A Contribution to the Knowledge of the Subfamily Panagaeinae Hope, 1838 from Asia. Part 2. East Palearctic and Oriental Species
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Studies and Reports Taxonomical Series 10 (2): 275-392, 2014 A contribution to the knowledge of the subfamily Panagaeinae Hope, 1838 from Asia. Part 2. East Palearctic and Oriental species of the genus Craspedophorus Hope, 1838, and the genus Tinoderus Chaudoir, 1879 (Coleoptera: Carabidae) Martin HÄCKEL1) & Erich KIRSCHENHOFER2) 1) Department of Game Management and Forestry Zoology, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 1176, CZ-165 21 Prague 6, Czech Republic e-mail: [email protected] 2) Otto Elsner Gasse 10 - 12, A 2380 Perchtoldsdorf, Austria e-mail: [email protected] Taxonomy, new species, Coleoptera, Carabidae, Craspedophorus, Tinoderus, Palearctic and Oriental Regions Abstract. Treated are Oriental and east Palearctic species of the genus Craspedophorus Hope, 1838 and the monotypic genus Tinoderus Chaudoir, 1879. Twenty new Oriental species of Craspedophorus are described: C. assamensis sp. nov. (India: Assam), C. austronesiensis (Indonesia: Nusa Tenggara, Maluku), C. buruensis sp. n. (Indonesia: Maluku), C. cenwanglao sp. nov. (China: Guangxi), C. chiangdaoensis sp. nov. (Thailand), C. chiangmaiensis sp. nov. (Thailand), C. facchinii sp. nov. (Thailand), C. freudeellus sp. nov. (Laos, Vietnam), C. hovorkai sp. nov. (Thailand), C. jakli sp. nov. (Laos), C. horaki sp. nov. (Vietnam), C. huensis sp. nov. (Vietnam), C. kerberos sp. nov. (Vietnam), C. khaoyai sp. nov. (Thailand), C. kiwlomensis sp. nov. (Thailand), C. lankaensis sp. nov. (Sri Lanka), C. phupanensis sp. nov. (Laos), C. punensis sp. nov. (India: Maharashtra), C. sikkimensis sp. nov. (India: Sikkim) and C. tamdaoensis sp. nov. (Vietnam). Four new subspecies are described: C. mandarinellus attapeuensis (Laos), C. mandarinellus malayensis (Western Malaysia), C. mannae sulawesiensis (Indonesia: Tanimbar I.) and C. sapaensis guangdongensis (China: Guangdong). The following new species groups are established: Craspedophorus basifasciatus group, C. elegans group, C. hexagonus group, C. kubani group, C. lykaon group, C. mandarinus group, C. obscurus group and C. sapaensis group (new status, formerly Dischissus sapaensis group). Craspedophorus microspilotus group is redefined. The following species are transferred from the genus Dischissus to the genus Craspedophorus: C. dehradunensis (Kirschenhofer, 2000) (India: Uttarakhand), comb. nov. and C. sapaensis (Kirschenhofer, 1994) (Thailand, Vietnam), comb. nov. Three species names are synonymized: Craspedophorus numitor Kirschenhofer, 2000 with C. tropicus (Hope, 1842), C. kachinensis Kirschenhofer, 2011 with C. laticornis Kirschenhofer, 2000, and C. louangnamthaensis Kirchenhofer, 2011 with C. saundersi (Chaudoir, 1869). INTRODUCTION This paper presents the results of our study of Palearctic and Oriental ground beetles of the subfamily Panagaeinae (Coleoptera, Carabidae). It is a continuation of Part 1 (Häckel & Kirschenhofer 2014) and concentrates on the systematics of the speciose and heterogeneous genus Craspedophorus Hope, 1838 and the monotypic genus Tinoderus Chaudoir, 1879. According to the most recent study (Häckel & Farkač 2013), the genus Craspedophorus comprises 151 species with mostly prevailing nocturnal activity, inhabiting tropical regions of the Old World and reaching into the eastern Palearctic (China, India, Japan, Nepal, Pakistan, Taiwan) and Australian regions (warmer areas of the Australian continent and New Guinea). Despite recent descriptions of numerous species by the second author (Kirschenhofer 2000- 2012), namely from southeastern Asia, the taxa of this subfamily and their bionomy in the 275 named faunistic regions remain inadequately known. Material from new localities comes primarily from local collectors, but mostly in small numbers, and neither representation in museum and large private collections provides substantially larger numbers of specimens, especially from southern China and the Oriental region. For instance, in the relatively recent taxonomic work “Carabidae from Vietnam” (Park, Trac et Will 2006) there is not a single species of Craspedophorus and the subfamily Panagaeinae is altogether absent. For that reason we include at the end of this paper a current checklist of the subfamily for Vietnam. As a continuation of our revision of the genus Dischissus Bates, 1873, we describe below 20 new species of closely related and possibly congeneric genus Craspedophorus from the eastern Palearctic and Oriental regions. Our current state of knowledge permitting, the described species are placed in homogeneous, but evidently not monophyletic groups, in an attempt to express mutual relations between the taxa. However, some of the taxa do not fit in the proposed groups, some others turn out to belong in the genus Dischissus Bates, 1873, and three names are deemed invalid: Craspedophorus numitor Kirschenhofer, 2000 is a new synonym of C. tropicus (Hope, 1842), C. kachinensis Kirschenhofer, 2011 is a new synonym of C. laticornis Kirschenhofer, 2000, and C. louangnamthaensis Kirschenhofer, 2011 is a new synonym of C. saundersi Chaudoir, 1869. MATERIAL AND METHODS Repositories: BMNH The Natural History Museum, London, United Kingdom (C. Gillett); MIZW Museum & Institute of Zoology - PAN, Warszawa, Poland (T. Huflejt); MNHB Museum of Natural History, Basel, Switzerland; MSNG Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy (R. Poggi); NMPC National Museum, Praha, Czech Republic (J. Hájek); NMWC Naturhistorisches Museum Wien, Austria (M. Jäch, H. Schillhammer); SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany (W. Schawaller); ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (B. Jaeger); ZMUC Statens Naturhistoriske Museum - Københavns Universitet, Denmark (O. Martin); ZSMC Zoologische Staatsammlung München, Germany (M. Balke, M. Baehr); CDW Private Collection of D. W. Wrase, Berlin, Germany; CMH Private Collection of M. Häckel, Praha, Czech Republic (property of NMPC); CRS Private Collection of R. Sehnal, Unhošť, Czech Republic; CSF Private Collection of S. Facchini, Piacenza, Italy. SYSTEMATIC PART Craspedophorus Hope 1838: 165; type species: Carabus reflexus Fabricius, 1781 (nec 1801): 302 [= Craspedophorus reflexus (Fabricius, 1781)] Eudema Laporte de Castelnau, 1840: 137; type species Panagaeus regalis Gory, 1833 Isotarsus LaFerté-Sénectere, 1851: 217; type species Panagaeus regalis Gory, 1833 Epicosmus Chaudoir, 1846: 512; type species Panagaeus tomentosus Vigors, 1825 [= Craspedophorus angulatus (Fabricius, 1781)] 276 Brachyonychus Chaudoir, 1878: 85; type species Epicosmus sublaevis Chaudoir, 1869 Acanthocosmus Jeannel, 1949: 855 (subgenus); type species Eudema nigrita Künckel d'Herculais, 1891 Brachycosmus Jeannel, 1949: 857 (subgenus); type species Panagaeus festivus Klug, 1833 This in our opinion basket genus contains a number of morphologically very different species as well as groups of mutually similar species that can be arranged into groups. In the regions covered by this study, the genus Craspedophorus is accompanied by other genera of the tribe Panagaeini Bonelli, 1810, namely Cintaroa Kasahara, 1989 (Taiwan), Panagaeus Latreille, 1802, Tinoderus Chaudoir, 1879 (eastern Palearct), Dischissus Bates, 1873, Euschizomerus Chaudoir, 1850, Microcosmodes Strand, 1936, Peronomerus Schaum, 1854 and Trichisia Motschulsky, 1865. Two other generic taxa used for Oriental species of the tribe, Eudema Laporte de Castelnau (1840: 137) and Epicosmus Chaudoir (1846: 512), based on e.g. the shape of the terminal palpomere or the transversity of the hind episternum (Chaudoir 1878: 85), were synonymized with Craspedophorus by Andrewes (1919). Species of some genera (Dischissus, Microcosmodes, Panagaeus, Tinoderus) can be due to elytral maculation easily confused with Craspedophorus. For that reason we include a key to the Palearctic and Oriental genera of the Panagaeinae (modified from Chaudoir 1878: 85). The character separating Craspedophorus from the otherwise very similar genus Dischissus is the absence of an excision that forms a cleft in the penultimate protarsomeres of both sexes. An exception is present in two groups of Craspedophorus, one of them being the C. sapaensis group (see Kirschenhofer 2000) that contains species originally placed in Dischissus Bates, 1873. Those species have the penultimate protarsomeres excised, but the cleft is shorter than half of tarsomere length whereas in Dischissus it exceeds half of the length. The second exception is the C. laevis group (formerly placed in the genus Brachyonychus Chaudoir, 1878: 86), in which a similar character was noted by Chaudoir (1878: 85). It is very variable, however, and lack of data including absence od DNA analyses makes any further division of the genus impossible. The geographic extent of the genus is very large, its species inhabit the entire Afrotropical and Oriental regions, most of the east Palearctic subregion, and the Sunda and Molucca islands in the Indo-Australian region. One species was described from Papua New Guinea, and some species inhabit the tropical and subtropical parts of Australia. In this study we list all species known from Asia (eastern Palearctic and Oriental species west to Lydekker‘s line, i.e. including the Moluccas), some of them hitherto placed in four species groups (one we transfer from another genus, see also Kirschenhofer 2000 and Häckel et Farkač 2012) and five more groups are proposed. In addition to descriptions of new species, those newly transferred to Craspedophorus are commented upon. This is