CLASSIFICATION OF AMPHIZOIDAE (COLEOPTERA) BASED ON IMMATURE STAGE CIiARACTERS A Thesis Submitted to the Facdty of Graduate Studies The University of Manitoba In Partial Fulfilment of the Requirements for the Degree of Master of Science in The Department of Entomology Winnipeg, Manitoba %opyright by Weiping Xie, February 2000 National Library Bibliothèque nationale 1*1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. rue Wellmgton OriawaON K1AW Octawa ON UlA ON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence aiiowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fkom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author' s ou autrement reproduits sans son permission. autorisation. THE UNIVERSITY OF MAlYITOBA FACULTY OF GRADUATE STUDIES ****t COPYRIGHT PERMISSION PAGE Classification of Amphizoidae (Coleoptera) Based on Immature Stage Characters Weiping Xie A Thesis/Practicum submitted to the Faculty of Graduate Studies of The University of Manitoba in partial fulflhnent of the requirements of the degree of Master of Science WEIPING XIE O 2000 Permission has been granted to the Librrry of The University of Manitoba to lend or seU copies of this thesis/practicum, to the National Library of Canada to microfilm this thesis/practicum and to lend or seU copies of the film, and to Dissertations Abstracts International to publish an abstract of this thesisfpracticum. The author resewes other publication rights, and neitber this thesis/practicum nor extensive extracts from it may be printed or otherwise reproduced without the author's written permission. 1 hereby declare that I am the sole author of this thesis. 1 authorize the University of Manitoba to lend this thesis to other institutions or individuals for the purpose of schoiarly research. Weiping Xie 1 funher authorize the University of Manitoba to reproduce this thesis by photocopying or by other means, in total or in pan, at the requcst of other institutions or individuals for the purpose of scholarly research Weiping Xie The University of Manitoba requires the signatures of al1 persons using or photocopying this thesis. Please sign below, and give address and date. WE~PMGXE, Department of Entomology, University of Manitoba, April, 1999. Classification of Amphizoidae (Coleoptera) based on immature stage characters. Major Advisor: Dr. RE-ROUGHLEY. .4mphizoidae is a family of the Suborder Adephaga of Coleoptera. There is only the single genus Ampltizoa. of which there are six species known in the world. There are three species in western North America: Amphboa insolens LECONTE 1 853, Amplzizoa leconrei MATII~EWS1872, and Amphizoa striata VAN DYKE1927, and three in China: Amphizoa davidi LUCAS 1882, Amphizoa sinica YU AND STORK 1991, and Amphizoa smeranai ROUGHLEY,XIE AND YU (in press). In this paper, the morphology, geographical distribution and habitat of each species in its larval stages are described, and certain characteristics of fom and structure are illustrated. Keys for identification of amphizoid Iarvae of hown species are presented for the first time. This study is based mainly on chaetotaxal analysis, a method of studying the serae and pores of immature stages. The chaetotaxal patterns of first instar larvae of five species of Amphizoa, and one species of Hygrobiidae (referred to as the out-goup) are described. The phylogenies of Hydradephaga and Amphizoidae were reconstnicted from these data. The origin of amphizoids is hypothesized to have occurred somewhere on Pacifica. a set of supposed continental masses, in the latc Permian or early Triassic. These areas would havc been highland areas with glaciations and ice cover during that tirne. As early as the Permian times, the amphizoid fauna originated within and adapted to high altitude, periglacial conditions, and cold lotic habitats. Then the arnphizoids drifted northward on thesc land masses of Pacifica, and split up into the Asian and North American branches. The Asian branch reached its present position, Sichuan Province, probably with the Yan-e block. The Nonh American lineage also speciared durkg the drifiing. One of these reached its location of Jilin Province with the Sino-Korean block, and others went to western Noah America with some Pacifica masses kom late Jurassic and early Cretaceous to the early Teniary. The present pattern of dismbution in geographical variation abo reflects a history of the development of the Himalayan Mountains and RocLy Mountain ranges, and of the repeated episodes of isolation and dispersal of these species during the various Pleistocme glaciations. Key words: Coleoptera; Adepbaga; Amphizoidae; immature stages; chaetotaxy; origin. 1 wish to express my sincere appreciation to my advisor, Dr. ROBERTE. ROUGHLEY, for his patience, continuous support and assistance in the completion of this research and preparation of this dissertation. I also thank the other members of my committee, Dr. TERRYD. GXLLOWAY,Deparnent of Entomology and Dr. BRUCEA. FORD.Department of Botany. University of Manitoba, for rheir conmburions to various aspects of my studies. My appreciation extends to the Department of Entomology of University of Manitoba for supporting this project and for providing the facilities necessary to cany out the research. 1 thank Prof. YU PEMJ,Institute of Zoology, Chinese Acaderny of Sciences, Beijing, for kindly providing advice and assistance during various phases of my research. I am gratefU1 to the Conservation Branch of Sichuan Provincial Forest Ministry (China) for assistance, Mr. HE HONGCHIand other colleagues of Feng Tongzai Nature Reserve for collecting the first instar larvae of A. davidi for me, and 1 also thank Mrs. LM FENGQM,Changbai Mountain Nature Reserve, for her assistance in the field observation of A. sinica. I thank Prof. LU BAOLIAN,Mr. LIU XLAOYAN,and Mrs. GAN YALN, Zoological Instimte, Chinese Academy of Sciences, Beijing for taking the scaming elcctron micro,gaphs. SpeciaJ thanks are exrended to Dr. DARRENPOLLOCK, Mr. DON HENNE, Mr. JASON DIEIILand Ms. WENDYGR~IIAM, Department of Entomology, University of Manitoba, and Mr. DANO'BWEN, Department of Zoology, University of Victoria, British Columbia, for their critical review and corrections to the English of the final manuscript. 1 ~vouldlike to thank Dr. PETERJ. CONEY,Department of Geosciences, University of Arizona, Tucson, Arizona; Dr. ZVI BEN-AVRAHAM,Department of Geophysics and Planetary Sciences, Te1 Aviv University, Israel; Dr. IAN DAUIEL, Institute for Geophysics, University of Texas, Austin, Texas; and Dr. JOHN HILLHOUSE, United States Geological Survey, Men10 Park, California, for providing information and comments on geology. I also thank the California Academy of Sciences, San Francisco, California; Field Museum of Natural History, Chicago, illinois; Depamnent of Entomology, Montana State University, Bozernan, Montana; Institute of Zoology, Chinese Academy of Sciences, Beijing, China; and Dr. A. SMETANA,ECORC, Agriculture and A@-Food Canada, Ottawa, Canada, for kindly providing valuable specimens. Funding for the field and laboratory work leading to this research was provided by R.E. ROUGHLEY'SNSERC (Canada) research grant A0428. vii .. - ABSTRACT ........................ ... ............................................................................................ iv ACKNO WLEDGMENTS..... .... .................................................................................... vi ... TABLE OF CONTENTS ................................................................................................. vtii LIST OF TABLES ............................................................................................................. x LIST OF FIGURES ............................................................................................................ xi INTRODUCTION ............................................................................................................... 1 LITERATURE REVIEW .................................................................................................... 4 Taxonornic History of Amphizoidae ......................................................................... 4 Bionomics of Amphizoa ............................................................................................ 6 Phylogeny of Hydradephaga and Amphizoidae ................... .... ........................... 7 OBJECTIVES ................................................................................................................... 13 MATERML AND METHODS ....................... .. ......................... ..,....................... 13 SY STEMATICS OF AMPHIZOIDAE............................................................................. 19 Generic diagnosis .................................................................................................... 19 Key to identifi various instars of Ampltizoa