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Cranial Morphological Distinctiveness Between Ursus Arctos and U East Tennessee State University Digital Commons @ East Tennessee State University Electronic Theses and Dissertations Student Works 5-2017 Cranial Morphological Distinctiveness Between Ursus arctos and U. americanus Benjamin James Hillesheim East Tennessee State University Follow this and additional works at: https://dc.etsu.edu/etd Part of the Biodiversity Commons, Evolution Commons, and the Paleontology Commons Recommended Citation Hillesheim, Benjamin James, "Cranial Morphological Distinctiveness Between Ursus arctos and U. americanus" (2017). Electronic Theses and Dissertations. Paper 3261. https://dc.etsu.edu/etd/3261 This Thesis - Open Access is brought to you for free and open access by the Student Works at Digital Commons @ East Tennessee State University. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Digital Commons @ East Tennessee State University. For more information, please contact [email protected]. Cranial Morphological Distinctiveness Between Ursus arctos and U. americanus ____________________________________ A thesis presented to the Department of Geosciences East Tennessee State University In partial fulfillment of the requirements for the degree Master of Science in Geosciences ____________________________________ by Benjamin Hillesheim May 2017 ____________________________________ Dr. Blaine W. Schubert, Chair Dr. Steven C. Wallace Dr. Josh X. Samuels Keywords: Ursidae, Geometric morphometrics, Ursus americanus, Ursus arctos, Last Glacial Maximum ABSTRACT Cranial Morphological Distinctiveness Between Ursus arctos and U. americanus by Benjamin J. Hillesheim Despite being separated by millions of years of evolution, black bears (Ursus americanus) and brown bears (Ursus arctos) can be difficult to distinguish based on skeletal and dental material alone. Complicating matters, some Late Pleistocene U. americanus are significantly larger in size than their modern relatives, obscuring the identification of the two bears. In the past, fossil bears have been identified based on differences in dental morphology or size. This study used geometric morphometrics to look at overall differences in cranial shape and used step-wise discriminant analysis to identify specific characters that distinguish cranial morphology between black and brown bears. Such differences could prove important in identifying fossil bears when crania are present but teeth are missing. Furthermore, being able to properly identify U. arctos and U. americanus crania is important in understanding evolutionary and ecological distinctions among both fossil and modern bears. Principal components, discriminant, and thin plate spline analyses indicated a clear morphological separation between the crania of U. americanus and U. arctos and highlighted key identifying features including a more convex forehead and a narrower, more elongate rostrum in U. arctos than U. americanus. 2 DEDICATION I dedicate this thesis to my mom Dana, my dad Bob, and my soon-to-be wife Kayla Sue 3 ACKNOWLEDGMENTS I would like to acknowledge Blaine Schubert, Steven Wallace, and Josh Samuels for serving on my committee and providing their guidance and insight. I would also like to acknowledge the Bell Museum of Natural History in St. Paul and the Mammal Collections at the Smithsonian for allowing me access to their collections of bear crania. Finally, I’d like to thank my family and all of the graduate students with whom I’ve had the pleasure of working with over the past two years. 4 TABLE OF CONTENTS Page ABSTRACT ................................................................................................................................ 2 Chapter 1. INTRODUCTION ................................................................................................................... 8 Phylogeny of North American Ursidae ........................................................................... 9 Ecology of North American Ursidae ............................................................................... 12 Cranial Morphology of Black and Brown Bears ………………………...……………. 16 Glacial Impact ................................................................................................................ 21 2. METHODS ............................................................................................................................ 24 Selection of Bear Crania ……………………………………………………………… 25 Photography ……………………………………………………………………........... 26 Landmark Placement …………………………………………………….…………… 26 Principal Components Analysis …………………………………………….………... 29 Discriminant Analysis ………………………………………………………………... 29 Thin Plate Splines ……………………………………………………………………... 30 Potential Complications ……………………………………………………………… 30 3. RESULTS ............................................................................................................................. 32 Principal Components Analysis .................................................................................... 32 PCA, Anterior View – Fig. 2a………………………………………………… 33 PCA, Dorsal View – Fig. 2b…………………………………………………... 33 PCA, Lateral View – Fig. 2c…………………………………………………… 34 PCA, Posterior View – Fig. 2d…………………………………………………. 34 PCA, Ventral View – Fig. 2e………………………………………………….... 35 Discriminant Analysis ................................................................................................... 35 DA, Anterior View – Fig. 2a………………………………………………… 36 DA, Dorsal View – Fig. 2b…………………………………………………... 36 DA, Lateral View – Fig. 2c…………………………………………………… 37 5 DA, Posterior View – Fig. 2d…………………………………………………. 38 DA, Ventral View – Fig. 2e………………………………………………….... 39 Thin Plate Splines ……………………………………………………………………... 39 TPS, Anterior View – Fig. 4a………………………………………………… 39 TPS, Dorsal View – Fig. 4a…………………………………………………... 40 TPS, Lateral View – Fig. 4a…………………………………………………… 40 TPS, Posterior View – Fig. 4b…………………………………………………. 42 TPS, Ventral View – Fig. 4b………………………………………………….... 42 4. DISCUSSION AND CONCLUSIONS................................................................................... 43 Conclusions…….……………………………………………………………………… 47 REFERENCES …...................................................................................................................... 48 APPENDIX ............................................................................................................................... 53 Supplemental Tables……………………………………………...…………………… 53 VITA …………………………………………………………………………………………. 58 6 LIST OF FIGURES Figure Page 1a. Landmark locations (Anterior, Dorsal, and Lateral Views) ……………………………. 27 1b. Landmark locations (Posterior and Ventral Views) ……………………………………. 28 2a-b. Principal components analysis (Anterior and Dorsal Views) ………………………….. 32 2c-d. Principal components analysis (Lateral and Posterior Views) …………………………. 33 2e. Principal components analysis (Ventral View) ………………………………………… 34 3a-b. Discriminant analysis (Anterior and Dorsal Views) …………………………………… 36 3c-d. Discriminant analysis (Lateral and Posterior Views) ………………………………….. 37 3e. Discriminant analysis (Ventral View) …………………………………………………. 38 4a. Thin plate splines (Anterior, Dorsal, and Lateral Views) ……………………………… 40 4b. Thin plate splines (Posterior and Ventral Views) ……………………………………… 41 7 CHAPTER 1 INTRODUCTION Paleontology provides an important foundation for understanding ecological changes over time. However, such studies are typically based on databases of compiled identifications, and overarching ecological studies are only good if original identifications are solid. As a result, ecological studies are only as good as the identification metrics with which taxa are categorized. One such case is the ability to identify North American black and brown bears through time. The most accurate and current paleontological method to distinguish these species is based on linear measurements of the second upper molar (Gordon 1977). Pleistocene black bears, however, have the potential to occasionally be significantly larger than their modern counterparts – potentially as large as modern brown bears (Wolverton 1996). This potential for size variability within fossil black bears renders current metrics of identification based solely on size untenable. This study used step-wise discriminant analysis to identify key shape differences between multiple cranial views of U. americanus and U. arctos so that these morphological features might be used as a new quantitative identification method for fossil specimens, which is not affected by the variable cranium size of black bears over time. In addition, determining what key differences separate brown and black bear crania may provide new insight into the evolution and ecology of black and brown bears. Previous studies have performed similar methods in analyses that included modern black and brown bear crania (Sacco and Van Valkenburgh 2004; Figueirido et al. 2009; Fuchs et al. 2015), but thus far study of the differences that exist in the crania of these bears has focused on ecomorphology, not identification. The present study has two main objectives. The first was to determine whether the amount of variation and phenotypic plasticity that occurs within U. americanus and U. arctos is 8 sufficient to obscure identification based on ursid crania alone. The present study used principal components analysis to compare black and brown bear crania at five different angles and examines the degree of morphological distinction found between the two bears. In doing so, the present study tested whether previous
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