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 Stygopholcus photophilus Senglet, 1971 Figs 4–5 , 9–11 , 113–144 , 151–152 , 156 Stygopholcus photophilus Senglet, 1971: 356 , figs 44–54. Hoplopholcus kratochvili – Brignoli, 1971: 259 , figs 1–6. — Deltshev 1985: 43 . Synonymized by Brignoli (1976) . Hoplopholcus photophilus – Brignoli 1976: 561 . Stygopholcus photophilus – Senglet 2001: 47 , 65, figs 15–20, 23–25, 28. Diagnosis Easily distinguished from known congeners by many characters, including: distinct dark marks on entire abdomen ( Figs 4–5 ), also ventrally; dark rings on femora and tibiae, dark lines on femora ( Figs 127–128 ); male chelicerae with strongly sculptured median margins ( Figs 130–131 ); larger number of modified hairs on male cheliceral base ( Fig. 162 ; with little overlap); male palpal femur distally more strongly widened ( Fig. 11 ); palpal tibia widest in distal half ( Fig. 11 ); procursus distally with strongly projecting dorsal sclerite ( Figs 113–114 ); dorsal bulbal process not divided into two separate tips ( Figs 115, 118 ); spines on male femur 1 in single row; male tibia 1 much longer than tibia 2 (tibia 1/tibia 2> 1.40; in other species <1.40; Fig. 165 ); strong transversal ridges on anterior part of epigynum ( Fig. 144 ); relatively longer epigynum (epigynum width/length: usually <1.7, in other species usually> 1.7, with little overlap; Fig. 163 ); female without stridulatory ridges on chelicerae ( Fig. 134 ), and without stridulatory apparatus between prosoma and abdomen. Type material of Stygopholcus photophilus Holotype GREECE – Thessaly • ♂ , examined; Larissa , Omolion (Tembe) [Omolio, Tempi Valley]; 39.900° N , 22.628° E ; 17 Jun. 1970 ; A. Senglet leg.; MHNG . Paratypes GREECE – Thessaly • 27 ♂♂ , 14 ♀♀ (3 vials), examined; same collection data as for holotype; MHNG . For further paratypes from other localities, see below. Type material of Hoplopholcus kratochvili Holotype GREECE – Epirus • ♂ , examined; Prosgoli [= Megálo Peristéri]; 39.723° N , 21.062° E ; date and collector unknown; ZMB ( 9077 ). Paratypes GREECE – Epirus • 1 ♂ , 1 juv. (2 vials), examined; same collection data as for holotype; ZMB ( 9077 ) • 1 ♂ , examined; same collection data as for holotype; MCVR . Other material examined GREECE – Thessaly • 2 ♂♂ , 3 ♀♀ ; Monte Ossa above Ampelakia , unnamed cave near church Profitis Elias , “The-76/13”; 39.862° N , 22.572° E ; 600 ma .s.l. ; 14 May 1976 ; B. Hauser leg.; MHNG • 3 ♂♂ , 2 ♀♀ ; same locality as for preceding, “Kri 78/27”; 10 Apr. 1978 ; B. Hauser leg.; MHNG • 1 ♂ ; Monte Ossa , near Spilia , Tsari Tripa cave on Mt. Psylodendron [= Spilaio Nymfon], “Kri-78/30”; 39.821° N , 22.641° E ; 11 Apr. 1978 ; B. Hauser leg.; MHNG • 2 ♀♀ ; ravine near Stomio (“ Ossa-Geb. , Stomio , Bachgraben ”), “TH 234”; 39.86° N , 22.72° E ; 22 Sep. 1995 ; K. Thaler and B. Knoflach leg.; ZFMK ( Ar 22202 ) . – Athos • 1 ♀ ; Dafni , “Ba 69-12”; 40.213° N , 24.223° E ; 10 ma .s.l. ; 4 May 1969 ; Ausobsky leg.; ZFMK ( Ar 22203 ) • 1 ♂ , 1 ♀ , 1 juv. ; Mount Athos , Agia Anna monastery , Lost Glasses Cave ; 40.14° N , 24.30° E ; 8 Sep. 2012 ; A. Zhalov leg.; NMNHS . – Central Macedonia • 1 ♂ ; Chalkididi , Sithonia , 1 km W of Zografou , humid stream valley with Osmunda , under logs and stones ; 40.1702° N , 23.8545° E ; 20 m a.s.l. ; 22 Aug. 2009 ; S. Huber and A. Schönhofer leg.; SMF . – Eastern Macedonia and Thrace • 2 ♂♂ , 5 ♀♀ ; Thasos , Panagia , Drakotrypa cave ; 40.732° N , 24.732° E ; 28 Dec. 1982 ; P. Beron and S. Andreev leg.; NMNHS • 1 ♂ ; Kavala , Zygos , Aghia Helleni cave [Spilaio Agias Elenis]; 41.018° N , 24.394° E ; 28 Dec. 1982 ; P. Beron and S. Andreev leg.; NMNHS . – Epirus • 4 ♂♂ , 2 ♀♀ ; Monodendri , dry valley before monastery, oak and hard-leafed forest, under stones and wood; 39.883° N , 20.749° E ; 1000 m a.s.l. ; 14 Aug. 2009 ; S. Huber and A. Schönhofer leg.; SMF . – Peloponnese • 1 ♂ ; Ag. Taxiarchi , 12 km Wof Leonidion , cellar of church ruin; 37.150° N , 22.760° E ; 29 Sep. 2006 ; H. Pieper leg.; ZFMK ( Ar 22204 ) • 6 ♂♂ , 7 ♀♀ , paratypes ; Laconia , Mystras / Sparti ; 37.067° N , 22.375° E ; 16 Aug. 1970 ; A. Senglet leg.; MHNG • 3 ♂♂ , 2 ♀♀ ; Laconia , Mystras ; 37.067° N , 22.375° E ; 23 May 1983 ; A. Senglet leg.; MHNG • 2 ♂♂ ; Kastoreio , Spilaio Skreti ; 37.1676° N , 22.3052° E ; 490 m a.s.l. ; 5 Dec. 2016 ; C. Cenni leg.; MCVR • 5 ♂♂ , 5 ♀♀ , 6 juvs , paratypes ; Messinia , Agios Dimitrios / Lefktro (“Leuktron”); 37.039° N , 22.102° E ; 20 Aug. 1970 ; A. Senglet leg.; MHNG • 1 ♀ , 4 juvs ; Arkadia , Kapsia , Spilaio Kapsias ; 37.623° N , 22.354° E ; 29 Sep. 1997 ; K. Paragamian leg.; HISR ( 1845 ) • 1 ♀ ; Arkadia , Leonidio , Spilaio Dionysou ; 37.1485° N , 22.8185° E ; 19 Sep. 1997 ; K. Paragamian leg.; HISR ( 1836 ) . – Crete • 10 ♂♂ , 15 ♀♀ , paratypes ; Rethymno , Choumerion / Pérama ; 35.355° N , 24.720° E ; 1 Jul. 1970 ; A. Senglet leg.; MHNG • 7 ♂♂ , 11 ♀♀ , 1 juv. , paratypes ; Rethymno , Keramoto / Pérama ; 35.333° N , 24.740° E ; 2 Jul. 1970 ; A. Senglet leg.; MHNG • 1 ♂ , 1 ♀ ; Rethymno , Archaia Eleftherna ; 35.324° N , 24.676° E ; 17 May 2006 ; H. Eikamp and U Kluge leg.; SMF ( 56590 ) • 1 ♂ ; Rethymno , Myloi , Spilaio Skoteinospilios ; 35.336° N , 24.505° E ; 1 Mar. 1987 ; K. Paragamian leg.; HISR ( 1640 ) • 1 ♂ , 1 ♀ ; Rethymno , Episkopi , along stream; 35.3417° N , 24.7656° E ; 230 m a.s.l. ; 7 Jul. 2015 ; B.A. Huber leg.; NHMC • 2 ♂♂ , 1 ♀ ; same collection data as for preceding; ZFMK ( Ar 22205 ) • 3 ♂♂ , 2 ♀♀ , in pure ethanol; same collection data as for preceding; ZFMK ( Cre19 ) • 2 ♂♂ , 2 juvs ; Irakleion , Zaros ; 35.13° N , 24.90° E ; 7 May 2009 ; K. Eckl and H. Eikamp leg.; SMF • 2 ♂♂ , 4 ♀♀ , 1 juv. ; Chania , Topolia , Spilaio Agias Sofias ; 35.417° N , 23.683° E ; 30 Dec. 2001 ; K. Paragamian leg.; HISR ( 2008 ) • 4 ♂♂ , 4 ♀♀ ; same locality as for preceding; 10 Oct. 1999 ; A. Senglet leg.; MHNG . – Central Greece • 2 ♂♂ , 5 ♀♀ ; Euboea , Artemisio / Istiaia ; 39.00° N , 23.22° E ( Senglet 2001 : 39.02° N, 23.25° E); 31 Aug. 1972 ; A. Senglet leg.; MHNG • 1 ♂ , 4 ♀♀ ; Euboea , Kato Steni ; 38.571° N , 23.825° E ( Senglet 2001 : 38.583° N, 23.850° E); 2 Sep. 1972 ; A. Senglet leg.; MHNG . – Ionian Islands • 3 ♂♂ , 4 ♀♀ , 2 juvs ; Lefkada , waterfalls 3 km Wof Nidri , rocky caves and mossy stones ; 38.7263° N , 20.6858° E ; 200 ma .s.l. ; 27 Mar. 2006 ; A. Schönhofer leg.; SMF • 1 ♂ , 1 ♀ ; Kefallonia , Lake Avithos , lakeside buildings ; 38.109° N , 20.528° E ; 28 May 1987 ; J. Murphy leg.; MMUE • 7 ♂♂ , 13 ♀♀ ; Corfu , Loutses ; 39.786° N , 19.880° E ; 20 Sep. 1972 ; A. Senglet leg.; MHNG • 1 ♂ ; Corfu , Sof Loutses, Spilaio Megali Grava / Peristerograva , on ground of cave, under stones ; 39.777° N , 19.890° E ; 350 m a.s.l. ; 22 Mar. 2006 ; A. Schönhofer leg.; SMF • 6 ♂♂ , 2 ♀♀ ; same locality as for preceding, (“Halbhöhle am Nordabhang des Pantokratorgebirges”), “Ko-60/11”; 19 Apr. 1960 ; B. Hauser leg.; MHNG • 6 ♂♂ , 13 ♀♀ ; Corfu , Pantocrator , Spilaio Katsampa (“Katsaba cave”) near Strinilas , “Pel74/19”; 39.744° N , 19.839° E ; 12 May 1974 ; B. Hauser leg.; MHNG . Figs 113–118. Stygopholcus photophilus Senglet, 1971 ; male from Episkopi, Crete (ZFMK (Ar 22205)). 113–114 . Left tarsus and procursus, prolateral and retrolateral views. 115–118 . Left genital bulb, retrolateral, ventral, prolateral-distal, and dorso-distal views; arrow points at weakly sclerotized process. Abbreviations: bs = basal sclerite; dp = dorsal process; ds = dorsal sclerite; rp = retrolateral process; vs = ventral sclerite. Scale bars: 0.5 mm. Figs 119–120. Stygopholcus photophilus Senglet, 1971 ; male from Episkopi, Crete (ZFMK (Ar 22205)), female from Choumerion/Pérama, Crete (MHNG). 119–120 . Male chelicerae, frontal and lateral views. 121–122 . Cleared female genitalia, ventral and dorsal views. Scale bars: 0.5 mm. Figs 123–128. Stygopholcus photophilus Senglet, 1971 ; epigyna, ventral views, and details of male femora 2 and 3, dorsal views. 123 . Thessaly, Omolio, Tempi Valley (MHNG). 124 . Corfu, Loutses (MHNG). 125 . Peloponnese, Agios Dimitrios/Lefktro (MHNG); arrows point at internal sclerites. 126– 128 . Crete, Episkopi (ZFMK (Ar 22205)). Scale bars: 123–126 = 0.5 mm; 127–128 = 0.2 mm. Figs 129–136. Stygopholcus photophilus Senglet, 1971 ; male and female from Choumerion/Pérama, Crete (MHNG). 129 . Male clypeus, frontal view; arrow points at modified median area. 130 . Male chelicerae, frontal view. 131 . Modified area medially on male chelicerae. 132 . Modified hairs frontally on male chelicerae. 133 . Stridulatory files on male chelicerae. 134 . Absence of stridulatory files on female chelicerae. 135 . Male ALS. 136 . Female anterior lateral spinneret (ALS) and posterior median spinneret (PMS). Scale bars: 129–130 = 100 µm; 131, 134–136 = 20 µm; 132–133 = 10 µm. Figs 137–144. Stygopholcus photophilus Senglet, 1971 ; male and female from Choumerion/Pérama, Crete (MHNG). 137–138 . Tip of left procursus, retrolateral (slightly ventral) and ventral views. 139 . Detail of previous figure. 140 . Dorsal bulbal process and transparent bulbal process (arrow); note hair-like structures on dorsal bulbal process. 141 . Female palpal tarsal organ. 142 . Tarsal organ on female tarsus 4. 143 . Male gonopore and epiandrous spigots. 144 . Anterior part of epigynum showing strong ridges; arrows point at pair of anterior bulges. Abbreviations: ds = dorsal sclerite; vs = ventral sclerite. Scale bars: 137–138, 140, 143–144 = 100 µm; 139, 141–142 = 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 157–160. Bulbal measures B–E for the three species of the ‘northern clade’ of Stygopholcus Kratochvíl, 1932 (cf. Figs 15–18). Bulbal measure Ais strongly correlated with measure Cand 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 Dis inapplicable). Asterisk in Fig. 160 = dubious specimen from Jama na Brezdanu (see text). ALBANIA – Gjirokastër • 1 ♂ , 1 ♀ ; Përmet , near Petran , artificial gallery; 40.208° N , 20.419° E ; 12 May 1995 ; B. Petrov and R. Stoev leg.; NMNHS . Credible published records (not examined) GREECE – Epirus • 1 ♀ ; 11 km SE of Konitsa , in gorge of river Voithomatis [“Bogsomatis”, side river of Aoos river], “Ep-73/48”; 40.019° N , 20.785° E ; 30 Apr. 1973 ; V. Mahnert leg.; ( Brignoli 1976 ); MHNG (?) . ALBANIA – (the following material could not be found, C. Deltshev pers. comm., 30 Sep. 2020 ) Durrës • ♂ ♀ (not specified); Durrës ; 41.3° N , 19.5° E ; 26 May 1993 ; P. Stoev and D. Zapryanova leg. ( Deltshev et al. 2011 ) . – Korcë • ♂ ♀ (not specified); Cave 2 near Leskovik ; 40.150° N , 20.600° E ; 1 Jun. 1994 ; T. Ivanova leg. ( Deltshev et al. 2011 ) . – Gjirokastër • ♂ ♀ (not specified), Dragot , Vijoshe River ; 40.288° N , 20.069° E ; 1 Jun. 1994 ; G. Blagoev leg. ( Deltshev et al. 2011 ) . 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 . Unidentified and unspecified localities GREECE • 1 ♀ ; Crete , “Chassia” [misspelling on copied label?], “RII/8”; May 1926 ; Roewer leg.; SMF • 1 ♂ , 1 ♀ , paratypes ; “7002”; no further data; MNHN ( Ar 10230 ) • 1 ♂ ; Peloponnese , “Mühlenbach”; 6 Oct. 1980 ; Malicky leg.; SMF . Redescription Male ( Crete ; ZFMK (Ar 22205)) MEASUREMENTS. Total body length 4.3, carapace width 1.5. Distance PME–PME 130 µm ; diameter PME 120 × 140 µm ; distance PME–ALE 30 µm ; diameter AME 60 µm ; distance AME-AME 30 µm . Leg 1: 40.4 (10.7 + 0.7 + 10.5 + 16.0 + 2.5), tibia 2: 6.9, tibia 3: 5.1, tibia 4: 5.7; tibia 1 L/d: 75. Fig. 167. Known distribution of Stygopholcus Kratochvíl, 1932 . For the ‘northern clade’, see Fig. 168. Abbreviations: Alb = Albania; BaH = Bosnia and Herzegovina; Cro = Croatia; Gre = Greece; Mon = Montenegro. COLOR (in ethanol). Carapace pale whitish to ochre-yellow, ocular area and clypeus slightly darker (light brown), ocular area posteriorly with darker brown V-mark; sternum very dark brown, almost black; legs ochre yellow, with darker (light brown) rings on femora (subdistally) and tibiae (proximally and subdistally), with short dark longitudinal lines dorsally on femora ( Figs 127–128 ); abdomen ochregray, with distinct dorsal pattern consisting of brown band around heart and dorsal and lateral marks in posterior half; with white internal marks; ventrally with wide dark brown band, interrupted at two thirds between gonopore and spinnerets. BODY. Habitus as in Fig. 4 . 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 with very indistinct modified area medially (arrow in Fig. 129 ) and sclerotized margin. Sternum wider than long (1.05/0.70), unmodified. Abdomen oval to short cylindrical. CHELICERAE. As in Figs 119–120 , with pair of frontal apophyses set with 2 and 3 modified hairs, respectively, with 23 modified hairs each on frontal face ( Figs 130, 132 ); with strongly sculptured median margins ( Fig. 131 ); with stridulatory ridges ( Fig. 133 ). PALPS. As in Figs 9–11 ; coxa with prominent retrolateral hump; trochanter barely modified; femur with distinct retrolateral process proximally, with stridulatory pick (modified hair) proximally on prolateral side, with transversal dark line retrolaterally, distally strongly widened and with rounded ventral protrusion; femur-patella joints shifted towards prolateral; tibia very large, thickest at two thirds, with two trichobothria; tibia-tarsus joints shifted towards retrolateral; procursus dorsally with ~5 weakly curved hairs, with strong hump on prolateral side. Procursus tip ( Figs 113–114 , 137–138 ) with scoop-shaped dorsal sclerite, strong hinged ventral sclerite (triangular in ventral view), and complex membranous central part with brush of hair-like processes on retrolateral side. Genital bulb ( Figs 115–118 ) with basal sclerite connecting to tarsus, small dorsal process weakly sclerotized, and main sclerite consisting of retrolateral process and dorsal process; retrolateral process distally widening, distal margin with pair of heavily sclerotized bulges on ventral side; dorsal process with single slightly curved tip, proximally with two small apophyses. Fig. 168. Known distribution of the ‘northern clade’ of Stygopholcus Kratochvíl, 1932 . Pale blueand pale orange marks denote specimens that are assigned tentatively to the respective species. Abbreviations: Alb = Albania; BaH = BosniaandHerzegovina; Cro = Croatia; Mon = Montenegro. LEGS. With ~12 short spines in one row ventrally on femur 1; without curved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with irregular platelets rather than with distinct pseudosegments. Male (variation) Color pattern consistent but individual elements slightly variable in size and distinctness; median section of ventral abdominal band sometimes divided into three parallel longitudinal bands; dark abdominal marks sometimes divided into many small round dots. Tibia 1 (N = 104): 9.7–15.3 (mean 12.7); tibia 2 (N = 41): 6.5–10.4 (mean 8.4); spines on femur 1 (N = 11): 0–34 (mean 19); modified hairs on cheliceral basis (N = 50): 19–44 (mean 30); modified hairs on cheliceral apophysis (N = 50): 3–4 (mean 3.3); chelicerae maximum width (N = 35): 0.60–0.81 (mean 0.70); procursus length (N = 35): 1.22–1.50 (mean 1.36). Female In general similar tomale ( Fig. 5 ), but femur 1 without spines, clypeus unmodified. Without modifications posteriorly on carapace and anteriorly on abdomen. Tibia 1 (N = 91): 8.9–13.7 (mean 11.2); tibia 2 (N = 46): 5.3–9.5 (mean 7.2); epigynum width (N = 25): 0.98–1.34 (mean 1.16); epigynum length (N = 25): 0.62–0.86 (mean 0.72); epigynal measure F (N = 25): 0.22–0.38 (mean 0.29). Epigynum anterior plate roughly triangular, weakly protruding, with very strong transversal ridges in anterior part ( Fig. 144 ); with internal structures visible in uncleared specimens: oval to drop-shaped anterior structure, short parallel (anteriorly slightly converging) sclerites posteriorly; posterior plate short but wide. Internal genitalia (Figs 121–122, 151–152) with distinctive ventral arc ( Fig. 156 ; with several transversal, crescent-shaped sclerites and slightly elongate ventral pocket), with large pore plates (Fig. 122). Distribution Widely distributed in Greece (including Crete and the Ionian Islands ) and southern Albania ; the species seems to be largely absent from the Aegean Islands (except Thasos) ( Fig. 167 ).