Liphistius malayanus Abraham, 1923

Peter J. Schwendinger, 2017, A revision of the trapdoor spider genus Liphistius (Mesothelae: Liphistiidae) in peninsular Malaysia; part 1, Revue suisse de Zoologie 124 (2), pp. 391-445 : 395-401

publication ID	https://doi.org/ 10.5281/zenodo.893555
DOI	https://doi.org/10.5281/zenodo.6042349
persistent identifier	https://treatment.plazi.org/id/4C30A452-FFE0-FFFD-BA9A-FAC53979F872
treatment provided by	Plazi
scientific name	Liphistius malayanus Abraham, 1923
status

Treatment

Liphistius malayanus Abraham, 1923 View in CoL

Figs 3B, D, F, H View Fig. 3 ; 4 View Fig. 4 - 5 View Fig. 5

Liphistius malayanus Abraham, 1923b: 770 View in CoL -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 View in CoL , 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. GoogleMaps – 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. GoogleMaps – SMF 7425 About SMF /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. GoogleMaps – 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. GoogleMaps – 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. GoogleMaps

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 View Fig. 4 ); distal edge of contrategulum carrying a series of small denticles ( Fig. 4 View Fig. 4 G-L), those at proventral end arranged in a V-shaped row ( Fig. 4C View Fig. 4 ), 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. 4 View Fig. 4 G-L); dorsal wall of embolus proper as wide as ventral wall ( Fig. 4 View Fig. 4 G-H); membranous part of embolus proper distally narrower than proximally ( Figs 3F, H View Fig. 3 , 4C View Fig. 4 ). Females distinguished by vulval plate ( Fig. 5 View 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 View Fig. 4 ), carrying four long apical megaspines ( Fig. 4 View Fig. 4 A-B). Paracymbium relatively small and shallow, about as deep as cymbium in retroventral view ( Fig. 4A View Fig. 4 ; 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. 4 View Fig. 4 D-F). Contrategulum with very indistinct, widely rounded ventral process ( Fig. 4 View Fig. 4 G-H); distal edge with a series of small denticles, those at proventral end arranged in a V-shaped row ( Fig. 4C View Fig. 4 ), 1-2 enlarged denticles or a sharp keel in prolateral to prodorsal section of distal edge ( Fig. 4 View Fig. 4 G-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. 4 View Fig. 4 G-L). Embolus complex with para-embolic plate developed as a more or less distinctly elevated, rounded edge ( Fig. 4 View Fig. 4 D-F); below it a second small edge present in some cases ( Fig. 4D View Fig. 4 ); sclerotised and membranous part of embolus proper in contact for almost their entire length ( Fig. 3D, F, H View Fig. 3 ); sclerotized part with numerous longitudinal ribs carrying tiny denticles distally ( Fig. 3B, D, F, H View Fig. 3 ), its dorsal and ventral walls equally wide and parallel to each other ( Figs 3D View Fig. 3 ; 4G-H), dorsal wall ending in indistinct rounded lobe ( Fig. 4 View Fig. 4 C-F); membranous part distally narrower than proximally ( Fig. 3D, F, H View Fig. 3 ), its proximal portion essentially unpigmented.

Additions to description of female: Vulval plate ( Fig. 5 View Fig. 5 ) always with few to many hairs laterally in genital atrium, rarely also posteromedially ( Fig. 5I View Fig. 5 ). 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. 5 View Fig. 5 A-C, E); anterior margin of poreplate straight to procurved; lateral and posterolateral margins on ventral side of pore plate bulging ( Fig. 5B, J View Fig. 5 ). CDO large and of variable shape, mostly longer than wide, rarely wider than long ( Fig. 5E View Fig. 5 ). Receptacular cluster large and strongly racemose, in most cases protruding beyond anterior margin of poreplate, rarely not ( Fig. 5G View Fig. 5 ).

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. 5 View Fig. 5 A-B cf. Fig. 5 View Fig. 5 C-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 View Fig. 4 ), which is also present in the second male from the Cameron Highlands ( Fig. 4F View Fig. 4 ). 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 View Fig. 4 ). 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. 4 View Fig. 4 A-B). The ventral megaspine is bent in the male from the Templer Park ( Fig. 4A View Fig. 4 ), whereas in the holotype of L. m. cameroni ( Fig. 4B View Fig. 4 ), 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 View Fig. 4 , I-L); in two males there is a sharp edge at the same place ( Fig. 4H View Fig. 4 ), 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 View Fig. 4 ), in a second male from the Cameron Highlands ( Fig. 4F View Fig. 4 ) 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 View Fig. 4 ). 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 View Fig. 4 ), but not in other males examined ( Fig. 4 View Fig. 4 E-F). The size of the embolus proper varies from relatively thick in the male from the Templer Park ( Fig. 4D View Fig. 4 ) to distinctly more slender in the holotype of L. m. cameroni ( Fig. 4E View Fig. 4 ); the other males are intermediate ( Fig. 4F View Fig. 4 ). The longitudinal ribs on the sclerotised part of the embolus proper vary from indistinct to distinct ( Fig. 3B, D, F, H View Fig. 3 ), 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 twothirds 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 View Fig. 5 cf. Fig. 5H View Fig. 5 ). In the largest female (from the Templer Park) hairs are also present posteromedially in the genital atrium ( Fig. 5I View Fig. 5 ). The shape of the posterior stalk is quite variable but mostly trapezium-shaped ( Fig. 5 View 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 View Fig. 5 , 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 View Fig. 5 ); only in one female examined (from the Genting Highlands) does it not protrude beyond the anterior margin of the poreplate ( Fig. 5G View Fig. 5 ). The anterior poreplate margin varies from more or less strongly invaginated (in most cases; Fig. 5 View Fig. 5 A-E, G-J) to slightly arched medially ( Fig. 5F View Fig. 5 ).

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 View 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.