Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula Author Huber, Bernhard A. 0000-0002-7566-5424 Zoological Research Museum Alexander Koenig, Bonn, Germany. b.huber@leibniz-zfmk.de Author Pavlek, Martina 0000-0001-6710-0581 Croatian Biospeleological Society, Zagreb, Croatia. & Ruđer Bošković Institute, Zagreb, Croatia. martina.pavlek@gmail.com Author Komnenov, Marjan 0000-0001-6830-1043 Blwd Kuzman Josifovski Pitu, 1000 Skopje, Macedonia. mkomnenov@gmail.com text European Journal of Taxonomy 2021 2021-06-11 752 1 60 journal article 6076 10.5852/ejt.2021.752.1391 f7f21cba-d005-486a-a383-fe6d09b81f04 2118-9773 4958808 F362301F-08C1-4660-8583-491BFEA32488 Stygopholcusmontenegrinus Kratochvíl, 1940 orig. stat. Figs 3 , 77–112 , 149–150 , 155 Stygopholcusmontenegrinus Kratochvíl, 1940: 20 , fig. 8. Stygopholcus skotophilus montenegrinus – Senglet 1971: 354 , figs 42–43; 2001: 65. — Naumova et al. 2016: 434 (part; see S. skotophilus ). Stygopholcus photophilus (misidentification) – Deltshev et al. 2011: 130 (specimen from near Kozhnja only). Hoplopholcus scotophilus (misidentification) –Tomić et al. 2000: 35P (specimens from near Velja cave). “S109 Stygopholcus absoloni ? Bal 11” – Eberle et al. 2018 . — Huber et al. 2018. Diagnosis Distinguished from other species in the ‘northern clade’ ( S. absoloni ; S. skotophilus ) by narrow, weakly sclerotized prolateral sclerite on procursus (arrows in Figs 77–78 , 100 ); from S. skotophilus also by bulb measures Aand C ( Fig. 158 ; larger than in S. skotophilus , no overlap), by epigynal measure F ( Fig. 164 ; smaller than in S. skotophilus , with little overlap); from S. absoloni also by straight procursus (rather than curved toward ventral; cf. Fig. 44 ), by longer procursus ( Fig. 161 ; with little overlap), by shape of retrolateral bulbal process in retrolateral view ( Fig. 79 ; large notch on ventral side), by bulb measures B and D ( Figs 157, 159 ; larger than in S. absoloni , no overlap) and bulb measure E ( Fig. 160 ; larger than S. absoloni , with little overlap). Type material Holotype MONTENEGRO – Nikšič • ♀ (without epigynum), near Nikšič , Studenačka pećina; 42.7680° N , 18.9312° E [ Naumova et al. 2019 : 42.826° N, 18.786° E]; 6 Sep. 1936 ; J. Kratochvíl leg.(?); NMP . Other material examined MONTENEGRO – Nikšič • 1 juv. ; same collection data as for holotype; NMP • 9 ♂♂ , 8 ♀♀ ; near Nikšić , Studenačka pećina, in cave near entrance and among rocks near cave ; 42.7680° N , 18.9312° E ; 640 m a.s.l. ; 27 May 2014 ; B.A. Huber and M. Komnenov leg.; ZFMK ( Ar 22201 ) • 1 ♂ , 1 ♀ , 1 juv. in pure ethanol; same collection data as for preceding; ZFMK ( Bal11 ) • 4 ♂♂ , 8 ♀♀ ; same locality as for preceding; 16 Sep. 1970 ; A. Senglet leg.; MHNG • 1 ♂ , 1 ♀ ; Trebjesa , Vojna pećina ; 42.765° N , 18.959° E ; 19 Aug. 2011 ; M. Komnenov leg.; CMK • 1 ♂ , 1 ♀ ; Velja pećina; 42.728° N , 18.943° E ; 16 Aug. 2011 ; M. Komnenov leg.; CMK • 1 ♂ , 3 ♀♀ , totally fragmented; unnamed cave above Velja pećina; 42.73° N , 18.94° E ; 6 Sep. 2000 ; B. Ćurčić leg.; IZUB • 2 ♀♀ ; Carev Most , Budoška pećina; 42.7303° N , 18.9498° E ; 26 Jul. 1969 ; C. Deeleman leg.; NMNL (ARA 19035) • 2 ♂♂ , 2 ♀♀ ; Slivlje , Golubešnica pećina, 42.724° N , 19.013° E ; 18 Aug. 2011 ; M. Komnenov leg.; CMK . – Podgorica • 2 ♀♀ ; Velje Brdo , Megara pećina; 42.463° N , 19.199° E ; 17 Feb. 2010 ; M. Komnenov leg; CMK • 2 ♂♂ , 3 ♀♀ ; Ljuboviči , artificial tunnel; 42.433° N , 19.255° E ; 21 Apr. 2011 ; M. Komnenov leg.; CMK • 2 ♂♂ , 2 ♀♀ ; Vranjina , Golubinja pećina (= Pećina na Vranjini); 42.2703° N , 19.1308° E ; 21 Apr. 2011 ; M. Komnenov leg.; CMK • 1 ♂ , 4 juvs ; same locality as for preceding; 23 Aug. 2017 ; A. Kirin leg.; CBSS ( Ar 5342 ) • 1 ♂ , 1 juv. ; same locality as for preceding; 23 Aug. 2017 ; M. Pavlek leg.; CBSS ( Ar 5329 ) • 1 juv. ; same locality as for preceding; 23 Aug. 2017 ; A. Kirin leg.; CBSS ( Ar 5334 ) • 10 juvs ; same locality as for preceding; 31 Jul. 1969 ; J. Kratochvíl leg.; NMNL (ARA 19041) . – Cetinje • 2 ♂♂ , 1 ♀ ; Cetinjska pećina; 42.389° N , 18.921° E [ Naumova et al. 2019 : 42.401° N, 18.935° E]; 10 Aug. 2011 ; M. Komnenov leg.; CMK • 1 ♂ ; Lipska pećina; 42.375° N , 18.955° E [ Naumova et al. 2019 : 42.375° N, 19.952° E]; 21 Apr. 2011 ; M. Komnenov leg.; CMK • 2 ♂♂ , 2 ♀♀ ; presumably same locality as for preceding (“Lipa Dobersko”); 16 Sep. 1970 ; A. Senglet leg.; MHNG • 1 ♂ ; Začir , Začirska pećina; 42.343° N , 18.992° E ; 5 Oct. 2010 ; M. Pavičević leg.; CMK • 1 ♂ ; same locality as for preceding; 21 Aug. 2017 ; T. Rožman leg.; CBSS ( Ar 5405 ) • 3 ♂♂ , 1 ♀ ; Rijeka Crnojevića , Obodska pećina ; 42.352° N , 19.005° E ; 19 Apr. 2011 ; M. Komnenov leg.; CMK • 1 ♀ ; same locality as for preceding; 23 Aug. 2017 ; M. Pavlek leg.; CBSS ( Ar 5484 ) • 1 ♂ , 4 juvs ; same collection data as for preceding; A. Kirin leg.; CBSS ( Ar 5485 ) • 3 ♂♂ , 1 ♀ ; Jankoviči Krš , Duruk pećina; 42.364° N , 19.023° E ; 16 Apr. 2011 ; M. Komnenov leg.; CMK • 1 ♀ ; near river Rijeka Crnojevića , Žmrljevica pećina; 42.361° N , 19.050° E ; 16 Apr. 2011 ; M. Komnenov leg.; CMK • 1 ♂ ; Vrela , Pećina u Vrelima ; 42.3247° N , 18.9233° E ; 730 ma .s.l. ; 27 Oct. 2015 ; M. Pavlek leg.; CBSS ( Ar 4169 ) • 2 ♂♂ ; same collection data as for preceding; CBSS ( Ar 3669 ) • 2 ♂♂ , 1 ♀ ; Trnovo , Babatuša pećina; 42.291° N , 19.034° E [ Naumova et al. 2019 : 42.229° N, 19.096° E]; 15 Feb. 2010 ; M. Komnenov leg.; CMK • 1 ♂ (palps and chelicerae missing); same locality as for preceding, (“Bobotuša pećina”); 1932(?); J. Kratochvíl leg.(?); NMP • 1 ♂ , 2 ♀♀ ; Trnovo , Grbočica pećina; 42.288° N , 19.036° E ; 15 Apr. 2011 ; M. Komnenov leg.; CMK • 1 ♂ , 1 juv. ; same locality as for preceding; 9 Nov. 2015 ; A. Kirin leg.; CBSS ( Ar 3727 ) . – Bar • 2 ♂♂ , 2 ♀♀ ; Virpazar , Donja Seoca , Ivanina pećina; 42.228° N , 19.136° E ; 100 m a.s.l. ; 3 Apr. 2007 ; M. Komnenov leg.; CMK • 1 ♀ ; Seoca , Golubova pećina, hillside forest , under stones and sieving ; 42.208° N , 19.131° E ; 440 m a.s.l. ; 12 Aug. 2006 ; B. Petrov and S. Lazarov leg.; NMNHS . – Kolašin • 1 ♂ , 1 ♀ ; Monastery Morača ; 42.7648° N , 19.3931° E ; 300 ma .s.l. ; 8 May 2006 ; A. Schönhofer leg.; SMF . Figs 77–84. Stygopholcus montenegrinus Kratochvíl, 1940 . 77–78 . Males from Studenačka pećina, Montenegro (ZFMK (Ar 22201)) and from Zeze Cave, Albania (CMK), tips of left procursi, prolateral (slightly ventral) views; arrows point at distinctive prolateral sclerites. 79–82 . Male from Studenačka pećina (ZFMK (Ar 22201)), left genital bulb, retrolateral, ventral, prolateral-distal, and dorso-distal views. 83–84 . Female from Studenačka pećina (ZFMK (Ar 22201)), cleared female genitalia, ventral and dorsal views. Scale bars: 0.5 mm. Figs 85–90. Stygopholcusmontenegrinus Kratochvíl, 1940 ; epigyna, ventral views. 85-86 . Studenačka pećina, Montenegro (ZFMK (Ar 22201)). 87 . Golubinja pećina, Montenegro (CMK). 88 . Ljuboviči, Montenegro (CMK). 89 . Nëngurit Cave, Albania (CMK). 90 . Petnjik Cave, Montenegro (CMK). Scale bar: 0.5 mm (all at same scale). Figs 91–98. Stygopholcus montenegrinus Kratochvíl, 1940 ; male and female from Studenačka pećina, Montenegro (ZFMK (Ar 22201)). 91 . Male carapace and ocular area, frontal-dorsal view. 92 . Male gonopore and epiandrous spigots. 93 . Male clypeus and chelicerae, frontal view. 94 . Modified hairs frontally on male chelicerae. 95–96 . Stridulatory files on male and female chelicerae. 97 . Tarsal organ on male tarsus 1. 98 . Transparent process on genital bulb. Scale bars: 91, 93 = 100 µm; 92, 95–96 = 20 µm; 94, 97–98 = 10 µm. Figs 99–106. Stygopholcus montenegrinus Kratochvíl, 1940 ; male and female from Studenačka pećina, Montenegro (ZFMK (Ar 22201)). 99 . Tip of right procursus, retrolateral view. 100–101 . Tip of left procursus, prolateral and dorsal views; arrow points at distinctive prolateral sclerite. 102 . Hair-like structures on tip of procursus. 103 . Stridulatory pick on male palpal femur. 104 . Male palpal tarsal organ. 105 . Retrolateral bulbal process, distal view. 106 . Possible glandular pores on retrolateral bulbal process (detail of previous photo). Abbreviations: dp = dorsal process; ds = dorsal sclerite; rp = retrolateral process; vs = ventral sclerite. Scale bars: 99–101, 105 = 100 µm; 102 = 20 µm; 103–104, 106 = 10 µm. Figs 107–112. Stygopholcusmontenegrinus Kratochvíl, 1940 ; male andfemalefrom Studenačka pećina, Montenegro (ZFMK (Ar 22201)). 107 . Female prosoma; arrows point at humps opposing modified areas on abdomen. 108 . Female anterior lateral spinneret (ALS) and posterior median spinneret (PMS). 109 . Tip of female palp, dorsal view. 110 . Tip of female tarsus 1. 111 . Epigynum, ventral view. 112 . Indentations (muscle attachment sites) between epigynum and spinnerets. Scale bars: 107 = 200 µm; 108 = 10 µm; 109–110 = 20 µm; 111–112 = 100 µm. MONTENEGRO – unknown municipality • 3 ♀♀ prosomata and 2 ♀♀ abdomens; “Južna Crna Gora , neka pećina” (= southern Montenegro, some cave); date unknown; P. Dabović leg.; NMP . ALBANIA – Shkodra • 2 ♂♂ , 1 ♀ ; Galeria Iliriana , Tuneli i Gencit ; 42.032° N , 19.474° E ; 24 Nov. 2000 ; A. Zhalov leg.; NMNHS . Credible published records (not examined) MONTENEGRO – Cetinje • 5 ♂♂ , 5 ♀♀ , 1 juv. ; Trnovo vill., Babatuša pećina, “109a”; 42.291° N , 19.034° E ; 24 Mar. 2006 ; S. Lazarov and B. Petrov leg. [ Naumova et al. 2016 ] • 1 ♂ , 1 ♀ , 1 juv. , Cetinje town , unnamed cave above monastery, “102”; 42.39° N , 18.92° E ; 25 Mar. 2006 ; B. Petrov and S. Lazarov leg. [ Naumova et al. 2016 ] . Material assigned tentatively The material listed here originates from SE Montenegro (Podgorica) and NW Albania . It strongly resembles S. montenegrinus but shows some small though consistent differences (see section on variation below). The material is listed apart to draw attention to this potential separate species. MONTENEGRO – Podgorica • 1 ♂ , 1 ♀ ; cave in Moraća canyon [“grotje langs de Moraća, ~ 14 km from Titograd ”]; 42.53° N , 19.34° E ; 22 Jul. 1972 ; C. Deeleman leg.; NMNL (ARA 19038) • 3 ♂♂ , 2 ♀♀ , 4 juvs ; Peuta , Dučica pećina; 42.4621° N , 19.3307° E ; 19 Jul. 1972 ; C. Deeleman leg.; NMNL (ARA 19039) • 1 juv. ; Peuta , Pećina od Zavora ; 42.46° N , 19.33° E ; 19 Jul. 1972 ; C. Deeleman leg.; NMNL (ARA 19040) • 2 ♂♂ , 2 ♀♀ ; Kuči , Ubalac , Petnjik pećina; 42.520° N , 19.382° E ; 565 m a.s.l. ; 23 Mar. 2013 ; M. Komnenov leg.; CMK • 3 ♂♂ , 1 ♀ ; Kuči , Liješta Karantin pećina; 42.483° N , 19.395° E ; 575 m a.s.l. ; 23 Mar. 2013 ; M. Komnenov leg.; CMK • 2 ♀♀ ; Kučke Mt. , Kučka korita, Radeča , beech forest ; 42.500° N , 19.534° E ; 22 Jul. 2010 ; M. Komnenov leg.; CMK • 1 juv. assigned tentatively; Kučka korita, Jama J008; 42.4747° N , 19.5247° E ; 1350 m a.s.l. ; 10 Nov. 2015 ; A. Kirin leg.; CBSS ( Ar 4178 ) . ALBANIA – Shkodra • 1 ♂ ; K2 Cave nr. Kozhnje [Kozhnja]; 42.457° N , 19.607° E ; 30 Aug. 1996 ; “P.B.” leg.; NMNHS . – Kukës • 1 ♂ ; Hekurave Mt. , Qereç-Mulaj , Zeze Cave ; 42.362° N , 19.930° E ; 18 Aug. 2010 ; M. Komnenov leg.; CMK • 2 ♂♂ ; below Hekurave Mt. , Qereç-Mulaj , small cave near River Kuqit ; 42.364° N , 19.933° E ; 17 Aug. 2010 ; M. Komnenov leg.; CMK • 1 ♀ ; Prokletije , Curaj i Epërm , Nëngurit Cave ; 42.339° N , 19.927° E ; 19 Aug. 2010 ; M. Komnenov leg.; CMK . Redescription Male ( type locality; ZFMK (Ar 22201)) MEASUREMENTS. Total body length 5.5, carapace width 2.15. Distance PME–PME 150 µm ; diameter PME 130 × 160 µm ; distance PME–ALE 30 µm ; diameter AME 60 µm ; distance AME-AME 20 µm . Leg 1: 50.6 (13.7 + 0.8 + 13.3 + 19.6 + 3.2), tibia 2: 9.7, tibia 3: 8.0, tibia 4: 8.5; tibia 1 L/d: 65. COLOR (in ethanol). Carapace pale whitish to ochre-yellow; ocular area, clypeus, and posteriorly between furrows slightly darker (light brown); sternum light brown, with dark brown margins and irregular dark triangular radial marks; legs ochre yellow to light brown, without dark rings, without dark lines; abdomen mostly monochromous pale gray, only above spinnerets and ventrally in gonopore area some dark marks. BODY. Habitus as in Fig. 3 . Ocular area slightly raised. Carapace with deep median pit and pair of shallow furrows diverging from pit to posterior margin of carapace ( Fig. 91 ). Clypeus unmodified. Sternum wider than long (1.40/1.05), unmodified. Abdomen oval, pointed at spinnerets. CHELICERAE. Very similar to S. skotophilus (cf. Figs 51–52 ; see also Fig. 93 ), with pair of frontal apophyses set with 3 and 4 modified hairs, respectively, with 16 and 18 modified hairs each on frontal face ( Fig. 94 ); with stridulatory ridges ( Fig. 95 ). 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 ( Fig. 103 ); femur-patella joints shifted towards prolateral; tibia very large, with two trichobothria; tibia-tarsus joints shifted towards retrolateral; tarsal organ exposed ( Fig. 104 ); procursus dorsally with ~8 curved hairs (distal third curved ‘upwards’), with strong hump on prolateral side. Procursus tip ( Figs 77 , 99–102 ) with slightly projecting dorsal sclerite, strong hinged ventral sclerite (flat, wider in ventral than in lateral view), and membranous central part with brush of hair-like processes on retrolateral side, with narrow and weakly sclerotized process on prolateral side bordering ventral sclerite. Genital bulb ( Figs 79–82 ) with basal sclerite connecting to tarsus, small transparent dorsal process ( Fig. 98 ), and main sclerite consisting of retrolateral process and dorsal process; retrolateral process with bifid tip, distal part on ventral side thickened and heavily sclerotized, possibly with glandular pores distally ( Figs 105–106 ); 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 in distal third, prolateral row ~19 spines, retrolateral row ~24 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) Some specimens with more and larger dark marks dorsally on abdomen (posterior third), few specimens without abdominal marks; dark pattern on sternum slightly variable. Tibia 1 (N = 29): 11.3–14.5 (mean 12.9); tibia 2 (N = 35): 8.1–10.8 (mean 9.6); spines on femur 1 (N = 23): 24–53 (mean 40); modified hairs on cheliceral basis (N = 70): 11–22 (mean 14.7); modified hairs on cheliceral apophysis (N = 69): 2–4 (mean 3.0); chelicerae maximum width (N = 35): 0.72–0.84 (mean 0.79); procursus length (N = 35): 1.22–1.46 (mean 1.37); bulbal measures A–E: A (N = 17): 0.34–0.43 (mean 0.37); B (N = 17): 0.10–0.15 (mean 0.13); C (N = 17): 0.24–0.27 (mean 0.25); D (N = 17): 0.25–0.29 (mean 0.27); E (N = 8): 0.82– 0.90 (mean 0.88); epiandrous spigots (N = 25): 5–9 (mean 6.2). Males from SE Montenegro (Podgorica) and NW Albania differ slightly from other specimens, most conspicuously in bulbal measure C (smaller: 0.19–0.22). Female In general similar to male ( Fig. 3 ), but femur 1 without spines, with pair of indistinct processes posteriorly on carapace ( Fig. 107 ) and pair of opposing poorly visible plates on abdomen, distance between processes/plates: 1.1. Tibia 1 (N = 28): 8.3–15.3 (mean 12.9); tibia 2 (N = 30): 6.0–10.9 (mean 9.3); epigynum width (N = 31): 0.94–1.50 (mean 1.27); epigynum length (N = 31): 0.50–0.70 (mean 0.62); epigynal measure F (N = 31): 0.22–0.40 (mean 0.30). Epigynum anterior plate semicircular, weakly protruding, with weak transversal furrows ( Figs 85–88 , 111 ), with internal structures visible in uncleared specimens: round anterior structure, short sclerites originating from posterior margin and slightly converging anteriorly; posterior plate short but wide. Internal genitalia ( Figs 83–84 , 149–150 ) with distinctive ventral arc ( Fig. 155 ; approximately as long as wide), with large pore plates ( Fig. 84 ). Females from SE Montenegro (Podgorica) and NW Albania tend to have a slightly larger epigynal measure F: 0.37–0.43 (N = 6) (cf. Figs 89–90 ). Distribution Known from Montenegro and NW Albania ( Fig. 168 ).