Bioacoustics of Artema

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 ) and a ventral pocket (vp: Fig. 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of taxa are ambiguous with respect to the positon of Artema (J. Eberle, A. Valdez-Mondragón, D. Dimitrov, B.A. Huber, unpublished data).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

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 , 76 , 121 ). 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 ); sometimes without any pattern (as in Figs 51–53 ). 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 , 68–69 ) 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 , 70 , 118 ). Abdomen globose and high (as in Fig. 5 ). Gonopore with 4–7 epiandrous spigots ( Fig. 63 ). Anterior lateral spinnerets with one extremely widened, one long pointed, and six cylindrically shaped spigots each ( Figs 26, 28 , 74 ); PMS with two spigots each ( Figs 25 , 75 ); PLS without spigots. PALPS. Coxa unmodified. Trochanter with short ventral projection (as in Figs 77–79 ). Femur with short, somewhat pointed retrolateral process proximally (as in Figs 180–182 ); 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 ); 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 , 27 , 39–40 , 64 ). 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 , 71 , 154 ). Epigynal plate usually consisting of two sclerotized lateral areas ( Fig. 15 ), sometimes with dark median sclerite anteriorly ( Figs 48 , 103 , 138 ; variably large and variably distinct); with distinctive pair of anterior epigynal projections (AEP: Figs 15 , 86 , 103 , 138 ). Cleared female epigyna are shown in dorsal views ( Figs 201–207 ) and in posterior dorsal views ( 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.

Notes This male specimen resembles A. kochi for its lack of bulbal processes c, d, and e but, unlike A. kochi , bulbal process b is pointed as in all other congeners and not flattened ( Fig. 175 ). Cheliceral processes with frontal row of about 12 modified hairs on each side, situated distally on chelicerae in “hook-like” arrangement ( Fig. 179 ). Stridulatory files laterally on chelicerae present ( Fig. 177 ). Although the male palp appears distinctive, this single male specimen was the only Artema specimen available to us from North Africa that was not A. atlanta . It was probably collected long ago (date is not indicated) impeding molecular analysis. We suggest that more individuals (incl. females) should be available before the formal description of this putative species.

Notes Female distinguished from all other Artema females by simple rectangular epigynum ( Fig. 170 ). Stridulatory files laterally on chelicerae present. Epigynum almost twice as wide as long, posterior margin almost straight, epigynal plate consisting of two gently sclerotized lateral areas that appear swollen posteriorly, pale median area wider anteriorly, dark median sclerite absent. Pair of projections anterior to epigynum (AEP) present but not prominent. Female genitalia in the genus Artema often show high intra-specific variation, e.g., as in A. nephilit sp. nov. ( Figs 45–54 ) and in A. doriae ( Figs 103–114 ), both of which occur close to Oman ( Yemen and the United Arab Emirates ). Therefore, we prefer not to describe this putative species formally until more individuals (females and males) are avilable.