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 (Mammalia) from

Florida

A thesis submitted

To Kent State University in partial

fulfillment of the requirement for the

degree of Master of Science

Andrew Gerwitz

August, 2016

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

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

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

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 from different localities in Florida…………………………...179-183

Table S5. Statistical comparison between left and right adult elements from similar geographic localities

and geochronologic times…………………………………………………………………..184-185

Table S6. Histologic measurements and data from sampled specimens……………………………186-188

Table S7. Individual APD measurements for individual LAGs from samples elements………...…189-190

Table S8. MAPDGR of sampled elements……...………………………………………………….……191

Table S9. mAPDGR of sampled elements…………………………………………………………….…192

Table S10. Anterior cortical LAG measurements ……………………………………………….…193-194

viii

Table S11. Anterior growth rates (M/mGRGP and MANTGR)……………………………………195-196

Table S12. Anterior growth rates (M/mGRST and mANTGR)………………………………….…197-199

Table S13. Individual single layer osteocyte lacunae count and calculated OD for each growth zone for

histologically sampled specimens………………………………………………………….200-202

Table S14. Stacked layer osteocyte lacunae counts and volume. …………………………….….…203-204

Table S15. Stacked image osteocyte lacunae density for each growth zone………………………..205-207

Acknowledgements

This project was financially supported by the Geological Society of America Graduate Research Grant,

Graduate Student Senate Research Grant and Travel Grant and Sigma Gamma Epsilon Travel Grant, and would not have been possible without access to the all vertebrate material at the Florida Museum of

Natural History provided by Richard Hulbert, Candace McCaffery, Verity Mathis and Irvy Quitmyer. I would like to thank all my friends and colleagues at Kent, especially those who kept me company through many hours of grinding thin-sections. I would especially like to thank my thesis committee members, Dr.

Joseph Ortiz who provided ample help and guidance regarding statistical testing and interpreting climatic and environmental information from stable isotopes; Dr. Rodney Feldmann for always demanding the best in me and imparting knowledge of paleontology attributable to all organisms. I would particularly like to thank my advisor Dr. Jeremy Green, for his mentorship, guidance, and unwavering patience, through all the trials and tribulations which arose during this project, through which I can regard him as both a colleague and a friend. Various other colleagues have provided helpful suggestions, including Dr.

Alison Smith, Dr. John Skedros, Dr. Mary Schweitzer, Ellen-Therese Lamm, and Carolyn Levitt-Bussian.

Preliminary results from this research were presented at the Annual GSA Meeting in Vancouver, British

Columbia in 2014 and Paleoecology Symposium Research Workshop at the Cleveland Museum of

Natural History, 2015. Finally I would like to thank my family for their continued support through all of this project and graduate school. I would like to especially thank my sister for providing me access to

Adobe Photoshop, through which none of the figures would have been made without, and my parents for their constant love and support provided each and every day.

Introduction

Populations are rarely evolutionarily stagnant; natural selection of traits constantly occurs and populations respond to changing conditions. Circumstances may arise as conditions change with different tempos where evolution may speed up or slow down. Organisms that have survived these different tempos of change are extremely useful for study in a broad context to determine the adaptations that have allowed them to succeed. The mammalian clade Cervidae is one such evolutionary success story, with diversification and rapid evolutionary adaption throughout the Pleistocene and well into the Holocene.

The purpose of this work is to provide new insights into how geologically rapid changes in climate and environment throughout the Pleistocene and Holocene may have influenced growth in Florida Odocoileus virginianus (Zimmermann, 1780), one of a few North American mid-large sized mammals to survive the end-Pleistocene extinction.

Background

Global Climate

The Quaternary period is characterized by cyclical shifts in global climate that is recognized through use of multiple proxies (e.g. stable isotopes, pollen, ice and sediment accumulation, glacially derived landscapes …etc). These climatic changes affected the environment of terrestrial and marine fauna alike. Stable isotopes from marine sediment cores reveal a gradual increasing trend in δ18O during the Pliocene and into the early Pleistocene, indicating a global cooling as more 16O was taken out of the oceans and incorporated into ice sheets in the polar regions (Sosdian and Rosenthal, 2009) (Fig. 1A).

Global ocean circulation patterns were already shifting through the Pliocene and into the

Pleistocene as the Central American Seaway and Indonesian Seaway were both closing, likely impacting world-wide climate through these time periods (Cane and Molnar, 2001; Woodburne, 2010). These changes in ocean circulation set up feedback loops which likely contributed to a global cooling trend from

Pliocene and into the Pleistocene (Woodburne, 2010; Fig. 1A). Specific climatic cycles during the

Pleistocene are also thought to be a direct response to changes in solar radiation due to cyclic orbital parameters such as precession, obliquity and eccentricity of the Earth’s axis resulting in ~23,000, 41,000,

100,000, and 400,000 year cycles respectively (Lisiecki and Raymo, 2007; Fig. 1).

Pliocene and early Pleistocene glacial and interglacial periods show up as symmetric curves in the

δ18O record about every 41,000 years (Lisiecki and Raymo, 2007) (Fig. 1). Around 900 ka to 700 ka, these cycles became much more asymmetric with increased amplitudes and wavelengths that fluctuate around 100,000 years, yet they have not been correlated to any change in precession, obliquity, or eccentricity (Lisiecki and Raymo, 2007) (Fig. 1). More recent glacials during the past 900 ka had longer

durations and created larger ice sheets on the continents, yet they also ended more abruptly and are interspersed with short interglacials (Fig. 1). Environmental, climatic, and oceanographic shifts associated with these glacial and interglacial cycles have been extensively documented in marine isotopes from ocean sediments, terrestrial pollen from lake sediments and faunal community make up throughout the

Pleistocene from the Caribbean and Florida (Emiliani et al, 1966; Imbrie et al., 1973; van Donk, 1976;

Joyce et al., 1990; Grimm et al., 1993, 2006; Willard et al., 1993; Koch et al., 1998, Lynch Steiglitz et al., 1999, 2011; DeSantis et al., 2009; Morgan and Emslie, 2010) making it an ideal locality for observing these changes in the fossil record. .

Florida Quaternary Climate and Ecosystems

The abundance of Quaternary fossils from Florida, representing organisms that lived during the glacial-interglacial cycles described above, offers the potential to analyze possible short and long term responses of ecosystems and species to climate change during this time. Late Pleistocene pollen records indicate changes in Florida’s vegetation, providing a proxy for environmental shifts between glacial and interglacial conditions (Grimm et al., 1993, 2006). Contrary to the cool, dry glacial and warm, wet interglacial climatic conditions experienced in most of North America, Florida interglacials were cool and dry with extensive oak-scrub prairie grasslands, whereas glacial climates were warm and wet with extensive pine dominated forests (Grimm et al., 1993, 2006). This was likely caused by a slowing of the

Gulf Stream and associated thermohaline circulation (which transports warm tropical waters to the North

Atlantic) during glacial intervals due to extent of sea ice cover in the North Atlantic (Lynch-Steiglitz et al., 1999, 2011; Grimm et al., 2006). As thermohaline circulation slowed or shut down, warmer waters accumulated in the Gulf of Mexico, resulting in a warmer and more humid climate in the surrounding area.

O (‰) O

18 δ

O (‰) C

Figure 1. A) Global Stacked benthic δ18O data modified from Lisiecki and Raymo (2005). Positive δ18O values indicate interglacial whereas higher values indicate interglacial periods. B) Planktic δ18O from van Donk, (1976) from tropical Atlantic core site V16-205. C) Planktic δ18O from Joyce et al. (1990) from off the western Florida platform in the Gulf of Mexico. D) Partial Pleistocene record from (A) (red box); numbers correlate with marine isotope stages; even numbers are glacial periods and odd numbers are interglacial periods. Localities for this study marked at approximate geochronologic range (see Fig. 2). Abbr.: North American Land Mammal Ages (NALMA; from Baronsky, 2014); Black: Inglis 1A (ING1A); Blue: Leisey Shell Pit 1A (LSP1A); Green: Coleman 2A (COL2A); Red: Nichol’s Hammock (NH).

In contrast, during interglacials the thermohaline circulation was strengthened and the Gulf Stream could transport warmer water to the North Atlantic, causing the climate around the Gulf of Mexico to be slightly cooler and more arid (Grimm, 2006). This hypothesis is supported by multiple paleoceanographic circulation models, as well as isotopic data from the Gulf of Mexico since the last glacial maximum, all of which reveal a correlation with Gulf Stream strength, sea surface temperatures, and glacial/interglacial cycles (Manabe and Stouffer, 1997; Rühlemann et al., 1999; Lynch-Steiglitz et al., 1999, 2011; Peterson et al., 2000). Grimm et al. (2006) suggested that seasonal differences in the amount of precipitation received during the winter months would be the greatest difference between glacial and interglacial cycles in Florida. Warmer water temperature during glacial intervals would insulate the Florida peninsula during winter months, creating more evaporation of sea water which would in turn correspond to more precipitation during the dry season (Grimm et al., 2006). Interglacial periods would have an opposite effect, producing rainfall amounts less than or similar to modern levels, especially during winter months

(Grimm et al., 2006). Changes in the availability of seasonal precipitation altered the environments on the

Florida peninsula between glacial and interglacial cycles over the past 50,000 years (Grimm et al., 2006).

Although the pollen record does not go back more than 50,000 years in Florida, changes in precipitation between glacial and interglacial times are in agreement with recorded stable isotope data from terrestrial mammal communities reflecting changes in the environment (DeSantis et al., 2009; Yann and DeSantis,

2014).

The impact of changes in precipitation and aridity on the Pleistocene terrestrial flora and fauna of

Florida has been extensively documented and studied (Grimm et al., 1993; Koch et al., 1998; Alroy et al.,

2000; Grimm, et al., 2006; Huang et al., 2006; Hoppe and Koch, 2007; DeSantis et al., 2009; Morgan and

Emslie, 2010; Yann et al., 2013; Yann and DeSantis, 2014). During glacials, increased humidity as well as the exposure of the continental margin around Florida (increasing its size three fold) created a “tropical paradise” with abundant food and space for a diverse ecosystem (Webb, 1974). Multiple mammalian species of intermediate to large size, along with some species of birds migrated into Florida during glacial times (Hoppe and Koch, 2007; Lister and Stuart, 2008; Morgan and Emslie, 2010), as low sea-levels

would have also exposed a shallow platform along the Gulf Coast for western and tropical organisms to move into Florida (Webb, 1974). Transitions to interglacials caused a major transgression and massive landmass reduction of the Florida peninsula, which resulted in migration of organisms out of Florida as well as increased the isolation of organisms within Florida (Morgan and Emslie, 2010).

However, some populations such as deer may have remained in Florida during transgressive sequences based on 87Sr/ 86Sr values from specimens recorded from before and after the late glacial maximum (Hoppe and Koch, 2007), supporting their ability to adapt to changing environmental conditions. Forests typically provide the browse which is the main component of deer diet (Geist, 1998;

Hoppe and Koch, 2007; DeSantis et al., 2009), yet the availability of browse (typically C3 plants which uptake CO2 and O2 in equal concentrations; thus are affected by changing CO2 concentrations; generally higher CO2 equals more C3) may have decreased during interglacials due to the decrease in precipitation and CO2 levels (DeSantis et al., 2009; Yann and DeSantis, 2014). This climatic change would have also increased C4 plant (utilize and extra metabolic step to concentrate CO2 gas used for photosynthesis, are not affected as much to changing CO2 concentrations) grassland communities through which would result in fewer habitats and quantity of browse for deer (DeSantis et al., 2009; Yann and DeSantis, 2014). Even so, deer were present and survived through the Pleistocene partially through their ability to opportunistically forage any resource available even if it was C4 plants (DeSantis et al., 2009; Yann and

DeSantis, 2014). Therefore, deer were able to survive in both arid and humid conditions in the Pleistocene

(as they continue to do today) even though their food availability may have diminished and changed during interglacials. Odocoileus virginianus is a well-known example of a Florida deer both in the modern and fossil records and was likely able to adapt to changing environmental and climatic regimes and enabling them to be resilient through the Pleistocene. I hypothesize that growth in deer populations in

Florida may reflect some adaptive capability to overcome these climatic and environmental shifts throughout the Pleistocene.

Evolutionary Success of Cervidae

Members from the clade Cervidae can be found globally especially within North America where diversification of Cervidae have produced five genera (Alces (Gray, 1821), Cervus (Linnaeus, 1758),

Mazama (Rafinesque, 1817), Odocoileus (Ranfinesque, 1832) and Rangifer (Smith, 1827); Gustafson,

2015). Generally Cervidae are rare and understudied within the fossil record due to lack of Neogene deposits, fragmentary preservation of material and lack of phylogenetically useful material (Gustafson,

2015). Odocoileus virginianus is the only example of a modern North American cervid with a fossil record that extends back into the Pre-Pleistocene geologic record (Gustafson, 2015). The earliest and geologic record of the genus Odocoileus was within the Palmetto fauna of Florida (5-4.7 Ma; Webb et al.,

2008). Around this time, other deer are known from around North America such as Eocoileus (Webb,

2000; Palmetto fauna) and Bretzia pseudalces (Fry and Gustafson, 1974; White Bluffs fauna), but lack the expansive chronologic and geographic range of Odocoileus. Due to the longevity of the geochronologic range of O. virginianus (Pliocene – Present), the species offers a unique chance to study the evolution of a single species within the fossil record. Further, due to the evolutionary success of O. virginianus through the present time this species provides an insight into how a species may respond to climatic and environmental upheaval that occurred through the Pleistocene.

Specimen Localities

Through collaboration with the Florida Museum of Natural History (FLMNH), four fossil localities that were deposited during either glacial (or transitional period into a glacial; i.e. Coleman 2A) or interglacial intervals, have constrained geochronological time frames beyond assignment to a broad biochronologic North American Land Mammal Age (NALMA) and contained ample associated deer limb material have been selected for study (Fig. 2).

Glacial sites:  Inglis 1A (ING1A; 2.0-1.6 Ma; Morgan and Hulbert, 1995; DeSantis et al., 2009)

 Coleman 2A (COL2A; 500-300 ka; transitional interglacial – glacial; Martin, 1969)

Interglacial sites:  Leisey Shell Pit 1A (LSP1A; 1.6-1.3 Ma; Morgan and Hulbert, 1995)

 Nichol’s Hammock (NH; 500 years; Hirschfeld, 1969)

Geologic Background

Glacial Localities

Inglis 1A (ING1A; 2.0 – 1.6 Ma; Table 1; Fig. 2) is a sinkhole deposit consisting of clay and sand that extends 5 m below current sea level in Inglis Formation (Eocene) in Citrus County, Florida

(Morgan and Hulbert, 1995). Morgan and Emslie (2010) describe the fauna from ING1A as lacking marine species, with a diverse, non-native assemblage of birds and mammals of western and South

American affinities, suggesting the site formed during low sea-level (Morgan and Hulbert, 1995). Based on the vertebrate faunal assemblage at this site, it was assigned to the Blancan NALMA, which extends through much of the Pliocene to the Early-Mid Pleistocene (Klein, 1971; Barnosky et al., 2014; Fig. 2).

Herpetological and avian fossils along with pollen samples, from this site, and contemporary (2.0

– 1.6 Ma) sites in Florida document an environment that consisted of a mix of long leaf pine and xeric oak pollen, indicating a mix of upland forests and low level grasslands (Meylan, 1982; Emslie et al.,

1996, 1998; Emslie, 1998). DeSantis et al. (2009) mentioned that ING1A was well forested with C3 mammalian browsers (mammals which contain lower concentrations of δ13C in their tooth enamel indicating diets rich in C3 plants; generally browsing animals) although there were also some C4 grazers

(mammals which contain higher concentrations of δ13C in their tooth enamel indicating diets rich in C4 plants; generally grazers) that occur in much lower abundance, which supports a mostly mesic environment.

Correlation between ING1A and dated North American glacial till deposits may further support the interpretation of this site as representing a glacial interval. Richmond (1986) described a glacial till within basalt flows from The Narrows of the Grand Canyon of the Yellowstone River, Yellowstone

National Park, Wyoming that are dated with the age range of ING1A. This glacial event may correspond to marine oxygen isotope stage 40 from van Donk (1976), (Richmond, 1986) (Fig. 1B). More recent δ18O data from a core taken off the Florida Platform by Joyce et al. (1990) in the NE Gulf of Mexico (ODP-

625B; Fig. 1C), and a stacked core (LR04), (Lisiecki and Raymo, 2005) shows at least 8-10 glaciations, correlating to marine isotope stages 56 - 76 (Fig. 1D). Although a specific glaciation event associated with the ING1A deposit is unknown, current evidence from faunal and environmental analyses support that this site as being deposited during a glacial interval (DeSantis et al., 2009; Morgan and Emslie,

2010).

Coleman 2A (COL2A; 500 – 300 ka, Table 1; Fig. 2) is also a sinkhole deposit consisting of sand and clays within the Crystal River Formation (upper Eocene) in Sumter County, Florida (Fig. 2).

Fauna found within the deposits are concentrated within a 3 ft layer, which may indicate rapid accumulation of animals within the sinkhole as revealed by excellent preservation and associated elements of individuals that extend beyond sedimentary boundaries (Martin, 1969). The fauna of COL2A are associated with a late Irvingtonian NALMA, placing it within the late middle Pleistocene (500-300 ka;

Figs. 1 and 2). Younger deposits directly above COL2A contain Bison, an index species for the

Rancholabrean (Morgan and Hulbert, 1995), the NALMA the age that directly follows the Irvingtonian.

COL2A fauna include a strong western and tropical influence among the avian and mammalian species, which suggests its correlation with a glacial interval (Morgan and Emslie, 2010). To acquire a paleoecological interpretation for this site, Martin (1969) used the habitat preference of living Floridian mammals as a comparison to fossil faunas. He observed that many species in COL2A would have lived in a savanna transitioning to a forested environment due to a mixture of both savanna-like species and some subtropical to tropical forest taxa; the latter being dominant in the environment post ~300 ka.

In North America, Richmond and Fullerton (1986) mention glacial deposits associated with a glaciation period between 610 – 302 ka, which falls within the time frame of COL2A. Deposits from this glacial interval represent at least two and maybe three glaciations from the mountain ranges of the western US and consists of till and/or moraine deposits constrained in age by volcanic rocks (Richmond and Fullerton (1986). The ages of these deposits correlate to isotope stages 10, 12, and 14 (Richmond and

Fullerton, 1986; Lisiecki and Raymo, 2005) (Fig 1D). The isotope record from van Donk’s (1976) and

Joyce et al. (1990) record multiple glaciations between 500 – 300 kyrs (Fig. 1B-C). Based on the faunal assemblage (western and tropical), environmental interpretation (transitioning savanna to forest) as well as isotopic and glacial advance deposit history this study will refer to COL2A as a rapidly accumulated fauna representing a transitional interglacial to glacial period.

Interglacial Localities

Leisey Shell Pit 1A (LSP1A; 1.6 – 1.3 Ma; Table 1; Fig. 2) consists of a bone bed in a dark organic lens of silt and clay in a shell bed, located stratigraphically in the uppermost Bermont Formation

(1.6-1.3 Ma) in Hillsborough County, Florida (Morgan and Hulbert, 1995). LSP1A consist of 203 species of vertebrates, including 42 mammals, and represents one of the most fossiliferous late Cenozoic sites from Florida. The mammal fauna correlates to the Irvingtonian NALMA, placing it within the middle

Pleistocene (Figs. 1-2). The abundance of shells in the site represents a mix of near-shore marine to estuarine environments with some freshwater and terrestrial influence (Webb et al., 1989). Many of the vertebrate and freshwater fossils are interpreted to have been washed into the environment based on taphonomic indicators, such as scratches on bones and fragmented shells (Morgan and Hulbert, 1995).

Therefore, the depositional environment was a near-shore, shallow, brackish to marine water, probably estuarine environment (Webb et al., 1989).The terrestrial environment of LSP1A is interpreted as a mix of mesic and xeric woodlands of non-tropical origin, with some wetland and shallow grassy embayments based on faunal composition, pollen, and macro-botanical specimens from this site (Pratt and Hulbert, 1995; Rich and Newsom, 1995). DeSantis et al. (2009) found that the δ18O concentration in mammalian teeth indicate an arid, grassland environment. Richmond and Fullerton (1986) concluded that

the time between 1.55 – 1.0 Ma as a period lacking any major glaciations. The LR04 core of Lisiecki and

Raymo (2005) reveals multiple interglacial events during this time (Fig. 1D). Foraminiferal δ18O records provided by van Donk (1976) and Joyce et al. (1990) reveal relatively smaller changes and broader transitions between glacial and interglacial events during this time period, corresponding with Richmond and Fullerton’s (1986) observations of few major glaciation events (Fig. 1). All the available data, therefore, indicates that LSP1A was deposited during an interglacial, with sea level higher than present, creating a swampy, estuarine depositional environment (Table 1). The vertebrate fauna represent a mixed mesic/xeric environment that occurred slightly upland of the depositional zone (Table 1).

Nichol’s Hammock (NH; ~500 years; Table 1; Fig. 2) is a sinkhole deposit in South Florida where the absence of Rattus and the presence of the now extinct Canis niger (Florida wolf; extinct by

1900 A.D) and Neotoma floridana (eastern woodrat; exterminated pre-1937) correspond to a pre- columbian age (Hirschfeld, 1969). Specifically, the age of Nichol’s Hammock is interpreted as being prior to 1900 and as far back as 500 yrs ago (Hirschfeld, 1969). An environmental interpretation similar to modern environments is used for NH which consists of mesic densely forested rockland hammock surrounded by xeric open pine rocklands (Hirschfeld, 1969). The fauna found within the sinkhole is a representative sample of the mammalian fauna currently found in South Florida (with the exceptions of extinct species noted above). Other organisms representing xeric environments may have been washed into this sink hole, while the mesic organisms most likely lived close to the sink hole (Hirschfeld, 1969).

Because of the recent age of this site, along with an interpreted mesic habitat surrounded by xeric habitats

(Hirschfeld, 1969), it will be considered an interglacial, but with a greater mesic influence than usual.

Table 1. Summary of environmental interpretation data from all Florida localities where histological specimens were taken.

Site Age Western and/or Environmental analysis Additional Comments Environmental Tropical interpretation Influenced fauna

Inglis 1A 2.0-1.6 Ma. Yes Mix of upland pine forests, and More C3 browsers than Glacial 2 low level grasses and oak C4 grazers scrub, lack of marine influence 1

Leisey Shell 1.6-1.3 Ma. No Mix of upland pine forests and Fauna dominated by C4 Interglacial Pit 1A shallow wetlands with grassy grazers 2 embayments 3

Coleman 2A ~500-300 ka. Yes Mixture of mammals found in Rapid accumulation of Transitional savannah and forests 3 fossils 3 Interglacial to Transitional savannah to Glacial forested 3

Nichol’s ~500 years No Similar to modern conditions, Greater mesic faunal Interglacial Hammock mixed, forest surrounded by influence 4 grasslands 4

1 – Morgan and Hulbert (1995)

2 – DeSantis et al. (2009)

3 – Martin (1969)

4 – Hirschfeld (1969)

Measuring Growth in Deer

Body/Limb Size

Overall skeletal/bone sizes can provide a broad measure of the body size and growth history of an organism. Although skeletal size cannot be directly calculated from disarticulated unassociated fossil specimens, articular length (AL) and mid-diaphysis circumference (MDC) of individual limb bone elements have been shown to be a good proxy for body size in mammals (Alexander et al., 1979;

Anderson et al., 1985; Purdue, 1987; Gingerich, 1990).

Skeletal size can be dependent on a variety of factors including: age, sexual maturity, gender, resource availability and geographic location. Therefore, it is important to control for and test as many of these factors as possible to determine the natural level of variations within and among different populations. Individuals within each population can be pooled into difference groups to account for these variables; such groups include ontogenetic stage and age (prenatal, juvenile, sub-adult, or adult, based on epiphyseal fusion of limb elements; Purdue, 1983; Flinn et al., 2013), sex (based on presence of antlers, if known), climatic/environmental regime (glacial vs. interglacial), geographic locality (i.e. north and southern Florida), and geochronology (i.e., from early, middle and late Pleistocene). Bone morphometric data (AL, and MDC) can be recorded to measure the natural variation that exists in these pooled groups among different populations. This analysis can provide a baseline test of whether changes in variables are statistically random or significant. Any changes in skeletal size due to any of the aforementioned factors can then be directly used in collaboration with the bone microstructure record to support changes in growth strategy among populations.

Bone Microstructure

Over the past 50 years, bone histology has been used to study the growth record of both modern and ancient vertebrates. These studies have focused primarily on living and extinct non-mammalian tetrapods such as dinosaurs (e.g. Horner et al., 1999, 2000; Padian et al., 2001) and non-mammalian therapsids (e.g. Chinsamy and Rubidge., 1993; Ray et al., 2004; Chinsamy-Turan, 2012), as these groups

routinely deposit Lines of Arrested Growth (LAGs) and/or annuli (zones of poorly vascularized cortical bone) that reflect a major stoppage or reduction, respectively, of somatic growth in their skeletal record

(Horner et al., 1999, 2000; Padian et al., 2001; ). Until recently, it was assumed that mammals grow continuously prior to sexual maturity and thus, rarely, if at all, exhibit LAGs during skeletal growth

(Chinsamy and Hillenius, 2004). Further, any such marks were presumed to have been removed by secondary remodeling of the cortex in adulthood (Chinsamy and Hillenius, 2004). This long-standing assumption was refuted by recent studies (Sander and Andrássy, 2006; Köhler et al., 2012; Jordana et al.,

In Press; Kolb et al., 2015a, b). Some studies even suggest that LAGs and cementum lines in teeth from mammals are deposited in response to seasonal changes in aridity (wet vs dry seasons; Meister, 1956;

Low and Cowan, 1963; Reimers and Norby, 1968; Chritz et al., 2009; Köhler et al., 2012). Further,

Köhler and Moya-Sola (2009) and Kolb et al. (2015a) found that island and some mainland artiodactyls can alter their growth and delay maturation due to limited resource availability. Thus deposition of LAGs can result due to changing climatic and environmental factors.

Most recently, many papers have addressed the evolution of growth strategies within artiodactyls

(Köhler and Moya-Sola, 2009; Marín-Moratalla et al., 2013; Jordana et al., 2014, In Press; Amson, In

Press; Kolb et al., 2015a, b), however, no histological study on fossil artiodactyl specimens has yet analyzed either spatial or temporal variation within an individual species. This study will address this problem by observing changes in growth strategies of Odocoileus virginianus through time in a geographically restricted region (i.e. Florida) from the Pleistocene to Holocene.

Many studies involving Pleistocene and Holocene terrestrial faunas of Florida have concentrated on isotopic and faunal distribution patterns as a means to interpret organismal responses to climatic change (e.g. Koch et al., 1998; Hoppe and Koch 2007; DeSantis et al., 2009; Higgins and MacFadden,

2009; Feranec et al., 2010; MacFadden et al., 2010; Yann et al., 2013; Yann and DeSantis, 2014).

However, little attention has focused on the analysis of growth signals in bone microstructure and their correlation with seasonality among Pleistocene mammals (Chritz et al., 2009) and modern mammals

(Köhler et al., 2012). Differences in seasonality between glacial and interglacial periods may have

influenced the timing, availability, and nutritional quality of plants for foraging, which could affect growth patterns of herbivorous animals. These growth patterns, if preserved, could be studied to learn about the physiological responses of organisms to these changes in seasonality. The proposed study will help address this deficiency in our knowledge.

For this study, I will test the hypothesis that late Cenozoic Odocoileus virginianus experienced frequent interruptions in growth during interglacial cycles and more sustained growth during glacial cycles in the Florida Peninsula. Growth variability in deer is primarily influenced by seasonal availability of browsing resources and environmental conditions (Geist, 1998), which would have been less favorable during times of high aridity. Thus, I predict that specimens collected from interglacial sites will have smaller limb sizes and lower annual growth rates recorded in frequent LAGs and/or annuli (i.e. deer had more periodic/seasonal interruptions in growth) due to unfavorable seasonality changes (cool and dry summers and winters) brought on by more arid climate that resulted in fewer, more dispersed forest environments. In contrast, glacial environments of Florida would have had much more abundant resources due to more rainfall and generally a warmer, wetter environment resulting in abundant forests.

Therefore, specimens collected from the glacial units should be larger in limb size and preserve histological features that would indicate a faster annual growth rate along with fewer LAGs and/or annuli

(i.e. more sustained growth). This hypothesis will be tested using two approaches: 1.) Measuring and analyzing changes in limb bone dimensions (used as a proxy for body size) to determine whether body size in O. virginianus fluctuated based on differences in environmental/climatic conditions (glacial vs. interglacial), chronology (early vs. middle, vs. late Pleistocene) and geographically (north vs. south

Florida); 2.) Measuring and analyzing changes in frequency and thickness of growth zones in O. virginianus limb bones is predicted to reflect changes in growth among the different conditions mentioned above.

Important Abbreviations in order of appearance:

FLMNH – Florida Museum of Natural History……………………………………………………7

NALMA – North American Land Mammal Ages…………………………….………………...…7

ING1A – Inglis 1A……………………………………………….…………………………..…….8

LSP1A – Leisey Shell Pit 1A………………………………………………………………………8

COL2A – Coleman 2A……………………………………………………………….……………8

NH – Nichol’s Hammock………………………………………………………………………….8

AL – Articular Length……………………………………………………………………………13

MDC – Mid-diaphyseal Circumference………………………………………………………….13

LAG(s) – Line(s) of Arrested Growth……………………………………………………………14

EFS – External Fundamental System…………………………………………………………….18

BGMs – Bone Growth Marks……………………………………………………………….……26

GZ – Growth Zone (ex. GZ1 = space between the innermost cortex and the first LAG)………..31

APD – Anterior Posterior Diameter…………………………………………………….………..31

APDGR – Anterior Posterior Diameter Growth Rate…………………………………………….31

M/mAPDGR – Maximum/minimum Anterior Posterior Diameter Growth Rate………..………32

ACT – Anterior Cortical Thickness……………………………………………………………....32

M/mGR – Maximum/minimum Growth Rate (Sanders and Tückmantel, 2003)…………….…..32

GRST – Growth Rate during a Standard Year (365 days)……………………………………….32

GRGP – Growth Rate during a Growth Period (260 days)………………………………...…….32

M/mANTGR – Maximum/minimum Anterior Growth Rate………………………….…………33

OD – Osteocyte Density………………………………………………………………………….35

DA – Diagenetic Alteration……………………………………………………………..………..57

EOAPD – Early Ontogenetic APD……………………………………………….………………97

EO – Early Ontogenetic…………………………………………………………...…………….110

EOGR – Early Ontogenetic Growth Rate………………………………………………….……111

Methods

Limb bone measurements

Overall comparison of body/skeletal size and bone measurements can provide a broad measure of an organisms’ growth through its lifetime. Although body/skeletal size cannot directly be calculated from disarticulated unassociated fossil specimens, articular length (AL) and mid-diaphysis circumference

(MDC) of individual limb bone elements have been shown to be a good proxy for body size in mammals

(Alexander et al., 1979; Anderson et al., 1985; Purdue, 1987; Gingerich, 1990).

Mid-diaphyseal circumference (MDC) and articular length (AL) were measured for 462 unassociated fossil and 481 associated modern deer limb elements: (humeri, radii, metacarpals, femora, tibiae and metatarsals) housed in the collections at the Florida Museum of Natural History (FLMNH) from various localities in Florida (Fig. 2 and S1). MDC was measured at the approximate mid-point of the shaft of an individual bone (if complete) using a plastic flexible measuring tape and recorded to the nearest .01 cm (Fig. 3). If the shaft was broken anywhere along the diaphysis then the closest possible measurement was taken to the mid-diaphysis. MDC was not recorded on any of the following: 1) Partial specimens that were broken too far proximally or distally along the shaft; 2) Radii that were fused to the ulna; 3) Shafts that partially broken, or contain a large crack or separation altering their MDC. AL measurements were recorded as the maximum length of the bone from one articular surface to the other due to partial breaks along proximal or distal projections (Fig. 3). Length was recorded using a caliper to the nearest .01 cm. Any pathological specimens observed were not included in MDC of AL analyses.

Three ontogenetic stages (juvenile, sub-adult, adult) were determined by the degree of epiphyseal fusion (Purdue, 1983; Flinn et al., 2013; Table 2); sex was only available for some modern specimens

(Table S2). Fossil sites were categorized into one of the three North American Land Mammal Ages

(NALMA); Blancan: (4.7 – 1.4 Ma), Irvingtonian: (1.4 Ma – 210 ka), Rancholabrean: (210-11.7 ka) (Fig.

1; sensu Barnosky et al., 2014). Elements from Nichol’s Hammock (~500 years old) were kept separate from modern (collected post 1900) as they likely represent a pre-Columbian age.

MDC and AL analyses were conducted on bones that were confirmed adults (i.e., had either fused epiphyses or an External Fundamental System (EFS) in thin-section, indicating skeletal maturity;

Woodward et al., 2013; Table 2), (150 fossil and 223 modern). Individual limb bone measurements were grouped by right or left side, geochronology (using NALMA and modern time references; described above), latitude, and glacial/interglacial/unknown designation.

Florida is separated into by four different Köppen-Geiger climate zones based on latitude, and are assumed to have been present in the past (Fig 4; Köttek et al., 2006). These modern differences were accessed to determine how to separate Florida into differing geographic regions via daily precipitation and temperature data from 12 latitudinally separate localities (www.ncdc.noaa.gov; Tables S3-S4) from the earliest record (late 1800s to early 1900s) – Dec. 2014. Localities were selected based on longevity of the data and general central location within the Florida peninsula (except for Tallahassee and

Jacksonville) to account for differences due to coastal proximity. It was assumed that latitudinal differences were more pronounced than longitudinal differences, although some longitudinal differences were observed in the data. In days which yielded any negative data values (ex. -9999) all data were excluded even if only one measurement was erroneous. Data was sorted; monthly averages were calculated and plotted using Microsoft Excel 2007, and yearly temperature and precipitation averages were recorded based upon monthly averages (Table S4).

Figure 2. All fossil localities and modern counties where material was obtained for morphometric and histological analyses. Regions of Florida color coded based on latitudinal similarities in climate over the past ~100 years for geographical comparisons. Localities 1 – 4 represent histologically sampled populations for this study. Grey squares and corresponding numbers represent other fossil sites of unknown environment sampled for measurement data. Modern sites and italicized numbers represent number of individual specimens from each county which were sampled for measurement data. NALMA for fossil sites as follows; Blancan (1 and 5); Irvingtonian (2 and 3); Rancholabrean (6 – 13); as well as geochronological distinctions: Holocene (4); Modern (all modern individuals). Abbreviations as follows:

Rainbow Springs Run Bear Site (RBS), Lecanto 2A Victoria Site (VS), all others see above text.

MDC

Figure 3. Example of methods for measuring Mid-Diaphysis Circumference (MDC) and Articular Length

(AL) from a humerus. MDC was measured at the approximate mid-point of the shaft. AL was measured from the distal most end of the element to the articular head, rather than the greater tubercule, as the latter was missing or damaged in some specimens. Measurements were repeated until consistent results were obtained.

Table 2. Age classes for deer based on epiphyseal fusion (Based on Purdue, 1983; Flinn et al., 2013). Abbreviations: Months = mths

Element Juvenile Sub-Adult Adult Humeri Unfused (0-8 mths) Fused distal, unfused proximal (12-26 mths) Both Fused (>38 mths) Radii Unfused (0-2 mths) Fused proximal, unfused/beginning to fuse distal (5-20 mths) Both Fused (>26 mths) Metacarpal Fused proximal (fusion occurs prior to birth)*, unfused distal (0-26 mths) Both Fused (>28 mths**) Femora Unfused (0-20 mths) Fused proximal unfused/beginning to fuse distal (20-26 mths) Both Fused (>26 mths) Tibiae Unfused (0-14 mths) Fused distal, unfused/beginning to fuse proximal (20-26 mths) Both Fused (>26 mths) Metatarsal Fused proximal (fusion occurs prior to birth)*, unfused distal (0-17 mths) Both Fused (>26 mths) *Based on Lewall and Cowan, 1963; Carden and Hayden 2006)

**Based on earliest timing of significant probability of fusion (p<.05) (see Table 2 of Flinn et al., 2013).

Figure 4. Köppen-Geiger climate zones of Florida (Modified from Donelan, 2012; based on data from

Köttek et al., 2006). White boxes are locations where climate data was collected and used in this study

(see Table S3).

The final grouping consisted of hypothesized glacial or interglacial climate based upon published ecological faunal composition and proximity to sea level, isotopic analysis of tooth enamel (a proxy for

C3 vs. C4 dominated-floral assemblages), and paleobotanical information (Fig. 2; Table 1). Presence of marine organisms within a terrestrial setting indicating a sea level rise with enriched mammalian enamel

δ13C suggests an interglacial stage (Morgan and Hulbert 1995; DeSantis et al., 2009); presence of migratory birds/mammals with western and South American affinities (Great American Biotic

Interchange; Stehli and Webb, 1985) due to a fall in sea level and depleted enamel δ13C (DeSantis et al.,

2009) suggests a glacial stage (Table 1).

Group means and standard deviations were plotted against geochronologic assignment in

Microsoft Excel 2007. Kruskal-Wallis tests were run using SPSS (Statistical Package for Social Science,

Inc. version 14 and 22) to test the following hypotheses: 1) limb bone size is significantly different

(p<.05) through time (geochronologic age); 2) limb bone size is significantly different among localities from different latitudes. Where Kruskal-Wallis tests revealed a significant difference (p<0.05), an

ANOVA contrast test was run along with post hoc Gabriel and Games-Howell tests. Groups with low sample size (<2) were excluded from Kruskal-Wallis tests and Mann-Whitney U Test were run against the two remaining groups. In ANOVA Contrasts the user creates different contrasts, assigning groups coefficients which add up to zero (i.e. three groups: [(-1, .5, .5); (.5, -1, .5); (.5, .5, -1); (0, -1. 1); (-1,0,1);

( -1,1,0)]. The test runs the contrasts against one another and outputs all the results in one file. Levene’s test for homogeneity was used to test for equal variances among grouping. If the null hypothesis of equal variance was not rejected (p>0.05) then resulting ANOVA homogeneous significance values were reported along with post hoc test Gabriel test, due to slight differences in values of data (Field, 2009). If

Levene’s test revealed a significant difference (p<0.05) then non-homogeneous values from the ANOVA contrasts were reported along with post-hoc Games-Howell test (Field, 2009). In most cases both Gabriel and Games-Howell tests were both run to control for low sample data present within each pooled group.

Climate stage (interglacial or glacial) was taken into account when interpreting significant differences

between individual localities, although this was not possible for the majority of fossil sites as this information could not be decoupled from geochronologic or geographic factors (Fig. 2; Table 1).

Bone Histology

For histological analysis, 54, unassociated left (where available) humeri, radii, femora and tibiae,

(elements that tend to best preserve growth patterns; Sander and Tückmantel, 2003; Kolb et al., 2015a;

Jordana et al., In Press), were selected from the FLMNH vertebrate paleontology collection. Sampled fossil localities were selected based on three criteria: 1) number of available specimens; 2) interpreted depositional environment; 3) age approximation for locality beyond general NALMA (Fig 2; localities

#1-4). Ontogenetic stage (juvenile, sub-adult, adult) for each individual was determined based on degree of epiphyseal fusion (Table 2) and presence of an (EFS). Each specimen was sketched and photographed using a Nikon D3100 with a 60 mm lens and a Canon Powershot D10, AL, MDC, and approximate location of thin section was recorded for each bone. Photographs recording anatomical directions for each specimen were taken during each stage of the sectioning process.

Molding and Casting Procedure

Molds of whole bones were created as requested by FLMNH, following Lamm (2013). A base of

“Plastalina” (Van Aken) clay was rolled out into a 1 cm thick sheet, large enough to provide ~12 cm of space between the bone and the edge of the base. The bone was placed horizontally on its side onto the base; clay was built up around the bone until it covered approximately half the bone. A moat was created approximately 2 cm from the bone on all sides but the proximal end, with perpendicular key troughs and circular key holes at each corner of the clay base to provide a secure fit to both halves of the mold. Two cone shaped pour spouts were constructed, positioned at the proximal epiphysis. Clay walls on the outside of the circular key holes were constructed slightly higher than the highest part of the bone. An acetone- based glue (Glyptol) was brushed onto the bone and inside the moat and troughs as a separator; vaseline was used as the separator on the rest of the clay base. A two part silicon material (Mold Max 20) was mixed in ~140 g batches, vacuumed in a Shel Lab SVAC2E economy vacuum chamber for 7 minutes at

23 mm Hg, and poured onto the base until the whole bone was covered by at least 2-3 cm of silicon. Any air bubbles forming at the top of the silicon were scraped away using a wooden stick prior to setting. The mold was allowed to set at room temperature for at least 24 hours. Once set, the mold was flipped over, the clay base and walls were removed and the process repeated for the other side of the bone.

To cast the bone, both halves of the two part mold were sprayed with mold release, allowed to dry, secured together and positioned vertically. A two part urethane casting plastic (TC-808) was gently mixed for 30 seconds and poured into one of the pour spouts on the mold, allowing any air bubbles to exit through the other spout. The cast was allowed to set (~30 minutes), then excess material was trimmed and cast was sanded down to smooth out edges.

Thin Sectioning Procedure

Thin sections were created by embedding a ~2 cm section of the diaphysis of each bone following standard procedure of Lamm (2013). The marked 2 cm section of the bone was removed from each bone using a Buehler Isomet low speed saw. A base (~1 cm) of resin (Silmar 41 clear polyester casting resin mixed with catalyst MEKP) was poured into a plastic cup and allowed to set overnight. This base was removed and the entire interior of the cup was sprayed with mold release and allowed to dry. The cut section was placed horizontally into the cup; Silmar 41 resin was mixed with catalyst and poured on top of the bone until it was 1-2 cm above the bone. The section was then placed into a Shel Lab SVAC2E economy vacuum at ~23 mm Hg for 7 minutes to allow the resin to infiltrate into pore spaces within the bone. The vacuum was slowly released and excess air bubbles on the surface of the resin were scraped to the side. The section was then allowed to polymerize slowly via refrigeration for 24 hours. The section was then removed and allowed to harden at room temperature for 24 more hours.

The embedded section was trimmed to form a rectangular block using a 10’ rock saw (Highland

Park Manufacturing), keeping at least a 1 cm spacing of resin surrounding the bone. Each bone was cut transversely at approximate mid-diaphysis (or as close as would provide an entire cross-section) using the low speed saw. Five minute epoxy was applied to both cut halves for stability and allowed to dry overnight. A ~2 mm slice was cut out from one of the block halves and ground down using increasingly

fine Struers MD-Piano magnetic grit discs on a Leco VP-50 grinding wheel to create a smooth surface.

Glass slides (3 x 2 with one side frosted) were cleaned using Kim Wipes; one was placed underneath and another on top (frosted size up) of a precut piece of wax paper (long enough to be folded over the slide).

The bone slice was air-dried and 2-Ton Epoxy was applied to the side to be glued. The slice was then placed onto the slide and pressed firmly down. Wax paper was folded over and two more glass slides were placed on top for stability. Clothes pins were used to apply equal pressure between the slides and the section and allowed to cure for 12 hours; clothes pins were removed and the section cured for another ~8 hours.

The glued section was placed onto a (Hillquist thin section machine) and was ground down using a diamond studded grinding wheel. Periodically the section was checked using a digital micrometer

(Mahr) until a thickness of ~1.2 – 1 mm was reached. The section was then manually ground down using increasingly fine magnetic discs and grit paper on grinding wheels (Leco VP 50/160), until either a thickness of ~200 - 100 microns was achieved or histological features were clearly visible via transmitted light. Slides were then polished to remove scratches and labeled with specimen number and anatomical direction, accordingly.

Histologic Descriptions

Histologic descriptions use terminology by Francillon-Vieillot et al. (1990) and Huttenlocker et al. (2013). Lines of arrested growth (LAGs) are defined as circumferentially oriented lines that represent a period of cessation of somatic growth, whereas annuli are circumferential zones of avascular lamellar or parallel-fibered tissue that represent a significant reductions (but not total stoppage) of growth

(Huttenlocker et al., 2013) (Fig. 5). These lines or zones may be cross cut by heavily remodeled regions caused by muscle attachments (ex. posterior of femora). Other identified growth lines that did not extend circumferentially around the entire cortex or were cut off by entering the medullary cavity were labeled as bone growth marks (BGMs).

Bone structure in modern and fossil vertebrates have been extensively studied and provide a basis for bone interpretation for the fossil record (Enlow and Brown, 1957, Enlow, 1963, 1969; de Ricqlés,

1975, 1991, Castanet et al., 1996, 2000). There are generally three types of bone matrix: woven-fibered, lamellar, and parallel fibered bone (Francillon-Vieillot et al, 1990; Huttenlocker et al., 2013; Fig. 5). The bone matrix from mammals, dinosaurs and birds is mostly composed of woven-fibered bone

(disorganized collagen fibers that are loosely and randomly arranged) and typically can contain numerous vascular canals (holes where blood vessels bisect the bone). Together this complex is known as fibrolamellar bone and can be identified by its isotropic nature (wave-like extinction) under cross polarized light (xpl; Fig. 5F). The orientation of the vascular canals can be broken down into five sub groups: radial

(“radiating” pattern from the medullary cavity), reticular (haphazardly oriented but well connected), plexiform (circumferential orientation with ‘step-like’ connections), laminar (circumferential oriented with few connections) and longitudinal (circular with no connections) (Huttenlocker et al., 2013; Fig. 5A-

D). Generally, larger sized canals with higher connectivity indicate the fastest growth (De Margerie et al.,

2002; Huttenlocker et al., 2013), although all types of vascular canals can pass into each other and be present within an individual section (Huttenlocker et al., 2013). Also, inter-element variation may occur between bones; thus, vascular canals may not exclusively indicate ontogenetic stage and/or whole organism growth rate (De Margerie et al., 2004) (i.e. compare Fig. 5B, C, D). This is why it is important to sample multiple elements from different individuals, rather than just one bone category.

Lamellar bone is comprised of closely packed collagen fibers formed into thin layers (lamellae) and is deposited in alternating layers resulting in light and dark lighting patterns as the microscope stage is rotated under cross polarized light (xpl) (Huttenlocker et al., 2013; Fig. 5E-F). Lamellar bone has the slowest apposition rate and frequently can contain LAGs (Huttenlocker et al., 2013). This type of bone is commonly found in non-archosaurian diapsids, but can be also be found among some mammals

(Huttenlocker et al., 2013) especially within annuli and or within the external cortex (Figs. 5-6). Parallel- fibered bone is composed of tightly packed parallel collagen fibrils and is anisotropic (Huttenlocker et al.,

2013; Fig. 5F-G). Typically osteocytes found within this bone matrix are oriented parallel to apposition, indicating a slowed growth rate (Huttenlocker et al., 2013). This bone type was also common within some humeri making up individual annuli (Fig. 5 D-E). It was also common along the outer cortex of some

individuals where an EFS had not yet developed (Fig. 5). Both lamellar bone and parallel-fibered bone typically do not contain many vascular canals and if present, they are longitudinally oriented.

Histological sections were photographed and measured using both Leica DM EP and Leica Z6

APO microscopes equipped with a camera attachment (Diagnostic Instruments, INC) and 2x lens, running

SPOT Advanced 4.6 software on a computer running Microsoft Windows XP operating system. Multiple images were photographed for each section; merged to form a composite image using Adobe Photoshop and Illustrator CS5 and printed onto 11 x 17 inch sheets of paper. LAGs were manually identified and traced onto printed sections, verified under a microscope, and traced onto a digital file using Adobe

Photoshop.

Figure 5. Typical bone structure in O. virginianus specimens for this study. A)UF300290: Juvenile humeri with woven-fibered bone with large longitudinal canals. B) UF300921: Juvenile-Subadult radii composed of fibro-lamellar bone and reticular vascularity. C)UF300295: juvenile femora composed of fibro-lamellar bone with mixed vascularity. D) UF300299: Sub-adult humeri with intermixed plexiform and laminar canals with annuli (red box, E) showing parallel fibered bone; note parallel oriented osteocytes. F) UF65962: sub-adult to adult humeri with both fibro-lamellar bone and annuli with lamellar bone; note laminae; taken under xpl. All Scale bars = 1 mm except G = .01 mm.

A B

Figure 6. Other histological features within O. virginianus specimens used in this study. A) UF57165:

Adult Humeri with an EFS along the outer cortex. B) UF300297: Posterior section of Juvenile femora with nutrient foramina and longitudinal to plexiform vascularity. C) UF45044: Posterior section of sub- adult to adult (8 years old but no EFS), showing multiple generations of secondary osteons. All scale bars are 1 mm.

Skeletochronology

Three separate methods were used from previous studies in artiodactyls (Kolb et al., 2015a;

Jordana et al., In Press) and sauropods (Sander and Tückmantel, 2003), to compare and contrast growth rates among different populations. Due to the nature of the non-associated fragmented fossil material, it was impossible to determine the exact age and sex of the individual animals; thus, a method of retrocalculation applied in previous artiodactyl studies (Marín-Moratalla et al., 2013; Jordana et al., In

Press) was used to determine the ontogenetic position of each LAG and growth zone (GZ; e.g. GZ2 = distance between LAG1 and LAG2). Previous research found that GZ1, LAG1 and portions of GZ2 can be missing in older individuals (Marín Moratalla et al., 2013; Kolb et al., 2015a; Jordana et al., In Press).

To resolve this issue the anterior-posterior diameter (APD) of successive LAGs for each specimen were recorded and plotted against other successive APDs from variously aged individuals (Fig. 7). The slopes of the growth curves were fitted to one another (retrocalculated) to determine the placement of the

“unknown” successive LAG sequences. This method of retrocalculating age has been supported using other age methods for artiodactyls based on tooth eruption and dental wear (Marín-Moratalla et al., 2013).

Based on these publications, this study assumed that the retrocalculation of APDs from individual LAGs would best represent the ontogenetic position of individual LAGs for unassociated specimens.

Using SPOT, APDs (in mm) were recorded on digitized merged photos and values were graphed vs GZ using Excel 2007. APDs were measured from all visible LAGs up until the start of the EFS where visible (Fig. 7). LAGs that were partially resorbed along the anterior/posterior axis (as in humeri) were not measured. Each APD measurement started and finished at the first visible successive LAG (even for multiple lined LAGs). For individuals with no visible LAGs and determined to be under 1 year of age

(based on epiphyseal fusion), whole section APD was taken and graphed representing growth within the first year (called “LAG0.5”).

The first method for determining growth rate used APD measurements (APDGR); the difference between each LAG (n) APD and the successive LAGn-1 APD was divided by 365 (standard yearly growth

rate; minimum (m) and 260 days (growth rate during seasonal growth period; maximum (M); Köhler et al., 2012). M/mAPDGRn was then plotted (per individual) against GZ in Excel as well as averages from each locality for each GZ. Average growth rates for GZs that were represented among multiple individuals (filled shapes) are best supported while other growth rates (opened shapes) are less supported.

All juvenile growth rates along with GZ1 and some GZ2 growth rates represent a minimum value due to resorption of cortical bone.

The second method developed by Sander and Tückmantel (2003) uses anterior cortical thickness

(ACT), number of LAGs, and distances between first and last LAG, in mid-diaphyseal sections of long bones to determine a lifetime growth rate (Fig. 7). Anterior (posterior in humeri) portions of each section containing LAGs were imaged using a Leica (DM EP) petrographic microscope with a 4x lens and distances between individual LAGs, ACT and distances between first and last LAG were measured out using SPOT 4.6 (Fig. 7). Maximum growth rate (MGR) was calculated using ACT and a minimum growth rate (mGR) using distance between 1st and last LAG; both values were then divided by the number of LAGs present (Fig. 7). Each growth rate was calculated over both a standard 365 day year

(M/mGRST) as well as a 260 day year (M/mGRGP; growth period) in µm/day.

The third method utilized individual growth rate per GZ in anterior (posterior in humeri) cortical sections (ANTGR) and were calculated by taking the distance (d) between each GZn and dividing it by

365 (m) and 260 (M) days (Marín Moratalla et al., 2013; Jordana et al.,In Press).

Because cervids, other mammals and dinosaurs do not have a linear growth curve and growth rates can vary greatly through ontogeny, M/mGRST/GP may represent a vast underestimate of growth rate. Rather than determine whole bone growth rate over the individual’s life, M/mANTGR per GZ are a more representative measure of a bone’s individual annual growth rate, and can reveal changes in annual growth (Köhler et al., 2009; Marín Moratalla et al., 2013; Jordana et al., In Press). Even so, GZ1 and GZ2 in some specimens may still represent a minimum value due to cortical resorption discussed earlier. For comparisons to other studies (Kolb et al., 2015a, b; Jordana et al., In Press) both M/mGRST/GP and

M/mANTGR are reported but only M/mANTGR are utilized to create growth curves and to compare maximum growth rates between localities. In humeri, anterior regions are highly remodeled while posterior regions are not; this trend is opposite among femora and tibiae. Due to these similarities, posterior regions of humeri were analyzed similar to that of the anterior regions of other bones.

M/mANTGR averages per population were plotted against GZ in Excel and comparisons were made between populations

Figure 7. Method for measuring anterior posterior diameter (APD) and anterior LAGs distances (d) for individual thin sections in limb bones. (AR) is anterior radius; (d1st-last) is distance between first LAG to last LAG. Based on methods developed by Sander and Tückmantel, (2003), Marín Moratalla et al.,

(2013), Kolb et al. (2015a), and Jordana et al. (In Press).

Osteocyte Density

Osteocyte density (OD) was determined by counting individual osteocytes from single and stacked images of both the femora and tibiae thin sections in a standard area following methodologies of

(Bromage et al., 2009; Stein and Werner, 2013; Jordana et al., In Press). For stacked images; multiple images (with standard area 381 µm2) were taken at the center of each GZ in the anterior region of each section, with a Leica DM EP microscope using a 40x lens (Fig. 8). To control for preservational issues, the sections in GZs were chosen based on best preservation. Each image was taken by adjusting the fine focus dial 6 µm and then merged together using Adobe Photoshop C5 to form a 3-dimensional stacked image of osteocytes occupying a volume of space within the bone. Along with the stacked images a “best representative” photograph was picked out of existing images to represent a single layer to record osteocytes for comparisons with previous research (Jordana et al., In Press). In both the stacked and single layer images three 0.1 x 0.1 mm boxes were created with an attempt to obtain “pure” bone OD representative (no overlap of individual osteons, areas of resorption or vascular canals which have been shown by previous studies to affect OD (D’Emic and Benson, 2013; Stein and Werner, 2013; Fig. 8).

These boxes were replicated in the same location on both the stacked and single images to control for inter-section variation. Recorded osteocytes were either touching or inside the created box and clear enough to see a well-defined ovate shape and/or individual caniculi; then were physically counted and digitally marked using Adobe Illustrator CS5 (Fig. 8). OD was then calculated by taking the individual counts in each box and dividing it by the area (single layer) or volume (stacked) of the standard square:

(OD of single layer = # of osteons/0.01 mm2; OD of volume = # of osteocytes/area x (# of stacked images x .006 mm)) and OD was averaged within each GZ. The OD was then plotted against individual GZs in

Excel and compared within and among populations.

Figure 8. Methodology for osteocyte counting. A) Anterior section of femur UF239925; boxes (381 x

381 µm) were created in each GZ. B) Stacked image of GZ3. Within each box three more boxes (100 x

100 µm) were created for osteocyte counts. C) Osteocytes with caniculi (faint hair-like protrusions from the osteocyte). Scale bars: A = 1 mm; B = 381 µm; C = 15 µm.

Results

Climate differences across the Florida Pennisula

Averages for climate data are presented in the Appendix Supplementary Tables S4.

Temperature Differences across Florida

Average annual temperatures across Florida vary along a North/South latitudinal gradient where north Florida (Tallahassee, Lake City, Jacksonville and Gainesville) experiences the lowest average temperature (<21°C) and the Everglades the highest (23-24°C) with central Florida (21-22°C; Ocala and

Crescent City) and southern Florida (>22-23°C; Orlando, Bartow, Avon Park, Arcadia and Royal Palm

Springs) in between (Fig.9A, Table S4).

Latitudinal precipitation changes across Florida

Average annual precipitation across the Florida peninsula is relatively constant and does not vary by latitude (~36 cm/ year) except for Tallahassee, which received well above the average (~43 cm/year;

Fig 9B, Table S4). Average precipitation by season does vary from high to low latitude; wet season: June

– September and dry season: October – May (Fig. 9C-D). These differences seem to coalesce around

Orlando, where localities south of and including Orlando experience less precipitation (<24 cm/yr) than locations north of Orlando (>25 cm/year) during the dry season, while areas north of and including

Orlando receive less precipitation (<60 cm/year) than areas south of Orlando during the wet season (>64 cm/year; Figs. 9C-D). Further, the difference between average dry vs wet seasonal precipitation reveals that locations north of Orlando received less precipitation (<33 cm/year) than the locations south (>38 cm/year; Fig 9E). Thus there seems to be a latitudinal difference in seasonal precipitation between northern and southern Florida.

These climate differences are assumed to have been present in Florida in the past although their specific placement and extent likely was different. Nevertheless, these modern climate differences provide a guide used to differntiate geographical sections of Florida (north and south; at 28.5°N; Orlando,

Florida; see Fig. 2) for use in statistical testing.

A Average Yearly Temperatures B Average Yearly Precipitation 25 45

C) ° 30 15 25

20 10

15 Average Temperature( Average

5 10 Average Precipitation Average (cm/Year) 5

0 Latitude 0 Latitude

Figure 9. Yearly average temperatures and precipitation across latitude (high to low; left to right) in Florida. A) Average yearly temperatures; B) Average Yearly Precipitation; C) Average “Dry Season” precipitation; D) Average “Wet Season” Precipitation; E) Average Precipitation difference between dry and wet seasons. All localities are in order of latitude from highest to lowest except for the Tallahassee, Lake City and Jacksonville which best represent longitudinal differences along a similar latitude. Red lines (A, C-E) represent placement of north-south latitudinal line marking a difference in climate.

Figure 9 Cont. C Average Dry Season (October-May)

Precipitation 40

35 30 25

15 10

5 Average Precipitation Average (cm/season) 0 D Average Wet Season (June-September) Precipitation 80

70 60 50 E 40 30 20

10 Average Precipitation Average (cm/season) 0 Difference in average precipitation between wet and dry seasons

10 Average Precipitation Average (cm)

0 Latitude

MDC and AL comparisons across time and geography

Measurement data, sex, ontogenetic stage and locality information from all fossil and modern individuals used in this analysis are presented in the Appendix Supplementary Tables S1-2.

Left vs. Right Elements

All fossil elements were found in sinkhole deposits and were labeled as unassociated with other material. There were no significant differences in AL or MDC between either left or right elements of similar geography and geologic age (Table S5). Thus, left and right elements of similar geography and geochronologic age were combined in order to provide larger sample sizes for statistical testing.

Northern Florida MDC and AL vs. geochronology

In northern Florida, adult Rancholabrean tibiae were significantly longer than modern tibiae, with no significant differences for any other element (Tables 3 and 4). Mean adult MDC of all elements decreased significantly over time (Fig. 10; Tables 5-7). The greatest observable chronologic change in average adult MDC came from radii (Fig. 10; Tables 5-6).

Among radii from northern Florida, mean MDC significantly decreased from Blancan and

Rancholabrean populations to Modern populations (p= 0.009; p= 0.038 respectively; Fig. 10; Tables 5-6), but not between Blancan to Rancholabrean populations (p= 0.293; Fig. 10; Tables 5-7).

Metatarsals from northern Florida provided the largest sample of individuals across all chronologies and a significant decrease was observed in adult MDC between fossil and Modern populations (p = 0.015; Table 5-7). These differences were most prevalent between Blancan and

Rancholabrean when compared with modern populations, while the Irvingtonian population showed no difference to the modern but also had a smaller sample size (n=3; Fig. 10; Table 5-7).

Adult MDC from north Florida metacarpals significantly decreased between fossil and modern specimens (Fig. 10; Tables 5-7). ANOVA contrasts revealed significant differences between each individual case except for those which compared Blancan and Rancholabrean against one another (Table

6). Further, mean MDC values from Rancholabrean overlap directly with Blancan populations (Fig. 10;

Table 7).

Significant differences were observed from northern Florida femora MDC among the Blancan populations when compared to Modern (p = 0.045; Figs. 5 and 7), although no significant difference was observed between Rancholabrean and modern populations (p = 0.068; Fig. 10; Tables 5-7). Pooled fossil populations revealed significant differences when compared to modern populations in ANOVA tests (p =

.021; Table 6) and mean MDC exhibits the greatest decrease following the Rancholabrean (Fig. 10; Table

7).

Table 3. Statistical comparison of limb bone AL from northern Florida O. virginianus over the past ~2 million years. Significant differences are in bold North Florida AL vs. Geochronology N (number of individuals) Kruskal-Wallis Mann Whitney U Element Blancan Irvingtonian Rancholabrean Modern χ² Asymp. p (2-tailed) Z Asymp. p (2-tailed) Femora 5 - 10 18 2.351 0.309 - - Tibiae - - 8 21 - - -2.489 0.013 Metatarsals 6 1* 4 16 .855 0.652* - - Humeri 4 - 6 18 3.914 0.301 - - Radii 2 1* 12 16 1.347 0.510* - - Metacarpals 4 - 5 15 2.402 0.301 - - * Blancan or Irvingtonian sample removed from statistical testing due to sample size <2.

Table 4. AL means and standard deviations for adult elements from northern Florida with significant differences among geochronology.

North Florida AL vs. Geochronology: Standard Deviation Element Time Mean (cm) Standard Deviation Tibiae Rancholabrean 266.53 5.77 Modern 260.60 14.86

Table 5. Comparison of limb bone MDC from northern Florida O. virginianus during the past 2 million years. Significant differences are in bold.

North Florida MDC vs. Geochronology N (number of individuals) Kruskal-Wallis Mann Whitney U ANOVA Contrast

Asymp. Asymp. Element Blancan Irvingtonian Rancholabrean Modern χ² Z p (2-tailed) p (2-tailed) Femora 9 1* 11 19 6.602 0.037* - - ** Tibiae 1* 1* 10 22 8.379 0.015 -2.797 .004* - Metatarsals 6 3 5 16 12.410 0.006 - - ** Humeri 6 1* 8 18 7.458 0.024* - - ** Radii 6 - 13 16 9.405 0.010* - - ** Metacarpals 4 - 6 16 10.710 0.005 - - ** * Blancan or Irvingtonian sample removed from statistical testing due to sample size <2. ** See Table 6

Table 6. P-values for ANOVA contrast tests for MDC of adult limb elements from northern Florida O. virginianus that showed significant differences in the Kruskal Wallis test. Levene’s Test of equal variances determined which values to use for the contrasts. Contrasts compared pooled geochronologic NALMAs with one another; B = Blancan; I = Irvingtonian; R= Rancholabrean; M = Modern. Significant values are bold.

North Florida MDC vs Geochronology: ANOVA Femora Levene’s Test = 0.263 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (B vs. R) 0.779 (B vs. R) 0.989 0.966 (B vs. M) 0.045 (B vs. M) 0.119 0.066 (R vs. M) 0.068 (R vs. M) 0.182 0.292 (B + R vs. M) 0.021

(B + M vs. R) 0.425

(B vs. R + M) 0.216

Metatarsals Levene’s Statistic = 0.076 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (B + I + R vs. M) 0.015 (B vs. I) 0.070 0.310 (B + I + M vs. R) 0.066 (B vs. R) 0.961 0.912 (B + R + M vs. I) 0.231 (B vs. M) 0.003 0.160 (I + R + M vs. B) 0.133 (I vs. R) 0.319 0.373 (I + R vs. M) 0.150 (I vs M) 1.000 1.000 (I + M vs. R) 0.028 (R vs M) 0.052 0.0 (R + M vs I) 0.398 (B + I vs. R) 0.402 (B + R vs I) 0.109 (I + R vs B) 0.185 (B + R vs M) 0.004 (B + M vs R) 0.297 (R + M vs B) 0.166 (B + I vs M) 0.110 (B + M vs I) 0.292 (I + M vs B) 0.053

Table 6. Cont.

Humeri Levene’s Statistic = 0.116 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (B + R vs. M) 0.031 (B vs. R) 0.883 0.402 (B + M vs R) 0.067 (B vs. M) 0.839 0.471 (R + M vs. B) 0.993 (R vs. M) 0.081 0.007 (R vs. M) 0.016 (B vs. M) 0.257 (B vs. R) 0.328 Radii Levene’s Statistic = 0.173 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (B vs. R) 0.293 (B vs. R) 0.626 0.531 (B vs. M) 0.009 (B vs. M) 0.021 0.013 (R vs. M) 0.038 (R vs. M) 0.107 0.148 (B + R vs. M) 0.005 (B + M vs. R) 0.703 (R + M vs. B) 0.046 Metacarpals Levene’s Statistic = 0.420 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (B + R vs. M) 0.001 (B vs. R) 0.647 0.622 (B + M vs. R) 0.664 (B vs. M) 0.007 0.121 (R + M vs. B) 0.034 (R vs. M) 0.063 0.011 (B vs. R) .303 (B vs. M) .003 (R vs. M) .026

Table 7. MDC means and standard deviations for adult elements from northern Florida with significant differences among geochronologic units.

North Florida MDC vs. Geochronology: Standard Deviations Element Time Mean Standard Deviation Element Time Mean Standard Deviation Femora Blancan 6.90 0.60 Humeri Blancan 6.09 0.43 Rancholabrean 6.81 0.95 Irvingtonian - - Modern 6.32 0.53 Rancholabrean 6.36 0.23 Modern 5.82 0.58 Tibiae Irvingtonian - - Radii Blancan 5.98 0.40 Rancholabrean 6.60 0.31 Rancholabrean 5.70 0.70 Modern 6.16 0.58 Modern 5.27 0.39 Metatarsals Blancan 5.8 8 0.47 Metacarpals Blancan 5.55 0.3 9 Irvingtonian 5.39 0.38 Irvingtonian - - Rancholabrean 5.76 0.10 Rancholabrean 5.34 0.17 Modern 5.39 0.18 Modern 5.00 0.31

Figure 10. Graphs of adult limb elements that show significant geochronologic changes in MDC of O. virginianus populations from northern

Florida. Error bars represent standard deviation (Table 7); No error bars represents a single specimen. Trend lines in red show decline in MDC.

Southern Florida MDC and AL vs. geochronology

Lower sample size was prevalent among southern Florida fossil populations. Only comparisons between Nichol’s Hammock (~500 yrs) to Modern radii AL revealed a significant increase across time

(Table 8-9). A significant decrease in MDC of metatarsals between Rancholabrean and Modern populations (p = 0.003) and also between pooled Irvingtonian and Rancholabrean vs. Modern populations

(p = 003; Fig. 11; Tables 10-12) was observed. Although other elements (humeri, metacarpals and tibiae) revealed non-significant p-values across chronology, mean MDC from Rancholabrean populations (n ≥ 2) show a higher mean MDC relative to the Modern populations (Fig. 11; Tables 11-12). This likely resulted in the observed p-values which were closer to 0.05 (Table 10). Unfortunately, adult femora and radii were only available from NH populations and modern which show no significant difference (Fig. 11; Tables

10, 12). A larger sample of southern Florida adult limb elements may provide additional evidence for these increases in MDC observed among Rancholabrean populations (Fig. 11).

Table 8. Statistical comparison of AL of southern Florida adult O. virginianus limb elements over the past ~1.5 mys. Significant values are in bold.

South Florida AL vs. Geochronology N (number of individuals) Mann Whitney U Kruskall-Wallis

Asymp. p-value Asymp. p-value Element Irvingtonian Rancholabrean Nichol’s Hammock Modern Z χ² (2-tailed) (2-tailed) Femora 1* 1* 3 16 -0.671 0.502* Humeri 4 2 4 16 - - 2.513 0.473 Radii 1* 1* 3 12 -2.021 0.043* 6.288 0.098 Metatarsals 2 3 - 15 - - 2.654 0.265 Metacarpals 2 2 2 16 - - 2.824 0.420 Tibiae 1 1 - 15 - - 1.579 0.454 * Irvingtonian and/or Rancholabrean samples removed from statistical testing due to sample size <2.

Table 9. MDC means and standard deviations for adult elements from southern Florida with significant differences among geochronologic units.

South Florida MDC vs Geochronology: Standard Deviations Element Time Mean Standard Deviation Radii Nichol’s Hammock 176.66 9.41 Modern 191.58 12.95

Table 10. Statistical comparison of MDC of southern Florida adult O. virginianus limb elements over the past ~1.5 mys.

South Florida MDC vs. Geochronology N (number of individuals) Mann Whitney U Kruskall-Wallis

Asymp. p-value Asymp. p- Element Irvingtonian Rancholabrean Nichol’s Hammock Modern Z χ² (2-tailed) value (2-tailed) Femora - 1* 3 16 -0.282 0.778* - - Humeri 4 2 4 16 - - 6.962 0.073 Radii 1* 1* 5 14 -1.022 0.307* - - Metatarsals 3 4 - 16 - - 8.541 0.014 Metacarpals 2 2 2 16 - - 6.942 0.074 Tibiae 1* 2 - 15 -1.944 0.052* - - * Irvingtonian and/or Rancholabrean samples removed from statistical testing due to sample size <2.

Table 11. P-values for ANOVA contrast tests for MDC of metatarsals from southern Florida O. virginianus that showed significant differences in the Kruskal Wallis test. Levene’s Test of equal variances determined which values to use for the contrasts. Contrasts compared pooled geochronologic NALMAs with one another; I = Irvingtonian; R= Rancholabrean; M = Modern. Significant values are bold.

South Florida MDC vs. Geochronology: ANOVA

Metatarsals Levene’s Statistic = 0.628 Post Hoc Individual comparisons ANOVA Contrasts Sig. (2-tailed) Individual comparisons Gabriel Games-Howell (I + R vs. M) 0.003 (I vs. R) 0.844 0.781 (I + M vs. R) 0.055 (I vs. M) 0.110 0.219 (R + M vs I) 0.576 (R vs. M) 0.007 0.104 (I vs R) .471 (I vs M) .054 (R vs M) .003

Table 12. MDC means and standard deviations for adult elements from southern Florida with potential differences between geochronologic units.

South Florida MDC vs Geochronologic: Standard Deviations Element Time Mean Standard Deviation Element Time Mean Standard Deviation Metatarsals Irvingtonian 6.90 0.60 Metacarpals Irvingtonian 5.16 0.11 Rancholabrean 6.81 0.95 Rancholabrean 5.60 0.42 Modern 6.32 0.53 N. Hammock 4.82 0.15 Modern 4.78 0.41 Humeri Irvingtonian 5.99 0.35 Rancholabrean 6.13 0.60 Tibiae Irvingtonian 5.85 - N. Hammock 5.33 0.33 Rancholabrean 6.70 0.35 Modern 5.46 0.53 Modern 5.8 0.43

Metacarpals 7.5 Humeri Metatarsals Tibiae

6.5

MDC (cm) MDC 5.5

4.5

4 1000 750 500 250 0 1000 750 500 250 0 1000 750 500 250 0 1000 750 500 250 0

Time (hundreds of years)

Figure 11. Graphs of limb elements that show geochronologic changes in MDC of O. virginianus populations from southern Florida. Error bars represent standard deviation; No error bars represents a single specimen.

MDC and AL vs. Geography

Adult fossil populations yielded few elements (humeri, metatarsals, and metacarpals) which could assess geographic change across a specific geochronologic time frame. Of these elements, all yielded non- significant geographic differences in both AL and MDC across north and south Florida during specific

NAMBLA time periods (Table 13). Modern adult femora and tibiae revealed significantly larger MDC and AL in northern Florida when compared with southern Florida (Tables 14-17).

Table 13. Statistical comparison of MDC and AL from Irvingtonian and Rancholabrean specimens across their geographic distribution in Florida.

Irvingtonian MDC vs. Geography Rancholabrean AL vs. Geography Mann Whitney U Mann Whitney U Element North South Z Asymp. p (2-tailed) Element North South Z Asymp. p (2- Metatarsals 3 3 -1.528 0.127 Humeri 6 2 0 1.000 Metatarsals 4 3 -1.061 tailed)0.289 Metacarpals 5 2 -1.549 0.121 Rancho labrean MDC vs. Geography Mann Whitney U Element North South Z Asymp. p (2-tailed) Humeri 8 2 1.525 0.599 Metatarsals 5 4 0 1.000 Metacarpals 6 2 -.849 .396 Tibiae 10 2 -.215 0.830

Table 14. Statistical comparison of MDC from Modern specimens across their geographic distribution in Florida. Bold numbers indicate significant differences. Modern MDC vs. Geography

N (number of individuals) Mann Whitney U

Element North South Z Asymp. p-value (2-tailed)

Femora 20 16 -2.296 0.022 Tibiae 26 15 -1.113 0.266 Metatarsals 20 16 -1.180 0.238 Humeri 21 16 -2.445 0.015 Radii 19 14 -1.500 0.133 Metacarpals 20 16 -1.857 0.063

Table 15. MDC means and standard deviations for elements of Modern populations of Florida O. virginianus with significant differences among geographic ranges.

MDC Modern populations of Florida Element Geography Mean Standard Deviation Femora North and Central 6.28 0.54 South 5.94 0.51

Humeri North and Central 5.71 0.65 South 5.46 0.53

Table 16. Statistical comparison of AL from Modern O. virginianus specimens across their geographic distribution in Florida. Bold numbers indicate significant differences.

AL of Modern populations of Florida N (number of individuals) Mann Whitney U

Element North South Z Asymp. p-value (2-tailed) Femora 18 16 -2.415 0.016 Tibiae 25 15 -1.048 0.295 Metatarsals 20 15 -0.100 0.920 Humeri 21 16 -3.112 0.002 Radii 17 14 -0.595 0.552 Metacarpals 18 16 -0.759 0.448

Table 17. AL means and standard deviations for elements of Modern populations of Florida O. virginianus across geographic ranges.

Element Geography Mean (cm) Standard Deviation Femora North 225.46 14.30 South 213.87 13.45

Humeri North 179.03 29.20 South 161.70 10.55

Potential Effects of Sexual Dimorphism

Sex could only be determined for modern populations (Tables 18 and S2). Adult specimens comprising the modern populations were mostly females with few males and many isolated bones of unknown gender (Table 18). The geographic differences observed within modern populations and differences observed when comparing modern with fossil populations may be a result of gender skewing if fossil populations contained more males within their pooled populations.

In summary, some significant chronologic differences in adult MDC exist among both northern and southern Florida populations during the past 2 million years. Generally the greatest chronologic of differences in adult MDC for these elements appears between Blancan and/or Rancholabrean populations when compared modern populations. Little to no significant chronologic difference was observed in AL across time except among northern tibiae and southern radii. Even with these observed differences, many elements lacked large enough sample sizes throughout all the chronologies tested (especially

Irvingtonian). Larger fossil sample sizes from this and other time frames could help assess whether changes in element dimensions were also apparent through geologic time. Potential gender skewing among the modern population may have impacted these results, more data (especially sexual identification) is needed to improve upon and support these trends.

Table 18. Gender of modern deer used in MDC and AL analyses

Gender North South Male 1 2 Female 10 7 ? 6 2

Histological Descriptions

The ontogenetic stage (OS), estimated age, presence/absence of an EFS, number of LAGs, bone measurements (MDC, AL), thickness and position of removed section, location of thin section along shaft, maximum anterior-posterior diameter (APD) and maximum lateral-medial diameter (LMD) of each thin section are provided in the Appendix: Table S6.

Femora

Inglis 1A

UF45040 is a partial left femur is broken along the distal shaft with a fused distal epiphysis (Fig. 12A).

The femur is composed of fibrolamellar bone with sub-reticular/plexiform (mid to outer cortex) to sub laminar canals (inner to mid cortex; Fig. 12B-C). Reticular canals are present in the posterior quadrant close to the linea aspera (Fig. 12B). Along the inner medial cortex there are numerous radial canals that continue postero-medially (Fig. 12B-D). Cortical drift is taking place medially and a zone of remodeling separates zones of radial (inner) and plexiform (outer) canals (Fig. 12B). There are LAGs and no EFS is present (Fig. 12B-C ; Table S6). Remodeling is present in the innermost postero-lateral cortex with one- two generations of secondary osteons (Fig. 12B). Some diagenetic alteration (DA) occurs along the inner and outermost cortical regions.

UF45044 is a partial left distal femur is broken mid to distal-mid diaphysis with a fused distal epiphysis

(Fig. 12E). This femur is composed of fibrolamellar bone with reticular (mid to outer cortex) to plexiform/laminar (inner to mid) canals (Fig. 12F-H). A small postero-medial section of bone along the innermost cortex has woven tissue, with some radial canals that extend from the inner to the mid-cortex and are interrupted by secondary remodeling (Fig. 12H). LAGs are present starting in the mid-cortex; the fifth LAG has multiple lines present (Fig. 12G; Table S6). No EFS is visible (Fig. 12G; Table S6). The posterior portion of the bone has undergone the most remodeling with at least two-three generations of secondary osteons (Fig. 12I). In other regions, remodeling is restricted to the innermost cortex. Laterally,

cortical drift is evident and remodeling exists between inner woven tissue and more mature outer fibrolamellar bone (Fig. 12F). A rim of DA obstructs the outermost bone.

UF300294 is a partial left femur broken proximally above the lesser trochanter with a laterally extended intertrochanteric crest with a fused distal epiphysis (Fig. 12J). This femur is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 12K-L). LAGs are present, of which the second outermost

LAG has multiple lines (Fig. 12K-L ; Table S6). What little remodeling is visible is concentrated in the posterior inner cortex and along the linea aspera (Fig. 12K). DA is prevalent along the external and internal cortex.

UF300295 is a complete left femur with unfused epiphyses (Fig. 12M). This femur is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 12N). There are no LAGs, EFS, or secondary osteons visible (Fig. 12N). Some DA is present along the outermost cortex.

UF300296 is a partial left proximal femur which is broken distally along the shaft along the medial flabella, with an unfused proximal epiphysis (Fig. 12O). This femur is composed of fibrolamellar bone with reticular (inner cortex) to plexiform/laminar (mid-outer cortex) canals (Fig. 12P-Q). A single LAG is visible along the outer mid cortex through most of the section but is obstructed by DA laterally in the outer cortex (Fig. 12P-Q; Table S6). Remodeling is very sparse throughout the section. The posterior region towards the linea aspera differs in bone type, composed of sub-reticular canals as compared to the plexiform to laminar vascular canals within the rest of the section (Fig. 12P). Some DA obscures tissue along the outer cortex.

UF300297 is a partial left femur, with unfused epiphyses (Fig. 12R). This femur is composed of fibrolamellar bone with longitudinal to reticular canals (Fig. 12S). The outermost cortex is heavily vascularized. There are no LAGs or EFS present. The outer and inner cortices are obscured by DA.

UF300298 is a partial left distal femur broken proximally just below the lesser trochanter with a lateral extension of the gluteal tuberosity (Fig. 12T). Distally the epiphyses are fused but the lateral condyle has been broken off (Fig. 12T). The femur is composed of fibrolamellar bone with sub-reticular to plexiform canals (Fig. 12U-V)). A single LAG is visible along the outer cortex but is obstructed by DA in certain

regions (Fig. 12V-U; Table S6). Secondary osteons are restricted to the posterior quadrant along the linea aspera. DA obscures any structure along the inner and outer cortices.

Figure 12. All femora from Inglis 1A sectioned for this study. A-D) UF45040; E-I) UF45044; J-L) UF300294; M-N) UF300295; O-Q)

UF300296; R-S) UF300297; T-V) UF300298. Anterior photo of whole bone prior to destruction (A, E, J, M, O, R, T); Thin section photograph with outlined LAGs (red) (B, F, K, N, Q, S, U); Anterior view of thin section showing bone structure and LAGs (arrows) (C, G, L, Q, V); Postero- medial view of primary bone with radial canals (D, H); Posterior remodeling close to linea aspera. All thin section photographs were taken under ppl. Red boxes indicate zoomed in areas with corresponding photographs. Scale bars are all 1 mm except for (A, E, J, M, O, R, T) which are 1 cm.

Leisey Shell Pit 1A

UF86378 is a partial left distal femur broken at the mid diaphysis with fused distal epiphysis and broken lateral condyle (Fig. 13A). The femur is composed of fibrolamellar bone with longitudinal vascular canals intermixed within a reticular to plexiform vascular complex (Fig. 13B-C). LAGs are present in the mid to outer cortex; the outermost LAGs may be a partial EFS (Fig. 13C; Table S6). Remodeling is occurring with one generation of secondary osteons present close to the linea aspera and in the innermost cortex of the medial quadrant. A very thin semi-transparent rim of DA makes up the outer cortex (Fig. 13B). The rest of the bone is fairly well preserved with some small patches of bacterial degradation and other DA.

UF239925 is partial right proximal femur, broken at the mid diaphysis, with a fused proximal epiphysis

(Fig. 13D). This femur is composed of fibrolamellar bone with reticular to laminar canals (Fig. 13E-F).

LAGs are present starting in the mid-cortex with no EFS (Fig. 13E-F; Table S6). Few if any secondary osteons are visible. There is some mineral/bacterial alteration sporadically throughout the bone, especially along cracks and along the outer and inner cortex.

Coleman 2A

UF300286 is a partial right distal femur, broken distally along the shaft with a fused distal epiphysis (Fig.

13G). The histological section was taken along the distal shaft (Fig. 13H). This femur has heavy DA and remodeling, which makes it difficult to observe any original tissue. There is one BGM present along the lateral outer cortex, but is obstructed by DA anteriorly and posteriorly; no EFS is present (Table S6). One

– two generations of remodeling is prevalent where original tissue is visible in the section.

UF300287 is partial right proximal femur, broken at mid diaphysis with an unfused proximal epiphysis

(Fig. 13I). The femur is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 13J). A potential BMG may be deposited on the external most medial cortex and can be partially followed anteriorly (Fig. 13J). There are no LAGs or EFS visible. Remodeling is occurring throughout the entire section. Most remodeling is taking place laterally towards the linea aspera and anteriorly evident by erosion cavities (Fig. 13J). First and second generation osteons are present throughout the section.

Preservation is moderate with heavy DA along the internal and external cortices.

UF300288 is partial right femur mid-shaft, broken proximally just below the lesser trochanter with a lateral extension gluteal tuberosity and distally broken along the suprapatellar bursa anteriorly and the supracondylar foramen posteriorly (Fig. 13K). Both epiphyses were missing, thus OS and age was indeterminate based on partial remains. The femur is composed of fibrolamellar bone with plexiform to laminar canals where visible (Fig. 13K). The internal and external cortices are heavily DA. There are 2

BGMs present postero-laterally, with no EFS visible (Fig. 13K). Secondary remodeling is visible posteriorly along the linea aspera.

Figure 13. All femora from Leisey Shell Pit 1A (A-F) and Coleman 2A (G-L) sectioned for this study. A-C) UF86378; D-F) UF239925; G-H)

UF300286; I-J) UF300287; K-L) UF300288. Anterior photos of whole bones prior to destruction, (A, D, G, I, K); Thin section photographs with outlined LAGs (red), (B, E, H, J, L); Anterior view of thin section showing bone structure and LAGs (arrows) (C, F). Red boxes indicate zoomed areas to corresponding photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, D, G, I, K) which are 1 cm.

Nichol’s Hammock

UF57620 is complete left femur with fused epiphyses (Fig. 14A). The femur is composed of fibrolamellar bone intermixed with parallel fibered bone and lamellar tissue along the external cortex (Fig. 14B-C).

Longitudinal vascular canals occur within the inner cortex and become smaller and more tightly packed plexiform/laminar canals within the mid-outer cortex. LAGs are present from the mid to outer cortex; the final LAG represents the beginning of an EFS (composed of ~3-4 BGMs) in a zone of lamellar bone (Fig.

14C; Table S6). Secondary osteons are most prevalent in the posterior part of the section along the linea aspera representing one-two generations of remodeling while few secondary osteons are present throughout the rest of the section.

UF57621 is a complete left femur with fused epiphyses (Fig. 14D). The femur is composed of fibrolamellar bone with small plexiform and longitudinal vascular canals (Fig. 14E-F). LAGs are visible along with the potential beginning of an EFS along the posterior margin (Fig. 14F; Table S6).

Remodeling is most prevalent as first generation secondary osteons visible posterior-laterally close to the linea aspera along the inner cortex.

UF57622 is a complete left femur with completely fused epiphyses (Fig. 14G). The femur is composed of fibrolamellar bone with small reticular to plexiform and vascular canals (Fig. 14H-I). LAGs are visible and no EFS is present (Fig. 14H-I; Table S6). Few secondary osteons are visible.

Figure 14. All femora from Nichol’s Hammock sectioned for this study. A-C) UF57620; D-F) UF57621; G-I) UF57622. Anterior photos of whole bones prior to destruction, (A, D, G); Thin section photographs with outlined LAGs (red), (B, E, H); Anterior view of thin section showing bone structure and LAGs (arrows) (C, F, I). Red boxes indicate zoomed areas to corresponding photographs. C was taken under xpl; all other thin section photographs were taken under ppl. Scale bars are all 1 mm except for (A, D, G) which are 1 cm.

Tibiae

Inglis 1A

UF45069 is a right distal tibia that is broken along the mid-diaphysis with a fused distal epiphysis (Fig.

15A). The tibia is composed of fibrolamellar bone with plexiform to laminar canals (Fig. 15B-C). LAGs are visible but no EFS is present (Fig. 15B-C; Table S6). All anterior LAG measurements were obtained from an antero-lateral region due to better preservation (Fig. 15C). Some first to second generation secondary osteons are visible in the antero-medial region along the tibial crest and in the posterior inner cortex.

UF45075 is a right distal tibia that is broken mid-diaphysis, with a fused distal epiphysis (Fig. 15D). The primary tissue in this tibia is obscured by heavy DA (Fig. 15E). A small portion of the outer cortex, visible around the majority of the circumference, contains fibrolamellar bone with reticular canals to avascular fibrolamellar tissue. Neither LAGs nor an EFS were visible. There may be a BGM present along the outer cortex wrapping around the lateral, posterior and medial quadrants of the bone. Secondary remodeling is present in the antero-medial portion of the bone with one-two generations of remodeling. In other regions, only one generation of secondary osteons is visible in the outer cortex. The entire section has heavy DA, except for parts of the outer cortex.

UF45087 is a partial right proximal tibia that is broken mid-diaphysis along the tibial crest with a fused proximal epiphysis (Fig. 15F). The tibia is composed of fibrolamellar bone with reticular to laminar canals (Fig. 15G). There is at least one BGM visible along the medial and lateral quadrants of the section.

Secondary osteons are present along the inner cortex posteriorly and around the tibial crest. DA obscures inner and outer cortices throughout the section.

UF276272 is a right tibia that is broken proximally above the tibial tuberosity with an extension to the lateral condyle with a fused distal epiphysis (Fig. 15H). The primary tissue in the tibia is heavily DA along the inner and outer cortices (Fig. 15I). A small section of fibrolamellar bone with plexiform canals is visible laterally. Neither LAGs nor an EFS are visible. Secondary remodeling is present in the posterior internal cortex.

UF302003 is a complete left tibia broken proximally along the tibial crest with unfused epiphyses (Fig.

15J). The tibia is composed of fibrolamellar bone with laminar canals where visible (Fig. 15K). Heavy

DA has obscured much of the tissue and neither LAGs nor an EFS are visible. No secondary remodeling is visible.

Figure 15. All tibia from Inglis 1A sectioned for this study. A-C) UF45069; D-E) UF45075; F-G)

UF47087; H-I) UF276272; J-K) UF302003; A,D, F, H and J). Anterior photos of whole bones prior to destruction; B, E, G, I and K) Thin section photographs with outlined LAGs (red); C) Anterior view of thin section showing bone structure and LAGs (arrows). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, D, F, H and J) which are 1 cm.

Leisey Shell Pit 1A

UF85251 is a partial left distal tibia broken along the distal shaft with fused distal epiphysis (Fig. 16A).

The tibia is composed of fibrolamellar bone with some laminar canals and much of the primary tissue structure is actively being resorbed and remodeled (Fig. 16B). No LAGs or an EFS are visible while one- two generations of secondary osteons are present.

UF226999 is a partial left tibia shaft and fused distal epiphysis that is broken above the tibial tuberosity with a portion extending proximally up toward the lateral condyle (Fig. 16C).The tibia is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 16D-E). LAGs are present but an EFS is not

(Fig. 16D-E; Table S6). There is remodeling present anteriorly along the tibial crest with two-three generations of secondary osteons, but these are sparse along the inner cortex of the section. Much of the section is well preserved, although the postero-lateral section was shattered.

UF239928 is a partial left distal tibia broken along the mid-distal shaft with a fused distal epiphysis (Fig.

16F). The tibia is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 16G). There is at least one BGM along the antero-medial external cortex but is lost in both directions and no EFS is visible (Fig. 16G). Remodeling is prevalent yet sporadic within the entire section with one-two generations of secondary osteons present along the inner to mid cortex. Much of the section has some DA comprised of an isotropic mineral deposited in splotches along the inner and outer cortices and along cracks (Fig. 16G).

Figure 16. All tibia from Leisey Shell Pit 1A sectioned for this study. A-B) UF85251; C-E) UF226999; F-G) UF239928. A,C and F) Anterior photos of whole bones prior to destruction; B, D, and G) Thin section photographs with outlined LAGs (red); E) Anterior view of thin section showing bone structure and LAGs (arrows). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl.

Scale bars are all 1 mm except for (A, C and F) which are 1 cm.

Coleman 2A

UF276488 is a partial left distal tibia which is broken along the proximal mid-shaft at the tibial crest, with a partially fused distal epiphysis (Fig. 17A). The tibia is composed of fibrolamellar bone with plexiform to laminar canals (Fig. 17B). No LAGs or an EFS are visible throughout the section. Resorption cavities are present, and are concentrated posteriorly and anteriorly towards the tibial crest (Fig. 17B). One – two generations of secondary remodeling are present anteriorly and posteriorly. The external cortex is obscured by DA.

UF276489 is a partial left tibia that is broken along the mid-shaft with a fused distal epiphysis (Fig. 17C).

The tibia is composed of fibrolamellar bone with plexiform to laminar canals (Fig. 17D-E). LAGs and a potential EFS are visible (Fig. 17E; Table S6). One-two generations of remodeling are visible throughout antero-lateral region along the tibial crest. Much of the internal and some of the external cortical bone are obscured by DA.

UF276490 is a partial left distal tibia which is broken along the mid-shaft with a fused distal epiphysis

(Fig. 17F). The tibia is composed of fibrolamellar bone with plexiform to sub-laminar canals (Fig. 17G-

H). LAGs are present starting in the mid-cortex best viewed from the antero-lateral and postero-medial regions of the bone (Fig. 17G-H; Table S1). No EFS or secondary osteons are visible. Much of the primary tissue is heavily DA; antero-lateral and postero medial regions offer the most visible primary structure.

Nichol’s Hammock

UF57185 is a left proximal tibia broken mid-shaft with a fused proximal epiphysis (Fig. 17I). The tibia is composed of fibrolamellar bone with mostly longitudinal, reticular, and plexiform canals, and some laminar canals along the posterior and lateral quadrants of the section (Fig. 17J-K). LAGs are present and composed of lamellar tissue, with the 5th LAG marking the start of an EFS (Fig. 17K; Table S6).

Anteriorly to the fifth LAG, there is a BGM visible, but it cannot be traced circumferentially (Fig. 17K).

Remodeling is prevalent anteriorly along the tibial crest with one-two generations of secondary osteons.

Overall preservation is great with little to no DA.

Figure 17. All tibiae from Coleman 2A (A-H) and Nichol’s Hammock (I-K) sectioned for this study. A-

B) UF276488; C-E) UF276489; F-H) UF276490; I-K) UF57185. A-B) UF85251; C-E) UF226999; A,C,

F and I) Anterior photos of whole bones prior to destruction; B, D, G and J) Thin section photographs with outlined LAGs (red); E, H and K) Anterior view of thin section showing bone structure and LAGs

(arrows). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, C, F and I) which are 1 cm.

Humeri

Inglis 1A

UF43597 is a partial left humerus with a fused distal and unfused proximal epiphysis (Fig. 18A). The primary tissue in the humerus is heavily DA except for portions of the outer cortex that are highly vascular and no EFS is present (Fig. 18B).

UF43600 is a partial left humerus broken midshaft with a fused distal epiphysis (Fig. 18C). The primary tissue in the humerus was heavily DA so no original microstructure was visible (Fig. 18D).

UF300299 is a partial left humerus shaft that is broken proximally at the deltoid tuberosity and distally through the coronoid and radial fossae, with unfused epiphyses (Fig. 18E). The humerus is composed of fibrolamellar bone with some plexiform canals (inner cortex) and laminar canals (mid-outer cortex).

LAGs are visible through most of the section, although the second LAG is comprised of multiple lines within a zone of parallel fibered tissue (annuli) rather than singular defined line (Fig. 5D-E; Fig. 18G-H;

Table S6) and the first LAG is partially resorbed posteriorly (Fig. 18F). The beginning of an EFS with two-three BGMs is visible along the lateral to postero-medial sections of the external most cortex.

Remodeling is visible throughout the lateral, antero-lateral and anterior quadrants with at least one-two generations of secondary osteons. Preservation for this section is exceptional except for a thin quasi transparent darkened area extending antero-laterally to antero-medially in the inner most cortex and a similar thin darkened area around the entire outer most cortex.

UF300300 is a partial left distal humerus that is broken proximally along the shaft above the pectoral ridge, with a fused distal epiphysis (Fig. 18I). The humerus is composed of fibrolamellar bone with reticular to laminar canals (Fig. 18J). Many LAGs are most apparent in the posterior most section, although only the first four and the final LAG can be traced circumferentially through the anterior region for an accurate APD measurement (Fig. 18J-K; Table S6). No EFS is visible. Some remodeling is present along the inner and outer cortex especially medially. DA is visible along the mid to outer cortex, providing only small windows of original microstructure in some regions.

UF302000 is a distal left humerus with unfused epiphyses (Fig 18L). The primary tissue in the humerus is heavily DA except for a postero-lateral outer cortical region that contains fibrolamellar bone with plexiform to laminar canals (Fig. 18M). There are no LAGs or EFS visible. Remodeling is visible through

DA antero-medially with two-three generations of secondary osteons present.

UF302001, 302002 are two halves creating a complete left humerus with a distal break above the radial and coronoid fossa, as well as a half broken distal shaft; epiphyses are fused (Fig. 18N). Primary tissue within the humerus is heavily DA except for a posterior mid to outer cortical region comprised of fibrolamellar bone with laminar canals (Fig. 18N). No LAGs, EFS or secondary osteons are visible (Fig.

18O).

Figure 18. All humeri from Inglis 1A sectioned for this study. A-B) UF43597; C-D) UF43600; E-H) UF300299; I-K) UF300300; L-M)

UF302000; N-O) UF302001 and UF302002. A,C, E, I, L and N) Anterior photos of whole bones prior to destruction; B, D, F, J, M and O) Thin section photographs with outlined LAGs (red); G and K) Posterior view of thin section showing bone structure and LAGs (arrows). H) 1st LAG is an annuli contains two lines with parallel fibered bone between (see Fig. 5D-E). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, C, E, I, L, N) which are 1 cm.

Leisey Shell Pit 1A

UF65962 is a partial left distal humerus broken along the proximal shaft at the top of the pectoral ridge, with a fused distal epiphysis (Fig. 19A; Table S3). The humerus is composed of fibrolamellar bone with plexiform to laminar canals (Fig. 19B-C). LAGs and annuli are present and composed of a mix of fibrolamellar and lamellar bone (Fig. 19D). The first LAG is partially resorbed posteriorly (Fig. 19B-C).

Remodeling of primary bone is prevalent along the anterior and medial sections of the bone with one-two generations of secondary osteons. Resorption cavities are visible within the anterior inner cortex (Fig.

19B). Overall preservation is very good, although mineral deposits along cracks, within the inner and outer cortex of the bone obstructs some structure.

UF81121 is a partial left humerus shaft distally broken above the radial and coronoid fossae and proximally along mid-shaft, with missing epiphyses (Fig. 19E). The humerus is composed of fibrolamellar bone with reticular to laminar canals (Fig. 19F-G). LAGs are present but no EFS is visible

(Fig. 19F-G, Table S6). Remodeling of primary bone occurs along the inner cortex throughout the section but is concentrated towards the anterior and antero-medially/laterally, although the number of generations is unknown due to DA. Some mineral deposits along cracks obscure structures, within the inner and outer cortex.

UF87904 is a partial left distal humerus broken proximally along the mid shaft with a fused distal epiphysis (Fig 19H). The humerus is composed of fibrolamellar bone with plexiform to laminar canals with visible LAGs but no EFS (Fig. 19I-J, Table S6). Remodeling is prevalent antero-medially and antero-laterally with one-two generations of secondary osteons. Deposition of an isotropic mineral and bacterial alteration has occurred along the bone edges, which obstructs some of the external cortical structure.

Coleman 2A

UF300283 is a partial distal left humerus that is broken along the mid-shaft with a fused distal epiphysis

(Fig. 19K). The primary tissue of the humerus is highly DA along the inner and outer cortices, while the mid cortex reveals some visible structure (Fig. 19L). The mid cortex is composed of fibrolamellar bone

with vascular canal partially obscured by DA. Posteriorly, vascular canals are laminar with some longitudinal and reticular canals. Neither LAGs nor an EFS are visible. Some resorption is occurring posteriorly while remodeling (with one-two generations of secondary osteons) is visible along the postero-lateral and postero-medial inner to mid cortex.

UF300284 is a partial distal right humerus that is broken along the proximal mid-shaft with a fused distal epiphysis (Fig 19M). Primary tissue within the humerus is heavily DA and highly remodeled (Fig. 19N).

An unobstructed view of bone structure medially reveals two-three generations of secondary remodeling.

There is an EFS present laterally in the section, but no LAGs are visible.

UF300285 is partial distal left humerus that is broken distally along the radial and coronoid fossae and proximally along the distal mid shaft with broken epiphyses (Fig. 19O). The humerus is composed of woven to fibrolamellar bone with longitudinal to laminar canals (Fig. 19O). No LAGs or an EFS are visible. Resorption is occurring throughout the antero-lateral region (Fig. 19P). Some secondary osteons are present throughout the internal cortex of the antero-lateral/medial regions with one-two generations of remodeling. Some DA of the internal and external cortex is present.

Figure 19. All humeri from Leisey Shell Pit 1A (A-J) and Coleman 2A (K-P) sectioned for this study. A-D) UF65962; E-G) UF81121; H-J)

UF87904; K-L) 300283; M-N) UF300284; O-P) UF300285. A, E, H, K, M and O: Anterior photos of whole bones prior to destruction; B, F, I, L,

N and P: Thin section photographs with outlined LAGs (red); C, F and I: Posterior view of thin section showing bone structure and LAGs

(horizontal arrows). D: Annuli composed of lamellar bone (red vertical arrows). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl except D. Scale bars are all 1 mm except for (A, E, H, K, M and O) which are 1cm.

Nichol’s Hammock

UF57164 is a complete left humerus with fused epiphyses (Fig. 20A). The humerus is composed of fibrolamellar intermixed with parallel fibered bone with longitudinal to laminar canals (Fig. 20B-C).

LAGs/annuli are present; the final two LAGs contain multiple lines with the final LAG possibly representing the formation of an EFS when viewed in the posterior region (Fig. 20C; Table S6). The first two LAGs are partially resorbed posteriorly (Fig. 20B). Two to three generations of remodeling are visible in the medial region and along the inner most cortex of the entire section. Overall preservation is excellent with little to no DA.

UF57165 is a complete left humerus that was broken post-burial along the shaft and is missing the greater tubercle and contains fused epiphyses (Fig. 20D). The humerus is composed of some fibrolamellar bone intermixed with parallel fibered bone with mostly longitudinal to small plexiform canals (Fig. 20E-F).

LAGs/annuli are present and the final LAG may be the beginning of an EFS (Fig. 20E-F, Table S6). Two- three generations of remodeling is prevalent medially while only one-two generations occur throughout the rest of the section. Resorption cavities are prevalent in the posterior and postero-lateral regions.

Overall preservation is excellent with little to no DA.

UF57166 is a complete left humerus with fused epiphyses (Fig. 20G). The humerus is composed of fibrolamellar bone within the inner to mid cortex and lamellar bone along the outer-mid to outer cortex

(Fig. 20H-I). Longitudinal intermixed with sub-plexiform canals occur within the inner to mid cortex and become sparse and more laminar within the mid to outer cortex (Fig. 20H-I). LAGs and annuli are present; the final LAG marks the potential beginning of an EFS (Fig. 20I, Table S6). One-two generations of remodeling are concentrated to the anterior and lateral inner most cortex. Overall preservation is excellent with little to no DA.

UF300289 is a complete left humerus with unfused epiphyses (Fig. 20J). The humerus is composed entirely of woven bone with large longitudinal canals (Fig. 20K). There are no LAGs, EFS, or secondary remodeling present.

UF300290 is complete left humerus with unfused epiphyses (Fig. 20L). The humerus is composed of woven to fibrolamellar bone with longitudinal canals within the inner cortex becoming reticular to plexiform towards the mid to outer cortex (Fig. 20M). No LAGs or EFS are present.

Figure 20. All humeri from Nichol’s Hammock sectioned for this study. A-C) UF57164; D-F) UF57165; G-I) UF57166; J-K) 300289; L-M)

UF300290. A,D, G, J and L) Anterior photos of whole bones prior to destruction; B, E, H, K and M) Thin section photographs with outlined LAGs

(red); C, F and I) Posterior view of thin section showing bone structure and LAGs (arrows). Red boxes indicate corresponding zoomed in photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, D, G, J, L) which are 1 cm.

Radii

Inglis 1A

UF45007 is a complete left radius with fused proximal and unfused distal epiphyses (Fig. 21A). The radius is composed of fibrolamellar bone with reticular to plexiform vascular canals (Fig. 21B). A single

BGM is present medially although DA and remodeling restricts tracing it laterally and posteriorly and no

EFS is visible. The outer medial cortex is heavily vascularized. Along the anterior and posterior inner cortexes primary fibrolamellar bone with some radial/plexiform canals are present. Remodeling represented by first and second generation secondary osteons occurs along the internal cortex both medially and laterally, and extends to the mid and outer cortex anteriorly. Posteriorly, DA obscures any microstructure.

UF45008 is a complete left radius with fused proximal and unfused distal epiphyses (Fig. 21C). The radius is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 21D). No LAGs or EFS are present. The outer cortex is vascularized with visible canal formation. Enlarged erosion cavities and first generation secondary osteons are present in the inner cortex anteriorly and posteriorly. DA obscures structure along the periosteal surface.

UF276267 is a partial left distal radius is broken proximally along the mid-shaft with a fused distal epiphysis (Fig. 21E). The radius is heavily DA and no internal bone structure is preserved (Fig. 21F).

UF300291 is a left radius with unfused epiphyses (Fig. 21G). The radius is composed of fibrolamellar bone with reticular vascular canals (Fig. 21H). No LAGs or EFS are present. Some resorption and remodeling (first generation) is visible posteriorly, although no other signs of remodeling are present elsewhere.

UF300292 is a left radius with fused proximal and unfused distal epiphyses (Fig. 21I). The radius is composed of fibrolamellar bone with reticular to plexiform vascular canals (Fig. 21J). No LAGs or EFS are present. Some resorption cavities are visible posteriorly, but no secondary osteons are visible. DA is concentrated towards the inner cortex and throughout the posterior region.

UF300293 is a left radius with a fused proximal and unfused distal epiphyses (Fig. 21K). The radius is composed of fibrolamellar bone with reticular to plexiform canals (Fig. 21L). Some radial vascular canals are present anteriorly along the inner cortex and separated from the outer cortical bone by a region of secondary remodeling. There is a single BGM present visible anteriorly and laterally, but no EFS is present. Remodeling is occurring with one-two generation of secondary osteons present in the anterior mid-cortex and throughout the posterior cortex. DA is present along the entire periosteal surface.

Figure 21. All radii from Inglis 1A sectioned for this study. A-B) UF45007; C-D) UF45008; E-F) UF276267; G-H) 300291; I-J) UF300292; K-L)

UF300293. A, B, D, F, G, I and K) Anterior photos of whole bones prior to destruction; B, D, F, H, J and L) Thin section photographs. All thin sections were taken under ppl. Scale bars are all 1 mm except for (A, D, G, J, L) which are 1 cm.

Nichol’s Hammock

UF57178 is a complete left radius with fused epiphyses (Fig. 22A). The radius is composed of regions of fibrolamellar bone with longitudinal to reticular canals interspersed between zones of lamellar bone, with the outer cortex entirely composed of parallel fibered bone (Fig. 22B). There are multiple BGMs; many expand laterally and medially but disappear posteriorly, due to remodeling and/or cortical drift, and merge anteriorly. Remodeling is prevalent along the inner to mid cortex. The anterior and posterior sections of the bone are most heavily remodeled with one-two generations of secondary osteons present. Overall preservation is excellent with little to no DA.

UF57179 is a complete left radius with fused epiphyses (Fig. 22C). The radius is composed of fibrolamellar bone with longitudinal to reticular vascular canals between zones of lamellar bone (Fig.

22D). There are multiple BGMs in the lamellar regions that expand medially but disappear posteriorly due to remodeling or merge together anteriorly and laterally (Fig. 22D). Many BGMs are composed of multiple lines but none are an EFS (Fig. 22D). Remodeling is prevalent along the inner to mid cortex throughout the bone. The anterior and posterior sections of the bone are most heavily remodeled with one-two generations of secondary osteons present. Some resorption cavities are present medially and posteriorly along the outer cortex as well. Overall preservation is excellent with little to no DA.

UF57180, 57182 are two halves of a complete left radius that are broken along the midshaft; each half having a fused epiphysis (Fig. 22E). The removed section was taken from the mid-diaphysis of UF57180.

The radius is composed of fibrolamellar bone with mostly longitudinal to reticular canals, and radial canals are present antero-laterally (Fig. 22F). Anteriorly along the outer cortex lamellar bone makes up multi-lined BGMs that expand medially but disappear posteriorly due to remodeling. An EFS may be developing antero-medial external cortex. The anterior and posterior sections of the bone are most remodeled with one-two generations of secondary osteons present. Overall preservation is excellent with little to no DA.

UF57181 is a partial left radius broken midshaft with a fused distal epiphysis (Fig. 22G). The radius is composed of fibrolamellar bone with longitudinal to reticular canals (Fig. 22H). BGMs contain multiple

lines that expand medially and laterally but disappear posteriorly due to remodeling. The outer BGMs may be the beginning of an EFS. Remodeling is present along the inner to mid cortex throughout the bone. The anterior and posterior sections of the bone are mostly remodeled, with one-two generations of secondary osteons. Some resorption cavities are present within the anterior and posterior cortex as well.

Overall preservation is excellent with little to no DA.

Figure 22. All radii from Nichol’s Hammock sectioned for this study. A-B) UF57178; C-D) UF571794;

E-F) UF57180 and UF57182; G-H) UF57181. A, B, D, F, G, I and K) Anterior photos of whole bones prior to destruction; B, D, F, H, J and L) Thin section photographs. All thin sections were taken under ppl.

Scale bars are all 1 mm except for (A, D, G, J, L) which are 1 cm.

Skeletochronology

Of the 54 thin sections created, nine femora, five tibiae and eight humeri contained LAGs that could be analyzed for skeletochronology. Possible LAGs/BGMs within radii were difficult to differentiate due to extensive cortical drift and extensive remodeling in the posterior region of the bone and were excluded from all skeletochronological analyses.

Comparison of successive anterior-posterior diameters (APDs) of LAGs among similar individual elements provided a baseline growth curve and revealed that resorption removed the first LAG in older individuals (as is common in artiodactyls; Marin-Moratalla et al., 2013; Kolb et al., 2015a; Jordana et al.,

In Press; Figs. 23, 26, 29; Tables 19, S7). Maximum and Minimum APD derived Growth Rate

(M/mAPDGR) were calculated based on the differences between successive individual LAG APDs (see methods; Tables 19-20, S7-9), and graphed per individual element and the average per locality (Fig. 24,

27, 30; Tables 19-20, S7-9). M/mAPDGR graphs resulted in the same trends but contained different values (see femora, Figs. 24B-C, Tables S8-9). Due to previous research that show that artiodactyls grow for approximately 260-280 days a year (Köhler et al., 2012), only MAPDGR (based on 260 day growth period; see methods) values were graphed for tibiae and humeri and are presented here (Figs. 27, 30,

Tables 20, S8).

Table 19. Average APD measurements for individual LAGs from femora, tibiae, and humeri.

Locality Element LAG 0.5* LAG 1 LAG 2 LAG 3 LAG 4 LAG 5 LAG 6 LAG 7 LAG 8 ING1A Avg. FE 17.44 16.79 19.49 21.63 22.03 20.55 20.96 21.26* - LSP1A Avg. FE - 17.87 19.35 20.01 19.47 19.73 - - - COL2A Avg. FE** 17.6 ------

NH Avg. FE - 13.34 15.44 15.99 16.61 17.78 18.28 - - ING1A Avg. TB** - - 18.64 19.28 19.95 - - - - LSP1A Avg. TB** - 15.33 16.82 17.65 - - - - - COL2A Avg. TB 16.06** 13.90 15.11 15.58 - 17.41 - - - NH Avg. TB** - - 14.60 15.34 15.79 16.45 16.81 - - ING1A Avg. HU - - 19.99 21.06 21.71 19.88** - - 20.56** LSP1A Avg. HU - 17.61 19.66 20.82 21.34 - - - - NH Avg. HU 9.77 - 15.30 16.81 17.33 18.04 18.47 18.11 - * indicates where measurement was taken from a juvenile < 1 year old (no LAG present). ** indicates where average was taken from a

single individual.

Table 20. Average Maximum APD growth rates (MAPDGR) (260 day growth period) based on APD measurements for each growth period. GZ1 cannot be accurately calculated due to unknown degree of resorption.

Locality Element GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 ING1A Avg. FE 7.34* 2.48 1.52 1.46 1.58* 1.15* - LSP1A Avg. FE 5.71 2.54 1.19 1* - - - NH Avg. FE 7.48 2.98 2.38 1.85* 1.92 - - ING1A Avg. TB* - 2.48 2.57 - - - - LSP1A Avg. TB* 5.72 3.21 - - - - - COL2A Avg. TB 4.63 1.80 2.61* 2.46* 2.22* - - NH Avg. TB* - 2.84 1.73 2.54 1.38 - - ING1A Avg. HU - 4.11 2.52 1.81* - - - LSP1A Avg. HU 7.87 2.30 1.98 - - - - NH Avg. HU - 2.98 2.01 2.73 1.46 0.96 0.54 * indicates where average was taken from a single individual.

Maximum and minimum growth rates for both a Standard (365 day) year (ST), and Growth

Period (GP) (260 days; M/mGRST/GP) based on methodology from Sander and Tückmantel (2003) represent a singular growth rate for the entire ontogenetic history of an animal (Table 21, S10-12), and can be used for comparison to other artiodactyl studies (Kolb et al., 2015a). Although, due to resorption of bone tissue and determinate growth of mammals (Lee et al., 2013), this would still likely grossly under estimate whole growth rates for an individual element. Instead, Maximum (260 day) and minimum (365 day) Anterior Growth Rates (M/mANTGR) were used from individual anterior growth zones and revealed similar trends to M/mAPDGRs with some minor differences (Figs. 24-25, 27-28, 30-31). Only individual and average MANTGR for each element were graphed for tibiae and humeri, rather than mAPDGR and mANTGR, due to similarities in the trends, (compare Figs. 25B with 25C) and based on published ~260 day growth period for artiodactyls (Köhler et al., 2012). Humeri MANTGRn was recorded from the posterior rather than anterior region due to better preservation and similarities to anterior femoral regions.

Table 21. Average anterior growth rates from methodologies of Sander and Tückmantel (2003) based on a 260 day growth period. See text for formulas for (MGRGP, mGRGP, MANTGRn). * = minimum growth rate values due to inner cortical resorption. ** = average growth rate is based on a single specimen. All rates are in (µm/day). Humeri MANTGRn was recorded from the posterior rather than anterior region due to better preservation and similarities to anterior femoral regions.

MANTGR n Specimen Mean Rate of Elmt. MGRGP mGRGP GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Number Growth ING1A Avg. FE 8.49 1.44 3.08 9.43* 8.00 1.42 1.11 0.84 0.65 1.11 0.56 LSP1A Avg. FE 4.12 1.09 2.61 9.98* 2.08 1.41 0.4 - - - - NH Avg. FE 3.81 1.60 2.71 6.62* 3.95 1.06 1.15 1.15 0.81 - - ING1A Avg.** TB 4.44 1.36 2.9 - 8.14* 2.08 1.98 - - - - LSP1A Avg. ** TB 6.39 2.86 4.63 9.01* 5.51 3.1 - - - - - COL2A Avg. TB 3.25 1.16 2.43 12.62* 2.93 1.86 2.15 1.03 1.25 2.75 - NH Avg.** TB 2.42 1.33 1.87 - 5.53* 2.58 1.18 1.92 0.59 1.72 - ING1A Avg. HU 2.80 2.54 2.67 3.52 5.35 2.02 1.06 0.80 0.79 0.28

LSP1A Avg. HU 4.17 2.85 3.04 6.64 6.30 2.08 1.59 - - - - NH Avg. HU 1.58 1.07 1.15 5.06 2.23 2.10 1.52 1.52 0.92 0.32 0.15

Femora

Retrocalculated femoral APDs reveal much larger diameters in both LSP1A and ING1A populations compared to NH (Fig. 23A; Tables 19, S7). There is no overlap in APD values of individuals from LSP1A or ING1A with NH (Fig. 23A; Table S7). Moreover, APDs from juveniles (based on epiphyseal fusion; see Table. 1) within their first year of growth (no LAGs present) from ING1A, LSP1A and COL2A show an APD size equal to at least that of a two year old NH individuals (Fig. 23A; Table

S7).

Average MAPDGR were similar at GZ2 between NH and ING1A populations and slightly higher than the LSP1A population (Fig. 24B-C; Tables 20, S8). After GZ2, average MAPDGR from ING1A and

LSP1A populations drop below NH and continue to decrease into GZ3 and beyond (Fig. 24B-C; Tables

20, S8). This is in contrast with NH population, which show a more gradual decrease after GZ3 and remains at a greater APDGR as compared to both ING1A and LSP1A throughout GZ6. Later growth zones and growth rates are based on single individuals (Figs. 24B-C; Tables 20, S8). The same relative pattern, but with higher values was observed when using mAPDGR 365 days (Fig. 24C; Tables S8-9).

Femoral average MGRGP and MGRST revealed a geochronologic trend with the greatest value in the ING1A followed by LSP1A and NH which differed only slightly (Tables 21, S11-12). mGRGP and mGRST both revealed the greatest growth rate within NH, but ING1A and LSP1A values were very similar in mGRGP, while ING1A had a greater value than LSP1A for mGRST (Tables 21, S11-12).

Overall average MANTGRs reveal greater early ontogenetic growth within ING1A and LSP1A populations as compared to NH populations (Fig. 25B-C; Tables 21, S11). Growth rates become mostly similar at GZ3 and beyond (Fig. 25B-C; Table S11). No specimens from COL2A yielded recordable

LAGs for use in these analyses.

A A Individual Femora APD B Average Femora APD 24 24

22 22

20 20

18 18

16 16

APD (cm) APD APD (cm) APD

14 UF45040 UF45044 14 UF300294 UF300295 UF300296 UF300297 Inglis 1A UF300298 UF86378 12 Leisey Shell Pit 12 UF239925 UF300287 Coleman 2A Single Indiv. UF57620 UF57621 Nichol's Hammock UF57622 10 10 0.5* 1 2 3 4 5 6 7 0.5* 1 2 3 4 5 6 7 LAG Number LAG Number

Figure 23. Femora APD measurements from all histological specimens showing LAGs. A) Individual femora APD from all specimens. B).

Average femora APDs from all localities. Unfilled shapes are averages based upon a single individual. All graphs are color code based on

chronology. Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman 2A, Red = Nichol’s Hammock. * indicates maximum APD from

juveniles with no LAGs representing minimum incomplete growth during their first year.

A 10 Individual Femora MAPDGR B 8 Average Femora MAPDGR C 6 Average Femora mAPDGR

9 7 5 8 6 7 UF 45040 UF 45044 4 5 6 UF 300294 UF 86378 5 UF 239925 4 3 4 UF 57620 Inglis 1A Inglis 1A UF 57621 3 Leisey Shell Pit Leisey Shell Pit 2 3 UF 57622 Nichol's Hammock Nichol's Hammock 2 Growth Rate (µm/day) Rate Growth 2 1

1 1

Average Growth Rate (µm/day ) (µm/day Rate Growth Average Average Growth Rate (µm/day ) (µm/day Rate Growth Average 0 0 0 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 Growth Zone Growth Zones Growth Zones A Figure 24. Maximum and minimum femora APD growth rates (M/mAPDGR) per growth zone (GZ; ex. GZ2 = distance between LAG1 to

LAG2). A) Individual MAPDGR per specimen. B) Average MAPDGR per population. C) Average mAPDGR; note no differences in trend but

rather in values between B and C. Unfilled shapes are averages based upon a single individual; color code based on chronology: Black = Inglis 1A,

Blue = Leisey Shell Pit 1A, Green = Coleman 2A, Red = Nichol’s Hammock.

A 10 8 A 14 Individual Femora MANTGR B Average Femora MANTGR C Average Femora mANTGR

12 8

6 10 UF45040 UF45044 6 8 UF300294 UF300295 4 UF300296 6 UF300297 4 UF300298 UF86378 Inglis Inglis 4 UF239925 Lesiey Shell Pit Lesiey Shell Pit Nichol's Hammock 2 Nichol's Hammock

UF57620 (µm/day) rate Growth

2 (µm/day) rate Growth Growth Rate (µm/day) Rate Growth 2 UF57621 UF57622

0 0 0 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Growth Zone Growth Zone Growth Zone Figure 25. Maximum/minimum femoral anterior growth rates (M/mANTGR) per growth zone (GZ; ex. GZ2 = distance between LAG1 to LAG2).

A) Individual MANTGR per specimen. B) Average MANTGR per population; open shapes represent when average growth rate was calculated

from a single individual. C) Average mANTGR per population; note different values but similar trends between B and C. Unfilled shapes are

averages based upon a single individual; color code based on chronology: Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman 2A,

Red = Nichol’s Hammock. * = minimum values. (Some individuals also have minimum values from GZ2; see Tables 21, S11-12).

Tibiae

Few tibiae provided sufficient preservation for observation of LAGs, thus most populations are represented by a single individual (ING1A, LSP1A and NH). Tibiae APD show distinct values from all localities, with a decreasing chronologic trend in overall APD values between the populations (Fig. 26;

Tables 19, S7). Average tibiae MAPDGRs show a similar pattern to femora with all the growth rates being more or less equal at GZ3 and beyond (Fig. 27; Tables 20, S8). The LSP1A individual has a slightly greater EOAPDGR (early ontogenetic APD growth rate) than the average COL2A and other population at GZ2 and 3 (Fig. 27; Tables 20, S8). MGRST and MGRGP were greatest in the LSP1A individual, decreasing within ING1A, COL2A and NH (Tables 21, S11-12). mGRST and mGRGP also put the LSP1A individual with the greatest growth rate but a different decreasing order: COL2A, ING1A,

NH (Tables 21, S11-12).

Overall tibiae MANTGR revealed similar trends as seen in femora with all Pleistocene populations having greater early ontogenetic growth rates when compared to NH (Fig. 28; Tables 21,

S11). A chronologic trend can be seen at GZ2 with ING1A having the greatest rate followed by LSP1A,

COL2A and NH, although mean rates for COL2A are almost identical to NH at GZ2. Growth rates become similar in all individuals at GZ3 and beyond (Fig. 28; Tables 21, S11).

A 21 Individual Tibiae APD B 21 Average Tibiae APD

20 20

19 19

18 18

17 17

16 16 APD (cm) APD 15 (cm) APD 15

Inglis 1A 14 UF45069 14 UF226999 Leisey Shell 13 UF276488 13 Pit UF276489 Coleman 2A UF276490 12 UF57185 12 Nichol's Hammock 11 11 0.5* 1 2 3 4 5 6 0.5* 1 2 3 4 5 6 LAG Number LAG Number Figure 26. Tibiae APD measurements per LAG from all specimens showing LAGs. A) Individual APD from all specimens. B). Average APD from all localities. Unfilled shapes at GZs represent when growth rate was calculated from a single individual. Color code based on chronology.

Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman 2A, Red = Nichol’s Hammock. * = APD from juvenile individuals with no

LAGs.

A 6 Individual Tibiae MAPDGR B 7 Average Tibiae MAPDGR

UF45069 5 4 UF226999 UF276489 4 Inglis 1A UF276490 3 Leisey Shell Pit UF57185 Coleman 2A 3 Nichol's Hammock

Growth Rate (µm/day) Rate Growth 2

1 Average Grwoth Rate (µm/day) Rate Grwoth Average 1

0 0 GZ2 GZ3 GZ4 GZ5 GZ6 GZ2 GZ3 GZ4 GZ5 GZ6 Growth Zone Growth Zone Figure 27. Maximum tibiae APD growth rates (MAPDGR) per growth zone (GZ; ex. GZ2 = distance between LAG1 to LAG2) based on a 260 day growth period. A) Individual growth rates per specimen. B) Average growth rate per population. Unfilled shapes are averages based upon a single individual, color code: Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman 2A, Red = Nichol’s Hammock.

100

14 A 20 Individual Tibiae MANTGR B Average Tibiae MANTGR

18 12

10 14

12 8

10 Inglis 1A Single Indiv. UF45069 6 Leisey Shell Pit 1A Single UF226999#2 8 Indiv. UF276489 Coleman 2A

UF276490 Growth Rate (µm/day) Rate Growth 6 (µm/day) Rate Growth 4 Nichol's Hammock Single UF57185 Indiv. 4 UF276488 2 2

0 0 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 Growth Zone Growth Zone Figure 28. Maximum tibiae anterior growth rates (MANTGR) per growth zone (GZ; ex. GZ2 = distance between LAG1 to LAG2) based on a 260 day growth period. A) Individual growth rates per specimen. B) Average growth rate per population. Unfilled shapes at GZs represent when growth rate was calculated from a single individual. Color coded based on chronology. Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green =

Coleman 2A, Red = Nichol’s Hammock. * = minimum values. (Some individuals also have minimum values from GZ2; see Tables 21, S11).

101

Humeri

Humeri APD values are on average greater in ING1A and LSP1A populations when compared to

NH, with the greatest values within the ING1A population (Fig. 29; Tables 19, S7). Average humeri

APDs between ING1A and LSP1A do not differ greatly from one another. Average MAPDGR among humeri show only marginal to no differences from GZ3 and beyond (Fig. 30; Tables 20, S8). Early ontogenetic growth rates reveal slightly greater values in ING1A at GZ2 followed by NH and LSP1A

(Fig. 30; Tables 20, S8).Humeri M/mGRST and M/mGRGP was greatest in the LSP1A population following with ING1A and NH populations (Table 21, S10-12).

Overall humeri average MANTGR revealed slightly different trends as seen in other elements; within GZ1 LSP1A has the greatest growth rate followed by NH and then ING1A populations, although at GZ1 ING1A is represented by a single individual (Fig. 31; Tables 21, S11). At GZ2, LSP1A has the greatest growth rate followed by ING1A and NH populations (Fig. 31; Tables 21, S11). At GZ3 and beyond all growth rates are generally consistent among the populations (Fig. 31; Tables 21, S11).

102

A 26 Individual Humeri APD B 26 Average Humeri APD

24 24

22 22

20 20

18 18

16 16

APD (cm) APD APD (cm) APD

14 14

UF300299 UF300300 12 12 Inglis 1A UF65962 UF81121 UF87904 UF57164 Leisey 10 10 Shell Pit UF57165 UF57166 Nichol's UF300289 UF300290 Hammock 8 8 0.5* 1 2 3 4 5 6 7 8

LAG Number LAG Number Figure 29. Humeri APD graphs from all localities. A) Individual APD from all specimens. B). Average APD from all localities. Unfilled shapes are averages based upon a single individual. Color coded based on chronology. Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman

2A, Red = Nichol’s Hammock.

103

A 10 Individual Humeri APDGR B 9 Average Humeri MAPDGR

9 8

7 6

6 UF300299 5 UF300300 5 UF65962 UF81121 4 4 UF87904 Inglis 1A UF57164 3 Leisey Shell 3 UF57165

Growth Rate (µm/day) Rate Growth Pit Nichol's UF57166 (µm/day) Grwoth Average 2 2 Hammock

1 1

0 0 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Growth Zone Growth Zone Figure 30. Humeri growth rates vs individual growth zone using APD. A) Individual growth rates per specimen B) Maximum average growth rate based on a 260 day growth period. Unfilled shapes are averages based upon a single individual. Color coded based on chronology. Black = Inglis

1A, Blue = Leisey Shell Pit 1A, Green = Coleman 2A, Red = Nichol’s Hammock.

104

A 9 Individual Humeri MANTGR B 7 Average Humeri MANTGR

8 6

5 6

5 UF300299 4 UF300300 UF65962 Inglis 1A 4 UF81121 3 UF87904 Leisey Shell Pit 3 UF57164

UF57165 Nichol's Growth Rate (µm/day) Rate Growth Growth Rate (µm/day) Rate Growth UF57166 2 Hammock 2 UF300289 UF300290 1 1

0 0 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Growth Zone Growth Zones Figure 31. Maximum humeri posterior growth rates (MANTGR) per growth zone (GZ; ex. GZ2 = distance between LAG1 to LAG2) based on a

260 day growth period. A) Individual growth rates per specimen. B) Average growth rate per population. Unfilled shapes represent when growth rate was calculated from a single individual. Color coded based on chronology: Black = Inglis 1A, Blue = Leisey Shell Pit 1A, Green = Coleman

2A, Red = Nichol’s Hammock. * = minimum values, (Some individuals also has minimum values from GZ2; see Tables 21, S11).

105

Osteocyte Density

Osteocyte counts and calculated densities per growth zone may offer an independent test for trends in growth derived from LAG data (Figs. 32-33; Tables 22-23, S13-15). Osteocytes were manually counted per anterior growth zone using both a single “representative” layer as done in previous studies on artiodactyls (Jordana et al., In Press), as well as using a stacked image methodology used in other histological studies (Bromage et al., 2009; Stein and Werner, 2013; Cullen et al., 2014). Humeri were not included in this analysis because remodeling in the anterior regions of the sections had destroyed any primary bone and osteocytes present. Posterior regions would not be comparable to other elements and osteocyte densities have been shown to differ in different regions of an individual thin section (Cullen et al., 2014).

Femora osteocyte single layer densities show generally no trend across all GZs (Fig. 32A-B;

Table 22, S13). Stacked layer osteocyte counts show a decreasing trend in all LSP1A and NH population

(Fig. 32C-D; Tables 23, S14-15). The ING1A population shows a decreasing trend until GZ5 and an increasing trend beyond. Osteocyte densities per GZ are similar among LSP1A and ING1A populations but much less than the NH population (Fig. 32; Tables 22-23, S13-15).

Tibiae osteocyte single layer and stacked layer densities show a decreasing trend in most GZs of individual elements (Fig. 33A-B; Tables 22-23, S13-15). The stacked NH osteocyte densities show an increase that differs from the decrease observed in the single layer osteocyte densities (Fig. 30A-B;

Tables 22-23, S13-15).

106

Table 22. Average single layer Osteocyte Lacunae (OL) density for each growth zone (GZ). El.= Element, * = Average based on a single individual

OL Density per GZ (lacunae/mm2) Locality El. GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 ING1A Avg. FE 1666.7 1388.9 1400 1600 1200 1666.7 1333.3 LSP1A Avg. FE 1583.3 1137.5 1333.3 - - - - NH Avg. FE 1733.3 1455.6 1383.3 1533.3 1766.7 1300 - ING1A Avg. TB* - 1600 1266.7 - - - - LSP1A Avg. TB* 1450 1366.7 - - - - - COL2A Avg. TB 1033.3 1300 1450 966.7 933.3 - - NH Avg. TB* - 1666.7 1600 1433.3 - - -

Table 23. Average stacked image osteocyte lacunae (OL) density for each growth zone El. = Element, * = Average based on a single individual

OL Density per GZ (lacunae/mm3)

Locality El. GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 ING1A Avg. FE 29850.1 36607.7 33585.9 27392.0 19934.6 21256.0 27381.0 LSP1A Avg. FE 40218.4 33611.1 22916.7 - - - - NH Avg. FE 61250.0 52268.5 46450.6 48625.1 40555.6 32098.8 - ING1A Avg. TB* - 31111.1 21296.3 - - - - LSP1A Avg. TB* - 44047.62 36507.9 - - - - COL2A Avg. TB 34343.4 42592.6 41666.7 36507.9 34127.0 - - NH Avg. TB* - 42857.1 46825.4 50000 - - -

107

A 2000 Femora Lacunae Density (Single Layer Image) B 1800 Average Femora Lacunae Density (Single Layer

1800 1600 Image)

1600 1400 1400 1200 1200 1000 1000 800 800 UF45040 UF45044 600 UF300294 UF300296 600

UF300297 UF300298 Inglis 1A

(osteocytes/mm²)

(osteocytes/mm²) Density Osteocyte Osteocyte Density Density Osteocyte UF86378 UF239925 400 400 Leisey Shell Pit 1A UF57620 UF57621 200 200 UF57622 Nichol's Hammock 0 0 GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 Growth Zone Growth Zone

C 80000 Femora Lacunae Density (Stacked Layer Image) D 70000 Average Femora Lacunae Density (Stacked Layer 70000 UF45040 Image)

UF45044 60000

60000 UF300294 UF300296 50000 50000 UF300297 UF300298 40000 40000 UF86378 UF239925 30000 30000 UF57620

UF57621 20000 (osteocytes/mm³)

Osteocyte Density Density Osteocyte 20000 (osteocytes/mm³) UF57622 Density Osteocyte Inglis 1A Leisey Shell Pit 1A 10000 10000 Nichol's Hammock 0 0 GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 Growth Zone Growth Zone

Figure 32. Osteocyte density (OD) per growth zone for all femora. A). Individual element OD based on single layer image. B) Average OD for single layer image. C) Individual element OD based stacked layers image. D) Average OD for stacked layer image.

108

A 2000 Tibiae Osteocyte Density (Single Layer Image) B 60000 Tibiae Osteocyte Density (Stacked Image)

1800 50000 1600

1400 40000 1200

1000 30000

800 UF45069 20000 600 UF57185 UF45069 UF276488 UF57185 400 UF276488 UF276489

10000 UF276489

Osteocyte Density (osteocytes/mm²) Density Osteocyte (osteocytes/mm³) Density Osteocyte 200 UF226999 UF226999

0 0 GZ1 GZ2 GZ3 GZ4 GZ5 GZ1 GZ2 GZ3 GZ4 GZ5 Growth Zone Growth Zone Figure 33. Osteocyte density (OD) per growth zone for all tibiae. A). Individual element OD based on single layer image. B) Individual OD for stacked layer image.

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Discussion

Studying populational growth dynamics within a single vertebrate fossil species requires analyzing a large sample of individuals from the same geologic time and space (locality). Unfortunately, poor preservation of vertebrate material in the fossil record had generally limited our ability to conduct such studies (Padian et al., 2013; Woodward et al., 2015). Woodward et al. (2015) were able to obtain a large enough sample size to observe populational histological variation amongst a single species

(Dinosauria: Maiasaura peeblesorum; Horner and Makela, 1979), but few if any studies have addressed populational variation in growth (using bone microstructure) in mammals, especially among Cervidae.

Most forays into mammalian bone microstructure have analyzed large-scale evolutionary differences between and among species in a clade, with each species being represented by only a few individuals

(Marín- Moratalla et al., 2013; Köhler et al., 2012, Kolb et al., 2015a, Jordana et al., In Press). Thus, such studies do not always account for possible intra-specific growth differences at the populational level

(Padian et al., 2013; Woodward et al., 2015). This study is unique because it addresses this problem directly by comparing growth differences in different populations of a single cervid species (Odocoileus virginianus) from a restricted geographic location over the course of ~2 million years. Histological thin sections reveal some differences in bone vascular canal networks and growth rates during early ontogeny.

In addition, chronologic, geographic, and environmental differences in growth rates across all populations were also present. Observed differences in growth history between populations could have numerous interconnected underlying extrinsic (quantity and quality of food, climate, intra-/interspecific competition, predation) and/or intrinsic (sexual dimorphism, phenology) causes, each of which will be discussed.

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Comparative Histology

Bone structure in modern and fossil vertebrates has been extensively researched and provide basis for interpretation of bones in the fossil record (Enlow and Brown, 1956-1958, Enlow, 1963, 1969; de

Ricqlés, 1975-1978, Castanet et al., 1996, 2000). The fastest growing bone matrix is woven-fibered and fibrolamellar bone which is commonly present in mammals, dinosaurs and birds, as well as comprised the majority of the bone microstructure observed in deer from this study (Huttenlocker et al., 2013; Figs. 5,

34). This bone contains numerous vascular canals in which larger and more connected canals equate to greater growth rates (Huttenlocker et al., 2013). Lamellar bone and parallel-fibered bone are both indicative of slower growth rates and typically have fewer vascular canals (Huttenlocker et al., 2013; Fig.

5). These types of bone microstructure are commonly found in non-archosaurian diapsids but can be found among some mammals as well as artiodactyls especially within the EFS, or individual annuli

(Köhler and Moya-Sola, 2009; Huttenlocker et al., 2013; this study).

Primary Bone

Bone histology generally reveals a conservative primary tissue structure between and among populations in this study. General fibrolamellar bone tissue from all localities (Figs. 5, 12-22, 34) correlate to high growth rates among all sampled deer, which agrees with previous studies of other cervids and artiodactyls (Köhler et al., 2012; Kolb. et al., 2015a; Jordana et al., In Press), supporting an evolutionarily conservative growth strategy within this group.

Juveniles from all localities had primary bone with the largest networks of vascular canals, supporting rapid growth (Figs. 24-25, 27-28, 30-31, 34). Although the largest and most interconnected vascular canals were present in early ontogenetic (EO) tissue for all localities, some differences in canal structure were observed among different populations. EO tissue (present as primary tissue in the outer cortex of juveniles or in the inner cortex of adult bones) for ING1A is highly connected reticular to sub plexiform (Fig 34B); LSP1A EO is reticular with some connectivity with some development of longitudinal canals (Fig. 34C); NH has EO bone with smaller sub-laminar to longitudinal canals with

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little direct connectivity (Fig. 34D-E). This suggests that juveniles form ING1A had the highest EO growth rates (EOGR), followed by LSP1A and then NH.

The outer cortex of adult elements corresponds to later growth, representing sub-adult to adult stages of ontogeny. A higher number of longitudinal vascular canals with fewer interconnections between them, forming a sub-reticular to laminar canal network, characterizes primary tissue in adult Odocoileus virginianus (Fig. 34B-E). Little to no observable variation exists between vascular networks among sub- adult to adult tissue in sampled populations (Fig. 34B-E). These vascular canal orientations indicate that species-wide reduction in growth slowed down during late ontogeny and that growth rates should be similar among all populations (Figs. 24-25, 27-28, 30-31). This ontogenetic reduction in growth rate is characteristic of all other sample cervids and mammals in general (Marín-Moratalla et al., 2013; Köhler et al., 2012; Kolb et al., 2015a, b; Jordana et al., In Press).

Femora, tibiae, and radii exhibited bone structure described above, yet there were some notable differences in humeri (Fig. 34F-J). Much of the vascularity within humeri consisted of a plexiform to laminar canal network (even among juveniles; Fig. 34F-H, J), although large longitudinal vascular canals in a prenatal humerus were present and surrounded by woven tissue (Fig. 34I). However, relative differences between canal network structures still suggest the same growth pattern as in other long bones, with juveniles growing fast than adults (Figs. 24-25, 27-28, 30-31).

Secondary Bone

Secondary remodeling was extensive in older specimens, but present in all but the very young.

Most remodeling starts in the inner to mid cortex and around muscle attachment points (ex. posterior femora along the linea aspera) and intensifies with age (Fig. 6), which agrees with previous results from other mammals (Kolb et al., 2015a). Some juvenile and presumed sub-adult specimens showed high degrees of resorption and secondary remodeling, evident from large resorption cavities as well as extensive presence of secondary osteons (Figs.13J; 16B; 17B), which may indicate particular difficulties obtaining ample nutrients. Lack of ample nutrients available in the environment (especially calcium for bucks, which is required for seasonal antler growth) may result in remodeling and resorption of bone for

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nutrient resources needed to sustain growth. However, remodeling of bones to release calcium for antler growth typically occurs in non-load bearing elements (ribs) but sometimes is also present in some load- bearing elements (metacarpals, metatarsals; Meister, 1956; Banks et al., 1968; Hillman et al., 1973; Baksi and Newbrey, 1989; Baxter et al., 1999), which were not sampled here.

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Figure 34. Comparison of bone vascularity among ontogenetic series and localities. A) UF300295: ING1A juvenile femur; B) UF45044: ING1A

Sub-Adult femur; C) UF239925, LSP1A Sub-Adult femur; D-E) UF57620, UF57621: Adult Femur (xpl). Top row all anterior views, anterior up medial right. F) UF300299: ING1A Sub-Adult Humerus; G) UF87904: Sub-Adult humerus LSP1A; H) UF300290 juvenile humerus I) UF300289 juvenile humerus; J) UF57166: Adult Humerus. Bottom row all posterior view but H (anterior). Scale bars are all 1mm; black arrows indicate

LAGs.

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Growth Lines and Skeletochronology

LAGs and/or annuli were present almost uniformly in adults among all sampled localities.

Spacing between individual LAGs among all elements, show little to no variation between each population among ontogenetically similar individuals (Tables S6, 7). LAGs were mostly composed of singular lines, although some in later ontogeny are visible as double or triple lines in some regions, indicating closely related timed periods of slowed or ceased growth in an individual. Some annuli in humeri were composed of a zone of lamellar bone with individual laminae, others contained parallel fibered matrix (Fig. 5). While LAGs indicate a total cessation of bone apposition, annuli indicate only a significant reduction in growth, although both may occur in the same individual (Huttenlocker et al.,

2013).

The attainment of skeletal maturity is represented by the formation of an EFS in outermost cortex and represents the complete cessation of lengthwise growth and complete fusion of epiphyses (Cormack

1987; Lee et al., 2013). Skeletal maturity was reached at a younger age in NH individuals (~4 – 5 years of age; Fig. 34E ) compared to ING1A individuals, with both individuals from ING1A (~4 and 8 years old) did not show any definitive formation of an EFS (UF45040 and UF45044; Fig. 12C and G; Table S1).

One LSP1A individual (UF86378) may have the beginning of an EFS present (2 years old) although the older individual from LSP1A (UF239925; 3 years old) does not show any EFS formation (Fig. 13C and

F).

Changes in the timing of skeletal maturity may be a product of sexual dimorphism, with females reaching skeletal maturity earlier than males (as in other deer; Jordana et al., In Press), or just intra- population variation. Previous studies on European ruminants have shown that the transition of fibrolamellar to lamellar bone in the outer cortex, signifying the start of an EFS may indicate the onset of sexual maturity (Marín-Moratalla et al., 2013; Jordana et al., In Press). However , this may not be true of

O. virginianus, as in UF45044 from ING1A was ~8 years old yet did not contain a definitive EFS and little change in bone microstructure (Fig. 34B; Table S1). It is unlikely that O. virginianus was not

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reaching sexual maturity until after 8 years of growth, since modern populations from Florida reach sexual maturity between 6 months (females) to 1.5 years (males; Garrison and Gedir, 2006). However, skeletal maturity among bucks in Florida O. virginianus occurs at 4-6 years and earlier among females

(~2.5 years; Garrison and Gedir, 2006; Gee et al., 2014), and agrees with timing observed in the most recent fossil population from NH (Fig. 31E). Jordana et al. (In Press) argues for potential predation resulting in early maturation. Earlier onset of skeletal maturity within the NH population likely does not indicate more predation since deer have relatively fewer potential predators in Florida (panther, bobcats, wolves) than were present during the Pleistocene (Morgan and Hulbert, 1995). Even so, intra-species variation and environmental conditions may account for these differences (Jordana et al., In Press).Thus, the presence of an EFS in Florida O. virginianus likely represents skeletal maturity rather than sexual maturity, although more specimens and data are needed to see if the trend of skeletal maturity in fossil and modern populations of O. virginianus represents environment pressures, predation pressures, or is merely a product of small sample size and intra-populational variation.

Growth Rates

The general chronologic trend of decreasing APD from the early Pleistocene to Holocene (Figs.

23, 26, 29; Table 19) is correlated with decreasing EOGR (as measured by ANTGR; Figs. 25, 28, 31;

Tables 21, S11). Thus, higher EOGR indicate larger juvenile size among ING1A specimens, which would result in larger adults as well. This same trend of larger size in chronologically older populations is supported by a statistically significant decrease in element MDC between fossil (ING1A and

Rancholabrean fossils) and modern populations (Tables 5-7; Fig. 10). More data from fossil localities especially Irvingtonian sites would help clarify this chronologic trend.

Even so, gender of the populations observed could have affected significance observed. The modern population used in this study consisted of mostly females with some individuals of unknown genders and very few males (Table 18). Thus, the chronological decreasing trend of limb size may have been affected by sexual dimorphism within populations and skewed modern measurement data. Skeletal size can also be affected by geography such as with Bergman’s Rule, the extent of this is discussed later.

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Differences between EO APDGR and ANTGR may be due to differences in exact positioning of thin section slices along the shaft (compare femora thin section shapes Figs. 7, 12-14, and tibiae thin section shapes Figs. 15-16). Changes in the position and curvature of the linea aspera, transverse shape of the femur from mid-to distal shaft and individual variability may reflect differences within APD measurements. Thin sections from two ING1A specimens (UF45040 and UF45044) were obtained at a more distal shaft position compare to mid-shaft from NH (UF57620, UF57621 and UF57622) (compare anterior posterior distances and shape of Fig. 12B, F with Fig. 14B, E). These differences in femoral and tibiae APD will not affect retrocalculation or APDGR trends because all measurements were taken along the same axis per element, but may have an effect on overall APD measurements and individual growth rates calculated from those measurements. Thus, APDGR may not be as useful as ANTGR unless bones can be sectioned at precisely the exact same spot along the mid-shaft and show little to no individual variability. Despite this possible source of error, some ING1A juvenile femora have with similar APD thin section shapes as those in NH, yet the ING1A individuals grew much larger in their first two years of growth compared to NH individuals (compare Figs. 12K, N, P, S, U; 14B, E, H; 23, 25, 31, 34). Thus, similarities in general growth trends from APDGR and ANTGR along with similarly shaped elements from ING1A and NH support the conclusion that ING1A population had greater EOGR.

Overall skeletochronologic results reveal a decreasing trend of EOGR through time and with latitude (Figs. 25, 28, 31). Larger EOGR could result in overall larger skeletal size, as in ING1A specimens used in this study (Fig. 10-11; Tables 5-8, 10-12, 14-16). Studies during the Holocene show that deer may have changed in overall body size influenced by resource availability due to climate and environmental changes (Purdue, 1989; Purdue and Reitz, 1993). Inter-populational variation may also relate to geographical differences observed in modern populations of O. virginianus (Tables 14, 16).

Bergman’s Rule hypothesizes that variability in body size across latitude is linked to a physiological response to availability of food resources due to differences in climate and environmental conditions

(Wolverton et al., 2009). It is this possibility that a physiological response to resource availability

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(quantity) and quality (nutritional value of resources obtained) may have caused changes observed in O. virginianus, with this response beginning in the early-mid Pleistocene for this species.

Osteocyte Density

Average osteocyte density (OD) per growth zone in both femora and tibiae show a slight decrease among all populations when using a single layer image (Fig. 32A-B and 33A; Tables 22, S13). This decrease is comparable to previous Cervidae studies that have shown a decrease in femoral ODs throughout GZs (Jordana et al., In Press), although the actual calculated densities (lacunae/mm2) in this study are three-times greater. Differences between the individual ODs between this study and other studies may be influenced by methodological/instrumentation differences and/or fossil/modern preservation. Subjective differences (e.g., human bias) in the identification and method of counting osteocytes likely played a significant role in differences between this and other studies, where different observers (with varying experience levels) may qualify variables in contrasting ways. A similar problem exists among dental microwear researchers (Green and Resar, 2012; Mihlbachler et al., 2012). The effect of subjectivity in two-dimensional (single-plane) OD studies should be further analyzed using controlled, blind experiments in the future.

A different methodology that used stacked images rather than a single plane was also applied.

Average OD among majority elements for z-stacked images show a decline between GZs, although an increase was observed from a single NH tibiae across GZs (Figs. 32C-D and 33B; Tables 23, S14-15).

This decline may directly relate to growth rate because previous studies have shown higher OD during periods of higher growth, such as early ontogeny (Bromage et al., 2009; Werning, 2012; D’Emic and

Benson, 2013; Stein and Werner, 2013; Cullen et al., 2014). This study supports previous studies that have used z-stacked images which provide a better representative of osteocytes present for morphometric and quantitative analyses than a single image (D’Emic and Benson, 2013; Fig. 35). Decreasing stacked

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OD among most bone samples independently supports the observed decrease in growth rate through ontogeny.

Femoral stacked OD from NH population were much greater and do not overlap with ING1A and

LSP1A populations across all elements sampled. Because NH experienced reduced early ontogenetic growth relative to other localities (Figs. 25, 32C-D), it stands to reason that growth rate must not be the only causation of increased OD. Other studies have also found inverse relationship between OD and body mass, (Cané et al., 1982, Bromage et al., 2009; D’Emic and Benson, 2013; Cubo et al., 2012; Stein and

Werner, 2013; Cullen et al., 2014). Although body mass was not directly known or calculated for individuals from any sampled population, adult limb elements from NH and modern individuals had significantly smaller MDC values when compared to fossil populations such as ING1A which supports a smaller skeletal size (Figs. 10-11;Tables. 5-7, 9-12).

Metabolism can vary between individuals and changes in metabolism can be observed within histology (Köhler and Moya Sola, 2009). Histologic studies in the Plio-Pleistocene bovid Myotragus show that changes in metabolism due to limited island resources can result in changes in bone microstructure and growth patterns (Köhler and Moya-Sola, 2009). Therefore, differences observed among ODs and growth rates within the NH population may have resulted from these individuals living in a more resource limited environment (explored further below).

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Figure 35. Comparison of the best representative single layer image (left) and stacked layer image (right) from anterior GZ1 from UF45040. To reduce subjectivity of counting osteocytes, this study shows that stacked images may be a better representative of osteocyte density rather than single images.

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Extrinsic factors influencing growth

Climate and environment

Glacial and interglacial plant and faunal communities within Florida fluctuated in response to climatic shifts through the Pleistocene (Grimm et al., 1993, 2006; De Santis et al., 2009; Yann and

DeSantis, 2014). These differences influenced the quantity and quality of food resources available for consumption by herbivorous animals, which in turn may have influenced a population’s growth history.

Environmental and climatic interpretations for fossil locations used in this study are presented in

Table 2. Isotopically depleted δ13C enamel from herbivores along with spikes in Pinus pollen from lake cores indicate that more glacial-like environments (ING1A, COL2A) were constructed of C3 plants within a mostly closed forest ecosystem (Grimm et al., 1993, 2006; DeSantis, et al., 2009, Yann and DeSantis,

2014). C3 plants may offer more protein rich forage for animals (Minson, 1990; Vendramini et al., 2010), which may result in the faster growth rates and larger limb/skeletal size seen within the ING1A population (Figs. 10, 23, 25-26, 28-31; Tables 5-7, 8-9, 11-12, S1). The hypothesis of faster growth in glacial environments would be further supported by elevated growth in other glacial environments relative to interglacial localities throughout Florida.

COL2A, with its transitional interglacial to glacial environment (Table 1), yielded adult metatarsal sizes that differed more when compared with ING1A, than when compared to chronologically separate central Florida populations (Fig. 10; Table 7). Further growth rates from COL2A tibiae also show more similarities to southern interglacial populations (LSP1A and NH; Figs. 27-28) than to ING1A especially during EO. Both ING1A and COL2A are very close in terms of geography (i.e., latitude; Fig.

2) but differ chronologically and in environmental reconstruction; (forest in ING1A vs. transitional savanna to forest in COL2A). The COL2A population lived about 1.5 million years after ING1A during intense glaciations on the NA continent (Fig. 1). Rising sea levels during interglacials would have flooded much of the Florida peninsula, creating fewer habitats for deer, reducing quantity of forage available and resulting in possible nutritional deficiency among the COL2A population. This nutritional deficience

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could manifest itself as reduced growth rates in the COL2A deer population. The COL2A population was transitioning out of an interglacial and thus it is possible that population genetics may have resulted in selection of for lower growth rates and smaller size to account for reduced environmental circumstances.

This is further supported by similarities of tibial growth and APD between COL2A and strict interglacial populations (LSP1A and NH; Figs. 23-24). Therefore, the influence of climate and environmental changes through time may explain some of the observed differences between glacial populations. More work is needed to help discern more specific environmental conditions at COL2A (isotopes, microwear and faunal assemblages). Also, available femora and humeri samples from COL2A were few (except for tibiae, which are underrepresented at other localities) and poorly preserved, to reveal a comparable sample with other Pleistocene populations. Thus, sample size cannot be ruled out for any similarities/differences observed between COL2A and other populations.

LSP1A is an interglacial locality with an interpreted warm arid environment dominated by C4 grazers (Table 1) (Pratt and Hulbert, 1995; Rich and Newson, 1995; DeSantis et al., 2009; Yann and

DeSantis 2014). Even so, the presence of Odocoileus and Tapirus (predominately C3 browsers) with

13 slightly enriched δ C, suggests that C3 plants were present but limited (DeSantis et al., 2009). This implies an environment consisting of open grassland with patchy forests (DeSantis et al., 2009). This type of habitat may provide the quantity of forage required by deer because they can be more generalized in their diet (Geist, 1998), but the inclusion of higher amounts of lower protein-rich C4 plants may have negatively impacted growth among deer living in interglacial environments (Minson, 1990; Vendramini et al., 2010).

However, average APDGRs and ANTGRs of limb elements from LSP1A do not completely support the hypothesis of reduced quality forage resulting in lower growth rates during interglacials (Figs.

25, 28, 31). Femora and humeri both reveal equal or greater MANTGRs in GZ1 in the LSP1A population when compared to ING1A (Figs. 25 and 31). Growth rates within GZ2 are consistently lower in LSP1A than ING1A (Figs. 24-25, 27-28). So perhaps the quality of forage obtained as a juvenile to sub-adult

(year 2) has a greater impact on growth rates. Even so little significant differences were observed in

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LSP1A adult MDC and AL, when compared to modern south Florida adult populations (Fig. 11; Tables

8-12). Although, environmental interpretations reveal significant changes in climate and environment and may be associated with changes in vegetation and faunal makeup, analyses of growth within the first year

(GZ1) among mid Pleistocene interglacial populations does not appear to be greatly impacted by dietary changes due to environmental changes. Data from DeSantis et al (2009) show that although deer

13 incorporate more C4 plants into their diets, their overall δ C value did not shift drastically and was still well in a range of a browsing organism. Thus, even if deer incorporated more C4 plants into their diet, it seemingly did not greatly affect their growth in the LSP1A population. Therefore, either food quality may not be the only driver in growth among mid Pleistocene populations, and/or environmental conditions may have not been drastically different enough between glacial and interglacials during the mid Pleistocene. Even so, interpreting physiological responses within a fossil population to changing climatic and environmental conditions in the past, likely involves the interplay of many complex and complicated systems.

The Holocene population from NH represents the current interglacial and has much lower APDs and EOGRs than all other populations (Figs. 23, 25-26, 28-31; Tables 19-21). These observed differences in the NH population could have multiple underlying causes discussed below, and the geographic, environmental and chronological differences between NH and other sampled populations may be affecting resulting data.

NH is a sinkhole deposit within a hammock surrounded by rocky pinelands (Table 1; Hirschfeld,

1969). A modern southern Florida faunal assemblage was found in the sinkhole suggesting a similar environment as modern southern hammocks with rocky pinelands that have high diversity of plants but poor soil fertility (Florida Natural Area Inventory (FNAI), 2010). Modern southern Florida soil fertility depends on availability of water, influx of nutrients from fires, and aeolian sediments transported by tropical storms (Myers, 1985, 1990; Garrison and Gedir, 2006; Lindemann, 2009; FNAI, 2010; Glaser et al, 2013). Changes in the position of the Bermuda High and southward shift in the jet stream caused by decreases in North Atlantic Oscillation (NAO) during the late Holocene (post ~2800 cal BP (calibrated

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radiocarbon age) may have shifted moisture and hurricanes away from Florida and into the Gulf of

Mexico, resulting in less moisture and nutrient rich aoelian sediments for southern Florida environments

(Glaser et al, 2013). Along with decrease in NAO, an increase in El Niño events after ~2700 cal BP may have decreased Atlantic hurricane frequency accentuating these environmental impacts through southern

Florida (Moreno et al., 2006; Glaser et al., 2013).

In contrast, modern El Niño events can produce more dry-season rainfall that may suppress the possibility of ignition and spreading of wildfires during the dry to wet transition (Beckage et al., 2003).

Along with this, prevalence of water in an environment such as southern Florida could result in lowland flooding leading to a reduction in environments and congregation of animals on higher ground that could result in overconsumption and reducing quality of forage (Garrison and Gedir, 2006). Therefore, changes in growth and limb sizes observed within NH may be a result of poorer quality of forage due to soil fertility and possible reduction of landscapes/overpopulation caused by climatic changes during late

Holocene. More work is needed to directly tie forage quality to growth differences especially among northern vs. southern Florida O. virginianus populations.

Similarities between modern and Pleistocene (Pinus) pollen records were observed from Lake

Tulane which may indicate that modern climate is similar to that of the Pinus phase which occurred during glacials, (warm and wet; Grimm et al., 1993, 2006). Although, pollen records may show similar vegetation types, patterns in δ18O variation from LSP1A also are similar to modern patterns of δ18O variation, which may infer similar aridity and seasonal changes in precipitation between modern and early

Pleistocene interglacial environments (DeSantis et al., 2009). Modern analogues for past climates can be further complicated, due in part to human influence; models show that summer convection and rainfall have both decreased in Florida since 1900 due to draining of the Everglades (Marshall et al., 2004). Thus similarities seen in modern and Pleistocene glacial δ18O records may be related but not directly correlative of modern environments and climate.

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Rather than short term climate and environmental differences (glacial/interglacial) affecting populations it appears that regional differences in collaboration with other climatic influences such as

NAO variability, rainfall/storm frequency, and orbital parameters through the past 2 MY may also explain some of the similarities and differences observe through these proxies. Interestingly southern Florida

Rancholabrean populations seem to show a dramatic increase in adult MDC among many all available elements (≥2) in relation to modern and Irvingtonian populations (Fig. 11; Table 12). More data can help differentiate whether this trend is due to lower sample sizes especially among Irvingtonian populations or an increase in the intensity of glacial and interglacial cycles post 900,000 years (Fig. 1).

Therefore, rather than specific environmental changes occurring through cycles of glaciations and interglaciation affecting quantity and quality of forage, long term climate and environmental changes through the entirety of the Pleistocene and early Holocene may have produced observed changes in limb bone histology, measurements and growth rates. Even so, there is an inherent complexity of understanding and correlating past with modern climate and environments, through which more data and work is needed.

Competition and predator-prey relationships

Predator-prey relationships and inter/intra-species competition could also influence growth within a population (Warren and Krysl, 1983, Jenks and Leslie, 2011; Sinclair et al., 2003; Cooper et al., 2008).

Currently only Felis concolor coryi (the Florida panther), a species listed as endangered by the U.S Fish and Wildlife Service, Felis rufus (bobcat) are the major predators among deer within southern Florida

(Schaefer and Main, 1997; Garrison and Gedir, 2006), excluding the impact of humans (known impact,

Garrison and Gedir, 2006) and naturalized Canis latrans (coyotes) who’s impact on Florida deer populations has not been extensively studied (Schaefer and Main, 2014). Many studies have shown a positive relationship between individual prey growth rate/body size and predation (Stearns, 1992; Gaillard et al., 2000; Sinclair et al., 2003; Festa-Bianchet et al., 2006; Ricklefs, 2007; Cooper et al., 2008). Modern

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populations revealed geographic decrease in limb size towards southern Florida (Tables 14-17), which does not fit these previous conclusions from other studies.

There were an abundance of predators present in Florida during the Pleistocene that may have preyed upon deer (Morgan and Hulbert, 1995). Larger size and faster growth of early Pleistocene deer may have been influenced by these predator dynamics. Even so, conflicting influences of prey growth due to predator-prey dynamics are present and likely represents a complexity within the predator-prey dynamics that are difficult to understand among fossil populations (Carpenter et al., 2005). Thus, predator-prey relationship may have affected growth but was likely not the only factor. More work is needed to test the possible impact of extinct predators on fossil deer and other prey populations.

Inter/Intra-species Competition

Competition with other herbivores could also explain changes in growth dynamics among

Pleistocene deer. Diversity of the Florida ecosystem grew substantially during glacials due to lower sea level increasing the surface area of the peninsula by three-fold (Stehli and Webb, 1985). Niche partitioning occurred within the Pleistocene glacial and interglacial habitats and multiple species of animals were able to survive together with environmental changes (DeSantis et al., 2009; Yann and

DeSantis, 2014). Interglacial sea level rise and subsequent decrease in Florida landmass could also have lead to competition between and among deer and other browsers due to reduction of resources. Carbon isotopes taken from enamel of herbivores from LSP1A show that both browsers and some mixed feeders competed with deer for resources during interglacials (DeSantis et al., 2009). Amidst competition, living deer typically will choose forage (such as grasses) not targeted by other herbivores even if it is less nutritious (Armstrong and Young., 1982; Geist, 1998). However, the results from this study do not suggest that growth was directly affected by these parameters, and much more research should be done within modern deer populations to understand the growth dynamics in populations under high competition and high predation pressures.

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Migration

It is important to note that animals can migrate into and out of environments during periods of stress, and there is some evidence of this occurring between glacial and interglacial periods in Florida

(Hoppe and Koch, 2007). Even so, strontium isotopic signatures from enamel of deer revealed that deer were not migrating outside of Florida between glacial and interglacial climatic shifts (Hoppe and Koch,

2007). Thus, changes observed in this study represent native populational changes rather than introduced population mixing.

Intrinsic factors

Phenology

Differences in early ontogenetic growth observed within populations may be affected by the availability/quality of forage acquired by the doe both during pregnancy and nursing (French et al., 1956;

Ullrey et al., 1967; Monteith et al., 2009). Changes in seasonal timing of breeding and fawning

(phenology) will affect what vegetation is available for all deer (bucks, does, fawns; Richter and Labisky,

1985; Garrison and Gedir, 2006). Geographically in Florida, deer show highly divergent breeding periods with as much as a six month lag between Panhandle (February – March) and Southern Florida. (July –

August) (Richter and Labisky, 1985, Garrison and Gedir, 2006).

The phenology of fawn births in southern Florida O. virginianus corresponds with the dry season and may have evolved with the hydroperiod of the Everglades, leading to fawning during February –

March, the driest time of year (Richter and Labisky, 1985). Fawning during the dry period in southern

Florida may lead to malnutrition for both the female and fawn if the region is unusually dry, but may also improve the chances for the fawn to become large enough to survive flooding events during the wet period later in the year than if they were born in spring (Garrison and Gedir, 2006). Although, flooding may also lead to further malnutrition by reducing land, congregating populations onto higher ground, exhausting the food supply and increasing predation and disease transmission (Garrison and Gedir, 2006).

Since the NH population is assumed to be least ~500 yrs old it is likely that the phenology of the

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population at NH did not differ from modern populations, which could have influenced observed differences in growth within NH in connection with environmental circumstances discussed above.

Further understanding of how timing of the rut and phenology influences growth dynamics within modern populations and improvements on understanding nuances in the interpretations of past environments can improve our knowledge of this effect on fossil populations.

Sexual dimorphism

Sexual dimorphism in cervids, like that of most mammals, has males growing larger in size than females within a given population (Geist, 1998; Hewitt, 2011). Histological studies on European cervids confirm that male deer grow faster and longer than females (Marín- Moratalla, et al., 2013; Jordana et al.,

In Press). Fusion of epiphyses also occurs much later in modern O. virginianus bucks, indicates delayed skeletal maturity compared to females (Purdue, 1983). Modern population of adult deer used in MDC and length measurements consisted of much higher percentages of females than males (Table 18), which may have skewed some of the data. Limited sample size available for sectioning plus the unassociated nature of the limb material from each fossil population means that it is not possible to identify sex in any fossil individual in this study. Thus, it is possible that either sex may be underrepresented in any of the population samples, which would influence population averages. Comparison of the intra-populational variations among femora and humeri APDs and ANTGRs from ING1A vs. NH show variation within localities (Fig. 23A, 25A, 29A, 30A and 31A), although it is unknown to what degree these differences are influenced by sex. Future detailed studies of histological differences between O. virginianus bucks and does may allow identification of sex in an isolated limb bone in the future, but such is not possible at this time.

Comparing growth between O. virginianus and other cervids

Kolb et al. (2015a) calculated femora and tibiae growth rates based on anterior GZ measurements for modern and some fossil European cervids (Fig. 36-37). When comparing femora growth rate per GZ,

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fossil O. virginianus populations from this study plot among the Cervus elaphus (red deer; Linnaeus,

1758) extinct Candiacervus (extinct deer from island of Crete; Kuss, 1975 ), and Dama dama (fallow deer; Linnaeus, 1758) (Fig 36). Megaloceros giganteus (Irish elk; Blumenbach, 1799) and Alces alces

(moose; Linnaeus, 1758) and exhibit greater femora growth rates within GZ2-3 and hypothetically a greater growth rate in GZ1, (Fig. 36) when compared to most other cervids, although, no measurements were available in GZ1 due to resorption (Kolb et al., 2015a). Smaller to moderate-sized cervids show equivalent femora growth rates within GZ3 and beyond; the largest taxa, M. giganteus and A. Alces, have equivalent rates that extend into GZ4 (Fig 36). This supports the hypothesis that the majority of active growth in cervids occurs within the first three years.

Tibiae growth rates show a different pattern among cervids with the greatest growth rates observed in ING1A O. virginianus population in GZ2, while M. giganteus growth rate is most similar to

LSP1A O. virginianus during early ontogeny and is similar to all O. virigianus populations later in ontogeny (Fig. 37). Procervulus praelucidus (Obergfell, 1957) has similar growth rates to COL2A and

NH O. virginianus populations during early ontogeny, but the former species diverges to lower growth rates during later ontogeny (Fig. 37). All other cervids (except Candiacervus sp. II) show tibiae growth rate that are more similar NH to O. virginianus population, but diverge to lower values than O. virginianus populations later in ontogeny (Fig 37). It is unlikely that O. virginianus tibiae grew larger than tibiae from Megaloceros, thus very small sample sizes of tibiae among O. virginianus populations may account for variation observed between this element and femora. Femora provide a larger sample size, and thus offers a more accurate comparison to other cervid species.

Kolb et al. (2015a) found similar growth rates among cervids of the same size and hypothesized that growth rates may relate to metabolism and body mass. They also found that growth strategies and remodeling follow phylogenetic relationships although on islands different growth strategies may be adapted (dwarfism, Candiacervus; also seen in Myotragus (Bate, 1909); Köhler and Moya-Sola, 2009).

Average mass of modern O. virginianus from Florida is between 57 to 43 kg (Garrison and Gedir, 2006), which is most comparable to Candiacervus sp. (59 kg) (Kolb et al., 2015a) The femora growth rate of

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Candiacervus sp., Canadiacervus ropalophorus (De Vos, 1979) are most similar to NH population in rate and slope (Fig. 36). This would support earlier conclusions that little to no change has occurred between the NH and modern Florida populations.

Femora growth rates and slopes among ING1A and LSP1A are most similar to D. dama and C. elephus (Fig. 36). Dama dama has a body mass size of 70 kg (Kolb et al., 2015a), which may not be completely unreasonable (~12 kgs larger than on average modern FL. deer). Femora EOGR among

ING1A deer is very similar to C. elaphus (Fig. 36); although, it is difficult to imagine deer in Florida with a weight of 200 kg, thus differences in environment and physiology likely is a larger factor in determining growth rate and body mass, although early to mid-Pleistocene deer may have been slightly larger in body size.

Kolb et al. (2015a) found that within Cervidae there were phylogenetic similarities among all histological features such as general bone tissue type, vascularity and secondary remodeling and any variation was due to island dwarfism in poor resource environment. Histological descriptions of fossil and modern deer from Europe, (Kolb et al. 2015a; Jordana et al. In Press) show similarities to O. virginianus from Florida with fibro-lamellar bone complexes along with well-connected networks of plexiform vascular canals within early GZs and less connected laminar to longitudinal vascular canals within later

GZs indicating a reduction in growth. This study also supports the earlier ideas that much of the variation in growth and histologic structures may relate to resource poor environments, resulting in lower metabolism (Kolb et al., 2015a; Köhler and Moya-Sola, 2009).

The divergence of the two main tribes within Cervidae; Cervinae and Capreolini (group of animals including all Capreolinae) occurred ~ 7.9-7.4 mya (Gilbert et al., 2006; Fig. 38). The conservation of a similar growth history for cervids through time suggests a common growth strategy for the most recent common ancestor of these two tribes. Even so, outside variables may have a larger effect than evolutionary relationships on growth strategy (Köhler and Moya-Sola, 2009). More histological studies on early cervids are needed to provide insight into how the growth evolved within this group.

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Jordana et al. (In Press) compared femora APDs of many bovids and some cervids that provide comparison with this study. In their study they looked at three species of modern European deer, Cervus elaphus hispanicus (red deer; Hilzheimer, 1909), Rangifer tarandus platyrhynchus (reindeer; Vrolik,

1829) and Capreolus capreolus (roe deer; Linnaeus, 1758). APD calculations show marked differences in early ontogenetic growth among these three species, as well as overall shape of the growth curve. Male individuals from C. elaphus show much more sustained growth all the way into GZ5; while R. tarandus individuals show sustained growth into GZ4 (Jordana et al. In Press). This is distinct from O. virginianus, where most active growth occurs within the first 3 years of life. This difference is likely a result of differences in timing of the onset of sexual maturity; ~ 1.5 years for male O. virginianus, after which time males begin placing more resources into seasonally growing antlers rather than their skeleton (Garrison and Gedir, 2006). Overall APD values were greater in C. elaphus as well as R. tarandus and is likely due to differences in body size (C. elaphus: ~200 kg; R. tarandus: ~ 112 kg) (Finstad and Prichard, 2000;

Kolb et al., 2015a).

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10 Average Anterior Femora Growth

Rates (260 day growth period) among Cervidae

8 O. virginianus (ING) O. virginianus (LSP) O. virginianus (NH) Megaloceros giganteus Dama dama 6 Candiacervus sp. Canadiacervus ropalophorus Procervulus praelucidus Alces alces 4 Cervus elephus Capreolus capreolus Muntiacus muntjak

2 Growth rate (µm/day) rate Growth

0 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Growth Zone

Figure 36. Femora anterior growth rates per growth zone for a 260 day growth period. O. virginianus growth rates are from this study; other growth rates obtained from Kolb et al. (2015a). Opened shapes represent data from a single individual. GZ1* is a minimal growth rate due to partial resorption of inner cortex.

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10 Average Anterior Tibiae Growth Rates (260 day growth period)

8 O. virginianus (ING) O. virginianus (LSP) O. virginianus (COL) O. virginianus (NH) Megaloceros giganteus 6 Dama dama Candiacervus sp. II Canadiacervus ropalophorus Procervulus praelucidus 4

Growth rate (µm/day) rate Growth 2

0 GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7

Growth Zone

Figure 37. Tibiae anterior growth rates per growth zone for a 260 day growth period. O. virginianus growth rates are from this study; other growth rates obtained from Kolb et al. (2015a). Opened shapes represent data from a single individual. GZ1* is a minimal growth rate due to partial resorption of inner cortex.

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Figure 38. Generalized Cervidae phylogeny based on mitochondrial DNA and skeletal morphology with added fossil genera *. Taxa included were choosen based on involvement with previous histological studies (Kolb et al., 2015a; Jordana et al., In Press). (Based on Hassanin et al.,

2012; with added fossil genera from Kolb et al., 2015a).

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Future Directions

A thorough study of bone microstructure of modern Florida deer could help elucidate patterns of growth observed within fossil deer. This type of study would also provide information regarding changes in growth and bone microstructure as it relates to sex, ontogenetic stage, geography, predators, competition, and environmental variables. There is a complication in that modern Florida deer and many southeastern deer populations have been repopulated by translocated individuals from other states

(Blackard, 1971). Populations from Wisconsin, Texas, Pennsylvania and Louisiana have all been transferred into Florida to help mitigate population restoration efforts (Blackard, 1971). Ellsworth et al

(1994) found that two populations in Florida (Citrus County and Three Lakes Wildlife Management Area) showed that males translocated had contributed to the genetic makeup of the population. Even so, most other populations throughout Florida and the southeast showed little to no effect of genetic diversity from translocated deer (Ellsworth et al., 1994). Thus, future studies should take this data into account when observing and interpreting “modern native” populations and comparing them to fossil populations.

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Conclusions

This project represents one of the most detailed studies of populational changes in bone microstructure through geochronologic and climatic/environmental changes of a single species from a geographically restricted area. Additionally, this is also the most detailed study to date of the microstructure of a single cervid species from the tribe Capreolinae. Results reveal that changes in growth rate and skeletal size of O. virginianus extend across the known chronologic range of this species in

Florida.

Bone microstructure reveals that vascular canals in early ontogenetic fibrolamellar tissue were both larger and more interconnected in chronologically older populations, corresponding to increased growth rates. All sampled adult elements had evidence of growth interruption in humeri consisting of zones of slowed apposition (annuli, lamellar and parallel-fibered tissue), whereas periodic stoppages of growth (LAGs) were common in femora and tibiae.

Retrocalculation revealed that many ontogenetically older specimens had resorbed LAG1

(corresponding to the entire first year of active growth), supporting conclusions from previous studies on artiodactyls. Growth rates (calculated through thickness differences in LAGs revealed more rapid growth in early ontogeny (the first 2 years) in the early-mid Pleistocene populations (ING1A compared to the

Holocene one (NH).

Osteocyte density can be used to support changes in growth rates as well as correlation to skeletal size, although there are inconsistencies in results generated using two different methodologies. It is important to choose specimens of similar size so that there is little influence of body mass on osteocyte density, as well as use consistent counting and measuring methods.

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Some differences were observed between environmentally similar populations (ING1A vs.

COL2A; LSP1A vs. NH) but external conditions alone cannot explain changes in growth rates. Rather than fluctuating in response to glacial and interglacial conditions, histological and skeletochronological analyses reveal a general trend of decreasing growth rate through time.

One of the most important factors that can influence bone growth in deer is resource availability

(quantity) and quality of forage. Environmental responses to changes in climate patterns (e.g., precipitation, flooding, fire frequency) can affect the fertility of soil and in turn the vegetation available to support growth in Florida O. virginianus, especially among southern Florida populations. Thus, a combination of changes in climate plus resource availability likely resulted in the general trend of decreasing growth rate through time, but intrinsic factors (such as differences in phenology) may have played a role as to what resources were available for the doe during pregnancy and the fawn in early ontogeny.

Overall, more histological data is needed especially from modern populations under controlled conditions to separate out the varying degrees to which intrinsic and extrinsic factors influence bone growth within this species.

Comparison of O. virginianus with other cervids reveals much faster early ontogenetic growth rates in the femora of early to mid-Pleistocene populations (ING1A and LSP1A) relative to similarly sized cervids (extinct Candiacervus; 59 kg). Pleistocene O. virginianus growth rates were mostly similar to the slightly larger Dama dama (70 kg) and much larger Cervus elephus (200 kg), although Pleistocene

Florida deer were likely not as large as C. elephus. These comparisons suggest a larger body size among

Pleistocene O. virginianus from Florida when compared to modern size (56 kg). Limb measurements

(MDC), bone microstructure and APD analyses also generally support this conclusion. Growth rates obtained here should be considered minimal estimates for the species based on the nature of unassociated, disarticulated specimen samples here (i.e. exact body size is unknown) and expressed differences in size by latitude observed in O. virginianus. All cervids seem to grow most rapidly during their first 3 years of life. Differences in APD between O. virginianus and other cervids is likely attributed to differences in

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relative body size, timing of sexual maturity, and shifting of resources from growth to reproducing (in females) and growing antlers (males).

This study provides a bounty of information on the populational dynamics of growth history in a mammal species, which up to this point has been sparely studied due to lack of sufficient sample sizes in the vertebrate fossil record. Fossil cervids, although relatively rare in their early evolutionary history, provide a key group to study and observe how evolution may impact a group through climate transitions such as the current one. Further, O. virginianus is unique because it represents both a significantly understudied fossil species and a wealth of knowledge from modern populations. Modern populations can provide potential information regarding how bone apposition is affected by responses to specific variables such as gender, geography, resource availability, ontogeny, environment and climate. This study addresses populational changes in growth as a function of time, geography, and climate/environment. It provides data that suggest bone microstructure and apposition rates may not be consistent within a species through time, space and environment. Odocoileus virginianus may provide a significant step forward toward correlating growth history of a fossil organism with specific climatic/environmental variables.

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Appendix

The following tables consist of all raw data collected and used in graphical and statistical analyses.

Table S1. All fossil specimens measured for this study. Organization of the table is geochronologic

(NALMA; refer to Fig. 1). Locality abbreviations: ING1A: Inglis 1A; COL2A: Coleman 2A; LSP1A, 3B:

Leisey Shell Pit 1A, 3B; DD: Devil’s Den; LEC2AVS: Lecanto 2A Veronica Site; LEC3A: Lecanto 3A;

Red1A: Reddick 1A; RSRBS: Rainbow Springs Run Bear Site; S. Cave: Sabertooth Cave; Vero: Vero

Beach Canal Site; WPBS: West Palm Beach Site; NH: Nichol’s Hammock. (All localities geographic positions given in Fig. 1). * = Uncatalogued specimens with temporary catalogue number for this study.

Assoc = Associated skeleton; FE = Femur; HU = Humerus; MC = Metacarpal; MT = Metatarsal; RA =

Radius; TB = Tibia. MDC = Mid-diaphysis circumference; OS = Ontogenetic Stage (based on Table 2;

Purdue, 1983; Flinn et al., 2013).

Fusion of Epiphyses

UF Number Locality Element Length (cm) MDC (cm) Distal Proximal OS UF10890 Haile 15A L/FE 208.34 5.5 yes Yes 3 UF10891 Haile 15A L/HU 182.48 6 yes Yes 3 UF10893 Haile 15A R/HU 172.27 - yes Yes 3 UF18246H/2d* ING1A ?/MC - 5.48 ? ? ? UF18246d* ING1A ?/MC - 5.4 no ? 1-2 UF276295 ING1A ?/MC - 5.2 yes ? 3

156

ING. Ial (unlabeled)* ING1A ?/MC - 5.2 yes ? 3 UF45062 ING1A L/FE - ? yes 2-3 UF45056 ING1A L/FE - 7.7 ? no 1 UF18248H/2h* ING1A L/FE - 6 ? no 1 UF300296 ING1A L/FE - 7 ? no 1 UF18248h/1a* ING1A L/FE - 6.55 no no 1 UF300297 ING1A L/FE - 6.2 no no 1 UF300295 ING1A L/FE - 5.5 no no 1 UF18248H/2i* ING1A L/FE - 6.1 no no 1 UF18248H/2j* ING1A L/FE - 6 no no 1 UF18248H/3b* ING1A L/FE - 5.65 no no 1 UF18248H/3c* ING1A L/FE - 6 no no 1 UF18248H/3d* ING1A L/FE - 5.9 no no 1 ING.1Aa (unlabeled)* ING1A L/FE - 7.6 no no 1 UF45064 ING1A L/FE - 6.5 no no 1 UF45065 ING1A L/FE 6.75 no no 1 UF18248H/1c* ING1A L/FE - 6.3 no no 1 UF45040 ING1A L/FE - 7.5 yes ? 3 UF45044 ING1A L/FE - 6.87 yes ? 3 UF45066 ING1A L/FE 245.85 7 yes yes 3 UF45067 ING1A L/FE 251.24 7.2 yes yes 3 UF300298 ING1A L/FE - 6.5 yes ? 3 UF300294 ING1A L/FE - 6.25 yes ? 3 UF300299 ING1A L/HU - 6.88 ? ? ? UF43599 ING1A L/HU - 6.63 no yes 1-3 UF302000 ING1A L/HU - 5.6 no no 1 UF43598 ING1A L/HU - 5.87 partial no 2 UF43597 ING1A L/HU - 6.05 partial no 2 UF43596 ING1A L/HU - 6.42 yes no 2 UF43600 ING1A L/HU - 5.35 yes ? 2-3 UF43595 ING1A L/HU - 6.9 yes no 2 UF300300 ING1A L/HU - 6.05 yes ? 2-3 UF18243c* ING1A L/HU - 6.08 yes ? 2-3 UF18243d* ING1A L/HU - 5.72 yes ? 2-3 UF18243h/1a* ING1A L/HU - 6.4 yes no 2 UF18243h/1b* ING1A L/HU 162.92 5.68 yes yes 3 UF18243h/1c* ING1A L/HU 201.82 7.8 yes yes 3 UF18243 (2)* ING1A L/HU - 6.35 yes ? 2-3 UF18243(H/3)a* ING1A L/HU - 5.89 yes no 2 302001+ 302002 ING1A L/HU 192.61 7.78 yes yes 3 UF18243(H/3)d* ING1A L/HU - 6.2 yes ? 2-3 UF18243g* ING1A L/HU - 6.5 yes ? 2-3 UF276290 ING1A L/MC - 6.05 ? yes 1-3 UF276361 ING1A L/MC - 5.3 ? yes 1-3 UF276364 ING1A L/MC - 5.25 ? yes 1-3

157

UF276365 ING1A L/MC - 4.75 ? yes 1-3 UF276366 ING1A L/MC - 5.9 ? yes 1-3 UF276367 ING1A L/MC - 5.4 ? yes 1-3 UF18246c* ING1A L/MC - 5.4 ? yes 1-3 UF18246H/1d* ING1A L/MC - 5.4 ? yes 1-3 ING IA H/1a (Martin, ING1A L/MC - 5.4 ? yes 1-3 1978)* ING 1Aa (Martin 1978)* ING1A L/MC - 5.1 ? yes 1-3 ING. IAh (unlabeled)* ING1A L/MC - 5.58 no yes 2 UF276288 ING1A L/MC 182.31 5.29 yes yes 3 UF276289 ING1A L/MC 185.72 5.32 yes yes 3 UF276373 ING1A L/MT - 6.7 ? yes 1-3 UF276376 ING1A L/MT - 5.75 ? yes 1-3 UF276374 ING1A L/MT - 5.77 ? yes 1-3 UF276372 ING1A L/MT - 5.95 ? yes 1-3 ING 1Ad (Martin 1978)* ING1A L/MT - 5.05 ? yes 1-3 UF276469 ING1A L/MT - 5.5 ? yes 1-3 UF276470 ING1A L/MT - 5.75 ? yes 1-3 UF276471 ING1A L/MT - 5.85 ? yes 1-3 UF18252a* ING1A L/MT - 4.65 no yes 1-2 UF18252b* ING1A L/MT - 5.65 no yes 1-2 ING. IAk (unlabeled)* ING1A L/MT - 5.35 no yes 1-2 UF18252d* ING1A L/MT - 5.68 no yes 1-2 UF18252H/1a* ING1A L/MT - 4.75 no yes 1-2 UF18252g* ING1A L/MT - 5.5 no yes 1-2 UF276298 ING1A L/MT 207.78 5.35 yes yes 3 UF276347 ING1A L/MT 225.71 5.5 yes yes 3 UF276300 ING1A L/MT 229.31 6.35 yes yes 3 UF276348 ING1A L/MT 216.75 6.15 yes yes 3 UF276299 ING1A L/MT 229.47 6.4 yes yes 3 UF276349 ING1A L/MT 201.48 5.55 yes yes 3 UF45011 ING1A L/RA - 5.79 ? yes 2-3 UF45006 ING1A L/RA - 5.52 ? yes 2-3 UF18245b* ING1A L/RA - 5.75 ? yes 2-3 UF45007 ING1A L/RA - 6.26 no yes 2 UF45008 ING1A L/RA - 5.74 no yes 2 UF18245H/1b* ING1A L/RA - 6.25 no ? 1-2 UF300292 ING1A L/RA - 5.25 no ? 1-2 UF18245(2A)* ING1A L/RA - 5.9 no ? 1-2 UF300293 ING1A L/RA - 5.39 no yes 2 UF300291 ING1A L/RA - 5.07 no ? 1-2 UF276260 ING1A L/RA 223.73 6.35 yes yes 3 UF276267 ING1A L/RA - 6.2 yes ? 3 UF18245d* ING1A L/RA - 5.9 yes ? 3 UF276266 ING1A L/RA 201.5 6.15 yes yes 3 UF45086 ING1A L/TB - 7.5 ? yes 3 ING. 1Ag (unlabeled)* ING1A L/TB - 6.65 ? no 1-2

158

UF18249H/2a* ING1A L/TB - 6.2 no no 1 UF302003 ING1A L/TB - 5.7 no no 1 UF45077 ING1A L/TB - 7.55 yes ? 2-3 UF45076 ING1A L/TB - 7.26 yes ? 2-3 UF276273 ING1A L/TB - 7 yes ? 2-3 UF276274 ING1A L/TB - 6.9 yes ? 2-3 UF276278 ING1A L/TB - 6.9 yes no 2 UF18249b* ING1A L/TB - 7.25 yes yes 3 UF276280 ING1A L/TB - 6.95 yes ? 2-3 UF45059 ING1A R/FE - 5.85 ? no 1-2 unlabeled (R/FE)* ING1A R/FE - 6.69 ? yes 2-3 ING.1Ab (unlabeled)* ING1A R/FE - 7 ? ? ? UF18248h/2a* ING1A R/FE - 7.3 no no 1 UF18248H/2d* ING1A R/FE - 6.1 no no 1 UF18248H/2e* ING1A R/FE - 5.5 no ? 1-2 UF18248H/2f* ING1A R/FE - 5.5 no no 1 UF18248H/2g* ING1A R/FE - 5.7 no ? 1-2 UF45068 ING1A R/FE - 6.45 no no 1 UF18248h/3a* ING1A R/FE 237.13 6.63 yes yes 3 UF45046 ING1A R/FE - 7.25 yes ? 3 UF18248a* ING1A R/FE 230.9 6.9 yes yes 3 UF18248h/1b* ING1A R/FE - 7.1 yes ? 3 UF18248H/3g* ING1A R/FE - 6.85 yes ? 3 UF18248b* ING1A R/FE - 7.65 yes ? 3 UF45004 ING1A R/HU - 5.65 ? yes 2-3 UF18243i* ING1A R/HU - 6.08 ? ? ? UF43591 ING1A R/HU - 5.12 no no 1 UF43584 ING1A R/HU - 5.09 partial ? 2-3 UF43581 ING1A R/HU - 6.15 yes ? 2-3 UF43589 ING1A R/HU - 5.83 yes no 2 UF43586 ING1A R/HU - 5.95 yes ? 2-3 UF43587 ING1A R/HU - 5.78 yes ? 2-3 UF18243a* ING1A R/HU 163.53 5.8 yes yes 3 UF18243e* ING1A R/HU - 7.08 yes ? 2-3 UF18243(H/3)b* ING1A R/HU 179.8 6.15 yes yes 3 UF18243 (H/3)c* ING1A R/HU 179.15 6.07 yes yes 3 UF18243f* ING1A R/HU - 6.01 yes ? 2-3 UF276287 ING1A R/MC - 6.28 ? yes 1-3 UF276285 ING1A R/MC - 5.98 ? yes 1-3 UF276286 ING1A R/MC - 5.3 ? yes 1-3 UF276358 ING1A R/MC - 5.15 ? yes 1-3 UF276360 ING1A R/MC - 4.95 ? yes 1-3 UF18246H/1c* ING1A R/MC - 5.3 ? yes 1-3 ING. IAf (unlabeled)* ING1A R/MC - 5.5 ? yes 1-3 UF18246H/1a* ING1A R/MC - 5.1 no yes 2-3 UF18246a* ING1A R/MC - 4.6 no yes 2-3 UF18246b* ING1A R/MC - 4.98 no no 1

159

UF18246H/3a* ING1A R/MC - 4.75 no yes 2-3 UF18246H/2a* ING1A R/MC - 4.7 no yes 2-3 UF18246H/1b* ING1A R/MC - 5.25 no yes 2-3 ING. IAi (unlabeled)* ING1A R/MC - 4.78 no yes 2-3 ING. IAj (unlabeled)* ING1A R/MC - 5.13 no yes 2-3 ING 1Ac (Martin 1978)* ING1A R/MC - 4.15 no yes 2-3 UF276284 ING1A R/MC 173.82 5.43 yes yes 3 UF276283 ING1A R/MC 189.03 5.05 yes yes 3 UF276282 ING1A R/MC 189.7 5.8 yes yes 3 UF276281 ING1A R/MC 188.23 5.9 yes yes 3 ING 1Ab (Martin 1978)* ING1A R/MC - 5.6 ? yes 1-3 UF18252f* ING1A R/MT - 5.2 ? yes 1-3 UF18252H/1b* ING1A R/MT - 5.55 ? yes 1-3 UF276378 ING1A R/MT - 6.23 ? yes 1-3 UF276371 ING1A R/MT - 6.15 ? yes 1-3 UF276370 ING1A R/MT - 6.2 ? yes 1-3 UF276368 ING1A R/MT - 6.6 ? yes 1-3 UF276369 ING1A R/MT - 6.57 ? yes 1-3 UF276466 ING1A R/MT - 6.3 ? yes 1-3 UF276467 ING1A R/MT - 6.15 ? yes 1-3 UF18252H/2a* ING1A R/MT - 5.3 ? ? ? UF18252h* ING1A R/MT - 6 ? yes 1-3 UF18252c* ING1A R/MT - 4.72 no yes 1-2 UF18252e* ING1A R/MT - 5.7 no yes 1-2 UF276296 ING1A R/MT 213 5.98 yes yes 3 UF276297 ING1A R/MT 218.8 5.9 yes yes 3 UF276463 ING1A R/MT 225.73 6.15 yes yes 3 UF18245a* ING1A R/RA - 5.58 ? yes 2-3 UF45005 ING1A R/RA - 6.11 no yes 2 UF18245H/2a* ING1A R/RA - 5.06 no ? 1-2 UF18245H/1a* ING1A R/RA - 5.69 no no 1 UF45318 ING1A R/RA - 4.9 no no 1 UF45009 ING1A R/RA - 6.06 yes ? 3 UF45010 ING1A R/RA - 6.01 yes ? 3 UF276262 ING1A R/RA 197.21 6.28 yes yes 3 UF276271 ING1A R/RA - 6.5 yes ? 3 UF45012 ING1A R/RA - 5.52 yes ? 3 UF276265 ING1A R/RA 195.53 5.5 yes yes 3 UF45087 ING1A R/TB - 7.7 ? yes 2-3 UF276272 ING1A R/TB - 6.7 no ? 1 UF45078 ING1A R/TB - 6.85 yes ? 2-3 UF45069 ING1A R/TB - 8.15 yes ? 2-3 UF45075 ING1A R/TB - 7.2 yes ? 2-3 UF45082 ING1A R/TB - 7.5 yes ? 2-3 UF18249H/1a* ING1A R/TB - 7.5 yes ? 2-3 UF45088 ING1A R/TB - 6.55 yes no 2 UF11952d* COL2A L/FE - 5.75 ? no 1

160

UF11952b* COL2A L/FE 213.39 6.15 yes yes 3 UF300285 COL2A L/HU - 4.95 ? ? ? UF300284 COL2A L/HU - 5.5 yes ? 2-3 COL2Ab* COL2A L/MC - 4.05 ? yes 1-3 COL2Aa* COL2A L/MT - 5.25 no yes 1-2 UF276497 COL2A L/MT 227.33 5.73 yes yes 3 UF11961a* COL2A L/RA - 3.52 no no 1 UF276489 COL2A L/TB - 6.55 yes ? 2-3 UF276490 COL2A L/TB - - yes ? 2-3 UF276488 COL2A L/TB - 5.7 yes ? 2-3 UF276502 COL2A L?/MT - 5.17 yes ? 3 UF276503 COL2A L?/MT - 5.26 yes ? 3 UF300287 COL2A R/FE - 5.85 ? no 1-2 UF300286 COL2A R/FE - - yes ? 3 UF300283 COL2A R/HU - 5.3 yes ? 2-3 UF11955b* COL2A R/MC - 4.55 no yes 2-3 UF276493 COL2A R/MT - 5.87 ? yes 1-3 UF276494 COL2A R/MT - 5.8 ? yes 1-3 UF276495 COL2A R/MT - 5.09 ? yes 1-3 UF276496 COL2A R/MT - 5.62 ? yes 1-3 UF11955a* COL2A R/MT - 5.4 no yes 1-2 UF276486 COL2A R/TB 254.83 5.85 yes yes 2 UF300288 COL2A R?/FE - 5.85 ? ? ? UF86378 LSP1A L/FE - 6.45 yes ? 3 UF81121 LSP1A L/HU - 5.73 ? ? ? UF65962 LSP1A L/HU - 6.35 yes ? 2-3 UF87904 LSP1A L/HU - 6.23 yes ? 2-3 UF87911 LSP1A L/MC 185.56 5.23 yes yes 3 UF81538 LSP1A L/MT - 5.45 ? yes 1-3 UF87914 LSP1A L/MT 225.69 6.12 yes yes 3 UF226999 LSP1A L/TB - 6.35 yes ? 2-3 UF239928 LSP1A L/TB - - yes ? 2-3 UF85251 LSP1A L/TB - - yes ? 2-3 UF239925 LSP1A R/FE - 6.9 yes ? 3 UF87644 LSP1A R/HU 159.74 6.25 yes yes 3 UF87894 LSP1A R/MC - 5.25 ? yes 1-3 UF88030 LSP1A R/MC 185.38 5.08 yes yes 3 UF87897 LSP1A R/MT 225.66 5.84 yes yes 3 UF87893 LSP1A R/RA 200.92 5.65 yes yes 3 UF102671 LSP1A3B L/HU 149.58 5.5 yes yes 3 UF226394 LSP1A3B L/MT - 5.3 yes ? 3 UF102658 LSP1A3B L/RA - 5.55 ? yes 2-3 UF130025 LSP1A3B R/HU 178.43 6.15 yes yes 3 UF219938 LSP1A3B R/HU 174.1 6.3 yes yes 3 UF219939 LSP1A3B R/MC - 4.88 ? yes 1-3 UF219941 LSP1A3B R/MT - 5.65 ? yes 1-3 UF219940 LSP1A3B R/TB 255.56 5.85 yes yes 2

161

UF51662 DD L/FE - 6.1 no no 1 UF9814h* DD L/FE - 6.45 no no 1 UF9814i* DD L/FE - 5.95 no no 1 UF9814j* DD L/FE - 5.1 no no 1 UF9814k* DD L/FE - 4.95 no no 1 UF9814l* DD L/FE - 5 no no 1 UF9814m* DD L/FE - 5.2 no no 1 UF9814n* DD L/FE - 4.15 no ? 1-2 UF9814o* DD L/FE - 4.95 no ? 1-2 UF9814c* DD L/FE 221.3 5.65 partial partial 2 UF9814f* DD L/FE - 6.7 partial partial 2 UF9814a* DD L/FE 198.19 5.92 yes yes 3 UF9814b* DD L/FE 232 6.8 yes yes 3 UF9814d* DD L/FE 225.22 - yes yes 3 UF9814e* DD L/FE - 6.6 yes no 2 UF9814g* DD L/FE - - yes no 2 UF9814p* DD L/FE - 6.05 yes partial 2 UF9814q* DD L/FE 219.91 6.5 yes yes 3 UF9834f* DD L/FE 228.5 - yes yes 3 UF9834g* DD L/FE 233.9 6.3 yes yes 3 UF9813m* DD L/HU - 5.5 no no 1 UF51659 DD L/HU - 4.5 no ? 1-3 UF9840c* DD L/HU - 7.15 partial ? 2-3 UF9840a* DD L/HU 176.56 6.4 yes ? 2-3 UF9813c* DD L/HU 183.24 6.4 yes yes 3 UF9813d* DD L/HU - 6.6 yes no 2 UF9813e* DD L/HU - 6 yes no 2 UF9813f* DD L/HU - 4.8 yes no 2 UF9813g* DD L/HU - 6.63 yes ? 2-3 UF9813j* DD L/HU 181.66 6.6 yes yes 3 UF51660 DD L/HU - 6.35 yes no 2-3 UF9813n* DD L/HU 181.5 6.3 yes yes 3 UF9790q* DD L/RA - 6.3 ? partial 1-2 UF51665 DD L/RA - 6.5 ? yes 2-3 UF9790 DD L/RA - 6.55 no yes 2 UF9790n* DD L/RA - 4.4 no no 1 UF9790o* DD L/RA - 5.9 no yes 2 UF9790p* DD L/RA - 4.35 no partial 2 UF9790r* DD L/RA - 5.6 no partial 2 UF9790a* DD L/RA 199.86 5.4 yes yes 3 UF9790b* DD L/RA 197.81 6.9 yes yes 3 UF9790c* DD L/RA 180.9 4.7 yes yes 3 UF9790d* DD L/RA 204.21 5.65 yes yes 3 UF9790e* DD L/RA 201.13 6.75 yes yes 3 UF9790f* DD L/RA 200.38 6.2 yes yes 3 UF9790m* DD L/RA - 5.75 yes ? 3 UF9832a* DD L/RA 203.7 - yes yes 3

162

UF9811l* DD L/TB - 5.8 ? no 1-2 UF9811n* DD L/TB - 5.6 ? no 1-2 UF9811h* DD L/TB - 6 no no 1 UF9811j* DD L/TB - 6.1 no no 1 UF9811k* DD L/TB - 5.25 no no 1 UF9811m* DD L/TB - 4.85 no ? 1-2 UF9811o* DD L/TB - 4.7 no no 1 UF9811f* DD L/TB - 6.1 partial no 1-2 UF9811* DD L/TB 261.94 6.65 yes yes 3 UF9811a* DD L/TB 269.84 7.1 yes yes 3 UF9811b* DD L/TB 266.28 - yes yes 3 UF9811c* DD L/TB 267.72 - yes yes 3 UF9811d* DD L/TB 240.8 - yes yes 3 UF9811e* DD L/TB 268.27 7 yes yes 3 UF9811g* DD L/TB - 7.2 yes no 2 UF9811i* DD L/TB - 6.6 yes ? 2-3 UF51664 DD L/TB 269.45 - yes yes 3 UF9833a* DD L/TB 262.2 - yes yes 3 UF9833b* DD L/TB 262.26 6.4 yes yes 3 UF9814d* DD R/FE - 5.65 ? yes 2-3 UF49884 DD R/FE - 5.15 no no 1 UF55257 DD R/FE - 5.2 no ? 1-2 UF9793 DD R/FE - 6.6 no no 1 UF9814e* DD R/FE - 5.6 no no 1 UF9814a* DD R/FE 232.11 8.8 yes yes 3 UF9814b* DD R/FE 233.43 8.5 yes yes 3 UF9814c* DD R/FE 213.89 - yes yes 3 UF9814f * DD R/FE - 6.8 yes no 2 UF9834a* DD R/FE 219.44 6.15 yes yes 3 UF9834b* DD R/FE 235.28 6.3 yes yes 3 UF9834c* DD R/FE 228.34 6.7 yes yes 3 UF9834d* DD R/FE - 6.65 yes no 2 UF49874 DD R/HU - 4.75 no ? 1-3 UF9793 DD R/HU - 6 no no 1 UF9840b* DD R/HU - 6.2 yes no 2 UF9813a* DD R/HU 181.76 6.4 yes yes 3 UF9813b* DD R/HU - 6.6 yes partial 2-3 UF9813h* DD R/HU 181.62 6.5 yes yes 3 UF9813i* DD R/HU - 6.8 yes no 2 UF9813k* DD R/HU - 5.83 yes no 2 UF9813l* DD R/HU - 6.13 yes ? 2-3 UF51661 DD R/HU - 6.5 yes no 2 UF9790s* DD R/RA - 4.55 no partial 1-2 UF9790t* DD R/RA - 5.85 no partial 1-2 UF9832c* DD R/RA - 5.4 no yes 2 UF9790g* DD R/RA 196.3 6.05 yes yes 3 UF9790h* DD R/RA 180.59 4.65 yes yes 3

163

UF9790i* DD R/RA 197.35 - yes yes 3 UF9790j* DD R/RA 148.98 - yes yes 3 UF9790k* DD R/RA - 5.75 yes yes 3 UF9790l* DD R/RA - 6.3 yes yes 3 UF9832b* DD R/RA 203.21 - yes yes 3 UF9811i* DD R/TB - 6.05 no no 1 UF9811j* DD R/TB - 4.8 no no 1 UF9811k* DD R/TB - 4.9 no ? 1 UF9811l* DD R/TB - 5.2 no no 1 UF51408 DD R/TB - 3.95 no no 1 UF9811a* DD R/TB 264.65 6.5 yes yes 3 UF9811b* DD R/TB 278.89 6.8 yes yes 3 UF9811c* DD R/TB 262.39 6.5 yes yes 3 UF9811d* DD R/TB 262.18 - yes yes 3 UF9811e* DD R/TB 195.89 6.4 yes yes 3 UF9811f* DD R/TB - 7.1 yes no 2 UF9811g* DD R/TB - 6.5 yes no 2 UF9811h* DD R/TB - 6.9 yes no 2 UF9811m* DD R/TB 264 6.7 yes partial 2-3 UF10255 Haile8A L/FE 208.8 6.1 yes yes 3 UF3239 Haile8A L/FE - 6.85 no no 1 UF126432 LEC2AVS L/HU - 5.95 yes no 2 UF126442 LEC2AVS L/MC 193.55 5.3 yes yes 3 UF126457 LEC2AVS L/MT 224.05 5.73 yes yes 3 UF126434 LEC2AVS L/RA 198.17 5.6 yes yes 3 UF126435 LEC2AVS L/RA - 5.15 yes yes 3 UF126447 LEC2AVS L/TB - 5.75 yes ? 2-3 UF126448 LEC2AVS L/TB - 6 yes ? 2-3 UF126431 LEC2AVS R/HU - 6.15 yes no 2 UF126458 LEC2AVS R/MT - 5.55 no yes 1-2 UF126456 LEC2AVS R/MT 222.85 5.7 yes yes 3 UF274030 LEC3A R/HU - 5.9 yes no 2 UF2441d* Red1A L/HU - 5.5 yes ? 2-3 UF2441e* Red1A L/HU - 4.68 yes ? 2-3 UF2441g* Red1A L/HU - 5.45 ? ? ? UF2441b* Red1A L/MC 180.73 5.45 yes ? 3 UF2441c* Red1A L/MT - 5.65 yes yes 3 UF2441a* Red1A L/RA - 5.05 ? yes 2-3 UF2441h* Red1A R/FE - 6.45 ? ? ? UF2402 (assoc) Red1A R/HU - 6.25 yes ? 2-3 UF2441f* Red1A R/HU - 5.8 no ? 1-2 UF2402 (assoc) Red1A R/MC - 5.5 partial yes 2-3 UF2402 (assoc) Red1A R/RA - 5.8 yes yes 3 UF266250 (assoc) RSRBS L/FE 230.4 6.3 yes yes 3 UF266250 (assoc) RSRBS L/HU 170.65 6.05 yes yes 3 UF266250 (assoc) RSRBS L/MC 191.08 5.15 yes yes 3 UF266250 (assoc) RSRBS L/MT 224.22 5.8 yes yes 3

164

UF266290 RSRBS L/RA - 5.85 ? yes 2-3 UF266250 (assoc) RSRBS L/RA 198.23 - yes yes 3 UF266250 (assoc) RSRBS L/TB - 6.28 yes ? 2-3 UF266248 RSRBS R/HU - 5.3 yes ? 2-3 UF266250 (assoc) RSRBS R/HU - 6 yes ? 2-3 UF266214 RSRBS R/MC 188.77 5.5 yes yes 3 UF266250 (assoc) RSRBS R/MC 191.5 5.15 yes yes 3 UF266250 (assoc) RSRBS R/MT 224.23 5.9 yes yes 3 UF266213 RSRBS R/RA 181.01 4.85 yes yes 3 UF266250 (assoc) RSRBS R/RA 198.23 5.8 yes yes 3 UF266250 (assoc) RSRBS R/TB - 6.1 yes ? 2-3 UF266296 RSRBS R/TB - 5 no no 1 Saber. Cave b* S. Cave R/HU - 6.15 yes ? 2-3 Saber. Cave a* S. Cave R/MC - 4.9 ? yes 2-3 Saber. Cave c* S. Cave R/TB - 6 ? yes 3 V8039 Vero L/FE - 5.95 ? no 1 V3906 Vero L/FE - 5.88 no ? 1-2 V92 Vero L/MC 202.22 5.9 yes yes 3 V420 Vero L/MT - 5.7 no yes 1-2 V422 Vero L/MT 228.5 6.23 yes yes 3 V93 Vero L/MT 251.96 6.7 yes yes 3 V2025 Vero L/RA - 5 ? yes 2-3 V1820 Vero L/RA - 5.28 no yes 2 V2059 Vero L/RA - 4.45 no no 1 V1816 Vero L/TB - 6.3 yes ? 2-3 V2762 Vero L/TB 183.4 4.8 yes yes 3 V3205 Vero L/TB - 5.85 yes no 2 V2188 Vero R/FE 237.4 6.95 yes yes 3 V684 Vero R/HU 161.51 5.7 yes yes 3 V685 Vero R/HU 183.5 6.55 yes yes 3 V3050 Vero R/HU - 5.4 yes no 2 V421 Vero R/MC 193.39 5.3 yes yes 3 V1824 Vero R/RA - 5.35 ? yes 2-3 V2904 Vero R/RA 200.34 5.5 yes yes 3 V1818 Vero R/TB - 6.25 yes ? 2-3 V3165 Vero R/TB - 6.95 yes ? 2-3 V2742 Vero R/TB 260.22 6.45 yes yes 3 UF51162 WPBS L/FE - 5.5 no ? 1-2 UF51163 WPBS L/MC - 4.85 ? yes 1-3 UF51166 WPBS L/MT 5.55 yes yes 3 UF51165 WPBS L/MT 223.43 5.55 yes yes 3 UF51161 WPBS L/RA - 5.4 yes ? 3

165

unlabeled NH Hutchison NH L/FE - 2.75 no no 1 1958b* unlabeled NH Hutchison NH L/FE - 4.4 no yes 2 1958c* UF57618 NH L/FE 211.12 6.15 yes yes 3 UF57620 NH L/FE 200.85 6.22 yes yes 3 UF57621 NH L/FE 213.1 5.6 yes yes 3 UF57622 NH L/FE 194.39 yes yes 3 UF300289 NH L/HU - 2.85 no no 1 UF300290 NH L/HU - 3.6 no no 1 UF57164 NH L/HU 167.68 5.81 yes yes 3 UF57165 NH L/HU 149.77 5.25 yes yes 3 UF57166 NH L/HU 163 5.12 yes yes 3 UF57169 NH L/HU 149.66 5.12 yes yes 3 UF 57627 NH L/MT 218.17 5.43 yes yes 3 UF57180 NH L/RA - 4.62 ? yes 2-3 UF57181 NH L/RA - 4.58 ? yes 2-3 UF57178 NH L/RA 175.02 4.67 yes yes 3 UF57179 NH L/RA 186.78 5.25 yes yes 3 UF57177 NH L/RA 168.18 4.98 yes yes 3 UF57182 NH L/RA - yes ? 3 UF57184 NH L/TB 220.43 5.84 yes yes 3 unlabeled NH Hutchison NH R/FE - 2.7 no no 1 1958a* UF57617 NH R/FE 194.03 5.58 yes yes 3 UF57619 NH R/FE 212.62 6.1 yes yes 3 unlabeled NH Hutchison NH R/HU - 3.7 no no 1 1960b* unlabeled NH Hutchison NH R/HU - 2.95 no no 1 1960c* unlabeled NH Hutchison NH R/HU - 5.82 no no 1 1960e* UF57167 NH R/HU 149.01 5.16 yes yes 3 UF57168 NH R/HU 160.37 5.55 yes yes 3 UF 37632 NH R/MC 175.13 4.71 yes yes 3 UF57631 NH R/MC 186.81 4.92 yes yes 3 UF 57626 NH R/MT - 4.95 no yes 1-2 UF 57628 NH R/MT 217.37 5.3 yes Yes 3 UF57176 NH R/RA 181.53 5.02 yes Yes 3 UF57186 NH R/TB - 3.8 no No 1 UF57185 NH R/TB - no partial 1-2

166

Table S2. All modern specimens measured for this study. Organization of the table is by UF number and county (All “Adult” specimens geographic positions are given in Fig. 1). FE =

Femur; HU = Humerus; MC = Metacarpal; MT = Metatarsal; RA = Radius; TB = Tibia. MDC =

Mid-diaphysis circumference; OS = Ontogenetic Stage (based on Table 1; Purdue, 1983; Flinn et al., 2013).

Fusion of

Epiphyses

Specimen # County Element AL MDC Distal Proximal Gender OS UF13489 Columbia L/FE 220.55 6.12 yes yes F 3 UF13489 Columbia L/HU 164.7 5.6 yes yes F 3 UF13489 Columbia L/MT 217.95 5.15 yes yes F 3 UF13489 Columbia L/RA 190.25 5.4 yes yes F 3 UF13489 Columbia L/TB 253.3 6 yes yes F 3 UF13489 Columbia R/FE 220.55 6.15 yes yes F 3 UF13489 Columbia R/MC 184.4 4.8 yes yes F 3 UF13489 Columbia R/MT 218.67 5.2 yes yes F 3 UF13489 Columbia R/RA 189.65 5.4 yes yes F 3 UF13489 Columbia R/TB 254.7 6 yes yes F 3 UF13747 Levy L/FE - 3.2 no no ? 1 UF13747 Levy L/HU - 3.1 no no ? 1 UF13747 Levy L/MC - 2.9 no no ? 1 UF13747 Levy L/MT - 2.8 no no ? 1 UF13747 Levy L/RA - 2.5 no no ? 1 UF13747 Levy L/TB - 2.9 no no ? 1 UF13747 Levy R/FE - 3.12 no no ? 1 UF13747 Levy R/HU - 3.1 no no ? 1 UF13747 Levy R/MC - 2.95 no no ? 1 UF13747 Levy R/MT - 2.8 no no ? 1 UF13747 Levy R/RA - 2.5 no no ? 1 UF13747 Levy R/TB - 2.82 no no ? 1 UF20752 Monroe L/FE 194.33 5.75 yes yes ? 3 UF20752 Monroe L/HU 151.9 5.48 yes yes ? 3 UF20752 Monroe L/MC 154 4.8 yes yes ? 3 UF20752 Monroe L/MT 176.67 4.6 yes yes ? 3 UF20752 Monroe L/RA 170.62 5 yes yes ? 3 UF20752 Monroe L/TB 213.8 5.3 yes yes ? 3 UF20752 Monroe R/FE 193.9 5.72 yes yes ? 3 UF20752 Monroe R/HU 151.95 5.55 yes yes ? 3 UF20752 Monroe R/MC 152.8 4.8 yes yes ? 3

167

UF20752 Monroe R/MT 175.65 4.6 yes yes ? 3 UF20752 Monroe R/RA 1701.3 5 yes yes ? 3 UF20752 Monroe R/TB 214.8 5.35 yes yes ? 3 UF24374 Monroe L/FE - 4.85 no yes ? 2 UF24374 Monroe L/HU 138.13 4.6 yes yes ? 3 UF24374 Monroe L/MC 145.6 4.1 yes yes ? 3 UF24374 Monroe L/MT 170.82 4.1 yes yes ? 3 UF24374 Monroe L/RA 156.71 yes yes ? 3 UF24374 Monroe R/HU 138.4 4.6 yes yes ? 3 UF24374 Monroe R/MC 146.7 4.33 yes yes ? 3 UF24374 Monroe R/MT 170.33 4.1 yes yes ? 3 UF24374 Monroe R/RA 157.9 yes yes ? 3 UF24374 Monroe R/TB - 5.75 yes yes ? 3 UF24374 Monroe L/TB 211 5.85 partially yes ? 2-3 UF24374 Monroe R/FE - 4.82 no yes ? 2-3 UF25430 Collier L/FE 226.9 5.6 yes yes F 3 UF25430 Collier L/HU 154.5 5.15 yes yes F 3 UF25430 Collier L/MC 190.3 4.75 yes yes F 3 UF25430 Collier L/MT - 5.4 yes yes F 3 UF25430 Collier L/RA 189.25 4.75 yes yes F 3 UF25430 Collier L/TB 241.68 5.7 yes yes F 3 UF25430 Collier R/FE 228.5 5.75 yes yes F 3 UF25430 Collier R/HU 154.2 5.23 yes yes F 3 UF25430 Collier R/MC 190.2 4.75 yes yes F 3 UF25430 Collier R/MT 225.54 5.4 yes yes F 3 UF25430 Collier R/RA 189 4.75 yes yes F 3 UF25430 Collier R/TB 241.1 5.75 yes yes F 3 UF3261 Marion L/FE 219.68 6.5 yes yes F 3 UF3261 Marion L/HU 177.02 5.75 yes yes F 3 UF3261 Marion L/MC 190.4 5 yes yes F 3 UF3261 Marion L/MT 224.15 5.45 yes yes F 3 UF3261 Marion L/RA 195.42 5.5 yes yes F 3 UF3261 Marion L/TB 261.91 5.9 yes yes F 3 UF3261 Marion R/FE 219.9 6.65 yes yes F 3 UF3261 Marion R/HU 177.85 5.75 yes yes F 3 UF3261 Marion R/MC 189.8 5 yes yes F 3 UF3261 Marion R/MT 223.85 5.45 yes yes F 3 UF3261 Marion R/RA 195.5 5.5 yes yes F 3 UF3261 Marion R/TB 261.26 6 yes yes F 3 UF3262 Putnam L/FE 249.31 7.3 yes yes M 3 UF3262 Putnam L/HU 195.55 6.73 yes yes M 3 UF3262 Putnam L/MC 212.74 5.7 yes yes M 3 UF3262 Putnam L/MT 246.25 5.9 yes yes M 3 UF3262 Putnam L/RA 218.31 yes yes M 3 UF3262 Putnam L/TB 295.17 6.92 yes yes M 3 UF3262 Putnam R/FE 249.07 7.3 yes yes M 3 UF3262 Putnam R/HU 195.55 6.79 yes yes M 3

168

UF3262 Putnam R/MC 211.6 5.8 yes yes M 3 UF3262 Putnam R/RA 220 yes yes M 3 UF3262 Putnam R/TB 294.66 6.92 yes yes M 3 UF3268 Levy L/HU 163.53 5.55 yes yes F 3 UF3268 Levy L/MC 185.75 5.1 yes yes F 3 UF3268 Levy L/MT 219.15 5.2 yes yes F 3 UF3268 Levy L/TB 254.25 6 yes yes F 3 UF3268 Levy R/HU 166.54 5.5 yes yes F 3 UF3268 Levy R/MC - 5.12 yes yes F 3 UF3268 Levy R/MT 220.73 5.3 yes yes F 3 UF3268 Levy R/TB 254.1 6.5 yes yes F 3 UF3270 Marion L/FE 216.47 5.85 yes yes F 3 UF3270 Marion L/HU 167.15 5.3 yes yes F 3 UF3270 Marion L/MT 222.46 5.3 yes yes F 3 UF3270 Marion L/TB 248.2 5.85 yes yes F 3 UF3270 Marion R/FE 215.92 5.95 yes yes F 3 UF3270 Marion R/MC 187.77 4.9 yes yes F 3 UF3270 Marion R/MT 221.75 5.35 yes yes F 3 UF3270 Marion R/RA 196 5.05 yes yes F 3 UF3270 Marion R/TB 249.7 5.85 yes yes F 3 UF3275 Marion L/HU - 3.65 no no M 1 UF3275 Marion L/RA - 3.25 no no M 1 UF3275 Marion L/TB - 3.7 no no M 1 UF3275 Marion R/FE - 4.1 no no M 1 UF3275 Marion R/HU - 3.65 no no M 1 UF3275 Marion R/RA - 3.2 no no M 1 UF3275 Marion R/TB - 3.7 no no M 1 UF3275 Marion L/MC - 3.3 no yes M 2 UF3275 Marion L/MT - 3.7 no yes M 2 UF3275 Marion R/MC - 3.35 no yes M 2 UF3275 Marion R/MT - 3.7 no yes M 2 UF3275 Marion L/FE - 4.2 partial no M 2-3 UF3276 Marion L/FE 228.1 6.15 yes yes F 3 UF3276 Marion L/HU 165.95 5.5 yes yes F 3 UF3276 Marion L/MC 186.45 4.8 yes yes F 3 UF3276 Marion L/MT 216.65 5.25 yes yes F 3 UF3276 Marion L/RA 194.75 5.15 yes yes F 3 UF3276 Marion L/TB 257.1 5.65 yes yes F 3 UF3276 Marion R/FE 228.55 6.18 yes yes F 3 UF3276 Marion R/HU 166.3 5.55 yes yes F 3 UF3276 Marion R/MC 187.3 4.8 yes yes F 3 UF3276 Marion R/MT 216.05 5.23 yes yes F 3 UF3276 Marion R/RA 194.53 5.1 yes yes F 3 UF3276 Marion R/TB 257.78 5.6 yes yes F 3 UF3277 Marion L/FE 223.12 6.15 yes yes F 3 UF3277 Marion L/MC 190.4 4.9 yes yes F 3 UF3277 Marion L/MT 221.22 5.5 yes yes F 3

169

UF3277 Marion L/RA 197.5 5.25 yes yes F 3 UF3277 Marion L/TB 262.35 6.1 yes yes F 3 UF3277 Marion R/FE 223.65 6.05 yes yes F 3 UF3277 Marion R/HU 170.56 5.75 yes yes F 3 UF3277 Marion R/MC 189.85 4.85 yes yes F 3 UF3277 Marion R/MT 221.3 5.52 yes yes F 3 UF3277 Marion R/RA 200.8 5.25 yes yes F 3 UF3277 Marion R/TB 261.8 6.1 yes yes F 3 UF3278 Marion L/FE - 6.15 yes ? F 3 UF3278 Marion L/HU 169.55 5.65 yes yes F 3 UF3278 Marion L/MC 190.15 4.8 yes yes F 3 UF3278 Marion L/MT 225.85 5.5 yes yes F 3 UF3278 Marion L/RA 188.5 5 yes yes F 3 UF3278 Marion L/TB 258.15 5.9 yes yes F 3 UF3278 Marion R/FE 221.65 6.2 yes yes F 3 UF3278 Marion R/HU 168.31 5.6 yes yes F 3 UF3278 Marion R/MC 190.93 4.78 yes yes F 3 UF3278 Marion R/MT 225.85 5.5 yes yes F 3 UF3278 Marion R/RA 189.5 5.05 yes yes F 3 UF3278 Marion R/TB 257.15 6 yes yes F 3 UF3279 Marion L/MC - 4.35 no yes F 2 UF3279 Marion L/RA - 4.38 no yes F 2 UF3279 Marion R/HU - 4.65 yes no F 2 UF3279 Marion R/MC - 4.35 no yes F 2 UF3279 Marion ?/MT - 4.8 no ? F 1-2 UF3279 Marion R/TB - 5.3 ? no F 1-2 UF3279 Marion L/HU - 4.75 yes partial F 2-3 UF3279 Marion L/TB - 5.25 ? partial F 2-3 UF3280 Levy R/FE - 5.4 no no F 1 UF3280 Levy R/TB - 5.13 no ? F 1 UF3280 Levy L/MC - 4.4 no yes F 2 UF3280 Levy L/MT - 4.8 no yes F 2 UF3280 Levy L/RA - 4.5 no yes F 2 UF3280 Levy R/MC - 4.4 no yes F 2 UF3280 Levy R/MT - 4.8 no yes F 2 UF3280 Levy R/RA - 4.6 no yes F 2 UF3280 Levy LTB - 5.2 partial no F 1-2 UF3280 Levy L/FE - 5.48 ? yes F 2-3 UF3280 Levy L/HU - 4.77 yes ? F 2-3 UF3280 Levy R/HU - 4.77 yes ? F 2-3 UF3281 Marion R/FE - 6 no no ? 1 UF3281 Marion L/HU - 5.6 yes no ? 2 UF3281 Marion L/TB - 6 yes no ? 2 UF3281 Marion R/HU - 5.75 yes no ? 2 UF3281 Marion R/TB - 5.95 yes no ? 2 UF3281 Marion L/MC - 5.1 yes no ? 2-3 UF3281 Marion L/MT - 5.5 yes no ? 2-3

170

UF3281 Marion L/RA - 5.1 yes no ? 2-3 UF3281 Marion R/MC - 5.1 yes no ? 2-3 UF3281 Marion R/MT - 5.55 yes no ? 2-3 UF3281 Marion R/RA - 5.15 yes no ? 2-3 UF4596 Monroe L/HU 154.5 5.2 yes no M 2 UF4596 Monroe R/HU 154.14 5.3 yes no M 2 UF4596 Monroe L/FE 209.15 5.4 yes yes M 3 UF4596 Monroe L/MC 151.7 4.55 yes yes M 3 UF4596 Monroe L/RA 173.8 4.65 yes yes M 3 UF4596 Monroe L/TB 223.9 5.12 yes yes M 3 UF4596 Monroe R/FE 210.9 5.45 yes yes M 3 UF4596 Monroe R/MT 182.45 4.42 yes yes M 3 UF4596 Monroe R/RA 174 4.75 yes yes M 3 UF4596 Monroe R/TB 224.45 5.2 yes yes M 3 UF5974 Citrus L/FE - 4.2 no no M 1 UF5974 Citrus L/TB - 4.2 no no M 1 UF5974 Citrus R/TB - 4.16 no no M 1 UF5974 Citrus L/HU - 3.93 yes no M 2 UF5974 Citrus L/MC - 3.7 no yes M 2 UF5974 Citrus L/MT - 4.1 no yes M 2 UF5974 Citrus L/RA - 3.55 no yes M 2 UF5974 Citrus R/HU - 3.9 yes no M 2 UF5974 Citrus R/MC - 3.7 no yes M 2 UF5974 Citrus R/MT - 4.15 no yes M 2 UF5974 Citrus R/RA - 3.58 no yes M 2 UF5974 Citrus R/FE - - ? no M 1-2 UF6671 Jackson L/FE - 2.8 no no ? 1 UF6671 Jackson L/HU - 4.75 no no ? 1 UF6671 Jackson L/MC - 2.1 no no ? 1 UF6671 Jackson L/RA - 2.95 no no ? 1 UF6671 Jackson L/TB - 2.72 no no ? 1 UF6671 Jackson R/HU - 4.85 no no ? 1 UF6671 Jackson R/MC - 2 no no ? 1 UF6671 Jackson R/RA - 2.85 no no ? 1 UF6671 Jackson R/TB - 2.7 no no ? 1 UF6671 Jackson R/FE - 2.8 ? no ? 1-2 UF7001 Walton L/HU 257.35 5.4 yes yes F 3 UF7001 Walton L/MC 188.95 4.6 yes yes F 3 UF7001 Walton L/MT 218.5 5.02 yes yes F 3 UF7001 Walton L/RA 190.5 - yes yes F 3 UF7001 Walton L/TB 243.55 5.5 yes yes F 3 UF7001 Walton R/HU 257.45 5.4 yes yes F 3 UF7001 Walton R/MC 188.55 4.6 yes yes F 3 UF7001 Walton R/MT 217.65 5.05 yes yes F 3 UF7001 Walton R/RA - - yes yes F 3 UF7001 Walton R/TB - 5.45 yes yes F 3 UF7001 Walton L/FE 209.1 5.9 yes partial F 2-3

171

UF7001 Walton R/FE - 5.8 yes no F 2-3 UF7003 Brevard L/FE - 6.4 no no M 1 UF7003 Brevard R/FE - 6.05 no no M 1 UF7003 Brevard L/HU - 6.35 yes no M 2 UF7003 Brevard L/MT - 5.75 no yes M 2 UF7003 Brevard L/RA - 6 no yes M 2 UF7003 Brevard L/TB - 6.65 yes no M 2 UF7003 Brevard R/HU - 6.3 yes no M 2 UF7003 Brevard R/MT - 6.3 no yes M 2 UF7003 Brevard R/RA - 6 no yes M 2 UF7003 Brevard R/TB - 6.7 yes yes M 3 UF7004 Volusia L/FE - 5.1 no no M 1 UF7004 Volusia L/TB - 5.2 no no M 1 UF7004 Volusia R/FE - 5.05 no no M 1 UF7004 Volusia R/TB - 5.15 no no M 1 UF7004 Volusia L/HU - 4.85 yes no M 2 UF7004 Volusia L/MC - 4.55 no yes M 2 UF7004 Volusia L/MT - 5 no yes M 2 UF7004 Volusia L/RA - 2.3 no yes M 2 UF7004 Volusia R/HU - 4.9 yes no M 2 UF7004 Volusia R/MC - 4.6 no yes M 2 UF7004 Volusia R/MT - 5 no yes M 2 UF7004 Volusia R/RA - 4.28 no yes M 2 UF8143 Leon L/FE - 4.95 no no F 1 UF8143 Leon L/HU - 4.55 no no F 1 UF8143 Leon L/TB - 4.95 no no F 1 UF8143 Leon R/FE - 5.03 no no F 1 UF8143 Leon R/HU - 4.5 no no F 1 UF8143 Leon R/TB - 4.9 no no F 1 UF8143 Leon L/MC - 3.8 no yes F 2 UF8143 Leon L/MT - 4.75 no yes F 2 UF8143 Leon L/RA - 4.2 no yes F 2 UF8143 Leon R/MC - 3.85 no yes F 2 UF8143 Leon R/MT - 4.85 no yes F 2 UF8143 Leon R/RA - 4.15 no yes F 2 UF8652 Marion L/FE - 6 no no M 1 UF8652 Marion R/FE - 6.02 no no M 1 UF8652 Marion L/HU - 5.5 yes no M 2 UF8652 Marion L/MC - 4.8 no yes M 2 UF8652 Marion L/RA - 5 no yes M 2 UF8652 Marion R/HU - 5.5 yes no M 2 UF8652 Marion R/MC - 4.82 no yes M 2 UF8652 Marion R/RA - 5 no yes M 2 UF8653 Marion L/HU 182.33 6.5 yes no M 2 UF8653 Marion R/HU 180.25 6.4 yes no M 2 UF8653 Marion L/FE 232.07 6.5 yes yes M 3 UF8653 Marion L/MT 231.44 5.65 yes yes M 3

172

UF8653 Marion L/RA 218 5.75 yes yes M 3 UF8653 Marion L/TB 270.79 6.15 yes yes M 3 UF8653 Marion R/FE 232.1 6.52 yes yes M 3 UF8653 Marion R/RA 218.52 5.75 yes yes M 3 UF8653 Marion R/TB 271.65 6.12 yes yes M 3 UF8654 Volusia L/FE - 5.4 no no M 1 UF8654 Volusia L/TB - 5.15 no no M 1 UF8654 Volusia R/FE - 5.3 no no M 1 UF8654 Volusia R/TB - 5.1 no no M 1 UF8654 Volusia L/HU - 4.9 yes no M 2 UF8654 Volusia L/RA - 4.3 no yes M 2 UF8654 Volusia R/HU - 4.85 yes no M 2 UF8654 Volusia R/RA - 4.35 no yes M 2 UF8741 Volusia R/FE - 5.1 no no F 1 UF8741 Volusia R/TB - 5.15 no no F 1 UF8741 Volusia L/HU - 4.9 yes no F 2 UF8741 Volusia R/HU - 5 yes no F 2 UF8741 Volusia R/RA - 4.3 no yes F 2 UF8747 Alachua L/FE - 6.62 no no M 1 UF8747 Alachua L/RA - 4 no no M 1 UF8747 Alachua R/FE - 6.5 no no M 1 UF8747 Alachua R/RA - 4 no no M 1 UF8747 Alachua L/HU - 4.4 yes no M 2 UF8747 Alachua L/MC - 4 no yes M 2 UF8747 Alachua R/HU - 4.35 yes no M 2 UF8747 Alachua R/MC - 4 no yes M 2 UF8747 Alachua TB? - 4.5 no ? M 1-2 UF9495 Duval L/FE - 5.3 no no F 1 UF9495 Duval L/TB - 5.25 no no F 1 UF9495 Duval L/HU - 4.7 yes no F 2 UF9495 Duval L/RA - 4.28 no yes F 2 UF9495 Duval R/HU - 4.65 yes no F 2 UF9495 Duval R/MT - 4.6 no yes F 2 UF9495 Duval R/RA - 4.23 no yes F 2 UF9495 Duval R/TB - 5.23 no partial F 1-2 UF9496 Duval L/RA - 6.25 no yes F 2 UF9496 Duval L/TB - 5 no yes F 2 UF9496 Duval R/RA - 6.3 no yes F 2 UF9496 Duval R/HU 138.83 4.65 yes yes F 3 UF9496 Duval L/FE 188.25 5.15 partial yes F 2-3 UF9496 Duval R/FE 188.75 5.2 partial yes F 2-3 UF9496 Duval R/MT - 4.45 partial yes F 2-3 UF9497 Duval L/TB - 4.9 no yes F 2 UF9497 Duval R/TB - 4.8 no yes F 1-2 UF9497 Duval L/FE - 5.15 yes partial F 2-3 UF9497 Duval R/FE - - ? ? F ? UF9498 Duval L/FE - 4.4 no no F 1

173

UF9498 Duval R/FE - 4.45 no no F 1 UF9498 Duval R/TB - 4.4 no no F 1 UF9498 Duval L/MT - 4.05 no yes F 2 UF9498 Duval L/TB - 4.5 no partial F 2 UF9498 Duval R/RA - 3.83 no yes F 2 UF9498 Duval L/RA - 3.8 ? yes F 1-2 UF9500 Duval R/HU - 4.2 yes no ? 2 UF9500 Duval L/HU - 4.2 yes yes ? 3 UF9500 Duval L/MC 171.55 3.85 yes yes ? 3 UF9500 Duval L/MT 200.05 4.3 yes yes ? 3 UF9500 Duval L/RA - 4.05 yes yes ? 3 UF9500 Duval L/TB 221.7 4.85 yes yes ? 3 UF9500 Duval R/RA 170.15 4.1 yes yes ? 3 UF9500 Duval R/TB 221.8 4.9 yes yes ? 3 UF9500 Duval R/FE - 4.83 no parital ? 1-2 UF9500 Duval L/FE 185.55 4.85 partial partial ? 2-3 UF9501 Duval R/RA - 4.52 no yes F 2 UF9501 Duval R/FE 206.5 5.3 yes yes F 3 UF9501 Duval R/HU 158.75 5 yes yes F 3 UF9501 Duval R/TB 234.9 5.3 yes yes F 3 UF9503 Duval L/FE - 5.4 no partial ? 2 UF9503 Duval L/HU - 5.1 yes no ? 2 UF9503 Duval L/TB - 5.48 yes no ? 2 UF9503 Duval R/HU - 5.1 yes no ? 2 UF9503 Duval R/MT - 4.95 no yes ? 2 UF9503 Duval R/RA - 4.75 no yes ? 2 UF9503 Duval L/MT 213.7 5 yes yes ? 3 UF9503 Duval R/FE 210.71 5.5 yes yes ? 3 UF9503 Duval R/MC - 4.5 yes yes ? 3 UF9503 Duval R/TB 247.8 5.55 yes yes ? 3 UF9503 Duval L/RA - 4.9 ? yes ? 1-2 UF9503 Duval L/MC - 4.55 ? yes ? 2-3 UF9505 Marion L/FE 249.58 7.05 yes yes M 3 UF9505 Marion L/HU 188.3 6.95 yes yes M 3 UF9505 Marion L/RA 210.55 6.05 yes yes M 3 UF9505 Marion L/TB 277.65 6.7 yes yes M 3 UF9505 Marion R/FE 248.3 7.05 yes yes M 3 UF9505 Marion R/HU 188.54 6.85 yes yes M 3 UF9505 Marion R/RA 211.25 6.05 yes yes M 3 UF9505 Marion R/TB 279.75 6.5 yes yes M 3 Z4569 Collier L/FE 203.11 5.6 yes yes F 3 Z4569 Collier L/HU 157.28 5.05 yes yes F 3 Z4569 Collier L/MC 185.08 4.4 yes yes F 3 Z4569 Collier L/MT 215.96 5.05 yes yes F 3 Z4569 Collier L/RA 190.93 4.5 yes yes F 3 Z4569 Collier R/FE 202.22 5.55 yes yes F 3 Z4569 Collier R/HU 156.32 5.05 yes yes F 3

174

Z4569 Collier R/MC 185.13 4.4 yes yes F 3 Z4569 Collier R/MT 216.95 5 yes yes F 3 Z4569 Collier R/RA 190.26 4.5 yes yes F 3 Z4569 Collier R/TB 247.53 5.4 yes yes F 3 Z4571 Collier L/FE 237.37 7.1 yes yes M 3 Z4571 Collier L/HU 185.66 6.7 yes yes M 3 Z4571 Collier L/MC 206.75 5.7 yes yes M 3 Z4571 Collier L/MT 241.1 5.9 yes yes M 3 Z4571 Collier L/RA 216.26 6.2 yes yes M 3 Z4571 Collier L/TB 276.39 6.6 yes yes M 3 Z4571 Collier R/FE 237.35 7.1 yes yes M 3 Z4571 Collier R/HU 185.73 6.8 yes yes M 3 Z4571 Collier R/MC 206.8 5.7 yes yes M 3 Z4571 Collier R/MT 241.36 6 yes yes M 3 Z4571 Collier R/RA 216.18 6.12 yes yes M 3 Z4571 Collier R/TB 278.11 6.7 yes yes M 3 Z4572 Collier L/FE 213.22 5.75 yes yes F 3 Z4572 Collier L/HU 157.55 5.3 yes yes F 3 Z4572 Collier L/MC 183.23 4.75 yes yes F 3 Z4572 Collier L/MT 215.4 4.95 yes yes F 3 Z4572 Collier L/RA 188.92 5.05 yes yes F 3 Z4572 Collier L/TB 249.97 5.55 yes yes F 3 Z4572 Collier R/FE 213.29 5.72 yes yes F 3 Z4572 Collier R/HU 158.6 5.3 yes yes F 3 Z4572 Collier R/MC 183.86 4.75 yes yes F 3 Z4572 Collier R/MT 215.57 4.95 yes yes F 3 Z4572 Collier R/RA 188.52 5.05 yes yes F 3 Z4572 Collier R/TB 249.41 5.55 yes yes F 3 Z4574 Collier L/HU - 6.05 yes no F 2 Z4574 Collier R/HU - 6.05 yes no F 2 Z4574 Collier L/FE 228.35 6.08 yes yes F 3 Z4574 Collier L/MC 196.13 52.8 yes yes F 3 Z4574 Collier L/MT 234.27 6 yes yes F 3 Z4574 Collier L/RA 208.91 5.5 yes yes F 3 Z4574 Collier L/TB 270.83 6 yes yes F 3 Z4574 Collier R/FE 229.28 6.12 yes yes F 3 Z4574 Collier R/MC 195.12 5.28 yes yes F 3 Z4574 Collier R/MT 234.68 6.02 yes yes F 3 Z4574 Collier R/RA 208.84 5.5 yes yes F 3 Z4574 Collier R/TB 272.32 6 yes yes F 3 Z4576 Collier L/FE 208.14 5.65 yes yes F 3 Z4576 Collier L/HU 157.26 5.1 yes yes F 3 Z4576 Collier L/MC 183.44 4.3 yes yes F 3 Z4576 Collier L/MT 220.35 4.7 yes yes F 3 Z4576 Collier L/RA 190.37 4.9 yes yes F 3 Z4576 Collier L/TB 248.69 5.6 yes yes F 3 Z4576 Collier R/FE 208.05 5.6 yes yes F 3

175

Z4576 Collier R/HU 157.33 5.2 yes yes F 3 Z4576 Collier R/MC 183.43 4.3 yes yes F 3 Z4576 Collier R/MT 220.86 4.7 yes yes F 3 Z4576 Collier R/RA 190.35 4.9 yes yes F 3 Z4576 Collier R/TB 248.77 5.55 yes yes F 3 Z4577 Collier L/FE - 5.63 no no F 1 Z4577 Collier R/FE - 5.63 no no F 1 Z4577 Collier L/HU - 5.1 yes no F 2 Z4577 Collier L/MC - 4.3 no yes F 2 Z4577 Collier L/MT - 4.75 no yes F 2 Z4577 Collier L/RA - 4.5 no yes F 2 Z4577 Collier L/TB - 5.5 yes no M 2 Z4577 Collier R/HU - 5 yes no F 2 Z4577 Collier R/MC - 4.35 no yes F 2 Z4577 Collier R/MT - 4.8 no yes F 2 Z4577 Collier R/RA - 4.45 no yes F 2 Z4577 Collier R/TB - 5.5 yes no M 2 Z4578 Collier L/FE - 5.45 no no M 1 Z4578 Collier R/FE - 5.4 no no M 1 Z4578 Collier L/HU - 4.95 yes no M 2 Z4578 Collier L/MC - 4.65 no yes M 2 Z4578 Collier L/MT - 5 no yes M 2 Z4578 Collier L/RA - 4.6 no yes M 2 Z4578 Collier L/TB - 5.3 yes no M 2 Z4578 Collier R/HU - 4.95 yes no M 2 Z4578 Collier R/MC - 4.65 no yes M 2 Z4578 Collier R/RA - 4.63 no yes M 2 Z4578 Collier R/TB - 5.25 yes no M 2 Z4578 Collier R/MT - 5.02 yes no M 2-3 Z4703 Collier L/FE 219.48 6.38 yes yes F 3 Z4703 Collier L/HU 169.01 5 yes yes F 3 Z4703 Collier L/MC 199.77 4.85 yes yes F 3 Z4703 Collier L/MT 231.68 5.5 yes yes F 3 Z4703 Collier L/RA 197.46 - yes yes F 3 Z4703 Collier L/TB 259.2 6.2 yes yes F 3 Z4703 Collier R/FE 219.08 6.35 yes yes F 3 Z4703 Collier R/HU 168.66 5.5 yes yes F 3 Z4703 Collier R/MC 199.76 4.85 yes yes F 3 Z4703 Collier R/MT 231.55 5.55 yes yes F 3 Z4703 Collier R/RA 197.55 - yes yes F 3 Z4703 Collier R/TB 260.19 6.15 yes yes F 3 Z4705 Collier L/FE 207.09 5.7 yes yes F 3 Z4705 Collier L/HU 160.91 5.5 yes yes F 3 Z4705 Collier L/MC 193.62 4.65 yes yes F 3 Z4705 Collier L/MT 225.3 5.2 yes yes F 3 Z4705 Collier L/RA 195.69 5.1 yes yes F 3 Z4705 Collier L/TB 259.62 5.85 yes yes F 3

176

Z4705 Collier R/FE 207.37 5.75 yes yes F 3 Z4705 Collier R/HU 160.35 5.45 yes yes F 3 Z4705 Collier R/MC 193.21 4.7 yes yes F 3 Z4705 Collier R/MT 224.79 5.2 yes yes F 3 Z4705 Collier R/RA 195.6 5 yes yes F 3 Z4705 Collier R/TB 259.5 5.75 yes yes F 3 Z5301 Levy L/FE 209.28 6 yes yes ? 3 Z5301 Levy L/HU 160.91 5.5 yes yes ? 3 Z5301 Levy L/MC 195.6 4.85 yes yes ? 3 Z5301 Levy L/MT 227.4 5.4 yes yes ? 3 Z5301 Levy L/RA 188.15 5.05 yes yes ? 3 Z5301 Levy L/TB 249.36 5.85 yes yes ? 3 Z5301 Levy R/FE 208.07 6.05 yes yes ? 3 Z5301 Levy R/HU 160.93 5.5 yes yes ? 3 Z5301 Levy R/MC 195.77 4.8 yes yes ? 3 Z5301 Levy R/MT - - yes yes ? 3 Z5301 Levy R/RA 187.55 5.05 yes yes ? 3 Z5301 Levy R/TB 249.3 5.85 yes yes ? 3

177

Table S3. Locations in Florida used for climate data. Data was collected through www.ncdc.noaa.gov.

Locality Global Historical Climatology Network Latitude/Longitude Tallahassee Regional USW00093805 30.433°N / 84.333°W Jacksonville International USW00013889(GHCN) location Number 30.417°N / 81.65°W LakeAirport City 2 E USC00084731 30.186°N / 82.594°W GainesvilleAirport University of USC00083316 29.65°N / 82.35°W Crescent City USC00081978 29.425°N / 81.5°W OcalaFlorida USC00086414 29.164°N / 82.078°W Orlando Executive Airport USW00012841 28.545°N / 81.333°W Bartow 1 SE USC00080478 27.817°N / 81.817°W Avon Park 2 W USC00080369 27.583°N / 81.5°W Arcadia USC00080228 27.233°N / 81.867°W Everglades USC00082850 25.85°N / 81.383°W Royal Palm Ranger Station USC00087760 25.383°N / 80.6°W

178

Table S4. Monthly climate averages from latitudinal localities based on records dating back to at least 1900 through to 2014 from www.ncdc.noaa.gov.

Avg. Monthly Temp Month Precip. (mm) Max. Temp (⁰C) Min. Temp (⁰C) Avg. Yearly Avg. Yearly Precip. (°C) Temp. (°C) (mm) Tallahassee

1 34.74 17.82 4.44 11.13 19.82 42.78493 2 43.79 19.51 5.62 12.56

3 48.09 22.99 8.66 15.82 Avg. Wet Season Precip. June- Sept. 4 33.01 26.76 11.92 19.34 58.22(mm) 5 33.43 30.46 16.64 23.55 Avg. Dry Season Precip. (Oct. – May) 6 60.14 32.66 20.74 26.70 35.07(mm) 7 67.05 33.05 22.12 27.58 Wet – Dry Season Precip. (mm) 8 59.55 32.89 22.17 27.53 36.60 9 46.13 31.23 20.27 25.75

10 25.33 27.24 13.93 20.59

11 27.75 22.42 8.24 15.33

12 34.41 18.71 5.20 11.95

Lake City

1 28.41 18.98 6.18 12.58 Avg. Yearly Avg. Yearly Precip. 2 33.81 20.35 7.25 13.80 Temp. (°C) (mm) 3 35.96 23.88 10.27 17.08 20.41 36.60 4 25.83 27.03 13.33 20.18 Avg. Wet Season Precip. June- Sept. (mm) 5 29.35 30.50 17.10 23.80 54.55 6 59.73 32.39 20.51 26.45 Avg. Dry Season Precip. (Oct. – May) 7 58.39 32.80 21.73 27.27 27.63(mm) 8 56.71 32.73 21.72 27.22 Wet – Dry Season Precip. (mm) 9 43.36 31.11 20.24 25.67 26.92 10 23.26 27.24 15.18 21.21

11 17.77 22.90 9.97 16.43

12 26.64 19.58 6.96 13.27

179

Jacksonville

1 25.55 18.37 5.79 12.08 Avg. Yearly Avg. Yearly Precip. 2 30.43 19.96 7.27 13.61 Temp. (°C) (mm) 3 30.54 23.17 10.10 16.63 20.43 36.51 4 24.85 26.57 13.40 19.98 Average Wet Season Precip. June- Sept. 5 28.65 29.91 17.57 23.74 57.22(mm) 6 51.82 32.13 21.20 26.67 Average Dry Season Precip. (Oct. – May) 7 53.05 33.25 22.64 27.94 26.16(mm) 8 58.34 32.71 22.63 27.67 Difference Wet – Dry Season Precip. (mm) 9 65.66 30.63 21.13 25.88 31.06 10 31.95 26.77 15.87 21.32

11 15.79 22.75 10.36 16.55

12 21.48 19.28 6.89 13.09

Gainesville, University of Florida

1 25.38 19.90 7.31 13.61 Avg. Yearly Avg. Yearly Precip. 2 27.60 20.52 7.74 14.13 Temp. (°C) (mm) 3 30.23 24.18 10.84 17.51 20.84 36.45 4 24.48 27.33 13.75 20.54 Avg. Wet Season Precip. June- Sept. (mm) 5 27.73 30.67 17.41 24.04 58.26 6 62.34 32.26 20.73 26.49 Avg. Dry Season Precip. (Oct. – May) 7 61.58 32.56 21.83 27.20 25.55(mm) 8 60.88 32.62 21.97 27.29 Wet – Dry Season Precip. (mm) 9 48.26 31.15 20.78 25.96 32.71 10 26.96 27.83 16.16 22.00

11 17.26 23.63 10.85 17.24

12 24.76 20.32 7.79 14.06

Crescent City

1 19.62 20.84 8.61 14.73 Avg. Yearly Avg. Yearly Precip. 2 27.17 22.10 9.75 15.93 Temp. (°C) (mm) 3 29.78 24.44 11.96 18.20 21.71 35.51 4 24.75 27.17 15.08 21.12 Avg. Wet Season Precip. June- Sept. (mm) 5 32.33 30.20 18.74 24.47 55.47 6 58.91 32.11 21.73 26.92 Avg. Dry Season Precip. (Oct. – May) 7 55.36 32.79 22.71 27.75 25.53(mm)

180

8 54.97 32.75 22.93 27.84 Wet – Dry Season Precip. (mm) 9 52.62 31.19 22.00 26.59 29.94 10 32.02 28.00 17.94 22.97

11 18.03 24.16 12.86 18.51

12 20.54 21.37 9.63 15.50

Ocala

1 19.62 21.33 7.68 14.50 Avg. Yearly Avg. Yearly Precip. 2 27.17 22.83 8.82 15.83 Temp. (°C) (mm) 3 29.78 25.72 11.41 18.57 21.71 36.65 4 24.75 28.67 14.02 21.34 Avg. Wet Season Precip. June- Sept. (mm) 5 32.33 31.72 17.74 24.73 58.55 6 58.91 33.01 21.02 27.02 Avg. Dry Season Precip. (Oct. – May) 7 55.36 33.40 22.04 27.72 25.69(mm) 8 54.97 33.28 22.09 27.68 Wet – Dry Season Precip. (mm) 9 52.62 32.08 20.92 26.50 32.86 10 32.02 28.89 16.62 22.76

11 18.03 25.07 11.84 18.45

12 20.54 22.02 8.64 15.33

Orlando

1 18.14 22.27 10.04 16.15 Avg. Yearly Avg. Yearly Precip. 2 22.45 23.16 10.66 16.91 Temp. (°C) (mm) 3 24.60 25.92 13.05 19.48 22.45 35.38 4 21.59 28.44 15.69 22.06 Avg. Wet Season Precip. June- Sept. (mm) 5 31.87 31.22 18.91 25.07 60.72 6 61.88 32.60 21.68 27.14 Avg. Dry Season Precip. (Oct. – May) 7 64.26 33.20 22.65 27.92 22.71(mm) 8 57.82 33.24 22.84 28.04 Wet – Dry Season Precip. (mm) 9 58.94 31.83 22.06 26.94 38.01 10 32.21 28.85 18.53 23.69

11 13.93 25.26 13.58 19.42

12 16.90 22.57 10.52 16.54

181

Bartow

1 20.24 22.76 9.93 16.34 Avg. Yearly Avg. Yearly Precip. 2 24.60 23.96 10.70 17.33 Temp. (°C) (mm) 3 26.84 26.58 13.01 19.80 22.60 37.44 4 22.95 29.03 15.44 22.24 Avg. Wet Season Precip. June- Sept. (mm) 5 35.37 31.65 18.65 25.15 64.89 6 67.98 32.88 21.39 27.14 Avg. Dry Season Precip. (Oct. – May) 7 67.81 33.32 22.29 27.80 23.72(mm) 8 63.40 33.47 22.45 27.96 Wet – Dry Season Precip. (mm) 9 60.37 32.21 21.80 27.01 41.18 10 26.09 29.48 18.23 23.85

11 15.69 25.84 13.52 19.68

12 17.97 23.22 10.68 16.95

Avon Park

1 17.98 23.18 10.15 16.66 Avg. Yearly Avg. Yearly Precip. 2 22.64 24.36 10.92 17.64 Temp. (°C) (mm) 3 22.52 26.76 13.10 19.93 22.76 36.66 4 21.75 29.21 15.53 22.37 Avg. Wet Season Precip. June- Sept. (mm) 5 34.37 31.68 18.55 25.11 65.06 6 73.02 32.79 21.24 27.02 Avg. Dry Season Precip. (Oct. – May) 7 66.03 33.25 22.17 27.71 22.46(mm) 8 61.40 33.33 22.40 27.86 Wet – Dry Season Precip. (mm) 9 59.80 32.18 21.86 27.02 42.60 10 29.57 29.55 18.60 24.08

11 15.56 26.24 14.24 20.24

12 15.31 23.82 11.21 17.52

Arcadia

1 16.45 23.58 9.72 16.65 Avg. Yearly Avg. Yearly Precip. 2 21.97 24.62 10.33 17.47 Temp. (°C) (mm) 3 23.15 26.93 12.38 19.66 22.51 36.77 4 19.97 29.43 14.62 22.03 Avg. Wet Season Precip. June- Sept. (mm) 5 32.49 31.96 17.56 24.76 67.37 6 73.56 32.87 20.63 26.75 Avg. Dry Season Precip. (Oct. – May) 7 66.67 33.22 21.68 27.45 21.46(mm)

182

8 66.78 33.21 21.95 27.58 Wet – Dry Season Precip. (mm) 9 62.48 32.18 21.42 26.80 45.91 10 27.06 29.64 17.96 23.80

11 14.57 26.28 13.47 19.87

12 16.02 24.01 10.66 17.34

Everglades

1 12.48 24.27 12.12 18.20 Avg. Yearly Avg. Yearly Precip. 2 15.59 24.95 13.01 18.98 Temp. (°C) (mm) 3 15.20 26.63 14.99 20.81 23.60 37.92 4 19.61 28.59 17.08 22.83 Avg. Wet Season Precip. June- Sept. (mm) 5 37.10 30.50 19.84 25.17 73.57 6 83.25 31.63 22.39 27.01 Avg. Dry Season Precip. (Oct. – May) 7 68.88 32.41 23.10 27.76 20.10(mm) 8 65.74 32.61 23.52 28.06 Wet – Dry Season Precip. (mm) 9 76.42 32.12 23.18 27.65 53.48 10 35.27 30.31 20.50 25.41

11 12.98 27.56 16.51 22.03

12 12.53 25.19 13.44 19.32

Royal Park Ranger Station

1 20.25 22.76 9.93 16.35 Avg. Yearly Avg. Yearly Precip. 2 24.60 23.96 10.70 17.33 Temp. (°C) (mm) 3 26.84 26.58 13.01 19.80 22.60 37.45 4 22.95 29.03 15.44 22.24 Avg. Wet Season Precip. June- Sept. (mm) 5 35.37 31.65 18.65 25.15 64.90 6 67.98 32.88 21.39 27.14 Avg. Dry Season Precip. (Oct. – May) 7 67.83 33.31 22.29 27.80 23.72(mm) 8 63.42 33.46 22.45 27.95 Wet – Dry Season Precip. (mm) 9 60.38 32.21 21.80 27.01 41.18 10 26.10 29.48 18.23 23.85

11 15.70 25.84 13.52 19.68

12 17.98 23.22 10.68 16.95

183

Table S5. Statistical comparison between left and right elements from similar geographic, and geochronologic times. Statistical tests were only performed when both left and right elements were present and n >1.

Blancan Northern Florida Length Blancan Northern Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora 3 2 0.800 3 6 0.606 Tibiae - - - 1 - - Metatarsals 3 - - 6 - Humeri 1 3 - 2 4 0.355 Radii - 2 - - 6 - Metacarpals - 4 - - 4 - Irvingtonian Northern Florida Length Irvingtonian Northern Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora - - - - 1 - Tibiae - - - 1 - - Metatarsals 1 - - 3 - - Humeri - - - - 1 - Rancholabrean Northern Florida Length Rancholabrean Northern Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora 6 4 0.171 7 4 0.104 Tibiae 4 4 0.564 5 5 0.690 Metatarsals 2 2 1.000 3 2 0.564 Humeri 4 2 0.643 4 4 0.770 Radii 8 4 0.062 8 5 0.558 Metacarpals 3 2 1.000 3 3 0.178 Modern Northern Florida Length Modern Northern Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora 8 10 Mo 0.625 9 11 0.703 Tibiae 12 13 0.828 11 14 0.783 Metatarsals 13 8 0.744 8 13 0.799 Humeri 10 13 0.828 11 12 0.477 Radii 8 10 0.859 9 11 0.848 Metacarpals 9 10 0.806 9 12 0.943

Irvingtonian South Florida Length Irvingtonian South Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora ------Tibiae - 1 - 1 - Metatarsals 1 1 - 2 1 - Humeri 1 3 - 2 3 0.083 Radii - 1 - - 1 - Metacarpals 1 1 - 1 1 -

184

Rancholabrean South Florida Length Rancholabrean South Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora - - - - 1 - Tibiae - 1 - - 2 - Metatarsals 3 - - 4 - - Humeri - 2 - - 2 - Radii - 1 - - 1 - Metacarpals 1 1 - 1 1 - Nichol’s Hammock South Florida Length Nichol’s Hammock South Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora 3 - - 3 - - Humeri 4 - - 4 - - Radii 3 - - 5 - - Metacarpals - 2 - - 2 - Modern South Florida Length Modern South Florida MDC Element Left Right Mann-Whitney U p-value Left Right Mann-Whitney U p-value Femora 11 8 0.869 8 8 0.958 Tibiae 7 8 1.000 7 8 0.727 Metatarsals 7 8 0.817 8 8 0.958 Humeri 12 8 0.758 8 8 0.371 Radii 7 7 0.565 7 7 0.747 Metacarpals 8 10 0.790 8 8 0.958

185

S6. All histologically sampled specimens from O. virginianus used in this study. Abbreviations: Elmt. = Element; Loc. = Location; ING =

Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s Hammock; OS = Ontogenetic Stage; MDC = Mid-diaphysis circumference; Partial lengths (PL) were taken on most specimens due to partial/broken elements, * indicated articular length (complete specimen); DD = thin section distance from distal epiphysis. DP = thin section distance from the proximal epiphysis; TOR = thickness of removed section; APD = Anterior – Posterior Diameter, LMD = Lateral – Medial Diameter; MAPD = Medullary APD; MLMD = Medullary LMD. L and

R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius. J = Juvenile; SA = Sub Adult; A = Adult; J. S-A; = Juvenile to

Sub-Adult; S-A-A = Sub-Adult to Adult. MDC, PL, DD, DP and TOSR are in cm. APD, LMD, MAPD, MLMD are all in mm.

Thin section dimensions (mm)

Age Specimen Elmt Loc. OS LAGs EFS MDC PL DD DP

(mths) APD

LMD

MAPD

MLMD TOR UF45040 LFE ING SA 8-26 3 No 7.5 14.1 10.8 - 2 24.86 20.68 17.09 14.46 UF45044 LFE ING SA 8-26 7 No 6.87 14.39 10.3 - 1.8 21.61 18.71 14.68 12.65 UF300294 LFE ING SA 8-26 2 ? 6.3 21.3 10.5 - 2.9 18.18 18.58 10.61 11.48 UF300295 LEF ING J >20 0 No 5.5 15.3 7.2 - 2.4 15.1 16.38 9.54 10.77 UF300296 LFE ING J >20 1 ? 7 16.7 - 9 1.8 17.91 18.89 12.41 12.41 UF300297 LEF ING J >20 0 No 6.2 20.6 10.2 - 2.6 19.56 19.35 11.51 11.9 UF300298 LEF ING SA 8-26 1 ? 6.5 21.1 12.6 - 1.5 19.46 17.71 11.79 11.4 UF86378 LFE LSP SA 8-26 5 No 6.5 15.9 12.9 - 3.7 20.1 18.19 13.1 12.65 UF239925 RFE LSP A >26 3 Partial? 6.9 13.1 - 10 5.3 21.08 20.64 14.85 12.84 UF300286 RFE COLA SA 8-26 ? ? 6.2 9.7 7.9 - 3.5 20.06 17.88 13.34 11.97 UF300287 RFE COLA J <20 ? ? 5.9 12.3 - 11.4 1.8 17.79 17.72 11.76 11.72 UF300288 RFE COL2A ? ? 2? ? 5.9 15.7 7.6 - 3.5 17.32 17.47 11.72 12.5 UF57620 LFE NH2A A >26 5 Yes 6.22 20.09* 10.4 - 1.6 18.74 18.69 12.51 12.9 2A

186

UF57621 LFE NH A >26 4 Partial 5.6 21.31* 10.8 - 2.6 16.07 18.21 10.05 11.88 UF57622 LFE NH A >26 2 No 5.4 19.4* 8.9 - 2.9 17.16 16.5 11.62 10.71 UF45069 RTB ING SA >20 3 No 8.2 17.7 14.9 - 2.3 21.15 26.41 12.87 17.58 UF45075 RTB ING SA >20 1? No 7.2 15.7 15.2 - 1.9 19.9 23.56 11.22 11.74 UF45087 RTB ING A >26 1? ? 7.5 20.3 - 14.4 2.7 19.75 24.37 10.29 15.02 UF276272 RTB ING SA 20-26 ? ? 6.7 26.8 11.9 3.7 18.34 21.08 10.51 12.64 UF302003 LTB ING J <14 ? ? 5.7 20.7 9.7 - 1.6 14.78 18.01 7.46 11.07 UF85251 LTB LSP SA >20 ? ? 5.8 10.7 6.2 - 2.2 14.79 18.55 9.87 12.25 UF226999 LTB LSP SA >20 3 No 6.4 22 13.7 - 2.8 18.66 20.06 9.36 11.88 UF239928 LTB LSPA SA >20 1? No 5.8 11.5 3.2 - 1.6 15.46 19.75 9.19 11.92 UF276488 LTB COLA J-SA 14-20 0 No 5.7 16.9 10.4 - 2.7 16.08 18.71 6.77 10.19 UF276489 LTB COLA SA >20 7 Partial? 6.6 17.4 11.1 - 2.8 18.89 21.56 9.76 11.47 UF276490 LTB COL2A SA-A >14 4 ? 5.7 9.3 8.4 - 2.2 16.14 18.92 8.5 11.27 UF57185 LTB NH2A SA 20-26 5-6? Yes 5.6 12.7 - 9.1 3.9 17.06 17.74 10 10.55 UF43597 LHU ING2A SA 12-26 ? No 6.05 15.4 8.4 - 1.7 20.96 16.85 15.75 11.77 UF43600 LHU ING SA 12-26 ? ? 5.4 8.7 8.1 - 2.7 18.64 14.29 13.4 9.57 UF300299 LHU ING ? ? 5 Partial? 6.9 13.9 7.5 - 1.6 24.75 18.98 16.61 10.81 UF300300 LHU ING SA 12-26 7 ? 6.1 13.6 8.7 - 1.4 21.24 17.25 14.83 11.11 UF302000 LHU ING J <8 ? No 5.6 15.5 7 - 1.8 18.6 15.38 11.94 8.53 UF302001, LHU ING A >38 ? No 7.8 19.3* 9.4 - 3.1 25.28 21.6 15.6 11.94 302002 UF65962 LHU LSP SA 12-26 4 No 6.3 14.5 8.2 - 2.3 22.11 17.83 15.37 11.34 UF81121 LHU LSP ? ? 2 No 5.7 10 5.5 - 3.1 19.03 15.92 12.34 9.19 UF87904 LHU LSPA SA 12-26 4 No 6.2 12.2 7.8 - 3.0 21.4 17.9 13.37 10.1 UF300283 LHU COLA SA 12-26 ? ? 5.5 10.1 7.6 - 3.7 19.4 14.4 13.94 9.45 UF300284 RHU COLA SA 12-26 ? Yes? 5.3 13.1 7.4 - 3.5 19.88 15.64 14.3 10.66 UF300285 LHU COL2A ? ? 0? ? 5 6.3 5.2 - 2.5 15.72 14.09 10.73 8.98 UF57164 LHU NH2A A <38 7 Yes 5.8 16.8* 8.7 - 1.4 20.92 16.46 15.32 10.48 UF57165 LHU NH2A A <38 7 Partial? 5.3 15.0* 7.4 - 2.7 18.59 15.06 14.32 10.44 UF57166 LHU NH A <38 5 Yes 5.12 16.3* 8.1 - 1.6 16.98 15.44 11.68 9.17

187

UF300289 LHU NH J <8 0 No 2.9 6.4 3.4 - 1.1 8.54 7.62 6.14 5.41 UF300290 LHU NH J <8 0 No 3.6 9.5 4.5 - 1.0 11.09 10.74 8.57 7.61 UF45007 LRA ING J-SA <26 ? No 6.2 19.5 7.5 - 2.2 13.86 23.23 6.75 13.04 UF45008 LRA ING J- SA <26 0 No 5.7 15.7 8 - 1.3 11.83 19.34 6.93 12.05 UF276267 LRA ING A >26 0 ? 6.2 12.7 10.9 - 1.7 13.76 23.23 8.08 13.65 UF300291 LRA ING J <5 0 No 5.1 15.6 7.7 - 1.9 10.94 18.55 6.12 11.43 UF300292 LRA ING J-SA <26 ? No 5.2 15 7 - 2.2 10.49 19.91 5.23 10.4 UF300293 LRA ING J-SA <26 ? No 5.4 15.7 8 - 1.2 10.72 19.66 5.53 10.55 UF57178 LRA NH A >26 ? Partial? 4.7 17.5* 9 - 1.2 10.76 17.2 5.65 9.94 UF57179 LRA NH A >26 ? Yes 5.3 18.7* 9.2 - 3.2 11.26 18.26 5.68 10.35 UF57180, LRA A ? 57182 NH >26 Partial? 4.6 18.7* 9.7 - 3.0 10.95 16.3 6.12 9.27 UF57181 LRA NH A >26 ? Yes 4.5 9.3 7.3 - 3.7 10.07 16.02 5.77 9.4

188

Table S7. Individual APD measurements for individual LAGs from femora, tibiae, and humeri. * indicate where measurement was taken from a juvenile < 1 year old (no LAG present). Location = Loc.; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s

Hammock. Elmt = Element; L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius.

APD of LAGs

Specimen Loc. Elmt LAG 0.5* LAG 1 LAG 2 LAG 3 LAG 4 LAG 5 LAG 6 LAG 7 LAG 8

UF 45040 ING LFE - - 22.78 23.44 23.88 - - - - UF 45044 ING LFE - - 19.19 19.82 20.17 20.55 20.96 21.26 - UF 300294 ING LFE - 14.60 16.51 ------UF 300295 ING LFE 15.58 ------UF 300296 ING LFE - 18.01 ------UF 300297 ING LFE 19.3 ------UF 300298 ING LFE - 17.75 ------UF 86378 LSP LFE - 17.01 18.63 19.16 19.47 19.73 - - - UF 239925 LSP RFE - 18.72 20.07 20.86 - - - - - UF 300287 COL RFE 17.6 ------UF 57621 NH LFE - 13.33 14.65 15.40 15.91 - - - - UF 57620 NH LFE - - 15.77 16.57 17.30 17.78 18.28 - - UF 57622 NH LFE - 13.34 15.91 ------UF45069 ING RTB - - 18.64 19.28 19.95 - - - - UF226999 LSP LTB - 15.33 16.82 17.65 - - - - - UF276488 COL LTB 16.06 ------UF276489 COL LTB - 14.82 16.08 16.69 17.37 18.01 18.59 - - UF276490 COL LTB - 12.99 14.14 14.46 - 16.82 - - - UF57185 NH LTB - - 14.6 15.34 15.79 16.45 16.81 - - UF300299 ING LHU - - 21.98 23.44 24.01 - - - - UF300300 ING LHU - - 17.99 18.67 19.41 19.88 - - 20.56

189

UF65962 LSP LHU - 18.87 21.26 21.54 22.03 - - - - UF81121 LSP LHU - 16.68 18.52 ------UF87904 LSP LHU - 17.28 19.19 20.11 20.65 - - - - UF57164 NH LHU - - 17.51 18.68 19.34 20.23 20.75 - - UF57165 NH LHU - - 16.08 16.74 17.03 17.52 17.86 18.11 18.25 UF57166 NH LHU - - 14.51 15.01 15.63 16.38 16.80

UF300289 NH LHU 8.61 ------UF300290 NH LHU 10.92 ------

190

Table S8. Maximum APD growth rates (MAPDGR) (260 day growth period) based on APD measurements for each growth period. GZ1 cannot be accurately calculated due to unknown degree of resorption.Abbreviations: L and R = Left or Right element; FE = Femur; TB = Tibia; HU =

Humerus; RA = Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s Hammock

MAPDGR per growth zone Specimen Locality Element GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 UF 45040 ING LFE - 2.54 1.69 - - - - UF 45044 ING LFE - 2.42 1.35 1.46 1.58 1.15 - UF 300294 ING LFE 7.35 ------UF 86378 LSP LFE 6.23 2.04 1.19 1 - - - UF 239925 LSP RFE 5.19 3.04 - - - - - UF 57621 NH LFE 5.08 2.88 1.96 - - - - UF 57620 NH LFE - 3.08 2.81 1.85 1.92 - - UF 57622 NH LFE 9.88 ------UF45069 ING RTB - 2.48 2.57 - - - - UF226999 LSP LTB 5.72 3.21 - - - - - UF276489 COL LTB 4.83 2.36 2.61 2.46 2.22 - - UF276490 COL LTB 4.43 1.25 - - - - - UF57185 NH LTB - 2.84 1.73 2.54 1.38 - - UF300299 ING LHU - 5.62 2.19 - - - - UF300300 ING LHU - 2.62 2.85 1.81 - - - UF65962 LSP LHU 9.19 1.07 1.89 - - - - UF81121 LSP LHU 7.08 ------UF87904 LSP LHU 7.35 3.54 2.08 - - - - UF57164 NH LHU - 4.5 2.54 3.42 2 - - UF57165 NH LHU - 2.54 1.12 1.88 1.31 0.96 0.54 UF57166 NH LHU - 1.91 2.38 2.88 1.62 - -

191

Table S9. Minimum APD growth rates (mAPDGR) (365 day growth period) for each growth period. GZ1 cannot be accurately calculated due to unknown degree of resorption. Abbreviations: L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius; ING =

Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s Hammock

mAPDGR per growth zone Specimen Locality Element GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 UF 45040 ING LFE - 1.81 1.21 - - - - UF 45044 ING LFE - 1.73 0.96 1.04 1.12 0.82 - UF 300294 ING LFE 5.23 ------UF 86378 LSP LFE 4.44 1.45 0.85 0.71 - - - UF 239925 LSP RFE 3.7 2.16 - - - - - UF 57621 NH LFE 3.62 2.05 1.4 - - - - UF 57620 NH LFE - 2.19 2 1.32 1.37 - - UF 57622 NH LFE 7.04 ------UF45069 ING RTB - 1.76 1.83 - - - - UF226999 LSP LTB 4.08 2.28 - - - - - UF276489 COL LTB 3.44 1.68 1.86 1.75 1.58 - - UF276490 COL LTB 3.16 0.89 - - - - - UF57185 NH LTB - 2.02 1.23 1.81 0.99 - - UF300299 ING LHU - 4 1.56 - - - - UF300300 ING LHU - 1.86 2.03 1.29 - - - UF65962 LSP LHU 6.55 0.76 1.35 - - - - UF81121 LSP LHU 5.04 ------UF87904 LSP LHU 5.23 2.52 1.48 - - - - UF57164 NH LHU - 3.21 1.81 2.44 1.42 - - UF57165 NH LHU - 1.81 0.79 1.34 0.93 0.68 0.38 UF57166 NH LHU - 1.36 1.7 2.05 1.15 - -

192

Table S10. Anterior LAG measurements and growth rate calculations. Italicized values are minimum values. ACT = Anterior Cortical

Thickenss; d 1st-LL = distance 1st to the last LAG. (*ACL for humeri was based on posterior cortical thickness) ** distance from 2nd LAG to last

LAG measured. Abbreviations: L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius.

Specimen Number Locality Element ACT (µm) d 1st-LL (µm)

GZ1 GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8

(um)

(µm) (µm) (µm) (µm) (µm) (µm) (µm)

UF45040 ING LFE 3663 923** resorbed 2740 556 368 - - - - UF45044 ING LFE 3517 1227** resorbed 2215 187 211 219 168 289 146 UF300294 ING LFE 3924 1281 1900 1281 ------UF300295 ING LEF 2989 - 2989 ------UF300296 ING LFE 2773 - 2164 ------UF300297 ING LEF 3679 - 3679 ------UF300298 ING LEF 4574 - 1528 ------UF86378 LSP LFE 3553 860 2693 626 127 105 - - - - UF239925 LSP RFE 3765 1054 2498 454 604 - - - - - UF57620 NH LFE 3105 1488** resorbed 1337 485 494 298 210 - - UF57621 NH LFE 3034 1062 1972 683 276 103 - - - - UF57622 NH LFE 3185 1373 1472 1373 ------UF45069 ING RTB 3465 1062** resorbed 2116 540 514 - - - - UF226999#2 LSP LTB 4982 2233 2343 1433 807 - - - - UF276488 COL LTB 5213 - 5213 ------UF276489 COL LTB 6192 3588 2522 1155 362 734 267 326 716 - UF276490 COL LTB 4021 1631 2109 368 606 386 271 - - - UF57185 NH LTB 3770 2072** Resorbed 1438 670 306 498 153 448 - UF300299 ING LHU 4046* 4123** Resorbed 2128 518 221 - - - - UF300300 ING LHU 3124* 2028 915 655 533 328 209 206 74 - UF65962 LSP LHU 2690* ** Resorbed 1715 239 447 - - - -

193

UF81121 LSP LHU 3043* 1510 1426 1510 ------UF87904 LSP LHU 4773* 2914 2028 1686 844 379 - - - - UF57164 NH LHU 2841* ** resorbed 950 734 262 329 357 - - UF57165 NH LHU 1852* 1300** resorbed 428 412 480 179 92 84 39 UF57166 NH LHU 2455* 1858** resorbed 362 476 441 675 265 - - UF300289 NH LHU 1281* - 1281 ------UF300290 NH LHU 1351* - 1351 ------

194

Table S11. Anterior growth rates from methodologies of Sander and Tückmantel, (2003) based on a 260 day growth period. See text for formulas for (MGRGP, mGRGP, MANTGRn). Bold values are average GRs, Italicized are when average GR is based on a single specimen. * = minimum growth rate values due to inner cortical resorption. All rates are in (µm/day). Abbreviations: L and R = Left or Right element; FE = Femur; TB =

Tibia; HU = Humerus; RA = Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s Hammock

MANTGR n

Specimen Mean Rate of

Locality Elmt. MGRGP mGRGP

GZ3 GZ4 GZ5 GZ6 GZ7 GZ8

Number Growth GZ2 GZ1*

UF45040 ING LFE 4.7 1.18 2.94 - 10.5* 2.14 1.42 - - - - UF45044 ING LFE 1.93 0.67 1.3 - 8.52* 0.72 0.81 0.84 0.65 1.11 0.56 UF300294 ING LFE 7.55 2.46 5 7.31* 4.93 ------UF300295 ING LEF - - - 11.5* ------UF300296 ING LFE 10.7 - - 8.32* ------UF300297 ING LEF - - - 14.2* ------UF300298 ING LEF 17.59 - - 5.88* ------ING Average - FE 8.49 1.44 3.08 9.43* 8.00 1.42 1.11 0.84 0.65 1.11 0.56

UF86378 LSP LFE 3.42 0.83 2.12 10.4* 2.41 0.49 0.4 - - - - UF239925 LSP RFE 4.83 1.35 3.09 9.61* 1.75 2.32 - - - -

LSP Average FE 4.12 1.09 2.61 9.98* 2.08 1.41 0.4 - - - -

UF57620 NH LFE 2.39 1.14 1.77 - 5.14* 1.87 1.9 1.15 0.81 - - UF57621 NH LFE 2.92 1.02 1.97 7.58* 2.63 1.06 0.4 - - - - UF57622 NH LFE 6.13 2.64 4.38 5.66* 5.28 ------

195

NH Average - FE 3.81 1.6 2.71 6.62* 3.95 1.06 1.15 1.15 0.81 - -

UF45069 ING RTB 4.44 1.36 2.9 - 8.14 2.08 1.98 - - - - UF226999#2 LSP LTB 6.39 2.86 4.63 9.01* 5.51 3.1 - - - - - UF276488 COL LTB - - - 20.05 *------UF276489 COL LTB 3.4 1.97 2.69 9.7* 4.44 1.39 2.82 1.03 1.25 2.75 - UF276490 COL LTB 3.09 1.25 2.17 8.11* 1.42 2.33 1.48 1.04 - - - COL Average - TB 3.25 1.61 2.43 12.62* 2.93 1.86 2.15 1.03 1.25 2.75 -

UF57185 NH LTB 2.42 1.33 1.87 - 5.53 2.58 1.18 1.92 0.59 1.72 -

UF300299 ING LHU 3.89 3.96 3.93 - 8.18* 2.000 0.9 - - - - UF300300 ING LHU 1.72 1.11 1.42 3.52 2.51 2.05 1.26 0.80 0.79 0.28 - ING Average * 2.80 2.54 2.67 3.52 5.35 2.02 1.06 0.80 0.79 0.28 - HU

UF65962 LSP LHU 2.07 1.85 1.96 - 6.60 0.92 1.72 - - - - UF81121 LSP LHU 5.85 2.90 4.38 5.48 5.80 ------UF87904 LSP LHU 4.59 2.80 3.70 7.8 6.48* 3.25 1.46 - - - - LSP Average HU 4.17 2.85 3.04 6.64 6.30 2.08 1.59 - - - -

UF57164 NH LHU 1.82 1.08 1.45 - 3.65 2.82 1.01 1.27 1.37 - - UF57165 NH LHU 1.02 0.71 0.87 - 1.65 1.58 1.85 0.69 0.35 0.32 0.15 UF57166 NH LHU 1.89 1.43 1.66 - 1.39* 1.83 1.70 2.60 1.02 - - UF300289 NH LHU - - - 4.93 *------UF300290 NH LHU - - - 5.20 *------NH Average HU 1.58 1.07 1.15 5.06 2.23 2.10 1.52 1.52 0.92 0.32 0.15

196

Table S12. Anterior growth rates from methodologies of Sander and Tückmantel, (2003) based on a 365 day growth period. See text for formulas for (MGRST, mGRST, mANTGRn). Italicized = Average GR based on a single specimen, *= minimum growth rate values due to inner cortical resorption.. Bold values are average GRs. Abbreviations: L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA =

Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman 2A, NH = Nichol’s Hammock. All rates are in (µm/day).

mANTGRn Specimen Mean Rate of Locality Element MGRST mGRST GZ1* GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 GZ8 Number Growth UF45040 ING LFE 3.35 0.84 2.09 - 7.51* 1.52 1.01 UF45044 ING LFE 1.38 0.48 0.93 - 6.07* 0.51 0.58 0.6 0.46 0.79 0.4 UF300294 ING LFE 5.38 1.75 3.57 5.21* 3.51 ------UF300295 ING LEF - - - 8.19* ------UF300296 ING LFE 7.6 - - 5.93* ------UF300297 ING LEF - - - 10.1* ------UF300298 ING LEF 12.53 - - 4.19* ------

ING Average - FE 6.05 1.03 2.2 6.72* 5.69 1.02 0.79 0.6 0.46 0.79 0.4

UF86378 LSP LFE 2.43 0.59 1.51 7.38* 1.72 0.35 0.29 - - - - UF239925 LSP RFE 3.44 0.96 2.2 6.84* 1.24 1.65 - - - -

LSP Average - FE 2.94 0.78 1.86 7.11* 1.48 1 0.29 - - - -

UF57620 NH LFE 1.7 0.82 1.26 - 3.66* 1.33 1.35 0.82 0.58 - - UF57621 NH LFE 2.08 0.73 1.4 5.4* 1.87 0.76 0.28 - - - - UF57622 NH LFE 4.36 1.88 3.12 4.03* 3.76 ------

NH Average - FE 2.71 1.14 1.93 4.72* 3.1 1.04 0.82 0.82 0.58 - -

197

UF45069 ING RTB 3.16 0.97 2.07 - 5.8* 1.48 1.41 - - - - UF226999#2 LSP LTB 4.55 2.04 3.29 6.42* 3.93 2.21 - - - - - UF276488 COL LTB - - - 14.28* ------UF276489 COL LTB 2.42 1.4 1.91 6.91 3.16 0.99 2.01 0.73 0.89 1.96 - UF276490 COL LTB 2.2 0.89 1.55 5.78 1.01 1.66 1.06 0.74 - - - COL Average - TB 2.31 1.15 1.73 8.99 2.09 1.33 1.53 0.74 0.89 1.96 -

UF57185 NH LTB 1.72 0.95 1.33 - 3.94* 1.84 0.84 1.36 0.42 1.23 -

UF300299 ING LHU 2.77 2.82 2.80 - 5.83* 1.42 0.61 - - - - UF300300 ING LHU 1.22 0.79 1.01 2.51 1.79 1.46 0.90 0.57 0.56 0.20 -

ING Average HU 2.00 1.81 1.90 2.51 3.81 1.44 0.75 0.57 0.56 0.20

UF65962 LSP LHU 1.47 1.32 1.40 - 4.70* 0.65 1.22 - - -

UF81121 LSP LHU 4.17 2.07 3.12 3.91 4.14 ------UF87904 LSP LHU 3.27 2.00 2.63 5.56 4.62 2.31 1.04 - - - -

LSP Average HU 2.97 2.03 2.38 4.73 4.48 1.48 1.13

198

UF57164 NH LHU 1.29 0.77 1.03 - 2.60* 2.01 0.72 0.90 0.98 - - UF57165 NH LHU 0.72 0.51 0.62 - 1.17* 1.13 1.31 0.49 0.25 0.23 0.11 UF57166 NH LHU 1.35 1.02 1.18 - 1.00* 1.30 1.21 1.85 0.73 - - UF300289 NH LHU - - - 3.51 ------UF300290 NH LHU - - - 3.70 ------

NH Average HU 1.12 0.76 0.94 3.61 1.59 1.48 1.08 1.08 0.65 .23 .11

199

Table S13. Individual Single Layer osteocyte lacunae (OL) count and calculated density for each growth zone. Abbreviations: El. = Element; L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL =

Coleman 2A, NH = Nichol’s Hammock Loc. Locality. “ = same value as above; italicized values are averages per individual and whole locality.

OL Density per GZ (lacunae/mm2)

OL OL OL OL OL OL OL Total Specimen # El. Loc. GZ GZ GZ GZ GZ GZ GZ Area GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 1 2 3 4 5 6 7 (mm2) UF45040 LFE ING - 13 14 14 - - - 0.01 - 1300 1400 1400 - - - UF45040 LFE ING - 11 10 17 - - - “ - 1100 1000 1700 - - - UF45040 LFE ING - 11 12 15 - - - “ - 1100 1200 1500 - - - AVG OD 1166.7 1200 1533.3 - - -

UF45044 LFE ING - 14 18 18 14 17 13 “ - 1400 1800 1800 1400 1700 1300 UF45044 LFE ING - 15 15 14 11 13 10 “ - 1500 1500 1400 1100 1300 1000 UF45044 LFE ING - 14 15 18 11 20 17 “ - 1400 1500 1800 1100 2000 1700 AVG OD 1433.3 1600 1666.7 1200 1666.7 1333.3

UF300294 LFE ING - 11 - - - - - “ - 1100 - - - - - UF300294 LFE ING - 18 - - - - - “ - 1800 - - - - - UF300294 LFE ING - 18 - - - - - “ - 1800 - - - - - AVG OD 1566.7

UF300296 LFE ING 13 ------“ 1300 ------UF300296 LFE ING 12 ------“ 1200 ------UF300296 LFE ING 14 ------” 1400 ------AVG OD 1300

UF300297 RFE ING 19 ------“ 1900 ------UF300297 RFE ING 21 ------“ 2100 ------UF300297 RFE ING 19 ------“ 1900 ------AVG OD 1966.7

UF300298 LFE ING 14 ------“ 1400 ------

200

UF300298 LFE ING 21 ------“ 2100 ------UF300298 LFE ING 17 ------“ 1700 ------AVG OD 1733.3

AVG OD FE 1666.7 1388.9 1400 1600 1200 1666.7 1333.3 ING UF86378 RFE LSP 18 12 - - - - - “ 1800 1200 - - - - - UF86378 RFE LSP 18 17 - - - - - “ 1800 1700 - - - - - 1A UF86378 RFE LSP 15 16 - - - - - “ 1500 1600 - - - - - 1A AVG OD 1700 1500 1A UF239925 RFE LSP 15 6 12 - - - - “ 1500 600 1200 - - - - UF239925 RFE LSP 14 10 14 - - - - “ 1400 1000 1400 - - - - 1A UF239925 RFE LSP 15 5 14 - - - - “ 1500 500 1400 - - - - 1A UF239925 RFE LSP - 10 - - - - - “ - 1000 - - - - - 1A AVG OD 1466.7 775 1333.3 1A AVG OD FE 1583.3 1137.5 1333.3 LSP UF57620 LFE NH 11 8 18 20 11 - - “ - 1100 800 1800 2000 1100 - UF57620 LFE NH 14 10 19 15 16 - - “ - 1400 1000 1900 1500 1600 - UF57620 LFE NH 8 20 16 18 12 - - “ - 800 2000 1600 1800 1200 - AVG OD 1100 1266.7 1766.7 1766.7 1300

UF57621 LFE NH 15 14 14 14 - - - “ 1500 1400 1400 1400 - - - UF57621 LFE NH 15 12 19 14 - - - “ 1500 1200 1900 1400 - - - UF57621 LFE NH 16 14 12 11 - - - “ 1600 1400 1200 1100 - - - AVG OD 1533.3 1333.3 1500 1300

UF57622 LFE NH 16 19 - - - - - “ 1600 1900 - - - - - UF57622 LFE NH 24 22 - - - - - “ 2400 2200 - - - - - UF57622 LFE NH 18 17 - - - - - “ 1800 1700 - - - - - AVG OD 1933.3 1933.3

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AVG OD FE 1733.3 1455.6 1383.3 1533.3 1766.7 1300 NH UF45069 RTB ING - 17 10 - - - - “ - 1700 1000 - - - - UF45069 RTB ING - 16 13 - - - - “ - 1600 1300 - - - - UF45069 RTB ING - 15 15 - - - - “ - 1500 1500 - - - -

AVG OD TB - 1600 1266.7 ING UF226999 LTB LSP 17 13 - - - - - “ 1700 1300 - - - - - UF226999 LTB LSP 12 13 - - - - - “ 1200 1300 - - - - - 1A UF226999 LTB LSP - 15 - - - - - “ - 1500 - - - - - 1A AVG OD 1A TB 1450 1366.7 LSP UF276488 LTB COL 12 ------“ 1200 ------UF276488 LTB COL 10 ------“ 1000 ------2A UF276488 LTB COL 9 ------“ 900 ------2A AVG OD 1033.3 2A UF276489 LTB COL - 13 11 6 7 - - “ - 1300 1100 600 - - - UF276489 LTB COL - 15 18 10 8 - - “ - 1500 1800 1000 - - - 2A UF276489 LTB COL - 11 - 13 13 - - “ - 1100 - 1300 - - - 2A AVG OD - 1300 1450 966.7 933.3 2A AVG OD TB 1033.3 1300 1450 966.7 933.3 COL UF57185 LTB NH - 16 15 13 - - - “ - 1600 1500 1300 - - - UF57185 LTB NH - 18 18 16 - - - “ - 1800 1800 1600 - - - UF57185 LTB NH - 16 15 13 - - - “ - 1600 1500 1400 - - - AVG OD 1666.7 1600 1433.3

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Table S14. Stacked layer osteocyte lacunae (OL) counts and calculated volume for each growth zone. Abbreviations: EL. = Element; Loc.

Locality; L and R = Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A,

COL = Coleman 2A, NH = Nichol’s Hammock cality. “ = same value as above; italicized values are averages per individual and whole locality.

Total Volume per GZ (x10-4 mm3)

Specimen OL OL OL OL OL OL OL El. Loc. GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 # GZ1 GZ 2 GZ 3 GZ 4 GZ 5 GZ 6 GZ 7 UF57621 LFE NH 19 19 15 18 - - - 3.0 3.6 5.4 5.4 - - - UF57621 LFE NH 24 14 18 14 - - - “ “ “ “ - - - UF57621 LFE NH 21 21 23 20 - - - “ “ “ “ - - - UF57620 LFE NH - 14 27 30 26 14 - - 3.0 3.6 6.6 6.0 5.4 - UF57620 LFE NH - 13 18 23 23 15 - - “ “ “ “ “ - UF57620 LFE NH - 21 18 31 24 23 - - “ “ “ “ “ - UF57622 LFE NH 22 21 - - - - - 4.8 4.8 - - - - - UF57622 LFE NH 28 32 - - - - - “ “ - - - - - UF57622 LFE NH 24 24 - - - - - “ “ - - - - - UF45040 LFE ING - 21 23 26 - - - - 6.0 6.6 9.6 - - - UF45040 LFE ING - 26 15 28 - - - - “ “ “ - - - UF45040 LFE ING - 25 19 30 - - - - “ “ “ - - - UF45044 LFE ING - 23 24 26 21 29 27 - 6.6 6.6 10.8 10.2 13.8 8.4 UF45044 LFE ING - 20 29 29 17 27 18 - “ “ “ “ “ “ UF45044 LFE ING - 21 23 28 23 32 24 - “ “ “ “ “ “ UF300294 LFE ING - 52 ------12.0 - - - - - UF300294 LFE ING - 42 ------“ - - - - - UF300294 LFE ING - 41 ------“ - - - - - UF300298 LFE ING 22 ------8.4 ------UF300298 LFE ING 34 ------“ ------UF300298 LFE ING 23 ------“ ------

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UF300296 LFE ING 20 ------7.2 ------UF300296 LFE ING 22 ------“ ------UF300296 LFE ING 28 ------“ ------UF300297 RFE ING 22 ------8.4 ------UF300297 RFE ING 21 ------“ ------UF300297 RFE ING 22 ------“ ------UF86378 RFE LSP 28 23 - - - - - 5.4 6.0 - - - - - UF86378 RFE LSP 23 29 - - - - - “ “ - - - - - UF86378 RFE LSPA 26 27 - - - - - “ “ - - - - - UF239925 RFE LSPA 25 7 23 - - - - 7.8 6.0 9.6 - - - - UF239925 RFE LSP 26 22 20 - - - - “ “ “ - - - - UF239925 RFE LSPA 26 14 23 - - - - “ “ “ - - - - UF239925 RFE LSPA - 13 ------“ - - - - - UF45069 RTB INGA - 22 15 - - - - - 6.0 7.2 - - - - UF45069 RTB INGA - 17 11 - - - - - “ “ - - - - UF45069 RTB ING - 17 20 - - - - - “ “ - - - - UF22699 LTB LSP 20 15 - - - - - 4.2 4.2 - - - - - UF22699 LTB LSP 17 18 - - - - - “ “ - - - - - UF22699 LTB LSPA - 13 - - - - - “ “ - - - - - UF276488 LTB COLA 26 - - - - - 6.6 ------UF276488 LTB COLA 24 ------“ ------UF276488 LTB COL 18 ------“ ------UF276489 LTB COL - 15 18 11 17 - - - 3.6 4.2 4.2 4.2 - - UF276489 LTB COL - 17 17 15 13 - - - “ “ “ “ - - UF276489 LTB COL - 14 - 20 13 - - - “ “ “ “ - - UF57185 LTB NH - 19 20 20 - - - - 4.2 4.2 4.2 - - - UF57185 LTB NH - 16 20 26 - - - - “ “ “ - - - UF57185 LTB NH - 19 19 17 - - - - “ “ “ - - -

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Table S15. Stacked image Osteocyte lacunae density for each growth zone. Bold values are averages per individual and population. Italicized values are averages per individual and whole locality. * = Osteocyte Density based on a single individual. Abbreviations: El. = Element; L and R

= Left or Right element; FE = Femur; TB = Tibia; HU = Humerus; RA = Radius; ING = Inglis1A, LSP = Leisey Shell Pit 1A, COL = Coleman

2A, NH = Nichol’s Hammock.

Density per GZ (lacunae/mm3)

Locality El. GZ1 GZ2 GZ3 GZ4 GZ5 GZ6 GZ7 UF45040 FE - 35000 34848.5 27083.3 - - - UF45040 FE - 43333.3 22727.3 29166.7 - - - UF45040 FE - 41666.7 28787.9 31250 - - - AVG OD 40000 28787.9 29166.7 - -

UF45044 FE - 34848.5 36363.6 24074.1 20588.2 21014.5 32142.9 UF45044 FE - 30303 43939.4 26851.9 16666.7 19565.2 21428.6 UF45044 FE - 31818.2 34848.5 25925.9 22549 23188.4 28571.4 AVG OD 32323.2 38383.8 25617.3 19934.6 21256 27381

UF300294 FE - 43333.3 - - - - - UF300294 FE - 35000 - - - - - UF300294 FE - 34166.7 - - - - - AVG OD - 37500 - - - - UF300296 FE 27777.8 ------UF300296 FE 30555.6 ------UF300296 FE 38888.9 ------AVG OD 32407.4 - - - - - UF300297 FE 26190.5 ------UF300297 FE 25000 ------UF300297 FE 26190.5 ------AVG OD 25793.7 - - - - - UF300298 FE 26190.5 ------

205

UF300298 FE 40476.2 ------UF300298 FE 27381 ------AVG OD 31349.2 - - - - -

AVG OD ING FE 29850.1 36607.7 33585.9 27392.0 19934.6 21256.0 27381.0

UF86378 FE 51851.9 38333.3 - - - - - UF86378 FE 42592.6 48333.3 - - - - - UF86378 FE 48148.2 45000 - - - - - AVG OD 47530.9 43888.9 - - - - UF239925 FE 32051.3 11666.7 23958.3 - - - UF239925 FE 33333.3 36666.7 20833.3 - - - - UF239925 FE 33333.3 23333.3 23958.3 - - - - UF239925 FE - 21666.7 - - - - - AVG OD 32906 23333.3 22916.7

AVG OD LSP FE 40218.4 33611.1 22916.7

UF57620 FE - 46666.7 75000 45454.6 43333.3 25925.9 - UF57620 FE - 43333.3 50000 63888.9 38333.3 27777.8 - UF57620 FE - 70000 50000 86111.1 40000 42592.6 - AVG OD 53333.3 58333.3 65151.5 40555.6 32098.8

UF57621 FE 63333.3 52777.8 27777.8 33333.3 - - - UF57621 FE 80000 38888.9 33333.3 25925.9 - - - UF57621 FE 70000 58333.3 42592.6 37037 - - - AVG OD 71111.1 50000 34567.9 32098.8

UF57622 FE 45833.3 43750 - - - - UF57622 FE 58333.3 66666.7 - - - - - UF57622 FE 50000 50000 - - - - - AVG OD 51388.9 53472.2 - - - - -

206

AVG OD NH FE 61250 52268.5 46450.6 48625.1 40555.6 32098.8

UF45069 TB - 36666.7 20833.3 - - - - UF45069 TB - 28333.3 15277.8 - - - - UF45069 TB - 28333.3 27777.8 - - - - AVG OD ING TB* 31111.1 21296.3

UF22699 TB - 47619.1 35714.3 - - - - UF22699 TB - 40476.2 42857.1 - - - - UF22699 TB - - 30952.4 - - - -

AVG OD LSP TB* 44047.62 36507.9

UF276488 TB 39393.9 ------UF276488 TB 36363.6 ------UF276488 TB 27272.7 ------AVG OD 34343.4

UF276489 TB - 41666.7 42857.1 26190.5 40476.2 - - UF276489 TB - 47222.2 40476.2 35714.3 30952.4 - - UF276489 TB - 38888.9 - 47619.1 30952.4 - -

AVG OD COL TB 34343.4 42592.6 41666.7 36507.9 34127.0

UF57185 TB - 45238.1 47619.1 47619.1 - - - UF57185 TB - 38095.2 47619.1 61904.76 - - - UF57185 TB - 45238.1 45238.1 40476.19 - - -

AVG OD NH TB* 42857.1 46825.4 50000

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