vas ; 22 Jul. 1999; L. Slana Novak and T. Novak leg. (108/1999, rev. 2007); PMSL • 2 ♂♂, 1 ♀; Vrčice ; 10 May 2008; L. Slana Novak and T. Novak leg. (110/2012); PMSL • 1 ♂, 3 ♀♀; Vrčice – Semič; 8 Oct. 2012; P. Kozel and T. Novak leg. (64/2015); PMSL. – WL16 • 2 ♀♀; Laze, Uršna Sela ; 10 May 2008; L. Slana Novak and T. Novak leg. (103/2012, rev. 2015); PMSL • 1 ♀; ibid .; (108/2012, rev. 2018); PMSL. – WL17 • 1 ♂, 1 ♀; Brod , Novo Mesto; 31 Aug. 2000; M. Mlakar leg. (811/2000, rev. 2007); PMSL • 1 ♂; Novo mesto; Karaman leg. (87/1983 microscopic preparation, JH det., TN rev. 2011); PMSL • 1 ♂, 1 ♀; ibid .; Karaman leg. (108/1983 microscopic preparation, JH det., TN rev. 2011); PMSL • 1 ♀; ibid .; (814/2000); PMSL • 1 ♂; ibid .; (153/2018 microscopic preparation, JH det., TN rev. 2018); PMSL • 1 ♀, 1 ♀; ibid .; 31 Aug. 2000; M. Mlakar leg. (811/2000, rev. 2007); PMSL • 1 ♂, 2 ♀♀; Ratež ; 29 Apr. 1995; S. Brelih leg. (750/1998, rev. 2007); PMSL. – WL18 • 6 ♂♂, 5 ♀♀; Dolenje Laknice ; 26 Oct. 2013; T. Novak leg. ( GR 4661 , 4662 , 4664 , 4666 , 4667 , 4669 , 4673 , 4676 , 4677−4679 ); PMSL. – WL19 • 1 ♀; Boštanj – Mokronog – Križ; L. Slana Novak and T. Novak leg. (70/2007, rev. 2011); PMSL • 1 ♂; Jelovec ; 15 Aug. 2007; L. Slana Novak and T. Novak leg. (56/2007); PMSL. – WL23 • 1 ♂; Zilje ; 18 Apr. 1996; S. Brelih leg. (767/1998, rev. 2007); PMSL • 1 ♂, 2 ♀♀; Zilje – Vinica ; 17 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (364/1998, rev. 2007); PMSL. – WL24 • 5 ♂♂, 1 ♀, 1 juv.; Gorenjci pri Adlešičih ; 15 Aug. 1999; M. Štangelj and T. Novak leg. (218/1999, rev. 2007); PMSL • 1 ♂; Dragoši, Griblje ; 18 Apr. 1994; S. Brelih leg. (733/1998, rev. 2007); PMSL • 4 ♂♂, 2 ♀♀; Griblje − Desinec ; 16 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (161/1997, rev. 2009); PMSL. – WL25 • 1 ♂; Bečka jama cave , Cad. No. 6275, Boldraž ; 9 Jun. 2004; V. Germovšek, T. Tkavc, A. Kocuvan and T. Novak leg. (21/2004, rev. 2007); PMSL • 1 ♂; Božakovo ; 3 May 2004; L. Slana Novak and T. Novak leg. (48/2004, rev. 2007); PMSL • 2 ♂♂, 3 ♀♀; Bušinja vas ; 18 Apr. 1996; S. Brelih leg. (743/1998, rev. 2007); PMSL • 1 ♂, 4 ♀♀; Kapljišče; 16 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (LSN 113/1986, TN rev. 2007); PMSL • 2 ♂♂, 1 juv.; ibid . (LSN 119/1986, TN rev. 2007); PMSL • 3 ♀♀; Metlika; 31 Aug. 2000; M. Mlakar leg. (816/2000, rev. 2007); PMSL • 1 ♂; ibid .; (818/2000, rev. 2015); PMSL • 7 ♂♂, 9 ♀♀; Trnovec , Metlika; 23 Jul. 1999; L. Slana Novak and T. Novak leg. (141/1999, rev. 2008); PMSL • 1 ♂, 2 ♀♀; Sv. Urban , Metlika; 18 Apr. 1996; S. Brelih leg. (784/1998, rev. 2007); PMSL. – WL26 • 1 ♂; Krašnji vrh; 9 Jun. 2004; V. Germovšek, T. Tkavc, A. Kocuvan and T. Novak leg. (37/2004, rev. 2007); PMSL • 1 ♂; Markučev Mlin, Krašnji vrh ; 1 Jul. 2007; L. Slana Novak and T. Novak leg. (21/2007); PMSL • 4 ♀♀; ibid .; (17/2007, rev. 2011); PMSL. – WL27 • 1 ♂; Gabrje , Novo Mesto; 11 Apr. 1984; I. Sivec leg. (875/1997, rev. 2018); PMSL • 1 ♀; Gabrje – Sv. Miklavž – Krvavi kamen , Gorjanci Mts ; 22 Jul. 1999; L. Slana Novak and T. Novak leg. (123/1999, rev. 2019); PMSL • 5 ♂♂, 6 ♀♀; Glažarica, Mt. Gorjanci ; 22 Jul. 1999; L. Slana Novak and T. Novak leg. (118/1999, rev. 2015); PMSL • 1 ♂, 1 ♀; ibid .; (134/1999, rev. 2018); PMSL • 1 ♀; Golo , Šentjernej; 29 Apr. 1995; S. Brelih leg. (757/1998, rev. 2007); PMSL • 1 ♀; Sv. Miklavž, Gorjanci Mts ; 22 Jul. 1999; L. Slana Novak and T. Novak leg. (128/1999, rev. 2019); PMSL. – WL28 • 3 ♂♂, 7 ♀♀; Brezje pri Raki ; 27 Jul. 2012; S. Samu, P. Kozel and T. Novak leg. (115/2012); PMSL • 1 ♂, 1 ♀; Podvrh, Raka ; 18 Jul. 2012; P. Kozel, T. Kozel and T. Novak leg. (55/2012); PMSL • 9 ♂♂, 10 ♀♀; ibid .; (GR 3734−3746, 3791, TN det.); PMSL • 4 ♂♂, 11 ♀♀; Škocjan; 26 Oct. 2013; T. Novak leg. (GR 4680−4694, TN det.); PMSL. – WL29 • 3 ♂♂, 1 juv.; Boštanj ; 5 Jul. 1987; L. Slana Novak, M. Slana Novak and T. Novak leg. (433/1998, rev. 2007); PMSL • 6 ♂♂, 12 ♀♀; Dolnje Brezovo ; 30 Jun. 2012; T. Novak leg. (GR 3670−3683, TN det.); PMSL • 1 ♂, 1 ♀; Sevnica; 29 Aug. 2000; M. Mlakar leg. (785/2000, rev. 2009); PMSL. – WL37 • 1 ♂, 1 ♀; Grič ; 29 Apr. 1995, S. Brelih leg. (763/1998, rev. 2008); PMSL • 2 ♂♂, 2 ♀♀; Velike Vodenice − Opatova gora ; 15 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (144/1997, rev. 2015); PMSL. – WL38 • 3 ♀♀; Krakovski gozd, Sajevce ; 5 May 1993; S. Brelih leg. (792/1998, rev. 2015); PMSL • 68 ♂♂, 59 ♀♀; Dolenje Radulje – Raški vrh ; 5 Sep. 2012; P. Kozel and T. Novak leg. (PK 98/2012); PMSL • 84 ♂♂, 78 ♀♀; ibid .; 17 Sep. 2012; P. Kozel and T. Novak leg. (PK 99/2012); PMSL • 73 ♂♂, 83 ♀♀; ibid .; 4 Oct. 2012; P. Kozel and T. Novak leg. (PK 100/2012); PMSL • 44 ♂♂, 38 ♀♀; ibid .; 29 Apr. 2013; P. Kozel and T. Novak leg. (PK 43/2013); PMSL • 74 ♂♂, 71 ♀; Podulce – Zabukovje pri Raki ; 20 Aug. 2012; P. Kozel and T. Novak leg. ( PK 96 /2012); PMSL. – WL39 • 1 ♂, 1 ♀; Arto, Brestanica ; 18 Nov. 1994; S. Brelih leg. (189/2001, rev. 2017); PMSL • 3 ♂♂, 3 ♀♀; ibid .; 1 Jul. 2007; L. Slana Novak and T. Novak leg. (28/2007); PMSL • 5 ♂♂, 3 ♀♀; Kremen ; 12 Oct. 2005; L. Slana Novak and T. Novak leg. (141/2005, rev. 2008); PMSL • 1 ♀; Krško; 11 Apr. 1978 (42/2019); PMSL • 1 ♂, 5 ♀♀; ibid .; (47/2019); PMSL • 2 ♀♀; ibid .; (63/2019); PMSL • 1 ♀; ibid .; M. Mlakar leg. (704/2000, rev. 2008); PMSL • 1 ♂; Stara vas ; 24 Aug. 2000; M. Mlakar leg. (709/2000, rev. 2007); PMSL • 2 ♂♂; Stolovnik–Anže ; 9 Oct. 2005; L. Slana Novak and T. Novak leg. (134/2005, rev. 2008); PMSL • 5 ♂♂, 7 ♀♀; ibid .; 14 Oct. 2005; L. Slana Novak and T. Novak leg. (154/2005, rev. 2007); PMSL. – WL48 • 1 ♂; Brežice; 30 Aug. 2000; M. Mlakar leg. (794/2000, rev. 2008); PMSL • 1 ♂; ibid .; 30 Aug. 2000; M. Mlakar leg. (792/2000, rev. 2007); PMSL • 1 ♀; Vitovec, Velike Malence ; Jul. 1994; M. Potokar leg. (289/1995, rev. 2008); PMSL. – WL49 • 6 ♂♂, 1 ♀; Podsreda castle ; 17 Jun. 2001; L. Slana Novak and T. Novak leg. (145/2001, rev. 2008); PMSL. – WL59 • 3 ♂♂, 4 ♀♀; Stara vas ; 14 Aug. 1985; L. Slana Novak, M. Slana Novak and T. Novak leg. (141/1997, rev. 2007); PMSL • 1 ♂; ibid .; 24 Aug. 2000; M. Mlakar leg. (709/2000, rev. 2007); PMSL • 1 ♂; ibid .; (714/2000, rev. 2007); PMSL • 1 ♂; ibid .; (716/2000, rev. 2007); PMSL. – WM01 • 3 ♀♀; Hrastnik; 30 Aug. 2000; M. Mlakar leg. (802/2000, rev. 2018); PMSL • 2 ♀♀; ibid .; (806/2000, rev. 2018); PMSL. – WM10 • 1 ♂, 1 ♀; Lokavec, Zidani most ; 26 May 2004; A. Kapla leg. (56/2004, rev. 2007); PMSL • 2 ♀♀; Radeče; 29 Aug. 2000; M. Mlakar leg. (796/2000, rev. 2015); PMSL • 1 ♀; ibid .; (801/2000, rev. 2015); PMSL. – WM20 • 1 ♂; Mt. Lisca ; 26 May 2004; A. Kapla leg. (54/2004, rev. 2007); PMSL • 1 ♂; ibid .; (62/2004, rev. 2007); PMSL. – WM30 • 7 ♂♂, 8 ♀♀, 1 juv.; Šentvid pri Planini ; 14 Aug. 1983; L. Slana Novak and T. Novak leg. (LSN 52/1983, TN rev. 2007); PMSL • 10 ♂♂, 4 ♀♀; Mt. Bohor ; 20 Aug. 1994, L. Slana Novak and T. Novak leg. (66/1997, rev. 2008); PMSL • 3 ♂♂, 2 ♀♀; Tovornik, Mt. Bohor; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1134/2002, TN rev. 2007); PMSL • 1 ♂, 1 ♀; ibid .; (1149/2002, rev. 2007); PMSL • 4 ♂♂, 1 ♀; Zgornji Reštanj, Senovo ; 20 May 2012; T. Novak leg. (53/2012); PMSL. – WM40 • 1 ♂; Bistri graben , Kozje; 11 Jul. 2003; T. Novak leg. (160/2003, rev. 2007); PMSL • 2 ♀♀; ibid .; (162/2003, rev. 2018); PMSL • 1 ♂; ibid .; (170/2003, rev. 2007); PMSL • 1 ♂; ibid .; (165/2003, rev. 2007); PMSL • 1 ♂; ibid .; (170/2003, rev. 2007); PMSL • 3 ♂♂, 4 ♀♀, 1 juv.; ibid .; 16 Jun. 2019; L. Slana Novak and T. Novak leg. (6/2019); PMSL • 1 ♂, 1 juv.; ibid .; (11/2019); PMSL • 1 ♀; ibid .; (12/2019); PMSL • 1 ♂, 1 ♀, 1 juv.; ibid .; (13/2019); PMSL • 2 ♂♂, 2 ♀♀; Kozje; 14 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 72/1983, TN rev. 2011); PMSL • 1 ♀; ibid .; 31 Jul. 1995; S. Polak leg. (177/2001, rev. 2015); NM • 1 ♀; ibid .; 1 Aug. 1995; S. Polak leg. (168/2001); NM • 3 ♂♂, 2 ♀♀; ibid .; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1150/2002, rev. 2007); PMSL • 3 ♂♂, 5 ♀♀; Podsreda ; 14 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 71/1983, TN rev. 2007); PMSL • 2 ♂♂, 3 ♀♀; Stara Glažuta ; 13 Jul. 2002; L. Slana Novak and T. Novak leg. (1145/2002, rev. 2007); PMSL. – WM41 • 1 ♂, 1 ♀; Podčetrtek; 10 Aug. 1996; L. Slana Novak and T. Novak leg. (64/1997, rev. 2018); PMSL. – WM50 • 4 ♂♂, 1 ♀; Bistrica ob Sotli; 13 Aug. 1983; T. Novak and L. Slana Novak leg. (LSN 46/1983, TN rev. 2007); PMSL • 1 ♂, 1 ♀; Svete gore ; 17 May 1997; L. Slana Novak and T. Novak leg. (2/1997, rev. 2008); PMSL • 1 ♂; ibid .; 29 Aug. 2009; C. Komposch leg.; PMSL • 11 ♂♂, 9 ♀♀; Orešje na Bizeljskem ; 10 Aug. 1996; L. Slana Novak and T. Novak leg. (82/1997, rev. 2009); PMSL .

Description

Male (holotype)

BODY. Length 1.99, body width 1.48.

CHELICERAE. Ch basal article (without Apo) length 0.60, relatively long, 2.4 times as long as wide at Ch max width at dorsal hump. Ch-Apo small, 0.3 times as high as basal article long, rounded-trapezoid, widest distally and highest posteriorly; Ch-Apo 0.18 high, max width 0.17, min width 0.10 (at the base), ventral margin straight, with 0.70 long and 0.02 deep indentation with 10–20 μm high tipped tubercles and denticles constituting a saw. Secretion field fronto-medially, apically. Distal article length 0.67, max width 0.24, movable finger length 0.29 ( Figs 4E , 6E , 9E ).

PEDIPALPS. Pa-Fe relatively thin (Pa-Fe min:max width ~1:2.8), with variable armament: either (the main population) with 4 subequidistant tubercles, spines or long setae, or their combination in the distal Pa-Fe half, or alternatively (marginal population in SW Slovenia), with 2–3 distal stout thorns. Pa-Pt relatively thin (length:max width ~ 1:4.2) ( Figs 5E , 6E , 10E ). Pa article lengths in Table 8 .

PENIS. Pe length 1.57, base 0.57, glans 0.18, stylus 0.07 ( Figs 7E , 8E ).

LEGS. Pseudoarticle leg-Fe formula I−IV: 0−1−1−2. Tarsomere leg-Fe formula I−IV: 10−16−9−10. Leg article lengths in Table 8 .

Female

BODY. Body length 2.49, body width 1.76.

CHELICERAE. Ch basal article length 0.65, 2.6 times as long as wide at dorsal hump, with low-arched dorsal margin in front of dorsal indentation, and low ventral round-triangular hump ( Fig. 13D ). Distal article length 0.67, max width 0.20, movable finger length 0.31.

PEDIPALPS. Pa-Tr with relatively high, symmetrically, slightly knee-like arched dorsal margin, Pa-Fe with conspicuous distal bonce, Pa-Ti elongated ellipsoid ( Fig. 13D ). Pa article lengths in Table 8 .

OVIPOSITOR. Ovipositor length 0.81, Rec sem with one tubular and 12 elongated balloon-like vesicles ( Fig. 11E ).

LEGS. Leg article lengths in Table 8 .

Variability

In eastern Slovenia, males mostly have four thin medio-ventral spines of similar size on the Pa-Fe, the proximal one placed approximately in the middle of the Pa-Fe. In south-eastern Slovenia, some spines can be very small or diminished to a tubercle with a seta. In contrast, in south-western Slovenia, males have three or rarely two large, stout thorns, sometimes resembling those in N. b. bidentatum . Further investigation in Croatian Istria is required to elucidate the whole range of morphological variation and the expected transition between the tender eastern and robust western armament of the Pa-Fe.

Remarks

Nemastoma b. gruberi ssp. nov. is the most variable subspecies within N. bidentatum . Prior to this study, a few collected individuals had been considered within the variability range of N. b. sparsum, e.g., individuals from Buje and Vižinada considered as paratypes of N. b. sparsum (NHMW 4700). Accordingly, Novak et al. (2002) and Novak (2004b, 2005b) misidentified specimens, and those from SW Slovenia were erroneously reported ( Novak 2005b) as a disjunctive population of N. b. sparsum in the Sub Mediterranean region in Slovenia. Near the border of its distribution area in SW Slovenia, N. b. gruberi ssp. nov. has relatively uniform, robust Pa-Fe, with 3 subequidistant, rarely 2, large, stout thorns, much different from the armament in the main population.

Distribution

Croatia, Italy, Slovenia. Vertical distribution in Slovenia: 4−1619 m a.s.l. Type locality: Dolenje Laknice, Mokronog (45.93° N, 15.21° E, 237 m a.s.l.), Slovenia.

Ecology

Nemastoma b. gruberi ssp. nov. inhabits various habitat types in lowland and hilly land, from thermophile shrub and Quercetalia pubescenti–petraeae Klika 1928 communities in southwestern regions to riverine, floodplain and submontane woods in southern and south-eastern Slovenia. Phenology: adults eurychronous.

Figures

Fig. 1. Representatives of the Nemastoma bidentatum Roewer, 1914 complex. A. The habitus of a male Nemastoma bidentatum gruberi Novak, Slana Novak, Kozel & Raspotnig ssp. nov.; Svete gore, Slovenia (photo: C. Komposch); inset: femur IV with two pseudo-articles (apparent articles, i.e., sections delimited with pale, indented rings on the proximal femur portion), and a saw on the latero-posterior margin. B. The body of a male Nemastoma relictum Gruber & Martens, 1968 stat. nov. showing the tuberculate surface of glittering scutum compositum, and position of the ocular mound (tuber oculorum), supracheliceral lamellae, chelicerae with cheliceral apophysis, and pedipalps and leg. The frontal portion of the scutum is smeared over with a hardened secretum of the integumental glands (cf. Wolff et al. 2016); Mt. Grosses Karr, the Koralpen Mts, Austria. C. The body of a male Nemastoma bidentatum gruberi ssp. nov., smeared over with the secretum; Kozje, Slovenia.

Fig. 2. The complete distributional area of the Nemastoma bidentatum Roewer, 1914 complex, according to Gruber & Martens (1968), Martens (2006), Schönhofer & Holle (2007) and own data, with marked positions of the type localities of the taxa.

Fig. 3. Distributional areas of six subspecies (circles) and hybrids (triangles, squares) within the Nemastoma bidentatum Roewer, 1914 in Slovenia and adjacent regions. Note that many localities overlap on the map.

Fig. 4. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Ch basal articles (medial, lateral views).

Fig. 5. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Pa (medial view).

Fig. 6. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Synchroscopic images of the male Ch basal article and Pa-Fe (medial views, except G, right figure, which is in dorso-medial view).

Fig. 7. Nemastoma bidentatum Roewer, 1914 complex. Penes (lateral view). Note: The penis of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 8. Nemastoma bidentatum Roewer, 1914 complex. Glans (dorsal and lateral views). Note: The penis of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 9. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Variation of the Ch basal article (medial view) in species, subspecies and hybrids.

Fig. 10. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. Variation of the Pa-Fe (A–B first image, C, D–E first image, F–G first image; H–J, L–M right, N–O: lateral view; the others in dorso-medial view to show the armament).

Fig. 11. Nemastoma bidentatum Roewer, 1914 complex, ♀♀; Rec sem (lateral view, right portion, except F left figure, which is in dorsal view). A. The frame encloses the ovipositor portion with Rec sem. Note: Female N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 12. Nemastoma bidentatum Roewer, 1914 complex, ♂♂. From left to right: Ch basal article (medial, lateral views), Pa (medial view) and Pe tip (dorsal, lateral views). Arrows indicate the most identifying characters. Note: Pe of N. pluridentatum (Hadži, 1973) stat. nov. is unknown.

Fig. 13. Nemastoma bidentatum Roewer, 1914 complex, ♀♀. From left to right: Silhouettes of the Ch basal article, Pa (both medial view) and Rec sem. Arrows indicate the most identifying characters. Note: female N. pluridentatum (Hadži, 1973) stat. nov. is unknown.