Artema atlanta Walckenaer, 1837

Aharon, Shlomi, Huber, Bernhard A. & Gavish-Regev, Efrat, 2017, Daddy-long-leg giants: revision of the spider genus Artema Walckenaer, 1837 (Araneae, Pholcidae), European Journal of Taxonomy 376 (376), pp. 1-57 : 8-15

Artema atlanta Walckenaer, 1837

Figs 2–3 , 15–34 , 201 , 208

Artema atlanta Walckenaer, 1837: 656–657 (♂ ♀, Brazil).

Artema mauriciana Walckenaer, 1837: 657–658 , pl. 15, fig. 1 (♂ ♀, Mauritius). Synonymized by Pickard-Cambridge 1902: 366.

Pholcus sisyphoides Doleschall, 1857: 408–409 ( Indonesia) . Synonymized by Pickard-Cambridge 1902: 366.

Artema convexa Blackwall, 1858: 332–334 (♀, Brazil). Synonymy with A. sisyphoides suggested by Thorell 1881: 179; synonymized with “ A. mauricia ” by Simon 1885: 19.

Pholcus borbonicus Vinson, 1863: 132–135 , 307, pl. 3, figs 4, 4a (♂ ♀, Réunion, Mauritius). Transferred and synonymized (with “ A. mauricia ”) by Simon 1885: 19.

Pholcus rotundatus Karsch, 1879: 106–107 (♀, Colombia). Transferred and synonymized (with “ A. mauricia ”) by Simon 1893: 465.

Crossopriza sexsignata Franganillo, 1926a: 49 ( Cuba) . Synonymized by Pérez González 1996: 431– 432.

Coroia magna González-Sponga, 2005: 102 , pl. 2, figs 1–11. Synonymized by Huber et al. 2014: 417.

Artema atlanta – Simon 1894: 519 ( A. atalanta – lapsus) ( St. Vincent). — Taczanowski 1874: 103 (French Guiana). — Banks 1898: 212 ( A. atlantica – lapsus) ( Mexico). — Moenkhaus 1898: 86 ( Brazil). — Pocock 1900: 238–239, fig. 81 ( India, Myanmar, Sri Lanka, Pakistan). — Pickard-Cambridge 1902: 366, pl. 34, figs 15, 15a–b, 16, 16a ( Mexico, Brazil, India, Pakistan). — Petrunkevitch 1911: 156; 1925: 66 ( Panama); 1929: 119–120, figs 102–104 ( Puerto Rico). — Mello-Leitão 1918: 89–91 ( Brazil); 1941: 244 ( Colombia); 1942: 382 ( Argentina); 1946: 33 ( Paraguay). — Sherriffs 1919: 228 ( India). — Chamberlin 1924: 632 ( Mexico). — Gertsch 1935: 10 ( USA). — Dyal 1935: 170– 171, pl. 15, figs 97–103 ( India). — Bryant 1940: 289 ( Cuba); 1948: 366 ( Dominican Republic). — Gertsch & Davis 1942: 7 ( Mexico). — Caporiacco 1948: 626–627 ( Guyana); 1949: 326 ( Kenya). — Chrysanthus 1967: 92–96, figs 15–19 ( Curaçao, New Guinea, Tanzania, “Sandwich Isl.” = Hawaii?). — Yaginuma 1970: 646 ( Japan). — Tikader 1977: 164–165 ( India). — Brignoli 1981: 92, figs 1–7 ( USA, Mexico, Brazil, Egypt). — Tikader & Biswas 1981: 18, fig. 12. — Majumder & Biswas 1993: 1 ( India). — Pérez González 1996: 431–432 ( Cuba). — Saaristo 1999: 2, 8 ( Seychelles); 2001: 15 –17, figs 16–22 ( Seychelles, India, Indonesia, Sri Lanka, Vietnam); 2010: 159, figs 25.1– 5. — Huber 2000: 342, figs 12–13, 48, 56–57, 99, 121, 145, 169, 195 (SEM); 2001: 135 –136 ( Australia). — Murphy & Murphy 2000: 246–247. — van Keer & van Keer 2001: 82 ( Belgium). — Lee 2005: 7 ( Great Britain, in container). — Colmenares-García 2008: 87, figs 1a–c ( Venezuela). — Beatty et al. 2008: 3–7, figs 15–16, 42–43 ( Micronesia, Polynesia, Indonesia). — Irie 2009: 106, figs (2-2-13) 1–2, pl. 4, fig. 1. — Gao & Li 2010: 11–13, figs 1–3 ( China). — Huber & Warui 2012: 3 , fig. 1 ( Kenya, Uganda, Somalia, Tanzania). — Huber & Kwapong 2013: 7 ( Guinea, Togo, Benin).

Artema mauriciana – Vinson 1863: 141–142, 307–308 ( A. mauricia – lapsus) ( Mauritius). — Simon 1885: 19 ( A. mauricia – lapsus) ( India); 1887: 453 ( A. mauricia – lapsus) ( Djibouti); 1893: 463–465 ( A. mauricia – lapsus); figs 441, 451–454; 1908: 426–427 ( Libya). — Kulczyński 1901: pl. 1, fig. 15 ( A. mauricia – lapsus). — Leardy Airaghi 1902: 348 ( A. mauricia - lapsus) ( India). — Tullgren 1910: 119, pl. 2, fig. 35 ( Tanzania). — Caporiacco 1928: 91 ( A. mauricia - lapsus) ( Libya); 1936: 86 ( Libya). — Lessert 1936: 233–234 ( Mozambique). — Millot 1941: fig. 1F only ( Guinea); 1946: 129, fig. 1 ( Madagascar). — Denis 1953: 320 ( Algeria). — Marples 1955: 465 ( Samoa).

Pholcus sisyphoides – Thorell 1881: 179–180 (“ins Elephanta”, Hawaii).

Artema sisyphoides – Karsch 1892: 276 ( Sri Lanka). — Thorell 1895: 69–70 ( Myanmar); 1898: 274 ( Myanmar).

Artema convexa – Blackwall 1866: 459–460 (“Equatorial Africa”); 1867: 394 ( India).

Pholcus borbonicus – Simon 1873: 47–48 ( Egypt). — Lenz 1886: 395 ( Madagascar); 1891: 173 ( Madagascar). — Bösenberg & Lenz 1895: 40 ( Mozambique).

Crossopriza sexsignata – Franganillo 1926b: 11; 1936a: 46; 1936b: 77.

Misidentifications

“ Artema atlanta ” – Dalmas 1920 (see A. nephilit sp. nov.). — Feng 1990: 45 (misidentification of Physocyclus globosus ).

“ Artema mauriciana ” – Bodenheimer 1937: 238 (see A. nephilit sp. nov.). — Millot 1941: 3–5, figs 1A–I (except fig. 1F; misidentification of A. bunkpurugu ; see Huber & Kwapong 2013).

“ Pholcus borbonicus ” – L. Koch 1875: 25–26. — Simon 1882: 234 (see A. kochi ).

“ Tibiosa coreana ” – González-Sponga 2006: pl. 3, figs 8–9 only (see Huber 2009).

Diagnosis

Males are easily distinguished from all known congeners by their bulbal processes: process c ( Fig. 20 ) with sclerotized ridge projecting prolaterally perpendicular to process b with small teeth prolaterally on round end of process ( Figs 27, 29 ); process d rounded ( Figs 19 , 27 ) rather than pointed as in A. doriae and A. transcaspica ; process e present and rounded. Males differ also by two deep dorsal notches on distal margin of procursus, with lighter cuticle than rest of procursus (arrows in Fig. 20 ). Females differ from other congeners by strongly indented posterior epigynal margin ( Fig. 15 ).

Types

BRAZIL: Artema atlanta Walckenaer, 1837 : syntypes ♂, ♀ probably lost (no further data).

MAURITIUS: Artema mauriciana Walckenaer, 1837 : syntypes ♂, ♀ probably lost [20.2° S, 57.5° E] (no further data).

INDONESIA: Pholcus sisyphoides Doleschall, 1857 : holotype ♂ probably lost; Amboina [3.6° S, 128.1° E] .

BRAZIL: Artema convexa Blackwall, 1858 : syntypes ♀♀ probably lost; Pernambuco [8.0° S, 35.0° W].

REUNION: Pholcus borbonicus Vinson, 1863 : syntypes ♂, ♀ probably lost [21.12° S, 55.52° E] (no further data).

COLOMBIA: Pholcus rotundatus Karsch, 1879 : syntype (s?) ♀ probably lost; Santa Marta [11.24° N, 74.20° W].

CUBA: Crossopriza sexsignata Franganillo, 1926 : syntypes ♂, ♀ probably lost; Habana, Luyanó , [23.10° N, 82.34° W].

VENEZUELA: Coroia magna González-Sponga, 2005 : holotype ♂ and 1 ♀ paratype; Falcón State, 10 km SE from Coro (11.20° N, 69.50° W), Mar. and Dec. 1999, E. Bravo and V. Wall leg., in Museo del Instituto de Zoología Agrícola of the Universidad Central de Venezuela 1440 a,b, not examined.

Material examined (arranged from west to east)

USA: 1 ♂, 1 ♀, Arizona, Tucson [~ 32.2° N, 110.9° W], Jul.–Aug. 1935, W.P. Steckler leg. ( AMNH); 3 ♀♀, same data (AMNH); 3 ♂♂, 4 ♀♀, same locality, Jul. 1937, collector not given ( NHMB).

HONDURAS: 1 juv., Amapala [13.28° N, 87.64° W], 1906, R. Paessler leg. ( ZMH).

HAITI: 1 ♀, Port au Prince [18.54° N, 72.33° W], 29 May 1901, G. Keitel Jr. leg. (ZMH); 1 ♀, Saint Marc [19.10° N, 72.70° W], 1905, C. Gazgo leg. ( ZMH).

VIRGIN ISLANDS: 1 juv., St. Thomas [18.33° N, 64.91° W], 28 Mar. 1898, L.D. Calwood leg. (ZMH).

PARAGUAY: 2 ♀♀, date and locality not given [~ 23.5° S, 58.1° W], Wiengreen leg. ( ZMH).

ALGERIA: 1 ♀, Djanet [24.55° N, 9.48° E], 25 Nov. 1989, Kechemir leg. (CRB).

TUNISIA: 7 ♂♂, 2 ♀♀, numerous juvs, Djerba [33.80° N, 10.88° E], date not given, Vibert leg. ( MNHN AR 10170).

LIBYA: 1 ♂, “ Gialo ” [= Jalu , 29.02° N, 21.54° E], Sep. 1934, Zavattari leg. (MZUF); 1 ♀, same locality, Jan. 1934, Di Caporiacco leg. ( MZUF); 1 ♂, same locality, 4–5 May 1931 (?), collector not given ( MSNG); 1 ♂, same locality (“Oajo di Gialo ”), Jul. 1931 (?), collector not given ( MSNG); 1 ♀, same locality (“ Gialo ”), Apr. 1931 (?), collector not given ( MSNG); 1 ♂, 1 ♀, same locality, 30 Apr. 1931, collector not given ( MSNG); 6 ♀♀, 6 juvs, “ Giarabub ” [= Al Jaghbub , 29.74° N, 24.51° E], Jun. 1926 – Mar. 1927, Confalonieri leg. ( MSNG); 1 ♀, with two locality labels: “ Giarabub ” [= Al Jaghbub , 29.74° N, 24.51° E], Jun. 1926 – Mar. 1927 and “ Porto Bardia ” [= Bardiyah , 31.75° N, 25.08° E], Mar. 1927 ( MZUF); 1 ♀, Tripoli [32.86° N, 13.17° E], 1906, Klaptocz leg. ( ZMH).

EGYPT: 1 ♂, 1 ♀, 1 juv., Sohag, Maragha [26.69° N, 31.60° E] 26 Feb. 2002, H. El-Hennawy leg. (ZFMK Ar 15215); 1 ♂, 7 ♀♀, numerous juvs, Alexandria [31.19° N, 29.91° E], and Suez [29.96° N, 32.54° E], date not given, E. Simon leg. ( MNHN AR 10155, 392); 2 ♀♀, near Cairo [~ 30.0° N, 31.2° E], 1911, E. Graeter leg. ( NHMB 1041a, 1402b); 3 ♂♂, Mansurah [31.04° N, 31.37° E], date not given, I. Sörensen leg. ( ZMUC).

CAPE VERDE: 1 ♀, Fogo, São Filipe [14.89° N, 24.49° W], in bathroom, on web, 9 Nov. 1998, W. Tavernier leg. ( MRAC 208404 ).

GUINEA-BISSAU (“ Portuguese Guinea ”): 1 ♂, 2 ♀♀, 1 juv., Bissao [11.86° N, 15.60° W], 4 Oct. 1898, H. Eberhardt leg. (ZMH); 1 juv., same data (ZMH).

GUINEA and BENIN: see Huber & Kwapong (2013).

TOGO: 1 ♂, 1 ♀, 2 juvs, “ Anecho ” [6.22° N, 1.57° E], 27 Jan. 1910, Günther leg. (ZMH); see also Huber & Kwapong (2013).

SÃO TOMÉ AND PRÍNCIPE: 1 ♀, “S. Thomé” [= São Tomé, 0.34° N, 6.73° E], 1879, R. Greef leg. ( ZMH no. 31).

CHAD: 3 ♂♂, 2 ♀♀, numerous juvs, between Bongor and Fort Lamy (“ Lancy ?”) [= N’Djamena , ~ 11.1° N, 15.2° E], Nov.–Dec. 1965, Y. Brandily leg. ( MRAC 132893 ).

SUDAN: 1 ♂, Khartoum [5.54° N, 32.53° E], 1960–1961, J.S. Cloudsley-Thompson leg. ( MRAC 120764 ); 1 ♂, same data ( MRAC 120766 ); 1 ♂, 1 ♀, 2 juvs, (South) Kordofan , Talodi [10.63° N, 30.38° E], 15 Oct. 1938, Werner leg. ( NHMW 356 ); 1 ♂, 2 ♀♀, 7 juvs, El-Obeid [13.18° N, 30.22° E], Mar. 1914, D.G. Mainhof leg. ( ZMH).

ERITREA: 1 ♂, 2 ♀♀, 1 juv., Massawa, 1870, Beccari leg. ( MSNG).

YEMEN-ERITREA: 1 ♂, 2 ♀♀, 1 juv., Aden [12.86° N, 44.98° E] and Massawa [15.61° N, 39.45° E], 1889, E. Simon leg. (“393” part) ( MNHN AR 10169 part).

SOMALIA: 1 ♀, 1 juv., locality unknown (“339.76”, “Mag. no. 866”), 1924, Stefanini and Puccioni leg. ( MZUF); 2 ♀♀, locality unknown (“1140 20”), 1924, Stefanini and Puccioni leg. ( MZUF); 1 ♀, Kismayo [0.35° S, 42.54° E], under stones, 16 Apr. 1993, Vercammen Gino leg. ( MRAC 177.415).

YEMEN: 1 ♀, 5 juvs, Aden [12.86° N, 44.98° E], 1880, Doria and Beccari leg. ( MSNG).

DR CONGO: 3 ♀♀, numerous juvs, Kivu, Uvira between Kolundu and Kavimvira [3.34° S, 29.15° E], Jun. 1961, R. Kiss leg. ( MRAC 119889 ); 2 juvs, same data ( MRAC 119917 ); 1 ♀, Mutelemuko, Uvira [~ 3.40° S, 29.12° E], 1954, J. Bouillon leg. ( MRAC 76923 ).

BURUNDI: 1 juv., “Usumbura” (= Bujumbura) (3.36° S, 29.36° E), Jan. 1926, H. Schouteden leg. ( MRAC 29993 ); 2 ♂♂, 1 ♀, same locality, 1948, A. Lestrade leg. ( MRAC 57974 -77); 1 ♀, Rumonge [3.97° S, 29.43° E], 1934, A. Lestrade leg. ( MRAC 24696 ).

UGANDA: 1 ♀, 1 juv., Jinja [0.43° N, 33.20° E], Apr. 1968, E. Vertriest leg. ( MRAC 134815 ); see also Huber & Warui (2012).

TANZANIA: 2 ♀♀, Dar es Salaam, UDSM campus [6.77° S, 39.20° E], 1970–1971, K.M. Howell leg. ( MRAC 159428 ); 1 ♀, “near Msala, Tanganyika Terr .” [~ 3.16° S, 30.98° E], date not given (“ Kuipper 64”) ( SMF); see also Huber & Warui (2012).

KENYA: 1 ♀, Rift Valley, Lodwar , along Turkwell River (3.11° N, 35.59° E), 10–12 Jun. 1999, W.J. Pulawski and J.S. Schweikert leg. (CAS); 1 ♂, Gabraland, Marsabit area [2.33° N, 37.98° E], 17 Jan. 1978 (from artifact in CAS anthropology collection) ( CAS); 1 ♂, “ Ostafrika Waboniland ” [2.07° S, 40.70° E], 15 Oct. 1938, Haessler leg. ( NHMW 355 ); see also Huber & Warui (2012).

ZAMBIA: 1 ♂, Mpulungu [8.76° S, 31.11° E], no date given (“ Kuipper 281”) ( SMF).

ZIMBABWE: 1 ♂, Victoria Falls (17.933° S, 25.833° E), 29 Nov. 1996, W. Pulawski and V. Ahrens leg. (CAS).

MOZAMBIQUE: 1 ♂, numerous juvs, “ Cucuta/ Brasilien ” [ Cucuta , 11.83° S, 40.24° E], 29 Sep. 1887, W. Bösenberg leg. (ZMH).

SOUTH AFRICA: 1 ♂, KwaZulu Natal, Durban (29.850° S, 30.016° E), 10 m a.s.l., 8 Jan. 2001, C. Whitmore leg. (DNSM); 3 juvs, Limpopo, Lajuma Mountain Retreat (23.037° S, 29.441° E), sifting leaf litter, 1325 m a.s.l., 10 Nov. 2012, J.A. Neethling leg. ( ZFMK Ar 15216).

COMOROS: 1 ♂, Grande Comore, Moroni [11.69° S, 43.25° E], inside house, 7 Jan. 2003, U. Dall ‘ Asta leg. ( MRAC 215035 ).

MADAGASCAR: 1 ♀, Tamatave [18.15° S, 49.40° E], 12 May 1885, A. O’Swald leg. ( ZMH 7791 ); 1 ♀, Nosy Be, Lokobe [13.30° S, 48.26° E], 5 Nov. 1885, A. O’Swald leg. (ZMH 12832); 1 ♀, Nosy Be, 13 Mar. 1908, collector not given ( ZMH).

INDIA: 3 ♀♀, 2 juvs, Bombay, “ Elephanta ” [= Gharapuri , 18.96° N, 72.93° E], 18 Sep. 1877, Beccari and E. D’Albertis (MSNG); 2 ♂♂, 9 ♀♀, Vellore [12.93° N, 79.13° E], date not given, Löwenthal leg. ( ZMUC); 1 ♀, 1 juv., Tuticorin [8.76° N, 78.13° E], collected in container arrived at Haifa port [32.82° N, 34.99° E], 5 Jun. 2012, E. Gavish-Regev leg. ( SMNH AR 50186); 1 ♂, Kerala, Malabar, Feroke [11.18° N, 75.85° E], date and collector not given ( SMF).

MYANMAR: 3 ♂♂, 14 ♀♀, “fra Rangoohn” [Rangoon, 16.84° N, 96.14° E], date not given, Lövendal leg. ( ZMUC).

THAILAND: 1 ♂, 2 ♀♀, Ratchaburi, Wat Huai Takaeng (13.587° N, 99.758° E), 30 m a.s.l., in cave, 15 Mar. 2015, B.A. Huber and B. Petcharad leg. (ZFMK Ar 15217); 1 juv., in pure ethanol, same data ( ZFMK Mal 374).

CHINA: 1 ♀, Canton [22.67° N, 113.60° E], 8 Aug. 1904, W. Helms leg. (ZMH); 1 ♂, “ Swatow ” [= Shantou , 23.33° N, 116.68° E], date and collector not given ( MNHN AR 10166).

TAIWAN: 1 ♀, 2 juvs, “Takao, Formosa” [= Kaohsiung, 22.77° N, 120.42° E], 1 Nov. 1908, W. Schwinghammer leg. (ZMH).

PHILIPPINES: 1 ♂, Manila [14.61° N, 121.04° E], 10 Jun. 1945, T. Aarons leg. (CAS).

INDONESIA: 1 ♂, Flores, Labuan Bajo (8.49° S, 119.87° E), 22 Mar. 2009, S. Huber leg. (SMF); 2 ♂♂, 4 ♀♀, 4 juvs, Sumatra, Medan [3.61° N, 98.67° E], 15 Oct. 1938, Fulmek leg. ( NHMW).

AUSTRALIA: Numerous specimens from Western Australia , Northern Territory , and Queensland were examined but not listed in detail by Huber (2001). We have not reexamined those specimens.

Locations not clear: 2 ♀♀, “El Bur” ( Somalia?), “Mag. no. 1220”, 15Aug. 1968 (“ SBS ”, “977”) ( MZUF); 1 ♂, 1 juv., “ Museat, Jayakan , 22” ( NHMUK); 1 ♀, 1 juv., “ Westindien ”, 3 Sep. 1906, C. Gazgo leg. ( ZMH).

Description

Male ( Myanmar: Rangoon, ZMUC)

MEASUREMENTS. Total body length 9.6, carapace width 4.0. Leg 1: 48.4 (16.0 + 1.6 + 16.6 + 22.5 + 2.6), tibia 2: 12.6, tibia 3: 9.5, tibia 4: 13.0; tibia 1 L/d: 42. Distance PME–PME 210 μm, diameter PME 210 μm, distance PME–ALE 110 μm, distance AME–AME 60 μm, diameter AME 180 μm.

COLOR. Carapace ochre beige, with light brown median band and small dots laterally; ocular area light brown; clypeus with light brown band fading towards edge, with brown rim (as in Fig. 3 ); legs light ochre, with dark rings on femora subdistally, patellae + tibiae proximally, and tibiae subdistally; sternum light ochre with two small light brown marks posteriorly, with narrow light brown margins; abdomen pale ochre with pale and dark dots forming large marks dorsally and stripes laterally.

BODY. Ocular area slightly elevated; carapace with median pit and distinct posterior furrow; clypeus unmodified; sternum wider than long (2.5/2.2); chelicerae as in Figs 22–24 , with frontal row of ~20 modified (cone-shaped; cf. Huber 2000: figs 12–13) hairs on each side in s-shaped pattern (as in Fig. 24 ); stridulatory ridges absent (as in Fig. 31 ); abdomen globose and high; gonopore with four epiandrous spigots (cf. Huber 2000: fig. 121); ALS with several cylindrical spigots in addition to widened and pointed spigot (as in Figs 26, 28 ) (cf. Huber 2000: fig. 169); PMS with two pointed spigots each (as in Fig. 25 ) (cf. Huber 2000: fig. 195).

PALPS. As in Figs 16–18 ; coxa unmodified; trochanter with short ventral projection; femur with short, somewhat pointed retrolateral process and small dorsal projection proximally; ventral membranous area proximally bordered on both sides by sclerotized ridges; femur-patella hinges close together dorsally; patella very short; procursus with proximal dorsal process and weakly developed ventral pocket; with two distal dorsal notches on sclerotized margin (arrows in Fig. 20 ); bulb with membranous embolus arising from base of processes a and b (as in Fig. 27 ; cf. Huber 2000: fig. 56), process a long, somewhat inflated; process b elongated and pointed, process c (as in Figs 20 , 27 ) with sclerotized ridge projecting prolaterally perpendicular to process b, with small teeth prolaterally on round end of process (as in Fig. 29 ) (cf Huber 2000: fig. 57), processes d and e low and rounded (as in Figs 19 , 27 ).

LEGS. Without spines, with long curved hairs, especially on tibiae and metatarsi; retrolateral trichobothrium on tibia 1 at 9%; prolateral trichobothrium present on all tibiae; pseudosegmentation not visible (cf. Huber 2000: fig. 99). Comb-hairs on tarsi 4 variant of complex ‘ Belisana -type’ (cf. Huber & Fleckenstein 2008: figs 17–18).

Male (variation)

Tibia 1 in 19 other males: 12.0–19.0 (mean 15.4); color pattern on abdomen varies from distinct stripes consisting of dark and pale dots on pale to brown-ochre background, to pale without any pattern; leg color varies from pale to ochre, prolateral ochre mark on patella 1 proximally sometimes absent; ocular area light brown to ochre; lateral dots on carapace vary from round radiating pattern to small indistinct marks; cheliceral modified hairs vary in numbers (~17–20), sometimes sclerotized ridge very dark and pattern of modified hairs hard to distinguish, one male with small process medial of sclerotized ridge (from Khartoum, Sudan; MRAC 120766); brown marks posteriorly on sternum variable; one male with procursus notches almost fused to one large notch without sclerotized line between them (from Shantou, China; MNHN AR 10166). No variation observed in numbers of epiandrous spigots.

Female

In general similar to male; tibia 1 in 20 females: 9.2–17.2 (mean 13.2); stridulatory files laterally on chelicerae present ( Fig. 30 ); epigynal plate trapezoidal (posteriorly wider), posterior margin with deep median indentation ( Figs 15 , 33 ); epigynal plate consisting of two sclerotized lateral areas that appear swollen posteriorly, usually light brown, sometimes very pale at lateral posterior edge, with light brown to pale median area, and dark sclerite anteriorly (variably large and variably distinct), posterior rim sometimes slightly sclerotized; pair of projections anterior to epigynum (AEP) ovoid, sometimes with variable light brown marks anteriorly.

Natural history

Like other species in the genus, A. atlanta inhabits caves as well as buildings and basements. Similar to several other pholcid genera, such as Crossopriza Simon, 1893 , Micropholcus Deeleman-Reinhold & Prinsen, 1987 , Modisimus Simon, 1893 , Pholcus Walckenaer, 1805 , Physocyclus Simon, 1893 and Smeringopus Simon, 1890 , it is not clear why this is the only species in the genus that has spread all over the world. Another surprising fact is that, to the best of our knowledge, no study on the biology of this widespread species has ever been conducted.

Distribution

Artema atlanta is the only species in this small genus with a worldwide distribution ( Fig. 2 ). It is widespread from the subtropical Americas through Africa and India, the Far East and Australia. It can be assumed from the genus distribution that A. atlanta was originally an Old World species, like the other species in the genus, which expanded its distribution worldwide ( Brignoli 1981). Intriguingly, A. atlanta is almost entirely absent from the large area where congeneric species occur (Arabian Peninsula, Middle East, Central Asia).

It seems that in several parts of the world, A. atlanta was more common a century ago than it is now. For example, Mello-Leitão (1918) reported the species to be very common along the Brazilian coast (“… muito commun em todo o littoral do Brazil …”). Nowadays, this is definitely not the case (not a single encounter during seven collecting trips along the coast ranging from the Brazilian states of Rio Grande do Sul to Rio Grande do Norte; B.A. Huber, unpublished data). Likewise, the species was reported frequently from the Caribbean until the mid 20 th century ( Simon 1894; Franganillo 1926a, 1926 b, 1936a, 1936b; Petrunkevitch 1929; Bryant 1940, 1948) but not thereafter. Intensive collecting in Cuba, Hispaniola, Jamaica, Puerto Rico, and the Lesser Antilles between 1999 and 2012 has not produced a single specimen (B.A. Huber and I. Agnarsson, unpublished data).

Figures

Figs 1–2. 1. Known distributions of species of Artema except A. atlanta Walckenaer, 1837. The Japanese record of A. doriae Thorell, 1881 is not shown as it is based on a single introduced specimen. 2. Known distribution of Artema atlanta. Green dots: specimens examined; black dots: specimens not examined (data from: Banks 1898; Beatty et al. 2008; Blackwell 1958; Bösenberg & Lenz 1895; Brignoli 1981; Bryant 1940, 1948; Caporiacco 1928, 1948, 1949; Chamberlin 1924; Chrysanthus 1967; Colmenares 2008; Denis 1953; Doleschall 1857; Dyal 1935; Franganillo 1926a; Gao & Li 2010; Gertsch & Davis 1942; Gonzalez-Sponga 2005; Karsch 1879, 1892; Leardi in Airaghi 1902; Lessert 1936; Majumder & Biswas 1993; Marples 1955; Mello-Leitão 1942; Millot 1946; Moenkhaus 1898; F.O.Pickard-Cambridge 1902; Petrunkevitch 1925, 1929; Pocock 1900; Saaristo 2001; Simon 1885, 1894; Taczanowsky 1874; Thorell 1895, 1898; Vinson 1863; Yaginuma 1970). Records from Israel, Belgium, and Britain are not shown as they are based on single introduced specimens.

Figs 3–8. Live specimens and habitats. 3. Artema atlanta Walckenaer, 1837 from Thailand, Ratchaburi. 4–5. A. nephilit sp. nov. from Israel. 6. Typical Artema web mass, in a cave in Petra, Jordan. 7–8. Caves populated by Artema nephilit sp. nov.: Oren Cave, Mount Karmel (7) and caves in the Eilat Mountains (8), Israel. Photos: BAH (3–4, 6–8), SA (5).

Figs 15–24. Artema atlanta. 15. Epigynum (MRAC 215041, Benin). 16–18. Male left pedipalp (MRAC 120766, Sudan), prolateral, dorsal, and retrolateral views. 19–21. Male left bulb and procursus, prolateral, dorsal, and retrolateral views, arrows in Fig. 20 show notches on distal margin of procursus. 22–24. Male chelicerae (same specimen), lateral and frontal views. Scale lines: 15, 19–24 = 0.5 mm; 16–18 = 1 mm.

Figs 25–29. Artema atlanta Walckenaer, 1837, ZFMK Gui 82 and ZFMK Gui 111. 25–26, 28. Male ALS and PMS (arrows: one of the six small cylindrically-shaped spigots wider than others). 27. Male left bulb, prolateral-distal view. 29. Small teeth prolaterally on round end of process c. Scale lines: 25 = 0.06 mm; 26, 29 = 0.03 mm; 27 = 0.2 mm; 28 = 0.04 mm.

Figs 30–34. Artema atlanta Walckenaer, 1837, ZFMK Ar 15217 (♀) and ZFMK Gui 82 (♂). 30. Female stridulatory ridges. 31. Absence of male stridulatory ridges. 32. Male chelicerae with frontal row of modified (cone-shaped) hairs. 33. Epigynal plate. 34. Modified (cone-shaped) hair on chelicerae process. Scale lines: 30–31 = 0.1 mm; 32 = 0.2 mm; 33 = 0.6 mm; 34 = 0.01 mm.

Figs 201–207. Cleared female genitalia, dorsal views. 201. Artema atlanta Walckenaer, 1837 (AMNH, USA, Arizona, Tucson). 202. A. bunkpurugu Huber & Kwapong, 2013 (ZFMK Ar 10516, Ghana, Bunkpurugu-Nakpanduri). 203. A. nephilit sp. nov. (HUJ Ara 16020, Israel, Samar). 204. A. kochi Kulczyński, 1901 (MNHN AR 10169 part). 205. A. doriae Thorell, 1881 (ZFMK Ar 15234, Iran, Tehran Prov., Tochal). 206. A. transcaspica Spassky, 1934 (ZIN type vials). 207. Artema sp. c. (ZMMU, Pakistan, Baluchistan, Muslim Bagh). Scale lines: 0.5 mm.

Figs 208–214. Cleared female genitalia, posterior dorsal views. 208. A. atlanta Walckenaer, 1837 (AMNH, USA, Arizona, Tucson). 209. A. bunkpurugu Huber & Kwapong, 2013 (ZFMK Ar 10516, Ghana, Bunkpurugu-Nakpanduri). 210. A. nephilit sp. nov. (HUJAra 16020, Israel, Samar). 211. A. kochi Kulczyński, 1901 (MNHN AR 10169 part). 212. A. doriae Thorell, 1881 (ZFMK Ar 15234, Iran, Tehran Prov., Tochal). 213. A. transcaspica Spassky, 1934 (ZIN type vials). 214. Artema sp. c (ZMMU, Pakistan, Baluchistan, Muslim Bagh). Scale lines: 0.5 mm.