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Adaptation and Exaptation in the Evolution of the Upper Molar Talon in Microbats

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Adaptation and Exaptation in the Evolution of the Upper Molar Talon in Microbats Adaptation and Exaptation in the Evolution of the Upper Molar Talon in Microbats (Suborder Microchiroptera) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Sarah Colleen Gutzwiller Graduate Program in Evolution, Ecology and Organismal Biology The Ohio State University 2015 Dissertation Committee: John P. Hunter, Advisor Debra Guatelli-Steinberg W. Scott McGraw Ian M. Hamilton Copyright by Sarah Colleen Gutzwiller 2015 Abstract The talon, the distolingual extension to the upper tribosphenic molar, has evolved convergently many times in therian mammals. In the form of the hypocone cusp, the talon has been identified as a key innovation allowing for the evolution of derived herbivory. However, the adaptive and exaptive significance of the talon prior to it being coopted in derived herbivores is not as well understood. This dissertation explores the potential function(s) of the talon, as it originated as a novel structure (small cingulum or cusp) and expanded into the diversity of shapes and sizes that we see today. This research uses extant microbats (suborder Microchiroptera) as a case study for therian talon evolution, due to the group’s diversity in molar shape and diet. The first study examines the role of the talon in the balance of crushing and shearing performed by the tribosphenic molar during food breakdown. This project tests the hypothesis that the talon increases crushing function of the molar. Using 3- dimensional computer renderings attained from microcomputed tomography scans of the upper first molars of 26 microbat species, crushing function was estimated using Relief Index. Relief Index quantifies the relationship between occlusal surface area and projected area, resulting in an estimate of the overall occlusal topography and crushing ability. Talons across all dietary (frugivore, insectivore, etc.) and hardness (hard or soft ii food) specialists were found to increase the crushing function performed by the tribosphenic molar, supporting the hypothesis. The crushing function of the talon may provide a benefit to a frugivorous mammal that relies greatly on crushing for the breakdown of fruit pulp. However, the benefit of a crushing-dominated talon to other dietary groups is not yet fully understood. The second study examines talon size evolution, primarily focusing on the role of the talon in mammalian body size evolution. This research tests the hypothesis that the talon functions to increase occlusal area in order to meet the metabolic demands of larger body size. Using a scaling approach that regressed projected and surface area datasets with body mass estimates attained from the literature, the results suggest that in many cases, talon size scales with body size. This relationship lends support to the hypothesis that increasing talon size helps to meet the occlusal requirements of a larger body size. The effects of dietary group and body mass range are discussed. The third study explores the role of the talon in the maintenance of tooth strength, testing the hypothesis that the talon functions in decreasing the likelihood of enamel cracking. Using a computer simulation technique called Finite Element Analysis, I measured the amplitude and distribution of maximum stress (an estimate of likelihood of tooth cracking) in theoretical and extant species models with a variety of talon morphologies. These analyses were performed for both hard food and soft food chewing simulations. Results suggest that the presence of the talon variably affects tooth strength, depending on talon size and simulation type. The talon provides the most benefit to tooth iii strength during the chewing of soft food, supporting the hypothesis that the talon may function in the maintenance of tooth strength. These studies suggest that both food breakdown and tooth strength functions are plausible for the talon, but depend on talon size, in addition to species’ dietary group and body mass. In the context of therian molar evolution, the convergent origins and elaborations of the talon likely represents a series of adaptations and exaptations reflecting changes in diet and body size. iv Dedication This dissertation is dedicated to my husband, Daniel Hill. Thank you for your never-ending support and love along the way. v Acknowledgments I would like to thank my advisor, John Hunter, for his patience, support, and general good nature throughout my PhD. His guidance and insight helped me through many a “crisis”, to ultimately be able to finish this dissertation. Thank you to my committee for always being happy to offer your time and feedback. Thank you to the Department of Evolution, Ecology, and Organismal Biology and to the College of Arts and Sciences. Thank you to my fellow graduate students for your intellectual support, not-so-intellectual good laughs, and general friendship. Thank you to David Gutzwiller for your advice and willingness to jump in and save the day. Thank you to Zachary Weisberg for help with data collection. Specimens were kindly provided by Angelika Nelson, curator of The Ohio State University, Museum of Biological Diversity; Cody Thompson, collections manager of the University of Michigan, Museum of Zoology; and Sue McLaren, collections manager of the Carnegie Museum of Natural History. Thank you to the staff of the Wright Center for Biomedical Imaging at The Ohio State University, with special consideration to Michelle Williams, who microCT scanned my specimens. vi Vita 2006................................................................Walnut Hills High School 2010................................................................B.S. Biology, Ohio University 2010 to present ..............................................Graduate Teaching Associate, Department of Evolution, Ecology, and Organismal Biology, The Ohio State University Publications Gutzwiller, S., O’Connor, P., and Su, A. 2013. Postcranial pneumaticity and bone structure in two clades of neognath birds. The Anatomical Record 296:867–876. Fields of Study Major Field: Evolution, Ecology and Organismal Biology vii Table of Contents Abstract ............................................................................................................................... ii Dedication ........................................................................................................................... v Acknowledgments.............................................................................................................. vi Vita .................................................................................................................................... vii List of Tables ..................................................................................................................... xi List of Figures ................................................................................................................... xii Chapter 1. Introduction: The Upper Molar Talon ............................................................... 1 Objectives ........................................................................................................................ 6 Figures ............................................................................................................................. 7 Literature Cited ............................................................................................................. 11 Chapter 2. Occlusal Relief and Function of the Upper Molar Talon ................................ 14 Introduction ................................................................................................................... 14 Materials and Methods .................................................................................................. 18 Results ........................................................................................................................... 24 Discussion ..................................................................................................................... 26 viii Figures ........................................................................................................................... 31 Tables ............................................................................................................................ 35 Literature Cited ............................................................................................................. 38 Chapter 3. Upper Molar Talon Size Evolution ................................................................. 42 Introduction ................................................................................................................... 42 Materials and Methods .................................................................................................. 49 Results ........................................................................................................................... 53 Discussion ..................................................................................................................... 56 Figures ........................................................................................................................... 62 Tables ............................................................................................................................ 66 Literature Cited ............................................................................................................
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