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Revision of the genus Aulacophora from Taiwan (Coleoptera: Chrysomelidae: Galerucinae)

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Ten species of the genus Aulacophora Chevrolat, 1836 are reported for Taiwan. Specimens of A. opacipennis Chûjô, 1962 collected from Southeast Asia were misidentified by Kimoto (1989) and should be identified as A. apicipes Jacoby, 1896. Color photos of habitus and drawings of male and female genitalia from eleven species (including A. apicipes) are presented. The following synonymies are proposed: Aulacophora analis (Weber, 1801) = Galeruca quadraria Olivier, 1808 (syn nov.); A. indica (Gmelin, 1790) = Rhaphidopalpa pubescens Allard, 1888 (syn. nov.); A. apicipes Jacoby, 1896 = A. nigripalpis Chen and Kung, 1959 (syn nov.). Lectotypes are designated for Crioceris abdominalis Fabricius, 1781, C. testacea Fabricius, 1787, Galleruca bicolor Weber, 1801, Galeruca quadraria Olivier, 1808, Aulacophora semiopaca Jacoby, 1886, A. frontalis Baly, 1888, A. lewisii Baly, 1886, A. intermedia Jacoby, 1892, A. semifusca Jacoby, 1892, A. dohrni Jacoby, 1899, A. almora Maulik, 1936, A. apicipes Jacoby, 1896, A. tenuicincta Jacoby, 1897, Rhaphidopalpa pubescens Allard, 1888, R. bengalensis Weise, 1892, R. ceramensis Weise, 1892, R. chinensis Weise, 1892, and Orthaulaca (Ceratia) cattigarensis Weise, 1892. A key to the Taiwanese species is provided.
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Accepted by M. Schoeller: 31 Mar. 2015; published: 24 Apr. 2015
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Revision of the genus Aulacophora from Taiwan
(Coleoptera: Chrysomelidae: Galerucinae)
CHI-FENG LEE
1
& RON BEENEN
2
1
Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung 413, TAIWAN. E-mail: chifeng@tari.gov.tw
2
Martinus Nijhoffhove 51, NL-3437 ZP Nieuwegein, THE NETHERLANDS. E-mail: r.beenen.wxs.nl
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .152
Key to species of the genus Aulacophora from Taiwan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Aulacophora analis (Weber, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .153
Aulacophora bicolor (Weber, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .156
Aulacophora frontalis Baly, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .160
Aulacophora indica (Gmelin, 1790) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
Aulacophora kotoensis Chûjô . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
[Aulacophora abdominalis (Fabricius, 1781)] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Aulacophora lewisii Baly, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
Aulacophora nigripennis Motschulsky, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .174
Aulacophora opacipennis Chûjô, 1962. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Aulacophora palliata (Schaller, 1783) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Aulacophora tibialis Chapuis, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Species excluded from Taiwan fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Aulacophora apicipes Jacoby, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .188
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .188
Abstract
Ten species of the genus Aulacophora Chevrolat, 1836 are reported for Taiwan. Specimens of A. opacipennis Chûjô, 1962
collected from Southeast Asia were misidentified by Kimoto (1989) and should be identified as A. apicipes Jacoby, 1896.
Color photos of habitus and drawings of male and female genitalia from eleven species (including A. apicipes) are pre-
sented. The following synonymies are proposed: Aulacophora analis (Weber, 1801) = Galeruca quadraria Olivier, 1808
(syn nov.); A. indica (Gmelin, 1790) = Rhaphidopalpa pubescens Allard, 1888 (syn. nov.); A. apicipes Jacoby, 1896 = A.
nigripalpis Chen and Kung, 1959 (syn nov.). Lectotypes are designated for Crioceris abdominalis Fabricius, 1781, C. tes-
tacea Fabricius, 1787, Galleruca bicolor Weber, 1801, Galeruca quadraria Olivier, 1808, Aulacophora semiopaca Jaco-
by, 1886, A. frontalis Baly, 1888, A. lewisii Baly, 1886, A. intermedia Jacoby, 1892, A. semifusca Jacoby, 1892, A. dohrni
Jacoby, 1899, A. almora Maulik, 1936, A. apicipes Jacoby, 1896, A. tenuicincta Jacoby, 1897, Rhaphidopalpa pubescens
Allard, 1888, R. bengalensis Weise, 1892, R. ceramensis Weise, 1892, R. chinensis Weise, 1892, and Orthaulaca (Ceratia)
cattigarensis Weise, 1892. A key to the Taiwanese species is provided.
Key words: taxonomy, new synonymy, lectotype designation, leaf beetles, Cucurbitaceae, Fabaceae
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Introduction
Species of Aulacophora had been intensively studied since some of them can cause serious damage of cucurbits.
Revisional works were mainly focused on regions, such as China (Chen and Kung 1959), Taiwan (Chûjô 1962),
China and Korea (Gressitt & Kimoto 1963), Indochina (Kimoto 1989), Malaysia (Mohamedsaid 1994), Sundaland
(Barroga & Mohamedsaid 2002), New Caledonia (Beenen 2008), and the Oriental species with yellow elytra
(Anand & Cox 1986) and metallic color (Medvedev 2001). However, most of them still caused taxonomic
confusion since type specimens have not been examined and both sexes of the same species were not associated
and studied. Thus, robust sample sizes are required for revision of this genus.
Another taxonomic problem occurs in the confusing status of Crioceris abdominalis Fabricius, 1781 and C.
testacea Fabricius 1787. Although types of these species exist and are deposited in the KIEL-collection at
Copenhagen, they are mixed together and with very few labels. Both species were treated as the same (Wilcox
1972) or redescribed based on single female syntype and one male respectively (Anand & Cox 1986) which were
not well represented. Five types deposited in the KIEL-collection have been studied and two are designated as
lectotype of C. abdominalis and C. testacea by considering nomenclatural stability and well represented by the
species at the present study.
The Taiwan Chrysomelid Research Team (TCRT) was founded in 2005 and is composed of 10 members. Most
of them are amateurs aiming to make an inventory of all species of Chrysomelidae in Taiwan. Specimens of the
genus Aulacophora have been extensively collected and studied, and host plants recorded. Diagnostic characters
were assessed and the status of all species was evaluated based on a large series of specimens.
Material and methods
More than 300 specimens have been examined. About half were collected by TCRT and deposited at the Taiwan
Agricultural Research Institute (TARI). The others are deposited in the historical collection at the same institute.
To prepare drawings of the adult reproductive systems, the abdomens of adults were separated and boiled in a
10% KOH solution, cleared in distilled water, and then mounted on microscope slides in glycerin for observation.
Specimens were examined and drawings were made using a Leica M165 stereomicroscope. Microscope slides
were examined and illustrated using a Nikon ECLIPSE 50i microscope. Body parts were then stored in glycerin
tubes with the dry mounted specimens.
Host plants are recorded by observing adult feeding behavior in the field. Plants were identified by Chih-Kai
Yang.
Specimens examined are deposited at the following institutes and museums.
BMNH The Natural History Museum, London, UK [Michael Geiser];
BPBM Bernice P. Bishop Museum, Hawaii, USA [Shepherd Myers];
IZAS Institute of Zoology, Academia Sinica, Beijing, China [Ruie Nie]
IRSB Institute Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium [Pol Limbourg] ;
JBCB Jan Bezděk collection, Brno, Czech Republic;
KIEL Universitetets Zoologiske Museum, Copenhagen, Denmark [Sree Gayathree Selvantharan] ;
MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy [Roberto Poggi];
MCZC Museum of Comparative Zoology, Harvard University, Massachusetts, USA [Philip D. Perkins] ;
MNHN Museum National d’Histoire naturelle, Paris, France [Antoine Mantilleri];
MNHUB Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der
Humboldt-Universität zu Berlin, Berlin, Germany [Joachim Willers];
PAHC Paul Aston collection, Hong Kong, China;
RBCN Ron Beenen collection, Nieuwegein, The Netherlands;
RMNH National Museum of Natural History, Leiden, the Netherlands [Hans Huijbregts];
TARI Taiwan Agricultural Research Institute, Taichung, Taiwan;
ZMUM Zoological Museum of Moscow Lomonosov State University, Moscow, Russian [Alexey Moseyko].
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REVISION OF AULACOPHORA FROM TAIWAN
Exact label data are cited for all type specimens of the described species; a double slash (//) divides the data on
different labels and a single slash (/) divides the data in different rows. Other comments and remarks are in square
brackets: [p]—preceding data are printed, [h]—preceding data are handwritten, [w]—white label, [y]—yellow
label, [b]—blue label, and [r]—red label.
Key to species of the genus Aulacophora from Taiwan
1. Elytron yellowish brown, some with black spots, or black elytron with yellow apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Whole elytron black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2. Whole elytron yellowish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Elytron yellowish brown with black spots or black elytron with yellow apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3. First antennomere enlarged and elytra with erect hairs behind elytra humerus in male; cone-like pygidium projecting elytral
apex in female; scutellum yellowish bown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Without such sexually secondary characters; scutellum blackish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tibialis
4. With a pair of tubercles on pronotum in male; pygidium slender and black in female . . . . . . . . . . . . . . . . . . . . . . . A. kotoensis
- Without tubercles on pronotum in male; pygidium wide and yellowish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. A. indica
5. Elytron yellow with two big black square patches, one at humerus and the other at apical 1/3 . . . . . . . . . . . . . . . . . . .A. analis
- Not such patterns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Ely tron yellow with one black spot on humerus; or two transverse rows of black spots, anterior row composed of two spots,
one at humerus, the other near suture, posterior row composed of transverse band (Figs 18, 20); some with posterior rows con-
nected with each other (Fig. 21); some with basal 1/4 black and one pair of small yellow spots at base near humerus, and pos-
terior rows connected with each other (Fig. 22) or separated bands (Fig. 24); some with anterior row composed of two spots
and apical half black (Fig. 23); some with basal 2/3 (Fig. 25) or most of elytra black (Fig. 26) but with one pair of small yellow
spots near humerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bicolor
- Basal 1/3 (Fig. 125), 2/3 (Fig. 126), or most of elytra (Fig. 123) black without yellow spots at base, meso- and metathoracic
and abdominal ventrites black; or black elytron with yellow apical margin extending into middle (Fig. 124), meso- and
metathoracic and abdominal ventrites yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tibialis
7. Meso- and metathoracic ventrites, and legs black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. nigripennis
- Whole ventral surface yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Antennomeres III to V filiform in both sexes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Antennomeres III to V triangular in male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9. Elytron opaque (Fig. 80); antenna of male more slender than female; apical margin of abdominal ventrite V truncate in female
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. opacipennis
- Elytron shining (Figs 73, 75); antenna of male wider than female; apical margin of abdominal ventrite V sinuate in female . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lewisii
10. Antennomere III longer than wide, and vertex with longitudinal groove (Fig. 111) in male; antennomeres VII to X relatively
wider than that of A. palliata (2.8–3.2 times longer than wide) in female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. frontalis
- Antennomere III long as wide, and ventex with transverse ridges (Fig. 112) in male; antennomeres VIII to X relatively slender
than that of A. frontalis (3.5–3.7 times longer than wide) in female. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. palliata
Aulacophora analis (Weber, 1801)
(Figs 1–3, 10–17)
Galleruca analis Weber, 1801: 55. (Sumatra).
Triaplatys analis: Fairmaire, 1883: 56.
Aulacophora analis: Baly, 1888: 176 (Indonesia: Sumatra, Sulawesi; Philippines, including Sulu islands); Allard, 1888: 322
(Adelaide); Chûjô, 1962: 85 (Taiwan: Lanyu island).
Ceratia analis: Miwa, 1931: 189 (Taiwan: Lanyu island).
Galeruca quadraria Olivier, 1808: 626 (Indes orientales). New synonym
Aulacophora quadraria: Allard, 1888: 322 (Adelaide); Barroga, 2001: 44 (Indonesia: Bali); Barroga & Mohamedsaid, 2002:
142 (Indonesia: Sumatra, Java; Malaysia: Sabah).
Type material. Galleruca analis: lectotype ♀ (KIEL), here designated, labeled: “Weber / analis. / 1743 [w, h] //
analis [w, h]”. This is the only specimen available in Fabricius’ collection. It is uncertain if there are other type
specimens.
Galeruca quadraria: lectotype ♀ (MNHN), here designated, labeled: “Galeruca / quadraria / Oliv. [h, w] //
SY6 / 36 [h, w, circle label] // TYPE [p, r] // M
USÉUM
P
ARIS
/ C
OLL
. G
ÉNÉRALE
[p, y] // G. quadraria / Oliv. / Java
/ Coll. Labillardiere [h, y] // SYNTYPE [p, r] // SYNTYPE / Aulacophora / quadraria (Olivier, 1808) [p, w] //
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FIGURES 19. Habitus of Aulacophora species. 1. A. analis, Lanyu Island, male, dorsal view; 2. Ditto, ventral view; 3. A.
analis, Lanyu Island, female, dorsal view 4. A. kotoensis, Lanyu Island, male, dorsal view; 5. Ditto, ventral view; 6. A.
kotoensis, Lanyu Island, female, dorsal view; 7. Ditto, ventral view; 8. Lectotype of Crioceris abdominalis, dorsal view; 9.
Ditto, ventral view.
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FIGURES 10–17. Diagnostic characters of Aulacophora analis (Weber). 10. Antenna, male; 11. Antenna, female; 12. Penis,
dorsal view; 13. Penis lateral view; 14. Gonocoxae; 15. Abdominal ventrite VIII, female; 16. Abdominal tergite VIII, male; 17.
Spermatheca.
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MNHN / EC4692 [p, w]”. This is the only specimen found at the MNHN, It is uncertain if there are other type
specimens.
Other specimens examined. INDONESIA. Java: 1♀, 1922, leg. Andrewes (BMNH); Sulawesi: 1♀,
Tondano, leg. Forsten (BMNH); Sumatra: 1♀, Dolok Mekangir, 5.VII.1974, leg. Diehl (MNHUB); 4♀, Sandaran
Agong, Korinchi Lake, 1914 (BMNH); 1♂, leg. Baly (BMNH); PHILIPPINES. 1925, leg. E. M. Ledyard
(BMNH); Luzon: 1♂, 1905, leg. Sharp (BMNH); 1♂, 1905, leg. Fry (BMNH); TAIWAN. Taitung: 2♀, Lanyu
Island, same locality, 26.VI.2009, leg. U. Ong (MNHUB); 4♂, 3♀, same locality, 4.IV.2011, leg. Y.-T. Wang
(BMNH, RBCN); 1♂, 1♀, same locality, 17.III.2012, leg. T. H. Lee (MCSN); 1♂, 1♀, same locality, 17.III.2012,
leg. M.-H. Tsou (TARI); 1♂, 1♀, same locality, 14.IV.2013, leg. B.-X. Guo (TARI); 4♀, same locality, 14.IV.2014,
leg. Y.-T. Chung (TARI).
Diagnosis. Aulacophora analis is similar to A. bicolor with yellowish brown body and black spots on the
elytron but A. analis display the characteristic and uniform patterns of arrangement of black spots on the elytron
which is different from A. bicolor.
Males. Length 5.5–7.0 mm, width 2.7–3.7 mm. General color (Figs 1–2) yellowish brown but elytron with two
large, black square patches at antero-lateral angles and behind middle respectively, apical 2/3 of tibia, and tarsus
black. Antenna (Fig. 10) filiform and extremely slender, antennomere I not enlarged; ratio of length of
antennomeres III to XI about 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 0.9 : 0.9 : 0.8 : 1.0; ratio of length to width from
antennomere III to XI about 4.3 : 4.3 : 4.6 : 4.9 : 4.7 : 4.7 : 5.0 : 4.7 : 5.9. First tarsomeres of front and middle legs
swollen. Apex of abdominal tergite VIII weakly emarginate (Fig. 16), convex at middle, with several long setae
along apical margin; base weakly sclerotized. Median lobe of abdominal ventrite V rectangular, disc weakly
depressed. Penis (Figs 12–13) slender, parallel-sided, abruptly narrowed from apical 1/6 to apex, apex pointed;
almost straight in lateral view, curved at apical 1/6; tectum well sclerotized, slender, apically widened, apex
rounded; endophallus with clusters of dense stout setae, and with one longitudinal sclerite, apex weakly recurved
and rounded, basally widened.
Females. Length 7.8–8.2 mm, width 4.0–4.5 mm. Similar to male (Fig. 3), but antennomeres III–V slender
(Fig. 11); ratio of length of antennomeres III to XI about 1.0 : 1.1 : 1.1 : 1.1 : 1.1 : 1.0 : 1.0 : 0.9 : 1.2; ratio of length
to width from antennomere III to XI about 3.5 : 3.9 : 4.1 : 4.2 : 4.3 : 4.2 : 4.6 : 4.5 : 5.6. Apical margin of abdominal
ventrite V truncate. Gonocoxae (Fig. 14) wide, apex of each gonocoxa with eight setae from apical 1/6 to apex;
gonocoxae connected at middle, base slender. Ventrite VIII (Fig. 15) weakly sclerotized; apex narrow, apical
margin a little emarginate at middle, surface with extremely dense short setae along apical margin, disc with dense
short setae; spiculum short. Spermathecal receptaculum (Fig. 17) a little swollen, hardly separated from pump;
pump strongly curved; spermathecal duct short, stout, shallowly projecting into receptaculum.
Host plant. Cucurbitaceae: Trichosanthes quinquangulata A. Gray (present study).
Distribution. Indonesia, Malaysia (Sabah), Philippines, Taiwan (only Lanyu Island).
Aulacophora bicolor (Weber, 1801)
(Figs 18–34)
Galleruca bicolor Weber, 1801: 56 (Indonesia: Sumatra); Fabricius, 1801: 482 (redescription).
Aulacophora bicolor: Baly, 1886: 19 (Indonesia: Sumatra, Java, Sulawesi, Gilolo, Flores, Lombok; Philippines; Sri Lanka);
Allard, 1888: 320 (Borneo); Allard, 1889: 308 (Cambodia: Pnomh Penh); Allard, 1891: 230 (Laos: Luang Prabang);
Chûjô, 1935b: 205 (Japan: Ryukyus); Maulik, 1936: 187 (India, including Andaman and Nicobar; Taiwan); Chen & Kung,
1959: 374 (China: Yunnan); Chûjô, 1962: 86 (Taiwan); Gressitt & Kimoto, 1963: 484 (China: Hainan, Vietnam); Kimoto,
1964: 305 (Japan: Ryukyu islands); Kimoto, 1989: 51 (Thailand, Laos); Mohamedsaid, 1994: 378 (Malaysia);
Mohamedsaid, 2000: 348 (Malaysia); Barroga, 2001: 40 (Indonesia: Bali).
Ceratia bicolor: Miwa, 1931: 189 (Taiwan).
Galleruca haemorrhoae Fabricius, 1803: 293 (new name for bicolor Fabricius, 1801).
Galeruca sexpunctata Olivier, 1808: 627 (Timor)
Aulacophora sexpunctata: Jacoby, 1884b: 213 (Sumatra); Allard, 1888: 322 (India); Schönfeldt, 1890: 174 (Japan: Ryukyus);
Duvivier, 1891: 146 (Java).
Ceratia (Orthaulaca) sexpunctata: Weise, 1922: 62.
Aulacophora bicolor ab. sexpunctata: Chûjô, 1935a: 83.
Aulacophora hemichroa Gistel, 1857 (Java): 540; Monrós & Bechyně, 1956: 1132 (as synonym of bicolor).
Aulacophora sexnotata Chapuis, 1876: 100 (Philippines); Weise, 1922: 62 (as synonym of sexpunctata).
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Aulacophora bicolor var. sexnotata: Baly, 1886: 19; Allard, 1888: 320 (Philippines);
Aulacophora semiopaca Jacoby, 1886: 51; Baly, 1887: 268 (as synonym of bicolor).
Type material. Galleruca bicolor: lectotype ♂ (KIEL), here designated, labeled: “Weber / bicolor. / 1723 [w, h] //
bico / lor [w, h] // Aulacophora bicolor (Weber) / det. by G.F. Barroga, 1999 [w, p]. There are two types of G.
bicolor in the KIEL collection. The other is attributed to A. ritsemae.
Galeruca sexpunctata: most of Olivier’s specimens were in very bad condition when given to MNHN in 1995.
The types should be destroyed by pests. (Mantilleri personal communication).
Aulacophora hemichora: Gistel’s specimens are in very bad condition and most of them didn’t bear any label.
Thus the types are either lost or not detectable anymore .
Aulacophora sexnotata: Chapuis’s types were originally deposited at the IRSB. They was studied and
borrowed by Barroga and Mohamedsaid (2002), and lost during return of the specimens (Limbourg personal
communication).
Aulacophora semiopaca: lecotype ♂ (MCSN), here designated, labeled: “Sumatra / Mto. Singalang / Luglio
1878 O. Beccari [w, p] // Typus [w, p, red letters and borders] // semiopaca / Jac. [w, h] // Aulacophora / semiopaca
Jac [b, h] // SYNTYPUS [p] / Aulacophora / semiopaca / Jacoby, 1886 [r, h] // Museo Civico / di Genova [w, p]”.
Paralectotypes: 4♂, 3♀ (MCSN), labeled: “Sumatra / Mto. Singalang / Luglio 1878 O. Beccari [w, p] //
SYNTYPUS [p] / Aulacophora / semiopaca / Jacoby, 1886 [r, h] // Museo Civico / di Genova [w, p]”; 2♀ (MCZC),
labeled: “Sumatra / Mto. Singalang / Luglio 1878 O. Beccari [w, p] // 1st Jacoby / Coll. [w, p] // ♀ [w, h] // Type [p]
/ 17568 [r, h] // semiopaca Jac. [b, h]”; 1♀ (BMNH), labeled: “Type / H. T. [w, p, round label with red border] //
Sumatra / Mto. Singalang / Luglio 1878 O. Beccari [w, p] // Jacoby Coll. / 1909-28a. [w, p] // semiopaca Jac [b, h]
// SYN- / TYPE [w, p, round label with blue border]”.
Other specimens examined. CHINA. Hainan: 1♂, Dwa Bi, 25.VII.1935, leg. L. Gressitt (BPBM);
INDONESIA. 1♂, Gilolo island, leg. Baly (BMNH); Java: 5♂, 2♀, Noesa Kembangan, leg. Drescher (MNHUB);
9♂, 17♀, Sukabumi, 1893, leg. H. Fruhstorfer (MNHUB); Sumatra: 1♂, Ajer Mantoior, Agosio, 1878, leg. O.
Beccari (BMNH); 2♀, Brastagi, 17.VII.1972, leg. Erber (MNHUB); 1♀, Bukkitinggi, Canyon, 7.IX.1979, leg.
Erber (MNHUB); 1♀, Dolok Meraugir, 5.VII.1974, leg. Diehl (MNHUB); 1♂, same locality, 5.X.1974, leg, Diehl
(MNHUB); 2♀, same locality, 15.XII.1974, leg. Diehl (MNHUB); 1♂, 1♀, Mt. Singalang, Luglio, 1878, leg.
Obeccari (BMNH); JAPAN. Okinawa: 4♂, 2♀, Yonaguni Island, 28.III.2014, leg. Y.-T. Wang (TARI); Nansei
Shotō: 1♂, 2♀, Iriomote, Funaura, 30 IV 1996, F.J. de Vries (RBCN); LAOS. 1♀, Luang Namtha env., 800–1200
m, V.1997, leg. native collector (JBCB); Vientiane: 1♂, Ban Van Eue, 15–31.V.1965, leg. native collector (BPBM);
1♂, same locality, 31.VII.1965, leg. native collector (BPBM); 1♀, same locality, 31.V.1966, leg. native collector
(BPBM); 1♂, same locality, 1–15.IX.1967, leg. native collector (BPBM); 1♀, Phou Kou Khouei, 31.V.1966, leg.
native collector (BPBM); PHILIPPINES. 2♀ (BMNH); TAIWAN . Chiayi: 2♂, Chungpu, 3.III.2010, leg. H.-T.
Shih (TARI); Ilan: 1♂, Chinyang, 23.X.2011, leg. C.-H. Hsieh (TARI); Kaoshiung: 3♂, Kosempo (= Chiasien), 1–
20.III.1908, leg. H. Sauter (BMNH); 1♂, Namahsia, 11.I.2013, leg. B.-X. Guo (TARI); 1♀, Tengchih, 2–
5.VI.2008, leg. C.-F. Lee (TARI); 1♂, same locality, 19.III.2013, leg. Y.-T. Chung (TARI); 1♀, same locality,
8.VI.2013, leg. W.-C. Liao (TARI); Nantou: 1♀, Hsitou, 15.VI.2011, leg. C.-F. Lee (TARI); 2♀, Lugu, 10.IX.2014,
leg. H.-T. Shih (TARI); 1♂, Wanfengtsun, 4.X.2007, W.-T. Liu (TARI); Pingtung: 1♀, Chiehchihlaishan,
14.II.2012, leg. J.-C. Chen (TARI); 1♂, 1♀, Kenting, 24.XI.2009, leg. C.-F. Lee (TARI); 1♂, Lilungshan,
13.III.2012, leg. J.-C. Chen (TARI); 1♀, same locality, 8.III.2014, leg. J.-C. Chen (TARI); 1♂, 2♀, Machia,
11.III.2013, leg. Y.-T. Chung (MNHUB); 1♀, Nanjenshan, 19.X.2008, leg. W.-C. Wang (TARI); 2♂, Neiwen,
24.II.2012, leg. J.-C. Chen (TARI); 1♂, Peitawushan, 4.IV.2013, leg. Y.-T. Chung (TARI); 4♂, Shouka, 24.II.2012,
leg. M.-H. Tsou (RBCN); 1♂, 1♀, Sulin, 15.IV.2009, leg. U. Ong (TARI); 1♂, Tahanshan, 28.VIII.2010, leg. Y.-L.
Lin (TARI); 1♂, 1♀, same locality, 25.II.2012, leg. J.-C. Chen (TARI); 1♀, 3.IX.2012, leg. Y.-T. Chung (TARI);
1♀, same locality, 29.IV.2014, leg. J.-C. Chan (TARI); 1♂, same locality, 23.V.2014, leg. Y.-T. Chung (TARI);
Tainan: 2♂, Meiling, 14.IV.2010, leg. B.-X. Guo (MNHUB); 2♀, same locality, 24.IV.2013, leg. B.-X. Guo
(MNHUB); Taipei: 1♀, Fushan, 1.IV.2011, leg. S.-F. Yu (TARI); 1♀, Jiajiuliao, 18.VIII.2009, leg. Y.-T. Wang
(TARI); 1♀, Taipei City Zoo, 12.VI.2007, leg. C.-F. Lee (TARI); Taitung: 1♀, Tajen, 18.X.2008, leg. W.-C. Wang
(TARI); Yulin: 1♀, Touliu, 4.I.2009, leg. H.-T. Shih (TARI); THAILAND. Mae Hong Son: 2♀, SE of Soppong,
1500 m, 23–27.V.1999, leg. M. Říha (JBCB); Phitsanulok: 1♀, Nam Muang vill., 450 m, 3.V.1999, leg. Říha
(JBCB); VIETNAM. 2♂, Buonlol, 1982, leg. L. Medvedev (MNHUB); 1♂, Hoa-Binh, leg. A. Cooman (BPBM);
1♀, Nha-Trang, 1987, leg. M. Opletal (JBCB).
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FIGURES 18–26. Habitus of Aulacophora bicolor (Weber). 18. Male, Taiwan, dorsal view; 19. Ditto, ventral view; 20.
Female, Taiwan, dorsal view; 21. Female, Java, dorsal view; 22. Male, Java, dorsal view; 23. Male, Java, dorsal view; 24.
Female, Sumatra, dorsal view; 25. Female, Sumatra, dorsal view; 26. Male, Vietnam, dorsal view.
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FIGURES 27–34. Diagnostic characters of Aulacophora bicolor (Weber), 27. Antenna, male; 28. Antenna, female; 29. Penis,
dorsal view; 30. Penis lateral view; 31. Gonocoxae; 32. Abdominal ventrite VIII, female; 33. Abdominal tergite VIII, male; 34;
Spermatheca.
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Males. Length 6.6–8.2 mm, width 3.3–4.3 mm. General color (Figs 18–19) yellowish brown but meso- and
metathoracic and abdominal ventrites black or blackish brown, middle and hind legs dark brown, outer margins of
tibia, and tarsi of front legs dark brown; elytron with two black spots near base, one at humerus, the other placed
behind scutellum, and with one transverse black band behind middle and usually intercepted by suture. Antenna
(Fig. 27) filiform and extremely slender, antennomere I not enlarged; ratio of length of antennomeres III to XI
about 1.0 : 1.0 : 0.9 : 0.9 : 0.9 : 0.9 : 0.8 : 0.8 : 0.9; ratio of length to width from antennomere III to XI about 3.8 :
3.8 : 3.5 : 3.5 : 3.4 : 3.5 : 3.3 : 3.3 : 4.0. Apex of abdominal tergite VIII truncate (Fig. 33), weakly convex at middle,
with several short setae along apical margin; base weakly sclerotized. Median lobe of fifth abdominal ventrite
rectangular, disc weakly depressed. Penis (Figs 29–30) slender, widest at apical 1/3, apically tapering, apex
pointed; narrowest at apical 3/5; almost straight in lateral view, curved at apical 1/3; tectum membranous, parallel-
sided, apex truncate; endophallus with clusters of several stout setae, and with one longitudinal sclerite, extremely
curved at apical 1/3, basally widened.
Females. Length 8.8–9.8 mm, width 4.3–4.9 mm. Similar to male (Fig. 20), but antennomeres III–V wider
(Fig. 28); ratio of length of antennomeres III to XI about 1.0 : 1.1 : 1.1 : 1.0 : 1.0 : 1.0 : 1.0 : 0.9 : 1.2; ratio of length
to width from antennomere III to XI about 3.3 : 3.7 : 3.6 : 3.3 : 3.3 : 3.2 : 3.3 : 3.3 : 4.5. Apical margin of abdominal
ventrite V truncate. Gonocoxae (Fig. 31) wide, apex of each gonocoxa with seven or eight setae from apical 1/6 to
apex; gonocoxae connected at middle, base slender. Ventrite VIII (Fig. 32) weakly sclerotized; apex narrow, apical
margin a little emarginate at middle, surface with dense short setae along apical margin; spiculum long.
Spermathecal receptaculum (Fig. 34) strongly swollen; pump strongly curved; spermathecal duct short, stout,
shallowly projecting into receptaculum.
Va ri at io n . Some specimens collected from China, Taiwan, and Japan have only one pair of black spots on
elytral humerus or even no black spots at all; specimens from China, and Southeast Asia have yellow brown legs.
Specimens collected from Java have their characteristic color patterns: in addition to typical form, some have well-
developed posterior spots and form transverse band (Fig. 21); some have well-developed anterior and posterior
spots and form transverse bands (Fig. 22); some have the black apical half (Fig. 23). Specimens collected from
Sumatra have also their characteristic color patterns: all have black middle and hind legs; some have black basal 1/
5 and one pair of transverse bands at apical 1/3 on elytra (Fig. 24); others have black basal 2/3 of elytra, with one
pair of yellow spots at base near humerus (Fig. 25). One specimen from Vietnam has black elytra with yellow
apices and one pair of small yellow spots at base near humerus (Fig 26).
Remarks. Some authors (such as Kimoto 1989; Barroga & Mohamedsaid 2002) thought that Galleruca
bicolor described by Fabricius (1801) was different from that by Weber (1801). Actually, Fabricius (1801)
redescribed Galleruca bicolor based on Weber’s types (Zimsen 1964).
Host plants. Cucurbitaceae: Zehneria mucronata (Bl.) Miq. (present study).
Distribution. Cambodia, China, India, Indonesia, Japan, Laos, Malaysia, Philippines, Sri Lanka, Taiwan,
Thailand, Vietnam.
Aulacophora frontalis Baly, 1888
(Figs 35–36, 44–51, 111)
Aulacophora frontalis Baly, 1888: 181 (Malaysia: Sarawak); Jacoby, 1896: 126 (Indonesia: Mentawai islands); Maulik, 1936:
183 (India); Chen & Kung 1959: 375 (China: Yunnan, Guandong); Chûjô, 1962: 96 (Taiwan); Gressitt & Kimoto, 1963:
487 (China: Guandong; Vietnam); Kimoto, 1989: 55 (Thailand, Laos, Cambodia, Vietnam); Mohamedsaid, 1994: 380
(Malaysia); Barroga, 2001: 42 (Indonesia: Bali); Barroga & Mohamedsaid, 2002: 97 (Malaysia, Indonesia: Sumatra, Java);
Mohamedsaid, 2000: 348 (Malaysia); Mohamedsaid & Constant, 2007: 166 (Thailand, Cambodia).
Aulacophora palliata: Allard, 1889: 308 (Vietnam: Hué, Saigon = Ho Chi Minh City, Qui Nhon, Tourane; Cambodia: Phnom
Penh); Allard, 1891: 230 (Laos: Luang Prabang); Laboissière, 1935: 140 (listed as frontalis).
Aulacophora (Ceratia) frontalis: Laboissière, 1929: 258 (Vietnam).
Type material. Lectotype ♂ (BMNH), here designated, labeled: “SYN- / TYPE [w, p, round label with blue
border] // Type [w, p, round label with red border] // SAR / 305 [w, h, round label] // Baly Coll. [w. p] //
Aulacophora / frontalis / Baly / Type [w, h] // This specimen was found without name attached, in Baly’s coll. in
close proximity to A. orientalis or A. palliata. It was the only specimen of this group of species having loc.
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Sarawak. It is undoubtedly the type. [w, h] // holotype [w, p]”. Paralectotypes: 1♂ (BMNH), labeled: “SYN- /
TYPE [w, p, round label with blue border] // Type [w, p, round label with red border] // [blue round label] // Borneo
[w, h] // Baly Coll. [w, p] // Aulacophora / frontalis / Baly (♂) [w, p] // not type [w, p]”. Although this specimen
was labeled as nontype, it cannot be excluded from the type series by original description. 1♂ (MCZC), labeled
“SAR. [w, h, round label] // A. frontalis Baly [w, h] // 1st Jacoby / Coll. [w, p] // TYPE / F. C. B. Coll. [pink label,
p] // Type [p] / 17571 [r, h]”; 1♂ (MCZC), labeled: “SUMAT [w, p] // Jacoby 2nd / Coll. [w, p] // dorhni / Jac [w, h]
// Type [p] / 17572 [w, h]”.
Other specimens examined. CAMBODIA. 1♂, Siem Reap, 7–11.IX.2002, leg. P. Kočárek (JBCB); INDIA.
Kamataka: 1♂, Coorg distr., 10 km SE of Virajpet, leg. Z. Kejval and M. Trýzna (JBCB); INDONESIA. Java: 2♂,
1♀, Batavia, Tanjong Priok, (BMNH); 1♂, Noesa Kembangan, leg. Drescher (MNHUB); Sumatra: 2♂,
Soekaranda, leg. Dohrn (MNHUB); 1♂, Nias island (BMNH); LAOS. Borikhane: 1♂, Paksane, 28.VIII.1965, leg.
native collector; 1♀, same locality, 1.XI.1965, leg. native collector (BPBM); Hua Phan: 1♂, Phu Loei N. P., Ban
Sakok, 23–26.V.2001, leg. J. Bezdĕk (JBCB); Khammouane: 2♀, Phon Tiou, 21–22.IX.1965, leg. native collector
(BPBM); 1♂, Ban Khoun Ngeun env., 200 m, 19–31.V.2001, leg. C. L. Peša (JBCB); Vientiane: 1♀, Ban Van Eue,
30.XI.1965, leg. native colletor (BPBM); 1♂, Tha Ngone, 6.IX.1965, leg. native collector (BPBM); MALAYSIA.
Sabah: 1♂, Tenompok, 15.II.1959, leg. T. C. Maa (BPBM); Sarawak: 1♂, Kapit Dist. Merirai,V., 1–6.VIII.1958,
leg. T. Maa (BPBM); 1♂, Mt. Matang, XII.1913, leg. G. E. Bryang (BMNH); 2♀, Shelford, 1900 (BMNH);
TAI WAN. Heitou (= Pingtung), 2♂, 1♀, 22.V.1930, leg. Y. Miwa (TARI); 4♀, same locality, V–VI.1931, leg. Y.
Miwa (TARI); THAILAND. Cholburi: 1♂, Sriracha Dist., 22.X.1966, leg. J. S. Burto (BPBM); VIETNAM. 1♂,
Minh Hoa, N. of Nha Trang, 28.XI.1960, leg. C. M. Yoshimoto (BPBM); 1♂, Saigon, X.1982, leg. Pokomý
(JBCB).
Diagnosis. Aulacophora frontalis is similar to A. palliata with the modified antennomeres III to V in male but
A. frontalis have the longitudinal groove on the vertex (Fig. 111) (the transverse ridges in A. palliata: Fig. 112) or
more slender antennomere III in male, and more wider antennomere VII to X in female.
Males. Length 5.7–6.7 mm, width 2.8–3.5 mm. General color (Figs 35–36) yellowish brown except elytron
black and shining. Vertical area of head with a longitudinal groove on each side (Fig. 111). Antenna (Fig. 44)
filiform, scape not enlarged, antennomere I swollen and curved; III–V triangular, III with lateral expansion
flattened, V with longitudinal groove on lateral expansion; ratio of length of antennomeres III to XI about 1.0 : 0.8
: 0.8 : 0.7 : 0.7 : 0.7 : 0.7 : 0.7 : 0.8; ratio of length to width from antennomere III to XI about 2.5 : 1.5 : 1.5 : 2.8 :
2.9 : 3.1 : 3.2 : 3.4 : 5.2. Apex of abdominal tergite VIII emarginate at middle (Fig. 50), apical margin sinuate and
with some setae, base weakly sclerotized. Median lobe of fifth abdominal ventrite short and rectangular, disc
depressed near apex, apical margin weakly emarginate at middle. Penis (Figs 46–47) apically tapering from apical
1/4; weakly curved in lateral view; tectum well sclerotized, apically tapering; endophallus with dense, irregularly
arranged setae, and with one longitudinal sclerite, apex rounded, basally widened and bifurcate.
Females. Length 6.5–6.9 mm, width 3.5–3.8 mm. Similar to male, but transverse ridge on vertical area absent;
antenna filiform and not modified (Fig. 45), ratio of length of antennomeres III to XI about 1.0 : 0.8 : 0.8 : 0.8 : 0.8
: 0.8 : 0.8 : 0.8 : 1.0; ratio of length to width from antennomere III to XI about 3.4 : 3.1 : 2.8 : 2.8 : 2.9 : 3.1 : 3.2 :
3.1 : 4.6. Apical margin of abdominal ventrite V not modified, rounded. Gonocoxae (Fig. 48) slender, apex of each
gonocoxa with eight or nine setae from apical 1/6 to apex; gonocoxae connected at middle, base slender. Ventrite
VIII (Fig. 49) weakly sclerotized; apex narrow, apical margin rounded, surface with dense short setae along apical
margin, and several long setae scattered at apical area; spiculum short. Spermathecal receptaculum (Fig. 51)
slightly swollen, hardly separated from pump; pump strongly curved; spermathecal duct short, stout, shallowly
projecting into receptaculum.
Host plant. Unknown
Distribution. Cambodia, China, India, Indonesia, Laos, Malaysia, Taiwan, Thailand, Vietnam. Only very old
specimens collected from Taiwan had been found. Chûjô (1962, 1963) reported this species for Taiwan based on
Miwa’s specimens collected in 1930s (examined at the present study), and Sauter’s ones in 1900s. No recent
specimens have been collected. It is supposed that this species is extinct in Taiwan.
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FIGURES 35–43. Habitus of Aulacophora species. 35. A. frontalis, Laos, male, dorsal view; 36. Ditto, ventral view; 37. A.
indica, Taiwan, male, dorsal view; 38. Ditto, ventral view; 39. A. indica, Taiwan, female, dorsal view; 40. Ditto, ventral view;
41. A. indica, China, female, dorsal view; 42. Lectotype of Crioceris testacea, dorsal view; 43. Ditto, ventral view.
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FIGURES 44–51. Diagnostic characters of Aulacophora frontalis Baly. 44. Antenna, male; 45. Antenna, female; 46. Penis,
dorsal view; 47. Penis lateral view; 48. Gonocoxae; 49. Abdominal ventrite VIII, female; 50. Abdominal tergite VIII, male; 51;
Spermatheca.
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Aulacophora indica (Gmelin, 1790)
(Figs 37–43, 52–61)
Crioceris testacea Fabricius, 1787: 87 (India); Fabricius, 1792: 4 (redescription).
Aulacophora testacea: Baly, 1879: 445 (India: Assam); Baly, 1886: 13; Allard, 1888: 320 (Indonesia: Celebes = Sulawesi);
Cryptocephalus (Crioceris) indica Gmelin, 1790: 1720. (replacement name for Crioceris testacea Fabricius, 1787)
Rhaphidopalpa indica: Laboissière, 1940: 10.
Aulacophora indica: Kimoto, 1970: 416 (Nepal); Kimoto, 1977: 354 (Bhutan); Kimoto, 1989: 57 (India, including Andaman
and Nicobar islands, Sri Lanka, Burma, Thailand, Cambodia, Laos, Vietnam, China, Taiwan, Philippines, Japan, including
Ryukyu islands, Korea, Russia, Indonesia: Sunda islands, Micronesia, New Guinea, Samoa, Fiji); Mohamedsaid, 1994:
381 (Malaysia); Mohamedsaid, 2000: 348 (Malaysia); Barroga, 2001: 42 (Indonesia: Bali); Barroga & Mohamedsaid,
2002: 102 (Indonesia: Sumatra, Nias, Buru, Java); Mohamedsaid & Constant, 2007: 166 (Thailand, Cambodia); Aston,
2009: 13 (China: Hong Kong).
Galeruca similis Olivier, 1808: 624 (Océan Indien); Kimoto, 1989: 56 (as synonym of indica).
Aulacophora similis: Baly, 1886: 16 (China, Japan, India, Malaysia); Jacoby, 1889: 206 (Burma); Duvivier, 1892: 430 (India:
Konbir); Chûjô, 1938: 152 (Taiwan); Gressitt, 1955: 28 (Micronesia).
Rhaphidopalpa similis: Allard, 1889: 308 (Vietnam: Hué, Saigon = Ho Chi Minh City, Mytho, Qui Nhon, Tourane; Cambodia:
Pnom Penh); Allard, 1891: 230 (Laos: Luang Prabang).
Orthaulaca similis: Weise, 1892: 393.
Ceratia similis: Miwa, 1931: 190 (Taiwan).
Raphidopalpa femoralis Motschulsky, 1857: 37 (Japan); Baly, 1886: 16 (as synonym of similis); Weise, 1892: 395
(Philippines); Ogloblin, 1936: 154 (Korea).
Aulacophora femoralis: Schönfeldt, 1890: 174 (Japan: Ryukyus); Chûjô, 1941: 157 (Japan, Korea); Gressitt & Kimoto, 1963:
486 (China, Vietnam); Kimoto, 1966: 28 (Taiwan).
Rhaphidopalpa coffeae: Allard, 1888: 319 (Japan; misidentification); Weise, 1892: 395 (as synonym of similis).
Aulacophora flavipes Jacoby, 1883: 202 (Indonesia: Sulawesi: Saleyar islands); Baly, 1886: 17 (as synonym of similis).
Rhaphidopalpa pubescens Allard, 1888: 306 (Annam). New synonym
Aulacophora pubescens: Kimoto, 1989: 62.
Rhaphidopalpa bengalensis Weise, 1892: 394 (India: Calcutta).
Rhaphidopalpa ceramensis Weise, 1892: 394 (Indonesia: Ceram island; Borneo); Laboissière, 1932: 153 (Indonesia:
Halmaheira); Barroga & Mohamedsaid, 2002: 103 (as synonym of indica).
Rhaphidopalpa niasiensis Weise, 1892: 394 (Indonesia: Nias and Sumatra); Barroga and Mohamedsaid, 2002: 103 (as
synony m of indica).
Rhaphidopalpa chinensis Weise, 1892: 395 (China: Shanghai); Ogloblin, 1936: 153 (Vietnam, Mongolia); Laboissière, 1940:
13 (India: Tetara); Gressitt & Kimoto, 1963: 486 (as synonym of femoralis).
Aulacophora femoralis chinensis: Chen & Kung, 1959: 375 (China: Hebei, Shaanxi, Shandong, Jiangsu, Zhejiang, Fujian,
Guandong, Guanxi, Hubei, Jiangxi, Sichuan, Yunnan).
Type material. Crioceris testacea: lectotype ♂ (KIEL), here designated, without labels (see remarks on A.
abdominalis).
Galeruca similis: most of Olivier’s were in very bad condition before given to MNHN in 1995. The types
probaly have been destroyed by pests. (Mantilleri personal communication).
Aulacophora flavipes: holotype ♀ (MCZC), labeled: “H.E.D. / Engelhard / Saleyer / 20/1, 81 [w, h, round
label] // 1st Jacoby / Coll. [w, p] // ♀ [b, h] // Type [p] / 17561 [r, h]”.
Rhaphidopalpa pubescens: lectotype ♂ (MNHN), here designated, labeled: “wchinchine [h, w] // Ex.Musæo /
1899 [vertical] / E.ALLARD [p, w] // M
USEUM
P
ARIS
/ 1952 / C
OLL
. O
BERTHUR
[p, w] // TYPE [p,r] //
Rhaphidopalpa / Pubescens All. / Cochinchine [p, y]”.
Rhaphidopalpa bengalensis: lectotype ♂ (MNHUB), here designated, labeled: “Calcutta / Hauser [w, h] // India or.
/ Calcutta [w, p] // ♂ [w, h] // SYNTYPUS / Rhaphidopalpa / bengalensis Weise, 1892 / labelled by MNHUB 2014
[r, p]”. Paralectotype: 1♀ (MNHUB), labeled: “Calcutta [w, h] / bengalensis / Ws. [w, h] // SYNTYPUS /
Rhaphidopalpa / bengalensis Weise, 1892 / labelled by MNHUB 2014 [r, p]”.
Raphidopalpa femoralis: not studied. They should be deposited at the ZMUM.
Rhaphidopalpa ceramensis: Lectotype ♀ (MNHUB), here designated, labeled: “Ceram / Jllo [w, h] // ex Coll. /
J. Weise [w, p] // ceramensis / Ws. [w, h] // Type [r, p] // SYNTYPUS / Rhaphidopalpa / ceramensis Weise, 1892 /
labelled by MNHUB 2014 [r, p]”. Paralectotypes: 1♂, 1♀ (MNHUB), labeled: “Ceram / Ribbe [w, h] // ex Coll. / J.
Weise [w, p] // Type [r, p] // SYNTYPUS / Rhaphidopalpa / ceramensis Weise, 1892 / labelled by MNHUB 2014 [r,
p]”; 1♂ (MNHUB), labeled: “Ceram / Jllo [w, h] // ex Coll. / J. Weise [w, p] // Type [r, p] // SYNTYPUS /
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Rhaphidopalpa / ceramensis Weise, 1892 / labelled by MNHUB 2014 [r, p]”; 1♀ (MNHUB), labeled: “ex Coll. / J.
Weise [w, p] // Type [r, p] // SYNTYPUS / Rhaphidopalpa / ceramensis Weise, 1892 / labelled by MNHUB 2014 [r,
p]”; 1♀ (MNHUB), labeled: “Ceram / Jllo / C. Ribbe 1884 [w, p] // ex Coll. / J. Weise [w, p] // Type [r, p] //
SYNTYPUS / Rhaphidopalpa / ceramensis Weise, 1892 / labelled by MNHUB 2014 [r, p]”.
Rhaphidopalpa niasiensis: Weise’s types were originally deposited at the MNHUB. They was studied and
borrowed by Barroga and Mohamedsaid (2002), and lost during return of the specimens (Willers personal
communication).
Rhaphidopalpa chinensis: lectotype ♂ (MNHUB), here designated, labeled: “Shanghai / Simon [w, h] // ♂ [w,
h] // chinensis / Ws. [w, h]”.This is the only specimen available in Weise’s collection. It is uncertain if there are
other type specimens.
Other specimens examined. CAMBODIA. 1♂, Siem Reap, 7–11.XI.2002, leg. P. Kočák (JBCB); CHINA.
Fujian: 1♀, Changting, Hotien, 19.IV.1941, leg. T. C. Maa (BPBM); 1♀, Chishui, 20.VI.2014, leg. Y.-T. Chung
(TARI); 1♀, Shaowu, Tachulan, 11.VI.1942, leg. T. Maa (BPBM); Guandong: 1♂, Lochang, Tintong,
18.VIII.1947, leg. L. Gressitt (BPBM); Hainan: 1♂, Fan Heang, 7.VI.1935, leg. J. L. Gressitt (BPBM); 1♂,
Tiaoloshan, 6.V.2011, leg. Y.-T. Chung (TARI); Hong Kong: 1♂, Lantau, Mui Wo, 24.IX.2005, leg. P. Aston
(PAHC); Hubei: 1♂, Wang-ga-ying, Lichuan, 23.IX.1948, leg. Gressitt & Djou (BPBM); Sichuan: 1♂, Wanhsien,
3.X.1948, leg. Gressitt & Djou (BPBM); INDONESIA. Sumatra: Riau: 2♂, Bukit Tigapuluh N. P., 18–25.I.2000,
leg. J. Bezdĕk (JBCB); JAPAN. Okinawa: 1♀, Okinawa island, 8.IV.2014, leg. Y.-T. Wang (TARI); 1♀, Yonaguni
island, 28.III.2014, leg. Y.-T. Wang (TARI); LAOS. Hua Phan: 1♂, Ban Kangpabong env., 25 km SE of Vient Xai,
14–18.V.2001, leg. J. Bezdĕk (JBCB); Khammouane: 1♂, Phon tiou, 6.VII.1965, leg. native collector (BPBM);
1♀, same locality, 11.IX.1965, leg. native collector (BPBM); Vientiane: 1♂, Tha Ngone, 20.XI.1965, leg. native
collector (BPBM); 1♂, Vientiane, 31.VII.1965, leg. native collector (BPBM); 1♀, same locality, 21.III.1966, leg.
native collector (BPBM); MALAYSIA. Johor: 1♂, 15 km NW of Kota Tinggi, Muntahak Mt., 13–14.V.2001, leg.
M. Řiha (JBCB); PHILIPPINES. Batanes: 4♂, 4♀, Itbayat island, 13.IV.2013, leg. Y.-F. Hsu (TARI); 1♀, Sabtang
island, 11.IV.2013, leg. Y.-F. Hsu (TARI); SOUTH KOREA. 2♂, Jeju-do, Jeju-si, Hwabuk-dong, 28.X.2009, leg.
H.-W. Cho (JBCB); THAILAND. Chiangmai: 1♂, Chiangdao, 5–11.IV.1958, leg. T. C. Maa (BPBM);
Nakornsawan: 1♀, 12.IX.1963 (BPBM); TAI WAN. Ilan: 1♂, Taipingthan, 3.VII.2010, leg. M.-H. Tsou (TARI);
1♂, Toucheng, 30.V.2009, leg. H.-J. Chen (TARI); Kaoshiung: 1♀, Meinung, 20.XII.2011, leg. J.-C. Chen (TARI);
4♂, 6♀, same locality, 17.II.2012, leg. J.-C. Chen (TARI); 5♂, 4♀, Hsiaokangshan, 9.III.2013, leg. B.-X. Guo
(TARI); 3♂, Yuangan, 16.IV.2013, leg. J.-C. Chen (TARI); Kinmen: 1♂, 1♀, Kinmen island, 11.VII.2001, leg. H.-
T. Shih (TARI); Pingtung: 1♂, Checheng, 6.II.2012, leg. J.-C. Chen (TARI); 1♂, Chiehchihlaishan, 14.II.2012, leg.
J.-C. Chen (TARI); 2♂, 3♀, Fengchuisha, 29.IV.2012, leg. J.-C. Chen (TARI); 1♂, Lilungshan, 10.III.2014, leg. Y.-
T. Chung (TARI); 1♂, Neiwen, 30.X.2012, leg. J.-C. Chen (TARI); Taipei: 3♀, Linshanpi, 15.IX.2007, leg. S.-F.
Yu (TARI); 1♀, Manyuehyuan, 9.IX.2011, leg. C.-F. Lee (TARI); 1♀, Menghuanhu, 28.VII.2007, leg. M.-H. Tsou
(TARI); 8♂, Pinglin, 8.VII.2008, leg. H.-J. Chen (TARI); 1♀, Sanchih, 11.III.2010, leg. W.-T. Liu (TARI); 1♂,
Wanli, 4.VII.2008, leg. W.-T. Liu (TARI); Taitung: 1♀, Lanyu island, 22–26.III.1998, leg. C.-F. Lee (TARI); 1♀,
same locality, 27.VI.2010, leg. H.-H. Lee (TARI); 1♀, same locality, 4.IV.2011, leg. Y.-T. Wang (TARI); 5♂, 5♀,
same locality, 17.III.2012, leg. M.-H. Tsou (RBCN, TARI); 5♂, 5♀, same locality, 17.III.2012, leg. T.-H. Lee
(MCSN, MNHUB); 1♀, same locality, 5.V.2012, leg. S.-F. Yu (TARI); 1♀, same locality, 14.IV.2013, leg. Y.-T.
Chung (TARI); 10♂, 12♀, Lutao island, 14.IV.2014, leg. Y.-T. Chung (TARI); 1♂, Taimali, 21.X.2009, leg. W.-T.
Liu (TARI); Taoyuan: 1♀, Fuhsing, 15.VII.2009, leg. H.-J. Chen (TARI); 2♂, 3♀, Lofu, 11.IV.2010, leg. M.-H.
Tsou (TARI); 1♂, Paling, 11.IV.2010, leg. M.-H. Tsou (TARI); THAILAND. Mae Hong Son: 1♂, 1♀, Soppong
env., 600 m, 28.V.–2.VI.1999, leg. M. Řiha (JBCB); VIETNAM. 1♀, Ap Hung-Lam, 21 km NW of Dilinh, 29.IX–
5.X.1960, leg. C. M. Yoshimoto (BPBM); 1♂, Nha Trang, 17–26.XI.1960, leg. C. M. Yoshimoto (BPBM).
Diagnosis. See diagnosis of Aulacophora abdominalis.
Males. Length 6.5–8.0 mm, width 3.3–4.0 mm. General color (Figs 37–38) yellowish brown but metathoracic
and abdominal ventrites black except apex of fifth abdominal ventrite; middle and hind legs black; outer margins of
femur and tibia of front legs black; labrum dark brown; antenna dark brown except three basal antennomeres
yellowish brown. Antenna (Fig. 52) filiform and slender, antennomere I enlarged; ratio of length of antennomeres
III to XI about 1.0 : 1.1 : 1.0 : 1.0 : 1.0 : 0.9 : 1.0 : 1.0 : 1.2; ratio of length to width from antennomere III to XI
about 2.6 : 3.0 : 3.2 : 3.0 : 3.1 : 3.0 : 3.1 : 3.7 : 4.4. Pronotum with deep, transverse groove. Elytra with cluster of
erect hairs behind humerus. Abdominal tergite VIII well sclerotized (Fig. 58), apical margin bifurcate and apices
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cute; with one small process at lateral margin. Median lobe of fifth abdominal ventrite rectangular, disc depressed
at left side. Penis (Figs 54–55) slender, parallel-sided, asymmetric and abruptly narrowed at apical 1/7 and apically
tapering, apex recurved; almost straight in lateral view; tectum medially sclerotized, apically tapering; endophallus
with clusters of short setae, and with one longitudinal sclerite, apically tapering, apex strongly curved in lateral
view, basally widened.
FIGURES 52–61. Diagnostic characters of Aulacophora indica (Gmelin). 52. Antenna, male; 53. Antenna, female; 54. Penis,
dorsal view; 55. Penis lateral view; 56. Gonocoxae; 57; Abdominal ventrite VIII, female; 58. Abdominal tergite VIII, male; 59.
Abdominal tergite VIII, paralectotype of Rhaphidopalpa ceramensis; 60. Apex of abdominal ventrite V, female; 61;
Spermatheca.
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Females. Length 6.8–8.2 mm, width 3.4–4.1 mm. Similar to male (Figs 39–40), but pronotum with transverse
groove shallow; cluster of erect hairs on elytra absent; antennomere I not enlarged (Fig. 53); ratio of length of
antennomeres III to XI about 1.0 : 1.0 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 1.1; ratio of length to width from
antennomere III to XI about 3.5 : 3.7 : 3.3 : 3.3 : 3.3 : 3.3 : 3.5 : 3.5 : 4.5. Pygidium projecting from elytral apex,
apically tapering, apex pointed, or truncate, or emarginate. Middle of apical margin of abdominal ventrite V (Fig.
60) emarginate. Gonocoxae (Fig. 56) slender, apex of each gonocoxa with eight setae from apical 1/6 to apex;
gonocoxae connected at middle, base slender. Ventrite VIII (Fig. 57) weakly sclerotized; apex narrowly rounded,
with dense short setae along apex; spiculum short. Spermathecal receptaculum (Fig. 61) a little swollen; pump
strongly curved; spermathecal duct short, stout, shallowly projecting into receptaculum.
Vari a t i o n. Although most of the females have pointed pygidia, few have truncate or emarginate ones. Color of
middle and hind legs are also variable among different populations. Black middle and hind legs occur in Taiwan,
Japan, Philippines, and Indonesia, while yellow ones (Fig. 41) in India, Laos, Thailand, and Vietnam. Both color
forms occur in China. Some exceptions could occur in small populations, such as adults with yellow legs occur in
Saleyar islands.
Remarks. The type(s) of Crioceris testacea is mixed with those of C. abdominalis in the Fabricius collection
(KIEL). One male is designated as the lectotype to serve nomenclatural stability. This specimen (Figs 42–43) fits
the current definition of Aulacophora indica and the color patterns (yellow legs) in India. For more details on the
designation of this lectotype see under remarks in the section on A. abdominalis. In addition, Kimoto (1989)
indicated that Rhaphidopalpa pubsecens should belong to another genus with dense hairs on the dorsal surface
although the type was not studied. Actually the character indicated by Kimoto (1989) is the erect haris behind the
humerus of the male since this type is deposited at the MNHN and regarded as a junior synonym of Aulacophor
indica after studying its aedeagus.
Host plants. Cucurbitaceae: including cultivated species.
Distribution. Bhutan, Burma, Cambodia, China, Fiji, India, Indonesia, Japan, Korea, Laos, Malaysia,
Micronesia, Nepal, Papua New Guinea, Philippines, Russia, Samoa, Sri Lanka, Taiwan, Thailand, Vietnam.
Aulacophora kotoensis Chûjô
(Figs 4–7, 62–65, 67, 69–72)
Aulacophora kotoensis Chûjô, 1962: 79 (Taiwan).
Type material. Aulacophora kotoensis: holotype ♂ (TARI), labeled: “KOTOSHO / (BOTEL-TOBAGO IS.) (=
Lanyu island) / FORMOSA / 20.VI–10.VII.1938 / COLL. M. CHUJO [w, p] // Ho / Type [w, p, round label, red
letters and border, but faded out] // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1813 [w, p]”.
Paratypes: 1♀ (TARI), labeled: “KOTOSHO / (BOTEL-TOBAGO IS.) (= Lanyu island) / FORMOSA / 20.VI–
10.VII.1938 / COLL. M. CHUJO [w, p] // ALLo / Type (w, p, round label, gray letters and border) // Aulacophora /
kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1811 [w, p]”; 1♂, 1♀ (TARI), labeled: “KOTOSHO / (BOTEL-
TOBAGO IS.) (= Lanyu island) / FORMOSA / 20.VI–10.VII.1938 / COLL. M. CHUJO [w, p] // Para / Type (w, p,
round label, green letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1820 & 676
[w, p]”; 1♂, 1♀ (TARI), labeled: “KOTOSHO / FORMOSA / IV.1936/ COLL. Y. CHUJO [w, p] // Para / Type (w,
p, round label, green letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1823 &
684 [w, p]”; 2♂, 2♀ (TARI), labeled: “Kotosho / 10 III–14.IV.1920 / Sonan [w, h] // Para / Type (w, p, round label,
green letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1422-1424, 2599 [w,
p]”; 2♂ (TARI), labeled: “Kotosho / 20.IX.1923 / Col. T. Okuni. [w, p] // Para / Type (w, p, round label, green
letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 1425 & 1426 [w, p]”; 1♂, 2♀
(TARI), labeled: “Formosa / Kotosho / III–IV.1932 / S. Hirayama [w, p] // Para / Type (w, p, round label, green
letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 681-683 [w, p]”; 2♂, 2♀
(TARI), labeled: “[Koutousyo] / FORMOSA / XII-1931-II-1932 / S. Hirayama // Para / Type (w, p, round label,
green letters and border) // Aulacophora / kotoensis / Chûjô [h] / DET. M. CHUJO [w, p] // 677-680 [w, p]”.
Other specimens examined. TA IWAN . Taitung: 1♂, Lanyu Is., 22–26.III.1998, leg. C.-F. Lee (TARI); 3♂,
same locality, 29.III.2009, leg. U. Ong (RBCN); 1, 1♀, same locality, 26.IV.2009, leg. U. Ong (BMNH); 1♂,
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same locality, 17.III.2012, leg. T.-H. Lee (MNHUB); 2♂, same locality, 17.III.2012, leg. M.-H. Tsou (MNHUB);
2♂, same locality, 5.V.2012, leg. S.-F. Yu (TARI); 2♂, same locality, 14.IV.2013, leg. B.-X. Guo (TARI); 2♂,
14.IV.2013, leg. B.-X. Guo (TARI); 1♂, 22.XI.2013, leg. Y.-T. Chung (TARI); 1♂, 2♀, Hsiaolanyu Is., 26.V.2009,
leg. Y.-T. Chung (TARI); 2♂, 1♀, same locality, 25.VI.2009, leg. Y.-T. Chung (TARI); 1♀, same locality,
27.VIII.2009, leg. Y.-T. Chung (TARI).
FIGURES 62–72. Diagnostic characters of Aulacophora abdominalis (Fabricius). 62. Antenna, male; 63. Antenna, female; 64.
Penis, dorsal view; 65. Penis lateral view; 66. Penis, lectotype of Crioceris abdominalis, details omitted; 67. Abdominal tergite
VIII, male; 68. Abdominal tergite VIII, lectotype of Crioceris abdominalis; 69. Apex of abdominal ventrite V, female; 70.
Gonocoxae; 71. Abdominal ventrite VIII, female; 72. Spermatheca.
Diagnosis. Aulacophora kotoensis is similar to A. indica with the enlarged first antennomere in male and cone-
like pygidium projecting from elytral apices in female, but A. kotoensis has a pair of tubercles on the pronotum in
male (without tubercles in A. indica), slender and black pygidium in female (wide and yellowish brown pygidium
in A. indica).
Males. Length 6.5–7.1 mm, width 3.5–3.8 mm. General color (Figs 4–5) yellowish brown but metathoracic
and abdominal ventrites, middle and hind legs black; outer margins of femur and tibia, and tarsi of front legs black;
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labrum dark brown; antenna dark brown except three basal antennomeres yellowish brown. Antenna (Fig. 62)
filiform and slender, antennomere I enlarged; ratio of length of antennomeres III to XI about 1.0 : 1.1 : 1.1 : 1.1 :
1.1 : 1.0 : 1.0 : 1.0 : 1.2; ratio of length to width from antennomere III to XI about 3.9 : 3.4 : 3.5 : 3.5 : 3.4 : 3.2 : 3.6
: 3.6 : 4.7. Pronotum with deep, transverse groove; one pair of tubercles behind groove. Elytra with cluster of erect
hairs behind humerus. Abdominal tergite VIII well sclerotized (Fig. 67), apical margin with wide triangular
incision and apices acute. Median lobe of abdominal ventrite V rectangular, with median, longitudinal, wide
groove and abbreviated near apex. Penis (Figs 64–65) slender, parallel-sided, abruptly narrowed at apical 1/7 and
apically tapering, apex recurved; slightly curved from middle to apex in lateral view; tectum laterally sclerotized;
endophallus with clusters of short setae, and with one longitudinal sclerite, apically tapering, apex strongly curved
in lateral view, basally widened.
Females. Length 7.2 mm, width 3.9 mm. Similar to male (Figs 6–7), but pronotum with transverse groove
shallow, and without tubercles; cluster of erect hairs on elytra absent; antennomere I not enlarged (Fig. 63); ratio of
length of antennomeres III to XI about 1.0 : 1.1 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.2; ratio of length to width from
antennomere III to XI about 3.0 : 3.2 : 3.2 : 3.3 : 3.3 : 3.2 : 3.2 : 3.2 : 4.4. Pygidium projecting from elytral apex,
parallel-sided, slender, and apex rounded. Middle of apical margin of abdominal ventrite V (Fig. 69) emarginate,
weakly convex at middle of emargination. Gonocoxae (Fig. 70) slender, apex of each gonocoxa with seven or eight
setae from apical 1/6 to apex; gonocoxae connected at middle, base slender. Ventrite VIII (Fig. 71) weakly
sclerotized; apex narrow, apical margin emarginate at middle, with dense short setae along apex; spiculum short.
Spermathecal receptaculum (Fig. 72) a little swollen; pump strongly curved; spermathecal duct short, stout,
shallowly projecting into receptaculum.
Host plant. Cucurbitaceae: Trichosanthes quinquangulata A. Gray (Chûjô 1962; present study).
Remarks. To exclude conspecifity between Aulacophora kotoensis and A. abdominalis (Fabricius, 1781) it
was necessary to study the type specimens of Crioceris abdominalis in the Fabricius-collection. The results of the
study of these specimens are treated extensively in the section on Aulacophora abdominalis and partly in the
section on A. indica. For the status of A. kotoensis it is relevant that we have found no arguments to treat this
species as conspecific with A. abdominalis.
Distribution. Lanyu and Hsiaolanyu islands (Taiwan). Hsiaolanyu is very small and deserted island which is
very close to Lanyu Island. The size of island is about 1.75 km2.
[Aulacophora abdominalis (Fabricius, 1781)]
(Figs 8–9, 66, 68)
Crioceris abdominalis Fabricius, 1781: 151 (Pacific islands); Fabricius, 1787: 87 (redescription).
Cryptocephalus abdominalis: Gmelin, 1790: 1719.
Galeruca abdominalis: Olivier, 1791: 590; Olivier, 1808: 623.
Galleruca abdominalis: Fabricius, 1792: 23; Fabricius, 1801: 483.
Rhaphidopalpa abdominalis: Dejean, 1837: 402; Weise, 1892: 395.
Aulacophora abdominalis: Baly, 1886: 14.
Type material. Crioceris abdominalis: lectotype ♂ (KIEL), here designated, labeled: “abdomina / lis [w, h] //
LECTO - / TYPE [w, h, circle label with blue border] // Lectotype / Crioceris ♂ / testacea Fab. [h] // N. A. Aslam
det. 19 [p] 71 [h, w]”. No paralectotypes designated (see under remarks).
Diagnosis. The lectotype of Crioceris abdominalis (Figs 8–9) is similar to Aulacophora. kotoensis but has
yellowish brown antenna and legs. Besides, the penis of the lectotype of C. abdominalis is distinctly asymmetric
near the apex and shows a lateral excavation at the middle (Fig. 66); abdominal tergite VIII has a semicircular
incision and lateral processes and basal margin convex with rounded corners (Fig. 68). A. kotoensis has antennae
(except basal segments) dark brown and middle and hind legs black. In A. kotoensis the penis is parallel sided,
narrowed towards apex which is symmetrical; abdominal tergite VIII has a broad triangular incision and basal
margin concave with sharp corners.
Remarks. To exclude conspecifity between Aulacophora kotoensis and A. abdominalis (Fabricius, 1781) it
was necessary to study the type specimens of Crioceris abdominalis in the Fabricius-collection.
Most of Fabrician types in Kiel had been transferred to the Zoological Museum of Copenhagen (KIEL)
(Zimsen 1964). Zimsen (1964) described the Coleoptera Collection as “There are no locality labels in Fabricius’
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own collection. Also peculiar are the very small name labels which Fabricius used: usually only a small scrap of
paper on which was written the name of the species; never any mention of the genus”.
In 1787 Fabricius described Crioceris testacea from India Orient., at present named Aulacophora indica
(Gmelin). Later, Fabricius (1801) synonymized Crioceris testacea with Crioceris abdominalis Fabricius, 1781 a
species he described from the South Pacific. Most probably he placed his specimens of both C. abdominalis and C.
testacea in his collection together. Because of the uncertainty around the taxa C. abdominalis and C. testacea (see
for example Baly 1879: 445 and Maulik (1936: 175, 197, 198) it was not only to exclude conspecifity between
Aulacophora kotoensis and A. abdominalis (Fabricius, 1781) that made it necessary for us to study the type
specimens of Crioceris abdominalis in the Fabricius-collection. The examination also could clarify the status of C.
testacea.
In the KIEL collection there are five specimens besides the name label “abdominalis, 1791”: one male
specimen carrying a label in Fabricius handwriting “abdominalis”; the other four specimens without labels. One of
them could be identified as Haplosomoides serena (Boheman) and one as Aulacophora coffeae Hornstedt. The two
remaining specimens are a male of Aulacophora indica (Gmelin, 1790) and an Aulacophora female, that could not
be identified.
The labeled specimen was designated as lectotype of Crioceris testacea Fabricius by Aslam in 1941. However,
designation of the lectotype is not valid since it was never published. Anand and Cox (1986) redescribed A.
abdominalis and A. testacea based on a female specimen collected in the South Pacific during the second voyage of
captain Cook (Banks collection) and a male specimen collected in India (Assam) respectively. At least the
redescriptions of A. abdominalis is not reliable since the female has fewer diagnostic characters. A redescription
based on the presumed syntypes in the Fabricius collection could finally end the uncertainty.
Because the series of syntypes is composed of several species we excluded the specimens of Haplosomoides
serena, Aulacophora coffaea and the female specimen from our examination. The two remaining male specimens
are clearly different. One male specimen that has no tubercles on the pronotum is identical with Aulacophora
indica (Gmelin,1790), the replacement name for Crioceris testacea Fabricius, 1787. This specimen is designated
lectotype of Crioceris testacea Fabricius, 1787. The other male specimen with a pair of tubercles on the pronotum
that unfortunately was labeled by Aslam as the lectotype of Crioceris testacea, is designated here as the lectotype
of Crioceris abdominalis. This is the specimen bearing the Fabricius label “abdominalis”. Both specimens are
labeled according these designations and serve nomenclatural stability.
Barroga & Mohamedsaid (2002) and Barroga (2002a) list a single male specimen as the type of Crioceris
testacea Fabricius they studied from the Fabricius-collection (KIEL). They mention it is labeled as Crioceris
abdominalis F., but do not list any further labels. In other type specimens they list all label information. Therefore
it is not likely they studied the specimen we designated the lectotype of C. abdominalis, because in that case they
would have listed the label attached by Aslam as well. It is very likely that they have studied the specimen we have
designated the lectotype of C. testacea. Because Barroga & Mohamedsaid (2002) and Barroga (2002a) do not
mention the presence of tubercles on the males pronotum, a character they unlikely would have left unnoticed,
makes it plausible that they have studied the male specimen herein designated the lectotype of Crioceris testacea
Fabricius, 1787.
Aulacophora lewisii Baly, 1886
(Figs 73–76, 82–90)
Aulacophora lewisii Baly, 1886: 24 (China); Baly, 1888: 179 (India, Malaysia); Duvivier, 1892: 430 (India: Mandar); Chen &
Kung, 1959: 375 (China: Hong Kong); Gressitt & Kimoto, 1963: 489 (China: Sichuan, Hubei, Anhui, Fujian, Guandong,
Guanxi, Hainan; Vietnam); Kimoto, 1964: 305 (Japan: Yakushima, Ryukyus); Kimoto, 1977: 354 (Bhutan); Kimoto, 1989:
59 (Thailand, Cambodia, Laos, Vietnam); Mohamedsaid, 1994: 382 (Malaysia); Mohamedsaid, 2000: 349 (Malaysia);
Barroga & Mohamedsaid, 2002: 116 (Malaysia); Mohamedsaid & Constant, 2007: 166 (Thailand, Cambodia); Aston,
2009: 14 (China: Hong Kong).
Ceratia lewisii: Miwa, 1931: 189 (Taiwan).
Orthaulaca (Ceratia) cattigarensis Weise, 1892: 397 (China: Shanghai; Japan); Gressitt and Kimoto, 1963: 489 (as synonym of
lewisii).
Aulacophora (Ceratia) cattigarensis: Laboissière, 1929: 258 (Vietnam); Chûjô, 1935b: 205 (Japan: Ishigaki island); Chûjô,
1935c: 160 (Taiwan).
Aulacophora cattigarensis: Ogloblin, 1936: 156 (Vietnam); Chen & Kung, 1959: 375 (China: Jiangsu, Zhejiang, Fujian,
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Guandong, Guanxi, Sichuan, Yunnan); Chûjô, 1962: 99 (Taiwan).
Aulacophora intermedia Jacoby, 1892: 942; Jacoby, 1896: 126 (Indonesia: Mentawai islands); Maulik, 1936: 181 (India);
Gressitt & Kimoto, 1963: 488 (as synonym of lewisii).
Type material. Aulacophora lewisii: lectotype ♀ (BMNH), here designated, labeled: “SYN- / TYPE [w, p, round
label with blue border] // Type / H. T. [w, p, round label with red border] // Baly Coll. [w, p] // China [w, h] //
Aulacophora / Lewisii / Baly / China [w, h] // BMNH(E) / #1024852 [w, p] // holotype [w, p]”. Paralectotypes: 2♂
(BMNH), labeled: “SYN- / TYPE [w, p, round label with blue border] // Hong / Kong / Lewis [w, h] // Aulacophora
/ lewisii / Baly (♂) [w, p] // not type [w, p] // BMNH(E) / #1024852 [w, p]”. Althought two paralectotypes were
labeled as nontype specimens, they cannot be ruled out of type series since the collecting data fits the original
description.
Orthaulaca (Ceratia) cattigarensis: lectotype ♀ (MNHUB), here designated, labeled: “Shanghai [w, h] //
cattigarensis / Ws [w, h] // SYNTYPUS / Orthaulaca / cattigarensis Wiese, 1892 / labelled by MNHUB 2014 [r, p]”.
Paralectotypes: 1♀ (MNHUB), labeled: “Shanghai [w, h] // SYNTYPUS / Orthaulaca / cattigarensis Wiese, 1892 /
labelled by MNHUB 2014 [r, p]”; 1♂ (MNHUB), labeled: “Shanghai / Simom ? [w, h] // ♂ [w, h] // SYNTYPUS /
Orthaulaca / cattigarensis Wiese, 1892 / labelled by MNHUB 2014 [r, p]”; 1♀ (MNHUB), labeled: “Shanghai /
Simom ? [w, h] // SYNTYPUS / Orthaulaca / cattigarensis Wiese, 1892 / labelled by MNHUB 2014 [r, p]”; 1♂, 1♀
(MNHUB), labeled: “Japon [w, h] // SYNTYPUS / Orthaulaca / cattigarensis Wiese, 1892 / labelled by MNHUB
2014 [r, p]”; 1♀ (MNHUB), labeled: “SYNTYPUS / Orthaulaca / cattigarensis Wiese, 1892 / labelled by MNHUB
2014 [r, p]”.
Aulacophora intermedia: lecotype ♂ (BMNH), here designated, labeled: “Type / H. T. [w, p, round label with
red border] // Rangoon/ Birmania / Fea 188 [w, p]7[h] // Jacoby Coll. / 1909-28a. [w, p] // Aulacop. / intermedia /
Jac. [b, h] // SYN- / TYPE [w, p, round label with blue border]”. Paralectotypes: 2♂ (MCZC), labeled: “Carin
Chebà / 1000–1100 m / L. Fea V XII-88 [w, p] // Aulacophora / intermedia / sp. n. [b, h] // Type [p] / 17560 [r, h]”.
Other specimens examined. CHINA. Guandong: 1♂, Tsha-jiu-san, V.-VI.1912, leg. S. V. Mell (BPBM); 1♂,
same locality, VIII.1913, leg. S. V. Mell (BPBM); Hainan: 1♂, 1♀, Dome Mountain, 12.VII.1935, leg. J. L.
Gressitt (BPBM); Hong Kong: 1♀, Lantau, Tung Chung, 31.VII.2007, leg. P. Aston (PAHC); Sichuan: 1♂, Lung-
chue-pa to Chang-tau-ging, Wanhsien, 30.IX.1948, leg. Gressitt & Djou (BPBM); 1♀, Suifu, leg. D. C. Graham
(BPBM); INDONESIA. Java: 1♀, leg. Chev (BMNH); Sulawesi: 1♀, leg. Wallace (BMNH); LAOS.
Khammouane: 1♂. 2♀, Ban Khoun Ngeun env., 250 m, 20–29.V.2004, leg. E. Jendek and O. Šauša (JBCB);
MALAYSIA. 1♀, MAL CA (= Malacca), leg. Wallace (BMNH); Pahang: 1♂, Tasik Bera, Pos Iskander, 4–
8.V.1993, leg. Sham, Razali and Saiful (JBCB); Penang: 1♂, Peng, leg. Wallace, Baly Coll. (BMNH); 2♂, 2♀, 63
47, leg. Bowring (BMNH); 2♀, Wellesley prov., Kg. Siam, Pinang Tunggal, 7.VIII.1979, leg. J. Pertanian
(BMNH); Perak: 1♂, 1905-313, leg. Sharp (BMNH); Sarawak: 1♀, Bidi, 1980-9, leg. C. J. Brooks (BMNH); 3♀,
Kuching, 1910, leg. J. E. A. Lewis (BMNH); PHILIPPINES. 1♂, 42 22 (BMNH); 1♂, 67 56 (BMNH); 1♂, leg. J.
Hano, 67 56 (BMNH); 1♀, 985 (BMNH); TA I WAN. Ilan: 2♂, 1♀, Yingshih, 31.VII.2009, leg. H.-J. Chen (TARI);
Kaoshiung: 2♀, Maolin, 6.VII.2011, leg. M.-H. Tsou (TARI); 5♂, 1♀, Meinung, 1.III.2012, leg. J.-C. Chen
(TARI); 1♀, same locality, 17.VII.2012, leg. J.-C. Chen (TARI); Miaoli: 1♀, Kuantaoshan, 3.XI.2009, leg. S.-F. Yu
(TARI); Nantou: 1♂, 2♀, Puli, 14.IX.2012, leg. Y.-P. Huang (TARI); Pingtung: 5♂, 2♀, Chuanfanshih,
26.VIII.2010, leg. J.-C. Chen (TARI); 1♂, Kueishan, 3.VIII.2012, leg. J.-C. Chen (TARI); 1♀, Tahanshan,
15.XI.2013, leg. Y.-T. Chung (TARI); Taipei: 1♀, Chihshanyan, 14.VI.2008, leg. M.-H. Tsou (TARI); 1♂, 1♀,
Chuwei, 2006, leg. H.-T. Cheng (TARI); 1♂, 2♀, Kuanyinshan, 28.VI.2008, leg. M.-H. Tsou (TARI); 1♀, Peitou,
6.XI.2006, leg. S.-F. Yu (TARI); 2♂, Taishan, 24.IV.2009, leg. H.-T. Cheng (TARI); 5♀, Taluntou, 23.VI.2009, leg.
S.-F. Yu (TARI); Taitung: 8♂, 4♀, Lanyu island, 17.III.2012, leg. T.-H. Lee (TARI); 7♂, 5♀, 17.III.2012, leg. M.-
H. Tsou (TARI); 6♂, 5♀, Lutao island, 14.IV.2014, leg. Y.-T. Chung (MNHUB, RBCN, TARI); 1♂, same locality,
16.IV.2014, leg. B.-X. Guo (TARI); Taoyuan: 1♀, Fuhsing, 28.III.2010, leg. M.-H. Tsou (TARI); THAILAND.
Kanchanaburi: 1♂, Phatad valley, 14–15.IX.2009, leg. V. Hula (JBCB); Nan: 1♂, Ban Huay Kon env., 27.V.–
10.VI.2002, leg. P. Prŭdek and M. Obořil (JBCB).
Diagnosis. Aulacophora lewisii is similar to A. frontalis and A. palliata with the yellowish brown body and
black shining elytron but easily identified by the filiform antenna in male and the sinuate apical margin of the
abdominal ventrite V in female (the modified and triangular antennomeres III to V in male and the rounded apical
margin of the abdominal ventrite V in female of A. frontalis and A. palliata).
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FIGURES 7381. Habitus of Aulacophora species. 73. A. lewisii, male, dorsal view; 74. Ditto, ventral view; 75. A. lewisii,
female, dorsal view; 76. Ditto, ventral view; 77. A. nigripennis, male, dorsal view; 78. Ditto, ventral view; 79. A. nigripennis,
female, dorsal view; 80. A. opacipennis, male, dorsal view; 81. Ditto, ventral view.
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FIGURES 8290. Diagnostic characters of Aulacophora lewisii Baly. 82. Antenna, male; 83. Antenna, female; 84. Penis,
dorsal view; 85. Penis lateral view; 86. Gonocoxae; 87; Abdominal ventrite VIII, female; 88. Apex of abdominal ventrite V,
female; 89. Abdominal tergite VIII, male; 90; Spermatheca.
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Males. Length 5.6–6.6 mm, width 2.8–3.1 mm. General color (Figs 73–74) yellowish brown except elytron
black and shining. Antenna (Fig. 82) filiform, antennomere I not enlarged, III–V rather stout; ratio of length of
antennomeres III to XI about 1.0 : 1.0 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 1.1; ratio of length to width from
antennomere III to XI about 2.6 : 2.6 : 2.3 : 2.7 : 2.5 : 2.6 : 3.0 : 3.0 : 4.7. Apex of abdominal tergite VIII forming
one pair of wide processes (Fig. 89), apex of process rectangular; weakly emarginate at middle; base weakly
sclerotized. Median lobe of abdominal ventrite V rectangular, with longitudinal groove from base to apex, apically
widened. Penis (Figs 84–85) slender, apically tapering, apex pointed; almost straight in lateral view; tectum well
sclerotized, apex pointed, disc ventrally covered with two longitudinal rows of fine setae; endophallus with two
longitudinal rows of stout setae at base of tectum, and two longitudinal rows of tingy setae above stout setae, and
with one longitudinal sclerite, apex curved and rounded, basally widened.
Females. Length 6.5–7.2 mm, width 3.2–3.6 mm. Similar to male (Figs 75–76), but antennomeres III–V
slender (Fig. 83); ratio of length of antennomeres III to XI about 1.0 : 1.0 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 1.1; ratio
of length to width from antennomere III to XI about 3.2 : 3.1 : 2.7 : 2.8 : 3.0 : 3.1 : 3.1 : 3.2 : 4.4. Apical margin of
abdominal ventrite V (Fig. 88) sinuate. Gonocoxae (Fig. 86) slender, apex of each gonocoxa with seven or eight
setae from apical 1/6 to apex; gonocoxae connected at middle, base widened. Ventrite VIII (Fig. 87) weakly
sclerotized; apex narrow, apical margin a little emarginate at middle, surface with dense long setae along apical
margin, spiculum short. Spermathecal receptaculum (Fig. 90) a little swollen, hardly separated from pump; pump
strongly curved; spermathecal duct short, stout, shallowly projecting into receptaculum.
Host plants. Cucurbitaceae: including cultivated species.
Distribution. Bhutan, Cambodia, China, India, Indonesia, Japan, Laos, Taiwan, Thailand, Vietnam.
Aulacophora nigripennis Motschulsky, 1857
(Figs 77–79, 91–99)
Aulacophora nigripennis Motschulsky, 1857: 38; Baly, 1886: 5 (China, Japan); Schönfeldt, 1890: 174 (Japan: Ryukyus);
Chûjô, 1935a: 83 (Taiwan, China: Heilongjian); Ogloblin, 1936: 156 (Russia); Chen & Kung, 1959: 375 (China: Hebei,
Shaanxi, Shandong, Jiangsu, Zhejiang, Fujian, Jiangxi, Sichuan); Chûjô, 1962: 91 (Taiwan); Gressitt & Kimoto, 1963: 489
(China: Hainan); Lee & An, 2001: 130 (South Korea); Aston, 2009: 14 (China: Hong Kong).
Ceratia (Orthaulaca) nigripennis: Weise, 1922: 62.
Ceratia nigripennis: Miwa, 1931: 189 (Taiwan).
Type material. Holotype ♂ (ZMUM), labeled: “Aulacophora / nigripennis / Motsch / Japonia [y, h] // Holotypus
[r, p]”. This specimen was in bad condition. The head and thorax was lost. Only elytra, hind legs, and abdomen
were attached to the pin.
Other specimens examined. CHINA. Fujian: 1♀, Houchai, 27.VI.2014, leg. Y.-T. Chung (TARI); 1♀,
Shawu, Tachufung, 1–3.VI.1943, leg. T. C. Maa (BPBM); 1♀, same locality, 4–9.VI.1943, let. T. C. Maa (BPBM);
Hong Kong: 1♂, Lantau, Tong Chung, 31.V.2008, leg. P. Aston (PAHC); Hubei: 1♂, Lichuan, Hsiaoho,
9.VIII.1948, leg. Gressitt & Djou (BPBM); 1♂, same locality, 11.VIII.1948, leg. Gressitt & Djou (BPBM);
Sichuan: 1♂, Yachow to Mupin, 23–27.VI.1929, leg. D. C. Graham (BPBM); JAPAN. Aomori: 1♂, Fukaura, 11–
13.VI.1999, leg. M. Hayashi (JBCB); TA IWAN . Hualien: 1♀, Huitouwan, 10.VII.207, leg. C.-F. Lee (TARI); 1♀,
Juisui, 27.VI.2013, leg. Y.-T. Chung (TARI); Kaoshiung: 1♀, Maolin, 6.VII.2011, leg. M.-H. Tsou (TARI); 1♂,
1♀, Meinung, 14.XII.2012, leg. J.-C. Chen (TARI); 1♂, 28.XII.2012, leg. J.-C. Chen (TARI); 1♂, Takangshan,
9.IV.2013, leg. B.-X. Guo (TARI); Miaoli: 1♂, Tunglo, 25.III.2008, leg. H.-T. Shih (TARI); Pingtung: 1♂,
Lilungshan, 2.III.2012, leg. J.-C. Chen (TARI); 1♂, same locality, 24.III.2014, leg. Y.-T. Chung (TARI); 1♂, same
locality, 26.V.2014, leg. Y.-T. Chung (TARI); 1♂, 1♀, Mutan, 27.VII.2011, leg. J.-C. Chen (TARI); 1♀,
Nanjenshan, 27.III–5.IV.2010, leg. M.-L. Jeng (TARI); 1♂, Peitawushan, 19.II.2014, leg. Y.-T. Chung (TARI); 1♀,
Shouka, 25.II.2009, leg. C.-F. Lee (TARI); 1♀, same locality, 5.VI.2012, leg. Y.-T. Chung (TARI); 1♀, same
locality, 28.II.2013, leg. B.-X. Guo (TARI); 1♂, 1♀, Tahanshan, 26.III.2013, leg. C.-F. Lee (TARI); 1♀, same
locality, 25.V.2013, leg. Y.-T. Chung (TARI); 1♂, Taiwu, 27.III.2014, leg. Y.-T. Chung (TARI); 1♂, 2♀, Wutan,
5.VIII.2014, leg. Y.-T. Chung (RBCN); Tainan: 1♀, Meiling, 12.III.2011, leg. M.-L. Jeng (TARI); 1♂, same
locality, 7.VI.2013, leg. B.-X. Guo (TARI); 1♀, same locality, 22.VI.2013, leg. W.-C. Liao (TARI); 1♀, Pichien
trail, 22.III.2010, leg. U. Ong (TARI); Taipei: 1♀, Chinshan, 9.V.2008, leg. W.-T. Liu (TARI); 1♂,
Chungchengshan, 17.II.2012, leg. S.-F. Yu (TARI); 2♀, Chuwei, 2006, leg. H.-T. Cheng (TARI); 1♂, Erhkoshan,
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FIGURES 9199. Diagnostic characters of Aulacophora nigripennis Motschulsky. 91. Antenna, male; 92. Antenna, female;
93. Penis, dorsal view; 94. Penis lateral view; 95. Gonocoxae; 96. Apex of abdominal ventrite V, female; 97; Abdominal
ventrite VIII, male; 98. Abdominal tergite VIII, female; 99; Spermatheca.
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26.XI.2006, leg. M.-H. Tsou (TARI); 1♂, Fengkueitsui, 29.VI.2007, leg. S.-F. Yu (TARI); 1♀, Guandu,
27.IV.2009, leg. H.-T. Cheng (TARI); 1♀, Kungliao, 24.IX.2009, leg. H. Lee (TARI); 2♂, 1♀, Taipei City Zoo,
13.XI.2006, leg. H.-T. Cheng (TARI); 1♀, Tatungshan, 3.IV.2008, leg. S.-F. Yu (TARI); 2♂, Tienhsiyuan,
15.IV.2011, leg. H. Lee (TARI); 1♂, 1♀, Wulai, 21.X.2006, leg. S.-F. Yu (TARI); 2♀, same locality, 26.V.2009,
leg. C.-F. Lee (TARI); 1♀, same locality, 15.VIII.2010, leg. S.-P. Kao (TARI); Taitung: 1♀, Liyuan, 19.VI.2013,
leg. C.-F. Lee (TARI); 2♂, 2♀, Motien, 5.X.2010, leg. C.-F. Lee (MNHUB); Taoyuan: 1♀, Houtzuhu, 6.IX.2011,
leg. S.-F. Yu (TARI); 1♂, Hsiaowulai, 18.X.1007, leg. S.-F. Yu (TARI); 1♂, 1♀, same locality, 19.IV.2008, leg. S.-
F. Yu (RBCN); 1♀, same locality, 19.IV.2008, leg. M.-H. Tsou (TARI); 1♀, Kaoi, 19.VI.2010, leg. H.-J. Chen
(TARI); 1♀, Tungyanshan, 15.V.2010, leg. H. Lee (TARI).
Diagnosis. Aulacophora nigripennis is similar to A. frontalis, A. lewisii, A. opacipennis, and A. palliata with
the black elytron but easily recognized by its black meso- and metathoracic ventrites and legs.
Males. Length 6.0–6.8 mm, width 3.3–3.9 mm. General color (Figs 77–78) black and shining except head,
prothorax, scutellum, and abdomen yellowish brown, antenna dark brown but three basal antennomeres paled.
Antenna (Fig. 91) filiform, antennomere I not enlarged; II apically enlarged ratio of length of antennomeres III to
XI about 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 0.9 : 1.1; ratio of length to width from antennomere III to XI about 2.6
: 2.9 : 2.9 : 3.1 : 3.2 : 3.3 : 3.6 : 3.3 : 4.1. Apex of abdominal tergite VIII forming shallow notch (Fig. 97), weakly
convex at middle; with several setae scattered on apical margin; base weakly sclerotized. Median lobe of
abdominal ventrite V long and rectangular, with longitudinal ridge along lateral margin. Penis (Figs 93–94) wide,
widest at apical 1/3, abruptly narrowed near apex, apex narrowly rounded; almost straight in lateral view, apex
curved and apical margin truncate; tectum absent; endophallus with scattered stout setae, and with one longitudinal
sclerite, apex curved and rounded, basally widened.
Females. Length 6.2–7.1 mm, width 3.5–4.2 mm. Similar to male (Fig. 79), but antennomeres II not enlarged
(Fig. 92), III–V slender; ratio of length of antennomeres III to XI about 1.0 : 1.2 : 1.2 : 1.2 : 1.2 : 1.2 : 1.2 : 1.0 : 1.3;
ratio of length to width from antennomere III to XI about 3.1 : 3.8 : 4.2 : 4.0 : 4.0 : 4.1 : 4.3 : 3.6 : 5.1. Apical
margin of abdominal ventrite V (Fig. 96) emarginate, convex and acute at middle. Gonocoxae (Fig. 95) wide, apex
of each gonocoxa with seven or eight setae from apical 1/6 to apex; gonocoxae connected at middle, base slender.
Ventrite VIII (Fig. 98) weakly sclerotized; apex narrow, apical margin a little emarginate at middle, surface with
extremely dense short setae along apical margin, some short setae scatter near apex; spiculum short. Spermathecal
receptaculum (Fig. 99) a little swollen, hardly separated from pump; pump strongly curved; spermathecal duct
short, stout, shallowly projecting into receptaculum.
Host plants. Cucurbitaceae: Trichosanthes cucumeroides (Ser.) Maxim. ex Fr. & Sav., Lagenaria siceraria
(Mol.) Standl.; Fabaceae: Callerya reticulata (Benth.) Schot.
Distribution. China, Japan, South Korea, Taiwan.
Aulacophora opacipennis Chûjô, 1962
(Figs 80–81, 100–107)
Aulacophora opacipennis Chûjô, 1962: 98 (Taiwan).
Type material. Holotype ♂ (TARI), labeled: “Kuraru [= Kenting, in Pingtung] / 31-VII-1931 / Col. T. Shiraki //
Ho / Type [w, p, round label, red letters and border, but faded out] // Aulacophora / opacipennis / Chûjô [h] / DET.
M. CHUJO [w, p] // 2312 [w, p]”. Paratypes: 1♀ (TARI), labeled: “Kuraru / 31-VII-1931 / Col. T. Shiraki [w, p] //
ALLo / Type [w, p, round label, gray letters and border] // Aulacophora / opacipennis / Chûjô [h] / DET. M.
CHUJO [w, p] // 2312 [w, p]”; 3♂, 2♀ (TARI), labeled: “Kuraru / 31-VII-1931 / Col. T. Shiraki [w, p] // Para / Type
(w, p, round label, green letters and border) // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] //
1463-1467 [w, p]”; 1♂, 1♀ (TARI), labeled: “KUARU (sic!) [h] / FORMOSA [p] / 20.VI.1937 [h] / COL. M.
CHUJO [w, p] // Para / Type [w, p, round label, green letters and border] // Aulacophora / opacipennis / Chûjô [h] /
DET. M. CHUJO [w, p] // 1468 & 1470 [w, p]”; 1♀ (TARI), labeled: “Shiiago [= Maopu] Chikuto [= Chutung] /
SHINCHIKU [= Hsinchu] / –27–30.VI.1930 / Col. M. CHUJO [w, p] // Para / Type [w, p, round label, green letters
and border] // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] // 1469 [w, p]”; 1♂ (TARI), labeled:
“Shizyukei (= Suchungchi, in Pingtung) / 1-VIII-1931 / Col. T. Shiraki [w, p] // Para / Type (w, p, round label,
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FIGURES 100107. Diagnostic characters of Aulacophora opacipennis Chûjô. 100. Antenna, male; 101. Antenna, female;
102. Penis, dorsal view; 103. Penis lateral view; 104. Gonocoxae; 105. Abdominal ventrite VIII, female; 106. Abdominal
tergite VIII, male; 107. Spermatheca.
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green letters and border) // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] // 1462 [w, p]”; 1♀
(TARI), labeled: “HEITOU [= Pingtung City] / 22.V.1939 / Y.MIWA [w, p] // Para / Type (w, p, round label, green
letters and border) // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] // 1461 [w, p]”; 1♀ (TARI),
labeled: “Formosa / Shinchiku (= Hsinchu), -18. / VII 1-30./ J. Sonan, [w, p] // Para / Type (w, p, round label, green
letters and border) // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] // 1460 [w, p]”; 1♂ (TARI),
labeled: “7.VI.1914 / Taitô (= Taitung) [h] / Col. I. Nitobe [w, p] // Para / Type (w, p, round label, green letters and
border) // Aulacophora / opacipennis / Chûjô [h] / DET. M. CHUJO [w, p] // 1460 [w, p]”.
Other specimens examined. TA I WAN . Hsinchu: 2♂, 3♀, Wufeng, 14–16.VII.1982, leg. K. C. Chou & C. C.
Pan (TARI, BMNH); Ilan: 1♂, Mt. Taiheizan (= Taipingshan), 23.VII.1940, leg. M. Chujo (TARI); Pingtung: 1♀,
Sheting, 4.XI.2009, leg. M.-H. Tsou (RBCN); 1♂, Suchunghsi, 13.V.2013, leg. Y.-T. Chung (RBCN).
Diagnosis. Aulacophora opacipennis is similar to A. apicipes with yellowish brown body, and black and
opaque elytron but A. opacipennis is easily separated from A. apicipes by the yellowish brown legs and black
elytron (black tibia and tarsi, yellow apical margin of elytron in A. apicipes).
Males. Length 6.2 mm, width 3.1 mm. General color (Figs 80–81) yellowish brown except elytron black and
opaque. Antenna (Fig. 100) filiform, antennomere I not enlarged, apico-lateral angles of antennomeres III–IV
produced anteriorly; ratio of length of antennomeres III to XI about 1.0 : 1.2 : 1.1 : 1.1 : 1.1 : 1.1 : 1.1 : 1.1 : 1.5;
ratio of length to width from antennomere III to XI about 2.7 : 3.3 : 2.7 : 2.7 : 2.9 : 3.2 : 3.5 : 3.7 : 5.4. Abdominal
tergite forming notch (Fig. 106), weakly convex at middle; base weakly sclerotized. Median lobe of fifth
abdominal ventrite rectangular, with median longitudinal ridge represented from middle extending beyong base,
median longitudinal internal ridge well developed; disc depressed at apical area. Penis (Figs 102–103) wide,
abruptly narrowed from apical 1/6 to apex, apex pointed, less widened at basal 2/5; curved in lateral view; tectum
well sclerotized, apex pointed, abruptly widened at basal 1/3, lateral margin with triangular sclerite recurved at
basal 1/3; endophallus with several stout setae scattered, and with one longitudinal sclerite, apex recurved and
rounded, basally widened.
Females. Length 7.5–7.9 mm, width 3.9–4.3 mm. Similar to male, but antenna more slender (Fig. 101), and
apico-lateral angles of antennomeres III–IV less produced anterior; ratio of length of antennomeres III to XI about
1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 0.9 : 0.9 : 0.9 : 1.0; ratio of length to width from antennomere III to XI about 3.7 : 3.8 : 4.2
: 4.2 : 4.2 : 4.2 : 4.4 : 4.2 : 5.5. Apical margin of abdominal ventrite V truncate. Gonocoxae (Fig. 104) slender, apex
of each gonocoxa with eight setae from apical 1/6 to apex; gonocoxae connected at middle, base widened. Ventrite
VIII (Fig. 105) weakly sclerotized; apex narrow, apical margin a little emarginate at middle, surface with dense
short setae inside and along apical margin, and with extremely long setae near apical margin; spiculum short.
Spermathecal receptaculum (Fig. 107) a little swollen, hardly separated from pump; pump strongly curved;
spermathecal duct short, stout, and strongly curved, shallowly projecting into receptaculum.
Host plant. Unknown.
Distribution. Endemic to Taiwan.
Aulacophora palliata (Schaller, 1783)
(Figs 108–110, 112–120)
Chrysomela palliata Schaller, 1783: 279 (India: Malayalam).
Crioceris palliata: Fabricius, 1787: 87.
Cryptocephalus palliatus: Gmellin, 1890: 1718.
Galleruca palliata: Fabricius, 1792: 22.
Galeruca palliata: Olivier, 1808: 625.
Aulacophora palliata: Jacoby, 1884: 41; Baly, 1888: 180 (India, Malaysia); Jacoby, 1889: 206 (Burma); Duvivier, 1892: 430
(India: Konbir); Weise, 1892: 398 (Sunda islands); Maulik, 1936: 182 (redescription); Chen & Kung, 1959: 375 (China:
Yunnan, Hainan); Chûjô, 1962: 94 (Taiwan); Gressitt & Kimoto, 1963: 490 (China: Hainan, Guandong; Vietnam); Barroga
& Mohamedsaid, 2002: 138 (Indonesia: Buru island); Aston, 2009: 13 (China: Hong Kong).
Orthaulaca (Ceratia) palliata: Weise, 1892: 398.
Aulacophora (Ceratia) palliata: Laboissière, 1929: 258 (Vietnam); Chûjô, 1935c: 160 (Taiwan).
Ceratia palliata: Miwa, 1931: 189 (Taiwan).
Type material. Schaller’s collection does no longer exist. It was lost in “Waisenhaus Halle”.
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FIGURES 108112. Habitus of Aulacophora species. 108. A. palliata, male, dorsal view; 109. Ditto, ventral view; 110. A.
palliata, female, dorsal view; 111. Head of A. frontalis, male; 112. Head of A. palliata, male.
Other specimens examined. CHINA. Guandong: 1♂, V.–VII.1911, leg. S. V. Mell (BPBM); Hong Kong: 1♂,
1♀, Lantau, Pui O., 27.V.2008, leg. P. Aston (PAHC); INDONESIA. Java: 2♂, 2♀, leg. Harsfield, 60-15 (BMNH);
2♂, 2♀, Noesa Kembangan, leg. Drescher (MNHUB); TA I WAN. Miaoli: 1♂, Sanyi, 6.VII.2013, Y.-T. Chung
(TARI); Kaoshiung: 1♂, Meinung, 14.XII.2012, leg. J.-C. Chen (TARI); 1♂, Tengchih, 4.VIII.2012, leg. J.-C.
Chen (TARI); Pingtung: 1♂, Lilungshan, 15.VIII.2009, leg. S.-F. Yu (TARI); 1♂, same locality, 23.XII.2009, leg.
J.-C. Chen (TARI); 1♂, Nanjenhu, 15.III.2010, leg. M.-H. Tsou (TARI): 1♂, Peitawushan, 18.VI.2012, leg. J.-C.
Chen (TARI); 1♂, 13.VIII.2013, leg. Y.-T. Chung (TARI); 1♂, same locality, 28.V.2014, leg. Y.-T. Chung (TARI);
1♂, Shouka, 13.VI.2013, Y.-T. Chung (TARI); 1♂, Shuangliu, 19.VII.2007, leg. S.-F. Yu (TARI); 4♂, 1♀,
Tahanshan, 19.XI.2012, leg. J.-C. Chen (TARI); 5♂, 1♀, same locality, 15.XI.2013, leg. Y.-T. Chung (TARI); 5♂,
4♀, Wutan, 5.VIII.2014, leg. Y.-T. Chung (BMNH, RBCN); Tainan: 1♂, Tanei, 25.VI.2007, leg. Y.-C. Chang
(TARI); 1♂, Tungshan, 14.III.2010, leg. M.-H. Tsou (TARI); Taipei: 1♂, Chungchengshan, 16.IX.2011, leg. S.-F.
Yu (TARI); 1♀, same locality, 9.X.2011, leg. M.-H. Tsou (TARI); Taitung: 5♂, 2♀, Yanping logging trail,
19.X.2012, leg. J.-C. Chen (MNHUB).
Diagnosis. See diagnosis of Aulacophora frontalis.
Males. Length 6.0–6.6 mm, width 3.1–3.5 mm. General color (Figs 108–108) yellowish brown except elytron
black and shining. Vertical area of head with a strongly thickened transverse ridge on each side (Fig. 112). Antenna
(Fig. 113) filiform, antennomere I a little swollen; III triangular, with semicircle groove at outer baso-lateral angles;
IV and V extremely widened; V with longitudinal groove on lateral expansion, groove without setae; ratio of length
of antennomeres III to XI about 1.0 : 0.8 : 0.9 : 0.8 : 0.9 : 0.8 : 0.9 : 0.8 : 1.0; ratio of length to width from
antennomere III to XI about 1.6 : 0.9 : 1.2 : 3.1 : 3.5 : 3.3 : 3.4 : 3.8 : 4.8. Apex of abdominal tergite VIII weakly
emarginate at middle (Fig. 119), apical margin with several setae; base weakly sclerotized. Median lobe of fifth
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FIGURES 113120. Diagnostic characters of Aulacophora palliata (Schaller). 113. Antenna, male; 114. Antenna, female; 115.
Penis, dorsal view; 116. Penis lateral view; 117. Gonocoxae; 118. Abdominal ventrite VIII, female; 119. Abdominal tergite
VIII, male; 120. Spermatheca.
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abdominal ventrite short and rectangular, disc flat and without longitudinal grooves, apical margin weakly
emarginate at middle. Penis (Figs 115–116) abruptly widened at apical 1/5, apex pointed; almost straight in lateral
view but curved at apical 1/5; ventral surface depressed at middle of apical 1/5; tectum well sclerotized, apex
widened; endophallus with dense, irregularly arranged setae, and with one longitudinal sclerite, apex rounded,
basally widened and bifurcate.
Females. Length 6.8–7.7 mm, width 3.2–4.4 mm. Similar to male (Fig. 110), but transverse ridge on vertical
area absent; antenna filiform and not modified (Fig. 114), ratio of length of antennomeres III to XI about 1.0 : 0.9 :
0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 0.9 : 1.1; ratio of length to width from antennomere III to XI about 3.2 : 3.2 : 3.3 : 3.0 : 3.7
: 3.6 : 3.7 : 3.5 : 4.5. Apical margin of abdominal ventrite V not modified, rounded. Gonocoxae (Fig. 117) slender,
apex of each gonocoxa with eight or nine setae from apical 1/6 to apex; gonocoxae connected at middle, base
widened. Ventrite VIII (Fig. 118) weakly sclerotized; apex wide, apical margin a little emarginate at middle,
surface with dense short setae along apical margin, and several long setae scattered at apical area; spiculum short.
Spermathecal receptaculum (Fig. 120) swollen, hardly separated from pump; pump strongly curved; spermathecal
duct short, stout, shallowly projecting into receptaculum.
Host plants. Cucurbitaceae: Trichosanthes cucumeroides (Ser.) Maxim. ex Fr. & Sav., and T. o vigera Bl.
(present study).
Distribution. Burma, China, India, Indonesia, Malaysia, Taiwan, Vietnam.
Aulacophora tibialis Chapuis, 1876
(Figs 121–126, 129–136)
Galleruca bicolor Weber, 1801: 56 (part).
Aulacophora tibialis Chapuis, 1876: 99 (Philippines: Mindoro, Luzon); Barroga and Mohamedsaid, 2002: 147 (as synonym of
ritsemae); Barroga, 2002b: 185 (corrected as valid name).
Aulacophora ritsemae Duvivier, 1884: 121 (Borneo; Indonesia: Java, Sumatra); Barroga & Mohamedsaid, 2002: 147
(Malaysia).
Aulacophora semifusca Jacoby, 1892: 942 (Burma); Maulik, 1936: 192 (redescription); Kimoto, 1989: 62 (Thailand, Laos,
Vietnam); Mohamedsaid, 1994: 385 (Malaysia); Mohamedsaid, 2000: 349 (Malaysia); Barroga & Mohamedsaid, 2002:
147 (as synonym of ritsemae); Mohamedsaid & Constan 2007: 166 (Thailand); Barroga & Mohamedsaid, 2002: 147 (as
synony m of ritsemae).
Aulacophora terminata Jacoby, 1899: 284 (Indonesia: Sumatra); Barroga & Mohamedsaid, 2002: 147 (as synonym of
ritsemae).
Aulacophora dohrni Jacoby, 1899: 285 (Indonesia: Sumatra); Barroga & Mohamedsaid, 2002: 147 (as synonym of ritsemae).
Aulacophora almora Maulik, 1936: 170 (India); Kimoto, 1989: 62 (as synonym of semifusca).
Aulacophora similis: Chen & Kung, 1959: 376 (misidentification; China: Zhejiang, Fujian, Sichuan, Yunnan); Gressitt &
Kimoto, 1963: 484 (as synonym of almora; China: Guandong, Hainan).
Aulacophora simplex Chûjô, 1962: 78 (Taiwan); Kimoto, 1966: 28 (as synonym of almora).
Type material. Galleruca bicolor: one paralectotype ♂ (KIEL), here designated, labeled: “Weber / bicolor. / 1723
[w, h] // bico / lor [w, h] // Aulacophora ritsemae Duviv. / det. by G.F. Barroga, 1999. [w, p]”. This specimen is
similar to the lectotype of A. bicolor in dorsal view with yellow apex on the black elytron. However, its yellow
meso- and metathoracic and abdominal ventrites differs from A. bicolor with black meso- and metathoracic and
abdominal ventrites. It is no doubt attributed to specimens of A. ritsemae collected from Java.
Aulacophora ritsemae: holotype ♀ (RMNH), labeled: “Dr. B. Hagen. / Tandjong Morawa. / Serdang / (N. O.
Sumatra). [w, p] // Aulacophora / ritsemae, Duv. / type [w, h] // ♀ [w, p] // Type [r, p]”.
Aulacophora semifusca: lectotype ♀ (BMNH), here designated, labeled: “Type / H. T. [w, p, round label with
red border] // Carin Chebà / 900–1100 m / L. Fea V VII-88 [w, p] // Jacoby Coll. / 1909-28a [w, p] // Aulacoph. /
semifusca / Jac. [b, h] // SYN- / TYPE [w, p, round label with blue border]”. Paralectotypes: 1♂ (MCSN), labeled:
“Carin Chebà / 900–1100 m / L. Fea V VII-88 [w, p] // semifusca / Jac. [w, h] // Aulacophora / semifusca / Jac. [b,
h] // SYNTYPUS [p] / Aulacophora / semifusca / Jacoby, 1892 [r, h] // Museo Civico / di Genova [w, p]”; 1♀
(MCSN): “Carin Chebà / 900–1100 m / L. Fea V VII-88 [w, p] // SYNTYPUS [p] / Aulacophora / semifusca /
Jacoby, 1892 [r, h] // Museo Civico / di Genova [w, p]”.
Aulacophora terminata: lectotype ♂ (BMNH), designated by Barroga and Mohamedsaid (2002), labeled:
“Dohrn / Sumatra / Soekaranda [w, p] // H. T. [w, p, round label with red border] // Jacoby Coll. / 1909-28a [w, p] /
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/ Aulacophora / terminata / Jac. [b, h] // Syntype. / A. terminata / Jac [w, h] // LECTOTYPE [p] / Aulacophora /
terminata Jacoby [h] / des. G. Barroga, 2000 [w, p]”. Paralectotype: 1♀ (BMNH), labeled: “Dohrn / Sumatra/
Soekaranda [w, p] // Jacoby Coll. / 1909-28a [w, p] // Syntype. / A. terminata / Jac [w, h] // PARALECTOTYPE /
Aulacophora / terminata Jacoby [h] / des. G. Barroga, 2000 [w, p]”.
Aulacophora dohrni: lectotype ♂ (BMNH), here designated, labeled: “Dohrn / Sumatra/ Soekaranda [w, p] //
Jacoby Coll. / 1909-28a [w, p] // Aulacophora [p] / dohrni Jac. [h] / det. G. Barroga, 2001 [w, p] // SYN- / TYPE [w,
p, round label with blue border]”. Paralectotypes: 1 (BMNH, abdomen lost, sex undetermined), labeled: “Dohrn /
Sumatra/ Soekaranda [w, p] // Type / H. T. [w, p, round label with red border] // Jacoby Coll. / 1909-28a [w, p] //
Aulacophora. / Dohrni Jac. [b, h] // Syntype / A. dohrni / Jacoby [w, h] // SYN- / TYPE [w, p, round label with blue
border]”.
Aulacophora tibialis: Chapuis’s types were originally deposited at the IRSB. They were studied and borrowed
by Barroga and Mohamdsaid (2002), and lost during return of the specimens (Limbourg personal communication).
Aulacophora almora: lectotype ♀ (BMNH), here designated, labeled: “W. Almora / Kumaon / India. H.G.C.
[w, h] // Type [w, p, round label with red border] // Aulacophora / almora Maulik [h] / des. S. Maulik 1933. [w, p] /
/ Brit. Mus. / 1939-[p] 149 [w, h] // SYN- / TYPE [w, p, round label with blue border]”. Paralectotypes: 1♀
(BMNH), labeled: “Ranikhet, / Kumaon, / India. H.G.C. [w, p] // Aulacophora / almora Mlk [h] / S. Maulik det. [p]
/ paratype 1936. [w, h] // SYN- / TYPE [w, p, round label with blue border]”; 1♂ (BMNH), labeled: “Haldwani
Dist. / Kumaon, / India. H.G.C. [w, p] // Aulacophora / almora Mlk [h] / S. Maulik det. [p] / paratype 1936. [w, h] /
/ SYN- / TYPE [w, p, round label with blue border]”; 1♂ (BMNH), labeled: “Sarju Valley, / Kumaon, 4000ft. /
India. H.G.C. / Aulacophora / almora Mlk [h] / S. Maulik det. [p] / paratype 1936. [w, h] // SYN- / TYPE [w, p,
round label with blue border]”.
Aulacophora simplex: holotype ♂ (TARI), labeled: “URAI [h] [= Wulai, in Taipei] / FORMOSA [p] /
20.IV.1930 [h] / COL. M. CHUJO [w, p] // Ho / Type [w, p, round label, red letters and border, but faded out] //
Aulacophora / simplex / C
HÛJÔ
[h] / DET. M. CHUJO [w, p] // 2371 [w, p]”. Paratypes: 1♀ (TARI), labeled:
“URAI [h] [= Wulai, in Taipei] / FORMOSA [p] / 18.IX.1932 [h] / COL. M. CHUJO [w, p] // ALLo / Type [w, p,
round label, gray letters and border] // Aulacophora / simplex / C
HÛJÔ
[h] / DET. M. CHUJO [w, p] // 2370 [w, p]”;
1♀ (TARI), labeled: “Formosa / Miwa [w, p] // Fushan [written in Japan] / 22.7.1929 [w, h] [on the back] // Para /
Type [w, p, round label, green letters and border] // Aulacophora / simplex / C
HÛJÔ
[h] / DET. M. CHUJO [w, p] //
1428 [w, p]”; 1♀ (TARI), labeled: “Formosa / Karenko [= Hualien], -19. / VII 20–VIII 4. / T. Okuni, [w, p] // Para /
Type [w, p, round label, green letters and border] // Aulacophora / simplex / C
HÛJÔ
[h] / DET. M. CHUJO [w, p] //
1427 [w, p]”.
Other specimens examined. CHINA. Fujian: 1♀, Chishui, 20.VI.2014, leg. Y.-T. Chung (TARI); 3♀,
Houchai, 25–27.VI.2014, Y.-T. Chung (TARI); 1♀, Shangzhai, 1.VII.2014, leg. Y.-T. Chung (TARI); 1♀, Shaowu,
Tachulan, 10.VI.1942, leg. T. Maa (BPBM); 1♂, same locality, 18.VI.1942, leg. T. Maa (BPBM); 1♂, same
locality, 9.XI.1942, leg. T. Maa (BPBM); INDIA. Meghalaya: 4♂, 2♀, 3 km E of Tura, 15–22.IV.1999, leg. J.
Rolčik (JBCB); INDONESIA. Sumatra: 2♂, 3♀, Lampung prov., Bukit Barisan Selatan N. P., 5 km SW of Liwa,
7–17.II.2000, leg. J. Bezdĕ, (JBCB); 11♂, 1♀, Noesa Kembangan, leg. Drescher (MNHUB); Java: 2♂, 1♀, 1909-
28a, leg. Jacoby (BMNH); 1♂, 2♀, leg. Haus. (MNHUB); 1♀, leg. Horsfield (BMNH); 1♂, Malang (MNHUB);
LAOS. 1♂, Ban Van Eue, SE of Phou Kow Kuei, 13.IV.1965, leg. J. L. Gressitt (BPBM); 2♀, same locality, 1–
15.V.1965, leg. J. A. Rondon (BPBM), 1♀, same locality, 15–31.V.1965, leg. native collector (BPBM); 1♂,
Phongsaly prov., 10 km SE of Boun Tai, 16–25.V.2004, leg. native collector (JBCB); 1♂, Sayaboury, 15.IV.1965,
leg. J. L. Gressitt (BPBM); 1♂, Uekinak, nr. PakKading, 22.IV.1965, leg. J. L. Gressitt (BPBM); MALAYSIA.
Pahang: 1♂, 1902-220, (Craddock), leg. Bingham (BMNH); TAIWAN . Taipei: 3♂, 2♀, Wulai, 7.VIII.2009, leg.
M.-H. Tsou (RBCN, TARI); 1♂, same locality, 6.IX.2014, leg. M.-H. Tsou (TARI); Taoyuan: 1♂, Fuhsing,
15.VII.2009, leg. H.-J. Chen (TARI); THAILAND. Chiangmai: 1♂, Doi Suthep, 28–31.III.1958, leg. T. C. Maa
(BPBM); Mae Hong Son: 1♂, San Huai Po, 9–16.V.1991, leg. L. Dembický (JBCB); VIETNAM. 1♀,
BanMeThaot, 16–18.V.1960, L. W. Quate (BPBM); 1♀, Blao (Balao), 14–21.X.1960, leg. C. M. Yoshimoto
(BPBM).
Diagnosis. Specimens of Aulacophora tibialis with the yellow elytra are similar to A. abdominalis and A.
indica but without sexually secondary characters and scutellum blackish brown. Those with the black base of the
elytron are similar to A. bicolor but lack the pair of small yellow spots at base.
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FIGURES 121128. Habitus of Aulacophora species. 121. A. tibialis, Taiwan, male, dorsal view; 122. Ditto, ventral view;
123. A. tibialis, Vietnam, female, dorsal view; 124. A. tibialis, Java, female, dorsal view; 125. Lectotype of A. dohrni, dorsal
view; 126. Paralectotype of A. dohrni, dorsal view; 127. A. apicipes, male, dorsal view; 128. Ditto, venral view.
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FIGURES 129136. Diagnostic characters of Aulacophora tibialis Duvivier. 129. Antenna, male; 130. Antenna, female; 131.
Penis, dorsal view; 132. Penis lateral view; 133. Gonocoxae; 134. Abdominal ventrite VIII, female; 135. Abdominal tergite
VIII, male; 136. Spermatheca.
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Males. Length 6.3–7.0 mm, width 3.3–3.6 mm. General color (Figs 121–122) yellowish brown but meso- and
metathoracic and abdominal ventrites black or blackish brown, middle and hind legs blackish brown but base of
tibia yellowish brown; outer margins of tibia, and tarsi of front legs dark brown; scutellum and labrum black;
antenna dark brown except two basal antennomeres yellowish brown. Antenna (Fig. 129) filiform and slender,
antennomere I not enlarged; ratio of length of antennomeres III to XI about 1.0 : 1.0 : 1.1 : 1.1 : 1.1 : 1.1 : 1.1 : 1.0
: 1.3; ratio of length to width from antennomere III to XI about 2.8 : 2.9 : 3.1 : 3.2 : 3.3 : 3.4 : 3.7 : 3.9 : 5.5. Apex
of abdominal tergite VIII truncate (Fig. 135), with several short setae along apical margin; base weakly sclerotized.
Median lobe of fifth abdominal ventrite small and rectangular, disc flat. Penis (Figs 131–132) slender, parallel-
sided, abruptly narrowed at apical 1/7 and apically tapering, apex pointed; moderately curved from middle to apex
in lateral view; tectum well sclerotized, apex weakly emarginate, sides parallel from apex to apical 1/4, with
angular process at apical 1/4; endophallus with dense setae on endophallic sac, and with one longitudinal sclerite,
apically tapering, curved at apical 1/3 in lateral view, basally widened.
Females. Length 6.5–6.8 mm, width 3.5–3.7 mm. Similar to male, but antennomeres III-V more slender (Fig.
130); ratio of length of antennomeres III to XI about 1.0 : 1.1 : 1.1 : 1.0 : 1.0 : 1.0 : 1.0 : 0.9 : 1.3; ratio of length to
width from antennomere III to XI about 3.2 : 3.6 : 3.8 : 3.4 : 3.4 : 3.7 : 3.5 : 3.6 : 5.5. Apical margin of abdominal
ventrite V rounded. Gonocoxae (Fig. 133) slender, apex of each gonocoxa with eight setae from apical 1/6 to apex;
gonocoxae connected at middle, base slender. Ventrite VIII (Fig. 134) weakly sclerotized; apex narrow, apical
margin widely rounded, with dense short setae along apical margin; spiculum short. Spermathecal receptaculum
(Fig. 136) a little swollen; pump strongly curved; spermathecal duct short, stout, shallowly projecting into
receptaculum.
Vari a t i o n. Specimens with yellow elytra occur in Taiwan, China, and Indochina, of which all but Fujian
(China) and Taiwan have yellow legs. Few specimens of Indochina have blackish brown elytra with narrow yellow
apical margin (Fig. 123). Most of specimens collected from Indonesia have black elytra with yellow apical margin
and extending into middle or basal 1/3 (Fig. 124), and thoracic and abdominal ventrite yellowish brown. Few
specimens have yellow elytra with basal 1/3 (Fig. 125) or half black (Fig. 126), thoracic and abdominal ventrites
blackish bown.
Host plants. Flowers of Lablab purpureus (L.) Sweet (Fabaceae) and Cucurbita moschata (Duch.) Poir.
(Cucurbitaceae) (present study).
Distribution. Myanmar, China, India, Indonesia, Laos, Malaysia, Philippines, Taiwan, Thailand, Vietnam.
Species excluded from Taiwan fauna
Aulacophora apicipes Jacoby, 1896
(Figs 127–128, 137–144)
Aulacophora apicipes Jacoby, 1896: 138 (Indonesia: Mentawei islands); Barroga & Mohamedsaid, 2002: 46 (Malaysia,
Indonesia: Sumatra, Java).
Aulacophora tenuicincta Jacoby, 1897: 406 (Indonesia: Batu islands); Barroga & Mohamedsaid, 2002: 46 (as synonym of
apicipes).
Aulacophora nigripalpis Chen & Kung, 1959: 378 (China: Yunnan). New synonym
Aulacophora opacipennis: Kimoto, 1989: 61 (Thailand, Laos). Misidentification
Type material. Aulacophora apicipes: lectotype ♂ (MCSN), here designated, labeled: “Mentawei / Si Oban IV
VIII / Modigliani 94 [w, p] // SYNTYPUS [p] / Aulacophora / apicipes / Jacoby, 1896 [r, h] // Museo Civico / di
Genova [w, p]”. Paralectotypes: 2♀ (MCSN), labeled: “Mentawei / Si Oban IV–VIII / Modigliani 94 [w, p] //
SYNTYPUS [p] / Aulacophora / apicipes / Jacoby, 1896 [r, h] // Museo Civico / di Genova [w, p]”; ♂ (BMNH),
labeled: “Type / H. T. [w, p, round label with red border] // Mentawei / Si Oban IV–VIII / Modigliani 94 [w, p] //
Jacoby Coll. / 1909-28a [w, p] // A. apicipes / Jac. [b, h] // SYN- / TYPE [w, p, round label with blue border]”; 1♀
(BMNH), labeled: “Mentawei / Si Oban IV-VIII / Modigliani 94 [w, p] // Museo Civ. / Genova [y, p] // Jacoby Coll.
/ 1909-28a [w, p] // SYN- / TYPE [w, p, round label with blue border]”; 1♂, 1♀ [still mating and glued on the same
card] (MCSN), labeled: “Mentawei / Si Oban IV–VIII / Modigliani 94 [w, p] // Typus [w, p, red letter, with red
border] // apicipes / Jac. [w, h] // Aulacophora / apicipes Jac [b, h] // SYNTYPUS [p] / Aulacophora / apicipes /
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Jacoby, 1896 [r, h] // Museo Civico / di Genova [w, p]”; 1♂ (MNHUB), labeled: “Mentawei / Si Oban IV–VIII /
Modigliani 94 [w, p] // apicipes / Jac. / Type. [w, h] // SYNTYPUS / Aulacophora / apicipes Jacoby, 1896 / labelled
by MNHUB 2014 [r, p]”; 1♂ (MNHUB), labeled: “Mentawei / Si Oban IV–VIII / Modigliani 94 [w, p] //
SYNTYPUS / Aulacophora / apicipes Jacoby, 1896 / labelled by MNHUB 2014 [r, p]”; 1♂ (MCZC), labeled:
“Mentawei / Si Oban IV–VIII / Modigliani 94 [w, p] // 209 [w, p] // Aul. / apicipes / n. sp. Jac. [w, h] // Type [p] /
17554 [r, h]”.
Aulacophora tenuicincta: lectotype ♂ (BMNH), here designated, labeled: “ISOLE BATU / 1896-97 / H. Raap
[w, p] // Jacoby Coll. / 1909-28a [w, p] // Syntype. / A. tenuicincta / Jac. [w, h] // ♂ [w, p] // MCZ [y, h] // SYN- /
TYPE [w, p, round label with blue border]”. Paralectotypes: 1♂ (BMNH), labeled: “Mentawei / Sipora / Sereinu
V–VI, 94 / Modigliani [w, p] // Jacoby Coll. / 1909-28a [w, p] // Syntype. / A. tenuicincta / Jac. [w, h] // SYN- /
TYPE [w, p, round label with blue border]”; 1♀ (BMNH), labeled: “Mentawei / Si Oban IV–VIII / Modigliani 94
[w, p] // Jacoby Coll. / 1909-28a [w, p] // Syntype. / A. tenuicincta / Jac. [w, h] // SYN- / TYPE [w, p, round label
with blue border]”; 1♂ (MCZC), labeled: “Is. Nias / Bawolovalani / Maggio 1886 / Modigliani [w, p] // 1st Jacoby
/ Coll. [w, p] // ♂ [w, p] // Type [p] / 17556 [w, h]”.
Aulacophora nigripalpis: holotype ♂ (IZAS), labeled: “ Ӂই : 㾯ৼ⡸㌽ ऀ㠈 [= Yunnan: Xishuangbanna
Mengla] / 620–650 ޜቪ (= 620–650 m) / ѝ഻、ᆨ䲒 [=Academia Sinica] [w, p] // 1959.v.31 [h] / 䟷䳶㘵 ᕥ Ⲭ 䋑 
(= collector Zhang Fa-Cai) [w, p] // HOLOTYPE [r, p] // Aulacophora / nigripalpis / [= Chen Kung] [h] / 䪁 
ᇊ㘵 : 䲣ц傔 [= determined by Sicien H. Chen] [w, p]”.
Other specimens examined. LAOS. Vientiane: 3♂, 2♀, Ban van Eue, 30.XI.1965, leg. native collector
(BPBM); 1♀, same locality, 15.I.1966, leg. native collector (BPBM); MALAYSIA. Penang, 1♂, leg. Pascoe
(BMNH); THAILAND. 1♂, Khao Yai, 10.IV.1963 (BPBM); 1♂, same locality, 12.II.1964 (BPBM); Trang: 1♂,
1♀, Khaophappha Khaochang, 10–11.I.1964, G. A. Samuelson (BPBM); 1♂, Loei prov., Phu Kradung N. P., 1000
m, 17.V.1999, leg M. Řiha (JBCB); VIETNAM. 1♂, Hoa-Binh, leg. A. Cooman (BPBM).
Diagnosis. See diagnosis of Aulacophora opacipennis.
Males. Length 6.3–7.0 mm, width 2.9–3.4 mm. General color (Figs 127–128) yellowish brown except elytron
black and opaque, tibia and tarsi darkened except base of tibia; apical margin of elytron yellowish brown. Antenna
(Fig. 137) filiform and extremely slender, antennomere I not enlarged; ratio of length of antennomeres III to XI
about 1.0 : 1.1 : 1.0 : 1.1 : 1.0 : 1.0 : 0.9 : 0.8 : 1.0; ratio of length to width from antennomere III to XI about 3.7 :
4.2 : 3.7 : 4.3 : 4.1 : 4.4 : 4.3 : 4.4 : 5.2. Apex of abdominal tergite VIII forming notch (Fig. 143), weakly convex at
middle, with several long setae along apical margin; base weakly sclerotized. Median lobe of fifth abdominal
ventrite rectangular, disc weakly depressed. Penis (Figs 139–140) slender, parallel-sided, abruptly narrowed from
apical 1/6 to apex, apex pointed; almost straight in lateral view, curved at apical 1/6; tectum well sclerotized,
extremely slender, apically widened, apex bifurcate; endophallus with clusters of dense stout setae, and with one
longitudinal sclerite, apex weakly recurved and rounded, basally widened.
Females. Length 6.7–7.5 mm, width 3.7–3.8 mm. Similar to male, but antenna shorter (Fig. 138), ratio of
length of antennomeres III to XI about 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.3; ratio of length to width from
antennomere III to XI about 3.5 : 3.8 : 3.7 : 3.5 : 3.3 : 3.5 : 3.6 : 3.8 : 5.6. Apical margin of abdominal ventrite V
truncate. Gonocoxae (Fig. 141) slender, apex of each gonocoxa with seven or eight setae from apical 1/6 to apex;
gonocoxae connected at middle, base widened. Ventrite VIII (Fig. 142) weakly sclerotized; apex narrow, apical
margin a little emarginate at middle, surface with dense short setae inside and along apical margin, and with
extremely long setae near apical margin; spiculum short. Spermathecal receptaculum (Fig. 144) a little swollen,
hardly separated from pump; pump strongly curved; spermathecal duct short, stout, shallowly projecting into
receptaculum.
Host plant. Unknown.
Remarks. Kimoto (1989) misidentified specimens of this species collected from Southeast Asia as
Aulacophora opacipennis.
Distribution. China, Indonesia, Laos, Malaysia, Thailand, Vietnam.
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REVISION OF AULACOPHORA FROM TAIWAN
FIGURES 137144. Diagnostic characters of Aulacophora apicipes Jacoby. 137. Antenna, male; 138. Antenna, female; 139.
Penis, dorsal view; 140. Penis lateral view; 141. Gonocoxae; 142. Abdominal ventrite VIII, female; 143. Abdominal tergite
VIII, male; 144. Spermatheca.
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Discussion
Although some species of Aulacophora display variable color patterns, the morphology of genitalia is constant and
in most cases largely different between the species. However, taxonomic problems still exist among species with
yellow elytra and pubescent humera in males, such as A. abdominalis, A. kotoensis and A. indica. The recent
descriptions of A. fauveli Beenen, 2008 and A. sulaksonoi Mohamedsaid, 2009 support the view that among species
with a wide distribution within the genus Aulacophora also species occur with very restricted ranges. Characters
hitherto not used might help to reveal more species. Secondary sexually characters, for example, the presence or
absence of paired tubercles on the males pronotum, as used in the present study, are suggested to be investigated in
other species too. Male specimens of A. foveicollis (Lucas, 1849) from Somalia showed paired tubercles, while
specimens for example from India do not, although the external morphology of the aedeagus is identical. Study of
the internal sac might reveal that more species are involved. Examination of large collection and applying other
characters seems necessary.
Acknowledgements
We would like to thank all curators listed above for giving us the opportunity to study the material from their
collections. We thank the Taiwan Chrysomelid Research Team for assistance in collecting material, including Jung-
Chang Chen, Hou-Jay Chen, Yi-Ting Chung, Bo-Xin Guo, Hsueh Lee, Ta-Hsiang Lee, Wen-Chuan Liao, Weiting
Liu, Mei-Hua Tsou, and Su-Fang Yu. We especially thank Ta-Hsiang Lee for photos of specimens, and Chih-Kai
Yang for identification of host plants. We acknowledge the help of Jan Bezděk (Brno) in providing some literature
and for discussion. This study was also supported by the Ministry of Science and Techonology MOST 103-2313-B-
055-001.
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Article
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Aulacophora indica has been intensively studied since it can cause serious damage to cucurbits. Here, we reported the complete mitochondrial genome of A. indica for the first time. The mitochondrial genome is 15,730 bp in length and contained 13 protein-coding genes (PCGs), 2 rRNAs, 22 tRNAs, and an AT-rich control region. Phylogenetic tree indicated that A. indica is more closely related to Aulacophora lewisii than other Galerucinae beetles.
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Based on a single well-preserved specimen from Eocene Baltic amber, Lilioceris groehni sp. n. is described and illustrated using phase-contrast X-ray microtomography. It is the first described species of Criocerinae (Coleoptera: Chrysomelidae) from Baltic amber. A check-list of fossil Criocerinae is provided. Placement of Crioceris pristiana (Germar, 1813) is discussed, this species is removed from Criocerinae and placed in Coleoptera incertae sedis.
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