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SYSTEMATICS
Revision of the Family Helotidae (Coleoptera: Cucujoidea) II:
The Vigorsii Group of the Genus Helota
CHI-FENG LEE
1
Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung, Wufeng 413, Taiwan
Ann. Entomol. Soc. Am. 101(4): 722Ð742 (2008)
ABSTRACT The vigorsii species group of Helota MacLeay is revised. Ten species and one subspecies
are recognized, including a new species H. brancuccii described. from Laos and Thailand. The
following new synonyms are proposed: H. vandepolli Ritsema, 1891 (⫽Helota fruhstorferi Ritsema,
1905; H. lugubris Ritsema, 1914); H. servillei Hope, 1840 (⫽Helota oberthueri Ritsema, 1889); H.
scintillans Olliff, 1884 (⫽Helota pasteuri Ritsema, 1893); Helota vigorsii vigorsii MacLeay, 1825 (⫽He-
lota vigorsii sumatrensis Ritsema, 1909). Lectotypes are designated for Helota oberthueri Ritsema, H.
feai Ritsema, H. vigorsii vigorsii MacLeay, H. vigorsii sumatrensis Ritsema, H. vigorsii borneensis
Ritsema, H. fairmairei Ritsema, and H. longipes Ritsema. A key and diagnosis for the Helota vigorsii
MacLeay species group are provided. Figures of protibia, penis, parameres, internal sac, eighth
abdominal tergite are given for all species.
KEY WORDS Helotidae, vigorsii species group, new species, taxonomic revision
Helotidae is a little-known, and originally monoge-
neric family. Wegrzynowicz (2000) presented data on
102 subspecies and species and indicated their type
depositories. Lee and Satoˆ(2006) described four new
species from Taiwan. Lee (2007) described two spe-
cies but three new synonyms were found. I consider
105 subspecies and species to be valid so at present.
Kirejtshuk (2000) divided the family into Þve gen-
era: Helota MacLeay, Neohelota Ohta, Afrohelotina
Kirejtshuk, Metahelotella Kirejtshuk, and Strophohe-
lota Kirejtshuk. Lee (2007) separated the genus Helota
into three species groups: the gemmata group, the
vigorsii group, and the thibetana group. Lee (2007)
deÞned each species group, provided a key to their
determinations and revised the gemmata species
group.
The vigorsii group is the most diverse group (13
species and two subspecies) within the genus. It has a
typical Oriental distribution. The group may be char-
acterized by each elytron having 10 striae lacking
additional punctures. All of members were described
⬎100 yr ago, and no revisionary works have been
written until the present. The modern research indi-
cates that synonyms are common within the family.
They may have existed due to individual variation and
polymorphism. Helotids are dimorphic sexually hav-
ing different shapes of the elytral apices and protobiae
in males and females. In the past, however, a number
of species were described on the basis on single spec-
imens only (including Helota rouyeri Ritsema, H.
vandepolli Ritsema, H. fruhstorferi Ritsema, H. servillei
Hope, H. scintillans Olliff, H. pasteuri Ritsema, H. feai
Ritsema, and H. vigorsii MacLeay). This often led to
description of opposite sexes as separate species. He-
lotidae possess diagnostic characters on the protibiae,
Þfth abdominal ventrite, eighth abdominal tergite, pe-
nis, parameres and internal sac. Helota thoracica Rit-
sema, one member of the group, was redescribed dur-
ing work on Taiwanese species (Lee and Satoˆ2006) by
using these diagnostic characters. Materials available
from various institutes and museums provided speci-
mens of undescribed species or the opposite sex of
species based solely on holotypes. There are now
sufÞcient diagnostic characters and materials available
so as to allow for a revision of this species group of
Helota.
Materials and Methods
The descriptive terminology follows Lee and Satoˆ
(2006). The types of all known species and 159 spec-
imens have been examined in this study. Codens of
museums or institutions and their curators are listed as
follows: BMNH, The Natural History Museum, Lon-
don, United Kingdom (M Barclay); BPBM, Bernice P.
Bishop Museum, Honolulu, HI, USA (G. A. Sameul-
son); CRS, Collection of Rudolf Schuh, Austria; EUMJ,
Ehime University, Matsuyama, Japan (N. Ohbayashi &
M. Sakai); HNHM, Hungarian Natural History Mu-
seum, Budapest, Hungary (O. Merkl); MHNG, Mu-
se´um dÕHistoire Naturelle, Geneva, Switzerland (G.
Cuccodoro); MNHN, Muse´um National dÕHistoire
Naturelle, Paris, France (A. Taghavian & T. Deuve);
MSNG Museo Civico di Storia Naturale “Giacomo
1
Corresponding author, e-mail: cßee@gate.sinica.edu.tw.
0013-8746/08/0722Ð0742$04.00/0 䉷2008 Entomological Society of America
Figs. 1–10. Dorsal habitus of species of Helota. (1) H. rouyeri, male. (2) Same, female. (3) H. vandepolli, male (lectotype
of H. fruhstorferi). (4) Same, female (lectotype). (5) H. brancuccii, male (holotype). (6) Same, female (paratype). (7) H.
servillei, male. (8) Same, female. (9) H. scintillans, male. (10) Same, female.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 723
Figs. 11–22. Dorsal habitus of species of Helota. (11) Color variation of H. scintillans, male (H. pasteuri). (12) Color
variation of H. scintillans, female (H. pasteuri). (13) H. feai, male. (14) Same, female. (15) H. vigorsii vigorsii, male. (16) Same,
female. (17) H. fairmairei, male. (18) Same, female. (19) H. thoracica, male (Hainan). (20) Same, female (Taiwan). (21) H.
longipes, male. (22). Same, female.
724 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
Doria,”Genova, Italy (R. Poggi); MTD, Museum fu¨r
Tierkunde, Dresden, Germany (O. Ja¨ger); NHMB,
Naturhistorisches Museum, Basel, Switzerland (M.
Brancucci); NMW, Naturhistorisches Museum Wien,
Austria (M. A. Ja¨ch); RMNH, Nationaal Natuurhisto-
rische Museum, Leiden, Netherlands (F. Assen);
SMNS, Staatliches Museum fu¨r Naturkunde, Stuttgart,
Germany (W. Schawaller); TARI, Taiwan Agricultural
Research Institute, Wufeng, Taichung, Taiwan; UMO,
University Museum of Natural History, Oxford, United
Kingdom (D. J. Mann); ZMHB, Museum fu¨r
Naturkunde der Humboldt-Universita¨t, Berlin, Ger-
many (B. Ja¨ger); and ZMUC, Zoological Museum,
University of Copenhagen, Copenhagen, Denmark.
Key to Species of the Vigorsii Species Group
1. Elytral apices serrate; proepisternum same color
as pronotum .....................2
Elytral apices smooth; proepisternum same color
as prosternum ....................4
2. Elytral apices strongly and remotely serrate
(Figs. 23 and 25); pronotum sparsely punctu-
ate (Figs. 1Ð4) ...................3
Elytral apices weakly and densely serrate (Fig.
27); pronotum densely punctuate (Figs. 5Ð6).
................ H. brancuccii sp. nov.
3. Dorsal surface golden bronze (Figs. 1Ð2); setae
on apical margin of parameres longer (Fig. 39);
apical and median sclerites of internal sac ab-
sent (Fig. 40) .........H. rouyeri Ritsema
Dorsal surface dark bronze (Figs. 3Ð4); setae on
apical margin shorter (Fig. 46); apical and me-
dian sclerites of internal sac present (Fig. 47) . H.
vandepolli Ritsema
4. Spots on elytra large (Figs. 9Ð12, 15Ð16) . . . 5
Spots on elytra small (Figs. 1Ð8, 13Ð14, 17Ð22).
............................. 7
Figs. 23–34. Elytral apices of Helota species. (23) H. rouyeri, male. (24) Same, female. (25) H. vandepolli, male. (26) Same,
female. (27) H. brancuccii, male. (28) Same, female. (29) H. servillei, male. (30) Same, female. (31) H. vigorsii vigorsii, female.
(32) H. fairmairei, male. (33) Same, female. (34). H. thoracica, female.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 725
5. Punctures on elytra smaller and less impressed;
elytral apices in females narrowly rounded
(Fig. 16) .......................6
Punctures on elytra larger and more impressed;
elytral apices in females acute (Figs. 10 and 12) . H.
scintillans Olliff
6. Posterior spots on elytra large, located between
striae 1Ð6 .....H. vigorsii vigorsii MacLeay
Posterior spots on elytra small, located between
striae 3Ð6 ....H. vigorsii borneensis Ritsema
7. Elytral apices in males with a small truncate pro-
cess (Figs. 11 and 13); elytral apices in females
acute (Figs. 12 and 14) ..............8
Elytral apices in males truncate without trun-
cate processes (Figs. 17 and 19, 21); elytral
apices in females rounded (Figs. 18 and 20,
22)...........................9
8. Dorsal surface reddish bronze, without pale
bands along lateral margins of pronotum (Figs.
11Ð12) ...............H. servillei Hope
Dorsal surface blackish bronze, with pale bands
along lateral margins of pronotum (Figs. 13Ð
14)..................H. feai Ritsema
9. Dorsal surface reddish bronze, without pale
bands along lateral margins of pronotum (Figs.
Figs. 35–40. Diagnostic characters of Helota rouyeri. (35) Eighth abdominal tergite. (36) Protibia. (37) Fifth abdominal
ventrite. (38) Penis. (39) Parameres. (40) Internal sac. Scale bar, 0.5 mm.
726 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
17Ð18, 21Ð22); carinae between striae 2 and 3
in males reduced (Figs. 17 and 21) ......10
Dorsal surface blackish bronze, with pale bands
along lateral margins of pronotum (Figs, 19Ð
20); carinae between striae 2 and 3 on elytra
in male well developed (Fig. 19) .......
................ H. thoracica Ritsema
10. Punctures on elytra smaller and denser (Fig.
80); elytral apices in females widely rounded
(Figs. 18 and 33) ....H. fairmairei Ritsema
Puncture on elytra larger and sparser (Fig. 81);
elytral apices in females narrowly rounded
(Figs. 22 and 31) .....H. longipes Ritsema
Helota rouyeri Ritsema
(Figs. 1, 2, 23, 24, 35Ð40)
Helota rouyeri Ritsema 1906: 131; Ritsema 1909: 181,
182; Ritsema 1911a: 106; Ritsema 1915a: 127; Ritsema
1915b: 229 [key].
Figs. 41–47. Diagnostic characters of Helota vandepolli. (41) Protibia. (42) Protibia, variation (lectotype of H. fruhstor-
feri). (43) Eighth abdominal tergite. (44) Fifth abdominal ventrite. (45) Penis. (46) Parameres. (47) Internal sac. Scale bar ⫽
0.5 mm.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 727
Helota rouyeri Wegrzynowicz 2000: 403; Kirejtshuk
2000: 27.
Diagnosis. H. rouyeri is similar to H. vandepolli with
strongly serrate elytral apices (Figs. 23 and 25), but it
differs by its golden bronze coloration (Figs. 1 and 2),
longer setae on the apical margin of the parameres
(Fig. 39), and the absence of median and apical scle-
rites of the internal sac (Fig. 40).
Male. Length 10.9Ð12.6 mm; width 3.9Ð4.7 mm. Dor-
sal surface (Fig. 1) golden bronze, antennae yellowish
brown, with last four antennomeres darkened; each
elytron with two small yellow spots between third and
sixth striae. Ventral surface yellow; proepisternum, head,
and elytral epipleuron golden bronze. Legs yellowish,
with trochanters, apical one third of femora, basal and
apical one third of tibiae, tarsomeres 1Ð4 and apical half
of tarsomere 5, blackish brown. Dorsal surface of head
with prominent punctures, denser at sides; ventral sur-
face medially reduced. Pronotum 0.74 Ð0.75 times longer
than wide, trapezoidal with weak crenulations on apical
one Þfth of lateral margins, surface rugose, with raised
patches and regular, prominent punctures. Elytra 1.83Ð
Figs. 48–53. Diagnostic characters of Helota brancuccii. (48) Eighth abdominal tergite. (49) Protibia. (50) Fifth abdominal
ventrite. (51) Penis. (52) Parameres. (53) Internal sac. Scale bar ⫽0.5 mm.
728 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
1.88 times longer than wide, gradually narrowed toward
apex, apex strongly serrate (Fig. 23); each elytron with
10 striae and three longitudinal carinae; Þrst carina be-
tween second and third striae, only visible near apices;
second weak, between sixth and seventh striae; third
between eighth and ninth striae. Thoracic ventrites with-
out punctures except for internal sides of proepisternum.
Protibia (Fig. 36) slightly curved, internal margin ven-
trally and apically depressed, with dense long setae along
apex of dorsal internal margin. Fifth abdominal ventrite
(Fig. 37) with apical margin very slightly sinuate, several
extremely long setae at sides, dense setae restricted to
apical margin. Eighth abdominal tergite (Fig. 35) square,
antero-lateral angle rounded. Penis (Fig. 38) slender,
gradually narrowed toward apex, apex rounded, mesal
margin truncate; dorsal lobes widest at apical one third,
gradually narrowed toward base, notch between con-
nection of dorsal lobes prominent; basal margin with
shallow notch. Parameres (Fig. 39) elongate, middle of
apical margin with a deep notch, with dense long setae
along apex, shorter posteriorly, ventral surface with
sparse long setae, only present near apex, basal margin
Figs. 54–59. Diagnostic characters of Helota servillei. (54) Eighth abdominal tergite. (55) Protibia. (56) Fifth abdominal
ventrite. (57) Penis. (58) Parameres. (59) Internal sac. Scale bar ⫽0.5 mm.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 729
truncate. Internal sac (Fig. 40) without apical and me-
dian sclerites, basal sclerites with lateral expansions.
Female (Fig. 2). Length 9.4Ð11.6 mm; width 3.8Ð4.7
mm. Similar to males, but protibiae without depression,
elytral apices (Fig. 24) tapering and weakly serrate.
Type Material. The holotype 씸of H. rouyeri Rit-
sema is labeled “Type/Cat No. 6a/Helota Rouyeri 씸;
type Rits./H. Rouyer Pajakombo Sumatra.”It is di-
rectly pinned and is in good condition. It is deposited
in the RMNH.
Type Locality. Indonesia: Sumatra (Pajakombo).
Materials Examined (5 Specimens). INDONESIA:
Sumatra: 1 씸; Brastagi, Tongo Nat-Park, 13-VIII-1992,
Barries & Cate, CRS; 1 么, Brastagi, III-1985, Arbaimun,
EUMJ; 1 么;2씸씸, Si-Rambe´, XII-1890⬇III-1891,
Modigliani, MSNG.
Distribution. Indonesia (Sumatra).
Helota vandepolli Ritsema
(Figs. 3, 4, 25, 26, 41Ð47)
Helota vandepolli Ritsema 1891a: 197; Ritema 1891b:
223, 231; Ritsema 1909: 181, 183; Ritsema 1911a: 106;
Ritsema 1915a: 127; Ritsema 1915b: 229.
Helota vandepolli Wegrzynowicz 2000: 405; Kirejtshuk
2000: 27.
Figs. 60–64. Diagnostic characters of Helota scintillans. (60) Eighth abdominal tergite. (61) Protibia. (62) Penis. (63)
Parameres. (64) Internal sac. Scale bar ⫽0.5 mm.
730 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
Helota fruhstorferi Ritsema 1905a: 117; Ritsema 1905b:
216; Ritsema 1911a: 105; Ritsema 1915a: 137; Ritsema
1915b: 229. syn. nov.
Helota fruhstorferi Wegrzynowicz 2000: 397; Kirejt-
shuk 2000: 27.
Helota lugubris Ritsema 1914a: 57; Ritsema 1914b: 165;
Ritsema 1915a: 137; Ritsema 1915b: 229; Wegr-
zynowicz 2000: 400; Kirejtshuk 2000: 27. syn. nov.
Diagnosis. See diagnosis of H. rouyeri.
Male. Length 9.8 mm; width 4.1 mm. Dorsal surface
(Fig. 3) dark bronze, with dark brown antenna; each
elytron with two small yellow spots between third and
sixth striae. Ventral surface yellow; proepisternum,
head, and elytral epipleuron blackish bronze. Legs
yellowish, with trochanters, apical one Þfth of femora,
basal one Þfth and apices of tibiae, and tarsi blackish
brown. Dorsal surface of head with prominent punc-
tures, denser at sides. Pronotum 0.77 times longer than
wide, trapezoidal, lateral margins with weak crenula-
tions, surface rugose, with raised patches and sparse
prominent punctures. Elytra 1.85 times longer than
wide, parallel, apex (Fig. 25) strongly serrate; each
elytron with 10 striae and three longitudinal carinae;
Þrst carina between second and third striae, only vis-
Figs. 65–69. Diagnostic characters of Helota feai. (65) Eighth abdominal tergite. (66) Protibia. (67) Penis. (68) Parameres.
(69) Internal sac. Scale bar ⫽0.5 mm.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 731
ible near apices; second between sixth and seventh
striae; third between eighth and ninth striae. Thoracic
ventrites without punctures except for proepisternum
and metepisternum and outer sides of metaventrite.
Protibia (Fig. 41) slightly curved, with angular process
near apex of internal margin; ventral internal margin
depressed at angular process, with dense long setae
along apex of dorsal internal margin. Fifth abdominal
ventrite with apical margin truncate, apico-lateral pro-
cesses projecting from lateral margins; several long
setae at sides, dense setae restricted to apical margin.
Eighth abdominal tergite (Fig. 48) trapezoidal, an-
tero-lateral angel rounded. Penis (Fig. 45) slender,
apically narrowed, apex rounded, mesal margin
emarginate; dorsal lobes widest at apical one third,
gradually narrowed toward base, notch between
connection of dorsal lobes shallow; notch on basal
margin shallow. Parameres (Fig. 46) elongate, mid-
dle of apical margin with a shallow notch, with dense
long setae along and inside apical margin, ventral
surface with sparse setae, basal margin truncate.
Internal sac (Fig. 47) with two apical, elongate and
slightly curved sclerites covered by teeth, basally
conjoined and dorsally covered by slightly wider
and elongate, toothed sclerite; three median scle-
rites, one elongate, pointed at apex and two smaller,
Figs. 70–74. Diagnostic characters of Helota vigorsii. (70) Eighth abdominal tergite. (71) Protibia. (72) Penis. (73)
Parameres. (74) Internal sac. Scale bar ⫽0.5 mm.
732 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
semicircular, toothed sclerites on its sides; one basal
sclerites with lateral extensions.
Female (Fig. 4). Length 9.8 Ð10.5 mm; width 3.9Ð 4.1
mm. Similar to male, but protibiae without depression,
elytral apices (Fig. 26) tapering and weakly serrate.
Variation. The male protibiae possess small rounded
tubercles and are bifurcate (Fig. 42). The male of this
species was previously described as H. fruhstorferi.
Type Material. The holotype 씸of H. vandepolli
Ritsema is labeled “Wahnes Borneo/Helota Vande-
polli, type 씸; Rits./Ex Musaeo Van de Poll 1909/TYPE
(red)/Muse´um Paris.”It is in good condition and is
deposited in the MNHN.
The holotype 么of H. lugubris Ritsema is labeled
“Laos Keng Kabao Prov. i.e., (Savannakhat)/Laos:
Kong Kahas prov. de Savannakit/Helota lugubris, Rits.
么;type/TYPE (red word)/MUSEUM PARIS A.
GROUVELLE 1914.”It is in good condition and is
deposited in the MNHN.
The holotype 么of H. fruhstorferi Ritsema is labeled
“Tonkin Than-Moi Juni-Juli H. Fruhstorfer/Helota
Fruhstorferi 么. type. Rits./TYPE (red)/Ritsema vidit
1905/Muse´um Paris.”It is in good condition and is
deposited in the MNHN.
Type Locality. Indonesia (Kalimantan).
Materials Examined (9 Specimens). MALAYSIA:
Sarawak: 1 씸, Cat No. 7a, Mt. Marapok Dent Province
North Borneo, Collector, RMNH; THAILAND: 1 씸,
MUSEUM PARIS LAKHON MARMANO 1878/M2
78/Helota Vandepolli, 씸, 1897, Rits./Helota lugubris씸,
1914, Rits., MNHN; VIETNAM: 1 씸, Tonkin P. Lome´e
1903Ð1906/Helota Fruhstorferi 씸. Rits, MNHN; 2 씸씸,
Tamdao, V-93, S. Nakamura, EUMJ; LAOS: Sedone: 1
씸, Pakge, 4-VI-65, native collector, BPBM; CHINA:
Guangdong: 1 씸, Lo-fau-zan, 14⬇19-V-12, Mell,
ZMHB); INIDA: 1 么,1씸, Assam Agri. Univ. 1985?, SE.
Phukan, BMNH.
Figs. 75–79. Diagnostic characters of Helota fairmairei. (75) Eighth abdominal tergite. (76) Protibia. (77) Penis. (78)
Parameres. (79) Internal sac. Scale bar ⫽0.5 mm.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 733
Distribution. China (Guangdong), India (Assam),
Laos, Thailand, Vietnam, Malaysia (Sarawak), Indo-
nesia (Kalimantan).
Helota brancuccii sp. nov.
(Figs. 5, 6, 27, 28, 48Ð53)
Diagnosis. This new species is similar to H. rouyeri
and H. vandepolli in possessing serrate elytral apices
and an unicolorous pronotum and proepisternum, but
it can be distinguished by the denser punctures on
pronotum (Figs. 5Ð6) and weakly and densely serrate
elytral apices (Fig. 27).
Male. Length 11.7 mm; width 4.3 mm. Dorsal surface
(Fig. 5) blackish bronze, with Þrst two antennomeres
yellowish brown; each elytron with two small yellow
spots between third and sixth striae. Ventral surface
yellow; proepisternum, head, and elytral epipleuron
blackish bronze. Legs yellowish, with trochanters, api-
cal one third of femora, basal one third and apices of
tibiae, and tarsi blackish brown. Dorsal surface of head
with prominent punctures, denser at sides. Pronotum
0.72 times longer than wide, trapezoidal, lateral mar-
gins with moderate crenulations, surface rugose, with
raised patches and dense prominent punctures. Elytra
1.92 times longer than wide, parallel, apex (Fig. 27)
weakly serrate; each elytron with 10 striae and three
longitudinal carinae; Þrst carina between second and
third striae, only visible near apices; second weak,
between sixth and seventh striae; third between
eighth and ninth striae. Thoracic ventrites without
punctures except for proepisternum and metepister-
num and outer sides of metaventrites. Protibia (Fig.
49) slightly curved, with angular process at apical one
third of internal margin; ventral internal margin de-
pressed at angular process, with dense long setae along
apex of dorsal internal margin. Fifth abdominal ven-
trite (Fig. 50) with apical margin very slightly bisinu-
ate, lateral margins apically tapering, several ex-
tremely long setae at sides, dense setae restricted to
apical margin. Eighth abdominal tergite (Fig. 48, bro-
ken) square, antero-lateral angle rounded. Penis (Fig.
51) slender, gradually narrowed toward apex; apex
rounded, mesal margin pointed; dorsal lobes widest at
apical one third, gradually narrowed toward base,
notch between connection of dorsal lobes prominent;
basal margin with very deep notch. Parameres (Fig.
52) elongate, middle of apical margin with a shallow
notch, with one pair of long setae near median notch,
ventral surface with sparse long setae, basal margin
truncate. Internal sac (Fig. 53) with two apical, square
sclerites covered by teeth, one pair of median,
toothed, curved sclerites and basal sclerite with lateral
expansions.
Female (Fig. 6). Length 10.6 mm; width 4.2 mm.
Similar to male, but protibiae without depression, ely-
tral apices (Fig. 28) tapering and weakly serrate.
Type Material. HOLOTYPE : 1 么, LAOS: Phong-
saly: Ban Sano Mai, 19⬇26-V-2004, 1,150 m, 21⬚21⬘N
102⬚03⬘E, P. Pachola´ko, NHMB. PARATYPES: 1 씸,
LAOS: E. Phonsavan 25 km, Xiang Khoang, 19-VI-
2005, J. Yamasako, TARI; 1 么: THAILAND: Chiang
Mai: Doi Suthep, 1⬇5-VI-1986, H. Hirasawa, EUHJ.
Type Locality. Laos: Phongsaly prov., Ban Sano Mai.
Etymology. It is named after Michel Brancucci who
provided specimens of this interesting new species for
study.
Distribution. Laos, Thailand.
Helota servillei Hope
(Figs. 7, 8, 29, 30, 54Ð59)
Helota servillei Hope, 1840: 187; Chapius 1876: 18;
Ritsema 1889: 111; Ritsema 1891b: 224, 231; Ritsema
1911a: 106; Ritsema 1915a: 127; Ritsema 1915b: 230.
Helota servillei Olliff 1883: 51; Wegrzynowicz 2000:
397; Kirejtshuk 2000: 27.
Helota oberthu¨ri Ritsema 1889: 100; Ritsema 1891b:
224, 231; Ritsema 1891c: 888; Ritsema 1893b: 140;
Ritsema 1894a: 97; Ritsema 1911a: 106 [world cat-
alog]; Ritsema 1915a: 128; Ritsema 1915b: 230.
syn. nov.
Helota oberthu¨ri Jakobson 1915: 900; Koˆno 1939: 157.
Helota oberthuri (sic!) Miwa 1931: 60; Kirejtshuk
2000: 27.
Helota oberthueri Wegrzynowicz 2000: 401.
Figs. 80–81. Left elytra of Helota species. (80) H. fair-
mairei, male. (81) H. servillei, male.
734 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
Diagnosis. H. servillei, H. scintillans, and H. feai can
be distinguished from other members of the genus by
the elytral apices of both sexes. In males the carinae
between second and third striae extend posteriorly,
from a truncate process (Fig. 29); in females elytral
apices strongly taper and extend to acute lobe (Fig.
30). H. servillei differs from both these species by its
reddish bronze coloration.
Male. Length 15.5Ð17.4 mm; width 6.1Ð 6.8 mm. Dor-
sal surface (Fig. 7): head black with granules reddish
bronze, antennae brown; pronotum reddish brown,
with raised patches black, granules inside patches red-
dish bronze; basal margins of pronotum and elytra
black, elytra reddish brown, each elytron with two
small yellow spots between third and seventh striae;
scutellum black. Ventral surface reddish brown, with
thoracic margins surrounding coxae black. Legs red-
dish brown, with bases and apices of coxae, trochan-
ters, femora, and tibiae black, apices of tarsomere 5
and claws black. Dorsal surface of head randomly and
densely punctate, ventral surface with more promi-
nent punctures, although medially reduced. Pronotum
0.77Ð0.80 times longer than wide, trapezoidal, lateral
margins with weak crenulations, narrowed at apex,
surface rugose, with raised patches and regular, prom-
inent punctures. Elytra 1.93Ð2.00 times longer than
wide, parallel, apex obliquely truncate, each elytron
with 10 striae and three longitudinal carinae; Þrst ca-
rina between second and third striae, only visible near
apices of elytra, apices of carinae projecting from
margins of elytra, second carina between sixth and
seventh striae, third between eighth and ninth striae.
Thoracic ventrites without punctures except for in-
ternal sides of proepisternum and outer sides of
metaventrite. A pair of setal clusters along median
longitudinal sulcus of metaventrite. Punctures on ab-
Figs. 82–86. Diagnostic characters of Helota longipes. (82) Eighth abdominal tergite. (83) Protibia. (84) Penis. (85)
Parameres. (86) Internal sac. Scale bar ⫽0.5 mm.
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 735
dominal ventrites very small. Protibia (Fig. 55) slightly
curved, internal margin ventrally and apically de-
pressed, with rows of short setae on dorsal apex. Fifth
abdominal ventrite (Fig. 56) with apical margin sinu-
ate, dense stout setae conÞned to semicircular area
along apical margin; three long setae on both sides at
this area. Eighth abdominal tergite (Fig. 54) elongate,
apical margin weakly emarginate at middle or trun-
cate. Penis (Fig. 57) slender, abruptly narrowed near
apex, apex truncate, mesal margin arcuate, dorsal lobes
widest at apical third, gradually narrowed toward base,
apices widely rounded, notch between connection of
dorsal lobes prominent. Parameres (Fig. 58) elongate
and apically narrowed, middle of apical margin with a
narrow notch, with several long setae along margin
behind apex, ventral surface with dense short setae,
basal margin truncate. Internal sac (Fig. 59) with a
longitudinal sclerite at apex, narrowed at apical third,
and with a longitudinal cluster of teeth at apical third,
covered posteriorly by a Y-shaped sclerite, with ven-
tral teeth on lateral expansions, a pair of median
curved sclerites provided with teeth at their apices
and bases, one basal sclerite with lateral expansions.
Female (Fig. 8). Length 14.1Ð16.1 mm; width 5.3Ð
6.0 mm. Similar to males, but protibiae without de-
pression, elytral apices (Fig. 30) abruptly narrowed,
forming acute process along Þrst carinae, metaventrite
without setae.
Type Material. The holotype 씸of H. servillei Hope
is labeled “Helota servillei F. W. H. col. man. 3 187 PL.
3 Þg. 4 Ind. or. W Poonah/TYPE COL: 174
1
/
3
Helota
servillei Hope HOPE DEPT. OXFORD/Type Co
(green)”. It is pinned directly and in good condition,
but one mandible, both maxillae, labium, and one
tarsus were dissected out and mounted on a card. It is
deposited at UMO. In addition to the holotype, two
female specimens (one represents H. longipes) are
found at the UMO. They should be excluded from the
type series due to lack of labels except type labels
(D. J. Mann, personal communication).
The lectotype 么of H. oberthueri Ritsema labeled by
Wegrzynowicz but not published, here designated to
preserve stability and make more universal use of this
name, is labeled “Darjeeling 1889/Compared with
Type/S. Olliff vidit 1884/REITTER/RITSEMA/He-
lota servillei Hope/Ritsema vidit 1896/Helota
Oberthuri, Rits. type 么/Oberthuri Rits. Type! India
(with yellow margin)/Museum Paris ex Coll.: R.
Oberthur/LECTOTYPUS Helota oberthueri Ritsema
des. P. Wegrzynowicz (with red margin, unpub-
lished).”It is pinned directly and is in good condition.
It is deposited at the MNHN. A female paralectotype
is labeled “India/Ex-Musaeo Mniszech/RITSEMA/
Ritsema 1896/Helota Oberthur, Rits. type 씸/TYPE
(red)/PARALECTOTYPUS Helota oberthueri Rit-
sema des. P. Wegrzynowicz (with red margin)/Mu-
seum Paris ex Coll.: R. Oberthur”(MNHN).
Type Locality. India (Darjeeling).
Materials Examined (36 Specimens). BHUTAN: 씸,
1898, L. Durel, MTD; 3 么么,2씸씸, same data, ZMHB;
1么, no date indicated, Sakiou, MTD; 1 么,1씸, no date
indicated, Maria Basti, MTD; INDIA: Assam: 1 么,no
year or collector indicated, MTD; 1 么, 1907, C.
Felsche, MTD; 1 么,2씸씸, July. Moser, ZMHB; 1 씸,
Khasia Hills, no date or collector indicated, MTD; 1 씸,
no date or collector indicated, MTD; 1 么, no date
indicated, T. Kricheldoff, ZMHB; Sikkim; 1 씸,
Kurseong, 1894, R. P. Bretaudeau, MTD; 1 씸, no date
indicated, July. Moser, ZMHB; 1 么, no date or collec-
tor indicated, ZMHB; 1 么, Chasseurs Indigenes, 1894,
R. P. Bretaudeau, MTD; 1 씸, Saharanpur, no date or
collector indicated, MTD; Darjeeling,: 1 么, no date
indicated, July. Moser, ZMHB; 1 么,1씸, no date or
collector indicated, ZMHB; 1 么, Kalimpong Mon-
shong Kaman, 20Ð21-IV-1987, B. Bhakta, NHMB; 3
么么,3씸씸, Pedong 1,300 m, 20-IV-1986, B. Bhakta,
NHMB; 1 么, Kalimpong 1,000 m Upper Bombusti,
5-V-85, D. Ch. J. Rai, NHMB; 1 씸, Panyung, 16Ð17-
V-1987, B. Bhakta, NHMB; 1 么, Bengale, VII-1957, no
collector indicated, MHNG; CHINA: Tibet (⫽Thi-
bet): 1 么, no date indicated, G. Hauser, ZMHB; NE-
PAL: 2 么么, Arun V., Khandbari 1,100- Arunthan 1,300
m, 29-V-1983, M. Brancucci, NHMB).
Distribution. India (Assam, Sikkim, Darjeeling),
Nepal, Bhutan, China (Tibet). Records from Fujian
(in China) (Mader 1955) and Taiwan (Miwa 1931) are
either misidentiÞcations (see below) or mistakenly
labeled (Lee et al. 2006).
Helota scintillans Olliff
(Figs. 9Ð12, 60Ð64)
Helota scintillans Olliff 1884: 99; Ritsema 1889: 111;
Ritsema 1891b: 223, 231; Ritsema 1909: 181, 182;
Ritsema 1911a: 106; Ritsema 1915a: 126; Ritsema
1915b: 229; Wegrzynowicz 2000: 403; Kirejtshuk
2000: 27.
Helota pasteuri Ritsema 1893a: 111; Ritsema 1893c: 160;
Ritsema 1909: 181, 182; Ritsema 1911a: 106; Ritsema
1915a: 127; Ritsema 1915b: 229. syn. nov.
Helota pasteuri Wegrzynowicz 2000: 402; Kirejtshuk
2000: 27.
Diagnosis. The elytral apices of H. scintillans are
similar to H. servillei and H. feai, but H. scintillans
differs by its larger spots on the elytra and being
greenish bronze or dark purple. H. scintillans resem-
bles H. vigorsii with the greenish bronze color and the
large spots on elytra, but it differs by having more
impressed punctures on elytra and acute elytral apices
in females.
Male. Length 13.3Ð15.7 mm; width 5.1Ð 6.3 mm. Dor-
sal surface (Fig. 9) metallic green, antennae yellowish
brown, and antero-lateral angles of pronotum yellow-
ish brown; each elytron with two large yellow spots,
anterior spot between second and sixth striae, poste-
rior spot between third and sixth striae; scutellum
black. Ventral surface yellowish brown, with thoracic
margins surrounding pro- and meso-coxae black; head
and elytral epipleuron metallic green. Legs yellowish
brown, with trochanters, bases and apices of femora
and tibiae black, tarsomeres 1Ð4 and apices of coxae
and tarsomere 5 black. Dorsal surface of head ran-
domly and densely punctate, ventral surface with
736 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
more prominent punctures, medially reduced. Prono-
tum 0.74Ð0.79 times longer than wide, trapezoidal,
lateral margins with weak crenulations, narrowed at
apex, surface rugose, with raised patches and regular,
prominent punctures. Elytra 1.94Ð2.00 times longer
than wide, parallel, apex obliquely truncate, each
elytron with 10 striae and three longitudinal carinae;
Þrst carina between second and third striae, only vis-
ible from posterior spots to apices of elytra, apices of
Þrst carina projecting from margins of elytra, second
carina between sixth and seventh striae, third between
eighth and ninth striae. Thoracic ventrites without
punctures except for internal sides of proepisternum
and outer sides of metaventrite, a pair of small setal
clusters along median longitudinal sulcus of metaven-
trite. Punctures on abdominal ventrites extremely
small. Protibia (Fig. 61) slightly curved, internal mar-
gin ventrally and apically depressed, with rows of
short setae on dorsal apex. Fifth abdominal ventrite
similar to H. servillei. Eighth abdominal tergite (Fig.
60) widest at basal one third, apically narrowed, apical
margin weakly emarginate at middle. Penis (Fig. 62)
slender, abruptly narrowed near apex, apex truncate
and recurved, mesal margin rounded; dorsal lobes
widest at apical one third, gradually narrowed toward
base, notch between connection of dorsal lobes prom-
inent; basal margin with deep notch, extending to half
length of dorsal lobes. Parameres (Fig. 63) elongate
and apically narrowed, middle of apical margin with a
small, narrow notch, with dense long setae along mar-
gin behind apex, shorter posteriorly, ventral surface
with dense, short setae, basal margin truncate. Internal
sac (Fig. 64) with apical, elongate sclerite provided
with longitudinal cluster of teeth at apical one third,
covered posteriorly by Y-shaped sclerite, which pos-
sess ventral teeth on lateral expansions; a pair of me-
dian longitudinal sclerites covered by teeth with in-
ternal margin possessing teeth near bases and apices,
one basal sclerite with lateral expansions.
Female (Fig. 10). Length 15.6 mm; width 5.8 mm.
Similar to males, but protibiae without depression,
elytra apically tapering, apices pointed; metaventrite
without setae.
Coloration Variation. Individuals have dark purple
coloration (Figs. 11 and 12), formerly described as H.
pasteuri Ritsema.
Type Material. The holotype 么of H. scintillans Olliff
is labeled “Type/Cat No. 4a/Helota scintillans Olliff/
Type/V. Medenbach de Rooy ?Java?”(RMNH). It is in
good condition and is deposited in the RMMH.
The holotype 씸of Helota pasteuri Ritsema, is la-
beled “Type/Cat No. 5a/Helota Pasteuri씸type Rits./
J. D. Pasteur Toegoe Java occ.”It is in good condition
and is deposited in the RMNH.
Type Locality. Indonesia (Java).
Materials Examined (5 Specimens). INDONESIA:
Java: 1 么, Mons Gede 4000⬘, 1893, H. Fruhstorfer,
RMNH; 1 씸, Pengalengan 4000⬘, 1893, H. Fruhstorfer,
RMNH; 1 么,1씸, no date indicated, Toegoe Pasteur,
RMNH; 1 么, no date or collector indicated, Ritsema
vidit 1900, ZMHB.
Distribution.Indonesia (Java).
Helota feai Ritsema
(Figs. 13, 14, 65Ð69)
Helota feae (sic!) Ritsema 1891c: 886; Ritsema 1891b:
223, 229; Ritsema 1894a: 103; Ritsema 1911a: 105;
Ritsema 1915a: 127; Ritsema 1915b: 229.
Helota feae (sic!) Kirejtshuk 2000: 27.
Helota feai Wegrzynowicz 2000: 397.
Diagnosis. H. feai resembles H. thoracica in general
color pattern, but it can be easily distinguished by
having different shapes of elytral apices.
Male. Length 14.7 mm; width 5.8 mm. Dorsal surface
(Fig. 13) blackish metallic, antennae blackish brown,
with Þrst three and last antennomeres pale; wide yel-
lowish brown longitudinal bands along lateral margins
of pronotum; each elytron with two small yellow spots
between third and sixth striae. Ventral surface yel-
lowish brown, with thoracic margins surrounding pro-
and meso-coxae black; head and elytral epipleuron
blackish metallic. Legs yellowish brown, with trochan-
ters, bases and apices of femora and tibiae black, tar-
someres 1Ð4 and apices of coxae and tarsomere 5
black. Dorsal surface of head randomly and densely
punctate, ventral surface with more prominent punc-
tures, medially reduced. Pronotum 0.77 times longer
than wide, trapezoidal, lateral margins with weak
crenulations, narrowed at apex, surface rugose, with
raised patches and regular, prominent punctures.
Elytra 1.90 times longer than wide, parallel, apex ob-
liquely truncate, each elytron with 10 striae and three
longitudinal carinae; Þrst carina between second and
third striae, only visible from posterior spots to apices
of elytra, apices of Þrst carina projecting from margins
of elytra; second carina between sixth and seventh
striae; third between eighth and ninth striae. Thoracic
ventrites without punctures except for internal sides
of proepisternum and outer sides of metaventrite, a
pair of small setal clusters along median longitudinal
sulcus of metaventrite. Punctures on abdominal ven-
trites very small. Protibia (Fig. 66) slightly curved,
internal margin ventrally and apically depressed, with
rows of short setae on dorsal apex. Fifth abdominal
ventrite similar to H. servillei. Eighth abdominal ter-
gite (Fig. 65) oval, apically narrowed, apical margin
weakly emarginate at middle. Penis (Fig. 67) slender,
abruptly narrowed near apex, apex truncate, mesal
margin extending into dorsal lobes; dorsal lobes widest
at apical one third, gradually narrowed toward base,
notch between connection of dorsal lobes prominent;
basal margin with deep notch, extending to half length
of dorsal lobes. Parameres (Fig. 68) elongate and api-
cally narrowed, middle of apical margin with a narrow
and small notch, with dense long setae along margin
behind apex, much shorter posteriorly, ventral surface
with dense short setae, basal margin truncate. Internal
sac (Fig. 69) with apical elongate sclerite provided
with longitudinal cluster of teeth at apical third, cov-
ered posteriorly by Y-shaped sclerite that possess ven-
tral teeth on lateral expansions and dense teeth along
basal four Þfths of lateral margin; a pair of median,
elongate and sinuate sclerites at middle, covered by
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 737
teeth, their internal margin with teeth near bases and
apices; one basal sclerite with lateral expansions.
Female (Fig. 14). Length 14.4Ð15.3 mm; width 5.4Ð
5.8 mm. Similar to males, but protibiae without de-
pression, elytral apex abruptly narrowed, forming
acute process along Þrst carina, similar to H. servillei;
metaventrite without setae.
Type Material. The lectotype 씸of Helota feai Rit-
sema, here designated to preserve stability and to
make more universal use of this name, is labeled “Co-
Type/Cat No. 8a/Carin Cheba`(in Burman) 900-1100
m L. FeaV.XII.88/Helota Feae Rits. type 씸/L. Fea
Burma.”It is in good condition and is deposited in the
RMNH. A female paralectotype is labeled: “Carin
Cheba`(in Burman) 900-1100 m L. FeaV.XII.88/Ty-
pus/Feae Rits./SYNTYPUS 씸Helota feae Ritsema,
1891 (red)”(MSNG).
Type Locality. Myanmar.
Materials Examined (12 Specimens). LAOS: Haut
Mekong: 1 么,1씸, Pou Hai Katoui, 6-IV-1918, R. V. de
Salvaza, RMNH; 么, Hieng Khouang, 22-IV-1919, Sal-
vaza, MSNG; Xamneua: 1 씸, Ban Saleui, 1,350 m,
21Ð24-V-2004, C.-F. Lee, TARI; 1 씸, Hieng Khouang,
24-IV-1919, Salvaza, MSNG; THAILAND: 1 么, Doi
Pui, 27-IV-1983, H. Kan, EUMJ; 1 么, 26-IV-1983, H.
Kan, EUMJ; Chiang Mai: 4 么么,1씸, Doi Suthep,1Ð5-
VI-1986, H. Hirasawa, EUMJ.
Distribution. Myanmar, Laos, Thailand.
Helota vigorsii vigorsii MacLeay
(Figs. 15, 16, 31, 70Ð74)
Helota vigorsii MacLeay 1825: 43; Chapuis 1876: 18;
Ritsema 1889: 111; Ritsema 1891b: 223, 229; Ritema
1891c: 885; Ritsema 1894a: 97; Ritsema 1909: 181,
183.
Helota vigorsii Skelley 1998: 38; Wegrzynowicz 2000:
405.
Helota vigorsi (sic!) Reitter 1876: 6; Ritsema 1911a: 106;
Ritsema 1915a: 126; Ritsema 1915b: 229.
Helota vigorsi (sic!) Olliff 1883: 51; Kirejtshuk 2000: 27.
Helota vigorsii sumatrensis Ritsema 1909: 183. syn. nov.
Helota vigorsi (sic!) sumatrensis Ritsema 1911a: 106;
Ritsema 1915a: 126; Ritsema 1915b: 229.
Helota vigorsi (sic!) sumatrensis Kirejtshuk 2000: 27.
Helota vigorsii sumatrensis Wegrzynowicz 2000: 405.
Diagnosis. H. v. vigorsii is similar to H. scintillans
with the greenish bronze color and large spots on
elytra, but it differs by its less-impressed punctures on
the elytra and rounded elytral apices in females (Fig.
31).
Male. Length 14.3Ð17.8 mm; width 5.3Ð 6.5 mm. Dor-
sal surface (Fig. 15) metallic green, antennae dark
brown with last antennomere yellowish brown; each
elytron with two large yellow spots between Þrst and
sixth striae; scutellum black. Ventral surface yellowish
brown, with thoracic margins surrounding pro- and
meso-coxae black; head and elytral epipleuron metal-
lic green. Legs yellowish brown, with trochanters,
bases and apices of femora and tibiae black, apices of
coxae and tarsomeres 5 black. Dorsal surface of head
randomly and densely punctate, ventral surface with
more prominent punctures, medially reduced. Prono-
tum 0.75Ð0.79 times longer than wide, trapezoidal,
lateral margins with very weak crenulations, narrowed
at apex, surface weakly rugose, with raised patches and
regular, prominent punctures. Elytra 2.04Ð2.07 times
longer than wide, parallel, apex obliquely truncate,
each elytron with 10 striae and three longitudinal
carinae; Þrst carina between second and third striae,
only visible near apices of elytra, apices of Þrst carina
projecting from margins of elytra, second carina be-
tween sixth and seventh striae, third between eighth
and ninth striae. Thoracic ventrites without punctures
except for internal sides of proepisternum and outer
sides of metaventrite, a pair of small setal clusters
along median longitudinal sulcus of metaventrite.
Punctures on abdominal ventrites very small. Protibia
(Fig. 71) slightly curved, internal margin ventrally and
apically depressed, with rows of short setae on dorsal
apex. Fifth abdominal ventrite similar to H. servillei.
Eighth abdominal tergite (Fig. 70) elongate, apically
narrowed, apical margin weakly emarginate at middle.
Penis (Fig. 72) slender, abruptly narrowed near apex,
apex truncate, mesal margin pointed; dorsal lobes wid-
est at apical one third, gradually narrowed toward
base, apices acute, notch between connection of dor-
sal lobes prominent. Parameres (Fig. 73) elongate and
apically narrowed, middle of apical margin with a
narrow and small notch, several setae of varying length
along margin behind apex, ventral surface with dense
short setae, basal margin truncate. Internal sac (Fig.
74) with apical elongate sclerite provided with longi-
tudinal cluster of teeth at apical one third, covered
posteriorly by Y-shaped sclerite possessing ventral
teeth on lateral expansions and teeth along basal half
of lateral margin and base; a pair of median sinuate
sclerites covered by teeth with internal margin pro-
vided with teeth near bases and apices; one basal
sclerite with lateral expansions.
Female (Fig. 16). Length 12.6Ð14.4 mm; width 4.9Ð
5.4 mm. Similar to males, but protibiae without de-
pression, elytra apically tapering with apices narrowly
rounded (Fig. 31); metaventrite without setae.
Variation. The shapes of the posterior spots on
elytra are variable. Thus, the subspecies H. v.
sumatrensis is regarded as a junior synonym of nom-
inate subspecies. But the populations from Borneo
have uniformly smaller spots which can be distin-
guished from other populations, thus H. v. borneensis
retains its taxonomic status.
Type Material. The holotype 씸of Helota vigorsii
MacLeay is labeled “Java (HorsÞeld)/60Ð15 E.I.C./
109/Vigorsii Mac./Helota Vigorsii MacLeay Type/
Type.”It is directly pinned and is in good condition.
It is deposited in the BMNH.
Although types of H. v. sumatrensis Ritsema were
not indicated in the original paper (1909), specimens
identiÞed by Ritsema (1915) should be regarded as the
type series. The lectotype 么, here designated to clarify
its taxonomic concept, is labeled “Cat No. 2b/Helota
vigorsii ML./sumatrensis么Rits./Tandjong Morawa.
Serdang (N. O. Sumatra)”(RMNH). Paralectotypes: 2
738 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
么么,3씸씸, same as lectotype but with Cat No. 2c⬇h
(RMNH); 1씸:“Cat No. 2a/Helota vigorsii ML./
sumatrensis씸Rits./Ludeking Sumatra”(RMNH); 1么:
“Cat No. 2i/Helota vigorsii ML./sumatrensis씸
Rits.A. L. v. Hasselt Sipirok Sumatra”(RMNH); 2么,
1씸“Cat No. 2j⬇l/Helota vigorsii ML./sumatrensis
Rits./H. Rouyer Pajakombo Sumatra”(RMNH);
13么么,13씸씸:“Cat No. 2m⬇rr/Helota vigorsii
sumatrensis Rits./P. O. Stolz Solok Sumatra occ.”
(RMNH); 1么:“Cat. No. 2ss/Padang Pandjang West-
Sumatra H. Rolle Berlin W./Helota vigorsii
sumatrensis么Rits./H. Rolle Padang Pandjang
Sumatra”(RMNH); 1么:“Cat. No. 2tt/Sumatra W.
Morton/Helota vigorsii sumatrensis么Rits./H. Rolle
Sumatra”(RMNH); 1씸:“Cat No. 2uu/Tamiang/He-
lota vigorsii sumatrensis Rits. 씸/H. Rolle Taming
Sumatra N. O.”(RMNH); 2씸씸:“Cat No. vv & ww/
Helota vigorsii sumatrensis Rits. 씸/H. Rolle Sumatra”
(RMNH); 1씸:“Cat No. xx/Coll. Thomson/Helota vig-
orsii sumatrensis Rits. 씸M. Se´dillot Sumatra.”
Type Locality. Indonesia (Java).
Materials Examined (23 Specimens). WEST MA-
LAYSIA: 1 么, Taiping, 1978Ð1980, no collector indi-
cated, NMW; 1 么, Gunon Bringchan, 22-VI-1985, M.
Satoˆ, TARI; 1 么,1씸, Cameron Highlands, 1986, no
collector indicated, EUMJ; 4 씸씸, bought from native
dealer, VII-1978, K. Ushijima, EUMJ; 1 么, III-1987, M.
Sato, EUMJ; Johor: 1 么, Kota Tinggi, 15-IX-1987, S.
Nagai, EUMJ; THAILAND: Trang: 1 씸, Khao Chong
Botanic Garden, rain forest, 22-XI-2004, M. Fo¨ldva´ri,
A. Orosz & L. Papp, HNHM; INDONESIA: Sumatra:
1씸, Ajer Mantcior, Agosio 1878, O. Beccari, MSNG; 1
씸, Palembang, no date indicated, W. Knappert, NMW;
1么,1씸, Solok, lack date, P. O. Stolz, MTD; 1 씸,
Silogiri, 7-VI-1983, EUMJ; Java: 1 씸, MT Ardjoe¨no
1,200Ð1,500 m, V-1891, W. Doherty, MSNG; 2 씸씸,
1,500⬘, 1891, H. Fruhstorfer, NMW; 1 么, Pengalengan
4,000⬘, 1893, H. Frustorfer, MTD; 2 么么,1씸, Mt. Kawi,
Reg. Passuruan, 1907, V. M. Duchon, Coll. C. Felsche
Geschenk, MTD.
Distribution. West Malaysia, Thailand, Indonesia
(Sumatra, Java). Olliff (1883) also reported it from
India. This record, however, requires conÞrmation.
Helota vigorsii borneensis Ritsema
Helota vigorsii borneensis Ritsema 1909: 183.
Helota vigorsi (sic!) borneensis Ritsema 1911a: 106;
Ritsema 1915a: 126; Ritsema 1915b: 229.
Helota vigorsi (sic!) borneensis Kirejtshuk 2000: 27.
Helota vigorsii borneensis Wegrzynowicz 2000: 405.
Diagnosis. This subspecies differs from the nomino-
typical subspecies by having small posterior yellow
spots on the elytra. They are restricted to area be-
tween third to sixth striae.
Type Material. As with H. v. sumatrensis, types of H.
v. borneensis Ritsema were not indicated in the orig-
inal paper (Ritsema 1909). Specimens identiÞed by
Ritsema (1915) should be regarded as the type series.
The lectotype 么, here designated to clarify its taxo-
nomic concept, is labeled “Cat No. 3a/Borneo occ.
1898 Pontianak/Helota vigorsii borneensis Rits.
么/Reni Oberthu¨r Borneo occ.”(RMNH). Paralecto-
types: 1么,5씸씸, same as lectotype but with Cat No.
3b-h (RMNH); 2씸씸:“CatNo.i&j,Borneo occ. Riv
Sintang, F. Buffat 1898 Helota vigorsii borneensis Rits./
Reni Oberthu¨r Borneo occ.”(RMNH); 1씸:“Cat No.
3k, Nord Borneo Mont Kina Balu 5Ð8-1903 John Wa-
terstradt/Helota vigorsii borneensis Rits./Reni
Oberthu¨r Borneo occ.”(RMNH).
Type Locality. Borneo
Materials Examined (12 Specimens). EAST MA-
LAYSIA: Sabah: 1 씸, 16 miles NW of Keningau, alt.
1,400 m, 20-IV-1984, S. Nagai, TARI; 1 么, same data but
with 23-IV-1984, TARI; 1 씸, 10 miles NW of Keningau,
alt. 900 m, 27-V-1980, S. Nagai, EUMJ; 1 씸, same data
but with 28-V-1980, EUMJ; 1 씸, same data but with
12-VI-1980, EUMJ; 1 씸, same data but with 14-VI-80,
EUMJ; 1 씸, Crocker Rang, 10.5 miles of Keningau,
11-II-1982, S. Nagai, EUMJ; 1 么, Sandakan, Gum Gum,
29-XI-1979, S. Nagai, EUMJ; 1 씸, Sandakan, Suan
Lamba, alt. 100 m, 25-XI-1979, S. Nagai, EUMJ; 1 么,
Mesilau, 16-IV-2006, A. Abe & R. Abe, EUMJ; 1 씸,
Bukndu Tuhan, 18-IV-2006, same collectors, EUMJ);
BORNEO: 1 씸, Poritianak, 1900, no collector indi-
cated, MSNG.
Distribution. Borneo
Helota fairmairei Ritsema
(Figs. 17, 18, 32, 33, 75Ð80)
Helota fairmairei Ritsema, 1889: 101; Ritsema 1891b:
224, 229; Ritsema 1893b: 140; Ritsema 1894b: 111;
Ritsema 1911a: 105; Ritsema 1915a: 128; 1915b: 230.
Helota fairmairei Jakobson 1915: 500; Wegrzynowicz
2000: 397; Kirejtshuk 2000: 27.
Diagnosis. H. fairmairei, H. thoracica, and H. longipes
are characterized by possessing truncate elytral apices
in males (Fig. 32). H. fairmairei bears a greater simi-
larity to H. longipes with its reddish bronze color, but
it differs by its denser and less impressed punctures on
the elytra (Fig. 80) and widely rounded elytral apices
in females (Fig. 18).
Male. Length 16.5Ð17.5 mm; width 6.0 Ð6.4 mm. Dor-
sal surface (Fig. 17): head black with granules reddish
bronze, antennae brown; pronotum reddish brown,
with raised patches black, granules inside patches red-
dish bronze; basal margins of pronotum and elytra
black, elytra reddish brown, each elytron with two
small yellow spots between third and seventh striae;
scutellum black. Ventral surface reddish brown, with
thoracic margins surrounding coxae black except that
of metaventrite, mesepisternum, and mesepimeron
black, metepisternum dark brown. Legs reddish
brown, with metacoxae and trochanters black, apices
of pro- and meso-coxae, bases and apices of femora,
and bases of tibiae black, claws black. Dorsal surface
of head randomly and densely punctate, ventral sur-
face with more prominent punctures, medially re-
duced. Pronotum 0.74Ð0.77 times longer than wide,
trapezoidal, lateral margins with weak crenulations,
narrowed at apex, surface rugose, with raised patches
and regular, prominent punctures. Elytra 1.93Ð2.00
July 2008 LEE:VIGORSII SPECIES GROUP OF Helota 739
times longer than wide, parallel, apex with a small
sutural tooth, truncate, each elytron with 10 striae and
three longitudinal carinae; Þrst carina between second
and third striae, only visible near apices of elytra,
reduced before margins of elytra, second carina be-
tween sixth and seventh striae, third between eighth
and ninth striae. Thoracic ventrites without punctures
except for internal sides of proepisternum and outer
sides of metaventrite, a pair of setal clusters along
median longitudinal sulcus of metaventrite, a few long
setae on abdominal ventrite 1. Punctures on abdom-
inal ventrites very small. Protibia (Fig. 76) moderately
curved, apex of internal margin expanded dorsally,
without rows of setae on dorsal apex. Fifth abdominal
sternite similar to H. servillei. Eighth abdominal tergite
(Fig. 75) transverse, apical margin truncate. Penis
(Fig. 77) wide, abruptly narrowed at apical one third,
apex truncate, mesal margin arcuate, sides narrowed at
apical one third, dorsal lobes widest near apices, grad-
ually narrowed toward base, apices acute, notch be-
tween connection of dorsal lobes prominent.
Parameres (Fig. 78) wide and apically narrowed, mid-
dle of apical margin with a narrow notch, with several
long setae along margin behind apex, ventral surface
with dense setae of varying length, basal margin ar-
cuate. Internal sac (Fig. 79) with apical elongate scle-
rite, apically narrowed, provided with longitudinal
cluster of teeth at apex, covered posteriorly by a bi-
furcate sclerite possessing ventral teeth on lateral ex-
pansions; a pair of median, curved sclerites covered
with dense teeth; one basal sclerite with lateral ex-
pansions.
Female (Fig. 18). Length 15.3Ð16.7 mm; width 6.0Ð
6.5 mm. Similar to males, but apices of protibiae with-
out expansion, elytra without carinae, elytral apex
(Fig. 33) widely rounded, setae on metaventrite and
abdominal ventrite one absent.
Type Material. The lectotype 么of Helota fairmairei
Ritsema, here designated to preserve stability and
make more universal use of this name, is labeled “Sik-
kim/64999/Helota么Fairmairei Rits./type/Type (or-
ange)/SYNTYPUS Helota fairmairei Ritsema, 1889 la-
beled by MNHUB 2005 (red)”. It is in good condition
and is deposited in the ZMHB. Paralectotypes: 1么:“Ex
Musaeo Van de Poll 1909/Coll. Janson/Helota Fair-
mairei type 么, Rits./TYPE (red)/Museum Paris ex
Coll. Oberthur/LECTOTYPUS Helota fairmairei Rit-
sema des. P. Wegrzynowicz”(MNHN); 1씸:“Sik-
kim/Ex Musaeo Van de Poll 1909/Helota Fairmairei
type 씸, Rits./TYPE (red)/Museum Paris ex Coll.
Oberthur/PARALECTOTYPUS Helota fairmairei Rit-
sema des. P. Wegrzynowicz”(MNHN). The designa-
tion of lectotype made by Wegrzynowicz is not valid
because it was not published. Ritsema (1889) indi-
cated that types were deposited at the Berlin Museum
(ZMHB), Paris Museum (MNHN), Leyden Museum
(RMNH), and Copenhagen Museum (ZMUC); but no
specimens bearing type labels are at the RMNH (Rit-
sema 1915a). Perhaps the author did not label them.
Twenty-three specimens at the RMNH, which were
labeled as 12a, b, x. Here, I select seven of them as
syntypes according to information provided in the
original paper, including 1 么and 2 씸씸 (12 g, h, i)
collected from Himalaya, which is one of the type
localities, and 1 么and 1 씸collected from Cherra
Ponjee and India bor., respectively and belong to
Se´dillotÕs collection as indicated by Ritsema (1889).
Thus, these Þve specimens are designated as paralec-
totypes. Some potential paralectotypes should be in
the ZMUC.
Type Locality. India (Sikkim).
Materials Examined (15 Specimens). NEPAL: 1 씸,
Arun Valley, Num-Tashigaon, 1,600Ð900-1,600 m,
10⬇12-VI-1992, J. & J. Probst, NMW; HIMALAYA: 2
么么,3씸씸, no other data, ZMHB; INIDA: 2 么么,4씸씸,
Sikkim, no date indicated, July. Moser, ZMHB; 1 么,1
씸, Darjeeling, lack date, July. Moser, ZMHB; MYAN-
MAR: Kachin: 1 씸, Mts. Imaw Bum, 3-VI-2002, A. Abe,
EUMJ.
Distribution. Nepal, India, Bhutan, and Myanmar.
Helota thoracica Ritsema
(Figs. 19, 20, 34)
Helota thoracica Ritsema 1895: 49; Ritsema 1905b: 217;
Ritsema 1911a: 106; Ritsema 1911b: 51; Ritsema
1915a: 127; Ritsema 1915b: 229; Miwa, 1931: 60;
Koˆno, 1939: 157; Wegrzynowicz, 2000: 404; Kirejt-
shuk, 2000: 27; Lee et al., 2006: 547 [redescription].
Helota feae (sic!) ab. mushana Ohta 1929: 109; Miwa,
1931: 59. (Synonymized by Kono 1939.)
Helota oberthu¨ri Mader 1955: 64 [misidentiÞcation].
Diagnosis. H. thoracica is similar to H. fairmairei and
H. longipes, which also have truncate elytral apices in
males, but the carinae between striae 2 and 3 are
developed so prominently that they appear as truncate
processes at the elytral apices in dorsal view. H. tho-
racica is easily distinguished from other two species by
its blackish bronze color.
Materials Examined (28 Specimens). CHINA: Si-
chuan (⫽Su-Tschuen): 1 么, Ya-Tcheou, 1897, no col-
lector indicated, ZMHB; 1 么, Emei Shan, Wannian
1,050 m, 19Ð30-III-1999, W. Schawaller, SMNS; Fujian
(⫽Fukien): 1 么,3씸씸, Kuatun, 15-VII-1946, Tschung-
Sen, Helota Oberthu¨ri Rits. Det. Mader, MHNG; 1 么,
Shaowu, TaChuLan, 1-VII-1946, T. C. Maa, BPBM;
Hainan: 1 么, Taipingshan, 16-V-1986, J. Okuma, TARI;
1么, same data but with 26-IV-1986, EUMJ; 1 么, same
data but with 5-V-1986, EUMJ; 1 씸, same data but with
25-V-1986, EUMJ; VIETNAM: 1 씸, Tonkin, Montes
Manson, no date or collector indicated, ZMHB; 1 么,1
씸, Tam Dao, lacking date, H. Perrot, MHNG; 1 씸, Tam
Dao, V-93, Nakamura, EUMJ; LAOS: Xamneua: 1 씸,
Ban Saleui, 1,350 m, 21Ð24-V-2004, C.-F. Lee, TARI;
THAILAND: 1 么, Doi Pui, 29-IV-1983, H. Kan, EUMJ);
1么,1씸, Chiang Mai, V-1989, H. Kan, EUMJ; Chiang
Mai: 6 么么,3씸씸, Doi Suthep, 1Ð5-VI-1986, H. Hira-
sawa, EUMJ.
Variation. The median sclerites of the internal sac
of individuals collected from Vietnam and Hainan are
longer and are covered with teeth.
Distribution. China (Tibet, Sichuan, Jiangxi, Fujian,
and Hainan), Vietnam, Laos, Thailand, Taiwan. No
740 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
obvious records of helotids existed for Hainan, China
previously. Those specimens collected from Hainan
have slightly greenish bronze elytra (Fig. 19). H.
oberthueri recorded from Fujian was misidentiÞed by
Mader (1955) and should be attributed to this species.
One additional specimen from Fujian was found at the
BPBM.
Helota longipes Ritsema
(Figs. 21, 22, 81Ð86)
Helota longipes Ritsema, 1889: 101; Ritsema 1891b: 224,
231; Ritsema 1893b: 140; Ritsema 1894a: 97; Ritsema
1911a: 105; Ritsema 1915a: 127; Ritsema 1915b: 230;
Jakobson 1915: 900; Wegrzynowicz 2000: 400; Kire-
jtshuk 2000: 27.
Diagnosis. H. longipes resembles H. fairmairei by
having similar color pattern, but it differs by its more
impressed and sparse punctures on the elytra (Fig. 81)
and the narrowly rounded elytral apices in females.
Male. Length 16.4 Ð16.8 mm; width 5.6Ð5.9 mm. Dor-
sal surface (Fig. 21): head black with granules reddish
bronze, antennae yellowish brown; pronotum reddish
brown, with raised patches black, granules inside
patches reddish bronze; basal margins of pronotum
and elytra black, elytra reddish brown, each elytron
with two small yellow spots between third and seventh
striae; scutellum black. Ventral surface reddish brown,
with thoracic margins surrounding coxae black,
mesepisternum and mesepimeron black, lateral mar-
gins of metaventrite black, bases and apex of metepis-
ternum dark brown. Legs reddish brown, with coxae
and trochanters black, bases and apices of femora and
tibiae black, claws and apices of tarsomere 5 black.
Dorsal surface of head randomly and densely punc-
tate, ventral surface with more prominent punctures
and medially reduced. Pronotum 0.77Ð0.79 times
longer than wide, trapezoidal, lateral margins with
weak crenulations, narrowed at apex, surface rugose,
with raised patches and regular, prominent punctures.
Elytra 2.00Ð2.02 times longer than wide, parallel, apex
with a small sutural tooth, truncate, similar to that of
H. fairmairei, each elytron with 10 striae and three
longitudinal carinae; Þrst carina between second and
third striae, only visible near apices of elytra, reduced
before margins of elytra, second carina between sixth
and seventh striae, third between eighth and ninth
striae. Thoracic ventrites without punctures except for
internal sides of proepisternum and outer sides of
metaventrite, a pair of setal clusters along median
longitudinal sulcus of metaventrite and a cluster of
setae on center of abdominal ventrite 1. Punctures on
abdominal ventrites small. Protibia (Fig. 83) slightly
curved, internal margin expanded dorsally at apical
one-fourths, apically and ventrally depressed, with
rows of short setae on dorsal apex. Fifth abdominal
ventrite similar to H. servillei. Eighth abdominal ter-
gite (Fig. 82) square, apical margin truncate. Penis
(Fig. 84) wide, abruptly narrowed near apex, apex
truncate, mesal margin arcuate, dorsal lobes widest at
apical one third, gradually narrowed toward base, api-
ces rounded, notch between connection of dorsal
lobes prominent. Parameres (Fig. 85) wide and api-
cally narrowed, middle of apical margin with a wide
notch, with rows of long setae along margin behind
apex, ventral surface with dense setae, basal margin
arcuate. Internal sac (Fig. 86) with apical elongate
sclerite provided with longitudinal cluster of teeth,
covered posteriorly by a bifurcate sclerite possessing
ventral teeth on basal half of margins; with a pair of
midian, long and twisted sclerites covered with dense
teeth; one basal sclerite with lateral expansions.
Female (Fig. 22). Length 13.9Ð16.1 mm; width 5.2Ð
5.8 mm. Similar to males, but apices of protibiae with-
out expansion, elytral apices narrowly rounded, like in
H. vigorsii; metaventrite and abdominal ventrite 1
without setae.
Type Material. The lectotype 么of Helota longipes
Ritsema, here designated to preserve stability and to
make more universal use of this name, is labeled
“Type/Cat No. 11a/Ind Bor/servillei/Moneuicurt/
Helota longipes Rits. type 么M. Sedillot India bor.”It
is in good condition and is deposited in the RMNH.
One male paralectotype in the MNHN is labeled “Sik-
kim/Ex Musaeo Van de Poll 1909/Helota longipes Rits.
type 么/TYPE (red)/Museum Paris ex Coll.
Oberthur/ECTOTYPUS Helota longipes Ritsema des.
P. Wegrzynowicz.”The label of the lectotype made by
Wegrzynowicz is not valid because the designation of
the lectotype was not published.
Materials Examined (14 Specimens). INDIA: 1 么,
Saharanpur, 2 么么,1么, MTD; 1 么, Assam, MTD; 1 么,
Assam, 1909, Gehr. W. Mu¨ller Verma¨ehr, MTD; 1 么,
Assam, 1907, C. Felsche Geschenk, MTD; Arunachal:
4么么,1씸, Drang vicinity 1,550 ⫾150 m, 27⬚21⬘-23⬘
N92⬚13⬘-16⬘, 1Ð9-VI-2004, E. L. Dembicky´, NHMB;
Darjeeling: 2 么么, Bengale, VII-1957, no collector in-
dicated, MHNG; MYANMAR: Kachin: 2 씸씸, Kang-
fang, 28-V-2002, A. Abe, EUMJ; 2 씸씸, same locality
and collector, 5-VI-2002, EUMJ.
Distribution. India, Bhutan, Myanmar.
Acknowledgments
I thank the curators mentioned previously for loans of type
specimens and unidentiÞed materials. Special thanks to C.
Carlton who read the Þrst draft. I am indebted to C.-C. Tsung
and D. Cheng for taking photographs of the specimens. This
study is supported Þnancially by the National Science Coun-
cil of Taiwan (NSC 96-2313-B-055-005-MY2).
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Received 25 October 2007; accepted 9 April 2008.
742 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 4
