Stygopholcus skotophilus Kratochvíl, 1940

Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, European Journal of Taxonomy 752, pp. 1-60 : 22-32

Stygopholcus skotophilus Kratochvíl, 1940

Figs 2 , 6–8 , 43–76 , 147–148 , 154

Stygopholcus skotophilus Kratochvíl, 1940: 16 , figs 1a, c, 2, 4c–d, 5c–d, 6, 7 (right).

Holocnemus (Hoplopholcus) Absolonii Kulczyński, 1914: 355 (part), pl. 16, fig. 25 only.

Holocnemus (Stygopholcus) absoloni (misidentification) – Kratochvíl 1935: 11.

Stygopholcus skotophilus – Senglet 1971: figs 40–41; 2001: 65, fig. 22.

Hoplopholcus scotophilus – Brignoli 1971: 261, fig. 19. — Tomić et al. 2000: 35P (specimen from Vidrovan cave).

Stygopholcus skotophilus montenegrinus (misidentification) – Naumova et al. 2016: 434 (all specimens except those from Cetinje, Golubova pećina, and Babatuša pećina).

“S112 Stygopholcus absoloni? Bal 15” & “S108 Stygopholcus absoloni? Bal 8” – Eberle et al. 2018. — Huber et al. 2018.

Misidentification

Hoplopholcus scotophilus – Tomić et al. 2000: 35P (specimens from near Velja cave; see S. montenegrinus ).

Diagnosis

Distinguished from other species in the ‘northern clade’ ( S. absoloni ; S. montenegrinus ) by bulb measures Aand C ( Fig. 158 ; smaller than other species, no overlap), by bulb measure D ( Fig. 159 ; larger than other species, with little overlap), and by epigynal measure F ( Fig. 164 ; larger than other species, with little overlap); from S. absoloni also by fewer hairs on male cheliceral base ( Figs 51 , 162 ; with little overlap), by smaller, weakly sclerotized prolateral sclerite on procursus (arrows in Figs 45–46 ), by straight procursus (rather than curved toward ventral; Fig. 44 ), by longer procursus ( Fig. 161 ; with little overlap), by shape of retrolateral bulbal process in retrolateral view ( Fig. 47 ; large notch on ventral side), and by bulb measures Band E ( Figs 157, 160 ; larger than in S. absoloni , with little overlap); from S. montenegrinus also by wider prolateral sclerite on procursus (arrows in Figs 45–46 );

Type material

MONTENEGRO – Kotor • 1 ♂, 1 ♀ (without epigynum), 5 juvs, 1 prosoma (juv.?), 1 damaged abdomen (juvenile?), one complete juvenile abdomen, putative syntypes (see Remark below), examined; Pećina u Ivici; 42.5384° N, 18.6207° E [ Naumova et al. 2019: 42.569° N, 18.662° E]; date and collector unknown; NMP .

Remark

According to Růžička et al. (2005), the type vial originally contained two labels saying “ Stygopholcus skotophilus sp. n. ” and “Pećina u Ivici, č. III”. This agrees with other labels in Kratochvíl’s handwriting in the NMP Stygopholcus material. Both labels are lost. Růžička et al. (2005) further state that the vial contained “ 1♂ 3♀ (one of them without epigyne) 4 juv. ”, suggesting that there were two epigyna. However, the vial does not contain any epigynum.

Other material examined

MONTENEGRO – Kotor • 7 ♂♂, 6 ♀♀, 1 juv.; near Zvečava , Pećina u Ivici, in cave; 42.5384° N, 18.6207° E; 960 m a.s.l.; 28 May 2014; B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22196 ) • 1 ♂, 1 ♀, 3 juvs , in pure ethanol; same collection data as for preceding; ZFMK ( Bal15 ) • 9 ♂♂, 7 ♀♀; near Zvečava, near Pećina u Ivici, under stones in forest; 42.5349° N, 18.6237° E; 1000 m a.s.l.; 28 May 2014; B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22197 ) • 2 ♀♀, 1 juv., in pure ethanol; same collection data as for preceding; ZFMK ( Bal16 ) • 1 ♀; Crkvice, Vilina pećina; 42.5546° N, 18.6624° E [ Naumova et al. 2019: 42.560° N, 18.629° E]; 855 m a.s.l.; 30 Oct. 2015; M. Pavlek leg.; CBSS ( Ar 3674 ) • 1 juv.; same locality as for preceding; 17 Jul. 1973; C. Deeleman leg.; NMNL (ARA 19032) • 1 ♂, 1 ♀; Crna Gora, Dragalj, Pećina kod Dvoriškog ždrijela ; 42.5881° N, 18.6737° E; 715 m a.s.l.; 3 Dec. 2014; M. Pavlek leg.; CBSS ( Ar 3482 ) • 1 ♂, 3 ♀♀; Crni Nugli , unnamed cave; 42.586° N, 18.697° E; 590 m a.s.l.; 26 Mar. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 5 ♂♂, 1 juv., Crni Nugli, Čora pećina ; 42.594° N, 18.712° E; 750 m a.s.l.; 26 Mar. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 1 ♂, 1 juv.; Crni Nugli, Dragalj, Pećina s mostom ; 42.5929° N, 18.7100° E; 725 m a.s.l.; 25 Aug. 2017; M. Pavlek leg.; CBSS ( Ar 5387 ) • 1 ♀; same locality as for preceding; 25 Aug. 2017; A. Kirin leg.; CBSS ( Ar 5391 ) • 1 ♀; Crni Nugli, Dragalj, Trokutasta pećina ; 42.5931° N, 18.7107° E; 710 m a.s.l.; 25 Aug. 2017; A. Kirin leg.; CBSS ( Ar 5395 ) • 1 ♂; 4 km SW Crni Nugli, Miljukovica pećina [ Kratochvíl 1935: “Mijukovica”]; 42.57° N, 18.67° E; 1932(?); J. Kratochvíl leg.(?); NMP • 1 ♀; Risan, Teveričke Polje, Teverička pećina; 780 ma.s.l.; 42.570° N, 18.676° E; 28 Jul. 2010; M. Komnenov leg.; CMK • 1 ♀; Risan, Kameno More, Knezlaz , Bukovica pećina ; 42.545° N, 18.683° E; 1 Aug. 2010; M. Komnenov leg.; CMK • 2 ♀♀; Risan, Kameno More, Gornja Pokljuka pećina; 42.542° N, 18.693° E; 580 m a.s.l.; 31 Jul. 2010; M. Komnenov leg.; CMK • 7 ♂♂, 5 ♀♀, 2 juvs; same locality as for preceding; 27 Mar. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 2 ♂♂; Risan distr., Crkvice vill., potholes near Shuto Blagojević monastery; 42.568° N, 18.623° E; 1045 m a.s.l.; 17 Aug. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 1 ♂; Blagojevići, Ovnova rupa, in cave; 42.5681° N, 18.6298° E; 1040 m a.s.l.; 25 Aug. 2017; M. Pavlek leg.; CBSS ( Ar 5384 ) • 2 ♂♂, 3 juvs; Strmena pećina, selo Han, 2 km Eof Bor pećina, near Veli Vrh; 42.564° N, 18.633° E; 28 Jul. 1969; J. Kratochvíl leg.; NMNL (ARA 19030) • 1 ♀; Risan, Sopot pećina ; 42.5137° N, 18.6816° E; 13 Jul. 1971; C. Deeleman leg; NMNL (ARA 19031) . – Nikšić • 1 ♂; Grahovo, Pećina Vojvode Dakovića ; 42.6558° N, 18.6758° E; 680 ma.s.l.; 6 Nov. 2015; M. Lukić leg.; CBSS ( Ar 3780 ) • 1 ♂; same locality as for preceding; 6 Nov. 2015; A. Kirin leg.; CBSS ( Ar 3776 ) • 2 ♂♂, 1 ♀; same locality as for preceding; 28 Mar. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 1 ♂, 4 ♀♀; Trubjela, Krstac, Pećina kod Krstaca ; 42.7659° N, 18.8031° E; 925 m a.s.l.; 21 Aug. 2017; M. Pavlek leg.; CBSS ( Ar 5355 ) • 1 ♂; same collection data as for preceding; CBSS ( Ar 5356 ) • 2 ♂♂ (2 vials); Trubjela, Kita, pećina Ledenica na Kiti ; 42.7729° N, 18.7843° E; 6 Aug. 1969 and 24 Jul. 1970; C. Deeleman leg.; NMNL ( ARA 19033–34 ) • 1 ♂, 2 ♀♀, 3 juvs; Podkita, Širbaba pećina; 42.786° N, 18.825° E; 940 m a.s.l.; 28 Mar. 2006; B. Petrov and S. Lazarov leg.; NMNHS • 1 ♂, 1 ♀, 1 juv.; Vidrovan, Vidrovanska pećina; 42.858° N, 18.937° E; 660 ma.s.l.; 29 Apr. 2016; T. Delić, D. Škufca and Zamolo leg.; ZCSL ( S227 ) • 2 ♂♂, 2 ♀♀; same locality as for preceding; 17 Aug. 2011; M. Komnenov leg.; CMK • 1 ♂; same locality as for preceding; 5 Sep. 2000; B. Ćurčić leg.; IZUB • 1 ♂, 1 ♀; Golija Mt., Donje Čarađe, Drljača pećina; 43.023° N, 18.674° E; 1035 m a.s.l.; 13 Aug. 2010; M. Komnenov leg.; CMK • 1 ♀; [near Javljem], Kerovačka pećina; 42.9981° N, 18.7273° E; 4Aug. 1969; C. Deeleman leg.; NMNL (ARA 19036) • 1 ♂, 1 ♀; Golija Mt., Stozi, Stoška pećina ; 42.952° N, 18.781° E; 1230 m a.s.l.; 10 Aug. 2010; M. Komnenov leg.; CMK • 1 ♂; Golija Mt., Jasikovice, Lazareva pećina ; 43.038° N, 18.699° E; 8 Aug. 2010; M. Komnenov leg.; CMK . – Plužine • 1 ♂; Zaborije, (Golija) Donja Brezna , Ðokova Ledenica pećina; 43.0333° N, 18.8528° E; 1 Aug. 1972; C. Deeleman leg.; NMNL (ARA 19037) .

BOSNIAANDHERZEGOVINA • 8 ♂♂, 6 ♀♀; near Trebinje, Kličanj brdo (hill), Ilijina pećina, in cave near eastern entrance; 42.7188° N, 18.3303° E; 550 m a.s.l.; 26 May 2014; B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22198 ) • 2 ♀♀, 2 juvs, in pure ethanol; same collection data as for preceding; ZFMK ( Bal8 ) • 2 ♂♂, 1 ♀; same locality as for preceding; 3 Apr. 2012; M. Lukić leg.; CBSS ( Ar 3376 , 3379 ) • 1 ♀, 1 juv.; Trebinje , Kličanj brdo, Vučja pećina; 42.716° N, 18.326° E; 425 m a.s.l.; 27 Jan. 2007; J. Bedek leg.; CBSS ( Ar 3383 ) • 2 ♂♂, 4 ♀♀; same locality as for preceding; 6 Dec. 2014; M. Pavlek leg.; CBSS ( Ar 3481 ) • 2 juvs; same locality as for preceding; 27 Jan. 2007; H. Bilandžija leg.; CBSS ( Ar 3375 ) • 3 ♂♂, 2 ♀♀; Trebinje , Arenstorfova pećina; 42.695° N, 18.364° E; 6 Sep. 2006; M. Komnenov leg.; CMK • 1 ♂, 1 ♀; Trebinje , Petrina, Matulićeva pećina; 42.694° N, 18.362° E; 6 Sep. 2006; M. Komnenov leg.; CMK • 2 ♂♂, 1 ♀; Trebinje , Zupci, Vilina pećina; 42.663° N, 18.364° E; 5 Sep. 2006; M. Komnenov leg.; CMK • 1 ♂, 2 ♀♀, 21 juvs (2 vials); same locality as for preceding; 22 Jul. 1969; J. Kratochvíl leg.; NMNL ( ARA 19025–6 ) • 3 ♂♂, 3 ♀♀; Trebinje , Ilijin vrh, Narančivi, Mitrušica pećina; 42.732° N, 18.268° E; 20 Sep. 2008; M. Komnenov leg.; CMK • 3 ♂♂, 2 ♀♀, 7 juvs; Trebinje , Dražin Do , Pećina kod Rupe [“pećina kod rupa, selo Dražin”]; 42.71° N, 18.30° E; 24 Jul. 1969; J. Kratochvíl leg.; NMNL (ARA 19029) • 1 ♀, 3 juvs; Trebinje , Dražin do, Tučevac, in cave; 42.7142° N, 18.3033° E; 21 Aug. 2017; A. Kirin leg.; CBSS ( Ar 5372 ) • 1 ♀, Trebinje , Taleža, Taleža pećina; 42.714° N, 18.231° E; 270 m a.s.l.; 7 Sep. 2006; M. Komnenov leg.; CMK • 1 ♂, 4 ♀♀; near Trebinje, Bihovo, Pavlova pećina [= Ilijina pećina near Bihovo; cf. Lukić et al. 2020], in cave near entrance; 42.6665° N, 18.3078° E; 420 m a.s.l.; 26 May 2014; B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22199–22200 ) • 1 ♀, 2 juvs, in pure ethanol; same collection data as for preceding; ZFMK ( Bal9 ) • 1 ♀, misidentified paralectotype of S. absoloni ; same locality as for preceding; 20 Jul. 1912; K. Absolon leg.; MIZW • 1 ♀, 3 juvs; same locality as for preceding (“Eliashöhle”); date unknown; Verhoeff leg.; ZMB ( 9067 ) • 1 ♀ [and 1 male palp, possibly from different locality, according to label by P. Brignoli]; same collection data as for preceding; MCVR • 1 ♂, 1 ♀; Zupci , Lemov do, Lemova pećina; 42.609° N, 18.399° E; 14 Sep. 2006; M. Komnenov leg.; CMK • 1 ♂; Orjen , Ubla, Čavčina jama; 42.598° N, 18.520° E; 18 Aug. 2007; M. Komnenov leg.; CMK • 1 ♀; Grebci , Debelin pećina [“Debeljin pećina, Osojnik”]; 42.7202° N, 18.0955° E; 420 m a.s.l.; 16 Jul. 1972; J. Kratochvíl leg.; NMNL (ARA 19023) • 1 ♂, 1 juv.; Grebci , Kali pećina; 42.7406° N, 18.0626° E; 525 m a.s.l.; 17 Jul. 1972; C. Deeleman leg.; NMNL (ARA 19024) • 1 ♀; Njemanica , unnamed cave below Jametina jama; 43.081° N, 18.482° E; 1025 m a.s.l.; 14 Aug. 2007; M. Komnenov leg.; CMK • 3 ♂♂, 5 ♀♀, 10 juvs (2 vials); Gacko , Korita , Pećina u Pejevim Torinama; 43.055° N, 18.493° E; 11 Aug. 1968; C. Deeleman leg.; NMNL ( ARA 19027–8 ) • 1 ♂, 1 ♀; Gacko , Dramešina, Studena pećina; 43.213° N, 18.609° E; 15 Aug. 2007; M. Komnenov leg.; CMK • 2 ♂♂; Gacko , Lukovice , Pećina u Lukovicama; 43.1838° N, 18.4458° E; 975 m a.s.l.; 12 Nov. 2015; M. Lukić leg.; CBSS ( Ar 3766 ) • 1 ♂; same locality as for preceding; 12 Nov. 2015; A. Kirin leg.; CBSS ( Ar 3764 ) • 1 ♂, 1 ♀; same collection data as for preceding; CBSS ( Ar 3799 ) .

CROATIA – Dubrovnik-Neretva • 1 ♂; Konavle, Sniježnica , “Jama V= 13”; 42.574° N, 18.352° E; 20 Nov. 2011; M. Kajinić leg.; CBSS ( Ar 2776 ) • 2 ♂♂, 1 ♀, 3 juvs; Konavle, Sniježnica, Jezero , in cave; 42.5804° N, 18.3561° E; 905 m a.s.l.; 16 Mar. 2002; R. Ozimec leg.; CBSS ( Ar 3356 ) • 1 ♂; same locality as for preceding; 1 Nov. 2016; A. Kirin leg.; CBSS ( Ar 5255 ) • 1 ♀, 5 juvs; same locality as for preceding; 1 Nov. 2016; T. Rožman leg.; CBSS ( Ar 4776 ) • 2 juvs; same locality as for preceding; 28 May 2004; M. Pavlek leg.; CBSS ( Ar 3360 ) • 1 ♂; Konavle , Sniježnica, Škrabuljica , in cave; 42.5670° N, 18.3851° E; 685 m a.s.l.; 1 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4772 ) • 1 ♂, Dubrovnik , Gromača, Špilja za Gromačkom vlakom; 42.7435° N, 18.0274° E; 605 m a.s.l.; 22 Feb. 2019; A. Kirin leg.; CBSS ( Ar 6819 ) • 1 ♀; same locality as for preceding; 11 Dec. 2019; M. Pavlek leg.; CBSS ( Ar 6833 ) • 1 ♂; Konavle , Gruda, Vilina špilja; 42.5066° N, 18.3913° E; 205 m a.s.l.; 6 Apr. 2010; R. Ozimec and A. Schönhofer leg. ; ROC (R260).

Localities unknown or unspecified

• 1 ♂, 1 ♀; “ Herzegowina, Tartarushöhle ”; Verhoeff leg.; ZMB ( 9073 ) • 1 ♂, 1 juv.; “c 86, 1935” [no further data]; NMP • 1 juv.; “ Wolfshöhle , Südherzegowina ”, “Vukovo jama bei Vrbanja”; Verhoeff leg.; ZMB ( 9069 ) .

Credible published records (not examined or juveniles only)

MONTENEGRO – Kotor • ♂ ♀ (?), material apparently lost; near Zvečava, Lopata pećina; 42.537° N, 18.622° E [ Naumova et al. 2019: 42.569° N, 18.662° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • 1 juv.; 2 km WČora pećina, Matjaševicapećina ; 42.595° N, 18.700° E [ Naumova et al. 2019: 42.569° N, 18.662° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940]; NMP • ♂ ♀ (?), material apparently lost; Dragaljsko polje, near Dvoriško ždrijelo (“Dvorečko zdrielo”), Tomova pećina; 42.603° N, 18.675° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Dragaljsko polje, above Crni Nugli , Lakičević pećina; 42.597° N, 18.700° E [ Naumova et al. 2019: 42.263° N, 19.033° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; 1 km above Lakičević pećina, Čora pećina ; 42.594° N, 18.712° E [ Naumova et al. 2019: 42.586° N, 18.697° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Jankovo selo, Pećina na Jankovom vrhu; 42.590° N, 18.658° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; 4 km Nof Pećina na Jankovom vrhu, Pećina za Jankovim vrhom; 42.620° N, 18.656° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near road Crkvice-Veli vrh, cave near Crkvice; 42.560° N, 18.630° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Crkvice, 1 km Wof Blagojević, Pećina kod Blagojevića ; 42.573° N, 18.620° E; 1932(?), J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Poljana, Pećina kod Šunjevca ; 42.577° N, 18.648° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Knezlac, Izeta pećina; 42.544° N, 18.689° E [ Naumova et al. 2019: 42.524° N, 18.646° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Knezlac, near Izeta pećina, Studena pećina; 42.544° N, 18.689° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Pokljuka donja [ Kratochvíl 1935: “dolnja”]; 4 km SE Knezlac; 42.542° N, 18.693° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Napode, near Risan-Crkvice road, Vilina (“ Vilna ”) pećina; 42.5546° N, 18.6624° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Vilina pećina, Pecinis kod Napode ; 42.555° N, 18.662° E [ Naumova et al. 2019: 42.519° N, 18.716° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; 1 km SW Risan, Golodražnica pećina; 42.503° N, 18.699° E [ Naumova et al. 2019: 42.535° N, 18.689° E]; 185 m a.s.l.; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940]. – Nikšič • ♂ ♀ (?), material apparently lost; near Jabukovac, Leskova pećina; 42.83° N, 18.47° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Leskova pećina, Golubnjačka pećina; 42.83° N, 18,47° E [ Naumova et al. 2019: 42.397° N, 18.841° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; near Grahovac, on Brdo Omutič mountain, Vodena pećina; 42.687° N, 18.617° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; above Osječenica, Bobjerska pećina; 42.699° N, 18.634° E [ Naumova et al. 2019: 42.674° N, 18.662° E]; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost, Elazov dô, near Zagora, Elazova pećina; 42.704° N, 18.655° E [ Naumova et al. 2019: 42.706° N, 18.673° E]; 1932(?); J. Kratochvíl leg. (?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Troškarica planina mountains ; Vranova jama; 42.838° N, 18.551° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] .

Possible record (not examined)

The following record in Kratochvíl (1940) is dubious because he assigned his specimen(s) to S. absoloni , but specimens from the closest caves we examined (Debelin pećina, Kali pećina, Špilja za Gromačkom vlakom) were all unambiguous S. skotophilus . We thus assume that Kratochvíl misidentified his specimen(s) from this cave.

BOSNIA AND HERZEGOVINA • ♂ ♀ (?), material apparently lost; near Grebci, Grabovica pećina ; 42.733° N, 18.081° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] .

Material assigned tentatively

The material listed here originates from caves around Sedlari in Bosnia and Herzegovina. This material has been studied before by Kratochvíl (1940) and Senglet (2001), and both authors assigned most of the specimens to S. skotophilus (only the female from Medja pećina was assigned to S. absoloni by Kratochvíl 1940). However, both males and females from these caves show some small but consistent differences to S. skotophilus (see section on variation below) so the material is here listed apart to draw attention to this potential separate species (of which no fresh material is available).

BOSNIAANDHERZEGOVINA • 2 ♂♂, 13 ♀♀, 3 juvs (2 vials); “Sedlari/Popovo-Polje” [according to Senglet 2001 the specimens were collected in Grabova pećina]; ~ 42.825° N, 18.055° E; 18 Sep. 1970; A. Senglet leg.; MHNG • 2 juvs; same locality as for preceding (“Grabova peć kod Sedlari”); 17 Jul. 1932; J. Kratochvíl leg.(?); NMP • 3 ♂♂, 6 juvs; “Trovrata peć kod Sedlari” (according to Kratochvíl 1940, this cave is near Grabova pećina); 17 Jul. 1932; J. Kratochvíl leg.(?); NMP • 6 juvs; “Vodena peć kod Sedlari” [according to Kratochvíl 1940, this cave is near Trovrata pećina and Grabova pećina]; 17 Jul. 1932; J. Kratochvíl leg.(?); NMP • 1 ♀, 2 juvs; “ Medja peć na Ilijina brdu , Grmljani ” [according to Kratochvíl 1940 this cave is located 2 km W of Grmljani]; ~ 42.84° N, 18.01° W; 1932(?); J. Kratochvíl leg.(?); NMP .

Redescription

Male (type locality; ZFMK (Ar 22196))

MEASUREMENTS. Total body length 5.5, carapace width 2.2. Distance PME–PME 160 µm; diameter PME 150 × 180 µm; distance PME–ALE 20 µm; diameter AME 65 µm; distance AME-AME 30 µm. Leg 1: 51.2 (13.7 + 0.9 + 13.6 + 20.3 + 3.2), tibia 2: 10.0, tibia 3: 8.2, tibia 4: 8.9; tibia 1 L/d: 62.

COLOR (in ethanol). Carapace pale whitish to ochre-yellow, posterior half, ocular area, and clypeus slightly darker (light brown); sternum light brown, with dark brown margins and three pairs of dark triangular radial marks; legs pale ochre-yellow to light brown, without dark rings, without dark lines; abdomen pale gray, with distinct dark dorsal marks in posterior third and lateral marks in posterior half; ventrally with light brown genital area, indistinct diffuse purplish marks in front and behind genital area and in front of spinnerets.

BODY. Habitus as in Fig. 2 . Ocular area slightly raised. Carapace with deep median pit ( Fig. 61 ) and pair of very shallow indistinct furrows diverging from pit to posterior margin of carapace. Clypeus unmodified. Sternum wider than long (1.50/1.05), unmodified. Abdomen oval, pointed at spinnerets; gonopore with six epiandrous spigots ( Fig. 62 ); ALS with two spigots each ( Fig. 69 ).

CHELICERAE. As in Figs 51–52 , with pair of large frontal apophyses set with 2 modified hairs each, with 12 and 13 modified hairs each on frontal face ( Figs 63–64 ); with stridulatory ridges ( Fig. 71 ).

PALPS. As in Figs 6–8 ; coxa with low retrolateral hump; trochanter barely modified; femur with distinct retrolateral process proximally, with stridulatory pick (modified hair) proximally on prolateral side ( Fig. 65 ); femur-patella joints shifted towards prolateral (arrows in Fig. 6 ); tibia very large, with two trichobothria; tibia-tarsus joints shifted towards retrolateral (arrows in Fig. 8 ); tarsus with exposed tarsal organ ( Fig. 66 ); procursus dorsally with curved hairs (distal third curved ‘upwards’), with strong hump on prolateral side; procursus tip with projecting dorsal sclerite, strong hinged ventral sclerite, and membranous central part with brush of hair-like processes on retrolateral side ( Figs 67–68 ), with weakly sclerotized cylindrical process on prolateral side bordering ventral sclerite (arrow in Fig. 45 ). Genital bulb ( Figs 47–50 ) with basal sclerite connecting to tarsus, small transparent dorsal process, and main sclerite consisting of retrolateral process and dorsal process; retrolateral process with bifid tip, distal part on ventral side thickened and heavily sclerotized; dorsal process with two finger-shaped tips, ventral tip simple, retrolateral tip with additional strong protrusion on ventral side.

LEGS. With short spines ventrally on femur 1 in two rows ( Fig. 73 ), prolateral row ~23 spines, retrolateral row ~28 spines; without curved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 4%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with irregular platelets rather than with distinct pseudosegments.

Male (variation)

Males from most other localities paler, without lateral dark marks on abdomen, with fewer median dark marks (restricted to area above spinnerets), rarely without abdominal marks. Tibia 1 (N = 35): 11.1–14.7 (mean 12.8); tibia 2 (N = 37): 8.1–10.8 (mean 9.6); spines on femur 1: 32–58 (mean 52); modified hairs on each cheliceral basis (N = 74): 9–16 (mean 12.3); modified hairs on each cheliceral apophysis (N = 74): 1–5 (mean 2.7); chelicerae maximum width (N = 36): 0.74–0.86 (mean 0.80); procursus length (N = 37): 1.28–1.48 (mean 1.39); bulbal measures A–E: A (N = 20): 0.28–0.31 (mean 0.30); B (N = 20): 0.08–0.10 (mean 0.09); C (N = 20): 0.13–0.15 (mean 0.14); D (N = 20): 0.28–0.33 (mean 0.31); E (N = 15): 0.82–0.90 (mean 0.87); epiandrous spigots (N = 26): 5–8 (mean 6.1).

Males from near Sedlari differ slightly in bulbal measure Asmaller (0.25–0.28) and bulbal measure B smaller (0.07–0.08); in addition, the smaller wart on the retrolateral bulbal process (arrow in Fig. 48 ) is more strongly developed.

Female

In general similar to male, but femur 1 without spines, with pair of indistinct processes posteriorly on carapace and pair of opposing poorly visible plates on abdomen; distance between processes/plates: 1.1. Tibia 1 (N = 36): 10.4–14.3 (mean 12.4); tibia 2 (N = 36): 7.2–10.3 (mean 9.0); epigynum width (N = 36): 1.02–1.46 (mean 1.21); epigynum length (N = 36): 0.54–0.68 (mean 0.61); epigynal measure F (N = 36): 0.42–0.58 (mean 0.47). Epigynum anterior plate semicircular, weakly protruding, with weak transversal furrows ( Figs 55–58 , 76 ), with internal structures visible in uncleared specimens: round anterior structure, short and indistinct parallel sclerites posteriorly wide apart; posterior plate short but wide. Internal genitalia ( Figs 53–54 , 147–148 ) with distinctive ventral arc ( Fig. 154 ; approximately as long as wide), with large pore plates ( Fig. 54 ).

Females from near Sedlari differ in having the parallel sclerites posteriorly in the female internal genitalia closer together (epigynal measure F: 0.32–0.36; Figs 59–60 ) and slightly more distinct.

Distribution

Known from the southern border area separating Bosnia and Herzegovina from Montenegro, and from the southernmost tip of Croatia ( Fig. 168 ).

Figures

Figs 1–5. Stygopholcus Kratochvíl, 1932; live specimens. 1. S. absoloni (Kulczyński, 1914); male from Vjetrenica, Bosnia and Herzegovina. 2. S. skotophilus Kratochvíl, 1940; male from Pavlova pećina, Bosnia and Herzegovina. 3. S. montenegrinus Kratochvíl, 1940; male and female with egg-sac from Studenačka pećina, Montenegro. 4–5. S. photophilus Senglet, 1971; male and female from Episkopi, Crete. Photos: BAH.

Figs 6–11. Stygopholcus Kratochvíl, 1932; left male pedipalps, prolateral, dorsal, and retrolateral views. 6–8. S. skotophilus Kratochvíl, 1940; from Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)). 9–11. S. photophilus Senglet, 1971; from Episkopi, Crete (ZFMK (Ar 22205)).Abbreviations: b = genital bulb; dp = dorsal process (of genital bulb); p = procursus; rp = retrolateral process (of genital bulb). Arrows point at joints shifted toward prolateral (femur-patella) and retrolateral (tibia-tarsus). Scale bars: 1 mm.

Figs 43–50. Stygopholcus skotophilus Kratochvíl, 1940. 43–44. Male from Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)), left tarsus and procursus, prolateral and retrolateral views. 45– 46. Males from Pećina u Ivici, Montenegro (ZFMK (Ar 22196)) and from Grabova pećina, Sedlari, Bosnia and Herzegovina (MHNG), tip of left procursus, prolateral (slightly ventral) views; arrows point at distinctive prolateral sclerites. 47–50. Male from Ilijina pećina (ZFMK (Ar 22198)), left genital bulb, retrolateral, ventral, prolateral-distal, and dorso-distal views; arrow points at wart that is more strongly developed in specimens from near Sedlari. ds = dorsal sclerite. Scale bars: 43–44, 47–50 = 0.5 mm; 45–46 = 0.25 mm.

Figs 51–54. Stygopholcus skotophilus Kratochvíl, 1940; male from Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)), female from Pećina u Ivici, Montenegro (ZFMK (Ar 22196)). 51– 52. Male chelicerae, frontal and lateral views. 53–54. Cleared female genitalia, ventral and dorsal views. Scale bars: 0.5 mm.

Figs 55–60. Stygopholcus skotophilus Kratochvíl, 1940; epigyna, ventral views. 55–56. Pećina u Ivici, Montenegro (ZFMK (Ar 22196)). 57. Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)). 58. Studena pećina, Bosnia and Herzegovina (CMK). 59–60. Specimens assigned tentatively from Grabova pećina, Sedlari, Bosnia and Herzegovina (MHNG). Scale bar: 0.5 mm (all at same scale).

Figs 61–68. Stygopholcus skotophilus Kratochvíl, 1940; male and female from Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)). 61. Male ocular area, frontal view; arrows point at ‘pseudoeyes’. 62. Male gonopore and epiandrous spigots. 63–64. Modified hairs frontally on male chelicerae. 65. Stridulatory pick on male palpal femur. 66. Male palpal tarsal organ. 67. Tip of left procursus, retrolateral view. 68. Hair-like structures on tip of procursus (detail of previous image). Abbreviations: ds = dorsal sclerite; vs = ventral sclerite. Scale bars: 61 = 100 µm; 62, 67 = 20 µm; 63, 66, 68 = 10 µm; 64–65 = 2 µm.

Figs 69–76. Stygopholcus skotophilus Kratochvíl, 1940; male and female from Ilijina pećina, Bosnia and Herzegovina (ZFMK (Ar 22198)). 69. Male ALS. 70. Female ALS and PMS. 71–72. Stridulatory files on male and female chelicerae. 73. Spines on male femur 1. 74. Tip of male right tarsus 4, ventral view; arrows point at comb-hairs. 75. Tarsal organ on male tarsus 4. 76. Epigynum, ventral view. Abbreviations: ALS = anterior lateral spinneret; PMS = posterior median spinneret. Scale bars: 69–70, 72, 74 = 10 µm; 71, 73 = 20 µm; 75 = 2 µm; 76 = 100 µm.

Figs 145–152. Stygopholcus Kratochvíl, 1932; cleared female genitalia, ventral and dorsal views. 145–146. S. absoloni (Kulczyński, 1914); from Zavala, Bosnia and Herzegovina (MHNG). 147– 148. S. skotophilus Kratochvíl, 1940; from Pećina u Ivici, Montenegro ((ZFMK Ar 22196)). 149– 150. S. montenegrinus Kratochvíl, 1940; from Golubinja pećina, Montenegro (CMK). 151–152. S. photophilus Senglet, 1971; from Choumerion/Pérama, Crete (MHNG).

Figs 153–156. Stygopholcus Kratochvíl, 1932; cleared female genitalia in dorsal views; dorsal sclerotized arcs tilted towards the back. 153. S. absoloni (Kulczyński, 1914); from Balićeva špilja, Croatia (ZFMK (Ar 22195)). 154. S. skotophilus Kratochvíl, 1940; from Pećina u Ivici, Montenegro (ZFMK (Ar 22196)). 155. S. montenegrinus Kratochvíl, 1940; from Studenačka pećina, Montenegro (ZFMK (Ar 22201)). 156. S. photophilus Senglet, 1971; from Choumerion/Pérama, Crete (MHNG). Abbreviations: da = dorsal arc; e = epigynal plate; v = ʻvalve’; va = ventral arc.

Figs 157–160. Bulbal measures B–E for the three species of the ‘northern clade’ of Stygopholcus Kratochvíl, 1932 (cf. Figs 15–18). Bulbal measure A is strongly correlated with measure C and therefore not shown. Bulbal measures sorted from smallest to biggest for each species of the ‘northern clade’. The bulb of S. photophilus was not measured (it differs strongly in shape, and measure D is inapplicable). Asterisk in Fig. 160 = dubious specimen from Jama na Brezdanu (see text).

Figs 161–166. Procursus length, number of modified hairs on each male chelicera, epigynum width/ length, epigynum measure F (cf. Fig. 21), and relationships between tibia 1 and tibia 2 lengths in males and females. Measures/counts sorted from smallest to biggest for each species. Abbreviations: abs. = S. absoloni; sko. = S. skotophilus; mon. = S. montenegrinus; pho. = S. photophilus.

Fig. 168. Known distribution of the ‘northern clade’ ofStygopholcus Kratochvíl, 1932. Pale blue and pale orange marks denote specimens that are assigned tentatively to the respective species. Abbreviations: Alb = Albania; BaH = Bosnia and Herzegovina; Cro = Croatia; Mon = Montenegro.