#ZooKeys ZooKeys 1240: 239-256 (2025) DOI: 10.3897/zookeys.1240.146399 Research Article Review of the wolf spider genus Acantholycosa Dahl, 1908 from China (Araneae, Lycosidae) Xiang-Yun Zhang™, Zhi-Sheng Zhang'™®, Lu-Yu Wang'® 1 Key Laboratory of Eco-environments in Three Gorges Reservoir Region (Ministry of Education), School of Life Sciences, Southwest University, Chongqing 400715, China Corresponding authors: Zhi-Sheng Zhang (zhangzs327@qq.com); Lu-Yu Wang (wangluyu1989@163.com) OPEN Qaccess Academic editor: Sarah Crews Received: 10 January 2025 Accepted: 23 April 2025 Published: 6 June 2025 ZooBank: https://zoobank.org/ E716A54F-CFF2-45F0-8ADB- 534EA97A6981 Citation: Zhang X-Y, Zhang Z-S, Wang L-Y (2025) Review of the wolf spider genus Acantholycosa Dahl, 1908 from China (Araneae, Lycosidae). ZooKeys 1240: 239-256. https://doi. org/10.3897/zookeys.1240.146399 Copyright: © Xiang-Yun Zhang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract The genus Acantholycosa Dahl, 1908 from China is reviewed, including five spe- cies: A. aborigenica Zyuzin & Marusik, 1988, A. lignaria (Clerck, 1757), A. sterneri (Marusik, 1993), A. zang sp. nov. (49) and A. zhangi sp. nov. (3°). Acantholycosa aborigenica and A. sterneri are recorded from China for the first time. Detailed de- scriptions are presented, along with photos of the habitus and copulatory organs, and a distribution map. Key words: Description, morphology, new record, new species, Pardosinae, taxonomy Introduction Acantholycosa Dahl, 1908 is a relatively large genus, comprising 37 species from the Holarctic (WSC 2025). Most species are endemics, restricted to the mountainous regions of South Siberia and Far East Russia (Marusik et al. 2004; Fomichev and Omelko 2020; Fomichev 2021). The genus is rela- tively well studied due to the existence of one global (Marusik et al. 2004) and several regional revisions (Fomichev and Marusik 2018; Fomichev and Omelko 2020; Fomichev 2021). In China, only two Acantholycosa species have been recorded: A. baltoroi (Caporiacco, 1935) and A. lignaria (Clerck, 1757). However, examination of the type specimen of A. baltoroi from India proved that it was a member of another wolf spider genus, Evippa Simon, 1882. The misidentification by Buchar (1976) — based on Roewer’s (1955) incorrect generic placement — and by the subse- quent authors indicated in the synonymy of the new species, means that all records of A. baltoroi were misidentified, leaving A. lignaria as the sole species of Acantholycosa reported from China (Sankaran and Caleb 2023). Based on the examination of newly collected specimens or those pre- viously identified as A. baltoroi from Hebei, Inner Mongolia, Jilin, Sichuan and Xizang, we reviewed the material and found two new species: A. zang sp. nov. and A. zhangi sp. nov., two species previously unknown in China (A. aborigenica Zyuzin & Marusik, 1988 and A. sterneri (Marusik, 1993)), and the previously reported species A. lignaria. 239 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China Material and methods All specimens were preserved in 75% ethanol and examined, illustrated, photo- graphed and measured using a Leica M205A stereomicroscope equipped with a drawing tube, a Leica DFC450 camera, and Leica Application Suite software (version 4.6). Male palps and epigynes were examined and illustrated after dis- section. Epigynes were cleared in a pancreatin solution (Alvarez-Padilla and Hormiga 2007). Leg measurements are shown as: total length (femur, patella + tibia, metatarsus, tarsus). All measurements are in millimetres. Specimens examined here are deposited in the spider collection at the School of Life Sci- ences, Southwest University, Chongqing of China (SWUC). Abbreviations used in the text: ALE, anterior lateral eye; AME, anterior medi- an eye; C, conductor; CO, copulatory opening; E, embolus; FD, fertilization duct; Ho, hood; HS, head of spermatheca; MA, median apophysis; Pa, paleal apoph- ysis; PLE, posterior lateral eye; PME, posterior median eye; Sb, septal base; Ss, septal stem; SS, stalk of spermatheca; St, subtegulum; TA, terminal apophysis; Te, tegulum. Terminology in the article follows Wang et al. (2025). Taxonomy Family Lycosidae Sundevall, 1833 Genus Acantholycosa Dahl, 1908 Type species. Lycosa sudetica L. Koch, 1875. Diagnosis. Members of Acantholycosa can be recognized by tibia | with 4, 5 or 6 pairs of ventral spines; palea modified, with a laminar or claw-like outgrowth referred to as a paleal apophysis; terminal apophysis long, with spine-like end; median apophysis with reduced apical arm (smaller than basal arm, except in A. oligerae Marusik, Azarkina & Koponen, 2004 and A. petrophila Marusik, Azarkina & Koponen, 2004); and presence of a spine-shaped or triangular out- growth at the base of the embolus in about half of the species (Marusik et al. 2004). Most females of this genus can be recognized by 4 to 6 pairs of ventral spines on tibia I, an elongate epigyne subdivided into an atrium (basal part) and upper flat part (exception baltoroi-group), hoods fused or almost fused (not separated by septum) and located far from atrium, and long spermathecae. Description. Medium sized (5.86-11.19). Carapace pear-shaped, with par- tially visible dark lateral bands, from moderately light colored to almost black. Fovea longitudinal and brown. Cervical groove and radial furrows indistinct. Eye region black. Chelicerae light brown to dark brown, with three promarginal and three retromarginal teeth. Endites and labium light brown to dark brown, longer than wide. Sternum pale brown to dark brown, shield shaped, covered with se- tae. Legs yellowish brown to dark brown, with black annulations. Spination vari- able, tibia | with 4 to 6 pairs of ventral spines. Leg formula: 4123, 4132 or 4312. Abdomen oval, dorsum light colored to almost black, without distinct pattern. Heart mark distinct or indistinct, lanceolate, light yellow to black. Venter light brown to brown. Spinnerets pale yellow to dark brown. Palp (Figs 1A, B, 2C-F, 3A, B, 4B-F, 5A, B, 6C-F, 7A-C, 8B-F, 10A-C, 11B-F) with uniformly brown to dark brown and droplet-shaped cymbium. Number of ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 240 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China claws variable, from 1 to 3. Palea modified, with three conformations: plate-like outgrowth, claw- or hook-like apophysis, and small triangular outgrowth. Tip of terminal apophysis with three modifications: small spine, strong conical spine- or claw-like outgrowth, and strong conical or cylindrical outgrowth bifurcate at tip or truncate. Median apophysis with two arms: apical arm and basal arm. Apical arm usually fully or partially reduced. Embolus wide, broader in terminal half. Tip of embolus with at least four variations: truncate, widened terminally and bifurcate, sharply curved in direction of bulb apically and slightly curved beyond bulb. Basal third of embolus in majority of east Palaearctic species with modifications: small spine, big conical spine, long outgrowth subdivided apically, and flat triangular lamina. Conductor membranous. Epigyne (Figs 1C, D, 2G, H, 5C, D, 6G, H, 7D, E, 9B, C, 10D, E, 12B, C) with three modifications of hood: undivided, two hoods fused to some extent, or two sep- arate hoods. Atrium well developed. In several species, septal base covering nearly almost entire atrium. Septum longer than wide, with variable base. Copu- latory openings located at anterolateral or posterolateral margin of septal base. Spermathecae long, without loops or sharp turns. Heads of spermathecae with wart-like projections in many species. Composition. 37 species. Biology. Almost all Acantholycosa species live in mountain scree (Marusik et al. 2004). Distribution. Widespread throughout the Holarctic. Acantholycosa aborigenica Zyuzin & Marusik, 1988 Figs -1,i2;.03 Common name. +3 lJ Acantholycosa sudetica Loksa, 1965: 16, fig. 23 (9). Pardosa aborigenica: Platnick 1993: 492. Acantholycosa aborigenica Zyuzin & Marusik, 1988: 1083, figs 1-6 (4'2); Marusik et al. 2004: 123, figs 108-114, 125-127, 147-151 (39); Marusik and Omelko 2011: 5, figs 22, 35-36 (3'2); Fomichev and Omelko 2020: 264, figs 31-40 (3'2). Material examined. CHINA: + 1 19, Inner Mongolia, Hulun Buir, Genhe City, Hanma National Nature Reserve, Abei Forest Farm, 51°48'42"N, 122°38'47'E, elev. 1023 m, 2.07.2018, R.B. Wu leg. Diagnosis. Acantholycosa aborigenica is similar to A. irinae Fomichev & Om- elko, 2020 (Fomichev and Omelko 2020, figs 1, 2, 7, 8, 13-15, 22-24, 48-49) in having the embolic tip bent anteriorly and median apophysis lacking apical arm (Figs 1A, B, 2C-F). The males of A. aborigenica can be distinguished from those of A. irinae by the paleal apophysis with broadened end ventrally (Figs 1A, 2C, E) (vs. sharp); median apophysis with triangular basal arm in ventral view (Figs 1A, 2C, E) (vs. semi-oval); absence of embolic spine near embolic base (Figs 1A, 2C, E) (vs. presence). Females can easily be distinguished from those of A. irinae by the septal base occupying 1/4 atrium (Figs 1C, 2G) (vs. 1/2), and septal stem reaching the apical pocket (Figs 1C, 2G) (vs. not reaching the apical pocket). Description. See Marusik et al. (2004). Habitus as shown in Fig. 2A, B, male palp as in Figs 1A, B, 2C-F, epigyne as in Figs 1C, D, 2G, H. ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 241 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China Figure 1. Acantholycosa aborigenica Zyuzin & Marusik, 1988, male (A, B) and female (C, D) A left male palp, ventral view B same, retrolateral view C epigyne, ventral view D same, dorsal view. Distribution. From Central Aimag in Mongolia to the upper reaches of the Kolyma River and south to Inner Mongolia (China). Comment. This species belongs to the A. /ignaria-group. Acantholycosa lignaria (Clerck, 1757) Figs 3, 4,13 Common name. Ai JRE Araneus lignarius Clerck, 1757: 90, pl. 4, fig. 4 (39). Acantholycosa lignaria: Dahl 1908: 367, 369, fig. 61 (3:2); Song et al. 1999: 316, fig. 186B (2); Marusik et al. 2004: 119, figs 27-29, 54, 115-121 (49); Alm- quist 2006: 184, figs 187a—-h (49); Marusik and Omelko 2011: 6, figs 23, 33, 34, 37 (69). Material examined. CHINA: * 3), Inner Mongolia, Hulun Buir, Ewenki Co., Yimin- he Town, Yimin River bank, 48°36'6'N, 119°47'40'E, elev. 669 m, 26-31.07.1972, J.M. He and D.R. Yang leg. * 163, Jilin Prov., Yanbian Pref., Antu Co., Erdaobaihe Town, 42°26'11"N, 128°6'1"E, elev. 728 m, 24.06.1979. Diagnosis. Acantholycosa lignaria is similar to A. zonsteini Marusik & Om- elko, 2017 (Marusik and Omelko 2017, figs 1-9) in having a rather large paleal apophysis, bilobated tip of embolus and flat and broad embolic spine (Figs 3A, 1 For complete list of 26 taxonomic references see WSC (2025). ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 242 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China \ ZZ Figure 2. Acantholycosa aborigenica Zyuzin & Marusik, 1988, male (A, C-F) and female (B, G, H) A male habitus, dor- sal view B female habitus, dorsal view C left male palp, ventral view D same, retrolateral view E palp bulb, ventral view F same, retrolateral view G epigyne, ventral view H same, dorsal view. B, 4B-F), but it can be differentiated by the terminal apophysis angled ventrally (Figs 3A, 4B, D, F) (vs. straight); the triangular embolic spine extending posteri- orly (Figs 3A, 4B, D, F) (vs. conical spine extending prolaterally); and the subdis- tal part of the embolus not tapering ventrally (Figs 3A, 4B, D, F) (vs. tapering). Description. See Marusik et al (2004). Habitus as shown in Fig. 4A, male palp as in Figs 3A, B, 4B-F. Distribution. From Germany and Sweden east to Kamchatka and south to Jilin and northeastern Inner Mongolia (China). Comment. This species belongs to the A. /ignaria-group. ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 243 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China A-B, 0.5mm Figure 3. Acantholycosa lignaria (Clerck, 1757), male A left male palp, ventral view B same, retrolateral view. Acantholycosa sterneri (Marusik, 1993) Figs 5, 6, 13 Common name. ff FOIA Pardosa sterneri Marusik, 1993: 77, figs 1-3 (@). Acantholycosa sterneri: Kronestedt and Marusik 2002: 67, figs 2, 7, 8, 12, 13, 15-18, 21, 25-27 (49); Marusik et al. 2004: 113, figs 66-72 (4@); Marusik and Omelko 2017: 599, fig. 12 (3). Material examined. CHINA: - 1, Inner Mongolia, Chifeng City, Balinyou Co., Saihanwula National Nature Reserve, top of Hanshan Mt., 44°10'27'N, 118°44'2°E, elev. 1828 m, 27.06.2015, Z.S. Zhang and L.Y. Wang leg. + 19, Saihanwula National Nature Reserve, 44°18'96'"N, 118°75'31'E, elev. 1941 m, 15.08.2023, S.T. Shi et al. leg. * 14, Balinyou Co., Wulanba Mt, 44°26'33'N, 118°42'8'E, elev. 1846 m, 11.07. 2023, K. Yu et al. leg. Diagnosis. Acantholycosa sterneri is similar to A. solituda (Levi & Levi, 1951) (Kronestedt and Marusik 2002, figs 1,3-6, 9-11, 14, 19, 20, 22-24) in having a similar conformation of the copulatory organs (Figs 54—D, 6C—H), but it can be differentiated by the median apophysis with a reduced apical arm (Figs 5A, 6C, E) (vs. without apical arm); embolus basal part wider than apical part (Figs 5A, 6C, E) (vs. embolus basal part as wide as apical part); embolus with pointed tip ventrally (Figs 5A, 6C, E) (vs. blunt). Females can be distinguished by the septal width measuring 1.3 times its length (Figs 5C, 6G) (vs. 2.3 times its length), and spermathecae separated (Figs 5D, 6H) (vs. spermathecae close together). Description. See Kronestedt and Marusik (2002). Habitus as shown in Figs 6A, B, male palp as in Figs 5A, B, 6C-F, epigyne as in Figs 5C, D, 6G, H. ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 244 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China 0.2 mm —= Figure 4. Acantholycosa lignaria (Clerck, 1757), male A male habitus, dorsal view B left male palp, ventral view C same, retrolat- eral view D palp bulb, ventral view E same, retrolateral view F paleal apophysis, terminal apophysis and embolus, ventral view. Distribution. China (Inner Mongolia), Russia (South Siberia), Mongolia. Comment. This species belongs to the A. solituda-group. Acantholycosa zang sp. nov. https://Zzoobank.org/2AF6ADA8-310B-48C8-B01E-910BD979CFAE Figs: 7—9,.1.3 Common name. jet ill Jk Acantholycosa baltoroi: Buchar 1976: 202, figs 1-3 (JQ); Chen et al. 1998: 72, figs 13-19 (49); Song et al. 1999: 310, figs 186A, M (<2); Marusik et al. 2004: 112, fig. 60 (3'); Marusik and Omelko 2017: 597, fig. 10 (<4) (misidentified). Type material. - Holotype 3 (SWUC-T-LY-25-01): CHINA, Xizang, Nyingchi City, Zayli Co., 64" km of Provincial Highway S201, 29°19'41"N, 97°8'8'E, elev. 3903 m, 25.06.2018, L.Y. Wang et al. leg. * Paratypes: 3 12 (SWUC-T-LY-25-02~05), same data as holotype * 14', Qamdo City, Riwogé Co., Riwogé Town, 31°23'26'N, ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 245 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China A-B, 0.5mm C—D, 0.5mm ———_ Dee ieee Figure 5. Acantholycosa sterneri (Marusik, 1993), male (A, B) and female (C, D) A left male palp, ventral view B same, retrolateral view C epigyne, ventral view D same, dorsal view. 96°32'51'E, elev. 3933 m, 22.05.2017, T. Lu and Q.Y. Wang leg. * 1< (SWUC- T-LY-25-06), Qamdo City, Markham Co., Quzika Township, Xiaochangdu ViII., 29°11'53"N, 98°38'42"E, elev. 3496 m, 11.05.2017, T. Lu and Z.S. Wu leg. * 23) (SWUC-T-LY-25-07~08), Nyingchi City, Zayi Co., Guyu Township, 29°5'32'N, 97°17'13'E, elev. 3196 m, valley scrub, 25.05.2019, L.Y. Wang et al. leg. » 2 (SWUC-T-LY-25-09~10), Sichuan Prov., Garze Pref., Dege Co., Que’er Mt., Wud- aoban, 31°56'33'N, 98°55'1'E, elev. 4707 m, 18.06.2016, T. Lu et al. leg. * 74 39 (SWUC-T-LY-25-11~20), Garze Pref., Daocheng Co., Haizi Mt., Yunsecuo River bank, 29°27'4"N, 100°11'30"E, elev. 4615 m, 12.06.2016, T. Lu et al. leg. Etymology. The specific name comes from the Chinese word “zang”, the name of one of the Chinese ethnic minorities that mainly live in Xizang and Sichuan provinces of China; noun in apposition. Diagnosis. This new species can be separated from other Acantholycosa species by the lanceolate end and retrolateral serrate margin of the terminal apophysis ventrally (Fig. 8D, arrow). The new species is most similar to A. zhangi sp. nov. in having a similar conformation of the copulatory organs (Figs 7A-E, 8B-F, 9B, C cf. Figs 10A—-E, 11B-F, 12B, C), but it can be dif- ferentiated by the triangular paleal apophysis with a pointed tip in ventral view (Figs 7A, C, 8B, D, F) (vs. rectangular with a wavy tip; Figs 10A, C, 11B, D, F); the median apophysis axe shaped without spine (Figs 7A, 8B, D) (vs. rectangular with hook-shaped basal arm and triangular spine; Figs 10A, 11B, D); embolus with subparallel margins abruptly tapering at tip (Figs 7A, C, 8B, D, F) (vs. tapering from base to tip, slightly curved towards tip; Figs 10A, C, 11B, D, F). Females can be distinguished by having an epigyne with a pair of distinct hoods (Figs 7D, 9B) (vs. a pair of less pronounced hoods; ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 246 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China i , \ " j Fi = ‘ iy “Fa i D.: ae, ; Spe a Rast 9, he fj J 05mm Wi /. —S WY) ———_—. Figure 6. Acantholycosa sterneri (Marusik, 1993), male (A, C-F) and female (B, G, H) A male habitus, dorsal view B fe- male habitus, dorsal view C left male palp, ventral view D same, retrolateral view E palp bulb, ventral view F same, retro- lateral view G epigyne, ventral view H same, dorsal view. Figs 10D, 12B); width of hoods equal to septal posterior width (Figs 7D, 9B) (vs. one-third septal posterior width; Figs 10D, 12B); septal base hexagonal (Figs 7D, 9B) (vs. trapezoidal; Figs 10D, 12B); spermathecal heads with wart- like projections (Figs 7E, 9C) (vs. smooth; Figs 10E, 12C). Description. Male holotype (Fig. 8A) total length 9.23. Carapace 4.87 long, 3.91 wide; opisthosoma 4.49 long, 2.99 wide. Carapace dark brown. Eye sizes and interdistances: AME 0.15, ALE 0.15, PME 0.48, PLE 0.42; AME-AME 0.17, AME-ALE 0.10, PME-PME 0.52, PME-PLE 0.55. Clypeus 0.3 high. Leg mea- surements: | 14.82 (3.83, 5.06, 3.88, 2.05); Il 14.71 (3.99, 4.84, 3.85, 2.03); IIl 14.46 (3.62, 4.16, 4.56, 2.12); IV 19.73 (4.67, 5.69, 6.59, 2.78). Femur | with 5 dorsal and 3 prolateral spines; tibia | with 2 prolateral, 2 retrolateral and 8 ven- tral spines; metatarsus | with 5 prolateral, 5 retrolateral and 3 ventral spines. Palp (Figs 7A—-C, 8B-F). Cymbium droplet-shaped, with two claws and black setae. Paleal apophysis triangular with acuminate tip. Terminal apophysis with lanceolate end and retrolateral serrate margin in ventral view. Median apophysis ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 247 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China A—B, 0.5mm A B Figure 7. Acantholycosa zang sp. nov., holotype male (A, B), paratype male (C) and paratype female (D, E) A left male palp, ventral view B same, retrolateral view C paleal apophysis, terminal apophysis and embolus, ventral view D epigyne, ventral view E same, dorsal view. Abbreviations: C = conductor; CO = copulatory opening; E = embolus; FD = fertilization duct; Ho = hood; HS = head of spermathecae; MA = median apophysis; Pa = paleal apophysis; Sb = septal base; Ss = sep- tal stem; SS = stalk of spermathecae; St = subtegulum; TA = terminal apophysis; Te = tegulum. with concave fold dorsally and serrated margins, no apical arm. Embolus 5.5 times as long as wide, uniform, tapering abruptly at tip and without basal spine. Female paratype (SWUC-T-LY-25-02, Fig. 9A) total length 11.19. Carapace 5.18 long, 4.12 wide; opisthosoma 6.45 long, 4.35 wide. Same as in male. Eye sizes and interdistances: AME 0.17, ALE 0.17, PME 0.51, PLE 0.41; AME-AME 0.24, AME-ALE 0.13, PME-PME 0.58, PME-PLE 0.64. Clypeus height 0.29. Leg measurements: | 15.33 (4.16, 5.41, 3.58, 2.18); Il 15.02 (4.05, 5.12, 3.67, 2.18); Ill 16.40 (4.02, 5.13, 4.67, 2.58); IV 20.47 (4.98, 5.99, 6.73, 2.77). Femur | with 5 dorsal and 2 prolateral spines; tibia | with 2 prolateral, 2 retrolateral and 8 ventral spines; metatarsus | with 4 prolateral, 4 retrolateral and 3 ventral spines. Epigyne (Figs 7D, E, 9B, C). Hoods distinct, clearly separated from each other. Atrium as long as wide. Septum with distinct stem and hexagonal base equal in length and width. Width of hoods equal to septal posterior width. Copulatory openings located at anterolateral margin of septal base. Spermathecae heads oval with small wart-like projections. Spermathecal stalks slightly curved. Fer- tilization ducts slender, width between two ducts longer than length of duct. Distribution. China (Xizang, Sichuan), Nepal. Comment. This species belongs to the A. solituda-group. Remark. Although the specimens of A. baltoroi from Nepal were not examined, it is clear from the descriptions in Buchar (1976), Marusik et al (2004), Marusik and Omelko (2017) that A. baltoroi from Nepal is identical to A. zang sp. nov. ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 248 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China wz | v Figure 8. Acantholycosa zang sp. nov., holotype male (A-C) and paratype male (D-F) A male habitus, dorsal view B left male palp, ventral view C same, retrolateral view D palp bulb, ventral view E same, retrolateral view F paleal apophysis, terminal apophysis and embolus, ventral view. a Acantholycosa zhangi sp. nov. https://zoobank.org/FD58FC89-BDA4-4248-87DB-8153F5D1FC9F Figs3l0—12)13 Common name. 4k EQ RIFE Acantholycosa baltoroi: Song et al. 2001: 226, figs 138A-D (32); Zhang et al. 2022: 119, figs 83A-J (32). (misidentified). Type material. * Holotype 3 (SWUC-T-LY-26-01) and paratype 2° (SWUC-T- LY-26-02): CHINA, Hebei Prov., Zhangjiakou City, Yu Co., Xiaowutai Nature Re- serve, Jinhekou, 39°56'17"N, 114°58'4'E, elev. 1334 m, 1.07.2012, F. Zhang leg. ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 249 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China Figure 9. Acantholycosa zang sp. nov., paratype female A female habitus, dorsal view B epigyne, ventral view C same, dorsal view. Etymology. The specific name comes from the family name of Prof. Feng Zhang of Hebei University (Hebei, China), who collected this new species. Diagnosis. The new species is similar to A. zang sp. nov. in having sim- ilar conformation of copulatory organs (Figs 10A-E, 11B-F, 12B, C; 7A-E, 8B-F, 9B, C), but differs by paleal apophysis rectangular with wavy tip in ventral view (Figs 10A, C, 11B, D, F) (vs. triangular with pointed tip; Figs 7A, C, 8B, D, F); median apophysis rectangular with hook-shaped basal arm and triangular spine (Figs 10A, 11B, D) (vs. axe shaped without spine; Figs 7A, 8B, D); embolus gradually tapering from base to tip and slightly curved (Figs 10A, C, 11B, D, F) (vs. with parallel margins except for tip suddenly taper- ing; Figs 7A, C, 8B, D, F). Females can be distinguished by two less pro- nounced (Figs 10D, 12B) (vs. distinct hoods, Figs 7D, 9B); width of hoods one-third septal posterior width (Figs 10D, 12B) (vs. width of hoods equal to septal posterior width; Figs 7D, 9B); septal base trapezoidal (Figs 10D, 12B) ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 250 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China A-B, 0.4mm Figure 10. Acantholycosa zhangi sp. nov., holotype male (A, B), paratype male (C) and paratype female (D, E) A left male palp, ventral view B same, retrolateral view C paleal apophysis, terminal apophysis and embolus, ventral view D epigyne, ventral view E same, dorsal view. Abbreviations: C = conductor; CO = copulatory opening; E = embolus; FD = fertilization duct; Ho = hood; HS = head of spermathecae; MA = median apophysis; Pa = paleal apophysis; Sb = septal base; Ss = sep- tal stem; SS = stalk of spermathecae; St = subtegulum; TA = terminal apophysis; Te = tegulum. (vs. base hexagonal without spine; Figs 7D, 9B); spermathecal heads smooth (Figs 10E, 12C) (vs. heads with wart-like projections; Figs 7E, 9C). Description. Male holotype (Fig. 11A) total length 9.47. Carapace 4.78 long, 3.86 wide; opisthosoma 4.61 long, 2.78 wide. Carapace black brown with dark margins. Eye sizes and interdistances: AME 0.16, ALE 0.14, PME 0.50, PLE 0.35; AME-AME 0.14, AME-ALE 0.10, PME-PME 0.45, PME-PLE 0.62. Cly- peus height 0.29. Leg measurements: | 14.56 (3.80, 5.20, 3.67, 1.89); Il 14.29 (3.53, 5.08, 3.70, 1.98); Ill 14.16 (3.48, 4.46, 4.22, 2.00); IV 18.94 (4.47, 5.58, 6.26, 2.63). Femur | with 5 dorsal and 2 prolateral spines; tibia | with 2 dorsal, 1 prolateral, 1 retrolateral and 8 ventral spines; metatarsus | with 2 dorsal, 4 prolateral, 5 retrolateral and 3 ventral spines. Palp (Figs 10A—-C, 11B-F). Cymbium with dense black setae and one claw. Paleal apophysis rectangular with wavy end. Terminal apophysis with a hook- shaped tip in retrolateral view. Median apophysis rectangular with hook-shaped basal arm and triangular spine. Embolus originating at 9 o'clock position, broad and almost straight, without basal spine, tapering from base to tip; distal end of embolus slightly curved, pointing posteriorly. Female paratype (Fig. 12A) total length 10.33. Carapace 5.16 long, 3.91 wide; opisthosoma 5.43 long, 3.72 wide. Eye sizes and interdistances: AME 0.16, ALE 0.15, PME 0.50, PLE 0.41; AME-AME 0.17, AME-ALE 0.10, PME-PME 0.49, ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 251 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China Figure 11. Acantholycosa zhangi sp. nov., holotype male (A-C) and paratype male (D-F) A male habitus, dorsal view B left male palp, ventral view C same, retrolateral view D right palp bulb, ventral view (mirror flip) E same, retrolateral view (mirror flip) F paleal apophysis, terminal apophysis and embolus, ventral view. PME-PLE 0.59. Clypeus 0.54 high. Leg measurements: | 14.89 (3.92, 5.47, 3.54, 1.96); Il 14.68 (4.04, 5.06, 3.58, 2.00); Ill 14.73 (3.88, 4.57, 4.31, 1.97); IV 20.63 (5.03, 6.18, 6.55, 2.87). Femur | with 5 dorsal and 2 prolateral spines; tibia | with 2 prolateral, 2 retrolateral and 8 ventral spines; metatarsus | with 5 prolateral, 4 retrolateral and 3 ventral spines. Except for the lighter body color, the other features are the same as those of the males. Epigyne (Figs 10D, E, 12B, C). Two hoods less pronounced and shallow. Atri- um rhomboid. Septum with trapeziodal base. Width of hoods one-third septal posterior width. Soermathecal heads clavate and smooth. Spermathecal stalks slightly curved. Fertilization ducts extending antero-laterally. Distribution. China (Hebei). ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 252 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China a re a \ ; / ae CT - = —— ie iy ii Cait” till t ~ = ». P a sl Figure 12. Acantholycosa zhangi sp. nov., paratype female A female habitus, dorsal view B epigyne, ventral view C same, dorsal view. Comment. This species belongs to the A. solituda-group. Remarks. Following the taxonomic revision by Sankaran and Caleb (2023) that transferred Acantholycosa baltoroi to the genus Evippa Simon, 1882 (currently recognized as E. baltoroi), and the subsequent recognition that all previous records of A. baltoroi represented misidentifications, the former "A. baltoroi-group” requires nomenclatural adjustment. We hereby propose the reclassification of this species group as the Acantholycosa solituda-group, based on taxonomic priority. In addition, because of the closer morphological similarity between A. lignaria and A. zonsteini, we reclassify A. zonsteini along with the similar species A. levinae Marusik, Azarkina & Koponen, 2004 into the A. lignaria-group. Therefore, the A. solituda-group comprises the following valid species: A. sergeevi Fomichev, 2021, A. solituda, A. sterneri, and the two newly described taxa (A. zang sp. nov. and A. zhangi sp. nov.). ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 253 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China 1 Acantholycosa aborigenica Zyuzin & Marusik, 1988 2 Acantholycosa lignaria (Clerck, 1757) 3 Acantholycosa sterneri (Marusik, 1993) 4 Acantholycosa zang sp. nov. r 5 Acantholycosa zhangi sp. nov. Ie N / 2) Ulaanbaatar A fs 44 en & - bit 80°E 85°E 90°E 95°E 100°E 105°E 110°E 115°E 120°E 125°E Figure 13. Distribution of Acantholycosa species in China. Acknowledgements Great thanks are given to the subject editor, Dr Sarah Crews, and four reviewers, Drs Yuri M. Marusik, Petr Dolejs, Galina N. Azarkina and Luis N. Piacentini for their helpful comments. Many thanks are given to Prof. Feng Zhang (College of Life Sciences, Hebei University, Baoding, China) for sending specimens as gifts. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding This research was supported by the Science & Technology Fundamental Resources In- vestigation Program (Grant No. 2022FY202100). ZooKeys 1240: 239-256 (2025), DOI: 10.3897/zookeys.1240.146399 254 Xiang-Yun Zhang et al.: Wolf spider genus Acantholycosa from China Author contributions All authors have contributed equally. Author ORCIDs Xiang-Yun Zhang © https://orcid.org/0009-0007-1627-3360 Zhi-Sheng Zhang ® https://orcid.org/0000-0002-9304-1789 Lu-Yu Wang ® https://orcid.org/0000-0002-5250-3473 Data availability All of the data that support the findings of this study are available in the main text. 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