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A Genus Level Revision of the Subfamily Cardiophorinae Candeze (Insecta

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A Genus Level Revision of the Subfamily Cardiophorinae Candeze (Insecta A genus level revision of the subfamily Cardiophorinae Candeze (Insecta: Coleoptera: Elateridae) By Hume B. Douglas BSc Trent University, Peterborough ON, 1996 MSc York University, Toronto ON, 2001 A Dissertation Submitted in Partial Fulfillment of the Requirement for the Degree of Doctor of Philosophy At Carleton University © Hume Douglas 2006 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Library and Bibliotheque et Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-18213-0 Our file Notre reference ISBN: 978-0-494-18213-0 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce,Canada de reproduire, publier, archiver, publish, archive, preserve, conserve,sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet,distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non­ sur support microforme, papier, electronique commercial purposes, in microform,et/ou autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in et des droits moraux qui protege cette these. this thesis. Neither the thesis Ni la these ni des extraits substantiels de nor substantial extracts from it celle-ci ne doivent etre imprimes ou autrement may be printed or otherwise reproduits sans son autorisation. reproduced without the author's permission. In compliance with the Canadian Conformement a la loi canadienne Privacy Act some supporting sur la protection de la vie privee, forms may have been removed quelques formulaires secondaires from this thesis. ont ete enleves de cette these. While these forms may be includedBien que ces formulaires in the document page count, aient inclus dans la pagination, their removal does not represent il n'y aura aucun contenu manquant. any loss of content from the thesis. i * i Canada Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Abstract Phytogenies are presented for Clicking Elateroidea and Elateridae: Cardiophorinae. Subfamilies Hypnoidinae Schwarz (or tribe Hypnoidini of Dendrometrinae) and Negastriinae Nakane and Kishii are placed in synonymy with Cardiophorinae. Tribe Nyctorini Semenov-Tian-Shanskij and Pjatakova of Cardiophorinae is synonymised under Cardiophorini. All 35 genera of higher Cardiophorinae are keyed and their genus level classification is revised. The genus Exoeolus Broun is transferred from Cardiophorinae to Hemiopinae; fossil genus Crioraphes Iablokoff-Khnzorian is transferred to Elaterinae; fossil genera, Pseudocardiophorites Dolin, and Protocardiophorus Dolin are transferred to clicking Elateroidea ihsertae sedis. The genera Lesnelater Fleutiaux (Physodactylinae, syn.nov of Pachyelater Lesne, Dendrometrinae), Margogastrius Schwarz (Physodactylinae), Pachyelater, Teslasena Fleutiaux (syn.nov of Globothorax Fleutiaux) and species Negastrius americanus (Horn) (as type of Floridelater gen.nov) are transferred to Cardiophorinae. Other new genera include Austrocardiophorus (type species = Cardiophorus humeralis Fairmaire and Germain), Chileaphricus gen.nov. (type species = Aphricus chilensis Fleutiaux). Paradicronychus Dolin and Grnjeva is found to be nomen nudem and not separate from Cardiophorus Eschscholtz . The genus Patriciella Van Zwaluwenburg is synonymised under Aphricus LeConte. The subgenera Diocarphus Fleutiaux (of Phorocardius), Metacardiophorus Guijeva (of Cardiophorus) and Zygocardiophorus Iablokoff-Khnzorian and Mardjanian (of Cardiophorus) are raised to Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. genus rank. Generic definitions of the genera Aptopus Eschscholtz, Cardiophorus Eschscholtz and Paracardiophorus Schwarz are restricted, and those of Blaiseus Fleutiaux, Cardiodontulus Van Zwaluwenburg and Horistonotus Canddze are expanded. Coptostethus Wollaston was found to belong at subgenus rank and Dicronychus Brulle and Platynychus Motschulsky at genus rank. Species Lesnelater madagascariensis Fleutiaux is synonymised under Pachyelater madagascariensis (Lesne). The hypothesis of a Jurassic origin of higher Cardiophorinae is supported by phylogenetic and biogeographic evidence, despite finding that previously published fossil evidence was invalid. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. iv Nomenclatural Disclaimer Although this dissertation contains phrases such as “new species,” “new synonymy,” and “new genus” (to name a few), it is not issued for the public and permanent scientific record or for the purposes of zoological nomenclature. A similar form of this work will soon be published in a scientific journal. That upcoming version will validate all of the nomenclatural acts seen within. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. v Acknowledgements I give thanks to Stewart Peck, Carleton University, for his encouragement, suggestions, and support during the lab, museum and field portions of this study. Thanks to R. Anderson (CMNC), S. Aranda (IMLA), S. Ashe (SEMC), P. Bouchard (CNCI), Y. Bousquet (CNCI), M. Barclay (NHM), R. Brett (CASC), R. Brooks (SEMC), R. Cave (EAPZ), S. Casari (MZSP), M. Caterino (SBMN), D. Drugmand (ISNB), Z. Falin (SEMC), M. Fitton (LSUK), C. Girard (MNHN), J. Grixti, C. von Hayek (NHM), L. Herman (AMNH), E. Hoebeke (CUIC), J. Holland, D. Kavanaugh (CASC), M. Kerley (NHM), A. Konstantinov (USNM), B. Korotyaev (ZMAS), S. Laplante (CNCI), R. Leschen (NZAC), P. Madaire (CNCI), S. Marshall (DEBU), H. Mendell (NHM), J. Mertlik, N. Nikitsky (ZMUM), J. Nunn (JNIC), L. Packer, G. Parsons (MSUC), P. Perkins (MCZC), G. Platia, G. Prinsloo (SANC), A. Rasnitsyn (PIN), E. Riley (TAMU), A. Rueda (EAPZ), A. Samuelson (BPBM), C. Scholtz (UPSA), D. Sikes (BDUC), P. Skelley (FSCA), C. Sole (UPSA), F. Stehr (MSUC), M. Thomas (FSCA), R. Tumbow (RHTC), M. Uhlig (ZMHB), J. Vandenberg (USNM), L. Vilhelmsen (ZMUC), A. Zayed, L. Zerche (DEIC), for specimen loans and help finding literature. I thank R. Anderson, E. Becker, J. Cook, P. Johnson, Jarmila Kukalova-Peck, L. Packer, D. Sikes, J. Skevington for systematics advice. Thanks to A. Drojjine, R. Tittler, J. Kukalova-Peck for translation help. I thank F. Chapleau, M. Smith for acting as advisory committee members. I also thank C. Elwood and B. Wilmore for acting as external examiners. I thank A.B.T. Smith for acting as external examiner and for his helpful suggestions. Thanks to fellow Carleton Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. vi University students B. Frei, N. Haghighi, H. Mirrashed, S. Mautner, M. Kotierk at Carleton University for help and guidance in the molecular laboratory; T. Sherratt and A. Rashed for use of their computer facilities. Financial support for this study and publication costs were provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to S. Peck, and an NSERC postgraduate scholarship, two years of Ontario Graduate Scholarships and a Scholarship from the Canadian Museum of Nature to H. Douglas. Special emphatic thanks to my wife Daphne Uras, for her proofreading, continual general assistance, and support. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. vii Table of Contents Chapter 1 - INTRODUCTION TO THE SUBFAMILY CARDIOPHORINAE 1.1 Introduction to the family Elateridae 1.1.1 Distribution, biology .................................................................................1 1.1.2 Taxonomic background and current status .......................................... 2 1.1.3 Previous phylogenetic research ............................................................. 5 1.1.4 Phylogeny: outgroup of Elateridae ...........................................................6 1.2 Introduction to the subfamily Cardiophorinae 1.2.1 Taxonomic background and current status ............................................ 7 1.2.2 Distribution, biology ............................................................................... 9 1.2.3 Phylogeny: outgroup of Cardiophorinae ................................................10 1.2.4 Phylogeny: monophyly of Cardiophorinae ............................................10 1.2.5 Phylogeny: within Cardiophorinae ........................................................ 12 1.3 Hypotheses Tested 1.3.1 Subdivisions within Elateridae: the position and membership of Cardiophorinae ............................................................................................... 13 1.3.2 Subdivisions within Cardiophorinae ......................................................14 1.3.3 Age of Cardiophorinae .......................................................................... 14 1.4 Research Objectives .......................................................................................... 16 Reproduced with permission of the
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