UNIVERSITY of CALIFORNIA Los Angeles Carnivory
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UNIVERSITY OF CALIFORNIA Los Angeles Carnivory in the Oligo-Miocene: Resource Specialization, Competition, and Coexistence Among North American Fossil Canids A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Biology by Mairin Francesca Aragones Balisi 2018 © Copyright by Mairin Francesca Aragones Balisi 2018 ABSTRACT OF THE DISSERTATION Carnivory in the Oligo-Miocene: Resource Specialization, Competition, and Coexistence Among North American Fossil Canids by Mairin Francesca Aragones Balisi Doctor of Philosophy in Biology University of California, Los Angeles 2018 Professor Blaire Van Valkenburgh, Chair Competition likely shaped the evolution of the mammalian family Canidae (dogs and their ancestors; order Carnivora). From their origin 40 million years ago (Ma), early canids lived alongside potential competitors that may have monopolized the large-carnivore niche—such as bears, bear-dogs, and nimravid saber-toothed cats—before becoming large and hypercarnivorous (diet of >70% meat) later in their history. This ecomorphological context, along with high phylogenetic resolution and an outstanding fossil record, makes North American fossil canids an ideal system for investigating how biotic interactions (e.g. competition) and ecological specialization (e.g. hypercarnivory) might influence clade evolution. Chapter 1 investigates whether ecological generalization enables species to have longer durations and broader geographic range. I developed and applied a carnivory index to 100+ ii species, processing 3708 occurrences through a new duration-estimation method accounting for varying fossil preservation. A non-linear relationship between duration and carnivory emerged: both hyper- and hypocarnivores have shorter durations than mesocarnivores. Chapter 2 tests the cost of hypercarnivory, quantified as elevated extinction risk. Large hypercarnivorous canids experienced extinction rates on par with other canids. However, in each canid subfamily, extinction rates rose after the first occurrence of large hypercarnivores, supporting the idea of hypercarnivory as a “macroevolutionary ratchet” for individual clades. Large hypercarnivores originated just over 10 Ma after the origin of Canidae, exhibiting constant diversification rates and peaking in richness around 12 Ma. Relationships emerge between diversification rates of canid subfamilies and temperature, suggesting future hypotheses to be investigated. Chapter 3 quantifies functional morphology in 114 fossil canids and non-canid competitors from 40 to 15 Ma, testing for ecological congruence between continental and regional scales and whether ecomorphological disparity reflects taxonomic diversity. Disparity appears inversely related to diversity. All four geographic regions examined preserved a narrower range of ecomorphology than at the continental scale. These results suggest that partitioning dietary as well as geographic resources, despite constraints to size, permitted canids to minimize competition and diversify. These macroecological and macroevolutionary studies of early canids and their potential competitors improve understanding of the resilience of the carnivore niche over long timescales, including periods of accelerated global change. iii The dissertation of Mairin Francesca Aragones Balisi is approved. Michael Edward Alfaro Paul Jeffrey Brantingham Blaire Van Valkenburgh, Committee Chair University of California, Los Angeles 2018 iv I dedicated my master’s thesis to the memory of my mother, Fe Aragones Balisi, who gave me my first biology lesson, and to the memory of my grandfather, Benjamin Aragones, who left me his National Geographic collection. I now dedicate my Ph.D. dissertation to my father, Mario Luczon Balisi, and my brother, John Paul Aragones Balisi, who have supported me in the pursuit of this degree even when memory faltered. v TABLE OF CONTENTS LIST OF TABLES .................................................................................................................. viii LIST OF FIGURES ................................................................................................................... ix ACKNOWLEDGEMENTS ........................................................................................................ x VITA ....................................................................................................................................... xiv I. DIETARY SPECIALIZATION IS LINKED TO REDUCED SPECIES DURATIONS IN NORTH AMERICAN FOSSIL CANIDS ............................................................................... 1 Abstract................................................................................................................................. 2 Background .......................................................................................................................... 3 Methods ................................................................................................................................ 7 Results ................................................................................................................................ 14 Discussion .......................................................................................................................... 18 Conclusion.......................................................................................................................... 23 Tables and Figures ............................................................................................................ 24 Supplementary Information ............................................................................................. 31 II. LARGE-BODIED HYPERCARNIVORY AS CONVEYOR BELT TO EXTINCTION? DIVERSIFICATION DYNAMICS OF NORTH AMERICAN FOSSIL CANIDS ................ 41 Abstract............................................................................................................................... 42 Background ........................................................................................................................ 43 Methods .............................................................................................................................. 46 Results ................................................................................................................................ 51 Discussion .......................................................................................................................... 55 Conclusions ........................................................................................................................ 59 Tables and Figures ............................................................................................................ 61 Supplementary Information ............................................................................................. 67 III. FUNCTIONAL DIVERGENCE IN BODY SIZE AND DIETARY ADAPTATIONS WITHIN NORTH AMERICAN FOSSIL CARNIVORAN COMMUNITIES ....................... 71 Abstract............................................................................................................................... 72 Background ........................................................................................................................ 73 Methods .............................................................................................................................. 77 Results ................................................................................................................................ 80 Discussion .......................................................................................................................... 82 Conclusions ........................................................................................................................ 86 vi Tables and Figures ............................................................................................................ 87 Supplementary Information ............................................................................................. 92 REFERENCES ......................................................................................................................... 93 vii LIST OF TABLES Table 1.1. Summary statistics and measures of phylogenetic signal.......................................... 24 Table 1.2. Quantile regression results at four levels.................................................................... 24 Table 1.3. Results of nested linear regressions........................................................................... 25 Table 2.1. Median posterior estimates of the parameters al and aµ........................................... 61 Table 2.2. Median posterior estimates of the parameters gl and gµ............................................. 62 Table 3.1. Subdivisions of North American Land Mammal Ages.............................................. 87 Table 3.2. Diversity metrics for each region............................................................................... 87 viii LIST OF FIGURES Figure 1.1. First two axes of principal component analysis......................................................... 26 Figure 1.2. Bivariate plots of emergent properties against intrinsic traits.................................... 27 Figure 1.3. Relationship between species duration and specialization........................................ 28 Figure 1.4. Bivariate plots of species duration as a function of the