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A Genus-Level Phylogenetic Analysis of Antilocapridae And

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A Genus-Level Phylogenetic Analysis of Antilocapridae And A GENUS-LEVEL PHYLOGENETIC ANALYSIS OF ANTILOCAPRIDAE AND IMPLICATIONS FOR THE EVOLUTION OF HEADGEAR MORPHOLOGY AND PALEOECOLOGY by HOLLEY MAY FLORA A THESIS Presented to the Department of EArth Sciences And the Graduate School of the University of Oregon in partiAl fulfillment of the requirements for the degree of MAster of Science September 2019 THESIS APPROVAL PAGE Student: Holley MAy Flora Title: A Genus-level Phylogenetic Analysis of AntilocApridae and ImplicAtions for the Evolution of HeAdgeAr Morphology and PAleoecology This Thesis has been accepted and approved in partiAl fulfillment of the requirements for the MAster of Science degree in the Department of EArth Sciences by: EdwArd Byrd DAvis Advisor SAmAntha S. B. Hopkins Core Member Matthew Polizzotto Core Member Stephen Frost Institutional RepresentAtive And JAnet Woodruff-Borden DeAn of the Graduate School Original Approval signatures are on file with the University of Oregon Graduate School Degree awArded September 2019. ii ã 2019 Holley MAy Flora This work is licensed under a CreAtive Commons Attribution-NonCommercial-NoDerivs (United States) License. iii THESIS ABSTRACT Holley MAy Flora MAster of Science Department of EArth Sciences September 2019 Title: A Genus-level Phylogenetic Analysis of AntilocApridae and ImplicAtions for the Evolution of HeAdgeAr Morphology and PAleoecology The shapes of Artiodactyl heAdgeAr plAy key roles in interactions with their environment and eAch other. Consequently, heAdgeAr morphology cAn be used to predict behavior. For eXAmple, lArger, recurved horns are typicAl of gregarious, lArge-bodied AnimAls fighting for mAtes. SmAller spike-like horns are more characteristic of small- bodied, paired mAtes from closed environments. Here, I report a genus-level clAdistic Analysis of the extinct family, AntilocApridae, testing prior hypotheses of evolutionary history And heAdgeAr evolution. I included 53 post-craniAl, craniAl, and heAdgeAr characters, expanding on previous analyses by developing 14 novel character traits. This phylogenetic analysis not only estAblishes ancestral heAdgeAr morphology of AntilocApridae but allows inferences of mAjor sociAl structure changes. These results confirm previous works inferred through comparison with Artiodactyl families that AntilocAprids evolved from small-bodied monogamous pairs to lArge-bodied gregarious herds. Our findings show multiple originations of herding sociAl behavior. iv CURRICULUM VITAE NAME OF AUTHOR: Holley MAy Flora GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED: University of Oregon, Eugene MontAna StAte University, BozemAn Boise StAte University, Boise, Idaho DEGREES AWARDED: MAster of Science, EArth Sciences, 2019, University of Oregon Bachelor’s of Science, EArth Sciences, 2015, MontAna StAte University AREAS OF SPECIAL INTEREST: Functional Morphology Evolutionary Processes PROFESSIONAL EXPERIENCE: PAleontologicAl AssistAnt, Geocorps, MAnti-LA SAl NAtional Forest, 2019 Graduate Employee, University of Oregon, Department of EArth Sciences, 2017- 2019 Preparing and teAching lAbs for introductory courses (GEOL 101 & 103), teAching AssistAnt (GEOL 304), and assisting the collections mAnager (Museum of NAtural And Cultural History) Graduate Employee, University of Oregon, 2017-2019 • Museum of NAtural and Cultural History o Coordinating volunteers, undergraduates, and supplies for collections. • GEOL 304 “Fossil Record” o Grading eXAms and running the online assignments. • GEOL 101 “Introductory Geology” and GEOL 203 “History of Life” v o TeAching multiple lAb sections on mineral and fossil identificAtion and geologicAl processes. Fossil Preparator, Geoscientists in the PArks, BadlAnds NAtional PArk, 2018 LAb TechniciAn, Horner PAleohistology LAboratory, Museum of the Rockies, 2014-2015 GRANTS, AWARDS, HONORS: DAvid S EAsly MemoriAl AwArd ($5,000) Spring 2019 University of Oregon, SpeciAl “Opps” Travel & ReseArch AwArd ($1,000) Spring 2019 University of Oregon, Department of EArth Sciences ($310) Winter 2018 University of Oregon Department of EArth Sciences Grant ($1,500) Spring 2018 Undergraduate ScholArs Program Travel Grant ($500) FAll 2015 Society of Vertebrate PAleontology JAckson Student Travel Grant ($500) FAll 2015 UndergrAduate ScholArs Program ReseArch Grant o Developing a Microfossil IdentificAtion Guide for LAb and Field Use ($1,200) Summer 2014 o Causes and Effects of LAndslides in Rural Morocco ($900) Spring 2015 Western Undergraduate Exchange ScholArship. 2012-2015 MontAna StAte University DeAn’s List. 2012 PUBLICATIONS: John P. Wilson; D. Cary Woodruff, JAcob D. GArdner; Holley M. Flora; John R. Horner; Chris Organ. (2016) Vertebral Adaptations to Large Body Size in Theropod Dinosaurs. PLoS ONE 11(7): e0158962. doi:10.1371/journal.pone.0158962 vi ACKNOWLEDGMENTS I would like to thank all of the institutions and collections mAnagers that welcomed datA collection for this project: Judy GAlkin at the AmericAn Museum of NAtural History, PAt Holroyd at the University of CaliforniA Museum of PAleontology, Dr. SAm McLeod And VAnessA Rhue at the NAtural History Museum of Los Angeles County, George Corner and Shane Tucker at the University of Nebraska StAte Museum. Thank you to all of the people that mAde visiting museum collections eAsier: Carol Baumbauer, Cathy Cline, The El Adlis, Devra Hock, ZAch Reid, and the Floras. I would like to acknowledge the grants from the University of Oregon department of EArth sciences, without which mAny of the aforementioned museum visits would not have happened. And finally, thank you to the Hopkins-DAvis vertebrate paleontology lAb and the Dueppen AfricAn archeology lAb for specimen and softwAre use and their continued general support and guidance. vii TABLE OF CONTENTS Chapter Page I . A GENUS-LEVEL PHYLOGENETIC ANALYSIS OF ANTILOCAPRIDAE AND IMPLICATIONS FOR THE EVOLUTION OF HEADGEAR MORPHOLOGY AND PALEOECOLOGY ............................................................................................................. 1 Introduction ....................................................................................................... 1 Methods ............................................................................................................. 6 Results ............................................................................................................. 10 SystemAtic PAleontology ........................................................................... 10 Phylogenetic Trait MApping ..................................................................... 18 Discussion ....................................................................................................... 19 Future Work .............................................................................................. 24 Conclusions ............................................................................................... 25 APPENDICES .................................................................................................................. 26 A. List of Species ................................................................................................. 26 B. Character List .................................................................................................. 26 HeAdgeAr Characters ........................................................................... 27 DentAl Characters ................................................................................ 32 CraniAl Characters ............................................................................... 36 Post-CraniAl Characters ...................................................................... 39 C. Character MAtriX .............................................................................................. 42 D. Specimen List ................................................................................................... 44 REFERENCES CITED ..................................................................................................... 50 viii LIST OF FIGURES Figure PAge Figure 1. Ottoceros Skull ................................................................................................... 9 Figure 2. AntilocApridae Phylogeny ................................................................................. 11 Figure 3. Behavioral Categories MApped ......................................................................... 21 Figure 4. Character #11 ..................................................................................................... 30 Figure 5. DentAl NomenclAture ......................................................................................... 32 Figure 6. Character #39 ..................................................................................................... 37 Figure 7. Character #53 ..................................................................................................... 41 ix LIST OF TABLES TAble PAge TAble 1. Behavior Category BreAkdown ............................................................................. 2 X INTRODUCTION Headgear are an important feature shared by almost every member of the Ruminantia (Davis et al., 2011). Bony ossicones and other protuberances are found in many ruminant families, although the evolution of the different forms of headgear remains enigmatic. Within a phylogenetic framework the ruminant history may be uncovered. Past work on Bovidae and Cervidae, families within Ruminantia, have examined correlation between headgear shape and behavior (Caro et al, 2003; Jarman, 1974;
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