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SYSTEMATICS
Revision of the Family Helotidae (Coleoptera: Cucujoidea): V. Species
Group Classification of the Genus Neohelota Ohta and Revisions of
N.laevigata and N.helleri Species Groups
C.-F. LEE
1
Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung, Wufeng 413 Taiwan
Ann. Entomol. Soc. Am. 103(4): 500Ð510 (2010); DOI: 10.1603/AN10003
ABSTRACT Species of the genus Neohelota are classiÞed into six species groups, and 60 species are
recognized. Taxonomy of the laevigata and helleri species groups is presented in detail. Two new
species are described: N.viklundi sp. nov. from Myanmar and N.satoi sp. nov. from Laos. A lectotype
is designated for N.tibialis Ritsema. Protibiae, genitalia, aedeagal internal sacs, eighth abdominal
tergites, and Þfth abdominal ventrites are illustrated for all species.
KEY WORDS Helotidae, Myanmar, Laos, taxonomic revision, new species
Helotidae is a little-known and originally monogeneric
family. Kirejtshuk (2000) divided the family into Þve
genera: Helota MacLeay, Neohelota Ohta, Afrohelotina
Kirejtshuk, Metahelotella Kirejtshuk, and Strophohe-
lota Kirejtshuk. Species of the genera Helota and Me-
tahelotella have been revised previously (Lee 2007,
2008, 2009a,b).
Members of Neohelota are similar to Helota in pos-
sessing two pairs of oval yellow spots on the elytra, but
they differ in having a ßat pronotum. Neohelota is the
most speciose genus within the family Helotidae, mak-
ing up 60 species on the Oriental and Eastern Palae-
arctic regions. RitsemaÕs key (Ritsema 1915b) to spe-
cies of Neohelota was mainly based on color patterns.
For the current study, the key to species groups of the
genus follows his methods and separates the genus
into six species groups.
Biological information of Neohelota was unknown
until Lee and Satoˆ(2006) studied Taiwanese species
of Helotidae. Adults of the genus were always found
on ßowers, suggesting that members of Neohelota are
anthophilous.
Materials and Methods
The descriptive terminology follows Lee and Satoˆ
(2006). The types of all known species and 69 ad-
ditional specimens were examined for this article.
Codens of museums or institutions and their cura-
tors are listed as follows: BPBM, Bernice P. Bishop
Museum, Honolulu, HI, USA (G. A. Sameulson);
EUMJ, Ehime University, Matsuyama, Japan (H. Yo-
shitomi); MHNG, Muse´um dÕHistoire Naturelle, Ge-
neva, Switzerland (G. Cuccodoro); MNHN, Mu-
se´um National dÕHistoire Naturelle, Paris, France
(A. Taghavian & T. Deuve); MTD Museum fu¨r
Tierkunde, Dresden, Germany (O. Ja¨ger); NHMB,
Naturhistorisches Museum, Basel, Switzerland (M.
Brancucci); NHRS, Naturhistoriska Riksmuseet,
Stockholm, Sweden (B. Viklund); NMW, Naturhis-
torisches Museum Wien, Austria (M. A. Ja¨ch);
RMNH, Nationaal Natuurhistorische Museum, Lei-
den, The Netherlands (F. Assen); SMNS, Staatliches
Museum fu¨r Naturkunde, Stuttgart, Germany (W.
Schawaller); and TARI, Taiwan Agricultural Re-
search Institute, Taichung, Taiwan.
Key to Species Groups of Neohelota
1. Pronotum totally bronze (Figs. 1 and 3) ....2
Pronotum bronze, except antero-lateral angles
yellowish brown (Figs. 5 and 7Ð10) ...... 3
2. Hypomeron bronze, same color as pronotum;
prosternum yellowish brown (Fig. 2) .....
.................. culta species group
Hypomeron and prosternum bronze, same color
as pronotum (Fig. 4) . . candezei species group
3. Legs totally yellowish brown (Figs. 11Ð16) . . .
............... laevigata species group
Distal and basal parts of legs bronze, same color
as pronotum (Figs. 1Ð10) .............4
4. Tibiae totally bronze, same color as pronotum
(Figs. 5 and 6) ...... guerinii species group
Distal/basal parts of tibiae yellowish brown
(Figs. 7Ð10) .....................5
5. Elytral apex shape uniform in both sexes (Figs. 7
and8) ..........attenuata species group
Elytral apex shape sexually dimorphic (Figs. 9
and10) ...........helleri species group
1
Corresponding author, e-mail: chifeng@tari.gov.tw.
0013-8746/10/0500Ð0510$04.00/0 䉷2010 Entomological Society of America
Neohelota culta Species Group
(Figs. 1Ð2)
Diagnosis. Pronotum totally bronze, same color as
elytra; proepisternum bronze, same color as prono-
tum, prosternum yellowish brown; leg yellowish
brown, but apex of femur and most of tibia bronze;
elytral apex uniform in shape in both sexes.
List of Neohelota culta species group (21 species):
Figs. 1–10. Habitus of species groups of Neohelota. (1) N.similis, male, dorsal (N.culta species group). (2) Same, ventral.
(3) N.montana, male, dorsal (N.candezei species group). (4) Same, ventral. (5) N.lini, male, dorsal (N.gueriniispecies group).
(6) Same, ventral. (7) N.sonani, male, dorsal (N.attenuata species group). (8) Same species, female, dorsal. (9) N.helleri,
male, dorsal (N.helleri species group). (10) Same species, female, dorsal. (Online Þgure in color.)
July 2010 LEE:SPECIES GROUP CLASSIFICATION OF Neohelota 501
Neohelota affinis (Ritsema, 1891c)
Neohelota boulei (Ritsema 1915c)
Neohelota boysii (Ritsema, 1889)
Neohelota bretaudeaui (Ritsema, 1894b)
Neohelota chinensis (Mader 1955)
Neohelota consanguinea (Ritsema 1915c)
Neohelota culta (Olliff, 1882)
Neohelota dubia (Ritsema, 1891e)
Neohelota fryi (Ritsema, 1894a)
Neohelota gestroi (Ritsema, 1891d)
Neohelota hopei (Ritsema 1915c)
Neohelota indicator (Ritsema 1905)
Neohelota jordani (Ritsema, 1899b)
Neohelota klapperichi (Mader 1955)
Neohelota notata (Ritsema, 1891e)
Neohelota olliffii (Ritsema, 1899b)
Neohelota pusilla (Oberthu¨r, 1883)
Neohelota severini (Ritsema, 1893a)
Neohelota sharpi (Ritsema 1915c)
Neoelota similis Lee & Satoˆ, 2006
Neoelota taiwana (Ohta 1929)
Neohelota candezei species group
(Figs. 3Ð4)
Diagnosis. Pronotum totally bronze, same color as
elytra; proepisternum and pronotum bronze, same
color as pronotum; leg yellowish brown, but apex of
femur, and tibia bronze; elytral apex uniform in shape
in both sexes.
List of Neohelota candezei species group (seven spe-
cies):
Neohelota candezei (Ritsema, 1899a)
Neohelota claudiae (Nguyen-Phung, 1985)
Neohelota jocelinae (Nguyen-Phung, 1985)
Neohelota krugeri (Ritsema 1900)
Neohelota margarethae (Ritsema 1900)
Neohelota mariae (Ritsema, 1899b)
Neohelota montana (Ohta 1929)
Neohelota guerinii species group
(Figs. 5Ð6)
Diagnosis. Pronotum bronze, same color as elytra,
except antero-lateral angles yellowish brown; proepis-
ternum and pronotum yellowish brown, proepister-
num occasionally bronze; leg yellowish brown, except
apex of femur and tibia bronze; elytral apex shape
sexually dimorphic.
List of Neohelota guerinii species group (23 species):
Neohelota babai Lee & Satoˆ, 2006
Neohelota brevis (Ritsema, 1891b)
Neohelota cereopunctata (Lewis, 1881)
Neohelota chunlini Lee & Satoˆ, 2006
Neohelota curvipes (Oberthu¨r, 1883)
Neohelota desgodinsi (Ritsema, 1893a)
Neohelota dohertyi (Ritsema, 1891e)
Neohelota elongata (Ritsema 1905)
Neohelota epipleuralis (Ritsema 1914)
Neohelota grouvellei (Ritsema 1910)
Neohelota guerinii (Hope, 1840)
Neohelota guttata (Ritsema, 1893a)
Neohelota imtermedia (Ritsema 1905)
Figs. 11–16. Dorsal habitus of species of Neohelota laevigata species group. (11) N.laevigata, male. (12) Same, female.
(13) N.tibialis, male. (14) Same, female. (15) N.viklundi, male. (16) Same, female. (Online Þgure in color.)
502 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4
Neohelota lini Lee & Satoˆ, 2006
Neohelota magdalenae (Ritsema 1910)
Neohelota mountoni (Ritsema 1905)
Neohelota neglecta (Ritsema 1915b)
Neohelota ocellata (Ritsema, 1881)
Neohelota pustulata (Ritsema, 1893a)
Neohelota renati (Ritsema 1905)
Neohelota serratipennis (Ritsema, 1891d)
Neohelota sumbavensis (Ritsema 1909)
Neohelota yezoana (Koˆno, 1939) com.nov.
Neohelota attenuata Species Group
(Figs. 7Ð8)
Diagnosis. Pronotum bronze, same color as elytra,
antero-lateral angles yellowish brown; proepisternum
and prosternum yellowish brown; legs yellowish
brown except apex of femora and parts of tibia bronze;
elytral apex uniform in shape in both sexes.
List of Neohelota attenduata species group (six spe-
cies):
Neohelota attenuata (Ritsema, 1894b)
Neohelota bhutanensis (Chuˆjoˆ, 1975)
Neohelota durelii (Ritsema 1905)
Neohelota lewisi (Ritsema 1915c)
Neohelota rotundata (Ritsema, 1891d)
Neohelota sonani (Ohta 1929)
Neohelota laevigata Species Group
(Figs. 11Ð16)
Diagnosis. Pronotum, bronze, same color as elytra,
antero-lateral angles yellowish brown; proepisternum
and pronotum yellowish brown; leg yellowish brown;
elytral apex shape sexually dimorphic.
Key to Males of Species of the N.laevigata Species
Group
1. Protibia with a prominent process near apex
(Figs. 22 and 27); pronotum bronze, with yel-
lowish brown band along lateral margin . . . 2
Protibia without a prominent process near apex
(Fig. 17); pronotum brown, antero-lateral and
postero-lateral angles yellowish brown ....
............... N.laevigata (Oberthu¨r)
Figs. 17–20. Diagnostic characters of Neohelota laevigata. (17) Protibia. (18) Penis; dl, dorsal lobes; dn, dorsal notch. (19)
Parameres. (20) Internal sac. Scale bar ⫽0.5 mm.
July 2010 LEE:SPECIES GROUP CLASSIFICATION OF Neohelota 503
2. Process near apex of protibia angular (Fig. 22);
penis and parameres wide (Figs. 24 and 25);
apical margin of parameres without a process.
................. N.tibialis (Ritsema)
Process near apex of protibia rounded (Fig. 27);
penis and parameres narrow (Figs. 28 and 29);
each apical margin of paramere with a small
process ......N.viklundi Lee, new species
Neohelota laevigata (Oberthu¨r)
Helota laevigata Oberthu¨r 1883: 59; Olliff 1884: 100;
Fleutiaux 1886: 220; Ritsema 1889: 111; Ritsema
1891a: 230; Ritsema 1893ax: 140; Ritsema 1894a: 98;
Ritsema 1894b: 111; Ritsema 1911a: 105; Ritsema
1915a: 133; Ritsema 1915b: 234; Jakobson 1915: 900;
Wegrzynowicz 2000: 400.
Neohelota laevigata: Kirejtshuk 2000: 28.
Diagnosis. This species differs from Neohelota tibi-
alis in possessing yellowish brown antero- and pos-
terolateral angles pronotum, the transverse anterior
spots on the elytra, and less prominent process on the
protibia.
Male. Length 7.6 Ð7.7 mm, width 2.7Ð3.0 mm. Dorsal
surface (Fig. 11) blackish bronze; antennae yellowish
brown, three distal antennomeres darker; antero-lat-
eral and postero-lateral angles of pronotum yellowish
brown, sometimes color occupying entire lateral mar-
gins; each elytron with two yellow spots between third
and seventh striae, anterior spots transversely oval.
Ventral surface pale yellow; head metallic green. Legs
yellowish brown with tarsal claws black.
Dorsal surface of head randomly and densely
punctate; ventral surface with more prominent
punctures. Pronotum 0.67Ð0.68 times longer than
wide, lateral margins with weak crenulations, sides
apically narrowed, surface randomly and sparsely
punctured, abbreviated at middle of pronotal base.
Elytra 2.03Ð2.22 times longer than wide, widest near
base, gradually narrowed toward apices, strongly
Figs. 21–26. Diagnostic characters of Neohelota tibialis. (21) Eighth abdominal tergite. (22) Protibia. (23). Fifth abdominal
ventrite. (24) Penis. (25) Parameres. (26) Internal sac. Scale bar ⫽0.5 mm.
504 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4
behind posterior spots; elytron without an apical
tooth, apex widely rounded. Thoracic ventrites with
random and prominent punctures, reduced medi-
ally and basally on metaventrite. Punctures on ab-
dominal ventrites small. Protibiae (Fig. 17) slightly
curved, internal edges with a small rounded process
near apex, slightly dorsally depressed at middle.
Fifth abdominal ventrite and eighth abdominal ter-
gite similar to N. tibialis in shape. Penis (Fig. 18)
wide, apex truncate or narrowly rounded, sides par-
allel; dorsal lobes with acute apices, projecting from
penis, a little wider than penis at middle, notch
between connection of dorsal lobes shallow; basal
margin with extremely deep notch reaching apex of
dorsal lobes, the latter anterior and moderately wid-
ened. Parameres (Fig. 19) wide, widest at middle;
middle of apical margin with an angular notch, a
small process at apex of each side, and several long
setae along apical margin; ventral surface with
dense short setae, basal margin truncate, a number
of long setae along lateral margin. Internal sac (Fig.
20) with apical sclerites composed of one pair of
longitudinal sclerites, apically conjoined, middle of
outer margin with teeth, dorsally covered with a
slender sclerite, surface with dense teeth; medial
sclerites composed of one wide, curved sclerite at
middle, and one pair of elongate triangular sclerites
at sides, mesal angle abruptly curved distally; basal
sclerites composed of three longitudinal sclerites,
lateral sclerites strongly sclerotized, anteriorly ar-
ticulated with medial sclerite.
Female. (Fig. 12). Length 8.4 Ð8.7 mm; width 3.2Ð3.3
mm. Differing from males by the straight protibiae and
tapering and diverging elytral apices.
Variation. Specimens from China have a yellowish
brown band along the lateral margin of the pronotum
as N. tibialis.
Type Material. The male holotype is labeled “Darj./
REITTER/RITSEMA/Ritsema vidit 1892/Ritsema
vidit 1894/A.S. Olliff vidit 1884/Helota laevigata R.
Oberthu¨r TYPE Col. novit. I. p. 59/TYPE/Museum
Paris ex Coll. R. Oberthur”. The holotype is glued on
the paper card, is in good condition, and it deposited
the MNHN.
Type Locality. India (Darjeeling).
Figs. 27–30. Diagnostic characters of Neohelota viklundi. (27) Protibia. (28) Penis. (29) Parameres. (30) Internal sac. Scale
bar ⫽0.5 mm.
July 2010 LEE:SPECIES GROUP CLASSIFICATION OF Neohelota 505
Material Examined (46 specimens). INDIA: Dar-
jeeling, West Bengal (British Bootang), 1乆, Basti &
Durel MTD; 1乆, Basti, RMNH; 1么,1乆, Padong, 1914,
Durel, RMNH; 1乆, Mt. Fedillat, Thomson RMNH; 1乆,
Kurseong, Bretaudeau RMNH; 1么, Arunachal, W of
Bomdila, 2,600 m, 27⬚16⬘N, 92⬚24⬘E, 17-V-2004,
Dembicky´, NHMB; 1么, 25 km SW of Darjeeling,
Goom, 2,100Ð2,400 m, 24Ð28-IV-1966, Sedlacek,
BPBM; NEPAL: 6乆乆, Koshi, Baasantapur, 2,300 m,
30-V-2VI-1985, Brancucci, NHMB; 1乆, Baasantapur,
2,300 m, Kathandu V., 30-V-2VI-1985, Brancucci,
NHMB; 1么,1乆, Koshi, Mutidhunga-Chitre, 2,200Ð
2,400 m, 28-V-1985, Brancucci, NHMB; 1么, Koshi,
Gufa-Gorza, 2,800Ð2,100 m, 4-VI-1985, Brancucci,
NHMB; 3乆乆, Dhawalagiri Mustang, Kali-G. Khola
Kalopani, 2,500Ð2,800 m, 21Ð25-VI-1986, Holzschuh,
NHMB; 1乆, Kali Gandaki Khola, Kalopani-Ko-
pchepani, 1,500Ð2,400 m, 20-V-1984, Rai, NHMB; 1么,
Kali Gandaki Khola, in K. Gand., 2,520 m, 28⬚40⬘20⬙N
83⬚35⬘36⬙E, 27-V-2002, Ja¨ger, MTD; 1么,2乆乆, Chitre
2,400 m, Ghar Khola, 26Ð31-V-1984, Rai, NHMB; 1乆,
same locality, 29-V-1985, Bhakta, NHMB; 1乆, Bagmati
Sindhupalchok, Dubhachaur-Sarmatang, 1,600Ð2,500
m, 3-VI-1989, Brancucci, NHMB; 1么, Bagmati, below
Tarke Ghyang, 2,600 m, 25-IV-1981, Lo¨bl & Smetana,
MHNG; 1么, Umg. Shermathang, Helambu, 1980,
Franz, NHMB; 1乆, Nawronkot b.Larjung Thakholage-
biet, 2,900Ð3,000 m, Franz, NMW; 1么, Ghorapani,
2,880 m, Pokhara-Jhomoson, 16-X-1981, Sakai, EUMJ;
3乆乆, Singrila Ridge, Gairibas, 2,600 m, 1-X-1983, Sa-
kai, EUMJ; 1乆, Lalitpur, Phulcoki, 2,650 m, 14-X-1983,
Smetana & Lo¨bl, MHNG; 1乆, Khandbari, below
Sheduwa, 2,100Ð2,550 m, Smetana, MHNG; 1乆, Mt.
Panchase 18 km W. Pokhara east slope, 2,300 m, 21Ð
22-V-1997, Ja¨ger, MTD; 1乆, Mts. Annapurna, Madi
Khola Talunterh. Sikis 1,500 m, 10-V-1996, Schmidt &
Ja¨ger, MTD; 1么, Dolakha, S slope of Khare Khola,
2,100 m, 2-VI-2000, Schawaller, SMNS; 1乆, Surkhet, N
Surkhet, 1,600 Ð2,000 m, 28-V-1998, Miksch, SMNS; 1乆,
Solukhumbu, Junbsei to Ringmo, 2,700Ð3,000 m, 12-
V-1997, Schawaller, SMNS; CHINA: 1么,3乆乆, Yun-
nan, 100 km W Baoshan, Gaoligongshan Nat. Res.,
14Ð21-VI-1993, Jendek and Sause NMW.
Distribution. India, Nepal, China (Yunnan).
Neohelota tibialis (Ritsema, 1893)
Helota tibialis Ritsema 1893a: 136; Ritsema 1893b: 160;
Ritsema 1894a: 98; Ritsema 1911a: 106; Ritsema
1915a: 134; Ritsema 1915b: 234; Jakobson 1915: 900;
Chuˆjoˆ1975: 290; Wegrzynowicz 2000: 405.
Neohelota tibialis: Kirejtshuk 2000: 30.
Diagnosis. This species is characterized presence of
a prominent angular process near the apex of the
protibia (Fig. 22), and a yellowish brown band along
lateral margin of the pronotum.
Male. Length 7.5Ð 8.4 mm, width 2.9Ð3.2 mm. Dorsal
surface (Fig. 13) bronze but pronotum darker; anten-
nae yellowish brown, three distal antennomeres dark-
er; yellowish brown band along lateral margin of
pronotum; each elytron with two yellow spots be-
tween third and seventh striae, anterior spots circular.
Ventral surface pale yellow; head metallic green. Legs
yellowish brown with tarsal claws black.
Dorsal surface of head randomly and densely punc-
tate; ventral surface with more prominent punctures.
Pronotum 0.69Ð0.71 times longer than wide, lateral
margins with weak crenulations, apically narrowed,
surface randomly and sparsely punctured, abbreviated
at middle of base, punctures bigger at sides. Elytron
2.03Ð2.08 times longer than wide, widest near base,
gradually narrowed toward apices; elytron without
apical teeth, apex widely rounded. Thoracic ventrites
with random, prominent punctures, reduced medially
and basally on metaventrite. Punctures on abdominal
ventrites small. Protibia (Fig. 22) slightly curved, in-
ternal margin with a triangular process near apex,
dorsally depressed at apical one third. Metatibia
slightly curved at apical one third. Fifth abdominal
ventrite (Fig. 23) with subtruncate apical margin, sur-
face with dense short setae conÞned to semicircle
along apical margin; several extremely long setae
among short setae. Eighth abdominal tergite (Fig. 21)
slightly transverse, medially emarginate. Penis (Fig.
24) wide, apex rounded or trunate, sides subparallel;
dorsal lobes with acute apices, slightly wider than
penis, notch between connection of dorsal lobes shal-
low; basal margin with extremely deep notch, almost
reaching apex of penis, apically widened. Parameres
(Fig. 25) wide, widest at middle; middle of apical
margin with angulate notch; ventral surface with
dense short setae, basal margin truncate, with some
long setae along apical and lateral margin. Internal sac
(Fig. 26) with apical sclerites composed of one pair of
longitudinal sclerites with apical half of outer margin
bearing teeth and dorsally covered with an oblong
sclerites, surface with dense teeth; an elongate medial
sclerite with apex bearing a pair of small processes at
sides; a transverse sclerite projecting from middle;
basal sclerites composed of a pair of longitudinal scle-
rites connected anteriorly with a narrow and trans-
verse sclerite; a pair of curved sclerites at sides of basal
sclerites with internal margin bearing teeth, dorsal
surface with fewer teeth.
Female. (Fig. 14). Length 8.7 mm; width 3.3 mm.
Differing from males by the straight protibiae and
tapering and divergent elytral apices. In addition to
both characters, yellowish brown bands on the prono-
tum narrower and broken at middle.
Type Material. The male holotype is labeled “Type/
Cat No. 48a/Darjeeling/Christie/Helota tibialis Rits.
1么,/Er. Wasman Darjeeling/Type (red)”. It is directly
pinned, is in good condition, and in the RMNH.
Type Locality. India (Darjeeling).
Material Examined (14 specimens). NEPAL: 3么么,
1乆, Arun V., Chichila, 1950 m, 31-V.1983, Brancucci,
NHMB; 1乆, Arun V., Arunthan-Chichila, 1,300Ð1,950
m, 29-V-1983, Brancucci, NHMB; 1乆, Arun V., Mure,
2000 m, 2Ð8-VI-1983, Brancucci, NHMB; 1乆, Arun V.,
Chichila-Mure, 2,050 m, Probst, NMW; 1乆, Arun V.,
Mure-Num, 1,550Ð2,000 m, 4Ð7-VI-1983, Brancucci,
NHMB; 1么, Modi Khola, Pothana, 1900 m, 5Ð7-V-1984,
Rai, NHMB; 1乆, same locality, 7Ð9-VI-1984, Bhakta,
506 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4
NHMB; 2么么, Kathmadu Tal Shivapuri Lekh,
Mulkharka sw. Bagmati, 27⬚47⬘04⬙N, 85⬚24⬘27⬙E,
22-V-2005, Ja¨ger, MTD; INDIA: 1么, Arunachal, be-
tween Dirang & Bomdila Pass, 2,200 m, 27⬚19⬘N, 92⬚
22⬘E, 15-V-2004, Dembicky´, NHMB; VIETNAM: 1么,
Sapa City, Xin Chai, 3-V-2009, Kusui, EUMJ.
Distribution. India, Nepal, Vietnam.
Neohelota viklundi Lee, new species
Diagnosis. This species is similar to N. tibialis based
on the prominent process on the apex of the protibia
(Fig. 15) and the yellowish brown band along the
lateral margin of the pronotum, but the tibial process
in this new species is rounded in contrast to the an-
gular process in N. tibialis.
Male. Length 7.6 Ð7.9 mm, width 2.7Ð2.8 mm. Dorsal
surface (Fig. 15) greenish bronze; antenna yellowish
brown, three distal antennomeres darker; yellowish
brown band along lateral margin of pronotum some-
times extending along entire length; elytron with two
yellow spots between third and seventh stria, anterior
ones transverse. Ventral surface pale yellow; head
metallic green. Legs yellowish brown with tarsal claws
black.
Dorsal surface of head randomly and densely punc-
tate; ventral surface with more prominent punctures.
Pronotum 0.70Ð0.72 times longer than wide, lateral
margins with weak crenulations, sides apically nar-
rowed, surface randomly and sparsely punctate, ab-
breviated at middle of base. Elytron 2.23Ð2.29 times
longer than wide, widest near base, gradually nar-
rowed toward apex, strongly narrowed adjacent to
apex; elytron without an apical tooth, apex widely
rounded. Thoracic ventrites with random, prominent
punctures, reduced medially and basally on metaven-
trite. Punctures on abdominal ventrites small. Protibia
(Fig. 27) slightly curved, internal margin with mod-
erate rounded process near apex, dorsally depressed at
apical one third. Metatibia with internal margin ßat-
tened at apical one third, ventral margin slightly ser-
rate. Fifth abdominal ventrite and eighth abdominal
tergite similar to N. tibialis. Penis (Fig. 28) narrow,
apex widely rounded, sides widest at middle; dorsal
lobes with acute apices, slightly wider than penis at
middle, notch between connection of dorsal lobes
deep; basal margin with extremely deep notch, almost
reaching apex of penis, slightly widened apically.
Parameres (Fig. 29) elongate, widest at middle; middle
of apical margin with an acute notch; a small process
dorsal present on apical margin; ventral surface with
dense, short setae, basal margin truncate, a number of
long setae at lateral and outer margins. Internal sac
(Fig. 30) with apical sclerites composed of one pair of
longitudinal sclerites apically conjoined, apical two
third with dense teeth; basal two longitudinal sclerites
connected anteriorly; a pair of curved sclerites lateral
to basal sclerites, internal margin with teeth, dorsal
surface with fewer teeth.
Female. (Fig. 16): Length 7.3Ð 8.3 mm; width 2.6Ð3.0
mm. Differing from males by the straight protibiae and
tapering, curved, and divergent elytral apices.
Type Material. Holotype 么:“N. E. BURMA Kam-
baiti, 7,000 feet 12/4 1934 R. MALAISE”; paratypes: 2
么么, same data but with 19/4; 1 乆, same data but with
25⫹2/4; 2 乆乆:“N. E. BURMA Kambaiti, 2,000 m
28/5,1934 Malaise”(all in NHRS); 1么:“(N. MYAN-
MAR) Pass between Panwa and Hpimaw Kachin St.,
6-VII-2003, A. Abe leg./alt. ⬇2,500 m/Collection of S.
Hisamatsu”(EUMJ).
Etymology. This new species is named after Bert
Viklund for providing materials of this interesting new
species.
Distribution. Myanmar.
Neohelota helleri Species Group
(Figs. 9Ð10)
Diagnosis. Pronotum elongate trapezoidal, bronze,
same color as elytra, antero-lateral angles yellowish
brown; proepisternum and pronotum yellowish
brown; legs yellowish brown, but apex of femora and
parts of tibiae bronze; elytral apices sexually dimor-
phic.
Neohelota helleri (Ritsema)
Helota Helleri Ritsema 1911b: 51; Ritsema 1915a: 133;
Ritsema 1915b: 234.
Helota helleri: Ohta 1929a: 109; Miwa 1931: 59; Koˆno
1939: 160; Hornig 1992: 70; Wegrzynowicz 2000: 399.
Neohelota helleri: Lee and Satoˆ2006: 534. [redescrip-
tion]
Neohelota tumaaka Ohta 1929b: 68 (synonymized by
Koˆno 1939)
Neohelota tsumaaka (sic!): Koˆno 1939: 160.
Material Examined (13 specimens). CHINA: 1么,
Guangdong, Lien-Ping, Kanlinshan, 700Ð900 m, 18-
IV-1940, Gressitt & To, BPBM; 1乆, same locality and
collector, 24-IV-1940, BPBM; 1么,3乆乆, Fujian, Sha-
owu, Tachuland, 30-IV-1942, Maa, BPBM; 1乆, same
locality and collector, 30-IV-1940, BPBM; 1乆, same
locality and collector, 30-IV-1943, BPBM; 1乆, Jiangxi,
Ginggangshan, 2Ð14-V-1994, NMW; 3乆乆, Guizhou,
Leigongshan, Xijiang, 1,200Ð1,900 m, 29-V-2-VI-1997,
Bolm, NHMB; 1乆, Guizhon, Fanchingshan,
27.VI.2009, TARI.
Distribution. Taiwan, China (Guangdon, Fujian,
Jiangxi, Guizhou).
Notes. Dorsal lobes of penis; parameres and internal
sacs are variable among Chinese specimens, but not in
consistent or obvious ways. These variations are not
regarded as sufÞcient to justify separate species status.
Neohelota satoi Lee, New Species
Diagnosis. This new species is distinguished easily
from N. helleri by its obliquely truncate elytral apices
(Fig. 31), truncate apex of the eighth abdominal ter-
gite (Fig. 33), and widely rounded apex of the penis
(Fig. 35).
Male. Length 8.8 mm, width 3.4 mm. Dorsal surface
purplish bronze; antennae yellowish brown, three dis-
July 2010 LEE:SPECIES GROUP CLASSIFICATION OF Neohelota 507
tal antennomeres darker; yellowish brown antero-lat-
eral angles of pronotum yellowish extending entire
length of lateral margin; elytron with two yellow spots
between third and sixth stria. Ventral surface pale
yellow. Head, outer, posterior, and anterior margins of
proepisternum surrounding procoxae, anterior margin
and margins of prosternum surrounding procoxae,
margins of mesoventrite surround mexocoxae, and
elytral epipleuron metallic green. Legs yellowish
brown; apices and bases of tibiae, and apices of femora
blackish brown; tarsi dark brown.
Dorsal surface of head randomly and densely
punctate; ventral surface with more prominent
punctures. Pronotum 0.75 times longer than wide,
trapezoidal, lateral margins with moderate crenu-
lations, sides apically narrowed, surface randomly
and sparsely punctured, abbreviated at middle of
pronotal base. Elytron 2.05 times longer than wide,
widest at basal one fourth, subparallel, apex (Fig.
31) obliquely truncate, moderately serrate near ob-
liquely truncate apex; surface ßat. Thoracic ven-
trites with random and prominent punctures, re-
Figs. 31–37. Diagnostic characters of Neohelota satoi. (31) Elytral apex, male. (32) Protibia. (33) Elytral apex, female. (34).
Fifth abdominal ventrite. (35) Penis. (36) Parameres. (37) Internal sac.
508 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 4
duced medially and basally on metaventrite.
Punctures on abdominal ventrites small. Protibia
(Fig. 32) slightly curved, apex angular, internal mar-
gin dorsally depressed at apical one third. Fifth
abdominal ventrite (Fig. 34) with truncate apical
margin, surface with short setae conÞned to apical
margin; several extremely long setae among short
setae. Eighth abdominal tergite transverse, apical
margin truncate. Penis (Fig. 35) wide, apex widely
rounded, sides parallel; dorsal lobes with acute api-
ces, slightly wider than penis at base, notch between
connection of dorsal lobes deep; basal margin with
deep notch, reaching middle of penis, apically ta-
pering. Parameres (Fig. 36) elongate, widest at mid-
dle; middle of apical margin with a deep notch;
ventral surface with dense short setae, basal margin
truncate, a number of long setae at apical margins.
Internal sac (Fig. 37): apical sclerites composed of
one oblong dorsal sclerite covered with teeth dor-
sally, and one transverse ventral sclerite narrowed
apically; medial sclerites paired, with inner rounded
process covered with teeth; basal sclerites compris-
ing one set of sclerites with lateral processes at apex
directed anteriorly, hollow basally.
Female. Length 11.9Ð12.6 mm; width 3.9Ð4.0 mm.
Differing from males by the straight protibiae and
rounded and divergent elytral apices (Fig. 33).
Type Material. Holotype 么:“[LAOS] Mt. Phu-
Pan, Ban Saleui, Xam Neua, 28Ð30-III-2005, J. Ya-
masako leg.”(EUMJ); paratypes: 2乆乆:“LAOS Nong
het Xienghouang Prov. 5-IV-2007 N. Ohbayashi leg.”
(EUMJ).
Etymology. This new species is named for late Dr.
Masataka Satoˆfor his contribution to taxonomy of
Coleoptera.
Distribution. Laos.
Acknowledgments
I thank the curators mentioned in the text for loans of type
specimens and unidentiÞed materials. Special thanks to C.
Carlton who read the Þrst draft. I am indebted to H.-T. Cheng
for taking photographs of the specimens. This study was
supported by the National Science Council of Taiwan (NSC
96-2313-B-055-005-MY2).
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Received 5 January 2010; accepted 1 April 2010.
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