Crossopriza lyoni ( Blackwall, 1867 )

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 : 193-204

Crossopriza lyoni ( Blackwall, 1867)

Figs 351 , 702–705 , 783–805

Pholcus Lyoni Blackwall, 1867: 392 .

Crossopriza brasiliensis Mello-Leitão, 1935: 94 , fig. 13a–c. Synonymized in Huber et al. 1999.

Crossopriza mucronata Mello-Leitão, 1942: 389 , figs 1–2. Synonymized in Huber et al. 1999.

Crossopriza françoisi Millot, 1946: 154 , figs 29a–b, 30b. Synonymized in Huber et al. 1999.

Crossopriza stridulans Millot, 1946: 156 , fig. 31a–b. Synonymized in Huber et al. 1999.

Crossopriza nigrescens Millot, 1946: 155 , fig. 30a. syn. nov.

Tibiosa caracensis González-Sponga, 2006: 11 , pl. 1 figs 1–10. Synonymized in Huber 2009b.

Tibiosa casanaimensis González-Sponga, 2006: 14 , pl. 2 figs 1–9. Synonymized in Huber 2009b.

Tibiosa coreana González-Sponga, 2006: 17 , pl. 3 figs 1–9. Synonymized in Huber 2009b.

Tibiosa guayanesa González-Sponga, 2006: 20 , pl. 4 figs 1–9. Synonymized in Huber 2009b.

Tibiosa moraensis González-Sponga, 2006: 23 , pl. 5 figs 1–9. Synonymized in Huber 2009b.

Smeringopus lyonii – Thorell 1895: 70; 1898: 274.

Crossopriza lyoni – Pocock 1900: 240. — Strand 1907: 125. — Simon 1909: 80. — Sherriffs 1919: 229. — Chamberlin 1924: 4. – Dyal 1935: 168, pl. 15 figs 90–96. — Zhu & Wang 1963: 462, figs 4, 7–8. — Chrysanthus 1967: 96, figs 20–24. — Tikader & Biswas 1981: 18, pl. 1 figs 13–15. — Yaginuma 1982: 15; 1986: 31, fig. 17.5a, e, p. — Kim 1988: 35, figs 1–6. — Chikuni 1989: 29, fig. 7. – Chen & Zhang 1991: 72, fig. 61.1–3. — Edwards 1993: 1, figs 1–2. — Majumder & Biswas 1993: 2. — Roth 1994: 145. — Huber et al. 1999: 1, figs 1–12; 2014a: 6. — Song et al. 1999: 52, fig. 22h–o. — Huber 2000: fig. 98; 2001: 136; 2009b: 65; 2014: 140; 2019: 51. — Murphy & Murphy 2000: 247. — Irie 2001: 7, figs 1–2(1–8); 2009: 106, figs (2-2-13) 3–5, pl. 4 fig. 2. — Van Keer & Van Keer 2001: 82; 2004: 79. — Guarisco & Cutler 2003: 105, fig. 1. — Van Keer 2007: 53 . — Jäger 2007: 34 . — Colmenares García 2008: 88, fig. 2a–c. — Beatty et al. 2008: 9, figs 17–18, 46–47. — Carvalho & Avelino 2010: 6. — Huber & Warui 2012: 8 . — Jäger et al. 2012: 80, fig. 3. — Yin et al. 2012: 163, fig. 30. — Huber & Kwapong 2013: 7 , fig. 16. — Tong 2013: 61. — Raychaudhuri & Saha 2015: 85, pl. 19 figs 514–518. — Bauer et al. 2016: 4, figs 1–7. — Huber & Villarreal 2020: 57 .

Crossopriza stridulans – Roth 1985: B33-1.

Justification of new synonymy

Even though the type material of C. nigrescens (a single juvenile specimen) is apparently lost, the name is here synonymized, for three reasons. First, all adult Madagascan specimens of Crossopriza studied are C. lyoni ( Huber et al. 1999; Irie 2001; herein); second, the type of C. nigrescens was collected in a house, suggesting a synanthropic species; third, the character that Millot (1946) considered diagnostic (the dark coloration) varies considerably among specimens, even within a locality.

Diagnosis

Distinguished from known congeners by details of male palp ( Figs 786–791 ; procursus ventral sclerite with transparent retrolateral branch; procursus tip with wide and rounded dorsal process; distal bulbal sclerite with distinctive row of apophyses on prolateral side); from the similar C. surobi sp. nov. (identical male chelicerae) also by shorter epigynum and elongate pore plates ( Figs 794–795 ); from the similar C. maculipes also by male chelicerae ( Fig. 793 ; lateral apophyses in lateral view long) and by female internal genitalia ( Fig. 795 ; pore plates elongated).

Type material

Syntypes of Pholcus lyoni

INDIA • ♂♂, ♀♀ (unspecified number); Delhi; 28.6° N, 77.2° E; Meerut; 28.97° N, 77.70° E; and Agra; 27.2° N, 78.0° E; dates and collector(s) unknown; possibly lost .

Syntypes of Crossopriza brasiliensis

BRAZIL • 1 ♂, 1 ♀, 1 juv. (examined); Bahia, Paraguassú ; 12.80° S, 38.87° W; date unknown; O. Leonardos leg.; MNRJ 42313 .

Syntypes of Crossopriza mucronata

ARGENTINA • 1 ♀, 1 juv. (examined); Santiago del Estero, Beltrán ; 27.83° S, 64.06° W; date unknown; M. Birabén leg.; MLP 15800 • 1 ♀ (not examined) (see Huber et al. 1999); same collection data as for preceding; with P. Brignoli`s (unpublished) lectotype designation; MLP .

Holotype of Crossopriza francoisi

MADAGASCAR • ♂ (examined); Mahajanga, Maevatanana ; 16.95° S, 46.83° E; Jul. 1945; J. Millot leg.; in bathroom; MNHN.

Holotype of Crossopriza stridulans

MADAGASCAR • ♀ (examined); Mahajanga, Majunga [= Mahajanga]; 15.71° S, 46.32° E; date and collector unknown; MNHN.

Holotype of Crossopriza nigrescens

MADAGASCAR • 1 juv.; Antsiranana, Ankarana Sud ; 12.97° S, 49.14° E; 1945; J. Millot leg.; apparently lost (not found in MNHN).

Tibiosa spp.

VENEZUELA. See Huber & Villarreal (2020).

Other material examined

Arranged from West to East and (within longitudes) from North to South.

USA – California • 1 ♀; Orange County, Orange ; 33.80° N, 117.85° W; 23 Feb. 2000; J. Coleman leg.; CAS 9027397 . – New Jersey • 1 ♀; Somerset County, Neshanic ; 40.498° N, 74.720° W; date unknown; E.S. Gaffney leg.; AMNH. – Texas • 1 ♂, 3 ♀♀; Brazos County, College Station ; 30.63° N, 96.33° W; date unknown; R.G. Breene leg.; AMNH .

NICARAGUA • 2 ♂♂, 1 ♀; Ciudad de León; 12.43° N, 86.88° W; 10 Aug. 1995; L.F. Armas leg.; in house; MELN • 1 juv.; Managua, Laguna Xiloa ; 12.209° N, 86.320° W; 13 Jul. 1995; L.F. Armas, J.M. Maes, and J.T. Goodwin leg.; MELN .

DOMINICAN REPUBLIC • 1 ♂, 1 ♀ (in pure ethanol); Barahona; 18.217° N, 71.100° W; 30 m a.s.l.; 9 Dec. 2007; B.A. Huber leg.; in building; ZFMK Hai 103 .

COLOMBIA • 1 ♂, 1 ♀; Meta, Puerto Lleras, Lomalinda ; 3.305° N, 73.364° W; 300 m a.s.l.; Mar. 1994; B.T. Carroll, V. and B. Roth leg.; CAS 9027460 • 1 ♂, 2 ♀♀; same locality as for preceding; date unknown; Roth leg.; CAS 9027261 • 1 ♀; same locality as for preceding; May 1987; B.T. Carroll leg.; CAS 9027465 .

VENEZUELA • approximately 40 ♂♂, 40 ♀♀; see Huber & Villarreal (2020) .

BRAZIL – Mato Grosso • 1 ♂; Corumba ; 19.01° S, 57.65° W; 21 Sep. 1956; C. Sincole leg.; under rocks; CAS 9027331 . – Mato Grosso do Sul • 1 ♀; Upper Paraguay River, Porto Esperança ; 19.61° S, 57.45° W; date and collector unknown; AMNH . – Minas Gerais • 1 ♂, 1 ♀, 1 juv.; Governador Valadares, Ibituruna ; 18.845° S, 41.943° W; 9 Sep. 1982; L. Sorkin et al. leg.; in building; AMNH • 1 ♀; Governador Valadares , northern part of city; 18.84° S, 41.95° W; 1 Sep. 1982; L. Sorkin and C.E. de Assis Bandeira leg.; AMNH • 1 ♂, 5 ♀♀; Governador Valadares ; 18.86° S, 41.95° W; 26 Mar. 1981; L. Sorkin and T. Spitzman leg.; ceiling in marketplace; AMNH • 2 ♀♀; Curvelo ; 18.75° S, 44.43° W; 23 Oct. 1943; F. Pough leg.; AMNH . – Pará • 1 ♂, 1 ♀; Itaituba ; 4.26° S, 56.00° W; 7 Dec. 1991; A.A. Lise leg.; MCN • 1 ♂, 3 ♀♀; same locality as for preceding; 7 Dec. 1991; F. Guilhow leg.; MCP 4629 , 4630 . – São Paulo • several ♀♀ and juvs (not counted); Jaboticabal ; 21.25° S, 48.32° W; 1979; W. and L. Miller leg.; MCZ .

PARAGUAY • 1 juv.; Boquerón, 19 km N of Filadelfia, Estancia Iparoma ; 22.150° S, 60.034° W; 5. Oct. 1978; K.L. Anderson leg.; AMNH .

ARGENTINA – Chaco • 1 ♀; Resistencia ; 27.45° S, 59.00° W; Oct. 1942; Freiberg leg.; MACN Ar 4347 • 1 ♀; Parque Nacional Chaco; 26.8081° S, 59.6075° W; 80 m a.s.l.; 11–13 Nov. 2007; C. Grismado et al. leg.; MACN Ar 13662. – La Rioja • 1 ♀; La Rioja ; 29.41° S, 66.86° W; Apr. 1914; E. and P. Boman leg.; AMNH • 1 ♂, 1 ♀, 1 juv.; same locality as for preceding; 15 Apr. 1914; Boman et al. leg.; MACN Ar 4345 • 1 ♂, 3 ♀♀, 3 juvs; “Guayapa”, Patquia ; 30.04° S, 66.88° W; Oct. 1963; E. Galiano leg.; MACN Ar 20118 • 3 juvs; same locality as for preceding (“ Guayapa ”); Oct. 1965; Maury leg.; MACN Ar 20033 . – Santiago del Estero • 2 ♀♀; Santiago del Estero ; 27.786° S, 64.266° W; Nov. 1966; E. Galiano leg.; MACN Ar 19949 • 4 ♀♀; same locality as for preceding; Aug. 1939; A. Prasen leg.; MACN Ar 4346 • 1 ♂; same locality as for preceding; 20 Sep. 1963; collector unknown; MACN Ar 20119 . – Tucumán • 1 ♂, 4 ♀♀; San Miguel de Tucumán ; 26.81° S, 65.22° W; 1–15 May 1950; M.L. Aczel leg.; AMNH .

GERMANY • 1 ♂, 2 ♀♀; see Bauer et al. (2016) .

SENEGAL • 1 ♀; see Huber & Kwapong (2013).

THE GAMBIA • 1 ♂, 4 ♀♀; see Huber & Kwapong (2013) .

MALI • 1 ♂; Bamako ; 12.6° N, 8.0° W; May–Jun. 1977; W.S. Settle leg.; CAS • 1 ♂, 3 ♀♀, 2 juvs; Gao ; 16.275° N, 0.050° W; 23–25 Nov. 1948; B. Malkin leg.; AMNH • 3 ♂♂, 9 ♀♀; see Huber & Kwapong (2013).

GUINEA • 6 ♂♂, 9 ♀♀; see Huber & Kwapong (2013) .

SIERRA LEONE • 1 ♀; see Huber & Kwapong (2013) .

IVORY COAST • 5 ♂♂, 14 ♀♀; see Huber & Kwapong (2013) .

BURKINA FASO • 12 ♂♂, 18 ♀♀; see Huber & Kwapong (2013) .

GHANA • 13 ♂♂, 13 ♀♀; see Huber & Kwapong (2013) .

BENIN • 5 ♂♂, 12 ♀♀; see Huber & Kwapong (2013) .

NIGER • 8 ♂♂, 28 ♀♀; see Huber & Kwapong (2013) .

NIGERIA • 1 ♂, 1 ♀; Oguta Lake ; 5.71° N, 6.80° E; 10–24 Feb. 1992; H. Segers leg.; motel; MRAC 174630 • 1 ♀; Ile-Ife, Obafemi, Awolowo University Campus ; 7.43° N, 3.89° E; 1991; H. Segers leg.; MRAC 174602 part • 1 ♀; Lagos, Lagos University ; 6.519° N, 3.392° E; 1973; Usua leg.; BMNH .

CHAD • 4 ♂♂, 1 ♀, 5 juvs; Bol ; 13.46° N, 14.71° E; 9–15 Dec. 1978; A. Spielman leg.; MCZ • approximately 6 ♂♂, 13 ♀♀, + juvs; between Bongor (“ Bougar ”) and N’Djamena (“ Fort Lamy ”), “depression inondée de la Karaska”; 11.1° N, 15.1° E; Oct.–Dec. 1965; Y. Brandily leg.; MRAC 132894 , 132896 , 132897 • approximately 9 ♂♂, 12 ♀♀, + juvs; N’Djamena , Gardolé (“ Fort Lamy ”); 12.12° N, 15.04° E; 1 Nov. 1965; Y. Brandily leg.; MRAC 132914 to 132916 • 1 ♂, 3 ♀♀, 3 juvs; Bébédja ; 8.68° N, 16.57° E; 1–10 Jul. 1977; G. Ruella leg.; MRAC 151455 .

CAMEROON • 3 ♂♂, 5 ♀♀; Extrême-Nord, Yagoua ; 10.34° N, 15.23° E; 1–5 Aug. 1971; F. Puylaert leg.; MRAC 143673 • 2 ♀♀; see Huber et al. (2014b).

GABON • 3 ♂♂, 1 ♀; see Huber et al. (2014b) .

SUDAN • 1 ♂, 1 ♀; Khartoum; 15.55° N, 32.55° E; 1 May 2016; M. Siyam leg.; in houses; ZFMK Ar 22451, Ar 22452 .

ETHIOPIA • 2 ♂♂, 8 ♀♀, 2 juvs (in pure ethanol); Southern Nations, Nationalities and People’s Region, Arba Minch, Nech Sar National Park ; 6.0002° N, 37.5429° E; 1390 m a.s.l.; 19 Jun. 2011; M. Moradmand, V. Hula, and J. Niedobová leg.; in houses; SMF .

UGANDA • 3 ♀♀; see Huber & Warui (2012).

KENYA • 1 ♀; see Huber & Warui (2012).

TANZANIA • 1 ♂; Dar es Salaam; 6.82° S, 39.27° E; 25–26 Oct. 1995; D. Ubick leg.; in house; CAS .

ZIMBABWE • approximately 8 ♂♂, 16 ♀♀; Kariba ; 16.52° S, 28.85° E; 10 Feb. 1995; W.J. Pulawski leg.; CAS • 4 ♂♂, 8 ♀♀, + juvs; Victoria Falls ; 17.93° S, 25.83° E; 1–8 Feb. 1995; W.J. Pulawski leg.; CAS (4 vials) • approximately 3 ♂♂, 8 ♀♀; same locality as for preceding; 19–22 Dec. 1995; W.J. Pulawski leg.; CAS • 6 ♂♂, 6 ♀♀; same locality as for preceding; 29 Nov. 1996; W.J. Pulawski and V. Ahrens leg.; CAS .

MADAGASCAR – Toliara • 1 ♂; Toliara ; 23.35° S, 43.67° E; 17 Sep. 1993; W.E. Steiner and R. Andriamasimanana leg.; USNM • 1 ♀; 20 km N of Toliara “on Ferme d’Auberge rd”; 23.429° S, 43.830° E (the coordinates are not N but SE of Toliara); 200 m a.s.l.; 15–16 Dec. 1999; E.I. Schlinger and M.E. Irwin leg.; CAS .

SEYCHELLES • 1 ♀; Mahé , Bel Ombre; 4.6187° S, 55.3971° E; 30 m a.s.l.; 6 Mar. 2013; C. Hoareau leg.; on wall near buildings; ZFMK Ar 22453 .

PAKISTAN • 2 ♂♂, 2 ♀♀; Lahore, University of the Punjab; 31.5° N, 74.3° E; Sep. 2013; collector unknown; ZFMK Ar 22454 .

INDIA – Andhra Pradesh • 1 ♀; Deccan highlands; 14° N, 77° E; date and collector unknown; SMF RII/3735 . – Delhi • 1 ♂, 3 ♀♀, 1 juv.; New Delhi, Russian Embassy ; 28.593° N, 77.186° E; 1–2 May 1999; Y.M. Marusik leg.; ZFMK Ar 5206 . – Haryana • 1 ♂; Kasauli ; 30.1210° N, 76.4075° E; 240 m a.s.l.; 9 Mar. 2011; P. Jäger leg.; in house and surroundings, farmland; SMF . – Karnataka • 2 ♂♂, 2 ♀♀; Mysore ; 12.30° N, 76.65° E; Dec. 1987; J. Murphy leg.; MMUE Murphy #15830, 15831 . – Maharashtra • 1 ♀; Pune, Agharkar Research Institute ; 18.520° N, 73.832° E; 2 Sep. 1992; G.W. Kendrick leg.; WAM 99 /1489 • 2 ♀♀, 2 juvs; “Kirkoskarvadt, Distr. South Sathra”, probably Kirloskarvadi ; 17.08° N, 74.42° E; date unknown; W. Bindermann leg.; SMF. – Punjab • 2 ♂♂, 4 ♀♀, 2 juvs; Patiala, University campus; 30.35° N, 76.45° E; 24–25 Jun. 1999; Y.M. Marusik leg.; ZFMK Ar 5205 • 1 ♀; same collection data as for preceding; 3–8 May 1999; ZFMK Ar 5201 • 7 ♀♀, 2 juvs; Patiala (“ Patriala ”); 30.35° N, 76.45° E; date and collector unknown; SMF RII/6617 • 1 ♂, 1 ♀; Balanda ; 31.0441° N, 75.4265° E; 220 m a.s.l.; 9 Mar. 2011; P. Jäger leg.; in house, courtyard, and farm; SMF . – Rajastan • 1 ♂, 1 ♀; Pushkar ; 26.49° N, 74.55° E; 11 Nov. 1989; V. Roth leg.; in building; CAS . – Tamil Nadu • 1 ♀; Chennai (“ Madras ”); 13.07° N, 80.25° E; Aug. 1924; collector unknown; SMF Roewer #1148 • 1 ♀; 10 mi SW of Gudiyattam ; 12.86° N, 78.80° E; 3 Apr. 1962; Ross and Cavagnaro leg.; CAS • 1 ♂, 1 ♀; Alagar [koil], 21 km NE of Madurai; 10.074° N, 78.215° E; 27–28 Dec. 1989; V. and B. Roth leg.; CAS • 1 ♂, 12 ♀♀; Coimbatore ; 11.017° N, 76.960° E; 430 m a.s.l.; 8 Mar. 1962; E.S. Ross and D.Q. Cavagnaro leg.; CAS • 1 ♀; 80 km S of Madras, Mahabalipuram ; 12.62° N, 80.19° E; 5 Mar. 1994; J.M. Waldock leg.; in house; WAM 99 /1802 • approximately 6 ♂♂, 11 ♀♀; Vellore ; 12.91° N, 79.13° E; date unknown; Löwenthal leg.; ZMUC • 1 ♀; Mandapam Camp ; 9.28° N, 79.12° E; 5 Oct. 1957; H. Lemche leg.; ZMUC . – Uttarakhand • 1 ♂, 1 ♀; Dehradun City, Forest Research Institute , campus and nearby; 30.342° N, 78.000° E; 660 m a.s.l.; 6–13 May 1999; Y.M. Marusik leg.; ZFMK Ar 5204 • 2 ♀♀; Dehradun Valley ; 30.34° N, 78.00° E; ~ 700 m a.s.l.; 4–13 Aug. 1978; collector unknown; ZMUC . – West Bengal • numerous ♂♂ and ♀♀ (not counted); Kanchrapara ; 22.93° N, 88.43° E; 1–12 Aug. 1944; M. Cazier leg.; AMNH .

NEPAL • 1 ♀; Kathmandu valley , Baneshwar, Ganaba-hal; 27.70° N, 85.32° E; 1350 m a.s.l.; 16–20 Aug. 1980; Martens and Schawaller leg.; SMF .

SRI LANKA • 4 ♂♂, 12 ♀♀; see Huber et al. (1999) and Huber (2019).

MYANMAR • approximately 10 ♂♂, 20 ♀♀; Rangoon; 16.806° N, 96.150° E; date unknown; Lövendal leg.; ZMUC .

THAILAND • 2 ♂♂, 1 ♀; Saraburi Province, 8 mi SE of Saraburi ; 14.45° N, 100.95° E; 100 m a.s.l.; 28 Jul. 1962; Ross and Cavagnaro leg.; CAS • 2 ♀♀, + juvs; Bangkok (= Krung Thep), Chulalong University ; 13.74° N, 100.53° E; 18–19 Oct. 1957; N. Meinkoth leg.; MCZ 76654 • 1 ♀; Lopburi Province, Khok Samrong District, Khao Wong Phrachan ; 14.9633° N, 100.7103° E; 600 m a.s.l.; 24 Jun. 2014; P. Schwendinger leg.; dry secondary forest with bamboo; MHNG • 1 ♂; Prachuap Khiri Khan Province, Reclining Buddha Cave ; 11.864° N, 99.823° E; 70 m a.s.l.; 13 Mar. 2015; B.A. Huber and B. Petcharad leg.; among rocks outside cave; ZFMK Ar 23880 • 1 ♂, 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mal367 .

LAOS – Vientiane • 1 ♀; Vang Vieng, Nam Song; 18.912° N, 102.456° E; 230 m a.s.l.; 12 Nov. 2009; P. Jäger and S. Bayer leg.; SMF • 1 ♂, 2 juvs; Sisathona Distr., Sapanthong Neua; 17.957° N, 102.629° E; 170 m a.s.l.; 26 Mar. 2011; L. Nophasead leg.; SMF. – Oudomxai • 1 ♂, 2 ♀♀; 4 km S of Oudomxai, Lipi provincial protected area; 20.653° N, 102.003° E; 870 m a.s.l.; 18 Apr. 2011; L. Nophasead leg.; SMF. – Bolikhamsai • 1 ♀, 1 juv.; Nam Kading National Protected Area; 18.341° N, 104.144° E; 23 Mar. 2011; P. Jäger and L. Nophasead leg.; disturbed secondary forest; SMF • 1 juv.; same collection data as for preceding; 25 Mar. 2011; SMF • 5 ♂♂, 13 ♀♀; see Jäger (2007) and Jäger et al. (2012).

CAMBODIA • 1 juv.; Banteay Meanchey, Sisophon, La Ang Chinchian 5; 13.5905° N, 102.9442° E; 16 Jan. 2017; H. Steiner leg.; ZFMK Ar 22455 .

VIETNAM • 2 ♂♂, 1 juv.; Kien Giang Prov., Ha Tien, Da Dung mountain ; 10.427° N, 104.476° E; 6 Jan. 2010; H. Steiner leg.; in cave; SMF • 1 ♂; Kien Giang Prov., Kien Luong, Nui Ca Danh mountain ; 10.283° N, 104.565° E; 9 Jan. 2010; H. Steiner leg.; SMF • 1 ♀; Ho Chi Minh (“Saigon”); 10.81° N, 106.64° E; Nov. 1966 – Feb. 1967; P. Fleischer leg.; MCZ .

CHINA • 2 ♂♂, 2 ♀♀; Guilin (“Kweilin”); 25.24° N, 110.18° E; 20 Jul. 1976; A. Jung leg.; CAS .

TAIWAN • 2 ♂♂, 1 ♀; Taichung, Tunghai University campus, near Tunghai Lake ; 24.1815° N, 120.6086° E; 140 m a.s.l.; 17 Jun. 2013; B.A. Huber leg.; at building; ZFMK Ar 23881 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Tai81 .

JAPAN • 2 ♂♂, 1 ♀; Kagawa Pref., Mitoyo-shi, Yamamoto-chô ; 34.18° N, 133.71° E; 7 Jan. 2000; Y. Ihara leg.; ZFMK Ar 5209 .

MALAYSIA • 1 ♂, 6 ♀♀, 2 juvs; Kedah, Kedah Peak (Gunung Jerai), at foot of mountain; 5.8° N, 100.4° E; 6 Jan. 1985; J.A. Beatty leg.; on buildings; BPBM .

SINGAPORE • 2 ♂♂, 1 ♀; Pulau Ubin , near park headquarters; 1.403° N, 103.967° E; 20 m a.s.l.; 16 Feb. 2015; B.A. Huber, J. Koh, and D. Court leg.; in building; ZFMK Ar 23882 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mal233 .

PHILIPPINES – Luzon • 1 ♂; Manila, Umber ; 14.37° N, 121.02° E; 27 Aug. 1950; collector unknown; MCZ 76679 • 1 ♂, 1 ♀; Batangas; 13.76° N, 121.06° E; 4 Aug. 1986; C.K. Starr leg.; in building; AMNH • 1 ♀; Rizal, Alabang ; 14.42° N, 121.04° E; Sep. 1945; B. Malkin leg.; AMNH • 1 ♀; Lingayen ; 16.02° N, 120.23° E; Jun.–Jul. 1945; R.B. Burrows leg.; AMNH • 1 ♂; Banaue ; 16.919° N, 121.058° E; Jan. 1980; P. Schwendinger leg.; MHNG. – Leyte • 2 ♂♂, 2 ♀♀; Baybay ; 10.68° N, 124.80° E; 3–6 Sep. 1984; C.K. Starr leg.; AMNH . – Mindanao • 2 ♀♀; Bukidnon Province, Central Mindanao University ; 7.859° N, 125.051° E; 350 m a.s.l.; 10 Feb. 2014; B.A. Huber and E. Mondejar leg.; in building; ZFMK Ar 22456 .

INDONESIA • 6 ♀♀ (2vials); Papua, Merauke (“Merambe”); 8.49° S, 140.40° E; 1956/57; M. Monulphus leg.; P. Chrysanthus ded.; SMF .

PAPUA NEW GUINEA • 1 ♂; National Capital District; 9.45° S, 147.21° E; 23 Jul. 1985; D. Court leg.; NMNL ARA 17485 .

AUSTRALIA – Northern Territory • 1 ♀; Darwin ; 12.45° S, 130.84° E; 20–31 Mar. 1945; B. Malkin leg.; AMNH • 1 ♀; Darwin, Alawa, Eileen St. ; 12.377° S, 130.874° E; 23 May 1992; J.M. Waldock leg.; WAM 99 /1672 • 1 ♂, 2 ♀♀; 70 mi S of Darwin, Adelaide River ; 13.24° S, 131.10° E; 20–27 Mar. 1945; B. Malkin leg.; AMNH • 1 ♀; 120 mi SE of Darwin, Edge Arnhemland Res., Ben Hole Billabong ; 13.8° S, 131.8° E; Jan.–Feb. 1972; J. Anderson leg.; AMNH • 1 ♀, 3 juvs; Alice Springs ; 23.70° S, 133.88° E; 29 Oct. 1983; E.I. Schlinger and M.E. Erwin leg.; CAS 9027208 • 2 ♂♂, 3 ♀♀, 1 juv.; Katherine ; 14.47° S, 132.26° E; Dec. 1980; R.R. Jackson leg.; in communal webs; QMB S49735 • numerous ♂♂ and ♀♀ (not counted); Kakadu National Park, Narramoor ; 12.696° S, 132.373° E; 25 m a.s.l.; 2 Jul. 1987; M.S. Harvey leg.; WAM 99 /1604 to 99/1613 • 1 ♂; Jabiru Residency ; 12.67° S, 132.84° E; 28 May 1992; Harvey and Waldock leg.; WAM 99 /1674 • 1 ♀; Nhulunbuy ; 12.18° S, 136.68° E; 8 Aug. 1991; A.F. Longbottom leg.; WAM 99 /1675 • 1 ♀; Berry Springs ; 12.7° S, 131.0° E; 26 May 1992; Harvey and Waldock leg.; woodland, on house; WAM 99 /1671 • 1 ♀; Humpty Doo, Marjerrison Rd ; 12.57° S, 131.1° E; 30 Oct. 1985; J.M. Waldock leg.; WAM 99 /1614 • 1 ♀; Uluru National Park ; 25.3° S, 131.0° E (label: 25.038° S, 131.070° E); 15 May 1999; Gray, Milledge, and Smith leg.; in house; AMS KS 56180 . – Queensland • 1 ♀, 1 juv.; Widgee , NW of Gympie; 26.170° S, 152.425° E; 16–20 Feb. 1987; J. Gallon leg.; QMB S50290 • 3 ♀♀, 1 juv.; Lake Broadwater, near Dalby ; 27.35° S, 151.10° E; 26 Jan. 1984; M. Bennie leg.; house; QMB S49726 • 1 ♀; Gatton Agricultural College ; 27.554° S, 152.336° E; 29 May 1986; Bayer leg.; QMB S50276 • 1 ♂; Proa Shi , 20 mi S of Nelia; 20.9° S, 142.2° E; 7–13 May 1978; J. Covacevich leg.; QMB S50262 • 1 ♀, 2 juvs; Mt Isa , township; 20.732° S, 139.485° E; 16–21 Apr. 1996; R. Raven leg.; QMB S33757 • ♂♂ (not counted); near Edungalba ; 23.72° S, 149.85° E; Jul. 1993; D. Walker leg.; QMB S22175 • 1 ♀; Torres Strait, Yam Island ; 9.90° S, 142.77° E; 28 Nov.–2 Dec. 1986; K. Saddler and J. Gallon leg.; under bark; QMB S12418 • 1 ♀, 1 juv.; Palm River Café; 10.96° S, 143.70° E; 21 Oct. 1974; V.E. Davies leg.; QMB S49730 • 1 ♂; Lockerbie Scrub Wilderness Lodge ; 10.78° S, 142.47° E; 9 Dec. 1980; J. Gallon leg.; QMB S13021 • 1 ♂, 1 ♀; Townsville ; 19.26° S, 146.82° E; Sep. 1986; collector unknown; QMB S49734 • 1 ♀; Boulia ; 22.91° S, 139.91° E; Jun. 1981; S. van Dyck and R. Molnar leg.; QMB S49727 • 1 ♀; Doondi , 50 km SW of St. George; 28.28° S, 148.56° E; 10 Sep. 1979; R. Raven leg.; QMB S49732 • 1 ♀, 1 juv.; The Lion’s Den Hotel [near Rossville ?]; 15.705° S, 145.223° E; 15 Apr. 1994; J. Thompson, M. Moulds, and F. Mac Killop leg.; AMS KS 45744 . – Western Australia • 2 ♀♀, 1 juv.; Kalumburu ; 14.295° S, 126.640° E; 19 Aug. 1969; A.R. Main leg.; WAM 99 /1754 to 99/1758 part • 2 ♀♀; Barrow Island , WAPET camp; 20.817° S, 115.433° E; 1–5 Nov. 1993; M.S. Harvey and J.M. Waldock leg.; indoors; WAM 99 /1739, 99/1740 • 2 ♂♂, 1 ♀; Kununurra ; 15.77° S, 128.74° E; 1962; K. Richards leg.; WAM 99 /1679 to 99/1683 • 1 ♂; same locality as for preceding; 8 Jun. 1992; M.S. Harvey and J.M. Waldock leg.; WAM 99 /1653 • 1 ♂, 4 juvs; same locality as for preceding; 25 Nov. 1978; D. Roberts and J. Staddart leg.; in hostel; WAM 99 /1598 to 99/1602 • 1 ♀; Derby ; 17.317° S, 123.633° E; May 1963; A.R. Main leg.; WAM 99 /1688 • 1 ♂; same locality as for preceding; 3 Nov. 1961; A.R. Main leg.; WAM 99 /1702 • 1 ♂, 1 ♀; Drysdale River Station ; 15.702° S, 126.380° E; 15 Jan. 1994; A.F. Longbottom leg.; WAM 99 /1747, 99/1748 • 1 ♂, 1 ♀; same locality as for preceding; 21 Jun. 1993; A.F. Longbottom leg.; WAM 99 /1745, 99/1746 • 1 ♀, 1 juv.; Geikie Gorge ; 18.08° S, 125.71° E; 24 Jul. 1987; B.Y. Main leg.; WAM 99 /1777, 99/1778 • 1 ♀; Geikie Gorge National Park ; 18.110° S, 125.697° E; 7 Jun. 1999; Gray, Milledge, and Smith leg.; under bark; AMS KS 56189 • 1 ♀; Care Spring ; 15.533° S, 128.833° E; 8 Jun. 1992; M.S. Harvey and J.M. Waldock leg.; WAM 99 /1637 • 1 ♂, 2 ♀♀; Fitzroy Crossing ; 18.19° S, 125.57° E; 11 Aug. 1991; J.M. Waldock leg.; outside of house; WAM 99 /1589 to 99/1591 • 1 ♀, 1 juv.; Meda Station ; 17.4° S, 124.0° E; 22 Aug. 1987; A.E. de Jong leg.; WAM 99 /1616, 99/1617 .

FEDERATED STATES OF MICRONESIA • 1 ♂, 1 ♀, 1 gynandromorphic specimen; Chuuk (“Truk”) Island ; 7.45° N, 151.85° E; 10 Mar. 1969; Sabath leg.; MCZ 76633, 76634 .

Redescription

Male ( Guinea, ZFMK Ar 10456)

MEASUREMENTS. Total length 6.5, carapace width 2.4. Distance PME–PME 140 µm; diameter PME 140 × 170 µm; distance PME–ALE 50 µm; diameter AME 130 µm; distance AME–AME 45 µm. Leg 1:

63.4 (18.7 + 0.9 + 17.6 + 22.7 + 3.5), tibia 2: 12.4, tibia 3: 9.2, tibia 4: 10.5; tibia 1 L/d: 70; femora 1–4 diameters: 0.36, 0.32, 0.28, 0.31.

COLOR (in ethanol). Carapace ochre-yellow; carapace pit anteriorly light brown; sternum dark brown; legs ochre-yellow, with indistinct darker rings subdistally on femora and tibiae, with black lines on femora and tibiae, patellae brown; abdomen ochre-gray, with whitish internal marks and some dark marks dorsally and laterally; ventrally with distinct black median band, partly disrupted, with three parallel longitudinal marks behind gonopore.

BODY. Habitus as in Figs 702–703 . Ocular area slightly raised. Deep thoracic pit and pair of furrows diverging from pit toward posterior margin. Clypeus unmodified, rim barely more sclerotized than in female. Sternum wider than long (1.7/1.0), unmodified. Abdomen slightly elongated, dorso-posteriorly angular.

CHELICERAE. As in Figs 792–793 , with two pairs of apophyses, lateral pair with 2–3 small processes (distinct in lateral view), median pair with one large modified cone-shaped hair each; distance between tips of modified hairs 80 µm; lateral stridulatory ridges fine but visible in dissecting microscope; distances between ridges: ~7.5 µm ( Huber et al. 1999).

PALPS. As in Figs 783–785 ; coxa with rounded retrolateral hump; trochanter barely modified; femur distally widened, with rounded ventral protrusion, proximally with prolateral stridulatory pick, retrolateral-ventral rim with row of projecting and sclerotized hair-bases, with barely visible retrolateral transversal line, without retrolateral proximal process; femur-patella joints slightly shifted toward prolateral side; tibia large relative to femur, tibia-tarsus joints shifted toward retrolateral side; tarsus without macrotrichia; procursus ( Figs 786–788 ) straight, densely set with long hairs dorsally, few hairs slightly curved upwards; proximally on prolateral side with strong hump set with numerous long hairs and followed distally by thick sclerotized ridge, procursus tip with several distinctive elements: ventral sclerite with transparent retrolateral branch (arrow in Fig. 788 ), rounded dorsal sclerite, semitransparent prolateral pointed process, and pair of crescent-shaped retrolateral elements; genital bulb ( Figs 789–791 ) with simple basal sclerite connected to distal (main) sclerite, sperm duct opening not seen; distal sclerite with retrolateral ridge and distinctive set of one large and four smaller rounded prolateral apophyses.

LEGS. Femur 1 with single row of ~23 ventral spines; without curved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 4%; prolateral trichobothrium absent on tibia 1, present on other leg tibiae; tarsal pseudosegments indistinct except 2–3 distally.

Male (variation)

Tibia 1 in 43 males: 11.5–18.0 (mean 14.5). Dark ventral band on abdomen variably distinct, section behind gonopore with 2–4 longitudinal bands. Small apophyses prolaterally on bulbal sclerite variably distinct, usually one large plus four small apophyses, rarely one large plus three or five small apophyses. Retrolateral branch on ventral sclerite of procursus slightly variable in shape.

Female

In general similar to male ( Figs 704–705 ) but without spines on legs, with smaller and less distinct stridulatory files on chelicerae (sometimes not visible in dissecting microscope; distances between ridges: ~8.5 µm, Huber et al. 1999), and with stridulatory organ consisting of pair of weakly sclerotized but distinct processes posteriorly on carapace and pair of light brown plates anteriorly on abdomen ( Fig. 797 ). Tibia 1 in 57 females: 9.2–15.5 (mean 12.5). Epigynum as in Figs 796–799 , main epigynal plate short and wide, weakly protruding; posteriorly laterally strongly sclerotized, anteriorly weakly sclerotized, with variably distinct light brown median area; with pair of pockets very close to median line on both sides of median ridge (distance between pockets 30 µm), posteriorly barely protruding; internal sclerotized arc and median round structure visible in uncleared specimens; posterior plate short but wide. Internal genitalia ( Figs 794–795 , 800–802 ) with large elongate pore plates converging anteriorly, dorsal arc wide but simple, ventral arc medially slightly modified, with asymmetric sclerites (three females cleared), with simple median pouch.

Gynandromorphic specimen

Specimens that combine male and female morphological characters (sometimes in a left-right asymmetry) are extremely rare in pholcid spiders. The only previously known case was reported by Blackwall (1867), who described a specimen of C. lyoni from India that had a male left palp and a male left chelicera, but a female right palp, female right chelicera, and incompletely developed female genitalia (epigynum) on the right side. He noted that this was the only case of gynandromorphism among “many thousands” of specimens of spiders studied. Surprisingly, the only gynandromorphic pholcid specimen I have ever seen (among roughtly 50,000 specimens) is also a representative of C. lyoni . This specimen (from Chuuk Island, Micronesia), has a half epigynum ( Figs 804–805 ; relatively normally developed on the left side), but an enlarged right palp that reminds of a penultimate instar male palp (arrow in Fig. 803 ). The right chelicera and the right femur are not modified, as would be expected from a penultimate instar male. Surprisingly, however, the female stridulatory apparatus between prosoma and abdomen appears fully and symmetrically developed on both sides (arrows in Fig. 805 ). Leg length is also symmetric.

Natural history

Surprisingly little is known about the biology of this large and widely distributed synanthropic spider. Strickman et al. (1997) give basic bionomic data for spiders collected in Thailand and held at 30°C: eggs were laid 4–6 days after copulation; a mean of 34 spiderlings (range: 5–54) hatched form each eggsac 11–13 days after egg-laying; maturity was reached after only 74 days. Females were significantly heavier than males (mean weight 28.6 mg vs 17.6 mg). As soon as spiderlings started to eat (after the first molt), they were capable of overpowering a mosquito; prey was wrapped by ‘throwing’ silk with the hind legs, without rotating the prey. The prey was not bitten until the moment the spider started feeding, sometimes days after wrapping. No female was observed to eat her own eggs (as reported by Downes 1987). Observations on US populations give slightly to significantly higher numbers of eggs per egg sac (or spiderlings accompanying the female): 53–58 in Edwards (1993), and 72 in Guarisco & Cutler (2003). In Florida, Edwards (1993) observed a wide variety of insect prey in C. lyoni webs. For further observations, see Nandi & Raut (1985) (publication only partly seen) and Irie (2001) (in Japanese).

Distribution

The closest known relatives of C. lyoni ( C. maculipes , C. surobi sp. nov., C. sengleti sp. nov.) live in an area that is mainly composed of Pakistan, Afghanistan, and Iran ( Fig. 353 ), suggesting that C. lyoni originated from this region. It seems to have expanded from its original area at a time from which few records are available, so the early expansion is difficult to reconstruct. The earliest records from supposedly non-native regions date from the late 19 th century ( Myanmar; Thorell 1895) and the early 20 th century ( China; Strand 1907). Towards the east, C. lyoni may have reached the Philippines and Australia not before the 20 th century (oldest records: 1940s). Towards the west, C. lyoni seems to have reached South America much sooner than North America: while the oldest record from Argentina dates from 1914, and the species was “not rare” in Bahia ( Brazil) by the early 1930s ( Mello-Leitão 1935), it may not have reached the USA before the early 1980s ( Roth 1985, Edwards 1993). In Central Europe, the species appears to be slowly spreading since approximately 2000, but it is still largely confined to buildings ( Van Keer 2007, Bauer et al. 2016).

Mysteriously, this synanthropic species is largely absent from the original area of distribution of Crossopriza ( Fig. 351 ). Such a pattern is mirrored in some other synanthropic pholcids but remains unexplained (see Discussion).

Figures

Fig. 351. Known distribution of Crossopriza Simon, 1893. Orange: C. lyoni (Blackwall, 1867); blue: all other species (cf. Figs 352–353).

Figs 698–705. Crossopriza Simon, 1893, live specimens. 698–701. C. miskin sp. nov., from Oman, Ibri-Rusdaq; male in dorsal and lateral views; female with and without egg-sac. 702–705. C. lyoni (Blackwall, 1867), male from Singapore and courting/mating pair from Guinea.

Figs 783–785. Crossopriza lyoni (Blackwall, 1867); male from Guinea, Kindia (ZFMK Ar 10456, see Huber & Kwapong 2013); left palp, prolateral, dorsal, and retrolateral views. Scale bar = 1.0 mm.

Figs 786–795. Crossopriza lyoni (Blackwall, 1867); male from Guinea, Kindia (ZFMK Ar 10456, see Huber & Kwapong 2013), female from Japan, Yamamoto-chô (ZFMK Ar 5209). 786–788. Left procursus, prolateral, dorsal, and retrolateral views; arrow: branch of ventral sclerite. 789–791. Left genital bulb, prolateral, dorsal, and retrolateral views; arrow: retrolateral ridge.792–793. Male chelicerae, frontal and lateral views. 794–795. Cleared female genitalia, ventral and dorsal views. Abbreviation: vs = ventral sclerite. Scale bars = 0.5 mm.

Figs 796–802. Crossopriza lyoni (Blackwall, 1867); females from Japan, Yamamoto-chô (796–798, 801–802; ZFMK Ar 5209) and from Guinea, Kindia (799–800; ZFMKAr 10456, see Huber & Kwapong 2013). 796–797. Abdomen, ventral and frontal views; arrows: stridulatory plates. 798–799. Epigyna, ventral views. 800–802. Cleared female genitalia, dorsal view with dorsal arc tilted backwards, ventral view, and regular dorsal view; arrow: asymmetric median sclerite. Abbreviations: da = dorsal arc; e = epigynum; va = ventral arc. Scale bars: 796–797 = 1.0 mm; 798–802 = 0.5 mm.

Figs 803–805. Crossopriza lyoni (Blackwall, 1867); gynandromorphic specimen from Micronesia, Chuuk Island (MCZ 76633). 803. Prosoma, frontal view; arrow: penultimate instar male palp. 804– 805. Abdomen, ventral and frontal views; arrows: female stridulatory plates.Abbreviation: e = epigynum. Scale bars = 0.5 mm.

Fig. 353. Known distribution of Crossopriza Simon, 1893; autochthonous species part 2 (cf. Fig. 352): species with main cheliceral apophyses in frontal position and epigynal pockets close together. The figure is split into two to avoid too much overlap; the split is not meant to reflect any natural grouping of species. The single record of C. maculipes from China is not shown (see text).