Holocnemus pluchei ( Scopoli, 1763 )

Huber, Bernhard A., 2022, Revisions of Holocnemus and Crossopriza: the spotted-leg clade of Smeringopinae (Araneae, Pholcidae), European Journal of Taxonomy 795 (1), pp. 1-241 : 9-29

publication ID

https://doi.org/ 10.5852/ejt.2022.795.1663

publication LSID

lsid:zoobank.org:pub:7394D45E-46E1-453C-BF7E-1FE1B2CEBB0A

persistent identifier

https://treatment.plazi.org/id/671CB865-FFA7-FFE7-FDA9-F93DFD878E6A

treatment provided by

Felipe

scientific name

Holocnemus pluchei ( Scopoli, 1763 )
status

 

Holocnemus pluchei ( Scopoli, 1763) View in CoL

Figs 2 View Fig , 5–6 View Figs 5–12 , 13–58 View Figs 13–19 View Figs 20–28 View Figs 29–35 View Figs 36–42 View Figs 43–50 View Figs 51–58

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 View in CoL – 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 View Cited Treatment . — 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 View Figs 29–35 ), by median process on female sternum ( Figs 31 View Figs 29–35 , 58 View Figs 51–58 ), and by pair of distinctive processes on male distal bulbal sclerite ( Figs 24, 26 View Figs 20–28 ); also by shape of procursus ( Figs 20–22 View Figs 20–28 ; distinct prolateral proximal process, short distal ventral sclerite), by female external and internal genitalia ( Figs 27–28 View Figs 20–28 , 36–42 View Figs 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 View Figs 29–35 ).

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 GoogleMaps .

Syntypes of Aranea rivulata EGYPT • Unknown number; Cairo; ~ 30.0° N, 31.2° E; probably 1762; P. Forsskål leg.; probably lost GoogleMaps .

Syntypes of Pholcus impressus GREECE • 4 ♂♂; Peloponnese, Nafplion (“Nauplion”); 37.57° N, 22.81° E; date unknown; [F.J.?] Schuch leg.; probably lost GoogleMaps .

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 GoogleMaps .

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 About CAS part GoogleMaps 1 ♀; Napa County, Lake Hennessey, below Lonn Dam; 38.48° N, 122.38° W; 11 May 1985; L. Vincent leg.; CAS 9027422 About CAS GoogleMaps 1 ♀; Los Angeles County, Diamond Bar ; 34.03° N, 117.80° W; 23 Sep. 1999; C. Yang leg.; CAS 9027418 About CAS GoogleMaps 1 ♀; Orange County, Yorba Linda ; 33.889° N, 117.812° W; 2 May 2000; S. Clayton leg.; CAS 9027393 About CAS GoogleMaps 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 GoogleMaps .

ARGENTINA – Mendoza • 1 ♀; Mendoza ; 32.89° S, 68.84° W; 900 m a.s.l.; 30–31 Mar. 1965; H. Levi leg.; MCZ GoogleMaps . – San Luis • 1 ♂; Naschel ; 32.915° S, 65.375° W; Feb.–Mar. 1962; A. Luchini leg.; MACN 19955 View Materials GoogleMaps . – La Pampa • 1 ♀; Rancul ; 35.07° S, 64.68° W; Jan. 1975; Maury leg.; MACN 20101 View Materials GoogleMaps 1 ♀; Santa Rosa [de Toay]; 36.62° S, 64.29° W; date and collector unknown; MACN 20039 View Materials GoogleMaps . – Buenos Aires • 1 ♂; Carlos Casares ; 35.62° S, 61.36° W; 27 Feb.–2 Mar. 1980; collector unknown; MACN 20087 View Materials GoogleMaps 1 ♂, 2 ♀♀, 2 juvs; Buenos Aires, Cerrito ; 34.60° S, 58.38° W; 19 Sep. 1972; C. Cesari leg.; in basement; MACN 20080 View Materials GoogleMaps .

PORTUGAL – Faro • 2 ♂♂, 1 ♀; Silves [Munic.], Norinha ; 37.198° N, 8.400° W; 14 Sep. 1969; A. Senglet leg.; MHNG GoogleMaps .

SPAIN – Extremadura • 1 ♂; Cáceres, Jarandilla [de la Vera]; 40.126° N, 5.658° W; 22–23 Aug. 1969; A. Senglet leg.; MHNG GoogleMaps . – Castilla-La Mancha • 2 ♂♂, 4 ♀♀, 9 juvs; Toledo, Escalona del Alberche ; 40.167° N, 4.400° W; 14 Jun. 1969; A. Senglet leg.; MHNG GoogleMaps . – Andalusia • 6 ♂♂, 7 ♀♀, 2 juvs; Córdoba, Palma del Río ; 37.70° N, 5.28° W; 26–27 Jun. 1969; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂, 2 ♀♀; Córdoba, Peñarroya ; 38.30° N, 5.27° W; 30 Jun. 1969; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀; Cádiz, Algodonales ; 36.88° N, 5.40° W; 19 Jul. 1969; A. Senglet leg.; MHNG GoogleMaps 3 ♂♂, 2 ♀♀, 5 juvs; Málaga, Estepona ; 36.43° N, 5.15° W; 24–26 Jul. 1969; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀; Málaga, Valle de Abdalajís ; 36.93° N, 4.68° W; 28 Jul. 1969; A. Senglet leg.; MHNG GoogleMaps 4 ♀♀; Málaga, Antequera ; 37.02° N, 4.56° W; 1–2 Aug. 1969; A. Senglet leg.; MHNG GoogleMaps 3 ♂♂, 5 ♀♀; Málaga, Torre del Mar ; 36.75° N, 4.10° W; 27 Jul. 1969; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀; Granada, Ugíjar ; 36.96° N, 3.05° W; 15 Jul. 1971; A. Senglet leg.; MHNG GoogleMaps 3 ♀♀; Granada, Huéneja ; 37.176° N, 2.948° W; 16 Jul. 1971; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps 1 ♀; San Pedro del Pinatar ; 37.83° N, 0.78° W; Sep. 1984; G. Hormiga leg.; USNM GoogleMaps 1 ♂; Fortuna, Cueva Las Magras ; 38.290° N, 1.143° W; 30 Jan. 2006; “ACP” leg.; CRBA 1146 GoogleMaps . – Valencian Community • 4 ♂♂, 3 ♀♀; Valencia, La Albufera ; 39.30° N, 0.33° W; 16 Jun. 1971; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps . – Catalonia • 1 ♂; Barcelona, Gelida ; 41.44° N, 1.86° E; 14 Jun. 1971; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂, 1 ♀, 3 juvs; near Barcelona, Cerdanyola [del Vallès]; 41.49° N, 2.14° E; 7 Dec. 2004; D. Dimitrov leg.; SMF 56746 GoogleMaps .

FRANCE – Pyrénées-Orientales • 1 ♂, 2 ♀♀; Banyuls ; 42.48° N, 3.13° E; date and collector unknown; MNHN Ar 13337 GoogleMaps 2 ♂♂, 3 ♀♀ (3 vials); same locality as for preceding; 9–20 May 1952; A. Comellini leg.; MHNG GoogleMaps 1 ♀; same locality as for preceding; no further data; MHNG GoogleMaps . • 1 ♀; Cerbère ; 42.44° N, 3.165° E; 11 May 1952; A. Comellini leg.; MHNG GoogleMaps . – Var • 1 ♂, 1 ♀; Bormes-les-Mimosas (“ Bornes ”); 43.15° N, 6.34° E; Jul. 1960; O. Kraus leg.; SMF 24800 GoogleMaps . – Gard • 3 ♀♀; Aigues-Mortes ; 43.57° N, 4.19° E; 8 May 1952; A Comellini leg.; MHNG. – Corsica GoogleMaps 1 ♂, 1 ♀; Propriano ; 41.67° N, 8.91° E; 29 May 1971; A. Senglet leg.; MHNG GoogleMaps 1 ♂, 7 ♀♀, 1 juv.; Ponte Leccia ; 42.463° N, 9.207° E; 19 Sep. 1952; Kahmann leg.; SMF 8799 About SMF , 8800 About SMF GoogleMaps 1 ♀; Biguglia ; 42.62° N, 9.44° E; 30 m a.s.l.; 2 May 1952; Kahmann leg.; SMF 8801 About SMF GoogleMaps .

BELGIUM – East Flanders • 1 ♀; near Gent; 51.10° N, 3.70° E; Jul. 2004; B.A. Huber leg.; ZFMK Ar 12608 GoogleMaps .

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 GoogleMaps . – 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 GoogleMaps 1 ♀; Mainz, Draisberghof ; 48.987° N, 8.203° E; 1 Feb. 1998; P. Jäger leg.; SMF 40411 GoogleMaps . – 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 GoogleMaps .

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 About ZFMK to G66 About ZFMK GoogleMaps .

ITALY – Sardinia • 1 ♂; S. Pietro ; 40.717° N, 8.550° E; 15 Aug. 1960; E. Kullmann leg.; ZFMK Ar 10705 GoogleMaps 1 ♀, 1 juv.; Castelsardo ; 40.91° N, 8.71° E; 11 Jun. 1958; E. Kullmann leg.; ZFMK Ar 10746 GoogleMaps 1 ♀; Lago Bunnari ; 40.716° N, 8.634° E; 11 Apr. 1952; collector unknown; SMF 9273 About SMF GoogleMaps 1 ♂, 2 juvs; Ozieri ; 40.58° N, 9.00° E; 23 Sep. 1968; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀, 1 juv.; Terralba ; 39.72° N, 8.63° E; 4 Aug. 1960; E. Kullmann leg.; ZFMK Ar 10696 GoogleMaps 1 ♂, 1 ♀, 1 juv.; Muravera, Nuraghe ; 39.318° N, 9.531° E; 19 Aug. 1960; E. Kullmann leg.; ZFMK Ar 8214 GoogleMaps 4 ♂♂, 5 ♀♀, 2 juvs; Sardinia , no further data; SMF RII 6384 About SMF . – 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 GoogleMaps . – Sicily • 1 ♂; Syracuse, Viale Ermocrate ; 37.07° N, 15.27° E; 18 Aug. 1993; B.A. Huber leg.; ZFMK Ar 5217 GoogleMaps 1 ♂, 1 ♀; Syracuse, Theatre ; 37.074° N, 15.279° E; 18 Aug. 1993; B.A. Huber leg.; ZFMK Ar 5218 GoogleMaps .

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 GoogleMaps 6 ♂♂, 1 ♀; Ploče ; 43.05° N, 17.43° E; 21 May 1972; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps .

BOSNIA AND HERZEGOVINA • 1 ♂, 2 ♀♀; Popovo-Polje , Zavala; 42.848° N, 17.979° E; 19 Sep. 1970; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂, 2 ♀♀; Popovo-Polje , Sedlari; 42.826° N, 18.056° E; 18 Sep. 1970; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂ (in pure ethanol); near Trebinje, near Pavlova pećina; 42.666° N, GoogleMaps

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 GoogleMaps 2 ♀♀; Podgorica , Ljuboviči; 42.433° N, 19.255° E; 21 Apr. 2011; M. Komnenov leg.; artificial tunnel; CMK GoogleMaps .

ALBANIA • 1 ♀; Kulla ë Lumës (“Kula Lums”); 42.098° N, 20.418° E; 1918; collector unknown; SMF RII 3726 About SMF GoogleMaps 1 ♀, 1 juv.; Durrës; 41.33° N, 19.45° E; 26 May 1993; P. Stoev and C. Deltshev leg.; NMNHS GoogleMaps 1 ♀; Përmet, near Petrani; 40.208° N, 20.419° E; 12 May 1995; B. Petrov and P. Stoev leg.; artificial gallery; NMNHS GoogleMaps .

NORTH MACEDONIA • 1 ♀; Gevgelija; 41.14° N, 22.50° E; 10 Jul. 1956; Lamel leg.; HECO GoogleMaps .

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 GoogleMaps 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 GoogleMaps 1 ♂, 2 ♀♀; Sandanski ; 41.566° N, 23.280° E; 28 Jul. 1972; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps . – Pazardzhik • 2 ♂♂, 1 ♀; Panagyuriste ; 42.505° N, 24.186° E; 720 m a.s.l.; 1 Jul. 1997; S. Lazarov leg.; NMNHS. – GoogleMaps Ruse • 1 ♂, 2 ♀♀; Byala ; 43.46° N, 25.74° E; 2013; collector unknown; MGAB GoogleMaps .

GREECE – Ionian Islands • 1 ♂; Corfu, Agios Stephanos ; 39.79° N, 19.81° E; 26 May 1999; A. Russell-Smith leg.; ZFMK Ar 22293 GoogleMaps 1 ♂, 1 ♀; Corfu, Loutses ; 39.786° N, 19.880° E; 20 Sep. 1972; A. Senglet leg.; MHNG GoogleMaps 3 ♂♂, 1 ♀; Corfu, Agios Matheos ; 39.495° N, 19.872° E; 18 Sep. 1972; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps 1 ♀; Pantocrator, Spilaio Katsouri ; 39.743° N, 19.925° E; 130 m a.s.l.; 29 Apr. 1975; D. Tzanoudakis leg.; MHNG GoogleMaps 1 ♂, 1 ♀; Lefkada , W of Mikros Gialos; 38.64° N, 20.69° E; 25 May 1995; A. Russell-Smith leg.; ZFMK Ar 22294 GoogleMaps 8 ♂♂, 3 ♀♀; Cephalonia, Sisia ; 38.10° N, 20.66° E; 29 May 1987; J.A. Murphy leg.; MMUE Murphy #A231 GoogleMaps 1 ♂, 2 ♀♀; same collection data as for preceding; reared from eggs; MMUE Murphy #15938 GoogleMaps 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 GoogleMaps . – Central Macedonia • 1 ♂; Yerakini (“ Gerakina ”); 40.27° N, 23.45° E; 14 Apr. 1978; J.A. Murphy leg.; stony area; MMUE Murphy #6830 GoogleMaps 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 GoogleMaps . – Thessaly • 2 ♂♂, 2 ♀♀; Larissa , “Omôlion (Tembe)”; 39.88° N, 22.58° E (coordinates between Omolio and Tempi); 17 Jun. 1970; A. Senglet leg.; MHNG GoogleMaps . – Central Greece • 1 ♂; Euboea, Lichada / Istiaia ; 38.9° N, 23.0° E (coordinates between Lichada and Istiaia); 30 Aug. 1972; A. Senglet leg.; MHNG GoogleMaps 3 ♂♂, 9 ♀♀, 2 juvs; Phthiotis, Tragána ; 38.62° N, 23.12° E; 21 May 1968; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps 1 ♂, 1 ♀; Phthiotis, near Malesina; 38.62° N, 23.22° E; 21 May 1968; A. Senglet leg.; MHNG GoogleMaps 1 ♂, 2 ♀♀; Boeotia , “Topôlia/Thiva”; 38.32° N, 23.32° E (coordinates of Thiva); 25 Aug. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♂; Euboea, Kato Steni ; 38.571° N, 23.825° E; 2 Sep. 1972; A. Senglet leg.; MHNG GoogleMaps 1 ♂, 1 juv.; Northern Sporades, Skyros Island ; 38.90° N, 24.56° E; 22 Mar. 1958; Schelkopf leg.; SMF GoogleMaps . – Attica • 5 ♂♂, 4 ♀♀ (partly used for SEM); Daphni / Athens ; 38.013° N, 23.636° E; 23 Jun. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♂, 1 ♀; same locality as for preceding; 19 Jun. 1968; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀, 1 juv.; Athens, Akropolis ; 37.971° N, 23.727° E; date and collector unknown; SMF RII 3724 About SMF GoogleMaps 4 juvs; same locality as for preceding; May 1926; F. Roewer leg.; SMF GoogleMaps 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 GoogleMaps 1 ♀; Vari, Spilaio Nympholyptou ; 37.858° N, 23.802° E; 25 Feb. 2018; N. Fytrou and E. Zenzefyli leg.; HISR 3042 GoogleMaps 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 GoogleMaps 1 ♂, 5 ♀♀; Gulf of Argolis, Kantia (“ Argolide , Kandia ”); 37.52° N, 22.96° E; 12 Aug. 1970; A. Senglet leg.; MHNG GoogleMaps 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 GoogleMaps . – 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 GoogleMaps . – South Aegean • 4 ♀♀; Paros, Naousa ; 37.12° N, 25.24° E; 21 Jun. 1968; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂; Paros , Parikia E , “Moni Aghii Anarghii”; 37.08° N, 25.16° E; 10 Aug. 2006; R. Bosmans leg.; on walls; CRB GoogleMaps 2 ♂♂, 2 ♀♀; Naxos, Chalkio (“ Chalkis ”); 37.063° N, 25.482° E; 27 Jun. 1968; A. Senglet leg.; MHNG GoogleMaps 2 ♀♀; Karpathos, near Pigadia; 35.51° N, 27.20° E; Apr. 1963; Kinzelbach leg.; SMF 15489 GoogleMaps 2 ♂♂, 1 ♀; Rhodes , “ rocca di Lindos ”; 36.091° N, 28.088° E; 2 Jul. 1996; F. Gasparo leg.; ZFMK Ar 22296 GoogleMaps . – Crete • 1 ♂, 2 ♀♀; Chania, Azogires Palaeochoras ; 35.272° N, 23.719° E; 4–6 Aug. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♂; Chania, Sougla ; 35.25° N, 23.81° E; 9 Oct. 1999; A. Senglet leg.; MHNG GoogleMaps 3 ♀♀; Chania, Halepa (“ Chalepa ”); 35.52° N, 24.05° E; date and collector unknown; SMF RII 3725 About SMF GoogleMaps 1 ♂, 1 ♀; Chania, Akrotiri, Kumaro Caves ; 35.579° N, 24.143° E; May 1926; F. Roewer leg.; SMF GoogleMaps 1 ♂; Chania, Georgioupoli ; 35.36° N, 24.26° E; 9 Jun. 2005; W. Schedl leg.; ZFMK Ar 22297 GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 1 ♂, 1 ♀; Rethymno, Choumeri / Perama ; 35.35° N, 24.72° E; 1 Jul. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♀; Rethymno, Anogia ; 35.24° N, 24.87° E; 11 May 2006; H. Eikamp and U. Kluge leg.; SMF 56588 GoogleMaps 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 GoogleMaps 1 ♀; same locality as for preceding; 15 May 2010; K. Eckl leg.; SMF GoogleMaps 1 ♂, 2 ♀♀; Heraklion, Rodia (“ Rogdia ”); 35.365° N, 25.020° E; 27 Jun. 1970; A. Senglet leg.; MHNG GoogleMaps 2 ♂♂, 2 ♀♀; Heraklion, Knossos ; 35.30° N, 25.16° E; 25 Jun. 1970; A. Senglet leg.; MHNG GoogleMaps 3 ♂♂, 5 ♀♀; Heraklion, Malia ; 35.29° N, 25.46° E; 8 Jul. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♂; same locality as for preceding; 11 May 2008; H. Eikamp and U. Kluge leg.; SMF GoogleMaps 2 ♂♂, 1 ♀; Heraklion, Charakas ; 35.01° N, 25.12° E; 24 Jul. 1970; A. Senglet leg.; MHNG GoogleMaps 1 ♂, 1 ♀; Heraklion, Marathos, Arkalospilios ; 35.245° N, 24.968° E; 24 Dec. 2000; K. Paragamian leg.; HISR 1971 part GoogleMaps 1 ♀, 6 juvs; Heraklion, Kastri ; 35.00° N, 25.37° E; 7 May 2008; H. Eikamp and U. Kluge leg.; SMF GoogleMaps 1 ♀; Heraklion, Kastri, Keratokambos ; 35.000° N, 25.375° E; 14 May 2006; H. Eikamp and U. Kluge leg.; SMF 56589 GoogleMaps 1 ♂; Lasithi, Plati ; 35.17° N, 25.44° E; 17 Dec. 2006; H. Eikamp and U. Kluge leg.; SMF GoogleMaps 1 ♂; Lasithi, Atsiganospilios ; 35.205° N, 25.618° E; 16 Apr. 2003; F. Gasparo leg.; ZFMK Ar 22299 GoogleMaps 1 ♂, 2 ♀♀, 1 juv.; Lasithi, Agia Fotia ; 35.02° N, 25.87° E; 6 Jul. 2007; H. Eikamp and U. Kluge leg.; SMF 57161 GoogleMaps 2 ♀♀; Lasithi, Bembonas, Tripti Mts ; 35.1° N, 25.9° E; 10 May 2009; K. Eckl and H. Eikamp leg.; SMF GoogleMaps 5 ♀♀; Lasithi, Exo Mouliana ; 35.17° N, 25.99° E; 18 Jul. 1970; A. Senglet leg.; MHNG GoogleMaps .

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 GoogleMaps 1 ♀; Truva , “84/15”; 39.95° N, 26.24° E; date and collector unknown; SMF GoogleMaps . – 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 GoogleMaps 1 ♂, 1 ♀; Çiğli (airport); 38.516° N, 27.013° E; 15 Apr. 1973; A Vigna leg.; MHNG GoogleMaps 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 GoogleMaps . – 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 GoogleMaps . – Muğla • 1 ♂, 1 ♀; Dalyan to Kaunos (“ Cavnos ”) path; 36.82° N, 28.61° E; 18 May 1997; A. Russell-Smith leg.; ZFMK Ar 22300 GoogleMaps 1 ♂; Marmaris ; 36.85° N, 28.26° E; Sep. 1973; C. and E. Supper leg.; MHNG GoogleMaps . – Antalya • 1 ♂; Xanthos ruins; 36.356° N, 29.318° E; 7 Mar. 1977; Kinzelbach leg.; SMF 37622 GoogleMaps 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 GoogleMaps 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 GoogleMaps 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Tur46 GoogleMaps 1 ♂, 3 ♀♀; Alanya Kalesi, above Kleopatra beach; 36.539° N, 31.992° E; 2 Nov. 2013; S. Huber leg.; ZFMK Ar 22302 GoogleMaps 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 GoogleMaps . – 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 GoogleMaps 1 ♀; NE of Silifke, above Narlıkuyu; 36.45° N, 34.10° E; 24 Apr. 2012; J. Altmann and J. Meier leg.; maquis; SMF GoogleMaps 2 ♀♀, 1 juv.; Içel Narlıkuyu ; 36.44° N, 34.11° E; 5 May 1994; “L.M.” leg.; in rubbish by restaurant; WML GoogleMaps 1 ♂, 1 ♀; Taş Obası (cave); 37.079° N, 34.928° E; 24 Aug. 2007; A. Topçu leg.; NOHUAM GoogleMaps . – Nevşehir • 2 ♂♂; Özkonak ; 38.81° N, 34.84° E; 16 Jun. 1993; C. Felton leg.; WML GoogleMaps . – Diyarbakır • 2 ♂♂; Çermik (“ Germik ”); 38.13° N, 39.45° E; 27 Jul. 2004; T. Danışman leg.; ZFMK Ar 5216 GoogleMaps .

CYPRUS – Limassol • 1 ♂, 1 ♀, 1 juv.; Pissouri ; 34.67° N, 32.70° E; 27 Apr. 1982; J. Murphy leg.; beach litter; MMUE Murphy #12617 GoogleMaps . – Famagusta • 2 ♀♀, 3 juvs; Fig Tree Bay , “Cave I”; 35.01° N, 34.06° E; 23 Oct. 1996; P. Strinati leg.; MHNG GoogleMaps .

GEORGIA – Tbilisi • 1 ♀; Tbilisi , Dighom I ; 41.778° N, 44.701° E; 17 Jul. 2019; Karalashvili, Seropian and Krammer leg.; ZFMK Ar 21279 GoogleMaps . – 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 GoogleMaps 1 ♂; Lagodekhi National Park ; 41.847° N, 46.284° E; 9 Jul. 2019; Karalashvili and Krammer leg.; forest; ZFMK Ar 21261 GoogleMaps 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 GoogleMaps .

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 GoogleMaps 4 ♀♀, 3 juvs; Jayrud (“Djeroud”); 33.807° N, 36.742° E; 1911; H. Gadeau de Kerville leg.; in house; MNHN GoogleMaps 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 GoogleMaps .

ISRAEL • 3 ♀♀, 1 juv.; Ahihud, E Akko[n]; 32.91° N, 35.17° E; 10 Apr. 1987; W. Heinz leg.; SMF GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Isr39 GoogleMaps 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 GoogleMaps 1 ♀; Nof Yam ; 32.19° N, 34.81° E; 17 Feb. 1968; collector unknown; AMNH GoogleMaps 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 GoogleMaps 1 ♀; near Ramla, Kibbuz Gezer ; 31.88° N, 34.92° E; 11 Aug. 1984; H. Feldmann leg.; indoors on ceiling; AMNH GoogleMaps 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 GoogleMaps 1 ♀, 2 juvs; near Ramallah, Bayt Ghur al-Tahta ; 31.895° N, 35.080° E; 6 Apr. 1987; W. Heinz leg.; SMF GoogleMaps 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 GoogleMaps 1 ♀; Jerusalem; 31.77° N, 35.21° E; Apr. 1952; Khuri leg.; MCZ 34064 GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 1 ♀ (in pure ethanol); same date as for preceding; ZFMK Isr59 GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 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 GoogleMaps 2 ♂♂, 4 ♀♀, + juvs; Sede Boqer ; 30.85° N, 34.78° E; Jan. 1987; V. and B. Roth; MCZ 34069 GoogleMaps 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 GoogleMaps 1 ♀ (in pure ethanol); same collection data as for preceding; SMF GoogleMaps 2 ♀♀; same collection data as for preceding; 3 Sep. 2011; SMF GoogleMaps 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 GoogleMaps 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 GoogleMaps 6 juvs; same locality as for preceding; 6 Sep. 2011; P. Jäger leg.; stream with water, reed, shrubs, under stones; SMF GoogleMaps 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 GoogleMaps 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 GoogleMaps .

JORDAN – Balqa • 1 ♂, 1 juv.; Fuhais [Fuhays]; 32.010° N, 35.768° E; Apr. 1980; F. Krupp and W. Schneider leg.; SMF GoogleMaps . – Madaba • 2 ♀♀, 2 juvs; Mt Nebo ; 31.77° N, 35.73° E; Jul. 1989; E. Hyazin leg.; MRAC 169974 View Materials GoogleMaps 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 GoogleMaps 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 GoogleMaps . – 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 GoogleMaps 2 ♂♂, 2 ♀♀ (in pure ethanol); same collection data as for preceding; ZFMK Isr53 GoogleMaps . – 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 GoogleMaps 1 ♀, 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Isr37 GoogleMaps . – 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 GoogleMaps 1 ♂, 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Isr52 GoogleMaps 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 GoogleMaps . – Drâa-Tafilalet • 2 ♂♂, 3 ♀♀, 1 juv.; Midelt ; 32.68° N, 4.73° W; 19 Dec. 1986; V. and B. Roth leg.; CAS GoogleMaps 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 . GoogleMaps

ALGERIA – Médéa • 2 ♂♂, 1 ♀; Médéa ; 36.26° N, 2.75° E; date unknown; P.-H. Lucas leg.; MNHN GoogleMaps . – Algiers • 1 ♀; Algiers ; 36.75° N, 3.05° E; 3 Nov. 1948; B. Malkin leg.; AMNH GoogleMaps 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 GoogleMaps 2 ♀♀; Tozeur ; 33.92° N, 8.12° E; Jul. 1972; E. and C. Supper leg.; MHNG GoogleMaps .

TUNISIA – Jendouba • 2 ♂♂, + juvs; “ Tabarca ( Khroumirie )”; ~ 36.9° N, 8.7° E; 1907; H. Gadeau de Kerville leg.; MNHN GoogleMaps 1 ♂, 3 ♀♀, 1 juv. (2 vials); Ayn Darahim (“ Ain-Draham ”); 36.78° N, 8.69° E; 1899; G. Seurat leg.; MNHN GoogleMaps . – Tunis • 1 ♂, 2 ♀♀, 1 juv.; Carthago ; 36.86° N, 10.33° E; 11–12 Oct. 1948; B. Malkin leg.; AMNH GoogleMaps 1 ♀; La Marsa ; 36.89° N, 10.32° E; Jul. 1972; E. and C. Supper leg.; MHNG GoogleMaps . – Gabès • 1 ♂; Gabès ; 33.88° N, 10.10° E; 20 Mar. 1961; Walch leg.; SMF 24801 GoogleMaps . – Beja • 1 ♂, 1 juv.; Beja ; 36.73° N, 9.19° E; 23 Mar. 2006; A. López and C. Ribera leg.; CRBA 1271 GoogleMaps . – Kairouan • 1 ♂, 2 ♀♀; Djebel-Trozza ; 35.56° N, 9.59° E; 1932; F. Santschi leg.; MHNG GoogleMaps . – Kef • 1 ♂, 1 ♀, 1 juv.; El Kef ; 36.17° N, 8.70° E; Jul. 1972; E. and C. Supper leg.; MHNG GoogleMaps .

LIBYA – Al Marquab • 1 ♀; Leptis Magna ; 32.63° N, 14.29° E; 18–26 Aug. 1948; B. Malkin leg.; AMNH GoogleMaps . – 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 GoogleMaps .

EGYPT – Dakahlia • 6 ♂♂, 6 ♀♀, 6 juvs; Mansoura (“ Mansurah ”); 31.04° N, 31.38° E; date unknown; I. Sörensen leg.; ZMUC GoogleMaps . – Beheira • 3 ♂♂, 8 ♀♀, + juvs; Bir Hooker ; 30.38° N, 30.35° E; 1901; J. Dewitz leg.; MNHN GoogleMaps . – Cairo • 1 ♀; Maadi ; 29.96° N, 31.26° E; further collection data on label unclear; MHNG GoogleMaps 1 ♀; Cairo [city]; 30.0° N, 31.2° E; date and collector unknown; SMF Roewer #4783 GoogleMaps . – Luxor • 2 ♂♂, + juvs; Luxor ; 25.70° N, 32.65° E; 2 Nov. 1996; P. Jäger leg.; in house; SMF GoogleMaps .

AUSTRALIA – Western Australia • 1 ♀, 1 juv.; Morawa Motel ; 29.217° S, 116.017° E; 23 May 1996; M.S. Harvey leg.; in building; WAM GoogleMaps 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 GoogleMaps 1 ♂; Maylands ; 31.93° S, 115.90° E; 7 Jan. 1992; J.M. Waldock leg.; WAM 99/1579 GoogleMaps 1 ♀; same collection data as for preceding; 27 Dec. 1991; WAM 99/1578 GoogleMaps 1 ♂; East Victoria Park ; 31.983° S, 115.900° E; 15 Jan. 1997; J.M. Waldock leg.; WAM 99/2097 GoogleMaps 1 ♂; same collection data as for preceding; 13 Sep. 1992; in house; WAM 99/1776 GoogleMaps 1 ♂; same collection data as for preceding; 19 Nov. 1994; WAM 99/1752 GoogleMaps . – South Australia • 1 ♂, 1 ♀; Pondanna Outstation ; 32.56° S, 135.55° E; 10 Dec. 1989; D. Hirst leg.; SAM 99/680 GoogleMaps 1 ♀; Flinders Range National Park, Oraparina ; 31.37° S, 138.73° E; Dec. 1984; B. Guerin leg.; SAM 99/679 GoogleMaps 1 ♀, 1 juv.; Erudine Station via Yunta; 31.43° S, 139.43° E; 22 Apr. 1980; J. McEntee leg.; SAM 99/678 GoogleMaps 1 ♂; Middleback Station ; 32.94° S, 137.40° E; Oct. 1983; B. Guerin leg.; SAM 99/682 GoogleMaps 1 ♀; Mt Lofty Ranges, Hawthorndene ; 35.02° S, 138.63° E; 18 Jan. 1992; L.N. Nicolson leg.; in garden; SAM 99/683 GoogleMaps 2 ♀♀, 2 juvs; Stoneleigh Park near Meningie; 35.70° S, 139.35° E; Mar. 1969; A.W. Forbes leg.; SAM 99/684 GoogleMaps 1 ♂, 2 juvs; Murray Range near Kingston; 34.2° S, 140.3° E; 17 Dec. 1978; M.R. Gray leg.; AMS GoogleMaps 2 ♂♂, 6 ♀♀, 4 juvs; near Pryap; 34.45° S, 140.50° E; 2–9 Jun. 1990; L.N. Nicolson leg.; SAM 99/670 GoogleMaps 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 GoogleMaps 1 ♂; Loxton ; 34.45° S, 140.57° E; 11 Jun. 1990; L.N. Nicolson leg.; in house; SAM 99/669 GoogleMaps . – Queensland • 1 ♀; south side of Lake Broadwater; 27.357° S, 151.098° E; 1–15 Nov. 1984; V. Wood leg.; buildings; QMB S49733 View Materials GoogleMaps 2 ♀♀; SW of Dalby, Lake Broadwater ; 27.35° S, 151.10° E; 9 Dec. 1987; J. Gallon leg.; cottage; QMB S14509 View Materials GoogleMaps 1 ♂; Texas ; 28.85° S, 151.17° E; date unknown; Harvey and Katzman leg.; in building; WAM GoogleMaps . – New South Wales • 1 ♂, 1 ♀; Duntroon Station ; 31.03° S, 143.04° E; Oct.–Nov. 1980; D. Hirst leg.; SAM 99/655 GoogleMaps 1 ♀, 2 juvs; Weinteriga ; 31.60° S, 142.95° E; 15 Apr. 1981; D. Hirst leg.; SAM 99/659 GoogleMaps 2 ♂♂; Menindee Lakes Caravan Park ; 32.354° S, 142.404° E; 12 Apr. 1981; D. Hirst leg.; SAM 99/657 , 99/658 GoogleMaps 3 ♂♂, 1 juv.; Sulcor ; 30.85° S, 150.81° E; 27 May 1995; S. Eberhard leg.; cave, twilight zone; AMS KS 49263 GoogleMaps . – Victoria • 1 ♂; 30 km N of Kaniva, near Broughton; 36.17° S, 141.35° E; 25 May 1988; D. Hirst leg.; SAM 99/661 GoogleMaps .

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 View Figs 5–12 . Ocular area slightly raised; each secondary eye with small accompanying elevation ( Fig. 43 View Figs 43–50 ). 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 View Figs 51–58 ). ALS with one widened spigot and one pointed spigot; PMS with two pointed spigots ( Fig. 56 View Figs 51–58 ).

CHELICERAE. As in Fig. 16 View Figs 13–19 ; 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 View Figs 51–58 ); distance between tips of modified hairs: 390 µm; without proximal frontal protrusion; lateral stridulatory ridges distinct ( Fig. 47 View Figs 43–50 ), distances between ridges ~12 µm.

PALPS. As is Figs 13–15 View Figs 13–19 ; 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 View Figs 51–58 ); tarsus without macrotrichia; procursus ( Figs 20–22 View Figs 20–28 ) straight, dorsal hairs not or only slightly curved upwards; proximally on prolateral-ventral side with prominent process free of hairs (arrow in Fig. 20 View Figs 20–28 ), 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 View Figs 43–50 ); genital bulb ( Figs 23–26 View Figs 20–28 ) 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 View Figs 43–50 ).

LEGS. With single ventral rows of spines on femur 1 (~31), tibia 1 (~38) and femur 2 (~14) ( Figs 17– 19 View Figs 13–19 ); 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 View Figs 29–35 ) except 2–3 distally; all tarsal organs capsulate with simple round rim ( Fig. 53 View Figs 51–58 ).

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 View Figs 5–12 ) but palp with strongly widened distal segments (tibia and tarsus; Figs 29–30 View Figs 29–35 ), without joints between tibia and tarsus; with strong median process posteriorly on sternum ( Figs 31 View Figs 29–35 , 58 View Figs 51–58 ) possibly acting against ventral process anteriorly on abdomen ( Figs 38 View Figs 36–42 , 57 View Figs 51–58 ; but see Discussion); with fewer but stronger stridulatory ridges ( Fig. 48 View Figs 43–50 ; 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 View Figs 29–35 ) 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 View Figs 29–35 ). 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 View Figs 36–42 and 57 View Figs 51–58 ; 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 View Figs 36–42 ). Internal genitalia ( Figs 27–28 View Figs 20–28 , 40–42 View Figs 36–42 ) 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 View Fig ). 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.

MNHN

France, Paris, Museum National d'Histoire Naturelle

SMF

Germany, Frankfurt-am-Main, Forschungsinstitut und Naturmuseum Senckenberg

USNM

USA, Washington D.C., National Museum of Natural History, [formerly, United States National Museum]

MMUE

United Kingdom, Manchester, The University, Manchester Museum

MGAB

Romania, Bucharest, Museum of Natual History "Grigore Antipa"

WML

UK, Liverpool, World Museum Liverpool

AMNH

USA, New York, New York, American Museum of Natural History

CAS

USA, California, San Francisco, California Academy of Sciences

ZMUC

Denmark, Kobenhavn [= Copenhagen], University of Copenhagen, Zoological Museum

AMS

Australia, New South Wales, Sydney, Australian Museum

QMB

Australia, Queensland, South Brisbane, Queensland Museum

MNHN

Museum National d'Histoire Naturelle

SMF

Forschungsinstitut und Natur-Museum Senckenberg

MCZ

Museum of Comparative Zoology

MHNG

Museum d'Histoire Naturelle

USNM

Smithsonian Institution, National Museum of Natural History

ZFMK

Zoologisches Forschungsmuseum Alexander Koenig

MMUE

Museum of Manchester University

ZMMU

Zoological Museum, Moscow Lomonosov State University

MGAB

Muzeul de Istorie Naturala "Grigore Antipa"

AMNH

American Museum of Natural History

CAS

California Academy of Sciences

ZMUC

Zoological Museum, University of Copenhagen

WAM

Western Australian Museum

QMB

Queensland Museum, Brisbane

Kingdom

Animalia

Phylum

Arthropoda

Class

Arachnida

Order

Araneae

Family

Pholcidae

Genus

Holocnemus

Loc

Holocnemus pluchei ( Scopoli, 1763 )

Huber, Bernhard A. 2022
2022
Loc

Holocnemus pluchii

Kritscher E. 1996: 123
Nicolic F. & Polenec A. 1981: 20
Timm H. 1976: 70
Senglet A. 1971: 354
Drensky P. 1939: 248
Kolosvary G. 1938: 65
Drensky P. 1936: 55
Kratochvil J. 1932: 2
Roewer C. F. 1928: 120
1928
Loc

Holocnemus pluchei

Kumada K. 2021: 17
Cargnelutti F. & Calbacho-Rosa L. & Unates D. & Costa-Schmidt L. E. & Cordoba-Aguilar A. & Peretti A. V. 2020: 536
Calbacho-Rosa L. & Cargnelutti F. & Cordoba-Aguilar A. & Peretti A. V. 2019: 1
Naumova M. V. & Lazarov S. P. & Deltshev C. D. 2019: 76
Ponomarev A. V. & Snegovaya N. Y. & Shmatko V. Y. 2019: 233
Cargnelutti F. & Calbacho-Rosa L. & Cordoba-Aguilar A. & Peretti A. V. 2018: 616
Lecigne S. 2018: 50
Barrientos J. A. & Bosco Febrer J. 2017: 22
Gajic I. & Grbic G. 2016: 54
Naumova M. V. & Lazarov S. P. & Petrov B. P. & Deltshev C. D. 2016: 434
Rozwalka R. & Rutkowski T. & Bielak-Bielecki P. 2016: 73
Reiser N. & Neumann J. 2014: 24
Deltshev C. & Komnenov M. & Blagoev G. & Georgiev T. & Lazarov S. & Stojkostka E. & Naumova M. 2013: 7
Calbacho-Rosa L. & Galicia-Mendoza I. & Dutto M. S. & Cordoba-Aguilar A. & Peretti A. V. 2013: 407
Bosmans R. & Van Keer J. & Russell-Smith A. & Kronestedt T. & Alderweireldt M. & Bosselaers J. & de Koninck H. 2013: 24
Turkes T. & Karabulut H. 2013: 619
Benhadi-Marin J. & Barrientos J. A. & Ferrandez M. A. & Henriques S. & Morano E. 2013: 75
Deltshev C. & Vrenosi B. & Blagoev G. & Lazarov S. 2011: 130
Draksic M. & Katusic L. 2011: 168
Dutto M. S. & Calbacho-Rosa L. & Peretti A. V. 2011: 1040
Vetter R. S. & Reierson D. A. & Rust M. K. 2011: 601
Kunt K. B. & Yagmur E. A. & Ozkutuk R. S. & Durmus H. & Anlas S. 2010: 32
Calbacho-Rosa L. & Cordoba-Aguilar A. & Peretti A. V. 2010: 1267
van Helsdingen P. J. 2010: 27
Laborda A. & Simo M. 2008: 262
Van Keer K. 2007: 48
Ferrandez M. A. & Morano E. & Fernandez de Cespedes H. & Camargo M. 2006: 77
Kovacs G. & Szinetar C. & Janos E. 2006: 9
Topcu A. & Demir H. & Seyyar O. & Turkes T. 2005: 289
Van Keer K. & Van Keer J. 2004: 79
Deltshev C. & Lazarov S. & Petrov B. P. 2003: 12
Gasparo F. 2003: 62
Hajer J. & Rehakova D. 2003: 345
Van Keer K. & Van Keer J. 2001: 82
Jager P. 2000: 51
Johnson S. A. & Jakob E. M. 1999: 957
Huber B. A. 1995: 291
Jager P. 1995: 20
Melic A. 1994: 15
Jakob E. M. 1991: 711
Heimer S. & Nentwig W. 1991: 40
Jakob E. M. & Dingle H. 1990: 95
Porter A. H. & Jakob E. M. 1990: 313
Barrientos J. A. & Ferrandez M. A. 1982: 82
Wunderlich J. 1980: 227
Constantini G. P. 1975: 90
Brignoli P. M. 1971: 82
Roewer C. F. 1959: 10
Caporiacco L. di 1948: 38
Kratochvil J. 1940: 7
Wiehle H. 1933: 242
Dalmas C. de 1920: 59
Simon E. 1914: 237
1914
Loc

Pholcus rivulatus

Koch L. 1875: 25
1875
Loc

Holocnemus rivulatus

Franganillo P. B. 1926: 70
Franganillo P. B. 1925: 35
Lessert R. de 1910: 89
Strand E. 1908: 93
Damin N. 1900: 21
Simon E. 1874: 256
Simon E. 1873: 49
1873
Loc

Pholcus pluchii

Simon E. 1866: 122
1866
Loc

Pholcus barbarus

Koch C. 1873: 113
Simon E. 1866: 123
1866
Loc

Pholcus ruralis

Simon E. 1873: 49
Blackwall J. 1858: 432
1858
Loc

Pholcus impressus

Simon E. 1866: 122
Koch C. L. 1850: 31
1850
Loc

Pholcus barbarus

Simon E. 1873: 49
Lucas H. 1846: 237
1846
Loc

Pholcus impressus C.L. Koch, 1837: 99

Simon E. 1873: 49
Koch C. L. 1837: 99
1837
Loc

Pholcus rivulatus

Koch L. 1875: 25
Walckenaer C. A. 1837: 653
Savigny J. - C. & Audouin V. 1826: 140
1826
Loc

pluchii

Rossi P. 1790: 134
1790
Loc

Pholcus pluchii

Simon 1866: 122
Forsskål 1775: 86
1775
Loc

pluchii

Scopoli J. A. 1763: 404
1763
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