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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 rugosopunc- tatum 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 relation- ships 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 © 2012 The Authors 332 Systematic Entomology © 2012 The Royal Entomological Society Revision of Ambrostoma and Parambrostoma 333 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.

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