Diet, Ecology, and Dental Morphology in Terrestrial Mammals – Silvia Pineda-Munoz – November 2015
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DIET, ECOLOGY, AND DENTAL MORPHOLOGY IN TERRESTRIAL MAMMALS Sílvia Pineda-Munoz, MSc Department of Biological Sciences Macquarie University Sydney, Australia Principal Supervisor: Dr. John Alroy Co-Supervisor(s): Dr Alistair R. Evans Dr Glenn A. Brock This thesis is submitted for the degree of Doctor of Philosophy April 2016 2 To my Little Bean; and her future siblings and cousins Al meu Fessolet; I als seus futurs germans i cosins i ii STATEMENT OF CANDIDATE I certify that the work in this thesis entitled “Diet, ecology and dental morphology in terrestrial mammals” has not previously been submitted for a degree nor has in been submitted as part or requirements for a degree to any other university or institution other than Macquarie University. I also certify that this thesis is an original piece of research and that has been written by me. Any collaboration, help or assistance has been appropriately acknowledged. No Ethics Committee approval was required. Sílvia Pineda-Munoz, MSc MQID: 42622409 iii iv Diet, ecology, and dental morphology in terrestrial mammals – Silvia Pineda-Munoz – November 2015 ABSTRACT Dietary inferences are a key foundation for paleoecological, ecomorphological and macroevolutionary studies because they inform us about the direct relationships between the components of an ecosystem. Thus, the first part of my thesis involved creating a statistically based diet classification based on a literature compilation of stomach content data for 139 terrestrial mammals. I observed that diet is far more complex than a traditional herbivore-omnivore-carnivore classification, which masks important feeding specializations. To solve this problem I proposed a new classification scheme that emphasizes the primary resource in a given diet (Chapter 3). This new classification was then contrasted with body mass (Chapter 4). I observed that there is a specific optimum body mass range for every dietary specialization, with the medium size range mostly composed of frugivorous species that inhabit tropical and subtropical rainforests. Thus, the near absence of medium-sized mammals in open environments can be linked to the decreasing density of fruit trees needed to support a pure frugivorous diet all year round. I then evaluated previous dietary proxies and observed that a relevant time scale needs to be determined before choosing a dietary proxy (Chapter 5). The main goal of my PhD research was to design quantitative and phylogeny-free method to infer the typical diet of each species. I therefore designed Multi-Proxy Dental Morphology Analysis (MPDMA) (Chapter 6). I three dimensionally scanned the dentitions of 138 extant mammals (28 marsupials and 110 placentals) and qualitatively classified their diets. Multiple variables were estimated from the 3D scans (i.e., orientation patch count, slope diversity, and relief index) and multivariate statistical analyses were used to test for discrimination power across dietary specializations (Chapter 7). MPDMA demonstrates significant morphological differences across diets (P < 0.05) and correctly discriminates diet for up to 82% of the specimens in the v dataset. Most remarkably, marsupials and placentals with the same dietary specializations overlap strongly in ecomorphospace, which suggests convergent phenotypic evolution across both clades. vi ACKNOWLEDGEMENTS I would like to thank the Higher Degree Research team at Macquarie University and the Office of Fellowships at the Smithsonian Institution for financial support. Additionally, I would like to thank all of the Paleobiology Database Workshop in Analytical Methods instructors for bringing me the opportunity to learn so much in such a short time. This thesis would have never materialized without the support I received from colleagues and friends from all around the world. Thank you to Kayla Friedman and Malcolm Morgan of the Centre for Sustainable Development, University of Cambridge, UK for producing the Microsoft Word thesis template used to produce this document. To my advisors: I would first like to thank John Alroy for believing in me when I first wrote you more than 4 years ago. For being available to chat any time I knocked your door. For teaching me how to approach science and for your invaluable advice during the whole journey of my Ph.D. For all the constructive comments on my manuscripts and for challenging me to keep improving my English skills. For saying “you go and figure it out” when I asked you how to write in R. For helping me when I first arrived in Australia. I have told you this so many times, but you will always be my advisor. I am also incredibly grateful to my co-advisor Alistair Evans. For opening the door to your lab when I first asked for your help, and teaching me how to think out of the box. vii For being patient with my curiosity and brainstorming and making me feel like home every time I visited Melbourne. For the long conversations about science and life. John and Alistair: I can’t think of a better duo of advisors, and I will always remember the time you both gave me advice about my Romer Session presentation at SVP in 2015. Siting next to each other, commenting on every single slide. That was the best criticism I have ever received. Thanks Glenn Brock. Although my research was so different from yours, you were always there for support and advice. I will always think of our lab as another home to me. My thesis was greatly improved by the opportunity Kay Behrensmeyer and Kate Lyons created for me by awarding me a pre-doc Research Fellowship at the National Museum of Natural History Smithsonian Institution. Thank you Kay for introducing me to so many researchers and believing in my work. For including me to your family plans for all the holidays – I can’t wait to run another race together! Thank you Kate for the long conversations and for making me see the importance of my work. But mainly, for trusting me and offering me such a rewarding post-doctoral position with the ETE program. During my Ph.D. program I was incredibly lucky to work in three amazing labs: The Paleobiology lab in Macquarie University. Thanks Graeme, Sarah, Bryony, Luke, Marissa, Sarah, Patrick, David, James, Julieta, Nick, Christian, Gabrielle and Matt. For the lab meetings and the paper reading clubs. You inspired me to keep learning about new topics every day. Special thanks to the Genes to Geosciences team. The Evans lab at Monash University. Thanks Matt, David, Travis, Angi, Lap, Alana and Kathleen. It was wonderful to hang out and discuss science. viii The National Museum of Natural History Smithsonian Institution. Thanks Matt, Andrew, Aniko, Advait, Amelia, Carlos, Andrew, Danielle, Laura, Arden, Scott, Rich, René, Thomas, and everybody who helped me during my stay at a wonderful institution. To my master's thesis advisor Isaac Casanovas-Vilar. I will soon lose my Padawan status, Mestre. And Mikael Fortelius, the first person in palaeontology to give me an opportunity. To my lab mates at the Institut Català de Paleontologia Miquel Crusafont. Special thanks to Daniel de Miguel, who taught me how to mould and cast. To my dear Maria: I miss our coffee breaks. Fortu, you’ve inspired me so much with your missal. Albert, your support has been crucial in so many ways. David, Marta, Miriam, Gretel, Salvador, Alba, Novella. To my friends at the University of Helsinki: Juha, Allu, Ellodie and Pierre. To my family: Thank you Mama. For encouraging me to give my best every day, and being there to share my laughs and wipe my tears. For our late night conversations and for picking up the phone at 3 a.m. pretending you were awake. For keeping me in touch with my most human side, for keeping my secrets and being my best friend in so many ways. Thank you Ferran (my dad). You used to tell me: “Aim for the sun, and you’ll reach the moon”. One of the last times I saw you, you told me: “I told you to aim for the sun so you could reach the moon; but you’ve gone beyond the sun already”. You made me so happy. But also thank you for the hikes in the mountains, for teaching me how to love nature. Thank you Tata-capu (my sister). For being there to cheer me up when I needed it the most, for being proud of me despite our having such different lives. But also, for giving me what I love the most in this world, my little bean, my niece Èlia, to whom I dedicate this thesis. ix Thank you to my grandmas and grandpas, uncles and aunties, and cousins. You are always there to support me although you sometimes struggle to understand my life as a scientist. I miss you tito Jose. To all my friends: Who have always been there to cheer me up and share the good and bad times. Thank you Zoraida, for being there to read my messages from Catalunya, the Czech Republic or Galicia. Julieta, for becoming my “sister” overseas and for our adventures together that have just started. Laura, for you becoming someone so important in my life in Australia. Peri, for so many important days together, for being my first climbing partner. Aniko, for your incredible friendship. Leo, for belaying me on the wall and in my personal life. Yuri, for mutual inspiration. Nick, the best lab-mate I can think of. Graeme, for dealing so well with all my quirkiness. Pere and Anna, you were so much like a little piece of home overseas. Thanks to my advisors’ families: Heather, Adi and Linus; Gudrun, Luke, David and Zoe; Bill, Sarah and Kristina; Kieran and Pete. For making me feel like home every time I visited.