Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Phylogeny of the Oxycoryninae sensu lato (Coleoptera : Belidae) and evolution of host-plant associations

A. E. Marvaldi A E , R. G. Oberprieler B , C. H. C. Lyal C , T. Bradbury C and R. S. Anderson D
+ Author Affiliations
- Author Affiliations

A CONICET, Instituto Argentino de Investigaciones de Zonas Áridas, Entomología, CC 507, 5500 Mendoza, Argentina.

B CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia.

C The Natural History Museum, Department of Entomology, Cromwell Road, London SW7 5BD, UK.

D Canadian Museum of Nature, PO Box 3443, Ottawa, ON K1P 6P4, Canada.

E Corresponding author. Email: marvaldi@lab.cricyt.edu.ar

Invertebrate Systematics 20(4) 447-476 https://doi.org/10.1071/IS05059
Submitted: 20 December 2005  Accepted: 29 June 2006   Published: 29 August 2006

Abstract

Phylogenetic relationships among the genera of the subfamily Oxycoryninae and other belids (Curculionoidea) were reconstructed by cladistic analysis using 21 terminals and 98 characters: 62 from imaginal morphology, 33 from larval morphology and three biological characters relating to host plants and larval feeding habits. Terminal taxa represent all extant genera of Oxycoryninae, two genera of each of the three tribes of Belinae plus two outgroup taxa used to root the tree. New information on the larvae and biology of the metrioxenines is used in phylogenetic reconstruction. In accord with the single optimal cladogram obtained, a revised classification of the Oxycoryninae is proposed. The subfamily is classified into three tribes (Oxycorynini, Metrioxenini and Aglycyderini), with the tribe Oxycorynini further classified into three subtribes (Oxycraspedina Marvaldi & Oberprieler, subtr. nov., Oxycorynina and Allocorynina) and the tribe Metrioxenini into two subtribes (Metrioxenina and Afrocorynina ( = Hispodini, syn. nov.)). Larval and adult unambiguous synapomorphies defining each clade are identified. Tracing the evolution of biological traits from the phylogenetic estimate indicates that drastic shifts to phylogenetically distant host plants occurred from the ancestral belid association with conifers. Structural, chemical and/or ecological similarities of the plant organs consumed apparently had a major influence in the colonisation of different plant taxa by this group of weevils.


Acknowledgments

An earlier version of this study was presented at the Phytophaga Symposium during the XXII International Congress of Entomology held in August 2004 in Brisbane, Australia, organised by Rolf Oberprieler and Catherine Duckett. A. E. M.’s work was funded in part by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET, Argentina) through a research grant (PIP-5766), by Fundación BBVA (‘Yámana’ project), and an Overseas Travel Grant by CSIRO Entomology to R. G. O. facilitated a visit by A. E. M. to the Zimmerman Weevil Research Laboratory in Canberra in February 2005 and the completion of this study. We thank Ben Boyd (CSIRO Entomology) for assistance with scanning electron microscopy and two referees for their constructive comments on the manuscript.


References


Alonso Zarazaga M. A., and Lyal C. H. C. (1999). ‘A World Catalogue of Families and Genera of Curculionoidea (Insecta: Coleoptera) (Excluding Scolytidae and Platypodidae).’ (Entomopraxis: Barcelona, Spain.)

Anderson R. S. (2005) New Oxycoryninae from Central and South America: phylogenetic and biogeographical implications (Coleoptera: Belidae). Systematic Entomology 30, 644–652.
CrossRef |

Anderson W. H. (1941) On some larvae of the genus Proterhinus (Coleoptera: Aglycyderidae). Proceedings of the Hawaiian Entomological Society 11, 25–35.

APG II (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141, 399–436.
CrossRef |

Arnoldi L. V. (1977). Eobelidae. In ‘Mesozoic Coleoptera’. (Eds L. V. Arnoldi, V. V. Zherikhin, L. M. Nikitrin and A. G. Ponomarenko.) pp. 195–241. (Nauka Publishers: Moscow, Russia.) [English translation, 1991, Oxonian Press: New Delhi, India.]

Böving A. G., and Craighead F. C. (1931). ‘An Illustrated Synopsis of the Principal Larval Forms of the Order Coleoptera.’ (Brooklyn Entomological Society: Brooklyn, New York, USA.)

Bruch C. (1912) Oxycorynus helleri, un nuevo gorgojo argentino. Anales del Museo Nacional de Buenos Aires 23, 265–267.

Bruch C. (1916) Un nuevo gorgojo del “Prosopanche” (Oxycorynus parvulus Bruch). Revista del Museo de La Plata 23, 231–232.

Bruch C. (1923) Coleópteros fertilizadores de “Prosopanche Burmeisteri” De Bary. Physis (Rio de Janeiro, Brazil) 7, 82–88.

Chamorro-Lacayo M. L., Konstantinov A. S. (2004) Morphology of the prothorax and procoxa in the New World Cryptocephalini (Coleoptera: Chrysomelidae: Cryptocephalinae). Zootaxa 676, 1–46.

Chevrolat L. A. A. (1832) Monographie de deux genres nouveaux dans la famille des Curculionites. Annales de la Société Entomologique de France 1, 210–220.

Chevrolat L. A. A. (1878) Diagnose d’un nouveaux genre de Curculionide de la division des Oxycorynides. Annales de la Société Entomologique de France, serie 5 8(3),

Cocucci A. E. (1965) Estudios en el género Prosopanche (Hydnoraceae) I. Revisión taxonómica. Kurtziana 2, 53–74.

Cowling R. M., Proches S., Vlok J. H. J. (2005) On the origin of southern African subtropical thicket vegetation. South African Journal of Botany 71, 1–23.

Crowson R. A. (1955). ‘The Natural Classification of the Families of Coleoptera.’ (N. Lloyd: London, UK.) [E. W. Classey: Hampton, UK, reprint 1967.]

Crowson R. A. (1960) The Phylogeny of Coleoptera. Annual Review of Entomology 5, 111–134.
CrossRef |

Crowson R. A. (1981). ‘The Biology of the Coleoptera.’ (Academic Press: London, UK.)

Crowson R. A. (1986) On the systematic position of Allocoryninae (Coleoptera: Allocorynidae). The Coleopterists Bulletin 40, 243–244.

Crowson R. A. (1991). The relations of Coleoptera to Cycadales. In ‘Advances in Coleopterology’. (Eds M. Zunino, X., Bellés and M. Blas.) pp. 13–28. (Asociación Europea de Coleopterología: Barcelona, Spain.)

Dransfield J. (1988). The palms of Africa and their relationships. In ‘Monographs in Systematic Biology Volume 25. Modern Systematic Studies in African Botany’. (Eds P. Goldblatt and P. P. Lowry.) pp. 95–103. (Missouri Botanical Garden Press: St Louis, MO, USA.)

Farrell B. D. (1998) “Inordinate Fondness” explained: why are there so many beetles? Science 281, 555–559.
CrossRef | PubMed |

Fauvel A. (1891) Les Coléoptères de la Nouvelle-Calédonie et dépendances avec descriptions, notes et synonymies nouvelles. Revue d’Entomologie 10, 148–182.

Fauvel A. (1904) Faune analytique des Coléoptères de la Nouvelle-Calédonie, 2e partie. Revue d’Entomologie 23, 113–208.

Fawcett P. K. S., and Norstog K. (1993). Zamia pumila in South Florida: a preliminary report on its pollinators R. slossoni, a snout weevil and P. zamiae, a clavicorn beetle. In ‘The Biology, Structure, and Systematics of the Cycadales: Proceedings of CYCAD 90, the Second International Conference on Cycad Biology’. (Eds D. W. Stevenson and K. Norstog.) pp. 109–119. (The Palm and Cycad Societies of Australia: Milton, Australia.)

Germishuizen G., and Meyer N. L. (Eds) (2003). ‘Plants of Southern Africa: an Annotated Checklist.’ (Strelitzia 14, National Botanical Institute: Pretoria, South Africa.)

Goloboff P., Farris J., and Nixon K. (2003). ‘T.N.T.: Tree Analysis Using New Technology.’ Program and documentation, available from the authors and online at: www.zmuc.dk/public/phylogeny [verified July 2006].

Gratshev V. G., and Zherikhin V. V. (2000). The weevils from the Late Cretaceous New Jersey amber (Coleoptera, Curculionoidea). In ‘Studies on Fossils in Amber, with Particular Reference to the Cretaceous of New Jersey’. (Ed. D. Grimaldi.) pp. 241–254. (Backhuys Publishers: Leiden, Holand.)

Gratshev V. G., and Zherikhin V. V. (2003). The fossil record of weevils and related beetle families (Coleoptera, Curculionoidea). In ‘Fossil Insects’. Proceedings of the 2nd Congress on Palaeoentomology: Krakow, Poland, 5–9 September, 2001. (Eds E. Krzeminska and W. Krzeminski.) pp. 129–138. Acta Zoologica Cracoviensia 46, Supplement.

Hansen B. (1980) A monograph of the neotropical Balanophoraceae. Flora Neotropica 23, 1–81.

Heller K. M. (1911) Übersicht der Oxycorynus-Arten. Wiener Entomologische Zeitung 30, 5–8.

Hill K. D., Chase M. W., Stevenson D. W., Hills H. G., Schutzman B. (2003) The families and genera of cycads: a molecular phylogenetic analysis of Cycadophyta based on nuclear and plastid DNA sequences. International Journal of Plant Sciences 164, 933–948.
CrossRef |

Howden A. T. (1995) Structures related to oviposition in Curculionoidea. Memoirs of the Entomological Society of Washington 14, 53–100.

IPNI (2004). ‘The International Plant Names Index.’ (The Royal Botanic Gardens, Kew, The Harvard University Herbaria and the Australian National Herbarium.) Available online at: www.ipni.org/index.html [verified July 2006].

Kasap H., Crowson R. A. (1977) A comparative study of the internal anatomy and abdominal structures of Curculionoidea. Hacettepe Bulletin of Natural Sciences and Engineering 6, 35–86.

Kolbe H. (1908) Mein System der Coleopteren (Schluss). Zeitschrift für Wissenschaftliche Insektenbiologie 4, 389–400.

Kukalová-Peck J., Lawrence J. F. (2004) Relationships among coleopteran suborders and major endoneopteran lineages: evidence from hind wing characters. European Journal of Entomology 101, 95–144.

Kuschel G. (1955) Nuevas sinonimias y anotaciones sobre Curculionoidea (1). Revista Chilena de Entomologia 4, 261–312.

Kuschel G. (1959) Nemonychidae, Belidae y Oxycorynidae de la fauna Chilena, con algunas consideraciones biogeográficas. Investigaciones Zoológicas Chilenas 5, 229–271.

Kuschel G. (1990). Beetles in a suburban environment: a New Zealand case study. The identity and status of Coleoptera in the natural and modified habitats of Lynfield, Auckland (1974–1989). DSIR Plant Protection Report No. 3, New Zealand Department of Scientific and Industrial Research, Auckland, New Zealand.

Kuschel G. (1992) Reappraisal of the Baltic Amber Curculionoidea described by E. Voss. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 73, 191–215.

Kuschel G. (1995a) A phylogenetic classification of Curculionoidea to families and subfamilies. Memoirs of the Entomological Society of Washington 14, 5–33.

Kuschel G. (1995b) Oxycorynus missionis spec. nov. from NE Argentina, with key to the South American species of Oxycoryninae (Coleoptera Belidae). Acta Zoológica Lilloana 43, 45–48.

Kuschel G. (2000) La fauna curculiónica (Coleoptera: Curculionoidea) de la Araucaria araucana. Revista Chilena de Entomologia 27, 41–51.

Kuschel G. (2003). ‘Nemonychidae, Belidae, Brentidae (Insecta: Coleoptera: Curculionoidea).’ Fauna of New Zealand, Volume 45. (Manaaki Whenua Press: Lincoln, New Zealand.).

Kuschel G., and Leschen R. A. B. (2003). Appendix 1. Phylogenetic relationships of the genera of Belinae. In ‘Nemonychidae, Belidae, Brentidae (Insecta: Coleoptera: Curculionoidea)’. (Author: G. Kuschel.) pp. 48–55. Fauna of New Zealand, Volume 45. (Manaaki Whenua Press: Lincoln, New Zealand.)

Labandeira C. C. (2002). Chapter 2. The history of associations between plants and animals. In ‘Plant–Animal Interactions: An Evolutionary Approach’. (Eds C. M. Herrera and O. Pellmyr.) pp. 26–74. (Blackwell Science: Oxford, UK.)

Lacordaire J. T. (1863). ‘Histoire Naturelle des Insectes. Genera des Coléoptères ou exposé méthodique et critique de tous genres proposés jusqu'ici dans cet ordre d'insectes. Tome sixième, contenant les famille des Curculionides.’ (Libraire Encyclopédique de Roret: Paris, France.)

Lawrence J. F., and Newton A. F. (1995). Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). In ‘Biology, Phylogeny and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson’. (Eds J. Pakaluk and S. A. Slipinski.) pp. 781–913. (Muzeum i Instytut Zoologii PAN: Warszawa, Poland.)

Marshall G. A. K. (1955) New South African Curculionidae (Col.). Annals and Magazine of Natural History (Historical Association (Great Britain)), Series 12 8, 1–24.

Marvaldi A. E. (2005) Larval morphology and biology of oxycorynine weevils and the higher phylogeny of the Belidae (Coleoptera, Curculionoidea). Zoologica Scripta 34, 37–48.
CrossRef |

Marvaldi A. E., Morrone J. J. (2000) Phylogenetic systematics of weevils (Coleoptera: Curculionoidea): a reappraisal based on larval and adult morphology. Insect Systematics & Evolution 31, 43–58.

Marvaldi A. E., Sequeira A. S., O’Brien C. W., Farrell B. D. (2002) Molecular and morphological phylogenetics of weevils (Coleoptera, Curculionoidea): do niche shifts accompany diversification? Systematic Biology 51, 761–785.
CrossRef | PubMed |

Marvaldi A. E., Lanfranco D., and Ruiz C. (2003). Hallazgo de larvas de Oxycraspedus (Coleoptera: Belidae: Oxycoryninae) en estróbilos femeninos de Araucaria araucana (Mol.) Koch. In ‘Resúmenes del XXV Congreso Nacional de Entomología’. p. 8. (Universidad de Talca: Talca, Chile.)

May B. M. (1993). ‘Larvae of Curculionoidea (Insecta: Coleoptera): A Systematic Overview’. Fauna of New Zealand, Volume 28. (Manaaki Whenua Press: Lincoln, New Zealand.)

May B. M. (1994). An introduction to the immature stages of Australian Curculionoidea. In ‘Australian Weevils (Coleoptera: Curculionoidea). Volume II – Brentidae, Eurhynchidae, Apionidae and a Chapter on Immature Stages by Brenda May’. (Author: E. C. Zimmerman.) pp. 365–728. (CSIRO Publishing: Melbourne, Australia.)

Menier J. J. (1974) Description d’une deuxiéme espèce du genre Aglycyderes Westwood: A. tavakiliani n. sp. (Col. Aglycyderidae). Bulletin de la Societe Entomologique de France 79, 14–16.

Montrouzier P. (1861) Essai sur la faune entomologique de la Nouvelle-Calédonie (Balade), et des îles des Pins, Art, Lifu, etc. Coléoptères (Fin) (1). Annales de la Société Entomologique de France, serie 4 1, 265–306.

Morrone J. J. (1996) Austral biogeography and relict weevil taxa (Coleoptera: Nemonychidae, Belidae, Brentidae and Caridae). Journal of Comparative Biology 1, 123–127.

Morrone J. J. (1997) The impact of cladistics in weevil classification with a new scheme of families and subfamilies (Coleoptera: Curculionoidea). Trends in Entomology 1, 129–136.

Morrone J. J. (2006) Biogeographic areas and transition zones of Latin America and the Caribbean Islands based on panbiogeographic and cladistic analyses of the entomofauna. Annual Review of Entomology 51, 467–494.
CrossRef | PubMed |

Morrone J. J., and Roig-Juñent S. (1995). ‘The Diversity of Patagonian Weevils. An Illustrated Checklist of the Patagonian Curculionoidea (Insecta: Coleoptera). Monografía 2.’ (Literature of Latin America: Buenos Aires, Argentina.)

Muñiz R., Barrera A. (1969) Rhopalotria dimidiata Chevrolat, 1878: estudio morfológico del adulto y descripción de la larva (Ins. Col. Curcul.: Oxycoryninae). Revista de la Sociedad Mexicana de Historia Natural 30, 205–222.

Nickrent D. L., Blarer A., Qiu Y.-L., Soltis D. E., Soltis P. S., Zanis M. (2002) Molecular data place Hydnoraceae with Aristolochiaceae. American Journal of Botany 89, 1809–1817.

Nickrent D. L., Der J. P., Anderson F. E. (2005) Discovery of the photosynthetic relatives of the “Maltese mushroom” Cynomorium. BMC Evolutionary Biology 5, 1–11.
CrossRef | PubMed |

Nixon K. C. (2002). ‘WinClada, version 1.00.08’. (Published by the author: Ithaca, NY, USA.) Available online at: www.cladistics.com [verified July 2006].

Norstog K. (1987) Cycads and the origin of insect pollination. American Scientist 75, 270–279.

Norstog K., Fawcett P. K. S. (1989) An insect-cycad symbiosis and its relation to the pollination of Zamia furfuracea (Zamiaceae) by Rhopalotria mollis (Curculionidae). American Journal of Botany 76, 1380–1394.
CrossRef |

Norstog K. J., and Nicholls T. J. (1997). ‘The Biology of the Cycads.’ (Cornell University Press: Ithaca, USA.)

Norstog K., Stevenson D. W., Niklas K. J. (1986) The role of beetles in the pollination of Zamia furfuracea L. fil. (Zamiaceae). Biotropica 18, 300–306.
CrossRef |

Norstog K., Fawcett P. K. S., Vovides A. P. (1992) Beetle pollination of two species of Zamia: evolutionary and ecological considerations. Palaeobotanist 41, 149–158.

Norstog K., Fawcett P. K. S., Nicholls T. J., Vovides A. P., and Espinosa E. (1995). Insect pollination of cycads: evolutionary and ecological considerations. In ‘Proceedings of the Third International Conference on Cycad Biology’. (Ed. P. Vorster.) pp. 265–285. (The Cycad Society of South Africa: Stellenbosch, South Africa.)

Oberprieler R. G. (1995a). The weevils (Coleoptera: Curculionoidea) associated with cycads. 1. Classification, relationships, and biology. In ‘Proceedings of the Third International Conference on Cycad Biology’. (Ed. P. Vorster.) pp. 295–334. (The Cycad Society of South Africa: Stellenbosch, South Africa.)

Oberprieler R. G. (1995b). The weevils (Coleoptera: Curculionoidea) associated with cycads. 2. Host specificity and implications for cycad taxonomy. In ‘Proceedings of the Third International Conference on Cycad Biology’. (Ed. P. Vorster.) pp. 335–366. (The Cycad Society of South Africa: Stellenbosch, South Africa.)

Oberprieler R. G. (2004). “Evil weevils” – the key to cycad survival and diversification? In ‘Proceedings of the 6th International Conference on Cycad Biology’. (Ed. A. J. Lindstrom.) pp. 170–194. (Nong Nooch Tropical Botanical Garden: Chonburi, Thailand.)

Pascoe F. P. (1868) Description of a new species of Curculionidae. Proceedings of the Entomological Society of London 1868, xiv.

Pascoe F. P. (1870) Contributions towards a knowledge of the Curculionidae. Part I. Journal of the Linnean Society of London 10, 434–458.

Paulian R. (1944) Les Aglycyderidae, une famille relicte. Revue Francaise d’Entomologie 10, 113–119.

Perkins R. C. L. (1900). Coleoptera. II. Coleoptera Rhynchophora, Proterhinidae, Heteromera and Cioidae. In ‘Fauna Hawaiiensis or the Zoology of the Sandwich (Hawaiian) Isles: Being Results of the Explorations instituted by the Joint Committee appointed by the Royal Society of London for Promoting Natural Knowledge and the British Association for the Advancement of Science. And carried on with the assistance of those Bodies and of the Trustees of the Bernice Pauahi Bishop Museum at Honolulu, Vol. 2(3)’. (Ed. D. Sharp.) pp. 117–270. (Cambridge University Press: Cambridge MA, USA.)

Perkins R. C. L. (1907) On a species of Proterhinus from Samoa. Proceedings of the Hawaiian Entomological Society 1, 87–88.

Perkins R. C. L. (1920) Some new Hawaiian Coleoptera. Proceedings of the Hawaiian Entomological Society 4, 341–359.

Perkins R. C. L. (1928) Notes on Proterhinus. Proceedings of the Hawaiian Entomological Society 7, 193–200.

Perkins R. C. L. (1931) Descriptions of new Hawaiian Coleoptera. Proceedings of the Hawaiian entomological Society 7, 509–515.

Perkins R. C. L. (1932) On two new species of Proterhinus from the Marquesas and the inclusion of this genus in the family Aglycyderidae. Bernice P. Bishop Museum Bulletin 98, 17–21.

Perkins R. C. L. (1936) A new species of Proterhinus from the Marquesas Islands. Entomologist's Monthly Magazine 72, 221–222..

Schneider D., Wink M., Sporer F., Lounibos P. (2002) Cycads: their evolution, toxins, herbivores and insect pollinators. Die Naturwissenschaften 89, 281–294.
CrossRef | PubMed |

Schoenherr C. J. (1840). ‘Genera et species curculionidum, cum synonymia hujus familiae. Species novae aut hactenus minus cognitae, descriptionibus a Dom. L. Gyllenhal, C. H. Boheman, O. J. Fahraeus et entomologis aliis, illustratae. Vol. 5, Part 2.’ (Roret: Paris, France.)

Scudder S. H. (1893) Tertiary Rhynchophorous Coleoptera of the United States. Monographs of the United Stated Geological Survey 21, 1–206.

Sharp D. (1878) Descriptions of some new species and a new genus of rhynchophorous Coleoptera, from the Hawaiian Islands. Transactions of the Entomological Society of London 1878, 15–26.

Sharp D. (1890). Subfam. Attelabinae, Pterocolinae, Allocoryninae, Apioninae, Thecesterninae, Otiorhynchinae (part, Apterae). In ‘Biologia Centrali-Americana, Insecta. Vol. 4, Part 3. Rhynchophora. Curculionidae. Attelabinae, Pterocolinae, Allocoryninae, Apioninae, Thecesterninae, Otiorhynchinae’. (Eds. D. Sharp and G. C. Champion.) pp. 41–80. (R. H. Porter, London, UK.)

Sharp D. (1899). Part II. Hymenoptera continued (Tubulifera and Aculeata), Coleoptera. Strepsiptera, Lepidoptera, Diptera, Aphaniptera, Thysanoptera, Hemiptera, Anoplura. In ‘Insects, the Cambridge Natural History, Vol. 6’. (Eds S. F. Harmer and A. E. Shipley.) 626 pp. (Macmillan and Co.: London, UK.)

Stevenson D. W., Norstog K., and Fawcett P. K. S. (1998). Pollination biology of cycads. In ‘Reproductive Biology in Systematics, Conservation and Economic Botany’. (Eds S. J. Owens and P. J. Rudall.), pp. 277–294. (Royal Botanic Gardens: Kew, UK.)

Swezey O. H. (1913) A leaf-mining Proterhinus. Proceedings of the Hawaiian Entomological Society 2, 212–214.

Swezey O. H. (1921) Notes on Proterhinus abnormis, a leaf-miner in leaves of Broussaisia arguta (Col.). Proceedings of the Hawaiian Entomological Society 4, 470–471.

Swezey O. H. (1927) Notes on the host plants of the species of Proterhinus in the Kokee region of Kauai. Proceedings of the Hawaiian entomological Society 6, 489–495.

Swofford D. L. (1999). ‘PAUP*: Phylogenetic Analysis Using Parsimony (* And Other Methods), version 4.0’. (Sinauer: Sunderland, MA, USA.)

Tang W. (1987) Insect pollination in the cycad Zamia pumila (Zamiaceae). American Journal of Botany 74, 90–99.
CrossRef |

Tang W. (2004). Cycad insects and pollination. In ‘Vistas in Palaeobotany and Plant Morphology: Evolutionary and Environmental Perspectives. Professor D. D. Pant Memorial Volume.’ (Ed. P. C. Srivastava.) pp. 383–394. (U.P. Offset: Lucknow, India.)

Thompson R. T. (1992) Observations on the morphology and classification of weevils (Coleoptera: Curculionoidea) with a key to the major groups. Journal of Natural History 26, 835–891.

Van Emden F. (1938) On the taxonomy of Rhynchophora larvae. Transactions of the Royal Entomological Society of London 87, 1–37.

Voss E. (1922) Monographische Bearbeitung der Unterfamilie Rhynchitinae (Curc.). 1. Teil. Nemonychini – Auletini. Archiv für Naturgeschichte 1922, 1–113.

Voss E. (1935). Subfam. Oxycoryninae (pp. 1–2). In ‘Coleopterorum Catalogus auspiciis et auxilio W. Junk editus a S. Schenkling. Pars 144: Curculionidae: Otidocephalinae, Ithycerinae, Belinae, Petalochilinae, Oxycoryninae’. (Eds K. W. von Dalla Torre and E. Voss.) (W. Junk: s’Gravenhage, The Netherlands.)

Voss E. (1943) Monographie der Rhynchitinen-Tribus Deporaini sowie der Unterfamilie Pterocolinae – Oxycoryninae (Allocorini). VII. Teil der Monographie der Rhynchitinae-Pterocolinae. Stettiner Entomologische Zeitung 104, 46–63.

Voss E. (1953) Einige Rhynchophoren der Bernsteinfauna (Col.). Mitteilungen aus dem Geologischen Staatsinstitut in Hamburg 22, 119–140.

Voss E. (1957) Archimetrioxena electrica Voss und ihre Beziehungen zu rezenten Formenkreisen (Col., Curc.). Deutsche Entomologische Zeitung (N.F.) 4, 95–102.

Voss E. (1965) Die Unterfamilie Camarotinae, ihre Beziehungen zur Familie Attelabidae sowie ein Versuch, die phylogenetischen Zusammenhänge innerhalb der letzteren zur Darstellung zu bringen (Coleoptera, Curculionidae). Entomologische Abhandlungen, Staatliches Museum für Tierkunde in Dresden 32, 223–244.

Vovides A. P. (1991) Insect symbionts of some Mexican cycads in their natural habitat. Biotropica 23, 102–104.
CrossRef |

Wagner W. L., Herbst D. R., and Sohmer S. H. (1999). ‘Manual of the Flowering Plants of Hawai’i. Revised Edition. Bishop Museum Special Publication 97’. (University of Hawai’i Press and Bishop Museum Press: Honolulu, Hawaii.)

Walters T., Osborne R., and Decker D. (2004). “We hold these truths... ”. In ‘Cycad Classification: Concepts and Recommendations’. (Eds. T. Walters and R. Osborne.) pp. 1–11. (CABI Publishing: Wallingford, UK and Cambridge, USA.)

Westwood J. O. (1864) Description of two new Coleoptera from the Canary Islands. Journal of the Proceedings of the Entomological Society of London (1863), Series 3) 1, 178–179.

Weitschat W., and Wichard W. (2002). ‘Atlas of Plants and Animals in Baltic Amber.’ (Verlag Dr. Friedrich Pfeil: München, Germany.)

Wollaston T. V. (1864). ‘Catalogue of the Coleopterous Insects of the Canaries in the Collection of the British Museum.’ (Taylor and Francis: London, UK.)

Zherikhin V. V. (2002). Ecological history of terrestrial insects. In ‘History of Insects’. (Eds A. P. Rasnitsyn and D. L. J. Quicke.) pp. 331–388. (Kluwer Academic Publishers: Dordrecht, The Netherlands.)

Zherikhin V. V., and Gratshev V. G. (1995). A comparative study of the hind wing venation of the superfamily Curculionoidea, with phylogenetic implications. In ‘Biology, Phylogeny and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson’. (Eds J. Pakaluk and S. A. Slipinski.) pp. 633–777. (Muzeum i Instytut Zoologii PAN: Warszawa, Poland.)

Zimmerman E. C. (1935) Mangarevan Expedition. Report of Elwood C. Zimmerman, Entomologist. Bernice P. Bishop Museum Bulletin 133, 68–71.

Zimmerman E. C. (1994). ‘Australian Weevils (Coleoptera: Curculionoidea). Volume I – Orthoceri, Anthribidae to Attelabidae. The Primitive Weevils.’ (CSIRO: Melbourne, Australia.)

Zimmerman E. C., Perrault G. H. (1989) Aglycyderidae of the Society Islands (Coleoptera: Curculionoidea). Bishop Museum Occasional Papers 29, 151–173.










Appendix 1.  Characters used in cladistic analysis of Belidae Oxycoryninae
In her preliminary phylogeny of Oxycoryninae, Marvaldi (2005) used 25 morphological characters from the adult, 30 from the larva and three from biology. We have expanded the dataset by 37 additional imaginal and three larval characters (indicated by an asterisk) and also changed the states of some of her characters to accord with the additional taxa and character studies here included.
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Appendix 1a. (continued)
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Appendix 1b. (continued)
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Appendix 2.  New classification of the subfamily Oxycoryninae (family Belidae)
General format as in the catalogue of Alonso-Zarazaga and Lyal (1999) to facilitate comparison and track changes
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