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The Dinaric Karst is a global hotspot for subterranean diversity, with two distinct peaks of species richness in the northwest and southeast, and an area of a lower species richness in the central part. In this article, we present a species list and describe the ecological conditions of the Lukina jama–Trojama cave system, located in the central part of the Dinaric Karst. This cave system is the deepest and one of the most logistically challenging cave systems sampled so far in the Dinaric Karst. Repeated sampling resulted in a list of 45 species, including 25 troglobionts, 3 troglophiles, 16 stygobionts, and 1 stygophile. Most of the recorded species are endemic to the Velebit Mountain, while three species are endemic to the Lukina jama–Trojama cave system. Within the system, species richness peaks in the deepest third of the cave, most likely reflecting the harsh ecological conditions in the upper parts, including ice, cold winds, and occasional waterfalls. Milder and more stable deeper parts of the cave contain a rich subterranean species community, part of which is associated with two very distinct aquatic habitats, the cave hygropetric and the phreatic zone. The newly recognized hotspot of subterranean biodiversity in the central Dinaric Karst, which has emerged between the two known centers of biodiversity, further highlights the species richness in large cave systems, but also challenges the diversity patterns in the Dinaric Karst overall.

In 2020 and 2021, biospeleological surveys were carried out in the wider area of Baraćeve špilje (eng. Barać caves), in cooperation with The Public Institution for the Management of Protected Natural Values on the Territory of the Municipality of Rakovica. The main goal was to make an inventory of the subterranean fauna in selected caves, and to conduct a two-year biospeleological monitoring of the Gornja Baraćeva špilja cave. We have undertaken eight field trips, one in each season (spring, summer, autumn and winter) over two years. In addition to the inventory of subterranean fauna, we measured microclimatic conditions, photographed interior of the caves and cave entrances, and collected, identified, and photographed fauna. In 2020, we investigated Božićeva špilja, Dumenčića špilja, Donja Baraćeva špilja and Gornja Baraćeva špilja caves, while in 2021 research was conducted in Velesova Jama, Jama pod Guvnom, Kunina špilja, cave system Ponorac-Jovina caves and biospeleological monitoring of Gornja Baraćeva špilja continued. With this research, we obtained the first lists of subterranean fauna for selected caves. By systematically studying the Gornja Baraćeva špilja cave, we updated its list of fauna which now contains 46 taxa—13 taxa more than were recorded in the literature before we started our research. The species Machaerites pavleki Hlaváč & Jalžić, 2010 was found for the first time in thirteen years, both in Donja Baraćeva špilja and in Gornja Baraćeva špilja. This two-year research was of great importance because it shows that interesting new fauna can be found even in well-studied caves with systematic approach.

Croatian Biospeleological Society and Biology Students Association – BIUS conducted biospeleological research at the Žumberak – Samobor hills Nature Park in 2020 and 2021. From a geotectonic point of view, this protected area is very interesting – it is situated at the contact zone between structural complexes of the Inner Dinarids and central Dinaric carbonate platform. The massif continues in southeast Slovenia under the name Gorjanci. More than 150 caves and pits are known from this massif (including the Slovenian part). Most of the biospeleological data concerning Žumberak and Samobor hills are scattered in technical reports, occasional publications or in abstracts of scientific conferences. Inventory of biodiversity of two groups (Coleoptera and Amphipoda) are more represented in scientific papers. Nevertheless, all literature data indicate diverse and unique subterranean fauna. This was also confirmed by our research, in which we recorded numerous taxa of Gastropoda, Arachnida, Crustacea, Myriapoda, Collembola and Coleoptera. From a speleological and biospeleological perspective several caves stand out: Provala (2161 m long and 55 m deep) and Bedara (1593 m long and 113 m deep) as the largest speleological objects in the Nature Park, together with Drobovnik and Pušina. All four caves have both dry passages and flowing water parts, with diverse troglobiotic and stygobiotic fauna. Drobovnik is a type locality for endemic coleopteran species Machaerites curvistylus Nonveiller & Pavićević, 2001. It is also very important to mention the Jamina pit, in which a dense population of Natura 2000 species Leptodirus hochenwartii Schmidt, 1832 was recorded. Thus, this pit needs to be regularly monitored. Furthermore, it is necessary to systematically investigate other, smaller caves and pits of Žumberak – Samobor hills Nature Park, which are also faunistically interesting.

The subterranean ecosystem exerts strong selection pressures on the organisms that thrive in it. In response, obligate cave-dwellers have developed a series of morphological, physiological, and behavioral adaptations, such as eye reduction, appendage elongation, low metabolic rates or intermittent activity patterns, collectively referred to as troglomorphism. Traditionally, studies on cave organisms have been hampered by the difficulty of sampling (i.e., small population sizes, temporal heterogeneity in specimen occurrence, challenges imposed by the difficult-to-access nature of caves). Here, we circumvent this limitation by implementing a museomics approach. Specifically, we aim at comparing the genetic population structures of five cave spider species demonstrating contrasting life histories and levels of troglomorphism across different caves in the northern Dinarides (Balkans, Europe). We applied a genome-wide hybridization-capture approach (i.e., HyRAD) to capture DNA from 117 historical samples. By comparing the population genetic structures among five species and by studying isolation by distance, we identified deeper population structuring and more pronounced patterns of isolation by distance in the highly troglomorphic Parastalita stygia and Stalita pretneri ground dwellers, while the three web-building Troglohyphantes species, two of which can occasionally be found in surface habitats, showed less structured populations compatible with higher dispersal ability. The spatial distribution of genetic groups revealed common phylogeographic breaks among lineages across the studied species, which hint at the importance of environmental features in driving dispersal potential and shaping underground diversity.

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 ��� Kratochv��l 1934: 181 (part; see Remarks below). Hoplopholcus absoloni ��� Brignoli 1971: 257, fig. 20. Stygopholcus absoloni ��� Kratochv��l 1932: 2; 1940: 10, figs 3, 4a���b, 5a���b, 7 (left); 1978: 28. ��� Absolon & Kratochv��l 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 ��� Kratochv��l 1934: 181 (part; see Remarks below); 1935: 11 (see Remarks below). Remarks For most of the localities listed in Kratochv��l (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). Kratochv��l���s (1934) ���Pec kod Ljubova��� [Ljubovo?] is not mentioned in the extensive list of localities in Kratochv��l (1940) and is thus not further considered herein. By contrast, all caves listed in Kratochv��l (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, Bjelu��ica, 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, Bjelu��ica (���Bjelu��ica pećina���); 42.8447�� N, 17.9786�� E; 350 ma.s.l.; 15���16 Jul. 1932; J. Kratochv��l 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. Kratochv��l 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. Kratochv��l leg.; NMP ��� 1 ♂, 1 ♀, with Kratochv��l���s original label ��� Stygopholcus Absoloni (Kulcz.), ♀ et ♂ type! 19 b��� [these are probably Kratochv��l���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-H��hle, leg. Kratochv��l 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. Kratochv��l leg.(?); NMP ��� 1 ♂, 1 ♀, 5 juvs; Sof Zavala, near Belenići, Bezdan pod Licem; 42.813�� N, 18.000�� E; 25 Jul. 1932; J. Kratochv��l 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. R��my and A. Schweitzer leg.; MNHN (Ar 10219) ��� 1 ♀, 3 juvs; Kovina pećina; 43.250�� N, 17.935�� E; 22 Aug. 1936; P. R��my 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 ��pilja 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 ��pilja (= 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, Bale��ina 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 ��pilja 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, Drinovču��a, 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 Turku��a; 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 ♀; Omi��, Kučiće, Nikolina ��pilja; 43.4253�� N, 16.8271�� E; 250 m a.s.l.; 23 Mar. 2003; M. Cukrov leg.; CBSS (Ar 2003) ��� 2 ♂♂, 1 ♀, 1 juv.; Omi��, 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. Sch��nhofer leg.; SMF ��� 2 ♂♂, 2 ♀♀; ��estanovac, 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 ��pilja u Solunu; 43.4589�� N, 17.2217�� E); date unknown; J. Kratochv��l 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.; ��estanovac, Svagu��e, Mužina ��pilja; 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 ��pilja 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, ��pilja 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 ��pilja; 43.3865�� N, 17.0199�� E; 360 ma.s.l.; 24 Nov. 2005; M. Pavlek leg.; CBSS (Ar 2005) ��� 1 ♂; Zagvozd, Čike��i, Jamau Čike��ima; 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, ��pilja u Gaju; 43.383�� N, 17.041�� E; 18 Oct. 2002; C. Deeleman leg.; CBSS (Ar 1993) ��� 2 ♂♂; Zagvozd, Prodani, Pružina ��pilja; 43.394�� N, 17.059�� E; 520 m a.s.l.; 15 Oct. 2002; C. Deeleman leg.; CBSS (Ar 1926) ��� 1 ♀; Zagvozd, Spilja near Katu��ići; 43.384�� N, 17.055�� E; 18 Apr. 1974; C. Deeleman leg.; NMNL (ARA 19012) ��� 1 ♀, 1 juv.; Zagvozd, Župa, Brikinjava ��pilja; 43.3452�� N, 17.0969�� E; 740 ma.s.l.; 17 Nov. 2005; M. Pavlek leg.; CBSS (Ar 2000) ��� 1 ♂; Zagvozd, Župa, Matija��eva 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 ��pilja; 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.; Ba��ka 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 Vla��ka, 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. Ćuku��ić leg.; CBSS (Ar 3370) ��� 1 ♀; same locality as for preceding; 24 Jun. 2017; M. Pavlek leg.; CBSS (5035) ��� 1 ♂, 2 ♀♀, 1 juv.; Tučepi, Ravna Vla��ka, 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 ��plija; 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, ��o��ići, 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, ��krapica, 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, Zastraži��če, Vela Stiniva cove, ��pilja 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 ��pilja; 43.1341�� N, 16.7537�� E; 200 m a.s.l.; 3 May 2011; A. Ćuku��ić leg.; CBSS (Ar 2512) ��� 1 ♂, 1 ♀ (epigynum missing); same locality as for preceding, ���Grabćina ��plija���; 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, ��olkina jama; 42.9665�� N, 17.6625�� E; 230 m a.s.l.; 27 Feb. 2016; B. Jalžić leg.; CBSS (Ar 3989) ��� 2 ♀♀; Zažablje, Dobranje, Kornjatu��a, 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 ♀; Pelje��ac, Ston, Ponikve, ��pilja 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 ♀; Pelje��ac, 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 ♀; Pelje��ac, Janjina, Vardi��ta, ��pilja 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 ♂; Pelje��ac, Orebić, Oskoru��no, ��pilja kod Zagruda u vinogradu; 42.9666�� N, 17.3227�� E; 295 m a.s.l, Published as part of Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, pp. 1-60 in European Journal of Taxonomy 752 on pages 12-22, DOI: 10.5852/ejt.2021.752.1391, http://zenodo.org/record/4958808, {"references":["Kulczynski W. 1914. Aranearum species novae minusve cognitae, in montibus Kras dictis a Dre C. Absolon aliisque collectae. Bulletin international de l'Academie des Sciences de Cracovie 1914: 353 - 387, pl. 16.","Kratochvil J. 1934. Liste generale des Araignees cavernicoles en Yugoslavie. Pregled pecinskih paukova u Jugoslaviji. Prirodoslovne Razprave 2: 165 - 226.","Brignoli P. M. 1971. Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae). Mitteilungen aus dem Zoologischen Museum in Berlin 47: 255 - 267. https: // doi. org / 10.1002 / mmnz. 1971047020","Kratochvil J. 1932. Prispevek k poznani Araneid stredni Slavonie. Sbornik Vysoke Skoly Zemedelske, Brno (C) 23: 1 - 16.","Kratochvil J. 1940. Etude sur les araignees cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1 - 26.","Absolon K. & Kratochvil J. 1932. Zur Kenntnis der hohlenbewohnenden Araneae der illyrischen Karstgebiete. Mitteilungen uber Hohlen- und Karstforschung 1932: 73 - 81.","Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grece. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345 - 359. https: // doi. org / 10.5169 / seals- 401666","Eberle J., Dimitrov D., Valdez-Mondragon A. & Huber B. A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https: // doi. org / 10.1186 / s 12862 - 018 - 1244 - 8","Kratochvil J. 1935. Araignees cavernicoles de Krivosije. Prace Moravske Prirodovedecke Spolecnosti 9: 1 - 25.","Ruzicka V., Kurka A., Buchar J. & Rezac M. 2005. Czech Republic - the type material of spiders (Araneae). Casopis Narodniho muzea, Rada prirodovedna 174: 13 - 64."]}

Stygopholcus skotophilus Kratochv��l, 1940 Figs 2, 6���8, 43���76, 147���148, 154 Stygopholcus skotophilus Kratochv��l, 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) ��� Kratochv��l 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 Babatu��a 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 Kratochv��l���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 Dvori��kog 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 [Kratochv��l 1935: ���Mijukovica���]; 42.57�� N, 18.67�� E; 1932(?); J. Kratochv��l 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. Kratochv��l leg.; NMNL (ARA 19030) ��� 1 ♀; Risan, Sopot pećina; 42.5137�� N, 18.6816�� E; 13 Jul. 1971; C. Deeleman leg; NMNL (ARA 19031). ��� Nik��ić ��� 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, ��irbaba 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. ��kufca 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, Sto��ka 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, ��okova 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. Kratochv��l leg.; NMNL (ARA 19025���6) ��� 3 ♂♂, 3 ♀♀; Trebinje, Ilijin vrh, Narančivi, Mitru��ica 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. Kratochv��l 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 (���Eliash��hle���); 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. Kratochv��l 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, Drame��ina, 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, ��krabuljica, 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, ��pilja 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 ��pilja; 42.5066�� N, 18.3913�� E; 205 m a.s.l.; 6 Apr. 2010; R. Ozimec and A. Sch��nhofer leg.; ROC (R260). Localities unknown or unspecified ��� 1 ♂, 1 ♀; ��� Herzegowina, Tartarush��hle ���; Verhoeff leg.; ZMB (9073) ��� 1 ♂, 1 juv.; ���c 86, 1935��� [no further data]; NMP ��� 1 juv.; ��� Wolfsh��hle, S��dherzegowina ���, ���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. Kratochv��l leg.(?) [Kratochv��l 1940] ��� 1 juv.; 2 km WČora pećina, Matja��evicapećina; 42.595�� N, 18.700�� E [Naumova et al. 2019: 42.569�� N, 18.662�� E]; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940]; NMP ��� ♂ ♀ (?), material apparently lost; Dragaljsko polje, near Dvori��ko ždrijelo (���Dvorečko zdrielo���), Tomova pećina; 42.603�� N, 18.675�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Jankovo selo, Pećina na Jankovom vrhu; 42.590�� N, 18.658�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; 4 km Nof Pećina na Jankovom vrhu, Pećina za Jankovim vrhom; 42.620�� N, 18.656�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near road Crkvice-Veli vrh, cave near Crkvice; 42.560�� N, 18.630�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Crkvice, 1 km Wof Blagojević, Pećina kod Blagojevića; 42.573�� N, 18.620�� E; 1932(?), J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Poljana, Pećina kod ��unjevca; 42.577�� N, 18.648�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Knezlac, near Izeta pećina, Studena pećina; 42.544�� N, 18.689�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; Pokljuka donja [Kratochv��l 1935: ���dolnja���]; 4 km SE Knezlac; 42.542�� N, 18.693�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Napode, near Risan-Crkvice road, Vilina (��� Vilna ���) pećina; 42.5546�� N, 18.6624�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 1940]. ��� Nik��ič ��� ♂ ♀ (?), material apparently lost; near Jabukovac, Leskova pećina; 42.83�� N, 18.47�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost; near Grahovac, on Brdo Omutič mountain, Vodena pećina; 42.687�� N, 18.617�� E; 1932(?); J. Kratochv��l leg.(?) [Kratochv��l 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. Kratochv��l leg.(?) [Kratochv��l 1940] ��� ♂ ♀ (?), material apparently lost, Elazov d&oc, Published as part of Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, pp. 1-60 in European Journal of Taxonomy 752 on pages 22-32, DOI: 10.5852/ejt.2021.752.1391, http://zenodo.org/record/4958808, {"references":["Kratochvil J. 1940. Etude sur les araignees cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1 - 26.","Kulczynski W. 1914. Aranearum species novae minusve cognitae, in montibus Kras dictis a Dre C. Absolon aliisque collectae. Bulletin international de l'Academie des Sciences de Cracovie 1914: 353 - 387, pl. 16.","Kratochvil J. 1935. Araignees cavernicoles de Krivosije. Prace Moravske Prirodovedecke Spolecnosti 9: 1 - 25.","Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grece. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345 - 359. https: // doi. org / 10.5169 / seals- 401666","Brignoli P. M. 1971. Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae). Mitteilungen aus dem Zoologischen Museum in Berlin 47: 255 - 267. https: // doi. org / 10.1002 / mmnz. 1971047020","Naumova M., Lazarov S. P., Petrov B. P. & Deltshev C. D. 2016. New faunistic data on the cave-dwelling spiders in the Balkan Peninsula (Araneae). Ecologica Montenegrina 7: 425 - 438.","Eberle J., Dimitrov D., Valdez-Mondragon A. & Huber B. A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https: // doi. org / 10.1186 / s 12862 - 018 - 1244 - 8","Naumova M., Lazarov S. & Deltshev C. D. 2019. Faunistic diversity of the spiders in Montenegro (Arachnida: Araneae). Ecologica Montenegrina 22: 50 - 89. https: // doi. org / 10.37828 / em. 2019.22.5","Ruzicka V., Kurka A., Buchar J. & Rezac M. 2005. Czech Republic - the type material of spiders (Araneae). Casopis Narodniho muzea, Rada prirodovedna 174: 13 - 64.","Lukic M., Delic T., Pavlek M., Deharveng L. & Zagmajster M. 2020. Distribution pattern and radiation of the European subterranean genus Verhoeffiella (Collembola, Entomobryidae). Zoologica Scripta 49: 86 - 100. https: // doi. org / 10.1111 / zsc. 12392","Senglet A. 2001. Copulatory mechanisms in Hoplopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxonomic data. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 74: 43 - 67. https: // doi. org / 10.5169 / seals- 402796"]}

Genus Stygopholcus Kratochvíl, 1932 Stygopholcus Kratochvíl, 1932: 2–3 (type species: Holocnemus absoloni Kulczyński, 1914; see below). Stygopholcus – Absolon & Kratochvíl 1932: 75. — Kratochvíl 1934: 182; 1940: 8; 1978: 27 — Senglet 1971: 354; 2001: 58, 65. — Brignoli 1971: 257; 1976: 561. Authority of Stygopholcus There has long been disagreement about the publication that has made the genus name Stygopholcus validly available. Some cataloguers such as Roewer (1942) and Brignoli (1983) considered the original description to be in Kratochvíl (1940), while Bonnet (1958) cited Absolon & Kratochvíl (1932) as the source for the name. Platnick (1993) first opted for Kratochvíl (1940), later (Platnick 2000) for Absolon & Kratochvíl (1932). The World Spider Catalog (2020) adopted Platnick's later view. A third option, namely Kratochvíl (1932), was offered by Kratochvíl himself (Kratochvíl 1940), and adopted by Senglet (1971). The name Stygopholcus is first mentioned in two papers in 1932: in one of these, Absolon & Kratochvíl (1932) explicitly refer to the other publication (Kratochvíl 1932) as the source of the name: "Mit dem Namen Stygopholcus n. g. hat J. Kratochvíl (siehe "Sur quelques Araignées de Slavonie central", Bulletin de l'Institut national agronomique, Sign. C. 23, Brno 1932) die Art Holocnemus (Hoplopholcus) absoloni Kulcz. als selbständige Gattung abgetrennt." [" With the name Stygopholcus n. g., J. Kratochvíl (see "Sur quelques Araignées de Slavonie central", Bulletin de l'Institut national agronomique, Sign. C. 23, Brno 1932) has moved the species Holocnemus (Hoplopholcus) absoloni Kulcz. to constitute a separate genus"]. Other than this statement, Absolon & Kratochvíl (1932) just say a few words about the habitat of S. absoloni. Absolon & Kratochvíl (1932) thus fail to satisfy a requirement of ICZN (1999) Article 13.1.1 for names published after 1930 to be available: "… be accompanied by a description or definition that states in words characters that are purported to differentiate the taxon." The fact that Absolon & Kratochvíl (1932) contains a bibliographic reference to such a published statement could be construed as satisfying ICZN Article 13.1.2. We do not know which 1932 paper was published first. However, if Kratochvíl (1932) was published first, it has priority for that reason; if it was published second, it has priority because Absolon & Kratochvíl (1932) refer to a statement that was not yet published at the time. Confusingly, the paper to which Absolon & Kratochvíl (1932) refer was originally also not intended to be the source of the name. Kratochvíl (1932) referred to " Stygopholcus n. g. in lit., jejž popíši na jiném mistě" [" Stygopholcus n.g. in lit., that Iwill describe elsewhere"], suggesting that the genus name was intended to be made available in an upcoming publication (possibly Kratochvíl 1934). However, Kratochvíl (1932) provides "characters that are purported to differentiate the taxon", satisfying ICZN Article 13.1.1; and he explicitly names " H. absoloni Kulcz. " as the only species to be included in the genus, satisfying the requirements of ICZN Article 13.3 regarding the fixation of a type species. From a formal perspective, Kratochvíl's (1932) intentions (of making the name available later) are as irrelevant as the confusion between genus rank (in the Czech text) and subgenus rank (in the French summary), and the minimalist nature of his diagnosis: he mentions that (1) the female sternum process present in Holocnemus pluchei is absent in Stygopholcus; (2) the abdomen shape differs between Holocnemus and Stygopholcus; (3) the female palp of Stygopholcus resembles that of Pholcus rather than that of Holocnemus pluchei (which is enlarged); and (4) Stygopholcus has cheliceral stridulation like Holocnemus (in contrast to Pholcus). We conclude that Stygopholcus was made available by Kratochvíl (1932). Identity of Stygopholcus absoloni (Kulczyński, 1914) In the original description of " Holocnemus (Hoplopholcus) Absolonii ", Kulczyński (1914) mentions two caves near Trebinje by name: "Mares et feminae lecta sunt prope Trebinje in speluncis: Ilijina pećina … et Zovica jama…" [" males and females were collected in caves near Trebinje: Ilijina cave … and Zovica pit … "]. He then notes that according to Absolon ("Teste Cel. C. Absolon"), this species has been found in many other caves in southeastern Herzegovina (southern region of modern Bosnia and Herzegovina). This strongly suggests that he had only seen the specimens from the two caves near Trebinje. Any specimens he had from these caves, and only these specimens, constitute the type series (syntypes). In Kulczyński's (1914) long original text, the morphology is described in considerable detail but complex shapes are notoriously difficult to understand from text alone. However, he was obviously aware of differences among the specimens available to him, starting a paragraph with "Maris palpi variant paululo formâ, etiam mandibularum armatura paulo mutabilis" [" the male palps vary slightly in shape, and also the armature of the chelicerae is slightly variable "]. He clearly interpreted this as intraspecific variation, but without specifying to which variant(s) his four figures referred. When Kratochvíl (1940) made a first careful and detailed revision of Stygopholcus, he realized that Kulczyński's (1914) figures represented the two variants, which Kratochvíl interpreted (correctly) as different species. He considered Kulczyński's (1914) fig. 25 to represent one species, figs 24, and 26– 27 the other species. This view was accepted by subsequent cataloguers and authors, also by Senglet (1971, 2001), the only author other than Kratochvíl who studied Stygopholcus in some detail. Since Kulczyński's (1914) figures of the male palps are in different views, we checked Kratochvíl's (1940) distinction of the two putative species by orienting palps of both species in exactly the same positions as in Kulczyński's (1914) figures. There is no doubt that Kratochvíl (1940) was right in this respect. Kratochvíl's (1940) error was that he felt either entitled or obliged (or both) to designate new types from a new type locality for S. absoloni, choosing a cave ~ 35 km NW of Trebinje, "Grabova peć kod Grabova dola". We can only speculate about his motivation, but it seems reasonable to assume that he had been postponing his detailed revision of the genus for many years (at least since 1932), hoping to eventually be able to see Kulczyński's type series. This hope was frustrated, as indicated in a footnote (Kratochvíl 1940: 16) where he explicitly stated that the specimens from Ilijina pećina and Zovica jama were not accessible to him. Unfortunately, among his rich new material there were no specimens from any of these two caves. We made a major effort to locate Kulczyński's types, but found only one vial, deposited in the Museum & Institute of Zoology, PAS, in Warsaw, Poland. The supposed types of S. absoloni deposited in Prague and listed in Růžička et al. (2005) are Kratochvíl's erroneous 'types' from "Grabova peć kod Grabova dola". Kulczyński's true type vial contains a label in his own handwriting, but the only locality information on this label is "Trebinje". A second, very detailed label says "Ilijina pećina", but other information on this label leaves no doubt that it was added much later. Fortunately, Kulczyński's original catalogue (in Warsaw) offers a clear hint about the origin of the specimens in the vial. It says "Jaskinie koło Trebinje" [" caves near Trebinje "], suggesting that Kulczyński joined the specimens from Ilijina pećina and Zovica jama. This type vial contains only three specimens, one male and two females. This is clearly only part of Kulczyński's material, because he obviously had at least two males available. However, both variants/ species are represented in this vial. Circumstantial evidence suggests that one female is from Ilijina pećina (it corresponds to more recently collected females from this cave). This forces us to conclude that the other specimens (one male and one female) are from Zovica jama. This is clearly a case where a lectotype should be designated for S. absoloni in order to provide stability and avoid further confusion about the basics (ICZN Recommendation 74G). However, it is not obvious which specimen would best serve as a lectotype. Selecting the male specimen has one major disadvantage: it makes Zovica jama the type locality, a cave that we were not able to locate, and that was even unknown to local cavers in Trebinje we met in 2014. On the other hand, Ilijina pećina is well known and easily accessible, providing a precise type locality. Selecting the female from Ilijina pećina would thus seem to better satisfy ICZN Recommendation 74E (Verification of locality). At the same time, selecting the male specimen as lectotype has also advantages. Most importantly, it preserves the prevailing usage of the name. This male is the same variant/species as Kratochvíl's (1940) S. absoloni ʻtypesʼ from "Grabova peć kod Grabova dola", and it also corresponds to what Senglet (1971) interpreted as S. absoloni. Thus, selecting the male satisfies ICZN Recommendation 74A (Agreement with previous restriction). Asecond (minor) advantage is the fact that species in Stygopholcus are slightly easier to distinguish by males than by females (as in most Pholcidae). Thus, we decided to select the male as lectotype, mainly for two reasons: it seems to better serve stability, one of the major guiding principles of the Code. Second, it is possible that the exact location of Zovica jama will eventually be determined, weakening the argument against selecting the male. Diagnosis Stygopholcus includes relatively large (total body length: ~5–6), long-legged spiders with oval abdomen (Figs 1–5). Males of Stygopholcus are easily distinguished from all other Smeringopinae by modified (club-shaped) hairs frontally on chelicerae (Figs 19, 51); also by unique transparent or weakly sclerotized dorsal bulbal process (arrows in Figs 17, 117) and by distinct brushes of hair-shaped retrolateral membranous processes at tip of procursus (Figs 29, 67, 99, 137). Males and females differ from other Smeringopinae by tarsal organs of legs with undulating rim (Figs 38, 75, 97, 142). Description Male BODY. Total body length ~5–6; carapace width ~1.5–2.5. Carapace with deep central pit and pair of shallow furrows diverging from posterior side of pit toward posterior rim (Figs 39, 61, 91, 107); ocular area slightly raised, eye triads relatively close together (distance PME–PME similar to PME diameter), each secondary eye (especially PME) accompanied by indistinct elevation (arrows in Fig. 61; "pseudoeyes"; cf. Huber 2009), AME relatively large (usually ~40–50% of PME small diameter). Clypeus high, unmodified or with indistinct modified area medially (arrow in Fig. 129). Abdomen oval, never elevated or pointed above spinnerets (Figs 1–5). Male gonopore with 4–8 epiandrous spigots (Figs 62, 92, 143), ALS with only two spigots each: one large widened spigot and one pointed spigot (Figs 69, 135); PMS with two spigots each (cf. female, Fig. 42); PLS without spigots. COLOR. In general ochre-yellow to brown, with distinct dark pattern in S. photophilus, without or with reduced dark pattern in other (troglophile) species. Carapace mostly pale, with darker ocular area and posterior triangle (Figs 1–2, 4); sternum either dark brown to black (S. photophilus), or light brown, with darker brown radial marks (other species). Legs in S. photophilus with dark rings on femora (subdistally) and tibiae (proximally and subdistally) and with short dark longitudinal lines dorsally on femora (Figs 127–128); in other species monochromous, without dark rings and longitudinal lines. Abdomen in S. photophilus with distinct dorsal and ventral patterns; in other species with fewer dorsal and lateral dark marks restricted to posterior part and without or with indistinct ventral pattern. CHELICERAE. Chelicerae with 1–4 modified (cone-shaped to club-shaped) hairs on each distal cheliceral apophysis and ~10–40 club-shaped hairs on frontal face (Figs 19, 27–28, 51, 63–64, 93–94, 119, 130, 132); in S. photophilus with strongly sculptured median edge (Fig. 131); with stridulatory ridges (Figs 35, 71, 95, 133), distances between ridges ~ 6–9 µm in 'northern clade', ~ 3 µm in S. photophilus. PALPS. In general as in Figs 6–11; coxa with rounded retrolateral hump; trochanter barely modified; femur widening distally, slightly curved towards dorsal, with proximal retrolateral process, indistinct transversal dark line on retrolateral side, and stridulatory pick (modified hair) on prolateral side (Figs 65, 103); femur-patella joints shifted toward prolateral side (arrows in Fig. 6); tibia-tarsus joints shifted toward retrolateral side (arrows in Fig. 8); palpal tarsus without dorsal macrotrichia, palpal tarsal organ exposed (Figs 30, 104); procursus dorsally with approximately five (S. photophilus) to ten (other species) weakly to strongly curved hairs; procursus without ventral 'knee', distally with strong ventral spine, with dorsal and prolateral processes of variable size, and distinctive brushes of hair-shaped retrolateral membranous processes (Figs 29, 67, 99, 137); genital bulb with basal sclerite connecting to tarsus (bs in Figs 15, 118), with small and weakly sclerotized (S. photophilus) or transparent (other species) dorsal process (arrows in Figs 17, 117), and main sclerite consisting of retrolateral and dorsal processes (rp and dp in Figs 15–18, 118); sperm duct opening directly on genital bulb at basis of dorsal process (arrow in Fig. 32). LEGS. Legs long and relatively thin, leg 1 length ~35–60, tibia 1 length ~10–15, tibia 2 longer than tibia 4 (1.1–1.2×). Tibia 1 L/d ~60–75. Femur 1 usually with spines ventrally in one row (S. photophilus) or two rows (other species) (Fig. 73), in very small males rarely without spines; spines proximally gradually transforming into regular setae; spines never present on femur 2 or on tibia 1; legs without curved hairs; with few short vertical hairs; retrolateral trichobothrium in proximal position (at 2–4% of tibia length in tibia 1), prolateral trichobothrium absent on tibia 1, present on other tibiae. Tarsal pseudosegments very indistinct, never regular rings but rather indistinct irregular platelets. Tarsus 4 with two rows of prolatero-ventral comb-hairs (Figs 34, 74; see also Huber & Fleckenstein 2008: fig. 13). Tarsal organs of legs capsulate, with weakly (S. photophilus) to strongly (other species) undulating rim (Figs 75, 97). Female In general very similar to male; chelicerae with less distinct or without stridulatory files (Figs 36, 72, 96, 134); legs slightly shorter than in male, without spines. All tarsal organs capsulate (i.e., also on palps; Figs 37, 109, 141). Representatives of the 'northern clade' with pair of indistinct processes posteriorly on carapace (arrows in Figs 39, 107) acting against pair of poorly visible plates on abdomen. Epigynum usually consisting of large, simple anterior plate and short but wide posterior plate (Figs 23, 55, 85, 123); in S. photophilus with pair of bulging areas in front of anterior plate (arrows in Fig. 144). Internal genitalia (Figs 145–156) with sclerotized arc that consists of dorsal and ventral parts (da and va in Figs 153, 156) and is variably visible in uncleared specimens; uterus externus with median ventral pouch, usually clearly visible as round or oval structure in untreated specimens (e.g., Figs 55–60); with pair of sclerites originating from posterior margin and either slightly converging anteriorly or parallel (arrows in Figs 25, 125), indistinct in S. skotophilus; pore plates large, flat, with homogeneously distributed pores (Figs 22, 54, 84, 122). Relationships Stygopholcus is morphologically very similar and geographically very close to Hoplopholcus (Huber 2020). This led to a debate between A. Senglet and P. Brignoli in the 1970s and 80s, on whether the two genera should be synonymized or not (Senglet 1971; Brignoli 1971, 1976, 1983). In the end, Senglet (2001) presented a series of morphological differences, and since version 2.5 of the World Spider Catalog (Platnick 2002) Stygopholcus has been listed as a valid genus. The most recent molecular phylogeny of Pholcidae (Eberle et al. 2018) has suggested that Stygopholcus and Hoplopholcus are not even sister taxa, placing Stygopholcus closer to Crossopriza than to Hoplopholcus (with a 'reasonable' bootstrap support of 87). Morphological characters that support a sister-group relationship between Stygopholcus and Crossopriza are the female stridulatory apparatus between prosoma and abdomen and the dark lines on the legs. However, a convincing morphological analysis will require a detailed study of Crossopriza (a revision is in preparation) and of Holocnemus. The type species of Holocnemus (H. pluchei) was not included in the molecular analysis of Eberle et al. (2018), and the two species of Holocnemus that were included (H. caudatus and H. hispanicus) may in fact be misplaced in Holocnemus and belong to Crossopriza (B.A. Huber, unpubl. data). The monophyly of Stygopholcus is strongly supported by molecular data (maximum bootstrap support; Eberle et al. 2018) and by several putative synapomorphies: (1) male chelicerae with club-shaped hairs on frontal face (Figs 19, 51); (2) unique undulating rim of leg tarsal organs (Figs 38, 75, 97, 142); (3) transparent to weakly sclerotized process dorsally on genital bulb (arrows in Figs 17, 117). Within Stygopholcus, there is strong evidence that S. photophilus is sister to all other species (the 'northern clade'). Both a six genes approach (Eberle et al. 2018) and a study using hundreds of universal single-copy orthologs (L. Dietz et al., unpublished data) consistently recovered this relationship, and numerous morphological characters are shared by all species except S. photophilus. Some of these are likely to be synapomorphies: (1) spines on male femur 1 arranged in two rows rather than in just one (Fig. 73); (2) dorsalbulbal processwithtwoseparatetips (Figs 17–18, 49–50, 81–82); (3) troglophile mode of life and corresponding pale coloration (Figs 1–2). Within the 'northern clade', there is equally strong support for the relationship absoloni + (skotophilus + montenegrinus). Several morphological characters shared by S. skotophilus and S. montenegrinus support this view, even though it is not clear yet if all of these shared similarities are synapomorphies: (1) procursus straight (rather than weakly curved toward femur as in S. absoloni) (Fig. 44); (2) distance betweentipsof dorsalbulbalprocess (bulbal measure D; Fig. 17)> 0.24 (S. absoloni) (Fig. 159); (3) sizeofbulb (bulbal measure E; Fig. 15)> 0.80 (S. absoloni) (Fig. 160). Natural history Stygopholcus, Published as part of Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, pp. 1-60 in European Journal of Taxonomy 752 on pages 4-12, DOI: 10.5852/ejt.2021.752.1391, {"references":["Kratochvil J. 1932. Prispevek k poznani Araneid stredni Slavonie. Sbornik Vysoke Skoly Zemedelske, Brno (C) 23: 1 - 16.","Kulczynski W. 1914. Aranearum species novae minusve cognitae, in montibus Kras dictis a Dre C. Absolon aliisque collectae. Bulletin international de l'Academie des Sciences de Cracovie 1914: 353 - 387, pl. 16.","Absolon K. & Kratochvil J. 1932. Zur Kenntnis der hohlenbewohnenden Araneae der illyrischen Karstgebiete. Mitteilungen uber Hohlen- und Karstforschung 1932: 73 - 81.","Kratochvil J. 1934. Liste generale des Araignees cavernicoles en Yugoslavie. Pregled pecinskih paukova u Jugoslaviji. Prirodoslovne Razprave 2: 165 - 226.","Kratochvil J. 1940. Etude sur les araignees cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1 - 26.","Kratochvil J. 1978. Araignees cavernicoles des iles Dalmates. Prirodovedne prace ustavu Ceskoslovenske Akademie Ved v Brne (N. S.) 12: 1 - 59.","Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grece. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345 - 359. https: // doi. org / 10.5169 / seals- 401666","Senglet A. 2001. Copulatory mechanisms in Hoplopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxonomic data. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 74: 43 - 67. https: // doi. org / 10.5169 / seals- 402796","Brignoli P. M. 1971. Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae). Mitteilungen aus dem Zoologischen Museum in Berlin 47: 255 - 267. https: // doi. org / 10.1002 / mmnz. 1971047020","Brignoli P. M. 1976. Ragni di Grecia IX. Specie nuove o interessanti delle famiglie Leptonetidae, Dysderidae, Pholcidae ed Agelenidae (Araneae). Revue suisse de Zoologie 83: 539 - 578. https: // doi. org / 10.5962 / bhl. part. 91452","Roewer C. F. 1942. Katalog der Araneae von 1758 bis 1940. 1. Band (Mesothelae, Orthognatha, Labidognatha: Dysderaeformia, Scytodiformia, Pholciformia, Zodariiformia, Hersiliaeformia, Argyopiformia). Natura, Buchhandlung fur Naturkunde und exakte Wissenschaften Paul Budy, Bremen.","Brignoli P. M. 1983. A Catalogue of the Araneae described between 1940 and 1981. Manchester University Press, Manchester.","Bonnet P. 1958. Bibliographia araneorum. Analyse methodique de toute la litterature araneologique jusqu'en 1939. Tome II. Systematique des araignees (Etude par ordre alphabetique) (4 me partie: N-S). Douladoure, Toulouse.","Platnick N. I. 1993. Advances in Spider Taxonomy 1988 - 1991, with Synonymies and Transfers 1940 - 1980. The New York Entomological Society, New York.","Platnick N. I. 2000. The World Spider Catalog, Version 1.0. Available from https: // wsc. nmbe. ch / archive / [accessed 8 May 2021].","World Spider Catalog. 2020. World Spider Catalog. Version 21.5. Natural History Museum Bern. Available from http: // wsc. nmbe. ch [accessed 5 Nov. 2020]. https: // doi. org / 10.24436 / 2","ICZN. 1999. International Code of Zoological Nomenclature. Fourth edition. London, U. K. International Trust for Zoological Nomenclature. Available from https: // www. iczn. org / the-code / [accessed 8 May 2021].","Ruzicka V., Kurka A., Buchar J. & Rezac M. 2005. Czech Republic - the type material of spiders (Araneae). Casopis Narodniho muzea, Rada prirodovedna 174: 13 - 64.","Huber B. A. 2009. Life on leaves: leaf-dwelling pholcids of Guinea, with emphasis on Crossopriza cylindrogaster, a spider with inverted resting position, pseudo-eyes, lampshade web, and tetrahedral egg-sac (Araneae: Pholcidae). Journal of Natural History 43: 2491 - 2523. https: // doi. org / 10.1080 / 00222930903207876","Huber B. A. & Fleckenstein N. 2008. Comb-hairs on the fourth tarsi in pholcid spiders (Araneae, Pholcidae). Journal of Arachnology 36: 232 - 240. https: // doi. org / 10.1636 / CSh 07 - 71.1","Huber B. A. 2020. Revision of the spider genus Hoplopholcus Kulczynski (Araneae, Pholcidae). Zootaxa 4726: 1 - 94. https: // doi. org / 10.11646 / zootaxa. 4726.1.1","Platnick N. I. 2002. The World Spider Catalog, Version 2.5. Available from https: // wsc. nmbe. ch / archive / [accessed 8 May 2021].","Eberle J., Dimitrov D., Valdez-Mondragon A. & Huber B. A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https: // doi. org / 10.1186 / s 12862 - 018 - 1244 - 8"]}

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 1.7, 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 [= Meg��lo Perist��ri]; 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. Sch��nhofer 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. Sch��nhofer 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 / P��rama; 35.355�� N, 24.720�� E; 1 Jul. 1970; A. Senglet leg.; MHNG ��� 7 ♂♂, 11 ♀♀, 1 juv., paratypes; Rethymno, Keramoto / P��rama; 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. Sch��nhofer 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. Sch��nhofer leg.; SMF ��� 6 ♂♂, 2 ♀♀; same locality as for preceding, (���Halbh��hle 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. ALBANIA ��� Gjirokast��r ��� 1 ♂, 1 ♀; P��rmet, 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) Durr��s ��� ♂ ♀ (not specified); Durr��s; 41.3�� N, 19.5�� E; 26 May 1993; P. Stoev and D. Zapryanova leg. (Deltshev et al. 2011). ��� Korc�� ��� ♂ ♀ (not specified); Cave 2 near Leskovik; 40.150�� N, 20.600�� E; 1 Jun. 1994; T. Ivanova leg. (Deltshev et al. 2011). ��� Gjirokast��r ��� ♂ ♀ (not specified), Dragot, Vijoshe River; 40.288�� N, 20.069�� E; 1 Jun. 1994; G. Blagoev leg. (Deltshev et al. 2011). 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, ���M��hlenbach���; 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 ��m; diameter PME 120 �� 140 ��m; distance PME���ALE 30 ��m; diameter AME 60 ��m; distance AME-AME 30 ��m. 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. 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. 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)., Published as part of Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, pp. 1-60 in European Journal of Taxonomy 752 on pages 41-53, DOI: 10.5852/ejt.2021.752.1391, http://zenodo.org/record/4958808, {"references":["Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grece. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345 - 359. https: // doi. org / 10.5169 / seals- 401666","Brignoli P. M. 1971. Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae). Mitteilungen aus dem Zoologischen Museum in Berlin 47: 255 - 267. https: // doi. org / 10.1002 / mmnz. 1971047020","Deltshev C. D. 1985. New data concerning cave spiders (Araneae) in Greece with description of a new Leptonetela (Araneae, Leptonetidae). Acta zoologica Bulgarica 27: 41 - 45.","Brignoli P. M. 1976. Ragni di Grecia IX. Specie nuove o interessanti delle famiglie Leptonetidae, Dysderidae, Pholcidae ed Agelenidae (Araneae). Revue suisse de Zoologie 83: 539 - 578. https: // doi. org / 10.5962 / bhl. part. 91452","Senglet A. 2001. Copulatory mechanisms in Hoplopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxonomic data. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 74: 43 - 67. https: // doi. org / 10.5169 / seals- 402796","Kratochvil J. 1932. Prispevek k poznani Araneid stredni Slavonie. Sbornik Vysoke Skoly Zemedelske, Brno (C) 23: 1 - 16.","Kulczynski W. 1914. Aranearum species novae minusve cognitae, in montibus Kras dictis a Dre C. Absolon aliisque collectae. Bulletin international de l'Academie des Sciences de Cracovie 1914: 353 - 387, pl. 16.","Kratochvil J. 1940. Etude sur les araignees cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1 - 26.","Deltshev C. D., Vrenosi B., Blagoev G. & Lazarov S. 2011. Spiders of Albania - faunistic and zoogeographical review (Arachnida: Araneae). Acta zoologica Bulgarica 63: 125 - 144."]}

Stygopholcusmontenegrinus Kratochv��l, 1940 orig. stat. Figs 3, 77���112, 149���150, 155 Stygopholcusmontenegrinus Kratochv��l, 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 ��� Nik��ič ��� ♀ (without epigynum), near Nik��ič, Studenačka pećina; 42.7680�� N, 18.9312�� E [Naumova et al. 2019: 42.826�� N, 18.786�� E]; 6 Sep. 1936; J. Kratochv��l leg.(?); NMP. Other material examined MONTENEGRO ��� Nik��ič ��� 1 juv.; same collection data as for holotype; NMP ��� 9 ♂♂, 8 ♀♀; near Nik��ić, 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, Budo��ka pećina; 42.7303�� N, 18.9498�� E; 26 Jul. 1969; C. Deeleman leg.; NMNL (ARA 19035) ��� 2 ♂♂, 2 ♀♀; Slivlje, Golube��nica 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. Kratochv��l 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 Kr��, 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, Babatu��a 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, (���Bobotu��a pećina���); 1932(?); J. Kratochv��l 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. ��� Kola��in ��� 1 ♂, 1 ♀; Monastery Morača; 42.7648�� N, 19.3931�� E; 300 ma.s.l.; 8 May 2006; A. Sch��nhofer 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., Babatu��a 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, Lije��ta 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. ��� Kuk��s ��� 1 ♂; Hekurave Mt., Qere��-Mulaj, Zeze Cave; 42.362�� N, 19.930�� E; 18 Aug. 2010; M. Komnenov leg.; CMK ��� 2 ♂♂; below Hekurave Mt., Qere��-Mulaj, small cave near River Kuqit; 42.364�� N, 19.933�� E; 17 Aug. 2010; M. Komnenov leg.; CMK ��� 1 ♀; Prokletije, Curaj i Ep��rm, N��ngurit 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 ��m; diameter PME 130 �� 160 ��m; distance PME���ALE 30 ��m; diameter AME 60 ��m; distance AME-AME 20 ��m. 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)., Published as part of Huber, Bernhard A., Pavlek, Martina & Komnenov, Marjan, 2021, Revision of the spider genus Stygopholcus (Araneae, Pholcidae), endemic to the Balkan Peninsula, pp. 1-60 in European Journal of Taxonomy 752 on pages 33-41, DOI: 10.5852/ejt.2021.752.1391, http://zenodo.org/record/4958808, {"references":["Kratochvil J. 1940. Etude sur les araignees cavernicoles du genre Stygopholcus Krat. Acta Societatis Scientiarum Naturalium Moraviae 12: 1 - 26.","Senglet A. 1971. Note sur les Pholcidae (Arachn.) de Grece. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 44: 345 - 359. https: // doi. org / 10.5169 / seals- 401666","Naumova M., Lazarov S. P., Petrov B. P. & Deltshev C. D. 2016. New faunistic data on the cave-dwelling spiders in the Balkan Peninsula (Araneae). Ecologica Montenegrina 7: 425 - 438.","Deltshev C. D., Vrenosi B., Blagoev G. & Lazarov S. 2011. Spiders of Albania - faunistic and zoogeographical review (Arachnida: Araneae). Acta zoologica Bulgarica 63: 125 - 144.","Eberle J., Dimitrov D., Valdez-Mondragon A. & Huber B. A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https: // doi. org / 10.1186 / s 12862 - 018 - 1244 - 8","Naumova M., Lazarov S. & Deltshev C. D. 2019. Faunistic diversity of the spiders in Montenegro (Arachnida: Araneae). Ecologica Montenegrina 22: 50 - 89. https: // doi. org / 10.37828 / em. 2019.22.5"]}

Prvi špiljski pauk na svijetu opisan je s Dinarida 1847. godine – vrsta Stalita taenaria iz porodice Dysderidae (na hrvatskom dizderidi). Od tada do danas broj poznatih vrsta na Dinaridima narastao je do 24, te su kao i većina špiljskih beskralješnjaka sve vrste endemi Dinarida. Dizderidi su karakteristični po tome što ne pletu mreže nego hodaju po podu i zidovima špiljskih kanala. Sve dinarske špiljske vrste su slijepe i depigmentirane te ih se ne može naći u vanjskih staništima. Rodovi Stalita i Parastalita rasprostranjeni su od Slovenije do južnog Velebita, a na južnom Dinaridima, od kanjona rijeke Krke pa sve do Crne Gore, možemo naći rodove Stalagtia, Folkia i Stalitella. Pauci iz ovih rodova su s veličinom tijela od 5 do 13 mm te nogama do 30 mm lako uočljivi. S druge strane, vrste iz rodova Rhode i Mesotalita rasprostranjene su po cijelim Dinaridima, ali u svega nekoliko špilja. Jako ih se rijetki nalazi, veličina tijela im je puno manja i pretpostavka je da im je primarno stanište sustav pukotina koji okružuje špilje. U zadnjih nekoliko godina sustavna biospeleološka istraživanja i upotreba DNA analiza rezultirala su otkrićem brojnih novih vrsta što pokazuje da Dinaridi, iako poznati kao svjetski centar podzemne bioraznolikosti, još uvijek kriju mnoge tajne.

A two-year research projects in the Krka National Park and Paklenica National Park were conducted aiming to test different methods for cave habitats and fauna monitoring in order to develop monitoring protocols for different species, show caves and cave habitats. Altogether, we investigated 16 caves and 28 target species from the following taxa: Serpulidae, Araneae, Isopoda, Amphipoda, Decapoda, Diplopoda, Chilopoda, Collembola, Diplura and Coleoptera. Here, we will present the results from the monitoring of fauna in three caves from the Krka National Park (Stara jametina, Miljacka II and show cave Oziđana špilja), and the show cave Manita peć in Paklenica National Park. We used two different methods: (1) invertebrate census of the whole cave (in Oziđana špilja), and (2) invertebrate census on plots (in Stara jametina and Miljacka II). The first method is suitable for small caves. For this method, it is best to divide the cave into 3-5 zones and conduct a “minimum- time census”, with a predefined number of people and the amount of time per zone. The second method is applicable for large caves. Again, it is necessary to define the number of people and the amount of counting time per plot. In Manita peć, we combined these two methods: the cave was divided into three zones and plots per zone (in triplicates) were established. With this approach, we were able to collect quantitative data sets and estimate the total abundance of the fauna as well as the abundance of individual taxa. Furthermore, this monitoring scheme is replicative and suitable for long-term monitoring of both fauna and cave habitats. A detailed species inventory of the investigated cave and the trained researchers that can identify species on sight, without collecting them, are crucial prerequisites for the successful implementation of the abovementioned methods.

The Dinaric karst, which covers the western part of the Balkan Peninsula, is a global hotspot of subterranean biodiversity. Spiders of the family Dysderidae are among the most conspicuous creatures in Dinaric caves, where they are represented by eight genera, two in the subfamily Harpacteinae (Stalagtia and Folkia) and five in Rhodinae, (Rhode, Stalita, Mesostalita, Parastalita and Stalitella), adding up to a total of 21 species. All these species are Dinaric endemics, and almost all are troglobiotic (obligatory cave dwellers). The two subfamilies show near disjunct distributions in the region, Harpacteine are restricted to the south part of the Dinarides and Rhodinae (with few exceptions) to the north part. With the aim of providing an explicit phylogenetic hypothesis of the family, we conducted a mutli-locus phylogenetic analysis combining mitochondrial and nuclear genes (COI, 16S, 28S and H3). Our results revealed a more complex taxonomic structure than currently recognized, and uncovered major overlooked diversity at the species and generic levels. Quantitative species delimitation methods confirmed the existence of many potentially new species, and revealed high levels of cryptic diversity within Harpacteinae. We further estimated divergence times using a combination of fossil and biogeographic node calibrations and reconstructed the biogeographic history of these spiders. All cave lineages showed a common pattern of long stem branches, which may hint at high levels of extinction in the history of these groups. We hypothesize that Miocene climatic changes may have played a crucial role in shaping the extant diversity of these cave-dwelling dysderids.

Subterranean genus Verhoeffiella occurs in five discontiguous karstic regions throughout Europe: Dinarides, Jakupica in North Macedonia, South-eastern Calcareous Alps, Southern Catalonia and Cordillera Cantabrica. This disjunct distribution challenges the monophyly of the genus and questions the scenarios of its origin and colonization. In the molecular phylogenetic framework we tested the monophyly of Verhoeffiella and explored its relationship with the surface and presumably related genus Heteromurus. Further, using species delimitation methods and time divergence estimation, we explored hidden diversity and temporal diversification of the genus. Our results show that both genera, Verhoeffiella and Heteromurus, are paraphyletic, while species delimitation methods revealed overlooked diversity and increased molecular divergence. We identified 79 MOTUs of Verhoeffiella in Dinarides, which is seven times more than the number of nominal species. In addition, we identified more than 30 MOTUs of troglophilic species Heteromurus nitidus. When placed on the time scale, events triggering the diversification were linked to the Messinian Salinity Crisis and Pleistocene climatic shifts. The combination of this extensive subterranean diversification and close evolutionary links with epigean relatives makes Verhoeffiella an exceptional case within the subterranean fauna of Dinarides.

Subterranean habitats are inhabited by numerous specially adapted taxa, which typically have limited distribution ranges on both specific and generic level. They are generally bound to narrow geographical areas, yet there are exceptional lineages exhibiting large and disjunct distributions. One of the cases is the subterranean collembolan genus Verhoeffiella that occurs in five remote karstic regions, from 150 to 1000 km apart (Dinarides, Jakupica in North Macedonia, South-eastern Calcareous Alps, Southern Catalonia and Cordillera Cantabrica). We tested the monophyly of Verhoeffiella and explored its relationship with the surface and presumably related genus Heteromurus. Further, using species delimitation methods and time divergence estimation, we explored hidden diversity and temporal diversification. Our results show that the genera Verhoeffiella and Heteromurus are paraphyletic, with lineages of Verhoeffiella intermixed with lineages of Heteromurus nitidus. Discovery of highly diversified subterranean group with closely related surface relatives represents an exceptional case within the subterranean fauna of temperate regions. Species delimitation methods revealed increased molecular divergence and overlooked diversity within both genera. The highest increase was noted in Verhoeffiella populations in the Dinarides, where the number of MOTUs (N=79) exceeded the number of described species by eight times. When put on the time scale, most of cladogenetic events coincide with Messinian salinity crisis (5.96-5.33 MA), while speciation events with the age of Pleistocene (2.5 MA - 11.7 k years ago), indicating that paleogeographic changes could be one of the main triggers of diversification within Verhoeffiella.

„Mali Šibenik - veliko bogatstvo podzemlja “(Mali Šibenik- great wonders of the underground) was a project that has been conducted during two expeditions in 2018. It was organised by the Breganja Association in conjunction with the SAK Ekstrem-Speleological Club from Makarska and the city of Vrgorac. The project was carried out in the hills above Vrgorac. During the expeditions, 21 new caves were discovered, thirteen of which have been speleologically investigated and eleven caves were surveyed. A great number of caves have shown a high biodiversity of subterranean fauna and pointed out a need for further exploration of this area. Although the deepest explored cave in this project is Bokšića Golubinka (-50.6 m), the more interesting fauna has been found in shallower caves. For diplopods and spiders, the most significant caves are Jama Malog Ulaza, Golubinka u Gaju, Antipatična, Rođendanska Jama, Ošlja Jama i Ančića Velika Golubinka in which potentially new species have been found. Results of this project were presented via a poster exhibition, the “Mali Šibenik- veliko bogatstvo podzemlja “at Croatian anannual Speleogathering in Ogulin (2018). Also, some educational presentations for high school students and interested public were held in the city of Vrgorac. The main goal of these presentations was to raise environmental awareness among general public, especially the local youth.

Geographic information collected for each locality, and local- and broad-scale predictors considered in the analyses.

Caves are one of the last uncharted territories on Earth. The Dinaric karst in the Balkan peninsula is home to many of them, but despite of darkness and harsh conditions, life thrives there. Spiders of the family Dysderidae are among their most abundant and conspicuous inhabitants. Yet, we know very little about their biology, ecology, and evolutionary history. As a first step towards improving our knowledge on cave dysderids, we here present an explicit phylogenetic hypothesis of the group based on an extensive biotic survey of the region. Seven Dysderidae genera with 21 species are known in the Dinaric caves: Stalagtia and Folkia in the Harpacteinae subfamily, and Rhode, Stalita, Mesostalita, Parastalita and Stalitella in the Rhodinae. Almost all species show high level of cave adaptation and are narrow endemics. We used a target multi-locus approach based on 4 concatenated genes to infer a phylogeny, delimit species boundaries and estimate divergence time combining fossil and biogeographic node calibrations. We uncovered major overlooked diversity at both species and genera levels. Quantitative species delimitation methods confirmed the existence of many new species, and revealed high levels of cryptic diversity within Harpacteinae. All cave lineages showed a common pattern of long stem branches, which may hint at high levels of extinction during the history of these groups. We hypothesise that Miocene climatic changes may have played a crucial role in shaping the extant diversity of these cave-dwelling spiders. Finally, we reconstructed the biogeographic scenario for the diversification of the Dinaric dysderid fauna.

Root-knot nematodes (genus Meloidogyne) are established as important model organisms of plant parasites. Their characteristics, such as holocentric chromosomes, polyploidy and parthenogenesis make them an ideal system to study genome evolution, speciation processes, and centromere structure and function.In this regard, our analysis of centromere specific histone H3 (CENH3) in the genome of M. incognitareveals three divergent CENH3 variants instead of the commonly present only one. All CENH3 variants are completely conserved in related species, and transcriptome data show their developmental stage- specific expression. Immunostaining of chromosomes with antibodies specific for CENH3 variants reveals holocentric distribution pattern for each of them. Given that the centromere tends to be established upon satellite DNA arrays wefurther focus on their characterization in related Meloidogyne genomes. Using next-generation sequencing combined with novel bioinformatics tools we detected a remarkable collection of satellite DNAs (satellitome) in each genome. Comparative analyses of satellitomes indicate the species evolutionary history in accordance with a recent hypothesis which suggests a multiple hybridization events in species origin. All results will be discussed in terms of coexistence and possible roles of different CENH3 variants in holocentric centromeres, and correlated with trends in satellitome evolution

In Dinarii iaves, dysderid spiders are aming the mist iinspiiuius animals. Their relatvely large size, irange iiliur and aitve firaging lifestyle makes them hard ti miss in the seemingly empty iave envirinment. In the subfamily Rhidine, 13 speiies frim 5 genera are iurrently reiignized in the Dinarides: genus Rhode with mistly epigean representatves in the ientral and western Mediterranean basin but with 4 iave adapted speiies in the study area, and 8 speiies frim Dinarii endemii and striitly iave-dwelling genera Stalita, Parastalita, Mesostalita and Stalitella. The debate remains in the plaiing eaih if these 8 related speiies in its iwn genus ir iimbine them all under Stalita. Rhidinae are distributed mistly thriugh the nirthern Dinarides, with a few remarkable exieptins reaihing far siuth areas. All but ine speiies have restriited distributin ranges as expeited under the assumptin if habitat fragmentatin and isilatin if the diferent karst areas. As tip predatirs, these speiies play a key rile in the fragile iave eiisystems. Here we present the preliminary results if the frst mutli-liius phyligeneti analysis if the subfamily Rhidinae. A siund phyligeneti hypithesis will represent a majir step firward ti iur iurrent understanding in the irigins and evilutin if these beautful ireatures and if the while Dinarii karst system - the Wirld’s hit spit fir iave biidiversity.

Krypyonesticus deelemanae gen. et sp. nov. urn:lsid:zoobank.org:act: BAC 5D31D-7A59-4FE7-BE9A-CB6167484170 Figs 1, 2A, 3 A���C, 4A, D, 5A, D Diagnosis The most prominent character of the male bulb is a large and conspicuous median apophysis which emerges from the middle part of the tegulum and reaches its apical part. The most obvious differences between the new species and the closest ones, K. fagei and K. arenstorffi, are: distal part of MA in the new species is massive and slightly thinner toward the top, while in K. arenstorffi the whole distal part is slender and the top is more narrowed; the proximal part of MA of the new species has a round process on the outer edge, while in K. fagei and K. arenstorffi it has no process. Another important feature is the tooth-like dorsomedian apophysis of the paracymbium���s dorsal process, which is not present in K. fagei and K. arenstorffi. The inner side of the distal part of MA, the curve between the proximal and distal parts of MA, the shape of the tegulum, the basal tegular apophysis and the dorsal paracymbium process are also diagnostic. The epigyne is most similar to that in K. fagei and K. arenstorffi, from which it differs in the outline of the posterior epigyne margin which is triangular in K. fagei and sinuous in other two species. The difference between the new species and K. arenstorffi is in the posterior epigyne border, lateral of the central chitinous arches, which is straight in the new species and slightly notched in K. arenstorffi. The vulva can be distinguished by the position of the spermathecae, whose apical part in the new species reaches over the vulvar chitinous structures, while it does not in K. fagei and K. arenstorffi. Etymology The specific name is a patronym in honor of Christa Laetitia Deeleman-Reinhold, an important Dutch arachnologist and a dear friend. Her work has vastly raised the knowledge of the cave spider fauna of Dinarides. The species name is in possessive genitive. Material examined Holotype CROATIA: 1 ♂, Biokovo Mt, Samogorska ��pilja, 43��19���5.71��� N, 17��7���29.9��� E, elevation 717 m, 23 Jan. 2016, leg. Marko Lukić (CBSS / AR 3934-1). Paratypes CROATIA: 1 ♀ (CBSS /AR 3934-2), 1 ♂, 6 ♀♀ (CBSS /AR 3934-5), leg. Marko Lukić; 1♂, 1♀, leg. Marko Lukić (SMF); 1♂, 7 ♀♀, leg. Tin Rožman (CBSS /AR 3932). All with same collection data as the holotype. Other material examined CROATIA: 1 ♂, Biokovo Mt, Kuna jama, 27 Aug. 1998, leg. Roman Ozimec (CBSS /AR 1495); 1 ♂, Biokovo Mt, Crna Ledenica, 24 Oct. 2006, leg. Marko Lukić (CBSS /AR 1277); 1 ♂, Biokovo Mt, Stonjska peć donja, 27 Oct. 2003, leg. Roman Ozimec (CBSS /AR 1287). Comparative material TURKEY: K. dimensis (L��pez-Pancorbo, Kunt & Ribera, 2013), 1 ♂, 1 ♀, Antalya Province, Alanya District, Kestel Town, Dim Valley, Dim Cave, 3 Dec. 2011, leg. Kadir Boğa�� Kunt. CROATIA: K. eremita (Simon, 1879), 2 ♀♀, 2 ♂♂, 2 juv.. (CBSS /AR 1160), Dubrovnik, Osojnik, Močiljska ��pilja, 21 June 2001, leg. Roman Ozimec. K. arenstorffi (Kulczyński, 1914) and K. fagei (Kratochv��l, 1933): see below. Description Male (holotype) COLORATION. Carapace uniform whitish, opisthosoma yellowish without marked darker patches. Sternum of the same color as carapace. Appendages slightly darker, especially the distal segments (Fig. 1A). PROSOMA. Approximately circular in dorsal view. Cephalic region not differentiated from rest of prosoma. Fovea and thoracic grooves visible. Eyes normally developed. Eye size and interocular distances: AM = 0.042; AL = 0.081; PM = 0.085; PL = 0.077; AM���AL = 0.07; AM���AM = 0.067; PM���PL = 0.075; PM��� PM = 0.129; PL���AL = 0.02. OPISTHOSOMA. Sub-elliptical in dorsal view. CHELICERA. Mesal part of promargin with three teeth of about the same size. MALE PALP. Paracymbium with a well-developed dorsal and very simple ventral processes. Dorsal process with two branches, apical branch parallel to cymbium���s longer axis and shorter basal branch pointing to cymbium. Apical branch with parallel rims in proximal part, slightly dilated in middle with distal part acuminated, dagger-shaped. Basal branch with convex upper rim and with distal part sharp and tilted upward (Figs 2A, 3B). Dorsomedian apophysis tooth-like, small with broad base. Ventral process (Figs 2A, 3A) broad, rounded, with transparent middle part and a membranous extension on rim (Figs 2A, 3A). Tegulum with a shallow, rounded and broad longitudinal groove (visible in apical view, Fig. 3C) and with a broad, roughly circular basal apophysis on base (in ventral view). Median apophysis remarkably well developed, V-shaped, distal arm reaching almost to apical part of tegulum. Proximal part of MA with wrinkled outer rim which in posterior view looks like a small process. Outer edge between proximal and distal part equally rounded, distal part of MA mildly narrowing toward top, bluntly ended, with slightly serrated upper edge. Inner side of distal part of MA slightly swollen. Conductor absent. TTA with two processes: TTA p1 and TTA p2 (homologous to processes p1���p6 of the conductor complex in Huber 1993). TTA p1 saddle-shaped, longer than wide, with slim, rounded and pale process on top and in central area with three small dark teeth in a row, middle one smallest and proximal one faintly rounded. TTA p2 located in an apical position, with one sharp and pointed process, the rest forming groove where tip of embolus lodges and which serves as a conductor. Filamentous embolus partially borders tegulum and reaches its apex in semicircular course (Figs 2A, 3A). Measurements: PL: 1.552; PW: 1.407; OL: 2.075; OW: 1.412; TL = 3.627. Leg formula: I> IV> II> III. Female (paratype CBSS/AR 3934-2) SOMATIC CHARACTERS (Fig. 1B) As in male, except that eye size and interocular distances are a bit different and body length a little larger: AM = 0.052; AL = 0.095; PM = 0.085; PL = 0.076; AM���AL = 0.061; AM���AM = 0.070; PM���PL = 0.128; PM���PM = 0.1112; PL���AL = 0.015. Epigyne slightly wider than long, in lateral view not much protruding (Fig. 5D). Lateral lobes with oval depressions in posterior part, reaching middle of epigyne. Posterior epigyne border made of two semicircular chitinous arches, flanking depressions, coming together in middle and forming a small, pale, inconspicuous convex tubercule. A ridge apically bordering tubercle and depressions faintly visible. Epigyne outline, laterally from semicircular chitinous arches, straight, with no notches (Figs 4A, 5 A���B). VULVA. Lateral sides of vulva made of compact dark chitinous structures enclosed in a firm transparent membrane (lateral pouches). Oval shaped spermathecae, with a shallow constriction in middle, located just above chitinous vulval structures (dorsal view). Narrower, upper part of spermathecae protrudes above chitinous vulval structures (Fig. 5C). Insemination ducts depart from spermathecae at bottom (Fig. 5C), make a sharp turn outwards and continue ventro-laterally in semicircular course to reach copulatory orifices (Fig. 4D). In ventral view, spermathecae slightly visible above lateral dark structures, while chitinous curves, through which insemination ducts run visible on both sides of epigyne (Fig. 4A). Two chitinous narrow, horn-like apophyses protrude from inner sides of dark chitinuos vulval structures, in apical direction, to empty middle part of vulva (Fig. 5C). Measurements: PL: 1,666; PW: 1,414; OL: 2,609; OW: 1,872; TL = 4,275. Leg formula: I> IV> II> III. Distribution The new species is endemic to Croatia; it is distributed on Biokovo Mt in central Dalmatia, a coastal region in Croatia. So far it has been recorded in 20 caves scattered through the whole mountain, from the south-west sea side to the north-east continental side, from the 310 to 1640 asl (Fig. 6A). Data on all records of K. deelemanae gen. et sp. nov. are given in Appendix 2. The distribution area of K. deelemanae gen. et sp. nov. is more than 80 km away from that of K. fagei and more than 100 km from that of K. arenstorffi (Fig. 6B). Natural history The type locality, Samogorska ��pilja, is a small cave with two entrances (Fig. 7). On the date of the last collection, 23 Jan. 2016, the air temperature in the cave was 0.5��C with cold air streaming through the cave, mostly near the cave floor. Spiders were found freely walking on the ceiling of the chamber (probably avoiding the cold air flow at the bottom) and on the side walls of the cave. The temperature in other caves where K. deelemanae gen. et sp. nov. is found ranges from 0 to 15��C. Some of those caves are very small and are greatly influenced by outside conditions (like the type locality), while the others are quite big, have a true cave microclimate and harbor diverse types of cave habitats (for example Pretnerova jama, a 254-meter deep pit). No other nesticid species are found in caves on Biokovo Mt., Published as part of Pavlek, Martina & Ribera, Carles, 2017, Kryptonesticus deelemanae gen. et sp. nov. (Araneae, Nesticidae), with notes on the Mediterranean cave species, pp. 1-27 in European Journal of Taxonomy 262 on pages 6-12, DOI: 10.5852/ejt.2017.262, http://zenodo.org/record/3823118, {"references":["Lopez-Pancorbo A., Kunt K. B., Blagoev G., Deltshev C. & Ribera C. 2013. Nesticus dimensis new species, a new troglobitic spider from Turkey (Araneae, Nesticidae), with comments on its phylogenetic relationships. Zootaxa 3721: 183 - 192. http: // dx. doi. org / 10.11646 / zootaxa. 3721.2.5","Kulczynski W. 1914. Aranearum species novae minusve cognate in montibus Kras dictis a Dre C. Absolon aliisque colectae. Bulletin international de l'Academie des Sciences de Cracovie 13: 353 - 387. Available from http: // biodiversitylibrary. org / page / 4553795 [accessed 17 Jan. 2017].","Kratochvil J. 1933. Evropske druhy celedi Nesticidae Dahl. Prace Moravske Prirodovedecke Spolecnosti 8: 1 - 69.","Huber B. A. 1993. Genital mechanics and sexual selection in the spider Nesticus - cellulanus (Araneae, Nesticidae). Canadian Journal of Zoology 71: 2437 - 2447. http: // dx. doi. org / 10.1139 / z 93 - 340"]}

Kryptonesticus arenstorffi (Kulczyński, 1914) comb. nov. Figs 2C, 3 G���I, 4C, 5I���L Nesticus arenstorffi Kulczyński, 1914: 378, pl. 16, fig. 50. Nesticus arenstorffi ��� Kratochv��l 1933: 42, 64, pl. 1, fig. 5, pl. 4, figs 34���36. ��� Le Peru 2011: 361, fig. 620. Ivesia arenstorffi ��� Lehtinen & Saaristo 1980: 51. Diagnosis Proximal part of MA without process on outer edge, inner side of distal part of MA with bulge in middle, edge between proximal and distal part of MA pointed. Tegulum with triangular basal apophysis. Dorsomedian apophysis of paracymbium absent. Posterior epigyne outline wavy, ridge apically bordering lateral depressions well visible. Spermathecae not reaching over vulvar lateral chitinous structures. Type material Not examined. The exact position of the type locality is unknown (for details see Distribution). Material examined MONTENEGRO: 1 ♂ (CBSS /AR 3288-1), 1 ♀ (CBSS /AR 3288-2), Kotorski zaljev, Golodražnica, 31 Mar. 2012, leg. Ana Komerički; 1 ♀ (CBSS /AR 3641-1), 1 ♀, 3 ♂♂, Cetinje, Cetinjska pećina, 28 Oct. 2015, leg. Martina Pavlek (CBSS /AR 3641-2). Redescription Male (CBSS/AR 3288-1) Paracymbium with well-developed dorsal and simple ventral process. Basal branch of dorsal process with slightly convex upper rim which ends in the shape of an indistinct minor blunt tooth. Longer apical branch similar to that in K. deelemenae gen. et sp. nov. but more robust, dilated in middle and attenuated towards top (Figs 2D, 3H). Ventral process broad and round, with transparent middle part and no membranous extension on rim. Dorsomedial apophysis absent (Fig. 3G). Tegulum with inconspicuous groove, low round swelling in apical part (Fig. 2B, in apical view in Fig. 3I), and with triangular broadbased basal apophysis. Median apophysis well developed, V-shaped, distal arm reaching almost to apical part of tegulum. Outer edge between proximal and distal part pointed, not equally rounded as in K. deelemanae gen. et sp. nov. and K. fagei. Distal part of MA narrowing toward top; top bluntly ended with serrated upper edge. Inner side of distal part of MA with bulge in middle. Embolus, TTA p1 and TTA p2 of TTA similar to those in K. deelemanae gen. et sp. nov. (Figs 2D, 3G). Female (CBSS/AR 3641-1) EPIGYNE. Slightly wider than long. In lateral view not protruding very much (Fig. 5L). Lateral lobes with oval depressions in posterior part, reaching middle of epigyne. Posterior epigyne border made of two semicircular chitinous arches, flanking depressions and coming together in middle, forming a small, pale, inconspicuous convex tubercule. Ridge apically bordering tubercle well visible. Epigyne outline, laterally of semicircular chitinous arches, with shallow notches. In ventral view, chitinous curves through which insemination ducts extend visible on both sides of epigyne (Figs 4C, 5 I���J). VULVA. Lateral sides of vulva made of compact dark chitinous structures enclosed in firm transparent membrane (lateral pouches). Spermathecae slender, oval-shaped, with shallow constriction in middle, located meso-apically behind dark chitinous lateral structures. Insemination ducts depart at bottom and extend to epigynal base. Two chitinous narrow, horn-like structures protrude from inner sides of dark chitinous lateral structures, in apical direction, to empty middle part of vulva (Fig. 5K). Distribution The type locality is Laketićeva pećina near the town of Trebinje in Bosnia and Herzegovina (Kulczyński 1914). Since the original description, it has not been collected at the type locality and its exact position is at the moment unknown. In the surroundings of the town of Trebinje no cave with such a name is known. The type material is deposited in the National Museum in Prague (Ružička et al. 2005). The species is also known from Čudna jama near Mostar (Fage 1931) and from 19 more caves in Montenegro (Kratochv��l 1933, 1935; Deeleman-Reinhold 1974). Recently, the species has been recorded in three more caves: Cetinjska pećina as reported by Deltshev et al. (2014) and personal observations, Pećina Vojvode Dakovića (pers. obs.) in Montenegro, and from cave Jezero on Sniježnica Mt which is the first record for Croatia (all records from the CBSS collection are listed in Appendix 3). All localities are marked on the map in Fig. 6B. In the last 20 years, researchers from CBSS visited many caves around the town of Trebinje, in the Popovo polje region and in other parts of Bosnia and Herzegovina, but recovered no K. arenstorffi. Čudna jama near Mostar is far away from all certain K. arenstorffi localities and we wanted to check that material, deposited in the National Museum in Prague, but unfortunately there are only two subadult males and a juv.enile inside the vial (personal communication with Petr Dolej��, curator of the spider collection) so the taxonomical status of the samples could not be confirmed. We consider that locality dubious. The citation of Serbia in ���The Spiders of Europe��� (Nentwig et al. 2016) is erroneous. Natural history K. arensorffi specimens were mostly found hanging upside down from the webs, in the entrance part but also deeper in the caves. The temperature in the caves where K. arenstorffi has recently been collected ranges between 5��C and 15��C. Generally, in all caves where K. arenstorffi was recorded no other nesticid species was present, except in Cetinjska pećina in the town of Cetinje in Montenegro. This cave is inhabited by the troglobiotic species Typhlonesticus absoloni as reported by Christa Deeleman-Reinhold (1974), and it has also been collected recently (Appendix 4). It is worth noting that in her 1974 paper, Christa Deeleman-Reinhold did not mention the presence of K. arenstorffi. Her collecting trip was in 1972, before Cetinjska pećina was adapted for touristic visits and before a tunnel was dug into the cave from the outside (in the 1980s). The tunnel has surely changed the microclimatic conditions in the cave by introducing a constant air flow, so it could have made some parts of the cave less suitable for the troglobiotic Typhlonesticus absoloni (found only in deeper parts), but more accommodating for the less troglomorphic K. arenstorffi., Published as part of Pavlek, Martina & Ribera, Carles, 2017, Kryptonesticus deelemanae gen. et sp. nov. (Araneae, Nesticidae), with notes on the Mediterranean cave species, pp. 1-27 in European Journal of Taxonomy 262 on pages 14-16, DOI: 10.5852/ejt.2017.262, http://zenodo.org/record/3823118, {"references":["Kulczynski W. 1914. Aranearum species novae minusve cognate in montibus Kras dictis a Dre C. Absolon aliisque colectae. Bulletin international de l'Academie des Sciences de Cracovie 13: 353 - 387. Available from http: // biodiversitylibrary. org / page / 4553795 [accessed 17 Jan. 2017].","Kratochvil J. 1933. Evropske druhy celedi Nesticidae Dahl. Prace Moravske Prirodovedecke Spolecnosti 8: 1 - 69.","Le Peru B. 2011. The Spiders of Europe, a synthesis of data. Volume 1 Atypidae to Theridiidae. Memoires de la Societe linneenne de Lyon 2: 1 - 522.","Lehtinen P. T. & Saaristo M. I. 1980. Spiders of the Oriental-Australian region. II. Nesticidae. Annales Zoologici Fennici 17: 47 - 66.","Ruzicka V., Kurka A., Buchar J. & Rezac M. 2005. Czech Republic - the type material of spiders (Araneae). Casopis Narodniho muzea, Rada prirodovedna 174: 13 - 64.","Fage L. 1931. Araneae cinquieme serie, precedee d'un essai sur l'evolution souterraine et son determinisme. Archives de Zoologie experimentale et generale, Biospeologica LV 71: 99 - 291.","Kratochvil J. 1935. Araignees cavernicoles de Krivosije. Prace Moravske Prirodovedecke Spolecnosti 9: 1 - 25.","Deeleman-Reinhold C. L. 1974. The cave spider fauna of Montenegro (Araneae). Glasnik Republickog Zavoda za Zastitu Prirode i Prirodnjackog Muzeja 6: 9 - 33.","Deltshev C., Curcic B., Wang C., Yao Z., Antic D., Curcic S. & Rada T. 2014. New data on the spiders (Araneae) in the caves of Balkan Peninsula. Archives of Biological Sciences 66: 465 - 472. http: // dx. doi. org / 10.2298 / ABS 1402465 D","Nentwig W., Blick T., Gloor D., Hanggi A. & Kropf C. 2016. Spiders of Europe. Version 02 [online]. Available from http: // www. araneae. unibe. ch /"]}

Kryptonesticus fagei (Kratochv��l, 1933) comb. nov. Figs 2B, 3 D���F, 4B, 5E���H Nesticus fagei Kratochv��l, 1933: 44, 64, pl. 1, fig 4, pl. 2, fig 22, pl. 4, figs 37���40. Nesticus fagei ��� Gasparo & Thaler 2000: 32, figs 3���4. ��� Le Peru 2011: 364, fig. 630. Ivesia fagei ��� Lehtinen & Saaristo 1980: 51. Diagnosis Proximal part of MA without process on outer edge, inner side of distal part of MA straight, curve between proximal and distal parts of MA equally rounded. Tegulum with longitudinal ridge stretching through entire surface, basal apophysis of tegulum triangular. Apical branch of dorsal paracymbium process curved in ���S��� shape, dorsomedian apophysis absent. Epigyne triangular in ventral view, protruding in lateral view. Spermathecae not reaching over vulvar lateral chitinous structures. Type material Not examined. The exact position of the type localities is not known and the surrounding area is at the moment inaccessible (for details see Distribution). Material examined BOSNIA AND HERZEGOVINA: Popovo polje, Bjelu��ica: 1 ♂ (CBSS /AR 3701-1), 1 ♀ (CBSS /AR 3701-2), 1 ♂ (CBSS /AR 3690), 26 Oct. 2015, leg. Martina Pavlek; 3 ♀♀, 1 juv.. ♂, 1 juv.., 26 Oct. 2015, leg. Vedran Sudar (CBSS /AR 3692); 1 ♀ (CBSS /AR 1510), 1 ♂, 1 juv.. ♂, 1 juv.. (CBSS /AR 1513), 16 Aug. 2004, leg. Roman Ozimec; 1 ♀, 4 Jun. 2014, leg. Martina Pavlek (CBSS /AR 3389). Redescription Male (CBSS/AR 3701-1) Paracymbium with a well-developed dorsal and simple ventral process. Basal branch of dorsal process with straight upper rim, distal part longer and less sharp than in K. deelemanae gen. et sp. nov., more pronounced. Apical branch slender, curved in ���S��� shape, distal part acuminated and sharp (Figs 2B, 3E). Ventral process larger than in K. deelemanae gen. et sp. nov., broad and round, most of its surface transparent and with small membranous extension on rim. Dorsomedial apophysis absent (Fig. 3D). Tegulum with longitudinal ridge (in apical view) stretching through entire surface and reaching broad-based triangular basal apophysis (Figs 2B, 3F). Median apophysis well developed, V-shaped, distal arm reaching almost to the apical part of tegulum. Outer edge between proximal and distal part equally rounded, distal part of MA narrowing toward top, which ends bluntly with slightly serrated upper edge. Inner side of distal part of MA straight. Embolus, TTA p1 and TTA p2 of TTA similar to in K. deelemanae gen. et sp. nov. (Figs 2B, 3D). Female (CBSS/AR 3701-2) EPIGYNE. Slightly wider than long. Lateral lobes with oval depressions in posterior part, reaching middle of epigyne. Posterior epigyne border triangular in ventral view, made of two straight chitinous arches which flank depressions, come together in middle and after sharp inward turn form broad and pale convex tubercle (Figs 4B, 5 E���F), protruding in lateral view (Fig. 5H). Insemination ducts visible on both sides of epigyne in a form of chitinous curves (Figs 4B, 5 E���F). VULVA. Lateral sides of vulva made of compact dark chitinous structures enclosed in firm transparent membrane (lateral pouches). Spermathecae oval-shaped, with shallow constriction in middle, narrower in upper part and less slender then in K. deelemanae gen. et sp. nov., located meso-apicaly behind dark chitinous lateral structures. Insemination ducts depart from bottom of spermathecae, make sharp turn outwards and continue to epigynal base. Two chitinous narrow, horn-like structures protrude from inner sides of dark chitinuos vulval structures, in apical direction, to empty middle part of vulva (Fig. 5G). Distribution This species was described based on the material collected by Kratochv��l in several caves in Herzegovina, on the north-west border of Popovo polje: Pećina kod Tamnicam and Jama za Jamskim vrhom near Belenići village and Pčelina pećina near Nevada village (Kratochv��l 1933). The type material is deposited in the National Museum in Prague (Ružička et al. 2005). The area where the type localities are is inaccessible at the moment because of the mine fields left after the civil war in Bosnia and Herzegovina in the 1990s. The species is known from only one more locality, Bjelu��ica (Dresco 1966; Kratochv��l 1936), a horizontal cave around 4 km north of Belenići village (Fig. 6B). The material for this redescription was collected there. The species is endemic for Bosnia and Herzegovina and the citation of ���Montenegro��� in ���The Spiders of Europe��� (Nentwig et al. 2016) is erroneous. Natural history Bjelu��ica is a small cave, around 150 m long. From the semi-dark entrance chamber departs a narrowing, 1 �� 2 meters wide passage that leads to the main channel which, after the first few meters that are slightly lighter, is always in complete darkenss. The spiders were found right after the narrowing and all along the cave. They were mostly hanging from their nets on the walls and between the big stones on the cave floor, or freely walking on the walls of the cave. The temperature in the cave is around 13.5��C. In his paper with the description of K. fagei (Kratochv��l 1933), Kratochv��l also listed K. eremita at one of the type localities, Jama za Jamskim vrhom. Since it is not common for two species of the same genus to inhabit the same cave, we wanted to check the original material, but that was unfortunately impossible since the specimens deposited in the museum in Prague are without pedipalps (personal communication with Petr Dolej��, curator of the spider collection). In the last 20 years, researchers from CBSS have visited many caves around the town of Trebinje, in the Popovo polje area, and in the hilly region between Popovo polje and Croatia, but have so far recovered no K. eremita., Published as part of Pavlek, Martina & Ribera, Carles, 2017, Kryptonesticus deelemanae gen. et sp. nov. (Araneae, Nesticidae), with notes on the Mediterranean cave species, pp. 1-27 in European Journal of Taxonomy 262 on pages 12-14, DOI: 10.5852/ejt.2017.262, http://zenodo.org/record/3823118, {"references":["Kratochvil J. 1933. Evropske druhy celedi Nesticidae Dahl. Prace Moravske Prirodovedecke Spolecnosti 8: 1 - 69.","Gasparo F. & Thaler K. 2000. I ragni cavernicoli della Venezia Giulia (Italia nord-orientale) (Arachnida, Araneae). Atti e Memorie, Comm. Grotte \" Eugenio Boegan \" 37: 17 - 55.","Le Peru B. 2011. The Spiders of Europe, a synthesis of data. Volume 1 Atypidae to Theridiidae. Memoires de la Societe linneenne de Lyon 2: 1 - 522.","Lehtinen P. T. & Saaristo M. I. 1980. Spiders of the Oriental-Australian region. II. Nesticidae. Annales Zoologici Fennici 17: 47 - 66.","Kulczynski W. 1914. Aranearum species novae minusve cognate in montibus Kras dictis a Dre C. Absolon aliisque colectae. Bulletin international de l'Academie des Sciences de Cracovie 13: 353 - 387. Available from http: // biodiversitylibrary. org / page / 4553795 [accessed 17 Jan. 2017].","Ruzicka V., Kurka A., Buchar J. & Rezac M. 2005. Czech Republic - the type material of spiders (Araneae). Casopis Narodniho muzea, Rada prirodovedna 174: 13 - 64.","Dresco E. 1966. Etude de quelques especes d'araignees du genre Nesticus (Fam. Nesticidae). Annales de Speleologie 21: 795 - 813.","Kratochvil J. 1936. Nouveau genre d'araignees cavernicoles en Yugoslavie. Typhlonyphia reimoseri n. gen. n. sp. Vestnik Ceskoslovenske Zoologicke Spolecnosti 3: 69 - 79.","Nentwig W., Blick T., Gloor D., Hanggi A. & Kropf C. 2016. Spiders of Europe. Version 02 [online]. Available from http: // www. araneae. unibe. ch /"]}

Kryptonesticus gen. nov. urn:lsid:zoobank.org:act: 19 EABFE 7-08 F 7-4 AB 8-860 B- 0 AB 2 E 59 EBC 03 Figs 1���5 Type species Kryptonesticus deelemanae gen. et sp. nov. Diagnosis Kryptonesticus gen. nov. differs from other nesticid genera by the morphology of both male and female copulatory organs. In males, the most conspicuous diagnostic character is the shape and size of the median apophysis (MA), which is large and conspicuous, highly developed and thickly sclerotized, emerging from the middle part of the tegulum and reaching its apical part. This characteristic is shared among a group of species including K. eremita, K. dimensis, K. henderickxi, K. fagei and K. arenstorffi (males of K. beshkovi and K. biroi are unknown). In other nesticid genera, the MA is absent, inconspicuous or poorly developed (Nesticella Lehtinen & Saaristo, 1980; Wraios Ballarin & Li, 2015; Nescina Ballarin & Li, 2015; Hamus Ballarin & Li, 2015; Gaucelmus Keyserling, 1884; Cyclocarcina Komatsu, 1942; Typhlonesticus Kulczyński, 1914; Canarionesticus Wunderlich, 1992); in the form of a small, curved and pointed hook (Carpathonesticus Lehtinen & Saaristo, 1980); compact, transverse and not very developed (Eidmannella Roewer, 1935); more conspicuous and transverse (Aituaria Esyunin & Efimik, 1998) or it is well developed (Nesticus cellulanus (Clerck, 1757) and Pseudonesticus Liu & Li, 2013), but in this case the MA is long, thin and extended laterally. In Kryptonesticus gen. nov., the MA extends laterally at its base and then stretches towards the apex of the tegulum, parallel to the embolus. The theridioid tegular apophysis (TTA) is more compact than in N. cellulanus or Pseudonesticus. This is a very important character, since the TTA and MA are actively involved in sperm transfer (Huber 1993). Kryptonesticus gen. nov. also differs from other nesticid genera in the shape, ramification and modifications associated with the paracymbium. Kryptonesticus gen. nov. shows a well-developed dorsal process and a poorly differentiated ventral one. Distal and paradistal apophyses are absent and the dorsomedian apophysis is short and pointed, inconspicuous or absent. Females of this new genus show characteristic slender spermathecae, oval-shaped, with a shallow constriction in the middle, located above or just behind the apical part of the internal chitinous structures of the vulva. Insemination ducts depart from the basal part of the spermathecae, make a sharp turn-out and continue ventro-laterally, reaching the copulatory orifices. Etymology The prefix ��� Krypto ���, from Ancient Greek ����υ������ς (krupt��s ���hidden���), alludes to the long time it took to diagnose this evolutionary line. Distribution From Bulgaria and Turkey to Croatia, including Montenegro, Bosnia and Herzegovina and Crete. All these species are known only from the type locality, or have small distribution ranges. K. eremita is an exception; this species is linked to human activities and can be found in hangars, cellars and cottages. It has been cited from France and Italy to Bulgaria and Turkey. It is a potentially invasive species, found in an abandoned air-raid tunnel in Auckland, New Zealand (Vink & Dup��rr�� 2011). Taxonomic remarks The morphology of both male and female copulatory organs of this group of species shows important differences with respect to N. cellulanus (the type species of the genus Nesticus) and the western Mediterranean Nesticus species, as well as the representatives of Carpathonesticus and Typhlonesticus. The asymmetrical development of the paracymbial ramification, the shape, size and arrangement of the median apophysis and some differences in the size and arrangement of the TTA p1 and p2 processes constitute the major differences in males. The number and position of spermathecae and the features of the insemination ducts are the most obvious characteristics in females. K. beroni and K. beshkovi are only known from females. Regardless, the shape and position of the spemathecae and the insemination ducts undoubtedly allow us to transfer these two species to the Kryptonesticus evolutionary lineage. Concerning species of Nesticus from other parts of the world (e.g., East Asia, North and South America) we can state that they do not show the diagnostic characters of the new genus described here. Surely this is a polyphyletic group that could be considered as a wastebasket group, and its inclusion within the genus Nesticus must be revised since the morphology of their genitalia is remarkably different from the type species of the genus (N. cellulanus). Of course, this affirmation must be tested in a phylogenetic framework including all species of Nesticus described so far. Preliminary results on the phylogeny of European nesticids, including some American and Asian representatives (results not shown), indicate the polyphyly of Nesticus species described from other parts of the world. Composition Kryptonesticus deelemanae gen. et sp. nov., K. eremita (Simon, 1879) comb. nov. (Southern and Central Europe, introduced in New Zealand, potentially invasive species), K. arenstorffi (Kulczyński, 1914) comb. nov. (Bosnia and Herzegovina and Montenegro), K. fagei (Kratochv��l, 1933) comb. nov. (Bosnia and Herzegovina), K. beroni (Deltshev, 1977) comb. nov. (Bulgaria), K. beshkovi (Deltshev, 1979) comb. nov. (Crete), K. henderickxi (Bosselaers, 1998) comb. nov. (Crete) and K. dimensis (L��pez- Pancorbo, Kunt & Ribera, 2013) comb. nov. (Turkey)., Published as part of Pavlek, Martina & Ribera, Carles, 2017, Kryptonesticus deelemanae gen. et sp. nov. (Araneae, Nesticidae), with notes on the Mediterranean cave species, pp. 1-27 in European Journal of Taxonomy 262 on pages 5-6, DOI: 10.5852/ejt.2017.262, http://zenodo.org/record/3823118, {"references":["Lehtinen P. T. & Saaristo M. I. 1980. Spiders of the Oriental-Australian region. II. Nesticidae. Annales Zoologici Fennici 17: 47 - 66.","Kulczynski W. 1914. Aranearum species novae minusve cognate in montibus Kras dictis a Dre C. Absolon aliisque colectae. Bulletin international de l'Academie des Sciences de Cracovie 13: 353 - 387. Available from http: // biodiversitylibrary. org / page / 4553795 [accessed 17 Jan. 2017].","Huber B. A. 1993. Genital mechanics and sexual selection in the spider Nesticus - cellulanus (Araneae, Nesticidae). Canadian Journal of Zoology 71: 2437 - 2447. http: // dx. doi. org / 10.1139 / z 93 - 340","Vink C. J. & Duperre N. 2011. Nesticus eremita (Araneae: Nesticidae): redescription of a potentially invasive European spider found in New Zealand. Journal of Arachnology 39: 511 - 514. http: // dx. doi. org / 10.1636 / a 11 - 57.1","Kratochvil J. 1933. Evropske druhy celedi Nesticidae Dahl. Prace Moravske Prirodovedecke Spolecnosti 8: 1 - 69.","Deltshev C. 1977. Genus Nesticus (Nesticidae, Araneae) from Bulgarian caves. Congres international de Speleologie 6 (Olomouc, 1973): 73 - 78.","Deltshev C. D. 1979. A contribution to the study of cave spiders (Araneae) in Greece. Four new species (Araneae, Nesticidae, Linyphiidae) from the islands of Crete and Thera. Acta Zoologica Bulgarica 13: 53 - 63.","Bosselaers J. 1998. Nesticus henderickxi (Araneae, Nesticidae), a new blind troglobitic spider from Crete. Bulletin of the British Arachnological Society 11: 9 - 14."]}

session 01 - Diversity and faunistics; World experts of different disciplines, from molecular biology to macroecology, recognize the value of cave ecosystems as ideal ecological and evolutionary laboratories. Among other subterranean taxa, spiders stand out as intriguing model organisms for their ecological role of top-predators, their unique adaptations to the hypogean medium and their sensitivity to anthropogenic disturbance. Here, we provide a general overview of the spider families recorded in hypogean habitats in Europe–20 families including nearly 500 species, most of them with restricted distributions. We also review the different adaptations of hypogean spiders to subterranean life and summarize the information gathered so far about their origin, population structure, ecology and conservation status. Taxonomic knowledge on subterranean spiders in Europe appears to be well, but not exhaustively documented. The origin of the European assemblages is mostly explained by past climate dynamics, although other factors are likely to be involved. Most of the macroecological issues related to spiders in European caves are based on qualitative assessments or have been quantified only at a sub-regional scale. In order to shed light on cave spiders’ biogeography and the macroecological patterns driving the diversity of European subterranean spiders we created the CAWEB network, a spontaneous collaboration between subterranean arachnologists from 30 different European countries. We here present the team and provide some preliminary results, which highlight Southern Europe as an important hot-spot for the European subterranean spider diversity.