Heterometrus petersii (Thorell, 1876), 1604

Prendini, Lorenzo & Loria, Stephanie F., 2020, Systematic Revision Of The Asian Forest Scorpions (Heterometrinae Simon, 1879), Revised Suprageneric Classification Of Scorpionidae Latreille, 1802, And Revalidation Of Rugodentidae Bastawade Et Al., 2005, Bulletin of the American Museum of Natural History 2020 (442), pp. 1-480 : 269-279

Heterometrus petersii (Thorell, 1876)

Figures 10 , 158 , 178 , table 2

Palamnaeus petersii Thorell, 1876a: 13 , nomen nudum; 1876b: 214–217; 1888: 335, 338, 339, 414; 1889: 588–590 (part).

Palamnaeum petersii: Thorell, 1876b: 164 .

Palamnaeus petersi: Simon, 1884a: 361 .

Palamnaeus petersii: Pocock, 1892: 38–40 (part); Kovařík, 2004: 2.

Palamnaeus spinifer: Pocock, 1892: 39–42 (misidentification, part); 1894b: 316 (misidentification, part).

Scorpio longimanus: Kraepelin, 1894: 34–39 (misidentification, part).

Palmmaeus petersii: Kraepelin, 1894: 35 .

Heterometrus longimanus: Kraepelin, 1899: 108 , 111 (misidentification, part).

Palamnaeus oatesii Pocock, 1900a: 84 , 98, 99 (misidentification, part); Flower, 1901: 34, 35 (misidentification, part); Kovařík, 2004: 41, fig. 27; syn. nov.

Heterometrus (Scorpio) longimanus: Kraepelin, 1901: 271 (misidentification, part).

Heterometrus oatesi: Kraepelin, 1913: 167 (misidentification, part).

Heterometrus longimanus petersi: Giltay, 1931: 4 (part).

Heterometrus oatesii: Giltay, 1931: 4 (part); Takashima, 1945: 94 (part).

Heterometrus longimanus petersi: Fage, 1933: 27 (part).

Heterometrus longimanus petersii: Takashima, 1945: 90 , 91 (part); 1952: 34 (part).

Heterometrus (Heterometrus) spinifer: Couzijn, 1981: 73 , 94, 96, 173, 175, 182, 187, 189, 190, figs. 62, 66b (misidentification, part); Fet, 2000: 437, 438 (part).

Heterometrus (Heterometrus) spinifer spinifer: Couzijn, 1981: 19 , 32, 75–77, 83, 89–93, 167, 174, 191 table 7, figs. 2b, 17b, 18, 19, 63 (misidentification, part); Fet, 2000: 438 (part).

Heterometrus longimanus: Gopalakrishnakone et al., 1995: 456–458 , figs. 3, 4 (misidentification).

Heterometrus spinifer: Robert, 1999: 19 (misidentification, part); Prendini et al., 2003: 192, 193, 202, 203, 205, 208, 209, 218, 222, 252– 259, tables 3, 4, figs. 4–7, appendices 1, 2 (misidentification, part); Kovařík, 2004: 9, 11, 23, 34, 40–42, 51, 53, table 2, fig. 27 (misidentification, part); 2009: 35, 44, 48, table 1 (misidentification, part).

TYPE MATERIAL: Palamnaeus oatesii : Syntypes?: 3 ♂, 2 ♀, 1 juv. ♂ (BMNH), Hardwicke [examined]. SINGAPORE: P. oatesii : Syntype: [sex?] (BMNH), Singapore. Palamnaeus petersii : Holotype ♀ (NMG 91), “India Orientalis, ex Singapore,” 22.iv.1852, J.F. Lijbeck [examined].

Six specimens (3 ♂, 2 ♀, 1 juv. ♂) in the BMNH, apparently transferred to ethanol from dry storage, with the determination label “ Palamnaeus oatesii ,” but without any other data, appears to be all that remains of the type material of P. oatesii . Although no other specimens so labeled have been

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found at BMNH, it is impossible to know whether these specimens really are syntypes, as noted by Kovařík (2004). Therefore, following Kovařík (2004), a lectotype is not designated. Whereas the six putative syntypes remaining at the BMNH are conspecific with H. petersii , with which P. oatesii is therefore synonymized below, the following syntypes, listed in the original description of P. oatesii , all of which appear to be lost, were probably conspecific with other species of Heterometrus : [sex?] (BMNH), “Bengal”; H. laevigatus : [sex?] (BMNH), Siam [ Thailand], 1♀? (BMNH), Mergui, Burma [ Myanmar: Tanintharyi Reg.: Myeik Archipelago, 12°27′N 98°37′E], Anderson; H. thorellii : 1 ♂? (BMNH), Rangoon, Burma [ Myanmar: Yangon Reg.: Yangon, 16°47′N 96°10′E], Oates.

DIAGNOSIS: Heterometrus petersii may be separated from other species of Heterometrus , except H. spinifer , as follows. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the male and female of H. petersii whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the male and female of H. glaucus and H. thorellii , and the interocular surface is entirely smooth in the male of H. laevigatus and the male and female of H. laoticus . The carapace anterolateral, mediolateral and posterolateral surfaces of the female are granular in H. petersii but smooth or nearly so in H. laoticus , and the posterolateral surfaces of the female smooth or nearly so in H. longimanus . The pedipalps of the adult male are short, with femur length: posterior carapace width ratio (FL:PCW) <0.77 and femur length: carapace length ratio (FL:CL) <0.74 in H. petersii but long, with FL:PCW ≥ 0.80 and FL:CL ≥ 0.76 in H. glaucus , H. laevigatus , H. longimanus , and H. thorellii . The pedipalp chela manus promedian carina of the female is present and granular in H. petersii but absent or obsolete in H. laevigatus . The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is similar to the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male and female of H. petersii whereas the DMC–DSC is less than the DSC–DC in the male of H. glaucus , H. laevigatus , and H. thorellii , greater than the DSC–DC in the male of H. silenus , and less than the DSC–DC in the male and female of H. longimanus . The chela manus dorsal intercarinal surface is smooth or nearly so in H. petersii but shallowly reticulate in H. glaucus , H. laevigatus , and, often, H. thorellii . The second (subproximal) lobe of the chela movable finger of the adult male is more strongly developed than other lobes on the finger, with a correspondingly well-developed notch in the fixed finger in H. petersii but similar to or barely larger than other lobes on the finger in H. glaucus , H. longimanus , and H. thorellii . Macroseta st on the proventral surface of the basitarsus of leg I is setiform in H. petersii but spiniform in H. laevigatus and H. thorellii ; st on the proventral surface of the basitarsus of leg II is setiform in H. petersii but spiniform in H. glaucus , H. laevigatus , H. thorellii , and, usually, H. longimanus ; sb on the proventral surface of the basitarsus of leg III is spiniform in H. petersii but setiform in all other species except H. glaucus and H. spinifer . The lateral surfaces of mesosomal tergites I–VI in the female are granular (fig. 178) in H. petersii but smooth in H. laevigatus , H. laoticus , and H. thorellii . The ventrosubmedian and ventrolateral carinae of metasomal segments I–IV are granular or costate-granular on segment IV only and costate on segments I–III in H. petersii but granular on segments I–IV, II–IV (costate on I), or III and IV (costate on I and II) in H. laoticus and H. silenus . The ventral intercarinal surfaces of segments III and IV of the female are smooth in H. petersii but granular in H. laoticus . The telson is dark reddish brown and paler than metasomal segment V in H. petersii but blackish and as dark as segment V in H. laevigatus , H. laoticus , H. silenus , and H. thorellii . The width of the telson vesicle is approximately equal to or less than the width of metasomal segment V in the male of H. petersii but greater than the width of segment V in the male of H. glaucus and H. longimanus . No consistent morphological differences between H. petersii and H. spinifer were identified in the present investigation.

DISTRIBUTION: The full extent of the distribution of H. petersii is currently unknown as many specimens have doubtless been mistaken for H. spinifer . The only confirmed records of this cryptic species are from Singapore and the Malaysian island of Penang (Penang state) (fig. 158, table 2). Its distribution overlaps that of H. longimanus in Singapore. Heterometrus laevigatus and H. spinifer appear to be allopatric with H. petersii in Thailand and peninsular Malaysia, respectively.

ECOLOGY: Heterometrus petersii inhabits wet primary and secondary rainforest. Recently collected specimens were found doorkeeping at burrow entrances and sometimes under or in logs, or under stones with UV detection at night. Burrows were constructed in open ground or at the base of logs, roots, or stones in clayey soils. The habitat and habitus of this species are consistent with the fossorial, pelophilous ecomorphotype ( Prendini, 2001b). The following scorpions were found in sympatry: the buthid Lychas scutilus and two chaerilids, Chaerilus rectimanus Pocock, 1899 , and an undetermined species of Chaerilus .

CONSERVATION STATUS: Material originating from Singapore, and misidentified as H. spinifer , has been offered in the exotic pet trade.

REMARKS: As noted by Fet (2000: 437), Palamnaeus petersii and Palamnaeus silenus were introduced as replacement names for a species based on specimens originating from Cochin China [ Vietnam] in the MNHN. Simon (1872b: 97, 98) misidentified these specimens as Heterometrus megacephalus , itself a junior synonym of Srilankametrus indus , that was originally described as Buthus megacephalus C.L. Koch, 1836 (see below). Thorell (1876a, 1876b) realized that the specimens described as H. megacephalus by Simon (1872b) were misidentified and assigned them to a new species, P. petersii . However, Thorell (1876a, 1876b) did not examine the MNHN material described by Simon (1872b), and based P. petersii on a specimen from Singapore, in the NMG ( Thorell, 1876b: 216). Simon (1884a) agreed with Thorell (1876a, 1876b) that the MNHN material previously examined ( Simon, 1872b) from Cochin China [ Vietnam] was not conspecific with B. megacephalus . However, Simon (1884a: 361) suggested that P. petersii was a different species, occurring on the Malaysian Peninsula and adjacent islands, and possibly the female of H. longimanus , and was not conspecific with the MNHN specimens from Cochin China, which were therefore described as another new species, P. silenus :

L’espèce que nous avons décrite (Etudes sur les Scorp., p. 3, pl. 6, f. 2, 1872) sous le nom de Het. megacephalus C. Koch , est sans doute differente de Buthus megacephalus C. Koch , comme le Dr. Thorell l’a déjà fait observer; le Dr. Thorell propose pour cette espèce le nom de P. petersi (Et. Scorpiol. p. 140) mais la description qu’il en donne ne s’applique aucunement au megacephalus Simon et désigne une espèce beaucoup plus grande, propre à la presqu’île de la Malacca et aux iles voisines. Chez petersi en effet la main est de moitié plus longue que large, sensiblement atténuée jusqu’à la base des doigts, en dessus lisse brillante très finement réticulée-ruguese non ponctuée, tandis que chez megacephalus Simon , qui habite Siam, la Cochinchine, et l’Annam, la main est à peine plus longue que large, plus convexe, nullement réticulée mais parsemée de points enfoncés. P. petersi est peut-être le femelle de P. longimanus Herbst , nous avons reçu les deux formes de l’Ile Bintang; quant au megacephalus Simon (non C. Koch) nous proposons pour lui le nom nouveau de P. silenus . (Simon 1884: 361)

Pocock (1892: 38–40; 1894b: 316) initially synonymized P. petersii with H. spinifer but, later ( Pocock, 1900a: 84, 98), synonymized P. silenus with P. petersii , noting:

The name petersii was originally applied by Thorell

in 1876 to the Cochin China form which Simon in

1872 had erroneously described and figured as

megacephalus, C. Koch , and which he subsequently

in 1882 renamed silenus .

Although Pocock (1892, 1900a) apparently did not examine the types of either, his synonymy of P. silenus with P. petersii has persisted until today ( Couzijn, 1981; Kovařík, 2004, 2009). Fet (2000: 437) mistakenly credited the synonymy to Couzijn (1981: 89), who also continued to list P. silenus in synonymy with H. petersii , despite examining the types of both taxa, and listing the holotype of H. petersii among the material examined for H. spinifer .

The types of P. petersii , P. silenus , and H. longimanus and its junior synonyms were examined during the present investigation. Palamnaeus petersii was not found to be conspecific with H. longimanus , as suggested by Simon (1884a) or H. silenus , as assumed ever since the latter was synonymized by Pocock (1900a), justifying the revalidation of H. silenus , below.

In agreement with Pocock (1892: 38–40; 1894b: 316) and Couzijn (1981: 89), the adult female holotype of P. petersii , which originated from Singapore, was morphologically indistinguishable from the neotype of H. spinifer , and nontype material, previously identified as H. spinifer , from Singapore and elsewhere (e.g., the Malaysian mainland and the islands of Penang and Tioman). However, nontype material from Singapore and Penang, assessed to be conspecific with the holotype of P. petersii based on morphological similarity, was genetically more closely related to H. longimanus than to morphologically indistinguishable material of H. spinifer from the Malaysian mainland and Tioman (Loria and Prendini, in press), leading to the conclusion that H. petersii is a cryptic species.

Palamnaeus oatesii was described from an unknown series of specimens from Rangoon and Mergui, Burma [ Myanmar], Siam [ Thailand], Singapore, and “Bengal” although Pocock (1900a: 99) doubted the accuracy of the latter. Pocock (1900a: 98) listed P. spinifer and P. petersii among the synonyms of P. oatesii . Couzijn (1981: 89) synonymized P. oatesii with H. spinifer , a decision upheld by Fet (2000) and Kovařík (2004, 2009). Reexamination of putative syntypes in the BMNH during the present investigation, taken together with the original description ( Pocock, 1900a: 98, 99), which mentions characters such as “vesicle generally reddish yellow and much paler … than the segments of the tail” and “ ♂ … with secondary sexual characters poorly developed” confirmed the morphological similarity of P. oatesii and H. spinifer which formed the basis of Couzijn’s (1981) synonymy. The morphology is also consistent with H. petersii , however, in which the males closely resemble the females. Pocock’s (1900a) original series evidently included several species, e.g., H. laevigatus from Mergui, H. petersii from Singapore, and H. thorellii from Rangoon. None of the type localities cited by Pocock (1900a) matches the known distribution of H. spinifer , however, and the sexually dimorphic species, H. laevigatus and H. thorellii , do not match the original description or the BMNH specimens, in which the males resemble the females. Therefore, by a process of elimination, and absent evident to the contrary, the following new synonym is proposed: Palamnaeus oatesii Pocock, 1900 = Heterometrus petersii (Thorell, 1876) , syn. nov.

MATERIAL EXAMINED: MALAYSIA: Penang: Penang [Penang Island, 05°24′N 100°14′E], i.1862, M.D. Parker, 1 ♂ (MCZ) ; Pulo Penang [Pulau Penang ( Penang Island )], vii.1860, W.H.A. Putnam, 1 ♂, 1 ♀ (MCZ) ; Penang Island, Bukit Bendera / Penang Hill , Moniot Road , 05°25′N 100°15′E, and grounds of Monkey Cup Cafe, 05°25′N 100°16′E, 16–27.x.2017, L. Esposito and S.F. Loria, night collecting with UV, 3 ♂, 2 ♀ (CAS 9080783), 1 ♂ [leg] (AMCC [LP 16458], 1 ♀ [leg] (AMCC [LP 16459]). SINGAPORE: Singapore [01°22′N 103°48′E], 1889, Dr. Hamlin, 1 ♂ (MCZ 14970), 1921, 1 subad. ♂ (LKC ZRC-ENT 7964), xii.1921, 1 subad. ♂ (LKC ZRC-ENT 7960), 1 subad. ♀ (LKC ZRC-ENT 7963), xi.2004, captive bred, 1 subad. ♀, 2 juv. ♀ (LKC ZRC-ARA 677) ; 8th Mile Old Upper Thomson Road /6th Mile Thomson Road [01°23′N 103°49′E], 16–17.ii.1963, K.O. Yee, 1 juv. ♀ (LKC ZRC- ENT 7229 [ZRC-ARA 731]) ; Gardens [Singapore Botanic Gardens, 01°19′N 103°49′E], 7.viii.1921, 1 ♂ (LKC ZRC-ENT 7961), xii.1921, 1 ♂ (LKC ZRC-ENT 7962), Botanical Gardens , 13.iv.1960, S.C. Keong, 1 ♀ (LKC ZRC-ARA 675), Botanic Gardens, National Orchid Garden, 2010, K. Lin, 1 ♂ (LKC ZRC-ENT 7239) ; Bukit Kallang , track near [01°22′N 103°49′E], 23.v.1994, C.H. Yeow and J. Choo P.S., 1 subad. ♀ (LKC ZRC-ARA 676) ; [probably Mandai , 01°25′N 103°47′E], ix.1998, C.K. Wee, 1 ♀ (AMNH [LP 1604]), legs (AMCC 101699 [LP 1604]) ; Mandai [01°25′N 103°47′E], 1998, C.K. Wee, 1 juv. ♀ (LKC RC-ARA 679) ; SADA Hill [Singapore Air Defence Artillery, 01°26′N 103°50′E], 1 ♂ (LKC ZRC-ENT 7965) .

Figures

FIGURE 10. Phylogeny of Asian forest scorpions (Scorpionidae: Heterometrinae Simon, 1879, stat. nov.), obtained by simultaneous phylogenetic analysis of 186 morphological characters and 4188 aligned DNA nucleotides from three mitochondrial and two nuclear gene loci (Loria and Prendini, in press). Maximum likelihood tree with unambiguous morphological synapomorphies optimized. Black circles indicate uniquely derived apomorphic states, white circles parallel derivations of apomorphic states. Numbers above circles indicate characters, numbers below indicate states (appendix 1).

FIGURE 158. Map of Southeast Asia, plotting known locality records of five species of Heterometrus Ehrenberg, 1828, based on material examined, and verified records from virtual museums and the literature: Heterometrus laevigatus (Thorell, 1876), comb. nov. et stat. rev.; Heterometrus longimanus (Herbst, 1800); Heterometrus petersii (Thorell, 1876); Heterometrus silenus (Simon, 1884), comb. nov. et stat. rev.; Heterometrus thorellii (Pocock, 1892).

FIGURE 178. Heterometrus petersii (Thorell, 1876), holotype ♀ (NMG 91), habitus: A. dorsal and B. ventral aspects. Scale bar = 10 mm.