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SYSTEMATICS
Revision of the Family Helotidae (Coleoptera: Cucujoidea) III: The
Thibetana Group and a Checklist of Helota MacLeay Species
CHI-FENG LEE
1
Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung, Wufeng 413, Taiwan
Ann. Entomol. Soc. Am. 102(1): 48Ð59 (2009)
ABSTRACT The thibetana species group of Helota MacLeay is revised. Five species are con-
sidered to be valid. The following synonyms are proposed: Helota sinensis Olliff, 1883 (⫽H.
tonkinensis Ritsema, 1905a); and H. thibetana Westwood, 1842 (⫽H. verrucosa Ritsema, 1894; H.
donckieri Ritsema, 1907; H. westwoodii Ritsema, 1907). Lectotypes are designated for Helota
sinensis Olliff, H. tonkinensis Ritsema, H. donckieri Ritsema, and H. westwoodii Ritsema. A key for
the thibetana species group is provided. Figures of the protibia, penis, parameres, internal sac, and
eighth abdominal tergite are given for each species. A checklist of the 23 valid species and
subspecies of the genus Helota MacLeay is provided.
KEY WORDS Helotidae, Helota, thibetana species group, taxonomic revision
Helotidae is a little-known, and originally monoge-
neric, family. Kirejtshuk (2000) divided the family into
Þve genera: Helota MacLeay, Neohelota Ohta, Afrohe-
lotina Kirejtshuk, Metahelotella Kirejtshuk, and Stro-
phohelota Kirejtshuk. Lee (2007) separated the genus
Helota into three species groups: gemmata group, vig-
orsii group, and thibetana group. This is the third
paper dealing with the genus, and the thibetana group
is the last species group requiring revision.
The thibetana group is characterized by the dense
punctures on the pronotum and serrate lateral mar-
gins of the elytra. This group is composed of nine
species, seven of which were described by Ritsema
(1894⬇1914). Species discrimination within this
group is difÞcult due to the subtle nature of the
diagnostic characters. Even Ritsema (1915b) could
not clearly separate species in his key. Moreover,
some species are rarely found in museums. This is
especially true of Helota acutipennis Ritsema, which
was described based on a single female specimen.
Fortunately, a male specimen was found at the
Naturhistorisches Museum, Basel, Switzerland
(NHMB), allowing revision of the thibetana species
group.
Materials and Methods
The descriptive terminology follows Lee and Satoˆ
(2006). The types of all known species and 39 addi-
tional specimens are examined for this article. Codens
of museums or institutions and their curators are listed
as follow: BMNH, The Natural History Museum, Lon-
don, United Kingdom (M. Barclay); EUMJ, Ehime
University, Matsuyama, Japan (N. Ohbayashi and M.
Sakai); MHNG Muse´um dÕHistoire Naturelle, Gene`ve,
Switzerland (G. Cuccodoro); MNHN, Muse´um Na-
tional dÕHistoire Naturelle, Paris, France (T. Deuve);
NHMB, Naturhistorisches Museum, Basel, Switzer-
land (M. Brancucci); NMW, Naturhistorisches Mu-
seum, Wien, Austria (M. Ja¨ch); RMNH, National
Natuurhistorische Museum, Leiden, Netherlands (F.
Assem); SMNS, Staatliches Museum fu¨r Naturkunde,
Stuttgart, Germany (W. Schawaller); and ZMHB, Mu-
seum fu¨r Naturkunde der Humboldt-Universita¨t, Ber-
lin, Germany (B. Ja¨ger).
Key to Species of Thibetana Group of Helota
1. Punctures very dense and tubercles small on
elytra (Fig. 11) ...................2
Punctures less dense and tubercles prominent
on elytra (Figs. 12 and 13) ...........3
2. Without yellowish brown spots on antero-lat-
eral angles of pronotum, black bands on bases
and middles of protibiae, and small rounded
processes near apices of ventral margins of
protibiae in males (Fig. 1); elytral apices in
females tapering (Fig. 2).............
................... H. lesnei Ritsema
With yellowish brown spots on antero-lateral
angles of pronotum, black bands on bases and
apices of protibiae, and prominent rounded
processes near apices of ventral margins of
protibiae in males (Fig. 3); elytra apices in
females acute (Fig. 4)...............
............... H. acutipennis Ritsema
3. With yellowish brown spots on antero-lateral
angles of pronotum (Figs. 7Ð10), black bands
1
Corresponding author, e-mail: cßee@gate.sinica.edu.tw.
0013-8746/09/0048Ð0059$04.00/0 䉷2009 Entomological Society of America
on middle of protibiae, and rounded pro-
cesses near apices of ventral margins of pro-
tibiae, tubercles on elytra oblong .......4
Without yellowish brown spots on antero-lat-
eral angles of pronotum (Figs. 5Ð6), without
black bands on protibiae, and angular pro-
cesses near apices of ventral margins of pro-
tibiae, tubercles on elytra elongate ......
................... H. sinensis Olliff
4. Metaventrites with one pair of clusters of stout
setae at sides in males...............
...............H. thibetana Westwood
Figs. 1–10. Dorsal habitus of species of Helota. (1) H. lesnei, male. (2) Same, female. (3) H. acutipennis, male. (4) Same,
female (holotype). (5) H. sinensis, male. (6) Same, female. (7) H. thibetana, male. (8) Same, female. (9) Same, variation, male
(lectotype of H. verrucosa). (10) H. jentinkii, male. (Online Þgure in color.)
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 49
Metaventrites without stout setae in males . . .
................. H. jentinkii Ritsema
Helota lesnei Ritsema
(Figs. 1Ð2, 14Ð19)
Helota lesnei Ritsema 1906a: 196; Ritsema 1911: 105;
Ritsema 1915a: 131; Ritsema 1915b: 232; Mader 1926:
714.
Helota lesnei: Jackobson 1915: 900; Wegrzynowicz
2000: 400; Kirejtshuk 2000: 27.
Diagnosis. Helota lesnei and H. acutipennis can be
distinguished from others of the species group by the
very dense punctures and small tubercles on the elytra
(Fig. 11). Helota lesnei differs from H. acutipennis by
the absence of yellowish brown spots on antero-lateral
angles of the pronotum and the presence of black
bands on bases and middles of protibiae (in contrast
to black bands on bases and apices in H. acutipennis);
the prominent processes near apices of the ventral
margins of protibiae in males; the pointed elytral api-
ces in females; and the narrower apex of the penis.
Male. Length 8.8 mm; width 3.7 mm. Dorsal sur-
face (Fig. 1) blackish bronze with antennae dark
brown; each elytron with two small yellow spots,
anterior ones circular, posterior ones oval. Ventral
surface metallic green with meso and metaventrites
medially brown; abdomen yellowish brown. Leg
yellowish brown, apical one third of femur, and
basal one third and middle to apical one third of tibia
dark brown, tarsus dark brown with tarsomere V
brown. Dorsal surface of head randomly and
densely punctate; ventral surface with more prom-
inent punctures. Pronotum 0.67 times longer than
wide; lateral margins rounded with strong crenula-
tions, widest at middle; surface rugose and densely
punctate, with raised patches, some of them covered
by punctures. Elytron 3.66 times longer than wide;
gradually narrowed toward apices, outer margin ser-
rate; with a tiny sutural tooth, apex moderately
rounded; disc densely punctate without visible
striae, with small and prominent tubercles (as in H.
Figs. 11–13. Left elytra of Helota species. (11) H. acutipennis, male. (12) H. sinensis, male. (13) H. thibetana, male.
50 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 1
acutipennis). Thoracic ventrites with random and
prominent punctures, those on metaventrite medi-
ally reduced, metaventrite without setae. Punctures
on abdominal ventrites very small. Protibia (Fig. 15)
slightly curved, slightly depressed dorsally near
apex of internal margin, apex forming an acute and
curved process, apex of internal margin with setae
along internal margin of hooked apex. Fifth abdom-
inal ventrite (Fig. 16) with apical margin slightly
sinuate, several pairs of long setae at sides, with
sparse short setae anteriorly and medially, apical
margin with dense long setae. Eighth abdominal
tergite (Fig. 14) transverse, apical margin truncate.
Penis (Fig. 17) slender, apex rounded, parallel and
narrowest from apex to apical one Þfth; basally wid-
ened; dense tiny setae around apices of dorsal lobes;
dorsal lobes widest near basal one Þfth, notch be-
tween connection of dorsal lobes prominent; dorsal
margin with a median notch very deep, reaching
behind apex of dorsal lobes. Parameres (Fig. 18)
elongate and gradually narrowed from base; with a
small, rounded process on middle of apical margin,
ventral surface with several moderately long setae at
apical margin near apical process, ventral surface
with dense setae; basal margin slightly convex. In-
ternal sac (Fig. 19) with one pair of slender curved
sclerites on basal one third; recurved at apical one
sixth; one basal sclerite elongate; with lateral margin
recurved.
Female (Fig. 2). Similar to males but protibiae
straight and not expanded, and elytral apices abruptly
tapering apically, forming acute apices.
Type Material. The male holotype is labeled
“MUSEUM PARIS Chine, Kouy-Tcheou Pe`re Cavale-
rie 1905/Helota Lesnei, Rits. 么type !/HOLOTYPE
(red)/Ritsema det. 1906/Muse´um Paris ex coll. R.
Oberthu¨r.”The holotype is directly pinned, is in good
condition, and is deposited in the MNHN.
Type Locality. China: Guizhou (⫽Kouy-Tcheou).
Material Examined (two specimens). CHINA.
Shandong, Qingdao (⫽Kiautschau) (1么, RMNH; 1씸,
ZMHB).
Distribution. China (Guizhou, Shandong).
Helota acutipennis Ritsema
(Figs. 3Ð4, 11, 20 Ð25)
Helota acutipennis Ritsema, 1914: 166; Ritsema 1915a:
138; Ritsema 1515b: 232; Mader 1926: 714; Wegr-
zynowicz 2000: 394; Kirejtshuk 2000: 27.
Diagnosis. See diagnosis of H. lesnei.
Male. Length 8.1 mm; width 3.4 mm. Dorsal surface
(Fig. 3) blackish bronze with antennae reddish brown
and antero-lateral angles yellowish brown; each
elytron with two small yellow spots, anterior one
larger and circular, posterior one oval. Ventral surface
metallic green with antero-lateral angles of proepis-
ternum yellowish brown, pro-, meso, and metaven-
trites medially yellowish brown; abdomen yellowish
brown. Leg yellowish brown, with apex of coxa, base
and apical one third of femur, and basal and apical one
third of tibia metallic green, tarsus dark brown.
Dorsal surface of head randomly and densely
punctate; ventral surface with more prominent
punctures. Pronotum 0.67 times longer than wide;
lateral margins rounded with strong crenulations,
widest at middle; surface rugose and densely punc-
tate, with raised patches, some of them covered by
punctures. Elytron 3.74 times longer than wide;
gradually narrowed toward apices, outer margin ser-
rate; with a tiny sutural tooth, apex moderately
rounded; disc densely punctate without visible
striae, with six longitudinal rows of carinae. Tho-
racic ventrites with random and prominent punc-
tures, medially reduced, one longitudinal cluster of
stout setae on each side of metaventrite. Punctures
on abdominal ventrites very small. Protibia (Fig. 21)
slightly curved, with a rounded process on dorsal
surface near apex of internal margin, apex forming
an acute process, apex of internal margin without
setae. Fifth abdominal ventrite (Fig. 22) with apical
margin slightly sinuate, one pair of long setae at
sides, anterior and medial surface with sparse, short
setae, apical margin with dense, long setae. Eighth
abdominal tergite (Fig. 20) transverse, apical mar-
gin truncate. Penis (Fig. 23) slender, apex truncate,
parallel and narrowest from apex to apical one
fourth; widest at basal one third; surface with dense
tiny setae, setae reduced medially and basally; dor-
sal lobes widest near basal one Þfth; notch between
connection of dorsal lobes prominent; basal margin
with a median notch very deep, reaching posterior
to apex of dorsal lobes. Parameres (Fig. 24) elongate
and parallel; a small, truncate process on middle of
apical margin, ventral surface with one pair of long
setae at apical margin near apical process, some
additional long setae on dorsal surface, ventral sur-
face with dense setae; basal margin slightly convex.
Internal sac (Fig. 25) with one pair of slender,
curved sclerites on basal third, one basal sclerite
elongate.
Female (Fig. 4). Length 8.2 mm; width 3.5 mm.
Similar to males but protibiae straight and not ex-
panded, and elytral apices tapering and acute.
Type Material. The female holotype is labeled “Yun-
nam-Sen 1911/Helota acutipennis 씸type Rits!/
HOLOTYPE/MUSEUM PARIS A. GROUVELLE
1914.”The holotype is directly pinned, is in good
condition and is deposited in the MNHN.
Type Locality. China: Yunnan.
Material Examined (one specimen). CHINA. Yun-
nan, Lijiang, Yufeng monast. 2500m, 7-VII-1990, leg.
Vit Kuba´n(1么, NHMB).
Distribution. China (Yunnan).
Helota sinensis Olliff, 1883
(Figs. 5Ð6, 12, 26 Ð31)
Helota sinensis Olliff, 1883: 54; Ritsema 1889: 111; Rit-
sema 1891: 226, 231; Ritsema 1911: 106; Ritsema
1915a: 131; Ritsema 1915b: 232; Jackobson 1915: 900;
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 51
Mader 1926: 714; Miwa 1931: 60; Koˆno 1939: 157;
Wegrzynowicz 2000: 403; Kirejtshuk 2000: 27.
Helota tonkinensis Ritsema, 1905a: 119; Ritsema 1905b:
217; Ritsema 1911: 106; Ritsema 1915a: 131; Ritsema
1915b: 232; Wegrzynowicz 2000: 405; Kirejtshuk
2000: 27. syn. nov.
Diagnosis. Helota sinensis is similar to H. thibetana
and H. jentinkii in possessing less dense punctures and
prominent tubercles on the elytra, but it differs by
lacking yellowish brown spots on anterio-lateral an-
gles of pronotum, the brown protibiae lack darker
bands and possess angular processes near the apices of
the ventral margins.
Male. Length 6.8 Ð7.6 mm; width 2.8Ð3.2 mm. Dorsal
surface (Fig. 5) blackish bronze with last three an-
tennomeres reddish brown; each elytron with two
small yellow spots oblong and obliquely directed. Ven-
tral surface dark reddish brown. Leg reddish brown or
yellowish brown, with apex and base of femur and tibia
blackish bronze.
Dorsal surface of head randomly and densely
punctate; ventral surface with more prominent
punctures. Pronotum 0.72Ð0.74 times longer than
wide; trapezoidal, lateral margins rounded with
strong crenulations, narrowed at apex; surface rug-
ose and densely punctulate, with raised patches
Figs. 14–19. Diagnostic characters of H. lesnei. (14) Eighth abdominal tergite. (15) Protibia. (16) Fifth abdominal ventrite.
(17) Penis. (18) Parameres. (19) Internal sac. Scale bar ⫽0.5 mm.
52 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 1
covered by dense punctures. Elytron 3.54- 3.72
times longer than wide; subparallel, outer margin
serrate; with a tiny sutural tooth, apex moderately
rounded; with two striae visible, stria one and two
visible from middle to apex, dense punctures out-
side stria 2, three carinae distinct, carina near suture
visible only near apex, other carinae broken by
prominent punctures. Thoracic ventrites with ran-
dom and prominent punctures; metaventrite with-
out setae. Punctures on abdominal ventrites very
small. Protibia (Fig. 27) slightly curved, with an
angular process on dorsal surface near apex of in-
ternal margin ventrally and apically depressed, apex
of internal margin with rows of very short setae.
Metatibia with rows of long setae at apex. Fifth
abdominal ventrite (Fig. 28) with apical margin
slightly trisinuate, several pairs of very long setae at
sides of semicircular area of dense long setae along
apical margin. Eighth abdominal tergite (Fig. 26)
slightly transverse, apically truncate. Penis (Fig. 29)
slender, apically tapering, apex rounded; internal
margin pointed between dorsal lobes, gradually ta-
pering medially; dorsal lobes widest near basal third,
projecting from penis; notch between connection of
dorsal lobes prominent; basal margin with a median
notch, reaching base of dorsal lobes. Parameres
Figs. 20–25. Diagnostic characters of H. acutipennis. (20) Eighth abdominal tergite. (21) Protibia. (22) Fifth abdominal
ventrite. (23) Penis. (24) Parameres. (25) Internal sac. Scale bar ⫽0.5 mm.
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 53
elongate (Fig. 30) and parallel; middle of apical
margin with a weak notch, apical processes very
widely rounded, with long setae at apical margin,
ventral surface with coarse setae; basal margin
slightly convex. Internal sac (Fig. 31) with one basal
sclerite expanded laterally.
Female (Fig. 6). Length 7.9 mm; width 3.1 mm.
Similar to males but protibiae straight and lacking
depression, and elytral apices apically tapering and
divergent, outer margin of elytra more weakly serrate.
Type Material. The male lectotype of Helota sinensis
Olliff, here designated to preserve and make more
universal use of this name, is labeled “Co-Type/Cat
No. 21a/China/Bowring/Helota sinensis Olliff, 么Co-
type”. It is pinned directly and in good condition, and
is deposited at the RMNH. A female paralectotype is
labeled “n. China. Chan-rong./Ex Musaeo Van de Poll
1909/Helota sinensis (Type) Olliff/Muse´um Paris ex
Coll. R. Oberthu¨r 1952/LECTOTYPUS Helota sinensis
Olliff det. P. Wegrzynowicz”(MNHN). The lectotype
designated by Wegrzynowicz is not valid because it is
not published.
The male lectotype of Helota tonkinensis Ritsema,
here designated to preserve and make more universal
use of this name, is labeled “H
t
Tonkin N. O. de Bao
Lac Dr Battrarel 1897Ð1898/Helota tonkinensis type 么
Rits./Muse´um Paris ex Coll. R. Oberthu¨r 1952/Rit-
sema vidit 1905/LECTOTYPUS Helota tonkinensis Rit-
sema det. P. Wegrzynowicz”. It is pinned directly and
in good condition, and is deposited at the MNHN. One
Figs. 26–31. Diagnostic characters of H. sinensis. (26) Eighth abdominal tergite. (27) Protibia. (28) Fifth abdominal
ventrite. (29) Penis. (30) Parameres. (31) Internal sac. Scale bar ⫽0.5 mm.
54 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 1
female paralectotype (MNHN) has the same data as
lectotype. The other female paralectotype is labeled
“Co-Type/Cat No. 22a/H
t
Tonkin N. O. de Bao Lac Dr
Battrarel 1897Ð1898/Helota tonkinensis type 씸; Rits./
Reni Oberthur Tonkin”(RMNH).
Type Locality. China: Shandong.
Materials examined (13 specimens). CHINA. Shan-
dong (⫽Kiautschau), (3么么, ZMHB; 1么,1씸, RMNH);
CHINA. Yunnan. Vallis Flumin, Soling-ho (4么么,1씸,
ZMHB; 1么, MHNG; 1么,1씸, NMW).
Distribution. China (Shandon, Yunnan), Vietnam.
The record from Taiwan (Miwa 1931) is false (see Lee
and Satoˆ2007).
Helota thibetana Westwood, 1842
(Figs. 7Ð9, 13, 32Ð37)
Helota thibetana Westwood, 1842: 123.
Helota thibetana Chapuis 1876: 18; Olliff 1883: 54; Rit-
sema 1891: 226, 231; Ritsema 1894: 104; Ritsema
1911: 106; Ritsema 1915a: 138; Ritsema 1915b: 232;
Jackobson 1915: 900; Mader 1926: 714; Wegrzynow-
icz 2000: 404; Kirejtshuk 2000: 27.
Helota Mellii Westwood 1848: 86; Ritsema 1889: 110.
Helota mellyi (sic!): Olliff 1883: 54; Kirejtshuk 2000: 27.
Helota verrucosa Ritsema, 1894: 98; Ritsema 1905b: 217;
Ritsema 1911: 106; Ritsema 1915a: 138; Ritsema
1915b: 232; Wegrzynowicz 2000: 405; Kirejtshuk
2000: 27. syn. nov.
Helota Donckieri Ritsema, 1907: 213; Ritsema 1911: 105;
Ritsema 1915a: 131; Ritsema 1915b: 232; Mader 1926:
714. syn. nov.
Helota donckieri: Jackobson 1915: 900; Wegrzynowicz
2000: 396.
Helota donkieri (sic!): Kirejtshuk 2000: 27.
Helota Westwoodii Ritsema, 1907: 216. syn. nov.
Helota Westwoodi (sic!): 1911: 106; Ritsema 1915a: 131;
Ritsema 1915b: 232 [key].
Helota westwoodi (sic!): Jackobson 1915: 900; Kirejt-
shuk 2000: 27.
Helota westwoodii: Wegrzynowicz 2000: 406 [world
catalog].
Diagnosis. Helota thibetana is very similar to H.
jentinkii in general appearance, but differs by the pres-
ence of one cluster of stout setae on each metaven-
trite.
Male. Length 7.7Ð7.8 mm; width 3.1Ð3.4 mm. Dorsal
surface (Fig. 7) blackish bronze with antennae red-
dish brown and antero-lateral angles yellowish brown;
each elytron with two small yellow spots, anterior ones
circular, posterior ones oval. Ventral surface yellowish
brown with proepisternum basally darkened, margins
surrounding coxae, metepisternum, and elytral epi-
pleuron darkened, mesepisternum and mesepimeron
metallic green. Leg yellowish brown, with apex of
femur and base of tibia metallic green, tarsomere IÐIV
and apical half of tarsomere V darkened, tibia with one
oblique dark band at middle.
Dorsal surface of head randomly and densely punc-
tate; ventral surface with more prominent punctures.
Pronotum 0.67Ð0.71 times longer than wide; lateral mar-
gins rounded with strong crenulations, widest at middle;
surface rugose and densely punctate, with raised patches
covered by dense punctures. Elytron 3.54Ð3.56 times
longer than wide; subparallel, outer margin serrate, with
a tiny sutural tooth, apex moderately rounded; with two
striae visible, stria one and two visible from middle to
apex, dense punctures outside stria 2, four carinae dis-
tinct, carina near suture visible only near apex, other
carinae broken by prominent punctures. Thoracic ven-
trites with random and prominent punctures, medially
reduced, one pair of longitudinal clusters of stout setae
on metaventrite. Punctures on abdominal ventrites very
small. Protibia (Fig. 33) slightly curved, with a small
rounded process near apex of ventral margin. Fifth ab-
dominal ventrite (Fig. 34) with apical margin slightly
sinuate, several long setae at sides, dense short setae
along apical margin. Eighth abdominal tergite (Fig. 32)
transverse, apical margin truncate. Penis (Fig. 35) wide,
abruptly narrowed near apex, apex rounded; internal
surface with dense tiny setae, medially and apically re-
duced; lateral margin medially narrowed; dorsal lobes
abruptly widened from basal fourth, widest near basal
third, projecting from penis; notch between connection
of dorsal lobes prominent; basal margin with a median
notch very deep, reaching posterior to apex of dorsal
lobes. Parameres (Fig. 36) elongate and parallel; a small
and rounded process on middle of apical margin, ventral
surface with several long setae at apical margin near
apical process, some additional long setae on dorsal sur-
face, ventral surface with dense setae; basal margin
slightly convex. Internal sac (Fig. 37) with one pair of
slender and curved sclerites on basal third, one basal
sclerite widened.
Female (Fig. 4). Length 8.3 mm; width 3.4 mm.
Similar to males but protibiae straight and lacking
expansion, and elytra apices abruptly and apically ta-
pering, divergent, apices narrowly rounded.
Variation. Size of punctures on the elytra is very
variable, some individuals with less impressed punc-
tures were described as H. westwoodii, and some in-
dividuals with more impressed punctures are de-
scribed as H. verrucosa.
Type Material. The female holotype of Helota mel-
lii Westwood is labeled “Typus (red)/Mellii West-
wood Simlah Thibet Iyfre/Helota thibetana Wes.
(⫽Melli Wes.) 씸”. It is pinned directly and in good
condition, and is deposited at the MHNG. Wegr-
zynowicz (2000) indicated that the type locality is
India, but that is incorrect. In addition, types of
Helota thibetana cannot be found at the BMNH as
indicated by Wegrzynowicz (2000). It is very pos-
sible that the type of Helota mellii and H. thibetana
are the same based to same label data indicated in
the original descriptions.
The male holotype of Helota verrucosa Ritsema (au-
tomatically Þxed by monotypy), is labeled “Birmah
Ruby M
es
/Doherty/Fry Coll. 1905.100/Helota verru-
cosa, 么type Rits./65151/Type (circle, with red mar-
gin)”. It is pinned directly and in good condition, and
is deposited at the BMNH.
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 55
The male lectotype of Helota donckieri Ritsema, here
designated to preserved and make more universal the use
of this name, is labeled “Type/Cat No. 24a/Helota don-
ckieri type 么Ritsema/H. Donckier Yunnan”. It is pinned
directly and in good condition, and is deposited at the
RMNH. A female paralectotye has the same data as the
lectotype, but with catalog no. 24b (RMNH).
The female lectotype of Helota westwoodii Ritsema,
here designated to preserved and make more universal
use of this name, is labeled “Co-Type/Cat No. 25a/
British Bootang Maria Basti L. Durel/Helota westwoo-
dii 씸type Rits./Reni Oberthur British Bootan/Type
(red)”. It is pinned directly and in good condition, and
is deposited at the RMNH). A female paralectotype is
labeled “British Bootang L. Durel 1898/Helota West-
woodii 씸type Rits./Muse´um Paris ex Coll. R.
Oberthu¨r 1952/TYPE (red)/LECTOTYPUS Helota
westwoodii Ritsema det. P. Wegrzynowicz”(MNHN).
The lectotype designated by Wegrzynowicz is not
valid because it is not published.
Type Locality. China: Tibet.
Material Examined (22 specimens). CHINA. Yun-
nan (8么么, ZMHB); CHINA. Yunnan. Kunming, Xis-
han (Huatingshan), 28-V-1986, leg. S. Koiwaya (1么,
1씸, EUMJ); LAOS. Louangphrabang pr., 20⬚33Ð4⬘N
102⬚14⬘E BanSong Cha (5km W), 1,200 m, 1Ð9-V-
1999, leg. Vit Kuba´nˇ(1씸, NHMB); WEST MALAY-
SIA. Cameron Highlands, 198? (1么,1씸, EUMJ); MY-
ANMAR. Kachin. Kangfang, 2-VI-2002, leg. A. Abe
(1么, EUMJ); MYANMAR. Kachin. Hpimaw, 28-VI-
Figs. 32–37. Diagnostic characters of H. thibetana. (32) Eighth abdominal tergite. (33) Protibia. (34) Fifth abdominal
ventrite. (35) Penis. (36) Parameres. (37) Internal sac. Scale bar ⫽0.5 mm.
56 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 1
2003, leg. A. Abe (1么, EUMJ); NEPAL. Arun V.
Khandbari 1100- Arunthan, 1,300 m, 29-V-83 leg. M.
Brancucci (3么么, NHMB); NEPAL. Surkhet. N.
Surkhet 1,600Ð2,000 m, 28-V-1999, leg. W. Schawaller
(2么么, SMNS); VIETNAM. Tonkin. Tam Dao, leg.
Perrot (2么么, MHNG).
Distribution. China (Tibet, Yunnan), Nepal, Laos,
Myanmar, Bhutan, Vietnam, West Malaysia.
Helota jentinkii Ritsema
(Figs. 10 and 38Ð43)
Helota Jentinkii Ritsema, 1906: 223; Ritsema 1909: 182.
Helota Jentinki (sic!): 1911: 106; Ritsema 1915a: 131;
Ritsema 1915b: 232.
Helota jentkini (sic!): Kirejtshuk 2000: 27.
Helota jentinkii: Wegrzynowicz 2000: 404.
Diagnosis. Helota jentinkii is very similar to H. thi-
betana but differs by lacking setae on the metaventrite
and possessing a more slender penis.
Male. Length 6.8 mm; width 2.8 mm. Dorsal surface
(Fig. 10) blackish bronze with antennae brown and
antero-lateral angles yellowish brown; each elytron
with two small yellow spots, anterior ones circular,
posterior ones oval. Ventral surface yellowish brown
with proepisternum basally darkened, margins sur-
rounding coxae, mesepisternum, mesepimeron, me-
tepisternum, and elytral epipleuron metallic green.
Leg yellowish brown, with apex of femur and base of
tibia metallic green, tarsomeres IÐIV and apical half of
tarsomere V darkened, tibiae each with one oblique
dark band at middle.
Dorsal surface of head randomly and densely punc-
tate; ventral surface with more prominent punctures.
Figs. 38–43. Diagnostic characters of H. jentinkii. (38) Eighth abdominal tergite. (39) Protibia. (40) Fifth abdominal
ventrite. (41) Penis. (42) Parameres. (43) Internal sac. Scale bar ⫽0.5 mm.
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 57
Pronotum 0.70 times longer than wide; lateral margins
rounded with strong crenulations, widest at middle;
surface rugose and densely punctulate, with raised
patches covered by dense punctures. Elytron 3.80
times longer than wide; subparallel, outer margin ser-
rate, each elytron with a tiny sutural tooth, apex mod-
erately rounded; with two striae visible, stria 1 and 2
visible from middle to apex, dense punctures outside
stria 2, four carinae distinct, carina near suture visible
only near apex, other carinae broken by prominent
punctures. Thoracic ventrites with random, promi-
nent punctures, medially reduced; metaventrite with-
out setae. Punctures on abdominal ventrites very
small. Protibia (Fig. 39) slightly curved, with a
rounded process on dorsal surface near apex of inter-
nal margin, apex forming an acute process, apex of
internal margins with setae; ventral margin internally
slightly wider at apical one third. Fifth abdominal
ventrite (Fig. 40) with apical margin slightly sinuate,
several pairs of long setae at sides, anterior and median
surface with sparse short setae, apical margin with
dense long setae. Eighth abdominal tergite (Fig. 38)
transverse, apical margin truncate. Penis (Fig. 41)
slender, apex rounded, gradually widened from apex
to apical one Þfth, abruptly widened from apical one
Þfth to apical one fourth, parallel from apical one
fourth to base, surface without setae; dorsal lobes as
wide as penis from apex to middle, notch between
connection of dorsal lobes prominent; basal margin
with a very deep median notch, surpassing dorsal
lobes. Parameres (Fig. 42) elongate and widest in basal
one third; a small and rounded process on middle of
apical margin, ventral surface with one pair of long
setae at apical margin near apical process, some ad-
ditional long setae on dorsal surface, ventral surface
with dense setae; basal margin slightly convex. Inter-
nal sac (Fig. 43) with one pair of slender, curved
sclerites.
Type Material. The male holotype is labeled: “Type/
Cat No. 26a/SUMATRA MEDAN Env. De Dolok-
Baros two semestre 1905 M. Moissinac/Helota jentkini
么type Rits./H. Dockier Medan Sumatra.”It is pinned
directly and in good condition, and is deposited at the
RMNH.
Type Locality. Indonesia: Sumatra (Medan).
Material examined (one specimen). INDONESIA.
Sumatra Medan, 1914, leg. A. Grouvelle (1么, MNHN).
Distribution. Indonesia (Sumatra).
Checklist of Species Belonging to the Genus Helota
MacLeay.Kirejtshuk (2000) listed 35 species and sub-
species in the genus Helota MacLeay, although some
species had been found to be junior synonyms. The
updated checklist catalogs 23 valid species and sub-
species, including three species that have been de-
scribed recently (Lee 2007, 2008).
Gemmata group
H. fulviventris Kolbe, 1886ÑChina, Japan, Korea,
USSR
H. distincta Ritsema, 1914
H. ritsemana Heller, 1923
H. fulviventris ab. awana Ohta, 1929
H. japonica Ohta, 1929
H. gemmata Gorham, 1874ÑJapan, Korea
H. gorhami Olliff, 1883ÑChina, South Korea
H. caudata Ritsema, 1889
H. kolbei Ritsema, 1889ÑChina
H. ohbayashii Lee, 2007ÑThailand, Laos
H. schuhi Lee, 2007ÑChina
H. ventralis Ritsema, 1891ÑMyanmar, Laos, Thailand,
Vietnam
Vigorsii group
H. brancuccii Lee, 2008ÑLaos, Thailand
H. fairmairei Ritsema, 1889ÑNepal, India, Bhutan,
Myanmar
H. feai Ritsema, 1891ÑMyanmar, Laos, Thailand
H. longipes Ritsema, 1889ÑIndia, Bhutan, Myanmar
H. rouyeri Ritsema, 1906ÑIndonesia (Sumatra)
H. scintillans Olliff, 1884ÑIndonesia (Java)
H. pasteuri Ritsema, 1893
H. servillei Hope, 1840ÑIndia, Nepal, Bhutan, China
H. oberthueri Ritsema, 1889
H. thoracica Ritsema, 1895ÑChina, Vietnam, Laos,
Thailand, Taiwan
H. feai ab. mushana Ohta, 1929
H. vandepolli Ritsema, 1891ÑChina, India, Laos, Viet-
nam, Malaysia, Indonesia
H. fruhstorferi Ritmsena, 1905
H. lugubris Ritsema, 1914
H. vigorsii vigorsii MacLeay, 1825ÑWest Malaysia,
Thailand, Indonesia (Sumatra, Java)
H. vigorsii sumatrensis Ritsema, 1909
H. vigorsii borneensis Ritsema, 1909ÑBorneo
Thibetana group
H. acutipennis Ritsema, 1914ÑChina
H. jentinkii Ritsema, 1906ÑIndonesia (Sumatra)
H. lesnei Ritsema, 1906ÑChina
H. sinensis Olliff, 1883ÑChina, Vietnam
H. tonkinensis Ritsema, 1905 syn. n.
H. thibetana Westwood, 1842ÑChina, Nepal, Laos,
Myanmar, Bhutan, Vietnam, West Malaysia
H. mellii Westood, 1848
H. verrucosa Ritsema, 1894 syn. n.
H. donckieri Ritsema, 1907 syn. n.
H. westwoodii Ritsema, 1907 syn. n.
Acknowledgments
I thank the curators mentioned in the text for loans of type
specimens and unidentiÞed materials. I am indebted to C.-C.
Tsung for taking photographs of the specimens. Special
thanks to C. Carlton who read the Þrst draft. This study was
supported Þnancially by the National Science Council of
Taiwan (NSC 96-2313-B-055-005-MY2).
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Received 27 June 2008; accepted 30 September 2008.
January 2009 LEE:THIBETANA SPECIES GROUP OF Helota 59
