Euscorpius

Occasional Publications in Scorpiology

A Review of the Genus Heterometrus Ehrenberg, 1828, with

Descriptions of Seven New Species (Scorpiones, Scorpionidae)

František Kovařík

December 2004 – No. 15

 Euscorpius
Occasional Publications in Scorpiology
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Publication date: 8 December 2004

 Euscorpius — Occasional Publications in Scorpiology. 2004, No. 15

A review of the genus Heterometrus Ehrenberg, 1828, with

descriptions of seven new species (Scorpiones, Scorpionidae)
František Kovařík1
1
P.O. Box 27, CZ-145 01 Praha 45, Czech Republic

Summary

The genus Heterometrus is revised and a key is presented to all species except H. tristis. Subgenera
Chersonesometrus Couzijn, 1978, Gigantometrus Couzijn, 1978, Javanimetrus Couzijn, 1981 and Srilankametrus
Couzijn, 1981 are synonymized with the subgenus Heterometrus Ehrenberg, 1828. Seven new species of
Heterometrus are described: H. beccaloniae sp. n., H. mysorensis sp. n., H. rolciki sp. n., and H. ubicki sp. n. from
India, H. nepalensis sp. n., from Nepal, H. sejnai sp. n. from Thailand, and H. cimrmani sp. n. from Thailand and
Vietnam. The following species and subspecies are synonymized: H. (Chersonesometrus) fastigiosus Couzijn, 1981
with H. bengalensis (C. L. Koch, 1841); H. (Srilankametrus) indus laevitensus Couzijn, 1981 with H. gravimanus
(Pocock, 1894); H. liophysa madoerensis Kopstein, 1921 and H. laevifrons Roewer, 1943 with H. liophysa (Thorell,
1888); H. (H.) longimanus belitungensis Couzijn, 1981 and H. (H.) longimanus tarawakanensis Couzijn, 1981 with
H. longimanus (Herbst, 1800); H. (Chersonesometrus) granulomanus Couzijn, 1981 with H. madraspatensis
Pocock, 1900; Palamnaeus phipsoni collinus Pocock, 1900 with H. phipsoni (Pocock, 1893); H. (H.)
malapuramensis Tikader & Bastawade, 1983 with H. scaber (Thorell, 1876); H. (Gigantometrus) swammerdami
titanicus Couzijn, 1981 with H. swammerdami Simon, 1872; and H. (Chersonesometrus) pelekomanus Couzijn,
1981 with H. wroughtoni (Pocock, 1899). H. thorellii (Pocock, 1892) comb. n. is recognized as valid species.
Lectotypes are designated for H. xanthopus (Pocock, 1897) and for H. bengalensis (C. L. Koch, 1841), which
invalidate Couzijn’s (1981) neotype designation. H. bengalensis (C. L. Koch, 1841) is for the first time recorded
from Orissa, H. fulvipes (C. L. Koch, 1837) from Mysore, Uttar Pradesh, and West Bengal, H. kanaraensis (Pocock,
1900) from Goa, H. phipsoni (Pocock, 1893) from Orissa, H. scaber (Thorell, 1876) from Mysore and Pondichery,
H. swammerdami Simon, 1872 from Mysore, and H. wroughtoni (Pocock, 1899) from Mysore (all states of India).

Introduction his opinion is concurred with only as to the importance

of the form and surface of the chela of pedipalps
Three studies can be considered essential for our present (granulation, punctation, carinae) and to some extent the
knowledge of the genus Heterometrus. The first granulation of the carapace. Of the characters neglected
unquestionably is a series of papers by Pocock (1892– by Couzijn,s the greatest emphasis is placed on sexual
1900), who described the most species and established dimorphism, but attention is paid also to the color of the
the basic criteria for their recognition. The second is the legs and development of carinae on the metasomal
work of Couzijn (1981), who based his analysis on as segments. For the sake of brevity, descriptive trivia of no
many as 131 characters. And the third is Tikader & direct value to species recognition are omitted. Tikader
Bastawade (1983), who treated the Indian species in & Bastawade (1983) attempted to differentiate species of
detail and drew attention to Couzijn’s errors by re- Heterometrus by means of trichobotriotaxy, which
validating several species synonymized by him (see also unfortunately cannot be done in this genus.
Fet, 2003). Although Couzijn’s extensive character set
gives the impression of a definitive work, at a closer ABBREVIATIONS. The institutional abbreviations listed
examination the inclusion of many of his characters and below and used throughout are mostly after Arnett,
the weight given to many others invite considerable Samuelson & Nishida (1993).
doubt. Surprisingly, Couzijn applied virtually none of
the characters established by Pocock. The approach used BMNH – The Natural History Museum, London, United
in this review differs markedly from that of Couzijn. Kingdom;
Most of his morphometric characters are abandoned CASC – California Academy of Sciences, San
because of their substantial intraspecific variation, and Francisco, California, USA;
2 Euscorpius — 2004, No. 15

FKCP – František Kovařík Collection, Praha, Czech = Heterometrus (Chersonesometrus) Couzijn, 1978: 330
Republic; (TS: Buthus fulvipes C. L. Koch, 1837); Couzijn, 1981:
HNHM – Hungarian Natural History Museum, 131; Tikader & Bastawade, 1983: 573; Fet, 2000: 438.
Budapest, Hungary; Syn. n.
MCSN – Museo Civico de Storia Naturale ”Giacomo = Heterometrus (Gigantometrus) Couzijn, 1978: 330;
Doria”, Genoa, Italy; (TS: Heterometrus swammerdami Simon, 1872);
MNHN – Muséum National d´Histoire Naturelle, Paris, Couzijn, 1981: 159; Tikader & Bastawade, 1983: 561;
France; Fet, 2000: 443. Syn. n.
MZUF – Museo Zoologico de ”La Specola”, Firenze, = Heterometrus (Javanimetrus) Couzijn, 1981: 124 (TS:
Italy; Buthus cyaneus C. L. Koch, 1836); Fet, 2000: 444. Syn.
NHRS – Naturhistoriska Riksmuseet, Stockholm, n.
Sweden. = Heterometrus (Srilankametrus) Couzijn, 1981: 120
NMPC – National Museum (Natural History), Praha, (TS: Scorpio indus DeGeer, 1778); Fet, 2000: 446. Syn.
Czech Republic; n.
NZSI – National Collection, Zoological Survey of India,
Calcutta, India; TYPE SPECIES. Buthus (Heterometrus) spinifer
SMFD – Forschungsinstitut und Naturmuseum Ehrenberg, 1828, by subsequent designation (Karsch,
Senckenberg, Frankfurt am Main, Germany; 1879: 20).
SOFM – National Museum of Natural History, Sofia,
Bulgaria; DIAGNOSIS: Pedipalp femur with three trichobothria, of
ZMHB– Museum für Naturkunde der Humboldt- them only one on internal surface. Patella of pedipalp
Universität zu Berlin, Germany; with 19 trichobothria, three on ventral and 13 on
ZMUH – Zoologisches Institut und Zoologisches external surface. Chela of pedipalp with 26 trich-
Museum, Universität Hamburg, Germany. obothria. Retrolateral pedal spurs absent. Lateroapical
margins of tarsi produced into rounded lobes.
Other abbreviations are: ♂: male; ♀: female; im.: Metasomal segments I to IV with paired ventral
immature; juv.: juvenile. submedian carinae. Stridulatory organ located on
opposing surfaces of pedipalp coxa and first leg. Total
Heterometrus Ehrenberg, 1828 length 60 to 180 mm.
(Figs. 1–7)
COMMENTS. Couzijn (1978, 1981) described four
Buthus (Heterometrus) Ehrenberg in Hemprich & subgenera but gave no characters that would permit to
Ehrenberg, 1828: pl. 1, figs 1–2 (part, only fig. 2); unequivocally distinguish them from the nominotypical
Hemprich & Ehrenberg, 1829: 351; Hemprich & subgenus. The invalidity of his subgenera is also clear
Ehrenberg, 1831: 2. from the below listed synonymizations of species across
Buthus: C. L. Koch, 1836: 73; C. L. Koch, 1837: 36; C. them, and I therefore regard all of them as synonyms.
L. Koch, 1837: 25; C. L. Koch, 1837: 45 (in part); The only subgenus that could possibly be upheld is
Agassiz, 1846: 7 (in part); C. L. Koch, 1841: 1 (in part); Gigantometrus, which comprises closely related species
C. L. Koch, 1850: 87 (in part). easily distinguishable from others. However, the genus
Scorpio Linné, 1758: 624 (in part). Heterometrus contains more of such groups (e.g. H.
Heterometrus: Ausserer, 1880: 466; Kraepelin, 1905: madraspatensis and H. gravimanus, or H. petersii and
344; Kraepelin, 1913: 165; Kopstein, 1921: 128; H. laoticus), and in my opinion they have the character
Werner, 1934: 277; Couzijn, 1978: 327; Couzijn, 1981: of species groups rather than of subgenera.
73; Tikader & Bastawade, 1983: 518; Sissom, 1990: I have found two characters in Heterometrus that
136; Fet, 2000: 431; Prendini, 2000: 44; Soleglad & Fet, could serve to justify subgeneric divisions. One of them
2003: 88. is sexual dimorphism, which Couzijn for some reason
Pandinus: Thorell, 1876b: 199 (in part); Karsch, 1892: did not include in his analysis. However, after
307. examination of a majority of the species it has become
= Centrurus Ehrenberg in Hemprich & Ehrenberg, 1829: clear that this character evolved in the genus more than
350 (TS: Centrurus galbineus C. L. Koch, 1838); C. L. once, because it differentiates otherwise closely related
Koch, 1837: 38; C. L. Koch, 1838: 110 (see Fet, 2000: species that definitely cannot be placed in separate
431). subgenera. As examples can serve H. liophysa, H.
= Palamnaeus Thorell, 1876a: 13 (TS: Palamnaeus longimanus, H. spinifer and H. cimrmani sp. n., whose
petersii Thorell, 1876); Karsch, 1892: 307; Pocock, females and juveniles are virtually indistinguishable and
1897: 114; Pocock, 1896: 77; Pocock, 1900: 84 (syn. by the species can be reliably identified only on males. The
Karsch, 1879: 20; Kraepelin, 1899: 107). other character is the genital operculum which comes in
 Kovařík: Review of Genus Heterometrus 3

Figures 1–7: Heterometrus wroughtoni (Pocock, 1899), specimen labeled as paratype of Heterometrus (Chersonesometrus)
pelekomanus Couzijn, 1981 syn. n. In Figs. 1 to 3 the first capital letters denote trichobothria situated on the manus, and the first
lower-case letters denote those situated on the fixed finger of pedipalp. Figs. 4 to 6 show the distribution of trichobothria on the
patella of pedipalp. Fig. 7 shows the distribution of trichobothria on the femur of pedipalp. Explanation: First letters: D, dorsal, E,
external, I, internal, V, ventral. Second or second plus third letters: b, basal, sb, suprabasal, m, medial, st, subterminal, t, terminal,
v, ventral. Numerals distinguish individual trichobothria of the same classification. Designation and description of trichobothria
after Vachon (1974). Morphological terminology after Stahnke (1970).
4 Euscorpius — 2004, No. 15

two basic shapes, but they cross some of the species- ETYMOLOGY. Named after Janet Beccaloni of the
group boundaries. I therefore believe that subgeneric Natural History Museum, London, in appreciation of her
divisions within the genus Heterometrus are not kind help.
justified.
DIAGNOSIS. Adult female holotype 115 mm long. Color
uniformly black, only telson reddish yellow. Pectinal
Heterometrus barberi (Pocock, 1900)
teeth number 14–16. Chela of pedipalp length to width
(Fig. 8) ratio 2.38 (holotype). Dorsal surface of manus granulate
and/or tuberculate, with carinae. Patella of pedipalp
Palamnaeus barberi Pocock, 1900: 95. granulate, without pronounced internal tubercle and with
Heterometrus barberi: Takashima, 1945: 94. five carinae. Femur of pedipalp unusually wide,
Heterometrus (Chersonesometrus) barberi: Tikader & granulated and convex. Carapace with smooth disc and
Bastawade, 1983: 614; Fet, 2000: 441. granulose margins. Mesosoma smooth. Telson hirsute,
Heterometrus (Chersonesometrus) phipsoni phipsoni: bulbous (in adults), vesicle longer than aculeus. Genital
Couzijn, 1981: 149 (in part). operculum large, in female about as wide as or slightly
wider than long.
TYPE LOCALITY AND TYPE REPOSITORY. Tinnevelly in S.
India; BMNH. DESCRIPTION: The adult female holotype is 115 mm
TYPE MATERIAL EXAMINED. India, Tinnevelly District, long. Measurements of the carapace, telson, segments of
in the western Gháts, Kannikatti evergreen forest, 2500 the metasoma and segments of the pedipalps, and
ft., 1♀, holotype, leg. C. A. Barber, BMNH No. numbers of pectinal teeth are given in Table 1. The male
1899.9.24.1. No other material. is unknown.
COLORATION: The color is uniformly black, only the
DIAGNOSIS. Adult holotype female 124 mm long. Base telson is reddish yellow (adults) or yellow (juveniles).
color uniformly black, only manus of pedipalps orangish Sternites and pectens are lighter-colored (brownish).
yellow. Pectinal teeth number 11–12. Chela not lobiform, MESOSOMA AND CARAPACE: The mesosoma lacks
its length to width ratio equals 2.2. Entire manus rugate carinae, is smooth, and the carapace is either entirely
(merging blunt granules) and without carinae. Patella of granulate or with the disc smooth. Sternites are smooth,
pedipalps without pronouced internal tubercle. Carapace entirely without granules, only the seventh sternite may
with disc smooth, at margins granulate. Spine formula of bear four smooth carinae. Pectinal teeth number 14–16.
third legs 4:6, of fourth legs 4:6–7. Telson bulbous, METASOMA AND TELSON: The metasoma is smooth. The
vesicle longer than aculeus. first to fourth segments bear eight smooth carinae. The
fifth segment bears seven carinae. The ventral carinae of
COMMENTS. Couzijn (1981: 149) considered this segment consist of pointed granules, whereas the
Palamnaeus barberi Pocock, 1900 a synonym of H. dorsal carinae are smooth. The telson is bulbous and
phipsoni (Pocock, 1893). However, these two species hirsute, dorsally without granules and ventrally with four
are very different (see diagnosis and key), which was rows of granules. The vesicle is longer than the aculeus,
pointed out already by Tikader & Bastawade (1983: in the adult female bulbous and in the juveniles elongate.
614) who regarded Heterometrus (Chersonesometrus) PEDIPALPS: The femur has three granulose carinae and
barberi (Pocock, 1900) as a valid species. The BMNH its dorsal surface is convex and granulate. The patella is
register lists the holotype of Palamnaeus barberi granulose, ventrally tuberculate, with five granulate
Pocock, 1900 as Palamnaeus flavimanus Pocock, 1900. carinae, and on the internal surface with scattered
pointed minute granules that do not form a conspicuous
internal tubercle. The dorsal surface of the manus is
DISTRIBUTION. India: Tamil Nadu (Pocock, 1900: 96). granulate or tuberculate, with six carinae, which are in
part smooth and in part granulate. The chela is sparsely
Heterometrus beccaloniae sp. n. hirsute and its external surface is granulate and
(Fig. 9) tuberculate, with two granulate carinae and a smooth
space between them. The movable finger bears six
straight rows of granules and internal and external
TYPE LOCALITY AND TYPE REPOSITORY. India, Tamil
granules.
Nadu, Javadu Hills, Vellore Dist., Kavalore; FKCP. LEGS: The legs bear solitary long and short setae. The
TYPE MATERIAL. India, Tamil Nadu, Javadu Hills, third and fourth leg spine formula is 5–6:6–7.
Vellore Dist., Kavalore, 2000 ft. TRSN coll., V.2003,
1♀ (holotype); Tamil Nadu, Nilgiri hills, Mudumalai n. AFFINITIES. The described features distinguish H.
p., 6.V.2000, 1im.7juvs. (paratypes), leg. J. Rolčík. beccaloniae sp. n. from all other species of the genus.
 Kovařík: Review of Genus Heterometrus 5

Figure 8: Heterometrus barberi, dorsal view, female holotype.

6 Euscorpius — 2004, No. 15

Figure 9: Heterometrus beccaloniae, sp. nov., dorsal view, female holotype.

 Kovařík: Review of Genus Heterometrus 7

H. becaloniae H. cimrmani H. mysorensis H. nepalensis H. rolciki H.sejnai H. ubicki

female male female male male male female male male female
HT HT AT HT HT HT AT HT HT AT
Total length 115 111.3 98 118.7 83 100 91 78 89 77
Carapace length 18.3 17.4 16.0 17.6 13.5 12.5 12.2 12.8 12.7 13.1
width 19.0 16.0 14.7 17.8 12.0 11.4 13.5 12.5 13.1 13.1
Metasoma
and telson length 57.8 61.2 52.0 63.7 43.0 47.0 47.0 42.7 46.8 40.9
segment I length 7.0 6.5 5.7 6.8 5.0 5.6 5.1 4.7 4.7 4.5
width 7.7 7.3 6.8 7.1 6.2 6.0 6.1 6.4 5.8 5.6
segment II length 7.4 7.0 6.2 7.7 5.5 6.1 6.1 5.5 5.5 4.8
width 6.9 7.1 6.4 6.7 6.0 5.7 5.6 5.8 5.6 5.2
segment III length 8.0 8.1 7.0 8.2 6.0 6.5 6.4 6.0 6.0 5.3
width 6.6 6.5 5.9 6.3 5.7 5.2 5.0 5.5 5.2 4.9
segment IV length 9.1 9.8 8.6 9.6 6.7 7.5 7.5 6.8 6.9 6.3
width 5.7 5.8 5.3 5.9 5.3 4.5 4.7 5.1 4.6 4.3
segment V length 13.0 14.1 12.4 13.6 9.5 10.4 10.3 9.8 9.7 9.0
width 5.1 5.3 4.8 5.2 4.5 4.4 4.2 4.6 4.1 3.9
telson length 13.3 15.7 12.0 13.8 9.6 10.5 10.6 9.7 11.0 8.8
Pedipalp
femur length 15.9 16.1 11.8 19.0 10.2 9.7 9.6 11.6 14.7 10.5
width 7.9 5.8 5.6 6.7 4.7 4.3 4.4 4.7 4.5 4.4
patella length 15.7 18.6 13.7 20.2 11.9 10.6 10.4 11.9 13.3 11.3
width 7.6 6.2 5.4 6.9 4.8 4.9 4.9 4.8 4.8 4.8
tibia length 32.5 35.5 28.5 35.5 22.2 20.2 20.1 22.6 25.7 20.8
width 13.6 12.5 12.1 12.5 9.8 8.7 8.6 10.2 7.6 9.2
finger mov. length 18.5 20.0 18.4 19.1 12.9 12.6 12.2 13.2 13.6 10.8

Pectinal teeth 15:16 16:17 15:16 17:16 14:15 11:11 10:10 13:14 16: ? 14: ?

Table 1: Measurements (in milimetres) of type specimens of seven new Heterometrus species.

They are recounted in the key below. H. beccaloniae sp. Heterometrus (Heterometrus) bengalensis: Couzijn,
n. is close to H. wroughtoni (Pocock, 1899) but has a 1981: 117 (in part); Tikader & Bastawade, 1983: 539;
less lobiform, more granulose and more tuberculate Kovařík, 1998: 137 (in part); Fet, 2000: 432 (in part);
chela. H. beccaloniae sp. n. is unique in the genus in Kovařík, 2002: 17.
having a very wide femur of pedipalp, which is convex = Palamnaeus costimanus glaucus Thorell, 1876b: 219
and only twice longer than wide. (syn. by Couzijn, 1981: 117).
= Heterometrus (Chersonesometrus) fastigiosus
Heterometrus bengalensis (C. L. Koch, 1841) Couzijn, 1981: 140; Tikader & Bastawade, 1983: 587;
(Fig. 10) Kovařík, 1998: 136; Fet, 2000: 438; ? Bastawade, 2002:
295; Kovařík, 2002: 17. Syn. n.
Buthus bengalensis C. L. Koch, 1841: 3, fig. 696; C. L.
Koch, 1850: 88. TYPE LOCALITY AND TYPE REPOSITORY. Ostindien,
Palamnaeus bengalensis: Pocock, 1892: 39; Pocock, Bengalen; ZMHB.
1900: 94; Kanungo, 1954: 536; Moritz & Fischer, 1980: TYPE MATERIAL EXAMINED. India, Bengal, 1♂
310. (lectotype hereby designated), ZMHB No. 70/2, 1♂
Scorpio bengalensis: Pocock, 1893: 310; Pocock, 1894: (paralectotype of Buthus bengalensis and paratype of
72; Kraepelin, 1895: 51. Heterometrus fastigiosus), ZMHB No. 70/1, 1♀
Heterometrus bengalensis: Kraepelin, 1899: 114; (paralectotype of Buthus bengalensis and paratype of
Werner, 1902: 601; Kraepelin, 1913: 166; Borelli, 1915: Heterometrus fastigiosus), ZMHB No. 70/3, 1♂
463; Lampe, 1918: 198; Roewer, 1943: 228; Takashima, (paralectotype of Buthus bengalensis), ZMHB No. 70/4,
1945: 93; Minnocci, 1974: 38; Brignoli, 1985: 415; 1♂ (paralectotype of Buthus bengalensis), ZMHB No.
Hjelle, 1990: 51; McCormick & Polis, 1990: 298. 70/5, 1♀ (paralectotype of Buthus bengalensis), ZMHB
Heterometrus (Scorpio) bengalensis: Kraepelin, 1901: No. 70/6, 1im. ♀ (paralectotype of Buthus bengalensis),
271. ZMHB No. 70/7; Assam, 1♂1♀1juv. (holotype and
8 Euscorpius — 2004, No. 15

Figure 10: Heterometrus bengalensis, dorsal view, male lectotype.

 Kovařík: Review of Genus Heterometrus 9

paratypes Heterometrus (Chersonesometrus) fastigiosus DISTRIBUTION. India: West Bengal (C. L. Koch, 1841: 5;
Couzijn, 1981), SMFD No. 8886/228. Pocock, 1894: 72), Orissa (first report). I presume also
occurrence in Bangladesh and other states of India.
OTHER MATERIAL EXAMINED. India, Mizgapur Banaras, However, the records listed by Tikader & Bastawade
1♂, CASC; Orissa, Nagpur-Keonjahr Dist., Daitari, (1983: 543) from Uttar Pradesh, Madhya Pradesh,
II.1974, 1♂1♀, FKCP; West Bengal, Midnapore, Assam, Meghalaya and Maharashtra are due to
19.III.1960, 1♂, leg. C. L. Kau, CASC; 30 m from confusion with other species doubtful and require
Kharpur, 28.IX.1960, 1juv., 20.VI.1960, 1♂1juv., verification.
28.IX.1960, 2♂2♀1juv., 5.X.1960, 2♀1im., 6.X.1960,
1im., 1.III.1961, 1juv., 24.III.1961, 1juv., 9.V.1961, Heterometrus cimrmani sp. n.
2juvs., leg. J. Singh, CASC; 30 m from Kharpur, (Fig. 11)
17.II.1961, 1♀, 27.II.1961, 1♀, leg. C. L. Kau, CASC; 5
m from Kharpur, roots of Polash tree, 9.V.1961, 1juv., Heterometrus (Heterometrus) spinifer: Kovařík, 1998:
leg. B. Singh, CASC; 4.III.1961, 1juv., VI.1961, 137 (in part); Fet, 2000: 438 (in part).
1♂2juvs., 18.VI.1962, 1♀ (im.) , CASC; before Heterometrus (Heterometrus) spinifer spinifer (in part):
13.III.1968, Zafer´s Export Centre, 5♂5♀2ims. (labeled ? Couzijn, 1981: 89; Kovařík, 1995: 203; ? Le Xuan Hue
as Heterometrus fastigiosus and Heterometrus et al., 1998: 7.
bengalensis), CASC, 14♂20♀ 20ims.7juvs. , CASC,
1♂1♀1im., FKCP; Calcutta, 17.VIII.1965, 1♀, leg. P. TYPE LOCALITY AND TYPE REPOSITORY. Thailand, Trang
K. Sina, CASC. env.; FKCP.

TYPE MATERIAL. Thailand, Trang env., 1♂1♀ (holotype

DIAGNOSIS. Adults 95–115 mm long. Color of adults
and allotype), FKCP. Holotype and allotype were bred
uniformly reddish brown to light brown. Pectinal teeth
by the author; the holotype died on 16.V.1992, and the
number 14–17 in both sexes. Male with chela, femur and
allotypic paratype gave birth, on 29.VIII.1992, to 18
patella of pedipalp relatively narrower and longer than in
juvs. of which paratypes are 2juvs. after 1st ecdysis,
female. Chela not lobiform, its length to width ratio
1juv. after 2nd ecdysis, 1juv. after 3rd ecdysis, 1juv.
about 3.0 in male and 2.2 in female. Much of manus
after 4th ecdysis, 2ims. after 5th ecdysis, and 4♀ after
sparsely tuberculate, uneven, but without discrete
6th ecdysis, reared by F. Kovařík, FKCP; Trang env.,
granules and carinae, especially in male. Patella of
1♂, 1980, FKCP; Khao Chong (Trang), X.1998, 4♀
pedipalp without pronounced internal tubercle. Carapace
(paratypes), leg. V. Šejna, FKCP; Trang-Kao Chong,
smooth, with granules only at margins. Third and fourth
1juv., 10.IX.1993, leg. M. Veselý, FKCP; Thaleban, 30
leg spine formula 4–5:5–6. Telson hirsute, bulbous,
km SW near Satun, 1♂ (paratype), 20.IX.1993, FKCP;
vesicle longer than aculeus.
Ban Huai Phan env., pr. Nakhon Si, Thammarat, 7–
12.VI.1996, 1♀ (im.) (paratype), leg. F. Kantner, FKCP;
COMMENTS. This species was based on seven syntypes 20 km E. of Trang, 1999, 1♀ (paratype – juv. born on 3-
deposited at ZMHB. Couzijn (1981: 120 and 142) stated IX-1999), leg. Komzík, FKCP. Vietnam, Saigon, 1940,
that these syntypes were destroyed during WW2, 1♀ (paratype), FKCP.
although he saw two of them and speculated that in
reality they were not the syntypes of Buthus bengalensis ETYMOLOGY. Named after Jára Cimrman, a well known
but a different species which he described as Czech renaissance man. Jára Cimrman bred large
Heterometrus fastigiosus. He thus designated the two Heterometrus from Thailand and Malaysia and in his log
examined syntypes of Buthus bengalensis as paratypes noted that they must be two species with different sexual
of Heterometrus fastigiosus, and for Heterometrus dimorphism (Heterometrus spinifer and the species
bengalensis designated a neotype deposited at MNHN. I described here). Regrettably, he has not published the
consider this taxonomic move entirely invalid and observation.
incomprehensible. All seven syntypes of Buthus
bengalensis are preserved at ZMHB and I hereby DIAGNOSIS. Adults 90–115 mm long. Base color of
designate one of them as lectotype to assure the stability adults uniformly black to greenish black. Pectinal teeth
of the species. In accordance with ICZN Article 75.8, it number 15–18 in both sexes. Male has longer but not
invalidates Couzijn‘s designation of the neotype. I narrower femur and patella of pedipalp, and longer and
examined the holotype and paratypes of Heterometrus narrower chela of pedipalp. Chela slightly lobiform, its
fastigiosus from SMFD and found them to be the same length to width ratio 2.8–3.0 in male and 2.1–2.4 in
species. Therefore, Heterometrus fastigiosus Couzijn, female. Male fixed fingers of pedipalp shorter than
1981 is a synonym of Heterometrus bengalensis (C. L. manus. Much of manus smooth, with punctae arranged in
Koch, 1841). rows. Patella of pedipalp with pronounced internal
10 Euscorpius — 2004, No. 15

Figure 11: Heterometrus cimrmani, sp. nov., dorsal view, male holotype.
 Kovařík: Review of Genus Heterometrus 11

tubercle. Carapace with disc smooth and margins Scorpio indicus: Pocock, 1893: 311 (in part); Pocock,
granulate. Telson hirsute, elongate, vesicle longer than 1894: 95; (in part) Kraepelin, 1895: 53; ? Kraepelin,
aculeus. 1898: 439.
Pandinus indicus: Thorell, 1876b: 208; Karsch, 1887:
DESCRIPTION: The adult male holotype is 111.3 mm 68; Thorell, 1888: 412 (in part); Thorell, 1893: 380 (in
long, other adults are 90–115 mm long. Measurements
part); Pavesi, 1898: 1.
of the carapace, telson, segments of the metasoma and
segments of the pedipalps, and numbers of pectinal teeth ? Scorpio afer (in part) Linné, 1758: 624; Latreille,
are given in Table 1. Sexual dimorphism is expressed in 1803: 267; (syn. by Thorell, 1893: 380).
the length of the femur, patella and chela of pedipalp, Buthus (Heterometrus) afer: Sundevall, 1833: 32.
and also in the width of the chela. Scorpio (Buthus) afer: Gervais, 1844b: 60 (in part).
COLORATION: The color is uniformly black to greenish Scorpio afer: Lönnberg, 1897: 178 (in part); Lönnberg,
black. 1898: 85 (in part).
MESOSOMA AND CARAPACE: The mesosoma lacks Heterometrus cyaneus: Simon, 1872: 98; Kraepelin,
carinae, is smooth, and the carapace has disc smooth and 1899: 115; Werner, 1902: 602; Domaniewski, 1913:
margins granulate. Sternites are smooth, entirely without 251; Borelli, 1915: 463; Lampe, 1918: 198; Kopstein,
granules, only the seventh sternite may bear smooth 1921: 133; Kopstein, 1923: 185; Kopstein, 1926: 111;
carinae. Pectinal teeth number 15–18 in both sexes. Kopstein, 1927: 102; Giltay, 1931: 6; Kästner, 1931:
METASOMA AND TELSON: The metasoma is smooth and 738; Werner, 1932: 576; Werner, 1934: 278; Giltay,
sparsely hirsute. The first to fourth segments bear eight 1935: 1; Fage, 1936: 181; Werner, 1936: 186; Fage,
carinae, of which the ventral are smooth and the dorsal 1944: 71; Takashima, 1945: 93; Roewer, 1943: 226;
consist of rounded or pointed granules. The fifth Bücherl, 1959: 269; Rosin & Shulov, 1963: 568;
segment bears five complete carinae and two additional Sreenivasa-Reddy, 1968: 760.
lateral carinae developed on only two-thirds of the Heterometrus (Scorpio) cyaneus: Kraepelin, 1901: 271.
segment. All carinae of this segment consist of minute Palamnaeus cyaneus: Kraepelin, 1898: 439.
rounded or pointed granules and the spaces between Heterometrus (Heterometrus) cyaneus: Couzijn, 1978:
them are finely granulate ventrally and smooth dorsally. 330.
The telson is elongate and hirsute, dorsally smooth and Heterometrus (Javanimetrus) cyaneus: Vachon & Abe,
ventrally with four rows of minute granules. The vesicle 1988: 27; Kovařík, 1992: 185; Kovařík, 1997: 183;
is longer than the aculeus. Kovařík, 1998: 137; Fet, 2000: 444; Kovařík, 2002: 17.
PEDIPALPS: The femur has three granulose complete Heterometrus (Javanimetrus) cyaneus cyaneus Couzijn,
carinae and its dorsal surface is smooth. The patella is 1981: 126; ? Le Xuan Hue et al., 1998: 8; Fet, 2000:
smooth, dorsally and externally slightly tuberculate, with 446.
five smooth carinae, and on the internal surface with Heterometrus (Javanimetrus) cyaneus sumatrensis
scattered pointed minute granules that form a Couzijn, 1981: 130; Kovařík, 1998: 137; Fet, 2000: 446.
conspicuous internal tubercle. The dorsal surface of the Heterometrus (Javanimetrus) cyaneus insulanus
manus is in internal parts tuberculate, without carinae. Couzijn, 1981: 131; Kovařík, 1998: 137; Fet, 2000: 446.
The chela is hirsute and its external surface is smooth, = Buthus defensor C. L. Koch, 1837: 3, fig. 254 (TL:
with two carinae composed of widely spaced granules. Mexico); C. L. Koch, 1850: 87 (syn. by Kraepelin, 1899:
The movable finger bears six or seven straight rows of 115; Couzijn, 1981: 126).
granules. Pandinus defensor: Karsch, 1887: 68.
LEGS: The legs bear solitary long and short setae. The = Buthus heros C. L. Koch, 1837: 1, fig. 253 (TL:
third and fourth leg spine formula is 4–5:6–7.
Africa); C. L. Koch, 1850: 87 (syn. by Kraepelin, 1899:
AFFINITIES. The described features distinguish H. 115; Couzijn, 1981: 126).
cimrmani sp. n. from all other species of the genus. They Pandinus (Buthus) heros: Karsch, 1879: 128.
are recounted in the key below. H. cimrmani sp. n. is = Buthus reticulatus C. L. Koch, 1837: 25, fig. 265 (TL:
close to H. spinifer, from which it differs primarily in Java); C. L. Koch, 1850: 88 (syn. by Simon, 1872: 98).
the expression of sexual dimorphism. = ? Buthus setosus C. L. Koch, 1841: Fig. 657; C. L.
Koch, 1841: 87 (TL: unknown); Gervais, 1844b: 458; C.
Heterometrus cyaneus (C. L. Koch, 1836) L. Koch, 1850: 88 (syn. by Kraepelin, 1899: 115).
(Fig. 12) Pandinus setosus: Karsch, 1887: 67.
= Heterometrus (Heterometrus) petersii luzonensis
Buthus cyaneus C. L. Koch, 1836: 75, fig. 225; C. L. Couzijn, 1981: 99; Kovařík, 1998: 137; Fet, 2000: 437
Koch, 1850: 88, (syn. by Kovařík, 2002: 3, 17).
12 Euscorpius — 2004, No. 15

Figure 12: Heterometrus cyaneus, dorsal view, female holotype of Heterometrus petersii luzonensis.
 Kovařík: Review of Genus Heterometrus 13

TYPE LOCALITY AND TYPE REPOSITORY. Java – holotype Islands (Couzijn, 1981: 130), Philippines (Couzijn,
lost, neotype from Indonesia, Java, Banjuwangi, Kali 1981: 99). A record from Vietnam (Le Xuan Hue et al.,
Baru designated by Couzijn (1981: 129); RMNH. 1998: 8) most likely concerns H. petersii, which was
TYPE MATERIAL EXAMINED. Philippines, Luzon, Mt. confused with H. cyaneus also by Couzijn (1981: 99 see
Maquiling, 1♂1♀ (holotype and paratype of Kovařík, 2002: 3, 17). The above list of other older
Heterometrus petersi luzonensis Couzijn, 1981), SMFD names and citations (see also Fet, 2000: 445) explains
No. 8882/224. many erroneous records from Africa and Asia.

OTHER MATERIAL EXAMINED. Indonesia, Java, 1♂1im.,

MZUF; Java, Matang, 1♀, SMFD No. 6705/112; Java, Heterometrus flavimanus (Pocock, 1900)
Idien-geb, 2♂2♀, SMFD No. 6743/150; Java, 1♂1♀, (Fig. 13)
col. V. Frič, NMPC, Java, Vulcan, 2♀, leg. Gurda,
NMPC; Sumatra, Fort de Kock, 1♀, SMFD No.
Palamnaeus swammerdami flavimanus Pocock, 1900:
8814/183; Sumatra, Deli, 2♀3ims., leg. Heyden, SMFD
87.
No. 5316; Java, Buitenzorg, 3♀, SMFD No. 8884/226;
Heterometrus (Gigantometrus) flavimanus: Tikader &
Java, 1906, 1♀, leg. Urbánková, NMPC; W. Java,
Bastawade, 1983: 568; Tikader, 1987: 36; Kovařík,
Djasinga, 1929, 1♂2♀, leg. Wolf, SMFD No. 5362;
1998: 137; Fet, 2000: 443.
Java, 1933, 1juv., leg. F. Ohaus, SMFD; Java, 4.X.1961,
Heterometrus (Gigantometrus) swammerdami
4ims., SMFD; Java, 1♀F, 1870, leg. J. Xantus, HNHM
flavimanus: Couzijn, 1981: 164.
Nos. 2005 and 2035; Java, 1♀, 1855 ?, leg. Doleschal,
HNHM No. 2033; Nias, 1896, 2♂1♀, leg. S. Raap and
Knuth, ZMHB No. 5705/37; Java, Buitenzorg, 1.I.– TYPE LOCALITY AND TYPE REPOSITORY. Coimbatore,
1.VII.1902, 4♂5♀2juvs., ZMHB; Java, 1♂, 1931, leg. India; BMNH.
Matysovszky, HNHM No. 2321; Sumatra, 1♀, CASC; TYPE MATERIAL EXAMINED. India, Tamil Nadu,
Pt Karya Tangan Indah, Br. Baturning, Desa Mambal, Coimbatore, 1♀ (lectotype), 1♂ (paralectotype), leg. W.
abian semal, 2♀, from Cynthia Hardy, CASC; Bogor, Brady, BMNH No. 1899.11.2.292-3.
purchase from Royal Botanical Garden, VI.1980, 1im.,
leg. Victoria Selmier, CASC; Java, 14♂10♀1im., 1993, OTHER MATERIAL EXAMINED. India, Tamil Nadu,
FKCP; Java, Bogor, 2♀, VIII.1994, leg. I. Gyerjdy, Coimbatore, VI.1954, 1im., VI.1955, 1juv.,17.XI.1957,
HNHM; Java, 1♂8♀3juvs., HNHM Nos. 2000, 2004, 1im., XI.1957, 1im.1juv., VII.1958, 1juv., XII.1958,
2011, 2013, 2020, and 2030; Sumatra, 1♂1im., leg. Dr. 1im.1juv., VI.1963, 1♂, leg. P. Susai Nathan, CASC;
Machik, HNHM No. 648/1883; Java, Bogor, The Madras, X.1971, 1juv., collector unknown, CASC;
Botanical Garden, ca 300 m, 26.VIII.1995, Coimbatore env., 1990, 1♂1♀, FKCP.
3♀5ims.4juvs., leg. P. Beron et T. Ivanova, SOFM.
Japan, Nagasaki, 1♀1juv., 1870, leg. J. Xantus (error or DIAGNOSIS. Adults 110–150 mm long. Base color
introduced), HNHM. Singapore, 1♂2♀2juvs., HNHM uniformly reddish brown, manus of pedipalp, legs and
Nos. 2001, 2009, 2018, 2019. ?, Usambara (loc. in telson reddish to yellow. Pectinal teeth number 19–22.
error), 1♀, SMFD No. 6707/114. Sexual dimorphism in proportions of pedipalps not
noticeable. Chela strongly lobiform, its length to width
DIAGNOSIS. Adults 95–117 mm long. Base color of ratio about 1.7 in both sexes. Entire manus covered by
adults uniformly black to greenish black, only telson may large, rounded granules that do not form true carinae.
be reddish brown. Pectinal teeth number 12–16 in both Patella of pedipalp without pronouced internal tubercle.
sexes. Sexual dimorphism in proportions of pedipalps not Carapace usually with disc smooth and all margins
noticeable. Chela lobiform, its length to width ratio in granulate and tuberculate (in some specimens entire
adults between 2.0 and 2.3 in both sexes. Dorsal surface carapace sparsely granulate). Fifth segment of metasoma
of manus tuberculate. Carinae weakly indicated. Patella longer than femur of pedipalp, fourth segment of
of pedipalp without pronounced internal tubercle. metasoma approximately as long as femur of pedipalp.
Carapace with disc smooth, at sides may be granulate. Telson bulbous, vesicle longer than aculeus.
Telson hirsute, bulbous, vesicle longer than aculeus.

COMMENTS. Couzijn (1981: 130 and 131) described two COMMENTS. Couzijn (1981: 164) designated the type
subspecies, H. c. sumatrensis and H. c. insulanus, which male as “allolectotype“ and the type female as
I have not studied and thus cannot discuss their status. “hololectotype“. Following the Code, I call these
specimens paralectotype and lectotype.
DISTRIBUTION. Indonesia (C. L. Koch, 1836: 78),
Borneo (Sabah) (Kopstein, 1921: 133), Lesser Nicobar DISTRIBUTION. India: Tamil Nadu (Pocock, 1900: 87).
14 Euscorpius — 2004, No. 15

Figure 13: Heterometrus flavimanus, dorsal view, male paralectotype.

 Kovařík: Review of Genus Heterometrus 15

Heterometrus fulvipes (C. L. Koch, 1837) Cavagnaro, CASC; 3 mi. W. Sakoli, 220 m., 30.I.1962,
(Fig. 14) 2juvs., leg. E. S. Ross and D. Q. Cavagnaro, CASC;
Bombay, 6.III.1967, 1♀, leg. S. R. Sane, CASC;
Buthus fulvipes C. L. Koch, 1837: 45, fig. 278; C. L. Mysore, 6 mi NE Ramanagaram, 750 m, 25.II.1962,
Koch, 1850: 87. 1im., leg. E. S. Ross & D. Cavagnaro, CASC;
Pandinus fulvipes: Karsch, 1887: 67; Thorell, 1888: 329; Rajasthan, Alwar district, Naranimata env., 27°05,46 N
Simon, 1905: 161. 76° 17,17 E, 380 m, VI–VIII.2002, 1♂, leg. P. Šrámek,
Scorpio fulvipes: Simon, 1885: 23; Kraepelin, 1895: 44; FKCP; Uttar Pradesh, Agra, 43 Kandhari Road,
Pocock, 1893: 304. 9.III.1964, 1im.6juvs., leg. S. Sina, CASC; Agra,
Palamnaeus fulvipes: Pocock, 1900: 87; Henderson, 9.III.1964, 19juvs., leg. S. Sina, CASC; West Bengal,
1919: 381; Cekalovic, 1982: 191. Calcutta, VI.1966, 1♂(im.), leg. N. Das, CASC.
Heterometrus fulvipes: Kraepelin, 1899: 112; Masi,
1912: 124; Kraepelin, 1913: 166; Kopstein, 1921: 133; DIAGNOSIS. Adults 70–100 mm long. Color of adults
Kopstein, 1923: 186; Kopstein, 1926: 111; Kopstein, uniformly reddish brown to black, legs and telson yellow
1927: 104; Giltay, 1931: 9; Werner, 1934: 277; to yellowish or reddish brown, always lighter colored
Takashima, 1945: 91; Brignoli, 1985: 415; Hjelle, 1990: than body. Pectinal teeth number 12–18 in both sexes.
26; Root, 1990: 354; Simard & Watt, 1990: 419; Sissom, Male has longer femur and patella of pedipalp. Chela
1990: 133; Warburg & Polis, 1990: 229; Pointer, 1991: lobiform, its ratio between 1.7 and 1.9 in both sexes.
24; Dupré, Lambert & Gérard, 1998: 13. Male and female chela of different shape but
Heterometrus (Scorpio) fulvipes: Kraepelin, 1901: 271. approximately same width. Entire manus covered by
Heterometrus (Chersonesometrus) fulvipes: Couzijn, large, rounded granules but devoid of carinae. Patella of
1978: 330; Couzijn, 1981: 133 (in part); Tikader & pedipalp without pronouced internal tubercle. Carapace
Bastawade, 1983: 599; Kovařík, 1997: 183; Kovařík, of juveniles and some females with disc smooth, at
1998: 136; Fet, 2000: 439; Bastawade, 2002: 296; margins granulate and anteriorly tuberculate. Males and
Kovařík, 2002: 17. some females with entire carapace sparsely granulate.
= Palamnaeus fulvipes bombayensis Pocock, 1900: 89 Telson hirsute, vesicle longer than or as long as aculeus.
(syn. by Couzijn, 1981: 133).

TYPE LOCALITY AND TYPE REPOSITORY. India, DISTRIBUTION. India: Andhra Pradesh, Bihar, Gujarat,
Maharashtra, Násik; BMNH. Karnataka, Madhya Pradesh, Maharashtra, Orissa,
TYPE MATERIAL EXAMINED. India, Maharashtra, Násik, Rajasthan, and Tamil Nadu (Pocock, 1900: 89, Couzijn,
1♀ (syntype of Palamnaeus fulvipes bombayensis 1981: 134, Tikader & Bastawade, 1983: 603); Mysore,
Pocock, 1900 and neotype of Buthus fulvipes C. L. Uttar Pradesh, and West Bengal (first report). The type
Koch, 1837, designated by Couzijn, 1981: 134), BMNH locality originally given as Java (C. L. Koch, 1837: 46)
No. 1913.1.6.27. is considered erroneous (see Fet, 2000: 439). Also
erroneous are records from Sri Lanka (Pocock, 1900: 88)
OTHER MATERIAL EXAMINED. India, 2♀(ims.), leg. and Myanmar (Kopstein, 1921: 133).
Kolle, ZMHB; east, X.1888, 1juv., HNHM; east, 1950,
2♀, FKCP; Tharigoppula, 8.II.1967, 1♀, 8.III.1967, Heterometrus gravimanus (Pocock, 1894)
1juv., leg. A. L. Slater, CASC; Andhra Pradesh, Podili, (Fig. 15)
9.I.1967, 2♂7♀4juvs., 16.I.1967, 1♀1♂, 13.II.1967,
1♂10♀1im., leg. D. E. Johnson, CASC; Gujarat, Scorpio gravimanus Pocock, 1894: 75.
Ahmedabad, 2♂, leg. Manning, BMNH No.
Pandinus gravimanus: Simon, 1905: 161.
1930.4.11.1-2; 4 mi. E. Godhra, 175 m., 12.I.1962, 1im.,
Palamnaeus gravimanus: Pocock, 1900: 90; Henderson,
CASC; Ahmedabad, 4.IV.1967, 3ims., 26.VII.1966, 1♂,
IX.1966, 1♂, 18.XI.1966, 1juv., 20.XII.1966, 1♂1♀, 1919: 381.
leg. M. S. Dubale, CASC, Ahmedabad, 1967, 2♂1♀A, Heterometrus gravimanus: Takashima, 1945: 94; Root,
FKCP; Karnataka, Hassan env., 2ims., 15–16.VI.1994, 1990: 354.
leg. R. Sauer, FKCP; Madhya Pradesh, Khaighat, 200 Heterometrus (Srilankametrus) gravimanus: Tikader &
m., 18.I.1962, 1♀, CASC, 3 miles from Damoh, Bastawade, 1983: 550; Kovařík, 1998: 137; Fet, 2000:
20.VII.1965, 1im., leg. S. N. Banerjee, CASC; 446.
Maharastra, 17 mi. NE. Malegaon, 400 m., 15.I.1962, Heterometrus (Srilankametrus) indus indus (in part):
1juv., leg. E. S. Ross and D. Q. Cavagnaro, CASC; Couzijn, 1981: 121.
Ajanta Caves, 500 m., 28.I.1962, 1♀(im.), leg. E. S. = ? Heterometrus (Srilankametrus) indus laevitensus
Ross and D. Q. Cavagnaro, CASC; 3 mi. W. Edalabad, Couzijn, 1981: 124; Kovařík, 1998: 137; Fet, 2000: 447.
240 m., 28.I.1962, 3juvs., leg. E. S. Ross and D. Q. Syn. n.
16 Euscorpius — 2004, No. 15

Figure 14: Heterometrus fulvipes dorsal view, female neotype.

 Kovařík: Review of Genus Heterometrus 17

TYPE LOCALITY AND TYPE REPOSITORY. India, Ceylon; Heterometrus afer: Simon, 1872: 99.
BMNH. Scorpio indus De Geer, 1778: 341; Lönnberg, 1898: 551.
Pandinus indicus: Pavesi, 1881: 556 ?; Thorell, 1888:
MATERIAL EXAMINED. India, south, Tanjore, 1♂, leg. E. 412 (in part); Thorell, 1893: 381 (in part).
P. Popert, BMNH No. 1896.7.30.121. Sri Lanka, Heterometrus indus: Kraepelin, 1899: 113; Werner,
Southern prov., Galle, IV.1994, 1♂, leg. P. Senft, FKCP. 1902: 601; Lampe, 1918: 198; Roewer, 1929: 612;
Giltay, 1931: 8; Werner, 1934: 277; Roewer, 1943: 226;
DIAGNOSIS. Adults 80–110 mm long. Color of adults Takashima, 1945: 92; Bücherl, 1959: 269; Root, 1990:
uniformly reddish brown, legs and telson usually lighter 354; Lourenço & Cloudsley-Thompson, 1996: 140.
colored than body. Pectinal teeth number 12–15 in both Heterometrus (Scorpio) indus: Kraepelin, 1901: 271.
sexes. Male has slightly longer femur and patella of Heterometrus (Heterometrus) indus: Couzijn, 1978: 330.
pedipalp. Chela lobiform, its length to width ratio Heterometrus (Srilankametrus) indus: Couzijn, 1981:
between 2.0 and 2.2. Entire manus covered by rounded 121; Kovařík, 1992: 185; ? Kovařík, 1997: 183;
granules and with five carinae mainly in anterolateral
Kovařík, 1998: 137; Fet, 2000: 446; Kovařík, 2002: 17.
part. Patella of pedipalp smooth, without pronouced
Heterometrus (Srilankametrus) indus indus: Couzijn,
internal tubercle. Carapace smooth and glossy. Third and
1981: 121; Vachon, 1982: 79; Fet, 2000: 447.
fourth leg spine formula 4:5–6. Telson elongate, vesicle
shorter than aculeus. Palamnaeus indus: Pocock, 1900: 96.
Scorpio indicus: Pocock, 1893: 311(in part); Kraepelin,
COMMENTS. This species is based on a male from Sri 1895: 53.
Lanka, which ought to be at BMNH but cannot be = Scorpio ceylonicus Herbst, 1800: 83; Kraepelin, 1895:
found. It was not seen by the previous revisers (Couzijn, 46; Lönnberg, 1897: 186 (syn. by Kraepelin, 1899: 113;
1981: 121, Tikader & Bastawade, 1983: 550) either. Pocock, 1900: 96).
Available is only a male from Tanjore, which was Scorpio (Buthus) ceylanicus: Gervais, 1844b: 62.
assigned to this species by its author (Pocock, 1900: 91). Scorpio ceylonicus: Lönnberg, 1898: 83.
Couzijn lists Scorpio gravimanus Pocock, 1894 as a Pandinus ceylonicus: Karsch, 1887: 69; Karsch, 1892:
synonym of Heterometrus (Srilankametrus) indus (Geer, 307.
1778) and says the specimen from Tanjore could be Palamnaeus ceylonicus: Kraepelin, 1898: 439.
Heterometrus (Srilankametrus) indus laevitensus = Buthus megacephalus C. L. Koch, 1836: 73, fig. 224;
Couzijn, 1981 (Couzijn, 1981: 125). Tikader & C. L. Koch, 1850: 88 (syn. by Thorell, 1876b: 168;
Bastawade (1983: 550–555) examined the specimen Lönnberg, 1898: 83).
from Tanjore and correctly concluded that it is Bathus megacephalus [sic]: C. L. Koch, 1836 (content).
Heterometrus gravimanus, which they considered valid Pandinus megacephalus: Thorell, 1876b: 203; Thorell,
(see also Fet, 2000: 447). Like Pocock, I have had an 1893: 381.
opportunity to examine the specimen from Tanjore as Scorpio megacephalus: Pocock, 1893: 311.
well as the one from Sri Lanka and am convinced that = Buthus caesar C. L. Koch, 1841: 6, fig. 697; C. L.
both populations belong to the same species, which Koch, 1850: 87; Moritz & Fischer, 1980: 311 (syn. by
differs from H. indus in smaller size, narrower chela of Simon, 1872: 99).
pedipalp and the presence of five carinae on the manus Pandinus caesar: Thorell, 1876b: 205; Karsch, 1887: 68.
of pedipalp. Scorpio caesar: Pocock, 1893: 311; Pocock, 1894: 73.
Heterometrus caesar: Kraepelin, 1899: 114; Roewer,
DISTRIBUTION. India: Tamil Nadu. Sri Lanka (Pocock, 1943: 229; Takashima, 1945: 92; Brignoli, 1985: 415.
1894: 77; Pocock, 1900: 91). Heterometrus (Scorpio) caesar: Kraepelin, 1901: 271.
Palamnaeus caesar: Pocock, 1900: 97.
= Scorpio crassimanus Becker, 1880: 140 (syn. by
Heterometrus indus (Geer, 1778) Kraepelin, 1899: 113; Couzijn, 1981: 121).
(Fig. 16) = Palamnaeus serratus Pocock, 1900: 97 (syn. by
Couzijn, 1981: 121).
? Scorpio indicus (in part) Linné, 1754 (syn. by Heterometrus serratus: Takashima, 1945: 94.
Kraepelin, 1899: 113). Heterometrus (Srilankametrus) serratus: Tikader &
= Scorpio afer Linné, 1758: 624 (in part); Linné, 1767: Bastawade, 1983: 555; Fet, 2000: 448.
1038; Linné, 1775: 1091; Fabricius, 1775: 399;
Fabricius, 1781: 550; Fabricius, 1787: 348; Fabricius, TYPE LOCALITY AND TYPE REPOSITORY. India; NHRS.
1793: 434; Herbst, 1800: 38; Latreille, 1804: 120 (syn. TYPE MATERIAL EXAMINED. Sri Lanka, Ceylon, 1♂
by De Geer, 1778: 341; Kraepelin, 1895: 53). (holotype of Palamnaeus serratus), leg. Dr. Ondaatje,
Scorpio (Buthus) afer: Gervais, 1844b: 60 (in part). BMNH No. 1888.55.
18 Euscorpius — 2004, No. 15

Figure 15: Heterometrus gravimanus, dorsal view, male from India (BMNH).
 Kovařík: Review of Genus Heterometrus 19

Figure 16: Heterometrus indus, dorsal view, male holotype of Palamnaeus serratus.
20 Euscorpius — 2004, No. 15

OTHER MATERIAL EXAMINED. Sri Lanka (Ceylon), TYPE MATERIAL EXAMINED. India, Kanara, Bombay,
Peradeniya, 2♂, leg. Uzel, NMPC; 1847, 1♀1juv., leg. 1♂1♀ (lectotype and paralectotype), leg. T. R. D. Bell,
G. Worms, SMFD No. 5319; Ceylon, 1912, 1♀1♂1juv., BMNH No. 1897.11.5.29-30.
leg. A. Hansen, SMFD No. 5317; Ceylon, IV.1914,
1♂2♀, leg. J. Mastbaum, SMFD No. 5318; Kandy, OTHER MATERIAL EXAMINED. India, 1♀ without
1.V.1982, 1♂, leg. R. C. Drewes, CASC; 1900, 1♀, locality, CASC; Goa, Portuguese India, 18.X.1943,
FKCP; 1994, 1♂, FKCP; Kandy, Paradeniya, 1im., leg. J. L. Gressitt, CASC; Maharashtra, Karwa,
30.III.1902, 1♂, IV.1994, 2♂1im.5juvs., 1♀ (gave birth North Kanara, 1980, 2♀, FKCP.
to 8 juvs. in 10–19.VII.1994, ♀ died 28.VI.1996, reared
by F. Kovařík), 1♀ after seventh ecdysis (5th DIAGNOSIS. Adults 120–150 mm long. Base color of
29.VII.1994, 6th 1.IV.1996, 7th ecdysis 1.IX.1996, adults uniformly reddish brown to black, only manus and
rearing F. Kovařík), 1♂ after seventh ecdysis (3rd telson may be lighter colored or occasionally black.
1.VIII.1994, 4th 26.V.1995, 5th 17.VIII.1995, 6th Pectinal teeth number 13–14 in male and 10–13 in
10.IV.1996, 7th 13.IX.1996, reared by F. Kovařík), leg. female. Male has as long but narrower chela of pedipalp
P. Senft, FKCP. ?, Hyderabad, Banjara Road, 1931, 1♀, than female. Other segments of pedipalps do not show
leg. S. Mirza, SMFD No. 5367; 1im., NMPC; ? Java sexual dimorphism. Chela length to width ratio 2.35 in
(loc. in error ?), Mus. Leiden, 1♀, SMFD No. 5321. male anda 2.1–2.25 in females. Chela lobiform and
covered with dense, long setae. Dorsal surface of
DIAGNOSIS. Adults 90–130 mm long. Color of adults pedipalp manus relatively smooth, with effaced tubercles,
uniformly reddish black to greenish black. Pectinal teeth without carinae. Patella of pedipalp without pronounced
number 11–15 in both sexes. Sexual dimorphism in internal tubercle. Carapace usually smooth and glossy,
proportions of pedipalps not noticeable. Chela hirsute, occasionally with sparse marginal granules and anterior
lobiform, its adult length to width ratio between 1.7 and tubercles. Third and fourth leg spine formula 4:6. Telson
2.0. Entire manus covered by rounded granules that may densely hirsute, elongate, vesicle shorter than the
merge and appear as rows. Patella of pedipalp without aculeus.
pronouced internal tubercle. Carapace usually smooth
and glossy, occasionally with granules at margins. DISTRIBUTION. India: Maharashtra (Pocock, 1900: 93),
Second metasomal segment longer than wide. Vesicle of Karnataka (Tikader & Bastawade, 1983: 641), Goa (first
telson usually longer than aculeus. report).

COMMENTS. Palamnaeus serratus Pocock, 1900 was Heterometrus keralaensis Tikader & Bastawade,
synonymized by Couzijn (1981: 121) with H. indus, 1983
whereas Tikader & Bastawade (1983: 555) considered it
a valid species. Examination of the holotype convinces Heterometrus (Heterometrus) keralaensis: Tikader &
me that it is a synonym of H. indus. Bastawade, 1983: 528; Kovařík, 1998: 137; Fet, 2000:
432; Bastawade, 2002: 295.
DISTRIBUTION. Sri Lanka (Herbst, 1800: 83, 84; Pocock,
1900: 96). This species has been repeatedly listed from TYPE LOCALITY AND TYPE REPOSITORY. India, Kerala,
India (see name and type locality), but it appears that in Meenmutty, New Amaranbalan; NZSI,
reality it is confined to Sri Lanka. It was also incorrectly
reported from Java (Lönnberg, 1897: 187) and COMMENTS. I was unable to examine this species and
Mozambique (Pavesi, 1881: 556). had to include it in the key solely on the data published
by Tikader & Bastawade (1983: 528–533). I believe it is
Heterometrus kanaraensis (Pocock, 1900) related to H. thorellii Pocock, 1892, from which it
differs (apart from the characters given in the key) in
Palamnaeus scaber kanarensis Pocock, 1900: 93. having the dorsal surface of the chela reticulated.
Heterometrus (Chersonesometrus) phipsoni kanaraensis
Couzijn, 1981:151; Kovařík, 1998: 136. DISTRIBUTION. India: Kerala (Tikader & Bastawade,
Heterometrus (Chersonesometrus) kanaraensis: Tikader 1983: 533), Maharashtra (Bastawade, 2002: 295).
& Bastawade, 1983: 636; Fet, 2000: 441; Bastawade,
2002: 296. Heterometrus laoticus Couzijn, 1981

TYPE LOCALITY AND TYPE REPOSITORY. India, Kanara; Heterometrus (Heterometrus) laoticus Couzijn, 1981:
BMNH. 94; Kovařík, 1995: 202; Kovařík, 1997: 183; Kovařík,
 Kovařík: Review of Genus Heterometrus 21

1998: 137; Le Xuan Hue et al., 1998: 8; Fet, 2000: 433; COMMENTS. This species is often mistaken for H.
Kovařík, 2002: 17. petersii (Thorell, 1876), whose distribution overlaps.
Heterometrus (Heterometrus) spinifer: Kovařík, 1992: Apart from sexual dimorphism, H. laoticus differs in
185. having a smooth carapace without granules and usually
also a smooth manus of pedipalp with more apparent
TYPE LOCALITY AND TYPE REPOSITORY. Laos; MNHN. punctae.

MATERIAL EXAMINED. Cambodia, Takeo, 1984, 2♀, leg. DISTRIBUTION. Cambodia, Laos, Thailand, Vietnam
Frühbauer, FKCP and NMPC. Laos, 1♀, FKCP; Karen- (Couzijn, 1981: 94).
Dorf, ca 8 km SE Luang Prabang, 18.XII.1964, 1♀,
SMFD; Bolikhamsai prov., Ban Nok env., alt. 220 ± Heterometrus latimanus (Pocock, 1894)
50m, N 18°08'7 E 104°28'1, 9–14.V.1998, 2♂, leg. E.
Jendek & O. Šauša. Thailand, Lampun N., Siam, leg. Scorpio latimanus Pocock, 1894: 74.
Daly, 1♂, BMNH; Tha-Manao, 30 km NW Palamnaeus latimanus: Pocock, 1900: 90.
Kanchanaburi, 145 km NW Bangkok, leg. W. Thielen, Heterometrus latimanus: Kraepelin, 1899: 113;
3♂3♀, ZMUH; Chiang Mai, leg. W. Schmidt, 1♂, Takashima, 1945: 92; Minnocci, 1974: 38.
ZMUH; Krachong For., near Trang, 100m, 2.VII.1962, Heterometrus (Srilankametrus) latimanus: Tikader &
1juv., leg. E. S. Ross and D. Q. Cavagnaro, CASC; Bastawade, 1983: 545; Fet, 2000: 448.
Khorat, 19.VI.1963, 1♂, leg. R. H. Nave, CASC; Heterometrus (Srilankametrus) indus: Couzijn, 1981:
Nakhon Phanom, 17.IX.1965, 3ims., leg. Jim Foote, 121 (in part).
CASC; Chiang Rai Prov., 4.V.1966, 1♂, leg. Paul
Soderberg, CASC; Chiang Mai Prov., VI.1966, TYPE LOCALITY AND TYPE REPOSITORY. India; BMNH.
1♀2juvs., leg. Terry W. Taylor, CASC; 50 miles N.
Bangkok, 1.XII.1968, 1♂, CASC; Udon Thani, MATERIAL EXAMINED. India oriental, 1960, 2♀,
24.XII.1970, 2♂2♀10juvs., leg. Jim Foote, CASC; collector unknown, FKCP.
Chain mai, 20.XI.1986, 1♀, leg. Leslie Saul; Doi Pui,
(near Chain mai), 1400 m, 20.VIII.1989, 1♂, leg. E. S. DIAGNOSIS. Adults 70–105 mm long. Color of adults
Ross, CASC; I.2002, 4♂7♀, CASC; 29.I.2002, died uniformly reddish brown to black. Legs may be lighter
22.VIII.2002, 1♂, CASC; Ban Saen near Chonburi, colored than body. Pectinal teeth number 13–15. Chela
3.VI.1991, 1♂, leg. D. Král (reared by F. Kovařík, 6th lobiform, rounded and convex, its adult female length to
ecdysis 12.X.1991); Khorat, 180 km NE of Bangkok width ratio 1.6–1.7. Entire manus covered by rounded
(purchase from Bangkok), VI.1991, 7♀4♂4juvs., 1994, granules, without carinae. Patella of pedipalp without
1♀2juvs. before first.ecdysis and 9 juvs. after first pronouced internal tubercle. Carapace usually smooth
ecdysis; Khao Yai nat. park, XII.1997, 1♂, leg. K. and glossy, but granules always present anteriorly and
Petrželka; Khao Khieo Wildlife and Nature Sanctury occasionally on other margins. Vesicle of telson usually
(Chan Buri), II.1994, 1juv. after 4th ecdysis, leg. S & L. shorter or as long as aculeus.
Mahunka, HNHM. Vietnam, QuiNhon, 5.VIII.1965,
1♂, leg. L. J. Barrier, CASC. COMMENTS. Scorpio latimanus Pocock, 1894 was
Specimens born and reared in captivity (Locality synonymized by Couzijn (1981: 121) with H. indus,
Thailand, Khorat), 1juv. after first ecdysis, 1juv. after whereas Tikader & Bastawade (1983: 545–550)
2nd ecdysis, 8juvs. after 3rd ecdysis, 1juv. after 4th considered it a valid species. H. latimanus differs from
ecdysis, 1im. after 5th ecdysis, 1M1F after 6th ecdysis, H. indus in the shape of the pedipalp chela, which is also
reared by F. Kovařík, FKCP. convex and lacks setae. In H. indus the anterior portion
of the carapace is not granulated.
DIAGNOSIS. Adults 90–125 mm long. Base color of
adults uniformly black, only manus and telson may be DISTRIBUTION. India (Pocock, 1894: 74). Unfortunately,
reddish brown. Pectinal teeth number 15–19 in both neither the types nor the specimens in my colection bear
sexes. Sexual dimorphism in proportions of pedipalps not precise locality data.
noticeable. Chela lobiform, its length to width ratio 2–2.3
in both sexes. Manus smooth, with or without smooth Heterometrus liophysa (Thorell, 1888)
carinae, sparsely and finely punctate. Patella of pedipalp (Fig. 17)
without pronounced internal tubercle. Carapace and
mesosoma, smooth without granules. Telson hirsute, Palamnaeus liophysa Thorell, 1888: 415.
elongate, vesicle longer than aculeus. Scorpio longimanus liophysa: Kraepelin, 1895: 41.
22 Euscorpius — 2004, No. 15

Figure 17: Heterometrus liophysa, dorsal view, male.

 Kovařík: Review of Genus Heterometrus 23

Heterometrus liophysa: Kraepelin, 1899: 112; Werner, Weh and H. l. spartanicus from Babi. I have not been
1916: 91; Lampe, 1918: 198; Kopstein, 1921: 131; able to verify the status of these two taxons.
Kopstein, 1923: 184; Takashima, 1945: 91. Couzijn (1981: 116) elevated H. l. madoerensis
Heterometrus (Heterometrus) liophysa: Kovařík, 1998: Kopstein, 1921 to species, although it is known only by
137; Fet, 2000: 433; Kovařík, 2002: 17. an immature female from Madura (Indonesia). I have not
Heterometrus liophysa liophysa: Giltay, 1931: 9. been able to study this specimen, but I have seen many
Heterometrus (Heterometrus) liophysa liophysa Couzijn, females of H. cimrmani sp. n., H. liophysa, H.
1981: 111; Fet, 2000: 433 longimanus, and H. spinifer and have to say that
= Heterometrus liophysa madoerensis Kopstein, 1921: distinguishing these valid taxa on solitary, lest immature,
132; Giltay, 1931: 9. Syn. n. females is very difficult and often impossible. For this
Heterometrus (Heterometrus) madoerensis Couzijn, reason I place H. madoerensis in synonymy of H.
1981: 116; Kovařík, 1998: 137; Fet, 2000: 436. liophysa.
= Heterometrus laevifrons Roewer, 1943: 228. Syn. n.
Heterometrus (Heterometrus) liophysa laevifrons: DISTRIBUTION. Indonesia – Sumatra and Mentawei,
Couzijn, 1981: 114; Kovařík, 1998: 137; Fet, 2000: 433; Nias, Babi, Batu and Weh Islands (Thorell, 1888: 415;
Kovařík, 2002: 17. Couzijn, 1981: 111–116; Fet, 2000: 433–434).
Heterometrus (Heterometrus) liophysa separatus
Couzijn, 1981: 115; Kovařík, 1998: 137; Fet, 2000: 433. Heterometrus liurus (Pocock, 1897)
Heterometrus (Heterometrus) liophysa spartanicus (Fig. 18)
Couzijn, 1981: 115; Kovařík, 1998: 137; Fet, 2000: 434.
Palamnaeus liurus Pocock, 1897: 114; Pocock, 1900:
TYPE LOCALITY AND TYPE REPOSITORY. Indonesia, 91.
Sumatra, Ajer Mancior – syntypes MCSN probably lost; Heterometrus liurus: Kraepelin, 1899: 115; Roewer,
neotype from Indonesia, Sumatra, Padang designated by 1929: 613; ? Fage, 1936: 181; Roewer, 1943: 228; Fage,
Couzijn (1981: 111); RMNH. 1944: 71; Takashima, 1945: 93; Minnocci, 1974: 38;
TYPE MATERIAL EXAMINED. Indonesia, Mentawei Kovařík, 2002: 20.
Islands, Siberut, 2juvs. (♂ holotype and ♀ paratype of Heterometrus (Chersonesometrus) liurus: Couzijn,
Heterometrus laevifrons Roewer, 1943), SMFD No. 1981: 153; Tikader & Bastawade, 1983: 604; Kovařík,
8883/225. 1998: 136; Fet, 2000: 442.

OTHER MATERIAL EXAMINED. Indonesia, Sumatra, TYPE LOCALITY AND TYPE REPOSITORY. Gwalior, Central
Padang, 2♂(ims.), SMFD No. 5278; Sumatra, Bunga India; BMNH.
Bondar, 1911, 1♂, 28.X.1911, 2♂, leg. Schütze, SMFD TYPE MATERIAL EXAMINED. India, Gwalior (Dane), 1♀
No. 5323 and 18158; Sumatra, Nias, 1♀, leg. P. Beyer, (lectotype), 2♀1im. (paralectotypes), BMNH No.
SMFD No. 5322; Padang env., 1♂1♀, 1980, FKCP; 1896.12.15.20-25, No. 995 (lectotype) and 992, 994, and
Mentawai islands, Suberut Island, Muarasiberut, 0–150 987 (paralectotypes).
m, 15–20.VIII.1995, 2juvs., leg. P. Beron & T. Ivanova,
SOFM Nos 180 and 182. DIAGNOSIS. Adults less than 80 mm long. Color
uniformly reddish brown, legs and telson yellow to
yellowish or reddish brown, always lighter colored than
DIAGNOSIS. Adults 110–140 mm long. Base color of body. Pectinal teeth number 10–14. Chela slightly
adults uniformly black to reddish black. Pectinal teeth lobiform, hirsute, its length to width ratio 2.2–2.5. Dorsal
number 13–18 in both sexes. Male with chela, femur and surface of pedipalp manus without carinae, with large,
patella of pedipalp narrower and longer than in female. rounded granules of uneven size, some of which may
Male fingers of pedipalp very long. Fixed finger longer merge. Patella of pedipalp without conspicuous internal
or at least as long as manus. Chela slightly lobiform, in tubercle. Carapace smooth and glossy, on margins
male widest basally and markedly narrowing toward sometimes sparsely granulose and anteriorly tuberculate.
fingers, its length to width ratio ca. 3.0 in male, ca. 2.2 in Third and fourth leg spine formula 4–5:5–6. Telson
female. Much of manus smooth. Patella of pedipalp with hirsute, bulbous, with vesicle markedly longer than
pronounced internal tubercle. Carapace granulate. Telson aculeus. Genital operculum large, approximately as wide
hirsute, elongate, vesicle longer than aculeus. as long.

COMMENTS. H. laevifrons Roewer, 1943 was based on COMMENTS. Couzijn (1981: 153) designated the female
juveniles (examined), which cannot be distinguished lectotype as „hololectotype“ and the male paralectotype
from the nominotypical subspecies. Couzijn (1981) as „allolectotype“. Those designation are brought into
described two more subspecies, H. l. separatus from compliance with the Code.
24 Euscorpius — 2004, No. 15

Figure 18: Heterometrus liurus, dorsal view, female paralectotype.

 Kovařík: Review of Genus Heterometrus 25

DISTRIBUTION. India: Madhya Pradesh (Pocock, 1897: Palamnaeus longimanus angustimanus: Simon, 1901:
114). The record for Sri Lanka (Roewer, 1943: 228) was 78.
a mistaken determination (see Kovařík, 2002: 20). Heterometrus (Heterometrus) longimanus angust-
imanus: Couzijn, 1981: 103; Kovařík, 1998: 137; Fet,
Heterometrus longimanus (Herbst, 1800) 2000: 435.
(Fig. 19) = Pandinus humilis Simon, 1877: 94 (syn. by Kraepelin,
1895: 34).
Scorpio longimanus Herbst, 1800: 42; Latreille, 1804: Heterometrus (Heterometrus) longimanus humilis
122; Kraepelin, 1895: 34; Kraepelin, 1897: 537; Couzijn, 1981: 109; Kovařík, 1998: 137; Fet, 2000: 435.
Lönnberg, 1897: 186; Kraepelin, 1908: 191. = Heterometrus (Heterometrus) longimanus belitung-
Buthus longimanus: C. L. Koch, 1840: Fig. 595; C. L. ensis Couzijn, 1981: 105; Kovařík, 1998: 137; Fet,
Koch, 1841: 1; C. L. Koch, 1850: 88. 2000: 435; Kovařík, 2002: 17. Syn. n.
Scorpio (Ischnurus) longimanus: Gervais, 1844b: 69. Heterometrus (Heterometrus) longimanus bengkalitensis
Palamnaeus longimanus: Simon, 1893: 328; Simon, Couzijn, 1981: 106; Kovařík, 1998: 137; Fet, 2000: 435.
1899: 79; Pocock, 1900: 97 (in part); Simon, 1901: 78; Heterometrus (Heterometrus) longimanus marmoratus
Schultze, 1927: 375; Auber, 1963: 273. Couzijn, 1981: 109; Kovařík, 1998: 137; Fet, 2000: 436.
Palamnaeus (Scorpio) longimanus: Pavesi, 1898: 2. Heterometrus (Heterometrus) longimanus paris Couzijn,
Heterometrus longimanus: Kraepelin, 1899: 111; 1981: 106; Kovařík, 1998: 137; Fet, 2000: 436.
Werner, 1902: 601; Masi, 1912: 123; Kraepelin, 1913: = Heterometrus (Heterometrus) longimanus tarawakan-
166; Lampe, 1918: 197; Kopstein, 1921: 129; Kopstein, ensis Couzijn, 1981: 108; Kovařík, 1998: 137; Fet,
1923: 186; Kopstein, 1926: 111; Kopstein, 1927: 104; 2000: 436. Syn. n.
Banks, 1928: 505; Werner, 1932: 576; Werner, 1934:
277; Werner, 1936: 186; Roewer, 1943: 228; Vachon, TYPE LOCALITY AND TYPE REPOSITORY. Africa –
1948: 57(1952: 37); ? Bücherl, 1959: 269; Rosin & incorrect type locality – holotype lost, neotype from
Shulov, 1963: 570; Vachon, Roy & Condamin, 1970: Indonesia, Sumatra, Talang, Gunung designated by
426; Stockmann, 1979: 405; Francke & Sissom, 1984: Couzijn (1981: 100); RMNH.
12; Casper, 1985: 281; McCormick & Polis, 1990: 301; TYPE MATERIAL EXAMINED. Indonesia, Bangka,
Polis & Sissom, 1990: 187; Pointer, 1991: 24; Soleglad 2♀2juvs. (paratypes of Heterometrus (Heterometrus)
& Fet, 2003: 8. longimanus belitungensis Couzijn, 1981), SMFD No.
Heterometrus (Scorpio) longimanus: Kraepelin, 1901: 5330.
271.
Heterometrus (Heterometrus) longimanus: Vachon & OTHER MATERIAL EXAMINED. Indonesia, Java, Idien-
Lourenço, 1985: 9; Kovařík, 1992: 185; Kovařík, 1994: geb, 3♀, SMFD No. 6743/150; Sumatra, Padang, 1♂,
197; Kovařík, 1997: 183; Kovařík, 1998: 137; Fet, 2000: SMFD No. 6702/109; Sumatra, 2juvs., leg. S. Auer,
434; Kovařík, 2002: 17; Soleglad & Fet, 2003: 5. SMFD No. 5291; Sumatra, Fort de Kock, 1♂1♀, SMFD
Heterometrus longimanus typicus: Masi, 1912: 123. No. 8814/183; Sumatra, Padang, 1♂, leg. Tiefsee,
Heterometrus longimanus longimanus: Giltay, 1931: 3; ZMHB No. 98/99 and 882/1914; Borneo, 1♂4♀3juvs.,
Giltay, 1935: 2; Takashima, 1945: 90. leg. Vráz, NMPC; Sumatra, 3♂, leg. Hildebrandt,
Heterometrus (Heterometrus) longimanus longimanus: NMPC; Java, 2juvs., 1870, leg. J. Xantus, HNHM;
Couzijn, 1981: 100; ?Tikader & Bastawade, 1983: 522; Borneo, Matang, 1870, 8♂1♀36juvs.17juvs. before first
ecdysis, leg. J. Xantus, HNHM Nos. 2024, 2031, 2032,
Fet, 2000: 435.
and 2039; Sumatra, Deli, 1879, 2♂1♀, leg. V. Schauler,
= Buthus costimanus C. L. Koch, 1837: 27, fig. 266; C.
SMFD No. 5328; Sumatra, 1882, 2♀, HNHM Nos. 2015
L. Koch, 1850: 88 (syn. by Kraepelin, 1895: 34).
and 2036, 1883, 2♂, leg. Dr. Machik, HNHM Nos. 2021
Palamnaeus costimanus: Thorell, 1876b: 217; Thorell, and 2026; Sumatra, 1886, 1♀, leg. Hochwiesner, SMFD
1888: 329. No. 5329; Sumatra, Deli, 1892, 2♂2♀1im., leg.
Palamnaeus costimanus borneensis Thorell, 1876b: 217; Benecke, SMFD No. 5348; West Borneo, Pontianak,
Thorell, 1888: 414. 1893, 2♂2♀6juvs., leg. F. Will, SMFD No. 5324;
Heterometrus (Heterometrus) longimanus borneensis: Halmahera, 1894, 5♂5♀4ims., leg. Kükenthal, SMFD
Couzijn, 1981: 104; Kovařík, 1997: 183; Kovařík, 1998: No. 5350; Borneo, Baramfluss, 1894, 3ims.13juvs., leg.
137; Fet, 2000: 435 Kükenthal, SMFD Nos 5325 and 5326; Nias, 1896, 2♂
= Centrurus galbineus C. L. Koch, 1838: 110; C. L. (ims.)1♀, leg. S. Raap and Knuth, ZMHB No. 5705/37;
Koch, 1850: 89; Pocock, 1902: 365; Braunwalder & Fet, Linga Island, VII.1899, 1♂, leg. W. L. Abbott, probably
1998: 33 (syn. by Kraepelin, 1895: 34). Pulau Lingga, 0º12'S 107º35'E, CASC; Sumatra, 1900,
= Palamnaeus angustimanus Thorell, 1876b: 211 (syn. 1♀1im., leg. V. Guer, SMFD No. 5327; E. Sumatra,
by Simon, 1901: 78). Kateman river, 20.VIII.1903, 1♂, 1.IX.1903, 1im.,
26 Euscorpius — 2004, No. 15

Figure 19: Heterometrus longimanus, dorsal view, male.

 Kovařík: Review of Genus Heterometrus 27

CASC; Sumatra, 9.II.1902, 1♂, ZMHB; Sumatra, DISTRIBUTION. Indonesia (L. Koch, 1837: 29), Malaysia
22.IV.1905, 1♂, leg. G. Studt, ZMHB No. 491/05; (Kraepelin, 1895: 39), Philippines (Simon, 1877: 95),
Sumatra, Kahat, X.1925, 1♂1im., CASC; Borneo, Singapore (Fet, 2000: 434). Apart from the original type
XI.1926, 1im. ♂ 2juvs., leg. Stebbing, BMNH No. locality (Africa), as erroneous must be regarded also all
1926.XI.18.10-11; Sumatra, Takengon, 21.I.1929, records for Cambodia (Kraepelin, 1895: 39), India
1♂1♀, leg. H. R. Rookmaaker, ZMHB No. 405D/29; (Kraepelin, 1901: 271; Tikader & Bastawade, 1983:
Mentaus, 3♀, ZMHB No. 1198/07; Sumatra, Hügelland 527), Laos (Kraepelin, 1901: 271), Myanmar (Kraepelin,
Boelaeh Blang-Cra, Lho Seumawe, Atjeh, 1929, 1895: 39, Kraepelin, 1913: 166), Thailand (Kraepelin,
1♂2♀1juv., leg. Rookmaaker, SMFD No. 5368; 1901: 271) and Vietnam (Kraepelin, 1901: 271).
Sumatra, Tandjong, Morava, 1931, 9♂3♀1juv., leg.
Kleinkay, HNHM No. 2316; Sumatra, Lippisches Heterometrus madraspatensis Pocock, 1900
Landes, 26.VII.1954, 4♂3♀5juvs., SMFD; 1992, (Fig. 20)
2♂1♀2juvs., FKCP; Timur Nunukan, rain forest, 15–
17.IX.1995, 1♀2juvs., leg. P. Beron & T. Ivanova, Palamnaeus fulvipes madraspatensis Pocock, 1900: 88.
SOFM No. 177; Sumatra, S. Medan, Brastago, VI.1997, Heterometrus (Chersonesometrus) madraspatensis:
1im., leg. R. Sauer, FKCP; Borneo, 1♂1♀, FKCP; Tikader & Bastawade, 1983: 630; Fet, 2000: 440.
Borneo, Nanga Pinoh distr, Tontang, 26.VII.1993, 1juv., Heterometrus (Chersonesometrus) fulvipes: Couzijn,
leg. J. Schneider, FKCP; W. Sumatra, Japannli bay, 1981: 133 (in part).
2.I.2002, 1♂, CASC. Malaysia, Sarawak, Kuching, = Heterometrus (Chersonesometrus) granulomanus
VIII.1993, 1♂, leg. J. Schneider, FKCP; Sarawak, Couzijn, 1981: 142; Tikader & Bastawade, 1983: 577;
Sebong, distr. Kapit, 9–20.III.1994, 1♂2juvs., leg. P. Kovařík, 1998: 136; Fet, 2000: 439; Kovařík, 2002: 17.
Bílek, FKCP. Sabah, Tamparuli, 3.V.1996, 2juvs., leg. Syn. n.
S. Snäll, FKCP. Philippines, Palawan, Puerto Princese,
1980, 1♀, FKCP; Mindanao Island, Misamis Occidental, TYPE LOCALITY AND TYPE REPOSITORY. India, Madras;
21.XII.1955, 2♀, leg. B. Marapao, CASC; Sulu island, BMNH.
Tawi Tawi, Tarawakan Farm District, 11.VII.1957, 1im., TYPE MATERIAL EXAMINED. India, Tamil Nadu,
CASC; Palawan island, Puerto Princesa, V–VI.1961, Madras, 2♂ (lectotype and paralectotype hereby
2♀, leg. Angel C. Alcala, CASC. Singapur, XII.1897, designated), BMNH; Palni-Hills, Kadai-canal, 1♂
3♂, leg. L. Biró, HNHM. (paratype of Heterometrus (Chersonesometrus)
granulomanus Couzijn, 1981), SMFD No. 5332.
DIAGNOSIS. Adults 90–140 mm long. Base color of
adults uniformly black, only manus and telson may be OTHER MATERIAL EXAMINED. India, Madireddy Palem,
reddish brown. Pectinal teeth number 12–18 in both 11.VII.1966, 1im., 20.XI.1966, 1♂1♀2juvs., 21.XI.1966,
sexes. Male with chela, femur and patella of pedipalp 1juv., 1.I.1967, 1♂2♀3ims., 16.I.1967, 1♂3juvs., leg.
narrower and longer than in female. Male fingers of Dorothy E. Johnson, CASC, 1.I.1967, 1♂1♀, FKCP;
pedipalp very long. Fixed fingers longer or at least as Andhra Pradesh, Addanki env., 1♂, FKCP; Podili,
long as manus of pedpalp. Chela not lobiform in male, VII.1966, 1juv., 6.VII.1966, 1juv., 20.VII.1966,
slightly lobiform in female, its length to width ratio 3.3– 1♂1im.1juv., 21.VII.1966, 1juv., 23.XI.1966, 1im.,
4.4 in male, ca. 2.4. in female. Much of manus smooth, 1.I.1967, 1♀, 13.II.1967, 1juv., leg. D. E. Johnson,
sparsely tuberculate. Patella of pedipalp with pronounced CASC. Tamil Nadu, Madras area, 1965, 2♂3♀, CASC;
internal tubercle. Carapace usually with disc smooth and Vellore, 11.VIII.1966, 2♂, 26.IX.1966, 1im.,
margins granulate, but sometimes entire surface 16.VIII.1967, 1♂, 15.IX,1966, 1♂, 24.IX.1966, 2♂,
granulate. Telson hirsute, elongate, vesicle as long as or 25.IX.1966, 1♂1♀, VIII.1967, 2juvs., leg. C.
longer than aculeus. Varadarajan, CASC; Madras (Chennai), II.1993, 2♀1im.,
leg. M. Veselý, FKCP, near Vellore, 30.X.1997, 1♂, leg.
COMMENTS. This species is known under a number of K. Werner, FKCP.
older names and as many as eight subspecies. I have
studied the paratypes of H. l. belitungensis Couzijn, DIAGNOSIS. Adults 70–100 mm long. Color of adults
1981 and specimens from nearly the entire area of uniformly reddish brown to black, legs and telson yellow
known distribution of the species, and have not found to yellowish or reddish brown, always lighter colored
any morphological differences indicating the existence than body. Pectinal teeth number 12–18 in both sexes.
of any subspecies. I have examined also CASC Male has longer femur and patella of pedipalp than
specimens from the type locality of H. l. tarawakanensis female. Chela lobiform, in male narrower than in female,
Couzijn, 1981 and have not found them to differ in any its length to width ratio 1.7–1.9 in females and 2.0–2.5 in
way from the nominotypical subspecies. However, I males. Entire manus covered by large, rounded granules
have not studied the types of all the subspecies. but devoid of carinae. Patella of pedipalp without
28 Euscorpius — 2004, No. 15

Figure 20: Heterometrus madraspatensis, dorsal view, male lectotype.

 Kovařík: Review of Genus Heterometrus 29

pronouced internal tubercle. Carapace of juveniles and COLORATION: The color is uniformly reddish brown,
some females with disc smooth and margins granulate only the telson and leg tarsomeres are reddish yellow.
and anteriorly tuberculate. Adult males and some females Also the sternites are lighter colored, and the pectens are
with entire carapace sparsely granulated. Third and yellow.
fourth leg spine formula 4:5–6. Telson hirsute, vesicle asMESOSOMA AND CARAPACE: The mesosoma lacks a
long as or longer than aculeus. carina. Both the mesosoma and carapace are smooth
except for minute granules on the margins and low
COMMENTS. Couzijn (1981: 133) suggested that tubercles on the anterior part of the carapace. The
Palamnaeus fulvipes madraspatensis Pocock, 1900 is a sternites are smooth, entirely without granules. The
synonym of H. fulvipes. However, Tikader & Bastawade pectinal teeth number 16 and 17.
(1983: 630) correctly elevated this subspecies to a METASOMA AND TELSON: The metasoma is smooth and
species. The two species are distinguishable on the shape hirsute. The first to fourth segments bear eight carinae,
of the male chela (see Fig. 25 in Pocock, 1900: 88), but of which the ventral are smooth and the dorsal consist of
problems may arise in identification of the females. minute, rounded granules. The fifth segment bears five
Couzijn (1981: 142) also described a new species, complete carinae and two additional lateral carinae
Heterometrus (Chersonesometrus) granulomanus, which developed only in the anterior half. The three ventral
in my opinion is a synonym of H. madraspatensis carinae of this segment consist of pointed granules and
Pocock, 1900. The lectotype of H. madraspatensis the spaces between them are finely granulate. The dorsal
Pocock, 1900 is hereby designated in order to stabilize carinae consist only of minute, rounded granules. The
the nomenclature. telson is hirsute, dorsally without granules, and ventrally
with four rows of granules. The vesicle is as long as the
DISTRIBUTION. India: Andhra Pradesh and Tamil Nadu aculeus.
(Pocock, 1900: 89). PEDIPALPS: The femur has granulose carinae and its
dorsal surface is slightly granulated. The patella is
Heterometrus mysorensis sp. n. smooth, dorsally and externally slightly tuberculate, with
(Fig. 21) four smooth carinae and on the internal surface with
scattered pointed granules substituting for two additional
TYPE LOCALITY AND TYPE REPOSITORY. India, Mysore, carinae; they do not form a conspicuous internal tubercle
Maddur env., cca 900 m; FKCP. otherwise characteristic for the genus. The chela is
TYPE MATERIAL. India, Mysore, Maddur env., ca. 900 slightly lobed and strongly hirsute. The dorsal side of the
m, 1983, 1♂ (holotype), collector unknown, FKCP. manus bears rounded granules of varying size, which do
not cover the entire surface and form elevated rows. The
ETYMOLOGY. Named after the state of occurrence. external surface of the chela is smooth, with two long
carinae. The movable fingers bear six straight rows of
DIAGNOSIS. Adult male 118 mm long. Female unknown. granules without internal and external granules.
Color uniformly reddish brown, only telson and leg LEGS: The legs bear solitary long and short setae. The
tarsomeres reddish yellow. Pectinal teeth number 16 and third and fourth leg spine formula is 5–6:6–7.
17. Male assumed to have femur, patella and chela of
pedipalp longer than female. Chela slightly lobed and AFFINITIES. The described features distinguish H.
strongly hirsute, its length to width ratio 2.8. Dorsal mysorensis sp. n. from all other species of the genus.
surface of pedipalp manus bears rounded granules of They are recounted in the key below. H. mysorensis sp.
varying size that form elevated rows. Patella of pedipalp n. is close to H. phipsoni, but has conspicuous rows of
without conspicuous internal tubercle. Disc of carapace granules (carinae) on the chela and only slightly
smooth and glossy, margins finely granulose and tuberculate patella of pedipalp, whereas in H. phipsoni
anteriorly slightly tuberculate. Telson hirsute, with the chela is strongly granulose.
vesicle as long as aculeus.

DESCRIPTION: The male holotype is an adult 118 mm Heterometrus nepalensis sp. n.

long. Measurements of the carapace, telson, segments of
(Fig. 22)
the metasoma and segments of the pedipalps, and
numbers of pectinal teeth are given in Table 1. Sexual
dimorphism is presumably expressed in the length of the TYPE LOCALITY AND TYPE REPOSITORY. Nepal, Royal
femur, patella and chela of pedipalp, and probably also Chitwan National Park, Island Jungle Resort; FKCP.
in the width of the chela. Although the female is not TYPE MATERIAL. Nepal, Royal Chitwan National Park,
known, morphology of other species makes this Island Jungle Resort, VI.1993, 1♂ (holotype), leg.
assumption fairly safe. Csorba; FKCP.
30 Euscorpius — 2004, No. 15

Figure 21: Heterometrus mysorensis, sp. nov., dorsal view, male holotype.
 Kovařík: Review of Genus Heterometrus 31

Figure 22: Heterometrus nepalensis, sp. nov., dorsal view, male holotype.
32 Euscorpius — 2004, No. 15

ETYMOLOGY: Named for the country of occurrence. It is AFFINITIES. The described features distinguish H.
the first species found in Nepal. nepalensis sp. n. from all other species of the genus.
They are recounted in the key below. H. nepalensis sp.
DIAGNOSIS. Adult male holotype 83 mm long. Color n. is close to H. indus but has shorter and wider
uniformly reddish black, only telson lighter colored. metasomal segments. Moreover, H. indus occurs only in
Pectinal teeth number 14–15. Sexual dimorphism in Sri Lanka, whereas H. nepalensis sp. n. is the first
proportions of pedipalps probably not noticeable. Chela species found in Nepal.
hirsute, lobiform, its adult length to width ratio 2.2.
Entire manus covered by rounded granules. Patella of
pedipalp without pronouced internal tubercle. Carapace Heterometrus petersii (Thorell, 1876)
smooth, with granules only at margins. Second (Fig. 23)
metasomal segment of male wider than long. Telson
bulbous, vesicle longer than aculeus.
Heterometrus megacephalus (nec Buthus megacephalus
C. L. Koch, 1836: 73): Simon, 1872: 97.
DESCRIPTION: The adult male holotype is 83 mm long.
Palamnaeus petersii Thorell, 1876a: 13; Pocock, 1892:
No other specimens are known. Measurements of the
39; Pocock, 1900: 84.
carapace, telson, segments of the metasoma and
Heterometrus (Heterometrus) petersi: Kovařík, 1998:
segments of the pedipalps, and numbers of pectinal teeth
137.
are given in Table 1. Sexual dimorphism in proportions
Heterometrus (Heterometrus) petersii: Fet, 2000: 437;
of pedipalps is probably not noticeable. Although the
Kovařík, 2002: 17.
female is not known, morphology of other species makes
Heterometrus petersii: Soleglad & Fet, 2003: 5, 8.
this assumption fairly safe.
Heterometrus petersii petersii: Warburg & Polis, 1990:
COLORATION: The base color of holotype is uniformly
225.
reddish black, only the telson is reddish brown. Sternites
Heterometrus (Heterometrus) petersii petersii: Couzijn,
are yellowish brown and pectens are yellow.
1981:96; Le Xuan Hue et al., 1998: 7; Fet, 2000: 437.
MESOSOMA AND CARAPACE: The mesosoma lacks
Heterometrus (Heterometrus) petersii mindanaensis
carinae. Both the carapace and mesosoma are smooth,
Couzijn, 1981: 97 (TL: Mindanao; MNHN); Kovařík,
with granules only at margins. The sternites are smooth,
1998: 137; Fet, 2000: 437.
without granules. Pectinal teeth number 14–15.
Palamnaeus spinifer (in part): Pocock, 1892: 40;
METASOMA AND TELSON: The entire surface of the
Pocock, 1893: 316.
metasoma is finely granulate and hirsute. The first to
Heterometrus longimanus (in part): Kraepelin, 1895: 34;
fourth segments bear eight carinae of which the ventral
Kraepelin, 1899: 111.
are partly smooth and the dorsal consist of conspicuous
Heterometrus longimanus petersi: Kraepelin, 1905: 199;
but sparse pointed granules. The fifth segment bears five
Giltay, 1931: 4; Fage, 1933: 25; Fage, 1936: 181; Fage,
carinae. All carinae on this segment consist of pointed
1944: 71; Takashima, 1945: 90.
granules. Apart from the just noted carinae, all
= Palamnaeus silenus Simon, 1884: 361; Flower, 1901:
metasomal segments bear two additional carinae present
35; Simon, 1904: 293 (syn. by Pocock, 1900: 84).
only in the anterior half on the fifth segment and the
Scorpio longimanus silenus: Kraepelin, 1895: 41;
posterior halves of the first to fourth segments. The first
Kraepelin, 1899: 111; Kraepelin, 1901: 271; Kraepelin,
and second segments are wider than long, whereas the
1905: 199; Masi, 1912: 123.
remaining segments are longer than wide. The telson is
hirsute and bulbous, dorsally without granules and
ventrally with four rows of granules; the vesicle is TYPE LOCALITY AND TYPE REPOSITORY. Cochinchina
longer than the aculeus. (Vietnam); MNHN.
PEDIPALPS: The femur has three inconspicuous
granulose carinae with spaces between them also bearing MATERIAL EXAMINED. Thailand, Bangkok, 1.VI.1957,
granules. The patella has four smooth carinae, is 1♂ (det. ?), leg. J. Hunyard, CASC; Prachin Buri,
ventraly smooth, dorsally and externally slightly 4.I.1966, 1♀, leg. Y. Siah, CASC. Vietnam, Quan Toi,
tuberculate, without granules or tubercles on the internal 29.IX.1969, 1♂, leg. W. Horan, CASC; Khank Hoa
surface. The dorsal surface of the manus is covered by prov., Nha Trang env., Xuan Son, 30 m, 23.I.1994, 1♀,
rounded granules. The chela is slightly lobiform and leg. K. Petrželka, FKCP; prov. Dong Nai, 80 km NE
strongly hirsute, and its internal surface is tuberculate, Saigon, valley Ma Da, Tri An dam, 10°56'N– 107°20'E,
with two carinae. The movable fingers bear six straight IV.1995, 1♂, VII.1995, 1juv., 27.IV.1996, 2ims.3juvs.,
rows of granules. 7.VI.1996, 1juv., 1997, 1♂, V.1998, 1juv., leg. K.
LEGS: The legs bear long and short solitary setae. The Petrželka, FKCP. ?, 1im., no location, CASC;
third and fourth leg spine formula is 4:5. Cochinchina, V.1872, 1♀, SMFD No. 5331.
 Kovařík: Review of Genus Heterometrus 33

Figure 23: Heterometrus petersii, dorsal view, male from Vietnam (FKCP).
34 Euscorpius — 2004, No. 15

DIAGNOSIS. Adults 90–125 mm long. Base color of TYPE MATERIAL EXAMINED. India, Tamil Nadu, Madras,
adults uniformly black, only manus and telson may be 1♂ (holotype), Sheverey Hills, 1♀ (paratype), BMNH
reddish brown. Pectinal teeth number 15–19 in both No. 1979.7.3.2,3; Nilgiri Hills, 2♀ (holotype and
sexes. Sexual dimorphism in proportions of pedipalps not paratype of Palamnaeus phipsoni collinus Pocock,
noticeable. Male has more pronounced tooth on movable 1900), leg. W. M. Daly, BMNH No. 1894.8.21.4-7.
fingers of pedipalp. Chela rounded, its length to width
ratio 2–2.4 in both sexes. Much of manus smooth, with
OTHER MATERIAL EXAMINED. India, Yercaud,
smooth carinae mainly on margins. Patella of pedipalp
6.III.1962, 1im.2juvs., leg. E. S. Ross & D. Cavagnaro,
without pronounced internal tubercle. Carapace with disc
CASC; Adhya Pradesh, 6 mi. W. Palmaner, 725 m.,
smooth and margins granulate. Telson hirsute, elongate,
26.II.1962, 1im. (det. ?) , CASC; Podili, 16.I.1967, 1♂
vesicle longer than aculeus.
(im.), leg. D. E. Johnson, CASC; Orissa, 13 mi SE
Pottangi, 550 m, 11.IV.1962, 1♂ (im.), leg. E. S. Ross &
COMMENTS. Couzijn (1981) described the Philippine D. Cavagnaro, CASC; Maharashtra, Bhiwandi, 15 mi
subspecies H. p. mindanaensis, whose holotype I have N. Bombay, XII.1964, 2♀3ims., leg. F. B. Steiner,
not been able to examine, and H. p. luzonensis, which CASC; Bombay, 1965, 2♂3♀, FKCP; Bombay,
has been synonymized with H. cyaneus (C. L. Koch, 6.III.1967, 4♂8♀12ims.24juvs., leg. S.R. Sane, CASC;
1836) (see Kovařík, 2002: 3, 17). Sexual dimorphism Tamil Nadu, south, Devala, V.1961, 5♂, leg. P. Sushai
(tooth on movable fingers of pedipalp more pronounced Nathan, CASC; Western Gháts, Pandalur env., 1998,
in male than in female) distinguishes this species from 1♂, FKCP.
H. laoticus Couzijn, 1981.
DIAGNOSIS. Adults 85–130 mm long. Color of adults
DISTRIBUTION. Cambodia (Kraepelin, 1901: 271), Laos uniformly reddish brown to black. Pectinal teeth number
(Kraepelin, 1901: 271), Philippines (Couzijn, 1981: 97) 12–16 in males and 10–15 in females. Male has longer
and Vietnam (Thorell, 1876a: 13, Kraepelin, 1899: 112). but not narrower femur and patella of pedipalp and
Thailand and Malaysia records (Kraepelin, 1901: 271) longer, usually narrower chela than female. Chela
are doubtful, as they could well pertain to H. laoticus slightly lobiform and strongly hirsute, its length to width
Couzijn, 1981 and H. spinifer (Ehrenberg, 1828). ratio 2.5–3 in males and 2.2–2.5 in females. Dorsal
surface of manus of pedipalp with large, rounded
granules of uneven size, which may merge but do not
Heterometrus phipsoni (Pocock, 1893) form carinae. Patella of pedipalp without internal
tubercle. Carapace smooth and glossy, only anterior part
Scorpio phipsoni Pocock, 1893: 307; Pocock, 1894: 73. slightly tuberculate and margins occasionally with sparse
Palamnaeus phipsoni: Pocock, 1900: 94. granules. Telson hirsute, elongate, vesicle as long as or
Heterometrus phipsoni: Kraepelin, 1899: 114; longer than aculeus.
Kraepelin, 1913: 166; Roewer, 1929: 612; Werner,
1936: 186; Roewer, 1943: 228; Takashima, 1945: 92; COMMENTS. Pocock (1900: 95) distinguished
Pointer, 1991: 24. Palamnaeus phipsoni collinus from Palamnaeus
Heterometrus (Scorpio) phipsoni: Kraepelin, 1901: 271. phipsoni by its shorter metasoma. The holotype female
Heterometrus (Chersonesometrus) phipsoni: Couzijn, makes this impression because it was apparently killed
1978: 330; Tikader & Bastawade, 1983: 646; Kovařík, shortly after feeding and thus has a relatively larger
1992: 185; Kovařík, 1998: 136; Fet, 2000: 442; mesosoma. However, another (paratype) female of the
Bastawade, 2002: 296. same size (metasoma and pedipalps) has a markedly
Heterometrus (Chersonesometrus) phipsoni phipsoni: shorter mesosoma.
Couzijn, 1981: 149 (in part). Couzijn (1981: 149) incorrectly regarded Palamnaeus
= Palamnaeus phipsoni collinus Pocock, 1900: 95. Syn. barberi Pocock, 1900 as a synonym of Scorpio phipsoni
n.
Pocock, 1893. These two species are very different (see
Heterometrus (Chersonesometrus) collinus: Couzijn,
diagnosis and key), as was pointed out by Tikader &
1981: 155; Tikader & Bastawade, 1983: 625; Kovařík,
Bastawade (1983: 614).
1998: 136; Fet, 2000: 441.
Palamnaeus phipsoni carnaticus (lapsus calami):
Pocock, 1900: 95. DISTRIBUTION. India: Tamil Nadu (Pocock, 1900: 95;
Pocock, 1893: 307), Kerala, Madhya Pradesh,
TYPE LOCALITY AND TYPE REPOSITORY. India, Madras; Maharashtra, West Bengal (Couzijn, 1981: 149; Tikader
BMNH. & Bastawade, 1983: 646), Orissa (first report).
 Kovařík: Review of Genus Heterometrus 35

Heterometrus rolciki sp. n. dorsally without granules and ventrally with four rows
(Fig. 24) of granules; the vesicle is as long as the aculeus.
PEDIPALPS: The femur has four granulose carinae with
TYPE LOCALITY AND TYPE REPOSITORY. India, Tamil spaces between them ventrally smooth, but other
Nadu state, Nilgiri hills, 15 km SE of Kotagiri, surfaces bear several granules. The patella is ventrally
Kunjappanai env., 76°56E 11°22N, ca 900 m; FKCP. smooth, dorsally and externally slightly tuberculate or
TYPE MATERIAL. India, Tamil Nadu state, Nilgiri hills, granulate, with four partly granulated carinae and
15 km SE of Kotagiri, Kunjappanai env., 76°56E several pointed granules on the internal surface in place
11°22N, ca 900 m, 7–22.V.2000, leg. J. Rolčík, 1♂ of granules, which do not form a conspicuous internal
(holotype) ecdysis 29.I.2001 and 30.V.2001, killed tubercle characteristic of other species. The dorsal
VIII.2004, reared by F. Kovařík, 22–30.V.1999, 1♂1im. surface of the manus is smooth and uneven to slightly
(paratypes), leg. Z. Kejval & M. Trýzna, FKCP; Tamil rugate, without granules and carinae. The chela is
Nadu, near Kotagiri, 27.X.1997, 1♀2juvs. (allotype and slightly lobiform and strongly hirsute, and its internal
paratypes), leg. Werner, FKCP. surface is tuberculate, with two carinae. The movable
fingers bear six straight rows of granules.
ETYMOLOGY: Named after Jakub Rolčík, who collected LEGS: The legs bear long and short solitary setae. The
the holotype. third and fourth leg spine formula is 4:6.

DIAGNOSIS. Adults 80–100 mm long. Base color of AFFINITIES. The described features distinguish H. rolciki
adults uniformly reddish brown to black; telson may be sp. n. from all other species of the genus. They are
lighter colored. Pectinal teeth number 10–12 in both recounted in the key below. H. rolciki sp. n. is close to
sexes. Sexual dimorphism in proportions of pedipalps not H. scaber, but its smaller size accounts for some
noticeable. Chela slightly lobiform and covered with long morphological differences.
setae, its adult length to width ratio 2.3 in both sexes.
Dorsal surface of manus of pedipalp smooth and uneven
to slightly rugate, without granules and carinae. Patella of Heterometrus scaber (Thorell, 1876)
pedipalp without conspicuous internal tubercle, but with (Fig. 25)
several small tubercles. Carapace and metasoma with
entire surface granulate. Telson hirsute and elongate, Heterometrus afer (err., non Linné, 1758): Simon, 1872:
vesicle as long as aculeus. 51 (see by Kraepelin, 1895: 58).
Pandinus scabro Thorell, 1876b: 202; new name for
DESCRIPTION: The adult male holotype is 100 mm long. Heterometrus afer sensu Simon, 1872.
Measurements of the carapace, telson, segments of the Scorpio scaber: Kraepelin, 1895: 58.
metasoma and segments of the pedipalps, and numbers Pandinus scaber: Simon, 1905: 161.
of pectinal teeth are given in Table 1. There is no Heterometrus scaber: Kraepelin, 1899: 116; Kraepelin,
noticeable sexual dimorphism in the proportions of 1913: 166; Lampe, 1918: 198; Takashima, 1945: 93;
pedipalps. Garnier & Stockmann, 1971: 13; Cloudsley-Thompson,
COLORATION: The base color of adults is uniformly 1990: 484; Hjelle, 1990: 54; Polis & Sissom, 1990: 166;
reddish brown to black, only the telson is reddish brown. Simard & Watt, 1990: 419; Bastawade, 1992: 221.
Sternites are also reddish brown to black, and the Palamnaeus scaber: Pocock, 1900: 93; Rosin & Shulov,
pectens are yellow. 1963: 566.
MESOSOMA AND CARAPACE: The mesosoma lacks Heterometrus (Scorpio) scaber: Kraepelin, 1901: 271.
carinae. Both the carapace and mesosoma have the entire Heterometrus (Chersonesometrus) scaber: Tikader &
surface granulate, more strongly in the mesosoma. The Bastawade, 1983: 619; Kovařík, 1998: 136; Fet, 2000:
sternites are smooth, without granules. Pectinal teeth 440.
number 10–12 in both sexes. Heterometrus (Chersonesometrus) scaber scaber:
METASOMA AND TELSON: The metasoma is hirsute and Couzijn, 1981: 144; Fet, 2000: 440.
has the entire dorsal surface granulate. The first to fourth Heterometrus (Chersonesometrus) scaber obscurus
segments bear eight carinae of which the ventral are Couzijn, 1981: 147 (TL: India, Maharashtra, Mathéran,
smooth and the dorsal consist of conspicuous but sparse near Bombay; MNHN); Kovařík, 1998: 136; Fet, 2000:
pointed granules. The fifth segment bears five carinae. 440.
All carinae on this segment consist of pointed granules. Heterometrus (Chersonesometrus) scaber rugosus
Apart from the just noted carinae, all segments bear two Couzijn, 1981: 146 (TL: India, Kerala, Malabar coast;
additional carinae present only in the anterior half of the MNHN); Kovařík, 1998: 137; Fet, 2000: 441.
fifth segment and the posterior halves of the first to = ? Heterometrus (Heterometrus) malapuramensis
fourth segments. The telson hirsute and elongate, Tikader & Bastawade, 1983: 533 (TL: Malapuram,
36 Euscorpius — 2004, No. 15

Figure 24: Heterometrus rolciki, sp. nov., dorsal view, male holotype.
 Kovařík: Review of Genus Heterometrus 37

Figure 25: Heterometrus scaber, dorsal view, male.

38 Euscorpius — 2004, No. 15

Poonus estate, Kozhikode distr., Kerala, India; NZSI); alive) 3ims. (paratypes – still-alive offspring of the
Kovařík, 1998: 137; Fet, 2000: 436. Syn. n. allotype), FKCP.

TYPE LOCALITY AND TYPE REPOSITORY. Bengal – type ETYMOLOGY: Named after Vladimír Šejna, who collected
locality uncertain – holotype lost, neotype from India, the types.
Kerala, Mahé designated by Couzijn (1981: 145–146);
MNHN. DIAGNOSIS. Adult male holotype 78 mm long. Other
adults 95–117 mm long. Base color of adults uniformly
MATERIAL EXAMINED. India, 18 mi.NE Mattanur, 75 m, reddish black with telson reddish brown; metasoma may
23.II.1962, 1♂, leg. E. S. Ross and D. Q. Cavagnaro, be lighter colored than body. Pectinal teeth number 13–
CASC; Karnataka, Malabar, 1♀, FKCP; Ghats, 14 in both sexes. No noticeable sexual dimorphism in
Mangalore, 3♂2♀, Battie col., BMNH No. proportions of pedipalps. Chela lobiform, its length to
1896.7.30.139-142; Mysore, 12 mi.N Balehonnur, 760, width ratio ca. 2.2 in adults of both sexes. Dorsal surface
19.II.1962, 1♀, leg. E. S. Ross and D. Q. Cavagnaro, of manus tuberculate, without carinae. Patella of pedipalp
CASC; 12 mi.E Virajpet, 850, 24.II.1962, 1♀, leg. E. S. without pronounced internal tubercle. Carapace either
Ross and D. Q. Cavagnaro, CASC; Jog Falls, 525 m., entirely granulate or disc smooth. Telson hirsute,
17.XI.1962, 1♀, leg. E. S. Ross and D. Q. Cavagnaro, bulbous, vesicle as long as aculeus.
CASC; Shimoga dist., Agumbe ghat, 2000 ft., T.R.S.N.
coll., V. 2001, 1♂2♀, VI.2004, 1♂3♀, FKCP; DESCRIPTION: The adult male holotype and female
Pondichery, Karaikal, T.R.S.N. coll., 2002, 1♀, FKCP. allotype are 78 mm long. Measurements of the carapace,
telson, segments of the metasoma and segments of the
DIAGNOSIS. Adults 100–130 mm long. Base color of pedipalps, and numbers of pectinal teeth are given in
adults uniformly reddish brown to black; manus and Table 1. There is no noticeable sexual dimorphism in the
telson may be lighter colored or manus may be black. proportions of pedipalps.
Pectinal teeth number 7–12 in both sexes. Sexual COLORATION: The base color of adults is uniformly
dimorphism in proportions of pedipalps not noticeable. reddish black, only the telson is reddish brown and the
Chela lobiform and covered with long setae, its adult metasoma may be lighter colored than the body.
length to width ratio 1.7–2.0 in both sexes. Dorsal Sternites are brown and pectens are yellow.
surface of manus of pedipalp relatively smooth, with MESOSOMA AND CARAPACE: The mesosoma lacks
effaced tubercles, without granules and carinae. Patella carinae. Both the mesosoma and carapace are slightly
of pedipalp without conspicuous internal tubercle. granulated. Sternites are smooth, without granules.
Carapace and mesosoma with entire surface granulate, Pectinal teeth number 13–14 in both sexes.
only sternites smooth. Telson hirsute, vesicle as long as METASOMA AND TELSON: The metasoma is generally
or longer than aculeus. smooth, hirsute, and without granules. The first to fourth
segments bear eight carinae of which the ventral are
COMMENTS. Couzijn (1981: 146 and 147) described two smooth and the dorsal consist of conspicuous but sparse
subspecies, H. s. obscurus and H. s. rugosus, which I pointed granules. The fifth segment bears five carinae,
have not been able to examine and therefore cannot with two additional carinae present only in its anterior
comment on their status. Neither have I been able to half. All carinae on this segment consist of pointed
study the types of H. (H.) malapuramensis Tikader & granules. The telson is bulbous, hirsute, dorsally without
Bastawade, 1983, but the characters included in their granules and ventrally with four rows of granules; the
description cause me to believe that this species is a vesicle is as long as the aculeus.
synonym of H. scaber. PEDIPALPS: The femur has three granulose carinae with
spaces between them smooth, but the dorsal surface may
DISTRIBUTION. India: Karnataka, Kerala, Maharashtra, bear several granules. The patella is smooth, dorsally
Tamil Nadu (Pocock, 1900: 93, Tikader & Bastawade, and externally slightly tuberculate, with four smooth
1983: 624; Fet, 2000: 440), Mysore, and Pondichery carinae and several pointed granules on the internal
(first report). surface in place of two additional carinae, none of which
forms a conspicuous internal tubercle characteristic of
Heterometrus sejnai sp. n. other species. The dorsal surface of the manus is
(Fig. 26) tuberculate, without carinae. The chela is slightly
lobiform and strongly hirsute, and its internal surface is
TYPE LOCALITY AND TYPE REPOSITORY. Thailand, smooth, with two carinae. The movable fingers bear six
Thaleban n. p., Satun; FKCP. straight double rows of granules.
TYPE MATERIAL. Thailand, Thaleban n. p., Satun, LEGS: The legs bear solitary long and short setae. The
X.1998, leg. V. Šejna, 1♂ (holotype), 1♀ (allotype – still third and fourth leg spine formula is 3:4.
 Kovařík: Review of Genus Heterometrus 39

Figure 26: Heterometrus sejnai, sp. nov., dorsal view, male holotype.
40 Euscorpius — 2004, No. 15

AFFINITIES. The described features distinguish H. sejnai m, 1983, 1♂, CASC; Pahang, Hutan Lipur Menchali, 16
sp. n. from all other species of the genus. They are km S of Merchang, 1VI.1983, 1♀, leg. E. S. Ross,
recounted in the key below. H. sejnai sp. n. is close to H. CASC; 16 km A of Kuala Rompin, cca 50 m.,
cyaneus but its smaller size causes some morphological 16.VII.1989, 2juvs., leg. E. S. Ross, CASC; Pulo
differences. The two species also differ in geographic Kundur, Rhio archipelago (S.E. end of Malay
distribution. Penninsula), 19.VI., 1im. , CASC; Gulf, Goh Samui
Island, w. side, coconut groves w. and s. of Aangtong
Heterometrus spinifer (Ehrenberg, 1828) Village, 9º31'15 N–99º56'22E to 9º31'15 N–99º56'37E,
(Fig. 27) 8.XI.1957, 1♂1♀1im., leg. H. A. Fehlman and R. R.
Rofen, CASC; Tung Song, 75 m., 4.VII.1962, 2juvs.,
Buthus (Heterometrus) spinifer Ehrenberg in Hemprich leg. E. S. Ross & D. Cavagnaro, CASC; Genting
& Ehrenberg, 1828: pl.1, fig. 2; Hemprich & Ehrenberg, Highland, 1100–1700 m, II–X.1990, 1♂, leg. P. Butti,
1829: 352; Hemprich & Ehrenberg, 1831: 3; Moritz & MZUF; Cameron Highlands, 1992, 2♀1juv., FKCP;
Fischer, 1980: 324; Braunwalder & Fet, 1998: 32. Pahang, Johor Endau-Rompin n. park, 100 m,
Buthus spinifer: C. L. Koch, 1837: 36. Salendang, 28.II–12.III.1995, 1♀, leg. M. Štrba & R.
Scorpio (Buthus) spinifer: Gervais, 1844a: 210; Gervais, Hergovits, FKCP; NE, Jelawang Jungle, near Dabong,
1844b: 60. 29.VI.1995, 1♀, leg. S. Bečvář, FKCP; Cameron
Palamnaeus spinifer: Pocock, 1892: 40; ? Pocock, 1894: Highlands, Ringlet, 1000 mnm, VII.2000, 1♂, leg. A.
96; Pocock, 1894: 316. Burda, FKCP. Thailand, 50 miles N. Bangkok (? error
Heterometrus longimanus: Kraepelin, 1895: 34 (in part); locality), 1.XII.1968, 1♂, CASC; Phang Nga,
Kraepelin, 1899: 111(in part); ? Kopstein, 1921: 129; 18.XI.1984, 1juv. after 3rd ecdysis, leg. P. Beron & St.
Takashima, 1945: 90 (in part). Andreev, SOFM No. 102; Phang Nga, 19.XI.1984,
Heterometrus longimanus longimanus: Giltay, 1931: 3 2juvs. after 2nd ecdysis, leg. P. Beron & St. Andreev,
(in part). SOFM No. 100; south, Betong, IV.1993, 1♂, leg. J.
Heterometrus spinifer: Karsch, 1879: 16; Ausserer, Strnad, FKCP.
1880: 466; Mahsberg, 1990: 271; Pointer, 1991: 24;
Delfosse, 1998: 30; Dupré, Lambert & Gérard, 1998: 63; DIAGNOSIS. Adults 100–135 mm long. Base color of
Robert, 1999: 19. adults uniformly black, only manus and telson may be
Heterometrus (Heterometrus) spinifer: Couzijn, 1978: reddish brown. In juveniles base color reddish brown,
330; Kovařík, 1998: 137 (in part); Fet, 2000: 438 (in telson may be yellow. Pectinal teeth number 15–19 in
part); Prendini, 2000: 44. both sexes. Sexual dimorphism in proportions of
Heterometrus (Heterometrus) spinifer spinifer (in part): pedipalps not noticeable. Chela slightly lobiform, its
Couzijn, 1981: 89; Kovařík, 1995: 203; Le Xuan Hue et adult length to width ratio 2.4–2.6 in both sexes. Much of
al., 1998: 7; Fet, 2000: 438. manus smooth, with smooth carinae forming irregular
Heterometrus (Heterometrus) spinifer solitarius reticulation. Patella of pedipalp with pronounced internal
Couzijn, 1981: 93; Vachon, 1982: 78; Kovařík, 1998:
tubercle. Carapace with disc smooth and margins
137; Fet, 2000: 438.
granulate. Telson hirsute, elongate, vesicle longer than
= Palamnaeus laevigatus Thorell, 1876b: 221;
aculeus.
Keyserling, 1885: 39 (syn. by Kraepelin, 1895: 34).
= Palamnaeus oatesii Pocock, 1900: 98; Flower, 1901:
34; Schultze, 1927: 375 (syn. by Couzijn, 1981: 89). COMMENTS. Couzijn (1981: 91) stated that the holotype
Heterometrus oatesi: Kraepelin, 1913: 167; Giltay, could not be found and had apparently been destroyed
1931: 4; Takashima, 1945: 94. during WW2. For that reason he designated a neotype.
However, the holotype was not destroyed, it is preserved
TYPE LOCALITY AND TYPE REPOSITORY. India (probably at ZMHB and I have been able to borrow it for study. It
incorrect); ZMHB. is an immature male. In accordance with ICZN Article
TYPE MATERIAL EXAMINED. India, 1♂ (im.) (holotype), 75.8, Couzijn’s neotype designation is invalid.
ZMHB No. 67. ?, 3♂2♀1 juv. without locality labels, Since it is impossible to ascertain that the BMNH
probably syntypes of Palamnaeus oatesii Pocock, 1900, specimens of Palamnaeus oatesii Pocock, 1900 really
BMNH. are syntypes, I do not designate a lectotype. The
specimens were apparently first kept dry and then
OTHER MATERIAL EXAMINED. Malaysia, Malay transferred to alcohol with the determination label
Penninsula, Pulo Aov, Endan River, Johare, 30.VI.1902, Palamnaeus oatesii, but without any other data. No
1im., CASC; Pahang, 400 ft., 1F, leg. Nevnay, BMNH other specimens so labeled could found at BMNH.
No. 1932.6.14.2 and No. 79; 1.X.1936, 1♂, Terry The distribution of this species can be verified only
Taylor, CASC; Perak, 10–20 km Ne of Tapah, 152–610 for Malaysia and southern Thailand. Other records are
 Kovařík: Review of Genus Heterometrus 41

Figure 27: Heterometrus spinifer, dorsal view, male, syntype of Palamnaeus (= Heterometrus) oatesii.
42 Euscorpius — 2004, No. 15

misidentifications that concern primarily H. cimrmani = Scorpio lucidipes Simon, 1885: 38 (syn. by Pocock,
sp. n. and H. petersii. 1890: 237).

DISTRIBUTION. ? Cambodia, Malaysia, Thailand, ? TYPE LOCALITY AND TYPE REPOSITORY. East Indies;
Vietnam (Couzijn, 1981: 89–90). The type locality MNHN.
„India“ must be regarded as erroneous. Couzijn (1981:
93) lists one female (the holotype of H. spinifer MATERIAL EXAMINED. India, Karnataka, Malabar,
solitarius Couzijn, 1981) from Sri Lanka, but also that Puddapoddy, 1♀, leg. O. Lotichius, SMFD No. 5333;
record should be regarded as doubtful. Malabar-Küste, 1♂1♀, SMFD No. 8885/227; Lanooli,
1911, 1♀1juv., leg. Low-Beck, SMFD No. 5320;
Madhya Pradesh, Muki Balagath, 27.VIII.1957, 1♀,
Deutsche Indien-Expedition 1955–57, ZMUH No. 688;
Heterometrus swammerdami Simon, 1872
Mysore, Shimoga dist., Agumbe ghat, 2000 ft., T.R.S.N.
(Fig. 28) coll., V. 2001, 1♀, FKCP; Orissa, Daitari, Jaypur-
Keonýahr District, 3.XII,1967, 1♀, leg. G. Topál,
Heterometrus swammerdami Simon, 1872: 56; HNHM No. 1011; Pondichery, Karaikal, III.1962, 1juv.,
Kraepelin, 1899: 112; Werner, 1902: 601; Kraepelin, IV.1962, 1♂2ims., VI,1962, 4juvs., VII,1963, 1juv., leg.
1913: 166; Borelli, 1915: 463; Lampe, 1918: 198; P. Sushai Nathan, CASC; Karaikal, T.R.S.N. coll., 2001,
Roewer, 1929: 612; Werner, 1934: 278; Werner, 1936: 5♂6♀, 2002, 2♂3♀6ims., 2003, 1♂1♀, X.2003–
186; Roewer, 1943: 228; Takashima, 1945: 91; V.2004, 2ims., FKCP. Tamil Nadu, Madras, 1889, 1♂,
Minnocci, 1974: 38; Hjelle, 1990: 49; Root, 1990: 367; leg. Th. Kolb, SMFD No. 5334; Madras, 2.II.1993, 1♀,
Warburg & Polis, 1990: 229. leg. M. Veselý, FKCP; Podhigai Hills, Papanasam,
Heterometrus swammerdammi: Pointer, 1991: 24. Tirunelveli Dist., 2000 ft., T.R.S.N. coll., V.2004, 1im.,
Scorpio swammerdami: Simon, 1885: 39; Pocock, 1890: FKCP. Sri Lanka, 1♂2ims., leg. Nonfried, NMPC; Kala
237; Kraepelin, 1895: 42; Pocock, 1893: 304; Lönnberg, Werva, 1896, 1im., leg. Madarász G. HNHM; Ceylon,
1897: 186. Paradenya, 2juvs., No. SMFD 8853/195; Ceylon, Estata
Heterometrus (Scorpio) swammerdami: Kraepelin, 1901: Beriturdi, 24.I.1914, 1juv., leg. J. Mastbaum, SMFD No.
271. 5304; North Western prov., Puttalam, IV.1994,
Pandinus swammerdami: Thorell, 1893: 379; Simon, 1♂1♀1juv. after 4th ecdysis, FKCP. ?: 1♂, MZUF;
1905: 161. Anuradhapura, I. 1889, 1juv., leg. Friasted, MZUF;
Pandinus (Heterometrus) swammerdami: Karsch, 1879: Ceylon, Kandy, 1♂, SMFD No. 6703/110;
127. Specimens born and reared in captivity (Locality Sri
Palamnaeus swammerdami: Pocock, 1900: 86; Tikader, Lanka, North Western prov., Puttalam), 1juv. after 3rd
1987: 37. ecdysis, 1juv. after 4th ecdysis, 1juv. after 5th ecdysis,
Heterometrus (Gigantometrus) swammerdami: Couzijn, 1im. after 6th ecdysis, 1♂ after 7th ecdysis, reared by F.
1978: 330; Tikader & Bastawade, 1983: 562; Sissom, Kovařík, FKCP.
1990: 131; Kovařík, 1992: 185; Kovařík, 1997: 183;
Kovařík, 1998: 137; Fet, 2000: 443; Prendini, 2000: 44; DIAGNOSIS. Adults 130–176 mm long. Base color
Bastawade, 2002: 295; Kovařík, 2002: 17; Lourenço & uniformly reddish brown to reddish black. Juveniles may
Huber, 2002: 273. be red with yellow telson. Pectinal teeth number 16–20.
Heterometrus (Gigantometrus) swammerdami swamm- Sexual dimorphism in proportions of pedipalps not
erdami: Couzijn, 1981: 162; Fet, 2000: 444. noticeable. Chela strongly lobiform, its length to width
= Heterometrus (Gigantometrus) swammerdami ratio 1.6– 1.8 in both sexes. Entire manus covered by
titanicus Couzijn, 1981: 165; Vachon, 1982: 79; large, rounded granulae that do not form true carinae.
Kovařík, 1997: 183; Kovařík, 1998: 137; Fet, 2000: 444. Patella of pedipalp without pronouced internal tubercle.
Syn. n. Carapace with disc smooth, margins and posterior
Heterometrus (Gigantometrus) titanicus Lourenço & portion granulate, and anterior portion granulate and
Huber, 2002: 273. tuberculate; occasionally entire surface sparsely
Buthus ceylonicus (non Herbst, 1800): C. L. Koch, 1841: granulate. Fifth segment of metasoma longer than femur
9, fig. 698 (Kraepelin, 1895: 42). of pedipalp, fourth segment of metasoma approximately
Buthus ceylanicus: C. L. Koch, 1850: 88. as long as femur of pedipalp. Telson bulbous, vesicle as
= Pandinus asper Thorell, 1876b: 199 (syn. by Pocock, long as or longer than aculeus.
1890: 237).
= Pandinus kochii Karsch, 1879: 127; Karsch, 1887: 68; COMMENTS. This species occurs in much of India and in
Karsch, 1892: 307; Moritz & Fischer, 1980: 316 (syn. by Sri Lanka. Couzijn (1981: 165) described the Sri Lankan
Pocock, 1890: 237). population as H. s. titanicus, and Lourenço & Huber
 Kovařík: Review of Genus Heterometrus 43

Figure 28: Heterometrus swammerdami, dorsal view, male.

44 Euscorpius — 2004, No. 15

(2002: 273) elevated this subspecies to species on COMMENTS. The taxonomy of this species has a rather
geographic grounds, without any morphological complex history. First Simon (1884: 36–38) believed it
justification. Examination of vast numbers of specimens to be P. bengalensis; then Pocock (1892: 40) described it
from both India and Sri Lanka convinces me that they all as P. thorellii and distinguished it from the very similar
are H. swammerdami. P. longimanus, but subsequently (1900) he placed it in
the synonymy of P. longimanus. This species can be
DISTRIBUTION. Sri Lanka; India: Andhra Pradesh, easily distinguished from H. bengalensis on having a
Karnataka, Madhya Pradesh, Maharashtra, Pondichery, pronounced internal tubercle on the patella of pedipalp.
Tamil Nadu, Uttar Pradesh, West Bengal (Karsch, 1892: It is more similar to H. longimanus, but has a different
307, Pocock, 1900: 87, Couzijn, 1981: 162; Tikader & geographic distribution. The morphological differences
Bastawade, 1983: 567, Fet, 2000: 444), Orissa (Kovařík, between these species are given in the key below.
1997: 183), and Mysore (first report). Although Couzijn (1981: 117) was aware of some of the
differences, he refused to accept the authenticity of the
original type series of H. bengalensis and based on it his
Heterometrus thorellii (Pocock, 1892) comb. n. new species, H. (Chersonesometrus) fastigiosus Couzijn,
(Fig. 29) 1981 (see comments under H. bengalensis). For that
reason most references to and identifications of H.
Palamnaeus bengalensis: Simon, 1884: 36–38 (360– bengalensis between 1981 and 2002 are mistaken and in
362); Simon, 1887: 112. reality pertain to H. thorellii (Pocock, 1892) comb. n.
Heterometrus (Heterometrus) bengalensis: Couzijn,
1981: 117 (in part); Kovařík, 1998: 137 (in part); Fet, DISTRIBUTION. Myanmar (Simon, 1884: 38; Pocock,
2000: 432 (in part); Kovařík, 2002: 17. 1892: 40).
Palamnaeus thorellii Pocock, 1892: 40; Laurie, 1896a:
193; Laurie, 1896b: 127; Weidner, 1959: 103.
Scorpio longimanus thorellii: Kraepelin, 1895: 40.
Heterometrus tristis (Henderson, 1919)
Palamnaeus longimanus: Pocock, 1900: 97 (in part).

TYPE LOCALITY AND TYPE REPOSITORY. Burma, now Palamnaeus tristis Henderson, 1919: 380.
Myanmar; BMNH. Heterometrus tristis: Takashima, 1945: 94.
Heterometrus (Chersonesometrus) tristis: Couzijn,
MATERIAL EXAMINED. Myanmar, Prome, 1♀ (det. 1981: 157; Tikader & Bastawade, 1983:
Couzijn, 1977 as Heterometrus bengalensis), SMFD No. 582; Kovařík, 1998: 137; Fet, 2000:442.
7986/160; Burma, Shayetunyo, 3ims. 13juvs, BMNH;
Birmania, Mandalay, 1885–1889, 1♂, leg. L. Fea, TYPE LOCALITY AND TYPE REPOSITORY. India, Tirupati
MZUF No. 1014; Bago Division, Bago Yoma, A Hills, North Arcot District, NZSI.
18.81670, E 96.08504, 10.XII.1998, 1♂1♀6ims.6juvs.,
leg. J. B. Slowinski, H. E. Robeck, and K. D. Wiseman,
COMMENTS. Unfortunately, I have not been able to
CASC; Malun, 19.VIII.1974, 1♂1♀, leg. J. Dostál,
examine this species and to include it in the key. Couzijn
FKCP; XI.1984, 1♂, FKCP; Division Magwe (Upper
(1981: 157) stated that it is similar to H. phipsoni and
Minhla), Malyn Camp., IX–X.1985, 1♂, leg. M. Veselý,
gave its size as up to 120 mm. Tikader & Bastawade
FKCP; W. Mandalay division, Bagan (Pagan)-Nyaung-
U, 29–31.V.1997, 1♂, leg. J. Rejsek, FKCP. (1983: 582) stated that the holotype male is 75.5 mm
long and the paratype female 98.5 mm long, however
DIAGNOSIS. Adults 90–120 mm long. Base color of according to Henderson (1919: 380) the holotype male is
adults uniformly brownish to black, only manus and 116 mm long and the paratype is also a male. Henderson
telson may be reddish brown. Pectinal teeth number 13– further considers H. tristis closely related to H. fulvipes
18 in both sexes. Male with chela, femur and patella of and H. gravimanus, which appears to be supported by its
pedipalp narrower and longer than in female. Chela not yellowish legs. From fig. 3 of Henderson (1919: 380) I
lobiform in male, slightly lobiform in female, its length suspect this species to be closest, or identical with, H.
to width ratio 3.8–5.5 in male, 2.6–3.1 in female. Much gravimanus Pocock, 1900, with which it shares the
of manus reticulate, sparsely punctate and with smooth shape of pedipalps and the presence of carinae on the
carinae, without granules. Patella of pedipalp with chela.
pronounced internal tubercle. Carapace usually with disc
smooth and margins granulate. Telson hirsute, elongate, DISTRIBUTION. India: Andhra Pradesh and Tamil Nadu
vesicle longer than aculeus. (Henderson, 1919: 380, Fet, 2000:442).
 Kovařík: Review of Genus Heterometrus 45

Figure 29: Heterometrus thorellii, dorsal view, male.

46 Euscorpius — 2004, No. 15

Heterometrus ubicki sp. n. carinae, of which the ventral are smooth and the dorsal
(Fig. 30) consist of minute, rounded granules. The fifth segment
bears five complete carinae and two additional lateral
TYPE LOCALITY AND TYPE REPOSITORY. India, carinae developed only in the anterior half. All carinae
Pondichery, Karaikal; FKCP. of this segment consist of minute rounded or pointed
granules and the spaces between them are finely
TYPE MATERIAL. India, Pondichery, Karaikal, T.R.S.N. granulate ventrally and smooth dorsally. The telson is
coll., 2003, 1♂1♀ (holotype and allotype), 2002 and bulbous and hirsute, dorsally without granules and
2003, 9♂2♀ (paratypes), X.2003–V.2004, 8♂15♀2ims. ventrally with four rows of granules. The vesicle is
1juv. (paratypes), FKCP; Karaikal Territory, Kurum- longer than the aculeus.
bargum P. O. Tanjore District, leg. P. Sushai Nathan, PEDIPALPS: The femur has three granulose carinae and
I.1954, 1♀ (paratype), VII.1954, 3juvs. (paratypes), its dorsal surface is slightly granulated. The patella is
12.VIII.1954, 1♀ (paratype), CASC; Karaikal, Madras, smooth, dorsally and externally slightly tuberculate, with
leg. P. Sushai Nathan, VIII.1962, 2♀ (paratypes), four smooth carinae (two of them on the external surface
IX.1962, 1♀ (paratype), X.1962, 1♀ (paratype), are often lacking), and on the internal surface with
VI,1963, 1♂ (paratype), VII,1963, 4♀ (paratypes), scattered pointed minute granules that do not form a
VIII,1963, 2♀ (paratypes), VIII,1964, 1♀ (paratype), conspicuous internal tubercle otherwise characteristic of
CASC; 1♀ (paratype) without data, CASC. the genus. The dorsal surface of the manus is
tuberculate, sometimes with indistinct smooth carinae.
ETYMOLOGY. Named after Darrell Ubick of the The chela is hirsute and its external surface is smooth,
California Academy of Sciences, San Francisco, USA, with two carinae. The movable finger bears five or six
in appreciation of his kind help. straight rows of granules and internal and external
granules.
DIAGNOSIS. Adult male holotype 89 mm long. Other LEGS: The legs bear solitary long and short setae. The
adults 77–90 mm long. Color uniformly reddish brown to third and fourth leg spine formula is 5–6:6–7.
black, legs yellow and telson yellow to reddish yellow.
Pectinal teeth number 14–17 in females and 15–19 in AFFINITIES. The described features distinguish H. ubicki
males. Male has femur, patella and chela of pedipalp sp. n. from all other species of the genus. They are
longer than female. Chela hirsute, its length to width recounted in the key below. H. ubicki sp. n. is close to H.
ratio ca. 3.3 in males and 2.2 in females. Dorsal surface thorellii (Pocock, 1892) comb. n. but has yellow legs
of manus tuberculate, without carinae. Patella of pedipalp and lacks the pronounced internal tubercle on the patella
without pronounced internal tubercle. Carapace either of pedipalp.
entirely granulate or disc smooth. Telson hirsute,
bulbous, vesicle longer than aculeus. Genital operculum
large, in female about as wide as long or slightly wider Heterometrus wroughtoni (Pocock, 1899)
than long. (Figs. 1–7, 31)

DESCRIPTION: The adult male holotype is 89 mm long, Palamnaeus wroughtoni Pocock, 1899: 745; Henderson,
other adults are 70–90 mm long. Measurements of the 1919: 381.
carapace, telson, segments of the metasoma and Heterometrus wroughtoni Pocock, 1900: 89; Takashima,
segments of the pedipalps, and numbers of pectinal teeth 1945: 93; Khatoon, 1999: 223.
are given in Table 1. Sexual dimorphism is expressed in Heterometrus (Chersonesometrus) wroughtoni: Couzijn,
the length of the femur, patella and chela of pedipalp, 1981: 158; Tikader & Bastawade, 1983: 593; Kovařík,
and also in the width of the chela. 1998: 137; Fet, 2000: 443.
COLORATION: The color is uniformly reddish brown to Heterometrus (Chersonesometrus) wrongtoni: Bast-
black, only legs are yellow and telson is yellow to awade, 2002: 296.
reddish yellow. Sternites and tergites are of the same = Heterometrus (Chersonesometrus) pelekomanus
color (reddish brown to black), and pectens are yellow. Couzijn, 1981: 137; Tikader & Bastawade, 1983: 641;
MESOSOMA AND CARAPACE: The mesosoma lacks Kovařík, 1998: 136; Fet, 2000: 440; Bastawade, 2002:
carinae, is smooth, and the carapace is either entirely 296; Kovařík, 2002: 17. Syn. n.
granulate or with disc smooth. Sternites are smooth,
entirely without granules, only the seventh sternite may TYPE LOCALITY AND TYPE REPOSITORY. India, Belgaum;
bear four smooth carinae. Pectinal teeth number 14–17 BMNH.
in females and 15–19 in males.
METASOMA AND TELSON: The metasoma is smooth and TYPE MATERIAL EXAMINED. India, Karnataka,
sparsely hirsute. The first to fourth segments bear eight Belgaum, 1♀ (lectotype) 3♂4juvs. (paralectotypes), leg.
 Kovařík: Review of Genus Heterometrus 47

Figure 30: Heterometrus ubicki, sp. nov., dorsal view, male holotype.
48 Euscorpius — 2004, No. 15

Figure 31: Heterometrus wroughtoni, dorsal view, male paralectotype.

 Kovařík: Review of Genus Heterometrus 49

W. A. Talbor, BMNH No. 1897.9.17.1-4.11.20; TYPE LOCALITY AND TYPE REPOSITORY. Kadas Tal,
Maharshtra, Deccan, Nilgiris, 2♂ (holotype and Satara, S. Dekhan; BMNH.
paratype of Heterometrus (Chersonesometrus) peleko- TYPE MATERIAL EXAMINED. India, S. Dekhan, Bombay,
manus Couzijn, 1981), SMFD No. 1088/19, Bombay, Kadas Tal (Satara), 1♂ (lectotype hereby designated)
Deccan, 1♂2♀18juvs. (paratypes of Heterometrus 1♀2juvs. (paralectotypes), leg. A.D.Wilkins, BMNH
(Chersonesometrus) pelekomanus Couzijn, 1981), No. 1896.9.26.88-981.
SMFD No. 329.
DIAGNOSIS. Adults 60–80 mm long. Color of adults
OTHER MATERIAL EXAMINED. India, Maharshtra, uniformly reddish brown, legs and telson yellow to
Decan, 1920, 2ims., FKCP; Mysore, 6 mi. NE yellowish or reddish brown, always lighter colored than
Ramanagaram, 750 m., 25.II.1962, 1im., leg. E. S. Ross body. Pectinal teeth number 13–16 in both sexes. Male
& D. Cavagnaro, CASC; Bandipur, 950 m, 18.III.1962, has longer femur and patella of pedipalp. Chela lobiform,
1im., leg. E. S. Ross & D. Cavagnaro, CASC. in male narrower, its length to width ratio 2.1 in males
and 1.6 in females. Entire manus covered by rounded
DIAGNOSIS. Adults 95–130 mm long. Color of adults minute granules that may merge but are devoid of
uniformly reddish brown. Juveniles lighter colored and carinae. Patella of pedipalp without pronouced internal
usually with yellow legs. Pectinal teeth number 13–17 in tubercle. Carapace in juveniles with disc smooth,
both sexes. Sexual dimorphism in proportions of margins granulate and anterior portion tuberculate; in
pedipalps not noticeable. Chela lobiform, its adult length adults entire carapace sparsely granulated. Vesicle of
to width ratio 1.8–2.0. Entire manus covered by rounded telson as long as or longer than aculeus.
granules which may merge and dorsally form up to five
usually incomplete rows. Patella of pedipalp without
pronouced internal tubercle. Carapace with smooth disc, COMMENTS. Couzijn (1981: 136) stated that the type
granulose margins and tuberculate anterior portion. series consists of the holotype, allotype and three
Telson hirsute, elongate, vesicle usually shorter than juveniles. However, he did not designate them, and they
aculeus. had not been designated by the original describer
(Pocock, 1897: 116) either. Consequently, in order to
COMMENTS. When describing Heterometrus peleko- stabilize the nomenclature of the species, I hereby
manus, Couzijn (1981: 139) compared it with H. fulvipes designate one of the examined syntypes as the lectotype.
which he regarded as similar to and considered H.
wroughtoni Pocock, 1900 as related to H. phipsoni (see DISTRIBUTION. India: Maharashtra (Pocock, 1897: 116).
1981: 158). He entirely diregarded differences in sexual
dimorphism of these species. The males of H. fulvipes Following is a key to the species of genus Hetero-
and H. phipsoni have longer femur, patella and chela of metrus. Consult Tables 2-3 for a synopsis of the
pedipalp than the females, whereas in H. wroughtoni this geographical distribution of these species.
dimorphism is absent. Examination of types of H.
wroughtoni and H. pelekomanus convinces me that there
is only one species involved, and that H. pelekomanus is Key to species of Heterometrus (except H. tristis)
a synonym of H. wroughtoni.
1. Dorsal surface of chela covered by pointed or rounded
DISTRIBUTION. India: Karnataka, Maharashtra (Pocock, granules .............. 2
1899: 745), Andhra Pradesh, Tamil Nadu (Couzijn, - Dorsal surface of chela smooth, may be punctate or
1981: 138, Fet, 2000: 440), Mysore (first report), and uneven but not granulate ............ 16
was cited also from Pakistan.
2. Fifth segment of metasoma longer than femur of
pedipalp, fourth segment of metasoma about as long as
Heterometrus xanthopus (Pocock, 1897) femur of pedipalps............... 3
(Fig. 32) - Fifth segment of metasoma about as long as femur of
pedipalp, fourth segment of metasoma shorter than
Palamnaeus xanthopus Pocock, 1897: 116; Pocock, femur of pedipalps ............... 4
1900: 92.
Heterometrus xanthopus: Kraepelin, 1899: 115; 3. Chela of pedipalp of adults colored similarly to body.
Takashima, 1945: 93; Khatavkar & More, 1990: 79. ................... H. swammerdami Simon, 1872
Heterometrus (Chersonesometrus) xanthopus: Couzijn, - Legs and chela of pedipalp of adults yellow to
1981: 136; Tikader & Bastawade, 1983: 609; Kovařík, yellowish or reddish brown, always lighter colored than
1998: 137; Fet, 2000: 441; Bastawade, 2002: 296. body .................. H. flavimanus (Pocock, 1900)
50 Euscorpius — 2004, No. 15

Figure 32: Heterometrus xanthopus, dorsal view, female paralectotype.

 Kovařík: Review of Genus Heterometrus 51

BO CA IN ID LA NI MA MY NE PA PH SI SL TH VI
H. barberi – – X – – – – – – – – – – – –
H. beccaloniae sp. n. – – X – – – – – – – – – – – –
H. bengalensis – – X – – – – – – – – – – – –
H. cimrmani sp. n. – – – – – – – – – – – – – X X
H. cyaneus X – – X – X – – – – X – – – –
H. flavimanus – – X – – – – – – – – – – – –
H. fulvipes – – X – – – – – – – – – – – –
H. gravimanus – – X – – – – – – – – – X – –
H. indus – – – – – – – – – – – – X – –
H. kanaraensis – – X – – – – – – – – – – – –
H. keralaensis – – X – – – – – – – – – – – –
H. laoticus – X – – X – – – – – – – – X X
H. latimanus – – X – – – – – – – – – – – –
H. liophysa – – – X – – – – – – – – – – –
H. liurus – – X – – – – – – – – – – – –
H. longimanus – – – X – – X – – – X X – – –
H. madraspatensis – – X – – – – – – – – – – – –
H. mysorensis sp. n. – – X – – – – – – – – – – – –
H. nepalensis sp. n. – – – – – – – – X – – – – – –
H. petersii – X – – X – – – – – X – – – X
H. phipsoni – – X – – – – – – – – – – – –
H. rolciki sp. n. – – X – – – – – – – – – – – –
H. scaber – – X – – – – – – – – – – – –
H. sejnai sp. n. – – – – – – – – – – – – – X –
H. spinifer – ? – – – – X – – – – – – X ?
H. swammerdami – – X – – – – – – – – – X – –
H. thorellii comb. n. – – – – – – – X – – – – – – –
H. tristis – – X – – – – – – – – – – – –
H. ubicki sp. n. – – X – – – – – – – – – – – –
H. wroughtoni – – X – – – – – – ? – – – – –
H. xanthopus – – X – – – – – – – – – – – –

Table 2: Geographical distribution of Heterometrus species: BO – Borneo (Sabah), CA – Cambodia, IN – India, ID –

Indonesia, LA – Laos, NI – Lesser Nicobar Islands, MA – Malaysia, MY – Myanmar, NE – Nepal, PA – Pakistan, PH –
Philippines, SI – Singapore, SL – Sri Lanka, TH – Thailand, VI – Vietnam.

4. Manus of pedipalp rounded and convex, transition of 6. Entire dorsal surface of adult femur and patella of
manus into fixed finger is an impressed smooth pedipalp covered by granules. Pedipalp femur of female
facet............H. latimanus (Pocock, 1894) very wide (only twice longer than wide) .................... H.
- combination not as in (4) .................. 5 beccaloniae sp. n.
- Pedipalp femur of female more than twice as long as
5. Chela with five or six carinae mainly in externolateral wide. Dorsal surface of femur with few granules, patella
part ................ 6 of pedipalp usually smooth. ........................ H.
- Chela without carinae but may bear rows of granules gravimanus (Pocock, 1894)
.................... 7
52 Euscorpius — 2004, No. 15

AP BI GO GU KA KE MP MA MY OR PO RA TN UP WB
H. barberi – – – – – – – – – – – – X – –
H. beccaloniae sp. n. – – – – – – – – – – – – X – –
H. bengalensis – – – – – – – – – X – – – – X
H. flavimanus – – – – – – – – – – – – X – –
H. fulvipes X X – X X – X X X X – X X X X
H. gravimanus – – – – – – – – – – – – X – –
H. kanaraensis – – X – X – – X – – – – – – –
H. keralaensis – – – – – X – X – – – – – – –
H. liurus – – – – – – X – – – – – – – –
H. madraspatensis X – – – – – – – – – – – X – –
H. mysorensis sp. n. – – – – – – – – X – – – – – –
H. rolciki sp. n. – – – – – – – – – – – – X – –
H. phipsoni – – – – – X X X – X – – X – X
H. scaber – – – – X X – X X – X – X – –
H. swammerdami X – – – X – X X X X X – X X X
H. tristis X – – – – – – – – – – – X – –
H. ubicki sp. n. – – – – – – – – – – X – – – –
H. wroughtoni X – – – X – – X X – – – X – –
H. xanthopus – – – – – – – X – – – – – – –

Table 3: Geographical distribution of Heterometrus species in India: AP – Andhra Pradesh, BI – Bihar, GO – Goa, GU –
Gujarat, KA – Karnataka, KE – Kerala, MP – Madhya Pradesh, MA – Maharashtra, MY – Mysore, OR – Orissa, PO –
Pondichery, RA – Rajasthan, TN – Tamil Nadu, UP – Uttar Pradesh, WB – West Bengal.

7. Legs of adults yellow to yellowish or reddish brown, 12. Male has markedly longer femur, patella and chela
always lighter colored than body……... 8 of pedipalp than female .............. 13
- Legs of adults colored similarly to body ......... 11 - Sexual dimorphism in proportions of pedipalps not
noticeable. ............... 14
8. Total length of adults less than 80 mm .............. 9
- Total length of adults more than 80 mm ............ 10 13. Granules on dorsal surface of chela do not form rows
(carinae) ..…..... H. phipsoni (Pocock, 1893)
9. Genital operculum always wider than long. Entire - Granules on dorsal surface of chela form elevated rows
adult carapace sparsely granulated. Pectinal teeth number (carinae) …….. H. mysorensis sp. n.
13–16. ............. H. xanthopus (Pocock, 1897)
- Genital operculum large, about as wide as long. Adult 14. Granules on chela do not form rows, pedipalp
carapace smooth and glossy, anteriorly may be slightly strongly hirsute. Adult color uniformly reddish black to
tuberculate and bear sparse granules at margins. Pectinal black. ..................... 15
teeth number 10–14. ........... H. liurus (Pocock, 1897) - Granules on chela form rows, pedipalp sparsely
hirsute. Adult color uniformly reddish brown.
10. Male and female chela differently shaped but of ...…........... H. wroughtoni (Pocock, 1899)
about the same width, its length to width ratio 1.7–1.9 in
both sexes............. H. fulvipes (C. L. Koch, 1837) 15. Second metasomal segment wider than long (only
- Chela narrower in male than in female, its length to male known). Nepal. ……...H. nepalensis sp. n.
width ratio 2.0–2.5 in males and 1.7–1.9 in females - Second metasomal segment longer than wide. Sri
............. H. madraspatensis Pocock, 1900 Lanka ……..... H. indus (De Geer, 1778)

11. Base color uniformly black, only manus of pedipalp 16. Male has longer and markedly narrower chela of
orangish yellow.….... H. barberi Pocock, 1900 pedipalp than female. Aso male femur and patella are
- Entire pedipalp uniformly colored .............. 12 longer. ........................... 23
 Kovařík: Review of Genus Heterometrus 53

- Sexual dimorphism in proportions of pedipalp not - Adults 110 mm long. Male chela length to width ratio
noticeable. ......................17 more than 2.5 ................................................…............
H. keralaensis Tikader & Bastawade, 1983
17. Entire carapace, mesosoma and dorsal surface of
metasomal segments granulate ..……..... 18 27. Dorsal surface of female chela smooth and punctate.
- Carapace with disc smooth or granulate, but no External surface of chela in both sexes smooth or
granules on mesosoma and dorsal surface of metasomal tuberculate. Male chela very narrow and long, its length
segments ......................19 to width ratio 3.8–5.5. .......... H. thorellii Pocock, 1892
- Dorsal surface of female chela tuberculate. External
18. Total length of adults 100 mm or less. Width of surface of chela in both sexes with pointed granules.
chela of pedipalp less than 10 mm. Adult chela length to Male chela narrow and long, its length to width ratio
width ratio 2.3 in both sexes ............... H. rolciki sp. n. 3.3–4.4. ........... 28
- Total length of adults 100–130 mm. Width of chela of
pedipalp in adults always greater than 10 mm. Chela 28. Male chela narrow and long, its length to width ratio
length to width ratio 1.7–2.0 in both sexes. ..….............. greater than 3.3. ........... H. longimanus (Herbst, 1800)
H. scaber (Thorell, 1876) – Male chela length to width ratio less than 3.1
........................... 29
19. Dorsal keels on fifth metasomal segment consist of
sparse large and pointed granules. Chela lobiform ..... 20 29. Male fingers of pedipalp very long. Fixed fingers
- Dorsal keels on fifth metasomal segment consist of longer, at least as long as manus of pedipalp
minute, less noticeable granules. Chela rounded. ..... 21 ................................ H. liophysa (Thorell, 1888)
- Male fixed fingers of pedipalp shorter than manus of
20. Dorsal surface of fingers tuberculate ............ 22 pedipalp ……... H. cimrmani sp. n.
- Much of dorsal surface of fingers smooth and punctate
............... H. spinifer (Ehrenberg, 1828)
Acknowledgments
21. Male has more pronounced tooth on movable fingers
of pedipalp than female. Carapace margins often I am grateful to the following individuals and
granulate. ...................... H. petersii (Thorell, 1876) institutions for making this study possible. Janet
- Movable fingers of pedipalp without noticeable sexual Becaloni (BMNH), Darrel Ubick and Charles Griswold
dimorphism. Carapace usually smooth, without granules. (CASC), Sarah Whitman (MZUF), Antonín Kůrka
...................... H. laoticus Couzijn, 1981 (NMPC), Ulrike Schreiber and Matt Grasshoff (SMFD),
22. Total length of adults less than 80 mm. Shahin Navai (ZMHB), and Hieronymus Dastych
..............…...... H. sejnai sp. n. (ZMUH) arranged for loans from collections in their
- Total length of adults more than 95 mm care. Stanislav Bečvář, Petr Bílek, Alexandr Burda,
.............................. H. cyaneus (C. L. Koch, 1836) František Kantner, David Král, Karel Petrželka, J.
Rejsek, Jakub Rolčík, Roman Sauer, Jan Schneider,
23. Legs of adults yellow or yellowish, always lighter Pavel Senft, Stanislav Snäll, Jan Strnad, Vladimír Šejna,
colored than body ....... H. ubicki sp. n. Petr Šrámek and Milan Veselý of the Czech Republic,
- Legs of adults colored similarly to body ............ 24 and Eduard Jendek and Ondrej Šauša of Slovakia,
passed specimens on to me.
24. Patella of pedipalp without pronouced internal tub- Jiří Zídek (Praha, Czech Republic) translated the
ercle ................... 25 text.
- Patella of pedipalps with pronouced internal tubercle The National Library of the Czech Republic
............................ 27 (International Loans Department) helped in obtaining
literature.
25. Entire dorsal surface of chelas tuberculate..................
H. bengalensis (C. L. Koch, 1841)
Dorsal surface of chelas smooth, may be slightly uneven References
and/or punctate ................... 26
AGASSIZ, L. 1846. Nomenclator zoologicus continens
26. Adults 120–150 mm long. Male chela length to nomina systematica generum arachnidarum tam
width ratio less than 2.4 ................... ............................... viventium quam fossilium. Soloduri, 1842–1846: 1–
H. kanaraensis (Pocock, 1900) 14.
54 Euscorpius — 2004, No. 15

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