Copyright by Katherine Hilary Bannar-Martin 2015 The Dissertation Committee for Katherine Hilary Bannar-Martin Certifies that this is the approved version of the following dissertation: Scale and Process: Primate and Non-Primate Mammal Community Composition and Diversity in Madagascar Committee: Rebecca J. Lewis, Supervisor Mathew A. Leibold Denné N. Reed Anthony Di Fiore Mariah E. Hopkins Scale and Process: Primate and Non-Primate Mammal Community Composition and Diversity in Madagascar by Katherine Hilary Bannar-Martin, B.Sc., M.Sc. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy The University of Texas at Austin May 2015 Dedication For my parents, Hilary and Mark. This would not be without you. Thank you for every sacrifice. Acknowledgements I have gotten a lot of results! I know several thousand things that won’t work. - Thomas A. Edison It is a tough proposition to acknowledge and thank all the people that have contributed to this labour of love, time, and brainpower. I would like to thank my advisor, Rebecca Lewis, who I met in 2006 in Ranomafana National Park, Madagascar, for sharing her love of lemurs, fieldwork, Madagascar and Anthropology with me. She has constantly reminded me to see the bigger picture, to be relevant, and to communicate effectively. Thank you to my committee: Rebecca Lewis, Mathew Leibold, Denné Reed, Anthony Di Fiore and Mariah Hopkins. I thank them for saying yes, for reading, for thinking and critiquing. I am honoured to have had them all contribute to this piece of work and honoured to have shared a room with them discussing ideas. I have been continually challenged and continually rewarded by the caliber of excellence that exists within my committee and at the University of Texas. A special thanks to Mathew Leibold for providing me with a home in the Biology department and the privilege to discuss ecological theory with his fantastic lab group. My eternal thanks are also due to my incredible crew of peers. Kim Valenta made my first year in Austin epic, and persuaded me that Canadians in Texas are always charming. Kelsey Ellis, Addison Kemp, and Lina Valencia Rodriquez have been wonderful friends and provided compelling discussions that ran the gamut of physical anthropology, ecology, academic life, alcohol, knitting, and a lot of et cetera. Andrew Barr, Carrie Veilleuz, Amber Heard-Booth, and Rick Smith were fountains of sage advice. Thank you also to Laura Abondano, Jaime Mata-Miguez, Gabrielle Russo, Brett Nachman, Angel Zeininger for sharing the journey. v My thanks are also due to the wonderful Anthropology department at the University of Texas. Liza Shapiro’s ability to question ideas until they were distilled to their simplest form was always appreciated. Chris Kirk’s attention to detail and engaging teaching style have made me a better teacher. Also thank you to Billy O’Leary, Chris McNett, Adriana Dingman, and Rolee Rios, who saved my ‘academic’ life on several occasions, especially when I decided to finish my dissertation from Canada. Thank you to all the people who helped along the way. Patricia Wright, Steig Johnson, and Summer Arrigo-Nelson provided data from Ranomafana National Park. Steve Goodman generously answered my emails and mailed me a pile of books, a.k.a. invaluable documents on Madagascar’s fauna that supplemented my data. Charlene Nielsen provided invaluable GIS and python advice. Jesse Sinclair was my savvy [R] coding guide. Thank you also to Mishka Jada King for help with data entry. Jason Kamilar, Lydia Beaudrot and Kaye Reed, and two anonymous reviewers provided valuable comments on chapter 2. Finally, none of this would have been possible without the support and love of my family and friends at home. My eternal gratitude is owed to my parents for instilling in me a desire for knowledge, the strength to ask questions, and the tenacity to find answers. My mother’s strength, generosity and fearlessness were an inspiration and an honour to have witnessed. My father’s patience has been a pillar; his quiet advice and work ethic encourage me daily to keep going. Sophie, and Emma, my incredibly talented, beautiful and wise sisters, consistently reminded me of which way is up. Thank you to my Grandma, Helen Cooper, and my godmother, Di Jones, who made my first journey to Madagascar possible. Thank you to Jenny Linton, Lindsay Horlor and Pia Nagpal for their friendship and unwavering support. My final thanks go to the love of my life, Neil McCallum. He was the voice of realism, hope and love that got me through the last go of it. vi This research was supported by a NSERC Postgraduate Doctoral Scholarship, an Explorer’s Club Research Grant, an AAUW Doctoral Scholarship, and a Rhonda L. Andrews Memorial Award. vii Scale and Process: Primate and Non-Primate Mammal Community Composition and Diversity in Madagascar Katherine Hilary Bannar-Martin, Ph.D. The University of Texas at Austin, 2015 Supervisor: Rebecca J. Lewis The study of community assembly, or the processes that shape the occurrence of species in an ecological community, is a fundamental area of inquiry in ecology. Patterns in community composition and diversity are attributed to the combined operation of deterministic (e.g., environmental sorting), stochastic (e.g., dispersal limitation), and biogeographic (e.g., dispersal barriers) processes. Environmental sorting results in communities composed of species that are ecologically adapted to their environment. Dispersal limitation results in communities shaped by the dispersal distance between sites. Biogeographic dispersal barriers prevent species dispersal between sites, and community membership is dependent upon site isolation. Community assembly is also dependent upon diversity type (taxonomic, functional, or phylogenetic) and spatial scale. I investigated the processes shaping the diversity of primate and nonvolant mammal communities using taxonomic, functional and phylogenetic diversity measures and a spatially explicit modelling approach. I described mammal diversity patterns at ecoregional, regional, and inter-regional scales within and across Madagascar and Australia. I tested the relationship of mammal community diversity to environmental, spatial, and biogeographic variables, indicating deterministic, stochastic, and viii biogeographic processes, in Madagascar and Australia. First, I found that arboreal mammal communities in Madagascar were more dispersal-limited than terrestrial mammal communities. Second, a combination of environmental sorting and dispersal limitation best explained primate taxonomic and functional diversity. Third, I tested for convergent diversity and assembly patterns in Madagascar and Australia, due to similar biogeographic and evolutionary histories, and found non-convergent patterns. Overall, biogeographic dispersal barriers were weak predictors of mammal diversity in Madagascar and Australia. Phylogenetic and functional diversity measures were weakly correlated, and phylogenetic diversity provided models with weak explanatory power. Environmental and spatial variables indicating the combined operation of environmental sorting and dispersal limitation variably shaped the taxonomic and functional diversity of mammal communities in Madagascar and Australia. Mammal community diversity was regionally specific, shaped by the unique historical and landscape components of each region, including ecoregional effects and the extinction of sympatric species. Macroscale studies of diversity should carefully investigate the influence of spatial scale and regional factors that can result in varied assembly patterns and unique ecological communities, such as those present for the nonvolant mammals of Madagascar and Australia. ix Table of Contents List of Tables .........................................................................................................xv List of Figures ...................................................................................................... xix Chapter 1: Introduction – Mammal Community Assembly ....................................1 Introduction .....................................................................................................1 Mammal Community Assembly .....................................................................5 Community Assembly Processes ....................................................................7 Deterministic processes .........................................................................8 Stochastic processes .............................................................................10 Biogeographic Processes .....................................................................13 Diversity and Community Assembly ............................................................16 Taxonomic Diversity ...........................................................................17 Phylogenetic Diversity .........................................................................18 Functional Diversity.............................................................................21 Community Assembly Across Spatial Scales ...............................................23 Dissertation Objectives .................................................................................26 Dissertation Study Areas...............................................................................27 Madagascar ..........................................................................................27