Diversity and Ecological Structure of Northern Biting Flies
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Diversity and Ecological Structure in Northern Biting Flies by Patrick Schaefer A thesis submitted in conformity with the requirements for the degree of Masters of Science Ecology and Evolutionary Biology University of Toronto © Copyright by Patrick Schaefer 2014 Diversity and Ecological Structure in Northern Biting Flies Patrick Schaefer Masters of Science Degree Ecology and Evolutionary Biology University of Toronto 2014 Abstract Contemporary climate change disproportionately impacts northern environments. Understanding these impacts requires knowledge of the ecological structure (e.g. diversity, abundance and distribution) of resident species. Biting flies are among the best-known groups of insects in northern Canada and are excellent sentinels for environmental change. The goals of this thesis are to determine how climate and geographical history affect the ecological structure of black flies and mosquitoes. Assessment of large-scale macroecological patterns revealed contrasts between Arctic and southern assemblages, while highlighting the importance of geographic history in structuring assemblages. Finer-scale sampling and analysis were used to assess the impacts of contemporary climate change on species ranges. A critical threshold temperature for the presence of biting species of black flies was found that approximates minimum requirements for flight and blood-feeding. Finally, mitochondrial DNA markers allowed the assessment of cryptic diversity in a widely distributed assemblage of northern black flies of the genus Stegopterna. ii Acknowledgments I would like to express my deepest gratitude to my supervisor, Dr. Douglas C. Currie, whose expertise, understanding, and patience have added considerably to my graduate experience. I appreciate his vast knowledge and skills in many areas, as well as his assistance in writing reports. I would also like to thank the other members of my supervisory committee, Dr. Donald Jackson, and Dr. Alan Baker for the assistance they provided at all levels of the research project. This research would not have been possible without the Northern Biodiversity Program. I acknowledge the contributions of the Northern Biodiversity Programs principle investigators: Dr. Chris Buddle (McGill University), Dr. Terry Wheeler (McGill University/Lyman Museum), Dr. Donna Giberson (Unviersity of Prince Edward Island) and Dr. Douglas C. Currie (University of Toronto/Royal Ontario Museum) for securing initial funding and providing logistical support for the duration of the project. I also acknowledge the collection and processing efforts of other Northern Biodiversity Program members: S. Laboda, K. Sim, L. Timms, A. Solecki, M. Blair and R. Cordero. I am very grateful for assistance provided by Royal Ontario Museum staff from the Entomology Department Brad Hubley and Antonia Guidotti as well as Kristen Choffe from the Laboratory of Molecular Systematics. I also thank Ida Conflitti (University of Toronto), Laura Timms (Royal Ontario Museum/Credit Valley Conservation), Peter Adler (Clemson University), Sally Adamowicz (Guelph University/Biodiversity Institute of Ontario) and Monty Wood (Canadian National Collection of Insects and Arachnids) for their input and guidance at various stages during this research. Finally, I am grateful towards my loving wife Heather for spending many long hours sorting flies with me in the lab and her patience and support in all aspects of my work. This work was supported by a National Science and Engineering Research Council of Canada (NCERC) Strategic Project Grant (Ecological Structure of Northern Arthropods: Adaptation to a Changing Environment) awarded to C. Buddle, T. Wheeler, and D. Currie, its supporting partners and collaborators, a NSERC Discovery Grant to D.C. Currie and a NSERC – Canada Graduate Scholarship and Ontario Graduate Scholarship awarded to P. Schaefer. iii Table of Contents Acknowledgments ......................................................................................................................... ii Table of Contents ......................................................................................................................... iv List of Tables ................................................................................................................................ vi List of Figures .............................................................................................................................. vii Chapter 1 - General Introduction ............................................................................................... 1 1.1 References ........................................................................................................................... 4 Chapter 2 - Environmental and historical determinants of mosquito (Culicidae) and black fly (Simuliidae) assemblage structure in northern Canada ...................................................... 6 2.1 Abstract ............................................................................................................................... 6 2.2 Introduction ......................................................................................................................... 6 2.3 Materials and Methods ........................................................................................................ 9 2.3.1 Biological Data ........................................................................................................... 9 2.3.2 Explanatory Variables .............................................................................................. 10 2.3.3 Analytical Methods ................................................................................................... 11 2.4 Results ............................................................................................................................... 13 2.5 Discussion ......................................................................................................................... 15 2.5.1 Assemblage Structure and the Importance of Predictor Variables .......................... 15 2.5.2 Macrodistributional Patterns .................................................................................... 17 2.5.3 Patterns of Species Richness ..................................................................................... 18 2.6 References ......................................................................................................................... 21 2.7 Tables ................................................................................................................................ 31 2.8 Figures ............................................................................................................................... 36 Chapter 3 - Climate-induced changes in Arctic black fly communities ................................ 45 3.1 Abstract ............................................................................................................................. 45 3.2 Introduction ....................................................................................................................... 46 3.3 Materials and Methods ...................................................................................................... 47 3.3.1 Study Sites ................................................................................................................. 47 iv 3.3.2 Climate and Species Assemblages ............................................................................ 48 3.4 Results ............................................................................................................................... 50 3.4.1 Historical and Contemporary species records ......................................................... 50 3.4.2 Temperature Thresholds for Bloodsucking Species .................................................. 51 3.4.3 Patterns of Species Richness ..................................................................................... 51 3.5 Discussion ......................................................................................................................... 52 3.5.1 Distinguishing Inadequate Baseline Data from Bona Fide Examples of Range Expansion. ............................................................................................................................. 52 3.5.2 Factors Limiting the Distribution of Bloodsucking Black Flies ............................... 54 3.5.3 Species Richness and Community Saturation ........................................................... 55 3.5.4 Implications of a Northward Migration of Bloodsucking Black Flies. ..................... 56 3.6 References ......................................................................................................................... 60 3.7 Tables ................................................................................................................................ 64 3.8 Figures ............................................................................................................................... 68 Chapter 4 - Cryptic diversity in northern Nearctic Stegopterna Enderlein (Diptera: Simuliidae). .................................................................................................................................. 75 4.1 Abstract ............................................................................................................................. 75 4.2 Introduction ......................................................................................................................