Holocnemus pluchei
Creators: name: Huber, Bernhard A.
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Araneae
Pholcidae
Holocnemus
Holocnemus pluchei

description: Holocnemus pluchei (Scopoli, 1763)
Figs 2, 5–6, 13–58
Aranea pluchii Scopoli, 1763: 404.
Aranea rivulata Forsskål, 1775: 86. Synonymized with Pholcus pluchii by Simon 1866: 122 (but: Simon 1873).
Pholcus impressus C.L. Koch, 1837: 99, pl. 137 fig. 313. Synonymized with H. rivulatus by Simon 1873: 49.
Pholcus barbarus Lucas, 1846: 237, pl. 15 fig. 1, 1a–f. Synonymized with H. rivulatus by Simon 1873: 49.
Pholcus ruralis Blackwall, 1858: 432. Synonymized by Simon 1873: 49.
Aranea rivulata – Forsskål 1776: 7, pl. 24 fig. f.
Aranea pluchii? (question mark in original publication) – Rossi 1790: 134.
Pholcus rivulatus – Savigny & Audouin 1826: 140, pl. 3 fig. 12. — Walckenaer 1837: 653. — Pickard- Cambridge 1872: 277; 1876: 566. — L. Koch 1875: 25 (specimens from Cairo).
Pholcus impressus – C.L. Koch 1850: 31. — Simon 1866: 122.
Pholcus pluchii – Simon 1866: 122, pl. 2 fig. 11.
Pholcus barbarus – Simon 1866: 123. — C. Koch 1873: 113.
Holocnemus rivulatus – Simon 1873: 49; 1874: 256; 1875: pl. 4 fig. 16; 1885: 28; 1893: 471; 1908: 427. — Damin 1900: 21. — Strand 1908: 93. — Lessert 1910: 89. — Franganillo 1925: 35; 1926c: 70.
Holocnemus pluchii – Roewer 1928: 120. — Kratochvíl 1932: 2, figs 1, 3–4. — Drensky 1936: 55; 1939: 248. — Kolosváry 1938: 65. — Senglet 1971: 354. — Timm 1976: 70, figs 1–2. — Nicolić & Polenec 1981: 20. — Kritscher 1996: 123.
Holocnemus pluchei – Simon 1914: 237. — Dalmas 1920: 59. — Wiehle 1933: 242, 247, figs 2a, 7c. — Kratochvíl 1940: 7, fig. 1b, d. — Caporiacco 1948: 38. — Roewer 1959: 10. — Brignoli 1971a: 82, figs 1–6; 1971b: 130; 1971c: 257; 1976: 559; 1977: 31; 1978: 486; 1979: 189; 1984: 289. — Constantini 1975: 90. — Wunderlich 1980: 227, figs 31–32. — Barrientos & Ferrández 1982: 82. — Roth 1985: B-33-1; 1994: 145. — Jakob & Dingle 1990: 95. — Porter & Jakob 1990: 313. — Jakob 1991: 711. — Heimer & Nentwig 1991: 40, fig. 80.1–3. — Melic 1994: 15. — Huber 1995: 291, figs 1a, 2, 4, 6–8, 10a; 2000: figs 14, 120, 176; 2001: 136; 2014: 140. — Jäger 1995: 20; 2000: 51. — Johnson & Jakob 1999: 957. — Van Keer & Van Keer 2001: 82; 2004: 79. — Deltshev et al. 2003: 12; 2011: 130; 2013: 7. — Gasparo 2003: 62. — Hajer & Řeháková 2003: 345. — Topçu et al. 2005: 289. — Ferrández et al. 2006: 77. — Kovács et al. 2006: 9, figs 1–3. — Van Keer 2007: 48. — Laborda & Simó 2008: 262. — Kunt et al. 2010: 32. — Calbacho-Rosa et al. 2010: 1267; 2013: 407; 2019a: 1, fig. 1; 2019b: 1, figs 1–3. — Van Helsdingen 2010: 27. — Drakšić & Katušić 2011: 168. — Dutto et al. 2011: 1040. — Vetter et al. 2011: 601. — Bosmans et al. 2013: 24. — Türkeş & Karabulut 2013: 619. — Benhadi-Marín et al. 2013: 75, fig. 2 (C1–3). — Reiser & Neumann 2014: 24. — Gajić & Grbić 2016: 54. — Naumova et al. 2016: 434; 2019: 76. — Rozwałka et al. 2016: 73. — Barrientos & Bosco Febrer 2017: 22. — Cargnelutti et al. 2018: 616; 2020: 536. — Lecigne 2018: 50, pl. 1 figs h–i, photo 3. — Ponomarev et al. 2019: 233, figs 2–4. — Kumada 2021: 17–19, figs 1–9.
Misidentifications
Pholcus rivulatus – L. Koch 1875: 25 (specimens from Massaua: Crossopriza sp. juveniles; see
Distribution below). Pholcus rivulatus – Leardi in Airaghi 1902: 348 (India; identity unclear; see Distribution below). Holocnemus pluchei – Roewer 1960: 40 (Afghanistan; juvenile from Rig-Revan: Pholcus sp.; for other
specimens, see Crossopriza ibnsinai sp. nov.).
Diagnosis
Easily distinguished from all other known Smeringopinae by enlarged female palps (Figs 29–30), by median process on female sternum (Figs 31, 58), and by pair of distinctive processes on male distal bulbal sclerite (Figs 24, 26); also by shape of procursus (Figs 20–22; distinct prolateral proximal process, short distal ventral sclerite), by female external and internal genitalia (Figs 27–28, 36–42; epigynum anteriorly with pair of sculptured areas; internal ventral arc with pair of strong pockets), and by heavily sclerotized female pedicel opposing pair of small sclerites on abdomen (Fig. 32).
Type material
Syntypes of Aranea pluchii SLOVENIA • Unknown number; “ Carniola ” [part of present-day Slovenia], possibly Idria [Scopoli’s place of residence at the time]; ~ 46.00° N, 14.03° E; 1759–1762; G.A. Scopoli leg.; probably lost.
Syntypes of Aranea rivulata EGYPT • Unknown number; Cairo; ~ 30.0° N, 31.2° E; probably 1762; P. Forsskål leg.; probably lost.
Syntypes of Pholcus impressus GREECE • 4 ♂♂; Peloponnese, Nafplion (“Nauplion”); 37.57° N, 22.81° E; date unknown; [F.J.?] Schuch leg.; probably lost.
Syntypes of Pholcus barbarus (examined) ALGERIA • 4 ♂♂, 4 ♀♀, + juvs (possible syntypes); near Algiers; 36.75° N, 3.05° E; 1839–1842; P.- H. Lucas leg.; with two labels: “Algerie, Lucas, Holocnemus rivulatus Forsk. sub barbarus Type” and “ Holocnemus rivulatus Forsk (sub barbarus) type, Algerie, Lucas”; MNHN.
Syntypes of Pholcus ruralis ALGERIA • Unknown number (possibly only 1 ♂); locality not specified; possibly 1856; Gray and Clarck leg.; probably lost.
Material examined
Arranged from West to East and (within longitudes) from North to South.
USA – California • 1 ♂; Sonoma County, Petaluma; 38.224° N, 122.626° W; 15 Jul. 2002; C.E. Griswold leg.; on house; CAS 9027116 part • 1 ♀; Napa County, Lake Hennessey, below Lonn Dam; 38.48° N, 122.38° W; 11 May 1985; L. Vincent leg.; CAS 9027422 • 1 ♀; Los Angeles County, Diamond Bar; 34.03° N, 117.80° W; 23 Sep. 1999; C. Yang leg.; CAS 9027418 • 1 ♀; Orange County, Yorba Linda; 33.889° N, 117.812° W; 2 May 2000; S. Clayton leg.; CAS 9027393 • 1 ♀; San Diego County, northern end of Harbison Canyon; 32.811° N, 116.844° W; 250 m a.s.l.; 7 Jul. 2012; under stones and wood panels in scrubland; A. Schönhofer leg.; SMF.
ARGENTINA – Mendoza • 1 ♀; Mendoza; 32.89° S, 68.84° W; 900 m a.s.l.; 30–31 Mar. 1965; H. Levi leg.; MCZ. – San Luis • 1 ♂; Naschel; 32.915° S, 65.375° W; Feb.–Mar. 1962; A. Luchini leg.; MACN 19955. – La Pampa • 1 ♀; Rancul; 35.07° S, 64.68° W; Jan. 1975; Maury leg.; MACN 20101 • 1 ♀; Santa Rosa [de Toay]; 36.62° S, 64.29° W; date and collector unknown; MACN 20039. – Buenos Aires • 1 ♂; Carlos Casares; 35.62° S, 61.36° W; 27 Feb.–2 Mar. 1980; collector unknown; MACN 20087 • 1 ♂, 2 ♀♀, 2 juvs; Buenos Aires, Cerrito; 34.60° S, 58.38° W; 19 Sep. 1972; C. Cesari leg.; in basement; MACN 20080.
PORTUGAL – Faro • 2 ♂♂, 1 ♀; Silves [Munic.], Norinha; 37.198° N, 8.400° W; 14 Sep. 1969; A. Senglet leg.; MHNG.
SPAIN – Extremadura • 1 ♂; Cáceres, Jarandilla [de la Vera]; 40.126° N, 5.658° W; 22–23 Aug. 1969; A. Senglet leg.; MHNG. – Castilla-La Mancha • 2 ♂♂, 4 ♀♀, 9 juvs; Toledo, Escalona del Alberche; 40.167° N, 4.400° W; 14 Jun. 1969; A. Senglet leg.; MHNG. – Andalusia • 6 ♂♂, 7 ♀♀, 2 juvs; Córdoba, Palma del Río; 37.70° N, 5.28° W; 26–27 Jun. 1969; A. Senglet leg.; MHNG • 2 ♂♂, 2 ♀♀; Córdoba, Peñarroya; 38.30° N, 5.27° W; 30 Jun. 1969; A. Senglet leg.; MHNG • 2 ♀♀; Cádiz, Algodonales; 36.88° N, 5.40° W; 19 Jul. 1969; A. Senglet leg.; MHNG • 3 ♂♂, 2 ♀♀, 5 juvs; Málaga, Estepona; 36.43° N, 5.15° W; 24–26 Jul. 1969; A. Senglet leg.; MHNG • 2 ♀♀; Málaga, Valle de Abdalajís; 36.93° N, 4.68° W; 28 Jul. 1969; A. Senglet leg.; MHNG • 4 ♀♀; Málaga, Antequera; 37.02° N, 4.56° W; 1–2 Aug. 1969; A. Senglet leg.; MHNG • 3 ♂♂, 5 ♀♀; Málaga, Torre del Mar; 36.75° N, 4.10° W; 27 Jul. 1969; A. Senglet leg.; MHNG • 2 ♀♀; Granada, Ugíjar; 36.96° N, 3.05° W; 15 Jul. 1971; A. Senglet leg.; MHNG • 3 ♀♀; Granada, Huéneja; 37.176° N, 2.948° W; 16 Jul. 1971; A. Senglet leg.; MHNG • 1 ♀; Sevilla; 37.39° N, 5.98° W; 16 Jun. 1953; A. Comellini leg.; MHNG. – Region of Murcia • 1 ♀; Caravaca [de la Cruz]; 38.104° N, 1.860° W; 5 Jul. 1971; A. Senglet leg.; MHNG • 1 ♀; San Pedro del Pinatar; 37.83° N, 0.78° W; Sep. 1984; G. Hormiga leg.; USNM • 1 ♂; Fortuna, Cueva Las Magras; 38.290° N, 1.143° W; 30 Jan. 2006; “ACP” leg.; CRBA 1146. – Valencian Community • 4 ♂♂, 3 ♀♀; Valencia, La Albufera; 39.30° N, 0.33° W; 16 Jun. 1971; A. Senglet leg.; MHNG • 1 ♂; Castellón, N of L’Alcora; 40.090° N, 0.199° W; 300 m a.s.l.; 30 May 2010; S. Huber and A. Schönhofer leg.; at stone walls; SMF. – Catalonia • 1 ♂; Barcelona, Gelida; 41.44° N, 1.86° E; 14 Jun. 1971; A. Senglet leg.; MHNG • 2 ♂♂, 1 ♀, 3 juvs; near Barcelona, Cerdanyola [del Vallès]; 41.49° N, 2.14° E; 7 Dec. 2004; D. Dimitrov leg.; SMF 56746.
FRANCE – Pyrénées-Orientales • 1 ♂, 2 ♀♀; Banyuls; 42.48° N, 3.13° E; date and collector unknown; MNHN Ar 13337 • 2 ♂♂, 3 ♀♀ (3 vials); same locality as for preceding; 9–20 May 1952; A. Comellini leg.; MHNG • 1 ♀; same locality as for preceding; no further data; MHNG. • 1 ♀; Cerbère; 42.44° N, 3.165° E; 11 May 1952; A. Comellini leg.; MHNG. – Var • 1 ♂, 1 ♀; Bormes-les-Mimosas (“ Bornes ”); 43.15° N, 6.34° E; Jul. 1960; O. Kraus leg.; SMF 24800. – Gard • 3 ♀♀; Aigues-Mortes; 43.57° N, 4.19° E; 8 May 1952; A Comellini leg.; MHNG. – Corsica • 1 ♂, 1 ♀; Propriano; 41.67° N, 8.91° E; 29 May 1971; A. Senglet leg.; MHNG • 1 ♂, 7 ♀♀, 1 juv.; Ponte Leccia; 42.463° N, 9.207° E; 19 Sep. 1952; Kahmann leg.; SMF 8799, 8800 • 1 ♀; Biguglia; 42.62° N, 9.44° E; 30 m a.s.l.; 2 May 1952; Kahmann leg.; SMF 8801.
BELGIUM – East Flanders • 1 ♀; near Gent; 51.10° N, 3.70° E; Jul. 2004; B.A. Huber leg.; ZFMK Ar 12608.
GERMANY – Nordrhein-Westfalen • 1 ♂, 4 ♀♀, 6 juvs; Bonn, Botanical Garden (?) (“Treibhaus”); 50.724° N, 7.091° E; Nov. 1967; E. Kullmann leg.; ZFMK Ar 5219. – Rheinland-Pfalz • 1 ♀; Hatzenport; 50.2265° N, 7.4108° E; 80 m a.s.l.; 26 Sep. 2014; B.A. Huber leg.; among wooden boards outside of building; ZFMK Ar 12304 • 1 ♀; Mainz, Draisberghof; 48.987° N, 8.203° E; 1 Feb. 1998; P. Jäger leg.; SMF 40411. – Hamburg • 1 ♂; Hamburg Mitte, Hamm; 53.5481° N, 10.0381° E; 25 m a.s.l.; 6 Jun. 2019; B.A. Huber leg.; in building; ZFMK Ar 22292.
AUSTRIA – Vienna • 1 ♂, 3 ♀♀ (in pure ethanol); Brigittenau; 48.2283° N, 16.3710° E; Aug. 2001; B.A. Huber leg.; at windows in attic (fifth floor); ZFMK G64 to G66.
ITALY – Sardinia • 1 ♂; S. Pietro; 40.717° N, 8.550° E; 15 Aug. 1960; E. Kullmann leg.; ZFMK Ar 10705 • 1 ♀, 1 juv.; Castelsardo; 40.91° N, 8.71° E; 11 Jun. 1958; E. Kullmann leg.; ZFMK Ar 10746 • 1 ♀; Lago Bunnari; 40.716° N, 8.634° E; 11 Apr. 1952; collector unknown; SMF 9273 • 1 ♂, 2 juvs; Ozieri; 40.58° N, 9.00° E; 23 Sep. 1968; A. Senglet leg.; MHNG • 2 ♀♀, 1 juv.; Terralba; 39.72° N, 8.63° E; 4 Aug. 1960; E. Kullmann leg.; ZFMK Ar 10696 • 1 ♂, 1 ♀, 1 juv.; Muravera, Nuraghe; 39.318° N, 9.531° E; 19 Aug. 1960; E. Kullmann leg.; ZFMK Ar 8214 • 4 ♂♂, 5 ♀♀, 2 juvs; Sardinia, no further data; SMF RII 6384. – Apulia • 2 ♂♂; south side of Monte Gargano near Manfredonia; 41.638° N, 15.890° E; 60 m a.s.l.; 7 Apr. 2006; A. Schönhofer and J. Hillen leg.; under stones in cultivated field, Opuntia field, native rock; SMF. – Sicily • 1 ♂; Syracuse, Viale Ermocrate; 37.07° N, 15.27° E; 18 Aug. 1993; B.A. Huber leg.; ZFMK Ar 5217 • 1 ♂, 1 ♀; Syracuse, Theatre; 37.074° N, 15.279° E; 18 Aug. 1993; B.A. Huber leg.; ZFMK Ar 5218.
CROATIA • 1 ♀; Karlovac, Ogulin; 45.266° N, 15.233° E; 320 m a.s.l.; 9 May 2010; J. Altmann and J. Meier leg.; in room; SMF • 6 ♂♂, 1 ♀; Ploče; 43.05° N, 17.43° E; 21 May 1972; A. Senglet leg.; MHNG • 1 ♀; Dubrovnik, Mt Srđ slopes; 42.65° N, 18.10° E; 0–400 m a.s.l.; 21 Apr. 1971; J. Murphy leg.; MMUE Murphy #5477.
BOSNIA AND HERZEGOVINA • 1 ♂, 2 ♀♀; Popovo-Polje, Zavala; 42.848° N, 17.979° E; 19 Sep. 1970; A. Senglet leg.; MHNG • 2 ♂♂, 2 ♀♀; Popovo-Polje, Sedlari; 42.826° N, 18.056° E; 18 Sep. 1970; A. Senglet leg.; MHNG • 2 ♂♂ (in pure ethanol); near Trebinje, near Pavlova pećina; 42.666° N, 18.308° E; 400 m a.s.l.; 26 May 2014; B.A. Huber and M. Komnenov leg.; under rocks; ZFMK Bal10, Bal19.
MONTENEGRO • 1 ♂, 4 ♀♀; Podgorica, Velje Brdo, Megara Cave; 42.463° N, 19.199° E; 17 Feb. 2010; M. Komnenov leg.; CMK • 2 ♀♀; Podgorica, Ljuboviči; 42.433° N, 19.255° E; 21 Apr. 2011; M. Komnenov leg.; artificial tunnel; CMK.
ALBANIA • 1 ♀; Kulla ë Lumës (“Kula Lums”); 42.098° N, 20.418° E; 1918; collector unknown; SMF RII 3726 • 1 ♀, 1 juv.; Durrës; 41.33° N, 19.45° E; 26 May 1993; P. Stoev and C. Deltshev leg.; NMNHS • 1 ♀; Përmet, near Petrani; 40.208° N, 20.419° E; 12 May 1995; B. Petrov and P. Stoev leg.; artificial gallery; NMNHS.
NORTH MACEDONIA • 1 ♀; Gevgelija; 41.14° N, 22.50° E; 10 Jul. 1956; Lamel leg.; HECO.
BULGARIA – Blagoevgrad • 1 ♂, 1 ♀, 1 juv.; Blagoevgrad; 42.014° N, 23.097° E; 380 m a.s.l.; 2 Aug. 2005; P. Jäger leg.; SMF 40643 • 6 ♀♀; N foothills of Krupnishna Planina Mt Range, ~ 0.2 km NNE of Krupnik; 41.850° N, 23.125° E; 375 m a.s.l.; 12 Aug. 2005; A.V. Gromov leg.; ZMMU • 1 ♂, 2 ♀♀; Sandanski; 41.566° N, 23.280° E; 28 Jul. 1972; A. Senglet leg.; MHNG • 1 ♂, 3 ♀♀; ~ 0.5 km S of Banya; 41.873° N, 23.528° E; 825 m a.s.l.; 11 Aug. 2005; A.V. Gromov leg.; ZMMU. – Pazardzhik • 2 ♂♂, 1 ♀; Panagyuriste; 42.505° N, 24.186° E; 720 m a.s.l.; 1 Jul. 1997; S. Lazarov leg.; NMNHS. – Ruse • 1 ♂, 2 ♀♀; Byala; 43.46° N, 25.74° E; 2013; collector unknown; MGAB.
GREECE – Ionian Islands • 1 ♂; Corfu, Agios Stephanos; 39.79° N, 19.81° E; 26 May 1999; A. Russell-Smith leg.; ZFMK Ar 22293 • 1 ♂, 1 ♀; Corfu, Loutses; 39.786° N, 19.880° E; 20 Sep. 1972; A. Senglet leg.; MHNG • 3 ♂♂, 1 ♀; Corfu, Agios Matheos; 39.495° N, 19.872° E; 18 Sep. 1972; A. Senglet leg.; MHNG • 1 ♂, 1 ♀; Pantocrator, Spilaio Katsampa near Strinilas; 39.744° N, 19.839° E; 650 m a.s.l.; 12 May 1974; B. Hauser leg.; MHNG • 1 ♀; Pantocrator, Spilaio Katsouri; 39.743° N, 19.925° E; 130 m a.s.l.; 29 Apr. 1975; D. Tzanoudakis leg.; MHNG • 1 ♂, 1 ♀; Lefkada, W of Mikros Gialos; 38.64° N, 20.69° E; 25 May 1995; A. Russell-Smith leg.; ZFMK Ar 22294 • 8 ♂♂, 3 ♀♀; Cephalonia, Sisia; 38.10° N, 20.66° E; 29 May 1987; J.A. Murphy leg.; MMUE Murphy #A231 • 1 ♂, 2 ♀♀; same collection data as for preceding; reared from eggs; MMUE Murphy #15938 • 1 ♀; Cephalonia, “Patra”; date unknown; Malicky leg.; SMF • 1 ♀; Cephalonia, “Castle Hill”; 19 May 1987; J.A. Murphy leg.; MMUE Murphy #15013 • 1 ♂, 1 ♀; Cephalonia, no further locality data; 21 May 1987; J.A Murphy leg.; MMUE Murphy #16161. – Epirus • 1 ♀, 1 juv.; Igoumenitsa; 39.50° N, 20.27° E; data unknown; F. Sauer leg.; SMF. – Central Macedonia • 1 ♂; Yerakini (“ Gerakina ”); 40.27° N, 23.45° E; 14 Apr. 1978; J.A. Murphy leg.; stony area; MMUE Murphy #6830 • 2 ♀♀; “ Halkidiki ”; 6 Apr. 1978; J.A. Murphy leg.; MMUE Murphy #6709. – Eastern Macedonia and Thrace • 1 ♀; Limni Vistonis (“ Limni Bouroú ”); 41.05° N, 25.10° E; Sep. 1984; collector unknown; SMF. – Thessaly • 2 ♂♂, 2 ♀♀; Larissa, “Omôlion (Tembe)”; 39.88° N, 22.58° E (coordinates between Omolio and Tempi); 17 Jun. 1970; A. Senglet leg.; MHNG. – Central Greece • 1 ♂; Euboea, Lichada / Istiaia; 38.9° N, 23.0° E (coordinates between Lichada and Istiaia); 30 Aug. 1972; A. Senglet leg.; MHNG • 3 ♂♂, 9 ♀♀, 2 juvs; Phthiotis, Tragána; 38.62° N, 23.12° E; 21 May 1968; A. Senglet leg.; MHNG • 2 ♂♂, 2 ♀♀, 1 juv.; Phthiotis, Theologus / Atalanti (Glyphada); 38.62° N, 23.10° E (coordinates between Theologos and Atalanti); 20 Jun. 1970; A. Senglet leg.; MHNG • 1 ♂, 1 ♀; Phthiotis, near Malesina; 38.62° N, 23.22° E; 21 May 1968; A. Senglet leg.; MHNG • 1 ♂, 2 ♀♀; Boeotia, “Topôlia/Thiva”; 38.32° N, 23.32° E (coordinates of Thiva); 25 Aug. 1970; A. Senglet leg.; MHNG • 1 ♂; Euboea, Kato Steni; 38.571° N, 23.825° E; 2 Sep. 1972; A. Senglet leg.; MHNG • 1 ♂, 1 juv.; Northern Sporades, Skyros Island; 38.90° N, 24.56° E; 22 Mar. 1958; Schelkopf leg.; SMF. – Attica • 5 ♂♂, 4 ♀♀ (partly used for SEM); Daphni / Athens; 38.013° N, 23.636° E; 23 Jun. 1970; A. Senglet leg.; MHNG • 1 ♂, 1 ♀; same locality as for preceding; 19 Jun. 1968; A. Senglet leg.; MHNG • 2 ♀♀, 1 juv.; Athens, Akropolis; 37.971° N, 23.727° E; date and collector unknown; SMF RII 3724 • 4 juvs; same locality as for preceding; May 1926; F. Roewer leg.; SMF • 3 ♂♂, 2 ♀♀; Athens, near Koropi; 37.900° N, 23.867° E; 19 Mar. 2007; A. Schönhofer leg.; under stones in olive grove and phrygana; SMF • 1 ♀; Vari, Spilaio Nympholyptou; 37.858° N, 23.802° E; 25 Feb. 2018; N. Fytrou and E. Zenzefyli leg.; HISR 3042 • 3 ♀♀, 1 juv.; near Var, “ Havara ”; 27 Jan. 1954; P. Strinati leg.; MHNG. – Peloponnese • 2 ♂♂, 4 ♀♀; Tripolis; 37.51° N, 22.37° E; Jun. 1926; F. Roewer leg.; SMF • 1 ♂, 5 ♀♀; Gulf of Argolis, Kantia (“ Argolide, Kandia ”); 37.52° N, 22.96° E; 12 Aug. 1970; A. Senglet leg.; MHNG • 1 ♂, 2 ♀♀, 1 juv.; NE of Gytheio (“ Yithion ”); 1977; Kinzelbach leg.; SMF 37616 • 1 ♀; Laconia, Pyrgos Dirou; 36.625° N, 22.380° E; 18 Aug. 1970; A. Senglet leg.; MHNG. – North Aegean • 2 ♂♂; Chios, Emboria Beach; 38.355° N, 26.143° E; 8 May 2006; A. Russell-Smith leg.; in public fountain; ZFMK Ar 22295. – South Aegean • 4 ♀♀; Paros, Naousa; 37.12° N, 25.24° E; 21 Jun. 1968; A. Senglet leg.; MHNG • 2 ♂♂; Paros, Parikia E, “Moni Aghii Anarghii”; 37.08° N, 25.16° E; 10 Aug. 2006; R. Bosmans leg.; on walls; CRB • 2 ♂♂, 2 ♀♀; Naxos, Chalkio (“ Chalkis ”); 37.063° N, 25.482° E; 27 Jun. 1968; A. Senglet leg.; MHNG • 2 ♀♀; Karpathos, near Pigadia; 35.51° N, 27.20° E; Apr. 1963; Kinzelbach leg.; SMF 15489 • 2 ♂♂, 1 ♀; Rhodes, “ rocca di Lindos ”; 36.091° N, 28.088° E; 2 Jul. 1996; F. Gasparo leg.; ZFMK Ar 22296. – Crete • 1 ♂, 2 ♀♀; Chania, Azogires Palaeochoras; 35.272° N, 23.719° E; 4–6 Aug. 1970; A. Senglet leg.; MHNG • 1 ♂; Chania, Sougla; 35.25° N, 23.81° E; 9 Oct. 1999; A. Senglet leg.; MHNG • 3 ♀♀; Chania, Halepa (“ Chalepa ”); 35.52° N, 24.05° E; date and collector unknown; SMF RII 3725 • 1 ♂, 1 ♀; Chania, Akrotiri, Kumaro Caves; 35.579° N, 24.143° E; May 1926; F. Roewer leg.; SMF • 1 ♂; Chania, Georgioupoli; 35.36° N, 24.26° E; 9 Jun. 2005; W. Schedl leg.; ZFMK Ar 22297 • 1 ♀; Chania, edge of Lake Kourna; 35.33° N, 24.28° E; 28 May 1993; A. Russell-Smith leg.; dense scrub, under stone; ZFMK Ar 22298 • 1 ♂, 3 ♀♀; Chania, Kournas; 35.327° N, 24.279° E; 10 m a.s.l.; 23 Mar. 2007; A. Schönhofer leg.; under stones, mostly open terrain; SMF • 1 ♂, 1 ♀; Chania, Grambousa, Spilaio Agiou Ioanni Prodromou; 5 Mar. 1995; K. Paragamian leg.; HISR 1773 • 1 ♂, 1 ♀, 2 juvs; Chania, Hora Sfakion (“ Sfakia ”); 35.20° N, 24.14° E; 17 Jul. 2009; H. Eikamp and U. Kluge leg.; SMF • 1 ♂, 1 ♀; Rethymno, Choumeri / Perama; 35.35° N, 24.72° E; 1 Jul. 1970; A. Senglet leg.; MHNG • 1 ♀; Rethymno, Anogia; 35.24° N, 24.87° E; 11 May 2006; H. Eikamp and U. Kluge leg.; SMF 56588 • 3 ♂♂, 3 juvs; Rethymno, Mpali, Anonymo Spilaio (National Road); 2 Nov. 2003; K. Foteinakis and K. Paragamian leg.; HISR 2428 • 1 ♂, 4 juvs; Heraklion, Gournes; 35.32° N, 25.28° E; 3 May 2008; H. Eikamp and U. Kluge leg.; SMF • 1 ♀; same locality as for preceding; 15 May 2010; K. Eckl leg.; SMF • 1 ♂, 2 ♀♀; Heraklion, Rodia (“ Rogdia ”); 35.365° N, 25.020° E; 27 Jun. 1970; A. Senglet leg.; MHNG • 2 ♂♂, 2 ♀♀; Heraklion, Knossos; 35.30° N, 25.16° E; 25 Jun. 1970; A. Senglet leg.; MHNG • 3 ♂♂, 5 ♀♀; Heraklion, Malia; 35.29° N, 25.46° E; 8 Jul. 1970; A. Senglet leg.; MHNG • 1 ♂; same locality as for preceding; 11 May 2008; H. Eikamp and U. Kluge leg.; SMF • 2 ♂♂, 1 ♀; Heraklion, Charakas; 35.01° N, 25.12° E; 24 Jul. 1970; A. Senglet leg.; MHNG • 1 ♂, 1 ♀; Heraklion, Marathos, Arkalospilios; 35.245° N, 24.968° E; 24 Dec. 2000; K. Paragamian leg.; HISR 1971 part • 1 ♀, 6 juvs; Heraklion, Kastri; 35.00° N, 25.37° E; 7 May 2008; H. Eikamp and U. Kluge leg.; SMF • 1 ♀; Heraklion, Kastri, Keratokambos; 35.000° N, 25.375° E; 14 May 2006; H. Eikamp and U. Kluge leg.; SMF 56589 • 1 ♂; Lasithi, Plati; 35.17° N, 25.44° E; 17 Dec. 2006; H. Eikamp and U. Kluge leg.; SMF • 1 ♂; Lasithi, Atsiganospilios; 35.205° N, 25.618° E; 16 Apr. 2003; F. Gasparo leg.; ZFMK Ar 22299 • 1 ♂, 2 ♀♀, 1 juv.; Lasithi, Agia Fotia; 35.02° N, 25.87° E; 6 Jul. 2007; H. Eikamp and U. Kluge leg.; SMF 57161 • 2 ♀♀; Lasithi, Bembonas, Tripti Mts; 35.1° N, 25.9° E; 10 May 2009; K. Eckl and H. Eikamp leg.; SMF • 5 ♀♀; Lasithi, Exo Mouliana; 35.17° N, 25.99° E; 18 Jul. 1970; A. Senglet leg.; MHNG.
TURKEY – Çanakkale • 1 ♀, 1 juv.; Kavak River, near bridge of road Keşan-Gallipoli (“ Gelibolu ”); 40.60° N, 26.87° E; date and collector unknown; SMF • 1 ♀; Truva, “84/15”; 39.95° N, 26.24° E; date and collector unknown; SMF. – Izmir • 1 ♀; Vişneli, “Fetrek-2 Cave”; 38.346° N, 27.421° E; 310 m a.s.l.; 5 Jun. 2009; Y.M. Marusik leg.; ZFMK Ar 5469 • 1 ♂, 1 ♀; Çiğli (airport); 38.516° N, 27.013° E; 15 Apr. 1973; A Vigna leg.; MHNG • 1 ♀; “ Agamemnon ”; 10 May 1975; C. Bisuchet and I. Löbl leg.; MHNG. – Aydin • 3 ♂♂, 4 ♀♀; Güzelçamli; 37.695° N, 27.163° E; 105 m a.s.l.; 7 Jun. 2009; Y.M. Marusik leg.; ZFMK Ar 5215. – Bursa • 2 ♀♀, 1 juv.; Görükle, Uludağ Univ. Campus; 40.226° N, 28.869° E; 420 m a.s.l.; 2–3 Jun. 2009; Y.M. Marusik leg.; ZFMK Ar 5468. – Muğla • 1 ♂, 1 ♀; Dalyan to Kaunos (“ Cavnos ”) path; 36.82° N, 28.61° E; 18 May 1997; A. Russell-Smith leg.; ZFMK Ar 22300 • 1 ♂; Marmaris; 36.85° N, 28.26° E; Sep. 1973; C. and E. Supper leg.; MHNG. – Antalya • 1 ♂; Xanthos ruins; 36.356° N, 29.318° E; 7 Mar. 1977; Kinzelbach leg.; SMF 37622 • 1 ♂ (in pure ethanol); Kemer District, Çıralı; 36.399° N, 30.475° E; 10 m a.s.l.; 3 Aug. 2016; H. Öztürk leg.; in building; ZFMK Tur64 • 1 ♂; İbradı District, Altınbeşik Mağarası; 37.039° N, 31.631° E; 660 m a.s.l.; 23 Jul. 2016; H. Öztürk leg.; among rocks near cave entrance; ZFMK Ar 22301 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Tur46 • 1 ♂, 3 ♀♀; Alanya Kalesi, above Kleopatra beach; 36.539° N, 31.992° E; 2 Nov. 2013; S. Huber leg.; ZFMK Ar 22302 • 1 ♂, 1 ♀; Alanya District, Cüceler Mağarası; 36.489° N, 32.276° E; 270 m a.s.l.; 31 Jul. 2016; H. Öztürk leg.; among rocks near cave entrance; ZFMK Ar 22303. – Mersin • 1 ♂; Anamur District, Köşekbükü Astım Mağarası; 36.127° N, 32.760° E; 130 m a.s.l.; 29 Jul. 2016; H. Öztürk leg.; in cave near entrance; ZFMK Ar 22304 • 1 ♀; NE of Silifke, above Narlıkuyu; 36.45° N, 34.10° E; 24 Apr. 2012; J. Altmann and J. Meier leg.; maquis; SMF • 2 ♀♀, 1 juv.; Içel Narlıkuyu; 36.44° N, 34.11° E; 5 May 1994; “L.M.” leg.; in rubbish by restaurant; WML • 1 ♂, 1 ♀; Taş Obası (cave); 37.079° N, 34.928° E; 24 Aug. 2007; A. Topçu leg.; NOHUAM. – Nevşehir • 2 ♂♂; Özkonak; 38.81° N, 34.84° E; 16 Jun. 1993; C. Felton leg.; WML. – Diyarbakır • 2 ♂♂; Çermik (“ Germik ”); 38.13° N, 39.45° E; 27 Jul. 2004; T. Danışman leg.; ZFMK Ar 5216.
CYPRUS – Limassol • 1 ♂, 1 ♀, 1 juv.; Pissouri; 34.67° N, 32.70° E; 27 Apr. 1982; J. Murphy leg.; beach litter; MMUE Murphy #12617. – Famagusta • 2 ♀♀, 3 juvs; Fig Tree Bay, “Cave I”; 35.01° N, 34.06° E; 23 Oct. 1996; P. Strinati leg.; MHNG.
GEORGIA – Tbilisi • 1 ♀; Tbilisi, Dighom I; 41.778° N, 44.701° E; 17 Jul. 2019; Karalashvili, Seropian and Krammer leg.; ZFMK Ar 21279. – Kakheti • 2 ♂♂, 3 ♀♀, 5 juvs; Sighnaghi; 41.621° N, 45.918° E; 9 Jul. 2019; Karalashvili and Krammer leg.; on houses; ZFMK Ar 21372, Ar 21373 • 1 ♂; Lagodekhi National Park; 41.847° N, 46.284° E; 9 Jul. 2019; Karalashvili and Krammer leg.; forest; ZFMK Ar 21261 • 1 ♂, 1 ♀, 2 juvs; Vashlovani National Park, bungalows near border to Azerbaijan; 41.111° N, 46.647° E; 26–27 Jul. 2019; Karalashvili and Krammer leg.; steppe; ZFMK Ar 21569, Ar 21570.
SYRIA • 1 ♀, 1 juv.; Latakia, ruins of Ugarit, Ra’s as-Samra, creek N of Tell Ugarit; 35.602° N, 35.783° E; 4 Mar. 1979; R. Kinzelbach leg.; SMF • 4 ♀♀, 3 juvs; Jayrud (“Djeroud”); 33.807° N, 36.742° E; 1911; H. Gadeau de Kerville leg.; in house; MNHN • 1 ♂; between Hamaa and Aleppo, “ Kaha al Suban ” (locality not identified); 500 m a.s.l.; 9 Apr. 1982; collector unknown; SMF.
LEBANON • 1 ♂; Antelias; 33.92° N, 35.59° E; May 1952; K. Christiansen leg.; MCZ 34059.
ISRAEL • 3 ♀♀, 1 juv.; Ahihud, E Akko[n]; 32.91° N, 35.17° E; 10 Apr. 1987; W. Heinz leg.; SMF • 1 ♀; Haifa Distr., Karmiya Ridge, Mt Karmel; 32.7185° N, 35.0065° E; 280 m a.s.l.; 17 Sep. 2013; B.A. Huber, S. Aharon and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22305 • 1 ♀ (in pure ethanol); Haifa Distr., Mt Karmel, Oren Cave; 32.7145° N, 34.975° E; 70 m a.s.l.; 17 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; in cave; ZFMK Isr57 • 2 ♂♂, 1 ♀; N of Teverya (“ Tiberias ”); 32.80° N, 35.52° E; stony meadow; 4 Mar. 1975; H. Levi, G. Levy, and G. Tsabor leg.; MCZ 34062 • 1 ♂, 1 ♀; Northern Distr., Mt Berenice, S of Teverya; 32.777° N, 35.541° E; - 120 m b.s.l.; 16 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22306 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Isr39 • 1 ♂, 1 ♀; Northern Distr., near Afula, HaGilbo’a Reserve; 32.52° N, 35.38° E; 300–500 m a.s.l.; 7 Apr. 1987; W. Heinz leg.; SMF • 1 ♀; Nof Yam; 32.19° N, 34.81° E; 17 Feb. 1968; collector unknown; AMNH • 1 ♂; Judea and Samaria Distr., Memorial Ha-Biq’a, NE Peza’el [Fatsa’el]; 32.0524° N, 35.4589° E; - 210 m b.s.l.; 15 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; in cave; ZFMK Ar 22307 • 1 ♀; near Ramla, Kibbuz Gezer; 31.88° N, 34.92° E; 11 Aug. 1984; H. Feldmann leg.; indoors on ceiling; AMNH • 1 ♀; Central Distr., Modi’in; 31.8951° N, 34.9618° E; 160 m a.s.l.; 8 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22308 • 1 ♀, 2 juvs; near Ramallah, Bayt Ghur al-Tahta; 31.895° N, 35.080° E; 6 Apr. 1987; W. Heinz leg.; SMF • 1 ♂; Judea and Samaria Distr., ’ En Perat, Nahal Perat [Prat]; 31.833° N, 35.303°– 35.311° E; 260–300 m a.s.l.; 15 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22309 • 1 ♀; Jerusalem; 31.77° N, 35.21° E; Apr. 1952; Khuri leg.; MCZ 34064 • 2 ♂♂, 2 ♀♀, 2 juvs; 21 km E of Jerusalem, Jericho Road, wadi in hills; 23 Feb. 1975; H.W. Levi and G. Levy leg.; MCZ 34067 • 1 ♀, 3 juvs; Hebron, Beit Kahil; 31.57° N, 35.07° E; ~ 800 m a.s.l.; 30 Apr. 1987; W. Heinz leg.; SMF • 6 ♂♂, 2 ♀♀, 3 juvs; Dead Sea, Ein Feshkha; 31.716° N, 35.452° E; 27 Feb. 1975; H. Levi, P. Amitai, and G. Levy leg.; MCZ 34061 • 1 ♂, 1 ♀; Southern Distr., Nahal Dawid (David), ’ En Gedi; 31.47° N, 35.39° E; -250 to - 300 m b.s.l.; 10 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22310 • 1 ♀ (in pure ethanol); same date as for preceding; ZFMK Isr59 • 1 ♂, 2 ♀♀; Southern Distr., Nahal Boqeq, W of ’En Boqeq; 31.1992° N, 35.3571° E; - 350 m b.s.l.; 8 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22311 • 1 ♂; Southern Distr., NW of ’En Tamar, Nahal Zin; 30.991° N, 35.347° E; - 340 m a.s.l.; 9 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; among rocks; ZFMK Ar 22312 • 1 ♂, 1 ♀; Negev, Mash’abei Sadeh, Golda Park; 31.0185° N, 34.7645° E; 340 m a.s.l.; 6 Sep. 2011; P. Jäger leg.; stony desert, under stones; SMF • 1 juv.; Negev, Mash’abei Sadeh, 1.5 air km S of Golda Park; 31.0026° N, 34.7576° E; 340 m a.s.l.; 7 Sep. 2011; P. Jäger leg.; sand dunes, stony desert, under stones; SMF • 2 ♂♂, 4 ♀♀, + juvs; Sede Boqer; 30.85° N, 34.78° E; Jan. 1987; V. and B. Roth; MCZ 34069 • 1 ♂, 3 ♀♀; Sede Boqer, 1.9 air km W of Midreshet Ben Gurion; 30.8554° N, 34.7642° E; 530 m a.s.l.; 8 Sep. 2011; P. Jäger leg.; stony desert, bushes, under stones; SMF • 1 ♀ (in pure ethanol); same collection data as for preceding; SMF • 2 ♀♀; same collection data as for preceding; 3 Sep. 2011; SMF • 1 ♂; Sede Boqer, 2.9 air km W of Midreshet Ben Gurion, Wadi Hawarim; 30.8476° N, 34.7550° E; 500 m a.s.l.; 4 Sep. 2011; P. Jäger leg.; under stones; SMF • 2 ♂♂, 1 ♀, 2 juvs; Sede Boqer, Ein Avedat, 3.3 air km W of Midreshet Ben Gurion; 30.8486° N, 34.7492° E; 400–480 m a.s.l.; 4 Sep. 2011; P. Jäger leg.; shrubs, under stones, in small caves; SMF • 6 juvs; same locality as for preceding; 6 Sep. 2011; P. Jäger leg.; stream with water, reed, shrubs, under stones; SMF • 1 ♂; Southern Distr., Har ’ Ayit; 30.0142° N, 35.0527° E; 470 m a.s.l.; 11 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; in deep crevice; ZFMK Ar 22313 • 1 ♂; Southern Distr., Samar; 29.8342° N, 35.0215° E; 100 m a.s.l.; 11 Sep. 2013; B.A. Huber, S. Aharon, and E. Gavish-Regev leg.; in building; ZFMK Ar 22314.
JORDAN – Balqa • 1 ♂, 1 juv.; Fuhais [Fuhays]; 32.010° N, 35.768° E; Apr. 1980; F. Krupp and W. Schneider leg.; SMF. – Madaba • 2 ♀♀, 2 juvs; Mt Nebo; 31.77° N, 35.73° E; Jul. 1989; E. Hyazin leg.; MRAC 169974 • 2 ♀♀, 1 juv.; Hammamat az-Zarqa’ Ma’in, hot springs, 150 m to Dead Sea; 31.608° N, 35.610° E; 14 Mar. 1977; R. Kinzelbach leg.; SMF • 1 ♂, 2 ♀♀; Wadi Mūjib; 31.465° N, 35.578° E; - 380 m b.s.l.; among rocks; 14 Sep. 2013; B.A. Huber leg.; ZFMK Ar 22315. – Kerak • 2 ♂♂, 2 ♀♀; Wadi Hasā; 31.004°–31.014° N, 35.494°–35.506° E; -330 to - 250 m b.s.l.; among rocks; 14 Sep. 2013; B.A. Huber leg.; ZFMK Ar 22316 • 2 ♂♂, 2 ♀♀ (in pure ethanol); same collection data as for preceding; ZFMK Isr53. – Ma’an • 2 ♂♂, 1 ♀; Petra; 30.324° N, 35.447° E; 900–950 m a.s.l.; 13 Sep. 2013; B.A. Huber leg.; among rocks; ZFMK Ar 22317 • 1 ♀, 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Isr37. – Aqaba • 1 ♂; Wadi Rām; 29.7405° N, 35.4574° E; 830 m a.s.l.; 12 Sep. 2013; B.A. Huber leg.; among rocks; ZFMK Ar 22318 • 1 ♂, 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Isr52 • 1 ♂; Wadi Rām; 29.683° N, 35.450° N; 9 Apr. 2004; J. Altmann leg.; SMF.
MOROCCO – Béni Mellal-Khénifra • 1 ♂, 1 ♀ (in pure ethanol); Imi n’Ifri; 31.724° N, 6.972° W; 1070 m a.s.l.; 26 Sep. 2018; B.A. Huber leg.; in building; ZFMK Mor105. – Drâa-Tafilalet • 2 ♂♂, 3 ♀♀, 1 juv.; Midelt; 32.68° N, 4.73° W; 19 Dec. 1986; V. and B. Roth leg.; CAS • 1 ♀; 22 km N of Errachida, between Sefrou and Tanout ou Filal; 32.04° N, 4.42° W; 24 Dec. 1986; V. and B. Roth leg.; CAS 9027138.
ALGERIA – Médéa • 2 ♂♂, 1 ♀; Médéa; 36.26° N, 2.75° E; date unknown; P.-H. Lucas leg.; MNHN. – Algiers • 1 ♀; Algiers; 36.75° N, 3.05° E; 3 Nov. 1948; B. Malkin leg.; AMNH • 4 ♂♂, 4 ♀♀, + juvs (possible syntypes of Pholcus barbarus Lucas); near Algiers; 36.75° N, 3.05° E; date unknown (between 1839 and 1842); P.-H. Lucas leg.; MNHN. – Tozeur • 2 ♀♀; Tozeur; 33.92° N, 8.12° E; Jul. 1972; E. and C. Supper leg.; MHNG.
TUNISIA – Jendouba • 2 ♂♂, + juvs; “ Tabarca (Khroumirie)”; ~ 36.9° N, 8.7° E; 1907; H. Gadeau de Kerville leg.; MNHN • 1 ♂, 3 ♀♀, 1 juv. (2 vials); Ayn Darahim (“ Ain-Draham ”); 36.78° N, 8.69° E; 1899; G. Seurat leg.; MNHN. – Tunis • 1 ♂, 2 ♀♀, 1 juv.; Carthago; 36.86° N, 10.33° E; 11–12 Oct. 1948; B. Malkin leg.; AMNH • 1 ♀; La Marsa; 36.89° N, 10.32° E; Jul. 1972; E. and C. Supper leg.; MHNG. – Gabès • 1 ♂; Gabès; 33.88° N, 10.10° E; 20 Mar. 1961; Walch leg.; SMF 24801. – Beja • 1 ♂, 1 juv.; Beja; 36.73° N, 9.19° E; 23 Mar. 2006; A. López and C. Ribera leg.; CRBA 1271. – Kairouan • 1 ♂, 2 ♀♀; Djebel-Trozza; 35.56° N, 9.59° E; 1932; F. Santschi leg.; MHNG. – Kef • 1 ♂, 1 ♀, 1 juv.; El Kef; 36.17° N, 8.70° E; Jul. 1972; E. and C. Supper leg.; MHNG.
LIBYA – Al Marquab • 1 ♀; Leptis Magna; 32.63° N, 14.29° E; 18–26 Aug. 1948; B. Malkin leg.; AMNH. – Al Jabal al Akh ḑar • 2 ♂♂, 2 ♀♀; Al Bayda (“El Baida”); 32.765° N, 21.743° E; 23 Jul. 1979; C. Goodnight and N. Barbash leg.; AMNH.
EGYPT – Dakahlia • 6 ♂♂, 6 ♀♀, 6 juvs; Mansoura (“ Mansurah ”); 31.04° N, 31.38° E; date unknown; I. Sörensen leg.; ZMUC. – Beheira • 3 ♂♂, 8 ♀♀, + juvs; Bir Hooker; 30.38° N, 30.35° E; 1901; J. Dewitz leg.; MNHN. – Cairo • 1 ♀; Maadi; 29.96° N, 31.26° E; further collection data on label unclear; MHNG • 1 ♀; Cairo [city]; 30.0° N, 31.2° E; date and collector unknown; SMF Roewer #4783. – Luxor • 2 ♂♂, + juvs; Luxor; 25.70° N, 32.65° E; 2 Nov. 1996; P. Jäger leg.; in house; SMF.
AUSTRALIA – Western Australia • 1 ♀, 1 juv.; Morawa Motel; 29.217° S, 116.017° E; 23 May 1996; M.S. Harvey leg.; in building; WAM • 1 ♂, 1 juv.; West Swan, behind Caversham Wildlife Park; 31.850° S, 115.983° E; 17 May 1992; J.M. Waldock leg.; on building; WAM 99/1790 • 1 ♂; Maylands; 31.93° S, 115.90° E; 7 Jan. 1992; J.M. Waldock leg.; WAM 99/1579 • 1 ♀; same collection data as for preceding; 27 Dec. 1991; WAM 99/1578 • 1 ♂; East Victoria Park; 31.983° S, 115.900° E; 15 Jan. 1997; J.M. Waldock leg.; WAM 99/2097 • 1 ♂; same collection data as for preceding; 13 Sep. 1992; in house; WAM 99/1776 • 1 ♂; same collection data as for preceding; 19 Nov. 1994; WAM 99/1752. – South Australia • 1 ♂, 1 ♀; Pondanna Outstation; 32.56° S, 135.55° E; 10 Dec. 1989; D. Hirst leg.; SAM 99/680 • 1 ♀; Flinders Range National Park, Oraparina; 31.37° S, 138.73° E; Dec. 1984; B. Guerin leg.; SAM 99/679 • 1 ♀, 1 juv.; Erudine Station via Yunta; 31.43° S, 139.43° E; 22 Apr. 1980; J. McEntee leg.; SAM 99/678 • 1 ♂; Middleback Station; 32.94° S, 137.40° E; Oct. 1983; B. Guerin leg.; SAM 99/682 • 1 ♀; Mt Lofty Ranges, Hawthorndene; 35.02° S, 138.63° E; 18 Jan. 1992; L.N. Nicolson leg.; in garden; SAM 99/683 • 2 ♀♀, 2 juvs; Stoneleigh Park near Meningie; 35.70° S, 139.35° E; Mar. 1969; A.W. Forbes leg.; SAM 99/684 • 1 ♂, 2 juvs; Murray Range near Kingston; 34.2° S, 140.3° E; 17 Dec. 1978; M.R. Gray leg.; AMS • 2 ♂♂, 6 ♀♀, 4 juvs; near Pryap; 34.45° S, 140.50° E; 2–9 Jun. 1990; L.N. Nicolson leg.; SAM 99/670 • 1 ♂, 6 ♀♀, 2 juvs; 7 km W of Loxton; 34.45° S, 140.50° E; 20 May 1990; L.N. Nicolson leg.; shed and house verandah; SAM 99/662 • 1 ♂; Loxton; 34.45° S, 140.57° E; 11 Jun. 1990; L.N. Nicolson leg.; in house; SAM 99/669. – Queensland • 1 ♀; south side of Lake Broadwater; 27.357° S, 151.098° E; 1–15 Nov. 1984; V. Wood leg.; buildings; QMB S49733 • 2 ♀♀; SW of Dalby, Lake Broadwater; 27.35° S, 151.10° E; 9 Dec. 1987; J. Gallon leg.; cottage; QMB S14509 • 1 ♂; Texas; 28.85° S, 151.17° E; date unknown; Harvey and Katzman leg.; in building; WAM. – New South Wales • 1 ♂, 1 ♀; Duntroon Station; 31.03° S, 143.04° E; Oct.–Nov. 1980; D. Hirst leg.; SAM 99/655 • 1 ♀, 2 juvs; Weinteriga; 31.60° S, 142.95° E; 15 Apr. 1981; D. Hirst leg.; SAM 99/659 • 2 ♂♂; Menindee Lakes Caravan Park; 32.354° S, 142.404° E; 12 Apr. 1981; D. Hirst leg.; SAM 99/657, 99/658 • 3 ♂♂, 1 juv.; Sulcor; 30.85° S, 150.81° E; 27 May 1995; S. Eberhard leg.; cave, twilight zone; AMS KS 49263. – Victoria • 1 ♂; 30 km N of Kaniva, near Broughton; 36.17° S, 141.35° E; 25 May 1988; D. Hirst leg.; SAM 99/661.
Redescription
Male (Israel, ZFMK Ar 22306)
MEASUREMENTS. Total length 6.0, carapace width 2.3. Distance PME–PME 220 µm; diameter PME 140 × 150 µm; distance PME–ALE 70 µm; diameter AME 125 µm; distance AME–AME 25 µm. Leg 1: 45.4 (12.9 + 1.1 + 12.7 + 16.4 + 2.3), tibia 2: 8.1, tibia 3: 6.0, tibia 4: 7.2; tibia 1 L/d: 42; femora 1–4 diameters: 0.46, 0.39, 0.37, 0.38.
COLOR (in ethanol). Carapace ochre-yellow, posterior part of ocular area brown; clypeus not darkened; sternum dark brown to black; legs ochre-yellow, with darker rings on femora (subdistally) and tibiae (proximally and subdistally), with oval to elongate black marks on femora and tibiae; abdomen ochregray, with dark and whitish marks dorsally and laterally; ventrally with distinct black median band, partly disrupted, with three parallel longitudinal marks behind gonopore.
BODY. Habitus as in Fig. 5. Ocular area slightly raised; each secondary eye with small accompanying elevation (Fig. 43). Deep thoracic pit and pair of shallow furrows diverging from pit toward posterior margin. Clypeus unmodified (contra Calbacho-Rosa et al. 2019b; see Discussion), only rim more sclerotized than in female. Sternum wider than long (1.5/1.1), unmodified (i.e., without indentations as in H. caudatus and H. reini). Abdomen cylindrical, dorso-posteriorly weakly angular. Gonopore with four epiandrous spigots (Fig. 55). ALS with one widened spigot and one pointed spigot; PMS with two pointed spigots (Fig. 56).
CHELICERAE. As in Fig. 16; see also Huber (1995: figs 1a, 4a; 2000: fig. 14); with pair of frontal lateral apophyses, each with one large modified cone-shaped hair (Fig. 54); distance between tips of modified hairs: 390 µm; without proximal frontal protrusion; lateral stridulatory ridges distinct (Fig. 47), distances between ridges ~12 µm.
PALPS. As is Figs 13–15; coxa with rounded retrolateral hump (not a distinct apophysis); trochanter barely modified (slightly protruding ventrally); femur curved towards dorsal, distally widened but without ventral protrusion, proximally with prolateral stridulatory pick, without retrolateral transversal line, with prominent retrolateral proximal process; femur-patella joints only slightly shifted toward prolateral side; tibia large compared to femur, tibia-tarsus joints shifted toward retrolateral side; tarsal organ capsulate (Fig. 51); tarsus without macrotrichia; procursus (Figs 20–22) straight, dorsal hairs not or only slightly curved upwards; proximally on prolateral-ventral side with prominent process free of hairs (arrow in Fig. 20), procursus tip with strong but short ventral sclerite, membranous elements on dorsal and prolateral side, and 4–5 hair-like transparent processes on retrolateral side (Fig. 49); genital bulb (Figs 23–26) with simple basal sclerite connected to distal (main) sclerite; distal sclerite with two distinctive processes; sperm duct opening on membranous area on prolateral side at basis of distal bulbal sclerite (arrow in Fig. 50).
LEGS. With single ventral rows of spines on femur 1 (~31), tibia 1 (~38) and femur 2 (~14) (Figs 17– 19); without curved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 4%; prolateral trichobothrium present on all tibiae; tarsal pseudosegments indistinct and irregular (cf. Fig. 35) except 2–3 distally; all tarsal organs capsulate with simple round rim (Fig. 53).
Male (variation)
Tibia 1 in 95 males: 7.9–15.5 (mean 11.6) (mean tibia 1 length in 204 males in Jakob & Dingle 1990: 10.67); body length ~3.5–7.5. Most specimens with more or less distinct dark median band on carapace; sternum sometimes with distinct black radial marks; abdomen sometimes without dark marks, but always with white marks in distinctive pattern. Smaller males with fewer spines, often without spines on tibia 1 and femur 2. Distance between tips of modified hairs on cheliceral apophyses: ~280–390 µm. Gonopore with 4–5 epiandrous spigots.
Female
In general similar to male (Fig. 6) but palp with strongly widened distal segments (tibia and tarsus; Figs 29–30), without joints between tibia and tarsus; with strong median process posteriorly on sternum (Figs 31, 58) possibly acting against ventral process anteriorly on abdomen (Figs 38, 57; but see Discussion); with fewer but stronger stridulatory ridges (Fig. 48; see also Huber 1995; Huber 2021a: fig. 12), distances between stridulatory ridges ~13.5 µm; without spines on legs; dark marks on femora and tibiae (Fig. 34) often more distinct than in males; with strong anterior sclerite dorsally on pedicel possibly acting against pair of sclerotized areas anteriorly on abdomen (Fig. 32). Without stridulatory apparatus between carapace and abdomen. Tibia 1 in 140 females: 7.2–14.1 (mean 10.5) (mean tibia 1 length in 172 females in Jakob & Dingle 1990: 10.05); body length: ~4.0–8.0 mm. Epigynum as in Figs 36–39 and 57; main epigynal plate wide, weakly protruding, posteriorly whitish, anteriorly with pair of depressions ~280–300 µm apart, with sculptured cuticle (cf. Huber 1995: fig. 6c), median internal round element and pair of lateral internal triangular elements usually visible in uncleared specimens; posterior epigynal plate large but simple; large anterior plate in front of epigynum with pair of low humps and distinct anterior elevated rim (arrow in Fig. 38). Internal genitalia (Figs 27–28, 40–42) with elongate pore plates in transversal position narrowing medially, dorsal arc relatively simple, ventral arc with distinctive pair of lateral sclerotized pockets, medially widened, with ventral median process (pocket?). Spigots as in male.
Natural history
Together with Pholcus phalangioides (Fuesslin, 1775), H. pluchei is probably the best studied pholcid spider with respect to its biology. Much of what we know about H. pluchei has been studied in the USA and Argentina, i.e., using non-native populations. Whether they differ in any significant way from Mediterranean populations has never been studied. Data on egg numbers (see below) suggest that such differences may exist.
Around the Mediterranean, H. pluchei is ubiquitous in almost any habitat suitable for its web, including places fully exposed to the sun. Occasionally, H. pluchei lives in peripheral parts of webs of the araneid Cyrtophora citricola (Forsskål, 1775), together with the cobweb spider Argyrodes argyrodes (Walckenaer, 1841) (Blanke 1972; Hajer 1995; Leborgne et al. 1998; Hajer & Řeháková 2003). In Central Europe, where H. pluchei has been massively spreading over the last decades, it may be displacing P. phalangioides, at least in warm/dry/light parts of buildings, close to windows (Jäger 2000; Van Keer 2007). In the southwestern USA, H. pluchei is often the dominant spider on outside surfaces in urban areas around homes and other structures (Vetter et al. 2011). It sometimes occurs is high densities: up to 600 spiders were estimated to inhabit a 3 × 15 m juniper bush in California (Blanchong et al. 1995).
The usual web of H. pluchei is dome-shaped, as in most space-dwelling pholcids. A skeleton made of 1.2 µm fibrils is filled with 0.8–0.9 µm fibrils (Hajer & Řeháková 2003). Occasionally, webs are densely covered with silk puffs, a common behavior in Smeringopinae that interferes with a clear view of the spider. The fact that silk puffs are produced 3–4 days before molting and before egg-laying supports the view that puffs may protect the spider from visual predators when it is most vulnerable (Hajer & Řeháková 2003). Unusual spherical webs are built by females before egg-laying. Females remain in
these small spheres (diameter ~ 5 cm) until the spiderlings hatch, and females do not prey during this time (Sedey & Jakob 1998; Jäger 2000; Hajer & Řeháková 2003).
Holocnemus pluchei is facultatively group-living, i.e., several conspecifics of all sizes may share a web (Jakob 1991). Groups are constantly forming and disintegrating as the spiders move frequently among webs, depending on size, feeding status, and presence or absence of conspecifics (Jakob 2004). Surprisingly, small spiders in a group seem to pay a high prize for group living: they feed less than solitary siblings and larger companions, they tend to lose contests, and they are sometimes cannibalized (Gerhardt 1927; Jakob 1991; Jakob et al. 2000). The reason they join conspecifics may be the cost of building an own web: it takes a spiderling at least 5 days to feed enough to build a web (Jakob 1991), and webs are not recycled (Jakob et al. 2000).
Sperm uptake was described by Gerhardt (1927). First, a line of silk between the tips of legs 3 is moved against the gonopore until the sperm droplet emerges. Then the line with the drop is brought to the chelicerae from where it is taken up by the genital bulbs. In alternating movements, each bulb contacts the drop three times for approximately 30 s each. Within the bulb, sperm remains viable at least for several weeks (99.6% viable after two weeks; Cargnelutti et al. 2020).
Basic aspects of courtship and copulation have been described by Gerhardt (1927), Huber (1995), and Dutto et al. (2011). Both males and females (and juveniles) use their chelicerae to stridulate, but in different contexts. While male stridulation seems to have a luring function, female stridulation conveys a negative message, mainly to males (signaling non-receptivity), but also to other females and maybe even to other species (e.g., the web-invading Pholcus phalangioides) (Huber 1995; Dutto et al. 2011). The stridulation of juveniles has never been studied. Female stridulatory behavior has been found to be consistent, i.e., individual females differ consistently in the frequency of stridulating during consecutive inter-sexual encounters (Calbacho-Rosa et al. 2019a).
In copulations with virgin females, sperm is transferred during a first phase that is characterized by rhythmic simultaneous movements of the male palps (Cargnelutti et al. 2018). This is followed by a second phase where the palps remain inserted but seemingly immobile. The significance of this second phase is only partly understood. It might be a form of mate-guarding, and the duration of the immobile phase positively affected sperm viability in females (which decreased more rapidly in females than in males) (Cargnelutti et al. 2020).
Copulations with non-virgin females are common; they take longer (~40 vs 30 min; Kaster & Jakob 1997), and they follow a slightly different pattern. They start with non-rhythmic alternating movements of only partially inserted palps (only procursus inserted) during which sperm of previous males is partially removed (Calbacho-Rosa et al. 2013). Second males fertilize an average of ~65–83% of the eggs, but the actual values range from 0 to 100% (Kaster & Jakob 1997). Second-male sperm precedence depends on timing: it is strongest in the few hours after copulation (Calbacho-Rosa et al. 2010) (explaining the pattern of mate guarding; see below).
The projection on the female sternum has been interpreted as a structure that controls the intensity and range of male palpal movements during copulation (Calbacho-Rosa et al. 2019b). These authors support their conclusion mainly by the fact that the projection contacts the male clypeus at the moment of maximum palpal contraction. They exclude a stidulatory function (with the “pre-epigynum”) because the structures do not contact each other during intersexual interactions (but see Discussion).
The biological significance of the dorsal female modification (pedicel, abdomen) has never been studied (it has apparently not even been described before). It may function during abdominal twitching, a component of low-level aggressive interactions (Jakob 1991, 1994). Abdomen twitching was the only antagonistic behavioral component that differed between sexes, but it was males (that lack pedicelabdomen modifications) that performed more twitches (Blanchong et al. 1995). Escalating fights involve the legs, and Johnson & Jakob (1999) found that 7–8% of spiders in natural populations were missing at least one leg, usually (in 86% of the cases) one of the anterior two pairs. However, leg loss does not significantly affect the male’s ability to compete over webs and prey (Johnson & Jakob 1999).
Reports on mate guarding are somewhat contradictory. For Californian populations, Kaster & Jakob (1997) report the absence of guarding; Sedey & Jakob (1998) found that most females with eggs were initially accompanied by at least one male. In Argentinean populations, Calbacho-Rosa et al. (2010) observed post-copulatory mate guarding: males stayed close to females for approximately 12–24 hrs after copulation, and actively defended females (or rather their own investment) from intruding males. This does not seem to imply chivalrous behavior: males do not cede prey to females, and they win interactions as often as females do (Blanchong et al. 1995).
Females produce several egg-sacs per season, and Hajer & Řeháková (2003) reported a maximum of eight clutches in the lab. As usual in pholcids (Huber & Eberle 2021), larger females lay more eggs (and thus produce heavier clutches), but egg weight is not affected by female body size (Skow & Jakob 2003). Kaster & Jakob (1997), working on Californian populations, reported a mean clutch size of 33 (9–70), while Ahmed (2021), working on Egyptian spiders, reported a mean of 89. While Ahmed’s (2021) publication is flawed in many respects, this number is not unfeasible. A single egg-sac of a female from Croatia reported in Huber & Eberle (2021) was estimated at having 77 eggs.
While caring for an egg-sac, females are reluctant to remate, but occasionally they do, especially at a late stage of embryo development (Calbacho-Rosa et al. 2017). In order to mate, they have to temporarily suspend the egg-sac in the web, which supposedly carries some risk for the eggs, especially from (conspecific) predators (Calbacho-Rosa et al. 2017). Egg-sac carrying has also been shown to protect the eggs from fungi (Calbacho-Rosa et al. 2017), but this is not likely to have a significant effect during the relatively short time of mating.
Developmental time depends on food level, and well-fed spiders often reach maturity after five molts, while poorly-fed spiders usually need six molts (Jakob & Dingle 1990). However, spiders maturing after six molts are on average larger, irrespective of food level (Jakob & Dingle 1990). When legs were removed at the third instar, they were not regenerated (Johnson & Jakob 1999).
When disturbed, H. pluchei starts to move vigorously like many long-legged pholcids. However, the movement has been described as ‘bouncing’, different from the ‘whirling’ of P. phalangioides (Jackson et al. 1993). When bouncing, H. pluchei lifts and lowers the body by flexing and extending the femurpatella and tibia-metatarsus joints at an amplitude of 2–20 cm and at a rate of 5–10/s (Jackson et al. 1993). This is thought to protect the spider from visual predators, and it may also interfere with a predator’s movement in the web (Jackson 1992; Jackson et al. 1993). An alternative strategy is to leap out of the web and ‘play dead’ (Jackson et al. 1993).
Distribution
The original distribution of H. pluchei is the Mediterranean or part of it. Compared to some other synanthropic and anthropophilic pholcids, it has established permanent colonies in other regions relatively recently (Fig. 2). Surprisingly, this seems to have happened within a relatively short period of time around the world. For the USA, Vetter et al. (2011) did a survey among regional arachnologists and concluded that H. pluchei had been introduced in the San Francisco Bay area in the 1950s. The oldest South American records date back to the early 1960s (Argentina; see Material examined). By that time, the species was present in at least two provinces, suggesting that it may well have been introduced in the 1950s as well. The species is now very common in central Argentina as well as in Uruguay (Laborda & Simó 2008).
The oldest confirmed Central European and Australian records are also from the 1960s (Germany: 1962; South Australia: 1969; see Material examined section). Older records from the Netherlands (19 th century) are dubious (van Helsdingen 2010). In Central Europe the species has been spreading massively since the 1990s (Van Keer & Van Keer 2001; Reiser & Neumann 2014) and has relatively recently reached countries like Denmark (oldest record 2006; https://www.danmarks-edderkopper.dk/), Great Britain (oldest record 2004; https://www.britishspiders.org.uk/), and the Caucasus (Ponomarev et al. 2019). The oldest known record for Japan is from 2008 (Kumada 2021), suggesting that the introduction to Japan happened relatively recently.
L. Koch’s (1875) record for Massaua (Eritrea) is based on misidentified juvenile specimens of Crossopriza, presumably C. pristina (photos kindly provided by M. Tavano, 30 Jan. 2014). Leardi in Airaghi’s (1902) record for Mahé (India) could not be checked but it here also considered to be based on misidentified specimens.
description: Published as part of Huber, Bernhard A., 2022, Revisions of Holocnemus and Crossopriza: the spotted-leg clade of Smeringopinae (Araneae, Pholcidae), pp. 1-241 in European Journal of Taxonomy 795 (1) on pages 9-29, DOI: 10.5852/ejt.2022.795.1663, http://zenodo.org/record/6299150