Liphistius cupreus Schwendinger & Huber

Schwendinger, Peter J., Huber, Siegfried, Lehmann-Graber, Christina, Ono, Hirotsugu, Aung, Mu Mu & Hongpadharakiree, Komsan, 2022, A taxonomic revision of the Liphistius birmanicus-group (Araneae: Liphistiidae) with the description of five new species, Revue suisse de Zoologie 129 (2), pp. 375-424 : 411-416

Liphistius cupreus Schwendinger & Huber , sp. nov.

Figs 1 , 2 D-I, 21-22

Holotype: MHNG-ARTO-0028341 (sample MT-14/21); male (matured 18.V.2017); Myanmar, Shan State, SE of Lashio, near Kong Nyaung Village (22°54’07”N, 97°47’27”E), 1130 m; 8.-9.VI.2014; leg. P.J. Schwendinger & S. Huber .

Paratypes: MHNG-ARTO-0028342 to MHNGARTO-0028347, BRCM (sample MT-14/21); 7 males (matured mid-VI.2015, two at end of VI.2015, 5.V.2016, 27.V.2016, 30.VI.2016, 1.VII.2016); collected together with the holotype. – MHNG-ARTO-0028349 to MHNG-ARTO-0028359, BRCM (sample MT-14/21); 12 females (allotype, MHNG-ARTO-0028349, last moulted 19.I.2019); collected together with the holotype.

Other material: MHNG; 2 small, badly preserved females, 1 juvenile; same data as for type specimens.

Etymology: The Latin adjective “cupreus” (= coppercoloured) refers to the colouration of the opisthosomal tergites in both sexes.

Diagnosis: Medium-sized spiders; both sexes with annulated legs (in females more distinct than in males) and without prefoveal setae. Most similar to L. nabang and L. platnicki sp. nov. by males sharing a distinctly elevated cumulus and a paracymbium with a distinctly conical distal side ( Fig. 21 A-B cf. Yu et al., 2021: fig. 3D-F and 23A-B); females sharing a similar vulval plate with more or less strongly reduced anterior poreplate lobes and with a short and wide posterior stalk widely connected to the popeplate ( Fig. 22 cf. Yu et al., 2021: fig. 4 and Fig. 24 ). Males distinguished from those of L. nabang by tegulum lacking a distinctly elevated distal edge ( Fig. 21K, M ; present in L. nabang, Yu et al., 2021 : fig. 3A-B, D-E) and having a widely rounded proximal edge ( Fig. 21K ; bent in L. nabang , giving the tegulum a triangular shape, Yu et al., 2021: fig. 3A); distal edge of contrategulum with an angular and finely dentate projection prodorsally ( Fig. 21A , E-F, H-I; in L. nabang widely arched and not dentate, Yu et al., 2021: fig. 3B); paracymbium without a retrolateral-proximal heel ( Fig. 21A cf. Yu et al., 2021: fig. 3E). Females distinguished from those of L. nabang by poreplates with 1-2 clusters of vesicles on each side of anterior margin (in L. nabang only 1) and without bulging posterolateral margins; posterior part of genital atrium not bent ventrad ( Fig. 22 cf. Yu et al., 2021: fig. 4). Males distinguished from those of L. platnicki sp. nov. by tegulum lacking a distinctly elevated distal edge ( Fig. 21K, M cf. Fig. 23 I-M), proximal tegular edge only slightly protruding ( Fig. 21J ; strongly protruding in L. platnicki sp. nov., Fig. 23M ); distal edge of contrategulum with an angular prodorsal projection ( Fig. 21A , E-F, H-I; no projection in L. platnicki sp. nov., Fig. 23A , E-F). Females distinguished from those of L. platnicki sp. nov. by poreplate lacking anterolateral processes and by a relatively shorter posterior stalk ( Fig. 22 cf. Fig. 24 ).

Description of male (holotype): Colour in alcohol (darker in life; as in Fig. 2 D-F): Carapace with orange background colour and dark lateral and anterior margins; pars cephalica grey-brown except for a pair of light brown lateral marks and an unpaired light posteromedian mark; eye mound black; pars thoracica with irregularly shaped dark marks on coxal elevations and with a light brown posteromedian mark. Chelicerae cream-coloured proximally and prolaterally, grey-brown medially, light orange-coloured distally. Ventral side of palpal coxae and of sternum light orange-coloured, contrasting with slightly darker labium and leg coxae. Legs and palps mostly dark brown on dorsal side, apart from distal half of tarsi I-II slightly lighter than proximal half and apart from an indistinct light median mark on leg tibiae; ventral side of legs with distinct light median mark on tibiae I-II and with an indistinct one on tibiae III-IV; ventral side of leg femora with distinct light median and proximal marks (remnants of annulations; see also Variation), plus scattered dark spots; ventral side of palpal tibia and femur light, with scattered dark spots; palpal tarsus and cheliceral claw dark reddish brown. Opisthosoma with light greyish brown membranous cuticle and with light orangecoloured sternites; tergite I entirely dark brown; tergite II mostly dark brown, with orange-coloured paramedian-posterior marks; tergites III-VII mostly light orange-coloured, with large dark marks, with a pair of dark paramedian-posterior marks and with scattered small dark spots; tergite VIII very small, entirely dark.

Setae on carapace: Few short setae (with wide gaps between them) on lateral margins; none on posterior margin; isolated setae on anterior coxal elevations, groups of 7-10 setae on posterior coxal elevations. No setae anterior to fovea.

Cheliceral teeth: 12 small teeth of different sizes on promargin of each cheliceral groove.

Scopula: Only distally divided by a median stripe on all tarsi; weak, posteriorly very thin and not clearly outlined in distal 4/5 of ventral side of leg tarsus I; equally thin and covering distal 5/6 of tarsus II; distinctly denser, more clearly outlined, covering 5/6 of tarsus III; equally dense and covering 4/5 of tarsus IV.

Tarsal claws: Paired claws with 4-5 teeth on tarsi I-III, 5 teeth on tarsus IV; unpaired claw of all leg tarsi without denticles.

Palp: Tibial apophysis short, triangular and basally wide in ventral view, not set back from distal margin of tibia ( Fig. 21C ), carrying 4 long apical megaspines, ventral 2 longer than dorsal 2; dorsal to apophysis a quite long spine on a slightly elevated base ( Fig. 21 C-D). Palpal tarsus with a pronounced subdistal suture on ventral side ( Fig. 21A ) and with a widely but shallowly invaginated distal margin ( Fig. 21N ). Paracymbium moderately deep ( Fig. 21B ), longer than wide, with a distinctly conical distal side and a widely arched retrolateral side in ventral view, lacking a retrolateral-proximal heel ( Fig. 21A ); cumulus quite large, distinctly elevated, carrying a densely packed group of about 10 long, strong bristles ( Fig. 21 A-B). Subtegulum without apophysis ( Fig. 21A, M ). Tegulum wide, axe-blade-shaped, its distal margin not elevated ( Fig. 21M ), its proximal edge short, widely rounded and finely dentate (see Fig. 21K showing paratype), only slightly bent and protruding, only little overhanging membranous contrategular area below it ( Fig. 21J ). Pigmented bridge between tegulum and contrategulum on retrodorsal side of palpal organ well developed ( Fig. 21E, M ). Contrategulum with moderately developed conical proventral process ending in a rounded apex in distal view ( Fig. 21 H-I); a few fine wrinkles on dorsal surface and a small proximal ledge on retrodorsal side ( Fig. 21H , L-N, see also Fig. 21 E-F for paratype); distal edge of contrategulum narrow, carrying a widely angular and finely dentate projection prodorsally ( Fig. 21A , H-I, see also Fig. 21 E-F for paratype), dorsal apex of contrategulum quite long and beak-like, very narrowly rounded ( Fig. 21 H-I, L, see also Fig. 21 E-G for paratype). Para-embolic plate short, about as long as retroventral edge of embolus complex and not visibly separated from it by an invagination ( Fig. 21A ). Membranous prolateral-proximal zone of embolus complex distinctly swollen ( Fig. 21A ); embolus proper narrowly divided, its sclerotised part strengthened by 2 thick and 1 thin longitudinal ribs reaching apex and carrying denticles distally ( Fig. 21A, M , see also Fig. 21 E-F, K for paratypes); membranous part of embolus proper distinctly shorter than sclerotised part, at its base a short, proximally wide and weakly pigmented area with numerous longitudinal wrinkles and with a concave distal margin; embolic folds short ( Fig. 21N ).

Measurements: Total length 15.55; CL 6.03, CW 5.55; opisthosoma 6.90 long, 5.55 wide; eye mound 0.86 long, 0.97 wide; palpal coxa 2.02 long, 1.43 wide; labium 0.52 long, 1.15 wide; sternum 3.06 long, 2.14 wide (1.03 on ventral surface); palp 9.65 long (2.94 + 1.79 + 3.21 + 1.71); leg I 16.82 long (4.68 + 2.38 + 3.41 +4.29 + 2.06); leg II 17.85 long (4.68 + 2.42 + 3.57 + 4.88 + 2.30); leg III 20.16 long (5.00 + 2.54 + 3.89 + 5.95 + 2.78); leg IV 24.64 long (5.99 + 2.70 + 4.88 + 7.58 + 3.49).

Description of female (allotype): Colour in alcohol (darker in life, as in Fig. 2 G-I): Mostly as in male, but generally slightly lighter (due to longer preservation in alcohol). Light distal area on chelicerae much smaller than in male and divided into a small median spot and a

► longitudinal retrodorsal stripe. Three light marks on pars cephalica and posteromedian light mark on pars thoracica of carapace less conspicuous. Palpal coxae darker than in males, almost of same colour as leg coxae I-II (leg coxae III-IV slightly lighter). Annulation of legs and palps more distinct than in males, clearly visible dorsally on all tibiae and on metatarsi III-IV, less clearly so on all femora and on tarsi III-IV. Distal 1/4 of palpal tarsus dark reddish brown.

Setae on carapace: Mostly as in male, but even fewer short setae on lateral margin; no setae anterior to fovea.

Cheliceral teeth: 11 strong teeth on promargin of right cheliceral groove; 11 strong teeth and 1 tiny denticle on left chelicera.

Claws: Palpal claws with 4 denticles each. Paired claws of legs I-II with 4 teeth, of leg III with 4-5 teeth, of leg IV with 5-6 teeth; unpaired claws of legs I-II with 1 denticle, of leg III with 0-1 denticle, of leg IV bare. All tarsi without scopulae.

Vulva ( Fig. 22J ): Membranous uterus externus with a distinct pair of lateral pockets (see Fig. 22 E-F for another female). Vulval plate wider than long; only one hair on lateral fold of one side. Poreplate with widely but shallowly invaginated anterior margin carrying a single small process or cluster of vesicles (probably reduced anterior lobes) on one side and two small processes on other side, with inconspicuous lateral and posterior margins and with small CDO; receptacular cluster racemose but rather simple, longer than wide, not reaching anterior margin of poreplate, very deep (see Fig. 22G, P for other females). Posterior stalk short, as wide as poreplate, axe-blade-shaped, anteriorly only slightly narrower than posteriorly, with a widely arched posterior margin.

Measurements: Total length 20.08; CL 7.62, CW 6.90; opisthosoma 8.65 long, 6.35 wide; eye mound 0.98 long, 1.12 wide; palpal coxa 2.54 long, 1.79 wide; labium 0.63 long, 1.67 wide; sternum 4.37 long, 2.62 wide (1.39 on ventral surface); palp 12.39 long (4.05 + 2.38 + 3.02 + 2.94); leg I 15.91 long (5.04 + 2.78 + 3.25 + 3.25 + 1.59); leg II 16.59 long (5.04 + 2.90 + 3.25 + 3.57 + 1.83); leg III 18.49 long (5.16 + 3.06 + 3.49 + 4.48 + 2.30); leg IV 25.04 long (6.83 + 3.29 + 4.92 + 7.06 + 2.94).

Variation: For carapace measurements and prefoveal setae counts see Table 1 . When still alive, at least one of the mature males examined showed light annulations on its leg femora, which are clearly visible not only on the ventral but also on the dorsal side ( Fig. 2D, F ); in alcohol these annulations are difficult to see. All specimens examined lack setae anterior to the fovea, and all specimens have both AME well developed. The extent of the scopulae differs among males: covering 3/4-5/6 of the ventral side of the leg tarsi (most often 4/5), rarely equally on all legs, in most specimens covering a larger portion on tarsi II-III that on tarsus I and tarsus IV. Variation in details of the male palp is given in Fig. 21 . The angular prodorsal protrusion at the base of the distal contrategular edge is more or less prominent but always clearly visible ( Fig. 21A , E-F, H-I, L). The proventral contrategular process in distal view is asymmetrically conical on the right palp of the holotype ( Fig. 21I ). Three male paratypes have a short subdistal ridge at the distoventral corner of their tegulum ( Fig. 21K ), which is very different in shape from but presumably homologous to the elevated distal tegular edge in L. nabang and L. platnicki sp. nov. ( Yu et al., 2021: fig. 3A-B, D-E and Fig. 23 I-M). All males examined have: three ridges (two thick and one thin) reaching the apex of the sclerotised embolus part; a swollen membranous area prolaterally-proximally on the embolus complex; a beak-like dorsal apex of the distal contrategular edge; a rather indistinct proximal ledge on the retrodorsal side of the contrategulum; a short and only slightly bent proximal tegular edge. In the largest female (the allotype) the annulation (a light median ring) of metatarsi I-II is completely broken on the dorsal side (being entirely dark); in medium-sized females there are small light spots on the dorsal side of metatarsi I-II (i.e. a partially broken annulation); in smaller females the annulation of metatarsi I-II is entire, as it is on metatarsi III-IV and on all tibiae. Variation in the shape of the vulval plate is considerable ( Fig. 22 ): the anterior poreplate margin bears one or two small processes on each side (often different numbers on both sides; these are probably reduced anterior lobes, as indicated by only little reduced lobes in two females, Fig. 22 A-B); the CDO is in the centre of the poreplate ( Fig. 22H ) or distinctly posterior to it ( Fig. 22N ); the shape of the receptacular cluster ranges from fairly small and simple ( Fig. 22O ) to quite large and complex ( Fig. 22C ), but it never reaches the anterior margin of the poreplate; most vulval plates have no hairs on their lateral folds, if present, there are only 1-2 and usually only on one side, in one specimen one of these hairs is unusually thick ( Fig. 22K ).

Relationships and biogeography: Liphistius cupreus sp. nov. is very similar and presumably most closely related to L. nabang . Considering how far the localities of both nominal species are apart, it is very unlikely that they are actually conspecific. All species in the birmanicus -group appear to have quite small geographical ranges. In Liphistius it is generally largesized species in the lowlands (e.g. L. yangae Platnick & Sedgwick, 1984 ; see Schwendinger et al., 2019: 333, fig. 2) that are known to have relatively large geographical ranges, not small or medium-sized species in the mountains. The type locality of L. nabang lies about 190 km north of that of L. cupreus sp. nov. (both in the mountains of the Shan-Yunnan Plateau), whereas the morphologically less similar but geographically much closer L. platnicki sp. nov. was found only 17 km north of the type locality of L. cupreus sp. nov. ( Fig. 1 ). These three species form a well-defined subgroup within the bristowei -group (see Discussion - Relationships).

Biology: The spiders were found on earth banks on the side of a rural road and on the sides of a stream in a disturbed evergreen hill forest. All burrows were closed by a single trapdoor and had up to eight signal lines attached to the entrance. Penultimate males had the front door 1.45-1.9 cm long and 2.2-2.5 cm wide. The largest female had a 2.1 cm long and 2.9 cm wide door.

Males matured in captivity between mid and end-June in the first year after capture (on June 8 and 9), between early May and the beginning of July in the second year, and in late May of the third year. The mating period in nature appears to be in May to June; possibly even much earlier, as indicated by the find of 3rd instar spiderlings in the burrow of a female when it was captured in early June. In captivity another female built an egg case, 2.8 cm long, 3.0 cm wide and 2.0 cm high, containing 39 light yellow eggs suspended on a layer of fine silk, about 2.5 months after being captured. Mature females usually moulted twice per year: between November and March and again between May and July. The female that built an egg case subsequently moulted in December.

Figures

Fig. 1. Localities of species in the Liphistius birmanicus-group in northern Thailand, eastern Myanmar and southern China. Blue lines indicate major rivers, black lines national borders, grey lines major roads.

Fig. 2. Habitus of some species of the Liphistius birmanicus-group. (A) Liphistius tung sp. nov., male holotype.(B) Liphistius pyinoolwin, male (matured 1.X.2014) from the Anisakan Falls; note exuviae of ectoparasitic mites (Ljunghia sp.) on the opisthosoma. (C) Liphistius platnicki sp. nov., female paratype (moulted 10.III.2021) from near Loi Kan Village. (D, F) Liphistius cupreus sp. nov., male paratype from near Kong Nyaung Village. (E) Same species, different male paratype. (G) Same species, female paratype (moulted 13.II.2021). (H-I) Same species, different female paratype (moulted 21.V.2021). Not to scale.

Fig. 21. Liphistius cupreus sp. nov., palps of three males; holotype (A-D, H-J, L-N), paratype matured end VI.2018 (E-G), paratype matured mid VI.2018 (K). (A) Left tarsus and palpal organ, ventral view. (B) Left paracymbium and distal margin of tibia, retrolateral view. (C) Distal part of left tibia, ventral view. (D) Same, retrolateral and slightly proximal view. (E) Left palpal organ, distal view (dorsal side up). (F) Dorsal part of left contrategulum and embolus proper, distal and slightly ventral view. (G) Dorsal part of left contrategulum, distal and slightly dorsal view. (H) Left contrategulum, distal and slightly ventral view. (I) Right contrategulum, same view. (J) Part of left palpal organ, retroventral view; slightly bent proximal edge of tegulum indicated by arrow. (K) Right palpal organ, retrolateral and slightly distal view. (L) Left palpal organ and distal margin of tarsus, dorsal and slightly proximal view. (M) Left palpal organ, retrolateral and slightly distal view. (N) Left palpal organ and distal margin of tarsus, prodorsal and slightly proximal view. Scale lines 1.0 mm (A-B, E-N; C-D). Abbreviation: a - angular prodorsal protrusion at base of distal edge of contrategulum.

Fig. 22. Liphistius cupreus sp. nov., vulval plates of ten females (all taken from exuviae), dorsal view (E with uterus externus attached, H, K, N), ventral view (A-D, I-J, L-M, O), ventral and slightly anterior view (P), posterior view (G), and isolated uterus externus (F). (A) Moult of 10.II.2019. (B) Moult of 12.V.2020. (C) Moult of 5.VII.2019. (D-F) Moult of 22.XI.2020. (G) Moult of VI.2019. (H-I) Moult of 13.II.2021. (J) Allotype; moult of 19.I.2019. (K-L) Moult of 14.II.2021. (M) Moult of 1.V.2021. (N-P) Moult of 23.X.2021. Arrows indicating paired lateral pockets of membranous uterus externus. Scale lines 1.0 mm.

Fig. 24. Liphistius platnicki sp. nov., vulval plates of ten females (all from exuviae), dorsal view (A, D, K, O), ventral view (B-C, E, H-J, L-N, P), anterior view (F), and isolated uterus externus (G). (A-B) Moult of 29.I.2021. (C, F) Moult of 10.III.2021 (al- lotype). (D-E) Moult of 25.IV.2020. (G, N) Moult of 14.X.2021. (H) Moult of 14.V.2020. (I) Moult of 20.XI.2021. (J) Moult of 13.X.2021. (K-L) Moult of 2.IV.2021. (M) Moult of 7.I.2021. (O-P) Moult of 3.X.2021. Arrows indicating paired lateral pockets of membranous uterus externus. Scale lines 1.0 mm

Fig. 23. Liphistius platnicki sp. nov., palps of three males; holotype (A-D, H-J, M), paratype matured 26.VII.2019 (E-G), paratype matured 8.VIII.2016 (K-L). (A) Left tarsus and palpal organ, ventral view. (B) Left paracymbium and distal margin of tibia, retrolateral view. (C) Distal part of left tibia, ventral view. (D) Same, retrolateral and slightly proximal view. (E) Left palpal organ, distal view (dorsal side up). (F) Dorsal part of left contrategulum and embolus proper, distal and slightly ventral view. (G) Dorsal part of left contrategulum, distal and slightly dorsal view. (H) Left palpal organ and distal margin of tarsus, prodorsal and slightly proximal view. (I) Same, dorsal and slightly proximal view. (J) Same, retrolateral and slightly distal view. (K) Left palpal organ, retrolateral view. (L) Tegulum and nearby parts of right palpal organ, same view. (M) Part of left palpal organ, retroventral view; elevated distal edge of tegulum indicated by short arrow, straight, salient proximal tegular edge indicated by long arrow. Scale lines 1.0 mm (A-B; C-D; E-M).