TAXONOMY, CLASSIFICATION, RECONSTRUCTED PHYLOGENY, BIOGEOGRAPHY, AND NATURAL HISTORY OF NEARCTIC SPECIES OF BRYC HIU S THOMSON (COLEOPTERA: HALIPLIDAE) By TONYA MOUSSEAU A thesis Submitted to the Faculty of Graduate Studies In Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE Department of Entomology University of Manitoba Winnipeg, Manitoba R3T 2N2 O Tonya Mousseau 2004 THE UNTVERSITY OF MANITOBA FACULTY OF GRADUATE STUDIES ***** COPYRIGHT PERMISSION PAGE TAXONOMY, CLAS SIFICATION, RE C ONSTRUCTED PHYLO GEI\IY' BIOGEOGRAPITY, AND NATURAL HISTORY OF NEARCTIC SPECIES OF' BRYCHIUS THOMSON (COLEOPTERA: HALIPLIDAE) BY TONYA MOUSSEAU A Thesis/Practicum submitted to the Faculty of Graduate Studies of The Univercity of Manitoba in partial fulfillment of the requirements of the degree of MASTER OF SCIENCE TONYA MOUSSEAU @ 2OO4 PermissÍon has been granted to the Library of The University of Manitoba to lend or sell copies of this thesis/practicum, to the National Library of Canada to microfilm this thesis and to lend or sell copies of the film, and to University Microfilm Inc. to publish an abstract of this thesis/practicum. The author reserves other publication rights, and neither this thesis/practicum nor extensive extracts from it may be printed or otherwise reproduced without the author's written permission. Brychius hornit Crotch Brychius hungerþrdi Spangler Brychius pacificus Carr Frontispiece Dedicated to my two loving grandfathers, Bernie Mousseau (Pépère) and William Both (Opa), who passed away during the time of this research. ACKNOWLEDGEMENTS First, f express my gratitude to my advisor, Dr. Robert E. Roughley, for his continuous guidance throughout my M. Sc. program. I thank him for his patience, enthusiasm, advice, and honesty. I also thank the members of my committee: Dr. Terry Galloway, Department of Entomology; Dr. Bruce Ford, Department of Botany; Dr. Brenda Hann, Department of Zoologyl Dr. Nathan Lovejoy, Department of Zoology, for their suggestions and guidance. I thank everyone in the Department of Entomology. Drs. Neil Holliday, Pat MacKay, and Rob Currie provided insightful feedback to the project. Lisa Babey and Dave Holder provided technical support. Fellow graduate students have been wonderful, I especially thank Lisa Capar, Michael Alperyn, David Wade, and Robyn Underwood for their feedback and supPort. Drs. Cheryl Podemski (Freshwater Institute, Department of Fisheries and Oceans, Winnipeg, MB) and Helena Shaverdo (Natural History Museum,'Wien, Austria) have given me inspiration for what I can achieve. Dr. Rex Kenner (Spencer Museum, University of British Columbia, Vancouver, BC) was thoughtful in pointing out the availability of Brychius specimens at various North American museums from which I had not borrowed specimens. Funding for the field and laboratory work necessary for this research was provided by R.E. Roughiey's NSERC (Canada) research grant and a Nature Discovery Grant from the Canadian Museum of Nature. I also thank Don Sutherland, Natural 11r Heritage Info¡mation Centre, Ontario Government of Natural Resources, for providing funding to produce a status report on Brychius hungerþrdi in Ontario. Finally I would like to thank my family and friends for thei¡ understanding, patience, and support through the past several years. 1V TABLE OF CONTENTS DEDICATION LIST LIST oF APPENDICES """""" """""""xrrr ABSTRACT """""'xiv FoREwoRD """"""" """""" xvi CHAPTER I GENERAL INTRODUCTION..... .".........." 1 CHAP|ER II LITERATURE REVIEW............. .'...'.........3 Taxonomic History of Haliplidae.'.....'..... .."""""""3 Taxonomic history of species of Brychi'us ."""""""'5 Nearctic species .'.""""""'5 Palaearctic species...... """'6 Classification of the Hydradephaga....'....... """""""7 Classification of the Haliplidae ...".""""'9 CHAPTER III TAXONOMY, CLASSIFICATION, RECONSTRUCTED PHYLOGENY AND BIOGEOGRAPHY OF NEARCTIC SPECIES OF BRYCHIUS THOMSON (COLEOPTERA: HALIPLIDAE). ....'.'.. """""""""" 16 Abstract """"""" 16 Introduction ............... """""I7 Materials and Methods.............. """""" 18 Specimens Examined """ 18 Pieparation of Specimens...'...'.'... """"""""""'20 Adults....... ..-.'...'...'.....20 Larvae """'2I Chaetotaxy ""'22 Drawings and Photographs........' """23 Quantitátive Analysis Phylogeny """24 Distribution............... """26 Species Treatments """"'27 Taxonomic characters and terms '.""""'27 Adults....... ..........-.......27 Larvae """' 30 Classification............. """"" 31 Genus Brychius Thomson, 1859......... ."""""""31 Diagnosis """' 31 Description ""'32 Adults....... ....-........'...'32 Larvae """'34 Key to the adults of Nearctic species of Brychius ."""""""35 Species Treatments """"'36 BRYCHIUS HORNII CROTCH '..................' 36 Diagnosis. """""""38 Description """""" 38 Adults """""""' 38 Larvae .......'."""41 Distribution............... """""""46 Remarks... ..""""""46 BRYCHIUS HUNGERFORDI SP4NGL8R.............. .....46 Diagnosis ."""""""47 Description """"""48 Adults .....'....""'48 Larvae """""""'51 Distribution............... ." """""' 53 BRYCHIUS PACIFICUS CARR...... .............53 Diagnosis .... "" """' 53 Description -"""""'54 Adults ......."'.""54 Distribution............... .""""""'57 0"""'ifii::":1iï::::::ï:::::: ::: : ::::: ::: : : ::: s8 Females ""'59 OverallTrends...... """59 Phylogeny """""'60 Discussion """""61 Species Classification............ """""'61 Biogeography............. ""'64 Conclusion """"" 68 Acknowledgements.... """""69 CHAIr|ER IV OBSERVATIONS ON THE NATURAL HISTORY OF A POPULATION OF BRYCHIUS HORNII CROTCH, 1873 (COLEOPIERA: HALIPLIDAE) FROM THE VI DUCK MOUNTAINS, MANITOBA ......113 Absrracr... .........113 Introduction............... """" 113 Materials and Methods...'.......... """"" 115 Specimen Collection ""' 115 Habitat DescriPtion """' 115 Rearing """" 116 Adults..'.... """""""" 116 Larvae ..... 116 Pupa......... """"""""117 Results...... """"717 Habitat DescriPtion """'ll7 Laboratory Obiervations .........'... "' 118 Reproduction............. """""""" 118 Pupation """""""""' 118 Life Cycle. """""""" 119 Behaviour. """""""" 119 Discussion """"I20 Habitat .........120 Life Cycle ""I23 4du1t................. """'123 Eggs......... """"""""124 Larva........ """"""""I25 Pupa......... """"""""126 Overwintering strategies...........'.. "'126 Nutrition and-Feeding """In Adults"""' """"""""127 Larvae ....129 Behaviour "" 130 Adults....... ................ 130 Larvae "" 131 conclusion ........131 Acknowledgements.... """"I32 CHAP|ER V GENERAL DISCUSSION........... """""' 138 Future Research """"""I4I Classification............. " """""""141 Surveys "'I41 Habitat. """"'142 Natural HistorY. """'I42 REFERENCES.......... """"""'r4 vll LIST OF TABLES Chapter II Table 1. The history of evolutionary hypotheses of Hydradephaga ...............................L2 Chapter III Table 1. Position of sensillae on legs of larvae of B. hornü Crotch (instar II and III and B. hungerfordi Spangler (instar III) (Coleoptera: Haliplidae) ..............70 Table 2. Character matrix used for phylogenetic analysis of species of BrychÌus Thomson (Coleoptera: Haliplidae)............ ........7I Table 3. Morphometric characters measured on adult specimens of Nearctic species of Brychius Thomson (Coleoptera: Haliplidae) ............. .........73 Chapter IV Table 1. Preliminary life cycle of B. hornii Crotch from the Duck Mountains, Manitoba, based on specimen collection and laboratory observations in in200112002........134 vlll List of Figures Chapter II Figure 1. Examples of phylogenetic hypotheses for members of the Hydradephaga: a) polyphyletic evolution (Beutel L995); b) monophyletic evolution (Rlbera et al. 2002). ..............r4 Figure 2. Cladogram of hypothesized evolution of members of Haliplidae (Beutel and Ruhnau 1990). ..................15 Chapter III Figure 1. Dorsal images of adult Nearctic Brychíus spp. (Coleoptera: Haliplidae): a) B. hornii Crotch; b) B. hungerþrdi Spangler; c) B. pacfficus Carr.... --."....74 Figure 2. Mandibles of adult female Nearctic Brychius spp. (Coleoptera: Haliplidae), ventral view: B. hornii Crotch: a) right, d) left; B. hungerþrdi Spangler: b) right, e) left;,8. pacificus Carr: c) right, f¡ 1eft............ -..-..75 Figure 3. Maxilla of adult Nearctic Brychius spp. (Coleoptera: Haliplidae); left side, ventral view: a) B. hornii Crotch; b) B. hungerþrdi Spangler; c) B. pacíficus Cart .... 76 Figure 4. Labium of adult Nearctic Brychius spp. (Coleoptera: Haliplidae), dorsal view: a) B. hornil Crotch; b) B. hungerþrdt Spangler; c) B. pacfficus Carr..........-...-.-.....77 Figure 5. Dorsal view of elytra of adult Nearctic Brychius spp. (Coleoptera: Haliplidae): a) B. hornü Crotch; b) B. hungerþrdi Spangler; c) B. pacificus Carr.................."..78 Figure 6. Ventral view of elytra of adult Nearctic Brychius spp. (Coleoptera: Haliplidae): a) B. hornil Crotch; b) B. hungerþrdi Spangler; c) B. pacificus Catr..--'..".-.........-.79 Figure 7. Hind wing of adult Nearctic Brychius spp. (Coleoptera: Haliplidae): a) B' hornii Crotch; b) B. hungerþrdi Spangler; c) B. pacíficus Carr.... ..'...."80 Figure 8. Prosternal and metasternal processes of adult Nearctic Brychius spp. (coleoprera: Haliplidae): a) B. hornii crotch; b) B. hungerþrdi spangler;c) B. pacificus Carr......... -........81 Figure 9. Prothoracic leg of adult male Nearctic Brychiu.t spp. (Coleoptera: Haliplidae), right, dorsal: a) B. hornii Crotch; b) B. hungerþrdi Spangler; c) B. pacificus Carr..... IX Figure 10. Mesothoracic Ieg of adult male Nearctic Brychius