Stygopholcusmontenegrinus 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 : 33-41

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 .

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 ).

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 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. Stygopholcus montenegrinus 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. Stygopholcus montenegrinus Kratochvíl, 1940; male and female from 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.

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.

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 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.