Zoosyst. Evol. 99 (2) 2023, 325-335 | DO! 10.3897/zse.99.103061 > PENSUFT. yee BERLIN A review of the genus Sernokorba Kamura, 1992 (Araneae, Gnaphosidae) Nikolett Gallé-Szpisjak', Rébert Gallé’:*, Tamas Szits? 1 ELKH Centre for Ecological Research, Lendiilet Landscape and Conservation Ecology Research Group, Alkotmadny ut 2-4. Vacratot, 2163, Hungary 2 MTA-SZTE ‘Momentum’ Applied Ecology Research Group, Kozép fasor 52, Szeged, 6726 Hungary 3 Department of Ecology, University of Veterinarian Medicine Budapest, Rottenbiller u. 50, Budapest, 1077, Hungary https://zoobank. org/8A BCBDD1-84C8-4761-SA6E-85EBF34D1507 Corresponding author: Nikolett Gallé-Szpisjak (szpisjak.nikolett@ecolres. hu) Academic editor: Danilo Harms # Received 6 March 2023 # Accepted 16 May 2023 Published 2 June 2023 Abstract The gnaphosid spider genus Sernokorba Kamura, 1992 is reviewed. While Sernokorba pallidipatellis (Bosenberg and Strand 1906) and Sernokorba fanjing Song, Zhu & Zhang, 2004, occur in the Far East and the Japanese archipelago, Sernokorba tescorum (Simon, 1914) is known from Europe. We here describe a fourth species, Sernokorba betyar sp. nov. (male and female) from the forest steppe vegetation in southern Hungary in Central Europe. Digital images, comparative drawings (except for S. fanjing) and a distribution map are provided for all the species, and an identification key is compiled. The cheliceral dentation as diagnostic character and its interpretation are discussed. Key Words Central Europe, forest steppe, identification key, new species, spider Introduction Kamura (1992) described the monotypic ground spider genus Sernokorba to accommodate Prosthesima pallidipatellis Bosenberg & Strand, 1906 from Japan, which was at that time placed in Ze/otes Gistel, 1848, since it lacks the preening comb on metatarsi HJ-I'V characteristic of the genus. The species occurs also in China (Song et al. 1999), Korea (Namkung 2002) and Russian Far East (Marusik 2009). According to Kwon et al. (2014), this ground-dwelling species occurs in a wide variety of habitats, in forests, vineyards and grasslands. Currently, three species of the genus are known. The second species, Sernokorba fanjing Song, Zhu & Zhang, 2004 was described from Mt. Fanjing, Guizhou, China. The third species occurs in Europe. It was originally described as Poechilochroa tescorum Simon, 1914 on the basis of a female specimen. In their recent study, Cornic and Ledoux (2013) revised the species, described the male and proposed a new combination, Sernokorba tescorum (Simon, 1914). The species has been collected in nine locations in southern France, in grasslands and pine forests on calcareous soil. Later, Hernandez-Corral et al. (2017) and Breitling (2018) provided new occurrence data from the Iberian Peninsula, from a Quercus rotundifolia, Lamarck forest and a dry grassland, respectively. Naumova et al. (2021) reported the species from Bulgaria, from the leaf litter of a mixed forest of Fagus sylvatica, Linnaeus and Pinus heldreichii, Christ. In this paper we report, describe and illustrate Sernokorba specimens from Hungary for the first time, belonging to a hitherto unknown species, which we hereby describe as new to science. We provide an identification key, and illustrate occurrences of the genus, except for S. fanjing. Copyright Gallé-Szpisjak, N. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 326 Materials and methods The specimens of the new species were collected in the cal- careous sand-dune area of the Kiskunsag, central Hungary. The region belongs to the forest steppe zone, the transi- tional biome between the temperate deciduous forest and the steppe zones in Eurasia (Gallé et al. 2022a). The re- gion has a semiarid continental climate, as the mean annual precipitation is between 500-550 mm and the annual tem- perature is between 10 °C and 12 °C (Gallé et al. 2022a). The calcareous soil is poor in organic matter (Tolgyesi et al. 2018). The forest steppe vegetation appears as a mo- saic of (1) open dry grasslands: brome sward (Brometum tectorum), caliciphilous festuca steppe (Festucetum vagi- nateae) and Pannonian sand grassland (Potentillo-Festu- cetum pseudovinae), (2) wind-grooves between the sand dunes: more humid, with dune-slack purple moorgrass meadow (Molinio-Salicetum rosmarinifoliae) as main vegetation type, and (3) small forest patches of native ar- boreal plants such as Populus alba, Crataegus monogyna and Juniperus communis. Spiders were sampled with fun- nel traps (Csaszar et al. 2018), near Fulophaza in an intact forest steppe fragment, approximately 3.3 km? (46°52'N, 19°24'E, Figs 1-4). A total of 10 sites were established, and in each site, three habitat types (grassland, forest and forest edge) were sampled with four traps in each habitat. Sam- plings were done between 18 May and 12 June 2014. We collected 21 out of the 28 specimens in the forest edges. Type material of the new species will be deposited in the Hungarian Museum of Natural History, Budapest (HMNH, curator: E. Deakné Lazanyi-Bacso). A male of S. tescorum has been kindly loaned to us by Antonio Mel- ic (Sociedad Entomologica Aragonesa: PCAM). Spec- imens of S. pallidipatellis have been kindly donated to HNHM (Hungarian Natural History Museum, Budapest, ) by Prof. Takahide Kamura (Otemon Gakuin University). We did not have access to specimens of S. fanjing. Results Key to the species bo | = an OD n NM | - Copulatory openings not clearly visible (Figs 40-45) Copulatory openings well visible (Figs 46, 47) ............... Gallé-Szpisjak, N. et al.: Review of the genus Sernokorba Specimens were photographed using a Nikon D300S camera and a Tucsen TrueChrome Metrics camera at- tached to a Nikon S800 stereomicroscope and a Nikon Eclipse E200 compound microscope at the Department of Ecology, University of Veterinarian Medicine Buda- pest. Digital multifocal images were assembled using HeliconFocus image stacking software. Epigynes were removed and illustrations of them were made after a day-long maceration in commercial pancreatic enzyme solution. Palps were examined immersed in methyl-sa- licylate and mounted in a slightly modified Coddington mount (Coddington 1983). We modified the mount as the coverslip is not horizontal, but touching the slide itself, thus creating a triangle allowing more precise manipula- tion and more stable fixation of the object. Illustrations of the new species, S. tescorum and S. pallidipatellis were redrawn from digital images, while for S. fanjing they were produced from the figures of the original de- scription (Song et al. 2004). We used Adobe Illustrator CS6 vector graphics software. We illustrated the distri- bution of the Sernokorba species with Google Earth sat- ellite images, and Adobe Photoshop CS6 software. Measurements are given in mm. Lengths of leg seg- ments and total length were measured on the dorsal view. Leg formula developed by Ono (1988) is used; lengths of leg segments are given as: total length (femur, patella, tibia, metatarsus, tarsus). Abbreviations AME _ anterior median eyes; d dorsal; p prolateral; r retrolateral; RTA _ retrolateral tibial apophysis; Vv ventral. 3 Spermathecae relatively small (i.e., ~50% of the height of the vulva); sperm ducts straight (Figs 44, 45)... Fete STIS 0G 18 Ee are Pee Net ae, Pech Sernokorba pallidipatellis - Spermathecae relatively large (i.e., over 70% of the height of the vulva); sperm ducts with a proximal characteristic SWitchback*(Eies AOA St cee. teat nh, tasoNs aur ok Je ono tac one oe 4 Lateral edge of spermatheca with an angular posterior edge (Figs 28, 40, 41)............ eee Sernokorba betyar sp. nov. on | - Embolar tip straight (Figs 20-22, 29-32) op) AD SeiMaCHiSSaS WOO) iit cw easement Aree. haseaue eins: Lateral edge of spermatheca round (Figs 42, 43) ........... Embolar tip slightly bent (Figs 23, 33, 34)..............:20 Beene acetate eta gaan Secunteen waaca de nauy acetate Sernokorba tescorum Hgts Suet te oe tr sed Cay rene ts ei Sernokorba pallidipatellis Conductor blunt (Figs 29, 30, 35, 36), spermophore strongly bent as seen in retrolateral view, ventral bump on the RTA WHEE Tiedt id shoes Pmd Bhan bvatae ier Peo ketene Sernokorba tescorum - Conductor with fin-like branches (Figs 17-19, 31, 32, 37, 38), spermophore almost straight as seen in retrolateral view, ventral bump on the RTA present (Figs 14, 15, 32) ......... zse.pensoft.net cA 8 7a RENE 1 Pasta Oa GA bn CELE ey BU es Sernokorba betyar sp. nov. Zoosyst. Evol. 99 (2) 2023, 325-335 Map data 2023 (C) Google Figures 1—4. The habitat and distribution of Sernokorba species; 1, 2. Habitat of S. betyar in Fulophaza (Hungary); 3. Occurrences in Europe, red dots: S. betvar, yellow dots: S. tescorum; 4. Occurrences in Asia, green dots: S. pallidipatellis. zse.pensoft.net 328 Taxonomy Subfamily: Herpyllinae Platnick, 1990 (type genus Herpyllus Hentz, 1832) Genus Sernokorba Kamura, 1992 Diagnosis. The genus is a members of the Herpylline group (Azevedo et al. 2018), with a conspicuous black and white abdominal pattern (Figs 5—7). Males can be recognized by the following combination of characters: a single RTA that is about as long as the cymbium, thin, slightly bent, evenly narrowing terminally, and with a hook—shaped end (Figs 13-16). The conductor is about as long or longer than the tegulum, not twisted around the embolus (Figs 9-11, 13-15, 29, 31, 33). Females can be recognized by the kidney—shaped spermathecae (Figs 40, 42) and the sperm ducts being as long as the height of the spermathecae (Figs 40, 42, 44, 46). Description. See Kamura (1992). Furthermore, males in all the three examined species have a characteristic apical depression on the retrolateral side of gnathocox- ae (Fig. 8), which females do not have (Kamura 1992: fig. 4, Murphy 2007: 296, 310). This sexually dimorphic character, was not mentioned in the original descriptions of S. pallidipatellis and S. tescorum, or in the description of the genus itself. Distribution. The genus has been reported from East Asia (Korea, China, Russia and Japan) and from Western, Central and Southern Europe. Gallé-Szpisjak, N. et al.: Review of the genus Sernokorba Type species. Prosthesima pallidipatellis Bosenberg & Strand, 1906 — by original designation by Kamura (1992); female holotype from JAPAN: Saga Pref., in Senckenberg Museum, Frankfurt am Main — not examined. Sernokorba pallidipatellis (Bésenberg & Strand, 1906) Figs 12, 16, 23, 33, 34, 39, 44, 45, 48-50 Remark. As mentioned by Kamura (1992), Herpyllus coreanus Paik, 1992 is most likely conspecific with S. pallidipatellis, however, we do not propose a formal synonymy as we did not examine the type specimen of H. coreanus. Material examined. JAPAN: Kyoto | male, Kyoto-shi, Sakyo-ku, Matsugasaki, 28. April —5. May 1982 pitfall trap T. Kamura leg. (HNHM Araneae-9237); JAPAN: Osaka 1 male, Ibaraki-shi, Nishi-Ai, 26. May 1995 T. Kamura leg. (HNHM Araneae-9233); JAPAN: Osaka | female, Ibara- ki-shi, Ai, 30. May 1997 T. Kamura leg. (HNHM Arane- ae-9219):; JAPAN: Osaka | female, Ibaraki-shi, Nishi-A1, 9. May 2003 T. Kamura leg. (HNHM Araneae-9007). Diagnosis. Males can be recognized by the slightly bent embolar tip (Figs 23, 33) and by the large conductor reaching the tip of the embolus (Figs 33-34). Distal tip of the conductor is blunt and triangular. RTA with a ventral bump and bearing a distal invagination, resulting in two subequal branches. Description. See Kamura (1992). Distribution. Japan, Korea and the Russian Federation. Figures 5-8. European Sernokorba species; 5. Sernokorba tescorum, male habitus, dorsal view; 6-8. Sernokorba betyar sp. nov.; 6. Female habitus, dorsal view; 7. Male habitus, dorsal view; 8. Male, gnathocoxae, ventral view. Scale bars: 1.0 mm. zse.pensoft.net Zoosyst. Evol. 99 (2) 2023, 325-335 329 10° ) ow y, Figures 9-19. Palp comparison of the Sernokorba species; 9, 13. Sernokorba tescorum 9. Ventral view; 13. Retrolateral view; 10, 11, 14, 15, 17-19. Sernokorba betyar sp. nov. 10. Ventral view; 11. Proventral view; 14, 15. Retrolateral view; 17. Partially ex- panded palp, showing the conductor details, retrolateral view; 18. Conductor closeup, retrolateral view; 19. Slightly different angle; 12, 16. Sernokorba pallidipatellis;, 12. Ventral view; 16. Retrolateral view. Scale bar: 0.25 mm. zse.pensoft.net 330 Gallé-Szpisjak, N. et al.: Review of the genus Sernokorba Figures 20-28. Cleared copulatory organs; 20, 24. Sernokorba tescorum, 20. Ventral view; 24. Retrolateral view; 21, 22, 25-28. Sernokorba betyar sp. nov.; 21, 22. Ventral view; 25. Retrolateral view; 26. Prolateral view; 27. Epigyne, ventral view; 28. Vulva, dorsal view; 23. Sernokorba pallidipatellis, ventral view. Scale bar: 0.25 mm. Sernokorba tescorum (Simon, 1914) Figs 5, 9, 13, 20, 24, 29, 30, 35, 36, 42, 43 Remark. This species was recently reported from the Bal- kans, Bulgaria (Naumova et al. 2021). Fortunately, the record is accompanied with high quality images, and it is clear that the specimen shows some differences (1.e., In size of the cymbium, shape of the cymbium, the absence of clear finger-like extensions, and the coil shape of the spermo- phore) from the new species. However, we do not suggest any identification without examining the specimen itself. zse.pensoft.net Material examined. Spain: Cantoblanco: | male, Monte de Valdelatas Madrid. UTM: 30TVK4287, 40°32'11.5"N, 3°41'05.0"W, 700 m, 2002 May 27. A. Jiménez leg. (PCAM 5949). Diagnosis. Abdominal pattern consists of three pairs of elongated white spots. Males have blunt conductor (Figs 9, 29, 35, 36). The female is very similar to that of the new species, but can be differentiated by the round spermathecae (Figs 42, 43). Description. See Hernandez-Corral et al. (2017) and Cornic and Ledoux (2013). Zoosyst. Evol. 99 (2) 2023, 325-335 331 Figures 29-39. Illustrations of the male copulatory organs; 29, 30, 35, 36. Sernokorba tescorum, 29. Male copulatory or- gan ventral view; 30. Male copulatory organ, retrolateral view; 35. Conductor ventral view; 36. Conductor retrolateral view; 31, 32, 37, 38. Sernokorba betyar sp. nov.; 31. Male copulatory organ ventral view; 32. Male copulatory organ retrolateral view; 37. Conductor ventral view; 38. Conductor retrolateral view; 33, 34, 39. Sernokorba pallidipatellis, 33. Male copulatory organ ventral view; 34. Male copulatory organ retrolateral view; 39. Conductor ventral view. zse.pensoft.net 332 Distribution. France, Spain and Bulgaria (Naumova et al. 2021; Nentwig et al. 2022). A combination of all the published occurrences until 2020 (Cornic and Ledoux 2013; Hernandez-Corral et al. 2017; Breitling 2018) is shown in fig. 3. Sernokorba betyar sp. nov. https://zoobank.org/FD527ED0-2148-47D0-8E28-22B555FC37E3 Figs 6-8, 10; 11, 14, 15, 17-19, 21, 22, 25-28, 31, 32, 37, 38, 40, 41, S152 Type material. Holotype: HuNGAry: Fiiléphaza: male (46°51'55.00"N, 19°24'27.18"E) forest edge, pitfall trap, 1-10. June 2014, R. Gallé & N. Gallé-Szpisjak leg. (HNHM Araneae-9230). Paratypes: HuNGary: Tazlar: 1 male, 46°30'27.62"N, 19°30'2.22"E, forest edge, pitfall trap, 1-10. June 2014, R. Gallé & N. Gallé-Szpisjak leg. (HNHM Araneae-9229); HunGary: Fiiléphaza: 1 female 46°52'47.57"N, 19°24'17.36"E, forest edge, pitfall trap, 1-10. June 2014, R. Gallé & N. Gallé-Szpisjak leg. GHNHM Arane- ae-9228): HUNGARY: Fiiléphaza: 2 male, 46°52'46.93"N, 19°24'43.59"E, forest edge, pitfall trap, 18-25. May 2014 R. Gallé & N. Gallé-Szpisjak leg. (HNHM Arane- ae-9231); HUNGARY: Fiiléphaza: | female 46°53'13.61"N, 19°24'33.89"E, forest edge, pitfall trap, 18-25. May 2014 R. Gallé & N. Gallé-Szpisjak leg. (HNHM Arane- ae-9241): HuNGary: Fiiléphaza: 1 male, 46°52'9.10"N, 19°24'56.25"E, forest edge, pitfall trap, 18-25. May 2014 R. Gallé & N. Gallé-Szpisjak leg. (HNHM Araneae-9238). Other material examined. HUNGAry: Tazlar: | male, 1 female, 46°31'7.83"N, 19°31'10.80"E, forest edge, pitfall trap 1-10. June 2014 R. Gallé & N. Gallé-Szpisjak leg.; HunGary: Fiiléphaza: 1 male, 1 female, 46°52'15.25"N, 19°24'29.06"E, forest edge, pitfall trap, 18-25. May 2014 R. Gallé & N. Gallé-Szpisjak leg. (HNHM Araneae-9208). Diagnosis. The male can be identified by the finger— like extensions on the tip of the conductor (Figs 37, 38), and by the almost straight spermophore as seen from the retrolateral side (Fig 25, 32). Also, the male has an api- cal ectal depression on the gnathocoxae (Fig. 8), similar to that of S. tescorum. The female can be distinguished by the edge of the spermathecae: the lateral edge is more or less straight (vs. rounded in S. tescorum) and the posterior edge is concave (vs. convex in S. tescorum). Also, it can be distinguished by the deep atrial pockets (Figs 27, 28, 40, 41) opposed to the shallow atrial pokets of the S. tescorum. Description. Male (Holotype; HNHM Araneae-9230). Colour. Carapace light brown with pale brown radiating stripes, covered with white fine setae (Fig. 7); thorac- ic groove dark brown (Fig. 7). Chelicerae pale brown (Fig. 7). Gnathocoxae brown with a dark brown outline; terminal part in ventral view pale yellow. Labium brown (Fig. 8). Sternum brown with radial light brown spots, posterior end dark brown. Trochanter I brown, all other trochanters pale yellow. All femora dark brown; all other leg segments pale yellow. Abdomen dark greyish brown with a reddish scutum; an anterior transverse line and two zse.pensoft.net Gallé-Szpisjak, N. et al.: Review of the genus Sernokorba pairs of whitish spots present in the area of the scutum; anterior pair of spots placed closer to the midline, pos- terior pair situated laterally. Venter light greyish yellow, with two thin longitudinal stripes starting at epigastric furrow and extending towards spinnerets. Epigastric area yellowish—brown. Sides of abdomen dark brown. Spin- nerets’ proximal segments dark brown/black, distal part pale yellow. Carapace suboval, cephalic region much narrower (about 40% of maximal width), posterior region truncated (Fig. 7) and elevated, about twice as high as in front. Chelicerae thin, with one tooth on promargin and two teeth on retro- margin. Gnathocoxae with an oblique depression on the lateral margin (Fig. 8). Sternum longer than wide. Labium as long as wide (Fig. 8), triangular. Clypeus low, about the diameter of AME high. Abdomen ovoid, longer than wide, truncated in front, with scutum covering 60% of the dor- sum (Fig. 5). Total length, not including spinnerets, 4.32. Carapace 1.35 long, 0.98 wide, 0.41 high, highest at coxae III, widest at coxae II. Abdomen 2.72 long, 0.87 wide, with large bristles on proximal margin. Clypeus low, 0.10. Leg measurements: I 3.05 (1.01, 0.42, 0.65, 0.51, 0.46); II 3.00 (0.99, 0.43, 0.63, 0.48, 0.47); IIT 2.89 (0.94, 0.39, 0.63, 0.47, 0.46); IV 3.99 (1.07, 0.57, 0.82, 1.0, 0.53). Leg formula IV-I-[-III. Leg spination: I: femur d 1-1-1, p 0-0-1; tibia v 0-1-1. II: femur d 1-1-1, p 0-0-1; tibia v 0-1-1. III: femur d 1-1- 1, p 0-0-1, r 0-0-1; patella pl, rl; tibia p 0-1-1 v 0-1-1, r 0-1-1; metatarsus p 0-1-2, r 0-0-2, v 0-0-1. IV: femur d 1-1-1, p 0-0-1, r 0-0-1; tibia d 0-1-0, p 1-1-1, v 1-1-1, r 0-1-1; metatarsus p 1-2-2, v 1-0-0, r 1-1-2. Palp: Tibia longer than wide, RTA about 70% at cym- bium’s length (Figs 10, 11, 21, 22, 25), thin, bent in its middle and with an apical hook (Figs 14, 15, 32); sper- mophore U-shaped, proximal part of tegulum is tight (Figs 10, 11, 21, 22) (vs. loose in that of S. tescorum, as shown in Figs 9, 20). Spermophore bent slightly in retro- lateral view (Figs 15, 25). Conductor membranous, with finger-like extensions (Figs 17-19, 31, 32, 37, 38). Female (Paratype; HNHM 9241). Coloration as in male, except carapace and abdomen lighter (Fig. 6). Epi- gastric area yellowish—dark grey. Shape of carapace and abdomen (Fig. 6) as in males, except for absence of ab- dominal scutum. Total length, not including spinnerets, 4.04. Carapace 1.98 long, 1.50 wide, 0.57 high. Abdomen 3.58 long, 2.03 wide, 1.19. Leg measurements: I 4.19 (1.45, 0.58, 0.82, 0.74, 0.60); II 4.13 (1.43, 0.57, 0.80, 0.75, 0.58); III 4.11 (1.30, 0.54, 0.81, 0.89, 0.57); IV 5.67 (1.65, 0.63, 1.21, 1.45, 0.73). Leg formula. IV-I-II-II. Leg spination: I: femur d 1-1-1, p 0-0-1; tibia v 0-1-1. II: femur d 1-1-1, p 0-0-1; tibia v 0-1-1. III: femur d 1-1- 1, p 0-0-1, r 0-0-1; patella pl, rl; tibia d 1-0-0, p 1-1-l v 1-1-1, r 0-1-1; metatarsus d 0-1-1, p 0-1-1, r 0-1-1, v 1-0- 1. IV: femur d 1-1-1, p 0-0-1, r 0-0-1; tibia d 1-0-0, p 1-1- 1, v 1-1-2, r 0-1-1; metatarsus p 1-2-2, v 1-1-1, r 1-2-2. Epigyne: copulatory openings positioned medially on anterior part; copulatory ducts short; spermathecae robust and pear-shaped. Zoosyst. Evol. 99 (2) 2023, 325-335 333 Figures 40—47. Illustrations of the female genitalia; 40, 41 Sernokorba betyar sp. nov.; 40. Dorsal view; 41. Ventral view; 42, 43. Sernokorba tescorum, 42. Dorsal view; 43. Ventral view; 44, 45. Sernokorba pallidipatellis, 44. Dorsal view, 45. Ventral view, 46, 47. Sernokorba fanjing, 46. Dorsal view; 47. Ventral view. zse.pensoft.net 334 Gallé-Szpisjak, N. et al.: Review of the genus Sernokorba 50 Figures 48-52. Male chelicerae; 48-50. Sernokorba pallidipatellis, 51, 52. Sernokorba betyar sp. nov. Etymology. The specific name is a Hungarian noun in apposition and refers to the outlaws “betyar” found in hiding places on the Hungarian Great Plain, just as this species has been avoiding its discovery so far. Distribution. Bacs-Kiskun county, Forest steppes, Hungary. Discussion In the genus description of Sernokorba, Kamura (1992) men- tioned concerning S. pallidipatellis: “J recognized that this Species 1s unique in having a serrated carina on the promar- gin of fang furrow of chelicera”, unequivocally illustrated (Kamura 1992: fig. 3), and used in this sense by subsequent authors (Song et al. 2004; Kim and Lee 2013) for S. fanjing and S. pallidipatellis. However, Cornic and Ledoux (2013) reports teeth, but only on the promargin: “Marge antérieure des chéliceres garnie de trois dents, marge postérieure mu- tique’”. We compared the dentation of specimens of S. pal- lidipatellis (Figs 48—50) to both literature images and to that of S. betyar sp. nov. (Figs 51, 52). The cheliceral dentation of S. pallidipatellis (Figs 48-50) looks almost identical to zse.pensoft.net literature sources (Kamura 1992: fig. 3). However, we also agree with Cornic and Ledoux (2013) and interpret it as teeth rather than a keel or carina. Azevedo et al. (2018) well illustrated the non-serrated cheliceral promargin (Azevedo et al. 2018: fig 20. e, g), which is diagnostic to the subfamily Herpyllinae. However, the keel on the promargin may not just “be a subtle projection or may appear as teeth with fused bases” (Kamura 1992). Despite Kamura’s (1992) descrip- tion and subsequent authors (Song et al. 2004; Kim and Lee 2013) implies the latter; we observed that the bases of the teeth are in fact separated as viewed from an oblique view (Figs 49, 50). We observed five teeth lumped together with a smaller one further placed, whereas there was one clear tooth on the retromargin. Such dentation has been observed (but not illustrated) in Latonigena Simon, 1893 by Ott et al. (2012), thus this character may require a second look with more genera involved. The dentation of S. betyar sp. nov. (1.e., two teeth on the promargin and one on the retromargin, Figs 51, 52) is different also from S. tescorum (1.e., three teeth on the promargin, none on the retromargin) which seems a good confirmation to separate the two species. Species of the genus occur in a wide variety of hab- itats including lowland forests and grasslands, however, Zoosyst. Evol. 99 (2) 2023, 325-335 specimens were collected mainly in mountainous regions (Kwon et al. 2014; Hernandez-Corral et al. 2017; Naumo- va et al. 2021). According to our data, S. betvar sp. nov. is a typical member of the ground-dwelling spider fauna of lowland forest steppes. The relatively understudied forest steppes of Central Europe are among the most complex ecosystems in the region, and their nature conservation value is high. Forest steppes are important biodiversity hotspots (Gallé et al. 2022b) and listed in the Habitats Di- rective of the European Union. They harbour numerous endemic and rare plant and animal species, including sev- eral recently described spiders such as Parasyrisca arra- bonica Szinetar & Eichardt, 2009 (Gnaphosidae) and A/o- pecosa psammophila Buchar, 2001 (Lycosidae). Although the abundance of S. betvar sp. nov. seems to be very low, this species presumably occurs in vast areas of forest steppes of Central and Eastern European forest steppes. Acknowledgements The study would have been impossible to carry out with- out the valuable help of colleagues who have sent com- parative material of S. pallidipatellis (Takahide Kamura; Otemon Gakuin University), and S. tescorum (Antonio Melic; Sociedad Entomologica Aragonesa). 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