Sixteen new species of the spider genus Althepus Thorell, 1898 (Araneae: Ochyroceratidae) from Southeast Asia Author Li, Fengyuan Author Liu, Chang Author Wongprom, Prasit Author Li, Shuqiang text Zootaxa 2018 2018-09-06 4471 3 401 445 journal article 29472 10.11646/zootaxa.4471.3.1 51adf720-0d97-45dd-ac98-c3c79ba4d3ce 1175-5326 1439790 294764A6-5EE6-40DC-B05C-BE23398A4B29 Genus Althepus Thorell, 1898 Type species. Althepus pictus Thorell, 1898 Emended diagnosis. The genus Althepus belongs to Psilodercinae and can be distinguished from other genera of the subfamily by the following combination of characters: cheliceral promargin with lamina and 1–2 teeth, retromargin with 1–2 small teeth; tarsus of male palp with lateral protrusion bearing a hook-shaped spine; short bulb with embolus; and female internal genitalia often with paired spermathecae (Deeleman-Reinhold 1995). Remarks. According to our observations, we used hook-shaped spine instead the "lanceolate apophysis" used by Deeleman-Reinhold (1995), and used lamina and 1–2 teeth instead the "3 teeth" used by Deeleman-Reinhold (1995). Althepus bamensis Li et Li sp. nov. Figs 1–2 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 33986), Thailand , Tak Province , Tha Song Yang District , Bam Thung Tham Sub District , unnamed cave, 17°16.603'N , 98°11.497'E , 169 m altitude, 19 November 2016 . Paratypes : 2 males and 2 females , same data as holotype (IZCAS-Ar 33987–33990). Etymology. The specific name is an adjective and refers to the type locality. Diagnosis. This new species can be easily distinguished from all the other species by their short, slightly sigmoid embolus ( Fig. 1A ) and their horn-shaped distal part of the conductor in males; and by two pairs of spermathecae with irregular margins in females ( Fig. 2A ). Description. Male ( holotype ): Total length 4.48; carapace 1.48 length, 1.52 width; abdomen 2.69 length, 1.62 width. Carapace round ( Fig. 2C ), yellow, with three longitudinal brown bands of different widths, the middle one shortened to form a triangular band around eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 20 small denticles ( Fig. 30A ). Labium brown. Sternum brown. Abdomen elongate, with complex pattern of white and dark-brown dorsally and ventrally. Legs brown, femur and tibia with white annulations. Leg measurements: I - (11.86, 0.63, 11.79, 21.15, -), II missing, III 17.32 (5.00, 0.57, 4.25, 6.28, 1.22), IV 24.76 (6.99, 0.59, 6.54, 9.04, 1.60). Male palp ( Figs 1A–D ): tarsus with three slightly curved, serrated bristles at the top of cymbial protrusion (two of them missing, Fig. 1D ), one hooked spine with tip directed proximally ( Fig. 1A ); bulb bright yellow, ovate; embolus arising distally from bulb, short, slightly sigmoid; conductor arising proximally from bulb, with two horn-shaped protrusions distally; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 2D–E ), but smaller. Internal genitalia with two pairs of spermathecae with irregular margins ( Fig. 2A ). Measurements: total length 3.90; carapace 1.25 length, 1.34 width; abdomen 2.44 length, 1.25 width. Leg measurements: I 40.95 (9.61, 0.56, 9.94, 17.63, 3.21), II 21.67 (5.44, 0.53, 5.77, 8.33, 1.60), III 14.14 (4.08, 0.51, 3.41, 5.00, 1.14), IV missing. Variation. Males: carapace 1.25–1.48 length, 1.30–1.52 width; femur I: 11.86, 12.31 (n = 3; leg I lost in one specimen). Females: carapace 1.10, 1.25 length, 1.02, 1.34 width; femur I: 9.61. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on a depression of the cave wall. The cave is located in the bamboo plant community of deciduous forest. Althepus changmao Li et Li sp. nov. Figs 3–4 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 33991), Thailand , Chiangmai , Tham (cave) Pha Daeng Cave , 19°30.187'N , 99°08.838'E , 577 m altitude, 27 June 2014 . Paratypes : 2 females , same data as holotype (IZCAS-Ar 33992–33993). Etymology. The specific name is derived from the Chinese pinyin 'cháng máo', which means 'long bristle', referring to the long bristle of the male palpal tibia ( Fig. 3A ); noun in apposition. Diagnosis. The species resembles A. tibiatus Deeleman-Reinhold, 1985 (Deeleman-Reinhold 1985) in having the upward curling part of the distal conductor and one distinctively long bristle on the male palpal tibia ( Fig. 3C ), and the spermathecae with many bifurcations in females ( Fig. 4A ). Males of the new species can be distinguished by the jagged distal part of the conductor ( Fig. 3B ), compared with the sharp distal part of the conductor in A. tibiatus . Females can be distinguished by two branches of the medial spermathecae ( Fig. 4A ), compared with 3–4 branches of the medial spermathecae in A. tibiatus . Description. Male ( holotype ): Total length 4.42; carapace 1.44 length, 1.62 width; abdomen 2.03 length, 1.10 width. Carapace round ( Fig. 4C ), yellowish, with three longitudinal brown bands of different width. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang provided with 21 small denticles ( Fig. 30B ). Labium brown. Sternum brown. Abdomen elongate, with complex patterns of white and brown dorsally and ventrally. Legs brown, femur and tibia with white annulations. Leg measurements: I 44.60 (10.83, 0.66, 11.22, 19.23, 2.66), II missing, III 26.58 (7.69, 0.64, 7.37, 9.28, 1.60), IV missing. Male palp ( Figs 3A–D ): tarsus distinct, extends distally, with three slightly curved, serrated bristles at the top of the cymbial protrusion (one of them missing, Fig. 3A ), one hooked spine with tip directed proximally ( Fig. 3A ); tibia with one distinctively long bristle; bulb yellow, ovate; embolus arising prolatero-distally from bulb, slender and proximal part curved; conductor arising proximally from bulb, curved, with the distal part curling upward forming a groove; embolus and conductor widely separated (distance almost equal to diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 4D–E ), but smaller. Internal genitalia with two pairs of spermathecae, curved, bifurcate, with a swollen distal part; the spermachecae left and right showing different number of branches ( Fig. 4A ). Measurements: total length 3.36; carapace 1.05 length, 1.20 width; abdomen 2.13 length, 1.34 width. Leg measurements: I missing, II missing, III 11.52 (3.24, 0.49, 2.88, 3.80, 1.11), IV missing. Variation. Females: carapace 1.05, 1.15 length, 1.19, 1.20 wide; leg I lost (n = 2). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the cave wall (high humidity in the cave). Remark. A. changmao Li et Li sp. nov. was labeled as “sp. 101 ” in the analysis of Li and Li (2018). Althepus dongnaiensis Li et Li sp. nov. Figs 5–6 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 33994), Vietnam , Dong Nai Province , Cat Tien National Park , Natural Forest , 11°27.005'N , 107°21.863'E , 173 m altitude, 0 5 September 2015 . Paratype : 1 female , same data as holotype (IZCAS-Ar 33995). Etymology. The specific name is an adjective and refers to the type locality. Diagnosis. This new species can be easily distinguished from all the other species by the wide, curved conductor and the arc-shaped margin of the distal conductor in males ( Figs 5A–D ) and by one pair of distinctively curved spermathecae in females ( Fig. 6A ). Description. Male ( holotype ): Total length 3.68; carapace 1.16 length, 1.13 width; abdomen 2.15 length, 0.90 width. Carapace round ( Fig. 6C ), yellowish, with three longitudinal brown bands of different widths, the middle one cleaved and shortened to form a triangular band around eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 20 small denticles ( Fig. 30C ). Labium brown. Sternum yellow, with two irregular brown lines each side. Abdomen elongate, with complex patterns of white and dark-brown dorsally and ventrally. Legs brown. Leg measurements: I 33.00 (8.00, 0.40, 8.00, 14.15, 2.45), II 21.45 (5.45, 0.40, 5.20, 8.70, 1.70), III 16.68 (5.00, 0.40, 4.49, 5.51, 1.28), IV 23.16 (6.86, 0.40, 6.41, 7.82, 1.67). Male palp ( Figs 5A–D ): tarsus distinct, extends distally, with three slightly curved, serrated bristles at the top of cymbial protrusion ( Fig. 5A ), one hooked spine with tip directed proximally ( Fig. 5A ); bulb yellow, pyriform; embolus arising proximally from bulb, slender and curved ( Figs 5A–B ); conductor arising distally from bulb, curved, distal part with a large lamina; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 6D–E ), but larger. Internal genitalia with one pair of spermathecae, distinctively curved at almost a right-angle, distal part swollen ( Fig. 6A ). Measurements: total length 3.84; carapace 1.19 length, 1.14 width; abdomen 2.33 length, 1.10 width. Leg measurements: I 16.40 (4.35, 0.40, 4.10, 6.05, 1.50), II 16.00 (4.25, 0.40, 4.00, 6.00, 1.35), III 10.75 (3.10, 0.40, 2.75, 3.50, 1.00), IV 19.57 (5.71, 0.40, 5.38, 6.41, 1.67). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected among tree buttresses at an altitude of 173 m . Remark. A. dongnaiensis Li et Li sp. nov. was labeled as “sp. 282” in the analysis of Li and Li (2018). Althepus guan Li et Li sp. nov. Figs 7–8 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 33996), Indonesia , Sumatra , West Sumatra Province , Payakumbuh , Koto Tinggi Village , Imam Bonjol Cave , 00°03.824'S , 100°20.695′E , 949 m altitude, 17 May 2014 . Paratypes : 1 male and 1 female , same data as holotype ( IZCAS Ar 33997–33998). Etymology. The specific name is derived from the Chinese pinyin 'guǎn', which means 'tube', referring to the tube-like embolus ( Fig. 7D ); noun in apposition. Diagnosis. The species can be easily distinguished from all the other species by the tube-like ( Fig. 7D ) and bifurcate ( Fig. 7B ) embolus in males and one single pair of small spermathecae in females. Description. Male ( holotype ): Total length 5.19; carapace 1.63 length, 1.64 width; abdomen 3.00 length, 1.38 width. Carapace round ( Fig. 8C ), yellow, with three longitudinal brown bands of different widths, the middle band wider than the other bands. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 24 small denticles ( Fig. 30D ). Labium brown. Sternum yellow, margin with a round of brown band. Abdomen elongate, with complex patterns of white and brown dorsally and ventrally. Legs brown, tibia with white annulations. Leg measurements: I missing, II 30.85 (7.69, 0.67, 7.69, 12.50, 2.30), III 18.70 (5.45, 0.65, 4.75, 6.60, 1.25), IV 27.35 (7.69, 0.67, 7.37, 9.62, 2.00). Male palp ( Figs 7A–D ): tarsus with three slightly curved, serrated bristles at the top of the cymbial protrusion ( Fig. 7A ); one hooked spine with tip directed proximally ( Fig. 7A ); bulb bright yellow, ovate; embolus arising distally from bulb, curved, distal part with a slender furcation; conductor arising prolatero-distally from bulb, shorter than embolus; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 8D–E ), but smaller. Internal genitalia with a pair of spermathecae, distal part swollen ( Fig. 8A ). Measurements: total length 4.05; carapace 1.25 length, 1.33 width; abdomen 2.72 length, 1.50 width. Leg measurements: I missing, II missing, III 13.83 (3.85, 0.48, 3.45, 4.65, 1.40), IV 19.57 (5.51, 0.50, 5.38, 6.54, 1.64). Variation. Males: carapace 1.24, 1.63 long, 1.28, 1.64 width; femur I: 8.97 (leg I lost in holotype ). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected at a cave entrance at an altitude of 949 m . Remark. A. guan Li et Li sp. nov. was labeled as “sp. 88” in the analysis of Li and Li (2018). Althepus maechamensis Li et Li sp. nov. Figs 9–10 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 33999), Thailand , Chiangmai Province , Mae Cham District , 18°31.677'N , 98°29.963'E , 1649 m altitude, 14 October 2014 . Paratypes : 1 male and 3 females , same data as holotype (IZCAS-Ar 34000–34003). Etymology. The specific name is an adjective and refers to the type locality. Diagnosis. The species resembles A. stonei Deeleman-Reinhold, 1985 (Deeleman-Reinhold 1985) and A. phadaengensis Li et Li sp. nov. ( Figs 16–17 ) in having a similarly shaped conductor and embolus in males and two pairs of spermathecae in females. Males of the new species can be distinguished by the shorter embolus and conductor, the shorter curved part of the conductor ( Fig. 9B ), compared to the longer embolus and conductor, the longer curved part of the conductor in A. phadaengensis ( Fig. 16B ), and by the sharp, upswept distal part of the conductor ( Fig. 9B ), compared with the blunt, pendulous distal part of the conductor in A. stonei . Females can be distinguished by a more coiled spermathecae ( Fig. 10A ), compared with the weakly twisted spermathecae in A. stonei , and by the lateral spermathecae having a more slender stalk than medial spermathecae, compared with the stalks of the spermathecae having the similar diameter in A. maechamensis ( Fig. 17A ). Description. Male ( holotype ): Total length 4.74; carapace 1.72 length, 1.70 width; abdomen 2.68 length, 1.51 width. Carapace round ( Fig. 10C ), yellowish, with three longitudinal brown bands of different widths, the middle shortened to form a triangular band around eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 23 small denticles ( Fig. 30E ). Labium brown. Sternum brown, with one irregular longitudinal yellow line in the middle. Abdomen elongate, with complex patterns of white and dark-brown dorsally and ventrally. Legs brown, femur and tibia with yellow annulations. Leg measurements: I 33.01 (8.46, 0.69, 8.33, 13.29, 2.24), II 22.00 (6.09, 0.69, 5.71, 8.14, 1.37), III 15.07 (4.25, 0.67, 3.84, 5.06, 1.25), IV 22.30 (6.41, 0.70, 6.15, 7.44, 1.60). Male palp ( Figs 9A–D ): tarsus distinct, extends distally and with three slightly curved, serrated bristles at the top of cymbial protrusion ( Fig. 9A ), one hooked spine with tip directed proximally ( Fig. 9A ); bulb yellow, ovate; embolus arising retrolatero-distally from bulb, proximal part flat and distinctly curved; conductor arising prolatero-distally from bulb, increasingly widening, curved at the distal part; embolus and conductor widely separated (distance almost equal to diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 10D–E ), but smaller. Internal genitalia with two spermathecae sitting on long, slender stalks on each side ( Fig. 10A ). Measurements: total length 3.72; carapace 1.19 length, 1.36 width; abdomen 2.18 length, 1.31 width. Leg measurements: I 23.42 (5.77, 0.53, 5.90, 8.97, 2.25), II 15.07 (4.00, 0.52, 3.75, 5.45, 1.35), III 10.92 (3.13, 0.50, 2.66, 3.48, 1.15), IV 15.33 (4.35, 0.53, 4.25, 5.00, 1.20). Variation. Males: carapace 1.34, 1.72 length, 1.27, 1.70 width; femur I: 5.32, 8.46. Females: carapace 1.19– 1.30 length, 1.34–1.40 width; femur I: 5.32–5.90. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected in a tropical forest at an altitude of 1649 m . Remark. A. maechamensis Li et Li sp. nov. was labeled as “sp. 170” in the analysis of Li and Li (2018). Althepus menglaensis Li et Li sp. nov. Figs 11–12 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 34004), China , Yunnan Province , Mengla County , Xiaolongha Village , Xishuangbanna Nature Reserve : Biological Diversity Corridor , Tropical Seasonal Rain Forest , 21°24.253′N , 101°36.324′E , 761 m altitude, 15 June 2013 , S. Li leg. Paratype : 1 female , same data as holotype (IZCAS-Ar 34005). Etymology. The specific name is an adjective refers to the type locality. Diagnosis. This new species can be easily distinguished from all the other species by a ventral protrusion of the palpal tarsus ( Figs 11A–D ) and the wave-shaped margin of the conductor ( Fig. 11B ) in males. Description. Male ( holotype ): Total length 3.60; carapace 1.47 length, 1.48 width; abdomen 2.10 length, 0.98 width. Carapace round ( Fig. 12C ), yellow, with three longitudinal brown bands of different widths, the middle one wider than the lateral ones. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 21 small denticles ( Fig. 30F ). Labium brown. Sternum brown with triangular yellow spots. Abdomen elongate, with complex patterns of white and dark-brown dorsally and ventrally. Legs brown. Leg measurements: I 28.83 (7.50, 0.55, 7.43, 11.35, 2.00), II 19.67 (5.38, 0.51, 5.04, 6.86, 1.88), III missing, IV 19.63 (5.51, 0.53, 5.64, 6.47, 1.48). Male palp ( Figs 11A–D ): tarsus distinct, extends distally,with three slightly curved, serrated bristles at the top of the cymbial protrusion ( Fig. 11D ), one hooked spine with tip directed proximally ( Fig. 11A ); bulb yellow, ovate; embolus arising retrolaterally from bulb, short and straight; conductor arising distally from bulb, with the wave-shaped margin; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 12E–F ), but larger. Internal genitalia with two pairs of spermathecae, long, curved, distal part spherical ( Fig. 12A ). Measurements: total length 3.80; carapace 1.05 length, 1.23 width; abdomen 2.50 length, 1.65 width. Leg measurements: I missing, II missing, III 10.03 (2.75, 0.31, 2.82, 3.13, 1.02), IV missing. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected among tree buttresses at an altitude of 761 m . Remark. A. menglaensis Li et Li sp. nov. was labeled as “sp. 47” in the analysis of Li and Li (2018). Althepus naphongensis Li et Li sp. nov. Figs 13–14 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 34006), Vietnam , Bac Kan Province , Ba Be National Park , Na Phong Cave , 22°23.392′N , 105°36.822′E , 179 m altitude, 20 May 2011 . Paratype : 1 female , same data as holotype (IZCAS-Ar 34007). Etymology. The specific name is an adjective and refers to the name of the cave where this species was found. Diagnosis. This new species can be easily distinguished from all other species by the slender male palp and the gradually widening conductor ( Figs 13C–D ) in males and five rounded spermathecae sitting on long, slender stalks on right side (left side has six spermathecae, Fig. 14A ) in females. Description. Male ( holotype ): Total length -; carapace 1.33 length, 1.42 width; abdomen missing. Carapace round ( Fig. 14C ), yellowish, with three longitudinal brown bands of different width, the middle shortened to triangular band around eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 16 small denticles ( Fig. 30G ). Labium brown. Sternum yellow, with two irregular brown lines on each side. Legs brown, femur and tibia with white annulations. Leg measurements: I 36.45 (8.97, 0.54, 8.78, 14.81, 3.35), II 22.67 (6.03, 0.53, 5.38, 8.65, 2.08), III 15.36 (4.24, 0.50, 3.97, 5.13, 1.52), IV 23.00 (6.73, 0.55, 6.28, 7.44, 2.00). Male palp ( Figs 13A–D ): tarsus with three slightly curved, serrated bristles at the top of the cymbial protrusion ( Fig. 13A ); one hooked spine with tip directed distally ( Fig. 13A ); tarsus and tibia with many bristles ( Figs 13C–D ); bulb bright yellow, ovate; embolus arising retrolaterally from bulb, short and slightly curved ( Fig. 13A ); conductor arising retrolatero-distally from bulb, distal part gradually widening ( Figs 13C–D ); embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Carapace similar to male in color and general features. Abdomen elongate ( Figs 14D–E ), brown, with a yellow herringbone streak dorsally and some longitudinal yellow bands of different widths and lengths ventrally. Internal genitalia with five pairs of spermathecae, slender, distal part swollen (one side has six spermathecae, Fig. 14A ). Measurements: total length 3.80; carapace 1.35 length, 1.48 width; abdomen 2.22 length, 1.28 width. Leg measurements: I 32.45 (7.95, 0.54, 8.15, 12.69, 3.12), II 20.83 (5.38, 0.53, 5.38, 7.56, 1.98), III 13.87 (3.85, 0.49, 3.65, 4.55, 1.33), IV 20.61 (5.77, 0.54, 5.83, 6.67, 1.80). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the cave wall. Remark. A. naphongensis Li et Li sp. nov. was labeled as “sp. 50 ” in the analysis of Li and Li (2018). Althepus natmataungensis Li et Li sp. nov. Figs 15 , 30 , 32 Type material. Holotype : male (IZCAS-Ar 34008), Myanmar , Chin State, Nat Ma Taung National Park , Road S of Nat Ma Taung Summit , Pristine Primary Forest , 21°12.479'N , 93°54.892'E , 2543 m altitude, P. Jäger leg., 16 May 2014 . Etymology. The specific name is an adjective refers to the type locality. Diagnosis. This new species can be easily recognized from all the other species by the bifurcate distal part of the conductor and the wavelike embolus ( Figs 15A–D ). Description. Male ( holotype ): Total length 3.85; carapace 1.30 length, 1.38 width; abdomen 2.13 length, 1.19 width. Carapace rounded, yellow, with three longitudinal brown bands of different width, the middle one wider than the lateral ones. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 19 small denticles ( Fig. 30H ). Labium brown. Sternum yellow, margin with brown band. Abdomen elongate, with complex pattern of white and brown dorsally and ventrally. Legs brown, femur and tibia with white annulations. Leg measurements: I 16.26 (4.50, 0.57, 4.10, 5.45, 1.64), II 15.88 (4.25, 0.56, 4.00, 5.45, 1.62), III 11.41 (3.20, 0.56, 3.00, 3.40, 1.25), IV missing. Male palp ( Figs 15A–D ): tarsus with three slightly curved, serrated bristles at the top (one of them missing, Fig. 15C ) and one large hooked spine with tip directed distally ( Fig. 15A ); bulb yellow, ovate; embolus arising distally from bulb, flat and wavy; conductor arising proximally from bulb, bifurcate; embolus and conductor widely separated (distance less than diameter of bulb). Female: Unknown Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected under leaf litter by hand and sieving. Remark. A. natmataungensis Li et Li sp. nov. was labeled as “sp. 210” in the analysis of Li and Li (2018). Althepus phadaengensis Li et Li sp. nov. Figs 16–17 , 31–32 Type material. Holotype : male (IZCAS-Ar 34009), Thailand , Mae Hong Son Province , Muang District , Jumpae Subdistrict , Tham (cave) Pha Daeng Cave , 19°25.395'N , 97°59.057'E , 293 m altitude, 21 November 2016 . Paratypes : 3 males and 2 females , same data as holotype (IZCAS-Ar 34010–34014). Etymology. The specific name is an adjective and refers to the cave in which the species was found. Diagnosis. The species resembles A. stonei Deeleman-Reinhold, 1995 (Deeleman-Reinhold 1995) and A. maechamensis Li et Li sp. nov. ( Figs 9–10 ) in having a similarly shaped conductor and embolus in males and two pairs of spermathecae in females, but can be distinguished by the longer embolus and conductor, the longer curved part of the conductor ( Fig. 16B ), compared with the shorter embolus and conductor, the shorter curved part of the conductor in A. maechamensis ( Fig. 9B ), by the sharp, upswept distal part of the conductor ( Fig. 16B ), compared with the blunt, pendulous distal part of the conductor in A. stonei in the males, and by the stalks of the spermathecae having the similar diameter, compared with the lateral spermathecae having a more slender stalk than medial spermathecae in A. stonei and A. maechamensis in the females ( Fig. 17A ). Description. Male ( holotype ): Total length 4.25; carapace 1.44 length, 1.45 width; abdomen 2.34 length, 1.28 width. Carapace round ( Fig. 17C ), yellowish, with three longitudinal brown bands of different widths, the middle one shortened to form a triangular band around the eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with one small tooth, posterior surface of fang with 24 small denticles ( Fig. 31A ). Labium brown. Sternum brown. Abdomen elongate, with complex pattern of white and dark-brown dorsally and ventrally. Legs brown. Leg measurements: I 45.23 (9.74, 0.62, 11.86, 19.42, 3.59), II 29.75 (7.37, 0.59, 7.37, 12.18, 2.24), III 18.81 (5.58, 0.59, 4.95, 6.41, 1.28), IV 27.65 (8.14, 0.59, 7.56, 9.42, 1.94). Male palp ( Figs 16A–D ): tarsus distinct, extends distally,with three slightly curved, serrated bristles at the top of the cymbial protrusion ( Fig. 16A ), one hooked spine with the tip directed towards proximally ( Fig. 16D ); bulb bright yellow, ovate; embolus arising retrolatero-distally from bulb, slender and slightly curved; conductor arising prolatero-distally from bulb, long, increasingly widening and curved at the distal part; embolus and conductor widely separated (distance almost equal to diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 17D–E ), but larger. Internal genitalia with two rounded spermathecae sitting on long stalks on each side ( Fig. 17A ). Measurements: total length 4.55; carapace 1.36 length, 1.42 width; abdomen 2.81 length, 1.48 width. Leg measurements: I 41.15 (9.62, 0.56, 10.45, 17.31, 3.21), II 25.81 (6.35, 0.56, 7.69, 9.23, 1.98), III 15.93 (4.70, 0.53, 4.10, 5.32, 1.28), IV 22.70 (6.41, 0.56, 6.41, 7.37, 1.95). Variation. Males: carapace 1.30–1.60 length, 1.42–1.50 width; femur I: 9.74–12.18. Females: carapace 1.36, 1.42 length, 1.33, 1.42 width; femur I: 9.62 (n = 2; leg I lost in the other specimen). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the wall near the entrance of cave. Althepus shanhu Li et Li sp. nov. Figs 18–19 , 31–32 Type material. Holotype : male (IZCAS-Ar 34015), Myanmar , Kachin State, Putao County , Hponkanrazi Wildlife Sanctuary Roadside , 27°36.550'N , 96°58.850'E , 2252 m altitude, 17 December 2016 . Paratypes : 1 male and 1 female , same data as holotype (IZCAS-Ar 34016–34017). Etymology. The specific name is derived from the Chinese pinyin 'shān hú', which means 'coral', referring to the female’s spermathecae looking like coral; noun in apposition. Diagnosis. This new species can be easily distinguished from all other species by many long bristles of the palpal tibia ( Figs 18C–D ), the petaloid structures on the distal conductor as well as the short embolus in males ( Fig. 18B ), and by many pairs of spermathecae in females ( Fig. 19A ). Description. Male ( holotype ): Total length 4.00; carapace 1.48 length, 1.48 width; abdomen 2.28 length, 1.42 width. Carapace round ( Fig. 19C ), yellow, with three longitudinal brown bands of different widths, the middle wider than the others. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang provided with 26 small denticles ( Fig. 31B ). Labium brown. Sternum yellow, with many brown bands. Abdomen elongate, with complex patterns of yellow and brown dorsally and ventrally. Legs brown. Leg measurements: I missing, II 17.45 (4.80, 0.53, 4.75, 6.28, 1.09), III 12.17 (3.60, 0.53, 3.13, 3.80, 1.11), IV 18.20 (5.19, 0.51, 5.19, 6.03, 1.28). Male palp ( Figs 18A–D ): tarsus with three slightly curved, serrated bristles at the top of the cymbial protrusion (one of them missing, Fig. 18A ), one hooked spine with tip directed proximally ( Fig. 18C ); bulb bright yellow, ovate; embolus arising retrolatero-distally from bulb, short and straight ( Fig. 18B ); conductor arising distally from bulb, wide, and some petaloid laminas arising distally; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 19D–E ) but larger. Internal genitalia with many pairs of spermathecae, short, distal part swollen ( Fig. 19A ). Measurements: total length 4.10; carapace 1.24 length, 1.41 width; abdomen 2.81 length, 2.05 width. Leg measurements: I missing, II 13.50 (3.60, 0.56, 3.68, 4.45, 1.21), III 10.31 (2.94, 0.54, 2.66, 3.00, 1.17), IV 14.57 (4.30, 0.56, 3.96, 4.35, 1.40). Variation. Males: carapace 1.35, 1.48 length, 1.42, 1.48 width (leg I lost in all specimens). Distribution. Known only from the type locality ( Fig. 32 ). Natural . Collected among tree buttresses at an altitude of 2252 m . Althepus suayaiensis Li et Li sp. nov. Figs 20 , 31–32 Type material. Holotype : male (IZCAS-Ar 34018), Thailand , Tak Province , Phop Phra District , Mae Ku Subdistrict , Tham Sua Yai Cave , , 14 November 2016 . Paratypes : 2 males , same data as holotype (IZCAS-Ar 34019–34020 ). Etymology. The specific name is an adjective and refers to the type locality. Diagnosis. This new species can be easily distinguished from all other species by the irregularly serrate margin of the distal conductor and a small triangular protrusion on the middle part of the conductor in males ( Figs 20A–B ). Description. Male ( holotype ): Total length 3.95; carapace 1.41 length, 1.36 width; abdomen 2.20 length, 1.13 width. Carapace round, light yellow, with three longitudinal brown bands of different widths, the middle one wider than the others. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 19 small denticles ( Fig. 31C ). Labium brown. Sternum yellow, with two irregular brown lines on each side. Abdomen elongate, with complex patterns of white and dark-brown dorsally and ventrally. Legs brown. Leg measurements: I 33.12 (7.69, 0.56 8.34, 14.29, 2.24), II 21.86 (5.90, 0.55, 5.64, 8.14, 1.63), III 19.93 (4.00, 0.52, 5.64, 8.14, 1.63), IV missing. Male palp ( Figs 20A–D ): tarsus with three slightly curved, serrated bristles at the top of the cymbial protrusion (two of them were missing, Fig. 20C ), one hooked spine with tip directed proximally ( Fig. 20A ); bulb yellow, ovate; embolus arising distally from bulb, proximal part distinctly wide and slightly curved ( Fig. 20A ); conductor arising distally from bulb, distinctly wide, middle and distal part with some small protrusions ( Figs 20A–B ); embolus and conductor widely separated (distance less than diameter of bulb). Female: Unknown Variation. Males: carapace 1.13–1.41 length, 1.17–1.36 width, femur I 7.69–8.21. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the cave wall. The cave is located in the bamboo plant community of deciduous forest. Althepus tadetuensis Li et Li sp. nov. Figs 21–22 , 31–32 Type material. Holotype : male (IZCAS-Ar 34021), Laos , Champasak Province , , 17 November 2012 . Paratypes : 1 male and 1 female , same data as holotype (IZCAS-Ar 34022–34023 ). Etymology. The specific name is an adjective refers to the type locality. Diagnosis. The species can be easily distinguished from all the other species by the four furcate distal part of the conductor in males ( Fig. 21D ) and by one pair of long, slender spermathecae in females ( Fig. 22A ). Description. Male ( holotype ): Total length 3.80; carapace 1.17 length, 1.30 width; abdomen 2.13 length, 1.06 width. Carapace round ( Fig. 22C ), yellow, with four longitudinal brown bands of different widths, the middle one shortened to form a triangular band around the eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 20 small denticles ( Fig. 31D ). Labium brown. Sternum yellow, with two irregular brown lines on each side. Abdomen elongate, with complex pattern of white and dark-brown dorsally and ventrally. Legs brown. Leg measurements: I 32.08 (8.01, 0.40, 7.69, 13.08, 2.95), II 30.26 (7.82, 0.40, 8.01, 11.15, 2.88), III 13.70 (3.95, 0.40, 3.50, 4.50, 1.35), IV 19.90 (5.71, 0.40, 5.45, 6.67, 1.67). Male palp ( Figs 21A–D ): tarsus with three slightly curved, serrated bristles at the top of the cymbial protrusion (one of them missing, Fig. 21C ); one hooked spine with tip directed proximally ( Fig. 21A ); bulb yellow, pyriform; embolus arising retrolaterally from bulb, slender and curved; conductor arising prolatero-distally from bulb, S-shaped, with a transparent lamina and three black divarications arising from distal part; embolus and conductor widely separated (distance almost equal to diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 22D–E ) but smaller. Internal genitalia with a pair of spermathecae, distinctively long, curved, distal part swollen ( Fig. 22A ). Measurements: total length 3.60; carapace 1.02 length, 1.33 width; abdomen 2.13 length, 1.09 width. Leg measurements: I 24.00 (6.41, 0.46, 6.54, 10.90, 2.69), II 17.92 (4.55, 0.46, 4.40, 6.09, 2.42), III 12.44 (3.38, 0.46, 2.97, 3.91, 1.72), IV 16.75 (4.81, 0.46, 4.49, 5.45, 1.54). Variation. Males: carapace 1.13, 1.17 length, 1.22, 1.30 width; femur I: 8.01 (n = 2; leg I lost in one specimen). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected in a pit of the hill at an altitude of 889 m . Remark . . tadetuensis Li et Li sp. nov. was labeled as “sp. 25” in the analysis of Li and Li ( 2018 ). Althepus tanhuang Li et Li sp. nov. Figs 23–24 , 31–32 ( IZCAS-Ar 34024 ), Thailand , Rayong Province , Kongdin Subdistrict , Phrayanak Cave , 12°46.000'N , 101°47.913'E , 0 m altitude, 0 9 November 2016 . Paratype : 1 female , same data as holotype (IZCAS-Ar 34025 ). Etymology. The specific name is derived from the Chinese pinyin 'tán huáng', which means 'spring', referring to coiled, spring-shaped spermathecae ( Fig. 24E ); noun in apposition. Diagnosis. This new species can be easily distinguished from all other species by the distinctly wide distal part of the folded conductor in male ( Fig. 23A ) and by the spring-shaped spermathecae in female ( Fig. 24A ). Description. Male ( holotype ): Total length 4.35; carapace 1.50 length, 1.70 width; abdomen 2.50 length, 1.70 width. Carapace round ( Fig. 24C ), yellowish, with three longitudinal brown bands of different width, the middle shortened to triangular band around eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang provided with 15 small denticles ( Fig. 31E ). Labium brown. Sternum brown. Abdomen elongate, with complex patterns of white and brown dorsally and ventrally. Legs brown. Leg measurements: I missing, II 32.32 (8.65, 0.71, 8.53, 12.18, 2.25), III 22.84 (6.54, 0.70, 6.09, 7.88, 1.63), IV 32.63 (8.97, 0.71, 9.29, 11.54, 2.12). Male palp ( Figs 23A–D ): tarsus with three slightly curved, serrated bristles at the top of cymbial protrusion ( Fig. 23C ), one hooked spine with tip directed proximally ( Fig. 23A ); bulb yellow, ovate; embolus arising prolatero-distally from bulb, slender and curved ( Figs 23C–D ); conductor arising retrolatero-distally from bulb, distinct wide and distal part gradually narrow ( Fig. 23A ); embolus and conductor widely separated (the distance almost equal to diameter of bulb). Female ( paratype ): Similar to male in general features ( Figs 24D–E ), but larger and darker. Internal genitalia with a pair of spermathecae, coiled, spring-shaped, the distal part swollen ( Fig. 24A ). Measurements: total length 4.36; carapace 1.38 length, 1.48 width; abdomen 2.50 length, 1.20 width. Leg measurements: I missing, II missing, III missing, IV 16.09 (6.73, 0.56, 6.54, 8.01, 5.45). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected around the entrance zone of a limestone cave. Althepus thanlaensis Li et Li sp. nov. Figs 25–26 , 31–32 Type material. Holotype : male (IZCAS-Ar 34026), Thailand , Phitsanulok Province , Nakhothai District , Ban Kaset Sut Village , , 17 October 2014 . Paratypes : 1 male and 2 females , same data as holotype (IZCAS-Ar 34027–34029 ). Etymology. The specific name is an adjective and refers to the type locality. Diagnosis. This new species can be easily distinguished from all other species by the blade-like conductor ( Figs 25A–D ) in males, and two pairs of similarly sized spermathecae in females ( Fig. 26A ). Description. Male ( holotype ): Total length 4.10; carapace 1.44 length, 1.53 width; abdomen 2.30 length, 1.22 width. Carapace rounded ( Fig. 26C ), yellow, with three longitudinal brown bands of different widths, the middle one wider than the others. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 20 small denticles ( Fig. 31F ). Labium brown. Sternum brown. Abdomen elongate, with patterns of white and brown dorsally and ventrally. Legs brown, femur and tibia with white annulations. Leg measurements: I 47.75 (10.90, 0.63, 11.54, 21.15, 3.53), II 26.97 (7.05, 0.63, 6.92, 10.13, 2.24), III missing, IV 26.17 (7.37, 0.60, 7.31, 8.97, 1.92). Male palp ( Figs 25A–D ): tarsus with three slightly curved, serrated bristles at the top of cymbial protrusion (one of them missing, Fig. 25A ), one hooked spine with tip directed proximally ( Fig. 25A ); bulb yellow, ovate; embolus arising distally from bulb, slender and slightly curved distally; conductor arising distally from bulb, blade-like and increasingly widening; embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 26D–E ) but larger. Internal genitalia with two pairs of spermathecae, long, curved, distal part swollen ( Fig. 26A ). Measurements: total length 4.50; carapace 1.36 length, 1.28 width; abdomen 2.78 length, 1.62 width. Leg measurements: I 38.06 (8.65, 0.57, 9.29, 16.34, 3.21), II 21.25 (5.77, 0.54, 5.64, 7.50, 1.80), III missing, IV - (6.03, 0.57, 5.96, 6.54, -). Variation. Males: carapace 1.42, 1.44 length, 1.50, 1.53 width; femur I: 10.90 (n = 2; leg I lost in one specimen). Prosoma: females 1.36, 1.48 length, 1.28, 1.44 width; femur I: 5.77, 8.65. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the cave wall. Remark . . thanlaensis Li et Li sp. nov. was labeled as “sp. 173” in the analysis of Li and Li ( 2018 ). Althepus tharnlodensis Li et Li sp. nov. Figs 27 , 31–32 , Thailand , Phang Nga Province , Tap Pud District , Tham Thong Lang Subdistrict , Tharn Lod Cave , 08°31.138'N , 98 °33.915'E , 116 m altitude, 27 October 2014 . Etymology. The specific name is an adjective and refers to the type locality. Diagnosis . ( Liu et al . 2017 ) and A. muangensis Li et Li, 2017 (Liu et al. 2017) in having one pair of spermathecae in females, but can be distinguished by the longer stalks of spermathecae ( Fig. 27A ), compared with the shorter stalks of spermathecae in A. muangensis , and by the larger, arch-shaped pore plates ( Fig. 27A ), compared with the smaller pore plates in A. lakmueangensis and A. muangensis in females.. Description. Female ( holotype ): Total length 3.80; carapace 1.18 length, 1.28 width; abdomen 2.13 length, 0.90 width. Carapace round ( Figs 27C–E ), yellow, with three longitudinal brown bands of different widths, the middle one wider than the others. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with one tooth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 24 small denticles ( Fig. 31G ). Labium brown. Sternum brown. Abdomen elongate, brown, with one longitudinal yellow band and two latitudinal yellow bands dorsally, and two narrow white bands ventrally. Legs brown, femur and tibia with white annulations. Leg measurements: I - (8.61, 0.53, 9.94, 22.44, -), II 22.72 (5.50, 0.50, 5.60, 9.62, 1.50), III 14.33 (4.00, 0.51, 3.75, 4.80, 1.27), IV 19.57 (5.77, 0.50, 5.51, 6.60, 1.19). Female epigyne as in Fig. 27A . Internal genitalia with a pair of spermathecae, curved, distal part swollen, each side with a pore plate at its base. Male: Unknown. Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the cave wall. Remark . . tharnlodensis Li et Li sp. nov. was labeled as “sp. 360” in the analysis of Li and Li ( 2018 ). Althepus viengkeoensis Li et Li sp. nov. Figs 28–29 , 31–32 ( IZCAS-Ar 34031 ), Laos , Vien Tiane Province , Vang Vieng District , North of Vieng Keo Village , Lom Cave , 18°57.458'N , 102°26.245'E , 314 m altitude, 0 2 December 2012 . Paratypes : 1 female , same data as holotype (IZCAS-Ar 34032 ). Etymology. The specific name is an adjective and refers to the type locality; adjective in genitive case. Diagnosis. This new species can be easily distinguished from all other species by the depigmented carapace and a sigmoid conductor ( Fig. 28A ) in males and one pair of duct-like spermathecae in female. Description. Male ( holotype ): Total length 4.60; carapace 1.62 length, 1.80 width; abdomen 2.50 length, 1.63 width. Carapace rounded ( Fig 29A ), light yellow, with three longitudinal brown bands, the middle shortened to form a triangular band around the eyes. Anterior margin of thoracic region distinctly elevated. Clypeus brown. Cheliceral promargin with two teeth, followed by a lamina, retromargin with two small teeth, posterior surface of fang with 22 small denticles ( Fig. 31H ). Labium brown, with two light, round areas basally. Sternum brown. Abdomen elongate, brown, with a yellow humanoid streak dorsally and two yellow bands ventrally. Legs brown. Leg measurements: I missing, II missing, III - (6.54, -, -, -, -), IV - (8.59, 0.68, 9.68, -, -). Male palp ( Figs 28A–D ): tarsus with three slightly curved, serrated bristles at the top ( Fig. 28A ) and one hooked spine slightly bent with tip directed proximally ( Fig. 28A ); bulb light yellow, ovate; embolus arising retrolatero-distally from bulb, slightly curved distally; conductor arising prolatero-distally from bulb, slightly curved, with distal part curling until it nearly closes ( Figs 28A–B ); embolus and conductor widely separated (distance less than diameter of bulb). Female ( paratype ): Similar to male in color and general features ( Figs 29D–E ) but larger. Internal genitalia with a pair of spermathecae, slender, transparent ( Fig. 29A ). Measurements: total length 4.84; carapace 1.50 length, 1.48 width; abdomen 2.60 length, 2.03 width. Leg measurements: I missing, II missing, III 18.22 (5.45, 0.65, 4.50, 6.09, 1.53), IV 24.47 (7.18, 0.67, 7.05, 7.69, 1.88). Distribution. Known only from the type locality ( Fig. 32 ). Natural history. Collected on the wall deep in the cave. Remark . . viengkeoensis Li et Li sp. nov. was labeled as “sp. 22” in the analysis of Li and Li ( 2018 ).