Liphistius desultor Schiødte, 1849.

Liphistius differs from the other seven liphistiid genera by the presence of signal lines radiating from the burrow’s entrance (Fig. 1B, C), by the male palp having a tibial apophysis (Figs 3A, 4A, 5A), and by the female genitalia having a sclerotized poreplate and a median receptacular cluster (Figs 3H–I, 4H–M, 5H–J).

Figure 3. 

Male palp and female genitalia of Liphistius hatyai Zhan & Xu, sp. nov. A palp, prolateral view B palp, ventral view C palp, retrolateral view D–G palp, distal views H vulva, dorsal view I vulva, ventral view A–G XUX-2017-492 (holotype) H, I XUX-2017-493. Abbreviations used: CDO = central dorsal opening; GA = genital atrium; PS = posterior stalk; RC = receptacular cluster; CT = contrategulum; Cu = cumulus; PC = paracymbium; PeP = paraembolic plate; ST = subtegulum; T = tegulum; TiA = tibial apophysis. Scale bars: 0.5 mm.

China (Yunnan Province), Indonesia (Sumatra), Laos, Peninsular Malaysia, Myanmar, and Thailand.

Holotype : Thailand • 1 ♂; Songkhla Province, Hat Yai District, Kho Hong, 7.04°N, 100.50°E; alt. 25 m; 13 November 2016; N. Warrit, V. Sivayyapram, N. Chatthanabun, P. Traiyasut leg.; XUX-2017-492. Paratype: Thailand • 1 ♀, same data as for the holotype; XUX-2017-493.

The male of L. hatyai sp. nov. resembles males of L. albipes Schwendinger, 1995 and L. yangae Platnick & Sedgwick, 1984 in having a rounded, scale-like paraembolic plate (Fig. 3A, E) but can be distinguished in having the tegulum with three transverse ridges in retrolateral view, while the latter two species have only one transverse ridge (Fig. 3C, G); from males of L. bicoloripes Ono, 1988, L. castaneus Schwendinger, 1995, and L. niphanae Ono, 1988 in having the cumulus slightly elevated (Fig. 3A–C); from the male of L. inthanon sp. nov. in having the tibial apophysis with fewer and longer setae (Fig. 3A–C), the cumulus slightly elevated (Fig. 3A–C), the subtegular apophysis absent (Fig. 3A, F), and the embolic parts detached (Fig. 3A–G); from the male of L. keeratikiati sp. nov. in having the sclerotised embolic part with three longitudinal ridges reaching the apex (Fig. 3A) and the tegulum with three transverse ridges distally (Fig. 3C, G); from males of other Liphistius species in having the cumulus slightly elevated, the sclerotised embolic part with three longitudinal ridges in prolateral view, and the subtegular apophysis absent (Fig. 3A, B, E).

The female of L. hatyai sp. nov. differs from females of L. albipes and L. castaneus in having a slightly narrower V-shaped posterior stalk (Fig. 3H, I); from females of L. bicoloripes and L. castaneus in having poreplate lacking anterolateral lobes (Fig. 3I); from the female of L. niphanae in having the anterior margin of the poreplate straight (Fig. 3H, I); from females of L. yangae, L. inthanon sp. nov., and L. keeratikiati sp. nov. in having the poreplate almost squared and with a slightly V-shaped posterior stalk (Fig. 3H, I); from females of other Liphistius species in having a hair at the center of posterior stalk dorsally (Fig. 3H).

Male. Carapace reddish-brown, with a few short, scattered bristles; opisthosoma brown, with 12 brown tergites, close to each other, 2–6 larger than others, fifth largest; chelicerae robust, promargin of cheliceral groove with 12 denticles of variable size; labium yellow and fused with sternum; sternum yellow, with a few short setae on the anterior tip and many long setae on the elongated posterior tip; legs yellowish brown, with strong hairs and spines, without distinct annulations, with 3 tarsal claws; 8 spinnerets. Measurements: BL 17.6, CL 8.81, CW 8.42, OL 8.33, OW 7.31; eye sizes and interdistances: AME 0.06, ALE 0.77, PME 0.47, PLE 0.59, AME–AME 0.13, AME–ALE 0.21, PME–PME 0.08, PME–PLE 0.09, ALE–PLE 0.12, ALE–ALE 0.17, PLE–PLE 0.39, AME–PME 0.12. Labium 0.55 long and 0.70 wide. Sternum 3.74 long and 1.04 wide. Leg I 22.55 (7.11 + 2.57 + 4.90 + 5.64 + 2.33), leg II 22.85 (6.76 + 1.73 + 5.56 + 6.06 + 2.74), leg III 26.62 (6.89 + 3.55 + 5.63 + 7.51 + 3.04), leg IV 30.54 (9.07 + 3.83 + 7.50 + 10.25 + 2.63).

Palp : tibial apophysis with four setae of same length, stouter basally and slender distally (Fig. 3A–C); paracymbium with many setae situated at tip (Fig. 3A–C); several tapering spines on slightly elevated cumulus (Fig. 3A–C); contrategulum with a triangular process, and an arched smooth edge with a sharp projection (Fig. 3B, C, G); tegulum with a serrated edge proximally and 3 transverse ridges distally (Fig. 3B, C, G); embolic parts detached (Fig. 3A–C, G), paraembolic plate semicircular, scale-like (Fig. 3A, E); embolus with 3 distinct longitudinal ridges reaching the tip prolaterally, with a few denticulations on apex, and with a flat membranous opening (Fig. 3A–C, G).

Female. Carapace orange, with few short, scattered bristles; opisthosoma gray, with 12 brown tergites, close to each other, with gray patches, 2–6 larger than others, fifth largest; 8 eyes on dark ocular tubercle; chelicerae robust, reddish brown; promargin of cheliceral groove with 10 denticles of variable size; labium yellow, fused with sternum; sternum yellow with several setae; legs with strong setae and spines, without distinct annulations, with 3 tarsal claws; 8 spinnerets. Measurements: BL 23.8, CL 11.42, CW 10.33, OL 11.03, OW 10.06; eye sizes and interdistances: AME 0.14, ALE 1.00, PME 0.52, PLE 0.61, AME–AME 0.12, AME–ALE 0.25, PME–PME 0.10, PME–PLE 0.12, ALE–PLE 0.15, ALE–ALE 0.10, PLE–PLE 0.60, AME–PME 0.15. Labium 2.45 long and 1.26 wide. Sternum 5.15 long, 1.81 wide. Palp 20.19 (7.41 + 3.65 + 4.72 +4.41), leg I 25.15 (8.71 + 4.49 + 4.59 + 4.74 + 2.62), leg II 26.62 (9.09 + 4.52 + 4.81 + 5.24 + 2.96), leg III 28.10 (9.25 + 4.28 + 5.56 + 5.89 + 3.12), leg IV 37.53 (11.06 + 4.80 + 7.70 + 9.97 + 4.00).

Genitalia : poreplate almost squared; posterior stalk slightly V-shaped, with a hair situated in the center dorsally; central dorsal opening small, situated in center of poreplate; receptacular cluster spherical (Fig. 3H, I).

The species epithet “hatyai” refers to the location of the type locality in Hat Yai District.

Southern Thailand (Songkhla Province) (Fig. 2).

Liphistius hatyai sp. nov. can be assigned to the trang-group according to the morphology of male palp and female genitalia. In males, the sclerotised part of embolus has two or three longitudinal ridges reaching the tip, the cumulus is slightly elevated, and the subtegulum is lacking an apophysis (Fig. 3A–G). In females, the poreplate has a small central dorsal opening and a receptacular cluster (Fig. 3H, I).

Specifically, L. hatyai sp. nov. belongs to the species complex D of the trang-group (sensu Schwendinger 1998; Schwendinger et al. 2019) based on male palp and female genital morphology. In males, the slightly elevated cumulus possesses long, needle-like spines (Fig. 3A–C), and the sclerotised part of embolus carries three longitudinal ridges reaching the tip (Fig. 3A). Female genitalia consist of a nearly squared poreplate, and a narrow, slightly V-shaped posterior stalk (Fig. 3H, I).

Species complex D includes species distributed in southern Thailand, western Peninsular Malaysia, and Sumatra. This species complex in Thailand includes L. albipes, L. bicoloripes, L. castaneus, L. niphanae, L. trang Platnick & Sedgwick, 1984, and L. yangae (for details see Schwendinger 1998; Schwendinger et al. 2019).

Holotype : Thailand • 1 ♂, Chiang Mai Province, Mae Chaem District, Doi Inthanon National Park, 18.52°N, 98.49°E; alt. 1700 m; 19 November 2017; F.X. Liu, D. Li, X. Xu, V. Sivayyapram leg.; XUX-2017-372A. Paratypes: Thailand • 1 ♂ 7 ♀♀, alt. 1700–1714 m, same data as for the holotype; XUX-2017-373A, XUX-2017-372, 374, 377, 378, 379, 380, 381.

The male of L. inthanon sp. nov. resemble males of L. bristowei Platnick & Sedgwick, 1984, L. lannaianus Schwendinger, 1990, L. maewongensis Sivayyapram, Smith, Weingdow & Warrit, 2017, L. marginatus Schwendinger, 1990 and L. yamasakii Ono, 1988 in having adjoining embolic parts (Fig. 4E, G) and a distinctly elevated cumulus (Fig. 4C), but it can be distinguished from the male of L. bristowei in having the tibial apophysis with more stouter spines (Fig. 4A–C) and a larger subtegular apophysis (Fig. 4D–G); from males of L. lannaianus, L. maewongensis, and L. marginatus in having the cumulus more elevated (Fig. 4C); from the male of L. yamasakii in having the elevated cumulus longer and with fewer spines (Fig. 4B, C); from males of L. hatyai sp. nov. and L. keeratikiati sp. nov. in having the tibial apophysis with shorter setae (Fig. 4A–C), the cumulus noticeably elevated (Fig. 4C), and a larger subtegular apophysis (Fig. 4F); from males of other Liphistius species in having adjoining embolic parts (Fig. 4E, G) and a strongly elevated cumulus (Fig. 4C).

Figure 4. 

Male palp and female genitalia of Liphistius inthanon Zhan & Xu, sp. nov. A palp, prolateral view B palp, ventral view C palp, retrolateral view D–G palp, distal views H–J vulva, dorsal view K–M vulva, ventral view A–C XUX-2017-372A (holotype) D–G XUX-2017-373A H, K XUX-2017-372 I, L XUX-2017-379 J, M XUX-2017-381. Abbreviations used: CDO = central dorsal opening; GA = genital atrium; RC = receptacular cluster; CT = contrategulum; Cu = cumulus; E = embolus; PC = paracymbium; PeP = paraembolic plate; ST = subtegulum; T = tegulum; TiA = tibial apophysis. Scale bars: 0.5 mm.

The female of L. inthanon sp. nov. differ from the female of L. bristowei in having the poreplate with an arched anterior margin (Fig. 4H–M); from females of L. lannaianus and L. yamasakii in having the central dorsal opening larger oval (Fig. 4H–J); from the female of L. maewongensis in having the genital atrium with a wider posterior margin (Fig. 4L, M); from females of L. hatyai sp. nov. and L. keeratikiati sp. nov. in having a wider posterior stalk (Fig. 4I, J), longer oval central dorsal opening (Fig. 4H–J), and larger receptacular cluster (Fig. 4K–M); from females of other Liphistius species in having a wider posterior stalk (Fig. 4H–M).

Male (holotype). Carapace reddish brown, with a few short, scattered bristles; opisthosoma olive-green, with 12 dark tergites, close to each other, 2–6 larger than others, fifth largest; chelicerae robust, promargin of cheliceral groove with 9 denticles of variable size; labium yellowish brown, separated from sternum; sternum yellowish brown, with a few weakly setae on the anterior tip and many long setae on the elongated posterior tip; legs dark brown, and with strong setae and spines, without distinct annulations and with 3 tarsal claws; 8 spinnerets. Measurements: BL 17.38, CL 9.25, CW 9.62, OL 7.42, OW 5.66; eye sizes and interdistances: AME 0.18, ALE 0.87, PME 0.49, PLE 0.63, AME–AME 0.09, AME–ALE 0.22, PME–PME 0.12, PME–PLE 0.15, ALE–PLE 0.10, ALE–ALE 0.15, PLE–PLE 0.51, AME–PME 0.13. Labium 1.00 long and 0.51 wide. Sternum 4.49 long and 1.35 wide. Leg I 28.45 (7.56 + 4.16 +6.37 + 6.50 + 3.86), leg II 31.13 (8.54 + 4.01 + 6.63 + 7.70 + 4.25), leg III 34.09 (8.54 + 4.24 + 6.58 + 9.61 + 5.12), leg IV 42.23 (10.62 + 3.78 + 8.60 + 12.70 + 6.53).

Palp : tibial apophysis with 4 stouter spines, and several strong spines on subterminal ledge (Fig. 4A–C); paracymbium with many short strong setae situated at the tip (Fig. 4B, C); cumulus distinctly elevated with several spines on tip (Fig. 4C); subtegular apophysis large, strongly developed (Fig. 4D, F); proximal edge of contrategulum elevated (Fig. 4F, G); tegulum lunate, with dentate margin (Fig. 4G); embolic parts adjoining (Fig. 4E, F, G); embolus with 2 longitudinal ridges reaching the tip distally (Fig. 4D, E, G).

Female (XUX-2017-372). Carapace reddish brown, with a few short, scattered bristles; opisthosoma olive-green, with 12 dark brown tergites, close to each other, 2–6 larger than others, fifth largest; 8 eyes on dark ocular tubercle; chelicerae robust, reddish brown; promargin of chelicerae groove with 12 denticles of variable size; labium reddish brown, fused with sternum; sternum reddish brown and with several setae; legs reddish brown, with strong setae and spines, without distinct annulations, with 3 tarsal claws; 8 spinnerets. Measurements: BL 30.6, CL 12.18, CW 11.74, OL 17.49, OW 17.38; eye sizes and interdistances: AME 0.14, ALE 1.02, PME 0.52, PLE 0.82, AME–AME 0.19, AME–ALE 0.25, PME–PME 0.09, PME–PLE 0.16, ALE–PLE 0.13, ALE–ALE 0.16, PLE–PLE 0.66, AME–PME 0.12. Labium 2.78 long and 1.85 wide. Sternum 5.15 long, 1.81 wide. Palp 22.82 (7.08 + 4.24 + 5.87 + 5.63), leg I 26.61 (8.27 + 4.51 + 5.71 + 4.80 + 3.32), leg II 26.59 (7.85 + 4.96 + 5.44 + 5.16 + 3.18), leg III 28.88 (8.66 + 4.70 + 5.41 + 6.41 + 3.70), leg IV 33.46 (9.37 + 3.76 + 7.23 + 8.79 + 4.31).

Genitalia : poreplate with a long, oval central dorsal opening, and with projecting posterior corners; receptacular cluster racemose and large; posterior stalk wide, lateral margins of genital atrium with some hairs (Fig. 4H–M).

The species epithet “inthanon” is a toponym referring to the type locality, Doi Inthanon National Park.

Northern Thailand (Chiang Mai Province) (Fig. 2).

The range of females’ measurements (N = 7): BL 16.92–30.6, CL 7.90–12.18, CW 7.53–11.74, OL 8.5–17.49, OW 6.63–17.38. The number of denticles on the promargin of cheliceral groove varies from 12–14 (N = 7). The examined female genitalia were found to differ in that the posterior margin of genital atrium can be narrow, slightly W-shaped (Fig. 4H, K), or wide and straight (Fig. 4I, J, L, M), and the shape of poreplate anterior margin can slightly vary (Fig. 4K–M).

Liphistius inthanon sp. nov. can be assigned to the bristowei-group based on the following characters: the male palp has a pronounced, elevated cumulus (Fig. 4C); the embolic parts are adjoining (Fig. 4D, G), the sclerotised part of the embolus bears two longitudinal ridges reaching the tip (Fig. 4D, G), and, except for L. marginatus, all have a large subtegular apophysis (Fig. 4F); the poreplate has a wide posterior stalk and a projecting posterior corner (Fig. 4H–M). The bristowei-group contains L. bristowei, L. lannaianus, L. maewongensis, L. marginatus, L. yamasakii (Schwendinger 1990), and L. inthanon sp. nov.

Holotype : Thailand • 1 ♂, Chumphon Province, Sawi District, Khao Thalu Subdistrict, Nam Lot Cave. 10.23°N, 98.94°E; alt. 30 m; 25 November 2017; F.X. Liu, D. Li, X. Xu, V. Sivayyapram leg.; XUX-2017-439. Paratypes: Thailand • 1 ♂, 3♀♀, same data as for the holotype; XUX-2017-439, XUX-2017-431, 436, 438.

The male of L. keeratikiati sp. nov. can be distinguished from the male of L. fuscus Schwendinger, 1995 in having the paraembolic plate scale-like and arched (Fig. 5A, E), and the tibial apophysis slightly wider basally (Fig. 5A–C), while in L. fuscus the paraembolic plate is broadly rounded; from the male of L. phuketensis Schwendinger, 1998 in having the tibial apophysis with four setae (Fig. 5B, C); from the male of L. schwendingeri Ono, 1988 in having a longer embolus (Fig. 5A, B, E), the contrategulum with fewer wrinkles proximally (Fig. 5G), and a smaller tegulum (Fig. 5B, C, G); from the male of L. hatyai sp. nov. in having the tibial apophysis with longer setae and the paracymbium narrower (Fig. 5A–C); from the male of L. inthanon sp. nov. in having the subtegular apophysis absent (Fig. 5B, F) and the paraembolic plate scale-like (Fig. 5E); from males of other Liphistius species in having the spines on the cumulus slightly separated from setae on the paracymbium (Fig. 5A–C).

Figure 5. 

Male palp and female genitalia of Liphistius keeratikiati Zhan & Xu, sp. nov. A palp, prolateral view B palp, ventral view C palp, retrolateral view D–G palp, distal views H vulva, dorsal view I vulva, ventral view J vulva, lateral view A–G XUX-2017-439 (holotype) H–J XUX-2017-431. Abbreviations used: CDO = central dorsal opening; GA = genital atrium; RC = receptacular cluster; CT = contrategulum; Cu = cumulus; E = embolus; PC = paracymbium; PeP = paraembolic plate; ST = subtegulum; T = tegulum; TiA = tibial apophysis. Scale bars: 0.5 mm.

The female of L. keeratikiati sp. nov. differs from the female of L. fuscus in having the anterior margin of the poreplate slightly curved (Fig. 5H, I); from the female of L. phuketensis in having the anterior margin of the poreplate slightly narrower (Fig. 5I); from the female of L. schwendingeri in having the receptacular cluster slightly larger (Fig. 5I); from females of other Liphistius species in having the arched poreplate lacking lateral edges (Fig. 5I, J) and much wider than long (Fig. 5H, I), and the central dorsal opening situated in the lower center of the poreplate (Fig. 5H).

Male (holotype). Carapace light yellow, with a few short, scattered bristles; opisthosoma yellow, with 12 tergites, with light brown patches; close to each other, 2–6 larger than others, fifth largest; chelicerae robust, promargin of cheliceral groove with 6 denticles of variable size; labium yellow and separated from sternum; sternum yellow, with a few short setae on anterior tip and many long setae on the elongated posterior tip; legs with strong setae and spines; with white annulations, with 3 tarsal claws; 8 spinnerets. Measurements: BL 15.61, CL 6.92, CW 6.72, OL 7.24, OW 5.07; eye sizes and interdistances: AME 0.09, ALE 0.74, PME 0.43, PLE 0.54, AME–AME 0.12, AME–ALE 0.15, PME–PME 0.03, PME–PLE 0.10, ALE–PLE 0.08, ALE–ALE 0.05, PLE–PLE 0.38, AME–PME 0.07. Labium 1.04 long and 0.76 wide. Sternum 3.02 long, 0.95 wide. Leg I 21.60 (5.41 + 1.81 + 4.68 + 4.79 + 4.91), leg II 23.29 (4.97 + 2.19 + 7.68 + 5.96 + 2.49), leg III 14.64 (missing metatarsus and tarsus) (6.17 +2.87 + 5.60 + NA + NA), leg IV 30.16 (7.69 + 3.08 + 5.97 + 9.31 + 4.11).

Palp : tibial apophysis pronounced elevated, with four tapering spines of similar length (Fig. 5A–C); paracymbium with short, strong setae situated at tip (Fig. 5C), and 5 tapering spines on elevated cumulus (Fig. 5A, B); subtegulum without apophysis (Fig. 5B, F); contrategulum with a process distally, and with several wrinkles proximally (Fig. 5B, G); tegulum with a dentate edge (Fig. 5B, G); embolic parts detached (Fig. 5B), paraembolic plate scale-like, semicircular (Fig. 5A, E); embolus slender, with a few denticulations at the tip (Fig. 5A, B, G).

Female (XUX-2017-431). Carapace light brown, with few short, scattered bristles; opisthosoma gray, with 12 brown tergites, close to each other, 2–6 larger than others, fifth largest; eight eyes on darkened ocular tubercle; chelicerae robust, brown, promargin of chelicerae groove with 11 denticles of variable size; labium yellow, separated from sternum; sternum yellow with several setae; legs with strong hairs and spines; with brown and yellow annulations and 3 tarsal claws; 8 spinnerets. Measurements: BL 16.9, CL 7.21, CW 6.67, OL 9.93, OW 7.57; eye sizes and interdistances: AME 0.07, ALE 0.64, PME 0.30, PLE 0.51, AME–AME 0.08, AME–ALE 0.16, PME–PME 0.06, PME–PLE 0.09, ALE–PLE 0.09, ALE–ALE 0.08, PLE–PLE 0.41, AME–PME 0.09. Labium 1.49 long and 0.76 wide. Sternum 3.40 long, 1.13 wide. Palp 13.37 (4.74 + 2.25 + 3.32 + 3.06), leg I 16.58 (5.65 + 2.72 + 3.62 + 2.92 + 1.67), leg II 19.13 (5.56 + 3.06 + 3.76 + 3.71 + 2.05), leg III 17.14 (4.99 + 2.83 + 3.91 + 3.92 + 2.29), leg IV 24.73 (6.82 + 2.31 + 5.39 + 7.03 + 3.18).

Genitalia : poreplate much wider than long, arched (Fig. 5J), lateral edges absent (Fig. 5I); central dorsal opening situated in the lower center of poreplate (Fig. 5H); receptacular cluster simple (Fig. 5I).

The specific name is dedicated to Mr Kaweesak Keeratikiat for providing information on the locality of the species.

South-central Thailand (Chumphon Province) (Fig. 2).

Range in female measurements (N = 3): BL 16.45–18.89, CL 7.15–7.49, CW 6.45–7.15, OL 8.79–10.46, OW 6.76–8.99. The number of denticles on the promargin of cheliceral groove varies from 6–13 (N = 3).

Liphistius keeratikiati sp. nov. can be assigned to the trang-group according to the morphology of male palp and female genitalia, see the remarks of hatyai sp. nov. The new species can be assigned to the species complex C of the trang-group. In males, the palp possesses the contrategulum with wrinkles proximally (Fig. 5B, G), the tegulum has a dentate edge (Fig. 5C, F, G), the spines on the elevated cumulus are slightly, distinctly separated from the setae on the paracymbium, and the apex of the embolus bears a few denticulations (Fig. 5A–C, E, G). In females, the poreplate is lacking lateral edges (Fig. 5I), arched (Fig. 5J), wider than long (Fig. 5H–I), and lacking a posterior stalk (Fig. 5H, I). Currently, the species complex C contains L. fuscus, L. phuketensis, L. schwendingeri (Schwendinger et al. 2019), and L. keeratikiati sp. nov.