Type species: Liphistius desultor Schiödte, 1849, described from female holotype deposited in the Natural History Museum of Denmark (= Zoological Museum of the University of Copenhagen).

Diagnosis and characterisation: See Platnick & Sedgwick (1984); habitus as in Fig. 2A-B.

Diagnosis: Distinguished from the trang-group by an indistinctly split embolus proper (Fig. 3B, D, F, H) and from all other species groups by distal edge of contrategulum proventrally ending in a V-shaped or U-shaped row of denticles (the clearest synapomorphy of this group; Figs 4C; 6A, C; 8A, D). Vulval plate strongly sclerotised, with a large, undivided CDO and with a large, undivided receptacular cluster (Figs 5, 7, 9).

Description: Small to large species (carapace length of males 4.36-12.04, carapace width 3.64-10.99); body colouration uniformly dark (in large species: L. malayanus and L. endau; Fig. 2A) or brown with annulated legs and palps (in smaller species: L. johore and L. gracilis sp. nov.). Male palp with membranous and sclerotised part of embolus proper in close contact with each other (both distinctly separated in tranggroup), together forming a closed tube; dorsal wall of sclerotized part of embolus proper straight and directed proventrad (L. malayanus; Fig. 4G-H), curved prodorsad (L. endau; Fig. 6F, H) or bent prodosad (L. gracilis sp. nov.; Fig. 8F-G); para-embolic plate short or absent (Figs 4D, 6E); tegulum with coarsely dentate proximal edge (corresponding to dorsal extension of terminal apophysis in Heptathelinae; see Schwendinger & Ono, 2011), distal margin (corresponding to marginal apophysis) indistinct, not drawn into a prominent edge (Figs 4D-F, 6D-E, 8E); contrategulum with indistinct ventral process or without, distal edge carrying several teeth, prolateral ones numerous, tiny and forming a serrate crest (L. gracilis sp. nov.; Fig. 8D-G), few and small (L. malayanus; Fig. 4G-L), or only represented by a single large triangular tooth (L. endau; Fig. 6F, H-O), proventral denticles of distal edge forming a downward-curved V-shaped (Fig. 4C) or U-shaped row (Figs 6A, C; 8D), dorsal apex of contrategulum widely to narrowly tongue-shaped (Figs 4G-L; 6F, H-O; 8F-G); subtegulum always without apophysis; paracymbium quite small and shallow to large and deep (Figs 4A, 6B, 8B); cumulus very low, indistinct, carrying several long, thick bristles; retrolateral tibial apophysis relatively short and carrying long apical megaspines (L. malayanus and L. endau; Figs 4A-B, 6A-B) or quite long and carrying distinctly shorter megaspines (L. gracilis sp. nov.; Fig. 8A-C, I-J); both apical lobes of cymbium weakly developed (Fig. 8D). Vulva (Figs 5, 7, 9, 10A): poreplate strongly pigmented and sclerotised, wider than long, its anterior margin more or less distinctly recurved, with or without anterolateral lobes or processes, never with mediolateral processes; CDO and ventral receptacular cluster large to very large, both undivided; posterior stalk wide (in L. endau even covering entire width of genital atrium; Fig. 7), somewhat trapezium-shaped or almost triangular (Fig. 10A); genital atrium with few to many lateral hairs and with no, few or numerous median hairs.

Species included: Liphistius endau, L. malayanus, L. gracilis sp. nov. and presumably also L. johore (male still unknown).

Relationships: Species of the malayanus-group are most similar and probably closely related to species of the tioman-group. The two large species, L. malayanus and L. endau, are closer to each other than to the much smaller L. gracilis sp. nov. (distinguished by serrate distal edge of contrategulum and by sharp bend in wide dorsal wall of sclerotised part of embolus proper). Small size, details of the vulval plate and geographical proximity suggest a close relationship between L. gracilis sp. nov. and L. johore, but this can only be confirmed when the male of the latter species is discovered.

Distribution: All described species of this group occur in the western and southern part of peninsular Malaysia (Fig. 1, localities 1-9); an undescribed species appears to be present in the northeastern corner of the country (Fig. 1, locality 13).

Liphistius malayanus Abraham, 1923b: 770-774, text-fig. 1a-b (description of female). – Abraham (1929: 674-676, pl. 1, figs 3-4, pl. 2, figs 11-16; first description of male). For synonymy and other taxonomically relevant publications see the World Spider Catalog, 2017.

Liphistius malayanus cameroni Haupt, 1983: 282, figs 3e, 4d, 5f, 6f (description of male and female). New synonym

Type material: BMNH; female holotype of L. malayanus (not examined); Negri Sembilan, Gunung Angsi near Seremban; XII.1922; leg. F. de la Mare Norris. – ZMH (A16/84); male holotype and female paratype of L. malayanus cameroni (examined); Pahang, Cameron Highlands, Berinchang; 4.III.1981; leg. J. Haupt & T. Dach.

Remark: A L. malayanus male (not examined) from Fraser's Hill was described by Abraham (1929) and deposited in the BMNH. Haupt (1983: 281, figs 3d, 4c) re-described this specimen and referred to it as the paratype, but it has no type status, having been collected in November 1928, after the publication of the original description of the species.

Material examined: ZMH (A16/84); male holotype [left palp detached, macerated and partly collapsed, cymbium and hematodocha cut open to expose tendons inside, all strong spines deformed; right palp without tegulum, contrategulum and embolus complex] (matured VII.1981) and female paratype of Liphistius malayanus cameroni; Pahang, Cameron Highlands, Berinchang; 4.III.1981; leg. J. Haupt & T. Dach. – Collection of Joseph Koh, n° JK.14.10.05.0001; 1 male; Pahang, Cameron Highlands, Taman Tringkap NE of Brinchang, 4°28′26″N, 101°22′58″E; 5.X.2014; leg. N. Bay. – MHNG; 1 female, 1 juv. male; Perak, Cameron Highlands, ca 1 km SW of Ringlet, 1060 m; 21.I.1995; leg. P.J. Schwendinger. – MHNG (sample TM-15); 1 juv. male; Pahang, Cameron Highlands, Tanah Rata, trail n° 9, 4°27.620′N, 101°23.400′E, 1210 m; 29.IX.2001; leg. L. Monod. – SMF 7425/2 (ex coll. C. F. Roewer); 2 juv. females (labelled as “1 male and 1 female, det. Roewer, 1935″); Pahang, “Ginting Kial Highlands” [probably a peak in the Cameron Highlands]. – SMF 40602; 1 male; Pahang, Fraser's Hill, roadside at Jalan Girdle; 19.XII.2001; leg. S. Huber. – SMF 56206; 1 female; Pahang, Fraser's Hill, roadside at Jalan Girdle; 19.XII.2001; leg. S. Huber. – MHNG (ex coll. S. Huber, sample O-4, C); 1 male (matured end March 2002), 1 female (moulted 24.VI.2002); Pahang, Fraser's Hill, Jalan Guillemard; 18.XII.2001; leg. S. Huber. – SMF 64093; 1 female; Pahang, Fraser's Hill, 1300 m, 3°43.105′N, 101°45.164′E; 17.VI.2013; leg. P. Jäger. – MHNG (sample MAL-04/02); 1 male (matured 18.IX.2004), 2 females (moulted 30.VII.2004, 24.I., 28.VIII., 8.XII.2005; 19.X.2004, 30.VII.2005), 4 juveniles; Pahang, Genting Highlands, 3°25′42″N, 101°47′41″E, 1650 m; 18.-19.V.2004; leg. P.J. Schwendinger. – SMF 21946/1; 1 female; without locality data; 10.XI.1933; ex coll. Wiehle, don. W. S. Bristowe. – SMF 60037; 1 female; “Selangor” [should be Pahang], Genting Highlands, 1800 m; 3.II.1989; leg. U. Maschwitz, don. H. Steiner. – SMF 40016; 1 female; Pahang, Genting Highlands, 1200 m; 27.VII.2001; leg. A. Kovac. – MHNG (sample SIM-01/14); 1 male (matured end VIII.2003), 1 female (moulted 31.XII.2001), 1 juvenile; Selangor, Templer Park, 3°17′55″N, 101°37′13″E, 230 m; 13.VII.2001; leg. P.J. Schwendinger.

Diagnosis: Large, dark-coloured species in both sexes. Males distinguished from those of other species in the malayanus-group by tibial apophysis situated distinctly lower (more proximal) than distal margin of palpal tibia (Fig. 4B); distal edge of contrategulum carrying a series of small denticles (Fig. 4G-L), those at proventral end arranged in a V-shaped row (Fig. 4C), 1-2 at prodorsal end isolated and slightly enlarged; dorsal apex of contrategulum large, wide, tounge-shaped and asymmetrical, its prolateral margin more strongly arched than retrolateral margin (Fig. 4G-L); dorsal wall of embolus proper as wide as ventral wall (Fig. 4G-H); membranous part of embolus proper distally narrower than proximally (Figs 3F, H, 4C). Females distinguished by vulval plate (Fig. 5) with widely trapezium-shaped (posteriorly widest) posterior stalk not connected to pigmented lateral patches in genital atrium; poreplate with distinct anterolateral lobes; CDO large, very variable in shape, mostly longer than wide; receptacular cluster very large, in most cases protruding beyond straight or procurved anterior margin of poreplate.

Additions to description of male: Scopula: See paragraph “Variation”.

Palp: Both apical lobes of cymbium equally short, dorsal one usually slightly more pointed than ventral one. Retrolateral apophysis of palpal tibia entire, situated at a clearly more proximal level than distal margin of article (Fig. 4B), carrying four long apical megaspines (Fig. 4A-B). Paracymbium relatively small and shallow, about as deep as cymbium in retroventral view (Fig. 4A; Platnick & Sedgwick, 1984: fig. 64); cumulus low, carrying a compact group of moderately long thick bristles. Subtegulum without apophysis. Tegulum with quite few small teeth only in retrodorsal portion of proximal edge, distal margin not drawn into an edge (Fig. 4D-F). Contrategulum with very indistinct, widely rounded ventral process (Fig. 4G-H); distal edge with a series of small denticles, those at proventral end arranged in a V-shaped row (Fig. 4C), 1-2 enlarged denticles or a sharp keel in prolateral to prodorsal section of distal edge (Fig. 4G-L); dorsal apex of contrategulum developed as a large, linguiform, strongly projecting horizontal plate, its prolateral margin being more strongly arched than its retrolateral margin (Fig. 4G-L). Embolus complex with para-embolic plate developed as a more or less distinctly elevated, rounded edge (Fig. 4D-F); below it a second small edge present in some cases (Fig. 4D); sclerotised and membranous part of embolus proper in contact for almost their entire length (Fig. 3D, F, H); sclerotized part with numerous longitudinal ribs carrying tiny denticles distally (Fig. 3B, D, F, H), its dorsal and ventral walls equally wide and parallel to each other (Figs 3D; 4G-H), dorsal wall ending in indistinct rounded lobe (Fig. 4C-F); membranous part distally narrower than proximally (Fig. 3D, F, H), its proximal portion essentially unpigmented.

Additions to description of female: Vulval plate (Fig. 5) always with few to many hairs laterally in genital atrium, rarely also posteromedially (Fig. 5I). Posterior stalk more or less distinctly trapezium-shaped (posteriorly wider than anteriorly), densely pitted in posterior portion. Poreplate wider than long, with a pair of anterolateral lobes projecting anteriad and slightly bent ventrad; several of these lobes more or less distinctly constricted at base (Fig. 5A-C, E); anterior margin of poreplate straight to procurved; lateral and posterolateral margins on ventral side of pore plate bulging (Fig. 5B, J). CDO large and of variable shape, mostly longer than wide, rarely wider than long (Fig. 5E). Receptacular cluster large and strongly racemose, in most cases protruding beyond anterior margin of poreplate, rarely not (Fig. 5G).

Taxonomic remarks: Liphistius malayanus cameroni has been the only subspecies in the genus Liphistius. As specimens later collected near the type locality show no relevant differences from specimens of the nominal subspecies, this case became doubtful. Thus a re-examination of the types of Liphistius malayanus cameroni was necessary, and it revealed that they differ considerably from the illustrations and explanations given in the original description of this subspecies. The vulva of the female paratype was cleared of tissue, but the ventral wall of the genital region (with bristles still attached) was not removed, thus not allowing a clear view of the ventral side of the poreplate. It therefore appears that Haupt's (1983) illustrations were made with the help of a compound microscope without using a drawing tube, which would explain why the poreplate in the original illustrations is wider than in reality and why the outlines of the posterior stalk are too rounded. The same is probably also the case in the dorsal view of the same poreplate, it being too wide, with a triangular CDO leading to the receptacular cluster and with a peculiar double posterior margin of the posterior stalk (see Haupt, 1983: figs 5f, 6f). In fact the vulva of the paratype of L. malayanus cameroni does not differ from the vulvae of the other L. malayanus females examined (Fig. 5A-B cf. Fig. 5C-J) in any way that would warrant a distinct taxonomic status.

The illustrated differences between the male holotype of L. malayanus cameroni and the male that Haupt incorrectly referred to as a paratype of L. malayanus (in BMNH, not examined) are mostly due to a slightly different view of the palp and due to the fact that the palpal organ of the BMNH male is expanded (turned clockwise by about 90°) (Haupt, 1983: figs 3D, 4C cf. figs 3E, 4D). None of the L. malayanus males examined has the tibial apophysis of the male palp situated as far distally as illustrated by Haupt (1983: figs 3D, 4C), therefore this may also be due to a different view. The embolus proper of the left palp of the L. m. cameroni holotype (that of the right palp is missing, together with tegulum and contrategulum) is slightly narrower than that of the second male from the Cameron Highlands and that of other conspecific males examined. This difference is presumably connected to size: the L. m. cameroni holotype is the smallest among the L. malayanus males examined. The character in which the holotype of L. m. cameroni most visibly differs from other conspecific males is a slightly elevated para-embolic plate (Fig. 4E), which is also present in the second male from the Cameron Highlands (Fig. 4F). However, an indistinctly elevated para-embolic plate is present in one of the two males from the Fraser's Hill, whereas all other conspecific males possess only a distinct edge at this place (Fig. 4D). Thus the holotype of L. m. cameroni differs from other conspecific males examined in possessing the most extreme states of three quite variable characters, but it is not representative for the population in the Cameron Highlands. I consequently place Liphistius malayanus cameroni in the synonymy of Liphistius malayanus which makes the species monotypic again.

Variation: Carapace lengths in males (n=6; including the holotype of L. m. cameroni) 7.90-11.23, carapace widths 7.04-9.69; in the largest female from each locality (n=4; not including the paratype of L. m. cameroni) 10.86-16.67 and 10.00-14.07, respectively. The holotype of L. m. cameroni is the smallest male examined, with carapace length 7.90, width 7.04; in the second male from the Cameron Highlands it is 9.63 and 8.40, respectively. The largest female is from the Templer Park, measuring 16.67 and 14.07; the second largest female is from the Fraser's Hill, measuring 16.54 and 14.07. The female paratype of L. m. cameroni measures 9.26 and 7.53, the second female from the Cameron Highlands 10.86 and 10.12.

The apical megaspines on the tibial apophysis are quite variable in length, but not as thin as illustrated in Murphy & Platnick (1981: figs 8, 11, 14, 17, 20) and in Platnick & Sedgwick (1984: figs 63-67); the ventral-most of them is mostly bent distad and has a thin, long apex (Fig. 4A-B). The ventral megaspine is bent in the male from the Templer Park (Fig. 4A), whereas in the holotype of L. m. cameroni (Fig. 4B), in a male from the Genting Highlands, and also in a second male from the Cameron Highlands it is slightly sigmoid.

The distal edge of the contrategulum carries one or two slightly enlarged dorsal denticles at some distance from the projecting dorsal apex (Fig. 4G, I-L); in two males there is a sharp edge at the same place (Fig. 4H), but only on one palp. These larger denticles are followed by a variable number of smaller proventral denticles.

The para-embolic plate is always low, most visibly elevated (but not with a sharp edge) in the holotype of L. m. cameroni (Fig. 4E), in a second male from the Cameron Highlands (Fig. 4F) and in a male from the Fraser's Hill. In the second male from the Fraser's Hill it is not elevated; the remaining males examined are intermediate (Fig. 4D). An indistinct second edge, situated below the indistinct para-embolic plate, is found in the only available male from the Genting Highlands and in the only available male from the Templer Park (Fig. 4D), but not in other males examined (Fig. 4E-F). The size of the embolus proper varies from relatively thick in the male from the Templer Park (Fig. 4D) to distinctly more slender in the holotype of L. m. cameroni (Fig. 4E); the other males are intermediate (Fig. 4F). The longitudinal ribs on the sclerotised part of the embolus proper vary from indistinct to distinct (Fig. 3B, D, F, H), so do the denticles on them.

There is considerable variation in the extent of the tarsal scopulae in males. The scopula on tarsus I is always thin and undivided, covering two-thirds of the ventral side in most males, only one half in the male from the Genting Highlands. The scopula on tarsus II is slightly denser than on tarsus I, undivided and covering two-thirds of the ventral side in all males. Tarsal scopula III is like tarsal scopula II in most males, in the non-type male from the Cameron Highlands it is light. Tarsal scopula IV is dense in all males except for the non-type male from the Cameron Highlands (light), undivided in all males except for the same male from the Cameron Highlands (distinctly divided) and for the male from the Genting Highlands (indistinctly divided), covering two-thirds of the ventral surface in most males except for a male from the Cameron Highlands (covering only half of the surface) and the male from the Templer Park (covering three-quarters). The male holotype of L. malayanus cameroni had already been returned to its depository when the tarsal scopulae were examined. Even if both males from the Cameron Highlands differed from those of other localities in shape and extent of their tarsal scopulae (especially on tarsus IV), this would not warrant a subspecific distinction. The tarsal scopula in this species is too variable to be of high taxonomic value.

The vulval plates of large females have more lateral hairs in the genital atrium than those of small females (Fig. 5I cf. Fig. 5H). In the largest female (from the Templer Park) hairs are also present posteromedially in the genital atrium (Fig. 5I). The shape of the posterior stalk is quite variable but mostly trapezium-shaped (Fig. 5). Even more variable is the size and shape of the CDO of the poreplate, ranging from (mostly) longer than wide to (rarely) wider than long, from quadrangular to pentangular and from near-triangular to near-circular (Fig. 5A, C-I). The receptacular cluster is always large, undivided and has a complex structure, covering most of the ventral side of the poreplate (Fig. 5B, J); only in one female examined (from the Genting Highlands) does it not protrude beyond the anterior margin of the poreplate (Fig. 5G). The anterior poreplate margin varies from more or less strongly invaginated (in most cases; Fig. 5A-E, G-J) to slightly arched medially (Fig. 5F).

Distribution: Liphistius malayanus is known from several lowland and upland localities in the western part of central Malaysia (Perak, Pahang, Selangor and Negeri Sembilan; Fig. 1, localities 1-5). See also Platnick & Sedgwick (1984: 24). Liphistius records from other localties near Kuala Lumpur (e.g. Klang Gates; Murphy & Murphy, 2000: plate 2.5) can also be attributed to L. malayanus. This species has a relatively wide geographical range (the northernmost localitiy, in the Cameron Highlands, is about 280 km away from the southernmost locality, Gunung Angsi) and a large vertical distribution (from 230 to 1800 m altitude).

Biology: Spiders were collected from quite different habitats: soil on exposed sides of roads and trails inside and outside forests (but never very far from a forest), sides of erosion gullies, sloping forest floor, and in decomposing wood of logs lying on the forest floor. Trapdoors of females were up to 3.8 cm long and 6.5 cm wide. Signal lines (not more than eight) were usually about 10 cm long, in one immature male even 24 cm long. The longest burrow measured was 20 cm long. Two penultimate males had trapdoors with a length of 1.9-2.2 cm and a width of 3.0-3.4 cm, which is probably normal. A juvenile male (moulted again but died before reaching maturity) from near Ringlet in the Cameron Highlands had a surprising 2.9 cm long and 5.0 cm wide trapdoor.

The female paratype of L. m. cameroni and two other females (from Fraser's Hill and Templer Park) show bite marks on their carapaces and chelicerae. These were probably caused by specimens of Ljunghia bristowi, a species of ectoparasitic laelapid mites originally described from L. malayanus (see Halliday & Juvara-Bals, 2016: 837-845).

Mature males were collected in the field in October and December; in captivity males matured between July and March. The mating period in L. malayanus appears to be much longer than in other congeneric species. I did not find any egg cases.

Description of male (matured 27.II.2003): Colour in alcohol (much darker in life; colouration as in female, Fig. 2A): All sclerotised parts uniformly brown, except for cream-coloured proximal portion of chelicerae, cream-coloured membranes of prosoma and light brown opisthosomal membranes.

Bristles on carapace: Short bristles along all margins (strongest on posterior margin, longest behind, on and in front of eye mound); none on coxal elevations; five short bristles anterior to fovea.

Scopula: Tarsus I with thin scopula in distal half of ventral side, divided for its entire length by narrow, pale, glabrous longitudinal median stripe and by some bristles; tarsus II with slightly denser scopula in distal three-quarters, only distally divided by median stripe; tarsi III-IV with dense scopula covering distal four-fifths, only distally divided by median stripe.

Cheliceral teeth: Eleven small ones on promargin of cheliceral groove on both sides.

Palp: Tibial apophysis situated distally, not clearly set back from distal margin of tibia, carrying four moderately long (dorsal ones shorter than ventral ones) megaspines (Fig. 6A-B). Both apical lobes of cymbium very short and equally rounded. Paracymbium basally very deep (Fig. 6B), its cumulus only slightly elevated, carrying stiff bristles reaching base of contrategulum (looking shorter in Fig. 6A because pointing ventrad rather than distad). Subtegulum without apophysis. Tegulum short and wide, coarsely dentate along entire proximal margin (Fig. 6D-E). Contrategulum with indistinct, widely arched proventral process (Fig. 6F); distal edge with denticles at proventral end distinctly elevated on a U-shaped ridge (Fig. 6A, C), with a single large triangular tooth prolaterally and with spatulate, quite symmetrical dorsal apex (Fig. 6F). Para-embolic plate only little elevated (Fig. 6A, C-E); sclerotised part of embolus proper strongly compressed dorso-ventrally, its dorsal wall distinctly wider than its ventral wall, curved prodorsad (Fig. 6F) and ending in a pronounced, prodorsad-directed, rounded lobe (Fig. 6C-E); membranous part of embolus proper distally much wider than proximally (Fig. 6C), its proximal portion slightly pigmented.

Measurements: Total length 22.30; carapace 9.26 long, 8.40 wide; opisthosoma 10.33 long, 7.62 wide; eye mound 1.34 long, 1.50 wide; palpal coxa 2.97 long, 1.98 wide; labium 0.69 long, 1.58 wide; sternum 4.06 long, 2.48 wide (1.29 on ventral surface); palp 15.83 long (4.43 + 2.79 + 5.74 + 2.87); leg I 27.21 long (7.21 + 3.61 + 5.90 + 7.05 + 3.44); leg II 28.11 long (7.21 + 3.61 + 6.15 + 7.62 + 3.52); leg III 30.99 long (7.54 + 3.77 + 6.56 + 9.02 + 4.10); leg IV 38.27 long (9.34 + 3.93 + 7.54 + 12.21 + 5.25).

Additions to description of female: Posterior margin of genital sternite more or less distinctly invaginated (Fig. 7A, C-H; 7I is a very small juvenile). Vulval plate (Fig. 7) strongly sclerotised and pigmented, roughly as long as wide, with a more or less distinct lateral constriction in posterior third. Genital atrium with many lateral hairs and in many cases also with additional median hairs; median zone of genital atrium clearly sunken below lateral zones (indistinct in small females). Posterior stalk wide, completely fused with poreplate and with strongly pigmented, buldging lateral parts of genital atrium (Fig. 7A, C-H). Poreplate entirely and strongly pigmented, its anterior margin slightly invaginated, with a pair of pronounced anterolateral lobes. CDO large to very large, its posterior margin not sunken, giving the opening the shape of a horseshoe (Fig. 7C-D, F) or of an open quadrangle (Fig. 7A, H); enlarged pores inside CDO leading to receptacular cluster (Fig. 7A, C-D, F); the latter large and complex but not or only slightly protruding beyond anterior margin of poreplate, divided into three more or less distinct subclusters (Fig. 7B, E, G).

Variation: Carapace lengths in males (n=7) 9.26-12.04, in the largest females (n=4) 13.33-14.94; carapace widths 8.09-10.99 and 10.74-12.84, respectively. The three males from Kota Tinggi have a shorter scopula on their anterior legs (I: very thin, medially divided, covering only distal quarter; II: slightly denser, only apically divided, covering distal half; III: dense, undivided, covering three-quarters; IV: dense, undivided, covering distal four-fifths) than males from the type locality and from an unknown locality (I: thin, divided, distal half; II: slightly denser, apically divided, distal tree-quarters; III-IV: dense, undivided, distal four-fifths).

Variation in the shape of the distal edge of the contrategulum is shown in Fig. 6F, H-O, variation in the shape of the vulval plate in Fig. 7. With age (and body size) the number of hairs in the genital atrium of females increases.

All females from Kota Tinggi have fewer hairs in the genital atrium than females from other localities. Among six medium to large-sized females from that locality only one large female has two hairs in a median position (Fig. 7C), all others (including another large female) have none.

The female holotype (in AMNH, not examined) has a carapace length of 7.5 and a width of 6.6, and is thus only about half the size of the three largest females examined. The illustrations of the holotype vulva (Sedgwick & Platnick, 1987: figs 1-2) differ from the vulvae examined by lacking distinct anterolateral lobes on the poreplate (also absent in the smallest juvenile female examined; Fig. 7I). The posterior stalk of the holotype vulva is not yet fused with the pigmented lateral zones of the genital atrium (as it is also the case in the two juvenile females examined; Fig. 7H-I) and it appears to lack lateral hairs (as it is the case in the smallest female examined; Fig. 7I). The holotype is therefore a juvenile female with a not fully developed vulval plate. As (according to the original description) its vulval plate corresponds quite well with vulval plates of juveniles examined from the same area (possibly even from the same locality), and as no other Liphistius species is known from that area, there is no doubt that these specimens are conspecific.

Distribution: Known from three localities in the northern and western part of Johor State (Fig. 1, localities 6-8).

Relationships: Large size, details of the male palp (e. g. shape of distal edge of contrategulum and its dorsal apex; shape of tibial apophysis) and details of the vulval plate (bulging lateral and posterolateral margins on ventral side of poreplate forming a distinct boundary between poreplate and posterior stalk) indicate that L. endau and L. malayanus are more closely related to each other than to L. johore and L. gracilis sp. nov.

Biology: The new specimens from the type locality or from very close to it (in the Endau-Rompin N.P.) were found on the banks of a stream in a rain forest; specimens from the Kota Tinggi Waterfall were collected from the sloping forest floor and soil banks on both sides of rain forest stream running over a series of falls. Most burrows were in the soil, but at the Kota Tinggi Waterfall two medium-sized burrows were constructed in the rotten wood of a fallen tree, with the signal lines spread over the wood surface. At the type locality the burrow entrance of a large female had nine signal threads, at the Kota Tinggi Waterfall four burrows (of three large females and one penultimate male) were equipped with nine signal lines; all other burrows had a maximum of eight lines running over rock, soil and tree roots. The longest signal line (of the largest female) was 34 m long, those of other burrows examined were not more than 21 cm long. Trapdoors of four penultimate males were 2.6-3.2 cm long and 4.0-4.7 cm wide; that of the largest female 3.3 and 5.9, respectively. The latter spider lived in a 35 cm long burrow. Males became mature in August, October and November (after 2-5 months in captivity) and in February (after one and a half years in captivity, therefore probably not corresponding to conditions in nature). One male (from the Kota Tinggi Waterfall) ate a half-dead cricket a few days after its last moult but not again later. I also observed this in newly matured males of L. dangrek Schwendinger, 1996, and it had been reported for males of L. desultor by Murphy & Platnick (1981: 46), but it occurs only rarely and probably only in large species. Usually Liphistius males cease feeding when becoming adult.

Egg cases were constructed (all of them in captivity or during transport) in June and July. One egg case built in Geneva by a female from the type locality was 5.0 cm long, 4.7 cm wide, 2.8 cm high and contained 453 eggs. Another female from the same locality built during the transport an egg case of tissue paper, with a 3 cm diameter, containing 331 eggs. A third, old and empty egg case at the Kota Tinggi Waterfall was 4.0 cm long, 54.6 cm wide, 1.8 cm high. A fourth female from the same locality built a tissue paper egg case of 4.2 cm diameter and 2.0 cm height, containing 395 light orange-coloured eggs.

Types: MHNG, sample SIM-01/07; male holotype (matured 13.VIII.2001), 5 male paratypes (matured 18.IX., 19.IX.2001, 5.I., 24.III., 3.VI.2002), 5 female paratypes; Malaysia, Johor, Kota Tinggi Waterfalls at foot of Gunung Muntahak, 170 m (evergreen rain forest); 24.-26.VI.2001, leg. P.J. Schwendinger. – SMF, sample TM-14/02; 1 male paratype (matured 5.XI.2016); same locality; 21.-22.VI.2014; leg. P.J. Schwendinger. – MHNG, SMF, sample Sum-00/01; 3 female paratypes [moulted 3.V. (allotype); early VIII.; 26.XII.2000; 16.V., 6.X.2001]; same locality; 4.II.2000; leg. P.J. Schwendinger.

Etymology: The species epithet is a Latin adjective meaning “gracile”, “slender”, “slim”, referring to the small body size of this species in comparison with L. malayanus and L. endau.

Diagnosis: Similar to L. malayanus and L. endau but much smaller in body size, with lighter body colouration and annulated legs and palps. Males distinguished from those of L. malayanus (Fig. 4) and L. endau (Fig. 6) by tibial apophysis of palp longer and directed more distad (Fig. 8A, C), carrying shorter megaspines, the dorsal one sitting clearly lower than the rest (Fig. 8B, H-J); paracymbium relatively deeper than in L. malayanus and shallower than in L. endau (Fig. 8A-C cf. Fig. 4A and Fig. 6B, respectively); distal edge of contrategulum serrate (Fig. 8A, D-G), its apex distinctly smaller (Fig. 8F-G cf. Fig. 4G-L and Fig. 6F, H-O); dorsal wall of sclerotised part of embolus proper ending in a sharply prodorsad-bent lobe (Fig. 8D-G) [both walls equally wide in L. malayanus (Fig. 4G-H); dorsal wall prodorsal-curved (not sharply bent) in L. endau (Fig. 6F, H)]. Vulva of L. gracilis sp. nov. (Fig. 9) different from vulvae of L. malayanus (Fig. 5) and L. endau (Fig. 7) by ventral side of poreplate without bulging lateral and posterolateral margins, thus transition between poreplate and posterior stalk much less marked; anterior margin of poreplate with less distinct anterolateral lobes; receptacular cluster wider than long, more distinctly protruding beyond anterior margin of poreplate. Vulva different from that of L. johore (Fig. 10A) by lacking pronounced anterolateral invatinations in the margin of the poreplate, by having a wide and indistinct transition between poreplate and posterior stalk (narrow and distinct in L. johore), any by possessing hairs in the genital atrium.

Description of male (holotype): Colour in alcohol (darker in life): Carapace mostly light brown, with darker patches along margins (medially broken on posterior margin), extending between coxal elevations; dark anterior margin enclosing very dark eye mound; dark W-shaped pattern behind eye mound indistinct; a few small dark spots on pars cephalica; a paramedian pair of dark patches anterior to fovea; chelicerae cream-coloured proximally, grey-brown distally; legs and palps mostly light brown, with darker zone proximally on patellae and tibiae of legs III-IV, and in proximal half of all metatarsi. Opisthosomal tergites cream-coloured, with dark marginal and central patches on tergite I, other tergites only with lateral and anterolateral patches; membranous part of opisthosoma cream-coloured. Sclerotised parts of ventral side of body light brown. Bristles on carapace: Short bristles along all margins (strongest on posterior margin, longest behind, on and in front of eye mound), on coxal elevations and in area behind fovea, none anterior to fovea.

Cheliceral teeth: Ten and eleven small ones on promargin of cheliceral groove of right and left chelicera, respectively.

Scopula: Tarsus I with very thin scopula confined to distal fifth of ventral side, divided for its entire length by narrow pale, glabrous longitudinal median stripe and by some bristles; tarsus II with very thin scopula in distal quarter, only distally divided by median stripe; tarsus III with dense scopula covering distal half, only distally divided by median stripe; tarsus IV with dense scopula covering distal three-fifths, only distally divided by median stripe.

Claws: Paired tarsal claws on anterior legs with 4-6 denticles, on posterior legs with 5-6 denticles; unpaired claws in most cases with a single denticle on anterior legs, none on posterior legs.

Palp: Tibial apophysis relatively long, situated at distal margin of tibia, inclined distad [Fig. 8A (showing paratype matured 5.I.2002), C], carrying four quite short megaspines, the dorsal one rising from a clearly more proximal position than the others [Fig. 8B, H (showing paratype matured 5.I.2002), I-J]. Both apical lobes of cymbium indistinct, prodorsal one more rounded than retrodorsal one (Fig. 8D). Paracymbium relatively large and basally deep [Fig. 8A-B (showing paratype matured 5.I.2002), C-D], its cumulus indistinctly elevated, carrying moderately long stiff bristles [Fig. 8A-B (showing paratype matured 5.I.2002), C]. Subtegulum without apophysis. Tegulum short and wide, coarsely dentate along entire proximal margin, not or only indistinctly connected to contrategulum on retrodorsal side (Fig. 8E). Contrategulum without ventral process; distal edge finely serrate, proventrally ending in a U-shaped row of denticles (Fig. 8D), dorsally ending in a spatulate asymmetrical apex (Fig. 8F). Para-embolic plate only little elevated (Fig. 8B, showing paratype matured 5.I.2002); sclerotized part of embolus proper without longitudinal keels or ribs and not carrying denticles (Fig. 8A-B, showing paratype matured 5.I.2002), its dorsal wall distinctly wider than its ventral wall, abruptly bent prodorsad approximately at a right angle (Fig. 8F) and ending in a pronounced angular lobe (Fig. 8D-E); membranous part of embolus proper apically widened and lying on lobe of dorsal wall of sclerotised part (Fig. 8D).

Measurements: Total length 11.32; carapace 4.67 long, 3.96 wide; opisthosoma 4.83 long, 3.64 wide; eye mound 0.68 long, 0.80 wide, AME well-developed; palpal coxae 1.38 long, 0.95 wide; labium 0.40 long, 0.79 wide; sternum 2.37 long, 1.50 wide (0.83 on ventral surface); palp 7.99 long (2.53 + 1.34 + 2.81 + 1.31); leg I 15.97 long (4.35 + 1.90 + 3.40 + 4.03 + 2.29); leg II 17.01 long (4.43 + 1.90 + 3.56 + 4.63 + 2.49); leg III 18.92 long (4.55 + 1.90 + 4.00 + 5.78 + 2.69); leg IV 23.89 long (5.38 + 2.06 + 5.10 + 8.03 + 3.32).

Description of female (allotype): Colour in alcohol (darker in life): As in male but with a more pronounced dark pattern on carapace, resulting in a more distinct W-shaped pattern behind eye mound, and with a pronounced, light brown, flower-shaped pattern on pars thoracica; all opisthosomal tergites with larger dark patches, tergites II-V additionally with dark anteromedian patches. Legs light brown, with dark proximal and subapical annulations on femora (also on palpal femur), tibiae, metatarsi (indistinct on posterior legs) and on tarsi (indistinct on posterior legs). Palpal tibia with dark proximal and apical annulation; tarsus mostly dark, proventral-distal area lighter.

Bristles on carapace: Slightly stronger and longer (especially in front of eye mound) than in male.

Cheliceral teeth: Eleven strong cheliceral teeth on promargin of cheliceral groove on both sides.

Claws: Paired tarsal claws on legs with mostly 4-5 denticles (one claw with only two); unpaired claws with two denticles on legs I-III, 1-2 on leg IV. Tarsi without scopula.

Vulva: Vulval plate (Fig. 9A-B) distinctly sclerotised and pigmented, wider than long. Genital atrium uniformly flat, its lateral parts not bulging on dorsal side, with several lateral and median hairs. Posterior stalk wide, indistinctly perforated with tiny micropores; transition to poreplate indistinct, only slightly constricted. Anterior margin of poreplate slightly to distinctly invaginated, with indistinct anterolateral lobes, without anterolateral invaginations; ventral side of poreplate without bulging lateral and posterolateral margins. CDO large, undivided, irregularly quadrangular, close to anterior margin of poreplate. Receptacular cluster clearly wider than long, deep, distinctly projecting beyond anterior margin of poreplate.

Measurements: Total length 13.53; carapace 5.34 long, 4.19 wide; opisthosoma 5.70 long, 4.15 wide; eye mound 0.74 long, 0.83 wide; palpal coxae 1.74 long, 1.27 wide; labium 0.63 long, 1.27 wide; sternum 2.53 long, 1.86 wide (0.91 on ventral surface); palp 8.98 long (3.01 + 1.58 + 2.14 + 2.25); leg I 11.82 long (3.56 + 1.94 + 2.73 + 2.25 + 1.34); leg II 12.22 long (3.60 + 1.98 + 2.73 + 2.49 + 1.42); leg III 12.74 long (3.60 + 1.98 + 2.45 + 3.05 + 1.66); leg IV 16.41 long (4.27 + 2.10 + 3.24 + 4.55 + 2.25).

Variation: Carapace lengths in males (n=7) 4.36-4.75, in females with a well-developed vulval plate (n=9) 4.29-5.34, in the smallest female with an egg case 4.66. Carapace widths 3.64-4.06, 3.11-4.22 and 3.51, respectively.

In four males (including the holotype) the scopula extends over the distal half of the ventral side of tarsus III and over the distal three-fifths of tarsus IV, in the other three males it extends over the distal three-fifths of tarsus III and over the distal two-thirds of tarsus IV; on legs I-II the extent of the scopula is the same in all males. Variation in the shape of the tibial apophysis of the male palp is shown in Fig. 8A-C, H-J; variation in the shape of the distal edge of the contrategulum in Fig. 8F-G; variation in the shape of vulval plates of the females examined in Fig. 9. Two female paratypes (Fig. 9C, E), but not the allotype (Fig. 9A), have a moderately pronounced anterolateral lobe followed by a slight anterolateral invagination on each side of the poreplate. The largest (oldest) female (Fig. 9A) has more hairs in the genital atrium than smaller (younger) females (Fig. 9E). The posterior margin of the genital sternite of females is almost straight to distinctly invaginated (Fig. 9A-F).

Distribution: The types were collected from the surroundings of the Kota Tinggi Waterfall, at the foot of Gunung Muntahak, north of Kota Tinggi, in Johor State (Fig. 1, locality 8). Burrows of probably the same species were seen along the trail to the nearby Pelepah Waterfall. Two juveniles (don J. Kral) from Gunung Belumut, about 32 km further northwest, may also belong to this species.

Biology: At the Kota Tinggi Waterfall L. gracilis sp. nov. occurs together with the much larger L. endau which belongs to the same species-group. Burrows of the two species were seen only a few centimetres from each other. Liphistius endau was exclusively found very close to the waterfall and to the stream, L. gracilis sp. nov. also further away, but both were absent from high earthbanks (old road sides) in a palm oil plantation adjacent to the rain forest. Most burrows were simple, equipped with a single trapdoor and dug into the soil; a few small ones were sac-like nests with two trapdoors, constructed on the moss-covered surface of rock (as also known from cave species and from juveniles of forest-dwelling species; see Schwendinger, 1987). The trapdoors of seven penultimate males were 1.1-1.3 cm long and 1.6-1.9 cm wide; that of the largest female 1.3 and 2.0, respectively. All burrows had a maximum of eight, relatively long signal lines running over soil, stones and tree roots; the longest signal line (of the largest female) was 21 cm long.

Four males collected in June became adult in August to November of the same year; two males reached maturity in January and February after over 1.5 years in captivity (thus probably not corresponding to conditions in nature). In early July and early August two females built egg cases in captivity, 1.7-2.0 cm in diameter and 1.1-1.4 cm high, containing 30 and 36 eggs suspended on a thin layer of fine silk strands. Adult females moulted twice per year, in May and again in August to November.

Liphistius johore Platnick & Sedgwick, 1984: 30, figs 79-80 (description of female).

Type material: Bernice Pauahi Bishop Museum, Honolulu, USA; female holotype (not examined); Malaysia, Johore [sic], Sungai Rengit, Pengarang; 19.XII.1961; leg. K.J. Kunchuna.

Material examined: None.

Diagnosis: Seemingly a fairly small species (carapace length of female holotype 5.15, carapace width 4.00). Female with annulated legs and yellow-coloured to orange-coloured carapace and opisthosomal tergites (Platnick & Sedgwick, 1984: 30); vulva with a wide poreplate with a distinct anterolateral invagination on each side of anterior margin, seemingly without bulging lateral and posterolateral margins on ventral side (Platnick & Sedgwick, 1984: fig. 80), with relatively large pores on dorsal side, with a large CDO (at its posterior margin level with the poreplate), and with a large receptacular cluster projecting far beyond anterior margin of poreplate; genital atrium without hairs, including a strongly trapezium-shaped (posteriorly wide, anteriorly very narrow, posterior margin almost straight) posterior stalk (Fig. 10A; Platnick & Sedgwick, 1984: figs 79-80).

Male: Unknown.

Female: This species was described on the basis of only the female holotype. For an easy comparison with other species treated here, the dorsal aspect of the vulval plate was re-drawn from Platnick & Sedgwick (1984: fig. 79) and is shown in Fig. 10A.

Relationships: The vulval plate (with a large CDO, with a huge receptacular cluster projecting far beyond the anterior margin of the well-sclerotised poreplate and with a well-sclerotised posterior stalk fully connected to the poreplate) indicates that L. johore is a member of the malayanus-group. Relatively small size, the presence of anterolateral invaginations on the poreplate (also present, but less distinct, in some females of L. gracilis sp. nov.; Fig. 9C, E) and the apparent absence of bulged lateral and posterolateral margins on the ventral side of the poreplate (not clearly evident from original illustrations of the vulval plate of the holotype, see Platnick & Sedgwick, 1984: fig. 80) indicate that L. johore and L. gracilis sp. nov. are more closely related to each other than to L. malayanus and L. endau.

Distribution and remarks: The type locality is given as “Sungai Rengit, Pengarang” (Fig. 1, locality 9). Kampung Sungai Rengit and the Pengarang ferry terminal and immigration post (with nearby remains of the “Pengarang Battery”, a British defence post overrun by the Japanese Army during the fall of Singapore in World War II) are about 15 km apart. I spent a full day searching in both the fairly undisturbed forest on the publicly accessible little hill next to the Pengarang ferry terminal (the nearby, distinctly higher and nicely forested Bukit Pengarang lies on the land of a large Malaysian naval base and is off limits to civilians and foreigners) and in the strongly disturbed forest of Bukit Pelali, about 5 km north of Kampung Sungai Rengit, but found no traces (not even empty burrows) of Liphistius.

A single female (in MHNG; leg. P.J. Schwendinger), large (carapace length 12.45, carapace width 11.02) and uniformly dark-coloured, was collected close to the Jeram Linang Waterfall (Fig. 1, locality 13), south of Kota Baharu in the Kelantan State, together with the types of L. linang sp. nov. It produced an egg case in late January. Judging from the shape of its vulval plate, this specimen belongs to an undescribed species in the malayanus-group,

Diagnosis: Medium-sized to large (carapace length of males 6.23-11.30, carapace length 6.01-10.04), uniformly dark-coloured spiders. Similar to species of the malayanus-group, distinguished by distal edge of contrategulum proventrally not ending in a V-shaped or U-shaped row of denticles; ventral process of contrategulum well-developed, basally wide (Figs 10B, 11F, 14C-D); para-embolic plate more elevated (Figs 11B-E, 14E-F); poreplate with CDO medium-sized and undivided (Fig. 12A, C, F) or large and divided by a more or less distinctly developed longitudinal bridge (Figs 13A, C-D, G; 14I); receptacular cluster medium-sized, more or less distinctly divided into two halves by a longitudinal trench (Figs 12B, D-E, G; 13B, E-F, H; 14J).

Species included: Liphistius negara sp. nov., L. panching and L. tioman.

Relationships: Species of the tioman-group are similar and probably closely related to species of the malayanus-group. Liphistius panching and L. negara sp. nov. from the mainland are more similar to each other than they are to the very distinct island-dwelling L. tioman.

Distribution: Eastern part of peninsular Malaysia and an offlying island (Fig. 1, localities 10-12).

Liphistius tioman Platnick & Sedgwick, 1984: 28-29, figs 81-87 (description of male and female).

Type material: AMHN; male holotype and female paratype (not examined); Malaysia, Pahang, Tioman Island, Gua Sinah and Gua Panah, 2600 ft. altitude; 29.VII.1982; leg. W.C. Sedgwick.

Material examined: MHNG, sample SIM-01/09; 1 male (matured 3.I.2002), 7 females (moulted 17.XII.2001; 2.IX.2001, 6.III., 4.X.2002, 7.VI.2003; 8.IX.2001, 16.II., 23.VI.2002), 1 penultimate male; Pahang, Tioman Island, Gunung Kajang (2°45′45″N, 104°08′43″E), 820-850 m; 30.VI.2001; leg. P.J. Schwendinger.

Diagnosis: Males distinguished from those of all other Liphistius species by a deeply and widely divided retrolateral apophysis on palpal tibia, both parts being equally long (Fig. 11A, H-I). Males distinguished from those of other species in the tioman-group additionally by a long and pointed para-embolic plate (Fig. 11C-E) and by a very narrow tegulum with few denticles on proximal edge, situated very close to dorsal apex of contrategulum (Fig. 11B, F). Females with poreplate widely fused with posterior stalk; receptacular cluster medially more or less distinctly divided; CDO undivided (Fig. 12).

Additions to description of male: Scopula: In distal two-thirds of ventral side of tarsus I, in distal three-quarters of tarsus II, in distal two-thirds of tarsus III, in distal half of tarsus IV (narrower than on other tarsi). All scopulae thin and only distally (behind the claws) divided by a glabrous longitudinal median stripe.

Palp: Tibial apophysis situated apically, not set back from anterior margin of palp (Fig. 11G), deeply divided: retrolateral part deeper, aligned with axis of tibia and carrying three very short pointed megaspines; retrodorsal part much narrower, pointing away from axis of tibia, carrying a single, slightly longer pointed magaspine (Fig. 11A, G-I). Distal margin of cymbium with very indistinct lobes (Fig. 11C). Paracymbium relatively short and moderately deep (Fig. 11A); cumulus indistinct, carrying a fairly compact group of long stiff bristles (Fig. 11A). Subtegulum without apophysis. Tegulum very narrow, its proximal edge carrying few overlapping scale-like denticles, situated close to dorsal apex of contrategulum (Fig. 11B, F). Contrategulum with large, conical ventral process being slightly constricted at base (Fig. 11F); distal edge of contrategulum distinctly elevated and composed of numerous parallel ribs instead of teeth (Fig. 11B-F), dorsal apex relatively small and narrowly rounded, only slighly elevated above underlying surface (Fig. 11B, F). Para-embolic plate long, narrow and asymmetrically triangular (Fig. 11B-E); embolus proper strongly inclined prolaterad (Fig. 11B), distally widened; dorsal wall of sclerotised part distinctly wider than ventral wall, ending in a quadrangular lobe (Fig. 11B-C, F), retrolateral wall not enforced by ridges but only with an indistinct, short distal keel (Fig. 11B, F); membranous part of embolus proper slightly widened distally (Fig. 11C).

Additions to description of female: Genital atrium of vulval plate (Fig. 12) without median hairs, few to several lateral hairs present in some females; lateral margin of genital atrium not forming flaps bending ventrad and inwards (as usual in Liphistius) but only indistinct mounds on the dorsal (not the ventral) side (Fig. 12A, C, F). Posterior stalk relatively short and wide, fused with poreplate without recognizable transition, reaching lateral margins of vulval plate in some females. Poreplate of variable shape, in most females examined anteriorly slightly narrower than posteriorly, its anterior margin indistinctly invaginated, without lobes; CDO quite large, undivided, anteriorly deep, posteriorly level with rest of poreplate (Fig. 12A, C, F; but see illustration of paratype in Platnick & Sedgwick, 1984: fig. 86); receptacular cluster quite large and wide, more or less distinctly divided into two paramedian subclusters (Fig. 12B, D-E, G).

Taxonomic remarks: The tibial apophysis of the male palp of L. tioman is unique and distinctly autapomorhic. A similar divided tibial apophysis is found in males of the linang-group, but that is considered to have evolved convergently (see also Discussion). The wide ventral process of the contrategulum, the wide posterior stalk and the divided receptacular cluster indicate a close relationship with L. panching and L. negara sp. nov.

Variation: Carapace lengths in males (n=2; values of the holotype are taken from Platnick & Sedgwick, 1984: 29) 6.23-6.93, carapace widths 6.01-6.34. In females with fully developed copulatory organs (n=7) the carapace length is 5.54-7.92 and the carapace width 5.15-7.38. Variation in the shape of poreplates of the females examined is shown in Fig. 12. The retrolateral tibial apophyses of the left and right palp of the examined male are shown in Fig. 11A and Fig. 11I, respectively.

Distribution: Known only from higher altitudes on Tioman Island (Fig. 1, locality 10).

Biology: Spiders of this species were found under and between huge granite boulders (some of them touching each other and forming recesses) in a lush rain forest. These recesses probably correspond to the Gua Sinah or Gua Panah which is the exact type locality and which were described as not being real caves (like their Malay names suggest) but just overhung caverns (Platnick & Sedgwick, 1984: 29). Burrows were mostly dug into loamy soil at the base and between the bolders (a few small spiders had silken nests on the surface of bolders just above the ground, as described for juvenile spiders of L. trang, see Schwendinger, 1987: fig. 5), and they had up to eight exceptionally long (up to 34 cm) signal lines spread over soil or rock surface. The trapdoor of the adult male (before its final moult) was 1.8 cm long and 3.2 cm wide, that of the largest female was 1.9 cm and 3.2 cm, respectively. When collected at the end of June, four females had egg cases: one was not opened, another was empty, the remaining two were 2.2-2.7 cm long, 2.8-2.9 cm wide and 1.9 cm high; they contained 3rd instar spiderlings (not collected, not counted).

Liphistius panching Platnick & Sedgwick, 1984: 27-28, figs 77-78 (description of female). – Sedgwick & Platnick (1986: 205-206, figs 1-8; description of male).

Type: AMHN; female holotype (not examined); Malaysia, Pahang, Gua Panching [= Gua Charas; see paragraph “Distribution”], 24 km N of Kuantan; 31.VII.1982; leg. W.C. Sedgwick.

Material examined: MHNG, sample MAL-04/07; 3 females (moulted 30.X.2004, 26.III., 7.IX.2005, 19.II.2006; 8.XII.2004, 30.XII.2005); Pahang, about 1 km north of Kampung Panching, Gua Charas, 3°54′41″N, 103°08′50″E, 120 m; 1.-2.VI.2004; leg. P.J. Schwendinger. – MHNG, sample SIM-01/11; 6 females (moulted 7.VII.2001, XII.2002; 8.VII.2001; 27.II., 16.X.2002), 4 juveniles; same locality; 7./8. VII.2001; leg. P.J. Schwendinger.

Diagnosis: Male (see Sedgwick & Platnick, 1986: figs 1-8 and Fig. 10B) distinguished by retrolateral apophysis of palpal tibia in ventral view very wide at base, only little set back from anterior margin of article (Sedgwick & Platnick, 1986: fig. 3); distal margin of cymbium with indistinct lobes (Sedgwick & Platnick, 1986: fig. 4); paracymbium quite shallow, without retrolateral-proximal heel; cumulus indistinct, with moderately long bristles (Sedgwick & Platnick, 1986: figs 1-5); tegulum with many teeth on proximal edge (Sedgwick & Platnick, 1986: figs 1-2); contrategulum with short, very widely arched ventral process pointing proventrad and with pointed dorsal apex (Fig. 10B; Sedgwick & Platnick, 1986: fig. 5); para-embolic plate short (Sedgwick & Platnick, 1986: figs 1-4); embolus proper apically wide (Sedgwick & Platnick, 1986: fig. 3; Fig. 10B), dorsal wall of sclerotized part apparently long and sharply bent prodorsad (Sedgwick & Platnick, 1986: fig. 5). Females (see Platnick & Sedgwick, 1984: figs 77-78 and Fig. 13) distinguished by vulva with few to many lateral and median hairs in genital atrium; posterior stalk wide, quadrangular to trapezium-shaped; poreplate anteriorly wider than posteriorly, with a more or less distinctly invaginated anterior margin and with bulging lateral and posterolateral margins on ventral side; two central dorsal openings (CDO) separated by a wide longitudinal bridge (Fig. 13A, C-D, G); two longitudinal receptacular clusters more or less completely separated from each other by a longitudinal trench (Fig. 13B, E-F, H).

Remarks: According to information from Lorenzo Prendini, the current curator of the arachnid collections at the AMNH, the normally developed male of L. panching (deposited together with a malformed male) cannot be found at the moment. To my knowledge, this is the only useful male specimen of L. panching in any public scientific collection, and a re-examination of details of the palpal organ of this species was therefore not possible. For an easy comparison with other species, the distal aspect of the palpal organ was re-drawn from Sedgwick & Platnick (1986: fig. 5) and is shown in Fig. 10B.

Unlike in any other Liphistius species examined, the vulval plates on the exuviae of L. panching females (much more than other sclerotised parts of the exuviae) have become partially depigmented; some have almost completely lost their pigmentation. This is probably due to alcohol preservation. In undissected females the vulval plates have retained their original pigmentation.

Variation: Sedgwick & Platnick (1986: 205) give the carapace length 9.8 and the carapace width 8.4 for the normally developed male, but no measurements for the teratological male with an incompletely developed palp. Carapace lengths in females with fully developed copulatory organs (n=8) 7.62-8.71, carapace widths 6.63-8.12.

Variation in the shape of the vulval plates examined is shown in Fig. 13. In one female the two receptacular clusters and the two central dorsal openings (CDO) are anteriorly connected with each other (Fig. 13G-H). The larger (older) a female is, the more hair it has in its genital atrium (Fig. 13D cf. Fig. 13A).

Distribution: The type locality is given as “Gua Panching [= Panching Cave], a cave 24 km north of Kuantan”. This is identical with Gua Charas (Fig. 1, locality 11) in Bukit Charas (= Charas Hill) near Kampung Panching (= Panching Village), which is actually a bit over 20 km northwest of Kuantan. This should not be confused with Bukit Panching, which did not have any caves, was situated about 1.5 km southwest of Bukit Charas and has been completely removed by quarrying in the 1990s. Today only a water-filled depression shows where the hill once stood (Liz Price, personal communication).

Biology: All spiders examined were collected from sloping loamy soil in the oligophic zone at the entrance area of the Charas Cave. In captivity one female built an egg case (2.7 cm long, 2.8 m wide and 1.4 cm high) in late July 2001. The eggs (more than 21) were found partially rotten when the egg case was opened.

Types: MHNG, sample MAL-04/14; male holotype (matured 9.VII.2006), 1 male paratype (matured end of July 2005), female allotype, 2 juv. males; Malaysia, Pahang, Taman Negara, trail from Nusa Camp to Abai Waterfall, 4°23′49″N, 102°25′50″E, 110 m; 16.-17. VI.2004; leg. P.J. Schwendinger.

Etymology: The species epithet, a name in apposition, is taken from the Malay name of the type locality: taman (= park), negara (= national).

Diagnosis: Large, uniformly dark-coloured species in both sexes. Similar to L. panching, distinguished by slightly larger size (carapace length of males 10.28-11.30, width 8.70-10.04 versus 9.8 and 8.4, respectively, in male of L. panching, see Sedgwick & Platnick (1986: 205). Males with tibial apophysis in ventral view narrower at base, more distinctly set back from anterior margin of tibia (Fig. 14A cf. Sedgwick & Platnick, 1986: fig. 3); paracymbium deeper (Fig. 14B cf. Sedgwick & Platnick, 1986: fig. 2); ventral process of contrategulum narrower and more pointed (Fig. 14C-D cf. Fig. 10B and Sedgwick & Platnick, 1986: fig. 5); distal edge of contrategulum with a large triangular tooth, dorsal apex rounded instead of pointed (Fig. 14C-D cf. Fig. 10B and Sedgwick & Platnick, 1986: fig. 5). Female distinguished from those of L. panching by vulval plate wider than long; poreplate anteriorly distinctly narrower than posteriorly instead of anteriorly wider than posteriorly (Fig. 14I-J cf. Fig. 13 and Platnick & Sedgwick, 1984: figs 77-78).

Description of male (holotype): Colour in alcohol (much darker in life): Most sclerotised parts uniformly chestnut-brown, on ventral side slightly lighter than on dorsal side; carapace reddish chestnut-brown; proximal portion of chelicerae cream-coloured; membranes of opisthosoma greyish cream, other membranes white. Bristles on carapace: Stiff bristles along all margins (longest anteriorly); many weaker and shorter ones on coxal elevations and behind fovea, fewer of them on both sides of eye mound and in a W-shaped arrangement behind it.

Cheliceral teeth: Eleven and twelve small ones on promargin of cheliceral groove on right and left chelicera, respectively.

Scopula: Tarsus I with thin scopula confined to distal third of ventral side, divided for its entire length by narrow pale, glabrous longitudinal median stripe and by some bristles; tarsus II with thin scopula in distal half, only distally divided by median stripe; tarsi III-IV with dense scopula covering distal four-fifths, only distally divided by median stripe.

Claws: Paired tarsal claws on anterior legs with 4-5 denticles, on posterior legs with 4-6 denticles; unpaired claws in most cases carrying a single denticle.

Palp: Tibial apophysis slightly set back from distal margin of tibia (Fig. 14A), carrying four medium-sized pointed megaspines (Fig. 14B, G). Both apical lobes of cymbium indistinct and equally rounded (Fig. 14E, H). Paracymbium basally deep (Fig. 14B), its cumulus indistinctly elevated, carrying moderately long stiff bristles (Fig. 14A-B). Subtegulum without apophysis. Tegulum short and wide, coarsely dentate along entire proximal margin (Fig. 14F). Contrategulum with distinct conical ventral process, its rounded apex directed proventrad (Fig. 14C); distal edge with several oblique ribs and with a large triangular tooth prodorsally, ending in spatulate asymmetrical dorsal apex (Fig. 14C). Para-embolic plate short (Fig. 14A, E-F); retrolateral side of sclerotised part of embolus proper without longitudinal keels or ribs and not carrying denticles, dorsal wall much wider than ventral wall and sharply bent prodorsad, thus opening of spermophor forming an L (Fig. 14A, C, E); membranous part of embolus proper distally wider than proximally, with only slightly pigmented proximal portion (Fig. 14E).

Measurements: Total length 27.94; carapace 11.30 long, 10.04 wide; opisthosoma 11.55 long, 9.04 wide; eye mound 1.51 long, 1.61 wide, AME well-developed; palpal coxae 3.68 long, 2.26 wide; labium 1.00 long, 2.01 wide; sternum 5.35 long, 3.01 wide (1.34 on ventral surface); palp 18.76 long (5.69 + 3.18 + 6.44 + 3.45); leg I 32.80 long (9.20 + 4.69 + 7.20 + 8.28 + 3.43); leg II 35.06 long (9.45 + 4.69 + 7.70 + 9.54 + 3.68); leg III 38.66 long (9.45 + 4.69 + 8.20 + 12.05 + 4.27); leg IV 47.53 long (11.55 + 4.85 + 9.79 + 15.48 + 5.86).

Description of female (allotype): Colour in alcohol (much darker in life) as in male. Carapace with stiff bristles as in male, plus a pair of bristles anterior of fovea. Eleven strong teeth on promargin of cheliceral groove on both sides. Paired tarsal claws on anterior legs with three denticles, on posterior claws with 3-4 denticles; unpaired claws in most cases with a single denticle (claws worn on all limbs). Tarsi without scopula.

Vulva (Fig. 14I-J): Posterior margin of genital sternite slightly and widely invaginated. Vulval plate distinctly sclerotised and pigmented, wider than long. Genital atrium with several lateral and median hairs. Posterior stalk wide, with micropores, fused with poreplate. Poreplate entirely and distinctly pigmented, its anterior margin indistinctly invaginated and clearly narrower than posterior margin. Two central dorsal openings (CDO) separated by a longitudinal bridge; two longitudinal receptacular clusters on ventral side completely separated from each other.

Measurements: Total length 22.00; carapace 9.62 long, 8.28 wide; opisthosoma 10.29 long, 8.62 wide; eye mound 1.22 long, 1.42 wide; palpal coxae 3.18 long, 2.01 wide; labium 1.17 long, 2.18 wide; sternum 4.35 long, 2.59 wide (1.59 on ventral surface); palp 16.23 long (5.35 + 3.01 + 4.02 + 3.85); leg I 19.76 long (5.86 + 3.60 + 4.27 + 4.02 + 2.01); leg II 20.57 long (6.02 + 3.68 + 4.35 + 4.43 + 2.09); leg III 22.25 long (6.19 + 3.68 + 4.52 + 5.35 + 2.51); leg IV 30.37 long (8.03 + 3.85 + 6.11 + 8.70 + 3.68).

Variation: Carapace lengths in males (n=2) 10.28-11.30, widths 8.70-10.04. The male paratype has the same reduced scopula on anterior legs (especially on tarsus I) as the holotype. Variation in the shape of the palpal organ, see Fig. 14C-D.

Distribution: Known only from the type locality in central peninsular Malaysia (Fig. 1, locality 12).

Biology: The types were collected from sloping soil and from decaying wood in a rain forest. The largest burrow (of juvenile male, not yet penultimate) had a 2.6 cm long ad 4.1 cm wide trapdoor. The trapdoors of old females thus must be of similar size as those of L. malayanus and L. endau. The female allotype of L. negara sp. nov. appears to be mature (judging from the degree of sclerotisation of its vulval plate), but not yet fully grown. Most burrow entrances had eight signal lines, one had nine. Both males became mature in July, after one and two years of captivity, respectively.

The male paratype has a pronounced pathological outgrowth on the right cheliceral fang, and bite marks of parasitic Ljungia bristowi mites on its carapace (many) and opisthosomal tergites (few), but not elsewhere.

Diagnosis: Small to medium-sized spiders (carapace length of males 4.50-6.83, carapace width 3.98-6.21) with a colour pattern on carapace, limbs and opisthosomal tergites (Fig. 2B). Distinguished from the malayanusgroup and the tioman-group by males possessing a large para-embolic plate with a coarsely serrate distal margin (Figs 15A-D, 17A-D), a narrow and pointed dorsal apex of the contrategulum (Figs 15E-F, 17G-H), and a retrolateral tibial apophysis deeply divided into a long, deep retroventral part and a very short, small retrodorsal part (Figs 15G-I, 17I-K) (tibial apophysis also deeply divided in L. tioman but in that species both parts equally long; Fig. 11A, G-I); three megaspines on retroventral part short and bent, single megaspine on retrodorsal part short, curved and weaker than other megaspines (Figs 15A, G-I; 17A, I-K). Females distinguished by a peculiar vulval plate with an only partially sclerotised poreplate widely separated from small remnant of posterior stalk situated at posterior margin of vulva (Figs 16A-E, 18); CDO small, in many cases slit-like (Figs 16A, C-D; 18A, C, E, G).

Species included: Liphistius linang sp. nov., L. indra sp. nov.

Relationships: The linang-group is morphologically intermediate between the tioman-group and the batuensis-group. Sharing with L. tioman a deeply divided tibial apophysis with the retrodorsal-apical megaspine distinctly separated from the other three megaspines, and sharing with this and other species in the tioman-group a distinctly elevated para-embolic plate. Females of the linang-group show strong resemblance with those of the batuensis-group by possessing an only partially sclerotised poreplate. However, females of L. linang sp. nov. have a divided receptacular cluster, as do females of all tioman-group species. The relationships of the linang-group are therefore not clear.

Distribution: Northeastern peninsular Malaysia and southern Thailand (Fig. 1, localities 13-14).

Types: MHNG; male holotype (matured end XII.2001), 4 male paratypes (matured 14.IX., 1.X.1999, early XI.2000, end XII.2001) and 4 female paratypes (including allotype, moulted 12.II.2002); Malaysia, Kelantan, about 15 km E of Machang, Jeram Linang Waterfall, 5°44′34″N, 102°22′29″E, 50 m; 12.I.1999; leg. P.J. Schwendinger.

Etymology: Name in apposition, taken from the Malay name of the type locality: jeram (= rapids), linang (tearful, weeping).

Diagnosis: Medium-sized, bicoloured species in both sexes. Similar to L. tioman, males distinguished by retroventral part of bipartite tibial apophysis carrying longer and bent megaspines; retrodorsal part of tibial apophysis developed as a small tubercle carrying a weaker megaspine (Fig. 15A, G-I cf. Fig. 11A, G-I); tegulum with wide, distinctly dentate proximal edge (Fig. 15D cf. Fig. 11B); contrategulum without (or with very indistinct) ventral process, its distal edge finely dentate instead of ribbed, prodorsally with a wide gap before reaching hook-shaped dorsal apex (Fig. 15B, E-F cf. Fig. 11C, F); para-embolic plate large, distally wide, with coarsely dentate margins (Fig. 15A-F cf. Fig. 11C-E); dorsal wall of sclerotised part of embolus proper not ending in protruding lobe, as wide as ventral wall (Fig. 15B, E-F cf. Fig. 11B-C, F); scopula on tarsi IV absent.

Description of male (holotype): Colour in alcohol (much darker in life): Carapace mostly light brown; pars cephalica brown except for indistinct grey-brown W-shaped pattern behind eye mound and light brown medial patch between eye mound and fovea; brown area on pars cephalica connected to brown flower-shaped area around fovea; long brown patches along lateral (wide) and posterior (very narrow) margins of pars thoracica and between coxal elevations. Chelicerae with proximal portion cream-coloured, distal portion dark brown. Palps in proximal half of femur and tibia and in distal half of patella light brown, mottled with dark spots, other parts dark brown; cymbium entirely very dark reddish brown. Legs mostly dark brown except for light brown distal half of all patellae and light brown proximal half of all femora (mottled with dark spots); leg tibiae (in contrast to those of palp) to tarsi entirely dark brown. Opisthosoma mostly light brown, mottled with dark spots laterally; tergites I-II almost entirely dark brown, tergite III light brown with extensive dark lateral and median patches, following tergites with distinct dark lateral spots and increasingly indistinct median spots, last two tergites tiny and entirely dark brown.

Bristles on carapace: Few short, weak bristles along all margins except posterior one, on coxal elevations, in front and behind fovea; longer, stronger bristles on, behind and in front of eye mound.

Cheliceral teeth: Ten small ones on promargin of cheliceral groove of each chelicera.

Scopula: Tarsi I-III with moderately dense scopula in distal two-thirds of ventral side, only distally divided by a short median stripe; tarsus IV entirely without scopula. Claws: Paired tarsal claws on anterior legs with 3-5 denticles, on posterior legs with 4-5 denticles; unpaired claws with one indistinct denticle or without.

Palp: Tibial apophysis situated distally, only slightly set back from anterior margin of palp (Fig. 15A), deeply divided: (a) retroventral part long and deep, pointing slightly away from axis of tibia, carrying three medium-sized, bent megaspines; (b) retrodorsal part much smaller and shorter, shaped like a low tubercle, carrying a single bent, relatively weak spine (i.e. a reduced megaspine); long bristle retroventrally below tibial apophysis weak (Fig. 15A, G-H). Distal margin of cymbium with indistinct lobes (Fig. 15B). Paracymbium quite long and moderately deep, with an indistinct retrolateral-proximal heel (Fig. 15A, G); cumulus indistinct, carrying a moderately compact group of long stiff bristles (Fig. 15A, G). Subtegulum without apophysis. Tegulum with wide, distinctly dentate proximal edge (Fig. 15D). Contrategulum without recognizable ventral process; prolateral surface with a few ribs, distal edge sharp in proventral part, finely dentate in prolateral part, with a deep gap in prodorsal part before reaching very narrow, slightly hook-shaped dorsal apex (Fig. 15B, E). Para-embolic plate large, its distal margin wide, strongly dentate and directed proventrad-distad (Fig. 15B-E); embolus proper relatively narrow, inclined prolaterad (Fig. 15A-D), dorsal and ventral walls of sclerotised part equally wide and lying close to each other, retrolateral wall enforced by five weak ridges (Fig. 15E); membranous part of embolus proper narrow, indistinct (Fig. 15B).

Measurements: Total length 14.22; carapace 5.83 long, 5.05 wide; opisthosoma 6.86 long, 4.31 wide; eye mound 0.91 long, 1.12 wide; palpal coxae 1.76 long, 1.23 wide; labium 0.49 long, 1.08 wide; sternum 2.55 long, 1.81 wide (1.03 on ventral surface); palp 9.66 long (2.55 + 1.91 + 3.53 + 1.67); leg I 17.64 long (4.80 + 2.30 + 3.82 + 4.61 + 2.11); leg II 18.67 long (4.80 + 2.30 + 4.02 + 5.20 + 2.35); leg III 20.48 long (4.95 + 2.35 + 4.26 + 6.27 + 2.65); leg IV 26.72 long (6.27 + 2.65 + 5.74 + 8.58 + 3.48).

Description of female (allotype): Colour in alcohol (much darker in life): As in male, except for carapace with more extensive dark areas, all of them interconnected; W-shaped pattern behind eye mound distinct; metatarsi of legs and palps mostly dark, with a small light zone at base; tibiae with dark proximal and subdistal annulations; light brown femora with indistinct (in comparison to tibiae) dark proximal and subdistal annulations; opisthosomal tergites III-VII with more extensive dark median patches; genital area darker than surrounding parts of genital sternite, with a white, bell-shaped posterior zone (Fig. 16F).

Bristles on carapace: Stiff bristles as in male, plus an additional pair just anterior to fovea.

Cheliceral teeth: Nine strong teeth on promargin of left cheliceral groove, ten on right side.

Claws: Palpal claw with three denticles on right side, none on left side. Paired claws with two denticles on anterior legs, 2-3 denticles on posterior legs; unpaired claws with 1-2 denticles on legs I-III, none on leg IV. All tarsi without scopula.

Vulva: Posterior margin of genital sterite invaginated (Fig. 16F). Vulval plate (Fig. 16A-E, illustrations of paratypes) with pigmentation and sclerotisation completely lost in median zone. Posterior stalk reduced to a small isolated, ventrad-bent posterior sclerite at some distance from or on posterior margin of genital sternite; posterior stalk and poreplate widely separated from each other. Genital atrium slightly sunken below level of poreplate, carrying numerous hairs on both sides of remnant of posterior stalk. Poreplate with indistinctly outlined lateral and posterior margins, these not bulging from ventral side of plate; anterior margin distinctly and widely invaginated, forming two more or less pronounced anterolateral lobes. CDO relatively small, wider than long, transversally slit-like or triangular, situated in unpigmented or weakly pigmented area of poreplate (Fig. 16A, C-D, paratypes). Ventral receptacular cluster quite large and racemose, longitudinally divided into two lateral subclusters, its individual vesicles globular, about as long as wide (Fig. 16B, E, paratypes).

Measurements: Total length 17.25; carapace 5.93 long, 5.00 wide; opisthosoma 7.89 long, 6.13 wide; eye mound 0.82 long, 1.04 wide; palpal coxae 1.91 long, 1.37 wide; labium 0.64 long, 1.37 wide; sternum 2.65 long, 2.11 wide (1.27 on ventral surface); palp 10.00 long (3.24 + 1.86 + 2.30 + 2.60); leg I 12.40 long (3.77 + 2.16 + 2.50 + 2.50 + 1.47); leg II 12.90 long (3.82 + 2.21 + 2.55 + 2.75 + 1.57); leg III 13.73 long (3.92 + 2.25 + 2.60 + 3.24 + 1.72); leg IV 19.71 long (5.10 + 2.55 + 4.07 + 5.54 + 2.45).

Remark: In the female allotype the long spinnerets (anterior and posterior laterals) are bent anteriad, which is an artefact resulting from having been pressed against the wall of the vial in that position for some time.

Variation: Carapace lengths in males (n=5) 4.91-6.15, carapace widths 4.19-5.09; in females with well-developed vulval plates (n=4) 5.53-5.90 and 4.29-5.03, respectively. All males lack a scopula on tarsus IV. Variation in the shape of the retrolateral tibial apophysis of the male palp is shown in Fig. 15G-I; variation in the shape of the prolateral-distal edge of the contrategulum in Fig. 15E-F; variation in the shape of the vulval plate in Fig. 16A-E.

Distribution: Known only from the type locality (Fig. 1, locality 13) in the northeastern corner of peninsular Malaysia. At the same locality, only metres away from burrows of L. linang sp. nov., a single female of another, much larger, dark-coloured Liphistius species was found. This female (mentioned under Liphistius sp. at the end of the presentation of the malayanus-group) appears to be more closely related to L. endau than to the geographically closer L. negara sp. nov.

Biology: Spiders of this species were quite abundant in a lush rain forest along the stream above the Jeram Linang Waterfall. The burrows had a single trapdoor and mostly nine signal lines (up to 4 cm long) spread over soil surface. Trapdoors of penultimate males (n=5) were 1.2-2.0 cm long and 1.8-2.3 cm wide, those of reproductive females (with egg case; n=5) 1.2-1.7 cm and 1.8-2.3 cm, respectively.

Two males matured in mid-September and at the beginning of October of the year when they were collected. The three other males were raised from eggs (hatched in late May); one of them matured in November of the following year, the other two one year later in December. Less than one and a half years from hatching to maturation is exceptionally short for a Liphistius male, and this may be due to conditions in captivity in Geneva. Three females built egg cases in captivity between early February and early March, two of them were 1.7-1.9 in diameter, 1.25-1.3 cm high, and contained 40 and 84 light beige-yellow eggs suspended on a thin mesh of fine silk threads. The first generation females in captivity moulted in April, July and September.

Types: MHNG; male holotype (matured 27.IX.2011), 8 male (matured 24.VIII., 7.IX., 10.IX., 17.IX., 25.IX., 26.X.2011, 30.IX., 4.X.2012) and 13 female paratypes (including allotype which did not moult); Thailand, Pattani Province, about 20 km NW of Yala, Sankalakhierie Mountains, 6°39′09″N, 101°05′55″E, 200 m; 12.VII.2011; leg. P.J. Schwendinger. – MHNG, SMF; 1 male (matured 17.XI.2000) and 5 female paratypes; same locality, 260 m; 22.X.1999; leg. P.J. Schwendinger.

Etymology: The species epithet refers to one of the two old names of the type locality: Indragiri (= Indra's mountain) and Bukit Besar (= big mountain) (Skeat, 1953: 21). Indra (“Phra In” in Thai mythology) is the king of the Vedic gods. Name in apposition.

Diagnosis: Medium-sized, light brown-coloured species in both sexes. Very similar to L. linang sp. nov., distinguished by males with a ventral scopula on tarsus IV; long bristle retroventrally below tibial apophysis much stronger, resembling a megaspine (Fig. 17A, I-K cf. Fig. 15A, G-I); paracymbium distinctly shorter, more globular (Fig. 17A, I cf. Fig. 15A, G); distal edge of contrategulum without dentate gap, ending in a beak-like rather than a hook-like dorsal apex (Fig. 17B, G-H cf. Fig. 15B, E-F). Females distinguished from those of L. linang sp. nov. by anterior margin of poreplate less deeply and less widely invaginated; very weak pigmentation (in contrast to none) in anterior part of genital atrium (corresponding to reduced anterior part of posterior stalk); CDO semicircular or longitudinally slit-like instead of transversally slit-like, flanked by an enlarged pore on each side (Fig. 18A, C, E, G cf. Fig. 16A, C-D); receptacular cluster larger, not divided into two lateral subclusters, its individual vesicles (especially posterior ones) longer (Fig. 18B, D, F, H cf. Fig. 16B, E).

Description of male (holotype): Colour in alcohol (much darker in life; see Fig. 2B for paratype): Carapace with wide grey-brown margin (broken posteromedially) including most of pars cephalica and connected to flower-shaped patch of same colour around fovea; three very light brown areas between eye mound and fovea (anterior two small, kidney-shaped and in a pair, posterior one larger, inverted lanceolate) and light patches between darker central and marginal areas. Chelicerae with proximal portion cream-coloured, distal portion grey-brown. Palps in proximal half of femur and tibia and in distal half of patella light brown, mottled with dark spots, other parts grey-brown; cymbium entirely reddish brown. Legs mostly grey-brown except for light brown distal half of all patellae and light brown proximal half of all femora (mottled with dark spots); leg tibiae (in contrast to those of palp) to tarsi entirely dark brown. Membranous parts of opisthosoma light brown; tergite I almost entirely grey-brown, tergite II with extensive dark lateral and median patches, following tergites with distinct dark lateral spots and increasingly indistinct median ones, last two tergites tiny and mostly grey-brown.

Bristles on carapace: Few short, weak bristles along most of its margins (completely absent posteriorly and largely absent laterally), on coxal elevations and in front of fovea (not behind); longer, stronger bristles on, behind and in front of eye mound.

Cheliceral teeth: Eleven small ones on promargin of cheliceral groove of each chelicera.

Scopula: Tarsi I-III with moderately dense scopula in distal half of ventral side, only distally divided by a short median stripe; tarsus IV with weak scopula in distal quarter, medially divided for its entire length.

Claws: Paired tarsal claws with 3-4 denticles on anterior legs, 2-4 denticles on posterior legs; unpaired claws with 1-2 indistinct denticles on tarsi I-III, none on tarsus IV. Palp: Tibial apophysis situated distally, only slightly set back from anterior margin of tibia (Fig. 17A), deeply divided: (a) long and deep retrolateral part pointing slightly away from axis of tibia, carrying three medium-long and bent megaspines; (b) short and small retrodorsal part carrying a single, slightly curved megaspine (Fig. 17I); long bristle retroventrally below tibial apophysis very strong, almost as thick as megaspines (Fig. 17A, I). Distal margin of cymbium with short lobes (Fig. 17B, showing paratype matured 4.X.2012). Paracymbium globular, quite short and shallow, without retrolateral heel (Fig. 17A, I); cumulus indistinct, carrying a group of long stiff bristles (Fig. 17A, I). Subtegulum without apophysis. Tegulum with moderately wide, strongly dentate proximal edge (Fig. 17C). Contrategulum without recognizable ventral process; prolateral surface with a few ribs; distal edge sharp throughout, with an indistinct, very short and wide invagination (not a gap) prodorsally (Fig. 17B, showing paratype matured 4.X.2012) before reaching very narrow, beak-shaped dorsal apex (Fig. 17G). Para-embolic plate large, its distal margin wide, strongly dentate (Fig. 17A, C, G); embolus proper slightly inclined prolaterad, relatively narrow (Fig. 17A, C-D), dorsal and ventral walls of sclerotised part equally wide and lying close to each other, retrolateral wall with few indistinct longitudinal ridges (Fig. 17G); membranous part of embolus proper narrow, indistinct (Fig. 17B, showing paratype matured 4.X.2012).

Measurements: Total length 11.57; carapace 5.08 long, 4.49 wide; opisthosoma 4.25 long, 2.91 wide; eye mound 0.80 long, 0.97 wide; palpal coxae 1.61 long, 1.10 wide; labium 0.43 long, 0.98 wide; sternum 2.24 long, 1.57 wide (0.87 on ventral surface); palp 9.10 long (2.72 + 1.65 + 3.31 + 1.42); leg I 15.36 long (4.13 + 1.97 + 3.31 + 3.98 + 1.97); leg II 15.83 long (4.17 + 2.01 + 3.31 + 4.33 + 2.01); leg III 17.32 long (4.41+ 2.05 + 3.46 + 5.04 + 2.36); leg IV 22.91 long (5.67 + 2.20 + 4.88 + 7.01 + 3.15).

Description of female (allotype): Colour in alcohol (much darker in life): Generally darker reddish-brown than male; light areas on carapace mottled with dark brown spots, flower-shaped area around fovea less clearly outlined; tarsi and metatarsi of legs and palps mostly dark, with a small light zone at base of palpal tarsi and leg metatarsi; all tibiae with dark proximal and subdistal annulations; femora with indistinct (in comparison to tibiae) and broken proximal and subdistal annulations; opisthosomal tergites II-V with larger dark median patches; genital area darker than surrounding parts of genital sternite, with a light posterior margin.

Bristles on carapace: As in male, plus several tiny bristles behind fovea.

Cheliceral teeth: Eleven mostly strong cheliceral teeth on promargin of left cheliceral groove, nine on right.

Claws: Palpal claws with two denticles. Paired claws with 2-3 denticles on anterior legs, 2-4 denticles on posterior legs; unpaired claws with two denticles on tarsi I-III, none on tarsus IV. All tarsi without scopula.

Vulva: Posterior margin of genital sternite slightly and widely invaginated. Vulval plate (Fig. 18, illustrations of paratypes) with pigmentation much reduced but still discernible in median zone. Posterior stalk reduced to a very weakly pigmented, wide to narrow anterior part and a small, strongly pigmented, ventrad-bent posterior sclerite at some distance from posterior margin of genital sternite. Genital atrium with numerous hairs on both sides of remnant of posterior stalk. Lateral and posterior margins of poreplate indistinctly outlined, not bulging from ventral side; anterior margin only slightly invaginated at midpoint, not forming anterolateral lobes. CDO relatively small, longitudinally slit-like or semicircular, lying in weakly pigmented area of poreplate (Fig. 18A, C, E, G, showing paratypes). Ventral receptacular cluster quite large and racemose, not divided into two lateral subclusters, its individual vesicles mostly digitiform, longer than wide (Fig. 18B, D, F, H, showing paratypes).

Measurements: Total length 19.45; carapace 6.61 long, 5.67 wide; opisthosoma 8.27 long, 6.69 wide; eye mound 0.95 long, 1.01 wide; palpal coxae 2.20 long, 1.54 wide; labium 0.75 long, 1.73 wide; sternum 3.15 long, 2.20 wide (1.26 on ventral surface); palp 11.61 long (3.78 + 2.20 + 2.83 + 2.80); leg I 14.25 long (4.49 + 2.48 + 2.95 + 2.87 + 1.46); leg II 14.68 long (4.49 + 2.56 + 2.95 + 3.11 + 1.57); leg III 15.43 long (4.57 + 2.56 + 2.99 + 3.58 + 1.73); leg IV 22.44 long (6.30 + 2.91 + 4.57 + 6.14 + 2.52).

Variation: Carapace lengths in males (n=10) 4.50-6.83, carapace widths 3.98-6.21; in females with well-developed vulval plates (n=18) 5.55-7.24 and 4.80-6.30, respectively. All ten males possess a weak ventral scopula on tarsus IV; it is medially divided in three of them (including the holotype), undivided in the others. Three males have more or less distinct remnants of “tibial spurs” (sensu Platnick & Goloboff, 1985) on legs I-III. A male lacks one of its AME; in all other specimens examined both AME are well-developed. Variation in details of the male palp, see Fig. 17G-H, I-K; variation in the morphology of the vulval plate, see Fig. 18. One of the larger females examined has a small knob-like sclerite in the membranous dorsal wall of its vulva which fits into the CDO of the ventral wall (= vulval plate). Such a structure was also observed in some (but only few) vulvae of large females of other species. This is an unlikely (because inefficient) plugging device and more probably a by-product of increased sclerotisation of the vulval plate in old females.

Distribution: Known only from the type locality (Fig. 1, locality 14) in the deep south of Thailand. This area lies in the former Sultanate of Pattani which was an independent Malay kingdom until 1785, then became a tributary of the kingdom of Siam, and in 1909 formally a Siamese province. A second species, Liphistius cf. thaleban Schwendinger, 1990, occurs at the same locality. That species is much larger, has orange-coloured femora in larger juveniles and adult females, and belongs to the trang-group.

Biology: The spiders examined were collected in an evergreen rain forest, close to a stream. Their burrows had a single trapdoor, usually opening downwards, and 8-10 (mostly nine) relatively short signal lines (up to 4 cm long) spread over soil surface. Trapdoors of penultimate males (n=8) were 1.25-1.8 cm long and 1.65-2.3 cm wide, those of females (n=17) up to 2.1 cm long and 3.0 cm wide.

Males matured in captivity between August and November, most (six out of ten) in September. Adult females in captivity usually moulted twice per year, in January to April and in June to November. As all females were collected in July and October (outside the breeding season), no egg cases were observed.

Upon being provoked, one mature male and one large female displayed “tiptoing” but without the pumping movements (raising and lowering the body above the surface) usually observed in large Liphistius spiders.

Two females carried mites of the genus Ljunghia (see Halliday & Juvara-Bals, 2016: 857) which left clearly visible dark bite marks on the carapace and on the light proximal portion of the chelicerae, but not on other parts of the body. The mites were seen to aggregate in the fovea, around the sternum, under the spinnerets and on the ventral side of the leg femora of the spiders. One mite was seen inserting its gnathosoma into one of the bite marks on the carapace, obviously taking in food.

Two immature males (with swollen palpal tarsi and thus possibly penultimate; not paratypes) had a swollen, light-coloured (like porcelain) opisthosoma when collected and died soon afterwards. This indicates an infection with Rickettsiales (see Haupt, 2003: 66-67, fig. 41D). Haupt (2003: 67, figs 24A, 25A) reported rickettsia-like microorganisms in the spermophore of the mesothelid Ryuthela nishihirai (Haupt, 1979) and assumed that infection can occur through copulation. This was certainly not the case in the two immature males of L. indra sp. nov.

At the type locality, L. indra sp. nov. occurs together with Liphistius cf. thaleban. Burrows of both species were found side by side.

Diagnosis: Small to medium-sized spiders (carapace length of males 4.42-6.46, carapace width 4.02-5.87). Distinguished from species of other groups by males with a strongly reduced proximal edge on the tegulum (Figs 19H, 21B, 23I) and with an elongated prodorsal-apical lobe on the cymbium (Figs 19C, 21H, 23J). Females similar to those of the linang-group in possessing a vulval plate with reduced pigmentation and sclerotisation, distinguished by a relatively wider vulval plate with the poreplate only partially connected to the large and wide posterior stalk or separated by a short distance (Figs 20, 22, 24).

Species included: Liphistius batuensis, L. tempurung and L. priceae sp. nov.

Relationships: The reduced pigmentation and sclerotisation of the vulval plate and the reduced posterior edge of the tegulum point to a close relationship with the linang-group, but a loss or reduction is only a weak indication for common ancestry. At present the relationships of the batuensisgroup with other groups are unclear, but there is strong morphological evidence for close relationship among the species of the batuensis-group. Moreover, all three species have a strong association with caves (much more so than in species of the tioman-group).

Distribution: Western and central part of peninsular Malaysia (Fig. 1, localities 15-18). All species are known exclusively from limestone caves.

Liphistius batuensis Abraham, 1923a: 15-21, pl. 1, figs 1-9 (description of male and female). Synonymy and taxonomic literature, see World Spider Catalog (2017).

Types: BMNH; male lectotype (designated by Haupt, 1983: 282) and 11 female plus 4 juvenile paralectotypes (all not examined); Malaysia, Selangor, Batu Caves; XII.1921-I.1922; leg. H.C. Abraham.

Material examined: MHNG; 1 male; Selangor, Kuala Lumpur, Batu Caves; 24.VII.1969; leg. R. Pilet. – MHNG; 2 females; Batu Caves, près de Kuala Lumpur; XI.1976 ; leg. B. Koepchen. – MHNG, sample SIM-01/13; 2 males (matured 18.X.2001, 8.II.2002), 2 females (moulted 13.I.2002; 24.I.2002), exuviae of 3rd female (spider not collected); Selangor, Batu Caves, Dark Cave, Caverns B, C, D (3°14′12.7″N, 101°41′00.0″E), 100 m; 12.VII.2001; leg. P.J. Schwendinger. – SMF-13907, n° 3; 2 females, 1 juv. male; Selangor, Batu Caves; leg. Clark, det. Roewer, 1962 (identified as 3 females). – SMF-13908, n° 4; macerated remains of 2 specimens (one a juvenile male, the other without opisthosoma); Selangor, Batu Caves; leg. Clark, det. Roewer, 1962. – SMF-21945/1; 1 female; no locality data; 19.XI.1933; det. W. S. Bristowe.

Diagnosis: Medium-sized, light brown-coloured species in both sexes. Males characterized by cumulus carrying a group of very long, thick bristles (Fig. 19A-B); tegulum with exceptionally small and narrow proximal edge carrying few denticles (Fig. 19H); distal edge of contrategulum with 2-4 quite long parallel ridges pointing towards dorsal apex, without denticles (Fig. 19D-E, I); para-embolic plate distinctly elevated (Fig. 19E-I), with sharp proventral angle at base (Fig. 19D-E, G; see arrows); apex of embolus proper wide in prolateral and retrolateral view (Fig. 19E-F, H), its dorsal wall much wider than its ventral wall, end of dorsal wall sharply bent proventrad (Fig. 19D-F, I). Females characterized by poreplate largely unpigmented in its posterior portion, more or less distinctly separated from short, wide posterior stalk; posterior portion of genital atrium not markedly bent ventrad; no anterolateral lobes on poreplate; CDO very wide, its anterior margin sunken and clearly outlined, its posterior margin at level of poreplate and not clearly outlined; genital atrium in most cases without hairs, its lateral folds weakly developed (Fig. 20).

Additions to description of male: Scopula: Tarsus I with thin scopula in distal half of ventral side, divided for its entire length by a narrow pale, glabrous longitudinal median stripe and by some bristles; tarsus II likewise, scopula only distally divided; tarsus III with thin scopula in distal three-fifths, only distally divided; tarsus IV with thin scopula (narrower than on other tarsi) in distal half, only distally divided.

Palp: Tibial apophysis basally wide in ventral view, moderately long, slightly set back from distal margin of tibia (Fig. 19A), its apical megaspines relatively long and thin (but not as thin as illustrated in Murphy & Platnick, 1981: figs 9, 12, 15, 21 and in Platnick & Sedgwick, 1984: figs 70-72, 74), dorsal ones longer than ventral ones. Prodorsal-apical lobe of cymbium distinctly longer and more pointed than proventral-apical one (Fig. 19C). Paracymbium of average size and depth (Fig. 19A-B), carrying very long (longest ones reaching base of embolus complex), thick bristles in a loose group on non-elevated cumulus (Fig. 19A). Subtegulum without apophysis. Tegulum with very small proximal edge carrying only few denticles (Fig. 19H). Pigmented bridge between tegulum and contrategulum on retrolateral side of palpal organ disconnected (Fig. 19H). Contrategulum with indistinct, short and wide ventral process; distal edge with 2-4 parallel ridges pointing towards tongue-shaped dorsal apex (Fig. 19D-E, I). Para-embolic plate distinctly elevated (Fig. 19E-I), with sharp proventral angle at its base (Fig. 19D-G, see arrows). Embolus proper with wide apex (Fig. 19E-F, H); dorsal wall of sclerotised part much wider than ventral wall, end of former sharply bent ventrad and partly overlapping membranous part of embolus proper (Fig. 19D-F, I); sclerotised part of embolus proper retrolaterally with a single longitudinal distal keel (Fig. 19D-F, G); membranous part of embolus proper unpigmented throughout.

Additions to description of female: Posterior margin of genital sternite straight or indistinctly invaginated (Fig. 20). Vulval plate (Fig. 20) short, wide; posterior portion and lateral margins of poreplate largely unpigmented, more or less distinctly separated from short, wide, well-pigmented posterior stalk; posterior part of genital atrium only slightly and gradually curved ventrad; anterior margin of poreplate slightly and widely invaginated, without anterolateral lobes; CDO very wide, its anterior margin sunken and clearly outlined, its posterior margin level with poreplate and not clearly outlined (Fig. 20A, C-D, F-G); receptacular cluster very large, strongly protruding anteroventrad, reaching beyond anterior margin of poreplate; genital atrium without lateral hairs and in most cases without median hairs, its lateral folds indistinctly developed (Fig. 20B, E, H).

Remarks: The apical megaspines on the tibial apophysis of the male palp are not as thin as illustrated by Platnick and Sedgwick, and the one situated most dorsally is not as long as shown by these authors (Platnick & Sedgwick, 1984: figs 70-72, 74).

It is not clear what Platnick & Sedgwick (1984: 25) mean with their diagnostic character “short, erect tegular apophysis”.

Variation: Carapace lengths in males (n=3) 5.24-5.99, in females with fully developed vulval plate (n=7) 4.76-6.67; carapace widths 4.60-5.56 and 4.09-5.56, respectively. The scopula on tarsus I is divided for its entire length in two males, only distally so in the third; the scopula on tarsus IV extends over the distal half in two males, over the distal three-fifths in the third. One mature male retains a proventral “tibial spur” (sensu Platnick & Goloboff, 1985) on its right leg I. Variation in the shape of the vulval plate of females examined is shown in Fig. 20 (see also Murphy & Platnick, 1981: figs 26-27 and Platnick & Sedgwick, 1984: figs 75-76). One female has a single median hair in its genital atrium (Fig. 20D); in all other females examined the genital atrium is completely devoid of hairs (Fig. 20A-C, E-H). Most specimens examined have fully developed anterior median eyes; only one small female has a reduced (but cornea present) AME on one side.

Distribution: First reported from the Batu Caves by Ridley (1899: 580), this specis is only known from the Dark Cave and the Rift Cave which are part of the Batu Cave system (Lim & Yussof, 2009: 127-128; Fig. 1, locality 15) north of Kuala Lumpur, and from the Anak Takun Cave in the Templer Park which lies about 9 km further north (Fig. 1, locality 2). The latter record is based only on females and juveniles (Platnick & Sedgwick, 1984: 26). The Gua Anak Takun population was reported to contain hundreds of burrows in 1961 (McClure et al., 1967), only one in 1986 (Yussof, 1987), three in 2006 (Lim & Yussof, 2009). Steiner (1998: 148) reported that the Gua Anak Takun population had probably become extinct, which fortunately proved to be incorrect. Neverthess the future of this population, which no longer receives legal protection, is bleak (Lim & Yussof, 2009: 131). Fortunately the Batu Cave populations are doing much better (Lim & Yussof, 2009: 125-126).

Biology: I collected spiders of this species only from sac-like nests on cave walls in the aphotic zone of the Batu Caves. No burrows were seen in the soil. The largest nest inspected (of a female) was 6 cm long and had a 1.9 cm long and 3.0 cm wide front door, as well as a 1.2 cm long and 1.9 wide back door. Penultimate males in captivity built nests with 1.6-1.8 cm long and 2.4 cm wide trapdoors. An egg sac of 1.8 cm diameter and 0.9 cm depth was found in the cave; it contained no eggs. One mature male was collected (by R. Pilet) in the cave in late July; two males matured in captivity (three and six and a half months after capture) in late October and early February. The mating period of L. batuensis appears to be relatively long, as it is the case in other congeneric species in peninsular Malaysia. Despite the fact that all specimens examined were found in the dark portions of caves, no troglobiomorphic adaptations are visible apart from a relatively light body colouration in living spiders.

An excellent and very detailed account of the biology of L. batuensis was given by Klingel (1967). Similar but shorter presentations of this species and its habitat appeared in Abraham (1923a), Bristowe (1933, 1952) and McClure et al. (1967). Lim & Yussof (2009) gave a detailed account of the state of the three known populations of this species.

Liphistius tempurung Platnick in Platnick, Schwendinger & Steiner, 1997: 4-6, figs 10-11 (description of female).

Type: AMNH; female holotype (not examined); Malaysia, Perak, Gua Tempurung; 18.V.1996; leg. H. Steiner.

Material examined: MHNG, sample MAL-04/17; 3 males (matured 25.IX., 26.IX.2004, 13.XI.2005), 6 females (moulted 21.X.2004; 9.XII.2004; 19.XI.2004 and 3.IV.2005; 30.XII.2004 and 17.X.2005 and 24.V.2006 and 22.I.2007; IX.2004 and 13.IX.2005 and 8.II.2006; 8.X.2004 and 1.III.2005); Perak, 25 km south of Ipoh, Gua Tempurung, 4°24′58″N, 101°11′15″E, 90 m (limestone cave, dark zone); 22.VI.2004; leg. P.J. Schwendinger. – MHNG, sample 200/01; 1 juvenile; Perak, Gunung Lanno, Gua Cicak, 80 m; 16.XI.2001; leg. H. Steiner. – MHNG, sample MAL-04/16; 3 males (matured 31.XII.2004, 23.I.2005, 17.VI.2005), 3 females (moulted 8.I.2005; 17.I.2005); Perak, Gunung Lanno, Gua Cicak; 21.VI.2004; leg. P.J. Schwendinger.

Diagnosis: Medium-sized, light brown-coloured species in both sexes. Similar to L. batuensis, males distinguished by much shorter and stouter megaspines on tibial apophysis of palp (Fig. 21E-G cf. Fig. 19A); proximal edge of tegulum without denticles (Fig. 21B cf. Fig. 19H); retrolateral bridge between tegulum and contrategulum unbroken (Fig. 21B cf. Fig. 19H); contrategulum with more pronounced, rather conical ventral process; distal edge with several short parallel ridges situated near base of ventral process and directed towards embolus; dorsal apex of contrategulum wider, without ridges (Fig. 21I-L cf. Fig. 19D); embolus proper more slender (Fig. 21A-B cf. Fig. 19E, H), dorsal wall of its sclerotised part without sharply bent end overlapping membranous part (Fig. 21C, I-L cf. Fig. 19D-F, I); base of embolus complex without sharp proventral angle (Fig. 21I-L cf. Fig. 19D-E, G), para-embolic plate less elevated (Fig. 21B cf. Fig. 19H). Females distinguished from those of L. batuensis by anterior and posterior margins of vulval plate distinctly invaginated (Fig. 22 cf. Fig. 20); posterior part of genital atrium more strongly curved ventrad, clearly visible on female specimen when viewed from posteroventrally (Fig. 22M-N); poreplate with much smaller CDO (Fig. 22A, C, E, G, I, K cf. Fig. 20A, C-D, F-G).

Description of male (matured 26.IX.2004): Colour in alcohol (slightly darker in life): Sclerotised parts light brown, except for cream-coloured proximal portion of chelicerae, whitish membranes of prosoma and cream-coloured membranes of opisthosoma.

Bristles on carapace: Short bristles along all margins (longest and strongest in front of eye mound); few on coxal elevations; four short bristles anterior to fovea; thick (spinelike) bristles on eye mound.

Scopula: Thin (weakest and narrowest on tarsus IV) and only distally divided by median stripe on all leg tarsi; covering distal half of ventral side of tarsi I-III, only distal one-third of tarsus IV.

Cheliceral teeth: Eleven small teeth of different sizes on promargin of right cheliceral groove, twelve on left cheliceral groove.

Palp (illustrations mostly of male matured 13.XI.2005): Tibial apophysis basally quite wide in ventral view, distinctly set back from distal margin of tibia (Fig. 21E), carrying four medium-long apical megaspines, the second from ventral situated clearly lower (more proximal) than others (Fig. 21E-F). Apical lobes of cymbium distinct, prodorsal one longer and narrower than proventral one (Fig. 21H). Paracymbium of average length but quite shallow (Fig. 21A, D), carrying long thick bristles in a widely spaced group on non-elevated cumulus (Fig. 21A, D). Subtegulum without apophysis. Tegulum with proximal edge indistinct and completely without denticles, distinguished from rest of tegulum only by a stronger pigmentation (Fig. 21B). Pigmented bridge between tegulum and contrategulum on retrolateral side of palpal organ unbroken (Fig. 21B). Contrategulum with distinct, somewhat conical (in distal view) ventral process, with several parallel ridges directed towards embolus and situated near base of ventral process; dorsal apex of contrategulum large and widely tounge-shaped (Fig. 21I). Para-embolic plate short (Fig. 21B); no sharp proventral angle on and no sharp edge below para-embolic plate. Embolus proper with moderately wide apex (Fig. 21A-B); dorsal and ventral walls of sclerotised part equally wide, without modification (Fig. 21I), retrolateral wall carrying a distinct distal keel (Fig. 21B); membranous part of embolus proper narrow, unpigmented throughout (Fig. 21C).

Measurements: Total length 14.55; carapace 5.69 long, 5.25 wide; opisthosoma 6.04 long, 5.15 wide; eye mound 0.67 long, 0.83 wide; palpal coxae 1.68 long, 1.19 wide; labium 0.45 long, 0.99 wide; sternum 2.52 long, 1.68 wide (on ventral surface 0.79); palp 9.80 long (2.97 + 1.73 + 3.47 + 1.63); leg I 16.78 long (4.85 + 2.13 + 3.61 +4.16 + 2.03); leg II 18.07 long (4.95 + 2.13 + 3.96 + 4.65 + 2.38); leg III 19.65 long (5.00 + 2.13 + 4.21 + 5.64 + 2.67); leg IV 25.45 long (6.04 + 2.38 + 5.25 + 8.12 + 3.66).

Additions to description of female: Posterior margin of genital sternite widely W-shaped (Fig. 22A-I), in slightly posteroventral view posterior edge of posterior stalk of vulval plate clearly visible between paramedian lobes (Fig. 22M-N). Vulval plate (Fig. 22A-I, K) moderately wide; large parts of poreplate unpigmented, particularly central area around CDO; CDO indistinct, composed of a small drop-shaped or longitudinally slit-shaped opening lying within a larger hollow with a distinct, sunken anterior margin and an indistinct (level with poreplate) posterior margin (Fig. 22A, C, E, G, I); receptacular cluster large and complex (Fig. 22B, D, F, H), slightly protruding beyond widely invaginated anterior margin of poreplate; posterior stalk strongly pigmented, more or less completely separated from pigmented areas of poreplate, its posterior portion finely pitted and bent ventrad at right angles; genital atrium with few or without lateral hairs, most vulvae without posterior hairs (Fig. 22A, C, E, G, I, K).

Variation: Carapace lengths in males (n=6) 5.51-6.46, carapace widths 4.92-5.87; in the largest females with a well-developed vulval plate (n=9) 5.94-7.87 and 4.80-6.69, respectively. In two females from Gua Tempurung both AME are completely absent, in other specimens they are reduced (without cornea), tiny (smaller than the pits from which nearby bristles arise) or normally developed. There is only minor variation in the tarsal scopula of males: covering distal two-thirds of tarsus II in two males (both from Gua Cicak), only distal half in all other males; being very thin (in two males from Gua Cicak) or thin (in remaining males) on tarsus IV, always weaker than on tarsi I-III.

Variation in the shape of the ventral process of the contrategulum is shown in Fig. 21I-L, variation in the shape of vulval plates in Fig. 22A-I, K. One male from the Gua Cicak has one of the four apical megaspines on its tibial apophysis weakly developed on both palps (Fig. 21G). The number of lateral hairs in the genital atrium of females ranges from a few to none (Fig. 22A-I); one female (from Gua Cicak) has additionally two paramedian hairs in the posterior part of its genital atrium (Fig. 22I). Females from Gua Cicak have more hairs in the genital atrium than females from the type locality. The CDO is relatively large and clearly discernible only in the two smallest females from each locality (Fig. 22I, K); in all larger females examined it is small and lies in a depression (Fig. 22A, C, E, G).

Biology: All specimens examined were collected from burrows in sandy or loamy soil on the floor in the aphotic zone of two caves. Unlike in L. batuensis, no nests were found on cave walls. Most burrows ran into the depth of the soil and were closed by a single trapdoor. Only one burrow ran under the soil surface and had a second door at the opposite end, about six centimetres away from the first door. From the main entrance up to eight quite long (as usual in cave-dwelling Liphistius) signal lines (the longest 20 cm) ran over the soil surface (in a few cases also over nearby rock). The trapdoor of the largest female was 1.8 cm long and 3.0 cm wide, in penultimate males trapdoors were 1.2-1.8 long and 2.0-2.8 wide.

Despite the fact that all spiders examined were found in the dark portions of these two caves, no noteworthy troglobiomorphic adaptations are visible, apart from a relatively light body colouration in all living spiders and a more or less distinct reduction of the AME in some spiders.

An illustrated presentation of L. tempurung (together with L. kanthan) and its habitat is given in Steiner (1998). An extensive bibliography on Lipistius and many other animals in caves of Malaysia can be found on Liz Price's website (Price, 2017).

Distribution: Known only from inside two limestone caves (Fig. 1, localities 16-17), about 13 km apart, southeast of Ipoh in the Perak State. A short description of Gua Tempurung and photographs of the cave and some spider nests are given in Platnick et al. (1997: 2-3, figs 1-3).

Remarks: First reported from a “cave in Gunong Tempurong” by Bullock (1972: 138) under L. batuensis, this is a narrowly endemic species restricted to caves. Like the other cave-living Liphistius species in Malaysia [L. batuensis, L. kanthan (currently on the IUCN list of critically endangered species;  http://www.iucnredlist.org/search?page=52) and L. priceae sp. nov.], L. tempurung is at great risk of becoming extinct if overcollected, or if their habitats change or are destroyed. Quarrying of limestone hills for cement production is widespread in Malaysia and elsewhere in Southeast Asia, and poses the greatest danger for these species.

Types: MHNG (most types, including the holotype), SMF (1 male and 1 female paratypes), sample MAL-04/10; male holotype (matured late VIII.2004), 8 male paratypes (matured 20.VII., 1.VIII., 18.IX. 2004, mid-VII.; 11.VIII.2005; 3 male paratypes collected mature at the site) and 11 female paratypes (including allotype, moulted 24.XI.2004); Malaysia, Kelantan, about 5 km S of Dabong, Gua Keris (= Kris Cave) and Gua Pagar (locally also called Gua King Kong), 130 m; 8.-9. VI.2004; leg. P.J. Schwendinger. The precise type locality is Gua Keris.

Etymology: The new species is named in honour of Liz Price (London, UK), a former long-time resident in Kuala Lumpur, and a very active speleologist who over 30 years explored and published on caves and cave faunas all over Southeast Asia. She was also involved in conservation and trying to save some caves from destruction by quarrying in Malaysia.

Diagnosis: Small to medium-sized, light-brown-coloured species, similar and closely related to L. tempurung. Both sexes slightly smaller than those of L. tempurung (carapace length of males 4.42-5.23 versus 5.51-6.46 in L. tempurung). Males distinguished by scopula on tarsus IV more extensive; palpal tibia with a longer retrolateral apophysis carrying much shorter megaspines (Fig. 23B, K-O cf. Fig. 21E-G); paracymbium larger, with a more prominent retrolateral-proximal part (Fig. 23A cf. Fig. 21A); tegulum with more clearly outlined proximal edge (Fig. 23I cf. Fig. 21B); contrategulum with series of oblique ribs on distal edge extending further prolaterally, dorsal apex much smaller and narrower, with a more pointed tip (Fig. 23D-G cf. Fig. 21I-L); para-embolic plate with more prominent, angular distal margin (Fig. 23A, D-E cf. Fig. 21A, I-L); sclerotised part of embolus proper with more pointed apex (Fig. 23A, I cf. Fig. 21A-B). Females distinguished from those of L. tempurung by larger and more prominent receptacular cluster (Fig. 24 cf. Fig. 22); CDO much wider and more distinctly outlined, not sitting in a hollow (Fig. 24A, C, E-F cf. Fig. 22A, C, E, G, I); anterior margin of poreplate not or only indistinctly invaginated; posterior stalk anteriorly narrower.

Description of male (holotype): Colour in alcohol (slightly darker in life): Body mostly light brown, carapace with indistinct cream-coloured and flower-shaped area around fovea. Chelicerae with proximal portion cream-coloured, distal portion light brown. Cheliceral fangs, palpal tarsi and sclerites of palpal organ reddish brown (darker than rest of body). Legs with indistinct annulations (light median rings) on all tibiae and metatarsi; legs and palps with a light distal zone on patellae. Opisthosomal tergite I uniformly light brown, following tergites with increasingly larger light areas in posterior part (except for bases of paired para-median spines); membranous part of opisthosoma cream-coloured.

Bristles on carapace: Stiff bristles on anterior and posterior margins, as well as on and behind eye mound; fewer weaker bristles on lateral margins and on posterior coxal elevations; no bristles anterior to fovea.

Cheliceral teeth: Twelve small teeth on promargin of cheliceral groove of each chelicera.

Scopula: All tarsi with indistinctly outlined scopula on roughly distal two-thirds of ventral side, only behind claw divided by a short median stripe; scopula on tarsus I very thin and its proximal limit difficult to identify; scopula on other tarsi denser and and more clearly delimited.

Claws: Paired tarsal claws with 3-4 denticles on anterior legs, 4-5 denticles on posterior legs; unpaired claws with one indistinct denticle or without.

Palp: Tibial apophysis well-developed and quite long, slightly set back from anterior margin of tibia, distinctly pointing away from axis of tibia (Fig. 23B), carrying four very short megaspines, the ventral one being longest, a median one set back from distal margin (Fig. 23K). Distal margin of cymbium with elongate prodorsal lobe (Fig. 23J). Paracymbium quite large and moderately deep, with a distinct retrolateral-proximal bulge (Fig. 23A, C); cumulus indistinctly elevated, carrying few long stiff bristles (Fig. 23A, C). Subtegulum without apophysis. Tegulum with wide, sharp, non-dentate proximal edge (Fig. 23I). Contrategulum with somewhat widely conical (its apex narrowly rounded) ventral process; distal edge with long row of oblique parallel ridges pointing towards embolus; dorsal apex asymmetrical, quite small and narrow, with narrowly rounded tip (Fig. 23D). Para-embolic plate low, its distal margin angular (Fig. 23A, D, I); embolus proper with slightly widened and obliquely truncate apex (Fig. 23A, I), dorsal and ventral walls of sclerotised part equally wide and lying close to each other (Fig. 23D), retrolateral side enforced by a long, distinct keel (Fig. 23D, I); membranous part of embolus proper narrow, indistinct (Fig. 23H).

Measurements: Total length 9.96; carapace 4.51 long, 4.24 wide; opisthosoma 3.96 long, 2.61 wide; eye mound 0.64 long, 0.76 wide, AME well-developed; palpal coxae 0.99 long, 0.95 wide; labium 0.40 long, 0.87 wide; sternum 2.02 long, 1.47 wide (0.75 on ventral surface); palp 7.77 long (2.46 + 1.39 + 2.69 + 1.23); leg I 14.69 long (3.96 + 1.82 + 3.17 + 3.64 + 2.10); leg II 15.61 long (4.08 + 1.82 + 3.29 + 4.08 + 2.34); leg III 17.18 long (4.16 + 1.86 + 3.56 + 4.91 + 2.69); leg IV 21.62 long (5.07 + 1.90 + 4.51 + 6.69 + 3.45).

Description of female (allotype): Colour in alcohol (slightly darker in life): Mostly as in male, annulations on tibiae and metatarsi of legs slightly more distinct; palpal tarsus reddish brown only at tip. Opisthosomal tergites generally darker, light patches smaller.

Bristles on carapace: Mostly as in male, additionally with very few tiny bristles on anterior coxal elevations.

Cheliceral teeth: Eleven strong teeth on promargin of left cheliceral groove, twelve on right side.

Claws: Each palpal claw with three worn denticles. Paired leg claws with 2-4 denticles; unpaired claws of legs I-III with 2-3 denticles, leg IV with 0-2. All tarsi without scopula.

Vulva: Posterior margin of genital sternite widely W-shaped (Fig. 24C-D), in slightly posteroventral view posterior edge of posterior stalk clearly visible between paramedian lobes. Vulval plate (Fig. 24C-D) wider than long; posterior part of poreplate partly unpigmented; CDO distinctly outlined, somewhat quadrangular, longer than wide (Fig. 24C), leading into large and complex receptacular cluster (Fig. 24D); the latter strongly protruding beyond slightly arched anterior margin of poreplate (Fig. 24C); posterior stalk strongly pigmented, trapezium-shaped, anteriorly much narrower than posteriorly, indistinctly separated from pigmented areas of poreplate, its posterior portion finely pitted and bent ventrad at right angles (Fig. 24B, showing paratype).

Measurements: Total length 15.38; carapace 6.18 long, 4.91 wide; opisthosoma 6.49 long, 4.83 wide; eye mound 0.69 long, 0.82 wide; palpal coxae 1.90 long, 1.43 wide; labium 0.63 long, 1.43 wide; sternum 2.85 long, 1.90 wide (1.11 on ventral surface); palp 10.26 long (3.33 + 1.82 + 2.65 + 2.46); leg I 12.91 long (4.12 + 2.06 + 2.81 + 2.61 + 1.31); leg II 13.42 long (4.12 + 2.10 + 2.85 + 2.81 + 1.54); leg III 14.30 long (4.20 + 2.14 + 2.93 + 3.33 + 1.70); leg IV 19.46 long (5.23 + 2.38 + 4.08 + 5.39 + 2.38).

Variation: Carapace lengths in males (n=9) 4.42-5.23, carapace width 4.02-4.76; in females with well-developed copulatory organs (n=9) 5.54-5.85 and 4.61-5.04, respectively. In a male and in a female one of the two AME is absent, in all other specimens both are distinct. In two males (including the holotype) the W-shaped marking behind the eye mound is indiscernible, in the other males it is more or less well-developed. No noteworthy variation in size and density of the tarsal scopula of males was detected. The arrangement of distal megaspines on the tibial apophysis of the male palp is quite variable (Fig. 23K-O). A male paratype has fewer bristles on the cumulus of both palps than the other males; one of the cumulus bristles of the illustrated holotype palp runs to the dorsal side of the palpal organ (Fig. 23A, C). The dorsal apex of the contrategulum is narrowly rounded to pointed (Fig. 23D-G). Variation in the shape of the vulval plates is shown in Fig. 24. The CDO is small to large, wider than long or longer than wide, circular, elliptical or quadrangular (Fig. 24A, C, E-F). The posterior stalk is more or less distinctly separated from the pigmented area of the poreplate. The lateral walls of the genital atrium are developed as narrow folds (Fig. 24D) or as simple trenches. Three of the females examined have 1-2 lateral hairs on only one side of the genital atrium (Fig. 24A-B, E-F), the allotype has none (Fig. 24C-D).

Distribution: Known only from two caves in the same limestone hill (Fig. 1, locality 18) in the north of peninsular Malaysia.

Biology: The spiders examined were all collected in different zones (from the euphotic entrance area to the aphotic interior) of both caves. No such spiders were found in the adjacent degraded rain forest. Despite apparently being confined to these caves, L. priceae sp. nov. shows no noteworthy cave adaptations in its morphology.

Quite short burrows with a single door and relatively long signal lines (no measurements were taken), as usual for cave-dwelling Liphistius, were built in horizontal or sloping loam at or near the cave entrance, as well as in cracks and holes of the walls at the entrance and in deeper portions of the caves. No sac-like retreats on rock surface, as constructed by some other cave-dwelling Liphistius, were seen. The trapdoor of the largest female was 1.4 cm long and 2.4 cm wide, in penultimate and adult males they were 1.15-1.6 long and 1.55-2.0 wide. Three males were adult when collected in early June 2004; four others matured not long afterwards, between late July and late September; the remaining two males matured in mid-July and late August of the following year. Adult females usually moulted twice per year: between March and August and again between September and January. No egg cases were found in the field and no eggs were laid in captivity, which means that the females had not yet mated when collected in early June.