Revision of the Eastern Asian Genera Ambrostoma Motschulsky and Parambrostoma Chen (Coleoptera: Chrysomelidae: Chrysomelinae)

Systematic Entomology (2012), 37, 332–345

Revision of the Eastern Asian genera Ambrostoma Motschulsky and Parambrostoma Chen (Coleoptera: Chrysomelidae: Chrysomelinae)

SI-QIN GE1, MAURO DACCORDI2, ROLF GEORG BEUTEL3, JING REN1, JUN-ZHI CUI1, WEN-ZHU LI1 and XING-KE YANG1

1Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China, 2Museo Civico di Storia Naturale, Verona, Italy and 3Institut fur¨ Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, FSU Jena, 07743 Jena, Germany

Abstract. The leaf beetle genera Ambrostoma Motschulsky, 1860 and Param- brostoma Chen, 1934 have been revised and now include 14 species. Two new species from Nepal are described, Parambrostoma kippenbergi sp.n. and P. medvedevi sp.n. Three new synonymies are established: Ambrostoma rugosopunctatum Chen = Ambrostoma (Parambrostoma) laosensis Kimoto & Gressitt, syn.n., Ambrostoma rugosopunctatum Chen = Ambrostoma daccordii Medvedev, syn.n., Ambrostoma fortunei (Baly) = Ambrostoma quadriimpressum chusanica Gruev, syn.n. One species was transferred from Chrysomela Linnaeus to Ambrostoma Motschulsky: A. superbum (Thunberg), comb.n. All the species now included are described and illustrated. Microcomputer tomography was applied for the ﬁrst time in a study on chrysomelid beetles. A cladistic analysis based on morphological characters of adults was conducted to reconstruct the intergeneric and interspeciﬁc phylogeny of Ambrostoma and Parambrostoma. The results show that the monophyly of both genera is well supported. Ambrostoma is widespread in East Asia, whereas Parambrostoma is restricted to the southern slope of the Himalayas, where a relatively recent and modest speciation took place.

Introduction from Taiwan, and A. daccordii Medvedev, 2007 from Viet- nam – were described in separate studies. The genus Ambrostoma (Fig. 1A) was erected by Motschul- Chen (1936) divided the genus into two subgenera, Ambro- sky in 1860. It included three species from the Amur stoma and Parambrostoma (Fig. 1B), based on setation region, eastern China and Nepal, respectively: A. quadri- patterns on the elytral epipleura, elytral punctures and the impressum, A. chinense (= A. fortunei Baly, 1860) and length of antennomeres II and IV. This taxonomic concept has A. nepalense (= A. mahesa Hope, 1831). Achard (1922) been accepted by other specialists (Gressitt & Kimoto, 1963; recorded A. fulgurans from China and Chen (1934) described Daccordi, 1977; Kimoto & Gressitt, 1981; Lobl¨ & Smetana, A. sublaeve from Korea. Chen (1936) revised the genus and 2010). Wang & Chen (1981) elevated these two subgenera recorded three other new species: A. fasciatum from Tien- to generic rank. Until now, ﬁve species of Parambro- Mo-Shan (China), A. ambiguum from Korea, and A. rugosop- stoma have been recorded: Parambrostoma ambiguum Chen, unctatum from China. Gressitt & Kimoto (1963) described P. mahesa (Hope), P. sublaeve (Chen), P. shuteae (Daccordi), A. omeishanum from Emeishan (Sichuan, China). Later, three P. montanum (Medvedev). new species – A. leigongshanum Wang, 1992 from Leigong- The purpose of this contribution is to provide accurate shan (Guizhou, China), A. chinkinyui Kimoto & Osawa, 1995 descriptions of species and to analyse the phylogenetic relationships of species assigned to Ambrostoma and Parambrostoma using morphological characters. Current taxonomic concepts Correspondence: Xing-Ke Yang, Institute of Zoology, Chinese (Daccordi, 1981; Kimoto & Gressitt, 1981; Wang & Chen, Academy of Sciences, 1 Beichen West Road, Chaoyang Districts, 1981) are evaluated with respect to their compatibility with Beijing 100101, China. E-mail: [email protected] the results of the parsimony analyses, in order to obtain a

A B

Fig. 1. Habitus: (A) Ambrostoma fortunei (Baly); (B) Parambrostoma mahesa (Hope). classification which truly reflects the phylogeny of the group Hawaii, USA (BPBM); California Academy of Sciences, San for the first time. The historical biogeography is discussed Francisco, California, U.S.A. (CAS); The Horst Kippenberg based on the phylogenetic results. We also provide a systematic collection, Herzogenaurach, Germany (HKPC); Hungarian review, redescriptions of the known species (see File S1), and Natural History Museum, Budapest, Hungary (HNHM); Insti- descriptions of two new species from Nepal. Dichotomous keys tute of Zoology, Chinese Academy of Sciences, Beijing, China to all the known species of the genera are presented separately. (IZAS); Kitakyushu Museum and Institute of Natural History, Another aim was to explore the usefulness of microcomputer Fukuoka, Kyushu, Japan (KMNH); Museo Civico di Storia tomography (μ-CT) in a taxonomic–phylogenetic study based Naturale ‘Giacomo Doria’, Genova, Italy (MCSN); Mauro on morphological data. Daccordi’s collection, Verona, Italy (MDC); Museum´ National d’Histoire Naturelle, Paris, France (MNHN); Naturkundesmu- seum, Erfurt, Germany (NMEG); Staatliches Museum fur¨ Natural history Naturkunde, Stuttgart, Germany (SMNS); United States National Museum (Natural History), Washington, District of Host plant records are only available for four species. The Columbia, U.S.A. (USNM); Uppsala University, Uppsala, three species Ambrostoma fortunei (Baly), A. leigongshanum Sweden (UZIU); Moscow State University, Moscow, Russia Wang and A. superbum (Thunberg) use the same host plant (ZMUM); and Zoological Institute, Russian Academy of Ulmus pumila Linnaeus (Ulmaceae). The host plant of Sciences, St. Petersburg, Russia (ZIN). Parambrostoma mahesa (Hope) is Nepeta sp. (Labiatae). Internal and external morphological characters of adults The larvae and adults of A. fortunei (Baly) and A. superbum form the basis of this work. Specimens were examined (Thunberg) feed on leaves and can cause heavy damage using a Leica microscope with a camera lucida (magnifi- (Yu et al., 1996). Ambrostoma superbum (Thunberg) has one cation 8–100×). Measurements were made using an ocular generation per year in northern China and A. fortunei Baly micrometer. Internal sclerotized structures were dissected in shows the same pattern in southern China (Yu et al., 1996). hot water. Heavily sclerotized parts were soaked in a diluted Ambrostoma superbum (Thunberg) is a major pest in northern solution (about 25%) of potassium hydroxide, then put in acetic China. Macquartia tenebricosa (Meigen, 1824) (Diptera) is acid and finally in ethanol. Species were characterized using: recorded as a predator of this species (Yu et al., 1996). colour; the shape of the clypeus, eyes, mouthparts, antennae, prothorax, scutellar shield, elytra, legs, mesoventrite, metaven- Materials and methods trite, abdomen; and punctures of the head, pronotum, elytra, and venter. One specimen of Ambrostoma fortunei (Baly) Taxa and terminology and one of Parambrostoma mahesa (Baly) were cleaned in a sonicator and sputter-coated with gold for scanning electron Specimens examined were obtained from: The Natural microscopy. Pictures were taken with an FEI (Philips) XL 30 History Museum, London, UK (BMNH); Bishop Museum, ESEM TMP. Digital photographs of the dorsal habitus were

© 2012 The Authors Systematic Entomology © 2012 The Royal Entomological Society, Systematic Entomology, 37, 332–345 334 S.-Q. Ge et al. taken with Automontage imaging systems by Syncroscopy with (Goloboff et al., 2008) (algorithm: traditional Search; starting a Leica M420 dissecting microscope. All pictures were evalu- trees: Wagner trees, 100 replicates; swapping algorithm: tree ated and assembled with Adobe Photoshop® and Illustrator® bisection reconnection). CS software. The terms mesoventrite and metaventrite replace the terms mesosternite and metasternite following Beutel & Haas (2000) and Lawrence et al. (2010). Characters (data matrix shown in Table 1) One specimen of Ambrostoma fortunei (Baly) was studied The selection of characters presented here was mainly based with high-resolution μ-CT (Skyscan 1142). The specimen for on features used in Ge et al. (2011), a study focused on the the μ-CT investigation was in 100% ethanol and critical point species-level phylogeny of the chrysomelid genera Suinzona, dried (EMS 850). The μ-CT data were obtained in Beijing Taipinus and Potaninia. Jishuitan Hospital, Beijing, China. The dataset has a resolution μ of 2.0 m in three dimensions. 1. Shape of body in lateral view: (0) strongly convex The following standards are used for characters: puncture (Fig. 3D); (1) not strongly convex. density, defined as dense if punctures are nearly confluent to 2. Surface structure of dorsum: (0) with metallic luster less than two puncture diameters apart, moderately dense if (Fig. 1A, B); (1) without metallic luster. punctures are between two to six puncture diameters apart, 3. Hind wing: (0) present (Fig. 7A); (1) absent. and sparse if punctures are separated by more than six punc- 4. Body colour: (0) multicoloured (Figs 1, 4); (1) unicoloured. ture diameters. Colour is described based on specimens viewed under magnification and with artificial illumination. Body Head length is measured from the anterior edge of the clypeus to 5. Frons: (0) concave (Fig. 5B); (1) not concave. the apex of the elytra. The body width is measured at the base 6. Frontal suture: (0) present; (1) absent (Fig. 5B). of the elytra. The pronotal length is the average length at mid- 7. Outer surface of mandibles: (0) with dense punctures; line. The width of the pronotum is the average width at the (1) with sparse punctures. middle region. The suture length is measured from the base 8. Outer surface of mandibles: (0) with sparse pubescence; of the elytral suture to the apex. The width of the elytra is (1) with dense pubescence. measured at the widest part. 9. Length of antennomere II :(0)II> IV (Figure S2A); (1) II = IV (Figure S10A). 10. Size ratio of maxillary palpomeres III and IV : (0) III > Taxon sampling IV; (1) III = IV; (2) III < IV (Fig. 5C). The taxon sampling includes all known species of Ambro- 11. Apex of maxillary palpomere IV : (0) truncate (Fig. 5C); stoma and Parambrostoma. Chrysolina mirabilis Daccordi was (1) pointed. selected as one of the outgroup taxa because Chrysolina was Thorax considered as closely related with the Ambrostoma lineage by 12. Trichobothrium of pronotum: (0) absent; (1) present on Daccordi (1994). Agrosteomela indica Hope, Humba cyani- anterior and posterior angles (Ge et al., 2011: fig. 29C). collis Hope, and Chrysomela populi Linnaeus were included as 13. Basal edge of pronotum: (0) immarginate; (1) marginate. they are likely closely related to the two genera under consid- 14. Lateral side of pronotum: (0) curved (Ge et al., 2011: eration (Daccordi, 1994) and presumably mainly characterised fig. 29C); (1) almost straight (Ge et al., 2011: fig. 28B). by unspecialised features compared to the ingroup species. 15. Lateral depression of pronotum: (0) present (Fig. 6A); (1) absent (Ge et al., 2011: fig. 27A). Cladistic analysis 16. Shape of scutellar shield: (0) longer than wide; (1) as long as wide; (2) shorter than wide. 47 adult morphological characters were coded and prepared 17. Humeral callus: (0) present; (1) absent. as a character state matrix using WINCLADA (Table 1). The 18. Transverse depression of elytra: (0) present (Figs 3D, 6A); cladistic analysis was performed in PAUP version 4.0b10 (1) absent. (Swofford, 2001) (branch and bound search) with all characters 19. Striae of elytra: (0) irregular (1) single striae; (2) double unordered and of equal weight. Inapplicable characters were striae. codes as “-”, and missing character states as “?” (Strong & 20. Elytral longitudinal stripes: (0) absent (Fig. 6C); (1) Lipscomb, 1999). Subsequently, the successive reweighting present (Fig. 6B, D). option implied in PAUP was applied. All outgroup species 21. Hypomera: (0) impunctate; (1) punctate. were treated as all other terminals in the analysis (simultaneous 22. Apex of prosternal process: (0) emarginate; (1) truncate. analysis; Nixon & Carpenter, 1993). Bremer support values 23. Size of apex of prosternal process: (0) strongly enlarged were calculated in PAUP using the ‘converse constraints (Fig. 5A); (1) not strongly enlarged (Fig. 9A). approach’ for branches supported in the strict consensus 24. Mesoventrite: (0) longer than wide; (1) not longer than trees with all characters equally weighted. All phylogenetic wide (Fig. 5A). trees were examined and manipulated in Dendroscope (Huson 25. Anterior edge of metaventrite: (0) immarginate (Fig. 5A); et al., 2007). Additional analyses were carried out using TNT (1) marginate.

Table 1. Character state matrix.

Character 0000000001 1111111112 2222222223 3333333334 4444444 Taxon 1234567890 1234567890 1234567890 1234567890 1234567 A. chinkinyui 0000000112 0100000010 0101011010 1001011001 0000??? A. fasciatum 0000000110 0100100010 0001011010 1001010000 1010000 A. fortunei 0000000112 0100000021 0001011010 0001110021 0010001 A. fulgurans 0000000112 0100000020 0001011010 1001100021 1110001 A. leigongshanum 0000000111 0100000021 0001011010 1001100000 0111001 A. omeishanum 0000000110 0100000011 0101011010 1001210010 1110??? A. superbum 0000000111 0100020021 0001011010 1001110021 0110001 A. rugosopunctatum 0000000110 0100020020 0001011010 1001100021 0110001 A. ambiguum 0010000012 0110000010 0111111010 11???????? ??????? P. kippenbergi 0010000011 0110010011 0111111010 1101111010 1110001 P. mahesa 0010001010 0110010001 0111011010 1101211000 1110001 P. medvedevi 0010001012 0110000011 0011011010 1101211000 1110001 P. montanum 0010001012 0110000010 0111011010 1101111000 1010000 P. shuteae 0010000012 0110020011 0011011010 1101111000 1110001 P. sublaeve 0010000012 0110100001 0011011010 ?1???????? ??????? Chrysomela populi 1101101001 1011000100 1000011010 0211110121 0100000 Agrosteomela indica 1101000002 0111001000 0111000101 1010110111 0000 – Chrysolina mirabilis 0010011002 0011001111 1111111011 0211111110 1000000 Humba cyanicollis 1101100012 1111121100 1011011111 0001111110 1010000

Inapplicable characters were codes as “-”, and missing character states as “?”

26. Shape of intercoxal process of metaventrite: (0) thickened; 41. Lateral knob of aedeagus: (0) present at each corner (1) not thickened. (Figure S10B); (1) absent (Figure S3B). 27. Shape of metaventrite: (0) convex, (1) not convex. 42. Lateral side of aedeagus before apex: (0) parallel-sided 28. Anterior part of metaventrite between coxae: (0) wider (Figure S3C); (1) not parallel-sided (Figure S4C). than long (Fig. 5A); (1) not wider than long. 43. Flagellum: (0) present (Figure S2C); (1) invisible 29. Metaventrite: (0) elongate, covering the mesoventrite; (1) (Figure S1C). not elongate, not covering the mesoventrite (Figs 5A, 9A). 44. Bifurcation of apex at flagellum: (0) absent (Figure S9C); 30. Metatarsomere III : (0) bilobed (Fig. 7B, C); (1) entire. (1) present (Figure S5C). Abdomen Female genitalia 31. Paired deep sinuation of last sternite of male: (0) absent; 45. Duct of spermatheca: (0) simple (Figure S5D); (1) convo- (1) present (Fig. 7F). luted (Ge et al., 2011: fig. 26H). 32. Setae of epipleuron: (0) present along the entire length 46. Length of base of spermatheca: (0) short (Figure S4D); (Fig. 8A); (1) present along the apical 1/3 (Fig. 8B); (1) long (Figure S10D). (3) absent. 47. Shape of spermatheca: (0) U-shaped (Figure S12D); Male genitalia (1) falciform (Figure S11D). 33. Shape of basal part of tegmen: (0) elongate (Ge et al., 2011; fig. 3A); (1) short (Ge et al., 2011; fig. 3B). 34. Basal portion of aedeagus: (0) sharply bent downward Results (Ge et al., 2011: fig. 13C); (1) not sharply bent downward (Figure S2B). Phylogenetic analysis 35. Length of basal foramen: (0) 1/3 of aedeagus; (1) less than 1/3 of aedeagus; (2) more than 1/3 of aedeagus. The analysis of the data matrix (Table 1) resulted in 62 36. Apical 1/3 part of aedeagus from lateral view: (0) much minimum-length trees with PAUP (CI = 0.464; RI = 0.614; narrower than basal part (Figure S5B); (1) not much RC = 0.285, 110 steps). The strict consensus tree is not well narrower than basal part (Figure S3B). resolved (Fig. 2A). However, only a single tree was obtained 37. Shape of apex of aedeagus: (0) anchor-shaped (Figure after applying the successive reweighting option in PAUP S10C); (1) not anchor-shaped (Figure S8B). (Fig. 2B). Using TNT the same number of steps (110) was 38. Apical orifice: (0) small; (1) large (Figure S7C). required, but only 30 minimum-length trees were obtained. 39. Apex of aedeagus: (0) rounded (Figure S1C); (1) subtri- Unambiguous apomorphies of clades confirmed in all trees angular (Figure S3C); (2) truncate (Figure S6C). with all characters unweighted: 40. Posterior edge of aedeagus: (0) flattened, curved ventrally (Figure S6B); (1) not flattened, not curved ventrally Ambrostoma. Outer surface of mandibles with dense pubes- (Figure S3B). cence (8:1); maxillary palpomere III > IV (10:0); basal edge of

B A Chrysolina mirabilis Chrysolina mirabilis Humba cyanicollis 1 Agrosteomela indica Chrysomela populi 51 Chrysomela populi Agrosteomela indica Humba cyanicollis A. fasciatum A. fasciatum 1 A. fortunei A. fortunei 1 1 A. fulgurans 1 A. fulgurans A. rugosopunctatum 2 A. rugosopunctatum A. superbum 2 1 50 A. superbum 61 A. leigongshanum Ambrostoma

A. leigongshanum Ambrostoma A. omeishanum A. omeishanum A. chinkinyui 3 A. chinkinyui 82 P. shuteae P. kippenbergi P. medvedevi P. ambiguum P. mahesa P. sublaeve 1 P. ambiguum P. mahesa P. montanum P. medvedevi P. sublaeve Parambrostoma outgroups

P. shuteae Parambrostoma outgroups P. kippenbergi P. montanum

Fig. 2. Phylogenetic reconstruction of Ambrostoma and Parambrostoma: (A) strict consensus tree of 62 most parsimonious trees (CI = 0.464, RI = 0.614, RC = 0.285) with 110 steps, branch support values (Bremer support) are shown above branches and bootstrap values (>49%) below. (B) tree obtained after applying the successive reweighting option implied in PAUP. pronotum immarginate (13:0); lateral side of pronotum curved Generic characters of Ambrostoma (14:0); apex of prosternal process strongly enlarged (23:0); Body. Elongate ovoid, large, 7–12 mm, convex dorsally; apex of aedeagus anchor-shaped (37:0); apical orifice small dorsal surface glabrous; metallic and multicoloured (Fig. 1A). (38:0); flagellum absent (43:1). Head. Subprognathous; clypeus trapezoid, base not raised; Parambrostoma. Hind wing absent (3:1); lateral side of frons concave (Fig. 5B); coronal suture and frontal suture pronotum curved (14:0); elytral longitudinal stripes present present. Compound eyes transversely oval or oblong, situated (20:1); setae of epipleuron present along apical 1/3 (32:1); api- obliquely at lateral margins of head. Labrum more or less cal orifice small (38:1); lateral side of aedeagus before apex not transverse, emarginated anteriorly, bearing a variable number parallel-sided (42:1); shape of spermatheca falciform (47:1). of setae, usually more numerous laterally. Antennae inserted at inner margin of eyes, entirely filiform or with the terminal segment slightly thickened (Fig. 5B); usually almost reaching basal transverse elytral impression posteriorly; scapus large, Review of genera thickened, sometimes slightly curved (Fig. 5B); pedicellus as long as or slightly shorter than fourth antennnomere; third Key to the genera of Ambrostoma and Parambrostoma distinctly longer than fourth; following segments subequal in length, often progressively elongated towards apex; mandibles Pronotum posteriorly immarginate; hindwings present; inner strong, bearing punctures and hairs; maxillary palpi with the margin of epipleuron with setae on whole length ...... terminal segment subtriangular, truncate apically (Fig. 5C)...... Ambrostoma Motschulsky Elytra with pronotum posteriorly marginate; hindwings absent; inner margin of epipleuron with setae only on apical part . . . . Pronotum. Transverse, convex; anteriorly marginate; setae ...... Parambrostoma Chen present along anterior margin; trichobothria present on anterior and posterior angles; lateral sides almost straight or more often narrowed in front and behind and widened and rounded Genus Ambrostoma Motschulsky, 1860 before midlength; anterior angles rounded, not strongly Ambrostoma Motschulsky, 1860: 205. Type species: Chryso- dilated anterior to middle (Fig. 5B); posterior angles almost mela superbum Thunberg, by subsequent designation. rectangular; posterior egde immarginate, broadly but slightly

C D Fig. 4. Habitus: (A) Ambrostoma superbum (Thunberg), holo- A. fortunei Parambrostoma kippenbergi sp.n. Fig. 3. Ambrostoma fortunei (Baly): volume rendering using μ-CT type; (B) (Baly); (C) ; P. medvedevi sp.n. image stacks. (A) dorsal view; (B) ventral view; (C) ventral-lateral (D) view; (D) dorsal-lateral view. of prosternal process more than half as wide as apex rounded in middle, distinctly sinuate on either side; pronotal (Fig. 5A); mesoventrite shorter than prosternum between coxae surface convex, punctate; close to lateral margin with more or (Fig. 5A), usually inclined and with triangular anteromedian less well-defined, broad, longitudinal depressed area, usually incision for reception of the prosternal process; metaventrite set with some coarse punctures (Fig. 6A). as long as prosternum; anteriorly immarginate (Fig. 5A). Legs moderately robust; tibiae simple, not armed with tooth-like Scutellar shield. Semicircular, ligulate, finely punctate or process, densely pubescent on apical half; metatarsomeres impunctate. Elytra: usually more or less glabrous, with humeral entirely pubescent ventrally; metatarsomere III bilobed, deeply callus; slightly widened posteriorly and rather deeply, trans- emarginate; claws simple (Fig. 7B). versely impressed before middle; transverse impression very well-defined (although somewhat shallower in the middle), Abdomen. With five distinct sternites. Aedeagus: tegmen Y- extending from lateral margin to near suture; slightly broader shaped; robust, simple in shape, slightly curved in lateral view. at base than prothorax; punctures either large or fine, densely Spermatheca: present, U-shaped or falciform, with simple duct. arranged or sparse; pattern often tending towards irregular arrangement of single or paired punctures; basal row of strong Secondary sexual characters. Male (1) first tarsomere of punctures from inner edge of humerus to transverse impres- fore and middle legs dilated and convex; (2) posterior border sion present. Epipleura flat and horizontal, inner margin with of last visible abdominal sternite more or less deeply sinuate setae entirely (Fig. 8A). Hindwings: present; Cu1a not con- on either side. Female: (1) first tarsomere of fore and middle nected with Cu1b by cross vein cv; apices of Cu1b and Pcu legs slightly less convex, not dilated, triangular and distinctly connected, separated (Fig. 7A). narrowed towards base; (2) posterior border of last abdominal sternite rounded, not sinuate. Thoracic venter. Anterior coxal cavities open posteriorly (Fig. 5A). Setae of anterior and posterior prosternal margin Remarks. The genus Ambrostoma can be characterised by present; prosternum elongate, slightly convex, dilated towards the following features: anterior coxal cavities open posteriorly, posterior margin where it is slightly excavated; more or inner margin of epipleuron with setae, claws simple, apex of less distinct, angular incision present medially; middle part process of metaventrite immarginate, well-defined transverse

AB1.0 mm md ant mxp ant mnt fr smnt e vx

2.0 mm hy pst

cx1 pn fem1 ppr tr1 cx2 v2 tr2 fem2 1.0 mm ep C v3 md ant

cx3 lbp tr3 lig mxp stI crd mnt fem3 smnt

pst

Fig. 5. SEM. Ambrostoma fortunei (Baly): (A) ventral side; (B) head, dorsal view; (C) head, ventral view. Abbreviations: ant, antenna; crd, cardo; cx1, fore-coxa; cx2, mid-coxa; cx3, hind-coxa; e, compound eyes; ep, epipleuron; fem1, fore-femur; fem2, mid-femur; fem3, hind-femur; fr, fron; hy, hypomeron; lbp, labial palpi; lig, ligula; md, mandible; mnt, mentum; mxp, maxillary palpi; pn, pronotum; ppr, prosternal process; pst, prosternum; smnt, submentum; stI, abdominal sternite I; tr1, fore-trochanter; tr2, mid-trochanter; tr3, hind-trochanter; v2, mesoventrite; v3, metaventrite; vx, vertex. depression at basal region of the elytra present. It is most – Elytra without well-defined post-median violaceous patch closely allied to the oriental genus Agrosteomela Hope, but this surrounded by green (Fig. 1A) ...... 4 genus differs from it in lacking the metallic lustre of the elytra 3. Arrangement of elytral punctures almost entirely irregular; and having a much shorter transverse elytral impression, which last abdominal sternite of male deeply emarginate on both sides is restricted to the region below the humerus. Moreover, the ...... A. fulgurans (Achard) process of the metaventrite of Agrostomela is very prominently – Elytral punctures arranged in simple rows at base before raised and produced, so that it covers a part of the mesoventrite. transverse impression (Fig. 4A), slightly irregular beyond The species of Ambrostoma display very similar color patterns, impression; last abdominal sternite moderately sinuate on both as has been mentioned above. Despite of this superficial sides...... A. fasciatum Chen uniformity they are sufficiently different in other structural 4. Pronotum slightly dilated in front of middle region; features as to be recognised as distinct species. interspaces between elytral striae finely and densely punctuate ...... A. superbum (Thunberg) Distribution. Eastern Asia (Fig. 10). – Pronotum strongly dilated in front of middle region; interspaces between elytral striae finely and very sparsely punctuate ...... 5 Key to the species of Ambrostoma Motschulsky 5. Elytra with double striae...... 6 – Elytral striae with single striae A. chinkinyui Kimoto & Osawa 1. Elytral punctures very distinctly impressed, at least as 6. Pronotum with dense and coarse punctures in lateral distinct as those of the lateral sides of pronotum ...... depression...... 7 ...... A. rugosopunctatum Chen – Pronotum with sparse punctures in lateral depression ...... – Punctures of elytra much finer than those of sides of ...... A. omeishanum Gressitt & Kimoto pronotum ...... 2 7. Elytra with longitudinal purple markings on media posterior 2. Elytra with large well-defined post-median violaceous patch surrounded by green; last segment in male shallowly sinuate. surrounded by green (Fig. 4B)...... 3 ...... A. leigongshanum Wang

AB slightly curved (Figure S11A); pedicellus as long as or slightly shorter than fourth antennnomere; third distinctly longer than fourth; following segments subequal, often increasing in length towards the apex; mandibles strong, bearing punctures and hairs; maxillary palpi with the terminal segment subtriangular, truncate apically (Fig. 5C).

Pronotum. Transverse, convex; anteriorly marginate; setae present along anterior margin; trichobothria present on anterior and posterior angles; lateral sides almost straight or more often narrowed in front and behind and widened and rounded before midlength; anterior angles rounded, not strongly dilated anterior to middle (Fig. 5B); posterior angles almost rectangu- CD lar; posterior edge marginate, broadly but slightly rounded in the middle, distinctly sinuate on either side; pronotal surface convex, punctate; close to the lateral margin with a more or less well-deﬁned, broad, longitudinal depressed area, usually set with some coarse punctures (Fig. 6D).

Scutellar shield. Semicircular, ligulate, finely punctate or impunctate. Elytra: usually more or less glabrous, with humeral callus; slightly widened posteriorly and rather deeply, transversely impressed before the middle; transverse impression very well-defined (although somewhat shallower in the middle), extending from the lateral margin to near the suture; slightly broader at base than prothorax; punctures either large or fine, densely arranged or sparse; pattern often tending towards irregular arrangement of single or paired punctures; Fig. 6. Habitus: (A) Ambrostoma leigongshanum Wang; (B) Parambrostoma kippenbergi sp.n.;(C)A. rugosopunctatum Chen; basal row of strong punctures from inner edge of humerus (D) P. medvedevi sp.n. to transverse impression present. Epipleura flat and horizontal, inner margin with setae only on apical part (Fig. 8B). Hindwings absent. – Elytra with four purple longitudinal stripes on centro- posterior part (Fig. 1A); last abdominal segment in male deeply sinuate(7F)...... A. fortunei (Baly) Thoracic venter. Procoxal cavities open posteriorly (Fig. 9A). Setae of anterior and posterior prosternal margin present; prosternum elongate, slightly convex, dilated towards pos- Genus Parambrostoma Chen terior margin where it is slightly excavated; more or less distinct, angular incision present medially; middle part of Ambrostoma (Parambrostoma) Chen, 1936: 718. Type species: prosternal process more than half as wide as apex (Fig. 9A); Ambrostoma sublaeve Chen, original designation. mesoventrite shorter than prosternum between coxae (Fig. 9A), Parambrostoma:Chenet Wang, 1981: 512. usually inclined and with triangular anteromedian incision for reception of the prosternal process; metaventrite as long Generic characters of Parambrostoma as prosternum; anteriorly immarginate (Fig. 9A). Legs mod- Body. Elongate ovoid, large, 5–10 mm, convex dorsally; erately robust; tibiae simple, not armed with tooth-like dorsal surface glabrous; metallic and multicoloured (Fig. 1B). process, densely pubescent on apical half; metatarsomeres entirely pubescent ventrally; metatarsomere III bilobed, deeply emarginate; claws simple (Fig. 7D). Head. Subprognathous; clypeus trapezoid, base not raised; frons concave (Fig. 9B); coronal suture and frontal suture present. Compound eyes transversely oval or oblong, situ- Abdomen. With five distinct sternites. Aedeagus: tegmen ated obliquely at the lateral margins of the head. Labrum Y-shaped; robust, simple in shape, slightly curved in lateral more or less transverse, emarginated anteriorly, bearing a view. Spermatheca: present, U-shaped or falciform, with variable number of setae, usually more numerous laterally. simple duct. Antennae inserted at inner margin of eyes, entirely filiform (Figure S10A); usually almost reaching basal transverse ely- Remarks. This genus is quite similar to Ambrostoma, tral impression posteriorly; scapus large, thickened, sometimes but differs in the following features: pronotum posteriorly

Pcu M F M3 Cu1a Cu1b

A G 0.30 mm 1.0 mm 1.0 mm

B C D E

Fig. 7. Ambrostoma fortunei (Baly): (A) hind-wing; (B) fore-leg, male; (C) fore-leg, female; (F) last abdominal segment, male; (G) last abdominal segment, female; (D and E), Parambrostoma mahesa (Hope): (D) fore-leg, male; (E) fore-leg, female.

A 0.2 mm B 0.5 mm

Fig. 8. SEM. Epipleuron, showing the setae on the inner side of epipleuron: (A) Ambrostoma fortunei (Baly); (B) Parambrostoma mahesa (Hope). Abbreviation: ep, epipleuron. marginate; hindwings absent; inner margin of epipleuron with 2. Elytral base with a short row of strong punctures extending setae only on apical part (Fig. 8B). from the inner edge of the humerus to the transverse impression ...... P. kippenbergi sp.n. – Elytral base without short row of strong punctures ...... Distribution. South slope of the Himalayas (Fig. 10)...... P. ambiguum Chen 3. Elytral punctures almost completely irregular (Fig. 1B) ...... P. mahesa Hope Key to the species of Parambrostoma Chen – Elytral punctures forming irregular striae ...... 4 4. Pronotum much broader than long, lateral margin straight, 1. Anterior part of metaventrite marginate ...... 2 without depression near lateral margin; lateral callus ﬂat, lateral – Anterior part of metaventrite immarginate (Fig. 5A) . . . . . 3 margin of pronotum visible ...... P. sublaeve Chen

AB1.0 mm 0.5 mm mxp md

lb md ant ant lig mxp mnt crd fr e smnt vx

hy pst

pn fem1 tr1 ppr cx1 C md ant v2 cx2 fem2 lbp

tr2 lig mxp 0.5 mm 0.5 crd v3 mnt ep smnt

cx3 stI hy tr3

Fig. 9. SEM. Parambrostoma mahesa (Hope): (A) ventral side; (B) head, dorsal view; (C) head, ventral view. Abbreviations: ant, antennae; crd, cardo; cx1, fore-coxa; cx2, mid-coxa; cx3, hind-coxa; e, compound eye; ep, epipleuron; fem1, fore-femur; fem2, mid-femur; fr, fron; hy, hypomeron; lb, labrum; lbp, labial palpi; lig, ligula; md, mandible; mnt, mentum; mxp, maxillary palpi; pn, pronotum; ppr, prosternal process; pst, prosternum; smnt, submentum; stI, abdominal sternite I; tr1, fore-trochanter; tr2, mid-trochanter; tr3, hind-trochanter; v2, mesoventrite;v3, metaventrite; vx, vertex.

◦ ◦ – Pronotum broader than long, lateral margin sinous or curved, 28 17N/84 27E, 8. 9. 2000, Hetzel leg. (IZAS). Paratypes: with depression near lateral margin; lateral callus convex, same data, one male and one female in MDC; one male and lateral margin of pronotum invisible (Fig. 4D) ...... 5 one female in IZAS; two females in HKPC. 5. Elytra with three longitudinal purple stripes on posteriome- dian area; pronotum with very dense punctures near depression; Etymology. Named after the German chrysomeline tax- apex of aedeagus elongate and narrow ...... 6 onomist Dr. Horst Kippenberg. – Longitudinal purple stripes absent; pronotum with very few punctures near depression, apex of aedeagus short and enlarged Diagnosis. Deﬁned by the unique shape of aedeagus...... P. montanum Medvedev 6. Lateral side of pronotum sinuous only at base, then straight Description. Body length 7.7–9.1 mm, width 4.0–4.6 mm. upward; lateral callus slightly swelling. Distal part of aedeagus Colouration of head metallic red bronze, with two dark slightly rounded (Figure S11C); basal part of spermatheca short markings surrounded by red bronze on central part; clypeus (FigureS11D)...... P. medvedevi sp.n. frons and posterior part of vertex metallic dark green; pronotum – Lateral side of pronotum curved in whole lenghth, Distal part dark green, with two subquadrate dark markings surrounded by of aedeagus rounded (Figure S13C); basal part of spermatheca red bronze, almost occupying entire central disc; two markings long(FigureS13D)...... P. shuteae Daccordi close to each other but not connected; lateral areas dark surrounded by red bronze; scutellar shield red bronze; elytra metallic dark green, with three dark stripes surrounded by Parambrostoma kippenbergi Daccordi & Ge, sp.n. reddish bronze; two of them close to elytral suture occupying (Fig. 4C, Figure S9A–D) entire length, the other one close to elytral margin interrupted by one green transverse marking on anterior 1/3 of elytra; ﬁrst Type specimens. Holotype: male, C-Nepal, Manaslu mas- two antennomeres, mouthparts, legs and venter reddish bronze, sif Barapokhari Lekh, 2650 m, 9 km NE Besisahar vill. eyes and apical antennomeres dark brown (Fig. 4C).

Head. Clypeus with sparse and moderately sized punctures Description. Body length 8.2–9.5 mm, width 4.0–4.5 mm. and sparse pubescence; frons with moderately sized and dense Colouration of head metallic green, with two large violaceous punctures; vertex with moderately sized and very sparse markings surrounded by green on central; pronotum greenish punctures; eyes kidney-shaped, average dorso-ventral diameter purple, with two violaceous markings separated from each 0.7 mm; aeverage interocular distance 1.5 mm; length of other on central disc; elytra with three broad violaceous stripes antennomeres (Figure S9A): 0.45 : 0.25 : 0.38 : 0.3 : 0.3 : 0.3 surrounded by green; interval of the patches, suture and lateral : 0.38 : 0.38 : 0.38 : 0.38 : 0.5 mm; outer surface of mandible margin green; antennae, mouthparts, scutellum, legs reddish with large and dense punctures and dense pubescence; green; eyes reddish green (Fig. 4D). maxillary palpomere IV as long as III, apex truncate. Head. Clypeus and frons with sparse and small punctures; Pronotum. 1.7–2.0 mm long, 3.7–4.3 mm wide; disc with clypeus additionally with dense pubescence; vertex with very moderately sized and sparse punctures, same as those of head; sparse and small punctures; eyes oblong, average interocular with shallow lateral longitudinal depression with large and distance 1.8 mm; average dorso-ventral diameter 0.6 mm; sparse punctures near lateral margin. Scutellar shield: ligulate, length of antennomeres (Figure S11A): 0.5 : 0.3 : 0.35 : 0.35 : smooth and with very fine and sparse punctures. Elytra: 0.35 : 0.4 : 0.35 : 0.35 : 0.35 : 0.35 : 0.6 mm; maxillary 5.0–6.7 mm long, 3.9–4.3 mm wide; row of strong punctures palpomere IV longer than III, apex truncate. Outer surface of from inner edge of humerus to transverse impression present; mandible with dense pubescence and with moderately sized with irregular single striae; interspace of striae with small and and sparse pubescence. sparse punctures. Pronotum. 2.0–2.1 mm long, 3.4–3.9 mm wide; disc with Venter. Hypomera and prosternum without punctures; pro- small and dense punctures, larger than those of head; with sternal process truncate apically; anterior part of metaventrite shallow lateral longitudinal depression, with dense and large margined and truncate; metanepisternum, mesepimeron and punctures in near lateral side. Scutellar shield: sub-triangular, metanepisternum with small and sparse punctures. with sparse and fine punctures. Elytra: 5.4–6.8 mm long, 4.4–5.6 mm wide; basal row of strong punctures from inner Aedeagus (Figure S9B, C). Slightly curved in lateral view; edge of humerus to transverse impression of elytra present; apex nearly as broad as aedeagus at the orifice; posterior edge with irregular single striae, as strong as those of side of flattened, curved ventrally, and at each corner bearing lateral pronotum; interspace with small and sparse punctures. lobe, which is sharply pointed and bent antero-ventrally; basal foramen less than one-third as long as aedeagus; flagellum Venter. Hypomera and prosternum without punctures; pro- visible, strongly developed. Spermatheca: present, U-shaped, sternum without punctures; prosternal process emarginate with simple duct (Figure S9D). apically; mesanepisternum, metanepisternum and mesepimeron with sparse and small punctures. Distribution. Nepal. Aedeagus (Figure S11B, C). Slightly curved in lateral view; Remarks. Similar to Parambrostoma ambiguum, but differ- apex wider than aedeagus at orifice; posterior edge flattened; ing in the following features: metallic markings on dorsum and curved ventrally, and at each corner possessing a lateral lobe, row of strong punctures at elytral base present. which is sharply pointed and bent antero-ventrally; basal foramen less than one-third as long as aedeagus; flagellum visible, strongly developed. Spermatheca: present, falciform, Parambrostoma medvedevi Daccordi & Ge, sp.n. with simple duct (Figure S11D). (Fig. 4D, Figure S11A–D) Distribution. Nepal. Type specimens. Holotype: Male, Nepal, Annapurna Mts., below Thulo Bugin, upper Dana 2500–2600 m, NN, 29.V. Remarks. Similar to Parambrostoma mahesa, but differing 2004, leg. J. Schmidt (NMEG). Paratypes: same data, two in the shape of the aedeagus. males and two females in IZAS; two males and two females in MDC; one male and one female (HKPC); two males and two females (BMNH).; eight males and five females in NMEG. Discussion

Phylogeny and taxonomy Etymology. Named after the Russian entomologist Dr. Lev N. Medvedev to acknowledge his work on Chrysomelidae. The strict consensus tree (Figure 2A) shows a well- supported clade formed by the genera Ambrostoma (eight Diagnosis. Deﬁned by the unique shape of the aedeagus. species) and Parambrostoma (seven species) (branch support

in Xishuangbanna (Yunnan, China), Laos and Thailand, and A. superbum in Siberia, Mongolia and North China. The distribution ranges of the other species are relatively limited. Ambrostoma fasciatum occurs in Tianmushan (Zhengjiang, China), A. fulgurans in Lo-Chouli-Tong (Yunnan, China) and Xianshan (Hunan, China) (also recorded from Annam, Viet- nam). Ambrostoma leigongshanum in Leigongshan (Guizhou, China), A. chinkinyui in Taiwan, and A. omeishanum in Emeishan (Sichuan, China). In clear contrast to this pattern, Parambrostoma is restricted to the southern slope of the Himalayas including Tibet and Nepal (Fig. 10). All the species of Parambrostoma are distributed east–west and display a zonal distribution pattern, living below an altitude of 3300 m in broadleaved deciduous forests and mixed conifer- ous/broadleaved forests. Moreover, all these species display morphological characters of alpine species, such as the absence of hind wings and a shiny metallic surface. The ancestor of the genera Ambrostoma and Parambrostoma was likely of tropi- cal origin. The splitting event resulting in two separate clades was likely related to the uplift of the earth’s crust and Quater- Fig. 10. Map showing the distribution of the genera Ambrostoma and nary climatic fluctuations. The species of Ambrostoma spread Parambrostoma: A. chinkinyui Kimoto & Osawa; A. omeishanum to the northeast as far as Siberia and gradually underwent Gressitt & Kimoto; A. fasciatum Chen; A. fortunei (Baly); speciation resulting in eight currently known extant species. A. fulgurans Achard; A. leigongshanum Wang; A. omeishanum Some of them are very widespread, such as A. superbum and Gressitt & Kimoto; A. superbum (Thunberg) ; A. rugosopunctatum Chen; Parambrostoma mahesa (Hope); P. medvedevi sp.n.; A. fortunei (see above). In clear contrast to the distribution P. montanum Medvedev; P. sublaeve Chen; P. shuteae Daccordi. pattern and evolution of Ambrostoma, high mountain barriers of the eastern and middle sections of the Himalayas and the rigorous climate of the northern plateau apparently confined value 3). Considering the distribution pattern (see below and Parambrostoma to a relatively limited area, where a modest Fig. 10) it would have appeared plausible to assume a subor- radiation took place. The relatively recent and rapid specia- dinate placement of Parambrostoma within the widely dis- tion is reflected by the poor resolution in the phylogenetic tributed genus erected earlier. However, the monophyly of analysis. both genera was confirmed with branch support values of 2 in thecaseofAmbrostoma (eight presumptive autapomorphies, see above), and 1 in the case of Parambrostoma. A place- Supporting Information ment of Parambrostoma ambiguum Chen in Chrysolina as sugested by Bienkowski (2007) can be clearly rejected based Additional Supporting Information may be found in the online on our phylogenetic results. An enforced clade Chrysolina version of this article under the DOI reference: + mirabilis Parambrostoma ambiguum requires seven addi- 10.1111/j.1365-3113.2012.00618.x tional steps. The poor resolution of the strict consensus tree reflects a high level of homoplasy as it is typical for Figure S1. Ambrostoma chinkinyui Kimoto et Osawa: A, morphology-based analyses on the species level (e.g., Komarek antennae, male; B, aedeagus, lateral view; C, aedeagus, & Beutel, 2007; Sikes et al., 2006; Ge et al., 2011). Notably, dorsal view. the phylogenetic relationships within Parambrostoma were not Figure S2. Ambrostoma fasciatum Chen: A, antennae, well resolved in the initial analyses. However, the analysis male; B, aedeagus, lateral view; C, aedeagus, dorsal view; after successive reweighting (implied in PAUP) yielded only D, spermatheca. one single tree (Fig. 2B). Figure S3. Ambrostoma fortunei (Baly): A, antennae, male; B, aedeagus, lateral view; C, aedeagus, dorsal view; Biogeography and evolution D, spermatheca. Figure S4. Ambrostoma fulgurans Achard: A, antennae, Ambrostoma and Parambrostoma show a typical southwest male; B, aedeagus, lateral view; C, aedeagus, dorsal view; and northeast disjunctive distribution pattern. The genus D, spermatheca. Ambrostoma has a very wide distribution area in the Far East, including eastern China, Korea, Vietnam, Laos, Thailand and Figure S5. Ambrostoma leigongshanum Wang: A, anten- Russia (Siberia) (Fig. 10). Ambrostoma fourtunei for example nae, male; B, aedeagus, lateral view; C, aedeagus, dorsal is widely distributed in South China, A. rugosopunctatum view; D, spermatheca.

Figure S6. Ambrostoma omeishanum Gressitt & Kimoto: (No. O529YX5105), the Knowledge Innovation Program of A, antennae, male; B, aedeagus, lateral view; C, aedeagus, Chinese Academy of Sciences (No. KSCX3-IOZ-1004), the dorsal view. National Science Fund for Fostering Talents in Basic Research (Special Subjects in Animal Taxonomy, NSFC-J0630964/ Figure S7. Ambrostoma rugosopunctatum Chen: A, J0109, J0930004). We used the Willi Hennig Society edition antennae, male; B, aedeagus, lateral view; C, aedeagus, of TNT which is gratefully acknowledged. dorsal view; D, spermatheca. Figure S8. Ambrostoma superbum (Thunberg): A, antennae, male; B, aedeagus, lateral view; C, aedeagus, dorsal References view; D, spermatheca. Achard, J. (1922) Descriptions de nouveaux Chrysomelini. Fragments Figure S9. Parambrostoma kippenbergi sp. n: A, anten- Entomologiques, 1–2, 13–28. nae, male; B, aedeagus, lateral view; C, aedeagus, dorsal Baly, J.S. (1860) Descriptions of six new species of Chrysomela from the East. The Journal of Entomology: Descriptive and Geographical, view; D, spermatheca. 1, 93–97. Figure S10. Parambrostoma mahesa (Hope): A, anten- Beutel, R.G. & Haas, F. (2000) Phylogenetic relationships of the nae, male; B, aedeagus, lateral view; C, aedeagus, dorsal suborders of Coleoptera (Insecta). Cladistics, 16, 103–141. Chen, S.H. (1934) On some species of Chrysomelidae in the British view; D, spermatheca. Museum. Stylops: A Journal of Taxonomic Entomology, 3, 66–78. Figure S11. Parambrostoma medvedevi sp. n.: A, anten- Chen, S.H. (1936) The chrysomelid genus Ambrostoma. Sinensia, 7, nae, male; B, aedeagus, lateral view; C, aedeagus, dorsal 713–729. Daccordi, M. (1977) Descrizione di Ambrostoma shuteae n. sp. del view; D, spermatheca. Nepal e considerazioni su Ambrostoma mahesa (Hope) (Coleoptera Figure S12. Parambrostoma montanum Medvedev: A, Chrysomelidae Chrysomelinae). Bollettino della Società Entomolog- antennae, male; B, aedeagus, lateral view; C, aedeagus, ica Italiana, 109, 133–138. Daccordi, M. (1981) Nuovi taxa di Chrysomelinae afrotropicali. dorsal view; D, spermatheca. Bollettino Museo Civico Storia Naturale Verona, 7, 181–195. Figure S13. Parambrostoma shuteae Daccordi: A, anten- Daccordi, M. (1994) Notes for phylogenetic study of Chrysomelinae, nae, male; B, aedeagus, lateral view; C, aedeagus, dorsal with descriptions of new taxa and a list of all the known genera (Coleoptera: Chrysomelidae: Chrysomelinae). Proceedings of the view; D, spermatheca. Third International Symposium on the Chrysomelidae, Beijing, 1992 Figure S14. Parambrostoma sublaeve Chen: A, antennae, (ed. by D.G. Furth), pp. 60– 84. Backhuys Publishers, Leiden. male; B, aedeagus, lateral view; C, aedeagus, dorsal view; Ge, S.Q., Daccordi, M., Beutel, R.G., Li, W.Z. & Yang, X.K. (2011) Revision of the chrysomeline genera Potaninia, Suinzona and D, spermatheca. 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(1981) Chrysomelidae (Coleoptera) of Thailand, Cambodia, Laos and Vietnam II. Clytrinae, Cryptocephali- nae, Chlamisinae, Lamprosomatinae and Chrysomelinae. Pacific Acknowledgements Insects, 23, 286–391. Kimoto, S. & Osawa, S. (1995) Descriptions of a new species of the We thank the following colleagues for the loan of mate- genus Ambrostoma Motschulsky from Taiwan, China (Coleoptera: rial: Dr. Horst Kippenberg (Herzogenaurach, Germany); Mrs. Chrysomelidae: Chrysomelinae). Entomological Review of Japan, Sharon Shute and Dr. Maxwell V. L. Barclay (BMNH); 50, 15–16. Dr. Allan G. Samuelson (BPBM); Dr. Otto Merkl (HNHM); Komarek, A. & Beutel, R.G. (2007) Problems in taxonomy and Dr. Wolgang Schawaller (SMNS); Dr. Alexander Konstantinov suggestions for a standardized description of new insect taxa. (USNM); Dr. Matthias Hartmann (NMEG); Dr. Roberto Poggi Entomological Problems, 36, 55–70. ´ (NMEG); Dr. Nicole Berti (MNHN); Dr. Hans Mejlon (UZIU); Lawrence, J.F., Beutel, R.G., Leschen, R.A.B. & Slipinski,´ A. (2000) Dr. Alexander Kirejtshuk (ZIN); Dr. David Kavanaugh (CAS). 2. Glossary of morphological terms. Coleoptera, Beetles, Vol 2: Mor- phology and Systematics (vol. ed. by R.A.B. Leschen, R.G. Beutel The project was supported by a grant from the National Sci- and J.F. Lawrence), Handbook of Zoology, Vol. IV: Arthropoda: ence Foundation of China to Xing-Ke Yang (PI, Grant No. Insecta Part 36 (ed. by N.-P. Kristensen and R.G. Beutel), 3010300101) and Si-Qin Ge (PI, Grant No. 30970390), and pp. 9–20. Walter de Gruyter, Berlin/ New York (New York). the grants from the Key Laboratory of Zoological System- Lobl,¨ I. & Smetana, A. (2010) Catalogue of Palaearctic Coleoptera, atics and Evolution of the Chinese Academy of Sciences Vol. 6: Chrysomeloidea. Apollo Books, Stenstrup.

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