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Songthela serriformis sp. nov. Figure 5 Holotype. Male (XUX-2012-245, matured on 13 June 2013), China, Chongqing Municipality, Nan���an District, Nanshan Botanic Garden, 29.56��N, 106.62��E, 554 m a.s.l., 18 November 2012, collected by D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-241A, matured on 22 March 2013) and 8 females (XUX-2012-237/238/239/240/2 41/242/243/244), same data as for holotype, 551���554 m a.s.l. Diagnosis. Males of S. serriformis sp. nov. can be distinguished from S. pluma, S. tianzhu, S. xiangnan and S. yuping by apical spine of conductor with a spinule basally, conductor forms a lamina with dentate edge proximally (Fig. 5A, B, E); from S. jinyun sp. nov., S. longbao sp. nov. and S. wangerbao sp. nov. by conductor with slender apical spine and dentate edge proximally (Fig. 5A, B, E); from other Songthela species by conductor with one apical spine and a dentate edge proximally (Fig. 5A, B, E). Females of S. serriformis sp. nov. differ from S. tianzhu by middle genital stalks longer and posterior part of genital area wider (Fig. 5H���M); from S. yuping by middle receptacular clusters lager than lateral ones and with longer genital stalks (Fig. 5H���J); from S. jinyun sp. nov., S. longbao sp. nov., S. wangerbao sp. nov. and other Songthela species by four receptacular clusters situated on dorsal wall of bursa copulatrix, and posterior margin of bursa copulatrix sclerotised (Fig. 5H���M). Description. Male holotype (XUX-2012-245). In alcohol, carapace yellowish-brown covered with a few hairs running over ocular area; opisthosoma brown with 12 dark brown tergites, close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, the fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 8 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 15.60, CL 6.95, CW 6.91, OL 8.90, OW 8.00; ALE> PLE> PME> AME; leg I 22.08 (6.11 + 2.80 + 4.55 + 5.82 + 2.80), leg II 22.79 (6.00 + 2.85 + 4.55 + 6.47 +2.92), leg III 24.88 (5.98 + 2.90 + 4.75 + 7.55 + 3.70), leg IV 31.00 (7.28 + 3.22 + 6.12 + 9.88 + 4.50). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines on tip (Fig. 5A��� C). Contrategulum with a regular denticulate margin (Fig. 5A, B, D). Tegulum with a helicoid marginal apophysis and dorsal extension of terminal apophysis, a platelike terminal apophysis retrolaterally (Fig. 5C, F, G). Conductor fused with embolus basally, gradually narrowing to a long apical spine with a spinule basally, forming a lamina with a dentate edge proximally (Fig. 5A, B, E). Embolus with a transparent, wide and flat opening, a distinct shrink near the opening ventrally, with numerous ribbed ridges retrolaterally (Fig. 5A, B, D, E). Female paratype (XUX-2012-244). In alcohol, carapace dark red-brown covered with a few hairs running over ocular area; opisthosoma light brown with 12 dark brown tergites, close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 12 denticles of variable size; legs with hairs/ spines; 8 spinnerets. Measurements: BL 14.05, CL 7.17, CW 5.91, OL 6.98, OW 5.68; ALE> PLE> PME> AME; palp 11.86 (4.10 + 1.95 + 2.53 + 3.28), leg I 13.49 (4.38 + 2.32 + 2.65 + 2.59 + 1.55), leg II 13.07 (4.27 + 2.22 + 2.47 + 2.53 +1.58), leg III 14.17 (4.05 + 2.33 + 2.51 + 3.31 + 1.97), leg IV 20.66 (5.58 + 2.87 + 3.75 + 5.53 + 2.93). Female genitalia: Four receptacular clusters situated on dorsal wall of bursa copulatrix. Middle pair with long genital stalks, larger than lateral ones. Lateral pair with obscure genital stalks situated more dorsally, posterior margin of bursa copulatrix sclerotised (Fig. 5H���M). Variation. Males and females vary in body size. The range of measurements in males as follow (N = 2): BL 13.51���15.60, CL 6.27���6.95, CW 6.12���6.91, OL 6.64���8.90, OW 5.60���8.00; females (N = 8): BL 10.19���18.12, CL 5.45���8.88, CW 5.07���7.68, OL 6.07���9.68, OW 4.26���7.05. The number of promargin of cheliceral groove varies from 8 to 12 (N = 10). There are 7 or 8 spinnerets. In addition, the conductor of males shows intraspecific variation: apical spine slightly curved with a spinule basally and forming a smooth lamina with a dentate edge proximally (Fig. 5A, B), or apical spine straight without spinule basally, and the formed lamina covered with several teeth (Fig. 5D, E). Female genitalia also show intraspecific variation: length of middle genital stalks is different (Fig. 5H���J); lateral receptacular clusters are distinctly smaller than middle ones (Fig. 5I) compared with the other samples (Fig. 5H, J), posterior margin of genital area trapezoid (Fig. 5K, L) or W-shaped (Fig. 5M). Etymology. The species epithet is an adjective from the combination of the Latin serri (serrate) and formis (shaped), and refers to the serrated margin of conductor proximally. Distribution. Chongqing (Nan���an), China. GenBank accession number. Holotype: XUX-2012-245: OL982297., Published as part of Chen, Zhaoyang, Liu, Fengxiang, Li, Daiqin & Xu, Xin, 2022, Four new species of the primitively segmented spider genus Songthela (Mesothelae, Liphistiidae) from Chongqing Municipality, China, pp. 546-558 in Zootaxa 5091 (4) on pages 553-555, DOI: 10.11646/zootaxa.5091.4.2, http://zenodo.org/record/5863681

Songthela jinyun sp. nov. Figure 3 Holotype. Male (XUX-2017-293, matured on 14 August 2019), China, Chongqing Municipality, Beipei District, Mt. Jinyun, 29.83ºN, 106.38ºE; 827 m a.s.l., 12 October 2017, collected by FX Liu, F Li, X Xu. Paratypes. 2 females (XUX-2017-290/291), same data as for holotype; 5 females (XUX-2012- 189/193/194/195/200), same data as for holotype, 29.84ºN, 106.40ºE, 713–729 m a.s.l., 14 November 2012, collected by D Li, FX Liu, X Xu; 2 females (XUX-2012-165/166), China, Chongqing Municipality, Beipei District, Mt. Jinyun, Banzigou, 29.86ºN, 106.38ºE, 253 m a.s.l. 29 November 2012, collected by D Li, FX Liu, X Xu; 2 females (XUX-2013-129A/133A), China, Chongqing Municipality, Beipei District, Mt. Jinyun, Banzigou, 29.86ºN, 106.38ºE, 266–283 m a.s.l. 29 March 2013, collected by D Li, X Xu, and LY Wang; 6 females (XUX-2017-282/2 84/285/287/288/289), China, Chongqing Municipality, Beipei District, Mt. Jinyun, 29.84ºN, 106.40ºE, 589–665 m a.s.l., 12 October 2017, collected by FX Liu, F Li, and X Xu. Diagnosis. Male of S. jinyun sp. nov. can be distinguished from S. ciliensis by conductor with wider base (Fig. 3B, E), and by marginal apophysis of tegulum incurved in middle portion (Fig. 3F, G); from S. pyriformis and S. longbao sp. nov. by apical spine of conductor slender and longer (Fig. 3A–G); from S. wangerbao sp. nov. by apical spine of conductor slightly thicker (Fig. 3A, B, D, E, G), and by middle portion of marginal apophysis of tegulum incurved (Fig. 3F, G); from S. serriformis sp. nov. by conductor with smooth margin (Fig. 3A–G); from other Songthela species by smooth conductor with one apical spine (Fig. 3A, D). Females of S. jinyun sp. nov. differ from S. ciliensis, S. longbao sp. nov. and S. wangerbao sp. nov. by distance between bases of middle genital stalks larger, middle genital stalks slightly slender (Fig. 3H, I); from S. bristowei by bases of four receptacular clusters separated from each other (Fig. 3H, I); from S. serriformis sp. nov. by middle receptacular cluster situated on anterior margin of bursa copulatrix, posterior margin of bursa copulatrix weakly sclerotised (Fig. 3H–K); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix and lateral receptacular clusters situated dorsolaterally (Fig. 3H–O). Description. Male holotype (XUX-2017-293) (Fig. 1E). In alcohol, carapace yellow brown with a few pointed hairs running over ocular area; opisthosoma pale yellow, with 12 brown tergites close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 10 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 14.78, CL 6.78, CW 6.44, OL 7.09, OW 4.44; ALE> PLE> PME> AME; leg I 21.74 (6.10 + 2.78 + 4.53 + 5.66 + 2.67), leg II 21.45 (5.91 + 2.68 + 4.29 + 5.77 + 2.80), leg III 22.96 (5.73 + 2.93 + 4.16 + 6.93 + 3.21), leg IV 30.45 (7.64 + 2.97 + 6.03 + 9.30 + 4.51). Palp: Paracymbium unpigmented and unsclerotised prolaterally, with numerous setae and spines distally (Fig. 3A–C). Contrategulum with denticulate margin, dentations sparse proximally and dense distally (Fig. 3A, D). Marginal apophysis of tegulum curved in middle portion, dorsal extension of terminal apophysis of tegulum with a helicoid edge, terminal apophysis of tegulum wide and lamellar retrolaterally (Fig. 3C, F, G). Conductor smooth, fused with embolus ventro-proximally, with an apical spine (Fig. 3B, E). Embolus with a transparent, wide and flat opening, slightly curved margin with numerous longitudinal ribs retrolaterally (Fig. 3A, B, D, E, G). Female paratype (XUX-2017-289). In alcohol, carapace dark red-brown with a few hairs running over ocular area; opisthosoma light brown, with 12 dark brown tergites close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 12 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 16.74, CL 7.46, CW 6.80, OL 8.02, OW 6.19; ALE> PLE> PME> AME; palp 13.59 (4.68 + 2.47 + 2.99 + 3.45), leg I 16.08 (4.98 + 2.70 + 3.13 + 3.23 + 2.04), leg II 15.45 (4.70 + 2.71 + 2.90 + 3.25 +1.89), leg III 16.74 (4.61 + 2.89 + 3.25 + 3.67 + 2.32), leg IV 23.15 (6.65 + 3.23 + 4.22 + 6.01 + 3.04). Female genitalia: Two pairs of receptacular clusters with tubular genital stalks and surrounded by transparent membrane. Middle pair of receptacular clusters situated on anterior margin of bursa copulatrix, separated from each other, smaller than lateral ones. Lateral ones situated slight dorsolaterally with short genital stalks, posterior margin of bursa copulatrix weakly sclerotised (Fig. 3H–O). Variation. Females vary in body size. The range of measurements in females (N = 17): BL 10.70–17.19, CL 4.83–6.75, CW 4.15–5.89, OL 5.56–9.02, OW 3.74–7.01. The number of promargin of cheliceral groove varies from 10–13 (N = 17). The number of spinnerets is 7 or 8. In addition, female genitalia show intraspecific variation: middle genital stalks tilted outward (Fig. 3H, L) or paralleled (Fig. 3I, J,); four receptacular clusters distinctly separated (Fig. 3H, I), or one of middle and lateral genital stalks fused basally (Fig. 3J, N), or only three receptacular clusters (Fig. 3K). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Beipei), China. GenBank accession number. Holotype: XUX-2017-293: OL982296.

Songthela jinyun sp. nov. Figure 3 Holotype. Male (XUX-2017-293, matured on 14 August 2019), China, Chongqing Municipality, Beipei District, Mt. Jinyun, 29.83��N, 106.38��E; 827 m a.s.l., 12 October 2017, collected by FX Liu, F Li, X Xu. Paratypes. 2 females (XUX-2017-290/291), same data as for holotype; 5 females (XUX-2012- 189/193/194/195/200), same data as for holotype, 29.84��N, 106.40��E, 713���729 m a.s.l., 14 November 2012, collected by D Li, FX Liu, X Xu; 2 females (XUX-2012-165/166), China, Chongqing Municipality, Beipei District, Mt. Jinyun, Banzigou, 29.86��N, 106.38��E, 253 m a.s.l. 29 November 2012, collected by D Li, FX Liu, X Xu; 2 females (XUX-2013-129A/133A), China, Chongqing Municipality, Beipei District, Mt. Jinyun, Banzigou, 29.86��N, 106.38��E, 266���283 m a.s.l. 29 March 2013, collected by D Li, X Xu, and LY Wang; 6 females (XUX-2017-282/2 84/285/287/288/289), China, Chongqing Municipality, Beipei District, Mt. Jinyun, 29.84��N, 106.40��E, 589���665 m a.s.l., 12 October 2017, collected by FX Liu, F Li, and X Xu. Diagnosis. Male of S. jinyun sp. nov. can be distinguished from S. ciliensis by conductor with wider base (Fig. 3B, E), and by marginal apophysis of tegulum incurved in middle portion (Fig. 3F, G); from S. pyriformis and S. longbao sp. nov. by apical spine of conductor slender and longer (Fig. 3A���G); from S. wangerbao sp. nov. by apical spine of conductor slightly thicker (Fig. 3A, B, D, E, G), and by middle portion of marginal apophysis of tegulum incurved (Fig. 3F, G); from S. serriformis sp. nov. by conductor with smooth margin (Fig. 3A���G); from other Songthela species by smooth conductor with one apical spine (Fig. 3A, D). Females of S. jinyun sp. nov. differ from S. ciliensis, S. longbao sp. nov. and S. wangerbao sp. nov. by distance between bases of middle genital stalks larger, middle genital stalks slightly slender (Fig. 3H, I); from S. bristowei by bases of four receptacular clusters separated from each other (Fig. 3H, I); from S. serriformis sp. nov. by middle receptacular cluster situated on anterior margin of bursa copulatrix, posterior margin of bursa copulatrix weakly sclerotised (Fig. 3H���K); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix and lateral receptacular clusters situated dorsolaterally (Fig. 3H���O). Description. Male holotype (XUX-2017-293) (Fig. 1E). In alcohol, carapace yellow brown with a few pointed hairs running over ocular area; opisthosoma pale yellow, with 12 brown tergites close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 10 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 14.78, CL 6.78, CW 6.44, OL 7.09, OW 4.44; ALE> PLE> PME> AME; leg I 21.74 (6.10 + 2.78 + 4.53 + 5.66 + 2.67), leg II 21.45 (5.91 + 2.68 + 4.29 + 5.77 + 2.80), leg III 22.96 (5.73 + 2.93 + 4.16 + 6.93 + 3.21), leg IV 30.45 (7.64 + 2.97 + 6.03 + 9.30 + 4.51). Palp: Paracymbium unpigmented and unsclerotised prolaterally, with numerous setae and spines distally (Fig. 3A���C). Contrategulum with denticulate margin, dentations sparse proximally and dense distally (Fig. 3A, D). Marginal apophysis of tegulum curved in middle portion, dorsal extension of terminal apophysis of tegulum with a helicoid edge, terminal apophysis of tegulum wide and lamellar retrolaterally (Fig. 3C, F, G). Conductor smooth, fused with embolus ventro-proximally, with an apical spine (Fig. 3B, E). Embolus with a transparent, wide and flat opening, slightly curved margin with numerous longitudinal ribs retrolaterally (Fig. 3A, B, D, E, G). Female paratype (XUX-2017-289). In alcohol, carapace dark red-brown with a few hairs running over ocular area; opisthosoma light brown, with 12 dark brown tergites close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 12 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 16.74, CL 7.46, CW 6.80, OL 8.02, OW 6.19; ALE> PLE> PME> AME; palp 13.59 (4.68 + 2.47 + 2.99 + 3.45), leg I 16.08 (4.98 + 2.70 + 3.13 + 3.23 + 2.04), leg II 15.45 (4.70 + 2.71 + 2.90 + 3.25 +1.89), leg III 16.74 (4.61 + 2.89 + 3.25 + 3.67 + 2.32), leg IV 23.15 (6.65 + 3.23 + 4.22 + 6.01 + 3.04). Female genitalia: Two pairs of receptacular clusters with tubular genital stalks and surrounded by transparent membrane. Middle pair of receptacular clusters situated on anterior margin of bursa copulatrix, separated from each other, smaller than lateral ones. Lateral ones situated slight dorsolaterally with short genital stalks, posterior margin of bursa copulatrix weakly sclerotised (Fig. 3H���O). Variation. Females vary in body size. The range of measurements in females (N = 17): BL 10.70���17.19, CL 4.83���6.75, CW 4.15���5.89, OL 5.56���9.02, OW 3.74���7.01. The number of promargin of cheliceral groove varies from 10���13 (N = 17). The number of spinnerets is 7 or 8. In addition, female genitalia show intraspecific variation: middle genital stalks tilted outward (Fig. 3H, L) or paralleled (Fig. 3I, J,); four receptacular clusters distinctly separated (Fig. 3H, I), or one of middle and lateral genital stalks fused basally (Fig. 3J, N), or only three receptacular clusters (Fig. 3K). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Beipei), China. GenBank accession number. Holotype: XUX-2017-293: OL982296., Published as part of Chen, Zhaoyang, Liu, Fengxiang, Li, Daiqin & Xu, Xin, 2022, Four new species of the primitively segmented spider genus Songthela (Mesothelae, Liphistiidae) from Chongqing Municipality, China, pp. 546-558 in Zootaxa 5091 (4) on pages 547-551, DOI: 10.11646/zootaxa.5091.4.2, http://zenodo.org/record/5863681

Songthela longbao sp. nov. Figure 4 Holotype. Male (XUX-2012-252, matured on 18 July 2013), China, Chongqing Municipality, Wanzhou City, Mt. Longbao, 30.76��N, 108.38��E, 226 m a.s.l., 18 November 2012, collected by D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-251A) and 10 females (XUX-2012-247/249/250/251/253/254A/255/256/257/2 58), same data as for holotype, 221���226 m a.s.l. Diagnosis. Males of S. longbao sp. nov. can be distinguished from S. ciliensis by conductor with a curved margin retrolaterally (Fig. 4B, E); from S. pyriformis by apical spine of conductor straight and slender (Fig. 4A, B, D, E), by marginal apophysis of tegulum semielliptic (Fig. 4C, G); from S. jinyun sp. nov. and S. wangerbao sp. nov. by apical spine of conductor slightly shorter and thicker (Fig. 4A, B), by marginal apophysis of tegulum helicoid (Fig. 4C, F); from S. serriformis sp. nov. by conductor with smooth margin (Fig. 4A���G); from other Songthela species by smooth conductor with one apical spine (Fig. 4A, B, E). Females of S. longbao sp. nov. differ from S. ciliensis by lateral genital stalks shorter (Fig. 4H���J); from S. jinyun sp. nov. by distance between bases of middle and lateral genital stalks larger (Fig. 4H���J); from S. wangerbao sp. nov. by middle genital stalks longer and close to each other (Fig. 4H���J); from S. bristowei by bases of four receptacular clusters separated from each other (Fig. 4H���J); from S. serriformis sp. nov. by middle receptacular clusters situated on anterior margin of bursa copulatrix, lateral receptacular clusters similar to or slightly larger than middle ones, posterior margin of bursa copulatrix weakly sclerotised (Fig. 4H���J); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix with long genital stalks, similar or smaller than lateral pair, lateral ones situated dorsolaterally (Fig. 4H���M). Description. Male holotype (XUX-2012-252). In alcohol, carapace reddish brown with a few pointed hairs running over ocular area; opisthosoma pale yellow, with 12 dark reddish brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 14.08, CL 6.15, CW 5.70, OL 7.20, OW 4.72; ALE> PLE> PME> AME; leg I 17.02 (4.88 + 1.97 + 3.52 + 4.20 + 2.45), leg II 17.58 (4.78 + 2.30 + 3.35 + 4.60 + 2.55), leg III 19.73 (5.00 + 2.23 + 3.55 + 5.85 + 3.10), leg IV 24.65 (6.20 + 2.60 + 4.60 + 7.20 + 4.05). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines distally (Fig. 4A���C). Contrategulum with denticulate margin, dentations sparse proximally and dense distally (Fig. 4A, D). Marginal apophysis of tegulum and dorsal extension of terminal apophysis of tegulum with helicoid edges, with a small terminal apophysis of tegulum (Fig. 4C, F, G). Conductor smooth, fused with embolus basally, with one apical spine (Fig. 4A, B, D, E). Embolus with a transparent, wide and flat opening and several ribbed ridges distally (Fig. 4A, B, D, E, G). Female paratype (XUX-2012-251). In alcohol, carapace slight red-brown covered with a few hairs running over ocular area; opisthosoma slight brown with 12 dark brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with hairs/ spines; 7 spinnerets. Measurements: BL 13.92, CL 5.25, CW 5.18, OL 7.32, OW 6.25; ALE> PLE> PME> AME; palp 8.60 (3.15 + 1.15 + 1.89 + 2.41), leg I 10.66 (3.40 + 1.90 + 2.07 + 1.98 + 1.31), leg II 10.50 (3.20 + 1.88 + 1.85 + 2.22 +1.35), leg III 11.00 (3.06 + 1.83 + 1.83 + 2.68 + 1.60), leg IV 16.74 (4.68 + 2.25 + 3.06 + 4.45 + 2.30). Female genitalia: Four receptacular clusters separated from each other basally; median pair situated on anterior margin of bursa copulatrix with long genital stalks, similar to or slightly smaller than lateral pair; lateral ones situated dorsolaterally; posterior margin of bursa copulatrix weakly sclerotised (Fig. 4H���M). Variation. Males and females vary in body size. The range of measurements in males (N = 2): BL 10.37���14.08, CL 5.04���6.15, CW 4.43���5.70, OL 4.95���7.20, OW 3.30���4.72; females (N = 10): BL 11.00���17.12, CL 4.72���6.63, CW 4.19���5.50, OL 6.88���9.75, OW 5.30���7.61. The number of promargin of cheliceral groove varies from 10 to 12 (N = 12). In addition, female genitalia show intraspecific variation: middle receptacular clusters separated from each other (Fig. 4H, I), or fused together basally (Fig. 4J); lateral ones similar to middle ones (Fig. 4H, I), or slightly larger than middle ones (Fig. 4J). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Wanzhou), China. GenBank accession number. Holotype: XUX-2012-252: OL982299., Published as part of Chen, Zhaoyang, Liu, Fengxiang, Li, Daiqin & Xu, Xin, 2022, Four new species of the primitively segmented spider genus Songthela (Mesothelae, Liphistiidae) from Chongqing Municipality, China, pp. 546-558 in Zootaxa 5091 (4) on pages 551-553, DOI: 10.11646/zootaxa.5091.4.2, http://zenodo.org/record/5863681

Genus Songthela Ono, 2000 Type species. Heptathela hangzhouensis Chen, Zhang & Zhu, 1981. Diagnosis. Males of Songthela can be distinguished from all other Heptathelinae genera by contrategulum with dense dentations (Figs 3A, 3D, 4A, 4D, 5A, 5D, 6A, 6D); by conductor with one or two apical spines, middle portion smooth or covered with several teeth (Figs 3A–E, 4A–4E, 5A–5E, 6A–E); by embolus with a wide and flat opening (Figs 3A, 3D, 4A, 4D, 5A, 5D, 6A, 6D). Songthela females differ from all other Heptathelinae genera by two pairs of receptacular clusters, median ones with obvious tubular genital stalks located on anterior margin of bursa copulatrix or slightly situated dorsally, lateral ones located relatively dorsolaterally (Figs 3H–O, 4H–M, 5H–M, 6H–M). Distribution. Southern China and northern Vietnam.

Songthela wangerbao sp. nov. Figure 6 Holotype. Male (XUX-2012-271, matured on 22 March 2013), China, Chongqing Municipality, Wanzhou City, Lishu Town, Tanjiashan, Wang���erbao Natural Reserve, 30.66��N, 108.72��E, 1172 m a.s.l., 20 November 2012, D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-263, matured on 13 June 2013) and 7 females (XUX-2012-259/260/261/262/2 65/267/268), same data as for holotype, 1149���1172 m a.s.l. Diagnosis. Males of Songthela wangerbao sp. nov. can be distinguished from S. pyriformis and S. longbao sp. nov. by conductor with narrower base and straight and slender apical spine (Fig. 6A, B, D, E), by contrategulum with an apophysis proximally and dense dentations distally (Fig. 6A, D); from S. jinyun sp. nov. by apical spine of conductor slender (Fig. 6A, B, D, E), by embolus with a distinct shrink near the opening ventrally (Fig. 6A, B, D, E), by contrategulum with dense dentations distally (Fig. 6A, D); from S. serriformis sp. nov. by conductor smooth (Fig. 6A���G); from other Songthela species by smooth conductor with one apical spine pointed to one fourth of embolus opening distally (Fig. 6A, B, D, E). Females of Songthela wangerbao sp. nov. differ from S. ciliensis by lateral receptacular clusters with shorter stalks (Fig. 6H���J); from S. jinyun sp. nov. by distance between middle receptacular clusters smaller and lateral genital stalks shorter (Fig. 6H���J); from S. longbao sp. nov. by distance between middle receptacular clusters larger, by posterior part of genital area wider (Fig. 6H���M); from S. serriformis sp. nov. by middle receptacular cluster situated on anterior margin of bursa copulatrix, posterior margin of bursa copulatrix weakly sclerotised (Fig. 6H���J); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix with distinct genital stalks, similar-sized lateral ones with obscure genital stalks, situated slight dorsolaterally (Fig. 6H���M). Description. Male holotype (XUX-2012-271). In alcohol, carapace reddish-brown with a few pointed hairs running over ocular area; opisthosoma brown, with 12 dark brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 9 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 16.18, CL 6.89, CW 6.55, OL 8.13, OW 5.70; ALE> PLE> PME> AME; leg I 22.36 (5.35 + 2.78 + 4.40 + 5.38 + 2.45), leg II 21.91 (5.95 + 2.90 + 4.56 + 6.10 + 2.40), leg III 22.63 (5.91 + 2.60 + 4.75 + 6.67 + 2.70), leg IV 29.53 (7.82 + 2.90 + 6.16 + 9.00 + 3.65). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines on the tip (Fig. 6A���C). Contrategulum with a denticulate margin and with an apophysis proximally (Fig. 6A, B, D). Marginal apophysis of tegulum and dorsal extension of terminal apophysis of tegulum with helicoid edges, with a small lamella terminal apophysis of tegulum retrolaterally (Fig. 6C, F). Conductor smooth and fused with embolus basally, gradually narrow to an apical spine (Fig. 6A, B, D, E). Embolus with a transparent, wide and flat opening (Fig. 6A, B, D, E, G). Female paratype (XUX-2012-262). In alcohol, carapace dark red-brown covered with a few hairs running over ocular area; opisthosoma slight brown with 12 dark brown and yellowish-brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 17.71, CL 7.70, CW 7.13, OL 8.73, OW 6.72; ALE> PLE> PME> AME; palp 14.50 (4.99 + 2.64 + 2.92 + 3.95), leg I 16.78 (5.65 + 3.06 + 3.10 + 3.13 + 1.84), leg II 16.01 (4.95 + 2.87 + 2.96 + 3.29 +1.94), leg III 16.60 (4.82 + 3.08 + 2.97 + 3.68 + 2.05), leg IV 23.63 (7.05 + 3.56 + 4.36 + 5.81 + 2.85). Female genitalia: Four receptacular clusters with tuber genital stalks separated from each other basally. The middle ones situated on anterior margin of bursa copulatrix. Lateral ones situated slight dorsolaterally, posterior margin of bursa copulatrix weakly sclerotised. Posterior margin of genital area wide trapezoid (Fig. 6H���M). Variation. Males and females vary in body size. The range of measurements in males as follow (N = 2): BL 15.65���16.18, CL 6.89���6.92, CW 6.55���6.68, OL 7.67���8.13, OW 5.70���6.42; females (N = 7): BL 14.56���20.88, CL 6.95���12.72, CW 6.48���8.38, OL 6.78���10.40, OW 4.90���10.42. The number of promargin of cheliceral groove varies from 9 to 11 (N = 9). In addition, male palps show intraspecific variation: contrategulum with a distinct apophysis (Fig. 6A) or not proximally (Fig. 6D). Female genitalia also show intraspecific variation: oval receptacular clusters (Fig. 6I) or irregular (Fig. 6H, J), and slender and longer middle genital stalks (Fig. 6H) or thicker and shorter middle genital stalks (Fig. 6I, J). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Wanzhou), China. GenBank accession number. Holotype: XUX-2012-271: OL982298. Remarks. The first Songthela species, S. bristowei, recorded from Wanzhou, Chongqing, was described based on one female specimen (Gertsch 1967). According to the original descriptions and illustrations (Gertsch 1967; Haupt 1983), S. bristowei has four receptacular clusters close to each other with short genital stalks, and lateral receptacular clusters larger than middle ones, all of which are different from those of the two new species, S. longbao sp. nov. and S. wangerbao sp. nov., described from Wanzhou in this study. The recorded male of S. bristowei was collected from Mt. Heng, Hengyang, Hunan Province, far away from the type locality of the species. Moreover, only two unclear illustrations were provided and lack of detailed descriptions of the male specimen (Yin et al. 1983). Therefore, we doubt that the male described from Mt. Heng by Yin et al. (1983) was S. bristowei as the female collected from Wanzhou, Chongqing according to the high endemism of liphistiids., Published as part of Chen, Zhaoyang, Liu, Fengxiang, Li, Daiqin & Xu, Xin, 2022, Four new species of the primitively segmented spider genus Songthela (Mesothelae, Liphistiidae) from Chongqing Municipality, China, pp. 546-558 in Zootaxa 5091 (4) on pages 555-556, DOI: 10.11646/zootaxa.5091.4.2, http://zenodo.org/record/5863681, {"references":["Gertsch, W. J. (1967). A new liphistiid spider from China (Araneae: Liphistiidae). Journal of The New York Entomological Society, 75, 114 - 118.","Haupt, J. (1983) Vergleichende Morphologie der Genitalorgane und Phylogenie der liphistiomorphen Webspinnen (Araneae: Mesothelae). I. Revision der bisher bekannten Arten. Zeitschrift Fur Zoologische Systematik Und Evolutionsforschung, 21, 275 - 293. http: // doi. org / 10.1111 / j. 1439 - 0469.1983. tb 00296. x","Yin, C. M., Wang, J. F. & Hu, Y. J. (1983) Essential types and the evolution of palpal organ of spiders. Journal of Hunan Teachers College (Natural Science Edition), 2, 31 - 46."]}

Songthela serriformis sp. nov. Figure 5 Holotype. Male (XUX-2012-245, matured on 13 June 2013), China, Chongqing Municipality, Nan’an District, Nanshan Botanic Garden, 29.56ºN, 106.62ºE, 554 m a.s.l., 18 November 2012, collected by D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-241A, matured on 22 March 2013) and 8 females (XUX-2012-237/238/239/240/2 41/242/243/244), same data as for holotype, 551–554 m a.s.l. Diagnosis. Males of S. serriformis sp. nov. can be distinguished from S. pluma, S. tianzhu, S. xiangnan and S. yuping by apical spine of conductor with a spinule basally, conductor forms a lamina with dentate edge proximally (Fig. 5A, B, E); from S. jinyun sp. nov., S. longbao sp. nov. and S. wangerbao sp. nov. by conductor with slender apical spine and dentate edge proximally (Fig. 5A, B, E); from other Songthela species by conductor with one apical spine and a dentate edge proximally (Fig. 5A, B, E). Females of S. serriformis sp. nov. differ from S. tianzhu by middle genital stalks longer and posterior part of genital area wider (Fig. 5H–M); from S. yuping by middle receptacular clusters lager than lateral ones and with longer genital stalks (Fig. 5H–J); from S. jinyun sp. nov., S. longbao sp. nov., S. wangerbao sp. nov. and other Songthela species by four receptacular clusters situated on dorsal wall of bursa copulatrix, and posterior margin of bursa copulatrix sclerotised (Fig. 5H–M). Description. Male holotype (XUX-2012-245). In alcohol, carapace yellowish-brown covered with a few hairs running over ocular area; opisthosoma brown with 12 dark brown tergites, close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, the fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 8 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 15.60, CL 6.95, CW 6.91, OL 8.90, OW 8.00; ALE> PLE> PME> AME; leg I 22.08 (6.11 + 2.80 + 4.55 + 5.82 + 2.80), leg II 22.79 (6.00 + 2.85 + 4.55 + 6.47 +2.92), leg III 24.88 (5.98 + 2.90 + 4.75 + 7.55 + 3.70), leg IV 31.00 (7.28 + 3.22 + 6.12 + 9.88 + 4.50). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines on tip (Fig. 5A– C). Contrategulum with a regular denticulate margin (Fig. 5A, B, D). Tegulum with a helicoid marginal apophysis and dorsal extension of terminal apophysis, a platelike terminal apophysis retrolaterally (Fig. 5C, F, G). Conductor fused with embolus basally, gradually narrowing to a long apical spine with a spinule basally, forming a lamina with a dentate edge proximally (Fig. 5A, B, E). Embolus with a transparent, wide and flat opening, a distinct shrink near the opening ventrally, with numerous ribbed ridges retrolaterally (Fig. 5A, B, D, E). Female paratype (XUX-2012-244). In alcohol, carapace dark red-brown covered with a few hairs running over ocular area; opisthosoma light brown with 12 dark brown tergites, close to each other, with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 12 denticles of variable size; legs with hairs/ spines; 8 spinnerets. Measurements: BL 14.05, CL 7.17, CW 5.91, OL 6.98, OW 5.68; ALE> PLE> PME> AME; palp 11.86 (4.10 + 1.95 + 2.53 + 3.28), leg I 13.49 (4.38 + 2.32 + 2.65 + 2.59 + 1.55), leg II 13.07 (4.27 + 2.22 + 2.47 + 2.53 +1.58), leg III 14.17 (4.05 + 2.33 + 2.51 + 3.31 + 1.97), leg IV 20.66 (5.58 + 2.87 + 3.75 + 5.53 + 2.93). Female genitalia: Four receptacular clusters situated on dorsal wall of bursa copulatrix. Middle pair with long genital stalks, larger than lateral ones. Lateral pair with obscure genital stalks situated more dorsally, posterior margin of bursa copulatrix sclerotised (Fig. 5H–M). Variation. Males and females vary in body size. The range of measurements in males as follow (N = 2): BL 13.51–15.60, CL 6.27–6.95, CW 6.12–6.91, OL 6.64–8.90, OW 5.60–8.00; females (N = 8): BL 10.19–18.12, CL 5.45–8.88, CW 5.07–7.68, OL 6.07–9.68, OW 4.26–7.05. The number of promargin of cheliceral groove varies from 8 to 12 (N = 10). There are 7 or 8 spinnerets. In addition, the conductor of males shows intraspecific variation: apical spine slightly curved with a spinule basally and forming a smooth lamina with a dentate edge proximally (Fig. 5A, B), or apical spine straight without spinule basally, and the formed lamina covered with several teeth (Fig. 5D, E). Female genitalia also show intraspecific variation: length of middle genital stalks is different (Fig. 5H–J); lateral receptacular clusters are distinctly smaller than middle ones (Fig. 5I) compared with the other samples (Fig. 5H, J), posterior margin of genital area trapezoid (Fig. 5K, L) or W-shaped (Fig. 5M). Etymology. The species epithet is an adjective from the combination of the Latin serri (serrate) and formis (shaped), and refers to the serrated margin of conductor proximally. Distribution. Chongqing (Nan’an), China. GenBank accession number. Holotype: XUX-2012-245: OL982297.

Songthela wangerbao sp. nov. Figure 6 Holotype. Male (XUX-2012-271, matured on 22 March 2013), China, Chongqing Municipality, Wanzhou City, Lishu Town, Tanjiashan, Wang’erbao Natural Reserve, 30.66ºN, 108.72ºE, 1172 m a.s.l., 20 November 2012, D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-263, matured on 13 June 2013) and 7 females (XUX-2012-259/260/261/262/2 65/267/268), same data as for holotype, 1149–1172 m a.s.l. Diagnosis. Males of Songthela wangerbao sp. nov. can be distinguished from S. pyriformis and S. longbao sp. nov. by conductor with narrower base and straight and slender apical spine (Fig. 6A, B, D, E), by contrategulum with an apophysis proximally and dense dentations distally (Fig. 6A, D); from S. jinyun sp. nov. by apical spine of conductor slender (Fig. 6A, B, D, E), by embolus with a distinct shrink near the opening ventrally (Fig. 6A, B, D, E), by contrategulum with dense dentations distally (Fig. 6A, D); from S. serriformis sp. nov. by conductor smooth (Fig. 6A–G); from other Songthela species by smooth conductor with one apical spine pointed to one fourth of embolus opening distally (Fig. 6A, B, D, E). Females of Songthela wangerbao sp. nov. differ from S. ciliensis by lateral receptacular clusters with shorter stalks (Fig. 6H–J); from S. jinyun sp. nov. by distance between middle receptacular clusters smaller and lateral genital stalks shorter (Fig. 6H–J); from S. longbao sp. nov. by distance between middle receptacular clusters larger, by posterior part of genital area wider (Fig. 6H–M); from S. serriformis sp. nov. by middle receptacular cluster situated on anterior margin of bursa copulatrix, posterior margin of bursa copulatrix weakly sclerotised (Fig. 6H–J); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix with distinct genital stalks, similar-sized lateral ones with obscure genital stalks, situated slight dorsolaterally (Fig. 6H–M). Description. Male holotype (XUX-2012-271). In alcohol, carapace reddish-brown with a few pointed hairs running over ocular area; opisthosoma brown, with 12 dark brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 9 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 16.18, CL 6.89, CW 6.55, OL 8.13, OW 5.70; ALE> PLE> PME> AME; leg I 22.36 (5.35 + 2.78 + 4.40 + 5.38 + 2.45), leg II 21.91 (5.95 + 2.90 + 4.56 + 6.10 + 2.40), leg III 22.63 (5.91 + 2.60 + 4.75 + 6.67 + 2.70), leg IV 29.53 (7.82 + 2.90 + 6.16 + 9.00 + 3.65). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines on the tip (Fig. 6A–C). Contrategulum with a denticulate margin and with an apophysis proximally (Fig. 6A, B, D). Marginal apophysis of tegulum and dorsal extension of terminal apophysis of tegulum with helicoid edges, with a small lamella terminal apophysis of tegulum retrolaterally (Fig. 6C, F). Conductor smooth and fused with embolus basally, gradually narrow to an apical spine (Fig. 6A, B, D, E). Embolus with a transparent, wide and flat opening (Fig. 6A, B, D, E, G). Female paratype (XUX-2012-262). In alcohol, carapace dark red-brown covered with a few hairs running over ocular area; opisthosoma slight brown with 12 dark brown and yellowish-brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than others, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with hairs/spines; 7 spinnerets. Measurements: BL 17.71, CL 7.70, CW 7.13, OL 8.73, OW 6.72; ALE> PLE> PME> AME; palp 14.50 (4.99 + 2.64 + 2.92 + 3.95), leg I 16.78 (5.65 + 3.06 + 3.10 + 3.13 + 1.84), leg II 16.01 (4.95 + 2.87 + 2.96 + 3.29 +1.94), leg III 16.60 (4.82 + 3.08 + 2.97 + 3.68 + 2.05), leg IV 23.63 (7.05 + 3.56 + 4.36 + 5.81 + 2.85). Female genitalia: Four receptacular clusters with tuber genital stalks separated from each other basally. The middle ones situated on anterior margin of bursa copulatrix. Lateral ones situated slight dorsolaterally, posterior margin of bursa copulatrix weakly sclerotised. Posterior margin of genital area wide trapezoid (Fig. 6H–M). Variation. Males and females vary in body size. The range of measurements in males as follow (N = 2): BL 15.65–16.18, CL 6.89–6.92, CW 6.55–6.68, OL 7.67–8.13, OW 5.70–6.42; females (N = 7): BL 14.56–20.88, CL 6.95–12.72, CW 6.48–8.38, OL 6.78–10.40, OW 4.90–10.42. The number of promargin of cheliceral groove varies from 9 to 11 (N = 9). In addition, male palps show intraspecific variation: contrategulum with a distinct apophysis (Fig. 6A) or not proximally (Fig. 6D). Female genitalia also show intraspecific variation: oval receptacular clusters (Fig. 6I) or irregular (Fig. 6H, J), and slender and longer middle genital stalks (Fig. 6H) or thicker and shorter middle genital stalks (Fig. 6I, J). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Wanzhou), China. GenBank accession number. Holotype: XUX-2012-271: OL982298. Remarks. The first Songthela species, S. bristowei, recorded from Wanzhou, Chongqing, was described based on one female specimen (Gertsch 1967). According to the original descriptions and illustrations (Gertsch 1967; Haupt 1983), S. bristowei has four receptacular clusters close to each other with short genital stalks, and lateral receptacular clusters larger than middle ones, all of which are different from those of the two new species, S. longbao sp. nov. and S. wangerbao sp. nov., described from Wanzhou in this study. The recorded male of S. bristowei was collected from Mt. Heng, Hengyang, Hunan Province, far away from the type locality of the species. Moreover, only two unclear illustrations were provided and lack of detailed descriptions of the male specimen (Yin et al. 1983). Therefore, we doubt that the male described from Mt. Heng by Yin et al. (1983) was S. bristowei as the female collected from Wanzhou, Chongqing according to the high endemism of liphistiids.

Songthela longbao sp. nov. Figure 4 Holotype. Male (XUX-2012-252, matured on 18 July 2013), China, Chongqing Municipality, Wanzhou City, Mt. Longbao, 30.76ºN, 108.38ºE, 226 m a.s.l., 18 November 2012, collected by D Li, FX Liu, X Xu. Paratypes. 1 male (XUX-2012-251A) and 10 females (XUX-2012-247/249/250/251/253/254A/255/256/257/2 58), same data as for holotype, 221–226 m a.s.l. Diagnosis. Males of S. longbao sp. nov. can be distinguished from S. ciliensis by conductor with a curved margin retrolaterally (Fig. 4B, E); from S. pyriformis by apical spine of conductor straight and slender (Fig. 4A, B, D, E), by marginal apophysis of tegulum semielliptic (Fig. 4C, G); from S. jinyun sp. nov. and S. wangerbao sp. nov. by apical spine of conductor slightly shorter and thicker (Fig. 4A, B), by marginal apophysis of tegulum helicoid (Fig. 4C, F); from S. serriformis sp. nov. by conductor with smooth margin (Fig. 4A–G); from other Songthela species by smooth conductor with one apical spine (Fig. 4A, B, E). Females of S. longbao sp. nov. differ from S. ciliensis by lateral genital stalks shorter (Fig. 4H–J); from S. jinyun sp. nov. by distance between bases of middle and lateral genital stalks larger (Fig. 4H–J); from S. wangerbao sp. nov. by middle genital stalks longer and close to each other (Fig. 4H–J); from S. bristowei by bases of four receptacular clusters separated from each other (Fig. 4H–J); from S. serriformis sp. nov. by middle receptacular clusters situated on anterior margin of bursa copulatrix, lateral receptacular clusters similar to or slightly larger than middle ones, posterior margin of bursa copulatrix weakly sclerotised (Fig. 4H–J); from other Songthela species by middle receptacular clusters situated on anterior margin of bursa copulatrix with long genital stalks, similar or smaller than lateral pair, lateral ones situated dorsolaterally (Fig. 4H–M). Description. Male holotype (XUX-2012-252). In alcohol, carapace reddish brown with a few pointed hairs running over ocular area; opisthosoma pale yellow, with 12 dark reddish brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with sturdy hairs/spines; 7 spinnerets. Measurements: BL 14.08, CL 6.15, CW 5.70, OL 7.20, OW 4.72; ALE> PLE> PME> AME; leg I 17.02 (4.88 + 1.97 + 3.52 + 4.20 + 2.45), leg II 17.58 (4.78 + 2.30 + 3.35 + 4.60 + 2.55), leg III 19.73 (5.00 + 2.23 + 3.55 + 5.85 + 3.10), leg IV 24.65 (6.20 + 2.60 + 4.60 + 7.20 + 4.05). Palp: Paracymbium unpigmented and unsclerotized prolaterally, with numerous setae and spines distally (Fig. 4A–C). Contrategulum with denticulate margin, dentations sparse proximally and dense distally (Fig. 4A, D). Marginal apophysis of tegulum and dorsal extension of terminal apophysis of tegulum with helicoid edges, with a small terminal apophysis of tegulum (Fig. 4C, F, G). Conductor smooth, fused with embolus basally, with one apical spine (Fig. 4A, B, D, E). Embolus with a transparent, wide and flat opening and several ribbed ridges distally (Fig. 4A, B, D, E, G). Female paratype (XUX-2012-251). In alcohol, carapace slight red-brown covered with a few hairs running over ocular area; opisthosoma slight brown with 12 dark brown tergites, close to each other with two pointed hairs on tergites from second to fourth, second to sixth larger than remaining, fourth largest; sternum narrow, much longer than wide; chelicerae robust with promargin of cheliceral groove with 11 denticles of variable size; legs with hairs/ spines; 7 spinnerets. Measurements: BL 13.92, CL 5.25, CW 5.18, OL 7.32, OW 6.25; ALE> PLE> PME> AME; palp 8.60 (3.15 + 1.15 + 1.89 + 2.41), leg I 10.66 (3.40 + 1.90 + 2.07 + 1.98 + 1.31), leg II 10.50 (3.20 + 1.88 + 1.85 + 2.22 +1.35), leg III 11.00 (3.06 + 1.83 + 1.83 + 2.68 + 1.60), leg IV 16.74 (4.68 + 2.25 + 3.06 + 4.45 + 2.30). Female genitalia: Four receptacular clusters separated from each other basally; median pair situated on anterior margin of bursa copulatrix with long genital stalks, similar to or slightly smaller than lateral pair; lateral ones situated dorsolaterally; posterior margin of bursa copulatrix weakly sclerotised (Fig. 4H–M). Variation. Males and females vary in body size. The range of measurements in males (N = 2): BL 10.37–14.08, CL 5.04–6.15, CW 4.43–5.70, OL 4.95–7.20, OW 3.30–4.72; females (N = 10): BL 11.00–17.12, CL 4.72–6.63, CW 4.19–5.50, OL 6.88–9.75, OW 5.30–7.61. The number of promargin of cheliceral groove varies from 10 to 12 (N = 12). In addition, female genitalia show intraspecific variation: middle receptacular clusters separated from each other (Fig. 4H, I), or fused together basally (Fig. 4J); lateral ones similar to middle ones (Fig. 4H, I), or slightly larger than middle ones (Fig. 4J). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Chongqing (Wanzhou), China. GenBank accession number. Holotype: XUX-2012-252: OL982299.

Genus Songthela Ono, 2000 Type species. Heptathela hangzhouensis Chen, Zhang & Zhu, 1981. Diagnosis. Males of Songthela can be distinguished from all other Heptathelinae genera by contrategulum with dense dentations (Figs 3A, 3D, 4A, 4D, 5A, 5D, 6A, 6D); by conductor with one or two apical spines, middle portion smooth or covered with several teeth (Figs 3A���E, 4A���4E, 5A���5E, 6A���E); by embolus with a wide and flat opening (Figs 3A, 3D, 4A, 4D, 5A, 5D, 6A, 6D). Songthela females differ from all other Heptathelinae genera by two pairs of receptacular clusters, median ones with obvious tubular genital stalks located on anterior margin of bursa copulatrix or slightly situated dorsally, lateral ones located relatively dorsolaterally (Figs 3H���O, 4H���M, 5H���M, 6H���M). Distribution. Southern China and northern Vietnam., Published as part of Chen, Zhaoyang, Liu, Fengxiang, Li, Daiqin & Xu, Xin, 2022, Four new species of the primitively segmented spider genus Songthela (Mesothelae, Liphistiidae) from Chongqing Municipality, China, pp. 546-558 in Zootaxa 5091 (4) on page 547, DOI: 10.11646/zootaxa.5091.4.2, http://zenodo.org/record/5863681, {"references":["Ono, H. (2000) Zoogeographic and taxonomic notes on spiders of the subfamily Heptathelinae (Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum Tokyo, Series A, Zoology, 33, 145 - 151."]}

Sinothela Haupt, 2003, nomen dubium Sinothela Haupt, 2003: 71, type species by original designation, Heptathela sinensis Crosby & Bishop; synonymized with Heptathela in Schwendinger & Ono, 2011: 601; Xu et al. 2015a: 139., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on page 152, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Haupt, J. (2003) The Mesothelae - a monograph of an exceptional group of spiders (Araneae: Mesothelae) (Morphology, behaviour, ecology, taxonomy, distribution and phylogeny). Zoologica, 154, 1 - 102.","Schwendinger, P. J. & Ono, H. (2011) On two Heptathela species from southern Vietnam, with a discussion of copulatory organs and systematics of the Liphistiidae (Araneae: Mesothelae). Revue suisse de Zoologie, 118, 599 - 637. https: // doi. org / 10.5962 / bhl. part. 117818","Xu, X., Liu, F., Chen, J., Ono, H., Li, D. & Kuntner, M. (2015 a) A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). Zookeys, 488, 121 - 151. https: // doi. org / 10.3897 / zookeys. 488.8726"]}

Sinothela sinensis (Crosby & Bishop, 1932), nomen dubium Sinothela sinensis: Haupt, 2003: 71. Heptathela sinensis Bishop & Crosby 1932: 5 (holotype female, from Shandong Christian University, Jinan City, Shandong Province, collected by A. Jacot in May 1930, deposited at the American Museum of Natural History, New York, not examined due to unable to loan or image the holotype from museum under the situation of COVID pandemic); Bristowe, 1933: 1055; Wen & Zhu, 1980: 39 (male description without the detail information on collection site and where it was deposited; presumed to be lost); Haupt 1983: 285; Hu, 1984: 26; Song & Haupt 1984: 445 (drawings of the male palp based on the specimen collected from Xingyang, Zhengzhou City, Henan Province, deposited in IZCAS, examined; Zhu & Shi, 1985: 48; Zhang, 1987: 44; Zhao, 1993: 53; Song, Zhu & Chen, 1999: 33; Song, Zhu & Chen, 2001: 46. Abcathela sinensis: Ono, 2000: 150. Sinothela sinensis: Haupt, 2003: 71; Xu et al., 2015a: 139. Songthela sinensis: Zhu & Zhang, 2011: 30. For the detailed description of holotype, see Bishop & Crosby 1932; for the illustration of holotype female and secondarily associated non-type male genital anatomy, see Haupt 1983 and S ong & Haupt 1984. Remarks. We were unable to redescribe the species, Sinothela sinensis, from the holotype female for a few reasons. We were unable examine the holotype female (lodged at the American Museum of National History). The type locality (Shandong Christian University) of the species no longer exists. While our collecting efforts in the type vicinity (Jinan, Shandong Province) did produce a single female specimen, the exact identity of the female specimen is doubtful without examining the holotype and due to highly intraspecific variation in female genital morphology. For the males collected from Huolu, Hebei Province (Wen & Zhu 1980) and from Xingyang, Zhengzhou City, Henan Province (Song & Haupt 1984), we doubt that they were corrected diagnosed as the species. Thus, Sinothela sinensis is here considered a nomen dubium., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 152-153, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Bishop, S. C. & Crosby, C. R. (1932) A new species of the spider family Liphistiidae from China. Peking Natural History Bulletin, 6, 5 - 7.","Haupt, J. (2003) The Mesothelae - a monograph of an exceptional group of spiders (Araneae: Mesothelae) (Morphology, behaviour, ecology, taxonomy, distribution and phylogeny). Zoologica, 154, 1 - 102.","Wen, Z. G. & Zhu, C. D. (1980) Four species of male spiders (Araneae) from China. Journal of the Bethune Medical University, 6, 39 - 43.","Haupt, J. (1983) Vergleichende Morphologie der Genitalorgane und Phylogenie der liphistiomorphen Webspinnen (Araneae: Mesothelae). I. Revision der bisher bekannten Arten. Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 21 (4), 275 - 293. https: // doi. org / 10.1111 / j. 1439 - 0469.1983. tb 00296. x","Song, D. X. & Haupt, J. (1984) Comparative morphology and phylogeny of Liphistiomorph spiders (Araneae: Mesothelae). 2. Revision of new Chinese Heptathelid species. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, 27, 443 - 451.","Ono, H. (2000) Zoogeographic and taxonomic notes on spiders of the subfamily Heptathelinae (Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum, 33, 145 - 151.","Xu, X., Liu, F., Chen, J., Ono, H., Li, D. & Kuntner, M. (2015 a) A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). Zookeys, 488, 121 - 151. https: // doi. org / 10.3897 / zookeys. 488.8726"]}

Luthela taian sp. nov. (Figs 12A���G, 13A���L) Holotype. Male (XUX-2014-143A, matured on 6 September 2014 at CBEE), Shandong Agricultural University, Taian City, Shandong Province, China, 36.155��N, 117.156��E, 131 m, collected on 17 July 2014, by F.X. Liu, C. Xu, and Z. T. Zhang, CBEE. Paratypes. 13 females (XUX-2014-133/134/135/136/137/138/140/141/141A/143/144/144A/144B/144C) and 3 males (XUX-2014-139, matured on 20 September 2014, XUX-2014-142 matured on 26 June 2015, XUX-2014- 142A matured on 24 August 2014 at CBEE), same data as for holotype; 2 females (XUX-2014-151/154) Confucius Cemetery, Qufu city, Shandong Province, China, 35.618��N, 116.991��E, 80 m, collected on 18 July 2014, by F.X. Liu, C. Xu, and Z. T. Zhang. All in CBEE. Diagnosis. Males of L. taian sp. nov. can be distinguished from those of L. badong sp. nov. by the conductor having two longer and thicker spines (Fig. 12A, 12G); from those of L. dengfeng sp. nov. by the conductor with longer lower spines (Fig. 12G); from the other Luthela species by the conductor lacking the middle spine (Fig. 12D��� G). Females of L. taian sp. nov. differ from those of L. badong sp. nov., L. luotianensis, and L. yuncheng sp. nov. by the middle receptacular clusters smaller than the lateral ones (Fig. 13A���C, 13G���I); from those of L. schensiensis and L. yiyuan sp. nov. by the middle receptacular clusters with slender and longer stalks (Fig. 13A���C); from those of L. handan sp. nov. by the middle receptacular clusters being separated from each other (Fig. 13A). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; a few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 8���9 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 12.50, CL 6.80, CW 7.30, OL 6.00, OW 5.60; ALE> PLE> PME> AME; leg I 18.00 (5.10 + 2.40 + 3.50 + 4.50 + 2.60), leg II 17.70 (4.60 + 2.25 + 3.25 + 4.60 + 3.00), leg III 19.00 (4.20 + 2.30 + 3.40 + 5.60 + 3.50), leg IV 25.90 (5.90 + 2.80 + 4.70 + 8.00 + 4.50). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 12A���C). Contrategulum with 7���10 teeth along margin, and distal tooth with 4 serrations (Fig. 12A, 12D). Marginal apophysis of tegulum with several serrations proximally and smooth distally, dorsal extension of terminal apophysis of tegulum with irregular serrations, and terminal apophysis smooth, triangle-shaped (Fig. 12E���G). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum (Fig. 12G). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 12A, 12G). Female (XUX-2014-136). Carapace and opisthosoma of the female similar to the male except for lighter colouration than the male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 15.00, CL 6.85, CW 5.70, OL 7.10, OW 4.90; ALE> PLE> PME> AME; palp 11.35 (3.75 + 1.95 + 2.45 + 3.20), leg I 13.40 (4.30 + 2.30 + 2.60 + 2.70 + 1.50), leg II 12.70 (3.85 + 2.20 + 2.25 + 2.80 + 1.60), leg III 13.45 (3.65 + 2.30 + 2.35 + 3.20 + 1.95), leg IV 19.05 (4.25 + 2.70 + 3.60 + 5.60 + 2.90). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters, middle ones smaller than lateral ones with stalks, separated from each other basally, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 13). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shandong Province (Taishan, Qufu) Remarks. Males and females vary in body size: males (N = 4): BL 12.50���16.60, CL 6.80���7.10, CW 6.50���7.30, OL 6.00���8.60, OW 5.60���6.00; females (N = 15): BL 12.50���20.50, CL 6.00���8.60, CW 4.50���7.20, OL 6.40���10.20, OW 4.80���8.35. There was intraspecific variation in female genital morphology and molecular markers. For the female genitalia, the middle receptacular clusters were weakly sclerotized with long stalks (Fig. 13A, 13B), or well sclerotized (Fig. 13C���I). Intraspecific genetic distance is between 0% and 0.6% based on the K2P substitution model and COI nucleotide sequences (N = 9). The GenBank accession code of the holotype (XUX-2014-143A) is MH172722., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 149-150, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635

Luthela schensiensis (Schenkel, 1953), comb. nov. (Figs 10A���K, 11A���P) Liphistius sinensis schensiensis Schenkel,1953: 1 (the subspecies was described based on one female collected from Tongyuanfang, Gaoling, Shaanxi Province, 8 October 1916, deposited at Museum Hoangho-Peiho in Tientsin [corresponding to Huanghebaihe in Tianjin and Yellow River-Haihe River in Tianjin, in Chinese and English, respectively], lost). Liphistius schensiensis: Gertsch, 1967: 115. Heptathela sinensis schensiensis: Haupt, 1983: 285. Heptathela xianensis: Zhu & Wang, 1983: 131; Wang & Zhu, 1984: 403. Heptathela schensiensis: Platnick & Sedgwick, 1984: 4; Song & Haupt, 1984: 447; Song, Zhu & Chen, 1999: 33. Liphistius heyangensis Zhu & Wang, 1984: 251 (holotype: female, from Heyang, Shaanxi Province, 30 June 1982, deposited at Bethune Medical University, now College of Life Sciences, Jilin University, type presumed to be lost). Syn. nov. Abcathela schensiensis: Ono, 2000: 150. Sinothela heyangensi s: Haupt, 2003: 71. Sinothela schensiensis: Haupt, 2003: 71; Xu et al., 2015a: 139. Heptathela heyangensis: Platnick, 1989: 57. Abcathela heyangensis: Ono, 2000: 150. Material examined. 3 males (XUX-2011-241/242/247, matured on 10 July 2012 at CBEE) and 7 females (XUX- 2011-243/245/248/249/251/253/254), Kangjiapo Village, Heyang County, Shaanxi Province, China, 35.242��N, 110.155E, 730 m, collected on 1 December, 2011; 3 males (XUX-2011-255/264 matured on 10 July 2012, XUX- 2011-263 matured on 13 June 2013, at CBEE) and 10 females (XUX-2011-256���262/265/266/269), Quyang Village, Tongyuan, Gaoling County, Shaanxi Province, China, 35.546��N, 109.040E, 390 m, collected on 2 December, 2011; 1 male (XUX-2011-273, matured on 10 July 2012 at CBEE), 34.493��N, 108.806E, 374 m, and 1 female (XUX- 2011-270), 34.498��N, 108.786E, 359 m, Taiping village, Jingyang County, Xianyang City, Shaanxi Province, China, collected on 3 December, 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu. All in CBEE. Diagnosis. Males of L. schensiensis resemble those of L. yiyuan sp. nov. and L. yuncheng sp. nov., but can be distinguished from those of L. yiyuan sp. nov. by the tegulum having a slightly smaller terminal apophysis (Fig. 10F, 10G), and the embolus with more longitudinal ribs retrolaterally (Fig. 10G, 10K); from those of L. yuncheng sp. nov. by the marginal apophysis and the dorsal extension of the terminal apophysis of the tegulum with irregular serrations (Fig. 10G, 10K); from those of L. dengfeng sp. nov. by the conductor with a longer lower spine (Fig. 10G, 10K); from those of L. handan sp. nov. by the contrategulum margin with slightly smaller teeth, and the dorsal extension of the terminal apophysis of the tegulum with irregular serrations (Fig. 10G, 10K); and from those of L. badong sp. n and L. taian sp. nov. by the conductor with small middle spines (Fig. 10A���E, 10H, 10K). Females of L. schensiensis differ from those of L. badong sp. nov. by the middle pair of receptacular clusters with slightly thicker and shorter stalks; from those of L. yiyuan sp. nov. by the middle pair of receptacular clusters with slightly longer stalks (Fig. 11A���D, 11I���L); from those of L. handan sp. nov., by the middle pair of receptacular clusters with slightly thicker stalks (Fig. 11A���D); from those of the other Luthela species by the bases of the middle stalks being close to each other (Fig. 11A���D). Redescription. Male (XUX-2011-273). Carapace reddish brown; opisthosoma brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 11 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, second to fifth larger than others and the others small; 8 spinnerets. Measurements: BL 14.95, CL 7.05, CW 6.60, OL 7.15, OW 4.90; ALE> PLE> PME> AME; leg I 20.65 (6.05 + 2.75 + 4.15 + 5.05 + 2.65), leg II 20.85 (5.95 + 2.65 + 3.95 + 5.25 + 3.05), leg III 21.30 (5.65 + 2.55 + 3.75 + 6.40 + 2.95), leg IV 29.00 (7.20 + 2.95 + 5.35 + 8.95 + 4.55). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 10A���C). Contrategulum with 7���10 teeth along margin, and distal tooth serrated (Fig. 10A, 10D, 10H, 10J). Tegulum with an irregular dentate marginal apophysis and an irregular dorsal extension of terminal apophysis, as well as smooth terminal apophysis (Fig. 10C, 10F, 10G, 10I���K). Conductor situated ventro-proximally on embolus, with two long spines, upper one parallel to the furrow of embolus distally, lower one pointed to contrategulum, and one or two small teeth situated at between two long spines (Fig. 10A���C, 10D, 10H, 10K). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 10H, 10K). Female (XUX-2011-243). Carapace and opisthosoma similar to male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 8 spinnerets. Measurements: BL 25.25, CL 10.35, CW 9.25, OL 14.75, OW 12.75; ALE> PLE> PME> AME; palp 16.30 (5.70 + 2.75 + 3.35 + 4.50), leg I 17.45 (4.85 + 3.05 + 3.55 + 3.75 + 2.25), leg II 19.15 (6.15 + 3.05 + 3.55 + 3.95 + 2.45), leg III 19.85 (6.05 + 3.15 + 3.45 + 4.45 + 2.75), leg IV 28.80 (7.65 + 3.75 + 5.35 + 8.00 + 4.05). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters, middle ones slightly smaller than lateral ones with stalks, close to each other, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 11A���P). Distribution. Shaanxi Province (Heyang, Gaoling, Jingyang) Remarks. Males and females vary in body size: males (N = 7): BL 14.95���18.55, CL 6.75���7.85, CW 6.25���7.30, OL 7.15���10.45, OW 4.90���7.55; females (N = 18): BL 17.85���25.65, CL 7.55���10.25, CW 6.90���9.10, OL 9.05���16.65, OW 5.90���13.25. There is intraspecific variation in male and female genital morphology and molecular markers. Specifically, for the male genitalia, there was considerable intraspecific variation in the number of the small middle spines of the conductor, either one (Fig. 10A) or two spines (Fig. 10D, 10K); in the position of the small spine, either close to the upper spine (Fig. 10K), close to the lower spine (Fig. 10D), or located at the middle of two long spines (Fig. 10A); in the tip shape of lower spine of the conductor, bifurcated (Fig. 10D���F) or not (Fig. 10A, 10H). Intraspecific variation was noted in the number of teeth (7���10) along the contrategulum margin, and the different shape of the serrated distal tooth (Fig. 10A, 10D, 10H, 10J). For intraspecific variation in female genitalia, the posterior margin of the genital sternite may be incurved (Fig. 11C, 11D, 11I���K) or straight (Fig. 11A, 11L), the middle receptacular clusters were situated at the anterior margin of the bursa copulatrix (Fig. 11A���J, 11L), or at slightly dorsal part of the bursa copulatrix (Fig. 11K, 11O). Intraspecific genetic distance ranges from 0% to 1.8% based on the K2P substitution model and COI nucleotide sequences (N = 18). We treated L. heyangensis as a junior synonym of L. schensiensis for several reasons. First, L. heyangensis was diagnosed and described based exclusively on one female (Zhu & Wang 1984). As liphistiid females exhibit considerable intraspecific variation in female genitalia (Xu et al. 2017, 2019), it is difficult to diagnose the species based exclusively on female characters of a single specimen. Second, we collected intensively from the type localities of both L. heyangensis and L. schensiensis, and we examined and compared the newly collected males and females with the original descriptions and illustrations of both L. heyangensis and L. schensiensis (Schenkel 1953; Song & Haupt 1984; Zhu & Wang 1984). All lines of evidence support the type female of L. heyangensis belonging to L. schensiensis. Third, there is very low genetic variation (1.8%) among the newly collected specimens from the type localities of both L. heyangensis and L. schensiensis. Thus, the specimens collected from Heyang County, Shaanxi Province should be considered the same species as those collected from both Gaoling and Jingyang Counties, Shaanxi Province. The GenBank accession code of XUX-2011-273 is MH172701., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 145-148, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Schenkel, E. (1953) Chinesische Arachnoidea aus dem Museum Hoangho-Peiho in Tientsin. Boletim do Museu Nacional do Rio de Janeiro, 119, 1 - 108.","Haupt, J. (1983) Vergleichende Morphologie der Genitalorgane und Phylogenie der liphistiomorphen Webspinnen (Araneae: Mesothelae). I. Revision der bisher bekannten Arten. Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 21 (4), 275 - 293. https: // doi. org / 10.1111 / j. 1439 - 0469.1983. tb 00296. x","Zhu, C. D. & Wang, Y. W. (1984) A new species of genus Liphistius (Araneae: Liphistiidae). Journal of the Bethune Medical University, 10, 251 - 253.","Song, D. X. & Haupt, J. (1984) Comparative morphology and phylogeny of Liphistiomorph spiders (Araneae: Mesothelae). 2. Revision of new Chinese Heptathelid species. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, 27, 443 - 451.","Ono, H. (2000) Zoogeographic and taxonomic notes on spiders of the subfamily Heptathelinae (Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum, 33, 145 - 151.","Haupt, J. (2003) The Mesothelae - a monograph of an exceptional group of spiders (Araneae: Mesothelae) (Morphology, behaviour, ecology, taxonomy, distribution and phylogeny). Zoologica, 154, 1 - 102.","Xu, X., Liu, F., Chen, J., Ono, H., Li, D. & Kuntner, M. (2015 a) A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). Zookeys, 488, 121 - 151. https: // doi. org / 10.3897 / zookeys. 488.8726","Xu, X., Liu, F., Ono, H., Chen, J., Kuntner, M. & Li, D. (2017) Targeted sampling in Ryukyus facilitates species delimitation of the primitively segmented spider genus Ryuthela (Araneae: Mesothelae: Liphistiidae). Zoological Journal of the Linnean Society, 181, 1 - 43. https: // doi. org / 10.1093 / zoolinnean / zlx 024","Xu, X., Ono, H., Kuntner, M., Liu, F. & Li, D. (2019) A taxonomic monograph of the liphistiid spider genus Heptathela, endemic to Japanese islands. Zookeys, 888, 1 - 50. https: // doi. org / 10.3897 / zookeys. 888.34494"]}

Luthela luotianensis (Yin et al., 2002) comb. nov. (Fig. 9A���D) Heptathela luotianensis Yin et al., 2002: 18 (holotype: female, from Luotian, Hubei Province, China, 3048���N, 11524���E, collected by J.Z. Zhao and F.X. Liu on 22���23 October 1978, deposited at College of Life Sciences, Hubei University, type presumed to be lost). Sinothela luotianensis: Xu et al., 2015a: 139. Material examined. 3 females (XUX-2012-079/080/082), Fengyi���ao Chinese chestnut plantation, Luotian County, Hubei Province, China, 30.750N, 115.397E, 123 m, collected on 8 July, 2012, by F.X. Liu, X.Y. Wang, and X. Xu. Diagnosis. Females of L. luotianensis can be distinguished from those of L. badong sp. nov. by the bases of the middle stalks being separated from each other (Fig. 9A���D); from those of L. dengfeng sp. nov., L. handan sp. nov., and L. taian sp. nov. by the similar-sized receptacular clusters (Fig. 9A, 9C); from those of L. schensiensis and L. yiyuan sp. nov. by the bases of middle stalks separated from each other and slightly slender (Fig. 9A���D); from those of L. yuncheng sp. nov. by obviously incurved posterior margin of genital sternite (Fig. 9B, 9D). Male. Unknown. Redescription. Female (XUX-2012-079). Carapace reddish brown; opisthosoma brown, with brown tergites; sternum narrow, much longer than wide; a few long pointed hairs running over ocular mound in a longitudinal row; chelicerae robust with promargin of groove bearing 9���10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 12.95, CL 5.95, CW 5.20, OL 6.10, OW 4.85; ALE> PLE> PME> AME; palp 12.70 (4.15 + 2.15 + 2.65 + 3.75), leg I 15.60 (5.05 + 2.65 + 3.05 + 3.05 + 1.80), leg II 15.65 (4.85 + 2.75 + 2.80 + 3.40 + 1.85), leg III 15.85 (4.65 + 2.70 + 2.70 + 3.75 + 2.05), leg IV 23.00 (6.25 + 3.35 + 4.00 + 6.15 + 3.25). Genitalia: Posterior margin of genital sternite obviously incurved (Fig. 9B, 9D), two pairs of receptacular clusters, middle ones situated at anterior margin of bursa copulatrix with slender stalks, separated from each other basally, lateral ones situated at slightly dorsolateral position of bursa copulatrix (Fig. 9A���D). Distribution. Hubei Province (Luotian) Remarks. Females vary in body size: (N = 3): BL 12.95���17.45, CL 5.95���7.55, CW 5.20���7.55, OL 6.10���8.85, OW 4.65���7.05. Since the holotype was lost, we identified the newly collected specimens as L. luotianensis because: (1) the new specimens were collected at the type locality; (2) we compared the newly collected specimens with the original descriptions and illustrations provided by Yin et al. (2002), and found that the body size and the receptacular clusters of newly collected specimens are similar to those of the holotype. Intraspecific genetic distance ranges from 0% to 0.4% based on the K2P substitution model and COI nucleotide sequences (N = 7, including juveniles). The GenBank accession code of XUX-2012-079 is KP229881., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 143-145, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Yin, C. M., Tang, G., Zhao, J. Z. & Chen, J. (2002) Two new species of the genus Heptathela from China (Araneae: Liphistiidae). Acta Arachnologica Sinica, 11, 18 - 21.","Xu, X., Liu, F., Chen, J., Ono, H., Li, D. & Kuntner, M. (2015 a) A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). Zookeys, 488, 121 - 151. https: // doi. org / 10.3897 / zookeys. 488.8726"]}

Luthela taian sp. nov. (Figs 12A–G, 13A–L) Holotype. Male (XUX-2014-143A, matured on 6 September 2014 at CBEE), Shandong Agricultural University, Taian City, Shandong Province, China, 36.155°N, 117.156°E, 131 m, collected on 17 July 2014, by F.X. Liu, C. Xu, and Z. T. Zhang, CBEE. Paratypes. 13 females (XUX-2014-133/134/135/136/137/138/140/141/141A/143/144/144A/144B/144C) and 3 males (XUX-2014-139, matured on 20 September 2014, XUX-2014-142 matured on 26 June 2015, XUX-2014- 142A matured on 24 August 2014 at CBEE), same data as for holotype; 2 females (XUX-2014-151/154) Confucius Cemetery, Qufu city, Shandong Province, China, 35.618°N, 116.991°E, 80 m, collected on 18 July 2014, by F.X. Liu, C. Xu, and Z. T. Zhang. All in CBEE. Diagnosis. Males of L. taian sp. nov. can be distinguished from those of L. badong sp. nov. by the conductor having two longer and thicker spines (Fig. 12A, 12G); from those of L. dengfeng sp. nov. by the conductor with longer lower spines (Fig. 12G); from the other Luthela species by the conductor lacking the middle spine (Fig. 12D– G). Females of L. taian sp. nov. differ from those of L. badong sp. nov., L. luotianensis, and L. yuncheng sp. nov. by the middle receptacular clusters smaller than the lateral ones (Fig. 13A–C, 13G–I); from those of L. schensiensis and L. yiyuan sp. nov. by the middle receptacular clusters with slender and longer stalks (Fig. 13A–C); from those of L. handan sp. nov. by the middle receptacular clusters being separated from each other (Fig. 13A). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; a few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 8–9 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 12.50, CL 6.80, CW 7.30, OL 6.00, OW 5.60; ALE> PLE> PME> AME; leg I 18.00 (5.10 + 2.40 + 3.50 + 4.50 + 2.60), leg II 17.70 (4.60 + 2.25 + 3.25 + 4.60 + 3.00), leg III 19.00 (4.20 + 2.30 + 3.40 + 5.60 + 3.50), leg IV 25.90 (5.90 + 2.80 + 4.70 + 8.00 + 4.50). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 12A–C). Contrategulum with 7–10 teeth along margin, and distal tooth with 4 serrations (Fig. 12A, 12D). Marginal apophysis of tegulum with several serrations proximally and smooth distally, dorsal extension of terminal apophysis of tegulum with irregular serrations, and terminal apophysis smooth, triangle-shaped (Fig. 12E–G). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum (Fig. 12G). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 12A, 12G). Female (XUX-2014-136). Carapace and opisthosoma of the female similar to the male except for lighter colouration than the male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 15.00, CL 6.85, CW 5.70, OL 7.10, OW 4.90; ALE> PLE> PME> AME; palp 11.35 (3.75 + 1.95 + 2.45 + 3.20), leg I 13.40 (4.30 + 2.30 + 2.60 + 2.70 + 1.50), leg II 12.70 (3.85 + 2.20 + 2.25 + 2.80 + 1.60), leg III 13.45 (3.65 + 2.30 + 2.35 + 3.20 + 1.95), leg IV 19.05 (4.25 + 2.70 + 3.60 + 5.60 + 2.90). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters, middle ones smaller than lateral ones with stalks, separated from each other basally, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 13). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shandong Province (Taishan, Qufu) Remarks. Males and females vary in body size: males (N = 4): BL 12.50–16.60, CL 6.80–7.10, CW 6.50–7.30, OL 6.00–8.60, OW 5.60–6.00; females (N = 15): BL 12.50–20.50, CL 6.00–8.60, CW 4.50–7.20, OL 6.40–10.20, OW 4.80–8.35. There was intraspecific variation in female genital morphology and molecular markers. For the female genitalia, the middle receptacular clusters were weakly sclerotized with long stalks (Fig. 13A, 13B), or well sclerotized (Fig. 13C–I). Intraspecific genetic distance is between 0% and 0.6% based on the K2P substitution model and COI nucleotide sequences (N = 9). The GenBank accession code of the holotype (XUX-2014-143A) is MH172722.

Luthela yuncheng sp. nov. (Fig. 14A–M) Holotype. Male (XUX-2011-235, matured on 10 July 2012 at CBEE), Wulipo Park, Linyi County, Yuncheng City, Shanxi Province, China, 35.157°N, 110.758°E, 441 m, collected on 30 November 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratypes. 7 females (XUX-2011-229/230/231/232/233/234/236, and 231, 232, 236 matured on 10 July 2012 at CBEE), same data as for holotype. All in CBEE. Diagnosis. The male of L. yuncheng sp. nov. can be distinguished from those of L. yiyuan sp. nov., and L. schensiensis by tegulum having a regular serrated marginal apophysis and a regular dorsal extension of terminal apophysis (Fig. 14G); from that of L. dengfeng sp. nov. by the conductor with a longer lower spine; from that of L. handan sp. nov. by the contrategulum with smaller marginal teeth (Fig. 14D); from those of L. badong s p. nov. and L. taian sp. nov. by the conductor with a small middle spine (Fig. 14E–G). Females of L. yuncheng sp. nov. differ from those of L. badong sp. nov., and L. yiyuan sp. nov. by the bases of middle receptacular clusters being separated from each other (Fig. 14H, 14I); from those of L. dengfeng sp. nov., L. handan sp. nov., L. schensiensis, and L. taian sp. nov. by the similar-sized receptacular clusters (Fig. 14H–M); from those of L. luotianensis by the posterior margin of genital sternite being slightly incurved (Fig. 14K, 14L). Description. Male (Holotype). Carapace dark brown; opisthosoma brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 16.85, CL 7.35, CW 6.95, OL 8.75, OW 6.35; ALE> PLE> PME> AME; leg I 21.00 (6.15 + 2.85 + 4.15 + 5.10 + 2.75), leg II 20.65 (5.65 + 2.75 + 3.95 + 5.35 + 2.95), leg III 22.70 (5.45 + 2.95 + 4.00 + 6.50 + 3.80), leg IV 29.70 (7.65 + 3.35 + 5.30 + 8.75 + 4.65). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 14A–C). Contrategulum with 8 teeth along the margin, distal tooth with 3 serrations (Fig. 14D, 14F). Marginal apophysis of tegulum with regular serrations, dorsal extension of terminal apophysis of tegulum with regular serrations basally and irregular serrations distally, and terminal apophysis of tegulum smooth (Fig. 14G). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum, and one small tooth situated at middle of two long spines (Fig. 14D–G). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 14D, 14G). Female (XUX-2011-229). Carapace reddish brown; opisthosoma brown, with dark brown tergites; chelicerae robust with promargin of groove with 10–11 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 25.55, CL 9.55, CW 8.00, OL 13.85, OW 11.65; ALE> PLE> PME> AME; palp 16.45 (5.65 + 2.55 + 3.50 + 4.75), leg I 18.45 (6.50 + 2.85 + 3.80 + 3.15 + 2.15), leg II 18.20 (5.85 + 2.80 + 3.35 + 3.85 + 2.35), leg III 18.90 (5.55 + 2.75 + 3.25 + 4.50 + 2.85), leg IV 26.90 (7.15 + 3.15 + 5.25 + 7.65 + 3.70). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters similar-sized, middle ones with stalks, separated from each other basally, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 14H–M). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shanxi Province (Yuncheng) Remarks. Females vary in body size: (N = 7): BL 15.65–25.55, CL 7.75–10.75, CW 6.95–9.55, OL 7.75–11.05, OW 5.90–11.65. Intraspecific genetic distance is between 0% and 3.4% based on the K2P substitution model and COI nucleotide sequences (N = 7, including juveniles). The GenBank accession code of the holotype (XUX-2011-235) is MH172738.

Luthela badong sp. nov. (Fig. 5A���I) Holotype. Male (XUX-2012-140), Quankou Village, Yanduhe Town, Badong County, Enshi, Hubei Province, China, 31.280��N, 110.318��E, 275 m, collected on 18 August 2012 by F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratype. Female (XUX-2012-141), Mashan Village, Yanduhe Town, Badong County, Enshi, Hubei Province, China, 31.286��N, 110.303��E, 450 m, collected on 18 August 2012 by F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Diagnosis. The male of L. badong sp. nov. can be distinguished from that of L. dengfeng sp. nov. by the conductor having two slender shorter spines, and a larger angle between lower spine and conductor (Fig. 5D���G), and the contrategulum with four larger teeth (Fig. 5D); from those of the other Luthela species by the conductor having only two large spines, upper and lower ones (Fig. 5A���G). The female of L. badong sp. nov. differs from those of L. dengfeng sp. nov., L. handan sp. nov., and L. taian sp. nov. by the two pairs of similar-sized receptacular clusters (Fig. 5H); from those of L. schensiensis and L. yiyuan sp. nov. by the middle receptacular clusters with slender stalks (Fig. 5H); from those of L. luotianensis and L. yuncheng sp. nov. by the bases of the middle receptacular clusters being close to each other (Fig. 5H). Description. Male (Holotype). Carapace reddish brown; opisthosoma brown, with brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 9 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and others small; 7 spinnerets. Measurements: BL 12.51, CL 6.32, CW 5.89, OL 6.10, OW 3.90; ALE> PLE> PME> AME; leg I 19.47 (5.45 + 2.55 + 3.92 + 4.90 + 2.65), leg II 19.50 (4.95 + 2.50 + 3.65 + 5.35 + 3.05), leg III 20.36 (5.00 + 2.28 + 3.70 + 6.18 + 3.20), leg IV 26.70 (6.45 + 2.65 + 5.25 + 8.40 + 3.95). Palp: Paracymbium unpigmented and unsclerotized on prolateral side, numerous setae and spines located at tip of paracymbium (Fig. 5A���C). Contrategulum with 4 distinct teeth along the margin (Fig. 5D), and distal one with 3 serrations (Fig. 5F). Marginal apophysis and dorsal extension of terminal apophysis of tegulum serrated, and terminal apophysis of tegulum smooth (Fig. 5E���G). Conductor situated ventro-proximally on embolus, with two large spines, upper one parallel to embolus, lower one slender and short (Fig. 5D���G). Embolus largely sclerotized, with flat opening and several longitudinal ribs distally (Fig. 5B���D). Female (XUX-2012-141). Carapace and opisthosoma similar to male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 16.50, CL 8.35, CW 6.90, OL 7.21, OW 5.25; ALE> PLE> PME> AME; palp 13.25 (4.45 + 2.45 + 2.80 + 3.55), leg I 15.25 (5.05 + 2.75 + 2.75 + 3.15 + 1.55), leg II 15.25 (4.65 + 2.70 + 2.55 + 3.45 + 1.90), leg III 15.87 (4.35 + 2.75 + 2.55 + 4.02 + 2.20), leg IV 24.25 (6.50 + 3.55 + 4.20 + 6.50 + 3.50). Genitalia: Posterior margin of genital sternite incurved (Fig. 5H, 5I), two paired receptacular clusters, middle ones with slender stalks, and bases of middle stalks close to each other, situated at anterior margin of bursa copulatrix, lateral ones situated at dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 5H). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Hubei Province (Badong). Remarks. There is no intraspecific variation between the COI nucleotide sequences of the male and female. The GenBank accession code of the holotype (XUX-2012-140) is KP229863., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 137-139, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635

Luthela yiyuan sp. nov. (Figs 3A–K, 4O–T) Sinothela sinensis: Xu et al., 2015a: 139; misidentification. Holotype. Male (XUX-2012-051, matured on 29 July 2012 at CBEE), Shiqiao Town, Yiyuan County, Zibo City, Shandong Province, China, 36.152°N, 118.334°E, 275 m, collected on 1 May 2012, by F.X. Liu, Z.L. Liu, and X. Xu, CBEE. Paratypes. 3 females (XUX-2012-046/049/050) and 1 male (XUX-2012-045, matured on 29 July 2012 at CBEE), same data as the holotype; 6 females (XUX-2012-034/035/036/037/040/042, XUX-2012-036 matured on 10 July 2012 at CBEE), Puji Town, Zhangqiu City, Shandong Province, China, 36.728°N, 117.611°E, 100 m, collected on 30 April 2012; 1 female (XUX-2016-096), Hushan forestry station, Zhangqiu City, Shandong Province, China, 36.674°N, 117.553°E, 184 m, collected on 5 August 2016; 2 females (XUX-2016-097/099), Puji Town, Zhangqiu City, Shandong Province, China, 36.728°N, 117.613°E, 106 m, collected on 7 August 2016; 10 females (XUX-2016-100/101/102/103/104/105/106/ 107/108/109), Zhangqiu City, Shandong Province, China, 36.669°N, 117.428°E, 103 m, collected on 8 August 2016, by F. Li, F.X. Liu, H. Liu, and Z. T. Zhang. All in CBEE. Diagnosis. Males of L. yiyuan sp. nov. resemble those of L. schensiensis and L. yuncheng sp. nov., but can be distinguished from those of L. schensiensis by the tegulum having a slightly larger terminal apophysis, and the embolus with less longitudinal ribs retrolaterally (Fig. 3J, 3K); from those of L. yuncheng sp. nov. by the marginal tegular apophysis and the dorsal extension of the terminal apophysis of the tegulum having irregular teeth (Fig. 3G, 3K); from those of L. dengfeng sp. nov. by the conductor having a longer lower spine (Fig. 3G, 3K); from those of L. handan sp. nov. by contrategulum margin having slightly small teeth (Fig. 3D, 3K); from those of L. badong sp. nov. and L. taian sp. nov. by the conductor with a small middle spine (Fig. 3A–G). Females of L. yiyuan sp. nov. differ from those of L. dengfeng sp. nov. by the bases of middle receptacular clusters being close to each other (Fig. 4). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs over ocular mound in longitudinal row; chelicerae robust, promargin of groove with 11 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 16.40, CL 6.65, CW 5.50, OL 8.80, OW 6.20; ALE> PLE> PME> AME; leg I 18.40 (5.45 + 2.30+ 3.65 + 4.70 + 2.30), leg II 18.10 (5.00 + 2.35 + 3.45 + 4.70 + 2.60), leg III 19.55 (4.95 + 2.35+ 3.50 + 5.60 + 3.15), leg IV 24.60 (6.25 + 2.50 + 4.80 + 7.55 + 3.50). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 3A–C). Contrategulum with 7 or 8 teeth along margin, distal tooth with 3 serrations (Fig. 3D, 3H). Tegulum with irregular dentate marginal apophysis, irregular dorsal extension of terminal apophysis, and smooth terminal apophysis (Fig. 3G, 3K). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum, and one small tooth situated at between two long spines, and close to upper one (Fig. 3A–G), or an inconspicuous tooth located at the middle of two long spines (Fig. 3H–J). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 3D, 3K). Female (XUX-2012-035). Carapace reddish brown; opisthosoma light brown, with brown tergites; chelicerae robust with promargin of groove with 10–11 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 21.70, CL 8.40, CW 8.10, OL 12.90, OW 12.20; ALE> PLE> PME> AME; palp 16.40 (5.20 + 2.90 + 3.90 + 4.40), leg I 17.90 (5.60 + 3.30 + 3.40 + 3.70 + 1.90), leg II 16.40 (4.50 + 3.20 + 3.10 + 3.50 + 2.10), leg III 18.40 (5.40 + 3.20 + 3.00 + 4.40 + 2.40), leg IV 27.60 (7.90 + 3.90 + 4.80 + 7.40 + 3.60). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters similar-sized, middle ones with stalks, close to each other (Fig. 4A–M), or fused together as one large receptacular cluster (Fig. 4N, 4O), situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 4). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shandong Province (Zhangqiu, Zibo) Remarks. Males and females vary in body size: males (N = 2): BL 14.75–16.40, CL 6.35–6.65, CW 5.50–5.85, OL 7.40–8.80, OW 5.00–6.20; females (N = 22): BL 16.70–24.90, CL 8.00–10.75, CW 7.10–8.90, OL 8.80–13.40, OW 6.90–13.20. There was intraspecific variation in male and female genital morphology and molecular markers. For male genitalia, the conductor has a small middle spine close to the upper spine (Fig. 3A–G), or an inconspicuous middle spine close to the lower spine (Fig. 3H). For female genitalia, the posterior margin of genital sternite was slightly incurved (Fig. 4F, 4G, 4I, 4J, 4P, 4Q) or straight (Fig. 4H, 4R–T); there were two pairs of receptacular clusters or the middle ones were fused as a large one (Fig. 4N, 4O). Intraspecific genetic distance is between 0% and 3.4% based on the K2P substitution model and COI nucleotide sequences (N = 21, including juveniles). The GenBank accession code of the holotype (XUX-2012-051) is MH172727.

Genus Luthela Xu & Li, gen. nov. Type species: Luthela yiyuan sp. nov. Diagnosis. Males of Luthela can be distinguished from those of Songthela Ono, 2000 by the contrategular margin having regular larger teeth (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); from those of the other genera of Heptathelinae by the conductor with a smooth surface, and with two long spines, upper and lower ones (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D). Luthela females differ from those of Songthela by the middle receptacular clusters having shorter stalks (Figs 4, 5H, 6H–J, 8, 9, 11, 13, 14H–M); from those of the other genera of Heptathelinae by the positions of two pairs of the receptacular clusters: middle ones situated at anterior margin of bursa copulatrix and lateral ones situated at dorsolateral position of bursa copulatrix (Figs 4, 5H, 6H–J, 8, 9, 11, 13, 14H–M). Description. Male body length, 12.5–18.6 (N = 18), female, 12.5–27.0 (N = 85); male palpal conductor with a smooth surface, and with two long spines, upper and lower ones (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); contrategular margin with regular teeth (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); tegulum with three distinct apophyses (Figs 3C, 5F, 6E, 7F, 10F, 12F, 14F); female genitalia with two paired receptacular clusters, middle ones situated at anterior margin of bursa copulatrix with tubular stalks, and lateral ones situated more dorsolaterally (Figs 4, 5H, 6H–J, 8, 9, 11, 13, 14H–M). Etymology. The genus name is created by a combination of ‘Lu-’, referring to Shandong Province, China, and ‘thela’, referring to spinnerets as the Greek word thele means nipple-like protuberance (Ono 2000). The gender is feminine. Composition. L. yiyuan sp. nov. (type species), L. luotianensis (Yin et al., 2002), comb. nov., L. schensiensis (Schenkel, 1953), comb. nov., L. badong sp. nov., L. dengfeng sp. nov., L. handan sp. nov., L. taian sp. nov., L. yuncheng sp. nov. Distribution. China (Hebei, Henan, Hubei, Shandong, Shaanxi, and Shanxi Provinces).

Genus Luthela Xu & Li, gen. nov. Type species: Luthela yiyuan sp. nov. Diagnosis. Males of Luthela can be distinguished from those of Songthela Ono, 2000 by the contrategular margin having regular larger teeth (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); from those of the other genera of Heptathelinae by the conductor with a smooth surface, and with two long spines, upper and lower ones (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D). Luthela females differ from those of Songthela by the middle receptacular clusters having shorter stalks (Figs 4, 5H, 6H���J, 8, 9, 11, 13, 14H���M); from those of the other genera of Heptathelinae by the positions of two pairs of the receptacular clusters: middle ones situated at anterior margin of bursa copulatrix and lateral ones situated at dorsolateral position of bursa copulatrix (Figs 4, 5H, 6H���J, 8, 9, 11, 13, 14H���M). Description. Male body length, 12.5���18.6 (N = 18), female, 12.5���27.0 (N = 85); male palpal conductor with a smooth surface, and with two long spines, upper and lower ones (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); contrategular margin with regular teeth (Figs 3D, 5D, 6D, 7D, 10H, 12D, 14D); tegulum with three distinct apophyses (Figs 3C, 5F, 6E, 7F, 10F, 12F, 14F); female genitalia with two paired receptacular clusters, middle ones situated at anterior margin of bursa copulatrix with tubular stalks, and lateral ones situated more dorsolaterally (Figs 4, 5H, 6H���J, 8, 9, 11, 13, 14H���M). Etymology. The genus name is created by a combination of ���Lu-���, referring to Shandong Province, China, and ���thela���, referring to spinnerets as the Greek word thele means nipple-like protuberance (Ono 2000). The gender is feminine. Composition. L. yiyuan sp. nov. (type species), L. luotianensis (Yin et al., 2002), comb. nov., L. schensiensis (Schenkel, 1953), comb. nov., L. badong sp. nov., L. dengfeng sp. nov., L. handan sp. nov., L. taian sp. nov., L. yuncheng sp. nov. Distribution. China (Hebei, Henan, Hubei, Shandong, Shaanxi, and Shanxi Provinces)., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 134-135, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Ono, H. (2000) Zoogeographic and taxonomic notes on spiders of the subfamily Heptathelinae (Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum, 33, 145 - 151.","Yin, C. M., Tang, G., Zhao, J. Z. & Chen, J. (2002) Two new species of the genus Heptathela from China (Araneae: Liphistiidae). Acta Arachnologica Sinica, 11, 18 - 21.","Schenkel, E. (1953) Chinesische Arachnoidea aus dem Museum Hoangho-Peiho in Tientsin. Boletim do Museu Nacional do Rio de Janeiro, 119, 1 - 108."]}

Luthela handan sp. nov. (Figs 7, 8) Holotype. Male (XUX-2011-214, matured on 10 July 2012 at CBEE), Qilidian, Yongnian County, Handan City, Hebei Province, China, 36.760��N, 114.476��E, 58 m, collected on 23 November 2011 by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratypes. 4 females (XUX-2011-206/207/208/215, 208 and 215 matured on 10 July 2012 at CBEE) and 1 male (XUX-2011-209, matured on 12 June 2013 at CBEE), same data as for holotype; 3 females (XUX-2011- 202/203/205), nearby Hebei University of Engineering, Yongnian County, Handan City, Hebei Province, China, 36.752��N, 114.471��E, 66 m, collected on 23 November 2011; 9 females (XUX-2011-217/218/219/220/221/222/ 223/224/227) Dongtumen Village, Luquan County, Shijiazhang City, Hebei Province, China, 38.074��N, 114.282��E, 160 m, collected on 25 November 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu. All in CBEE. Diagnosis. Males of L. handan sp. nov. can be distinguished from those of L. dengfeng sp. nov., L. schensiensis, L. yiyuan sp. nov., and L. yuncheng sp. nov. by the conductor with a longer lower spine, and the dorsal extension of terminal apophysis of tegulum with regular serrations (Fig. 7E���G); from those of L. badong sp. nov. and L. taian sp. nov. by the conductor with small middle spines (Fig. 7B, 7D), and lower spine thicker and longer (Fig. 7E, 7G); Females of L. handan sp. nov. differ from those of L. badong sp. nov., L. luotianensis, L. yiyuan sp. nov., and L. yuncheng sp. nov. by the middle receptacular clusters smaller than lateral ones (Fig. 8A���C, 8G���I); from those of L. schensiensis by the lateral receptacular clusters with slightly longer stalks (Fig. 8J); from those of L. dengfeng sp. nov. and L. taian sp. nov. by the middle receptacular clusters with short stalks and close to each other basally (Fig. 8A���F, 8H, 8I). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 15.55, CL 7.15, CW 6.85, OL 8.35, OW 5.65; ALE> PLE> PME> AME; leg I 19.95 (5.95 + 2.65 + 4.05 + 4.95 + 2.35), leg II 19.95 (5.80 + 2.60 + 3.55 + 5.05 + 2.95), leg III 20.55 (5.05 + 2.70 + 3.55 + 5.90 + 3.35), leg IV 26.65 (6.65 + 3.00 + 5.00 + 8.05 + 3.95). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 7A���C). Contrategulum with 6 teeth along margin, and distal one with 3 serrations (Fig. 7D, 7F). Marginal apophysis of tegulum with few serrations proximally and smooth distally, dorsal extension of terminal apophysis of tegulum with regular serrations, and terminal apophysis of tegulum smooth, thumb-shaped (Fig. 7E���G). Conductor situated ventro-proximally on embolus, with two long spines, upper one parallel to embolus, lower one pointed to proximal of contrategulum, and one or two inconspicuous, small spines situated at middle between two long spines (Fig. 7D���G). Embolus largely sclerotized, with flat opening and several longitudinal ribs retrolaterally (Fig. 7D, 7G). Female (XUX-2011-224). Carapace and opisthosoma light brown; chelicerae robust with promargin of groove with 9���12 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 17.35, CL 8.05, CW 6.95, OL 8.65, OW 6.30; ALE> PLE> PME> AME; palp 13.75 (4.65 + 2.40 + 3.10 + 3.60), leg I 15.85 (5.25 + 2.85 + 3.15 + 3.20 + 1.40), leg II 14.75 (4.70 + 2.85 + 2.60 + 3.25 + 1.40), leg III 16.30 (4.50 + 2.65 + 2.75 + 3.95 + 2.45), leg IV 22.45 (4.85 + 3.35 + 4.35 + 6.45 + 3.45). Genitalia: Posterior margin of genital sternite slightly incurved (Fig. 8A���L), two pairs of receptacular clusters, middle ones smaller than lateral ones with slender stalks, separated from each other (Fig. 8B, 8F, 8G���I), situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with short stalks (Fig. 8C, 8G���I). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Hebei Province (Handan, Shijiazhuang) Remarks. Males and females vary in body size: males (N = 2): BL 14.75���16.20, CL 6.35���7.25, CW 5.50���6.85, OL 7.40���8.55, OW 5.00���6.15; females (N = 16): BL 17.35���26.95, CL 8.05���10.15, CW 6.95���9.45, OL 8.65���16.16, OW 6.30���15.35. Intraspecific variation was found in female genital morphology and molecular markers. For female genitalia, the middle receptacular clusters are sclerotized well (Fig. 8A���C), or weakly (Fig. 8G���I); the bases of the middle stalks are fused together (Fig. 8A, 8D), or separated from each other (Fig. 8B, 8C, 8G���L). Intraspecific genetic distance ranges from 0% to 3.5% based on the K2P substitution model and COI nucleotide sequences (N = 14, including juveniles). The GenBank accession code of the holotype (XUX-2011-214) is KP229810., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 141-142, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635

Luthela luotianensis (Yin et al., 2002) comb. nov. (Fig. 9A–D) Heptathela luotianensis Yin et al., 2002: 18 (holotype: female, from Luotian, Hubei Province, China, 3048’N, 11524’E, collected by J.Z. Zhao and F.X. Liu on 22–23 October 1978, deposited at College of Life Sciences, Hubei University, type presumed to be lost). Sinothela luotianensis: Xu et al., 2015a: 139. Material examined. 3 females (XUX-2012-079/080/082), Fengyi’ao Chinese chestnut plantation, Luotian County, Hubei Province, China, 30.750N, 115.397E, 123 m, collected on 8 July, 2012, by F.X. Liu, X.Y. Wang, and X. Xu. Diagnosis. Females of L. luotianensis can be distinguished from those of L. badong sp. nov. by the bases of the middle stalks being separated from each other (Fig. 9A–D); from those of L. dengfeng sp. nov., L. handan sp. nov., and L. taian sp. nov. by the similar-sized receptacular clusters (Fig. 9A, 9C); from those of L. schensiensis and L. yiyuan sp. nov. by the bases of middle stalks separated from each other and slightly slender (Fig. 9A–D); from those of L. yuncheng sp. nov. by obviously incurved posterior margin of genital sternite (Fig. 9B, 9D). Male. Unknown. Redescription. Female (XUX-2012-079). Carapace reddish brown; opisthosoma brown, with brown tergites; sternum narrow, much longer than wide; a few long pointed hairs running over ocular mound in a longitudinal row; chelicerae robust with promargin of groove bearing 9–10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 12.95, CL 5.95, CW 5.20, OL 6.10, OW 4.85; ALE> PLE> PME> AME; palp 12.70 (4.15 + 2.15 + 2.65 + 3.75), leg I 15.60 (5.05 + 2.65 + 3.05 + 3.05 + 1.80), leg II 15.65 (4.85 + 2.75 + 2.80 + 3.40 + 1.85), leg III 15.85 (4.65 + 2.70 + 2.70 + 3.75 + 2.05), leg IV 23.00 (6.25 + 3.35 + 4.00 + 6.15 + 3.25). Genitalia: Posterior margin of genital sternite obviously incurved (Fig. 9B, 9D), two pairs of receptacular clusters, middle ones situated at anterior margin of bursa copulatrix with slender stalks, separated from each other basally, lateral ones situated at slightly dorsolateral position of bursa copulatrix (Fig. 9A–D). Distribution. Hubei Province (Luotian) Remarks. Females vary in body size: (N = 3): BL 12.95–17.45, CL 5.95–7.55, CW 5.20–7.55, OL 6.10–8.85, OW 4.65–7.05. Since the holotype was lost, we identified the newly collected specimens as L. luotianensis because: (1) the new specimens were collected at the type locality; (2) we compared the newly collected specimens with the original descriptions and illustrations provided by Yin et al. (2002), and found that the body size and the receptacular clusters of newly collected specimens are similar to those of the holotype. Intraspecific genetic distance ranges from 0% to 0.4% based on the K2P substitution model and COI nucleotide sequences (N = 7, including juveniles). The GenBank accession code of XUX-2012-079 is KP229881.

Luthela yuncheng sp. nov. (Fig. 14A���M) Holotype. Male (XUX-2011-235, matured on 10 July 2012 at CBEE), Wulipo Park, Linyi County, Yuncheng City, Shanxi Province, China, 35.157��N, 110.758��E, 441 m, collected on 30 November 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratypes. 7 females (XUX-2011-229/230/231/232/233/234/236, and 231, 232, 236 matured on 10 July 2012 at CBEE), same data as for holotype. All in CBEE. Diagnosis. The male of L. yuncheng sp. nov. can be distinguished from those of L. yiyuan sp. nov., and L. schensiensis by tegulum having a regular serrated marginal apophysis and a regular dorsal extension of terminal apophysis (Fig. 14G); from that of L. dengfeng sp. nov. by the conductor with a longer lower spine; from that of L. handan sp. nov. by the contrategulum with smaller marginal teeth (Fig. 14D); from those of L. badong s p. nov. and L. taian sp. nov. by the conductor with a small middle spine (Fig. 14E���G). Females of L. yuncheng sp. nov. differ from those of L. badong sp. nov., and L. yiyuan sp. nov. by the bases of middle receptacular clusters being separated from each other (Fig. 14H, 14I); from those of L. dengfeng sp. nov., L. handan sp. nov., L. schensiensis, and L. taian sp. nov. by the similar-sized receptacular clusters (Fig. 14H���M); from those of L. luotianensis by the posterior margin of genital sternite being slightly incurved (Fig. 14K, 14L). Description. Male (Holotype). Carapace dark brown; opisthosoma brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 16.85, CL 7.35, CW 6.95, OL 8.75, OW 6.35; ALE> PLE> PME> AME; leg I 21.00 (6.15 + 2.85 + 4.15 + 5.10 + 2.75), leg II 20.65 (5.65 + 2.75 + 3.95 + 5.35 + 2.95), leg III 22.70 (5.45 + 2.95 + 4.00 + 6.50 + 3.80), leg IV 29.70 (7.65 + 3.35 + 5.30 + 8.75 + 4.65). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 14A���C). Contrategulum with 8 teeth along the margin, distal tooth with 3 serrations (Fig. 14D, 14F). Marginal apophysis of tegulum with regular serrations, dorsal extension of terminal apophysis of tegulum with regular serrations basally and irregular serrations distally, and terminal apophysis of tegulum smooth (Fig. 14G). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum, and one small tooth situated at middle of two long spines (Fig. 14D���G). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 14D, 14G). Female (XUX-2011-229). Carapace reddish brown; opisthosoma brown, with dark brown tergites; chelicerae robust with promargin of groove with 10���11 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 25.55, CL 9.55, CW 8.00, OL 13.85, OW 11.65; ALE> PLE> PME> AME; palp 16.45 (5.65 + 2.55 + 3.50 + 4.75), leg I 18.45 (6.50 + 2.85 + 3.80 + 3.15 + 2.15), leg II 18.20 (5.85 + 2.80 + 3.35 + 3.85 + 2.35), leg III 18.90 (5.55 + 2.75 + 3.25 + 4.50 + 2.85), leg IV 26.90 (7.15 + 3.15 + 5.25 + 7.65 + 3.70). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters similar-sized, middle ones with stalks, separated from each other basally, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 14H���M). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shanxi Province (Yuncheng) Remarks. Females vary in body size: (N = 7): BL 15.65���25.55, CL 7.75���10.75, CW 6.95���9.55, OL 7.75���11.05, OW 5.90���11.65. Intraspecific genetic distance is between 0% and 3.4% based on the K2P substitution model and COI nucleotide sequences (N = 7, including juveniles). The GenBank accession code of the holotype (XUX-2011-235) is MH172738., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 150-152, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635

Luthela handan sp. nov. (Figs 7, 8) Holotype. Male (XUX-2011-214, matured on 10 July 2012 at CBEE), Qilidian, Yongnian County, Handan City, Hebei Province, China, 36.760°N, 114.476°E, 58 m, collected on 23 November 2011 by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratypes. 4 females (XUX-2011-206/207/208/215, 208 and 215 matured on 10 July 2012 at CBEE) and 1 male (XUX-2011-209, matured on 12 June 2013 at CBEE), same data as for holotype; 3 females (XUX-2011- 202/203/205), nearby Hebei University of Engineering, Yongnian County, Handan City, Hebei Province, China, 36.752°N, 114.471°E, 66 m, collected on 23 November 2011; 9 females (XUX-2011-217/218/219/220/221/222/ 223/224/227) Dongtumen Village, Luquan County, Shijiazhang City, Hebei Province, China, 38.074°N, 114.282°E, 160 m, collected on 25 November 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu. All in CBEE. Diagnosis. Males of L. handan sp. nov. can be distinguished from those of L. dengfeng sp. nov., L. schensiensis, L. yiyuan sp. nov., and L. yuncheng sp. nov. by the conductor with a longer lower spine, and the dorsal extension of terminal apophysis of tegulum with regular serrations (Fig. 7E–G); from those of L. badong sp. nov. and L. taian sp. nov. by the conductor with small middle spines (Fig. 7B, 7D), and lower spine thicker and longer (Fig. 7E, 7G); Females of L. handan sp. nov. differ from those of L. badong sp. nov., L. luotianensis, L. yiyuan sp. nov., and L. yuncheng sp. nov. by the middle receptacular clusters smaller than lateral ones (Fig. 8A–C, 8G–I); from those of L. schensiensis by the lateral receptacular clusters with slightly longer stalks (Fig. 8J); from those of L. dengfeng sp. nov. and L. taian sp. nov. by the middle receptacular clusters with short stalks and close to each other basally (Fig. 8A–F, 8H, 8I). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 15.55, CL 7.15, CW 6.85, OL 8.35, OW 5.65; ALE> PLE> PME> AME; leg I 19.95 (5.95 + 2.65 + 4.05 + 4.95 + 2.35), leg II 19.95 (5.80 + 2.60 + 3.55 + 5.05 + 2.95), leg III 20.55 (5.05 + 2.70 + 3.55 + 5.90 + 3.35), leg IV 26.65 (6.65 + 3.00 + 5.00 + 8.05 + 3.95). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 7A–C). Contrategulum with 6 teeth along margin, and distal one with 3 serrations (Fig. 7D, 7F). Marginal apophysis of tegulum with few serrations proximally and smooth distally, dorsal extension of terminal apophysis of tegulum with regular serrations, and terminal apophysis of tegulum smooth, thumb-shaped (Fig. 7E–G). Conductor situated ventro-proximally on embolus, with two long spines, upper one parallel to embolus, lower one pointed to proximal of contrategulum, and one or two inconspicuous, small spines situated at middle between two long spines (Fig. 7D–G). Embolus largely sclerotized, with flat opening and several longitudinal ribs retrolaterally (Fig. 7D, 7G). Female (XUX-2011-224). Carapace and opisthosoma light brown; chelicerae robust with promargin of groove with 9–12 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 17.35, CL 8.05, CW 6.95, OL 8.65, OW 6.30; ALE> PLE> PME> AME; palp 13.75 (4.65 + 2.40 + 3.10 + 3.60), leg I 15.85 (5.25 + 2.85 + 3.15 + 3.20 + 1.40), leg II 14.75 (4.70 + 2.85 + 2.60 + 3.25 + 1.40), leg III 16.30 (4.50 + 2.65 + 2.75 + 3.95 + 2.45), leg IV 22.45 (4.85 + 3.35 + 4.35 + 6.45 + 3.45). Genitalia: Posterior margin of genital sternite slightly incurved (Fig. 8A–L), two pairs of receptacular clusters, middle ones smaller than lateral ones with slender stalks, separated from each other (Fig. 8B, 8F, 8G–I), situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with short stalks (Fig. 8C, 8G–I). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Hebei Province (Handan, Shijiazhuang) Remarks. Males and females vary in body size: males (N = 2): BL 14.75–16.20, CL 6.35–7.25, CW 5.50–6.85, OL 7.40–8.55, OW 5.00–6.15; females (N = 16): BL 17.35–26.95, CL 8.05–10.15, CW 6.95–9.45, OL 8.65–16.16, OW 6.30–15.35. Intraspecific variation was found in female genital morphology and molecular markers. For female genitalia, the middle receptacular clusters are sclerotized well (Fig. 8A–C), or weakly (Fig. 8G–I); the bases of the middle stalks are fused together (Fig. 8A, 8D), or separated from each other (Fig. 8B, 8C, 8G–L). Intraspecific genetic distance ranges from 0% to 3.5% based on the K2P substitution model and COI nucleotide sequences (N = 14, including juveniles). The GenBank accession code of the holotype (XUX-2011-214) is KP229810.

Luthela badong sp. nov. (Fig. 5A–I) Holotype. Male (XUX-2012-140), Quankou Village, Yanduhe Town, Badong County, Enshi, Hubei Province, China, 31.280°N, 110.318°E, 275 m, collected on 18 August 2012 by F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Paratype. Female (XUX-2012-141), Mashan Village, Yanduhe Town, Badong County, Enshi, Hubei Province, China, 31.286°N, 110.303°E, 450 m, collected on 18 August 2012 by F.X. Liu, X.Y. Wang, and X. Xu, CBEE. Diagnosis. The male of L. badong sp. nov. can be distinguished from that of L. dengfeng sp. nov. by the conductor having two slender shorter spines, and a larger angle between lower spine and conductor (Fig. 5D–G), and the contrategulum with four larger teeth (Fig. 5D); from those of the other Luthela species by the conductor having only two large spines, upper and lower ones (Fig. 5A–G). The female of L. badong sp. nov. differs from those of L. dengfeng sp. nov., L. handan sp. nov., and L. taian sp. nov. by the two pairs of similar-sized receptacular clusters (Fig. 5H); from those of L. schensiensis and L. yiyuan sp. nov. by the middle receptacular clusters with slender stalks (Fig. 5H); from those of L. luotianensis and L. yuncheng sp. nov. by the bases of the middle receptacular clusters being close to each other (Fig. 5H). Description. Male (Holotype). Carapace reddish brown; opisthosoma brown, with brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 9 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and others small; 7 spinnerets. Measurements: BL 12.51, CL 6.32, CW 5.89, OL 6.10, OW 3.90; ALE> PLE> PME> AME; leg I 19.47 (5.45 + 2.55 + 3.92 + 4.90 + 2.65), leg II 19.50 (4.95 + 2.50 + 3.65 + 5.35 + 3.05), leg III 20.36 (5.00 + 2.28 + 3.70 + 6.18 + 3.20), leg IV 26.70 (6.45 + 2.65 + 5.25 + 8.40 + 3.95). Palp: Paracymbium unpigmented and unsclerotized on prolateral side, numerous setae and spines located at tip of paracymbium (Fig. 5A–C). Contrategulum with 4 distinct teeth along the margin (Fig. 5D), and distal one with 3 serrations (Fig. 5F). Marginal apophysis and dorsal extension of terminal apophysis of tegulum serrated, and terminal apophysis of tegulum smooth (Fig. 5E–G). Conductor situated ventro-proximally on embolus, with two large spines, upper one parallel to embolus, lower one slender and short (Fig. 5D–G). Embolus largely sclerotized, with flat opening and several longitudinal ribs distally (Fig. 5B–D). Female (XUX-2012-141). Carapace and opisthosoma similar to male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 16.50, CL 8.35, CW 6.90, OL 7.21, OW 5.25; ALE> PLE> PME> AME; palp 13.25 (4.45 + 2.45 + 2.80 + 3.55), leg I 15.25 (5.05 + 2.75 + 2.75 + 3.15 + 1.55), leg II 15.25 (4.65 + 2.70 + 2.55 + 3.45 + 1.90), leg III 15.87 (4.35 + 2.75 + 2.55 + 4.02 + 2.20), leg IV 24.25 (6.50 + 3.55 + 4.20 + 6.50 + 3.50). Genitalia: Posterior margin of genital sternite incurved (Fig. 5H, 5I), two paired receptacular clusters, middle ones with slender stalks, and bases of middle stalks close to each other, situated at anterior margin of bursa copulatrix, lateral ones situated at dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 5H). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Hubei Province (Badong). Remarks. There is no intraspecific variation between the COI nucleotide sequences of the male and female. The GenBank accession code of the holotype (XUX-2012-140) is KP229863.

Luthela schensiensis (Schenkel, 1953), comb. nov. (Figs 10A–K, 11A–P) Liphistius sinensis schensiensis Schenkel,1953: 1 (the subspecies was described based on one female collected from Tongyuanfang, Gaoling, Shaanxi Province, 8 October 1916, deposited at Museum Hoangho-Peiho in Tientsin [corresponding to Huanghebaihe in Tianjin and Yellow River-Haihe River in Tianjin, in Chinese and English, respectively], lost). Liphistius schensiensis: Gertsch, 1967: 115. Heptathela sinensis schensiensis: Haupt, 1983: 285. Heptathela xianensis: Zhu & Wang, 1983: 131; Wang & Zhu, 1984: 403. Heptathela schensiensis: Platnick & Sedgwick, 1984: 4; Song & Haupt, 1984: 447; Song, Zhu & Chen, 1999: 33. Liphistius heyangensis Zhu & Wang, 1984: 251 (holotype: female, from Heyang, Shaanxi Province, 30 June 1982, deposited at Bethune Medical University, now College of Life Sciences, Jilin University, type presumed to be lost). Syn. nov. Abcathela schensiensis: Ono, 2000: 150. Sinothela heyangensi s: Haupt, 2003: 71. Sinothela schensiensis: Haupt, 2003: 71; Xu et al., 2015a: 139. Heptathela heyangensis: Platnick, 1989: 57. Abcathela heyangensis: Ono, 2000: 150. Material examined. 3 males (XUX-2011-241/242/247, matured on 10 July 2012 at CBEE) and 7 females (XUX- 2011-243/245/248/249/251/253/254), Kangjiapo Village, Heyang County, Shaanxi Province, China, 35.242°N, 110.155E, 730 m, collected on 1 December, 2011; 3 males (XUX-2011-255/264 matured on 10 July 2012, XUX- 2011-263 matured on 13 June 2013, at CBEE) and 10 females (XUX-2011-256–262/265/266/269), Quyang Village, Tongyuan, Gaoling County, Shaanxi Province, China, 35.546°N, 109.040E, 390 m, collected on 2 December, 2011; 1 male (XUX-2011-273, matured on 10 July 2012 at CBEE), 34.493°N, 108.806E, 374 m, and 1 female (XUX- 2011-270), 34.498°N, 108.786E, 359 m, Taiping village, Jingyang County, Xianyang City, Shaanxi Province, China, collected on 3 December, 2011, by H.Z. Li, F.X. Liu, X.Y. Wang, and X. Xu. All in CBEE. Diagnosis. Males of L. schensiensis resemble those of L. yiyuan sp. nov. and L. yuncheng sp. nov., but can be distinguished from those of L. yiyuan sp. nov. by the tegulum having a slightly smaller terminal apophysis (Fig. 10F, 10G), and the embolus with more longitudinal ribs retrolaterally (Fig. 10G, 10K); from those of L. yuncheng sp. nov. by the marginal apophysis and the dorsal extension of the terminal apophysis of the tegulum with irregular serrations (Fig. 10G, 10K); from those of L. dengfeng sp. nov. by the conductor with a longer lower spine (Fig. 10G, 10K); from those of L. handan sp. nov. by the contrategulum margin with slightly smaller teeth, and the dorsal extension of the terminal apophysis of the tegulum with irregular serrations (Fig. 10G, 10K); and from those of L. badong sp. n and L. taian sp. nov. by the conductor with small middle spines (Fig. 10A–E, 10H, 10K). Females of L. schensiensis differ from those of L. badong sp. nov. by the middle pair of receptacular clusters with slightly thicker and shorter stalks; from those of L. yiyuan sp. nov. by the middle pair of receptacular clusters with slightly longer stalks (Fig. 11A–D, 11I–L); from those of L. handan sp. nov., by the middle pair of receptacular clusters with slightly thicker stalks (Fig. 11A–D); from those of the other Luthela species by the bases of the middle stalks being close to each other (Fig. 11A–D). Redescription. Male (XUX-2011-273). Carapace reddish brown; opisthosoma brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs running over ocular mound in longitudinal row; chelicerae robust with promargin of groove with 11 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, second to fifth larger than others and the others small; 8 spinnerets. Measurements: BL 14.95, CL 7.05, CW 6.60, OL 7.15, OW 4.90; ALE> PLE> PME> AME; leg I 20.65 (6.05 + 2.75 + 4.15 + 5.05 + 2.65), leg II 20.85 (5.95 + 2.65 + 3.95 + 5.25 + 3.05), leg III 21.30 (5.65 + 2.55 + 3.75 + 6.40 + 2.95), leg IV 29.00 (7.20 + 2.95 + 5.35 + 8.95 + 4.55). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 10A–C). Contrategulum with 7–10 teeth along margin, and distal tooth serrated (Fig. 10A, 10D, 10H, 10J). Tegulum with an irregular dentate marginal apophysis and an irregular dorsal extension of terminal apophysis, as well as smooth terminal apophysis (Fig. 10C, 10F, 10G, 10I–K). Conductor situated ventro-proximally on embolus, with two long spines, upper one parallel to the furrow of embolus distally, lower one pointed to contrategulum, and one or two small teeth situated at between two long spines (Fig. 10A–C, 10D, 10H, 10K). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 10H, 10K). Female (XUX-2011-243). Carapace and opisthosoma similar to male; chelicerae robust with promargin of groove with 10 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 8 spinnerets. Measurements: BL 25.25, CL 10.35, CW 9.25, OL 14.75, OW 12.75; ALE> PLE> PME> AME; palp 16.30 (5.70 + 2.75 + 3.35 + 4.50), leg I 17.45 (4.85 + 3.05 + 3.55 + 3.75 + 2.25), leg II 19.15 (6.15 + 3.05 + 3.55 + 3.95 + 2.45), leg III 19.85 (6.05 + 3.15 + 3.45 + 4.45 + 2.75), leg IV 28.80 (7.65 + 3.75 + 5.35 + 8.00 + 4.05). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters, middle ones slightly smaller than lateral ones with stalks, close to each other, situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 11A–P). Distribution. Shaanxi Province (Heyang, Gaoling, Jingyang) Remarks. Males and females vary in body size: males (N = 7): BL 14.95–18.55, CL 6.75–7.85, CW 6.25–7.30, OL 7.15–10.45, OW 4.90–7.55; females (N = 18): BL 17.85–25.65, CL 7.55–10.25, CW 6.90–9.10, OL 9.05–16.65, OW 5.90–13.25. There is intraspecific variation in male and female genital morphology and molecular markers. Specifically, for the male genitalia, there was considerable intraspecific variation in the number of the small middle spines of the conductor, either one (Fig. 10A) or two spines (Fig. 10D, 10K); in the position of the small spine, either close to the upper spine (Fig. 10K), close to the lower spine (Fig. 10D), or located at the middle of two long spines (Fig. 10A); in the tip shape of lower spine of the conductor, bifurcated (Fig. 10D–F) or not (Fig. 10A, 10H). Intraspecific variation was noted in the number of teeth (7–10) along the contrategulum margin, and the different shape of the serrated distal tooth (Fig. 10A, 10D, 10H, 10J). For intraspecific variation in female genitalia, the posterior margin of the genital sternite may be incurved (Fig. 11C, 11D, 11I–K) or straight (Fig. 11A, 11L), the middle receptacular clusters were situated at the anterior margin of the bursa copulatrix (Fig. 11A–J, 11L), or at slightly dorsal part of the bursa copulatrix (Fig. 11K, 11O). Intraspecific genetic distance ranges from 0% to 1.8% based on the K2P substitution model and COI nucleotide sequences (N = 18). We treated L. heyangensis as a junior synonym of L. schensiensis for several reasons. First, L. heyangensis was diagnosed and described based exclusively on one female (Zhu & Wang 1984). As liphistiid females exhibit considerable intraspecific variation in female genitalia (Xu et al. 2017, 2019), it is difficult to diagnose the species based exclusively on female characters of a single specimen. Second, we collected intensively from the type localities of both L. heyangensis and L. schensiensis, and we examined and compared the newly collected males and females with the original descriptions and illustrations of both L. heyangensis and L. schensiensis (Schenkel 1953; Song & Haupt 1984; Zhu & Wang 1984). All lines of evidence support the type female of L. heyangensis belonging to L. schensiensis. Third, there is very low genetic variation (1.8%) among the newly collected specimens from the type localities of both L. heyangensis and L. schensiensis. Thus, the specimens collected from Heyang County, Shaanxi Province should be considered the same species as those collected from both Gaoling and Jingyang Counties, Shaanxi Province. The GenBank accession code of XUX-2011-273 is MH172701.

Luthela dengfeng sp. nov. (Fig. 6A–M) Holotype. Male (XUX-2012-031, matured on 29 July 2012 at CBEE), Shaolin Temple, Dengfeng City, Henan Province, China, 34.511°N, 112.950°E, 568 m, collected on 27 April 2012 by F.X. Liu, Z.L. Liu, and X. Xu, CBEE. Paratypes. 2 females (XUX-2012-023/030), same data as for holotype; 1 female (XUX-2016-111) same data as for holotype, collected on 23 August 2016 by F. Li, F.X. Liu, H. Liu, and Z. T. Zhang, CBEE. Diagnosis. The male of L. dengfeng sp. nov. can be distinguished from those of L. badong sp. nov. and L. taian sp. nov. by the conductor having one inconspicuous middle spine (Fig. 6D–G), and from that of L. badong sp. nov. by the conductor with two thicker spines; from those of all the other Luthela species by the conductor with a short lower spine, and a smaller angle between lower spine and conductor (Fig. 6D–G). Females of L. dengfeng sp. nov. differ from those of L. badong sp. nov., L. luotianensis, and L. yuncheng sp. nov. by the middle receptacular clusters being smaller than lateral ones (Fig. 6H–J); from those of L. schensiensis and L. yiyuan sp. nov. by the bases of the middle receptacular clusters separated from each other (Fig. 6H); from those of L. handan sp. nov. and L. taian sp. nov. by the middle receptacular clusters with slender and slightly longer stalks (Fig. 6H–M). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; a few long pointed hairs running over ocular mound in a longitudinal row; chelicerae robust with promargin of groove with 9–10 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 12.50, CL 6.00, CW 5.40, OL 6.90, OW 5.00; ALE> PLE> PME> AME; leg I 17.35 (5.10 + 2.35 + 3.55 + 4.05 + 2.30), leg II miss, leg III 18.10 (4.40 + 2.35 + 3.25 + 5.20 + 2.90), leg IV 23.95 (5.55 + 2.45 + 4.45 + 7.65 + 3.85). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 6A–C). Contrategulum with 6 teeth along the margin (Fig. 6D), and distal one with 4 serrations. Marginal apophysis of tegulum with several serrations proximally and smooth distally, dorsal extension of terminal apophysis of tegulum with irregular serrations, and terminal apophysis of tegulum smooth, triangle-shaped (Fig. 6F, 6G). Conductor situated ventro-proximally on embolus, with two large spines, upper one parallel to embolus, the angle between lower one and conductor about 30 ventrally (Fig. 6B, 4E), and one inconspicuous spine located at between two large spines (Fig. 6D–G). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 6A, 6D, 6G). Female (XUX-2012-030). Carapace and opisthosoma similar to male; chelicerae robust with promargin of groove with 10–11 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 17.80, CL 6.90, CW 6.60, OL 11.30, OW 10.20; ALE> PLE> PME> AME; palp 11.20 (3.90 + 1.80 + 2.60 + 2.90), leg I 13.70 (4.60 + 1.90 + 2.80 + 2.70 + 1.70), leg II 12.90 (4.00 + 2.20 + 2.40 + 2.50 + 1.80), leg III 13.90 (4.30 + 2.20 + 2.40 + 3.10 + 1.90), leg IV 21.00 (5.90 + 2.80 + 3.80 + 5.70 + 2.80). Genitalia: Posterior margin of genital sternite slightly incurved (Fig. 6H–J), two pairs of receptacular clusters, middle ones smaller than lateral ones with slender stalks, separated from each other basally, and situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with short thick stalks (Fig. 6H–M). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Henan Province (Dengfeng) Remarks. Female variation (N = 3): BL 17.80–21.30, CL 6.90–8.40, CW 6.60–7.80, OL 11.30–12.60, OW 10.20–11.20. No intraspecific variation was found among the COI nucleotide sequences of seven specimens (including juveniles). The GenBank accession code of the holotype (XUX-2012-031) is MH172686.

Luthela yiyuan sp. nov. (Figs 3A���K, 4O���T) Sinothela sinensis: Xu et al., 2015a: 139; misidentification. Holotype. Male (XUX-2012-051, matured on 29 July 2012 at CBEE), Shiqiao Town, Yiyuan County, Zibo City, Shandong Province, China, 36.152��N, 118.334��E, 275 m, collected on 1 May 2012, by F.X. Liu, Z.L. Liu, and X. Xu, CBEE. Paratypes. 3 females (XUX-2012-046/049/050) and 1 male (XUX-2012-045, matured on 29 July 2012 at CBEE), same data as the holotype; 6 females (XUX-2012-034/035/036/037/040/042, XUX-2012-036 matured on 10 July 2012 at CBEE), Puji Town, Zhangqiu City, Shandong Province, China, 36.728��N, 117.611��E, 100 m, collected on 30 April 2012; 1 female (XUX-2016-096), Hushan forestry station, Zhangqiu City, Shandong Province, China, 36.674��N, 117.553��E, 184 m, collected on 5 August 2016; 2 females (XUX-2016-097/099), Puji Town, Zhangqiu City, Shandong Province, China, 36.728��N, 117.613��E, 106 m, collected on 7 August 2016; 10 females (XUX-2016-100/101/102/103/104/105/106/ 107/108/109), Zhangqiu City, Shandong Province, China, 36.669��N, 117.428��E, 103 m, collected on 8 August 2016, by F. Li, F.X. Liu, H. Liu, and Z. T. Zhang. All in CBEE. Diagnosis. Males of L. yiyuan sp. nov. resemble those of L. schensiensis and L. yuncheng sp. nov., but can be distinguished from those of L. schensiensis by the tegulum having a slightly larger terminal apophysis, and the embolus with less longitudinal ribs retrolaterally (Fig. 3J, 3K); from those of L. yuncheng sp. nov. by the marginal tegular apophysis and the dorsal extension of the terminal apophysis of the tegulum having irregular teeth (Fig. 3G, 3K); from those of L. dengfeng sp. nov. by the conductor having a longer lower spine (Fig. 3G, 3K); from those of L. handan sp. nov. by contrategulum margin having slightly small teeth (Fig. 3D, 3K); from those of L. badong sp. nov. and L. taian sp. nov. by the conductor with a small middle spine (Fig. 3A���G). Females of L. yiyuan sp. nov. differ from those of L. dengfeng sp. nov. by the bases of middle receptacular clusters being close to each other (Fig. 4). Description. Male (Holotype). Carapace brown; opisthosoma light brown, with dark brown tergites; sternum narrow, much longer than wide; few long pointed hairs over ocular mound in longitudinal row; chelicerae robust, promargin of groove with 11 denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, the second to fifth larger than others and the others small; 7 spinnerets. Measurements: BL 16.40, CL 6.65, CW 5.50, OL 8.80, OW 6.20; ALE> PLE> PME> AME; leg I 18.40 (5.45 + 2.30+ 3.65 + 4.70 + 2.30), leg II 18.10 (5.00 + 2.35 + 3.45 + 4.70 + 2.60), leg III 19.55 (4.95 + 2.35+ 3.50 + 5.60 + 3.15), leg IV 24.60 (6.25 + 2.50 + 4.80 + 7.55 + 3.50). Palp: Prolateral paracymbium unpigmented and unsclerotized, numerous setae and spines at tip of paracymbium (Fig. 3A���C). Contrategulum with 7 or 8 teeth along margin, distal tooth with 3 serrations (Fig. 3D, 3H). Tegulum with irregular dentate marginal apophysis, irregular dorsal extension of terminal apophysis, and smooth terminal apophysis (Fig. 3G, 3K). Conductor situated ventro-proximally on embolus, with two long spines, upper one fits to furrow of embolus, lower one pointed to contrategulum, and one small tooth situated at between two long spines, and close to upper one (Fig. 3A���G), or an inconspicuous tooth located at the middle of two long spines (Fig. 3H���J). Embolus largely sclerotized, with a flat opening and several longitudinal ribs retrolaterally (Fig. 3D, 3K). Female (XUX-2012-035). Carapace reddish brown; opisthosoma light brown, with brown tergites; chelicerae robust with promargin of groove with 10���11 strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, similar to male; 7 spinnerets. Measurements: BL 21.70, CL 8.40, CW 8.10, OL 12.90, OW 12.20; ALE> PLE> PME> AME; palp 16.40 (5.20 + 2.90 + 3.90 + 4.40), leg I 17.90 (5.60 + 3.30 + 3.40 + 3.70 + 1.90), leg II 16.40 (4.50 + 3.20 + 3.10 + 3.50 + 2.10), leg III 18.40 (5.40 + 3.20 + 3.00 + 4.40 + 2.40), leg IV 27.60 (7.90 + 3.90 + 4.80 + 7.40 + 3.60). Genitalia: Posterior margin of genital sternite slightly incurved, two pairs of receptacular clusters similar-sized, middle ones with stalks, close to each other (Fig. 4A���M), or fused together as one large receptacular cluster (Fig. 4N, 4O), situated at anterior margin of bursa copulatrix, lateral ones situated at slightly dorsolateral position of bursa copulatrix with inconspicuous stalks (Fig. 4). Etymology. The species epithet, a noun in apposition, refers to the type locality. Distribution. Shandong Province (Zhangqiu, Zibo) Remarks. Males and females vary in body size: males (N = 2): BL 14.75���16.40, CL 6.35���6.65, CW 5.50���5.85, OL 7.40���8.80, OW 5.00���6.20; females (N = 22): BL 16.70���24.90, CL 8.00���10.75, CW 7.10���8.90, OL 8.80���13.40, OW 6.90���13.20. There was intraspecific variation in male and female genital morphology and molecular markers. For male genitalia, the conductor has a small middle spine close to the upper spine (Fig. 3A���G), or an inconspicuous middle spine close to the lower spine (Fig. 3H). For female genitalia, the posterior margin of genital sternite was slightly incurved (Fig. 4F, 4G, 4I, 4J, 4P, 4Q) or straight (Fig. 4H, 4R���T); there were two pairs of receptacular clusters or the middle ones were fused as a large one (Fig. 4N, 4O). Intraspecific genetic distance is between 0% and 3.4% based on the K2P substitution model and COI nucleotide sequences (N = 21, including juveniles). The GenBank accession code of the holotype (XUX-2012-051) is MH172727., Published as part of Xu, Xin, Yu, Li, Liu, Fengxiang & Li, Daiqin, 2022, Delimitation of the segmented trapdoor spider genus Luthela gen. nov., with comments on the genus Sinothela from northern China (Araneae, Mesothelae, Liphistiidae), pp. 131-154 in Zootaxa 5091 (1) on pages 135-137, DOI: 10.11646/zootaxa.5091.1.5, http://zenodo.org/record/5840635, {"references":["Xu, X., Liu, F., Chen, J., Ono, H., Li, D. & Kuntner, M. (2015 a) A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae). Zookeys, 488, 121 - 151. https: // doi. org / 10.3897 / zookeys. 488.8726"]}

GENUS GANTHELA XU & KUNTNER, 2015 For genus diagnosis and description, see Xu et al., 2015a. Type species by original designation, G. yundingensis Xu, 2015. For diagnosis and description, see Xu et al., 2015a.

Xu, Xin, Liu, Fengxiang, Chen, Jian, Li, Daiqin, Kuntner, Matjaž (2015): Integrative taxonomy of the primitively segmented spider genus Ganthela (Araneae: Mesothelae: Liphistiidae): DNA barcoding gap agrees with morphology. Zoological Journal of the Linnean Society 175 (2): 288-306, DOI: 10.1111/zoj.12280, URL: http://dx.doi.org/10.1111/zoj.12280

GANTHELA VENUS XU SP. NOV. (FIG. 9) Holotype Female (XUX-2013-160), Mount Wangjiang, Huangyang Village, Zhuangbian Town, Hanjiang District, Putian City, Fujian Province, China, 25.66°N, 118.88°E, 459 m a.s.l., 11 July 2013, collected by F.X. Liu, X. Xu, and Z. T. Zhang. No male found. Etymology Named after the Greek goddess of love, Venus. Diagnosis Females of G. venus sp. nov. can be distinguished from G. wangjiangensis sp. nov. by details in the genitalia, which have separated stalks on the anterior part (Fig. 9A, B). Ganthela venus sp. nov. furthermore differs from all other Ganthela species by the following unique nucleotide substitutions in the standard DNA barcode alignment: C (36), A (80), T (227), C (333), C (341), G (347), T (354), C (431), and T (539). Description Female (holotype). Carapace and opisthosoma light brown; tergites slightly dark brown; sternum narrow, length more or less twice of width; a few long pointed hairs running over ocular mound in a longitudinal row; chelicerae robust, with promargin of cheliceral groove with ten strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, with tergites 2–6 larger than others, and with tergite 4 being largest; seven spinnerets. Measurements: BL 9.80, CL 4.38, CW 4.20, OL 5.6, and OW 5.00; ALE> PLE> PME> AME; palp 7.63 (2.50 + 1.31 + 1.65 + 2.17), leg I missing, leg II 8.49 (2.71 + 1.50 + 1.68 + 1.50 + 1.10), leg III 9.34 (2.52 + 1.58 + 1.56 + 2.30 + 1.38), and leg IV 13.70 (3.75 + 1.60 + 2.55 + 3.68 + 2.12). Female genitalia Posterior part of genital area W-shaped (Fig. 9A, B), with pair of receptacular clusters with short stalks on the anterior part of genital separated from one another (Fig. 9A, B). Distribution Fujian (Putian) Province, China. Remarks See comments under G. wangjiangensis sp. nov., Published as part of Xu, Xin, Liu, Fengxiang, Chen, Jian, Li, Daiqin & Kuntner, Matjaž, 2015, Integrative taxonomy of the primitively segmented spider genus Ganthela (Araneae: Mesothelae: Liphistiidae): DNA barcoding gap agrees with morphology, pp. 288-306 in Zoological Journal of the Linnean Society 175 (2) on pages 302-303, DOI: 10.1111/zoj.12280, http://zenodo.org/record/5338452

GANTHELA VENUS XU SP. NOV. (FIG. 9) Holotype Female (XUX-2013-160), Mount Wangjiang, Huangyang Village, Zhuangbian Town, Hanjiang District, Putian City, Fujian Province, China, 25.66°N, 118.88°E, 459 m a.s.l., 11 July 2013, collected by F.X. Liu, X. Xu, and Z. T. Zhang. No male found. Etymology Named after the Greek goddess of love, Venus. Diagnosis Females of G. venus sp. nov. can be distinguished from G. wangjiangensis sp. nov. by details in the genitalia, which have separated stalks on the anterior part (Fig. 9A, B). Ganthela venus sp. nov. furthermore differs from all other Ganthela species by the following unique nucleotide substitutions in the standard DNA barcode alignment: C (36), A (80), T (227), C (333), C (341), G (347), T (354), C (431), and T (539). Description Female (holotype). Carapace and opisthosoma light brown; tergites slightly dark brown; sternum narrow, length more or less twice of width; a few long pointed hairs running over ocular mound in a longitudinal row; chelicerae robust, with promargin of cheliceral groove with ten strong denticles of variable size; legs with strong hairs and spines; opisthosoma with 12 tergites, with tergites 2–6 larger than others, and with tergite 4 being largest; seven spinnerets. Measurements: BL 9.80, CL 4.38, CW 4.20, OL 5.6, and OW 5.00; ALE> PLE> PME> AME; palp 7.63 (2.50 + 1.31 + 1.65 + 2.17), leg I missing, leg II 8.49 (2.71 + 1.50 + 1.68 + 1.50 + 1.10), leg III 9.34 (2.52 + 1.58 + 1.56 + 2.30 + 1.38), and leg IV 13.70 (3.75 + 1.60 + 2.55 + 3.68 + 2.12). Female genitalia Posterior part of genital area W-shaped (Fig. 9A, B), with pair of receptacular clusters with short stalks on the anterior part of genital separated from one another (Fig. 9A, B). Distribution Fujian (Putian) Province, China. Remarks See comments under G. wangjiangensis sp. nov.