Artema 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 : 3-8

publication ID

https://doi.org/10.5852/ejt.2017.376

DOI

https://doi.org/10.5281/zenodo.6524915

persistent identifier

https://treatment.plazi.org/id/516F87BB-130E-BC70-FD50-B9E2FAF2ED91

treatment provided by

Carolina

scientific name

Artema Walckenaer, 1837
status

 

Artema Walckenaer, 1837

Artema Walckenaer, 1837: 656 ; type species: Artema atlanta (by subsequent monotypy).

Coroia González-Sponga, 2005: 102; type species: Coroia magna González-Sponga, 2005 ; synonymized by Huber et al. 2014: 416.

Diagnosis

Artema is easily distinguished from other pholcids by its large body and strong legs (body length 5.5– 9.5 mm; leg span up to 15 cm; tibia 1 L/d: 34–42); also by distinctive pattern on globose and high abdomen (dark dots dorsally, arranged in stripes from dorsal to lateral, sometimes absent; Figs 3–5 View Figs 3–8 , 51– 53 View Figs 45–54 ); by male pedipalp with its unique bulbal processes and short but massive procursus with proximal dorsal process (dp: Fig. 89 View Figs 83–89 ) and weakly developed ventral pocket (vp: Fig. 89 View Figs 83–89 ); by armature of male chelicerae (frontal row of cone-shaped hairs on each side, situated on elevated processes or ridges; Figs 23 View Figs 15–24 , 44 View Figs 35–44 ); and by pair of low to high projections in front of large anterior epigynal plate (AEP: Fig. 15 View Figs 15–24 ).

Description

Male

MEASUREMENTS. Total body length 5.5–9.5, carapace width 3.0–4.3. Leg 1: 39–61, tibia 1: 6.0–22.2; tibia 1 L/d: 34–42. Distance PME–PME 150–220 μm, diameter PME 170–230 μm, distance PME–ALE 90–140 μm, distance AME–AME 30–60 μm, diameter AME 180–210 μm.

COLOR. In general variable, even within species. Carapace pale to light brown, sometimes with brown radial marks, with light to dark brown median band (as in Figs 35 View Figs 35–44 , 76 View Figs 76–82 , 121 View Figs 121–127 ). Clypeus with wide light brown band. Legs pale yellow to ochre with dark rings on femora subdistally, patellae + tibiae proximally, and tibiae subdistally; tips of femora and tibiae whitish. Abdomen pale gray to light brown, usually with dark dots dorsally, arranged in distinctive stripes from dorsal to lateral side of abdomen (as in Fig. 4 View Figs 3–8 ); sometimes without any pattern (as in Figs 51–53 View Figs 45–54 ).

BODY. Ocular area slightly elevated. Carapace with median pit and distinct posterior furrow. Clypeus unmodified. Sternum wider than long. Chelicerae with frontal row of ~16–25 modified (cone-shaped; Figs 34 View Figs 30–34 , 68–69 View Figs 67–75 ) hairs on pair of elevated processes or ridges, variable in shape among species; lateral field of stridulatory ridges on chelicerae present or absent (variable even within species; Figs 31 View Figs 30–34 , 70 View Figs 67–75 , 118 View Figs 115–120 ). Abdomen globose and high (as in Fig. 5 View Figs 3–8 ). Gonopore with 4–7 epiandrous spigots ( Fig. 63 View Figs 55–66 ). Anterior lateral spinnerets with one extremely widened, one long pointed, and six cylindrically shaped spigots each ( Figs 26, 28 View Figs 25–29 , 74 View Figs 67–75 ); PMS with two spigots each ( Figs 25 View Figs 25–29 , 75 View Figs 67–75 ); PLS without spigots.

PALPS. Coxa unmodified. Trochanter with short ventral projection (as in Figs 77–79 View Figs 76–82 ). Femur with short, somewhat pointed retrolateral process proximally (as in Figs 180–182 View Figs 180–185 ); ventral membranous area proximally bordered on both sides by sclerotized ridges. Patella very short. Procursus with proximal dorsal process and weakly developed ventral pocket ( Fig. 89 View Figs 83–89 ); with brush-like distal modifications of filiform cuticular projections (cf. Huber 2000: fig. 48). Bulb with several distinctive processes (here called ‘processes a–e’; processes c–e sometimes absent) and membranous embolus arising from base of process a (e.g., Figs 19–20 View Figs 15–24 , 27 View Figs 25–29 , 39–40 View Figs 35–44 , 64 View Figs 55–66 ).

LEGS. Without spines; with long curved hairs, especially on tibiae and metatarsi; retrolateral trichobothrium on tibia 1 at 8–10%; prolateral trichobothrium present on all tibiae; tibia 2/ tibia 4 length usually <1.0, only in Artema doriae Thorell, 1881 >1.0; pseudosegmentation of tarsi not visible; tarsi 4 with distinct comb-hairs of the plesiomorphic, complex ‘ Belisana -type’ ( Huber & Fleckenstein 2008).

Female

In general similar to male, but legs shorter (tibia 1: 5.3–16.5), and stridulatory files laterally on chelicerae always present and distinct ( Figs 30 View Figs 30–34 , 71 View Figs 67–75 , 154 View Figs 146–154 ). Epigynal plate usually consisting of two sclerotized lateral areas ( Fig. 15 View Figs 15–24 ), sometimes with dark median sclerite anteriorly ( Figs 48 View Figs 45–54 , 103 View Figs 103–114 , 138 View Figs 134–145 ; variably large and variably distinct); with distinctive pair of anterior epigynal projections (AEP: Figs 15 View Figs 15–24 , 86 View Figs 83–89 , 103 View Figs 103–114 , 138 View Figs 134–145 ). Cleared female epigyna are shown in dorsal views ( Figs 201–207 View Figs 201–207 ) and in posterior dorsal views ( Figs 208–214 View Figs 208–214 ); the diagnostic value of the female internal genitalia in Artema is unclear so we do not put further emphasis on this in the species descriptions.

Natural history

Like most other non-forest pholcids, representatives of the genus Artema live in dark and sheltered habitats such as caves, holes and crevices, and spaces under large rocks. However, they also occur in man-made habitats like bunkers, buildings and basements, and under concrete bridges (cf. Huber & Kwapong 2013). In a large cave survey we conducted in Israel ( Aharon 2016), we found most individuals of A. nephilit sp. nov. in caves in a very hot and arid zone with low mean annual precipitation (50–150 mm). Other species in the genus were also collected partly from very dry regions and arid deserts with less than 50 mm of annual precipitation, e.g., A. doriae at the Arava Valley, Israel, and Artema sp. b at El Goléa in the Algerian Sahara.

The web is made of non-sticky silk and often appears irregular. This may partly be due to the fact that webs accumulate over time to form huge masses of old silk full of exuviae, while the spider occupies only a small but more regular slightly domed sheet within this mass ( Fig. 6 View Figs 3–8 ). As usual in pholcids, females carry their egg sacs, but the silk cover is much denser than in most other pholcid genera observed ( Fig. 3 View Figs 3–8 ). For further data on natural history, see under A. nephilit sp. nov. description below.

Distribution

The type species A. atlanta Walckenaer, 1837 has a worldwide distribution, presumably as a result of human-aided transport ( Fig. 2 View Figs 1–2. 1 ). All other species are restricted to the Old World, ranging from the African Sahel to Central and South Asia ( Fig. 1 View Figs 1–2. 1 ).

Relationships

Together with eight other genera, Artema is a member of the subfamily Arteminae Simon, 1893 , which currently contains 99 nominal species. Other genera included are: Wugigarra Huber, 2001 ; Trichocyclus Simon, 1908 and Holocneminus Berland, 1942 in Australia and Southeast Asia; Chisosa Huber, 2000 ; Aucana Huber, 2000 and Physocyclus Simon, 1893 in the New World, and the monotypic genera Tibetia Zhang, Zhu & Song, 2006 and Nita Huber & El-Hennawy, 2007 in Central Asia and northeastern Africa ( Huber 2011 b, 2016). The subfamily is characterized by two (functionally closely related) morphological synapomorphies, which constitute a pair of distinctive structures on the procursus: a dorsal process (dp: Fig. 89 View Figs 83–89 ) and a ventral pocket (vp: Fig. 89 View Figs 83–89 ) ( Huber & Eberhard 1997; Huber 2011b). Previous molecular studies supported the subfamily (e.g., Dimitrov et al. 2013), but current analyses of a much larger sample of taxa are ambiguous with respect to the positon of Artema (J. Eberle, A. Valdez-Mondragón, D. Dimitrov, B.A. Huber, unpublished data). Morphological data suggest that Artema is ‘basal’ within Arteminae , i.e., sister to all other genera. This is supported by the fact that all other Arteminae genera lack epiandrous spigots ( Huber 2016). Internal relationships among species remain largely obscure. Some of the similarities described below (e.g., between A. transcaspica and A. doriae ) are likely synapomorphic, while others (e.g., among A. nephilit sp. nov., A. bunkpurugu , A. atlanta and A. magna ) may turn out to be homoplastic.

Composition

As construed here, the genus Artema includes eight nominal species: the type species A. atlanta Walckenaer, 1837 ; A. bunkpurugu Huber & Kwapong, 2013 ; A. kochi Kulczyński, 1901 (revalidated); A. nephilit sp. nov.; A. doriae Thorell, 1881 ; A. transcaspica Spassky, 1934 ; A. magna Roewer, 1960 ; and the dubious A. ziaretana ( Roewer, 1960) (which is based on a single juvenile specimen and might be a synonym of A. magna ; see Notes under redescription of A. magna ).

Based on known distribution patterns (no small-scale endemism), we do not expect a significantly higher species richness. However, some of our taxonomic decisions may prove wrong as more material becomes available and this may affect species numbers. In addition, specimens from northern Africa ( Algeria, Sudan), the Arabian Peninsula ( Yemen, Oman), the Middle East ( Iran), and South Asia ( Pakistan, India) that we do not formally describe may all represent additional species.

Identification key

Males

1. Genital bulb without process c ( Figs 81 View Figs 76–82 , 175 View Figs 170–179. 170 ) ……………………………………………………2

– Genital bulb with process c ( Figs 20 View Figs 15–24 , 40 View Figs 35–44 ) ………………………………………………………3

2. Genital bulb with bulbal process b flattened, rectangular distally ( Fig. 81 View Figs 76–82 ); cheliceral processes strongly projecting proximally with frontal rows of modified hairs ( Fig. 83 View Figs 83–89 ); with several coneshaped, sclerotized processes situated frontally near median line (arrows in Figs 83, 85 View Figs 83–89 ) ……… ………………………………… ……………………………………… Artema kochi Kulczyński, 1901

– Genital bulb with bulbal process b rather pointed ( Fig. 175 View Figs 170–179. 170 ); cheliceral processes not strongly projecting proximally and without sclerotized processes near median line ( Figs 177–178 View Figs 170–179. 170 ) …… ………………………………………………………………………………… Artema sp.b( Algeria)

3. Genital bulb process c curved and pointing towards prolateral ( Figs 40 View Figs 35–44 , 95 View Figs 90–96 ) …………………4

– Genital bulb process c different (either membranous and pointing towards process b, or with sclerotized ridge projecting perpendicular to process b and with small teeth prolaterally; Huber & Kwapong 2013: fig. 58; Figs 20 View Figs 15–24 , 27, 29 View Figs 25–29 ) …………………………………………………8

4. Genital bulb without process d ( Fig. 39 View Figs 35–44 ) or with low rounded projection on ventral side of bulb ( Fig. 159 View Figs 155–161 ). Cheliceral process with median projection ( Figs 43–44 View Figs 35–44 , 164 View Figs 162–169 ) ………………5

– Genital bulb with distinct process d ( Figs 94 View Figs 90–96 , 125 View Figs 121–127 ). Cheliceral process without median projection ( Figs 98 View Figs 97–102 , 129 View Figs 128–133 , 192–194 View Figs 186–194 ) ……………………………………………………………………………6

5. Genital bulb without process d ( Fig. 39 View Figs 35–44 ). Each chelicera with frontal row of about 20 modified hairs that splits distally, with outer branch ending at tip of process, inner branch ending at tip of median projection ( Fig. 44 View Figs 35–44 ) ………………………………………… Artema nephilit sp. nov.

– Genital bulb process d indistinct, low rounded projection on ventral side of bulb ( Fig. 159 View Figs 155–161 ). Chelicerae modified hairs on median projection not linked (by row of hairs) to main modified hairs ridge (as in A. nephilit sp. nov.) ( Fig. 163 View Figs 162–169 ) …………………… Artema magna Roewer, 1960

6. Chelicerae in lateral view without or with very indistinct ridge or process above modified hairs ( Figs 128, 131–133 View Figs 128–133 ) …………………………………………… Artema transcaspica Spassky, 1934

– Chelicerae in lateral view with distinct ridge or process above modified hairs ( Figs 100–102 View Figs 97–102 , 186– 191 View Figs 186–194 ) …………………………………………………………………………………………………7

7. Chelicerae in lateral view with small ridge or process above modified hairs ( Figs 100–102 View Figs 97–102 ) ……… ………………………………………………………………………… Artema doriae Thorell, 1881

– Chelicerae in lateral view with robust ridge or process above modified hairs ( Figs 186–191 View Figs 186–194 ) …… …………………………………………………………… Artema sp. c ( India, Pakistan, Iran, Sudan)

8. Genital bulb process c partly membranous and directed towards process b; processes d and e absent. Chelicerae with frontal row of about ten modified hairs in straight line on each side (cf. Huber & Kwapong 2013: figs 59–60) ……………… Artema bunkpurugu Huber & Kwapong, 2013

– Genital bulb process c with sclerotized ridge projecting prolaterally perpendicular to process b, with small teeth prolaterally on round end of process ( Figs 20 View Figs 15–24 , 27, 29 View Figs 25–29 ); processes d and e present (both low and rounded) ( Fig. 19 View Figs 15–24 ). Chelicerae with frontal rows of about 20 modified hairs each in s-shaped pattern ( Fig. 24 View Figs 15–24 ) …………………………………… Artema atlanta Walckenaer, 1837

Females

1. Anterior epigynal projections (AEP) sclerotized and elongated ( Fig. 86 View Figs 83–89 ); epigynal plate wide rectangular, with triangular anterior projection between AEP …… Artema kochi Kulczyński, 1901

– Anterior epigynal projections (AEP) not or only weakly sclerotized and not elongated; epigynal plate different ………………………………………………………………………………………2

2. Epigynal plate rectangular, without dark median sclerite and without prominent pale median area ( Fig. 170 View Figs 170–179. 170 ) ……………………………………………………………… Artema sp. a ( Oman)

– Epigynal plate different, either with dark median sclerite and/or prominent pale median area …3

3. Epigynal plate posterior margin strongly indented ( Fig. 15 View Figs 15–24 ) …… Artema atlanta Walckenaer, 1837

– Epigynal plate posterior margin either straight ( Figs 46 View Figs 45–54 , 105 View Figs 103–114 ), gently indented ( Figs 135, 143 View Figs 134–145 ) or protruding ( Fig. 169 View Figs 162–169 ) ………………………………………………………………………………4

4. Anterior epigynal projections (AEP) distinctively prominent and pointed (cf. Huber & Kwapong 2013: figs 49, 53–54) ……………… Artema bunkpurugu Huber & Kwapong, 2013

– Anterior epigynal projections (AEP) not prominent and not pointed ……………………………5

5. Epigynal plate ~2.5 × wider than long; median posterior margin distinctively protruding ( Fig. 169 View Figs 162–169 ) ………………………………………………………………………… Artema magna Roewer, 1960

– Epigynal plate less than ~2.5 × wider than long, median posterior margin not protruding ………6

6. Epigynal plate with long dark median sclerite fused at posterior epigynal margin with lateral sclerotized plates ( Figs 134–145 View Figs 134–145 ) …………………………… Artema transcaspica Spassky, 1934

– Epigynal plate long with dark median sclerite not fused at posterior epigynal margin with lateral sclerotized plates ( Figs 45–50 View Figs 45–54 , 103–114 View Figs 103–114 , 195–199 View Figs 195–200 ) ………………………………………………7

7. Epigynal plate pale median area very inflated and prominent ( Figs 195–200 View Figs 195–200 ) …………………… ………………………………………………………… Artema sp. c ( India, Pakistan, Sudan, Iran)

– Epigynal plate pale median area slightly inflated or not inflated …………………………………8

8. Epigynal plate posterior margin almost straight, gently undulating. Pale median area with distinct dark median sclerite ⅓–½ the length of epigynal plate ( Figs 45–50 View Figs 45–54 ) …… Artema nephilit sp. nov.

– Epigynal plate trapezoidal, i.e., wider posteriorly; pale median area slightly inflated posteriorly; dark median sclerite ⅓–⅔ the length of epigynal plate; dark median sclerite not fused at posterior epigynal margin with lateral sclerotized plates ( Figs 103–114 View Figs 103–114 ) … Artema doriae Thorell, 1881

Kingdom

Animalia

Phylum

Arthropoda

Class

Arachnida

Order

Araneae

Family

Pholcidae

Loc

Artema Walckenaer, 1837

Aharon, Shlomi, Huber, Bernhard A. & Gavish-Regev, Efrat 2017
2017
Loc

Artema

Walckenaer C. A. 1837: 656
1837
Loc

Coroia González-Sponga, 2005: 102

Coroia González-Sponga, 2005: 102