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NOTE TO USERS This reproduction is the best copy available. UMI nm u Ottawa L'Universite canadienne Canada's university urn FACULTE DES ETUDES SUPERIEURES 1=1 FACULTY OF GRADUATE AND ET POSTDOCTORALES U Ottawa POSTDOCTORAL STUDIES L'Universite canadienne Canada's university Adam Conan Algar AUTEUR DE LA THESE / AUTHOR OF THESIS Ph.D. (Biology) GRADE/DEGREE Department of Biology FACULTE, ECOLE, DEPARTEMENT / FACULTY, SCHOOL, DEPARTMENT Climatic Influences on the Evolution and Diversity of Regional Species Assemblages TITRE DE LA THESE / TITLE OF THESIS D. Currie DIRECTEUR (DIRECTRICE) DE LA THESE / THESIS SUPERVISOR J.Kerr CO-DIRECTEUR (CO-DIRECTRICE) DE LA THESE / THESIS CO-SUPERVISOR EXAMINATEURS (EXAMINATRICES) DE LA THESE/THESIS EXAMINERS G. Blouin-Demers R. Kassen A. Gonzalez A. Simons Gary W. Slater Le Doyen de la Faculte des etudes superieures et postdoctorales / Dean of the Faculty of Graduate and Postdoctoral Studies Climatic influences on the evolution and diversity of regional species assemblages Adam C. Algar Thesis submitted to the Faculty of Graduate and Postdoctoral Studies University of Ottawa in partial fulfillment of the requirements for the PhD degree in the Ottawa-Carleton Institute of Biology, These soumise a Faculte des etudes superieures et postdoctorales Universite d'Ottawa en vue de l'obtention du doctorat L'institut de biologie d'Ottawa-Carleton © Adam C. Algar, Ottawa, Canada, 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 OttawaONK1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-61378-8 Our file Notre reference ISBN: 978-0-494-61378-8 NOTICE: AVIS: The author has granted a non L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives 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 le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non support microforme, papier, electronique et/ou commercial purposes, in microform, 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 this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extra its substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 14-1 Canada Acknowledgements Many people, in one way or another, have made contributions to this work. Without their help, this thesis would have remained a loose collection of scattered ideas and half- formed hypotheses. I am indebted to each and every one of them. In particular, I must thank my supervisors, Jeremy Kerr and David Currie. Together, they have offered me guidance when I needed a guide, challenged me when I needed to be challenged, and encouraged me when I needed encouragement. Just as importantly, they allowed me to make mistakes along the way and gave me time to find the solutions. More than anything else, they gave me the freedom to explore any question that I found interesting and for that I am extremely grateful. They are both more than advisors and mentors; they are colleagues and friends. The impact that Tom Nudds has had on my life cannot be understated. He was my first mentor and, much to the frustration of many I'm sure, first introduced me to the philosophy of science. But more than that, he inspired me to be passionate about the scientific endeavour. Without Tom's influence I would not be pursuing a career in science. In addition to Tom, Darren Sleep also helped shape my earliest scientific years by making time for a wide-eyed undergrad despite the pressures of his own PhD; I can only hope I have provided one-tenth of the help to the undergrads I have since encountered. My advisory committee, Gabriel Blouin-Demers and Andrew Simons have greatly improved my research by having an uncanny ability to cut to the quick of an issue and to ask the right questions. Rees Kassen and Howard Rundle have also been sources of advice and both have been successful gadflies for evolution in the ear of a young macroecologist. ii Over the last five years, I have seen many lab-mates and colleagues come and go and all have taught me something. In particular Peter, April, Chris, and Lana made the first months of grad school enjoyable. Tony acted as an advisor and a bottomless pool of knowledge over many a breakfast. Justin, undoubtedly one of the brightest scientists I have ever met, taught me more about evolution than anyone else and was always up for a pint. Matt made long rides less lonely and with no one else could I have sat and discussed the applications of quantitative genetics to community ecology while watching suffering and glory on the roads of France. Nora spent hours searching the literature for data on hylid frogs - and, to her credit, did not complain once. Julie and Rachelle translated the abstract; any awkwardness in the French is not theirs, but is certainly due to the English material with which they had to work. Special thanks go to Heather, Erin and Rachelle. Heather, in addition to being a friend, has been, and hopefully will continue to be, a much appreciated collaborator and someone with whom to share my frustrations. Erin, more than anyone else, reminded me that the numbers in my spreadsheets represent real living things. Plus, she was willing to get up at 5 am and drive all over Ottawa searching for birds. Rachelle, who has been there almost from the beginning, your contributions are too many to count. I couldn't have asked for a better lab-mate or friend to sit beside every day for 5 years. NSERC provided much of the funding for my research. I can only hope that they support fair and equitable funding for the sciences in Canada in the future, despite pressure from current or future governments that do not understand or value what we do. My family's contribution goes far beyond the last five years. My mother was the first to teach me to enjoy and value learning. My father was the first to show me the wonder in 111 the diversity of living things. And my brother was always there when I needed him. Also, Marc and Christine have provided continual support. Gordon has been a source of much joy - much to the disappointment of a certain mother and mother-in-law who were hoping for human grandchildren. To quote Stanley Coren (MacLean's Dec. 2008) on Cairn Terriers: "When you spend your whole life in a research laboratory or at a keyboard writing, having a dog that makes you laugh is better for stress reduction than a psychotherapist." Without question, the person who has offered the most support and help along the way has been my beautiful wife, Leslie. She has held me up when life has threatened to push me down, and picked me up when it succeeded. She has shared in every joy and has been the source of most of them. She has sacrificed more than anyone can imagine and through it all, she has believed in me and my work, even when I have not. To do science is to ask questions - but no matter which questions I ask in the future, I have already asked the most important one. And Leslie answered yes. Leslie, throughout my university career, many people - Tom, Darren, Jeremy, David - have taught me to love reason. But you gave me a reason to love. I dedicate this thesis to you. IV Abstract It has long been recognized that climate can influence the diversity and dynamics of communities and regional assemblages. Within this thesis, I ask three questions: 1) which processes are most important in mediating climate-species richness relationships; 2) are predictions of spatial climate-richness models temporally consistent, and 3) is local community structure determined primarily by regional or local processes. Metabolic theory proposes that climate-richness relationships arise due to the temperature dependence of metabolic rate. I tested the theory's predictions for six taxa in North America. Contrary to the theory's predictions, temperature-richness relationships were curvilinear and their slopes deviated from the predicted value. This suggests that the mechanism proposed by metabolic theory does not underlie climate-richness relationships. If climate determines species richness, then climate should predict how species richness will change over time. To test this, I compared alternative methods (regression and niche modelling) of forecasting shifts in species richness given global climate change. Models were trained on butterfly richness data from the early 20th century and their predictions were compared to observed changes throughout the 20th century. Overall, regression-based approaches that incorporated spatial autocorrelation outperformed other methods Broad-scale richness gradients could arise from climatic niche conservatism.