Artema doriae Thorell, 1881

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 : 30-35

Artema doriae Thorell, 1881

Figs 1 , 90–120 , 205 , 212

Pholcus doriae Thorell, 1881: 179–180 (in footnote) (♀, Teheran, Iran). Synonymized with Artema atlanta by Pickard-Cambridge 1902: 366; revalidated by Brignoli 1981: 92.

Artema doriai (unjustified emendation; see Etymology below) – Simon 1893: 465. — Tabrizi et al. 2014: 35, figs 1M–N, 3L–M. — Malek Hosseini et al. 2015: 92.

Misidentifications

“ Artema transcaspica ” – Denis 1958: 112 ( Afghanistan). — Roewer 1955: 752 ( Iran). — Gharari & Marusik 2009: 4 ( Iran). See Note below.

Etymology

Even though the species was named for a man (Marchese Giacomo Doria, 1840–1913), the ICZN (1999: article 31.1) clearly states that the correct patronym has to be doriae, not doriai. The latter is thus an unjustified emendation.

Diagnosis

Artema doriae is easily distinguished from other congeners (except A. transcaspica and Artema sp. c.) by the arrangement of modified hairs situated frontally on cheliceral processes ( Fig. 98 ), by the triangular bulbal process c, usually curved and pointing towards prolateral ( Fig. 95 ), and by the distinct ventral process d ( Fig. 94 ). Males are possibly distinguishable from A. transcaspica and Artema sp. c by the cheliceral proximal incline usually with small ridge or process above modified hairs in lateral view (arrows in Figs 100–102 ) ( A. transcaspica usually with smooth incline proximally of modified hairs, see arrows in Figs 131–133 ; Artema sp. c with robust ridge or process above modified hairs, see arrows in Figs 190–191 ). Females differ more easily from other congeners (except A. transcaspica and Artema sp. c) by their quadrilateral epigynal shape. Females appear distinguishable from A. transcaspica by their more trapezoidal epigynal plate rather than rectangular to square-shaped, by median sclerite not reaching posterior margin, and by more distinct and inflated pale median area posteriorly (compare Figs 103–114 with Figs 134–145 ); Artema sp. c differs by wider epigynal plate, and by more inflated and prominent pale median area of epigynal plate ( Figs 195–200 ).

Material examined

Syntypes

IRAN: 1 ♂, 1 ♀, Tehran [35.70° N, 51.42° E], 1862, G. Doria leg. ( MSNG); 1 ♂, 1 ♀, same data ( MNHN AR 10180) (E. Simon collection number 397).

Other material

ISRAEL: 1 ♀, Upper Galilee, Ramot Naftali (33.08° N, 35.54° E), 15 May 2012, S. Zonstein leg. ( CSZ); 2 ♂♂, Yavni’el [32.70° N, 35.50° E], Sep. 1961, collector not given ( HUJ Ara 16030-31); 1 ♂, Poriyya slope [32.73° N, 35.54° E], Sep. 1970, Pener et al. leg. ( HUJ Ara 16032); 1 ♀, Nachal Chagal, N of Gesher (32.634° N, 35.534° E), 3 Dec. 2010, L. Friedman and C. Drees leg. ( CSZ); 1 ♀, Memorial Ha-Biq’a, NE Peza’el (32.052° N, 35.458° E), in cave, 10 Mar. 2014, S. Aharon and E. Gavish-Regev leg. ( HUJ Ara 16033); 2 ♀♀, 1 juv., Memorial Ha-Biq’a, NE Peza’el (32.0524° N, 35.4589° E), - 245 m b.s.l., at base of stonewall, 15 Sep. 2013, B.A. Huber, S. Aharon and E. Gavish-Regev ( ZFMK Ar 15243-44); 1 ♀, 2 juvs, in pure ethanol, same data ( ZFMK Isr 40); 1 ♀, Jericho [31.89° N, 35.42° E], 25 Feb. 1968, G. Tsabar leg. ( HUJ Ara 16034); 1 ♀, same locality, 12 Dec. 1967, P. Amitai leg. ( HUJ Ara 16035); 1 juv., same locality, 20 Oct. 1969, P. Amitai ( HUJ Ara 16036); 6 ♀♀, Modi’in (31.893° N, 34.971° E), pitfall trap in shrub habitat, Apr. 2012, I. Berenstein et al. leg. ( SMNH); 1 ♂, Hazeva [30.76° N, 35.27° E], 17 Jun. 2010, L. Friedman and C. Drees ( CSZ); 1 ♀, near Hazeva [30.76° N, 35.27° E], 11 Aug. 2014, Akiva Topper leg. ( HUJ Ara 16037).

TURKEY: 2 ♀♀, Şanlıurfa Prov., Urfa , 15 km Urfa - Gaziantep [37.10° N, 38.64° E], webs under rocks on dry rocky hills, 12 Sep. 1956, collector not given ( XUM); 1 ♀, 1 juv., Diyarbakır Prov., Çermik [38.14° N, 39.45° E], 700 m a.s.l., 27 Jul. 2004, T. Danisman leg. ( ZFMK Ar 15931) .

IRAN: 1 ♂, Tehran, 15 Oct. 1938, collector not given ( NHMW 354 ); 1 ♂, 1 ♀, Tehran Prov., Tochal (35.827° N, 51.406° E), 6 Jun. 2013, A. Zamani leg. ( ZFMK Ar 15233–34); 1 ♂, Alborz Prov., Karaj (35.84° N, 50.95° E), date not given, Ressl leg. ( ZFMK Ar 15930); 2 ♂♂, 2 ♀♀, 1 juv., Lorestan Prov., Ma’amulan (33.33° N, 47.90° E), 6 Aug. 1973, A. Senglet ( MHNG); 2 ♂♂, Esfahan Prov., Esfahan (32.56° N, 51.51° E), 23Aug. 1973, A. Senglet leg. ( MHNG); 1 ♀, Esfahan, 18–20Aug. 1934, H. Field and R.A. Martin leg. ( FMNH); 1 ♂, Hamedan Prov., Ganj nameh (34.76° N, 48.44° E), 2100 m, 4 Jul. 1974, A. Senglet leg ( MHNG); 1 ♂, Fars Prov., Allabad (30.01° N, 53.00° E), 9 Jun. 1974, A. Senglet leg. ( MHNG); 1 ♂, 1 ♀, Fars Prov., Bishapour (29.78° N, 51.58° E), 28 May 1974, A. Senglet leg. ( MHNG); 1 ♂, Kermanshah Prov., SE of Berendjan (34.55° N, 47.03° E), 12 Sep. 1975, A. Senglet leg. ( MHNG); 1 ♀, Yazd Prov., 10 km NE of Bafq (31.70° N, 55.53° E), 1260 m, 10 Apr. 2004, V. Vignoli and P. Crucitti leg. ( SMF); 1 ♂, 1 ♀, Yazd Prov., Mehriz, Gode Morghun Cave (31.4° N, 54.4° E), 1760 m a.s.l., date not given, M.S. Tahami leg. ( ZMSU); 2 ♂♂, Khuzestan Prov. [ Ahvaz , 31.31° N, 48.68° E], 17 Nov. 1958, W. Frank leg. ( SMF); 2 juvs, Fars Province, Shiraz (29.607° N, 52.533° E), 18–26 May 2000, Y.M. Marusik leg. ( ZFMK Ar 15235); 1 ♂, 1 ♀, 2 juvs, Fars Province, 50 km NNE of Shiraz (29.75° N, 52.75° E), 18–28 May 2000, Y.M. Marusik leg. ( ZFMK Ar 15236); 1 ♂, Fars Province, “ Hosseinie House ” [29.63° N, 52.51° E], 27 May 1981, collector not given ( ZFMK Ar 15237); 1 ♀, University of Shiraz [29.63° N, 52.51° E], 29 May 2000, F. Hosseinie leg. ( ZFMK Ar 15238).

UNITED ARAB EMIRATES: 2 ♀♀, Sharjah Desert Park (25.28º N, 55.69º E), hand-collected, 24 Feb. 2005, C. Griffioen leg. ( ZFMK Ar 15239); 2 ♀♀, same locality, in pitfall traps, 6–28 Dec. 2006, A. van Harten leg. ( ZFMK Ar 15240); 1 ♀, same locality, hand-collected, 6 Oct. 2004, C. Griffioen leg. ( ZFMK Ar 15241); 1 ♀, Um al-Quwain (25.51º N, 55.53º E), in pitfall traps, 1–30 Nov. 2008, A. van Harten leg. ( ZFMK Ar 15242); 2 ♀♀, Dubai, Deira, inside Al Quisais Mosque (25.277° N, 55.373° E), 7 May 2014, N. Iqbal leg. ( NHMUK); 1 ♂ (palps missing), 1 juv., Dubai, Falcon City, in building (25.087° N, 55.360° E), 22 May 2014, Morshid leg. ( NHMUK).

AFGHANISTAN: 1 ♂, Herat (34.33° N, 62.21° E), 15 Aug. 1975, A. Senglet leg. ( MHNG).

JAPAN: 1 ♀, Okayama Pref., Kasaoka Port [34.5° N, 133.5° E], Kasaoka-shi, 23 Dec. 1995, K. Nojima leg. ( ZFMK Ar 5203).

Description

Male (MNHN AR 10180)

MEASUREMENTS. Total body length 8.5, carapace width 4.1. Leg 1: 60.8 (16.0 + 1.9 + 18.2 + 21.2 + 3.5), tibia 2: 14.2, tibia 3: 10.0, tibia 4: 13.8; tibia 1 L/d: 36. Distance PME–PME 220 μm, diameter PME 200 μm, distance PME–ALE 140 μm, distance AME–AME 50 μm, diameter AME 190 μm.

COLOR. Carapace ochre with light brown median band; ocular area light brown, clypeus with subtle brown rim and light brown band below AME enlarged at base of clypeus to triangular shape (as in Fig. 90 ); legs ochre yellow to ochre without dark rings on femora subdistally but with light brown rings on patellae + tibiae proximally, and tibiae subdistally, tips of femora and tibiae not whitish; sternum ochre with narrow brown margins; abdomen beige with indistinct pale stripes from dorsal to posterior of abdomen with several marks dorsally.

BODY. Ocular area slightly elevated; carapace with distinct posterior furrow and distinct median pit; clypeus unmodified; sternum wider than long (3.0/2.2); chelicerae, with frontal row of ~20–25 modified (cone-shaped) hairs on each side, situated on elevated process (see Figs 97–99 ); cheliceral proximal incline in lateral view with distinct small process above modified hairs (see Fig. 97 ); with stridulatory ridges laterally (as in Fig. 118 ); abdomen globose and high; gonopore with four epiandrous spigots.

PALPS. Coxa unmodified, trochanter with short ventral projection, femur with distinct retrolateral process proximally, ventral membranous area proximally bordered on both sides by heavily sclerotized ridges, and small dorsal projection proximally; femur-patella hinges close together dorsally; patella very short; procursus with proximal dorsal process and weakly developed ventral pocket, and distal dorsal notch on prolateral sclerotized margin (as in Fig. 95 ); bulb with membranous embolus rising from base of processes a and b; process a with subdistal hump; process b narrow, elongated, and pointed; process c small, triangular (as in Fig. 95 ); process d is a distinct small ventral projection distally (as in Figs 94, 96 ).

LEGS.Without spines;with long curved hairs, especially on tibiae and metatarsi; retrolateral trichobothrium on tibia 1 at 8%; prolateral trichobothrium present on all tibiae; pseudosegmentation not visible.

Male (variation)

Tibia 1 in 19 other males: 8.0–22.2 (mean 14.5); color pattern on abdomen varies from pale without any marks to beige with lateral stripes and large marks dorsally; leg color varies from light brown to ochre; ocular area usually light brown; carapace pattern varies from sub-marginal brown marks to pale carapace with median band only; clypeus sometimes without dark band; process c varies in length, sometimes curved and pointing towards prolateral; lateral stridulatory ridges usually present and easy to notice ( Fig. 118 ), sometimes invisible; cheliceral proximal incline (in lateral view) usually with ridge or small process above modified hairs, sometimes absent or indistinct ( Figs 97, 100–102 ).

Female

In general similar to male; tibia 1 in 20 females: 5.3–14.3 (mean 8.3); stridulatory files laterally on chelicerae more distinct than in males; epigynal plate trapezoidal, consisting of two sclerotized lateral areas that are wider posteriorly, gently swollen posteriorly, and depressed medially anteriorly, pale median area inflated posteriorly with dark median sclerite ⅓–⅔ the length of epigynal plate and not fused at posterior epigynal margin with lateral sclerotized plates, epigynal posterior margin usually straight, sometimes with small median indentation (i.e., Figs 111, 113–114 ); anterior epigynal projections oval, not prominent in lateral view. Intraspecific variation of epigynum as shown in Figs 103–114 .

Distribution

This species is widespread, ranging from Israel and eastern Turkey in the west, to Afghanistan in the east ( Fig. 1 ). The single specimen from Japan is most probably a human introduction.

Note

We have not seen the material identified by Denis (1958), Roewer (1955), and Ghahari & Marusik (2009) as A. transcaspica (from Pirzada, Afghanistan, and various localities in Iran, respectively). We assume that these authors were all dealing with the very similar A. doriae , judging from its known distribution (we examined one male of A. doriae that was collected from Herat, Afghanistan, and the Iranian localities in the papers above also suggest A. doriae rather than A. transcaspica ).

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 90–96. Artema doriae Thorell, 1881. 90. Female, habitus (HUJ Ara 16033, Israel, Memorial Ha- Biq’a, NE Peza’el). 91–93. Male left pedipalp (ZFMK Ar 15233, Iran, Tehran): prolateral, dorsal, and retrolateral views. 94–96. Male left bulb and procursus (same specimen): prolateral, dorsal, and retrolateral views. Scale lines: 91–93 = 1 mm; 94–96 = 0.25 mm.

Figs 97–102. Artema doriae Thorell, 1881, male chelicerae in lateral and frontal views. 97–99. Iran, Tehran (MNHN Ar 10180). 100–101. Iran, Tehran (ZFMK Ar 15233 and SMF). 102. Israel, Yavni’el (HUJ Ara 16030). Arrows = projections proximally on frontal process. Scale lines: 97–99 = 0.5 mm; 100–102 = 0.2 mm.

Figs 103–114. Artema doriae Thorell, 1881, intraspecific variation of epigynum. 103. Iran, Tochal (ZFMK Ar 15233). 104. Iran, Esfahan (FMNH). 105. Iran, Lorestan, Ma‘amulan (MHNG). 106. Iran, Tehran (MNHN AR 10180). 107. Iran, 10 km NE of Bafq (SMF). 108–109. Turkey, Urfa-Gaziantep (XUM). 110. Turkey, Diyarbakır Prov., Çermik (ZFMK Ar 15931). 111. Israel, Upper Galilee, Ramot Naftali (CSZ). 112. Israel, Jericho (HUJ Ara 16035). 113. Israel, Memorial Ha-Biq’a, NE Peza’el (HUJ Ara 16033). 114. Israel, Modiin (SMNH). Scale lines: 103, 105–113 = 0.5 mm.

Figs 115–120. Artema doriae Thorell, 1881, MHNG, Ma’amulan (Iran). 115–116. Male chelicerae with frontal rows of modified (cone-shaped) hairs. 117. Male left bulbal processes. 118. Male stridulatory ridges. 119. Male left bulbal processes; arrow points at modified hair on palpal femur used for stridulation. 120. Apophysis c with putative pores. Scale lines: 115 = 0.4 mm; 116 = 0.06 mm; 117, 119 = 0.2 mm; 118 = 0.1 mm; 120 = 0.02 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.

Figs 128–133. Artema transcaspica Spassky, 1934, male chelicerae in lateral and frontal views. 128– 131. ♂♂, Uzbekistan, Zarafshon (ZMMU). 132. ♂, Turkmenistan, Badkhyz, Kushka (NHMW 13.677). 133. ♂, Uzbekistan, 36.5 km SSW of Uchkuduk (ZMMU). Arrows = absence of proximal projections on frontal processes. Scale lines: 128–130 = 0.5 mm; 131–133 = 0.2 mm.

Figs 186–194. Artema sp. c, male chelicerae, lateral and frontal views. 186, 190, 192. Pakistan Baluchistan Province, Quetta (ZMMU). 187, 193. India Rajastan, Durjana (SMF). 188–189, 191, 194. Sudan, Khartoum (MRAC 121706). Arrows = robust projection proximally on frontal process. Scale lines: 186–189, 192–194 = 0.5 mm; 190 = 0.2 mm; 191 = 0.25 mm.

Figs 134–145. Artema transcaspica Spassky, 1934, intraspecific variation of epigynum. 134. Uzbekistan, SE Bukhara (ZMMU). 135. Uzbekistan, Ak-Mechat (ZMMU). 136–137. Uzbekistan, SE Zarafshon (ZMMU). 138. Turkmenistan, Badkhyz, Kushka (NHMW 13.677). 139–145. Unspecified data: Tajikistan\Turkmenistan\Uzbekistan (ZIN type vials). Scale lines: 0.5 mm.

Figs 195–200. Artema sp. c, ♀♀, epigyna, ventral (195–199) and lateral (200) views. 195. India, Rajastan, hill S Pali (CAS). 196–198, 200. Pakistan, Baluchistan, Muslim Bagh (ZMMU). 199. Sudan (AMNH). Scale lines: 0.5 mm.