Revision of the Genus Diaphorocera Heyden, 1863 (Coleoptera, Meloidae, Cerocomini)
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Contributions to Zoology, 76 (2) 63-85 (2007) Revision of the genus Diaphorocera Heyden, 1863 (Coleoptera, Meloidae, Cerocomini) Federica Turco and Marco A. Bologna Dipartimento di Biologia, Università “Roma Tre”, viale Marconi 446-00146 Roma, Italy, [email protected], bologna@ uniroma3.it Key words: Coleoptera, Meloidae, Diaphorocera, taxonomy, phylogeny, morphology, faunistics, biogeography, Palaearctic region. Abstract Introduction Diaphorocera, a Saharo-Sindian genus belonging to the tribe The family Meloidae (Coleoptera, Tenebrionoidea) was Cerocomini, is revised and a new synonymy is proposed. A cla- recently revised cladistically (Bologna and Pinto, 2001) distic classifi cation is proposed as well, on a set of adult morpho- and four subfamilies were recognised: Eleticinae, logical characters. The available bionomical records, both origi- Meloinae, Tetraonycinae and Nemognathinae (Pinto nal and from literature, concerning phenology, elevation, habitat preference, and host plants, are summarised. Adult morphology and Bologna, 1999; Bologna and Pinto, 2002). Blister of all species is described and fi gured, the catalogue of localities beetles are phytophagous, feeding usually on leaves with maps of distribution is reported, and a biogeographical and/or fl owers of several plant families, and are char- analysis is proposed. acterised by cantharidin production and hypermeta- morphic development, except in the primitive sub- family Eleticinae (Pinto et al., 1996; Bologna et al., Contents 2001). Larvae feed on the provisions and larvae of Hymenoptera Apoidea, or on grasshopper (Acridoidea) Introduction ...................................................................................... 63 eggs (Bologna, 1991). Material and methods ..................................................................... 64 Within the subfamily Meloinae, our interest was Results ............................................................................................... 65 recently addressed to the Old World tribe Cerocomini, Bionomics ................................................................................... 65 which is composed of fi ve genera (Bologna and Pinto, Classifi cation ............................................................................. 65 Key to the species of Diaphorocera ........................................... 67 2002): Cerocoma Geoffroy, 1762 (about 27 species), Taxonomy ......................................................................................... 68 Diaphorocera Heyden, 1863 (8 spp.), Anisarthrocera Genus Diaphorocera ................................................................ 68 Semenov, 1895 (2 spp.), Rhampholyssa Kraatz, 1863 Group obscuritarsis .................................................................. 69 (2 spp.) and Rhampholyssodes Kaszab, 1983 (1 sp.). Diaphorocera carinicollis ................................................. 69 Bologna and Pinto (2001) used larval and adult char- Diaphorocera johnsoni ...................................................... 70 acters to reconstruct the phylogeny of Meloidae, but Diaphorocera obscuritarsis ............................................. 71 the placement of the tribe Cerocomini, then repre- Group hemprichi ....................................................................... 73 Diaphorocera chrysoprasis .............................................. 73 sented only by Cerocoma schreberi, remained unre- Diaphorocera hemprichi ................................................... 75 solved. The authors recognised two larval synapomor- Diaphorocera promelaena ................................................ 76 phies for the tribe, then confi rmed for other species of Diaphorocera sicardi ......................................................... 78 Cerocoma (Di Giulio et al., 2002) and also for Diapho- Diaphorocera peyerimhoffi ............................................... 78 rocera chrysoprasis (Turco et al., 2006). Moreover, the Biogeography ................................................................................... 80 monophyly of the tribe is strongly supported by adult Acknowledgements ........................................................................ 83 synapomorphies, as the position of antennae (sockets References ........................................................................................ 83 Appendix 1 ....................................................................................... 85 distant from eyes, placed below or on the frontal su- Appendix 2 ....................................................................................... 85 ture), the epigamic modifi cations of male head, anten- Appendix 3 ....................................................................................... 85 nae, maxillary palpi and fore legs, the labrum elongate Downloaded from Brill.com09/29/2021 11:40:14AM via free access 64 F. Turco and M.A. Bologna – Taxonomy and biogeography of Diaphorocera (meloid beetles) and longitudinally furrowed or carinate, the shape of Material and methods mouthparts and the endophallic structure (Bologna, 1991; Turco et al., 2003). During our fi eld researches we obtained ecological The biology of the tribe is poorly known with the information on host plants and phenology on three exception of some notes on host plants (Bologna, 1991), species (D. chrysoprasis, D. hemprichi of both subspe- on sexual behaviour of the genus Cerocoma (Turco et cies, D. johnsoni). We reared these species and we al., 2003), and on larval hosts of few Cerocoma species obtained the triungulin of D. chrysoprasis, described (see Bologna, 1991 for a synthesis). Pre-imaginal stages in a separate paper (Turco et al., 2006). of Cerocomini are known only for six species of Cero- For the taxonomical study we examined 396 speci- coma (Di Giulio et al., 2002 for a review; Turco and mens representing all the Diaphorocera species, with Bologna, unpublished), and one Diaphorocera (Turco at least one male of each species, including typical et al., 2006). Eggs are laid in holes dug by the female material, as reported in Table 1. The examined speci- in the ground as in other Meloinae, and the fi rst instar mens are preserved in the following collections (as- larvae (“triungulins”) are not phoretic, actively reaching sociated acronyms reported in the text): CB = M. A. the host’s nest where they develop to adult stage. As far Bologna coll., Università “Roma Tre”, Roma; CBA = as is known, triungulins feed on larvae and honey or on J. Batelka coll., Prague; CK = S. Krejcik coll., Unicov, eggs and food stored by Hymenoptera Aculeata (Bolog- Czech Republic; CP = J. D. Pinto coll., University of na, 1991 for a review). California, Riverside; CR = J. C. Ringenbach coll., After the fi rst studies on the larval morphology (Di Paris; CS = D. Sechi coll., Cagliari; ISR = Institut Giulio et al., 2002; Turco et al., 2006), our project was Sciéntifi que, Rabat; UCD = University of California addressed to the revision of the tribe Cerocomini as a Museum, Davis; MCSN = Museo Civico di Storia whole. The present paper is focused on the study of the Naturale G. Doria, Genova; MHNG = Muséum genus Diaphorocera; another contribution is addressed d’Histoire Naturelle, Genève; MNHN = Muséum Na- to the classifi cation of the fi ve genera and to the revision tional d’Histoire Naturelle, Paris; MVE = Museo di of Anisarthrocera, Rhampholyssa and Rhampholyssodes Storia Naturale, Venezia; HNHM = Hungarian Natural (Turco and Bologna, unpublished); the third study will History Museum, Budapest; OUM = Oxford Univer- be devoted to the revision of the more speciose genus sity Museum, Oxford. Cerocoma (Turco and Bologna, unpublished). For the phylogenetical analysis of the eight species The genus Diaphorocera is a typical Saharo-Sindian of Diaphorocera we used the software PAUP 4.0 (Swof- element, distributed in the whole Northern Africa, ford, 2002) and 24 adult characters available for all southern Middle East (Palestine and Israel), Arabian species, selected from a larger set previously identifi ed; Peninsula, and in southern Iran. This genus was never character 24 was not observed in D. peyerimhoffi be- revised: Bedel (1895) published the fi rst key to the spe- cause the dissection of types was not possible. The cies, updated by Kaszab (1951; taken up by Dvorˇak, character matrix with the related list are given in Ap- 1989), after which one new species and one new subspe- pendices 1 and 2. According to Bologna and Pinto cies were described (Kocher, 1954; Kaszab, 1983). (2002), the single taxon constrained as outgroup is “Anisarthrocera” semirufa (Fair- maire, 1882) (see “Classifi cation”). Table 1. Types and other specimens examined. Autapomorphic characters were uti- Species Types and Other lized for the diagnoses and key only. location specimens As expected, characters displayed D. carinicollis lectotype, 5 paralectotypes MNHN 1 some homoplasies, as indicated by D. chrysoprasis 90 relatively low consistency index (CI) D. hemprichi hemprichi 53 (Appendix 3). Both binary and multi- D. hemprichi saudita 4 paratypes HNHM 47 state characters were employed, all D. johnsoni 3 paratypes HNHM 8 elaborated as unordered and equally D. kerimii (syn. of D. chrysoprasis) holotype MCSN D. obscuritarsis holotype, 12 paratypes MNHN, HNHM 76 weighted, and processed by the D. obscuritarsis var. rufi cornis syntype MNHN branch-and-bound algorithm using D. peyerimhoffi holotype, 'allotype', 1 'cotype' ISR 17 ‘parsimony’ as optimality criterion D. promelaena 9 paratypes HNHM