Stygopholcus absoloni ( Kulczyński, 1914 )

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 : 12-22

Stygopholcus absoloni ( Kulczyński, 1914)

Figs 1 , 12–42 , 145–146 , 153

Holocnemus (Hoplopholcus) Absolonii Kulczyński, 1914: 355 (part), pl. 16, figs 24, 26–27 (not fig. 25).

Holocnemus (Stygopholcus) absoloni – Kratochvíl 1934: 181 (part; see Remarks below).

Hoplopholcus absoloni – Brignoli 1971: 257, fig. 20.

Stygopholcus absoloni – Kratochvíl 1932: 2; 1940: 10, figs 3, 4a–b, 5a–b, 7 (left); 1978: 28. — Absolon & Kratochvíl 1932: 75. — Senglet 1971: figs 37–39; 2001: 65, fig. 21.

“S110 Stygopholcus Bal 3 ” – Eberle et al. 2018. — Huber et al. 2018.

Misidentifications

Holocnemus (Stygopholcus) absoloni – Kratochvíl 1934: 181 (part; see Remarks below); 1935: 11 (see Remarks below).

Remarks

For most of the localities listed in Kratochvíl (1934), the presence of S. absoloni could either be confirmed or appears very probable judging from the distribution of confirmed specimens. Exceptions are the following: Ilijina pećina and “Pećina Napode” (= Vilina pećina) (see S. skotophilus ), Medja peć na Ilijina brdu, Vodena peć kod Sedlara, Grabova peć kod Sedlara, and Trovrata peć (tentatively assigned to S. skotophilus ), and Jamutina [pećina] (see S. montenegrinus ). Kratochvíl’s (1934) “Pec kod Ljubova” [Ljubovo?] is not mentioned in the extensive list of localities in Kratochvíl (1940) and is thus not further considered herein.

By contrast, all caves listed in Kratochvíl (1935) probably contain S. skotophilus ; this is either confirmed by new material or very likely judging from the distribution of confirmed specimens.

Diagnosis

Distinguished from other species in the ‘northern clade’ ( S. skotophilus ; S. montenegrinus ) by large prolateral sclerite on procursus (distally narrowing; arrow in Fig. 14 ), by procursus slightly curved toward femur (rather than straight; Fig. 13 ), by shorter procursus ( Fig. 161 ; with little overlap), by shape of retrolateral bulbal process in retrolateral view ( Fig. 15 ; small notch on ventral side), and by bulb measures: measure B smaller ( Fig. 157 ; with little overlap), measure D smaller ( Fig. 159 ; no overlap), and measure Esmaller ( Fig. 160 ; no overlap). From S. skotophilus also by more hairs on male cheliceral base ( Fig. 162 ; with little overlap) and by larger bulb measure C ( Fig. 158 ; no overlap), and by smaller epigynal measure F ( Fig. 164 ; little overlap).

Type material

Lectotype

BOSNIAANDHERZEGOVINA • ♂ (designated herein, examined); “near Trebinje, Zovica jama”; ~ 42.7° N, 18.3° E; date unknown (1912?); K. von Arenstorff leg.; MIZW.

Paralectotype

BOSNIA AND HERZEGOVINA • ♀, examined; same collection data as for lectotype; MIZW ,

Other material examined

BOSNIAANDHERZEGOVINA • 1 ♂, 2 ♀♀; Vjetrenica pećina; 42.8459° N, 17.9837° E; 280 ma.s.l.; 27 May 2014; B.A. Huber and M. Komnenov leg.; in cave near entrance and outside cave, hole in ground; ZFMK ( Ar 22192 ) • 1 ♀, 1 juv., in pure ethanol; same collection data as for preceding; ZFMK ( Bal12 ) • 1 ♂, 1 ♀; Popovo polje, Zavala , Bjelušica , in cave; 42.8447° N, 17.9786° E; 350 ma.s.l.; 2 Dec. 2014, M. Pavlek leg.; CBSS ( Ar 3480 ) • 1 ♂; same locality as for preceding; 26 Oct. 2015; M. Pavlek leg.; CBSS ( Ar 3687 ) • 3 ♀♀, 5 juvs; southern limit of Zavala village, Bjelušica (“Bjelušica pećina”); 42.8447° N, 17.9786° E; 350 ma.s.l.; 15–16 Jul. 1932; J. Kratochvíl leg.(?); NMP • 15 ♂♂, 10 ♀♀; Zavala; 42.849° N, 17.978° E; 19 Sep. 1970; A. Senglet leg.; MHNG • 1 ♀; near monastery of Zavala, Pećina kod Monastira ; 42.850° N, 17.979° E; 16 Jul. 1932; J. Kratochvíl leg.; NMP • 1 ♂; Zavala monastery , small cave behind monastery; 42.8499° N, 17.9795° E; 300 m a.s.l.; 27 May 2014: B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22193 ) • 1 ♂, 1 ♀; Zavala , Orlica pećina; 42.846° N, 17.973° E; 23 Oct. 1968; G. and A. Vigna leg.; MCVR • 1 ♂, 2 ♀♀; 2 km NW of Zavala, Grabova peć kod Grabova dola ; 42.858° N, 17.960° E; 20 Jul. 1932; J. Kratochvíl leg.; NMP • 1 ♂, 1 ♀, with Kratochvíl’s original label “ Stygopholcus Absoloni (Kulcz.) , ♀ et ♂ type! 19 b” [these are probably Kratochvíl’s (1940) ‘emended types’ and the supposed syntypes in Růžička et al. (2005)]; presumably from same locality as preceding; NMP • 1 ♂, 1 ♀; “ Inv. No 409”; same collection data as as for preceding (“ Grabowa-Höhle , leg. Kratochvíl 1932 ”); NHMW • 1 ♀, 1 juv.; Čvaljina , Zavala , Baba pećina ; 42.8608° N, 17.9722° E; 270 m a.s.l.; 27 Aug. 2017; M. Pavlek leg.; CBSS ( Ar 5509 ) • 1 ♂; same locality as for preceding; 2 Dec. 2014; M. Pavlek leg.; CBSS ( Ar 3479 ) • 1 ♂, 1 ♀, 5 juvs; between Grabova peć and Duga peć, Vukotina peć kod Čvaljinskih staja ; 42.86° N, 17.96° E; date unknown; J. Kratochvíl leg.(?); NMP • 1 ♂, 1 ♀, 5 juvs; Sof Zavala , near Belenići, Bezdan pod Licem ; 42.813° N, 18.000° E; 25 Jul. 1932; J. Kratochvíl leg.; NMP • 1 ♂, 2 ♀♀, 1 juv.; Njavre, Donje Hrasno , Markova pećina ; 42.9779° N, 17.8269° E; 400 ma.s.l.; 2 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4682 ) • 17 ♂♂, 8 ♀♀, 1 juv. (2 vials); Počitelj/Čapljina ; 43.11° N, 17.73° E; 20 Sep. 1970; A. Senglet leg.; MHNG • 4 ♂♂, 6 ♀♀, 3 juvs; Studenci (“grotte”); 43.161° N, 17.638° E; 21 Sep. 1970; A. Senglet leg.; MHNG • 1 ♂, 4 juvs; Marica pećina; 43.255° N, 17.905° E; 25 Jul. 1936; P. Rémy and A. Schweitzer leg.; MNHN ( Ar 10219 ) • 1 ♀, 3 juvs; Kovina pećina; 43.250° N, 17.935° E; 22 Aug. 1936; P. Rémy and A. Schweitzer leg.; MNHN ( Ar 10235 ) • 2 ♂♂; Drinovci, Majića ponor ; 43.3600° N, 17.2834° E; 260 m a.s.l.; 5 Dec. 1999; M. Cukrov leg.; CBSS ( Ar 3372 ) • 1 ♂, 1 ♀; Grude, Peć Mlini , Ravlića pećina; 43.3390° N, 17.3233° E; 250 m a.s.l.; 17 Jul. 1974; C. Deeleman leg.; NMNL (ARA 19022).

CROATIA – Split-Dalmatia • 1 ♂; Lećevica, Kladnjice, Jama 7; 43.6918° N, 16.2915° E; 470 ma.s.l.; 28 Apr. 2020; T. Rožman leg.; CBSS ( Ar 6962 ) • 1 ♂; same locality as for preceding; 17 Apr. 2020; N. Kuharić leg.; CBSS ( Ar 6959 ) • 1 ♀; Lećevica, Kladnjice, Trasirana jama ; 43.6913° N, 16.2873° E; 460 m a.s.l.; 17 Apr. 2020; T. Rožman leg.; CBSS ( Ar 6980 ) • 1 ♂; Muć, Neorić, Velika špilja kod Neorića ; 43.686° N, 16.553° E; 705 m a.s.l.; 31 Jul. 2007; M. Pavlek leg.; CBSS ( Ar 3374 ) • 3 ♂♂, 2 ♀♀; Dugopolje, Balići, Balićeva špilja (= Kraljeva pećina); 43.5728° N, 16.5738° E; 405 m a.s.l.; 31 Aug. 2000; F. Gasparo leg.; ZFMK ( Ar 22195 ) • 1 ♂; same locality as for preceding; 12 May 2012; M. Pavlek leg.; CBSS ( Ar 3189 ) • 1 ♀; same locality as for preceding; 5 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4802 ) • 1 juv.; same locality as for preceding; date unknown; T. Delić et al. leg.; ZCSL ( S19 ) • 1 ♀; Dugopolje, Balešina jama ; 43.579° N, 16.581° E; 310 m a.s.l.; 28 Apr. 2012; M. Pavlek leg.; CBSS ( Ar 2759 ) • 1 ♂; Dugopolje, Kotlenice, Maklutača , in cave; 43.5693° N, 16.6205° E; 400 m a.s.l.; 5 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4978 ) • 1 ♂, 2 ♀♀; Dugopolje, Kotlenice, Golubinka pećina; 43.56° N, 16.65° E; 2 May 1978; C. Deeleman leg.; AMNH • 1 ♀; same collection data as for preceding; NMNL (ARA 19016) • 1 ♂, 2 ♀♀, 3 juvs; Dugopolje, near Kotlenice, Vranjača špilja kod Dugopolja (“ Vranjača pećina”); 43.5622° N, 16.6482° E; 30 Jul. 1967; C. Deeleman leg.; NMNL (ARA 19020) • 1 juv., assigned tentatively; Dugopolje, Bradarići, Drinovčuša , in cave; 43.5485° N, 16.6386° E; 795 m a.s.l.; 2 Nov. 2017; T. Delić leg.; CBSS ( Ar 5698 ) • 2 ♀♀ (2 vials); Trilj, Ugljane, pećina Turkuša ; 43.5789° N, 16.7363° E; 430 m a.s.l.; 16 Apr. 1974; C. Deeleman leg.; NMNL ( ARA 19017–18 ) • 1 ♂, 2 ♀♀; Ugljane, Jamau Krčevini ; 43.57° N, 16.76° E; 15 Apr. 1974; C. Deeleman leg.; NMNL (ARA 19019) • 2 ♂♂, 1 ♀; Omiš, Kučiće, Nikolina špilja ; 43.4253° N, 16.8271° E; 250 m a.s.l.; 23 Mar. 2003; M. Cukrov leg.; CBSS ( Ar 2003 ) • 2 ♂♂, 1 ♀, 1 juv.; Omiš, Cetina (“ Cajetina ”)- Canyon , small valley with stream, dense Fagus forest with Hedera sp. , with big rocks, in crevices, under stones; 95 m a.s.l.; 43.433° N, 16.762° E; 13 May 2006; A. Schönhofer leg.; SMF • 2 ♂♂, 2 ♀♀; Šestanovac, Gornji Balići, Zlatna peć (” Balica zlatna pećina, Balići ”); 43.490° N, 16.905° E; 17 Jun. 1974; C. Deeleman leg.; NMNL (ARA 19021) • 5 ♂♂, 7 ♀♀, 1 juv.; Imotski, “ N of Imotski near Čočić, Pećina u Solunu ” (possibly Špilja u Solunu; 43.4589° N, 17.2217° E); date unknown; J. Kratochvíl leg.(?); NMP • 1 ♀: Brela, Dupci, Svetica , in cave; 43.4011° N, 16.8945° E; 300 m a.s.l.; 16 Nov. 2005; M. Pavlek leg.; CBSS ( Ar 2012 ) • 1 ♂; Brela, Nevistine stine, Jama iznad Prozorčića ; 43.3838° N, 16.9320° E; 615 m a.s.l.; 24 Mar. 2004; H. Bilandžija leg.; CBSS ( Ar 1995 ) • 1 ♂, 1 juv.; Šestanovac, Svaguše, Mužina špilja; 43.4170° N, 17.0076° E; 505 m a.s.l.; 18 Oct. 2002; M. Lukić leg.; CBSS ( Ar 1928 ) • 5 ♂♂, 2 ♀♀; same locality as for preceding; 18 Apr. 1974; C. Deeleman leg.; NMNL (ARA 19011) • 2 ♂♂, 1 ♀; Zagvozd, Sridnja gora, Dedići , Mala špilja na Bukovcima ; 43.4037° N, 17.0217° E; 465 m a.s.l.; 21 Nov. 2005; M. Pavlek leg.; CBSS ( Ar 2007 ) • 1 ♀; Zagvozd, Rastovac, Dedići , Špilja u Radinovcima ; 43,3873° N, 17.0103° E; 400 m a.s.l.; 25 Oct. 2006; M. Pavlek leg.; CBSS ( Ar 2009 ) • 1 ♂; Zagvozd, Rastovac, Dedići , Raputinova špilja; 43.3865° N, 17.0199° E; 360 ma.s.l.; 24 Nov. 2005; M. Pavlek leg.; CBSS ( Ar 2005 ) • 1 ♂; Zagvozd, Čikeši, Jamau Čikešima ; 43.4161° N, 16.9670° E; 200 m a.s.l.; 23 Nov. 2005; M. Pavlek leg.; CBSS ( Ar 1996 ) • 1 ♀; Zagvozd, cave near Zagvozd; 43.40° N, 17.05° E; 13 Oct. 2002; C. Deeleman leg.; CBSS ( Ar 1930 ) • 1 ♂, 5 ♀♀; Zagvozd, Brnasi, Špilja u Gaju ; 43.383° N, 17.041° E; 18 Oct. 2002; C. Deeleman leg.; CBSS ( Ar 1993 ) • 2 ♂♂; Zagvozd, Prodani, Pružina špilja ; 43.394° N, 17.059° E; 520 m a.s.l.; 15 Oct. 2002; C. Deeleman leg.; CBSS ( Ar 1926 ) • 1 ♀; Zagvozd, Spilja near Katušići; 43.384° N, 17.055° E; 18 Apr. 1974; C. Deeleman leg.; NMNL (ARA 19012) • 1 ♀, 1 juv.; Zagvozd, Župa, Brikinjava špilja ; 43.3452° N, 17.0969° E; 740 ma.s.l.; 17 Nov. 2005; M. Pavlek leg.; CBSS ( Ar 2000 ) • 1 ♂; Zagvozd, Župa, Matijaševa peć; 43.3206° N, 17.1223° E; 585 m a.s.l.; 23 Jan. 2016; M. Lukić leg.; CBSS ( Ar 3954 ) • 1 ♀; same locality as for preceding; 23 Jan. 2016; M. Pavlek leg.; CBSS ( Ar 3960 ) • 1 ♂; Zagvozd, Župa, Gradska spila; 43.3180° N, 17.1176° E; 755 m a.s.l.; 23 Jan. 2016; M. Pavlek leg.; CBSS ( Ar 3980 ) • 1 ♂; Zagvozd, Župa, Samogorska špilja; 43.3182° N, 17.1211° E; 715 m a.s.l.; 23 Jan. 2016; M. Pavlek leg.; CBSS ( Ar 3935 ) • 1 ♂; same locality as for preceding; 23 Jan. 2016; M. Lukić leg.; CBSS ( Ar 3962 ) • 1 juv.; same locality as for preceding; 23 Jun. 2017; R. Ozimec leg.; CBSS ( 5477 ) • 1 ♂, 1 ♀; Zagvozd, Župa, Stonjska peć donja; 43.3152° N, 17.1307° E; 610 m a.s.l.; 22 Jan. 2016; M. Pavlek leg.; CBSS ( Ar 3975 ) • 1 ♂, 1 ♀, 1 juv.; Baška Voda, Bast, Kukor , in cave; 43.3556° N, 16.9917° E; 425 m a.s.l.; 2 Jan. 1999; R. Ozimec leg.; CBSS ( Ar 3367 ) • 2 ♂♂; same locality as for preceding; 21 Jan. 2016; M. Lukić leg.; CBSS ( Ar 3952 ) • 1 ♂; same locality as for preceding; 21 Jan. 2016; M. Pavlek leg.; CBSS ( Ar 3968 ) • 3 ♀♀, 1 juv.; same locality as for preceding; 21 Jan. 2016; T. Rožman leg.; CBSS ( Ar 3971 ) • 4 juvs, assigned tentatively; Makarska, Stipkovac, Jama pod Stipkovcem ; 43.3090° N, 17.0651° E; 1360 m a.s.l.; 21 Jun. 2017; A. Kirin leg.; CBSS ( Ar 5467 ) • 1 ♀, 1 juv.; Tučepi, Ravna Vlaška, Tučepska Vilenjača (“ Pećina Vilenjača ”); 43.2854° N, 17.0842° E; 1180 m a.s.l.; 28 Aug. 2000; F. Gasparo leg.; ZFMK ( Ar 22194 ) • 1 ♀; same locality as for preceding; 30 Apr. 2012; A. Ćukušić leg.; CBSS ( Ar 3370 ) • 1 ♀; same locality as for preceding; 24 Jun. 2017; M. Pavlek leg.; CBSS ( 5035 ) • 1 ♂, 2 ♀♀, 1 juv.; Tučepi, Ravna Vlaška, Jama kod Tučepske Vilenjače ; 43.2847° N, 17.0850° E; 1195 m a.s.l.; 17 Jun. 2002; R. Ozimec leg.; CBSS ( Ar 2001 ) • 1 ♂, 1 juv.; Vrgorac, Kozica, Jujnovići , Jujnovića šplija; 43.2666° N, 17.1980° E; 405 m a.s.l.; 27 Apr. 2002; R. Ozimec leg.; CBSS ( Ar 1999 ) • 3 ♀♀; same locality as for preceding; 14 Oct. 2002; C. Deeleman leg.; CBSS ( Ar 1927 ) • 1 ♀; same locality as for preceding; 14 Oct. 2002; R. Ozimec leg.; CBSS ( Ar 2006 ) • 1 ♀; same locality as for preceding; 1 May 2012; M. Pavlek leg.; CBSS ( Ar 2773 ) • 1 ♀; Vrgorac, Saranač, Šošići , Jama iznad Saranača ; 43.2490° N, 17.1747° E; 770 m a.s.l.; 25 Mar. 2003; R. Ozimec leg.; CBSS ( Ar 1998 ) • 1 ♂, 1 ♀, 7 juvs; probably same locality as for preceding; [“grotje bij Saranač op berg Skok”, Gornje Igrane]; 7 Jul. 1974; C. Deeleman leg.; NMNL (ARA 19010) • 1 ♂, 1 ♀; Vrgorac, Zavojane, Signalka , in cave; 43.2691° N, 17.2380° E; 1030 m a.s.l.; 29 Aug. 2018; T. Rožman leg.; CBSS ( Ar 6573 ) • 1 ♂, 3 juvs; Vrgorac, Zavojane, Rođendanska jama; 43.2686° N, 17.2389° E; 1045 m a.s.l.; 29 Apr. 2018; F. Belak leg.; CBSS ( Ar 6420 ) • 1 ♂, 2 juvs; Vrgorac, Zavojane, Škrapica , in cave; 43.2558° N, 17.2679° E; 720 m a.s.l.; 30 Apr. 2018; F. Belak leg.; CBSS ( Ar 6578 ) • 2 ♀♀; Vrgorac, Baletići, Ančića velika Golubinka , in cave; 43.2584° N, 17.2842° E; 875 ma.s.l.; 8 Oct. 2018; N. Kuharić leg.; CBSS ( Ar 6553 ) • 1 ♂, 1 ♀; Vrgorac, Zavojane, Drž gače 2, in cave; 43.2590° N, 17.2828° E; 870 m a.s.l.; 8 Oct. 2018; N. Sudar and N. Kuharić leg.; CBSS ( Ar 6395 ) • 1 ♂, 2 ♀♀; Vrgorac, Stilja, Jama pod Gažnovcem ; 43.2297° N, 17.3142° E; 510 m a.s.l.; 4 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4283 ) • 3 ♀♀, 1 juv.; same locality as for preceding; 31 Mar. 2018; F. Belak leg.; CBSS ( Ar 6587 ) • 1 ♂, 1 ♀, 1 juv. (2 vials); Vrgorac, Orah [hill], Ružovača pećina; 43.229° N, 17.417° E; 390 m a.s.l; 23 Oct. 1980 and 26 Apr. 1981; C. Deeleman leg.; NMNL ( ARA 19008–9 ) • 1 ♀, 1 juv.; Hvar Island, Jelsa, Viskoka , Franetova jama; 43.14° N, 16.93° E; 28 Feb. 1999; R. Ozimec leg.; CBSS ( Ar 3363 ) • 1 ♀; Hvar Island, Jelsa, Zastražišče , Vela Stiniva cove, Špilja u Grliću ; 43.1580° N, 16.8136° E; 3 m a.s.l.; 3 May 2011; D. Hmura leg.; CBSS ( Ar 2513 ) • 2 ♂♂; same locality as for preceding; 19 Aug. 2016; M. Pavlek leg.; CBSS ( Ar 3996 ) • 1 ♀; Hvar Island, Jelsa, Humac , Grapčeva špilja; 43.1341° N, 16.7537° E; 200 m a.s.l.; 3 May 2011; A. Ćukušić leg.; CBSS ( Ar 2512 ) • 1 ♂, 1 ♀ (epigynum missing); same locality as for preceding, “Grabćina šplija”; 20 Aug. 1968; P. Brignoli leg.; MCVR • 2 ♂♂, 1 juv. (2 vials); same locality as for preceding; 26 Oct. 1980 (one vial without date); C. Deeleman leg.; NMNL ( ARA 19013–14 ). – Dubrovnik- Neretva • 1 ♀, 1 juv.; Ploče, Gradac, Ričevica , in cave; 43.0999° N, 17.3909° E; 525 m a.s.l.; 5 Nov. 2019; T. Rožman leg.; CBSS ( Ar 6430 ) • 1 ♀, 1 juv.; Kula Norinska, Krvavac, Kokanova jama; 43.0273° N, 17.5849° E; 90 m a.s.l.; 12 Dec. 2018; B. Jalžić leg.; CBSS ( Ar 6481 ) • 1 ♀; Zažablje, Goračić, Vidonje , Šolkina jama; 42.9665° N, 17.6625° E; 230 m a.s.l.; 27 Feb. 2016; B. Jalžić leg.; CBSS ( Ar 3989 ) • 2 ♀♀; Zažablje, Dobranje, Kornjatuša , in cave; 42.9790° N; 17.6740° E; 610 m a.s.l.; 2 Nov. 2013; T. Čuković leg.; CBSS ( Ar 3384 ) • 1 ♀; Dubrovačko Primorje, Slano, Točionik , Vranja peć; 42.8683° N, 17.8124° E; 230 ma.s.l.; 2 Nov. 2016; T. Rožman leg.; CBSS ( Ar 5148 ) • 1 ♀; same locality as for preceding; 2 Nov. 2006; M. Pavlek leg.; CBSS ( Ar 4877 ) • 1 ♂, 1 ♀; Pelješac, Ston, Ponikve , Špilja Crno jezero; 42.8473° N, 17.6365° E; 415 m a.s.l.; 15 Nov. 2018; N. Kuharić leg.; CBSS ( Ar 6276 ) • 1 ♂; same collection data as for preceding; CBSS ( Ar 6284 ) • 1 ♀; Pelješac, Ston, Putniković , Kameni kolačić , in cave; 42.8731° N, 17.4664° E; 420 m a.s.l.; 28 Mar. 2019; N. Kuharić leg.; CBSS ( Ar 6161 ) • 1 ♀; Pelješac, Janjina, Vardišta , Špilja u Žarmi ; 42.9354° N, 17.4382° E; 35 m a.s.l.; 28 Apr. 2019; T. Rožman leg.; CBSS ( Ar 6258 ) • 1 ♂, 1 juv.; same collection data as for preceding; M. Pavlek leg.; CBSS ( Ar 6261 ) • 1 ♂; Pelješac, Orebić, Oskorušno , Špilja kod Zagruda u vinogradu ; 42.9666° N, 17.3227° E; 295 m a.s.l.; 13 Dec. 2019; M. Pavlek leg.; CBSS ( Ar 6464 ) • 1 ♀; Pelješac, Orebić, below monastery, Čagjina jama ; 42.9771° N, 17.1552° E; 115 m a.s.l.; 14 Nov. 2018; F. Šarc leg.; CBSS ( Ar 6247 ) • 1 ♀; Pelješac, Orebić, Kućište , Šimunkovića špilja ; 42.9786° N, 17.1291° E; 50 ma.s.l.; 3 Nov. 2016; M. Pavlek leg.; CBSS ( Ar 4692 ) • 1 ♀, 2 juvs; same locality as for preceding; 26 Apr. 2019; T. Rožman leg.; CBSS ( Ar 6781 ) • 1 ♂; Pelješac, Trpanj, Duba Pelješka , Spilja Rupine ; 43.0053° N, 17.1580° E; 460 m a.s.l.; 6 Apr. 2019; T. Dražina leg.; CBSS ( Ar 5980 ) • 1 ♀; same locality as for preceding; 24 Apr. 2019; M. Pavlek leg.; CBSS ( Ar 5986 ) • 1 ♂; same locality as for preceding; 24 Apr. 2019; T. Rožman leg.; CBSS ( Ar 6197 ) • 1 ♂; Pelješac, Trpanj, Donja Vrućica , Špilja u Privoru ; 43.0092° N, 17.2133° E; 180 ma.s.l.; 26 Mar. 2019; N. Kuharić leg.; CBSS ( Ar 6038 ) • 1 ♀, 1 juv.; Pelješac, unnamed cave near Trpanj ; 43.01° N, 17.27° E; 18 Jun. 1975; C. Deeleman leg.; NMNL (ARA 19015) • 1 ♂; Korčula, Žrnovo, Deukalion špilja; 42.9557° N, 17.0976° E; 110 m a.s.l.; 21 Mar. 2010; R. Ozimec leg. ; ROC (R97).

Assigned tentatively (see Variation)

CROATIA – Split-Dalmatia • 1 ♂; Baška Voda, Bast, Topići , Jama na Brezdanu ; 43.3670° N, 16.9732° E; 915 m a.s.l.; 20 Mar. 2004; collector unknown; CBSS ( Ar 1994 ) .

Credible published records (not examined or juveniles only)

BOSNIAANDHERZEGOVINA • 3 juvs; near Čvaljina, Duboka peć; 42.87° N, 17.97° E; date unknown; J. Kratochvíl leg.(?); NMP • ♂ ♀ (?), material apparently lost; between Timor brdo and Čvaljina village, Duga peć; 42.87° N, 17.95° E; date unknown; J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Baba pećina; 42.8608° N, 17.9722° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; 500 m Wof “Bjelušica pećina”, env. Zavala, Orlica pećina; 42.846° N, 17.973° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • 1 juv.; Sof Zavala, near Belenići, Jama za Jamskim vrhom; 42.81° N, 18.00° E; 28 Jul. 1932; J. Kratochvíl leg.(?); NMP .

CROATIA – Split-Dalmatia • ♂ ♀ (?), material apparently lost; Muć, near Sinj , Pećina kod Sinja ; 43.70° N, 16.64° E; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Dugopolje, near Kotlenice, Vranjača špilja kod Dugopolja (“Vranjača pećina”); 43.5622° N, 16.6482° E; 445 m a.s.l.; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), materialapparently lost; Dugopolje, near Dugopolje, Kraljeva pećina [ Kratochvíl 1940 used this present-day synonym of Balićeva špilja for a cave “near” Balićeva špilja]; 43.573° N, 16.570° E, 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Dugopolje, near Dugopolje, Balićeva špilja ; 43.573° N, 16.570° E; 405 m a.s.l.; 1932(?); J. Kratochvíl leg.(?) [ Kratochvíl 1940] • ♂ ♀ (?), material apparently lost; Hvar Island, Jelsa, Grapčeva špilja (“Grabćina šplija”); 43.1341° N, 16.7537° E; 1932(?); J. Kratochvíl leg. (?) [ Kratochvíl 1940].

Redescription

Male (Zavala, MHNG)

MEASUREMENTS. Total body length 5.8, carapace width 2.2. Distance PME–PME 150 µm; diameter PME 140 × 180 µm; distance PME–ALE 30 µm; diameter AME 65 µm; distance AME-AME 25 µm. Leg 1: 55.2 (14.8 + 1.1 + 14.0 + 22.1 + 3.2), tibia 2: 10.8, tibia 3: 8.7, tibia 4: 9.5; tibia 1 L/d: 62.

COLOR (in ethanol). Carapace pale whitish to ochre-yellow, ocular area and clypeus slightly darker (light brown); sternum light brown, with darker brown radial marks; legs ochre yellow, without dark rings, without dark lines; abdomen monochromous pale gray, only above spinnerets with darker mark.

BODY. Habitus as in Fig. 1 . Ocular area slightly raised. Carapace with deep median pit and pair of very shallow indistinct furrows diverging from pit to posterior margin of carapace. Clypeus unmodified. Sternum wider than long (1.50/1.00), unmodified. Abdomen oval, pointed at spinnerets.

CHELICERAE. As in Figs 19–20 , with pair of large frontal apophyses set with 2 and 3 modified hairs, respectively, with 17 and 18 modified hairs each on frontal face; with distinct stridulatory ridges ( Fig. 35 ).

PALPS. In general as in S. skotophilus (cf. Figs 6–8 ); coxa with retrolateral hump; trochanter barely modified; femur with distinct retrolateral process proximally, with stridulatory pick (modified hair) proximally on prolateral side; femur-patella joints shifted towards prolateral; tibia very large, with two trichobothria; tibia-tarsus joints shifted towards retrolateral; procursus ( Figs 12–13 ) slightly curved towards ventral, dorsally with curved hairs (distal third curved ‘upwards’), with strong hump on prolateral side (asterisk in Fig. 12 ). Procursus tip with evenly curved dorsal sclerite and strong hinged ventral sclerite (ds and vs in Figs 13 , 29 ); with strong conical process on prolateral side bordering hinged sclerite (arrow in Fig. 14 ); membranous central part on retrolateral side with brush of hair-like processes ( Fig. 29 ). Genital bulb ( Figs 15–18 ) with basal sclerite connecting to tarsus (bs in Fig. 15 ), small transparent dorsal process (arrow in Fig. 17 ; Fig. 33 ), and main sclerite consisting of long retrolateral process and short dorsal process (rp and dp in Figs 15–18 , 32 ); retrolateral process with bifid tip, distal part on ventral side thickened and heavily sclerotized; dorsal process with finger-shaped ventral tip slightly hooked, retrolateral tip wider, with additional strong protrusion on ventral side.

LEGS. With short spines ventrally on femur 1 in two rows, prolateral row ~19 spines, retrolateral row ~24spines; withoutcurved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 3%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with irregular platelets rather than with distinct pseudosegments.

Male (variation)

Most males with monochromous abdomen with one or few dark marks above spinnerets; few males with additional dark marks laterally in posterior third. Tibia 1 (N = 60): 10.4–15.1 (mean 12.8); tibia 2 (N = 64): 7.6–11.5 (mean 9.6); spines on femur 1 (N = 27): 0–48 (mean 38); modified hairs on cheliceral basis (N = 122): 11–23 (mean 17.4); modified hairs on cheliceral apophysis (N = 122): 1–4 (mean 2.6); chelicerae maximum width (N = 61): 0.66–0.85 (mean 0.78); procursus length (N = 58): 1.10–1.28 (mean 1.21); bulbal measures A–E: A (N = 28): 0.31–0.37 (mean 0.35); B (N = 28): 0.06–0.08 (mean 0.07); C (N = 20): 0.22–0.28 (mean 0.26); D (N = 20): 0.18–0.23 (mean 0.21); E (N = 22): 0.68–0.77 (mean 0.73); epiandrous spigots (N = 20): 5–7 (mean 6.1).

In the single male from Jama na Brezdanu ( Croatia), the bulb differs in several respects from all other material examined, but it is unclear if this is teratological or not (even though it is symmetric). Most conspicuously, the heavy sclerite connecting the basal and distal bulbal sclerites (asterisk in Fig. 18 ) is much smaller and it is not connected to the basal bulbal sclerite; bulb measure Ealso deviates strongly (asterisk in Fig. 160 ).

Female

In general similar to male, but femur 1 without spines, with pair of indistinct processes posteriorly on carapace (arrows in Fig. 39 ) and pair of opposing poorly visible plates on abdomen; distance between processes/plates: 1.1. Tibia 1 (N = 55): 8.2–15.1 (mean 12.0); tibia 2 (N = 59): 4.4–11.1 (mean 8.6); epigynum width (N = 57): 0.72–1.22 (mean 1.05); epigynum length (N = 57): 0.42–0.68 (mean 0.54); epigynal measure F (N = 57): 0.21–0.37 (mean 0.30). Epigynum ( Figs 23–26 ) anterior plate semicircular, weakly protruding, with weak transversal furrows, with internal structures visible in uncleared specimens: round anterior structure and parallel sclerites posteriorly; posterior plate short but wide. Internal genitalia ( Figs 21–22 , 145–146 ) with distinctive ventral arc (va in Fig. 153 ; relatively shorter and wider than in other species), with large pore plates ( Fig. 22 ).

Distribution

Known from southern Croatia and southern Bosnia and Herzegovina ( 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 12–18. Stygopholcus absoloni (Kulczyński, 1914); male from Balićeva špilja, Croatia (ZFMK (Ar 22195)).12–13. Left tarsus and procursus, prolateral and retrolateral views; asterisk in Fig. 12 marks prolateral hump. 14. Tip of left procursus, prolateral (slightly ventral) view; arrow points at distinctive prolateral sclerite. 15–18. Left genital bulb, retrolateral, ventral, prolateral-distal, and dorso-distal views, showing bulb measures A–E; arrow in Fig. 17 points at transparent bulbal process; asterisk in Fig. 18 marks connecting sclerite that is absent in specimen from Jama na Brezdanu. Abbreviations: bs = basal sclerite; dp = dorsal process; ds = dorsal sclerite; rp = retrolateral process; vs = ventral sclerite. Scale bars: 12–13, 15–18 = 0.5 mm; 14 = 0.25 mm.

Figs 19–22. Stygopholcus absoloni (Kulczyński, 1914); male from Balićeva špilja, Croatia (ZFMK (Ar 22195)), female from Zavala, Bosnia and Herzegovina (MHNG). 19–20. Male chelicerae, frontal and lateral views. 21–22. Cleared female genitalia, ventral and dorsal views, showing epigynal measure F. Scale bars: 0.5 mm.

Figs 23–26. Stygopholcus absoloni (Kulczyński, 1914); epigyna, ventral views. 23. Balićeva špilja, Croatia (ZFMK (Ar 22195)). 24. Biokovo, Kukor, Croatia (CBSS (Ar 3367)). 25. Studenci (grotte), Bosnia and Herzegovina (MHNG); arrows point at distinctive internal sclerites. 26. Vjetrenica, Bosnia and Herzegovina (ZFMK (Ar 22192)). Scale bar: 0.5 mm (all at same scale).

Figs 27–34. Stygopholcus absoloni (Kulczyński, 1914); male from Počitelj/Čapljina, Bosnia and Herzegovina (MHNG). 27. Modified hair frontally on male chelicerae. 28. Left male cheliceral apophysis with modified hairs. 29. Tip of left procursus, retrolateral view. 30. Male palpal tarsal organ. 31. Retrolateral bulbal process, distal view. 32. Bulbal processes; arrow points at opening of sperm duct. 33. Transparent bulbal process. 34. Comb hairs on left tarsus 4, prolateral view. Abbreviations: dp = dorsal process; ds = dorsal sclerite; rp = retrolateral process; vs = ventral sclerite. Scale bars: 27, 30, 33, 34 = 10 µm; 28, 31 = 20 µm; 29, 32 = 100 µm.

Figs 35–42. Stygopholcus absoloni (Kulczyński, 1914); male and female from Počitelj/Čapljina, Bosnia and Herzegovina (MHNG). 35–36. Stridulatory files on male and female chelicerae. 37. Female palpal tarsal organ. 38. Tarsal organ on female tarsus 2. 39. Female prosoma, oblique dorsal view; arrows point at posterior humps acting against modified areas on abdomen. 40. Epigynum, ventral view. 41. Female ALS. 42. Female PMS. Scale bars: 35 = 30 µm; 36 = 20 µm; 37–38 = 10 µm, 39 = 200 µm; 40 = 100 µm; 41–42 = 10 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 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.

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 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.

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.