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Evaluating Potential Growth Strategies Using Bone Histology in Pleistocene-Holocene Odocoileus Virginianus

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Evaluating Potential Growth Strategies Using Bone Histology in Pleistocene-Holocene Odocoileus Virginianus Evaluating potential growth strategies using bone histology in Pleistocene-Holocene Odocoileus virginianus (Mammalia) from Florida A thesis submitted To Kent State University in partial fulfillment of the requirement for the degree of Master of Science By Andrew Gerwitz August, 2016 © Copyright Thesis written by Andrew Nathan Gerwitz B.A., State University of New York at Geneseo, Geneseo, N.Y., 2010 M.S. Kent State University, 2016 Approved by Jeremy L. Green , Masters Advisor, Department of Geology Daniel Holm , Chair, Department of Geology James Blank , Dean College of Arts and Sciences TABLE OF CONTENTS TABLE OF CONTENTS………………………………………………………….………….…iii-iv LIST OF FIGURES……………………………..……..….………………….…….…..……...…v-vi LIST OF TABLES.………………………………….….….…….……….……….….….......…vii-ix ACKNOWLEDGEMENTS……………………………….…….…………….……….….....…..…x INTRODUCTION……………………………..………………….…..…………….…..….1 BACKGROUND………………………………...……………….....……………….….….2 Global Climate……………………………………………………………………..2 Florida Quaternary Climate and Ecosystems………………………………………3 Evolutionary Success of Cervidae…………………………………………………7 Specimen Localities……….……………………………….………........................7 Geologic Background.…...…………...…………………..……………..….….…..8 Measuring Growth in deer...……………………………………………………...13 Bone Microstructure………………..…………………...……...…………….......13 Abbreviation Key……………………………………………..…………..….…..16 METHODS……………………………....………………………...….…..….…………...18 Limb Bone Measurements……....……………………….….….……..……..…...18 Bone Histology………………...………………...………...……………..………25 Skeletochronology……….…….…………..………...…………………..…….....32 Osteocyte Density…………..……………..…………………….………..………36 RESULTS……………………….………..…………..…………………...…………........38 Climate differences across the Florida Penninsula…..……………….………......38 MDC and AL Comparisions Across Time and Geography…….……….…..……42 Histologic Descriptions…………………………………….…..……………..…..58 Skeletochronology………………………….………...….……………………….89 Osteocyte Density…………………………………...……………………..…....106 iii DISCUSSION……………………………………………………………………..……..110 Comparative Histology…………………………..……………………...……....111 Growth Lines and Skeletochronology….…...………………….......................…115 Osteocyte Density……………...………….......……………………..…..…...…118 Extrinsic Factors Influencing Gro.…………....……..……………...……...……121 Intrinsic Factors Influencing Growth……..……………...………………...........127 Comparing Growth Between O. virginianus and Other Cervids…...……......….129 Future directions……………....……………..……………….……..……...……135 CONCLUSIONS…………………..………….………………………..…...……....……136 REFERENCES CITED……......……...…….….….….…..……..………...…….....…….139 APPENDIX...…………………...………………….....…..……………...……......….….156 iv LIST OF FIGURES Figure 1. Pliocene to Pleistocene Marine δ18O isotope curves……………………….….………………....4 Figure 2. Map of localities in Florida where specimens were obtained for this study……….…………...20 Figure 3. Methodology for measuring limb bones AL and MDC……………………..……………….…21 Figure 4. Köppen-Geiger climate zones of Florida and locations for climate data……………….………23 Figure 5. Description of bone microstructures I……………………………………………………....…..30 Figure 6. Description of bone microstructures II……………………………………….………….…...…31 Figure 7. Methodology for APD and anterior derived growth rates……….……………..…………....….35 Figure 8. Methodology for osteocyte counting.……………………………………………….……….….37 Figure 9. Average temperature and precipitation data from Florida……………………………….….40-41 Figure 10. Chronologic MDC trends in limb elements from Northern Florida….…………….…..….…..48 Figure 11. Chronologic MDC trends in limb elements from Southern Florida….……………..….……...53 Figure 12. All histologically sampled femora from ING1A……………………………………....……....61 Figure 13. All histologically sampled femora from LSP1A and COL2A……………………...........……64 Figure 14. All histologically sampled femora from NH………………………..…………………....……66 Figure 15. All histologically sampled tibiae from ING1A……………………………………….…....….69 Figure 16. All histologically sampled tibiae from LSP1A……………………………………….…...…...71 Figure 17. All histologically sampled tibiae from COL2A and NH………………………………….…...73 Figure 18. All histologically sampled humeri from ING1A…………………………………….…..….....76 Figure 19. All histologically sampled humeri from LSP1A and COL2A……………………………...…79 Figure 20. All histologically sampled humeri from NH…………………………………….………...…..82 Figure 21. All histologically sampled radii from ING1A..…………………………….…………….....…85 Figure 22. All histologically sampled radii from NH……………………………..………………..…..…88 Figure 23. Average Femora APD measurements across all fossil localities………………..………….....95 Figure 24. Average Femora M/mAPDGR measurements across all fossil localities ………………….....96 Figure 25. Average Femora M/mANTGR measurements across all fossil localities ……………….....…97 v Figure 26. Average tibiae APD measurements across all fossil localities ……………………………..…99 Figure 27. Average tibiae MAPDGR measurements across all fossil localities ………………………...100 Figure 28. Average tibiae MANTGR measurements across all fossil localities………………………...101 Figure 29. Average humeri APD measurements across all fossil localities ………………………….…103 Figure 30. Average humeri MAPDGR measurements across all fossil localities ………………..…..…104 Figure 31. Average humeri MANTGR measurements across all fossil localities…………………….…105 Figure 32. Femora Osteocyte Density per GZ……………………………………………………….…..108 Figure 33. Tibiae Osteocyte Density per GZ………………………………………………………….....109 Figure 34. Ontogenetic and chronological changes in anterior and posterior bone microstructure.….…114 Figure 35. Comparison of Osteocyte density from single and stacked images………………………….120 Figure 36. Comparison of cervidae femoral growth rates……………………………………………….132 Figure 37. Comparison of cervidae tibial growth rates…………………………………………….…….133 Figure 38. Cervidae phylogeney. ………………………………………………………………………..134 vi LIST OF TABLES Table 1. Environmental summary of sites based on previous research…………………………………...12 Table 2. Age classes for deer used in this study based on degree of epiphyseal fusion…………………..22 Table 3. Statistical comparison of AL of adult limb elements from northern Florida populations over the past ~2 million years.......................................................................................................................44 Table 4. Means and standard deviations from AL of adult limb elements from northern Florida populations over the past ~2 million years….................................................................................44 Table 5. Statistical comparison of MDC of adult limb elements from northern Florida populations over the past ~2 million years……………………………………………………………………….…44 Table 6. ANOVA contrast and Post-Hoc results from comparison of MDC of adult limb elements from northern Florida populations over the past ~2million years……………………………..……45-46 Table 7. Means and standard deviations from MDC of adult limb elements from northern Florida populations over the past ~2 million years……...………………………………………………..47 Table 8. Statistical comparision of AL of adult limb elements from southern Florida populations over the past ~1.5 million years…………………………………………………………………………....50 Table 9. Means and standard deviations from MDC of adult radii from southern Florida populations over the past ~500 years……………………………………………………………………….…50 Table 10. Statistical comparison of MDC of adult limb elements from southern Florida populations over the past ~1.5 million years…...................……………………………………………..….............51 Table 11. ANOVA contrast and Post-Hoc results from comparison of metatarsal MDC from southern Florida over the past 1.5 million years.…………………………………………………….……..51 Table 12. Means and standard deviations from MDC of adult elements from southern Florida over the past 1.5 million years.………………………………………………………………………….....52 Table 13. Statistical comparison of MDC and AL of adult elements from Irvingtonian and Rancholabrean populations across their geographic distribution in Florida..…………………………………….55 vii Table 14. Statistical comparison of MDC of adult elements from Modern populations across their geographic distribution in Florida……………………………………………..………………….55 Table 15. Means and standard deviations from MDC of adult elements from modern populations across their geographic distribution in Florida………..............................................................................56 Table 16. Statistical comparison of AL of adult elements from modern populations across their geographic distribution in Florida……………………………………………………………..…56 Table 17. Means and standard deviations from AL of adult elements from modern populations across their geographic distribution in Florida……………………………………………………………..…56 Table 18. Gender from modern populations……………………………………………………………....57 Table 19. Average APD measurements per growth zone among each locality with limb elements containing measurable LAGs…….…………………………………………………………..….90 Table 20. Average MAPDGR for each element per locality containing measurable LAGs…………..…91 Table 21. Average MANTGR for each element per locality containing measurable LAGs………..........93 Table 22. Average single layer osteocyte density……………………………………………………..…107 Table 23. Average stacked layer osteocyte density…………………………………………………...…107 Table S1. Measurement data and age determination of all fossil specimens……………………….156-166 Table S2. Measurement data, age determination and gender of all modern specimens….………....167-177 Table S3. Locations in Florida used for climate data……………………………………………...…….178 Table S4. Monthly climate averages
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