Pinelema bailongensis Wang & Li, 2012 from Guangxi, China.

Pinelema can be distinguished from Telema by the following: belt-shaped tibial glands (arrows on Fig. 1B) (vs. plate-shaped), the presence of a cymbial apophysis (vs. absent), and the triangular, trapezoidal, or tube-like embolus (vs. duckbill shaped). The endogyne is extended distally (vs. no extension).

Figure 1. 

Tibial glands on leg III of telemids in East and Southeast Asia (Zhuanlema gen. nov. is not included). A Telema guihua B Pinelema bailongensis C Mekonglema xinpingi comb. nov. D Siamlema changhai sp. nov. E Sundalema anguina comb. nov. F Apneumonella jacobsoni G Telemofila samosirensis.

Total length: 1.07–1.77 (male), 1.20–2.02 (female). Carapace 0.48–0.82 long. Sternum with several long setae. Six eyes ringed with black, vestigial, or completely absent in some species. Legs thin, long relative to body. Leg formula: 1-2-4-3. Tibia I 0.94–2.20 long, glands of legs belt-shaped (Fig. 1B). Cymbial apophysis distinct, bulb large relative to body, oval, kidney-shaped, droplet-shaped, etc.; embolus triangular, tube-shaped, needle-shaped, etc., embolus short, medium, or long relative to cymbium. Receptacle with several membranous tubes inside and typically swollen distally.

China (Guangxi, Guizhou, Yunnan, Hainan), Laos (Vien Tiane), and Vietnam (northern and central Vietnam) (Figs 30–32).

All 26 new combinations are supported by morphological characters, such as the presence of a cymbial apophysis and the extended tip of the receptacle. Including the new species described in this paper increases the total number of Pinelema species to 54, making it the most speciose genus in Telemidae.

According to the morphological characters of the male palp, 54 Pinelema species have been divided into seven species groups as well as six species not attached to a species group. The composition of species groups is discussed below.

This group resembles the pacchanensis-group by having a long bulb but can be distinguished by the bulb is protruding ventro-basally and concave dorso-sub-basally (Fig. 2A) (vs. concave ventro-sub-basally and protruding dorso-mesially).

Body length 1.40–1.98. Carapace 0.60–0.82 long. Six eyes ringed with black or absent. Tibia I 1.40–2.20. Ratio of bulbal length/width 1.96–2.35, bulb protruding ventro-basally and concave dorso-subbasally, embolus membranous (except in P. tortutheca (Lin & Li) comb. nov.), length ratio of embolus/bulb 0.38–0.71. Receptacle J-shaped.

China (Guangxi, sites 1–5 in Fig. 30).

Pinelema adunca (Wang & Li, 2010) comb. nov., P. qingfengensis Zhao & Li, 2017, P. renalis (Wang & Li, 2010) comb. nov., P. tortutheca comb. nov., and P. yashanensis (Wang & Li, 2010) comb. nov.

Figure 2. 

Pinelema spp., palp, retrolateral view. A The adunca-group B the bailongensis-group C the cunfengensis-group.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Pingguo County, Bafeng Mountain, Guandi Cave, 23.5670N, 107.6794E, elevation ca. 285 m, 3.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema adunca comb. nov. resembles P. yashanensis comb. nov. but can be distinguished by the following: the smaller length ratio of the embolus/bulb (0.64, Fig. 2A, and cf. Wang and Li 2010b: fig. 1A, B) (vs. 0.85), the cymbial apophysis which is as long as the cymbial base (cf. Wang and Li 2010b: fig. 1B) (vs. ¼ times shorter); the receptacle tip is 3.20 times wider than the neck (cf. Wang and Li 2010b: fig. 3C) (vs. 1.20 times).

See Wang and Li (2010b).

China (Guangxi, site 1 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Chongzuo Prefecture, Tiandeng County, Qingfeng Cave, 23.1720N, 107.1565E, elevation ca. 444 m, 26.XII.2012, Z. Chen and Z. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema qingfengensis resembles P. renalis comb. nov. but can be distinguished by the following: the absence of eyes (cf. Song et al. 2017: fig. 5A) (vs. present); the ventrally bent embolus, and the obtuse-angled bulb (Fig. 2A, and cf. Song et al. 2017: fig. 5C, D) (vs. right-angled), the bulb is not protruding ventro-distally (Fig. 2A, and cf. Song et al. 2017: fig. 5C, D) (vs. protruding), and the width ratio of the basal embolus/distal bulb is 1.0 (cf. Song et al. 2017: fig. 5C, D) (vs. 0.5).

See Song et al. (2017).

China (Guangxi, site 2 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hezhou Prefecture, Zhongshan County, Guanyin Cave, 24.5233N, 111.3242E, elevation ca. 196 m, 25.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema renalis comb. nov. resembles P. qingfengensis but can be distinguished by the following: six eyes ringed with black (cf. Wang and Li 2010b: fig. 24A) (vs. no eyes); the embolus bent ventrally at a right angle (Fig. 2A, and cf. Wang and Li 2010b: fig. 24C, D) (vs. obtuse angle), the bulb protrudes ventro-distally (Fig. 2A, and cf. Wang and Li 2010b: fig. 24C, D) (vs. not protruding), and the width ratio of the basal embolus/distal bulb is 0.5 (cf. Wang and Li 2010b: fig. 24C, D) (vs. 1.0).

See Wang and Li (2010b).

China (Guangxi, site 3 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Nanning Prefecture, Mashan County, Guling Town, Yangyu Village, Jinlun Cave, 23.5663N, 107.2622E, elevation ca. 1490 m, 6.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema tortutheca comb. nov. can be distinguished from all four other species in this group by the sclerotized embolus (Fig. 2A, and cf. Lin and Li 2010: fig. 16C) (vs. membranous).

See Lin and Li (2010).

China (Guangxi, site 4 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Laibin Prefecture, Yashan County, Yashan Cave, 23.6025N, 108.9124E, elevation ca. 115 m, 16.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema yashanensis comb. nov. resembles P. adunca comb. nov. but can be distinguished by the following: the larger length ratio of the embolus/bulb (0.85, Fig. 2A, and cf. Wang and Li 2010b: fig. 28A, B) (vs. 0.64); the cymbial apophysis ¼ as long as the cymbial base (cf. Wang and Li 2010b: fig. 28A) (vs. equal length); and the receptacle tip is 1.20 times wider than the neck (cf. Wang and Li 2010b: fig. 30C) (vs. 3.20 times).

See Wang and Li (2010b).

China (Guangxi, site 5 in Fig. 30), known only from the type locality.

This group can be distinguished from other species groups in Pinelema by the following: the length ratio of the embolus/bulb is 1.19–1.80 (Fig. 2B) (vs. 0.37–0.84), the junction of the bulb and cymbium is located ventro-mesially on the bulb (Fig. 2B) (vs. ventro-basally, except the cunfengensis-group).

Body length 1.11–1.75. Carapace 0.50–0.82 long. Tibia I 0.90–2.18 long. Six eyes ringed with black or absent. Embolus longer than bulb, length ratio of embolus/bulb is 1.19–1.80, junction of bulb and cymbium located ventro-mesially on bulb (Fig. 2B), bulb with papillae proximo-retrolaterally (except P. curcici Wang & Li, 2016, P. huoyan Zhao & Li, 2018, and P. xiushuiensis Wang & Li, 2016). Endogyne U-shaped, J-shaped, or spiralled.

China (Guangxi, Guizhou, Yunnan, sites 6–18 in Fig. 30).

Pinelema bailongensis, P. cheni Zhao & Li, 2018, P. cordata (Wang & Li, 2010), P. curcici, P. huoyan, P. liangxi (Zhu & Chen, 2002), P. lizhuang Zhao & Li, 2018, P. strentarsi (Lin & Li, 2010), P. wangshang Zhao & Li, 2018, P. wenyang Zhao & Li, 2018, P. xiushuiensis, P. yunchuni Zhao & Li, 2018, and P. zhewang (Chen & Zhu, 2009).

For the diagnoses and descriptions of this group (except P. bailongensis and P. curcici), see Zhao et al. (2018b).

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Pingguo County, Bailong Cave, 23.3182N, 107.5731E, elevation ca. 111 m, 1.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

For differences between P. bailongensis and P. curcici, see Wang and Li (2016); for differences between P. bailongensis and the other eleven species in this group, see Zhao et al. (2018b).

Tibial glands belt-shaped (Fig. 1B), the arrangement of secretory orifices linear within a smooth, striped tegument (arrows on Fig. 1B). For a more detailed description, see Wang and Li (2012), Song et al. (2017), and Zhao et al. (2018b).

China (Guangxi, site 6 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Qiubei County, Shuanglongying Town, Fengwei Cave, 24.3361N, 104.2862E, elevation ca. 1372 m, 20.VIII.2010, Z. Yao, C. Wang and X. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema curcici resembles P. liangxi but can be distinguished by the following: the presence of eyes (vs. absent); the embolic tip is much narrower than the embolic base (Fig. 2B) (vs. equal), the smaller ratio of the bulbal length/width (1.13, Fig. 2B, and cf. Wang and Li 2016: fig. 1C, D) (vs. 1.27), the greater length ratio of the embolus/bulb (1.31, Fig. 2B, and cf. Wang and Li 2016: fig. 1C, D) (vs. 1.19); the J-shaped receptacle (vs. C-shaped), with the distal part of the receptacle 1.75 times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 2C) (vs. three times).

See Wang and Li (2016).

China (Yunnan, site 9 in Fig. 30), known only from the type locality.

This group resembles the feilong-group by having a triangular and short embolus relative to the bulb length but can be distinguished by the following: the bulb is bent at a right angle dorso-distally (Fig. 2C) (vs. not bent), and the junction of the bulb and cymbium is located dorso-mesially on the bulb (Fig. 2C) (vs. dorso-basally); the receptacle is U-shaped or coiled (vs. bag-like or globular).

Body length 1.33–1.48. Carapace 0.48–0.64 long. Tibia I 1.15–1.98 long. Six eyes ringed with black (P. cunfengensis Zhao & Li, 2017) or absent (P. spirae (Lin & Li, 2010) comb. nov.). Bulb bent dorso-distally at a right angle (Fig. 2C), ratio of bulbal length/width 1.22–1.39, length ratio of embolus/bulb 0.38–0.58, embolus triangular. Receptacle U-shaped (P. cunfengensis) or coiled (P. spirae comb. nov.).

China (Guangxi, sites 19–20 in Fig. 30).

Pinelema cunfengensis and P. spirae comb. nov.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Nanning Prefecture, Long’an County, Nanxu Town, Nawan Village, Feng Cave, 23.2098N, 107.5906E, elevation ca. 115 m, 13.V.2015, Z. Chen and Y. Li. leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema cunfengensis resembles P. spirae comb. nov. but can be distinguished by the following: the presence of eyes (cf. Song et al. 2017: fig. 1A) (vs. absence); the different shape of the embolus (Fig. 2C, and cf. Song et al. 2017: fig. 1C, D), the ratio of bulbal length/width is 1.22 (vs. 1.39), and the length ratio of the embolus/bulb is 0.58 (Fig. 2C, and cf. Song et al. 2017: fig. 1C, D) (vs. 0.38); the U-shaped receptacle (cf. Song et al. 2017: fig. 2C) (vs. coiled).

See Song et al. (2017).

China (Guangxi, site 19 in Fig. 30), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Bama County, Poyue Town, Poyue Village, Dawan Cave, 24.3016N, 107.1155E, elevation ca. 438 m, 16.VIII.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema spirae comb. nov. resembles P. cunfengensis but can be distinguished by the following: the absence of eyes (vs. presence); the different shape of the embolus (Fig. 2C), the ratio of bulbal length/width is 1.39 (vs. 1.22), and the length ratio of the embolus/bulb is 0.38 (Fig. 2C) (vs. 0.58); the coiled receptacle (cf. Lin and Li 2010: fig. 13B, C) (vs. U-shaped).

See Lin and Li (2010).

China (Guangxi, site 20 in Fig. 30), known only from the type locality.

This group resembles the xiezi-group by the short embolus relative to the bulb but can be distinguished by the triangular shape of the embolus (Fig. 3A) (vs. trapezoidal).

Body length 0.98–1.85. Carapace length 0.47–0.76. Tibia I length 0.85–2.20. Six eyes ringed with black (P. bella (Tong & Li, 2008) comb. nov., P. damtaoensis Zhao & Li, 2018, and P. spina comb. nov.) or absent (other species in this group), bulb oval, the junction of the bulb and the cymbium is located ventro-basally, the embolus is triangular, and the length ratio of the embolus/bulb is 0.33–0.49. The receptacle is stick-shaped (P. bella comb. nov., P. claviformis comb. nov., and P. feilong (Chen & Zhu, 2009) comb. nov.), boot-shaped (P. circularis (Tong & Li, 2008) comb. nov.), or globular.

China (Guangxi, Yunnan, Hainan) and Vietnam (Vinh Phuc) (Fig. 31).

Pinelema bella comb. nov., P. circularis comb. nov., P. claviformis comb. nov., P. damtaoensis, P. feilong comb. nov., P. huobaensis Wang & Li, 2016, P. spina comb. nov., P. vesiculata (Lin & Li, 2010) comb. nov., and P. yaosaensis Wang & Li, 2016.

Figure 3. 

Pinelema spp., palp, retrolateral view. A The feilong-group B the pacchanensis-group C the podiensis-group. P. claviformis comb. nov. and P. spina comb. nov. are modified from Tong and Li 2008b and 2008a, respectively.

Holotype: ♂ (IZCAS), China, Hainan Province, Dongfang County, Datian National Natural Reserve, Mihou Cave, 18.9114N, 109.0639E, elevation ca. 293 m, 28.V.2005, Y. Song, X. Han, G. Deng and Y. Tong leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema bella comb. nov. resembles P. spina comb. nov. but can be distinguished by the following: the larger ratio of the bulbal length/width (1.75, Fig. 3A, and cf. Tong and Li 2008a: fig. 2C) (vs. 1.45); the obtuse-angled ventral bend between the embolus and bulb (Fig. 3A, and cf. Tong and Li 2008a: fig. 2B, C) (vs. right-angled), and the blunt tip of the embolus (Fig. 3A, and cf. Tong and Li 2008a: fig. 2B, C) (vs. sharp).

See Tong and Li (2008a).

China (Hainan, site 1 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guizhou Province, Anshun Prefecture, Guanling County, Shangguan Town, Ganzhishu Cave, 25.8120N, 105.6716E, elevation ca. 987 m, 8.V.2005, Y Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema circularis comb. nov. resembles P. claviformis comb. nov. but can be distinguished by the following: the sharp tip of the embolus (Fig. 3A, and cf. Tong and Li 2008b: fig. 2B, C) (vs. blunt); the boot-shaped receptacle (cf. Tong and Li 2008b: fig. 2D, E) (vs. stick-shaped).

See Tong and Li (2008b).

China (Guizhou, site 2 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guizhou Province, Qianxinan Buyei and Miao Autonomous Prefecture, Xingyi County, Maling Town, Qiuxiang Cave, 25.2000N, 104.8833E, 12.V.2005, Y. Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS). Examined.

Pinelema claviformis comb. nov. resembles P. circularis comb. nov. but can be distinguished by the following: the blunt tip of the embolus (Fig. 3A) (vs. sharp); the stick-shaped receptacle (cf. Tong and Li 2008b: fig. 3D, E) (vs. boot-shaped).

See Tong and Li (2008b).

China (Guizhou, site 3 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), Vietnam, Vinh Phuc Province, Dam Tao National Park, leaf litter, 21.4600N, 105.6480E, elevation ca. 999 m, 1.XI.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema damtaoensis resembles P. bella comb. nov. but can be distinguished by the following: a black spot and radial stripes on the carapace (cf. Zhao et al. 2018a: fig. 1A) (vs. carapace without pattern); the spiral ridge of the embolus is sclerotized (Fig. 3A, and cf. Zhao et al. 2018a: fig. 1B) (vs. membranous); the receptacle is globular (cf. Zhao et al. 2018a: fig. 3C) (vs. stick-shaped).

See Zhao et al. (2018a).

Vietnam (Vinh Phuc, site 4 in Fig. 31), known only from the type locality.

Holotype: ♂ (MLR), China, Guizhou Province, Qianxinan Buyei and Miao Autonomous Prefecture, Xingyi County, Feilong Cave, 24.9166N, 104.8833E, elevation ca. 1335 m, 25.V.2004, H. Chen and Y. Zhang leg. Paratypes: 3♂ and 1♀ (MHBU), same data as holotype. Not examined.

1♂ and 2♀ (including molecular voucher, IZCAS), same data as the type locality, 8.III.2011, C. Wang and L. Lin leg.

Pinelema feilong comb. nov. resembles P. vesiculata comb. nov. but can be distinguished by the following: the smaller ratio of bulbal length/width (1.31, Fig. 3A, and cf. Chen and Zhu 2009: fig. 2C,) (vs. 1.44), the smaller length ratio of the embolus/bulb (0.36, Fig. 3A, and cf. Chen and Zhu 2009: fig. 2C, D) (vs. 0.48); the unmodified stick-shaped receptacle (cf. Chen and Zhu 2009: fig. 2J) (vs. receptacle with a vesicle ventro-distally).

See Chen and Zhu (2009).

China (Guizhou, site 5 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Qiubei County, Shuanglongying Town, Puzhehei Village, Huoba Cave, 24.1385N, 104.1126E, elevation ca. 1457 m, 18.VIII.2009, Z. Yao, C. Wang and X. Wang leg. Paratypes: 1♂ and 1♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema huobaensis resembles P. yaosaensis but can be distinguished by the following: the bulb protrudes ventro-subdistally (Fig. 3A, and cf. Wang and Li 2016: fig. 5B, C) (vs. protrudes dorso-distally); the distal part of the receptacle is five times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 6C) (vs. eight times).

See Wang and Li (2016).

China (Yunnan, site 6 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Hainan Province, Wuzhishan County, Wuzhi Mountain, leaf litter, 18.8167N, 109.6500E, Y. Song leg. Examined.

Pinelema spina comb. nov. resembles P. bella comb. nov. but can be distinguished by the following: the smaller ratio of the bulbal length/width (1.45, Fig. 3A) (vs. 1.75); the obtuse-angled ventral bend between the embolus and bulb (Fig. 3A) (vs. right-angled), the sharp tip of the embolus (Fig. 3A) (vs. blunt).

See Tong and Li (2008a).

China (Hainan, site 7 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Yunnan Province, Qujing Prefecture, Luoping County, Luoxiong Town, Pingtian Village, Laobie Cave, 24.8419N, 104.2678E, elevation ca. 1490 m, 20.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema vesiculata comb. nov resembles P. feilong comb. nov. but can be distinguished by the following: the larger ratio of the bulbal length/width (1.44, Fig. 3A, and cf. Lin and Li 2010: fig. 18D) (vs. 1.31), the larger length ratio of the embolus/bulb (Fig. 3A, and 0.48, cf. Lin and Li 2010: fig. 18E) (vs. 0.36); the receptacle with a vesicle ventro-distally (cf. Lin and Li 2010: fig. 20B–D) (vs. unmodified).

See Lin and Li (2010).

China (Yunnan, site 8 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Xichou County, Niuchangba Township, Mosa Village, Yaosa Cave, 23.5072N, 104.9023E, elevation ca. 1297 m, 6.VIII.2009, Z. Yao, C. Wu and X. Wang leg. Paratypes: 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema yaosaensis resembles P. huobaensis but can be distinguished by the following: the bulb does not protrude ventro-subdistally but protrudes dorso-distally (Fig. 3A, and cf. Wang and Li 2016: fig. 13B, C) (vs. protruding ventro-subdistally but not dorso-distally); the distal part of the receptacle is eight times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 14C) (vs. five times).

See Wang and Li (2016).

China (Yunnan, site 9 in Fig. 31), known only from the type locality.

This group resembles the adunca-group by having a long bulb but can be distinguished by the bulb being slightly concave ventrally and protruding dorso-mesially (Fig. 3B) (vs. protrude ventro-basally and concave dorso-subbasally).

Body length 1.33–1.60. Carapace 0.56–0.71 long. Eyes absent. Tibia I 1.33–2.00 long. Ratio of bulbal length/width 1.80–2.11, bulb slightly concave ventrally and protruding dorso-mesially (Fig. 3B), embolus triangular or tube-like. Receptacle swollen distally and globular.

China (Guizhou) and Vietnam (Bac Kan) (sites 10–11 in Fig. 31)

Pinelema daguaiwan sp. nov. and P. pacchanensis Zhao & Li, 2018.

Holotype: ♂ (IZCAS), China, Guizhou Province, Qianxinan Buyi and Miao Autonomous Prefecture, Wangmo County, Sanglang Town, Xinghe Village, Daguaiwan Cave. 25.2706N, 106.4328E, elevation ca. 886 m, 29.XII.2010, Z. Zha and Z. Chen leg. Paratypes: 1♂ and 5♀ (IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

Pinelema daguaiwan sp. nov. resembles P. pacchanensis by the bulb being concave ventrally but can be distinguished by the following: the bulb is more deeply concave (arrow in Fig. 4C, D) (vs. weaker), the smaller length ratio of the embolus/bulb (0.48, Figs 3B, 4C, D) (vs. 0.71), the triangular embolus (Figs 3B, 4C, D) (vs. tube-shaped); the diameter ratio of the receptacle tip/neck is 7.0 (Fig. 5A) (vs. 2.0).

Figure 4. 

Pinelema daguaiwan sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Male (holotype). Total length 1.60. Carapace 0.71 long, 0.67 wide. Abdomen 0.85 long, 0.76 wide. Carapace light brown (Fig. 4A). Eyes absent (Fig. 4A). Chelicerae, legs, labium, and endites brown. Sternum light brown with sparse setae. Leg measurements: I 6.25 (1.80, 0.26, 2.00, 1.41, 0.78); II 5.46 (1.56, 0.26, 1.74, 1.19, 0.71); III (1.20, 0.25, 1.14, –, –); IV 4.96 (1.52, 0.24, 1.44, 1.13, 0.63). Abdomen pale brown (Fig. 4A).

Palp. Tibia two times longer than patella, cymbium 2.47 times longer than tibia, 1.88 times longer than femur, cymbial apophysis finger shaped (Fig. 4C); bulb concave ventrally, and shaped as in Fig. 4C, D, length ratio of bulb/cymbium about 0.9; embolus triangular and membranous, 1/5 as long as bulb (Fig. 4C, D), spiral ridge brown (Fig. 4B).

Female. Total length 1.72. Carapace 0.66 long, 0.59 wide. Abdomen 1.04 long, 0.87 wide. Coloration as in male (Fig. 5B, C). Leg measurements: I 5.33 (1.56, 0.24, 1.68, 1.15, 0.70); II 4.59 (1.33, 0.24, 1.41, 0.99, 0.62); III 3.37 (1.05, 0.22, 0.94, 0.67, 0.49); IV 4.15 (1.25, 0.22, 1.21, 0.91, 0.56). Abdomen pale brown (Fig. 5B, C). Neck of receptacle membranous, as long as diameter of receptacle (Fig. 5A); receptacle tip globular with several membranous tubes, much wider than neck (Fig. 5A).

China (Guizhou, site 10 in Fig. 31), known only from the type locality.

Figure 5. 

Pinelema daguaiwan, sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Holotype: ♂ (IZCAS), Vietnam, Bac Kan Province, Cho Don District, Pac Chan Cave, 22.3790N, 105.6130E, elevation ca. 225 m, 18.X.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema pacchanensis resembles P. daguaiwan sp. nov. but can be distinguished by the following: the ventrally concave part of the bulb is weaker (Fig. 3B, and cf. Zhao et al. 2018a: fig. 11A, B) (vs. stronger), the larger length ratio of the embolus/bulb (0.71, Fig. 3B, and cf. Zhao et al. 2018a: fig. 11A, B) (vs. 0.48), and the triangular embolus (cf. Zhao et al. 2018a: fig. 11 A–D) (vs. tube-shaped); the diameter ratio of the receptacle tip/neck is 2.0 (cf. Zhao et al. 2018a: fig. 12C) (vs. 7.0).

See Zhao et al. (2018a).

Vietnam (Bac Kan, site 11 in Fig. 31), known only from the type locality.

This group resembles the feilong-group by the embolus which is short relative to the bulb but can be distinguished by the following: the length ratio of the embolus/bulb ranges from 0.57 to 0.79 (Fig. 3C) (vs. 0.33 to 0.49), and the shape of the embolus is a long isosceles triangle (Fig. 3C) (vs. equilateral triangle).

Body length 1.22–1.75. Carapace 0.48–0.75 long. Tibia I 1.15–1.87 long. Six eyes ringed with black (P. biyunensis (Wang & Li, 2010) comb. nov., P. shiba sp. nov., and P. zonaria (Wang & Li, 2010) comb. nov.), vestigial (P. podiensis Zhao & Li, 2017), or absent (P. bifida (Lin & Li, 2010) comb. nov.). Ratio of bulbal length/width 1.31–1.67, embolus shorter than bulb, length ratio of embolus/bulb 0.57–0.79, embolic shape long isosceles triangle (Fig. 3C). Receptacle J-shaped, slightly swollen distally.

China (Guangxi, sites 10–14 in Fig. 31).

Pinelema bifida comb. nov., P. biyunensis comb. nov., P. podiensis, P. shiba sp. nov., and P. zonaria comb. nov.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Dahua County, Qibainong Town, Qiaoxu Village, Qiaoxu Cave, 24.0761N, 107.6706E, elevation ca. 550 m, 9.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

. Pinelema bifida comb. nov. resembles P. zonaria comb. nov. but can be distinguished by the following: the absence of eyes (vs. present); the bulb does not protrude ventro-distally (Fig. 3C, and cf. Lin and Li 2010: fig. 2C, D) (vs. protruding), the ratio of bulbal length/width is larger (1.43, cf. Lin and Li 2010: fig. 2C, D) (vs. 1.31).

See Lin and Li (2010).

China (Guangxi, site 12 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hezhou Prefecture, Zhongshan County, Biyun Cave, 24.357N, 111.1923E, elevation ca. 131 m, 25.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema biyunensis comb. nov. resembles P. zonaria comb. nov. but can be distinguished by the following: the larger ratio of bulbal length/width (1.59, Fig. 3C, and cf. Wang and Li 2010b: fig. 7C, D) (vs. 1.31), the smaller length ratio of the embolus/bulb (0.57, Fig. 3C, and cf. Wang and Li 2010b: fig. 7C, D) (vs. 0.76); the tip of the receptacle is 2.10 times wider than the neck (cf. Wang and Li 2010b: fig. 8C) (vs. 1.10 times).

See Wang and Li (2010b).

China (Guangxi, site 13 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Debao County, Podi Cave, 23.3919N, 106.6400E, elevation ca. 578 m, 4.VIII.2011, C. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema podiensis resembles P. shiba sp. nov. but can be distinguished by the following: the weakly concave dorsum of the bulb (Fig. 3C, and cf. Song et al. 2017: fig. 3C, D) (vs. strongly concave), the larger bulbal length/width ratio (1.75, Fig. 3C, and cf. Song et al. 2017: fig. 3C, D) (vs. 1.33); the tip of the receptacle is 2.20 times wider than the neck (cf. Song et al. 2017: fig. 4C) (vs. 4.10 times).

See Song et al. (2017).

China (Guangxi, site 14 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS); China, Guangxi Zhuang Autonomous Region, Chongzuo Prefecture, Daxin County, Hucheng Town, Baoxian Village, Shiba Cave. 22.8133N, 107.1632E, elevation ca. 157 m. 17.XI.2011, C. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

Pinelema shiba sp. nov. resembles P. podiensis but can be distinguished by the following: the strongly concave dorso-mesial part of the bulb (Fig. 6C, D) (vs. weakly concave), the ratio of the bulbal length/width is 1.33 (Fig. 6C, D) (vs. 1.75); the tip of the receptacle is 4.10 times wider than the neck (Fig. 7C) (vs. 2.20 times).

Male (holotype). Total length 1.22. Carapace 0.51 long, 0.48 wide. Abdomen 0.71 long, 0.63 wide. Carapace brown (Fig. 6A). Six vestigial eyes (Fig. 6A). Chelicerae, legs, labium, and endites light brown (Fig. 6A). Sternum light brown with sparse setae. Leg measurements: I 5.00 (1.47, 0.21, 1.60, 1.06, 0.66); II 4.26 (1.28, 0.21, 1.33, 0.88, 0.56); III 3.05 (0.97, 0.18, 0.89, 0.56, 0.45); IV 3.55 (1.23, 0.17, 1.00, 0.65, 0.50). Abdomen pale yellow with a few long setae.

Palp. Tibia 2.30 times longer than patella, cymbium 1.91 times longer than tibia, 1.90 times longer than femur, cymbial apophysis short, as long as 1/4 width of cymbium base (Fig. 6C); bulb strongly concave dorsally (arrow in Fig. 6C, D); embolus triangular and half as long as cymbium (Fig. 6C, D), spiral ridge brown (Fig. 6B).

Figure 6. 

Pinelema shiba sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Female. Total length 1.39. Carapace 0.54 long, 0.50 wide. Abdomen 0.82 long, 0.73 wide. Six eyes ringed with black (Fig. 7B). Coloration similar to male (Fig. 7B, C). Leg measurements: I 4.94 (1.52, 0.21, 1.56, 1.00, 0.65); II 4.18 (1.31, 0.20, 1.27, 0.85, 0.55); III 3.08 (1.00, 0.19, 0.91, 0.53, 0.45); IV 3.92 (1.25, 0.19, 1.19, 0.78, 0.51). Abdomen brown (Fig. 7B, C). Receptacle U-shaped with several membranous tubes, distally swollen (Fig. 7A).

Figure 7. 

Pinelema shiba sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

China (Guangxi, site 15 in Fig. 31), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Yizhou County, Xiannv Cave, 24.4887N, 108.5701E, elevation ca. 205 m, 28.VII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema zonaria comb. nov. resembles P. bifida comb. nov. but can be distinguished by the following: eyes are present (vs. absent); the bulb protrudes ventro-distally (Fig. 3C, and cf. Wang and Li 2010b: fig. 33C, D) (vs. not protrude), the ratio of the bulbal length/width is smaller (1.31, Fig. 3C, and cf. Wang and Li 2010b: fig. 33C, D) (vs. 1.43).

See Wang and Li (2010b).

China (Guangxi, site 16 in Fig. 31), known only from the type locality.

This group resembles the feilong-group by the short embolus relative to the bulb but can be distinguished by the trapezoidal shape of the embolus (Fig. 8) (vs. triangular).

Body length 0.98–2.05. Carapace 0.51–0.82 long. Tibia I 0.81–2.13 long. Six eyes ringed with black, vestigial (P. exiloculata (Lin, Pham & Li, 2009) comb. nov.), or absent, bulb oval, junction of bulb and cymbium located ventro-basally on bulb, embolus trapezoidal, length ratio of embolus/bulb 0.30–0.59. Receptacle J-shaped.

China (Guangxi, Guizhou, Hainan) and Vietnam (Hai Phong, Ninh Binh, Phu Tho, Quang Binh) (Sites 1–12 in Fig. 32).

Pinelema breviseta (Tong & Li, 2008) comb. nov., P. conglobare (Lin & Li, 2010) comb. nov., P. cucphongensis (Lin, Pham & Li, 2009) comb. nov., P. cucurbitina (Wang & Li, 2010) comb. nov., P. dongbei (Wang & Ran, 1998) comb. nov., P. exiloculata comb. nov., P. grandidens (Tong & Li, 2008) comb. nov., P. laensis Zhao & Li, 2018, P. oculata (Tong & Li, 2008) comb. nov., P. pedati (Lin & Li, 2010) comb. nov., P. spinafemora (Lin & Li, 2010) comb. nov., and P. xiezi Zhao & Li, 2018.

Figure 8. 

Pinelema spp., palp, retrolateral view. The xiezi-group. P. spinafemora comb. nov. is modified from Lin and Li 2010.

Holotype: ♂ (IZCAS), China, Hainan Province, Dongfang County, Donghe Town, Yalong Village, Yalong Cave, 18.9770N, 108.8935E, elevation ca. 273 m, Y. Song, X. Han, G. Deng and Y. Tong leg. 1.IV.2005. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema breviseta comb. nov. resembles P. laensis but can be distinguished by the following: the ventro-distal bulb does not protrude (Fig. 8, and cf. Tong and Li 2008a: fig. 3B, C) (vs. protrude), the embolus lacks modification retrolaterally (Fig. 8, and cf. Tong and Li 2008a: fig. 3B, C) (vs. embolus with a vertical groove retrolaterally); the tip of the receptacle is three times wider than the neck (cf. Tong and Li 2008a: fig. 3H) (vs. four times).

See Tong and Li (2008a).

China (Hainan, site 1 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Fengshan County, Fengcheng Town, Songren Village, Xi’an Cave, 24.5657N, 107.0411E, elevation ca. 574 m, 12.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema conglobare comb. nov. resembles P. pedati comb. nov. but can be distinguished by the following: the larger ratio of bulbal length/width (1.77, Fig. 8, and cf. Lin and Li 2010: fig. 4D, E) (vs. 1.60), the ventral bend of the embolus and bulb is right-angled (Fig. 8, and cf. Lin and Li 2010: fig. 4D, E) (vs. acute-angled); the distal part of the receptacle is globular (cf. Lin and Li 2010: fig. 5B–E) (vs. boot-shaped).

See Lin and Li (2010).

China (Guangxi, site 2 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), Vietnam, Cuc Phuong National Park, Prehistoric Man Cave, 20.2930N, 105.6670E, elevation ca. 256 m, 19.VII.2008, S. Li leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema cucphongensis comb. nov. resembles P. xiezi but can be distinguished by the following: a pair of lateral scutae on abdomen (cf. Lin et al. 2009: fig. 5H, I) (vs. absent); the different shape of the embolus, the larger ratio of the bulbal length/width (1.49, Fig. 8, and cf. Lin et al. 2009: fig. 5A–C) (vs. 1.28), and the smaller length ratio of the embolus/bulb (0.36, Fig. 8, and cf. Lin et al. 2009: fig. 5A–C) (vs. 0.63); the distal part of the receptacle is seven times wider than the neck of the receptacle (cf. Lin et al. 2009: fig. 6F) (vs. five times).

See Lin et al. (2009).

. Vietnam (Cuc Phuong National Park, site 3 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Guilin Prefecture, Lingui County, Shuixian Cave, 25.2137N, 110.2008E, elevation ca. 161 m, 18.VII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema cucurbitina comb. nov. resembles P. spinafemora comb. nov. but can be distinguished by the following: the bulb does not protrude ventro-distally (Fig. 8, and cf. Wang and Li 2010b: fig. 16C, D) (vs. protrude), the ventral bend between the embolus and the bulb is acute-angled (Fig. 8, and cf. Wang and Li 2010b: fig. 16C, D) (vs. right-angled).

See Wang and Li (2010b).

China (Guangxi, site 4 in Fig. 32).

Holotype: ♂ (AMNH), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Libo County, Yuping Town, Dongbei Cave, 25.4804N, 107.8959E, elevation ca. 812 m, 13.IX.1996, J. Ran leg. Paratype: 1♀ (AMNH), same data as holotype. Not examined.

1♂ and 2♀ (including molecular voucher, IZCAS), from the type locality, 18.III.2011, C. Wang and L. Lin leg.

Pinelema dongbei comb. nov. resembles P. exiloculata comb. nov. and P. oculata comb. nov. but can be distinguished by the following: eyes are absent (vs. vestigial); the embolus protrudes dorsally (Fig. 8, and cf. Wang and Ran 1998: figs 1, 2) (vs. not protruding); the receptacle is slightly swollen distally (cf. Wang and Ran 1998: fig. 5) (vs. distinctively swollen).

See Wang and Ran (1998).

China (Guizhou, site 5 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), Vietnam, Hai Phong Province, Cat Ba National Park, Trung Trang Cave, 20.8000N, 106.9833E, elevation ca. 256 m, 16.VII.2008, S. Li leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema exiloculata comb. nov. resembles P. dongbei comb. nov. but can be distinguished by the following: vestigial eyes (vs. absent); the embolus does not protrude dorsally (Fig. 8, and cf. Lin et al. 2009: fig. 7B) (vs. protruding); the receptacle is distinctively swollen distally (cf. Lin et al. 2009: fig. 8D–G) (vs. swollen slightly).

See Lin et al. (2009).

Vietnam (Cat Ba National Park, site 6 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Dushan County, Xiasi Town, Xinhe Village, Bayoudadong Cave, 25.4457N, 107.4316E, elevation ca. 929 m, 21.V.2005, Y. Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema grandidens comb. nov. resembles P. oculata comb. nov. but can be distinguished by the following: the eyes are absent (vs. present); the dorsal bend between the embolus and bulb is ca. 100° (Fig. 8, and cf. Tong and Li 2008b: fig. 4C) (vs. ca. 180°); the distal part of the receptacle is two times wider than the receptacle neck (cf. Tong and Li 2008b: fig. 4E) (vs. three times).

See Tong and Li (2008b).

China (Guizhou, site 7 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), Vietnam, Phu Tho Province, Tan Son District, Xuan Dai, Xuan Son National Park, La Cave, 21.1380N, 104.9390E, elevation ca. 424 m, 27.X.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema laensis resembles P. breviseta comb. nov. but can be distinguished by the following: the bulb does not protrude ventro-distally (Fig. 8, and cf. Zhao et al. 2018a: fig. 7C, D) (vs. protrude), the embolus with a retrolateral vertical groove (Fig. 8, and cf. Zhao et al. 2018a: fig. 8B) (vs. embolus without any structure retrolaterally); the tip of receptacle is four times wider than the neck (cf. Zhao et al. 2018a: fig. 9C) (vs. three times).

See Zhao et al. (2018a).

Vietnam (Phu Tho, site 8 in Fig. 32), known only from the type locality.

Paratypes: 11 ♀ (IZCAS), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Dushan County, Xiasi Town, Guojiafen Cave, 25.4833N, 107.4500E, 24.V.2005, Y. Tong and Y. Lin leg. Examined.

1♂ and 2♀ (including one molecular voucher, IZCAS), same data as holotype.

Pinelema oculata comb. nov. resembles P. grandidens comb. nov. but can be distinguished by the following: the eyes are present (vs. absent); the dorsal bend between the embolus and bulb is ca. 180° (Fig. 8) (vs. ca. 100°); the distal part of the receptacle is three times wider than the receptacle neck (cf. Tong and Li 2008b: fig. 5C) (vs. two times).

See Tong and Li (2008b).

China (Guizhou, site 9 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Nandan County, Chengguan Town, En Village, Encun Cave, 25.0693N, 107.6033E, elevation ca. 605 m, 3.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema pedati comb. nov. resembles P. conglobare comb. nov. but can be distinguished by the following: the smaller ratio of the bulbal length/width (1.60, Fig. 8, and cf. Lin and Li 2010: fig. 8A, B) (vs. 1.77), the ventral bend of the embolus and bulb is acute-angled (Fig. 8, and cf. Lin and Li 2010: fig. 8A, B) (vs. right-angled); the distal part of the receptacle is boot-shaped (cf. Lin and Li 2010: fig. 9B–D) (vs. globular).

See Lin and Li (2010).

China (Guangxi, site 10 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Lingyun County, Sicheng Town, Shuiyuan Cave, 24.3464N, 106.5579E, elevation ca. 573 m, 14.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema spinafemora comb. nov. resembles P. cucurbitina comb. nov. but can be distinguished by the following: the bulb protrudes ventro-distally (Fig. 8) (vs. not protrude), the ventral bend between the embolus and bulb is right-angled (Fig. 8) (vs. acute-angled).

See Lin and Li (2010).

China (Guangxi, site 11 in Fig. 32), known only from the type locality.

Holotype: ♂ (IZCAS), Vietnam, Quang Binh Province, Phong Nha-Ke Bang National Park, Tien Son Cave, 17.5800N, 106.2820E, elevation ca. 102 m, 17.V.2016, Z. Chen and Q. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♂ (molecular voucher, IZCAS), same data as holotype.

Pinelema xiezi resembles P. cucphongensis comb. nov. but can be distinguished by the following: the lateral scuta on the abdomen is absent (cf. Zhao et al. 2018a: figs 16A, 17A) (vs. present); the different shape of the embolus, the smaller ratio of the bulbal length/width (1.28, Fig. 8, and cf. Zhao et al. 2018a: fig. 16C, D) (vs. 1.49), the larger length ratio of the embolus/bulb (0.63, Fig. 8, and cf. Zhao et al. 2018a: fig. 16C, D) (vs. 0.36); the distal part of the receptacle is five times wider than the neck of the receptacle (cf. Zhao et al. 2018a: fig. 18C) (vs. seven times).

See Zhao et al. (2018a).

Vietnam (Quang Binh, site 12 in Fig. 32), known only from the type locality.

Apneumonella oculata Fage, 1921 from Tanzania, Africa; Simon and Fage 1922: 528, fig. II.

This genus currently includes three species: two from Africa and one from Sumatra. The Sumatran species, A. jacobsoni Brignoli, 1977, is known by the female only and is most likely misplaced in this genus (see comments below).

Holotype: ♀ (RMNH), Indonesia, Sumatra, West Sumatra Province, Fort de Kock, 0.2484S, 100.4832E, elevation ca. 920 m, 1926, E. Jacobson leg. Not examined.

3♀ (including molecular voucher, IZCAS) from the type locality: I. 2014, H. Zhao leg.

Indonesia (Sumatra, West Sumatra, site 1 in Fig. 33).

The placement of this species in Apneumonella is doubtful because the males of both A. oculata Fage, 1921 (the type species of Apneumonella) and A. jacobsoni are unknown, and females of the above two species provide little information regarding their generic belonging. To test the relationship of A. jacobsoni to A. oculata, molecular data of A. oculata is necessary.

Mekonglema bailang sp. nov. from Yunnan, China.

The generic name is a combination of “Mekong” referring to the Mekong-Lancang River which encompasses the distributional range of the genus, and “-lema”, a convention used because it is part of the genus Telema, which was the first genus described in Telemidae. The gender is feminine.

Mekonglema gen. nov. resembles Pinelema but can be distinguished by the following: Males of Mekonglema gen. nov. have a bulbal apophysis (M. bailang sp. nov., M. xinpingi comb. nov., and M. yan sp. nov.) (Fig. 12D) (vs. absent), or the tip of the embolus is directed dorsally (M. kaorao sp. nov. and M. walayaku sp. nov.) (Figs 14C, D, 16C, D) (vs. tip of embolus directed ventrally). Females of Mekonglema gen. nov. can be distinguished from those of Pinelema by the receptacle lacking tubes inside (Fig. 13A) (vs. receptacle with several membranous tubes inside).

Figure 12. 

Mekonglema bailang sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Total length: 1.06–1.50 (male), 1.15–1.70 (female). Carapace without pattern in troglobitic species or with radial striae in rainforest species, i.e. M. xinpingi comb. nov. Sternum with sparse setae, milky white or light brown in troglobitic species, or nearly black in M. xinpingi comb. nov. Eyes ringed with black, vestigial, or absent; Leg formula: 1-2-4-3, tibial glands belt-shaped (Fig. 1C). Abdomen of males with three different types of scutae (except in M. yan sp. nov.): the first scuta connects to pedicel, dorsal (arrow 1 on Fig. 12A); the second scuta is posterior to the first one, dorsal (arrow 2 on Fig. 12A); and the third scutae are paired, lateral (arrows 3 in Fig. 12A). Abdomen of females without scuta. Male palp: cymbial apophysis cone shaped and located medially (Fig. 12C); bulb ellipsoid or nearly ellipsoid, bulbal apophysis present (except M. kaorao sp. nov. and M. walayaku sp. nov.); embolus short relative to cymbium, sclerotized, directed outward from the cymbium in M. bailang sp. nov., M. xinpingi comb. nov., and M. yan sp. nov.; embolus long relative to cymbium, unsclerotized, directed toward cymbium in M. kaorao sp. nov. and M. walayaku sp. nov. Endogyne: composed of single tube-like or globular receptacle with short neck, membranous tubes absent.

Figure 13. 

Mekonglema bailang sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Figure 14. 

Mekonglema kaorao sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Mekonglema bailang sp. nov., M. kaorao sp. nov., M. walayaku sp. nov., M. xinpingi comb. nov., and M. yan sp. nov. (Fig. 34).

China (Yunnan) and Laos (Luang Prabang) (sites 2–6 in Fig. 33).

Holotype: ♂ (IZCAS), China, Yunnan Province, Baoshan Prefecture, Shidian County, Bailang Town, Xianren Cave, 24.6536N, 99.2645E, elevation ca. 1987 m, 29.VII.2010, C. Wang and Q. Zhao leg. Paratypes: 5♂ and 1♀ (IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles M. yan sp. nov. but can be distinguished by the following characters: abdominal scutae present in the male (vs. absent); the embolus is sclerotized (Fig. 12B–D) (vs. unsclerotized), the bulb length/width ratio is ca. 1.2 (Fig. 12C, D) (vs. ca. 1.7); the receptacle is short and almost straight (Fig. 13A) (vs. long and U-shaped). This species also resembles M. xinpingi comb. nov. but can be differentiated by the following: the absence of eyes (vs. presence), the nearly ellipsoid shape of the bulb (Fig. 12C, D) (vs. droplet shaped), the fin-like embolus (Fig. 12C) (vs. cone shaped), and the receptacle is short and not swollen distally (vs. receptacle long and swollen distally).

Male (holotype). Total length 1.50. Carapace 0.65 long, 0.61 wide. Abdomen 0.85 long, 0.79 wide. Carapace brown (Fig. 12A). Four vestigial eyes (Fig. 12A). Chelicerae, legs, labium, and endites light brown. Sternum bright brown with sparse setae (Fig. 12A). Leg measurements: I 5.05 (1.45, 0.23, 1.60, 1.13, 0.64); II 4.54 (1.36, 0.23, 1.41, 0.95, 0.59); III 3.20 (0.98, 0.21, 0.92, 0.62, 0.47); IV 4.17 (1.30, 0.20, 1.24, 0.88, 0.55). Abdomen pale brown (Fig. 12A).

Palp. Tibia 2.12 times longer than patella, cymbium 1.74 times longer than tibia, cymbial apophysis length 2/3 as wide as cymbial base (Fig. 12C); bulb shaped as shown in Fig. 12C, D; bulbal apophysis sclerotized and semi-circular with claw-like tip, embolus sclerotized (Fig. 12B–D).

Female. Total length 1.48. Carapace 0.65 long, 0.63 wide. Abdomen 0.81 long, 0.80 wide. Coloration as in male (Fig. 13B, C). Leg measurements: I 4.78 (1.47, 0.24, 1.45, 1.00, 0.62); II 4.35 (1.34, 0.22, 1.33, 0.88, 0.58); III 3.04 (0.96, 0.22, 0.87, 0.54, 0.45); IV 4.05 (1.28, 0.19, 1.18, 0.85, 0.55). Abdomen pale grey. Insemination entrance membranous, two times thinner than receptacle (Fig. 13A); receptacle tube-like and straight (Fig. 13A).

China (Yunnan, Baoshan, site 2 in Fig. 33), known only from the type locality.

Holotype: ♂ (SMF), Laos, Luang Prabang Province, Vieng Phoukha town, Ban Nam Eng, Kao Rao Cave, 20.7251N, 101.1541E, elevation ca. 729 m, P. Jäger leg. Paratype: 1♀ (SMF), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles M. walayaku sp. nov. but can be distinguished by the following characters: the bulb is nearly globular (Fig. 14C, D) (vs. ellipsoidal), the tip of the embolus is not sclerotized (Fig. 14B) (vs. well sclerotized). The tip of receptacle is 1.50 times wider than the neck (Fig. 15A) (vs. three times).

Figure 15. 

Mekonglema kaorao sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Male (holotype). Total length 1.13. Carapace 0.49 long, 0.44 wide. Abdomen 0.58 long, 0.45 wide. Carapace brown (Fig. 14A). Six vestigial eyes (Fig. 14A). Chelicerae, legs, labium, and endites light brown. Sternum bright brown with sparse setae. Leg measurements: I 3.06 (0.80, 0.18, 0.98, 0.62, 0.48); II 2.84 (0.85, 0.17, 0.84, 0.53, 0.45); III 2.06 (0.63, 0.15, 0.56, 0.37, 0.35); IV 2.73 (0.81, 0.15, 0.81, 0.56, 0.40). Abdomen light brown (Fig. 14A).

Palp. Tibia 2.71 times longer than patella, cymbium 1.65 times longer than tibia, length of cymbial apophysis as wide as cymbial base (Fig. 14C); bulb nearly globular (Fig. 14C, D); embolus 2/3 as long as cymbium, directed dorsally (Fig. 14C, D).

Female. Total length 1.18. Carapace 0.46 long, 0.45 wide. Abdomen 0.65 long, 0.52 wide. Eyes vestigial (Fig. 15B). Coloration as in male (Fig. 15B, C). Leg measurements: I 2.95 (0.88, 0.18, 0.88, 0.56, 0.45); II 2.62 (0.79, 0.17, 0.75, 0.50, 0.41); III 1.92 (0.59, 0.14, 0.53, 0.34, 0.32); IV 2.62 (0.79, 0.15, 0.75, 0.53, 0.40). Abdomen grey. Receptacle membranous, without tubes inside, neck 1.50 times thinner than tip (Fig. 15A).

Laos (Luang Prabang, site 3 in Fig. 33), known only from the type locality.

Holotype: ♂ (IZCAS), China, Yunnan Province, Nujiang Lisu Autonomous Prefecture, Lushui County, Daxingdi Township, Walayaku Cave. 26.1215N, 98.8581E, elevation ca. 895 m, 24.VI.2016, Y. Li and J. Liu leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles M. kaorao sp. nov. but can be distinguished by the following characters: the bulb is ellipsoidal (Fig. 16C, D) (vs. nearly globular), the tip of the embolus is well sclerotized (arrow on Fig. 16B) (vs. unsclerotized); the neck of the receptacle is ca. three times thinner (vs. 1.50 times thinner) than the distal part of the receptacle (Fig. 17A).

Figure 16. 

Mekonglema walayaku sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Male (holotype). Total length 1.06. Carapace 0.49 long, 0.43 wide. Abdomen 0.56 long, 0.55 wide. Carapace light brown (Fig. 16A). Eyes vestigial (Fig. 16A). Chelicerae, legs, labium, and endites light yellow. Sternum bright brown with sparse setae. Leg measurements: I 3.65 (1.03, 0.18, 1.13, 0.78, 0.53); II 3.20 (0.96, 0.17, 0.95, 0.65, 0.47); III 2.26 (0.67, 0.16, 0.61, 0.44, 0.38); IV 2.86 (0.85, 0.16, 0.83, 0.59, 0.43). Abdomen dark blue (Fig. 16A).

Palp: Tibia 1.88 times longer than patella, cymbium 1.34 times longer than tibia, cymbial apophysis length as wide as cymbial base (Fig. 16C); bulb ellipsoidal as in Fig. 16C, D; embolus 5/7 as long as cymbium, directed dorsally (Fig. 16C, D).

Female. Total length 1.15. Carapace 0.50 long, 0.47 wide. Abdomen 0.63 long, 0.58 wide. Coloration as in male (Fig. 17B, C). Leg measurements: I 3.36 (0.97, 0.19, 1.03, 0.69, 0.48); II 3.00 (0.89, 0.19, 0.88, 0.59, 0.45); III 2.15 (0.65, 0.17, 0.59, 0.40, 0.34); IV 2.80 (0.85, 0.18, 0.79, 0.56, 0.42). Abdomen blue. Receptacle membranous, neck much longer than tip, and tip three times wider than neck (Fig. 17A).

China (Yunnan, Nujiang, site 4 in Fig. 33), known only from the type locality.

Figure 17. 

Mekonglema walayaku sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Paratypes: 1♂ and 1♀ (IZCAS), China, Yunnan Province, Xishuangbanna Autonomous Prefecture, Mengla County, Menglun Town, rainforest, leaf litter, 21.9001N, 101.1833E, V–VII.2005, G. Zheng leg. Examined.

3♀ (including one molecular voucher, IZCAS), same data as paratypes.

Mekonglema xinpingi comb. nov. resembles M. bailang sp. nov. but can be distinguished by the following characters: eyes are present (vs. absent); carapace with distinct radial striae (vs. no pattern); bulb is droplet-shaped (cf. Lin and Li 2008: fig. 2) (vs. nearly ellipsoidal).

See Lin and Li (2008).

China (Yunnan, Xishuangbanna, site 5 in Fig. 33), known only from the type locality.

This species is transferred to Mekonglema gen. nov. because it shares a similar shape of the copulatory organs with M. bailang sp. nov., the type species of the genus. This placement is also supported by molecular phylogenetic analyses (Fig. 34).

Holotype: ♂ (IZCAS), China, Yunnan Province, Baoshan Prefecture, Tengchong County, Diantan Town, Lianzu Village, Yan Cave, 25.5501N, 98.4452E, elevation ca. 1867 m, 27.XI.2013, Y. Li and J. Liu leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

The species refers to the type locality; noun in apposition.

Mekonglema yan sp. nov. resembles M. bailang sp. nov. but can be distinguished by the following characters: an abdominal scuta is absent in the male (vs. present); the tip of the embolus is unsclerotized (Fig. 18B–D) (vs. sclerotized), the bulbal length/width ratio is approximately 1.7 (Fig. 18C, D) (vs. 1.2); the receptacle is long and U-shaped (Fig. 19A) (vs. short and nearly straight).

Figure 18. 

Mekonglema yan sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Male (holotype). Total length 1.48. Carapace 0.69 long, 0.57 wide. Abdomen 0.77 long, 0.63 wide. Carapace brown (Fig. 18A). Four vestigial eyes (Fig. 18A). Chelicerae, labium, and endites brown, legs yellow. Sternum bright brown with sparse setae. Leg measurements: I 4.86 (1.44, 0.23, 1.52, 1.00, 0.67); II 4.33 (1.34, 0.23, 1.25, 0.91, 0.60); III 3.20 (1.00, 0.21, 0.88, 0.61, 0.50); IV 4.04 (1.25, 0.20, 1.16, 0.87, 0.56). Abdomen light brown (Fig. 18A).

Palp: Tibia 2.05 times longer than patella, cymbium 1.73 times longer than tibia, length of cymbial apophysis as wide as cymbial base (Fig. 18C); bulb shaped as in Fig. 18C, D; bulbal apophysis sclerotized and looped about 270° (Fig. 18B–D); embolus membranous and finger-like (Fig. 18C, D).

Female: Total length 1.70. Carapace 0.71 long, 0.61 wide. Abdomen 0.97 long, 0.90 wide. Coloration lighter than in male (Fig. 19B, C). Leg measurements: I 4.72 (1.44, 0.23, 1.45, 0.95, 0.65); II 4.24 (1.30, 0.23, 1.25, 0.85, 0.61); III 3.49 (1.33, 0.20, 0.89, 0.59, 0.48); IV 4.05 (1.25, 0.20, 1.19, 0.85, 0.56). Abdomen grey. Receptacle membranous, U-shaped, neck as wide as distal part (Fig. 19A).

China (Yunnan, Baoshan, site 6 in Fig. 33), known only from the type locality.

Figure 19. 

Mekonglema yan sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Siamlema changhai sp. nov. from Trang Province, Thailand.

The generic name is derived from “Siam”, referring to the old name of Thailand, and “-lema” is a convention from Telema, the type genus of the family. Feminine in gender.

Siamlema gen. nov. can be distinguished from Telema by the following: belt-shaped tibial glands (Fig. 1E) (vs. plate-shaped); males can be distinguished from those of Telema by having a cymbial apophysis (Fig. 20C) (vs. absent), a distinct dorsal spine present on the palpal tibia (Fig. 20C, D) (vs. absent), and the palpal femur is longer than the cymbium (Fig. 20C, D) (vs. shorter). Receptacle without tubes inside (Fig. 21A) (vs. receptacle with membranous tubes).

Figure 20. 

Siamlema changhai sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Total length: 0.92–1.11 (male), 1.04–1.20 (female). Carapace 0.41–0.48 long. Sternum with sparse setae. Eyes normally developed, vestigial, or absent. Leg formula: 1-2-4-3, tibia I 0.97–1.05 long, leg glands belt shaped (arrows on Fig. 1D). Male palp: cymbium, tibia, patella, and femur robust relative to bulb, femur longer than tibia and cymbium, dorsal spine present on distal part of palpal tibia, distinct cymbial apophysis located prolaterally on mesial part; bulb droplet-shaped or nearly globular; embolus sclerotized. Endogyne: receptacle tube-like or globular, without tubes inside (Fig. 21A).

Thailand (Trang, Yala, sites 7, 8 in Fig. 33).

Siamlema changhai sp. nov. and S. suea sp. nov. (Fig. 34).

Holotype: ♂ (IZCAS), Thailand, Trang Province, Noyong District, Chang Hai Cave, 7.5893N, 99.6688E, elevation ca. 32 m, XI.2015, Z. Chen, G. Zhou, and Q. Zhao leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

1♀ (molecular voucher, IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles S. suea sp. nov. but can be distinguished by the following: the absence of eyes (Figs 20A, 21B) (vs. presence), the presence of a bulbal apophysis (Fig. 20B–D) (vs. absence), the presence of a dorsal spine on the palpal femur distally (arrows on Fig. 20C, D) (vs. spine absent), and the absence of a spine on the palpal tibia retrolaterally (Fig. 20D) (vs. spine present). Receptacle is tube-shaped (Fig. 21A) (vs. globular).

Figure 21. 

Siamlema changhai sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Male (holotype). Total length 0.92. Carapace 0.41 long, 0.40 wide. Abdomen 0.50 long, 0.44 wide. Carapace brown (Fig. 20A). Eyes absent (Fig. 20A). Chelicerae, legs, labium, and endites brown. Sternum bright brown. Leg measurements: I 3.28 (0.95, 0.17, 1.00, 0.69, 0.47); II 2.93 (0.86, 0.16, 0.85, 0.63, 0.43); III 2.17 (0.66, 0.14, 0.58, 0.43, 0.36); IV 2.73 (0.86, 0.14, 0.75, 0.56, 0.42). Abdomen brown with a few long setae (Fig. 20A).

Palp. Tibia 1.57 times longer than patella, cymbium 1.22 times longer than tibia, femur 1.06 times longer than cymbium (Fig. 20C, D); cymbial apophysis present at sub-distal part (Fig. 20C), one femoral distal spine (Fig. 20C, D), a semi-round extension on patella proximally (Fig. 20D); bulb shaped as in Fig. 20C, D; bulbal apophysis nearly transparent, with 4 sclerotized teeth (Fig. 20B–D); embolus sclerotized, finger shaped (Fig. 20B–D).

Female. Total length 1.04. Carapace 0.48 long, 0.41 wide. Abdomen 0.56 long, 0.50 wide. Carapace, sternum, legs milky white (Fig. 21B, C). Leg measurements: I 3.31 (1.00, 0.17, 0.99, 0.67, 0.48); II 2.97 (0.90, 0.16, 0.86, 0.60, 0.45); III 2.18 (0.67, 0.14, 0.59, 0.42, 0.36); IV 2.73 (0.86, 0.15, 0.75, 0.56, 0.41). Abdomen dark brown with some long setae (Fig. 21B, C). Receptacle tube-shaped, neck two times thinner than distal part (Fig. 21A).

Thailand (Trang, site 9 in Fig. 33), known only from the type locality.

Holotype: ♂ (IZCAS), Thailand, Yala Province, Mueang District, Suea Cave, 6.5226N, 101.2311E, elevation ca. 43 m, XI.2015, Z. Chen, G. Zhou, and Q. Zhao leg. Paratypes: 1♂ and 4♀ (IZCAS), same data as holotype.

1♀ (molecular voucher, IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles S. changhai sp. nov. but can be distinguished by the following: six eyes ringed with black or eyes vestigial (Figs 22A, 23B) (vs. absent); the lack of a bulbal apophysis (Fig. 22B–D) (vs. present), having a spine on the palpal tibia retrolaterally (Fig. 22D) (vs. absent), the absence of a femoral dorsal spine (Fig. 22C, D) (vs. present); and a globular receptacle (Fig. 23A) (vs. tube-shaped).

Figure 22. 

Siamlema suea sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Male (holotype). Total length 1.11. Carapace 0.46 long, 0.41 wide. Abdomen 0.60 long, 0.50 wide. Carapace light brown (Fig. 22A). Six vestigial eyes (Fig. 22A). Chelicerae, legs, labium, and endites brown. Sternum bright brown with sparse setae. Leg measurements: I 3.46 (1.03, 0.16, 1.05, 0.74, 0.48); II 2.92 (0.90, 0.16, 0.83, 0.60, 0.43); III 2.07 (0.63, 0.13, 0.59, 0.40, 0.32); IV 2.77 (0.85, 0.16, 0.77, 0.59, 0.40). Abdomen brown with a few long setae (Fig. 22A).

Palp. Tibia 1.38 times longer than patella, cymbium 0.94 times shorter than tibia, femur 1.17 times longer than cymbium (Fig. 22C, D); cymbial apophysis stout and cone shaped (Fig. 22C); tibia with 2 spines, one dorsal spine distally, the other bifurcate, located retrolaterally medially (Fig. 22D); patella with retrolateral semi-round extension mesially (Fig. 22D); bulb droplet shaped as in Fig. 22C, D; embolus strongly sclerotized, and tiny in comparison to bulb.

Female. Total length 1.20. Carapace 0.45 long, 0.41 wide. Abdomen 0.70 long, 0.66 wide. Coloration as in male (Fig. 23B, C). Six eyes ringed with black (Fig. 23B). Leg measurements: I 3.15 (0.94, 0.16, 0.97, 0.61, 0.47); II 2.79 (0.84, 0.16, 0.81, 0.55, 0.43); III 2.02 (0.62, 0.13, 0.56, 0.38, 0.33); IV 2.73 (0.85, 0.14, 0.76, 0.56, 0.42). Abdomen brown (Fig. 23B). Neck of receptacle umbrella-shaped, and distal part of receptacle globular (Fig. 23A).

Thailand (Yala, site 8 in Fig. 33), known only from the type locality.

Figure 23. 

Siamlema suea sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Sundalema bonjol sp. nov. from West Sumatra Province, Indonesia.

The generic name is derived from “Sunda”, referring to Sundaland (distributional range of this genus), and “-lema” is a convention from the type genus of the family. Feminine in gender.

Females belonging to Sundalema gen. nov. can be distinguished from those of other genera by the following: a sclerotized and spiral receptacle (Fig. 24A) (vs. unsclerotized or not spiral). Males belonging to Sundalema gen. nov. resemble species in the bailongensis-group by having a long embolus but can be distinguished by the embolus lacking a spiral ridge and being nearly L-shaped (Fig. 25B–D) (vs. a distinct spiral ridge present on the nearly straight embolus).

Figure 24. 

Sundalema bonjol sp. nov., female holotype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Total length: 0.98–1.10 (male), 1.05–1.25 (female). Carapace 0.44– 0.58 long. Sternum light brown or milky white, with several sparse setae. Six vestigial eyes or ringed with black. Leg formula: 1-2-4-3, tibia I 0.90–1.27 long, leg glands belt-shaped (Fig. 1E). For females, neck of receptacle membranous, receptacle spiral, its dorsal part sclerotized, coiled into 1.25 to 2.5 loops. For males, cymbial apophysis tiny, present meso-prolaterally; embolus long relative to bulb, nearly L-shaped, lacking spiral ridge. Abdomen blue or pale brown with several long setae.

Sundalema acicularis comb. nov., S. anguina (Wang & Li, 2010) comb. nov., S. bonjol sp. nov., and S. khaorakkiat sp. nov.

Southeast Asia (Thailand and Indonesia, sites 9–12 in Fig. 33).

The females of this genus are easily distinguished from all other genera, as their receptacles are long, spiral, and sclerotized; thus, a female has been chosen to represent the holotype.

Holotype: ♀ (IZCAS), Indonesia, Sumatra, West Sumatra Province, Payakumbuh, Koto Tinggi Village, Imam Bonjol Cave, 0.0637S, 100.3451E, elevation ca. 962 m, III.2014, Z. Yao leg. Paratypes: 2♂ and 3♀ (IZCAS), same data as holotype.

1♀ (molecular voucher, IZCAS), same data as holotype.

The species refers to the type locality; noun in apposition.

The species resembles S. anguina comb. nov. but can be distinguished by the following characters: the eyes are vestigial (Figs 24B, 25A) (vs. ringed with black); the receptacle is coiled into 1.5 loops (Fig. 24A) (vs. 2 loops, cf. Wang and Li 2010a: fig. 4C); the embolus is thin and bent at an obtuse angle (Fig. 25B–D) (vs. wide and bent at a right angle, cf. Wang and Li 2010a: fig. 4A, B).

Figure 25. 

Sundalema bonjol sp. nov., male paratype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Female (holotype). Total length 1.09. Carapace 0.53 long, 0.45 wide. Abdomen 0.64 long, 0.56 wide. Carapace light brown (Fig. 24B). Four vestigial eyes (Fig. 24B). Chelicerae, legs, labium, endites, and sternum bright brown. Sternum with sparse setae (Fig. 24B, C). Leg measurements: I 3.45 (1.04, 0.18, 1.06, 0.67, 0.50); II 3.10 (0.96, 0.17, 0.92, 0.60, 0.45); III 2.17 (0.68, 0.17, 0.58, 0.40, 0.34); IV 2.89 (0.90, 0.16, 0.79, 0.61, 0.43). Abdomen brown (Fig. 24B, C). Receptacle coiled into 1.5 loops, and neck as wide as distal part (Fig. 24A).

Male. Total length 1.03. Carapace 0.47 long, 0.40 wide. Abdomen 0.53 long, 0.44 wide. Coloration as in female (Fig. 25A). Leg measurements: I 3.34 (0.94, 0.16, 1.04, 0.71, 0.49); II 3.05 (0.90, 0.16, 0.91, 0.63, 0.45); III 2.16 (0.67, 0.14, 0.58, 0.42, 0.35); IV 2.82 (0.87, 0.15, 0.79, 0.59, 0.42). Palp: tibia 1.80 times longer than patella, cymbium 1.71 times longer than tibia, 1.69 times longer than femur; length of cymbial apophysis as wide as 1/2 width of cymbial base (Fig. 25C); bulb ellipsoidal (Fig. 25C, D); embolus tube-like, as long as major-axis of bulb (Fig. 25B–D).

Indonesia (Sumatra, site 11 in Fig. 33), known only from the type locality.

Holotype: ♀ (IZCAS), Thailand, Songkhla Province, Rattaphum District, Khao Rak Kiat Cave, 7.0724N, 100.2502E, elevation ca. 52 m, XI.2015, Z. Chen, G. Zhou and Q. Zhao leg. Paratypes: 2♂ and 2♀ (IZCAS), same data as holotype.

1♀ (molecular voucher, IZCAS), same data as holotype.

The species name refers to the type locality; noun in apposition.

This species resembles S. acicularis comb. nov. but can be distinguished by the following characters: the eyes are vestigial (Figs 26B, 27A) (vs. ringed with black); the receptacle is coiled into 1.25 loops (Fig. 26A) (vs. 2.5 loops); the embolus is longer than the cymbium (Fig. 27C, D) (vs. shorter), and the palpal tibia/cymbium ratio is 0.4 (Fig. 27C, D) (vs. 0.45).

Figure 26. 

Sundalema khaorakkiat sp. nov., female holotype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Female (holotype). Total length 1.25. Carapace 0.58 long, 0.50 wide. Abdomen 0.63 long, 0.55 wide. Carapace pale white (Fig. 26B). Six vestigial eyes (Fig. 26B). Chelicerae, legs, labium milky white, endites brown (Fig. 26B). Sternum milky white with sparse setae (Fig. 26B). Leg measurements: I 3.93 (1.21, 0.19, 1.21, 0.80, 0.52); II 3.52 (1.10, 0.19, 1.06, 0.72, 0.45); III 2.45 (0.73, 0.18, 0.69, 0.49, 0.36); IV 3.28 (1.03, 0.18, 0.92, 0.70, 0.45). Abdomen pale brown with a few long setae (Fig. 26B, C). Receptacle coiled into 1.25 loops, and neck much thinner than mesial part (Fig. 26A).

Male. Total length 1.17. Carapace 0.54 long, 0.48 wide. Abdomen 0.63 long, 0.50 wide. Eyes and coloration as in female (Fig. 27A). Leg measurements: I 4.08 (1.24, 0.19, 1.27, 0.86, 0.53); II 3.70 (1.13, 0.19, 1.15, 0.75, 0.48); III 2.67 (0.81, 0.16, 0.76, 0.56, 0.39); IV 3.48 (1.06, 0.17, 1.00, 0.76, 0.49). Palp: Tibia 1.65 times longer than patella, cymbium 2.06 times longer than tibia, 1.50 times longer than femur; cymbial apophysis dark brown and cone shaped (Fig. 27C); bulb ellipsoidal; embolus longer than cymbium (Fig. 27C, D).

Figure 27. 

Sundalema khaorakkiat sp. nov., male paratype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Thailand (Songkhla, site 12 in Fig. 33), known only from the type locality.

Holotype: ♂ (RMNH), Thailand, Prachuap Khiri Khan Province, Hua Hin, Tham Kaew Cave, 12.5947N, 99.9571E, 3.I.1989, C.L. Deeleman-Reinhold leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

This species resembles S. khaorakkiat sp. nov. but can be distinguished by the following characters: six eyes ringed with black (vs. eyes vestigial); receptacle coiled into 2.5 loops (cf. Wang and Li 2010a: fig. 1B) (vs. 1.25 loops); embolus shorter than cymbium (cf. Wang and Li 2010a: fig. 1A) (vs. longer), and the palpal tibia/cymbium length ratio is 0.45 (cf. Wang and Li 2010a: fig. 1A) (vs. 0.4).

Palpal tibia/cymbium ratio 0.45 in the male. Receptacle coiled into 2.5 loops (cf. Wang and Li 2010a: fig. 1B). For a more detailed description, see Wang and Li (2010a).

Thailand (Prachuap Khiri Khan, site 9 in Fig. 33), known only from the type locality.

Holotype: ♂ (RMNH), Thailand, Krabi Province, Diamond Cave, 8.0670N, 98.9164E. 3.I.1989, C.L. Deeleman-Reinhold leg. Paratypes 1♂ and 1♀ (RMNH), same data as holotype. Examined.

3♀ (including one molecular voucher, IZCAS) from the type locality, X.2014, Z. Chen and Y. Li leg.

This species resembles S. bonjol sp. nov. but can be distinguished by the following characters: the eyes are ringed with black (cf. Wang and Li 2010a: fig. 6B) (vs. vestigial); the receptacle is coiled into two loops (cf. Wang and Li 2010a: figs 4C, 7B) (vs. 1.5 loops); the embolus is thin and bent at a right angle (cf. Wang and Li 2010a: fig. 4A, B) (vs. much wider and obtuse angled).

See Wang and Li (2010a).

Thailand (Krabi, site 10 in Fig. 33), known only from the type locality.

Telema tenella Simon, 1882 from southern France.

Telema resembles Telemofila but can be distinguished by the following: plate-shaped tibial glands (Fig. 1A; cf. Emerit 1984: fig. A) (vs. belt-shaped); lacking a cymbial apophysis (cf. Wang et al. 2012: fig. 2C) (vs. cymbial apophysis present); the endogyne is walking-stick shaped (cf. Wang et al. 2012: fig. 3C, D) (vs. bean-shaped or globular).

Total length: 1.24–1.48 (male), 1.06–1.90 (female). Carapace 0.45–0.78 long. In T. auricoma Lin & Li, 2010 and T. nipponica, carapace with radial shaded pattern, six eyes ringed with black, sternum dark brown, abdomen blue; in T. tenella, T. guihua Lin & Li, 2010, and T. wunderlichi Song & Zhu, 1992, carapace without pattern, eyes absent, sternum and abdomen bright brown. Leg formula: 1-2-4-3, tibia I 1.32–2.45, tibia with many plate-shaped glands (Fig. 1A; cf. Emerit 1984: fig. A). In male, length of cymbium > femur > tibia > patella; bulb ellipsoid, large relative to body; embolus short relative to bulb, duckbill shaped. Receptacle walking-stick shaped, with a few membranous tubes inside.

Telema auricoma, T. guihua, T. nipponica, T. tenella and T. wunderlichi. The composition is supported not only by morphological characters but also by molecular phylogenetic analyses (Fig. 34; T. nipponica is not included).

Eurasian disjunctive range, known from Southern Europe (France and Spain, one species) and East Asia (Japan and southwestern China, four species, sites 13–16 in Fig. 33).

Holotype: ♀ (IZCAS), China, Guizhou Province, Dafang County, Xiaotun Town, Shiqiang Village, Daxiao Cave, 27.0925N,105.5551E, 1168m, 4.V.2007, J. Liu and Y. Lin leg. Paratypes 2♀ (IZCAS), same data as holotype. Examined.

1♀ (molecular voucher, IZCAS), same data as holotype; 2♀ (IZCAS), Guizhou Province, Bijie County, Xiaoyan Cave, 27.1181N, 105.2367E; 1♀ (IZCAS) Guizhou Province, Dafang County, Shilongshang Cave, 27.0925N, 105.5551E; 3♀ (IZCAS) Guizhou Province, Hezhang County, Tanjiayan Cave, 27.2003N, 104.5910E; 1♀ (IZCAS) Guizhou Province, Weining County, Banbianshan Cave, 33.9061N, 104.5396E; 2♀ (IZCAS), Yunnan Province, Xuanwei County, Jianjiao Cave, 33.3085N, 104.3900E; 3♀ (IZCAS), Yunnan Province, Xuanwei County, Fengchao Cave, 33.3909N, 104.2093E.

See Lin and Li (2010).

China (Yunnan-Guizhou Plateau).

All known specimens of this species are female, and molecular barcoding data from all eight populations examined shows no differences (unpublished data). This indicates that the species may be parthenogenetic, a character that may allow it to easily disperse more broadly than gametogenetic species.

Holotype: ♂ (IZCAS), China, Guizhou Province, Suiyang County, Wenquan Town, Guihua Village, Mahuang Cave, 28.2437N,107.2891E, elevation ca. 730 m, 4.V.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

1♀ (molecular voucher, IZCAS), same data as holotype.

This species resembles T. wunderlichi but can be distinguished by the following characters: the small body size (vs. larger size); the embolus is membranous (vs. sclerotized), and the tip of the embolus is blunt (cf. Lin and Li 2010: fig. 6E) (vs. sharply pointed).

See Lin and Li (2010).

China (Guizhou, site 14 in Fig. 33), known only from the type locality.

Holotype: ♀, Japan, Yamanashi Prefecture, Narusawa-mura, Karumizu, Shoiko-daini-fuketsu Cave, 22.IX.1969, S. Ueno and K. Kato leg. Not examined.

This species resembles T. tenella but can be distinguished by the thumb-like shape of the embolus (cf. Yaginuma 1974: fig. 4), whereas in T. tenella, the embolus is short and triangular (cf. Wang et al. 2012: fig. 2C, D).

See Yaginuma (1972, 1973).

Japan (Site 15 in Fig. 33).

Holotype: ♀ (IZCAS), China, Hunan Province, Zhangjiajie Prefecture, Sangzhi County, Wulingyuan Scientific and Historic Interest Area, 29.1171N, 110.4792E, elevation ca. 280 m, 27.XI.1992. D. Wang leg. Paratypes: 1♂ and 1♀ (IZCAS), same data as holotype. Examined.

1♂ and 2♀ (including molecular voucher, IZCAS) from the type locality: XII.2015, Z. Chen leg.

This species resembles T. guihua but can be distinguished by the following characters: the larger body size (vs. smaller), the slightly sclerotized embolus (cf. Song and Zhu 1994: fig. 1E) (vs. membranous), the sharply pointed embolus tip (cf. Song and Zhu 1994: fig. 1E) (vs. blunt).

See Song and Zhu (1994), and Lin and Li (2010).

China (Hunan, site 16 in Fig. 33).

Telemofila samosirensis Wunderlich, 1995 from Sumatra, Indonesia.

Telemofila can be distinguished from Telema by the following characters: leg formula is 1-4-2-3 (vs. 1-2-4-3), tibial glands are belt-shaped (Fig. 1G) (vs. plate-shaped); a cymbial apophysis is present (vs. absent), the embolus is sickle shaped or claw-like (vs. duckbill-shaped), and the length of the embolus is three times shorter than the diameter of the bulb (vs. two times shorter); the distal part of the receptacle is swollen (vs. not swollen).

Total length: 0.90–1.27 (male), 0.98–1.11 (female). Carapace 0.37–0.50 long. Sternum 0.20–0.25 long, with several sparse setae. Tibia I 0.51–0.82 long, leg formula: 1-4-2-3, belt-shaped glands present (Fig. 1G). Six eyes ringed with black, body blue or brown. For males, cymbial apophysis present mesially, length as wide as cymbial base (cf. Wunderlich 1995: fig. 16), embolus sickle-shaped, length of embolus 1/3 as long as diameter of bulb (cf. Wunderlich 1995: fig. 17). For females, receptacle bean shaped or globular, neck narrower than distal part (cf. Wang and Li 2010a: figs 11A, B, 15A, B).

Telemofila fabata (Wang & Li, 2010) comb. nov., T. malaysiaensis comb. nov., T. pecki (Brignoli, 1980), and T. samosirensis Wunderlich, 1995.

Rainforests in Southeast Asia (Indonesia, Singapore, Malaysian Borneo, sites 17–19 in Fig. 33) and a cave in New Caledonia.

The placement of T. pecki in this genus is dubious because its leg formula is 1-2-4-3 (Brignoli 1980) and the embolus is triangular (cf. Brignoli 1980: figs 1, 2). These characters are inconsistent with the genus characters of Telemofila. However, we have been unable to examine the types of T. pecki, and molecular data from this species is lacking.

Holotype: ♂ (RMNH), Singapore, Bukit Timah Nature Reserve, Seraya Loop, 1.3521N, 103.8198E, 2.II.1983, P.R. Deeleman leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

1♂ and 3♀ (including molecular voucher, IZCAS) from the type locality, VIII.2015, S. Li and Y. Tong leg.

This species resembles T. malaysiaensis comb. nov. but can be distinguished by the following: bigger body size, bean-shaped receptacle (cf. Wang and Li 2010a: fig. 8C, D) (vs. globular); claw-like embolus (cf. Wang and Li 2010a: fig. 8A, B) (vs. sickle-shaped). This species can be distinguished from T. samosirensis by the embolus being three times shorter than the bulb (cf. Wang and Li 2010a: fig. 9A, B) (vs. two times shorter).

See Wang and Li (2010a).

Singapore (Site 17 in Fig. 33).

Holotype: ♂ (RMNH), Malaysian Borneo, Sarawak Province, swampy lowland rainforest of Bako National Park, 1.7167N, 110.4667E, 29.III.1985, P.R. & C.L. Deeleman-Reinhold leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

This species resembles T. fabata comb. nov. but can be distinguished by the following: the smaller body size, the shape of the receptacle is globular (cf. Wang and Li 2010a: fig. 15B) (vs. receptacle bean-shaped); the embolus is sickle-shaped (cf. Wang and Li 2010a: fig. 12A–C) (vs. claw-like embolus).

See Wang and Li (2010a).

Malyasia (Borneo, Sarawak, site 18 in Fig. 33).

Holotype: ♂ (SMF), Indonesia, Sumatra, North Sumatra Province, Lake Toba, Samosir Village, 2.7424N, 98.7699E, elevation ca. 916 m, VIII. 1994, J. Wunderlich leg. Not examined.

2♀ (including molecular voucher, IZCAS) from the type locality: I.2014, H. Zhao leg.

This species can be distinguished from T. fabata comb. nov. and T. malaysiaensis comb. nov. by the length of the embolus which is equal to the radius of the bulb (cf. Wunderlich 1995: fig. 17) (vs. length of embolus 1/3 as long as the diameter of the bulb in two similar species).

See Wunderlich (1995).

Indonesia (Sumatra, North Sumatra, site 19 in Fig. 33).

Zhuanlema peteri sp. nov. from Luang Prabang Province, Laos.

The generic name is derived from “Zhuan”, referring to the Chinese pinyin “zhuan”, indicating that the apex of the embolus is twisted, and “-lema” which is a convention from the type genus of the family. Feminine in gender.

The new genus resembles species in the bailongensis-group but can be distinguished by the following characters: the apex of the embolus is twisted (Fig. 28C) (vs. tube-like embolus), the cymbial apophysis is located basally and four times shorter than the width of the cymbial base (Fig. 28C, D) (vs. cymbial apophysis located mesially or sub-basally, and longer or 1–3 times shorter than the width of the cymbial base). Females can be distinguished by the sclerotized receptacle (Fig. 29A) (vs. membranous).