Evippinae Zyuzin, 1985

Steinpress, Igor Armiach, Alderweireldt, Mark, Cohen, Mira, Chipman, Ariel & Gavish-Regev, Efrat, 2021, Synopsis of the Evippinae (Araneae, Lycosidae) of Israel, with description of a new species, European Journal of Taxonomy 733, pp. 87-124 : 90-94

Subfamily Evippinae Zyuzin, 1985

Diagnosis

Zyuzin defined this subfamily by the character “The embolus enters the tegular apophysis, forming a single structural complex with it” ( Zyuzin 1985: 48). Roewer recognized this distinguishing feature for Evippinae (at the time, tribe Evippeae ): Metatarsus IV shorter than patella IV + tibia IV ( Roewer 1959). Roewer's character is invalid for identification, as Evippa amitaii sp. nov. deviates from these proportions.

Description

Small- to medium-sized lycosids ( Figs 1–5 ), superficially similar to Pardosinae and Lycosinae in somatic appearance, but distinct in genital structure ( Figs 6–13 ). Anterior eye row narrower than second eye row (PME). Tibia I with 2–6 pairs of ventral spines (apart from apical pair). Spinnerets not elongated. Body densely covered with short (often specialized) setae (Alderweireldt 1992) ( Figs 1 , 14 ). Sexual dimorphism usually weak ( Figs 4–5 ) (but see section about Evippomma ( Figs 1–2 , 4–5 )). Zyuzin (1985) described the Evippinae as having an embolus encased in a sheath of transparent tissue ( Figs 6–8 ); embolus base in mesoapical position, partly flattened, lying against cymbium; bent sharply near tip, inserted into tegular apophysis, forming single structure. Epigyne atria very shallow, pale, conspicuous; septum with narrow base, often covered with setae; spermathecae usually large and conspicuous ( Figs 10–11 , 13 ). Coloration variable; carapace often with median band ( Figs 1 – 3 ). Certain genera ( Evippa , Zenonina ) have a derived habitus, clearly discernible from other lycosid genera ( Roewer 1959; Alderweireldt 1991, 1992; Marusik Kovblyuk & Koponen 2011).

Natural history

Most species inhabit deserts, steppes and savannas, but species of open patches in and along forests ( Xerolycosa spp.) are known. Zyuzin (1985) described Xerolycosa as found in dry conifer forests, steppes and arable land, while Evippa inhabits flood plains, semideserts and deserts, often as a sole representative of Lycosidae . They are largely presumed vagrant ( Alderweireldt & Jocqué 2017), although burrow construction behavior is known from species of Xerolycosa ( Marusik et al. 2011) and Evippomma ( Bayer et al. 2017) . Both diurnal and nocturnal activity has been observed ( Zyuzin 1985;

* Direct submission to Gen Bank by Astrin J., Hoefer H., Spelda J., Holstein J., Bayer S., Hendrich L., Huber B.A., Kielhorn K.-H., Krammer H.-J., Lemke M., Monje J.C., Moriniere J., Rulik B., Petersen M., Janssen H. and Muster C.

I. Armiach Steinpress, pers. obs.). As in all lycosids, the female carries the egg sac attached to her spinnerets and carries the young on her body for some time after hatching (I. Armiach Steinpress, pers. obs.).

Distribution

The subfamily is restricted to the Old World where it is widely distributed, except in the polar region, wet tropics and wet subtropics. Xerolycosa is found across the Palearctic, from Europe to Japan (with a doubtful species from Zanzibar). Proevippa , Pseudevippa and Zenonina are restricted to sub-Saharan Africa. Evippomma is found in Africa and India (Indian species might belong to a different genus). Evippa is found across arid and semiarid climates throughout Africa and Eurasia, from Tanzania to Siberia ( World Spider Catalog 2020).

Relationships

Molecular studies that included Xerolycosa tend to place it in a rather basally branching position in Lycosidae , away from the Lycosinae ( Park et al. 2007; Zehethofer & Sturmbauer 1998). Murphy et al. (2006) have placed Xerolycosa as a sister taxon of the venoniin genera Aulonia (C.L. Koch, 1847) and Hygrolycosa (Dahl, 1908) (Venoniinae was not recovered in the study). A recent molecular phylogeny

included for the first time three representatives of Evippinae : Proevippa , Xerolycosa , and Evippomma , and recovered Evippinae as monophyletic ( Piacentini 2019).

Composition

Evippinae consists of 67 species in six genera: 38 species in Evippa Simon, 1882 ; seven species in Evippomma Roewer, 1959 ; eleven species in Proevippa Purcell, 1903 ; one species in Pseudevippa Simon, 1910 ; four species in Xerolycosa Dahl, 1908 ; and six species in Zenonina Simon, 1898 .

Figures

Fig. 1. Evippomma simoni Alderweireldt, 1992. Live ♂ (HUJ INV-Ar 16579), Mamshit, Apr. 2018. A. Lateral view. B. Frontal view. C. Dorsal view. D. Cephalic region in dorsal view, right side. Leafshaped setae discernible. Photos by I. Armiach Steinpress.

Fig. 2. Evippomma simoni Alderweireldt, 1992, a live ♀, (HUJ INV-Ar 20421), Mash’abbim sands, 16 June 2020. A. Lateral view. B. Frontal view. C. Dorsal view. D. Burrow in the field. Photos A–C by I. Armiach Steinpress; photo D by S. Aharon.

Fig. 3. ♀♀, live specimens. A. Evippa arenaria (Audouin, 1826) (Ze’elim, 21 Jul. 2015), dorsal view. B. E. praelongipes (O. Pickard-Cambridge, 1871) (Nahal Ashalim, 13 Apr. 2018), dorsal view. C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 20314, Biq’at ’Uvda (’Uvda valley), 13 Mar. 2018), dorsal view. D. E. onager (HUJ INV-Ar 20314, Biq’at ’Uvda (’Uvda valley), 13 Mar. 2018), frontal view. Photos by I. Armiach Steinpress.

Fig. 4. ♂♂, habitus. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16051). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16508). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 20316). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16041). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16049). Scale bars = 10 mm. Photos by A. Uzan.

Fig. 5. ♀♀, habitus. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16048). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16507). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 16510). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16044). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16544). Scale bars = 10 mm. Photos by A. Uzan.

Fig. 6. ♂♂, left palp, ventral view. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16051). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16508). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 16510). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16531). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16538). Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.

Fig. 7. ♂♂, left palp, retrolateral view. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16051). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16508). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 16510). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16531). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16538). Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.

Fig. 8. ♂♂, left palp, prolateral view. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16051). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16508). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 16510). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16531). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16538). Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.

Fig. 9. Evippa amitaii sp. nov., ♂ (HUJ INV-Ar 16051), bulb of left palp, proximal view. Photo by I. Armiach Steinpress. Scale bar = 0.3 mm.

Fig. 10. ♀♀, genitalia, ventral view. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16053). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16058). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 20314). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16521). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16543). Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.

Fig. 11. ♀♀, genitalia, dorsal view. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16053). B. E. arenaria (Audouin, 1826) (HUJ INV-Ar 16058). C. E. onager Simon, 1895 sensu Šternbergs 1979 (HUJ INV-Ar 20314). D. E. praelongipes (O. Pickard-Cambridge, 1871) (HUJ INV-Ar 16521). E. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16543). Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.

Fig. 12. Evippa amitaii sp. nov., ♂ (HUJ INV-Ar 16051), drawings of left palp. A. Ventral view. B. Retrolateral view. C. Proximal view.D. Prolateral view. Scale bars = 0.5 mm. Drawings by I. Armiach Steinpress.

Fig. 13. ♀♀, genitalia, drawings. A. Evippa amitaii sp. nov. (HUJ INV-Ar 16053), ventral view. B. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16543), ventral view. C. Evippa amitaii sp. nov. (HUJ INV-Ar 16053), dorsal view. D. Evippomma simoni Alderweireldt, 1992 (HUJ INV-Ar 16543), dorsal view. Scale bars = 0.5 mm. Drawings by I. Armiach Steinpress.

Fig. 14. Cephalic region (right side, dorsal view), showing covering setae. A. Evippa praelongipes (O. Pickard-Cambridge, 1871), round-profile setae. B. Evippomma simoni Alderweireldt, 1992, leafshaped setae. Scale bars = 0.5 mm. Photos by I. Armiach Steinpress.