THE OHIO PLEISTOCENE MAMMAL DATABASE (OPMDB): CREATION AND PRELIMINARY TAPHONOMIC AND SPATIAL ANALYSES. Ina M. Terry. A Thesis

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1 THE OHIO PLEISTOCENE MAMMAL DATABASE (OPMDB): CREATION AND PRELIMINARY TAPHONOMIC AND SPATIAL ANALYSES Ina M. Terry A Thesis Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE December 2013 Committee: Margaret M. Yacobucci, Advisor Peter V. Gorsevski Jeff Snyder

2 2013 Ina M. Terry All Rights Reserved

3 iii ABSTRACT Margaret M. Yacobucci, Advisor The Late Pleistocene-Early Holocene of Ohio was a period of dynamic changes in climate, flora, and fauna. Climate and flora studies have been greatly aided by palynology research in Ohio s prevalent peat deposits but faunal dynamics, particularly for large (> 44 kg) mammals, are less certain. Available Pleistocene-aged fossils are limited and existing databases are largely incomplete. This study adds to the available data through the creation of the Ohio Pleistocene Mammal Database (OPMDB). The database is composed of fossil finds within Ohio of probable Pleistocene age that have been collected from historic sources, i.e., period newspapers, science journals, etc., and compiled into a geographically referenced database. Within this thesis, I describe the scope and breadth of the OPMDB and present preliminary taphonomic and geospatial analyses using the OPMDB. Initial results are consistent with those previously described in the scientific literature, supporting the view that historical reports can be reliable sources for information about fossil mammal occurrences. Clear differences in spatial distribution and preservational potential exist among Late Pleistocene-Early Holocene mammals. Analysis of the distributions of preserved species and individual skeletal elements by sedimentary context revealed that the greatest variety of taxa is preserved in peats. Mastodons dominate Ohio fossil localities, making up 56% of occurrences in the OPMDB, with sites spread throughout the state. Mastodons are found in a variety of depositional contexts, from peats and clays to gravels. In

4 iv contrast, mammoths are relatively rare in peats and clays, and none are known from the lake plain of northwest Ohio. The peccary record is notably rich, with many complete skeletons; peccaries are most likely to be found in fluvial sands and silts. Other ungulates, including equids, cervids, and bovids, are most often represented in the OPMDB by isolated cranial material (teeth, horns/antlers, partial crania); a collecting bias against unremarkable postcranial material may be a factor for these taxa. Cluster and geospatial analyses delineate two mammal species associations in Ohio that have also been reported elsewhere, the Mastodon and Mammoth Faunas. Additional faunal associations derived from the OPMDB may be unique to the Great Lakes region. Integrating data on early human occupation sites in Ohio did not reveal a strong association between humans and any one potential prey species; rather, humans may have tracked suitable habitat space. While the OPMDB has proven to be a useful tool for investigating Late Quaternary faunal dynamics in Ohio, inclusion of additional data on depositional environments and evidence of human modification of skeletal material, as well as more radiocarbon dates, would increase its utility. In particular, future work should expand on taphofacies analysis, exploration of spatial relationships among mammal taxa and humans, and changes in faunal composition and spatial distribution throughout this interval.

5 v No person exists in a vacuum. There are many people who have assisted me in my path. I would like to thank my children Michael, Caitlynn, and Nathan who supported me wholeheartedly, my brother Mike and his beautiful family for their encouragement, and my parents Mike and Linda Lambert who knew I would finish. To my sister and best friend Lynn, I made this. To my little granddaughter Lauren, Science is cool. To everyone else who ever assisted me in any manner possible, you know who you are, I thank you. Last, but most certainly not least, my husband Joseph who listened and believed I could complete this work even when I did not.

6 vi ACKNOWLEDGMENTS I would like to acknowledge the tremendous support of my advisor Dr. Peg Yacobucci. I cannot thank her enough. I would also like to thank my committee members Dr. Peter Gorsevski and Dr. Jeff Snyder for their help and constructive criticism. I would like to acknowledge and thank Bob Glotzhober, Senior Vertebrate Curator of the Ohio Historical Society, for his assistance. Many thanks go to Betsy Zunk who assisted me with GIS issues. Additionally, I thank the Department of Geology at Bowling Green State University for their support through the years.

7 vii TABLE OF CONTENTS Page CHAPTER I. INTRODUCTION AND OBJECTIVES... 1 Objectives... 2 CHAPTER II. BACKGROUND... 3 Glacial and Interglacial Intervals... 3 Beringia... 4 Refugia... 5 Dynamic Environment of the Ohio Late Pleistocene... 6 Fauna... 7 North American People CHAPTER III. METHODS Database Database Statistics Temporal Difficulties NISP vs. MNI Richness and Dominance Skeletal Element Analyses Skeletal Element vs. Taxa Herding vs. Non-Herding Groups Skeletal Element vs. Sedimentology Geographic Information Systems Paleohuman Data... 23

8 viii Raup-Crick Cluster Analyses Paleohuman-Mammal Associations Sedimentological Analyses Sedimentology vs. Taxa Sedimentology vs. Skeletal Element Taphonomy Voorhies Groups Taphofacies CHAPTER IV. RESULTS Database Statistics Skeletal Element Analyses Skeletal Elements by Taxon Herding vs. Non-Herding Skeletal Element Analyses Sedimentological Context of Skeletal Elements GIS Analyses Spatial Distribution of Mammal Taxa Spatial Distribution of Paleohumans Spatial Distribution of Faunal Associations Raup-Crick Cluster Analyses Paleohuman-Mammal Associations Sedimentological Analyses Sedimentology vs. Taxa Sedimentology vs. Skeletal Element... 55

9 ix Taphonomy Voorhies Groups Taphofacies CHAPTER V. DISCUSSION Database Statistics Skeletal Element Analyses Skeletal Element by Taxon Bison spp Caprinae Carnivora Castoroides ohioensis Cervidae Equus spp Pilosa Proboscideans Rangifer tarandus Tapirus spp Tayassuidae Herding vs. Non-Herding Groups Sedimentological Context of Skeletal Elements GIS Analyses Spatial Distribution of Mammal Taxa Bison spp

10 x Caprinae Castoroides ohioensis Cervalces scotti Cervus elaphus Equus spp Megalonyx jeffersonii Proboscideans Rangifer tarandus Tayassuidae Spatial Distribution of Paleohumans Spatial Distribution of Faunal Associations Raup-Crick Cluster Analyses Paleohuman-Mammal Associations Sedimentology Sedimentology vs. Taxa Sedimentology vs. Skeletal Element Taphonomy Voorhies Groups Taphofacies CHAPTER VI. CONCLUSIONS AND FUTURE AVENUES OF RESEARCH CHAPTER VII. FIGURES AND TABLES REFERENCES APPENDIX A. ADDITIONAL BACKGROUND TEXT AND FIGURE

11 APPENDIX B. ADDITIONAL FIGURES AND TABLES xi

12 xii LIST OF FIGURES Figure Page 1 Quaternary timescale as adapted from the Subcommission on Quaternary Stratigraphy North American glacial maximum as adapted from Dyke et al., Glacial lobes into the Great Lakes region Glacial map of Ohio Some of the oldest known radiocarbon dated sites within the Americas State of Ohio digital elevation map within Ohio state border outline in white Polygon created by the intersection of the dated paleohuman site directional ellipse and all paleohuman site ellipse Number of individual listings per taxon from the Ohio Pleistocene Mammal Database (OPMDB) Taxon representation in the OPMDB Whittaker plot and statistics for relative species abundance in the OPMDB Rarefaction curve for OPMDB assemblage Relative frequency of skeletal elements in OPMDB Relative frequency of whole skeletons (A) and partial skeletons (B) by taxon Relative frequency of skeletal elements for Bison spp. (A) and for Subfamily Caprinae (B) Relative frequency of skeletal elements for Order Carnivora (A) and for Castoroides ohioensis (B)

13 xiii 16 Relative frequency of skeletal elements for Family Cervidae (A) and for Equus spp. (B) Relative frequency of skeletal elements for Mammut americanum (A) and for Mammuthus primigenius (B) Relative frequency of skeletal elements for unidentified Proboscidea (A) and for Order Pilosa (B) Relative frequency of skeletal elements for Rangifer tarandus (A) and for Tapirus sp. (B) Relative frequency of skeletal elements for Family Tayassuidae Relative frequency of skeletal elements for herding versus non-herding mammals Relative frequency of skeletal elements for herding mammals (A) and for non-herding mammals (B) Relative frequency of sedimentologies represented in the OPMDB Relative frequency of sedimentologies in which crania (A), mandibles (B), teeth (C), and tusks (D) were recovered Relative frequency of sedimentologies in which horns, horn cores, and antlers (A), shoulder girdles (B), ribs (C), and vertebrae (D) were recovered Relative frequency of sedimentologies in which hip girdles (A), long bones (B), and manus/pes (C) were recovered Relative frequency of sedimentologies in which whole (A) and partial (B) skeletons were recovered Map showing localities of all taxa in the OPMDB

14 xiv 29 Kernel density overlain by localities of fossil mammals (in blue) from the OPMDB Kernel density map derived from OPMDB locality data excluding all mastodon point data Kernel density map derived from OPMDB locality data excluding all proboscidean point data Bison spp. kernel density map overlain by directional deviation ellipse Caprinae kernel density map overlain by directional deviation ellipse Castoroides ohioensis kernel density map overlain by directional deviation ellipse Cervalces scotti kernel density map overlain by directional deviation ellipse Cervus elaphus kernel density map overlain by directional deviation ellipse Equus spp. kernel density map overlain by directional deviation ellipse Mammut americanum kernel density map overlain by directional deviation ellipse Mammuthus primigenius kernel density map overlain by directional deviation ellipse Megalonyx jeffersonii kernel density map overlain by directional deviation ellipse Rangifer tarandus kernel density map overlain by directional deviation ellipse Tayassuidae kernel density map overlain by directional deviation ellipse Unidentified Proboscidea kernel density map overlain by directional deviation ellipse Map showing localities of all paleohuman site localities in the Ohio Historical Society (OHS) database

15 xv 45 Directional deviation ellipse created from all paleohuman localities in OHS database Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 15,460-14,030 cal yr Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 13,840-12,950 cal yr Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 12,900-12,000 cal yr Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 11,255-10,245 cal yr Kernel density map created from all OHS paleohuman localities Kernel density map created from all paleohuman localities with radiocarbon dates Kernel density map created from paleohuman localities from the 15,460-14,030 cal yr interval Kernel density map created from paleohuman localities from the 13,840-12,950 cal yr interval Kernel density map created from paleohuman localities from the 12,900-12,000 cal yr interval Kernel density map created from paleohuman localities from the 11,255-10,245 cal yr interval Kernel density map for all Proboscidea localities in the OPMDB Kernel density map for the Mastodon Fauna Kernel density map for the Mammoth Fauna

16 xvi 59 Kernel density map for all hoofed mammals Dendrogram of taxa using full dataset Directional deviation ellipses for Cluster Directional deviation ellipse for Cluster Directional deviation ellipses for Cluster Directional deviation ellipses for Cluster Directional deviation ellipse for Cluster Directional deviation ellipse for Cluster Dendrogram of taxa using simplified site and taxon data Dendrogram of taxa excluding M. americanum M. americanum kernel density map overlain by paleohuman occupation ellipse Mastodon Fauna kernel density map overlain by paleohuman occupation ellipse M. primigenius kernel density map overlain by paleohuman occupation ellipse Mammoth Fauna kernel density map overlain by paleohuman occupation ellipse Megalonyx jeffersonii kernel density map overlain by paleohuman occupation ellipse Castoroides ohioensis kernel density map overlain by paleohuman occupation ellipse Cervalces scotti kernel density map overlain by paleohuman occupation ellipse Equus spp. kernel density map overlain by paleohuman occupation ellipse Bison bison kernel density map overlain by paleohuman occupation ellipse Platygonus compressus kernel density map overlain by paleohuman occupation ellipse

17 xvii 79 Relative frequency of taxa found within the human occupation ellipse Relative frequency of taxa found within the human occupation ellipse for sites dated to 15,460-14,030 cal yr (A) and 13,840-12,950 cal yr (B) Relative frequency of taxa found within the human occupation ellipse for sites dated to 12,900-12,000 cal yr (A) and 11,255-10,245 cal yr (B) Relative frequency of taxon occurrences in gravel (A), sand (B), sand-mixed (C), and silt (D) settings Relative frequency of taxon occurrences in clay (A), peat (B), and mixed (C) settings Relative frequency of skeletal elements preserved in gravel (A), sand (B), sand-mixed (C), and silt (D) settings Relative frequency of skeletal elements preserved in clay (A), peat (B), and mixed (C) settings Cluster analyses of skeletal elements Simplified taphofacies dendrogram Cluster A Glacio-facies One-Ice Contact Deposition (A & B) and Cluster B Glacio-facies Three-Late and Post Glacial Deposition (C & D) Cluster C Glacio-facies 2 Proglacial Lake Deposition (A & B)

18 xviii LIST OF TABLES Table Page 1 Key to Americas map with reference numbers, sites, and average radiocarbon age Skeletal element categories and explanation of contents within each category Sedimentology categories and description of sediment listings within each category Total fossil skeletal element content per sedimentology category Skeletal element composition of Voorhies groups as defined by M. Voorhies (1969) Statistical representation of taxa in the Ohio Pleistocene Mammal Database (OPMDB) Distribution of feeding modes for taxa in OPMDB Richness and diversity indices for the OPMDB assemblage Total number of fossil specimens per skeletal element category Taxonomic composition of each group defined by cluster analysis on localities Taxonomic composition of each group defined by cluster analysis on localities using simplified dataset Taxonomic contents and taxon percentage of each group for Figure 68 dendrogram Correlation table between kernel density rasters of paleohumans and relevant prey mammals Correlation table between kernel density rasters of paleohumans dated 15,460-14,030 cal yr and relevant prey mammals Correlation table between kernel density rasters of paleohumans dated 13,840-12,950 cal yr and relevant prey mammals

19 xix 16 Correlation table between kernel density rasters of paleohumans dated 12,900-12,000 cal yr and relevant prey Correlation table between kernel density rasters of paleohumans dated 11,255-10,245 cal yr and relevant prey mammals Taxon occurrences in each sedimentological category Key to groups shown in Figure

20 1 CHAPTER I. INTRODUCTION AND OBJECTIVES The Late Quaternary encompasses approximately the last 130 ka and is a period full of climatic and environmental changes. It includes part of the Pleistocene and all of the Holocene epoch (Figure 1). The present research project focuses on the last portion of the Rancholabrean or Sheridanian North American Land Mammal Age ( ka). Of particular interest is the Late Pleistocene/Early Holocene boundary, when there was a mass extinction event(s) affecting a significant segment of the large vertebrate fauna of North America. As a consequence of this extinction, by the Middle Holocene faunal and floral associations were mostly modern in construction and distribution. The end-pleistocene extinction of North American megafauna has long been a point of interest among geologists, paleontologists, anthropologists, and archaeologists. Thirty-five genera of large mammals (body mass of over 44 kg) became extinct during this time period (Haynes, 2002; Yansa & Adams, 2012). Many researchers consider the extinction to be the result of global climate change at the onset of the Younger Dryas chronozone, about 12,800-11,700 cal yr (Woodman & Beavan Athfield, 2009). Still additional investigations have posed the role of other mammals, such as Canis domesticus, in the depletion of megafaunal populations, or that of disease-driven megadeath of Pleistocene mammals (Fiedel, 2005). A popular culprit in the megamammal extinction has been the implied over-kill by humans, specifically the Clovis culture. Paul Martin, a University of Arizona paleoecologist, first proposed this idea about thirty years ago (Levy, 2006; Martin, 1984). The end Pleistocene extinction(s) transpired around 13,060-11,610 cal yr, which overlaps with the Clovis Culture complex, 13,200-12,420 cal yr (Woodman & Beavan Athfield, 2009). The timing of the spread of Clovis culture and the extinction of large vertebrates is generally suggestive of a link between

21 2 the two as many academics also believe the Clovis culture was specialized in large mammal hunting. Questions still remain as to the role that individual events may have played in the end- Pleistocene extinction. This is especially true as regional differences in extinction rates are becoming more apparent and more refugia areas are discovered for previously presumed extinct fauna. Objectives My initial objective is to determine whether historical sources can be utilized to augment the paucity of Pleistocene vertebrate data for Ohio. I have gathered as many available sources previously disregarded as non-scientific or too vague, e.g., historical newspapers and periodicals, older science journals, etc., in order to create a database, the Ohio Pleistocene Mammal Database (OPMDB), of probable Pleistocene-aged large mammals. My second objective is to assess the relative completeness of the OPMDB by exploring taxonomic, taphonomic, and sedimentology patterns that reveal differences in preservational potential among Late Quaternary mammals in Ohio. Third, I hypothesis the OPMDB will depict associations between Ohio Late Quaternary fauna consistent with other North American fossil assemblages and therefore, will statistically and spatially analyze the OPMDB for such relationships. Fourth, I predict paleohumans do not depend largely upon Mammut americanium and Mammuthus primigenius as generally hypothesized and therefore, my fourth objective will incorporate paleohuman site data from the Ohio Historical Society (OHS), the Paleoindian Database of the Americas (PIDBA), and The Kentucky, Ohio, and West Virginia Radiocarbon Database from Cultural Resource Analysts, Inc (CRA) within the GIS to compare spatial patterns between likely human occupation and probable ranges of large mammal taxa from the OPMDB.

22 3 CHAPTER II. BACKGROUND Glacial and Interglacial Intervals The Late Wisconsinan glaciation of North America generally is considered to span the period between 35,000 and 11,150 cal yr (Bluemle et al., 1999; Fullerton & Bush, 2004). Two main ice sheets covered the northern portions of the continent, the Laurentide and Cordilleran ice sheets (Figure 2). Together they covered nearly all of Canada and portions of the United States as well, an area of some 16 million km². When added to the ice on Greenland, these sheets constituted roughly 40% of total global ice volume (Dawson, 1992). Sea level was approximately 130 m lower than today because of the sheer volume of fresh water locked up in ice pack (Bluemle et al., 1999). Directly to the east of the Rockies was a relatively ice-poor to ice-free corridor created by a scarcity of precipitation and adiabatically warmed air in the lee of the mountains (Dawson, 1992). Worldwide efforts to determine global ice extents began in 1976 with the CLIMAP project. The Last Glacial Maximum period is generally considered to coincide with a period of low global sea level and relatively stable climate approximately 24,700 to 23,600 cal yr (Dyke et al., 2002; Fullerton & Bush, 2004). Optical dating from sediments deposited in small icemarginal lakes in Baraboo Hills, WI place the maximum expanse of glaciations at 18,500 cal yr. The persistence of these lakes is taken to indicate no significant ice volume thinning at the glacial margins (Attig et al., 2011). According to Dyke et al. (2002), using evidence from Heinrich events, a large portion of the Laurentide Ice Sheet actually reached its greatest extent before the Last Glacial Maximum by several thousand years and remained at this limit until beginning its retreat around 17,000 cal yr (Dyke et al., 2002). In central Iowa, the late Wisconsin glacial limit occurred about 16,500 cal yr (Fullerton & Bush, 2004).

23 4 During the Quaternary, ice lobes would advance and retreat across Ohio, fluctuating before ultimately retreating around 16,700 cal yr into the Great Lake basins (Figure 3) (Dawson, 1992; Dyke et al., 2002). The fluctuations of the ice lobes coupled with melt waters from decaying ice sheets created ice-dammed proglacial lakes that covered portions of the state at different time periods before the final retreat into the present day Lake Erie basin (Dawson, 1992). These periods of advance and retreat are captured in the surficial deposits of Ohio (Figure 4). The Laurentide margin to the south of the Great Lakes has been extensively studied and dated. Fortuitously, this portion of the sheet traversed across a landscape full of trees and vegetation, which created regions of sub-till forest and peat deposits available for radiometric dating techniques. The southern Laurentide margin reached 39 N in the Midwest region of the United States (Bromwich et al., 2005; Dyke et al., 2002). Radiocarbon dates show the advance into the area around 28,700 cal yr and peaking at 23,950-22,490 cal yr before sharply declining after 21,290 cal yr (Dyke et al., 2002). The retreating ice sheet left lakes, ponds, and rivers behind filling in topographical lows. Some lakes and ponds underwent siltation and expanded plant growth at their margins, eventually in-filling and eliminating open water conditions. A general succession of open water to mire to bog transpires. These wetlands became areas of peat deposition due to reduced vegetative decay under increasingly anaerobic conditions. Basins differ from each other in sediment accumulation, vegetation content, and pollen suites according to geographic locale and permit the determination of successive floral regimes through time (Dawson, 1992). Beringia

24 5 The term Beringia was first proposed by the Swedish botanist Eric Hultén in Since that time, researchers have speculated on the role of the Bering Land Bridge as a biotic interchange between Asia and North America. The known geographic range of Beringia has expanded considerably since Hultén first proposed its designation (Elias & Crocker, 2008). A hiatus of fossil material from central Alberta between ka indicates that the icefree corridor through Beringia did not exist during the full glacial interval. During the last postglacial, late-surviving proboscideans, muskoxen, horses, camels, and other mammals facing extinction followed preferred habitats northward while other mammals such as moose (Alces alces) and elk (Cervus elaphus) pushed southward towards midcontinental North America (Burns, 2010). Refugia The scenario of a near-synchronized extinction wave crossing continents is inaccurate as evidenced by the existence of late survivors within refugia (Woodman & Beavan Athfield, 2009). Refugia are presently the subject of much study, in part because of concerns about current climate change. Geographic ranges and genetic diversity of species are frequently linked to the cycles of glacial and interglacial environmental parameters (Klütsch et al., 2012). This pattern included the persistence of species subgroups well beyond their established time ranges within protected refugium areas. These areas present opportunities to learn about evolutionary processes such as genetic diversity, speciation, adaption, and extinction (Klütsch et al., 2012). Major refugia for both plant and animal life have been located south of the Laurentide ice sheet placement within North America. The Gulf of Mexico coastal region between the Mississippi River and the Apalachicola River-Appalachian Mountain system was a refugium, as was the area south of the Appalachian Mountains in Eastern North America. Extensive mountain chains, e. g.,

25 6 the Rocky Mountains, and areas adjacent to them along the western coastline of North and Central America also provided refugia far longer than previously supposed. For example, woodland caribou (Rangifer tarandus caribou) persisted in Alabama until 11,820 cal yr, far south of the retreating ice sheets and later than many other regions of North America (Klütsch et al., 2012). Dynamic Environment of the Ohio Late Pleistocene Climate during the Pleistocene was marked by warming and cooling trends tied to glacial and interglacial periods. While some have argued that climate change may have been the primary trigger for the megafauna s extinction (Broughton et al., 2008; Haynes, 2002; Woodman & Beavan Athfield, 2009), it is important to note that the animals that went extinct at the end of the Pleistocene had been exposed to climate fluctuations before that were no more severe in nature than the ones preceding. The fauna weathered these climatic shifts with apparently little difficulty (Levy, 2006; Scott, 2010). Ohio was located alongside the southern margin of the Laurentide ice sheet. At one point, about 2/3 of Ohio was covered with ice (Hostetler, 1997). Beetle data from the Quillin site (northeast Ohio) from a leaf litter layer dated 18,005-16,849 cal yr provides a July temperature of 15 C. Climate models suggested precipitation was cm per yr. Even at the coolest extremes, humans could reside within ice-free zones. There is evidence of a warming phase in Ohio around 15,639-15,102 cal yr and by this time, precipitation has a marked east-west gradient with increased rainfall in the east and decreased to the west (Shane, 1994). The altering climate conditions produced changes within Ohio flora, which effectively altered faunal habitats. Mapping of paleovegetation is based upon fossil occurrence data and extrapolation of information about modern flora/environment/climate relationships. This

26 7 approach can be particularly useful for Quaternary studies as the flora from this time is identical, or nearly so, to modern vegetation. Therefore, using known tolerances allows greater robustness of inferences made about past environments. Shane (1994) used pollen assemblages from stratigraphic sections to evaluate climatic conditions within Ohio and subdivided records into four major time increments. From the beginning of ice retreat to ca. 13,690 cal yr, arboreal pollen assemblages were dominated by spruce pollen (>50%) and indicating spruce parkland or forest, which varied in density across the state (Shane, 1994). From 13,690-12,900 cal yr, conditions began to transform as the diversity of the tree population expanded and maple, elm, hickory and beech appeared in the fossil record, indicating progressively more open conditions (Shane, 1994). The period from 12,900-11,700 cal yr saw rapid changes in vegetation. Spruce pollen spiked in portions of the state before crashing and then all conifer populations collapsed at the end of the period. Deciduous pollen occurrences declined as conifer pollen increased (Shane, 1994). Between 11,700-10,200 cal yr, practically all conifer taxa were eradicated. Over the state, oak and hickory became the dominant taxa as did walnut to a lesser degree (Shane, 1994). These types of trees presented different foraging opportunities and more open woodland environments. Fauna By the end of the Pleistocene, North America lost some 35 genera of mostly large mammals, primarily herbivores. These taxa may have become globally extinct (29 genera) or extirpated only from North America (6 genera). Of the 35 genera that went extinct in the Pleistocene, 16 genera survived somewhere within North America until about 13,800-11,400 cal yr. However, it is not precisely known when the other 19 genera became extinct. Their extinctions may have been time-transgressive as they were in Eurasia. Mammal populations were

27 8 not the only groups affected. Approximately 19 genera of birds also became extinct, about half of which were predators or scavengers (Grayson, 2007). Other mammals diverted from prior geographic ranges and developed entirely new ranges. Caribou (Rangifer tarandus) and musk ox (Subfamily Caprinae) roamed across Tennessee and even down to Mississippi, whereas presently they reside within the Arctic Circle (Grayson, 2007). Members of the extinct fauna of Ohio are most closely related to species common to the eastern United States. The Great Lakes region, as well as other portions of North America, have yielded great quantities of Pleistocene age fauna sites. Frequently, fossil assemblages are dominated by one, or a few very common taxa. Few specimens or sites have been radiocarbon dated, thus hampering efforts at constraining the timing of extinction. Additionally, at least some of those fossils with dates are likely incorrect as dates were obtained using antiquated radiocarbon methodology (Woodman & Beavan Athfield, 2009). Therefore, it becomes imperative that previous data are reexamined under new stringent methods before possible timing, patterns, and causes of the end Pleistocene extinctions can be revealed. Pleistocene fauna are often dominated by Equus spp., Bison spp., and Mammuthus spp. (Jass et al., 2011). Examples of this include a locality from Dawson, Alaska in which 46% of the individuals were steppe bison, 19% were Yukon horse, 11% each were mammoth and Dall sheep, and 3% were caribou. Other sites in Alaska are dominated nearly equally by Bison sp. and Equus sp. with lesser amounts of other taxa (Harington, 2011). This relationship between taxa allows paleontologists to define typical faunal associations, like that of the Mammoth Fauna assemblage. The Mammoth Fauna consists of the co-occurring steppe bison (Bison priscus), horse (Equus spp.), caribou (Rangifer tarandus), and Mammuthus spp. The association was prevalent throughout the British Isles, northern Europe, northern Asia, and through Alaska, the

28 9 Yukon, and Northwest Territories (Boeskrorov, 2006; Harrington, 2011; Sommer & Nadachowski, 2006; Zazula et al., 2009). Other species are associated with Mammoth Fauna but not consistently or commonly enough to be included as members of the group. Musk-oxen (Caprinae) and cave lion (Panthera leo spp.) are associated with the Mammoth Fauna (Boeskrorov, 2006). Other species also often co-occur with the Mammoth Fauna, such as giant deer (Megaloceros sp.) and the woolly rhinoceros (Coelodonta antiquitatis), but are not useful for the purpose of this study as they are not found in North America (Boeskrorov, 2006; Rivals et al., 2010). Giant sloths are tentatively associated with Mammuthus spp. as they share many of the same habitats. The North American mylodont sloth (Paramylodon harlani) frequently occurs in the same distributions as mammoths, primarily M. columbi, as an association typical for the southwestern United States and Mexico (McDonald & Pelikan, 2006). Mammuthus spp. often had associations with large Holocene species, presumably resulting from shared habitat and dietary requirements. For instance, in central Europe, mammoth fossils are often found with both roe deer (Capreolus capreolus) and red deer (European Cervus elaphus). This seems to be a typical refugia association from the Last Glacial Maximum. Such communities of mixed Pleistocene and typical Holocene faunal elements are known only from the last ka (McDonald & Pelikan, 2006; Sommer & Nadachowski, 2006). Mastodons (Mammut americanum), too, have their typical faunal assemblage. The giant beaver (Castoroides ohioensis) and elk-moose (Cervalces scotti) are the common members of the Mastodon Fauna and share equivalent habitat requirements (Miller et al., 2000). These animals share habitat requirements that overlap. Slight changes in environmental parameters

29 10 adjust the proportions of the members of the association. Intriguingly, the data presented below also suggest correlations between the Mammoth and Mastodon Faunas. Many more Late Quaternary mammals were included in the database presented below. Specific information about each relevent taxon is available within Appendix A. North American People The arrival of the Clovis culture peoples about 12 ka with their specialized fluted projectile points appears to coincide with the extinction of many mammalian species in North America (Haynes, 2002). However, there is some question as to the accuracy of this date. Fiedel (1999) pointed out an inherent problem with radiocarbon dating is that of young radiocarbon dates. Adjusting radiocarbon dates would place humans on the North American continent for a more extended timeframe than previously supposed and would amplify the anthropogenic impact upon mammalian fauna as the human population increased. Alternatively, pushing back the arrival of humans farther in time would also make any anthropogenically-caused extinction appear more gradual and less catastrophic. Furthermore, evidence suggests Pre-Clovis cultures lived in North America. Fiedel (2000) claimed the Monte Verde, Chile occupation site was pre-clovis in origin and therefore must be the result of a more complex migratory pattern. It has been hypothesized humans were on the Great Plains at 22,500-21,300 cal yr (Holen, 2006). Many very old Clovis and Clovisrelated sites within North, Central, and South America have been accurately dated up to 13,840 cal yr (Faught, 2008) (Figure 5 and Table 1). The Laurentide ice sheet blocked entrance into lower North America from around 25,400-13,830 cal yr as indicated by Pleistocene fossils in central Alberta before and after this time along with a hiatus of faunal evidence during the time period. Cosmogenic chlorine dates of

30 11 glacial erratics from central Alberta concur. An Upper Paleolithic population would, then, have had to migrate overland from Siberia, across Beringia, and southward onto the central Great Plains at some point between 44,100-25,400 cal yr, before glaciations closed the route through Canada (Holen, 2006). Possible evidence for such an event exists in the Old Crow Basin, Yukon, Canada, dated 44,100-29,650 cal yr, where mammoth bone technology seems to predate that of Clovis culture. Evidence suggests the bones were cracked for removal of bone marrow, cut when meat was removed, and altered into tools by early humans of the Yukon. Additionally, bison bones dated 45,400-41,550 cal yr from the Yukon exhibit cut marks from stone tools (Harington, 2011; Holen, 2006). The Bluefish caves and Dawson City areas have artifacts made from bison, mammoth, and caribou bones, which suggest people had been there at least by 25 ka, if not as early as 40 ka (Harrington, 2011). Patterns of bone modification do seem to appear within North American mid-wisconsinan bones. The La Sena mammoth and Lovewell mammoth sites of the Great Plains exhibit this same mammoth bone technology (Harington, 2011; Holen, 2006). The oldest documented paleohuman site within Ohio is Paleo Crossing, about 15,590 cal yr, in the northeastern region of the state.

31 12 CHAPTER III. METHODS Database Sources for the Ohio Pleistocene Mammal Database (OPMDB) were varied but consisted primarily of historically published newspapers, older science journals and publications, and a smaller, Cervalces scotti specific database provided by the Ohio Historical Society s senior vertebrate curator, Bob Glotzhober. It was Mr. Glotzhober who expressed the idea to use such older sources to create a database as, to our knowledge, no one had attempted to do so before. Many of the fossil finds were from publications from the 1870 s-1920 s, coinciding within increased drainage of lowland wet meadows and increasing utilization of gravel and sand pits throughout Ohio. A second phase appeared during the 1950 s-1970 s coinciding with increased public interest in Ohio archaeology and paleontology. However, the recorded fossil finds are not limited to those two phases. The oldest find in the database is from 1812, the newest are from Newspapers were often extremely useful. Older papers frequently provided illustrations of finds so identification of specific taxa was relatively confident. Where illustrations were missing, descriptions of finds permitted placement of specific fossils into taxa. Also, most often scientists and naturalists from local universities and colleges would make positive identification of the find. Newspapers were frequently the only available, or at least the most common, source of information for Americans of the s. Fossil finds made headlines and would draw crowds much like a new art or history museum display does today. Additional reports in successive papers would increase the amount of information available about each find. Fossils finds from papers, whether popular or scientific, were frequently missing specific geographic coordinates. To make sites consistent, location latitudes and longitudes were found

32 13 first by employing the freeware Google Earth program, and then converted in Excel to decimal degrees with the formula: DD = D+( m / 60 )+( s / 3600 ), where DD equals decimal degrees, D is degrees, m is minutes, and s is seconds. Point coordinates from Google Earth use a Simple Cylindrical projection with the WGS84 datum; these were later converted in ArcGIS 10.0 to NAD 1983, zones 16 and 17. Historic site names were recorded, though if the modern nomenclature differed, modern names were put in parentheses. Township information was often listed but if not, it was determined from site location. However, in some cases, such as that of larger cities, township designations no longer apply and this was indicated by NA. If the specific site name was not known, township center coordinates were used, as they represent the next finest level of location detail. When such information was not available, county information was chosen to represent the location data. The latitude and longitude of the county center was applied in such cases, based on the idea that the large vertebrates in this study would have had little trouble ranging over several kilometers within a county. Hence, the exact locale was unnecessary. County data was generally available. The few sites without county-level data were excluded from the study, though they were included within the database contingently, on the hope that more data will become available in the future. Geological data were recorded whenever possible and in such detail as available. Many times, multiple sources for the same fossil find revealed more data, which would then be noted in the database. Sedimentological information was sparse as not even half of all points contain any such information. Due to its fragmentary nature, only the most basic of sedimentology categories were used. Depositional environment was often ill-defined but was recorded in such detail as

33 14 available. Frequently, the horizon at which a find occurred was recorded. Inches and feet designations were converted to metric in Excel through the formulas: m = 0.305*f, where m equals meters and f equals feet. m = *in, where m equals meters and in equals inches. cm = 2.54*in, where cm equals centimeters and in equals inches. The preservational state of each fossil was categorized as disarticulated, partially articulated, or articulated to give some idea of how much transport, disturbance, etc., a site may have undergone. When available, dates of fossils were recorded and labeled as to radiocarbon, biostratigraphic, or stratigraphic method. However, very few finds have reliable dates. For radiocarbon dates, the identification number as well as lab number and location were recorded. The taxon was noted to at least the genus level. Most fossils could be identified down to species. The total number of skeletal elements and the type of bones found were recorded in the database. Sex and age were noted when data were available, although few specimens had determined sex or age. If fossils displayed any alteration that was likely to be made by paleohumans, this was recorded and categorized as art, food, or tool. Whenever possible, the discoverer of the find was noted. Often they were unnamed but any information about them was recorded. Additionally, academic persons or groups that identified or excavated fossils were noted. The date of the discovery was documented as specifically as possible, as this was often used to correlate between finds and eliminate duplicates. If the repository of the fossil was listed, this too was recorded. Often, the fossil went into private collections or those of small town libraries, museums, and even mercantiles. In recording this historic information, it may eventually be possible to trace provenance. If the find went to an official repository, such as a museum or college, the identification number was

34 15 documented. Finally, all publications used were cited as to author, source, and date. If multiple sources were found for the same fossil, sources were recorded with most recent publication first and older sources in descending numerical date order. A total of 549 individual fossil finds were placed into the database. A total of 520 finds were incorporated into the GIS. However, only 513 were used in population and diversity analyses. Finds excluded were those with inadequate or questionable taxon identifications, as well as those with insufficient site location data. Various statistical analyses were performed on population, taxon, sedimentary, and fossil element data (see below). All viable data were used to create graphical representations of total assemblage composition, total skeletal element composition, percentage of whole skeletal finds, percentage of partial skeletal finds, skeletal element assemblages by taxon, skeletal element occurrences within herding vs. non-herding taxa, skeletal element occurrences within specific sedimentary contexts, and taxon occurrences within specific sedimentary contexts. A table of the total number of species and individuals was created in Excel and imported into the free software package PAST to calculate diversity indices and confidence levels, test for normality, and create Whittaker and Individual Rarefaction plots (Hammer et al., 2001). Further, the database was uploaded into ArcGIS 10.0 (ESRI) for geospatial analyses. Database Statistics Temporal Difficulties As so few specimens have reliable geologic dates attached to them, refining the temporal component of the OPMDB was impossible. Like many assemblages, it must therefore be heavily time-averaged. This is most unfortunate as determining large vertebrate and paleohuman relationships relies heavily on the temporal and spatial resolution between these taxa. However,

35 16 some inferences may be made relying upon only spatial relationships, as the large mammals persisted in some relative abundance throughout the end-pleistocene. The Great Lakes region represented a mosaic of the last favorable habitats for these mammals (Yansa and Adams, 2013). As the areas of human habitation are reliably time resolved, taxon densities for possible prey species may still be useful, as Ohio represents a relatively geographically restricted area as defined by Jass et al. (2011). Czaplewski (2012) advocates that even generic identifications can add to the partial depiction of Pleistocene vertebrate associations in regions, lend a deeper time perspective, and support studies and projects that are addressing biological communities today. NISP vs. MNI Generally when a fossil assemblage is examined, researchers attempt to determine the relative population sizes of taxa and the number of taxa present, using rarefaction techniques on abundance data. The number of identifiable specimens (NISP) and minimum number of individuals (MNI) for each genus (or species) is ascertained through a formula such as: MNI = N i, max /N element i where N i, max is the number of bone element i recovered for a given species and N element i is the number of bone element i that is found in a single complete individual of the species (Cannon, 2004; Marshall & Pilgrim, 1993; Smith & Polly, 2013; Springer, 2012). The element used in these calculations is the most common one recovered for the species and can therefore vary between species. For example, if the most common fossil element found for Canis dirus was the atlas vertebra (N element i = 1) but that for Cervus elaphus was the scapula (N element i = 2), the researcher would use each element particular to the species. MNI was developed to create a conservative count of the number of individuals within the ecological community. Criticisms of MNI estimates involve its sensitivity to sample size,

36 17 unequal distributions of elements across locales, and difficulties in calculating MNI (Springer, 2012). The danger of NISP is the possibility of an exaggerated count as it tends towards multiple counting of body parts within the assemblage. This is not a problem if the researcher is using descriptive comparisons of skeletal element representation or in statistical analyses where NISP are individual data points, but it does indicate NISP should not be used for inferential statistics because statistical significance will be overvalued (Marshall & Pilgrim, 1993). Generally, MNI and NISP taxonomic abundance estimates concur except in instances of high levels of skeletal fragmentation (Springer, 2012). In the case of the OPMDB, NISP =MNI as most sites contain one or two skeletal elements. Those which do have more than a few bones are listed separately as a new site location when specimens are separated stratigraphically, temporally, or taxonomically and also in the case where multiple finds were from the same large urban area, such as the city of Columbus, but not from the same street, lot, etc. As the entire database was pooled into one fossil assemblage and not examined per site, this approach was permissible. It is possible some few individuals have been overrepresented but that is unlikely. Richness and Dominance Biologic communities vary in time and space. Biodiversity is an important element of biotic systems and quantitatively determining the amount of diversity within a community is an important component of any paleontological study. Species richness is, most generally, the number of species present within a given sample. Richness is easy to calculate, even when only presence/absence data are available. This count will normally be an underestimate of the actual richness of the total life assemblage.

37 18 Generalities about biodiversity exist, such as the observation that tropical ecosystems are taxonomically richer than Arctic ones. There are definite differences in biodiversity composition due to latitude and elevation gradients. Determining taxonomic richness in paleocommunities is often trickier, though it is assumed many of the generalities that exist today likely existed in the past as well. North American Pleistocene animal communities, regardless of local or regional scale, seem to have been richest in the eastern mid-latitudes of the continent or, at minimum, as rich as those found in the western portion of the continent, and large herbivore richness waned from south to north (Cannon, 2004). Richness will usually increase as the sample size increases. Richness for the database was calculated for the whole state but not for individual sites as few contained more than one taxon or individual mammal. A few sites, such as Sheriden Cave, do contain several taxa and perhaps more than one individual of each but this is the exception rather than the rule within the database. Formulas such as Menhinick s richness index and Margalef s richness index account for sample size and assist in quantifying assemblages. Menhinick s richness index uses the formula: Menhinick's R I = S / n where S equals the number of taxa and n equals sample size. Margalef s richness index uses the formula: Margalef s R I = (S-1) / ln n where S equals the number of taxa and n equals the sample size. Neither index is considered universally correct. Both Menhinick s richness and Margalef s richness were calculated for sites in the OPMDB. Another important characteristic of a community is dominance, put simply as the prevalence of one or more species within the total sample. Samples with more uniformly

38 19 distributed numbers amongst multiple taxa are considered more even and thus, have minimum dominance. Samples with large numbers of individuals of just one or two species show high dominance and are considered to be less diverse. The simplest measure is the Berger-Parker Index of Dominance, which divides the number of individuals within the most common taxon by the total sample size. The Berger-Parker Index of Dominance was here calculated using the American mastodon (M. americanum) as the most abundant taxon. Skeletal Element Analysis Element vs. Taxa All data for all sites in the state of Ohio were pooled into a single assemblage. A skeletal element matrix was created to determine what fossil elements were most common in the assemblage as well as the division of the material amongst the taxa. The data were divided into 14 categories based upon the basic structure of the vertebrate skeleton (Table 2). These categories were chosen to best represent the entirety of the assemblage. Totals for skeletal elements were then tabulated in Excel as percent of category occurrence per taxon. Bone elements were also tabulated for each taxonomic grouping separately, as members of a particular taxonomic group did not always have comparable bone elements preserved within the total fossil assemblage. Skeletal element counts were compiled for all taxonomic groups. Herding vs. Non-Herding Groups Taxa were further divided into herding and non-herding mammals to reveal any differences in the amounts of fossil material available within each group. Mammals within the herding group are as follows: Bison sp. (except B. latifrons), Caprinae, Cervidae, Equus sp., M. americanum, M. primigenus, Tayassuidae, unidentified proboscidae, and Rangifer tarandus.

39 20 Mammals within the non-herding group include: carnivores, Castoroides ohioensis, Pilosa, Tapirus sp., and B. latifrons. Skeletal Element vs. Sedimentology Of the entire database of 549 sites, only 191 sites (less than 35%) reported any sedimentological data. Most data recorded were inadequate for meaningful comparison based upon the typical sediment characters of mineral composition, grain size, etc. However, some observations were made on the most basic of characteristics, described or inferred, such as grain size and combination of grain sizes. Eight classes in total were created for standardizing the assessment between sites with sediment records (Table 3). A total of 310 fossil elements had some sedimentological context (Table 4). The total fossil assemblage was analyzed for the percentage of skeletal elements found in each sedimentological category. Each skeletal element was individually broken down into percentage of occurrence within individual sedimentology classes. For example, mandibles appeared in some percentage within six different sedimentologies while manus/pes occurred in some percentage in only three different sedimentologies. Geographic Information Systems Geographic Information Systems (GIS) allow for the input of vast amounts of data into software (e.g., ArcGIS 10.0) that converts the information into a visual format capable of revealing patterns across space and time. This software permits the quantitative comparison of data between specific time frames for a single location and the detection of spatial correlations in variables. Data for integration into ArcGIS 10.0 include that of the OPMDB and paleohuman site information from the Ohio Historical Society (OHS), Paleoindian Database of the Americas (PIDBA), and radiocarbon data from Cultural Resource Analyst, Inc. (CRA) sources (see below).

40 21 The OPMDB site data were loaded into ArcGIS 10.0 as point data, with the attribute table including key variables. All GIS projections were in NAD 1983 UTM zone 17N and 16N. Points were loaded as an Excel file and then geographically projected onto an Ohio Digital Elevation Map (DEM) available as quadrant rasters from the National Elevation Dataset from the USGS website (Gesch, 2007; Gesch et al., 2002; U.S. Department of the Interior-U.S. Geological Survey, 2006). A DEM was chosen as the base map for its availability, aesthetics, and futures research that may incorporate elevation analyses. Relevant rasters were downloaded and then combined into one base map using the mosaic function in the ArcGIS toolbox. The Ohio DEM was classified by 50 m elevation intervals from m, a total of 30 classes. The Ohio State border was loaded as a feature class, available as a download from Geospatial Gateway, and projected onto the Ohio DEM to create the borders of this study (Figure 6) (USDA/NRCS- National Geospatial Management Center, 2009). The Ohio county polygons were loaded as a single shape file, also available as a download from Geospatial Gateway, onto the Ohio DEM (USDA/NRCS-National Geospatial Management Center, 2009). Point data for taxa of interest were used to construct mammal density layers as shape files within ArcGIS 10.0, using the Kernel Density function in the Spatial Statistics Analysis toolbox. Kernel Density method creates a smoothly tapered raster surface using a kernel function that calculates an interpolated magnitude per unit area from vector data, in this case from point data of fossil localities. The search radius may be set or a default radius will be chosen by the program. The default is the shortest of the width or height of the extent of the input features in the output spatial reference. For the first analyses, data were run with the default search radius setting equaling about 16,600 m. These analyses produced density rasters that were too tightly concentrated around known sites. To create a broader density surface with a more smoothed

41 22 output, a search radius of 50,000 m was input. Classification of the kernel density layers were based on one standard deviation with exclusion of zero values. To create smoother transitions between color classes, 1 / 3 standard deviation breaks were used. The color ramp selected ranged from green (least dense) to red (most dense). Densities and directional deviation ellipses were produced for each single taxon, for taxon associations, and for paleohuman data. PAST-defined dendrogram clusters, i.e., groups of points clustered by their Raup-Crick similarity (see below), were input as an attribute query in ArcGIS 10.0 so that statistics of each association could be determined. Average nearest neighbor statistics were compiled for each cluster. The Average Nearest Neighbor tool determines a Nearest Neighbor Index based upon the average distance from each feature (e.g., point) to its nearest neighboring feature. Nearest neighbor statistics were compiled for each PAST dendrogram cluster. Both 95% and 90% confidence values were used to evaluate p-values in order to assess if clusters were random, dispersed, or clustered spatial associations. P-values are probabilities, in this case, the probability that an observed spatial pattern is created as a random process. The smaller the numeric value, the more unlikely the pattern is from random processes and the more likely the association reflects a quantifiable relationship (ESRI, 2012). The Nearest Neighbor Index (NNRatio) reflects the degree of clustering vs. random spatial patterns. It is expressed as a ratio of the nearest neighbor observed distance (NNObserved) divided by the nearest neighbor expected distance (NNExpected). The expected distance is the average distance value between neighbors (points in this case) if it were a hypothetically random pattern. NNRatios that have values of more than one show a trend of dispersion, while those which are less than one depict a clustering pattern (ESRI, 2012). These

42 23 ratios were calculated for each cluster of each PAST dendrogram and put into an Excel table. All tables of Nearest Neighbor statistics may be found in the appendices. Directional deviation ellipses were also created in ArcGIS for each PAST dendrogram cluster. The Directional Distribution tool generates a standard deviation ellipse encapsulating the spatial characteristics of central tendency, dispersion, and directional tendency. The default setting is one standard deviation, meaning 68% of all site (points) values were contained within the ellipse. The directional deviation ellipse has the greatest concentration of points within the center of the ellipse and near the periphery of the ellipse border (ESRI, 2012). Ellipses depict the pattern of taxon distribution across the known range area in both area and trend direction. Directional deviation ellipses require at minimum three points. For groups such as Tapirus sp. possessing only two known occurrences in Ohio, a Standard Deviation ellipse was created instead. Standard deviation ellipses do not require more than two points to construct. However, ellipses created with so few points have very low confidence and is at best, a suggestion of range. The standard deviation ellipse includes 68% of sites, as in the directional ellipse, but no directional tendency is indicated so the trend of the taxon range cannot be determined. Paleohuman Data Site name data were obtained from two sources, the Ohio Historical Society (OHS) and the Paleoindian Database of the Americas (PIDBA) (Anderson et al, 2010; Ohio Historical Society On-line Mapping). Dates of paleohuman sites were obtained from the Kentucky, Ohio and West Virginia Radiocarbon Database available at Cultural Resource Analyst, Inc. (CRA) and PIDBA (Anderson et al, 2010; Maslowski et al., 1995). Data from PIDBA and CRA do not have geographical location. Location data were obtained from the OHS with permission and are not available to the general public. Geographical locations were downloaded as an Excel file and

43 24 loaded into ArcGIS No geographic coordinate system alterations were necessary as locales were already in UTM coordinates. Points from UTM zone 16 and zone 17 were combined into one dataset using merge in ArcGIS 10.0 toolset. Archeologists and geologists have alternate meanings for the word Paleo. As a result, the OHS and PIDBA have more human sites labeled as paleohuman than those which qualify as paleohuman to geologists. However, all sites were included to reveal patterns of human behavior within the Ohio landscape. Often, paleohuman sites are re-inhabited or in near continuous habitation through time, so utilizing all sites is not unreasonable. Locales with radiocarbon dates listed on the CRA website were further labeled with appropriate time periods. Total point data were analyzed to produce a directional deviation ellipse in ArcGIS Human point data were further analyzed within a temporal context and direction deviation or standard deviation ellipses were created for each time period: 15,460-14,030 cal yr, 13,840-12,950 cal yr, 12,900-12,000 cal yr, and 11,255-10,245 cal yr. The time intervals were chosen from natural breaks in radiocarbon dates from the CRA data. These dates were converted into approximate calendar years for ease of interpretation. Time intervals are roughly 1000 yr in length. Changes in ellipse shape and locale are interpreted to be alterations in total paleohuman range through time. Additionally, kernel density layers were created for the total dataset and for the temporal subsets using the same parameters as the kernel density analyses described above. Raup-Crick Cluster Analyses Data were converted into a presence/absence matrix that was then used to create dendrograms in PAST expressing the distance (or conversely, similarity) between items of interest, in this case, taxa, bone types, and localities. Clustering is a common method of multivariate data analysis; both R-mode and Q-mode hierarchical clustering are possible in

44 25 PAST. The similarity matrix upon which the clusters are based is computed and used to produce a simple correlation dendrogram based upon single linkage. In this method, clusters are joined based upon the smallest distance (nearest neighbor analysis) between the two groups. The analysis continues through all pairs in like manner until the dendrogram with the highest cophenetic correlation coefficient, i.e., most faithful representation of the original pairwise distances, is produced (Hammer et al., 2001). Lengths of branches and clustering patterns indicate degree of distance, or degree of similarity, between end members. Taxon data were input into PAST in order to create dendrograms showing similarities between taxa and site attributes, using the Raup-Crick Similarity Index as the distance measure. Dendrograms produced by this analysis work best to mitigate the effect of one or two dominant taxa upon a whole fossil assemblage. The Raup-Crick Similarity Index is limited to presence/absence data with 1=present and 0=absent. The algorithm compares the observed number of species or other entities co-occurring in both assemblages to the distribution of cooccurrences seen in 200 random replicates (Hammer et al., 2001). The Raup-Crick similarity metric was chosen over simple correlations or Euclidean distance based upon a similar study published by Smith and Polly (2013). Smith and Polly (2013) used cluster analysis to create both Simpson and Raup-Crick dendrograms with Canis dirus, Platygonus vetus, and Panthera onca data from Indiana, Idaho, Missouri, Arkansas, Pennsylvania, Ohio, Arizona, Texas, Canada, Oregon, Colorado, California, Virginia, and Wyoming. They determined the Raup-Crick metric to be the most robust approach, so this method was used here. The default setting of single pair linkage within PAST was used. Clusters and branching patterns in the resulting dendrograms revealed relationships between groups. The initial dendrogram was redrawn by this researcher to simplify its structure, and clusters were assigned a

45 26 distinguishing number for ease of understanding and interpretation. Note that the lengths of branches in the simplified dendrograms are arbitrary and do not represent numerical distances, unlike those produced by the PAST program initially. Three Raup-Crick dendrograms with the most robust cophenetic correlation coefficients were kept. The first dendrogram contained every individual site location and each taxon as individual listings. This dendrogram produced groupings of locations at which a particular taxon was found. The second dendrogram was derived from taxon data simplified to site, meaning locations (such as the city of Columbus) were represented by a single site containing all available taxa at that geographic location. This dendrogram created groupings of locations in which cooccurring taxa were found, i.e., clusters of taxa associations. As mastodon (M. americanum) are the single most common taxon in the OPMDB, input data were trimmed by the removal of all mastodon listings to create the third dendrograms, which can be used to assess the effect this taxon had upon the total association. Individual clusters from the three dendrograms were projected in ArcGIS 10.0 to permit visualization of spatial relationships between ranges of specific taxa and taxa associations. Paleohuman-Mammal Associations General associations of taxa and H. sapiens were determined by first intersecting the total human site directional deviation ellipse and the total human dated site directional deviation ellipse to create a more temporally constrained human polygon (Figure 7). This human occupation polygon was then projected over taxon and taxon association density layers best fitting the configured human range. Unfortunately, few mammal sites are radiocarbon dated so faunal associations are approximate. Additionally, the directional deviation ellipses previously created and defined as 15,460-14,030 cal yr, 13,840-12,950 cal yr, 12,900-12,000 cal yr, and

46 27 11,255-10,245 cal yr paleohuman ranges were compared to taxon point data in ArcGIS 10.0 using the intersect tool in the Analyst toolset. Intersect created a table of all features (taxon point data) within the paleohuman polygons (directional deviation ellipses). Taxa were then totaled and a percentage of each taxon within the polygon obtained. This metric offers an approximate idea of available prey taxa for paleohumans in specific time intervals. Associations of paleohumans and other taxa were further examined through Multivariate Band Collection in ArcGIS Classified kernel density rasters were used in this analysis. Multivariate Band Collection creates statistics for the analysis of raster bands, i.e., those of mammal and paleohuman kernel density raster layers. By selecting the compute covariance and correlation matrices option, the software outputs covariance and correlation matrices as well as basic statistics such as mean, minimum, maximum, and standard deviation for each layer. Multiple rasters may be compared at one time but rasters should all have the same scale. Relevant prey taxa and taxon association density rasters were compared to the total paleohuman density raster and to paleohuman density rasters divided into time slices. Pairwise comparisons of cell values between raster layers, e.g., paleohuman layer and M. americanum layer, create the covariance and correlation matrices, expressed as single numbers. The covariance reflects the relationship between two sets of data, in this case, the cell values of the kernel density rasters. Because covariance has the units of the data from which it is derived, these values are more difficult to interpret than correlations, which are scaled, unitless expressions of the covariances. Specifically, the correlation is the ratio of the covariance between the two layers divided by the product of their standard deviations (ESRI.com, 2012). As the correlation is a unitless number, the correlation matrix is used here for determining associations between paleohuman, individual taxon, and taxa associations.

47 28 There are limitations to covariance and correlation matrices produced by this method. Because the values are derived from spatial data, they are subject to spatial autocorrelation, which means the standard tests of statistical significance cannot be used. As a result, I was limited to interpreting the rank order of correlation values. This does not devalue the entire analysis, as important relationships between all taxa are still observed. Sedimentological Analysis Sedimentology vs. Taxa To document which taxa occurred most often within each sedimentology category, taxon data were counted and graphs of occurrences were created per individual sedimentology class. Sedimentology vs. Skeletal Element Each sedimentology category contained different types of skeletal elements. There were patterns of skeletal types that appeared in each individual sediment class. For example, the peat category had the greatest variety of skeletal elements while the sand category had the lowest variety of skeletal elements. To document which skeletal element groups occurred most often within each sedimentology category, bone data were counted and graphs of occurrences per individual sedimentology category were created. Taphonomy Voorhies Groups Voorhies (1969) used coyote and sheep skeletons in flume experiments. By altering flow rate and recording changes in skeletal element position within the flume, he was able to determine how different skeletal elements were transported within a fluvial situation. Voorhies determined there were three main groups of skeletal elements: Group One, highly transportable skeletal elements usually by flotation or saltation; Group Two, consisting of intermediate

48 29 transport potential by tumbling or traction along the bottom of the stream; and Group Three, having low transport potential. Some skeletal elements qualify for either Group One or Two (Table 5). These groups were converted into a presence/absence matrix to create a dendrogram in PAST and baseline cophenetic correlation coefficient value to compare with similar dendrograms created from OPMDB skeletal element data. Bone elements in the OPMDB were classified as either present or absent in one of the Voorhies groups, 0-absent, 1-present, or 2- could be present in Group One and Group Two. The hierarchical clustering routine in PAST was used to produce dendrograms of county, site location, taxa, and sedimentology data versus skeletal element occurrence data. After examination of all dendrograms, the two dendrograms with the highest cophenetic correlation coefficients were chosen. These were the taxon/skeletal element and sedimentology/skeletal element dendrograms; each had cophenetic correlations greater than the Voorhies baseline value. Taphofacies Taphofacies are sedimentary rock units that are distinguished by the mode(s) of fossil preservation within the package (Allaby & Allaby, 1999; Zabini et al., 2010). Using taphofacies decouples taxon-specific effects on taphonomic processes. A taphofacies unit could be defined by the large percentage of robust bones as opposed to fragile bones, or as one having only planar bone types, such as the mandible, as opposed to cylindrical bones, such as long bones. The best assemblages to use for taphofacies analysis are ones that have multiple taxa, which limits bias based upon the preservational characteristics of specific species (Yesares-García & Aguirre, 2004). Increased heterogeneity in sedimentology, which is often the case in terrestrial systems, can create difficulties in defining taphofacies. This effect may be minimized when shorter time

49 30 frames are used (Moore, 2012). The Pleistocene in Ohio would likely make a good candidate for these types of analyses as the time span is well constrained. Furthermore, many environments that are common to the Pleistocene persist into the modern day and therefore the method may be field tested. Multivariate analysis of skeletal elements in this case was problematic. Generally, taphofacies studies utilize characters of the bone such as cracks, abrasion, etc. to determine the amount of transport and weathering that occurred before burial and fossilization. However, these data are not available, as yet, within this database. Consequently, as with the sedimentology data, element data was simplified and based solely upon presence/absence. Ohio s surficial deposits are largely due to the actions of glaciers, which progressed and retreated repeatedly; these glacial deposits have been mapped in great detail (Figure 4). Detailed stratigraphic data for the OPMDB are practically non-existent so distinctive features such as lamination, fining upwards trends, etc. are unavailable. The only data present are subjective descriptions of sediments, from which sediment grain size may be inferred. These inferred grain sizes are what the initial taphofacies analysis was based upon. Miller (2009) denoted three main glacial depositional facies within New York. As New York and Ohio have similar glacial histories, these designations were chosen for taphofacies analysis. Facies One is Ice Contact Deposition (ICD) characterized by large percentages of gravel and sand sized grains transported great distances through subglacial streams or by ice marginal streams, likely higher energy settings. Facies Two is Proglacial Lake Deposition (PLD) characterized by fine sand, silt, and clay disgorged in meltwaters into subaquatic fans and deltas of proximal glacial lakes and lowlands. Facies Three is Late and Post Glacial Deposition (LPGD), which consists of gravels and sands, but finer than those of facies one, deposited in shallow ponded water or along stream

50 31 channels late in glaciations or postglacially (Miller, 2009). Miller did not discuss peat content, which appears frequently within this study. Hypothetically, peat percentage within Facies Two should be high. The division between Facies One and Three is a gradation that becomes somewhat subjective in the face of so little quantifiable sediment data. It is less certain how great a percentage of peat will occur within Facies One and Three, though it is more likely Facies Three with its lower energy would have more peat than Facies One. As with the Voorhies analyses, taphofacies analysis used Q-mode hierarchical cluster analysis but instead of simple correlation, a Euclidean distance is used to maximize differences instead of commonalities. This generally means the clusters themselves represent specific taphofacies (Moore, 2012; Yesares-García & Aguirre, 2004). The sites within the OPMDB that contained sufficient sedimentary data for analysis were chosen and loaded into PAST. A skeletal element presence/absence matrix was used to produce the results. Site localities were numbered in no particular order to render them anonymous and to avoid personal bias when interpreting the clusters in the dendrogram produced in PAST and in producing a more simplified dendrogram. The resulting distance dendrogram shows localities grouped by the similarities of their recovered skeletal elements. The dendrogram was simplified and each branch was assigned a group number. Separate graphs of both skeletal element occurrence data and sedimentological occurrence data as percentages were created for each individual group. Sediment graphs were integrated into the simplified dendrogram to discover correlations and distinguish taphofacies based upon criteria from Moore (2012). The skeletal and sedimentology composition for each cluster was also determined.

51 32 CHAPTER IV. RESULTS Database Statistics The OPMDB has a total of 527 separate listings. However, only 513 are considered usable for any meaningful statistics. Entries omitted include those from the category giants, as neither genus nor species may be established, and Homo sapiens, as this is not a verifiable paleohuman find (Figure 8). A further three points, all mastodon, were excluded from GIS analyses as they contained errors in projection which could not be rectified. It is probable their geographic locations were inaccurately recorded, which later produced projection error. A total of 21 species have been recorded. However, Bison bison and Bison sylvestris, while divided historically, are likely the same species and were combined. The number of individuals, mode of feeding, and percentage of total assemblage were tabulated for each taxon (Table 6). Three modes of feeding are represented: omnivore, herbivore, and carnivore (Table 7). Most taxa (76.2%) were herbivores, with smaller proportions of carnivores (14.3%) and omnivores (9.5%). The proportional representation of each taxon in the total assemblage was calculated (Figure 9). All muskoxen were pooled into their subfamily (Caprinae) due to their meager representation. Species of Bison, Cervidae (deer), Equus, and Tayassuidae (peccaries) were similarly combined into families. The ground sloths Megatherium sp. and Megalonyx sp. were grouped under Order Pilosa. All carnivores were grouped together because of their scant representation. Mastodons dominate the assemblage with 56% of the total. The next largest group is the related mammoth at 12% and then unidentified proboscidae at 10%. All other groups do not exceed 5% of the total assemblage. The Menhinick s richness index for the database is and the Margalef s richness index is The Berger-Parker Index of Dominance is (Table 8). The Whittaker plot of

52 33 rank abundance reveals the OPMDB is dominated by a single taxon (M. americanum, the American mastodon) above all others by a considerable margin (Figure 10). The rarefaction curve for the OPMDG assemblage begins to level off at approximately specimens, signifying that species richness is unlikely to increase with the addition of new specimens (Figure 11). Skeletal Element Analysis The total skeletal element count for all categories is 723 (Table 9). The unidentified and tooth categories are the most numerous with 184 and 158, respectively. Figure 12 shows the relative frequency of skeletal elements in the OPMDB. Unidentified bone material comprises 25% of the total assemblage. Teeth are the next greatest component with 22%. Surprisingly, 7% of the total assemblage falls into the whole skeleton category and 4% composes the partial skeleton category. Skeletal Elements by Taxon The relative frequency of whole and partial skeletons by taxon were calculated (Figure 13A and Figure 13B). M. americanum again dominate the whole and partial skeleton categories in nearly equal number, 56% and 57% respectively. Tayassuidae comprise 30% of whole skeleton finds. Castoroides, Cervidae, Pilosa, M. primigenius, and unidentified proboscidae each contain less than 5% of whole skeleton finds. Partial skeleton representation is more equitable, with Castoroides and mammoth at 14% each, Cervidae at 7%, and Tayassuidae and mammoth at 4% each. Graphs of taxonomic groups depict differing element assemblages. In all groups but Caprinae, Pilosa, and Tapirus sp., unidentified bone elements composed a significant proportion

53 34 of bone element categories, varying from 11% to 86% of the total. Details for each taxonomic group are discussed in turn below. Bison are represented mainly by cranial material (Figure 14A). Forty-six percent of bone elements have not been identified within the source material. Of the assemblage that remains, crania at 20%, horn at 13%, and mandible at 7% are all from the cranial complex. The only postcranial elements that appear are vertebrae and long bones, each with 7%. Subfamily Caprinae have total element numbers equal to Bison (Figure 14B). However, the skeleton is even more poorly represented. While all material recorded for this group is identified, there is no post-cranial material recorded. Rather, 60% of bone elements are crania and 40% are horn elements. There is a definitive paucity of carnivore material only seven elements and bones that are present are too fragmentary in nature to be identified (Figure 15A). Only 14% of bone elements are identified and these are all teeth. C. ohioensis has the smallest percentage of unidentified material among the OPMDB taxa (Figure 15B). The majority of identified bone elements are from the cranial complex. Crania represent 22%, teeth also 22%, and the mandible rounds out the total with 11%. Partial and whole skeletons are also relatively common with 22% and 6%, respectively. A final 6% of fossils are in the vertebrae group. Other than as whole or partial skeletal finds, little post-cranial bone elements have been recorded. Cervidae is another group that has relatively good representation in all element categories (Figure 16A). Unidentified specimens comprise 23% of the total assemblage, long bones 19%, crania and horn categories each have 13%, and teeth with 10%. Whole (7%) and partial skeletons

54 35 (6%) also have large contributions. The mandible, vertebrae, and manus/pes each have 3% of the total. Equus is a group with relatively poor skeletal representation despite being nearly equal in total element numbers to Castoroides and Pilosa (Figure 16B). Unidentified elements characterize 19% of the total material. Teeth and mandible are 38% and 31% respectively. Very little post-cranial bone material has been identified. Only the groups of long bone, 6%, and vertebrae, 6%, are represented in the total assemblage. Mastodons are among the taxa with the best representations of all element categories (Figure 17A). Teeth comprise the greatest proportion of all elements found with 27%. Tusks at 13% are the second most common identified element. All other skeletal elements are relatively equal in frequency, ranging in between 1-7%. Unidentifiable bone elements are 19% of the total. Mammoth bone elements are also well represented within the fossil assemblage (Figure 17B). However, teeth and unidentified bones are equal in proportion at 32% of the total assemblage. As with mastodons, tusks, 9%, are the second most common identifiable bone element. Other categories are relatively equal in frequency, ranging between 1-6%. Overall, the mammoth proportions are very similar to those of the mastodon. Unidentified proboscidae are those for which the fossil could be either mammoth or mastodon (Figure 18A). A large percentage, 54%, of these bone elements are unidentified. Tusks (13%), long bones (10%), and teeth (9%) are otherwise the largest contributors. Partial and whole skeletons are 6% and 1% of the total, respectively, with ribs, crania, vertebrae, and mandible ranging from 1% to 3%. All Pilosa bone elements recorded within the database have been identified (Figure 18B). However, they do not represent all types of skeletal element. Peculiarly, the elements represented

55 36 have relatively equal frequencies. Long bones, manus/pes, and vertebrae comprise 17% each. Whole skeletons, ribs, and miscellaneous groups each have 11% of the total. Teeth and tusks (which in this case probably refers to the large claws of sloths) each have 5% of the total assemblage, and the shoulder girdle has 6%. For Rangifer, only 29% of bone material has been identified and those are all antlers and antler fragments (Figure 19A). Tapirs are represented in the OPMDB by just two mandibles (Figure 19B). The relative frequency of skeletal elements for Family Tayassuidae is distinctive (Figure 20). Unidentified bones comprise 40% of the total. However, 46% of all element occurrences are whole skeletons. Partial skeletons comprise a further 3%. Individual skeleton elements are relatively rare: mandibles at 5% and teeth and long bone at 3% each. Herding vs. Non-Herding Skeletal Element Analyses There are 674 skeletal element occurrences within the herding group. The herding group is dominated by M. americanum; M. americanum has over four times the total number of skeletal elements in the OPMDB as the next highest group, M. primigenius with 85. M. americanum also has over five times the total of the third ranked group, unidentified proboscidae with 70. Other groups are less well-represented, as totals for each do not exceed 35 skeletal elements. R. tarandus is the least represented, with only seven elements. A total of 49 elements are present in the non-herding group. C. ohioensis and Pilosa comprise most of the total, with 18 elements for each class. All other taxa do not exceed seven elements per class. The least represented is Tapirus, with two element occurrences. Raw counts were converted to percentages within each of the two categories, herding and non-herding, to allow comparison between skeletal element categories (Figure 21). Both groups

56 37 have similar numbers of elements within the categories of shoulder girdle, ribs, long bones, and whole skeleton finds. Total unidentified skeletal elements are dominant percentages for both groups. However, non-herding dominated every other category except tooth and tusk by at least 3%. Tooth finds in the herding group are almost double those of non-herding, and tusk finds are five times more abundant in the herding classification than in the non-herding classification. Graphs of herding and non-herding categories were created to permit description of the relative frequency of skeletal elements in each group. Herding mammal fossils are dominated by unidentified elements at 26% and teeth at 23% (Figure 22A). Tusk (10%), skeleton (7%), and long bone (6%) are the next most abundant categories. All other skeletal elements are at or below 5% of the total assemblage. Non-herding mammal fossils are dominated by unidentified elements at 17%, crania at 13%, and teeth at 12% (Figure 22B). All other categories are more equally represented than those of the herding group. Many classes are between 4-8%; only tusks and shoulder girdles are lower, with both at 2% each. Sedimentological Context of Skeletal Elements The relative frequencies in which sedimentological classes occur in the OPMDB were determined (Figure 23). The clay class contains the majority of fossil elements at 30% of total. Peat (22%) and gravel (20%) are the next largest classes. All other classes have less than 12% of the total assemblage per class. The sedimentological context for each type of skeletal element was then determined. Miscellaneous and unidentified categories were excluded from this analysis. The cranium-upper palate is best represented within the gravel (42%) and sand-mixed (25%) classes (Figure 24A). Peat has 17% of the total. Clay and silt contain 8% each. Mixed and sand classes are not represented.

57 38 Mandibles occur most frequently within clay settings, at 47% (Figure 24B). Mixed sedimentology is not represented. Peat, sand, and sand mixed each have 13% of the total. Gravel and silt classes each have 7%. Teeth are most likely to be found within gravel (34%) and peat (24%) (Figure 24C). However, clay (21%) is also significant. Other categories have 9% or less per class. Tusks are most common within gravels at 39% (Figure 24D). A further 29% occur within the clay class. All other classes have between 7-9% of total except mixed, which has 2%. Horns, horn cores, and antlers occur 57% of the time within the gravel category (Figure 25A). Mixed, sand, and silt have no representation. All other categories are nearly equal at 14-15% each. Shoulder girdles are only present within three sedimentological classes; peat, clay, and silt (Figure 25B). Mixed, gravel, sand and sand-mixed are not represented. 50% of these elements are found within clay, 33% within peat, and 17% within silt. Ribs tend to be found within clays (40%; Figure 25C). Peat (25%) and sand-mixed (15%) are also significant. Other characterized groups have 10% or less. The sand class is not represented. Vertebrae are very common within the clay category at 48% (Figure 25D). Peat includes 21% and sand-mixed 16% of vertebra finds. The remaining classes contain 5% each. No vertebrae are found within the sand class. Hip girdles are equally divided amongst two main groups, with peat and clay having 37% and 38% of occurrences, respectively (Figure 26A). The gravel and sand-mixed classes have 12% and 13%, respectively. Sand and mixed sedimentologies are not represented.

58 39 The incidence of long bones is similarly limited, with no representation within mixed or sand classes (Figure 26B). Thirty-eight percent of all finds appear within peat, 29% within clay, and 21% within gravel class. The sand-mixed and silt classes have 8% and 4%, respectively. Manus/pes elements are best represented in the clay category at 67% (Figure 26C). Only two other categories have occurences of this element category, peat with 25% and mixed sedimentology with 8%. Whole and partial skeletons were also grouped within the sedimentological categories. Whole skeleton finds appear most often within the sand-mixed sedimentology class at 38% (Figure 27A). Clay (25%) and peat (19%) are the next best represented classes. Sand (12%) and gravel (6%) comprise the rest of the total. No whole skeletons are found within the classes of mixed, aquatic, and silt. Partial skeleton finds occur most often within peat (50%) and clay (29%) sedimentologies (Figure 27B). Silt at 14% and gravel at 7% comprise the rest. No partial skeletons occurrences appear within mixed, sand, or sand mixed classes. GIS Analyses Spatial Distribution of Mammal Taxa Figure 28 shows the spatial distribution of all taxa in the OPMDB. The kernel density map computed from the raw point data shows that fossil localities are concentrated in the western half of the state. A northeast to southwest trend is apparent when all data are included. This trend mimics the location of the till plains in Ohio. The lake plain in the northwest corner of Ohio and the area of high elevation within southeast Ohio are the least dense. Figure 29 displays the overlay of individual localities on the kernel density map. It should be noted that while the northeast portion of Ohio appears as densely populated by point data as more central locations,

59 40 the area does not show a marked density when a density layer was utilized. This situation results from multiple stacked point data (that is, multiple fossil specimens recovered) within a single location. The OPMDB data are dominated by proboscidean sites, especially mastodon finds. To explore the possible impact of this mammal group on the overall results, kernel density maps excluding mastodon data and total proboscidean data were compiled. Removal of just mammoth finds did not affect the resulting maps significantly and is therefore not discussed here. When mastodon sites are removed, density narrows across the state, further clearing off the lake plains to the north and high elevations to the south (Figure 30). Removing mastodon points effectively halves the available point data from the data set. Parts of the state in the extreme northwest corner and southeast corner are now barren. The overall northeast to southwest trend of the data is still apparent. Removal of all proboscidean data results in a different density distribution within the state (Figure 31). The greatest density within the state now occurs slightly west of central Ohio. The eastern half of the state has very low densities of taxa compared to the western half. Portions of lake plain and high elevations are further denuded, however a secondary area of density occurs along the southern margin of modern Lake Erie and at the extreme southwest corner of Ohio. The spatial distribution of each mammal taxon is described below. All taxon figures include the kernel density raster overlain by the directional deviation ellipse, in order to express the suggested taxa range. Bison bison and B. latifrons were combined into one grouping as the total numbers were low and the species are relatively closely related. The kernel density map shows that Bison spp. falls within two main clusters, one along Lake Erie s southern margin and two proximal lesser

60 41 concentration areas in the southwestern corner of the state (Figure 32). An outlying cluster is located near the eastern border of Ohio. The directional deviation ellipse trends northeast to southwest. Bootherium, Ovibos, and Symbos were all combined into one group (Subfamily Caprinae) as the numbers of individuals is low. Density pockets occur within four distinctive areas arranged linearly along a northeast to southwest transect of the state (Figure 33). Two areas show the greatest density, one at the northeastern corner and one at the southwestern corner of Ohio. Sites are located so nearly along the northeast to southwest trend that the directional deviation ellipse is extremely vertically constrained and extends nearly from the northeast corner to the southwest corner of Ohio. Order Carnivora numbers were so low and localities had such close proximity that meaningful density and directional deviation analyses for this group were not possible. Castoroides ohioensis density is greatest in the west-central portion of Ohio (Figure 34). Density hubs are loosely connected, somewhat like beads upon a string, with two major concentrations. The directional deviation ellipse trends east to west from the center of Ohio to its western border. Cervalces scotti density concentrations fall into two discrete areas (Figure 35). Both are irregularly shaped triangles. The largest is located within east central Ohio and the smaller within the northeastern corner of the state. The directional deviation ellipse is more vertically compressed than prior groups and trends northeast to southwest. Cervidae elaphus density falls within four distinct concentrations (Figure 36). Two areas are of substantially greater density than the others. Most areas are in the northern half of Ohio. However, one intense density area is in the south central portion of the state. The directional

61 42 deviation ellipse is more ovoid than previous groups but trends northeast to southwest, as for many of the other taxa. Equus density is similarly arranged to that of Castoroides but shifted further southward; two areas of density are discrete from the others (Figure 37). The greatest density occurs within the center of the state with three lesser areas in the western half of the state. One area of density is located at the extreme eastern edge of Ohio. The directional deviation ellipse trends northeast to southwest, with the southern edge in the southwest corner of the state. By far the largest number of specimens belongs to the mastodon, Mammut americanum. The kernel density map for mastodons covers the entire state, except the southernmost tip (Figure 38). The west-central to southwest portion of Ohio has the greatest centers of concentration but significant centers of density also occur in north-central and northeastern Ohio. The directional deviation ellipse trends northeast to southwest as for many other taxa, but the total area of the ellipse is much greater. The woolly mammoth Mammuthus primigenius is the second most abundant identifiable taxon in the OPMDB. However, no mammoth sites occur on the northwestern lake plain. The density map reveals a narrower statewide distribution than mastodons (Figure 39). The entire northwest corner of Ohio is outside of the projected range. Concentration centers are spread arclike along the southern to eastern border of the state. When a directional deviation ellipse is created, the mammoth ellipse falls within state boundaries but farther east and south than mastodons. The ellipse is also more vertically constrained than that of mastodon. The Megalonyx density map reveals five discrete areas of equal intensity (Figure 40). Most are within the northern half of the state. The directional deviation ellipse trends east to west within Ohio.

62 43 Rangifer density is concentrated mainly in the northern half of the state (Figure 41). The area of greatest density is along the southern margin of Lake Erie, with lesser density extending southward into central Ohio. Discrete areas of density are also located in the northeastern corner of the state and on the south-central border. The directional deviation ellipse trends northnortheast to south-southwest in central Ohio. Family Tayassuidae (that is, the peccary Platygonus) density covers more of the state than many other groups but areas of concentration are greatest within central Ohio and the southwestern margin of Lake Erie (Figure 42). The directional deviation ellipse is more ovoid than many groups and trends only marginally northeast to southwest. The Tapirus numbers were so low and had such close proximity between sites that a meaningful density and directional deviation analyses for this group was not possible. Unidentified proboscideans are the second largest group within the data set. The density layer covers nearly the entire state (Figure 43).The lake plain and areas of higher elevation along the eastern and southern borders are exempted. The greatest concentration by far is at the southwestern corner of Ohio. The directional deviation ellipse trends from northeast to southwest. The ellipse s southern end falls within the southwestern corner of the state. Spatial Distribution of Paleohumans Figure 44 shows the spatial distribution of all paleohuman sites in the Ohio Historical Society database. Points cover most of the state but appear less dense on the eastern plateau regions of the state. When nearest neighbor analyses were conducted on the total data set and on each available time slice, localities were determined to be non-randomly distributed, except for the 11,255-10,245 cal yr time slice, to a 95% confidence level. Localities in the 11,255-10,245 cal yr interval were established to be randomly distributed at the 95% confidence level. The total

63 44 data set and data from the 13,840-12,950 cal yr, and 12,900-12,000 cal yr intervals were found to be a clustered pattern, while localities in the 15,460-14,030 cal yr and 11,255-10,245 cal yr intervals were determined to be a dispersed pattern. The directional deviational ellipse for all the paleohuman localities trends northeast to southwest within the central portion of Ohio, beginning at the southern margin of present Lake Erie (Figure 45). Dividing the data into narrower time intervals results in suggestive temporal ranges for paleohumans. The 15,460-14,030 cal yr time range does not have enough data points for a directional deviation ellipse, so a standard direction ellipse was created (Figure 46). The ellipse is in the northeast corner of Ohio and small in area. The 13,840-12,950 cal yr time range also did not have sufficient data for a directional deviation ellipse and is portrayed as a standard deviation ellipse (Figure 47). The 13,840-12,950 cal yr ellipse is larger than that for the previous time frame and is farther southwest of the initial range, moving more towards central Ohio. The 12,900-12,000 cal yr directional deviation ellipse moves northward and trends nearly east-west, with the ellipse exceeding the western border of the state (Figure 48). The 11,255-10,245 cal yr directional deviation ellipse shifts southward and trends northeast to southwest once again (Figure 49). All ellipses, standard and directional, are anchored in the northeast corner of Ohio. The kernel density map for all paleohuman localities covers nearly the entire state but shows the greatest concentration of sites in the north-central to western half of the state (Figure 50). The kernel density for all radiocarbon-dated paleohuman sites is similarly concentrated in the north-central region to the western border of Ohio, although the overall density covers less area (Figure 51). The 15,460-14,030 cal yr time range kernel density is obviously concentrated in the northeast corner of Ohio (Figure 52). The 13,840-12,950 cal yr density layer has concentrations in the northeast and central Ohio (Figure 53). The northeastern concentration is

64 45 secondary to the one located in the center of the state. The density map for 12,900-12,000 cal yr shows three areas of concentration in Ohio, in the northeast corner, southwest corner, and westcentral region along the state border (Figure 54). The northeastern concentration is slightly broader than the other two. The west-central border region is the densest of all areas. Areas of greatest density for the 11,255-10,245 cal yr interval occur in a linear fashion from the northeast corner to southwest corner of Ohio, with the highest density occurring in the southwest corner (Figure 55). Spatial Distribution of Faunal Associations Creating density layers from associations of taxa can reveal ecological links. As a consequence, kernel density maps for several such faunal associations were created. Figure 56 shows the kernel density map for all proboscideans, including both unidentified specimens and specimens identified to genus or species. Such is the commonness of proboscideans within Ohio that all but the extreme southern tip of the state is covered. The southwestern quarter of the state contains the greatest density but the eastern portion of Ohio also has areas of increased density. The kernel density map for taxa belonging to the Mastodon Fauna (i.e., Mammut, Castoroides, Cervalces) is shown in Figure 57. The density for this faunal association is greatest in the southwest to central portions of Ohio, with an additional area of significant density in the northeast portion of the state. Figure 58 shows the kernel density map for the Mammoth Fauna (i.e., Mammuthus, Bison, Equus, Rangifer). The spatial distribution of this faunal group is very different from that of the Mastodon Fauna. The areas of greatest density fan in an arc across the state from the

65 46 northeastern corner, down the eastern border, along the southern border, and ending in the southeast of Ohio. A further analysis pooling all hoofed mammals revealed yet another pattern of density across Ohio (Figure 59). The lake plain and high eastern elevations have no or low density values. The highest density is located in the central portion of the state. Further significant areas are along the southern margin of Lake Erie, the southwest corner, and the northeast corner of Ohio. Raup-Crick Cluster Analyses Three Raup-Crick cluster analyses were designed to test the reliability and robustness of intragroup associations between members of the Mastodon Fauna and Mammoth Fauna. The clusters defined by these analyses were then projected into ArcGIS to create directional ellipses, where overlap of ellipses signifies congruent spatial distributions. All three cluster analyses will be presented here. However, only GIS analyses based on the results of the most strongly supported dendrogram (Figure 60) will be illustrated as figures; the GIS figures based on the other two dendrograms are very similar and may be found in the appendices. The dendrogram shown in Figure 60 was derived from a cluster analysis in which all the taxon occurrences in the OPMDB were grouped according to how similar their localities were. The dendrogram output by PAST was simplified by pooling data that resulted in defining six large clusters and 12 smaller groupings within them. Each of the six clusters and 12 groups were assigned numbers as labels for ease of communication. The taxonomic composition of each of the 12 numbered groups is listed in Table 10. Most groups were classified as 100% of a particular taxon.

66 47 Localities for these 12 groups were mapped separately in ArcGIS. Nearest neighbor statistics were compiled for each group (available in the appendices). At 95% confidence, seven groups showed a non-random spatial pattern while five showed a random pattern. Non-random pattern Groups include 300, 303, 304, , and 311. Groups determined to be random include 301, 302, 305, 309, and 310. Relaxing the confidence level to 90% confidence, eight groups showed a non-random pattern. Group 302 now qualifies as non-random pattern. Directional deviation ellipses for each group were compiled in order to depict the spatial distributions of associated taxa. Cluster 1 (Groups ) has relatively medium to large sized ellipses that generally trend from northeast to southwest, except for Group 300 which trends in an east-west manner (Figure 61). Taxa within this branch include M. jeffersonii, S. cavifrons, R. tarandus, Tapirus sp., and P. compressus. All groups are located in the northern two-thirds of the state except S. cavifrons, which descends past the southeastern state border. Cluster 2 contains just one group (304), which creates a large directional ellipse trending northeast to southwest and generally within the central portion of Ohio (Figure 62). The included taxon is M. americanum. Cluster 3 (Groups 305 and 308) contains the taxa C. ohioensis, Equus spp., and O. moschatus (Figure 63). The directional ellipse for C. ohioensis trends northeast to southwest, within the southern two-thirds of the state, and touching the southwest border. Equus spp. and O. moschatus (Group 308) trend east to west within the western half of the state The directional ellipses for Cluster 4 (Groups 306, 307, and 309) trend northeast to southwest and are located in central Ohio (Figure 64). Bison spp., Megatherium sp., and M. nasutus (309) is northernmost and touches the Lake Erie border. C. scotti (306) is a smaller,

67 48 narrower ellipse located within central Ohio. Unidentified Proboscidae (307) is southernmost and touches the southeast border of Ohio. Cluster 5 contains only one group (311), which itself only contains the taxon M. primigenius (Figure 65). The ellipse is large and relatively narrow. It trends northeast to southwest but primarily covers the southeastern half of Ohio. Cluster 6 also contains only one group (310) that includes Family Carnivora and taxon C. elaphus (Figure 66). The ellipse is medium sized and near circular. The trend is somewhat northeast to southwest and located centrally within the western half of Ohio. The other two cluster analyses were conducted in a similar fashion to that described above, but used trimmed datasets. The second cluster analysis used simplified site and taxon data; as described in the Methods section, this simplified data set combined nearby sites into one location and pooled all taxa found at each location. The dendrogram produced from this simplified dataset has the lowest cophenetic correlation coefficient of the cluster analyses performed (0.8993). The resulting dendrogram may be broken down into 13 groups, numbered (Figure 67). Relationships between all groups may be further simplified into six main clusters. Table 11 lists the taxonomic composition of each group. Cluster 1 contains the taxa Bison spp., R. tarandus, Equus sp., O. moschatus, Tapirus sp., and M. jeffersonii (Groups ). Cluster 2 includes taxa C. ohioensis, M. americanum, and M. primigenius (group ). Cluster 3 contains the fauna Equus spp., M. americanum, Tayassuidae, Carnivora, C. ohioensis, and S. cavifrons (groups , 107, and 112). Cluster 4 includes Equus spp., M. americanum, M. primigenius, and unidentified Proboscidae (groups ). Cluster 5contains the taxa C. ohioensis, C. scotti, M. americanum, and M. jeffersonii (group 110). Cluster 6 includes Carnivora, C. ohioensis, M. americanum, Tayassuidae, O. virginianus, and unidentified

68 49 Proboscidae (group 111). Nearest neighbor statistics were compiled for individual groups. At 95% confidence, eight groups have a non-random spatial pattern while five are randomly distributed. Non-randomly distributed Groups include 101, 103, 104, 106, 107, and Groups Groups 100, 102, 105, 108, and 112 were determined to have a random pattern. Relaxing the confidence level to 90%, 11 groups are likely to be non-randomly distributed. Groups 100, 105, and 112 are no longer determined to show a random spatial pattern. The third cluster analysis excised mastodon occurrences from the dataset, to see whether their abundance was swamping the data signal. The resulting dendrogram may be broken down into 12 groups ( ) falling into four larger clusters (Figure 68). Table 12 lists the taxonomic composition of each group. Cluster 1 includes the fauna C. elaphus, M. jeffersonii, O. virginianus, C. ohioensis, unidentified Proboscidae, Tapirus sp., and M. primigenius (groups ). Cluster 2 contains the taxa C. scotti, Equus spp., U. americanus, O. moschatus, P. compressus, and S. cavifrons (Groups and 611). Cluster 3 includes the taxa B. latifrons, Megatherium sp., M. nasutus, and R. tarandus (Group 610). Cluster 4 contains A. simus and B. bison (group 609). Nearest neighbor statistics showed that, at 95% confidence values, six groups had non-random spatial patterns while six are considered randomly distributed. Non-random spatial pattern groups include Groups and Group 611. Randomly distributed groups include Groups 600, 601, and Groups Relaxing the confidence level to 90% obtained the same results. Paleohuman-Mammal Associations Recall that the polygon defined by the intersection of dated paleohuman sites and all paleohuman sites in the OHS database covers the central portion of the state and avoids the lake plain as well as much of the plateau region (Figure 7). We can use this occupation ellipse as a

69 50 representation of the likely spatial distribution of paleohumans in Ohio, and compare it to the spatial distributions of other mammal taxa. While the distributions of all taxa were compared to the human range, most taxa were not significantly represented within the human range. Therefore, only taxa with the greatest concentration within the human range and those which are possible prey for humans are discussed here. When the paleohuman occupation ellipse is layered over the kernel density map for the mastodon, M. americanum, the greatest density concentrations overlap along the southern margin of the paleohuman range and the entire human range is within the limits of the mastodon density layer (Figure 69). Comparing the entire Mastodon Fauna with paleohumans, the human range overlaps slightly more north than the mastodon-only density map (Figure 70). The M. primigenius density map and paleohuman occupation ellipse overlap the most along the southeastern edge of the human range, although portions of the human range have no mammoth density at all (Figure 71). The Mammoth Fauna density maps overlaps most of the human range, though some areas are still open and overall density is lessened (Figure 72). The ground sloth M. jeffersonii shows an interesting pattern of density that mirrors the outline of the human polygon range (Figure 73). The density map for the giant beaver C. ohioensis is concentrated in the western part of the state, extending eastward across the human occupation ellipse (Figure 74). The stag-moose s (C. scotti) density layer is almost entirely within the human range (Figure 75). Horses (Equus spp.) are concentrated within the southern portion of the human range (Figure 76). On the other hand, the American bison (B. bison) is found only marginally within the human range (Figure 77). The peccary P. compressus is most densely distributed in the center of the human range, although there are areas of concentration outside the human distribution as well (Figure 78).

70 51 For each taxon, the relative frequency of occurrence within the human occupation ellipse was calculated (Figure 79). As with the entire OPMDB, M. americanum dominates the total with 45.81%. All other taxa have less than 11% representation. Unknown Proboscidae are the second most common class with 10.57%, as was true for the OMPDB overall. However, peccaries move up in rank frequency, with M. primigenius and P. compressus equally represented with 10.13%. Next come C. scotti with 5.73%, C. ohioensis with 3.52%, and Equus spp. with 3.52%. Similar calculations were made for each time slice of dated paleohuman occupations (note that, without radiocarbon dates for most mammal site, all members of taxon were included in each time interval). All of these time slices are dominated by M. americanum, comprising between 50-54% of the total taxon occurrences found within that time interval s human occupation ellipse. For the oldest paleohuman time interval (15,460-14,030 cal yr), the human occupation range is dominated by members of the Mastodon Fauna, specifically by M. americanum and C. scotti (Figure 80A). The human occupation ellipse for 13,840-12,950 cal yr has a mix of Mastodon and Mammoth Fauna with lesser percentages of P. compressus and M. jeffersonii (Figure 80B). The human occupation ellipse for 12,900-12,000 cal yr has the widest range of taxa within it (Figure 81A). Mastodons specifically and the Mastodon Fauna more generally dominate the total percentage; however, the Mammoth Fauna percentage is significant and P. compressus has also increased. The time interval between 11,255-10,245 cal yr shows the highest percentage of Mammoth Fauna in the paleohuman occupation ellipse, and the second highest occurrence of M. primigenius of all time ranges (Figure 81B). Other than M. americanum, members of the Mastodon Fauna are at the lowest representation of all time ranges. P. compressus, on the other hand, is at its highest occurrence with 17% of the total.

71 52 The Multivariate Band Collection tool in ArcGIS allows one to compute correlation coefficients for pairs of rasters. In this case, kernel density rasters for mammal taxa and paleohumans were compared to assess the relative degree of spatial correlation between them (Table 13). For paleohumans, the Mastodon Fauna, and the Mammoth Fauna, all correlative faunal associations will be discussed below. For all other taxa, only the most prominent associations will be reported. Subfamily Caprinae, C. elaphus, and M. nasutus have very low correlations with all other taxa and therefore will not be discussed as separate groups. Mammal taxa spatially correlate with paleohuman occupation sites in the following rank order: Mastodon Fauna, M. americanum, Mammoth Fauna, B. bison, M. primigenius, M. jeffersonii, P. compressus, R. tarandus, C. scotti, Equus spp., C. ohioensis, Caprinae, C. elaphus, and M. nasutus (Table 13). The Mastodon Fauna correlates most strongly with the Mammoth Fauna and paleohumans; in rank order: M. americanum, Mammoth Fauna, M. primigenius, paleohumans, C. ohioensis, Equus spp., C. scotti, Caprinae, M. jeffersonii, C. elaphus, P. compressus, R. tarandus, B. bison, and M. nasutus. The Mammoth Fauna correlates most strongly with the Mastodon Fauna; in rank order: M. primigenius, M. americanum, Mastodon Fauna, Equus spp., paleohumans, B. bison, R. tarandus, P. compressus, Caprinae, C. elaphus, C. ohioensis, C. scotti, M. jeffersonii, and M. nasutus. As one would expect, mastodons (M. americanum) are closely correlated with other members of the Mastodon Fauna, specifically C. ohioensis and C. scotti. M. americanum also correlates strongly with M. primigenius, paleohumans, Equus spp., Caprinae, and M. jeffersonii.

72 53 Woolly mammoth (M. primigenius) are closely correlated with their faunal members, Equus spp., R. tarandus, and B. bison, although B. bison is relatively lower. M. primigenius also correlate highly with M. americanum, paleohumans, C. ohioensis, Caprinae, and C. elaphus. Other highly correlated taxon associations include R. tarandus with B. bison and with C. elaphus, P. compressus with Equus spp., M. jeffersonii with M. americanum, M. americanum with M. primigenius, Equus spp. and with C. elaphus, Equus spp. with Caprinae, and C. scotti with C. elaphus. Dividing the data up into temporal slices changes the rank order of correlations between paleohuman and mammal taxa. The spatial distribution of paleohumans from 15,460-14,030 cal yr correlates with mammals in the following rank order: C. scotti, Mastodon Fauna, M. americanum, M. jeffersonii, M. primigenius, Mammoth Fauna, R. tarandus, and C. elaphus (Table 14). Humans in this time interval are actually negatively correlated to some mammals; in rank order; C. ohioensis, M. nasutus, P. compressus, B. bison, Caprinae, and Equus spp. Paleohumans from 13,840-12,950 cal yr correlate to mammals in the following rank order: C. scotti, Mastodon Fauna, M. primigenius, M. americanus, Mammoth Fauna, C. ohioensis, P. compressus, M. jeffersonii, Caprinae, Equus spp., R. tarandus, and C. elaphus (Table 15). Humans in this time are negatively correlated to two mammal species, M. nasutus and B. bison. Paleohumans from 12,900-12,000 cal yr correlate to mammals in the following rank order: C. elaphus, Mastodon Fauna, M. americanum, M. jeffersonii, C. scotti, C. ohioensis, M. primigenius, Mammoth Fauna, M. nasutus, R. tarandus, B. bison, Equus spp., and Caprinae (Table 16). Humans in this time are negatively correlated to P. compressus.

73 54 Paleohumans from 11,255-10,245 cal yr correlate to mammals in the following rank order: Mastodon Fauna, M. americanum, C. scotti, Caprinae, Equus spp., Mammoth Fauna, M. primigenius, R. tarandus, C. elaphus, M. nasutus, C. ohioensis, P. compressus, and M. jeffersonii (Table 17). Humans in this time are negatively correlated to B. bison. Sedimentological Analysis Sedimentology vs. Taxa Taxon occurrences in each sedimentological category are tabulated in Table 18. The peat category contains the most diverse set of fossil occurrences, with ten taxa represented. The next most populous categories are the mixed class with eight taxa, clay with seven taxa, and sandmixed with seven taxa. Only mastodons were found in uncategorized sediments; given this, it was left out of further analyses. Gravel settings are dominated by mastodon occurrences, at 58% (Figure 82A). However, mammoths are most commonly found within gravels (16%). Unidentified Proboscidae also compose a further 12% of the total. Subfamily Caprinae with 8%, Castoroides with 4%, and Equus with 2% make up the remainder of occurrences in gravel settings. Only four taxonomic groups appear within sand settings (Figure 82B). This class is further unusual in that Tayassuidae and mastodon are nearly equal in value, with 36% and 37% respectively. Equus is at 18% and unidentified proboscidae at 9%. The sand-mixed category is completely different from any other sediment class (Figure 82C). Unlike every other category, mastodon, 9%, is not a prominent component. Instead, the most abundant taxon are peccaries (Tayassuidae) with 57%. Mammoth is also abundant at 14%. Bison, Caprinae, Equus, and unidentified Proboscidae each make up 5% of the total.

74 55 As with sand, silt-dominated localities only contain four taxonomic groups (Figure 82D). Mastodon (37%) dominates but not exceedingly so over Tayassuidae and Castoroides, each at 25%. Mammoth is also relatively high at 13%. Clays are also dominated by mastodon at 79% (Figure 83A). Mammoth is the next most abundant at 6%. Pilosa, unidentified Proboscidae, Rangifer, Tapirus, and Cervidae each have 3%. As noted above, peats appear to preserve the most diverse set of taxa. The class is dominated by mastodons at 52% (Figure 83B). Cervidae at 11%, Castoroides at 9%, unidentified Proboscidae at 9%, and mammoth at 4% compose the remainder of more dominant taxa groups. Rangifer, Carnivora, Caprinae, and Equus each have 2% of the total. Mixed sedimentology category is dominated by predator with 33% and Cervidae with 27% occurrence (Figure 83C). Mammoth, Pilosa, Tayassuidae, and Rangifer each have 7% of total occurrences. Castoroides and mastodon have 6% each as well. Sedimentology vs. Skeletal Element The occurrence of skeletal elements in each sedimentological category was also calculated. Unidentified elements composed between 6-20% of each sediment class, with an average of 10.2%. The exception is the mixed sedimentology class, in which fully 55% of all skeletal elements were unidentified. The range of skeletal elements found in gravel settings is larger than that found in clay or peat settings (Figure 84A). Tooth and tusk groups dominate in near equal numbers, 28% and 27% respectively. Shoulder girdles, manus/pes, and miscellaneous elements have no representation. Most skeletal elements show 1-3% occurrences, except horn with 6% and cranium with 8%.

75 56 The sand class is very different from the other sediment categories (Figure 84B). Only five skeletal element groups are represented: tusk 25%, tooth 17%, mandible 17%, unidentified 8%, and most unusually, whole skeleton at 33%. Sand-mixed settings contain a more variable assortment of skeletal elements when compared to pure sands (Figure 84C). While shoulder girdles, manus/pes, and partial skeletons have no representation, whole skeletons represent 33%, as in the sand class. All other skeletal elements occur at frequencies between 3-8%. Tooth and tusk occurrences are surprisingly rare in sand-mixed settings, compared to other sedimentological categories, with only 8% each. Silt settings do not contain several skeletal element groups (Figure 84D). Whole skeletons, horns, manus/pes, hip girdles, and miscellaneous elements are not represented at all. Dominant groups include tusks and unidentified, both with 20% of the total. Partial skeletons make up 13% of fossil occurrences in silt. All remaining skeletal elements comprise 6-7%. Clay settings also include a broad range of skeletal element groups (Figure 85A). Tusks (14%) and teeth (12%) are the largest contributors. Most skeletal element class values are between 3-10%. The cranium, horn, and miscellaneous categories each only have 1%. All skeletal element groups have been found in peats (Figure 85B). The most abundant component found, at 19%, is the tooth class, followed by the long bone group at 13%. All other classes comprise between 1-10%. Mixed sedimentology is the most variable of the sediment classes. The types of sediment included were perhaps less constrained than other groups (Figure 85C). Aside from unidentified skeletal elements, the remaining percentage is dominated by the tooth category with 23%. All other skeletal element classes are between 4-9%. However, only the classes of tusk, rib, vertebrae, and manus/pes are represented. All other element categories are absent.

76 57 Taphonomy Voorhies Groups Voorhies groups were defined in Table 5. Cluster analyses produced dendrograms showing relationships between skeletal element occurrences and taxon, sedimentology, and Voorhies group (Figure 86). The Voorhies dendrogram contains six groups within four larger clusters. The taxon dendrogram also has six groups but five larger clusters. The sedimentological dendrogram has seven groups and six main branches. Neither the taxon nor sedimentological dendrograms match the Voorhies dendrogram. However, the sedimentological dendrogram generally has the same relationships between elements as the Voorhies dendrogram (Table 19). Taphofacies Figure 87 has graphs of skeletal element occurrence as the bottom row and sedimentology composition as the top row of graphs and Figure 120 is the key for skeletal element and sedimentology graphs. The initial dendrogram was created using skeletal element occurrence for fossil site localities. The OMPDB dataset was trimmed and only localities which had attendant sedimentology were used. Sites were grouped into branches using Euclidean distance analysis. They were then pooled together to produce one graph of skeletal occurrence for each branch. Three obvious clusters of similar skeletal element graphs were produced. Using the same skeletal element dendrogram, sedimentary data were reintegrated to sites. Sedimentary data for each branch was pooled to produce one graph of sedimentology for each branch. The total skeletal and element composition for Clusters A, B, and C were determined. Cluster A contains mostly robust skeletal elements and six skeletal element classes overall (Figure 88A). The total skeletal assemblage is dominated by teeth, 42%, and tusks, 29%.

77 58 Cluster B contains a mix of fine and robust skeletal elements within seven skeletal element groups (Figure 88C). Unidentified bones compose the largest segment of the fossil assemblage at 51%; this is the largest percentage of any cluster. Most of the bone elements are robust, with crania-upper palate and horn-horn core-antler having significant representation over other categories. Cluster C contains a large variety of fine and robust elements in which no single bone category dominates the total percentage (Figure 89A). The standard deviation for this cluster, 4.77, is far lower than Cluster A, 13.25, and Cluster B, Of the skeletal element groups, only the horn-horn core-antler category is missing. While the original dendrogram was analyzed using elements and not sedimentological data, some interesting sedimentological patterns emerged. Groups of sedimentological settings appeared which seem closely related. Accordingly, the dendrogram was divided into the three glaciofacies, based upon qualifiers from Miller (2009) and the average peat content of the cluster. Individual graphs for element and sedimentary data are available in the appendices. Using these measures, Cluster A represents Ice Contact Deposition, Cluster B represents Late and Post Glacial Deposition, and the remaining Cluster C represents Proglacial Lake Deposition. Cluster A-Ice Contact Deposition has very high (42%) gravel content and low percentages of smaller grains such as clay, 9%, and silt, 1%. Peat content is 24% for this cluster (Figure 88A). Cluster B-Late and Post Glacial Deposition has a greater mix of sediments than any other group. The total gravel content, 22%, is much lower than Cluster A but the percentage of smaller to mid ranged grain sizes is greater. Overall peat content has dropped to 16% (Figure 89A).

78 59 Cluster C-Proglacial Lake Deposition has the largest percentage of clay, 45%, of any groups as well as other small to mid ranged grain sizes (Figure 88C). This cluster also has the lowest amount of gravel, 5%, of any other cluster. Peat content for this cluster, 20%, is greater than Cluster B but less than Cluster A.

79 60 CHAPTER V. DISCUSSION Database Statistics There are 13 listings for giants within the database. Initially, they were to be excluded from the database entirely, as well as the analyses. However upon reflection, it was decided to include them because later positive identification may be possible using the attributes from the database. Overall, I presume these listings are in fact proboscidean in nature, though some might prove to be Pilosa (ground sloths) as well. People of the late 1800 s and early 1900 s were in the initial stages of scientific thought and preconceived notions of biblical teachings played a large part in investigations. This does not make their observations invalid, merely a product of their time. Therefore, the observations made may prove useful as further details appear. Late Quaternary mammals are not evenly represented in the OPMDB. This is largely the result of the extremely high number of proboscidean finds compared to the entire fossil assemblage. The overall species richness is fair but mastodons dominate the finds. There are many reasons why other taxa may not be as well represented as those of proboscideans. Most probably, collection bias plays a large role, as mastodon teeth, tusks, and bones are large, conspicuous, and unusual. The bones of smaller, hoofed mammals may easily be overlooked. However, there are some puzzling trends which appeared in the database that may not be explainable merely by collection bias. Skeletal Element Analyses Some trends in skeletal element abundance are related largely to differential preservation potential of different bone materials. Unidentified bone elements composed a quarter of all fossil finds and occur in nearly every faunal group (Figure 12). Some of these are unidentified because they were too degenerated or fragmented to be identified but this group also has bones not

80 61 adequately identified in sources. Teeth were the greatest identified category and this makes sense given their composition. While bones and teeth are both made from calcium phosphate, teeth are considerably harder as the ratio of mineral to soft tissue is much higher in tooth enamel than in bone. Most other skeletal element categories appear to be similar in their preservation potential. However, not all taxa have equal representation in each skeletal element group. As a result, examining skeletal element occurrences reveals interesting patterns. Whole skeleton and partial skeleton finds are not uncommon within the database, making up about 11% of recorded finds (Figure 13A and Figure 13B). Taxa preserved in this way are an interesting mix of herding, non-herding, browsing, and grazing animals. Presumably, certain habitats would have had higher preservation potential for intact carcasses, but not all of the taxa preserved as whole or partial skeletons share the same, or even similar, habitats. Rather, a variety of habitats permitted good fossil preservation. Skeletal Elements by Taxon Bison spp. For Bison spp., there exists an issue of misidentification, as species are hard to distinguish from each other and even from domesticated cattle. Bison spp. appear in similar numbers as Equus spp. within this database and both are common components of the Pleistocene fauna elsewhere in North America (Burns, 2010; Czaplewski, 2012; Jiménez-Hidalgo et al., 2013; Harington, 2011; Rivals et al., 2007; Scott, 2010; Velivetskaya et al., 2011; Zazula et al., 2009). Cranial elements comprise 40% of identified Bison bones (Figure 14A). The upper cranium is one of the more robust bones within the skeleton. This heightened resistance to postmortem damage could result in more crania persisting in the fossil record. Cranial features are what best distinguish bison from other bovines, such as domesticated cattle. Without the

81 62 distinguishing cranium and mandible complex, identifying bison with postcranial bones is difficult. Furthermore, B. bison appeared historically within Ohio until the early 1800 s. Therefore, Pleistocene aged Bison spp. could easily be mistaken for historic bison, or even domestic cows. Caprinae Muskoxen (Subfamily Caprinae) are similar in numbers to Equus spp. and Bison spp. within the database, i.e. 2%. Interestingly, muskoxen are not common within the Pleistocene fossil record despite their geographically large range (MacPhee et al., 2005). Caprinae are generally associated with mammoths (Mammuthus spp) rather than mastodons (M. americanum). As Pleistocene Ohio presented less mammoth habitat than mastodon, it is probable there were fewer muskoxen in Ohio than in other areas of North America. Elements found in Ohio are crania and horn-horn cores; this seems to be the case wherever muskoxen are found (Jass et al., 2011; Ray, 1966; Zazula et al., 2009). These skeletal elements are more robust and certainly, more recognizable (Figure 14B). Perhaps, like bison, without any key cranial elements, such as horns, there is misidentification of caprine postcranial fossils. Carnivora Perhaps the most puzzling discovery of this research is the obvious dearth of Carnivora finds. Simply put, mammalian carnivores (dogs, cats, bears, etc.) must have lived here but are conspicuous in their absence from the fossil record (Figure 15A). Identified bone elements consist only of a few teeth from Ursus spp. Carnivora are naturally rarer in most fossil associations, barring predator traps like La Brea or some karst deposits, since they would have existed in lower numbers than their prey species. Many large predators are also solitary with expansive ranges, which also contributes to the absence of finds. However, they appear in much

82 63 greater abundance within states bordering Ohio. Canis dirus, dire wolves, are particularly widespread geographically and yet, none have been found in the state (Dundas, 1999; MacFadden, et. al., 2012; Rincon, et al., 2011). It is not clear why carnivores are so rare in the Ohio record. Perhaps there are fewer opportunities to preserve predators. The Ohio carnivore fossils discovered are generally associated with the pit caverns of the state. There could also be misidentification of taxa. Bone elements could be mistaken for common carnivores like domestic dog (Canis familiarus), or even gray wolf (Canis lupus) and black bear (Ursus americanus), as these species persisted within Ohio into historic periods. Nonetheless, Ohio is unusually deficient in large Carnivora fossil finds. Castoroides ohioensis Castoroides ohioensis (giant beaver) represent only 3% of the total database but interestingly, 22% of finds are of partial skeletons and a further 6% by whole skeleton finds (Figure 15B). Like the elk-moose C. scotti, C. ohioensis shared the same habitat as mastodon. It is probable this is the reason the taxa is so well-represented by near-complete skeletal finds. Interestingly, very little other postcranial material has been attributed to C. ohioensis. Crania, mandibles, and teeth comprise 55% of the total element assemblage. This predominance of skeletal material from the head region is consistent with other occurrence within the North American C. ohioensis fossil record. Incisors and crania are common while postcranial bones are very rare (Lanken, 1993; Swinehart & Richards, 2001). Perhaps the postcranial material is considerably less robust than the outsized crania and large incisors. Or perhaps the postcranial material is just not recognized without the distinctive cranium in attendance. Cervidae

83 64 Deer (Family Cervidae) are reasonably represented within the database, although that is largely due to the inclusion of the C. scotti (elk-moose) database provided by Senior Vertebrate Curator of the Ohio Historical Society Bob Glotzhober (Figure 16A). Elk (Cervus elaphus) are less well represented and most of their finds are unidentified. C. scotti comprises most of the whole and partial skeleton occurrences. C. scotti existed within the same habitat as mastodons while C. elaphus did not. It is probable that this factor relates to why C. scotti has better representation than that of C. elaphus. The remainders of the identifiable elements are skull material such as mandible, teeth, crania, and antler beams. This pattern is consistent with other finds from North America (Laub, 2003; Long & Yahnke, 2011, Schubert, 2004), and is expected, as these skeletal components are distinctive and easily recognizable. The paucity of postcranial bone elements is likely the result of misidentification of species and the persistence of C. elaphus into the present day. While elk do not roam Ohio today, they did historically. Furthermore, white-tailed deer persist and are geographically widespread. Therefore, identification of postcranial cervid material to Pleistocene species and age would probably be problematic. Equus spp. Equus spp. are not as common in Ohio as would be presumed. In North America, horses are a frequently appearing member of the Pleistocene fauna (Jass et al., 2011). In this database, they are represented only by four identified elements: teeth, mandible, long bone, and vertebrae (Figure 16B). Additionally, they compose only 2% of the total assemblage. The problem may be solely an identification issue. The specimens within the OPMDB attributed to the Pleistocene are definitively dated by researchers. Pleistocene horses are nearly identical to modern horses. Again (as with the elk, elk-moose, and peccary finds), any Equus spp. find may not be identified as

84 65 Pleistocene without radiocarbon dating. Instead, it would simply be viewed as domesticated horse bone, hardly a rare occurrence in rural Ohio. Pilosa Ground sloths (Order Pilosa) have an unusual pattern of preservation in which the forequarters seem to have better representation than the hindquarters (Figure 18B). Also, 11% of occurrences are as whole skeletons. Globally, ground sloths are relatively common as whole or nearly whole skeletons and are well represented within all skeletal element groups (Borrero, 2012; Brandoni, 2011; De Iuliis et al., 2009; Hoganson & McDonald, 2007; McDonald, 2000). Ground sloth skeletons could be preferentially preserved because the animals were large, similar in size to the proboscideans M. americanum and M. primigenius. Additionally, ground sloths have very robust bones that are difficult to disarticulate, particularly the vertebrae and forearm connections (Borrero & Martin, 2012). This could explain why the forequarters might be more common within Ohio. However, I have not found this preservational pattern to be noted anywhere else. There are only a few specimens within Ohio, though, so this pattern may be an artifact of sampling. Proboscideans Overall, proboscideans (mastodons and mammoths) have by far the best record in the database. Of the 513 distinct listings, over 250 were mastodons alone. There could be several reasons for this, related both to the wetter habitat where they lived and to the nature of their bones. Generally, the larger bones of proboscideans seem to preserve well globally. Mastodon bone elements are highly recognizable as different from any modern fauna and thus, are preferentially collected by the public. M. americanum teeth, in particular, are very large, robust elements that do not resemble those of any modern mammal and are attractive keepsakes for an

85 66 average person. Moreover, based upon literature, much of Ohio was prime habitat for these proboscideans (Feranec & Kozlowski, 2012; Yansa and Adams, 2012). Low-lying wet meadows, patchy woodlands, and plentiful small water bodies would present the perfect area for cultivating sedges and browse desired by mastodons. These areas also create excellent fossil preservation with their attending peat and clay layers. Animals dying within vegetated wetlands are quickly buried within a relatively anaerobic environment and thus, preserved. The woolly mammoth, M. primigenius, is also well represented in the database, though not nearly at the rate of mastodons (Figure 17B). This difference probably results from a few important factors. Mammoth habitat was not as prevalent in Ohio, thus limiting their geographic range. Preservation potential of the more open, grassy areas preferred by M. primigenius within higher elevations is also not as great as that of mastodon habitats. However, mammoths are still a significant percentage, 12%, of the total fossil assemblage. Like mastodon, mammoth also have the virtue of size, which could be a prime factor for skeleton preservation. Their larger, robust bones, like those of mastodon, preserve well and are easily recognized as different so, again, public bias plays a role in the collection of fossil mammoth material. Mastodon are well-represented within all bone element groups, though teeth and tusks comprise a majority of the finds (Figure 17A). This seems to be the case in other areas where mastodon bone elements are found, as well (Feranec & Kozlowski, 2012; Robinson et al., 2005; Teale & Miller, 2012; Woodman & Athfield, 2009). Mammoth are similarly well represented within all bone element categories. Globally, mammoth skeletons are some of the most abundant Pleistocene fossils (Czaplewski, 2012; Feranec & Kozlowski, 2012; Harington, 2011; Holen, 2006; MacFadden, 2012; Van Kolfschoten et al., 2011; Whittecar et al., 2007). In many skeletal element classes, both mastodons and mammoths have nearly identical representation. The

86 67 reasons for this are likely similarity of size and bone robustness. Although the mammoth is larger than the mastodon, it is not significantly larger and mastodons are considered to have thicker, more robust bones. Therefore, it is likely that they have equal preservation potentials. This idea is supported by the equal number of unidentified proboscidean to mammoths and the fact that they comprise a significant proportion of total finds. Surprisingly, teeth comprise 9% of total unidentified proboscidean occurrences. This is noteworthy because mastodon and mammoth teeth are very different from one another, and informed discoverers presumably should have been able to tell the difference between the teeth (Figure 18A). Percentages of whole and partial skeletal finds are also unusually high for an unidentified group. One explanation is that people of the day used mammoth and mastodon interchangeably within the same article. The problem lies in the term mammoth, as it is both a description of size and a common name for species of the genus Mammuthus. Some publications often did so and in those cases, barring a depiction of the tooth or skeleton, I labeled it as unidentified. Furthermore, whole or partial skeletons without crania could have been problematic to identify given the similar nature of mastodon and mammoth postcranial skeletons. Unidentified tusk fragments are also more easily understood, as fragmented tusks from either mastodon or mammoth greatly resemble each other. Rangifer tarandus The listings in the database for caribou, Rangifer tarandus, are largely unidentified skeletal material, as specific bone elements were not recorded in publications (Figure 19A). Of the identified skeletal material, all were antler beams and fragments. Consequently, the available bone data are sparse. Caribou fossils are globally rare in associations and finds elsewhere often only consists of scant postcranial material, skull pieces, and antler fragments (van Kolfschoten et

87 68 al., 2011; Zazula et al., 2009). Therefore, the lack in caribou fossil material in Ohio could be explained merely as a general preservational issue for this species. The paucity of fossil finds could also be the result sampling bias. R. tarandus antlers are very different from elk or deer and they would thus be more noticeable and likely to be collected, but other portions of the skeleton resemble those of deer or even domestic pig, which promotes misidentification or dismissal of fossil material. Another reason for the rarity of R. tarandus fossils could relate to caribou s association with M. primigenius. Similar scarcities of other Mammoth Fauna species in the OPMDB have already been noted. Tapirus spp. Tapirus spp. in Ohio are extremely rare only two occurrences have been recorded. Of those occurrences, both are from the eastern border of the state and the material found is two mandible fragments (Figure 19B). Fossil tapirs are generally quite rare in the global fossil record despite their expansive geographic range, and generally consist of mandible fragments and teeth (Eberle, 2005; Hollanda et al., 2011; Hollanda & Ferrero, 2013; Scherler et al., 2011). The reasons for this rarity may be the solitary nature of the family. There could also be issues of preservation intrinsic to the species or related to environmental constraints as most extant tapirs prefer forested, tropical habitats. Tayassuidae Tayassuids (peccaries) are an unusual group almost 50% of all finds are of whole or partial skeletons (Figure 20). This occurrence likely results from some biological or sedimentological feature, rather than any unusual feature of their bones. Tayassuids are not exceptionally large; peccaries are roughly the size of a domestic meat pig. They do travel in groups, however, which could be a factor in this case since most peccary finds within Ohio

88 69 include multiple skeletons. This clustering of fossilized individuals suggests that their social behavior permits better preservation. Perhaps if one peccary becomes injured or trapped, the other peccaries linger until they too are ensnared. Of the remaining recorded skeletal elements for this group, 40% are unidentified. This large amount of unidentified material resulted from inadequate information within publications. Often, specific bone elements were not described and therefore were not identified. Of identifiable skeletal elements, mandibles and teeth compose the majority. This is consistent with peccary finds globally (Czaplewski, 2012; Frailey & Campbell, 2012; Gasparini et al., 2010; Gasparini & Ubilla, 2011; Gasparini, 2013; MacFadden et. al., 2012). Mandibular and tooth characteristics are often useful for distinguishing between mammal taxa. It is therefore probable that the paucity of postcranial material, other than whole skeletal finds, results from misidentification of bones as domestic pig. Ohio was and still is largely rural. A person familiar with domestic farm animals would likely ignore bones that resembled pigs, or even deer. Herding vs. Non-Herding Groups Initially, I hypothesized herding mammals would have better fossil representation than non-herding taxa. This seems a reasonable assumption as herding mammals tend to have higher populations than non-herding mammals and a greater density of individuals per unit area. Social behaviors inherent in herding suggest skeletal elements would be present in greater numbers within a fossil assemblage. This increased number of available bones would presumably allow a better representation within each skeletal element category. However, this proved not to be the case. In many instances, non-herding mammals were better represented by a range of skeletal elements (Figure 21).

89 70 The giant beaver C. ohioensis and ground sloths (Order Pilosa) are both part of the nonherding group. These two taxa have good representation among all skeletal element categories in the OPMDB. It is likely this factored greatly in the total representation of non-herding mammals. However, the non-herding group also includes members such as Tapirus spp., B. latifrons, and Carnivora that are poorly represented in nearly all skeletal element categories. Herding members such as mastodon and mammoth are very well represented in total. Additionally, Cervidae and Tayassuidae are reasonably well represented. Nonetheless, the results overall show that herding behaviors do not correlate with increased representation in skeletal element categories. In some individual classes, the results were quite surprising. I had presumed teeth for both groups would be equal, or nearly so. When the results were compiled and compared, herding mammal teeth were represented at nearly twice the numbers as non-herding (Figure 22A and Figure 22B). One likely reason for this difference is the significant amounts of mastodon teeth in the database. Intrinsic physical parameters of the cranium/mandible, such as if herding mammal teeth disarticulate from the dental complex more readily than those of non-herding mammals, could also contribute. C. ohioensis teeth do not seem to separate readily from the jaw, nor do those of Tapirus sp. This is not surprising, as Pilosa, Carnivora, Tapirus spp., and C. ohioensis all consumed tougher materials. If non-herding teeth were preserved within the crania/mandible, they were not counted in the separate tooth category. Suggestively, the percentage of mandibles and crania preserved for non-herding mammals is higher than that of herding mammals. Smaller elements such as the manus/pes and vertebrae also have higher percentages of occurrence in non-herding mammals. There seems to be no obvious explanation for this observation. It is possible these smaller elements from herding group members were consumed

90 71 by predators and scavengers in greater quantity than those of non-herding group members. Such preferential consumption may adequately explain the manus/pes, however, vertebrae have been shown to disarticulate late in the process of reducing a carcass (Hill and Behrensmeyer, 1984). Additionally, why would herding group small skeletal elements be consumed in greater quantity than those of non-herding mammals? A possible explanation could be preservational bias in ground sloths, which tend to have a higher percentage of those skeletal elements conserved and could therefore bias the total non-herding sample. More easily explained is the category of horn, horn core, and antler. Despite herding group members having a greater likelihood of this element occurrence, the non-herding group has twice the representation. This difference likely results from preservational bias in the taxon B. latifrons, as this taxon is only known in the database from horn cores and crania. All other members of the herding group have species that are represented by the horn category and other elements as well. Also obvious is the dominance of the tusk category in the herding group, due to the large number of proboscideans. Sedimentological Context of Skeletal Elements When considering the relative frequency of sediment types in the OPMDB (Figure 23), three categories are conspicuously greater than all other categories: clay, peat, and gravel. Seventy-two percent of fossil occurrences with sedimentological data are found within these three classes. Most of the fossils are found in clay (30%). This makes sense as clay creates very good preservation potential. Peat is only marginally better than gravel, 22% versus 20%. This was somewhat surprising as gravel deposition is associated with high-energy flow, in which one would not expect good preservation of fossil material. However, many of these gravel finds may

91 72 actually represent moraine deposits rather than water-transported material. Under these circumstances, the fossils may be more apt to be preserved at rates closer to peat preservation. According to Moore (2012), crania preservation is highly variable and dependent upon the specific taxa considered. Certainly this is seen within the OPMDB, although this could be an artifact of my classification. Crania were not tallied as present unless specifically listed as such. Therefore, the categories of whole skeleton and partial skeleton may have crania which were not counted separately within the database. Very few taxa are represented within the crania category. Only M. primigenius, M. americanum, S. cavifrons, and C. ohioensis occur in this category. Certainly every member of this group has distinctive skull features and robust skull physiology. Proportionally, most cranial material appears to have been transported some distance. What material is described generally is fragmented and the tougher braincase is the portion remaining. Of all crania preserved, 42% are from gravel settings (Figure 24A). In nearly all cases, these are braincases. Cranial preservation in the sand-mixed category (25%) appears similar. As clay and peat are generally assumed to result from low energy hydrologic conditions, it would be safe to assume crania preserved here would be more complete. However, this seems not to be the case. Other than one complete skull of S. cavifrons from peat, the other representatives are braincases and portions of upper palate. Overall, cranial preservation depends as much on biology as on environmental factors. Mandibles are preserved within nearly every sedimentological category (Figure 24B). According to Voorhies (1969) and Hill and Behrensmeyer (1984), taxonomic identity and transport both affect mandible preservation in slightly different manners. Voorhies grouping places the mandible with the crania with regards to transport potential (that is, can be transported intact great distances), although if the mandible disarticulates from the crania, perhaps its blade-

92 73 like morphology and smaller size would cause it to behave more similarly to hip and shoulder elements. Hill and Behrensmeyer (1984) found that the type of taxon involved may have a role in how quickly mandibles become separated from the crania. These factors could both account for the differences between cranial and mandible sedimentology within this study. Clay and peat housed 47% of mandibles in the database, which are likely in situ based upon sediment type. The other categories of gravel, sand, sand-mixed, and silt possibly indicate transport, although siltdeposited mandibles may be in situ as well. Although teeth are ubiquitous in the OPMDB, they are most commonly found in gravel, peat, and clay classes (Figure 24C). These three categories contain 79% of all tooth finds. A certain portion of these occurrences could be interpreted as in situ or minimally transported, such as those in peat and clay. Certainly the low flow energy associated with clay deposition is inconsistent with the transport of teeth. Teeth are more robust and heavier than many elements. They are rooted in the crania and mandible until tissue and bone deteriorate enough to dislodge teeth. Mandibles are shown to travel some distance through flotation. Crania generally move through saltation, rolling and bumping along the stream bed. Therefore, it is quite possible the gravel category contains tooth elements which may have traveled distances before final deposition. Tusks (in reality, modified front teeth) are limited to just a few taxa within the OPMDB. Tusks occur within every sedimentological category, implying they are robust elements that are highly conservable (Figure 24D). They are well represented within peats and clays, as is the case with most skeletal elements, but they are equally preserved in gravel settings. Interestingly, the sedimentological context of tusks seems to follow the glacial taphofacies model proposed in its representation. Facies One, Ice Contact Deposition, would correspond to the 38% of tusk

93 74 occurrences in gravels. Facies Two, Proglacial Lake Deposition, would be characterized by peats and clays, in which 38% of tusks were found. Facies Three, Late and Post Glacial Deposition, would be characterized by pure and mixed sands and silts, in which 20% of tusks occurred (Miller, 2009). It may be that if tusks within the database were better described, fragmentation and erosion patterns on tusks could delineate specific glaciotaphofacies. The horn, horn-core, and antler category is another taxon-limited skeletal class, as not all large mammals have them, but unlike tusks, very few of these skeletal elements are included in the OPMDB. Peat has 15%, represented by a single Rangifer tarandus antler (Figure 25A). Occurrences of horns/antlers in clay are nearly identical to peat. Both of these possibly derive from lacustrine deposits created by infilling glacial kettle lakes and lowlands. They are also probably in situ or at least minimally transported, as antlers detach seasonally and therefore, discoveries of single beams have little consequence regarding the amount of transport the skeletal element has undergone. The remainders of the sedimentology classes are likely representative of transported skeletal elements. Muskox horn cores (both Ovibos moschatus and Symbos cavifrons, separated from the crania) found in gravel comprise 57% of the horn, horncore, and antler category. Most probably, these gravels are representative of glaciofluvial deposits, as Quaternary muskoxen were periglacial taxa. The sand-mixed category is represented by S. cavifrons horn cores attached to a cranium. Given that the sedimentology was sand-mixed, specifically sand and gravel, and the taxon is known to be periglacial, this was likely a glaciofluvial deposit as well. Shoulder girdles are clustered with vertebrae and hip girdles in Voorhies groups and are presumed to be similarly preserved within fluvial conditions. There are similarities between shoulder, hip, and vertebrae, though the shoulder girdle is preserved under fewer sedimentology

94 75 conditions than either of the other elements (Figure 25B). This difference actually makes sense as presumably the shoulder complex would preserve at least slightly differently and have a different transportation potential. Like many fossil elements, the shoulder girdle has its greatest representation, 83%, within the clay and peat categories, suggesting that these elements preserve very well within lower energy environments. An additional 17% of shoulder girdles are found in silt settings, implying greater transportation potential than the hip girdle (see below). Perhaps because the shoulder bones are lighter in mass but similar in their blade-like construction to hip bones, they have greater flotation potential than the more robust hip girdle. Rib skeletal elements are preserved in many sedimentologies but in the greatest quantity in clay, with 40%, and peat, with 23% (Figure 25C). The likelihood of fossils in these two sedimentologies representing in situ occurrences has been discussed above. However, the occurrence of ribs in the remaining sediment types implies that ribs have a high potential of transportation. Voorhies (1969) related rib and vertebrae performance in fluvial systems. The present results concur as both groups have similar distributions within sediment types. Vertebrae are less common as fossils in the OPMDB. Vertebrae are harder to scavenge from the carcass and tend to disarticulate late in the decomposition process, according to Hill and Behrensmeyer (1984). When transported, the processes that protrude from the core of the vertebra often break off, leaving the round disc of the vertebral body intact. As a result, if the vertebra is clearly identifiable and is found within finer-grained sediment, there has probably been minimal or no transportation. Of the vertebrae found, 69% are from peats and clays and are likely in situ (Figure 25D). The physical conditions of the remaining fossil vertebrae are unknown; it is therefore difficult to interpret the depositional setting.

95 76 The hip girdle is related to other trunk components, such as vertebrae, ribs, and the shoulder girdle, in Voorhies groups and is presumed to be similarly transported within fluvial settings (Voorhies, 1969). Additionally, Hill and Behrensmeyer (1984) noted the trunk complex of shoulder-spine-hip tends to disarticulate last and often preserve together. There are similarities between hips and vertebrae in this analysis as well, though the hip girdle is preserved under fewer sedimentological conditions than vertebrae are. Hips are also similarly preserved to shoulders, although shoulders are present in even fewer sedimentological categories (Figure 26A). This pattern actually makes sense as presumably the larger hip complex would behave differently in preservation and transport potential. Like many fossil elements, the hip girdle has its greatest representation within clay and peat settings. These occurrences are likely in situ based upon the rationale discussed previously. Occurrences of hip elements in the sand-mixed and gravel categories are nearly equal at 13% and 12%, respectively. These occurrences are likely to represent marginally transported elements based upon the larger sediment sizes and Voorhies analysis. The larger blade-like bones would catch the water and probably induce flotation, movement through the water column, or tumbling along the bed at least to the point where stream energy no longer resists gravitational pull or the bone becomes lodged. Moore (2012) reported that long bone elements are most likely to be derived from multiple species and therefore, should be used as the best representative of assemblage species composition. Certainly a number of taxa are represented by long bones in the OPMDB but still less than 30% of all taxa within the database. Long bones are common in peat and clay (Figure 26B); these are probably in situ occurrences. Based on occurrences in the other sediment classes, long bone elements are at least moderately transportable; this factor may supersede taxonomic factors in at least some cases. Voorhies (1969) grouped long bones with shoulder girdles, a

96 77 connection supported by my results as well. While not conclusive, this similarity suggests that environmental factors play a significant role in the preservation of long bones, which must be considered in using fossil material for ancient population reconstructions. Mani/pes (hands and feet) are rarely preserved in the fossil record. In all instances in the OPMDB, these skeletal elements were found in situ, and only within three sedimentological categories (Figure 26C). Furthermore, few taxa are represented in the class. This dearth of hand and foot elements is not unexpected. Both Voorhies (1969) and Hill and Behrensmeyer (1988) reported that the bones of the manus/pes are lost first in transportation and disarticulation. The likelihood these bones will be conserved in fossil assemblages is small. The in situ nature of the OPMDB occurrences stems from the nature of the sedimentology. Animals becoming mired in clay will have their digits preserved upon death. As a significant portion of all finds in the OPMDB were whole and partial skeletons, the sedimentological context for each of those categories was determined as well. With these two categories, transport is expected to be minimal. Fifty percent of whole skeletons were preserved in pure or mixed sand settings (Figure 27A). Interestingly, all of these whole skeletons found in sands were from one taxon, the peccary Platygonus compressus. Only four other taxa are known as whole skeletons. It would seem that while sedimentology does influence whole skeleton preservation, it is additionally controlled by some biologic factors intrinsic to certain taxa, most probably habitat preference, although there could be some social element as well. This conclusion is supported by data on partial skeletons. In this group, 50% are from peats and 29% are from clays (Figure 27B). Within this peat-clay group, only two identified taxa occur, M. americanum and C. ohioensis, and the unidentified taxon is a proboscidean, so it may well be M. americanum as well. In other words, most of the partial skeletons are mastodons,

97 78 regardless of sedimentological class, and the remainders are the giant beaver, C. ohioensis. Both M. americanum and C. ohioensis are members of the Mastodon Fauna. As previously discussed, the habitat preferred by the Mastodon Fauna also produces high preservation potential for fossil assemblages. The high incidence of P. compressus could reflect either their habitat or their social nature. GIS Analyses Spatial Distribution of Mammal Taxa A total of 510 distinct fossil mammal occurrences were geographically located onto an Ohio DEM within ArcGIS 10.0 (Figure 28). Coverage across the state is good and a few obvious clusters can be observed. Because multiple fossil occurrences at the same coordinates appear as one point on the map, a kernel density map derived from all occurrences more accurately shows areas with a preponderance of Late Quaternary mammal sites (Figure 29). In particular, areas of the map, especially in the north, that initially appeared to have fewer localities, actually had a greater concentration of sites. Fossil sites are concentrated within the southwestern quarter of Ohio. There is a northeast to southwest band of higher site density that follows the till plain. Conversely, the northwestern corner of the state and higher elevations of the foothills of the Appalachian Mountains in the southeast contain fewer sites. This makes sense for a few reasons. Higher elevations do not preserve fossil materials as well since rapid burial rarely occurs on slopes and streams transport skeletal material down to lower elevations. The northwestern corner of Ohio was glaciated or lake bed for much of the Quaternary, so terrestrial vertebrates would not have lived in the area nearly as long as in other portions of the state. The largest glacial lobes, the Miami and Scioto,

98 79 covered all of northwestern Ohio and extended further south than lobes over the eastern half of the state (Figure 3). Some of the density pattern may also be explained through historic and present land use patterns. High site concentrations in the southwest along the present Ohio River valley and some of the higher densities in southern and central Ohio correspond to larger cities such as Cincinnati and Columbus. Construction in these cities in the late 1800s-early 1900s uncovered fossil material, and further draining of lowlands and wetlands in west-central Ohio unearthed many fossils as well. Curiously, Cleveland, along present Lake Erie s southern margin in northeast Ohio did not experience any similar boom in fossil discoveries. The northeastern portion of the state would have been covered by the Killibuck and Grand River glacial lobes, which could partially explain the lack of fossil sites. Given that the database is dominated by mastodon (M. americanum) occurrences, point data from this taxon were removed and another kernel density layer was produced (Figure 30). Removing mastodon clearly depicts how much this single taxon directs density values. With mastodon gone, the northwest and southeast corners of the state are clear and there are now three areas in the state with no fossil occurrences. The density in the southwestern corner of the state is still high but not manifestly more so than that of central Ohio. Removing only mammoth (M. primigenius) did not significantly alter density values in any part of the state. Removing all identified and unidentified proboscideans shifts site density distinctly north and east towards central Ohio (Figure 31). The total density within the state is now concentrated within the middle of the state; areas of increased density also appear more prominently along the Lake Erie margin and eastward. The original northeast to southwest trend nearly disappears. Clearly taxa are not

99 80 equally distributed across the landscape and certain taxa may affect the total faunal assemblage in unexpected manners. Kernel densities and directional deviations do not conclusively depict the geographic ranges of taxa. Factors such as patchy habitat spaces, large individual ranges, and temporal changes in range will all affect the appearance of the analyses. However, these maps provide some insight into the overall geographic range of each taxon. Almost all mammal taxa have their greatest density within the glacial till plains. Despite these similarities, each taxon is unique in both its distribution and density within the state. Some taxa follow the till plains very closely, while others move north or south to follow the margins of either the lake plain or plateau regions of Ohio. Details of each mammal group s spatial distribution are discussed below. Bison spp. Bison spp. density is surprisingly patchy considering the genus was so indicative of the Pleistocene (Figure 32). There are two main concentrations, one at the northern end of the till plain and the other at the southern edge. A lesser area of density appears within the glaciated plateau region of Ohio. This area perhaps hints at a wider distribution of the genus that is not captured by data available at this time. No sites within the database occur on the higher elevations of the unglaciated plateau of Ohio. Modern B. bison are grazers for the most part. Fossil bison included in the database were mostly B. bison. It is likely the flat till plains provided better grazing opportunities. B. latifrons was both more solitary than B. bison and less flexible in dietary requirements, preferring woodland and forest habitats. B. latifrons only appears twice within the database, both in Brown County, southwestern Ohio. Because little statistical relevance can be obtained from only two samples, all bison were placed within one group. The B. latifrons sites are on the edge of the till plain and the beginning of the unglaciated plateau to

100 81 the east. B. bison could have grazed within the till plains, avoiding low wet meadows, marshes and river drainage in the center of the state, while B. latifrons occupied the higher elevations of the plateau. The temporal extent of B. latifrons is not well known but it is thought to become extinct about ka (Scott, 2010). It may be that these two Bison species are temporally separate within Ohio and not subject to species partitioning. Caprinae Muskoxen (Subfamily Caprinae) have an unusual distribution in Ohio although there are few incidences of the group in Ohio. Species that do appear are Bootherium bombifrons, Ovibos moschatus, and Symbos cavifrons. B. bombifrons and S. cavifrons are the most common taxa. It has been suggested by some researchers that Symbos is the male and Bootherium the female of the genus Bootherium (Campos et al., 2010). Within this database, both recognized species do co-occur spatially. However, sites are not temporally constrained and could also represent a time-averaged replacement of one species for another. The Caprinae range plots from the northeastern border to the southwestern tip of the state and is very narrow, much more so than any other group encountered (Figure 33). Pleistocene muskoxen, especially O. moschatus, were periglacial (Campos et al., 2010; MacPhee et al., 2005). The distribution of fossil occurrences mimics that of the glacial lobes within Ohio (Figure 3). B. bombifrons commonly occurred in broad intermontane valleys and low plains (Campos et al., 2010). The portions of Ohio where B. bombifrons are found were those suitable and most likely to attract muskoxen. Evidence suggests the muskox diet is very similar to that of the caribou (R. tarandus; Barboza & Reynolds, 2004; Kazmin & Abaturov, 2011; Kazmin et al., 2011). My research supports this interpretation, as occurrences of muskoxen in Ohio are along

101 82 the southern margin of the caribou range, though caribou also appear farther southeast than muskoxen. Castoroides ohioensis The giant beaver C. ohioensis occupies the till plains, for the most part. However, the taxon is noticeably different in distribution from bison. Giant beaver are most common within the lowest elevations of Ohio, but not within the lake plain (Figure 34). Most sites are marginal to the lake plain and drainage of the ancient Teays River within central Ohio. Density trends and the directional ellipse run east to west, with greatest site concentrations along the western border of the state. The ellipse is much smaller and vertically constrained than that of many other groups, indicating a smaller range for the taxon, which may stem from narrower selection of habitat and smaller personal ranges required by individuals of this group. While C. ohioensis has not been proven to build dams or lodges like its smaller modern relative Castor canadensis, it does prefer wetland habitats bordering open water; this portion of Ohio would have provided this setting (McDonald & Bryson, 2010; Swinehart & Richards, 2001). However, there is one unusual area of density within the higher elevations of the unglaciated plain, although still marginal to watercourses, suggesting that elevation was not a factor if sufficient water habitats were available. Cervalces scotti Elk-moose C. scotti are members of the Mastodon Fauna and their range overlaps that of M. americanum and C. ohioensis. The elk-moose is another group very constrained in both areas of density and in its directional ellipse (Figure 35). Most areas of density are within the till plain, although edges extend into lake plain and plateau regions. Elk-moose are believed to have inhabited an environment with no true modern analog (Schubert et al., 2004). The common

102 83 element in all habitats in which they appeared was related more to periglacial climate conditions than to specific flora (Long & Yahnke, 2011). The density pattern presented here seems to suggest such a connection, as it appears to mimic the position of the Wisconsin glacier lobes within Ohio (Figure 3). The deviation ellipse is more vertically constricted than many other groups. As these are larger, long-legged animals that likely had broad home ranges, the narrow distribution in Ohio could indicate less available preferred habitat. These samples might therefore be more temporally constrained than other groups in the OPMDB, as the receding glacier altered habitat space. Cervus elaphus Examples of the elk, C. elaphus, in the database are few. Consequently while it was possible to construct kernel density and directional deviation for the taxon, it is not as informative as other groups. Elk density is patchy in its distribution and subsequently has a large deviation ellipse (Figure 36). Marginally, it seems to be restricted to the till plain but areas of density also occur elsewhere on the plateau regions. Perhaps a more evocative observation would be that the taxon occurs most often at flatter mid-elevations of the state. Research has shown elk are flexible in habitat requirements but choose ones that are proximal to good cover (Sawyer et al., 2007). It is probable these wooded higher plains of Ohio provided the grass and browse they preferred with the cover they required. Equus spp. Equus spp. density is primarily contiguous and on the till plain, though there are two separate areas of density located in the north of the state (Figure 37). There are few samples of horse within the database and so even one site can greatly affect the directional ellipse. The site in northeastern Ohio pulls the overall range slightly to the east and to the border of the till plain

103 84 to plateau region. Most Equus spp. density is in the southwest. Horses are considered members of the Mammoth Fauna (Boeskrorov, 2006; Harrington, 2011; Sommer & Nadachowski, 2006; Zazula et al., 2009). The highest density of mammoth occurs towards the southern borders of Ohio so it is reasonable to expect horse density to be higher there as well. Interestingly, the density layer created suggests a common theme within grazing communities. Bison spp. and Equus spp. have been shown to be codependent where one group dominates, usually Bison spp., the other overlaps its range (Jass et al., 2011). The data similarly show bison density increasing on the till plains where horse density decreases, although not in the entire range and not always to the same degree. Jass et al. (2011) suggested that bison typically replace horse in the later Pleistocene. It may be that occurrences of these two groups are depicting a temporal pattern of faunal change. Megalonyx jeffersonii As there was only one Megatherium sp. and five M. jeffersonii, I created the kernel density and directional deviation for M. jeffersonii alone (Figure 40). M. jeffersonii, C. scotti, and C. ohioensis have been linked together as sharing similarities in habitat (McDonald et al., 2000). However, other researchers have linked ground sloth occurrences to Mammuthus spp. (McDonald & Pelikan, 2006). Data from this analysis concurs with findings from McDonald et al. (2000). The range of the giant sloth encircles large portions of both the elk-moose and giant beaver ranges. However, the density of M. jeffersonii is patchier than that of either taxon. This could result from several different factors. Research about preferred vegetation for sloths is mixed. Some authors claim it preferred flora found within areas of high soil moisture (Borrero & Martin, 2012; Hoganson & McDonald, 2007; McDonald & Pelikan, 2006; Schubert et al., 2004), while others suggested mixed spruce-hardwood forests associated with river systems (Hoganson

104 85 & McDonald, 2007). My research agrees with both to some degree. Sites are lower to mid elevation and proximal to water sources. The considerable overlap with both C. scotti and C. ohioensis suggests shared habitat requirements. I surmise the patchy nature of the ground sloth density could reflect the more solitary nature and expanded range of M. jeffersonii versus the restricted range of C. scotti and of C. ohioensis. Proboscideans M. americanum density is greatest in the southwestern quarter of Ohio and, more importantly perhaps, all of Ohio barring the extreme south shows a significant density of mastodon occurrences (Figure 38). This indicates that mastodon were a geographically widespread species. Mastodons were thought to favor lowlands and wetter areas than mammoths. Certainly the area of greatest distribution lay within portions of Ohio that include these conditions. Furthermore, according to many researchers the Great Lakes region is considered a refugium with Late Pleistocene fauna persisting far longer here than elsewhere in North America (Laub, 2003; Teale & Miller, 2012; Yansa & Adams, 2012). These would help explain some of the density hotspots within Ohio. However, mastodons are also present in areas of higher elevation within Ohio, areas not known for the low, wet meadows preferred by mastodons. Some authors have suggested M. americanum expansion into less preferred habitat could result from increased interspecies competition with mammoth for dwindling resources (Yansa & Adams, 2012). This is certainly plausible. Other researchers have noted mastodon are seemingly less choosy about browse than previously thought and clearly exploit the flora available in a region (Teale & Miller, 2012). Perhaps Ohio is a case where all of these factors apply. There is such a widespread

105 86 obvious persistence of mastodon that the taxon could be exploiting the Great Lakes as a refugium and capitalizing on resources mammoths were reluctant to utilize. Woolly mammoth (M. primigenius), by constrast, have markedly different occurrences in the database, both in number and in geographic distribution. Mammoth do not occur at all in northwestern Ohio and seem to follow areas of higher elevation. Mammoth are geographically restricted to the southern and eastern border of the state with decreasing density towards central Ohio (Figure 39). It is presumed they would similarly continue past the border but as data are confined to Ohio, density decreases at state boundaries. McDonald and Pelikan (2006) noted a decrease in mammoth density eastward in North America as mastodon density increased. Arguably this east-west gradient does not exist within Ohio. However, mammoth density does increase where mastodon density decreases in a north to south pattern. The areas of greatest density within Ohio are also at higher elevation. The directional deviation ellipse is located farther south and east of that from mastodon. This range is consistent with data reported by Yansa and Adams (2012) in their study of mammoths within the entire Great Lakes region. Mammoth are known to be grazing animals. Areas of higher elevation would be cooler, drier, and more conducive to their favored diet. Superficially, the pattern of mammoth s highest density mimics that of the glacial lobe pattern across the state (Figure 3). Areas proximal to the glacier would be cooler than those distal. They would likely also be more open as woodlands required time to achieve any concentration. However with available data, it is not known if mammoth density is related to temporal glacial ice patterns, or interspecies competition with mastodon, or simply favored habitat distribution. Initially, it was hoped that determining ranges for M. americanum and M. primigenius would aid in classifying some of the unidentified Proboscidae within the Ohio fossil record. This

106 87 cannot be accomplished, however, because of the extremely cosmopolitan range of mastodons in the state. The density and directional ellipse of unidentified Proboscidae are more restricted than either the mastodon or mammoth (Figure 43). However, there is still too much overlap of ranges between all the proboscideans. It may be that some inferences could be made about specimens occurring on the fringes of M. americanum range but the certainty of identification of unknown Proboscidae would be suspect. It could be surmised that specimens occurring to the extreme southeast of Ohio would be M. primigenius but again, it would not be conclusive because of the expansive mastodon range. At this point with the data available, classifying unidentified Proboscidae as either M. americanum or M. primigenius is not possible. Rangifer tarandus The caribou (R. tarandus) range is farther north than most other groups. The area of greatest density is at the southern tip of Lake Erie (Figure 41). Caribou were a member of the Mammoth Fauna. However, the sites with R. tarandus are at the extreme northern extent of mammoth in Ohio. All the finds are at lower elevations for the state. Mammoth prefer higher elevations, which may explain why most occurrences of caribou are north of the center of mammoth density. The density of R. tarandus occurrences is roughly comparable to the areas of highest density for peccaries (see below). Fossil assemblages with both caribou and P. compressus do occur in New York (Robinson et al., 2005). However as most sites are located within river valleys, caribou density and range may also be an artifact of transport from higher elevations to lower. Most fossils of this taxon are not identified as to what skeletal element they represent and therefore it is not possible to determine which sites may be in situ and which may not.

107 88 Caribou reportedly migrated northward following the retreating Wisconsinan ice sheets and its attendant boreal habitats (Long & Yahnke, 2011). The pattern of R. tarandus density within Ohio could result from a time transgressive movement of the mammals from higher ice marginal ranges to lower ice marginal elevations as the glacier retreated across the state. The area of highest density for caribou is at the Lake Erie shore. The lake would have proved a barrier for a time and the mammal population could have built up here before they continued northward along the lake shore. Without more temporal data, it is hard to draw any certain conclusions. Tayassuidae MacFadden et al. (2010) reported peccaries are not common members of Quaternary faunal assemblages. This is not the case in this database. Next to the Proboscidae, Tayassuidae is the most common family. Accordingly, the density pattern and directional ellipse is broader than other groups (Figure 42). Platygonus compressus and Mylohyus nasutus both appear in Ohio but not in the same quantity. P. compressus is far more common than M. nasutus and, as a result, the two taxa were combined into one group, Tayassuidae. The areas of greatest density are in the central portions of Ohio but secondary areas of density appear across the south and in the northeast. The greater density areas result from multiple skeletons occurring proximal to each other. In fact, in this database it was very common to have multiple occurrences of peccaries at the same locality. Despite the extensive geographic area covered by Tayassuidae, certain habitat types seem to be preferred. Research describes open, arid, grassy habitats with scattered trees as typical habitat for Platygonus spp. (Gasparini, 2013; Gasparini & Ubilla, 2011; Smith and Polly, 2013). Other evidence suggested they frequent a greater diversity of habitats such as mangroves,

108 89 marsh, woodlands, and even dense canopied forest (MacFadden et. al., 2010). The results here support the latter assessment. The areas where Tayassuidae were found in Ohio were not arid, tropical, or likely to be heavily forested. Apparently within Ohio, peccaries frequented many diverse habitats. However, they do all have one common element that agrees with most peccary research: marginal and coastal water environs. Globally, peccary fossils are regularly located in river lag deposits, beaches, creek beds, and coastal deposits from deltaic to nearshore (Czaplewski, 2012, Frailey & Campbell, 2012, MacFadden et. al., 2010; MacFadden et. al., 2012). Nearly every peccary find within this database is from some type of deposit proximal to a former source of moving water. The pattern of density follows former stream valleys and lake margins. Most peccary density follows the valley of an ancient Teays River tributary. The unconnected density area in the northeast of Ohio follows a similar pattern along the present-day Mahoning River drainage and sources closest to the present Lake Erie are from ancient beach deposits. Spatial Distribution of Paleohumans Paleohuman archeological sites are scattered over most of Ohio, though the plateau regions appear to have the fewest sites overall (Figure 44). Unlike all other taxa, paleohuman sites are abundant over the lake plain area in the state. There are no radiocarbon dates from lake plain sites, so the time of occupation is limited only by when the lake plain first became habitable as proto-lake Erie retreated north. The general trend of human occupation mimics that of most other taxa, from northeast to southwest (Figure 45). However, the range is generally more northern than other taxa and taxa groups. This may result from the inclusion of sites that do not qualify as paleosites in the paleontological sense.

109 90 Considering all paleohuman occupations sites together, density was greatest in the northcentral to central portion of Ohio, with density rapidly falling off to the east and decreasing gradually to the west (Figure 50). As this layer included sites that were not paleo by the geologic definition, a density layer of all radiocarbon-dated sites was created (Figure 51). When the two density layers are compared, the dated sites density layer is similar to that of total sites density pattern but with less land area coverage. Temporal slices of radiocarbon-dated sites reveal changes in paleohuman density across Ohio, probably reflecting shifts in total range and increasing populations, as the northeast portion of the state is consistently occupied in all density layers. The implied geographic range of the earliest dated human sites within Ohio (from 15,460-14,030 cal yr) is small and located in the northeastern portion of the state on the glaciated plateau (Figure 46, 52). It is possible older western site(s) may still be discovered. It is also possible that historic land development could have destroyed any evidence of older sites, if indeed they existed. The western portion of Ohio was largely glaciated and probably less hospitable for paleohuman occupation. The plateau regions to the east and southeast of the state were areas most available for habitation. The oldest paleohuman site is just north of the furthest reaches of the Grand River and Killibuck glacial lobes. When considering the temporal range of 13,840-12,950 cal yr, the paleohuman range moves south and westward following the glacial ice margin and still within the plateau regions (Figures 47, 53). However, the northeast area of density is still significantly high. This shift could indicate a population increase and subsequent range increase rather than just a range shift. By the 12,900-12,000 cal yr time slice, humans had shifted northward onto the glacial till plain and lake plain (Figures 48, 54). This pattern matches the timing of glacial retreat, as

110 91 according to Glover et al. (2011), the area was unglaciated by 14.6 ka. The center of Ohio has no area of density but there is an additional area of density to the far southwest border signifying another extension in overall paleohuman range. This may signify an abandonment of central Ohio for the lake plain or perhaps a migration westward following the habitable zones revealed as the ice retreated. The pattern of human temporal ranges replicates the pattern of receding glacial lobes and may indicate the earliest time frames in which areas became habitable by humans. There are many more dated paleohuman sites within the latest 11,255-10,245 cal yr range and their spatial distribution is dispersed at a 90% confidence level. The human range within this temporal slice shifts southward, trending from the northeastern corner to the southwestern corner of the state and narrowing considerably (Figure 49, 55). This change could indicate that paleohumans became decoupled from the ice margin and diffused throughout the state, expanding to multiple localized areas of density. Perhaps at this time the paleohuman populations were sufficient to support multiple groups, or preferred prey ranges similarly shifted southward. Data seems to suggest the human presence within Ohio was linked to glacial retreat and the subsequent spread of habitable zones into formerly glaciated areas, at least prior to 11,255-10,245 cal yr range. Spatial Distribution of Faunal Associations The kernel density map for all proboscideans covers the entire state of Ohio, except a small portion of the extreme south (Figure 56). This data includes all identified and unidentified Proboscidae. It is most influenced by M. americanum, hardly surprising as mastodons comprise so great a proportion of the total sample. However, M. primigenius also contributes to the overall distribution of the density hot spots. Southwestern Ohio is the area of greatest occupation for

111 92 both groups. However, including mammoth influenced the total density of Proboscidae by moving centers of concentration to the east-southeast. As these two mammals are different in habit and have distinct associations with other taxa, it is best to consider them separately. Composition of the Mastodon Fauna was discussed within the text previously. The kernel density map for the Mastodon Fauna is similar to that of mastodon alone (Figure 57). The only real difference is the dilution of intensity of strong density centers, except the extreme southwest area and the one directly to its north. Those two occupation centers were marginally strengthened in concentration. The close match between the Mastodon Fauna and mastodons alone results from the proportionally larger representation of M. americanum within the Mastodon Fauna when compared to the other member species, C. ohioensis and C. scotti. Density of occurrences resides mostly across the till plains, with lesser areas to the east and south onto the glaciated and unglaciated plateaus. All taxa associated with the Mastodon Fauna tend to utilize the wetter lowlands of Ohio, so these results are consistent with previously published data from other regions. The kernel density map for the Mammoth Fauna was similarly affected by the proportionally greater number of included M. primigenius fossils compared to the other member species, Bison spp., Equus spp., and R. tarandus (Figure 58). However, the spatial distribution of the Mammoth Fauna is clearly different from that of mammoth alone. In both, the greatest density is in the south. However within the inclusion of other Mammoth Fauna members, overall density shifts northward and slightly west. This shift pulls the Mammoth Fauna density away from the higher elevations of the plateaus and more onto the till plains of Ohio. This pattern makes sense, as other Mammoth Fauna members are not known to prefer higher elevations as mammoth do, but frequent the more flat till plains.

112 93 When comparing the Mastodon and Mammoth Faunas, there is considerable overlap of ranges within southwest Ohio. The Mastodon Faunal range is broader over the state from northwest to southeast. The Mammoth Fauna has a greater range to the northeast and avoids the lake plain entirely. Instances of Bison spp. and Equus spp. occurring near the lake shore to the north of Ohio influence the density of Mammoth Fauna, forcing it northwest when contrasted to woolly mammoth (M. primigenius) density alone. Perhaps this signifies Mastodon Fauna have a greater habitat connection than Mammoth Fauna. To understand the degree to which the hoofed members of the Mastodon and Mammoth Faunas may influence overall density, a Hoofed Mammal Fauna density layer was constructed. This layer did not include only members of either faunal association but, rather incorporated all hoofed mammals in the database. I thought this acceptable because both Mammoth and Mastodon Fauna do share considerable range. Additionally, all other hoofed mammals, such as peccaries and muskoxen, occurred within both ranges. The hoofed mammal association is most concentrated within the center of Ohio (Figure 59). The density is very similar to that of all taxa without Proboscidae. Both have areas of greatest density in the central portion of Ohio and secondary ones to the southwest, northcentral, and northeast. However, there are distinct differences. The Hoofed Mammal Fauna largely avoided the unglaciated plateau region. There are some areas of low density but most of that portion of Ohio is bare. The entire lake plain is also empty, with no hoofed mammal occurrence. The greatest density occurs within the central portion of the till plain. Hoofed mammal density increases more to the southwest than that of all taxa excluding Proboscidae. As the Mammoth Fauna includes three hoofed mammals, Bison spp., Equus spp., and R. tarandus, it makes sense that this group has a density substantially different from that of mammoth alone. The Mastodon Fauna, on the other hand, has only one

113 94 hoofed member, C. scotti, and their distribution is therefore little different from that of mastodons alone. Raup-Crick Cluster Analyses The first dendrogram created used all available data from the OPMDB to cluster taxa by the similarity of their spatial occurrences (Figure 60). I hypothesized this dendrogram would both categorize each taxon as a group with branching patterns suggesting links between taxa. Taxa were generally sorted into distinct groupings (Table 10). When these groups and clusters were plotted as directional deviation ellipses, closely linked dendrogram groups and clusters had largely overlapping ranges. Hence, associations defined by cluster analysis are significant in their taxonomic and spatial context. Cluster analysis linked Equus spp. to O. moschatus, and Bison spp. to Megatherium sp. Mammuthus spp. have also been suggestively linked to Caprinae and Pilosa (Boeskrorov, 2006; McDonald & Pelikan, 2006). These connections may indicate a common ecological context for the Mammoth Fauna, muskoxen, and ground sloths Other groupings could denote different associations as well. The connection between Carnivora and C. elaphus within this analysis could result from a predator-prey relationship. R. tarandus was linked with Tapirus sp. in this analysis. This connection is less explainable but Tapirus sp. has only two occurrences within the OPMDB. It is possible any associations between Tapirus sp. and other tax are spurious. Bison spp. were similarly linked to M nasutus. Like Tapirus sp., M. nasutus is very poorly represented and therefore, this association may be spurious. Indeed, O. moschatus, Megatherium sp., Carnivora, Tapirus sp., and M. nasutus all have very poor representation within the dataset. The paucity of data could have created these implied relationships.

114 95 The second dendrogram used simplified site and taxon data (Figure 67). I hypothesized this analysis should show the distinction between Mammoth Fauna and Mastodon Fauna, as well as the connections between individual taxa. Clusters of taxa were assumed to suggest faunal associations. Two taxa separated into groups containing 100% of a taxon, M. americanum and S. cavifrons (Table 11). However, most taxa were in some fashion combined with two or more taxa into a group. M. americanum tended to dominate all results and was most often linked with fellow proboscideans. However, mastodons were also decisively combined with fellow Mastodon Fauna members, C. ohioensis and to a lesser extent, C. scotti. Interestingly, M. americanum was also grouped with Equus spp., S. cavifrons, and Tayassuidae. This cluster could result from the wide distribution of mastodons; there will likely be associations between mastodons and other taxa that are not strictly part of the Mastodon Fauna. Furthermore, the OPMDB is time averaged. Therefore, the apparent association between Equus spp. and M. americanum could result from temporal overlap and not range sharing. Tayassuidae do not have a definitive faunal association but it may be they are more likely to appear with M. americanum versus M. primigenius. Mastodon Fauna members were often associated with each other. C. scotti is grouped with C. ohioensis and M. americanum. C. ohioensis has strong connections to M. americanum and tentatively to Tayassuidae. Tayassuidae were associated with the Mastodon Fauna even when M. americanum occurrence is low, which may signify peccaries in Ohio are more connected to mastodon habitat than mammoth habitat. M. primigenius were only linked with other proboscideans. Members of the Mammoth Fauna were linked together but not including M. primigenius. Bison sp., Equus sp., and R. tarandus were all distinctly grouped together as a separate group. Further, O. moschatus was

115 96 included as was C. elaphus, though to a much smaller degree. Muskoxen have been related to the Mammoth Fauna in fossil associations and in Europe, Mammuthus spp. are linked to larger herding deer such as Megaloceros sp. (Boeskrorov, 2006; Rivals, et al., 2010). Therefore, it is probable the dendrogram relationships are actual associations within Ohio as well. Bison spp. and R. tarandus are only associated with the Mammoth Fauna. Equus spp. are also linked with Tayassuidae and to a lesser degree, M. americanum. This pattern could result from a couple of factors. Equus spp. could be more flexible in habitat than other Mammoth Fauna members. Equus spp. are known to have lived along marginal beach environments in the past, much like those favored by Ohio Tayassuidae, and therefore, the connection between horse and peccary could signify habitat sharing (Harington, 2011; MacFadden, et. al., 2012; Zazula et al., 2009). The co-occurrence with M. americanum could relate to the larger fossil record of the mastodon (i.e., many sites could be near each other just by chance), or result from temporal changes in the Ohio fauna. Intriguingly while O. moschatus were grouped with Mammoth Fauna, S. cavifrons were not. It is not immediately clear why this may be the case. Perhaps they had very different habitat requirements or the paucity of the muskoxen record is creating division where it does not exist. The Mammoth Fauna were also grouped with Tapirus spp. though it is uncertain as to why this association appears. In all dendrograms created, Tapirus spp. associations varied. This variability may result from the poor representation of Tapirus spp. in the dataset. The third dendrogram created used site and taxon data excluding M. americanum since mastodons are overwhelmingly represented within the dataset (Figure 68). I hypothesized this would show closer relationships between taxa without the influence of the rich mastodon record. The dataset without M. americanum is considerably smaller, so Megatherium sp. and M. nasutus were included in the analysis as separate entities instead of lumping them with other Pilosa and

116 97 Tayassuidae, respectively. Bison taxa were also separated into B. bison and B. latifrons. Carnivora was separated into Ursus americanus and Arctodus simus. Many dendrogram clusters consisted of a single taxon (Table 12). However, some taxa were split into groups that were not composed of single taxon. C. elaphus and M. jeffersonii were grouped together. Both taxa have tenuous connections to Mammuthus spp. according to other research, although analyses presented here finds M. jeffersonii more closely linked to M. americanum. Possibly, the connection between C. elaphus and M. jeffersonii is reflecting connections to of both to mammoths. O. moschatus and U. americanus were connected as were B. bison and A. simus in this cluster analysis. Why this is the case is uncertain. Both U. americanus and A. simus have very poor representation in the OPMDB, which may result in relationships that are insupportable as they are sample biased. Finally, R. tarandus, M. nasutus, Megatherium sp., and B. latifrons were linked together. R. tarandus and B. latifrons are in the Mammoth Fauna. Megatherium sp. is another ground sloth, like M. jeffersonii, and therefore, may also be tentatively linked to Mammuthus spp. Peccaries, like M. nasutus, have been related to Equus spp. as a member of the Mammoth Fauna. Hence these taxa all have more or less tenuous connections to the Mammoth Fauna. Paleohuman-Mammal Associations Revealing associations between paleohumans and other large mammal taxa can assist in determining likely prey species for Homo sapiens. As some researchers relate extinction to overhunting by paleohumans of Pleistocene large mammal taxa, such associations could indicate the role of humans in this context. Therefore, probable human range, i.e., kernel density and directional or standard ellipses, was compared to those of possible hunting targets and to taxonomic associations.

117 98 Mastodon (M. americanum) extinction is frequently attributed to paleohumans and their spatial distribution maps show considerable overlap, especially within the southwestern portion of the paleohuman range (Figure 69). However, the paleohuman southern range limits are largely driven by radiocarbon dated sites from 11,255-10,245 cal yr temporal range. A temporally similar M. americanum has not yet been found in the state. Much of the Ohio M. americanum material has not been radiocarbon dated so it may be that young specimens do exist. The latest persisting mastodon dated thus far in the region is from Indiana and dated to 11,515 cal yr and 11,500 cal yr (Woodman and Athfield, 2009). Therefore, it would seem that paleohumans range and the greatest concentration of mastodons have little correlation. Comparing the Mastodon Fauna with probable human range reveals similar pattern of density within the paleohuman range although concentrations differ (Figure 70). Examining other members of Mastodon Fauna separately was necessary as M. americanum were likely not the prey available in the area of greatest overlap between the faunal assemblage and humans. C. ohioensis density and paleohuman range have considerable overlap though the area of greatest density occurred west of the human range (Figure 74). Interaction between C. ohioensis and paleohumans are inconclusive. Beaver bones have been found in association with paleohuman sites but no evidence of predation/scavenging behavior by humans exists on the bones themselves (Miller, 1982). C. scotti density and human range have even more overlap between them (Figure 75). Areas of greatest concentrations are within and immediately adjacent to the paleohuman range. However, like C. ohioensis, there is no evidence paleohumans preyed upon or even scavenged C. scotti, though bones and tools have been found in association at Sheriden Caves, Ohio (Long & Yahnke, 2011). ). Certainly, C. ohioensis and C. scotti would be attractive as a source of meat and pelts to early humans.

118 99 Woolly mammoth (M. primigenius) density and paleohuman range overlap in the greatest concentration within the southern border of the human range (Figure 71). As with mastodon, the overlap may be misleading, as southern paleohuman sites are also the youngest radiocarbon dated locations. The youngest Mammuthus spp. from the Great Lakes region date to 13,500 to 13,000 cal yr (Boeskrorov, 2006; Harrington, 2011; Zazula, 2009). Comparing Mammoth Fauna density and paleohuman range expands the area of overlap, although the concentration decreases slightly (Figure 72). When each member of the Mammoth Fauna is examined, the influence of each taxon on the whole association can be seen as well as their range interaction with the paleohuman range. Equus spp. kernel density and paleohuman range have significant overlap; the greatest concentration of Equus spp. occurred midrange for humans (Figure 76). There is an additional area of slightly lesser concentration in the southwest corner of the state. If this figure is compared to Figure 55 for paleohuman sites dated 11,255-10,245 cal yr, the location of high Equus spp. density is near an area of kernel density concentration for paleohumans. Horses were common prey for paleohumans globally (Jass et al., 2011). It is possible paleohumans, especially later humans, preyed upon Equus spp. as their ranges and density distributions are similar. B. bison density and apparent paleohuman range have some overlap, especially to the north (Figure 77). Although much of B. bison density remains on the margins of paleohuman range, bison have also been common food items for native peoples throughout prehistory into historical times. It seems probable Equus spp. and B. bison were on the menu for Ohio paleohumans. One other taxon has a notable interaction with the apparent paleohuman range, P. compressus. When flat-headed peccary density is overlain with the paleohuman range, much of the peccary s range is within it (Figure 78). Areas that appear outside the human range are

119 100 congruent with areas of density for paleohumans within different temporal ranges. This pattern is suggestive of a connection between P. compressus as prey and H. sapiens as predator or scavenger. The relative frequency of peccaries within the paleohuman range bears this idea out (Figure 79). As in nearly all taxon analyses, M. americanum dominates. However, P. compressus and M. primigenius are tied for third with 10.13%, behind the second most common group, unidentified Proboscidae with 10.57%. No other taxon is close to this value. Hence, it seems reasonable to propose that the peccary P. compressus may be a preferred hunting target, at least at some time in paleohuman history. The relative frequencies of taxa found within the paleohuman occupation ellipse for each defined time slice were also calculated. From 15,460-14,030 cal yr time range, the dominating taxon is M. americanum (Figure 80A). This is more significant when considering only three taxa categories appear and two of them, M. americanum and C. scotti, are members of the Mastodon Fauna. There is no evidence that elk-moose were preferred prey but this preponderance of the Mastodon Fauna is likely significant. It seems reasonable to suggest paleohumans of this time were utilizing one or more members of the Mastodon Fauna. However, only two paleohuman sites exist for this time range, so any conclusion must be tentative. More sites are dated to 13,840-12,950 cal yr and a corresponding increase in the diversity of available prey taxa occurs (Figure 80B). Both Mastodon and Mammoth Faunas are significant proportions of the total. M. americanum and the Mastodon Fauna are dominant but M. primigenius is at its highest percentage and other members of the Mammoth Fauna are present. It is probable that humans were utilizing both Mastodon and Mammoth Fauna as prey items. P. compressus and M. jeffersonii also appear. Ground sloths, like M. jeffersonii, have not yet been documented as prey items (Borrero & Martin, 2012). However, P. compressus and other

120 101 peccaries are notable food sources in many American paleosites and likely qualify as prey for this time range. The time range of 12,900-12,000 cal yr has the greatest variety of taxa available (Figure 81A). The percentage of available M. americanum and M. primigenius both decreased from the prior time range, while Mastodon and Mammoth Fauna numbers held steady. These results imply the population of mastodon and mammoth were falling but their associated fauna were not similarly in decline. The falling percentage of mastodons and mammoths are small. This could be an actual decrease or it could be the result of biases in sampling and processing. There are few dates for large fauna so temporal comparisons are naturally suspect. The M. jeffersonii percent is unchanged from the 13,840-12,950 cal yr time range but P. compressus doubled from 2% to 4% and an additional peccary species, M. nasutus, appeared. The increase in peccary occurrence signifies increasing overlap of peccary and paleohuman range. It may be that peccaries were becoming more of a food preference but whether that is due to decreases in mastodons, mammoths, and associated faunas is not clear. There are many sites from 11,255-10,245 cal yr but taxonomic diversity decreased from 12,900-12,000 cal yr (Figure 81B). The highest percentage of Mammoth Fauna and second highest relative frequency of M. primigenius are within this time slice. However, it is very unlikely M. primigenius was a prey source, though other associated fauna may have been, as mammoth are unknown from this time period in Ohio. M. americanum and associated fauna are at their lowest percentage of all time ranges. When these results are compared to those of previous time segments, it is suggestive that the mastodons, mammoths, and associated fauna populations are decreasing, or at least shifting in range. The paleohuman range of 11,255-10,245 cal yr is narrow but still covers a considerable portion of Ohio so it is not likely that a smaller

121 102 human range is the reason for diminished mastodon/mammoth returns. M. jeffersonii does not occur within the human range. However, the P. compressus percentage has continued to increase and peccaries are now 17% of the total number of mammal occurrences found within the paleohuman extent; however, M. nasutus was not found. The paleohuman range has shifted southward and narrowed in this interval, yet the P. compressus percentage increased, suggesting a prey preference for this taxon. M. nasutus is not common in the database so its absence is likely an artifact. Alternatively, P. compressus and other peccary species often exchanged dominance in regions, as P. compressus preferred drier areas than peccaries like M. nasutus (Gasparini, 2013; Gasparini & Ubilla, 2011). Correlations between the kernel density rasters of mammal and paleohuman occurrences also suggest relationships between paleohumans and other taxa (Table 13). Paleohumans correlate well with Mastodon and Mammoth Fauna groups but are not equally associated with M. americanum and M. primigenius. M. americanum was second in rank order but M. primigenius was fifth. This implies that early humans were closely linked with these faunal groups but not necessarily to M. americanum and M. primigenius alone. There are many more representatives of M. americanum and M. primigenius in the OPMDB than any other taxon. This naturally biases the results in favor of these two taxa. However, correlations must not be entirely dependent on the total number of occurrence because the fourth ranked group, B. bison, has fewer representatives than those of lower ranked taxa such as C. scotti, C. ohioensis, and others. Mastodons ranked second in correlation to the paleohuman occurrences. As the other Mastodon Fauna members are ranked much lower, it is probable the high ranking of the group is related to M. americanum. Mastodons are so dominant in the OPMDB, this rank is certainly influenced by the total occurrence of the taxon. When the paleohuman data set was divided into

122 103 temporal slices, M. americanum consistently places in the top fifth in all analyses (Tables 14-17). The consistent abundance of M. americanum may result from the high percentage of total finds. The Mastodon Fauna contain two members not known to be utilized as prey by paleohumans, C. scotti and C. ohioensis. C. scotti ranks ninth in correlation to paleohumans and between R. tarandus, eighth, and Equus spp., tenth. While there is no direct evidence as yet that elk-moose were part of the paleohuman diet, considering its placement between caribou and horse as well as its correlative rank, this taxon was certainly available and likely on the menu. Furthermore when examining different time slices, C. scotti ranks first in the 15,460-14,030 cal yr and 13,840-12,950 cal yr time frames, sixth in the 12,900-12,000 cal yr time, and third in the 11,255-10,245 cal yr. C. ohioensis ranks low, eleventh, in the order of correlations with paleohuman occurrences. This low correlation is plausible as the taxon has been determined to live in environments less suitable for human habitation. When all time ranges are examined, C. ohioensis never ranks above sixth in correlation, and most often falls below the top ten. The Mammoth Fauna has the third highest correlation with paleohuman occupation. Members of the Mammoth Fauna vary in their ranking: M. primigenius is fifth ranked, B. bison is fourth ranked, and Equus spp. is tenth ranked. The rank order of M. primigenius fluctuates little in all temporal analyses, ranking third in 15,460-14,030 cal yr, fifth in 13,840-12,950 cal yr, and seventh in both 12,900-12,000 cal yr and 11,255-10,245 cal yr time intervals. By the later two temporal ranges, the presence of mammoth in Ohio is suspect. It is unlikely M. primigenius remained in the state during this time. Mammoth remains are the second most frequently found Quaternary fossils in Ohio. Certainly this contributes to some of the total abundance within the analyses. However many other taxa with fewer numbers are ranked higher than mammoth in all

123 104 analyses, temporal and pooled. It may be that, like mastodons, the consistent abundance of M. primigenius may result from the high percentage of total fossil finds. This cannot be the case for B. bison, Equus spp., and R. tarandus as their fossil find numbers are considerably lower. B. bison are fourth ranked in correlation with the paleohuman occupation range. When examined in separate time ranges, this is not the case. Generally, B. bison decrease in correlation through time. In the earliest time ranges, bison are negatively correlated with paleohumans. They do not become positively correlated until the 12,900-12,000 cal yr time segment when they rank eleventh and then become negatively correlated again in 11,255-10,245 cal yr. The reasons for the fluctuating rank order are not clear but may result from several key factors. The overall abundance of bison fossils is low and, furthermore, they are not temporally constrained. Additionally, the number of radiocarbon-dated paleohuman sites is 37 of the 533 data points, about 7%. That bison are so highly ranked overall indicates some significance between the taxon and paleohumans but it is not possible to determine changes in importance through time. Equus spp. ranked tenth in correlation with paleohuman occupation. Generally, there is a trend of increasing correlation through time between Equus spp. and paleohumans. This situation changes through time as Equus spp. is negatively correlated and ranked last in 15,460-14,030 cal yr segment, but positively correlated and ranked eleventh for 13,840-12,950 cal yr and 12,900-12,000 cal yr. Then in 11,255-10,245 cal yr time frame, Equus spp. ranked fifth. Like B. bison, changes in rank could result from a few causes. However, it is important to note that overall ranking and correlation increased through time. This increase could signify a growing importance in the paleohuman diet or perhaps an increase in Equus spp. population as habitat conditions favorably altered for horses. The decreasing B. bison correlation supports this. Research suggests that when Equus spp. dominate a landscape, B. bison decrease in prominence

124 105 and vice versa. The link between horses and paleohumans could also be the result of horse populations south of Ohio migrating northward and becoming increasingly utilized by paleohumans. R. tarandus correlation with paleohuman sites is ranked eighth overall. This ranking changes insignificantly through all time ranges. Unlike M. americanum and M. primigenius, the constant ranking is not likely due to overall abundance as total numbers of R. tarandus fossils are low. The consistency in rank probably indicates caribou and paleohumans at a minimum shared similar habitat. As R. tarandus has been a source of prey prehistorically, historically, and in the present, it seems likely this correlation signifies paleohumans were utilizing caribou in Ohio. Interestingly, the ground sloth M. jeffersonii is fifth ranked in association with paleohumans. As these are not common prey mammals for early humans, it is likely the environmental parameters that ground sloths favor are similar to those attractive to paleohumans. Ground sloths were consistently ranked through most of the time ranges except the last, 11,255-10,245 cal yr time interval, when they fall to thirteenth in rank. M. jeffersonii are not common fossils in Ohio and the latest time frame would be the right before the end-pleistocene extinction. Perhaps ground sloth populations were declining and this contributed to the drop in correlation between humans and sloths. Certainly paleohuman range shifted southward and away from areas where M. jeffersonii have been found. It may be that this range shift best explains the drop in correlation, rather than a decrease in ground sloth populations. P. compressus is sixth in correlation ranking with all paleohuman sites, but its rank changes when examined through time ranges. Initially it is negatively correlated and tenth in rank but increased to sixth and positively correlated in the next time range, 13,840-12,950 cal yr. By the 12,900-12,000 cal yr, this peccary is negatively correlated and slips to last. However, M.

125 106 nasutus is eighth in ranking. This time frame is the highest correlation for this peccary species. Research has shown that M. nasutus and P. compressus do not share territories (Gasparini, 2013; Gasparini & Ubilla, 2011). When P. compressus leaves an environment, other peccaries, such as M. nasutus, move into the area. While it is not conclusive, this evidence suggests M. nasutus, rather than P. compressus, was the prey species available during this temporal range. M. nasutus continues to dominate in the 11,255-10,245 cal yr time interval, placing ninth above the eleventh ranked P. compressus. There is one additional paleohuman site available from the 11,255-10,245 cal yr time slice than from 12,900-12,000 cal yr interval, but the spatial distribution of humans has decreased. It seems unlikely the additional site played an important role in increasing M. nasutus correlation when the overall paleohuman range decreased. Evidence instead suggests that as P. compressus moved from the area, M. nasutus immigrated into the abandoned habitat space and therefore, became the preferred prey of paleohumans in this time. Several mammal taxa showed strong correlations with each other, unrelated to paleohumans. P. compressus is correlated with Equus spp. Presumably, this may be an indication of habitat sharing. Both Equus spp. and P. compressus have some affinity for larger water bodies and this close correlation reflects this situation. Both peccaries and horses are common prey animals for paleohumans of the Pleistocene and the relationship between Equus spp. and P. compressus could indicate paleohumans utilized both taxa because of shared ranges. M. americanum and M. primigenius have a strong relationship as well. This is somewhat surprising as mastodons and mammoths reportedly have different diets and habitat preferences. These results could indicate that one or both taxa are more flexible in dietary requirements than assumed; in particular, evidence exists that M. americanum had greater flexibility in diet than previously thought. Research suggests that mastodons adapted to eat regionally abundant foods

126 107 (Teale & Miller, 2012). However, this correlation may also result from the sheer abundance of both taxa in the OPMDB. M. americanum and Equus spp. share a substantial correlation as well. This connection is interesting as Equus spp. are members of Mammoth Fauna and not Mastodon Fauna. It is further surprising as Equus spp. shares stronger correlations with other Mammoth Fauna and with M. americanum than with M. primigenius. Caprinae are poorly correlated with almost every taxon and taxonomic grouping but had a strong relationship with Equus spp. It is apparent from the data that the Mammoth Fauna has marginally more significant correlations between faunal members than the Mastodon Fauna has between its faunal members. Therefore, relationships between M. americanum and Mammoth Fauna taxa may indeed simply result from the overabundance of M. americanum in the dataset. R. tarandus and B. bison have a robust relationship that may be explained through their relationship as members of the Mammoth Fauna. R. tarandus and C. elaphus also have an equally robust connection. However, C. elaphus is not known to be a member of either the Mammoth or Mastodon Fauna, although large European deer have been linked to Mammuthus spp. C. elaphus has a correlation with C. scotti, which is marginally less than that it has with R. tarandus. Furthermore, R. tarandus has a higher correlation with C. scotti than with its fellow Mammoth Fauna members, except B. bison. All of these mammals have different habitat preferences. It is unclear what connection these taxa may share to produce such strong spatial correlations. M. jeffersonii are best correlated with M. americanum. Ground sloths have been associated with the Mammoth Fauna in previous research (McDonald & Pelikan, 2006; McDonald et al, 2000). However in this analysis, the taxon is best related to Mastodon Fauna

127 108 members M. americanum and C. ohioensis over any member of Mammoth Fauna. The relationship between M. jeffersonii and M. americanum could result from the larger population of mastodons in the database. The relationship between ground sloths and C. ohioensis is likely more complicated. The numbers of giant beaver are not significantly more than those of ground sloths in the database. Both animals are likely solitary, or at least members of small family groups but they do not share habitat preferences. Like the R. tarandus-c. elaphus-c. scotti relationships, there may be some connection between these taxa that is not immediately clear. Sedimentology Sedimentary data are sparse for all taxa and skeletal element groups in the OPMDB. However, all taxa are represented in some manner (Table 18). M. americanum dominates every sedimentological category, likely due to its predominance within the database. M. primigenius also occurs in nearly every category, but notably in greater quantity within the larger grain sizes. Most of the finds with sedimentological information are from marshes and swamps, stream banks, and sand and gravel pits. Marshes and swamps with their clay and peat deposition would tend to preserve material better than that of other sedimentological categories. Stream banks will naturally bias the record due to postmortem transport. Sand and gravel pits in Ohio are the result of ancient glaciofluvial channels and tills. Skeletal elements within each sedimentological category varied greatly and are suggestive of the environment in which they were deposited. Unidentified bones tend to compose a significant proportion of each sedimentological class. Sedimentology vs. Taxa M. primigenius appear at greatest abundance within gravel settings (Figure 82A). M. americanum dominate this class, as it does in most, but if mastodons were removed, the profusion of mammoth in gravel sites would be even more apparent. Most of the attendant taxa

128 109 within gravel settings are similarly members of the Mammoth Fauna. It is in this category that unidentified proboscideans occur the most. Likely this results because larger sediment size indicates increased water flow and bones that would distinguish between M. americanum and M. primigenius have been transported away. Alternatively, mammoth bones appear to be conserved in greater quantity within the larger sediment size groups, so perhaps some inference of taxon may be made based upon sediment type. M. primigenius purportedly prefer higher elevations than M. americanum. Higher elevations favor more energy in hydrologic systems. Perhaps unidentified proboscideans occurring in gravels and mixed sand settings are likely M. primigenius while those found in clay and peat are more likely to be M. americanum. More sediment data than currently available may refine this distinction. Sand and sand-mixed categories are similar in the mammal taxa they preserve. Tayassuidae composes 36% of sand category and 57% of sand-mixed (Figure 82B and Figure 82C). Tayassuidae have been associated with beach deposits in the fossil record. Some of the peccaries within the database were collected from ancient dune deposits, so this certainly appears consistent. It is possible that Tayassuidae that did not have depositional environment identified but did have sediment data consistent with sand and sand-mixed categories are also from beach deposits. A further collaborating fact is the amount of Equus spp. material found in these two categories. Globally, Equus spp. are frequently found on beach margins and shorelines of water bodies. Both sand and sand-mixed have the least amount of M. americanum occurrences. M. primigenius specimens do not appear within sand but do occur within sand-mixed in considerable quantity. Furthermore, most of the attendant taxa within both sedimentological categories are members of the Mammoth Fauna. It is probable these two sedimentological

129 110 classes represent open areas, perhaps grasslands marginal to significant bodies of water, rather than small streams. It may also be that some of the areas are floodplain. Very few occurrences are documented from silt settings, and perhaps unsurprisingly then, this sedimentological category is unusual in its dominant taxa (Figure 82D). M. primigenius and Tayassuidae appear in large quantity. With such a proportionally large percentage of M. primigenius, it may be assumed this would be a habitat favorable to mammoth. Additionally if M. americanum were removed from consideration, the assemblage would appear to be primarily Mammoth Fauna. As mastodons dominate nearly every category, they may be discounted somewhat in this case. However, the appearance of C. ohioensis is noteworthy. C. ohioensis consistently appears within sediment classes associated with slower moving, but not stagnant, bodies of water. Giant beaver behavior cannot be determined definitively but it has been proposed they behaved much like muskrats (Swinehart & Richards, 2001). The sedimentary data in the OPMDB supports this conclusion. Peat was the most diverse of all sediment classes with ten taxa (Figure 82B). The occurrences for peat were all from swamps and marshes within Ohio, particularly from the western half of the state. Linked to peat deposition were clays (Figure 82A). Generally, the clay sedimentological class contains sites from marshes and swamps. Very few incidences were clay without an attendant wetland component. As a result, I will discuss them together. Peat and clay have the highest preservational potential of all the sedimentological categories. This is one reason why these classes are so dominated by mastodon. If the dominant taxa lived within the wetland environments, the potential for preservation increases further. Both peat and clay contain the largest percentage of the Mastodon Fauna assemblage. This makes sense as the conditions to produce these classes are preferred habitat by this

130 111 assemblage. M. americanum, C. ohioensis, and Cervidae all occur in quantity. Cervidae here includes both Cervalces scotti and Cervus elaphus. However, only one C. elaphus, in clay, had sedimentological data so it contributes little to the total Cervidae category. Notably, the giant beaver C. ohioensis does not occur in clay. This is unexpected as they are prominent members of peat class and are considered part of the Mastodon Fauna. The absence of beaver from clay deposits could be a consequence of their preferred water conditions. Additionally surprising, Pilosa appear within these categories in the greatest quantity. Research suggests these mammals to be more akin to mammoths in their habitat preferences and less likely to live within the wetter areas that produced the peat and clay deposits. These results would seem to suggest otherwise although there are few occurrences so this could be a sampling bias related to the enhanced preservational potential of peat and clay. Tapirus sp. appear only in clays. It is difficult to draw a meaningful conclusion from this fact as the group is so scantly represented in the database. They could occur in clays as a result of a preference for warmer and more humid settings. As Tapirus sp. sites are on the extreme east of Ohio, perhaps climate conditions were different from those in the western part of the state. The mixed sedimentology category also contains a diverse assemblage (Figure 82C). However, it is probable this results from the characteristic nature of the class. It is the most variable of all classes, containing everything from cave deposits to coal basin deposits. Yet the analysis is still valuable since most of the category is composed of karst deposits such as those from Sheriden Cave, Wyandot County. Carnivora appear in greatest quantity in this class. They represent 33% of the total and perhaps tellingly, Cervidae represent 27%. This could be indicative of a predator trap situation. Furthermore, other species from Sheriden Cave are prey species such as R. tarandus and P. compressus, both at 7%. Occurrences of ground sloth,

131 112 mastodon, and mammoth within the mixed sedimentology category are all ones with other circumstances, like coal basin deposits. Sedimentology vs. Skeletal Element Teeth and tusks are nearly equal in proportion and compose over half of all skeletal finds in gravel settings (Figure 84A). Given the parameters discussed previously about transport within a fluvial system, this makes sense. Similarly, the related Voorhies element groups are conserved. Crania, mandibles, and horns-horn cores appear likely grouped. The proportion of mandibles was small and there is a small component of partial skeletons. Additionally, whole skeletons do occur within gravels. Both of these are single occurrence of the dominant taxon M. americanum. One of the specimens was from a gravel pit and the other from gravel within a swamp. The second is related to glacial deposition but the precise depositional environment of former is less clear. When skeletal element occurrences within the sand and sand-mixed classes are compared, they have very different components. Sands have only a few more robust elements, except for the notable occurrence of 33% whole skeletons (Figure 84B). Mandibles, teeth, and tusks are the majority of all finds in sand. This may be explainable if this category represented depositional environments near a larger body of water such as a beach. A majority of the assemblage could be from elements transported by wave action onto shore. The large number of whole skeletons comes from reported dune deposits and represents a single taxon, P. compressus. Modern peccaries travel in small groups. It is therefore likely these animals comprise a small group that were killed or trapped in some fashion near a shoreline. Perhaps a dune collapsed upon them and the animals asphyxiated.

132 113 The sand-mixed category has a greater diversity of skeletal elements (Figure 84C). With the exception of whole skeletons, the rest of the bone elements are the most robust of the skeleton. Tooth, tusk, ribs, mandibles, etc have similar transportation potential based upon Voorhies group analysis. It is probable this sedimentological category also represents deposition marginal to a larger body of water, or perhaps a river system with attending floodplains and deltas rather than a lake margin. Of the elements found within silts, most are more robust bones less likely to be transported great distances (Figure 84D). Most of the elements are ones that require higher energy to transport, i.e. teeth, tusk, and long bones, or ones which tend to disarticulate from each other late in taphonomic processes, i. e., vertebrae and ribs, cranium and mandible. However, there is also one partial skeleton recovered from silt. This skeleton is a single specimen of C. ohioensis. When considered with the high percentage of isolated C. ohioensis skeletal elements finds within silt, this suggests a preferred environment of slowly moving or circulating water, and explains the apparent lack of the large beaver within categories preserving other members of the Mastodon Fauna. Peat and clay have the most diverse types of bone elements (Figure 85B and 85A). Additionally, the two categories have high percentages of partial and whole skeleton occurrences. This likely result from several factors, chief among them that preferred mastodon habitat is likely to contain these two sediment classes. Additionally, properties of these sediments also promote preservation. Clay deposition requires slow-moving to stagnant water conditions. Peat develops when small ponds and lakes begin to fill with vegetation until no open water exists. As the water bodies fill, little mixing will result within the water column. Conditions become increasingly anaerobic within the area, promoting the preservation of fossil

133 114 material by slowing bacterial decay. Furthermore, marshy settings may discourage active scavenging and therefore whole and partial skeletons, as well as small bone elements, are preserved in situ. Furthermore, the sticky, slippery clay found in these marshy areas prevents escape by animals that wander into boggy settings. Essentially, skeletal elements become stuck in place with little to no transport, abrasion, or active weathering. Despite its diverse taxonomic assemblage, the mixed sedimentology category has very few skeletal element types preserved within it. Most skeletal elements are unidentified (Figure 85C). Teeth and tusks dominate the total. The tooth percentage is likely higher because of predator representation. Why there are more predator teeth than others is not as clear. It could be that many predators have high amounts of tooth breakage and loss when hunting and scavenging. Ribs and vertebrae are also significant proportions. Ribs and vertebrae are some of the last elements to disarticulate from a carcass, according to Hill and Behrensmeyer (1984). As the majority of sites within the mixed sedimentology class represent karst deposition, the preferential preservation of the above elements could be the result of a predator trap. The occurrence of manus/pes is a single occurrence of Megatherium sp. within mixed gravel, sand, and clay. This locality is not likely to have been a karst deposit, but the sedimentology was too vague to place this listing within another category. Most probably, the element is from a fluvial setting and was transported. Taphonomy Voorhies Group Analyses Voorhies analysis of the skeletal elements present within the database was hindered by some inconsistencies. Voorhies (1969) used coyote and sheep skeletons within his flume experiments. While some breeds of sheep do possess horns, none were referred to within

134 115 Voorhies results. Additionally, teeth disarticulated from the cranium and mandible were similarly not discussed in his study. All of these skeletal elements occur within the OPMDB. Furthermore, I also have a tusk category since there are proboscideans within the database. Presumably, all three elements would be linked in some fashion to crania and mandibles. Cluster analyses (Figure 86) show that certain skeletal elements tend to co-occur. Groups colored brown, tan, green, and blue in Figure 86 are equally connected in all dendrograms but not all skeletal elements are grouped in the same order as Voorhies sequence. Clustering skeletal elements by similarities in their sedimentological context produces the most congruence with Voorhies groups. Voorhies (1969) treated elements as sediment particles within a fluvial system. Some of the skeletal elements within the OPMDB are likely in situ with no postmortem transport, which may explain differences in the clusters shown in Figure 86. Grouping skeletal elements by taxon does not produce clusters that match well with Voorhies groups. There are only two groups that match and, again, there are skeletal elements in the OPMDB that Voorhies did not consider. Interestingly, the two matching groups have skeletal elements that are consistent on all dendrograms but the groups are not clustered the same on all dendrograms. The skeletal element by taxon dendrogram resembles that of the sedimentology dendrogram to a greater degree than that of the Voorhies. This likely implies that the taphonomy of skeletal elements in this database is affected by transport factors and taxonomic identity in a similar fashion. The Voorhies dendrogram has two clusters; the crania, mandible, and miscellaneous bone in one cluster and another cluster containing all other bones. Only in this analysis are the skull bones separated to such a degree from postcranial material. This could mean that the skull complex behaves decidedly differently in fluvial systems than in any other depositional setting.

135 116 As the cranium and mandible are not connected within the other dendrograms but are closely grouped within the taxa dendrogram, this could imply taxonomic identity affects cranial preservation more than anticipated by Voorhies analyses (1969). As further suggestive evidence, the skeletal elements affecting clustering in the taxon and sedimentology dendrograms are the teeth, tusk, and horn core classes, all of which are skull components. Presumably, teeth would be linked to the cranium and mandible, and tusks would be linked to the cranium, as would the horn-horn core-antler category. When the analyses were completed, both taxon and sedimentology dendrograms clustered teeth proximal to mandible, cranium, and horn core classes. However, the sedimentology dendrogram shows mandibles generally less associated with other skull complex members than either the Voorhies or taxon dendrograms. As Voorhies (1969) did not use loose teeth in his experiment, it is hard to fully interpret the results of these cluster analyses. However, Hill and Behrensmeyer (1984) conducted disarticulation studies and noted teeth disarticulate after the cranium-mandible complex has separated from the animal. This would lead me to suspect taxonomic identity has less of a role than depositional environment and post-depositional factors. Tusks were unexpectedly inconsistent in clustering. The taxon dendrogram grouped tusks with ribs and linked them to shoulder and hip girdles and miscellaneous bones, while the sedimentology dendrogram placed tusks with hip girdles and linked them to teeth and mandibles. The taxonomic clusters could result from the low representation of taxa in the database, only two, that possess tusks. Horn core is also an underrepresented class resembling tusk, but they were grouped with crania and linked to teeth and mandibles in both dendrograms as expected, although less proximally in the sedimentology dendrogram. There are more horn/antler-bearing taxa than tusk-bearing taxa. Probably, tusks and horns are elements whose taphonomy is taxon

136 117 dependent and independent of sedimentology or depositional setting. This makes sense as both classes are largely composed of durable biological materials, although horn cores are somewhat more fragile. Two other skeletal element groups are dissimilarly clustered on all dendrograms, miscellaneous and rib elements. Miscellaneous element class is a mixed bone category and this explains some of the differences between dendrograms. The Voorhies dendrogram placed it in an individual class closely linked to the independent cluster of crania and mandibles. Both the sedimentology and taxonomic dendrograms have miscellaneous elements grouped differently. The sedimentology dendrogram categorized these bones separately but linked with long bones, mandibles, tusks, and hip girdles. The taxonomic dendrogram places miscellaneous with shoulder and hip girdle elements, which may imply a taxonomic component to preservation. Hip and shoulder girdles are rarer elements in the database. Vertebrae are similarly rare. I speculate that whether a vertebra persists long enough to be incorporated into sediments is more taxondependent but once the element is buried, it has equal or near equal preservation probability within all environments. Ribs were grouped with vertebrae on the Voorhies dendrogram and closely linked with hip girdles. This is explainable as Hill and Behrensmeyer (1984) reported the trunk-torso of mammals persisted intact longer than the limbs did. Vertebrae and ribs tend to disarticulate last and likely experience similar transport potential as a result. The sedimentology dendrogram associates ribs with vertebrae and mani/pes with links to shoulder girdles on one side and to long bones and mandibles on the other side. The taxon dendrogram places ribs with tusks and links them to shoulder and hip girdles, and miscellaneous. The connections of all three dendrograms are similar but not the same. Perhaps more than any other skeletal element class, shape

137 118 contributes substantially to rib preservation. Mandibles, shoulder and hip girdles, and ribs are all thinner, planar bones that could be preserved in like fashion. The connection with tusks is less certain, though they have comparable shape, which might be a factor. In all dendrograms, shoulder girdles, long bones, and manus/pes elements all appear similarly placed and closely linked. I would surmise that while these may not be the most common elements found but when they do occur as fossils, they are the skeletal elements least affected by specific taxonomic identity but most likely to represent depositional conditions consistent with Voorhies experiments (1969). Taphofacies Facies One, Ice Contact Deposition, corresponds to Cluster A (Figure 87). The bottom graphs depict each group as clearly dominated by a single robust skeletal element. Pooled skeletal data, Figure 88A, additionally display this as 71% of skeletal elements are teeth and tusks. When examining sediment data, the singleton nature of each group makes sense, as this facies has an explicitly defined sedimentological type within the study conducted by Miller (2009). Ice Contact Deposition is characterized by large percentages of gravel and very little percentages of smaller sediments (Figure 88B). Interestingly, this facies also has the largest percentage of peat, which was not expected. Facies Two, Proglacial Lake Deposition, is consistent with Cluster C. Each group is composed of a variety of bone elements, robust and fragile. Pooled skeletal element data indicate this facies has the best and greatest representation of all skeletal elements in near equal proportions within each group (Figure 89A). This makes sense as this sedimentology would represent fossil assemblages most apt to be in situ with little to no transportation of skeletal material. Proglacial lake deposition is characterized by sand-mixed, silt, and clay. These three

138 119 groups are characterized by a greater percentage of very fine sediments than any other cluster. The average clay content is 45%, sand- mixed and silt add a further 20% (Figure 89B). This facies had the second-greatest abundance of peat. Peat within this facies is largely associated with marls and clays. Facies Three, Late and Post Glacial Deposition, was determined to correspond to Cluster B. While a variety of fine and robust bone elements are preserved, each individual group only has a few element classes, in contrast to Cluster C (Facies Two). Pooled skeletal data depict these skeletal groups are also ones at least moderately transportable in fluvial situations as well as a high percentage of unidentified bones (Figure 88C). The large percentage of unidentified skeletal elements indicates the higher transport potential of this facies versus Facies One and Two. Late and Post Glacial Deposition consists of less gravel and more fine sediments than those of Facies One but coarser material than in Facies Two (Figure 88D). Facies Three has gravel but in a much lesser quantity, 22%, than Facies One. Facies Three also has clay and silt, 30% total, but less than Facies Two. The mix of sediment types in Facies Two is more even in percentage with no clearly dominant sediment type. The peat content of this facies is the lowest of all, with 16%. I initially hypothesized that Facies Two (Proglacial Lake Deposition) would have the greatest percentage of peat, with Facies Three (Late and Post Glacial Deposition) second and Facies One (Ice Contact Deposition) as likely to have the lowest percentage. After analyses were completed, however, Facies One had the greatest percentage with 24%, Facies Two was second with 20%, and Facies Three ranked last with 16%. When comparing all these values within one standard deviation, however, their differences are nonexistent, which may reflect the limited

139 120 amount of data. Alternatively, perhaps peat is not as important in glaciofacies as I hypothesized. I was unable to discover any facies research with peat as a component of the study. While the results of this study are not conclusive they do suggest that taphofacies analyses are meaningful for terrestrial settings. Perhaps with more data they may be as useful as those conducted in marine situations. In particular, incorporating the Ohio geology map (Figure 4) into spatial analyses would help pinpoint the associations between particular taxa or skeletal elements and the variety of glacial deposits found across the state.the analyses conducted here are temporally imprecise but regionally constrained. It may be that recent time periods, such as the Pleistocene, are not as subject to the vagaries of fossil preservation for terrestrial vertebrates encountered in the Permian and Cretaceous, and therefore this study should be seen as a rough template for further research. If definitive results are available with the rudimentary data available within the OPMDB, then this area of research in the Ohio Pleistocene would be a profitable avenue of exploration in other regions with similar glacial history such as other states from the Great Lakes region.

140 121 CHAPTER VI. CONCLUSIONS AND FUTURE AVENUES OF RESEARCH Fossil records are generally incomplete and, despite its recent nature, the Pleistocene record is similarly deficient. Scott (2010) proposed that a concerted effort needs be made to document and radiometrically date fauna, quantify abundances, and carefully account for taphonomic factors among faunal assemblages from dissimilar circumstances where possible in order to understand how the late Pleistocene fauna interacted and responded to changing environmental conditions. He asserted this would be a more constructive use of research rather than continuing to repeat established arguments with the aim to prove one side over the other (Scott, 2010). As a useful corollary, it would also behoove researchers to reexamine bones, especially mammoth bones fragmented and otherwise, for evidence of human alterations (Harington, 2011). The research presented in this thesis was conducted to add to the understanding of the Pleistocene fauna. My initial objective was to determine whether historical sources can be utilized to augment the paucity of Pleistocene vertebrate data for Ohio and to create a database, the Ohio Pleistocene Mammal Database (OPMDB), of probable Pleistocene-aged large mammals. There was some concern that the historical sources used to create the database would not be useful for scientific research. However, historical documents proved to be helpful and significant as sources of fossil research from time periods inadequately documented in the scientific record. The record of Ohio fossil occurrences has been expanded and collected into a database available for further research. My second objective was to demonstrate the relative completeness of the database. The OPMDB, despite the historic nature of its sources, has proved to produce results consistent with current data. It has been statistically analyzed and has yielded data pertaining to taxonomic,

141 122 taphonomic, and sedimentological patterns among Late Quaternary mammals in Ohio, data that are both consistent with prior research and reveal new avenues for further research. My third objective was to determine if the OPMDB depicted associations between Ohio Late Quaternary fauna consistent with other North American fossil assemblages in the fossil record as I hypothesized would be the case. For this objective, the data are mixed. Both cluster analyses and spatial analyses suggested associations and geographic ranges for Pleistocene age fauna that are congruent with previous studies, most particularly in intragroup relationships between Mastodon Fauna and Mammoth Fauna members. However, research has also suggested connections between taxa that have not been reported in other studies, which may result from the overall abundance of proboscideans in the database and the cosmopolitan nature of M. americanum, in particular, within Ohio. Alternatively, the Great Lakes area has been determined to be a refugium for Pleistocene species and associations within refugia may be different from those frequently appearing elsewhere. More research into the Great Lakes and other refugia may resolve some puzzling associations. This could be particularly important as alterations of climate create changes in habitat space and mapping previous responses to range shifts may aid in revealing faunal response in the future. One of the factors proposed in the end Pleistocene extinction has been over-hunting by paleohumans, most especially hunting of proboscideans. Yet, data on paleohuman-large fauna interaction in Ohio is sparse. I hypothesized for my fourth objective that paleohumans in Ohio did not depend upon greatly upon proboscideans as has been suggested in literature. Again as with my third objective, the results are mixed. Ohio s large vertebrate record was too poorly dated to produce definitive temporal patterns but geospatial patterns can be suggestive of relationships between predators and their prey. Range shifts of paleohumans and alterations in

142 123 prey availability within that range suggest that humans may have contributed in some part to the Pleistocene extinction in Ohio. There are suggestions of consistent range overlap between paleohumans and proboscideans that could have contributed to M. americanum and M. primigenius extinction in Ohio. However, there is also some suggestion that humans may have been more attached to habitat space rather than specific taxa. Additionally, it should be noted that the spatial extent of the paleohuman and taxa density rasters were not equal, which could influence the multivariate band collection, although the taxa layers were congruent with each other as were the paleohuman layers. Further avenues of research should include the expansion of the OPMDB with more sedimentological data for fossil occurrences already recorded within the database, more taphonomic data for skeletal elements, additional dates for fossil finds, and the integration of new fossil discoveries. Also, further spatial statistical analyses should be completed, perhaps on an even smaller regional level than the entire state of Ohio. The basic sedimentological and taphonomic (skeletal element types) data available from the OPMDB has produced promising results in vertebrate taphofacies research. Increasing the available data will help refine Pleistocene taphofacies models in particular but may also aid in improving the area of terrestrial taphofacies research more generally. As more faunal data are incorporated, greater temporal and geographical constraints will be placed upon relevant fauna, paleohumans, and their interactions during the latest Pleistocene.

143 124 Age (Ma) Age (Ma) CHAPTER VII. FIGURES AND TABLES SubChron/ Excursion Events Figure 1 Quaternary timescale as adapted from the Subcommission on Quaternary Stratigraphy. Time period of interest is outlined in red. Figure 2 North American glacial maximum as adapted from Dyke et al., Cordilleran ice sheet, Laurentide ice sheet, and Great Lakes region are labeled.

144 Figure 3 Glacial lobes into the Great Lakes region, from 125

145 Figure 4 Glacial map of Ohio, as published by the Ohio Department of Natural Resources, Division of Ohio Geologic Survey (2005). 126

146 127 Figure 5 Some of the oldest known radiocarbon dated sites within the Americas, as adapted from Faught, See Table 1 for key locality numbers. Table 1 Key to Americas map with reference numbers, sites, and average radiocarbon age. Table key to Americas map Reference Site Average ¹⁴C Age Reference Site Average ¹⁴C Age 1 Broken Mammoth CZ-4C Hedden Broken Mammoth CZ-4B Vail Broken Mammoth CZ-4A Page/Ladson Swan Point Aubrey Mead Blackwater Draw Walker Road Lubbock Lake Dry Creek Domebo Vermillion Lakes Murray Springs Charlie Lake Cave Lehner Ranch Lange/Ferguson Jake Bluff Oklahoma Anzick (Bone rods) Tequendama RS Anzick (Skeletal) Pedra Pintada Dent Lapa do Boquet Kanorado Kansas RS I Colby Cerro La China (Site 1) Indian Creek Cerro El Sombrero Mill Iron Monte Verde Hell Gap Quereo II Agate Basin Quereo I 11041

147 Lindenmeier Tagua Tagua Paisley Caves Quebrada Jaguay Fishbone Cave Quebrada Jaguay Bonneville Estates Rockshelter Quebrada Santa Julia Bonneville Estates Rockshelter Quebrada Tacahuay Marmes Rockshelter Casa del Minero Smith Creek Cave Piedra Museo Hiscock Cerro Tres Tetas Paleo Crossing Fell Cave Sheriden Cueva del Medio Sheriden Lago Sofía Shawnee-Minisink Tres Arroyos Debert Table 2 Skeletal element categories and explanation of contents within each category. Category Cranium upper palate Mandible Tooth Tusk Horn, horn core, antler Shoulder girdle Rib Vertebrae Hip girdle Long bone Manus/pes Misc Unidentified Skeleton Partial skeleton Items in category full skull or any portion of, save mandible mandible, with or without teeth tooth/teeth, any portion or type, unattached to mandible or cranium tusk, in portion or whole horn, horn core, and antler, in portion or whole any bone, or portion, of the shoulder anatomy any rib or portion any vertebrae or portion any bone, or portion, of the hip anatomy any long bone, or portion, of hind or forelimb; "leg bones" any bone, or portion, of manus or pes any bone which does not fit into other category any unnamed or unidentified bone, or portion near complete skeleton or whole skeleton partial skeleton and partial remains which are not included in skeleton category

148 129 Table 3 Sedimentology categories and description of sediment listings within each category. Category clay silt sand sand mixed gravel mixed peat uncategorized Items in category any predominately clay sedimentology including; "marl", "shelly marl", etc. without regard to coloration any predominately silt sedimentology without regard to color any predominately sand sedimentology without regard to color any sedimentology in which sand is a major component mixed with another such as clay, silt, etc. any predominately gravel sedimentology without regard to color or origin any mixed sedimentology including; "alluvium" without other qualifiers, diamicton, clay and sand, etc. any predominately peat sedimentology including; "bog", "boggy", "muck", etc. any sedimentology which does not suit any other category including: "strip mine", "coal basin", etc Table 4 Total fossil skeletal element content per sedimentology category. Fossils per Sedimentology Mixed Sedimentology 22 Peat sedimentology 70 Clay Sedimentology 92 Gravel Sedimentology 63 Sand Sedimentology 12 Sand Mixed Sedimentology 36 Silt Sedimentology 15 Total 310

149 Figure 6 State of Ohio digital elevation map within Ohio state border outline in white. Data available from USGS and compiled within ArcGIS

150 Figure 7 Polygon created by the intersection of the dated paleohuman site directional ellipse and all paleohuman site ellipse. Dated paleohuman site directional ellipse is upper left. All paleohuman site ellipse is upper right. Combined polygon is below. Central feature is white circle. Ohio outline in white. 131

151 132 Table 5 Skeletal element composition of Voorhies groups as defined by M. Voorhies (1969). Voorhies (1969) Groups based upon Transport in a Fluvial System Group 1 Groups 1 or 2 Group 2 Group 3 Ribs Phalanges Long bones Crania Vertebrae Scapula Pelvis Mandible Sternum Sacrum Ulna Taxon Ursus arctos americanus Ursus americanus Symbos cavifrons Rangifer tarandus Proboscidae Platygonus compressus Ovibos moschatus Mylohyus nasutus Megatherium sp. Megalonyx jeffersonii Mammuthus primigenius Mammut americanum Equus sp. Cervus elaphus Cervalces scotti Casteroides ohioensis Bootherium bombifrons Bison sylvestris Bison latifrons Bison bison Arctodus simus Individuals Figure 8 Number of individual listings per taxon from the Ohio Pleistocene Mammal Database (OPMDB). Excludes listings with inadequate data, as discussed in text.

152 133 Table 6 Statistical representation of taxa in the Ohio Pleistocene Mammal Database (OPMDB). Taxon # of Individual Listings % of Total Mode "giant" NA Homo sapiens (human) NA Arctodus simus (short-faced bear) C Bison bison (American bison) H Bison latifrons (giant broad-headed bison) H Bison sylvestris (syn. B. bison) H Bootherium bombifrons (woodland muskox) H Castoroides ohioensis (giant beaver) H Cervalces scotti (stag-moose) H Cervus elaphus (red deer / elk) H Equus sp. (horse) H Mammut americanum (American mastodon) H Mammuthus primigenius (woolly mammoth) H Megalonyx jeffersonii (Jefferson s ground sloth) H Megatherium sp. (giant ground sloth) H Mylohyus nasutus (long-nosed peccary) O Ovibos moschatus (muskox) H Platygonus compressus (flat-headed peccary) O Proboscidae (unspecified ) H Rangifer tarandus (caribou) H Symbos cavifrons (helmeted muskox) H Ursus americanus (black bear) C Ursus arctos americanus (American brown bear) C total # of species (w/o "giants" and human) 21

153 134 Table 7 Distribution of feeding modes for taxa in OPMDB. Mode of feeding # of taxa % of taxa C carnivore H herbivore O omnivore Total R.tarandus 1% Unidentified Proboscidea 10% Tayassuidae 5% Order Pilosa 2% Tapirus sp 0% Carnivora 1% Bison spp 2% Caprinae 2% Cervidae 4% Equus spp 2% C. ohioensis 3% M. primigenius 12% M. americanum 56% Figure 9 Taxon representation in the OPMDB.

154 135 Table 8 Richness and diversity indices for the OPMDB assemblage. Menhinick's R I Margalef s R I Berger-Parker I of D Abundance Rank Log Series alpha x chi² p (same) 2.53E-16 Figure 10 Whittaker plot and statistics for relative species abundance in the OPMDB.

155 136 Taxa (95% confidence) Specimens Figure 11 Rarefaction curve for OPMDB assemblage. Table 9 Total number of fossil specimens per skeletal element category. Element Number of Specimens Cranium-upper palate 35 Mandible 40 Tooth 158 Tusk 71 Horn, horn core, & antler 14 Shoulder girdle 10 Rib 30 Vertebrae 29 Hip girdle 9 Long bone 44 Manus/pes 13 Misc. 5 Skeleton 53 Partial skeleton 28 Unidentified 184 Total of taxa elements 723

156 137 Cranium-upper palate Mandible 5% 6% Unidentified 25% Tooth 22% Partial skeleton 4% Misc. 1% Skeleton 7% Tusk 10% Manus/pes 2% Long bone 6% Hip girdle 1% Rib 4% Vertebrae 4% Horn, horn core, & antler Shoulder girdle 2% 1% Figure 12 Relative frequency of skeletal elements in OPMDB. Unidentified Proboscidea 2% Tayassuidae 30% C. ohioensis 2% Cervidae 4% Tayassuidae 4% M. primigenus 4% Unidentified Proboscidea 14% C. ohioensis 14% Cervidae 7% Order Pilosa 4% M. primigenus 2% M. americanum 56% M. americanum 57% A B Figure 13 Relative frequency of whole skeletons (A) and partial skeletons (B) by taxon.

157 138 Unidentified 46% Cranium-upper palate 20% Mandible 7% Horn, horn core, & antler 40% Horn, horn core, & antler 13% Cranium-upper palate 60% Long bone 7% Vertebrae 7% A B Figure 14 Relative frequency of skeletal elements for Bison spp. (A) and for Subfamily Caprinae (B). Tooth 14% Unidentified 11% Cranium-upper palate 22% Partial skeleton 22% Mandible 11% Unidentified 86% Skeleton 6% Vertebrae 6% Tooth 22% A B Figure 15 Relative frequency of skeletal elements for Order Carnivora (A) and for Castoroides ohioensis (B).

158 139 Unidentified 23% Cranium-upper palate 13% Mandible 3% Unidentified 19% Mandible 31% Partial skeleton 6% Skeleton 7% Tooth 10% Horn, horn core, & antler 13% Long bone 6% Vertebrae 6% Manus/pes 3% Long bone 19% Vertebrae 3% Tooth 38% A A B Figure 16 Relative frequency of skeletal elements for Family Cervidae (A) and for Equus spp. (B). Cranium-upper palate 3% Cranium-upper palate 4% Unidentified 19% Mandible 6% Mandible 2% Partial skeleton 4% Skeleton 7% Tooth 27% Unidentified 32% Tooth 32% Misc. 1% Vertebrae 4% Long bone 6% Manus/pes 2% Hip girdle 1% Rib 5% Shoulder girdle 2% Tusk 13% Partial skeleton 1% Skeleton 1% Misc. 1% Long bone 1% Hip girdle 4% Vertebrae 5% Rib 6% Shoulder girdle 2% Tusk 9% A B Figure 17 Relative frequency of skeletal elements for Mammut americanum (A) and for Mammuthus primigenius (B).

159 140 Cranium-upper palate 2% Tooth 9% Mandible 1% Skeleton 12% Tooth 6% Shoulder girdle 6% Tusk 13% Misc. 12% Rib 12% Unidentified 54% Long bone 10% Rib 3% Vertebrae 1% Manus/pes 18% Vertebrae 17% Partial skeleton 6% Skeleton 1% Long bone 17% A B Figure 18 Relative frequency of skeletal elements for unidentified Proboscidea (A) and for Order Pilosa (B). Horn, horn core, & antler 29% Unidentified 71% Mandible 100% A B Figure 19 Relative frequency of skeletal elements for Rangifer tarandus (A) and for Tapirus sp. (B).

160 141 Mandible 5% Tooth 3% Long bone 3% Unidentified 40% Skeleton 46% Partial skeleton 3% Figure 20 Relative frequency of skeletal elements for Family Tayassuidae. Percentage of Elemental Occurrence Herding Non-herding Skeletal Element Figure 21 Relative frequency of skeletal elements for herding versus non-herding mammals.

161 142 Cranium-upper palate 4% Unidentified 26% Tooth 23% Mandible 5% Partial skeleton 8% Unidentified 17% Cranium-upper palate 13% Mandible 8% Partial skeleton 4% Skeleton 6% Tooth 12% Misc. 1% Manus/pes 2% Long bone 6% Hip girdle 1% Skeleton 7% Vertebrae 4% Rib 4% Shoulder girdle 1% Tusk 10% Horn, horn core, & antler 2% Misc. 4% Manus/pes 6% Long bone 6% Vertebrae 8% Rib 4% Shoulder girdle 2% Tusk 2% A B Figure 22 Relative frequency of skeletal elements for herding mammals (A) and for non-herding mammals (B). 7% 5% 30% 22% Mixed Sedimentology Peat sedimentology Gravel Sedimentology Sand Sedimentology Sand Mixed Sedimentology Silt Sedimentology Clay Sedimentology 12% 20% 4% Figure 23 Relative frequency of sedimentologies represented in the OPMDB

162 143 25% 8% 8% 17% Mixed Peat Gravel Sand Mixed Sand 47% 13% 7% 13% Mixed Peat Gravel Sand Mixed Sand 0% 42% Silt Clay 7% 13% Silt Clay A B 2% 21% 9% Mixed Peat 29% 9% Mixed Peat 2% 6% 24% Gravel Sand Gravel Sand 4% 34% Mixed Sand Silt Clay 7% 7% 7% 39% Mixed Sand Silt Clay C D Figure 24 Relative frequency of sedimentologies in which crania (A), mandibles (B), teeth (C), and tusks (D) were recovered

163 144 14% 15% Mixed Mixed 14% Peat Gravel Sand 50% 33% Peat Gravel Sand Mixed Sand Mixed Sand Silt Silt 57% Clay 17% Clay A B 10% Mixed 5% Mixed 40% 25% Peat Gravel Sand 48% 21% Peat Gravel Sand Mixed Sand 5% Mixed Sand 5% 15% 5% Silt Clay 5% 16% 0% Silt Clay C D Figure 25 Relative frequency of sedimentologies in which horns, horn cores, and antlers (A), shoulder girdles (B), ribs (C), and vertebrae (D) were recovered.

164 145 Mixed Mixed 38% 37% Peat Gravel 29% 38% Peat Gravel Sand Sand 13% 12% Mixed Sand Silt Clay 4% 8% 21% Mixed Sand Silt Clay A B 8% Mixed Peat 25% Gravel Sand 67% Mixed Sand Silt Clay C Figure 26 Relative frequency of sedimentologies in which hip girdles (A), long bones (B), and manus/pes (C) were recovered

165 146 25% 19% Mixed Peat 29% Mixed Peat 6% Gravel Sand 50% Gravel Sand 12% Mixed Sand Mixed Sand Silt 14% Silt 38% Clay 7% Clay A B Figure 27 Relative frequency of sedimentologies in which whole (A) and partial (B) skeletons were recovered. 50 km Figure 28 Map showing localities of all taxa in the OPMDB. Ohio outline in white.

166 147 Legend Per 100 km² 50 km Figure 29 Kernel density overlain by localities of fossil mammals (in blue) from the OPMDB. Legend Per 100 km² 50 km Figure 30 Kernel density map derived from OPMDB locality data excluding all mastodon point data. Ohio outline in black.

167 148 Legend Per 100 km² 50 km Figure 31 Kernel density map derived from OPMDB locality data excluding all proboscidean point data. Ohio outline in white. Legend Per 100 km² 50 km Figure 32 Bison spp. kernel density map overlain by directional deviation ellipse. Map depicts probable Bison spp. range in state. Central feature is white circle. Ohio outline in white.

168 149 Legend Per 100 km² 50 km Figure 33 Caprinae kernel density map overlain by directional deviation ellipse. Map depicts probable Caprinae range in state. Central feature is white circle. Ohio outline in white. Legend Per 100 km² 50 km Figure 34 Castoroides ohioensis kernel density map overlain by directional deviation ellipse. Map depicts probable Castoroides ohioensis range in state. Central feature is white circle. Ohio outline in white.

169 150 Legend Per 100 km² 50 km Figure 35 Cervalces scotti kernel density map overlain by directional deviation ellipse. Map depicts probable Cervalces scotti range in state. Central feature is white circle. Ohio outline in white. Legend Per 100 km² 50 km Figure 36 Cervus elaphus kernel density map overlain by directional deviation ellipse. Map depicts probable Cervus elaphus range in state. Central feature is white circle. Ohio outline in white.

170 151 Legend Per 100 km² 50 km Figure 37 Equus spp. kernel density map overlain by directional deviation ellipse. Map depicts probable Equus spp. range in state. Central feature is white circle. Ohio outline in white. Legend Per 100 km² 50 km Figure 38 Mammut americanum kernel density map overlain by directional deviation ellipse. Map depicts probable M. americanum range in state. Central feature is white circle. Ohio outline in white.

171 152 Legend Per 100 km² 50 km Figure 39 Mammuthus primigenius kernel density map overlain by directional deviation ellipse. Map depicts probable M. primigenius range in state. Central feature is white circle. Ohio outline in white. Legend Per 100 km² 50 km Figure 40 Megalonyx jeffersonii kernel density map overlain by directional deviation ellipse. Map depicts probable M. jeffersonii range in state. Central feature is white circle. Ohio outline in white.

172 153 Legend Per km² 50 km Figure 41 Rangifer tarandus kernel density map overlain by directional deviation ellipse. Map depicts probable R. tarandus range in state. Central feature is white circle.ohio outline in white. Legend Per km² 50 km Figure 42 Tayassuidae kernel density map overlain by directional deviation ellipse. Map depicts probable Tayassuidae range in state. Central feature is white circle. Ohio outline in white.

173 154 Legend Per 100 km² 50 km Figure 43 Unidentified Proboscidea kernel density map overlain by directional deviation ellipse. Map depicts probable unidentified Proboscidea range in state. Central feature is white circle.ohio outline in white. 50 km Figure 44 Map showing localities of all paleohuman site localities in the Ohio Historical Society (OHS) database. Points are deliberately oversized to prevent disclosure of the precise geographic locations of archeological sites, in keeping with OHS requirements. Ohio outline in white.

174 km Figure 45 Directional deviation ellipse created from all paleohuman localities in OHS database. Central Feature is white circle Ohio outline in white. 50 km Figure 46 Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 15,460-14,030 cal yr. Central feature is white circle. Ohio outline in white.

175 km Figure 47 Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 13,840-12,950 cal yr. Central feature is white circle. Ohio outline in white. 50 km Figure 48 Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 12,900-12,000 cal yr. Central feature is white circle. Ohio outline in white.

176 km Figure 49 Directional deviation ellipse created from paleohuman localities with radiocarbon dates from 11,255-10,245 cal yr. Central feature is white circle. Ohio outline in white. Legend Per km² 50 km Figure 50 Kernel density map created from all OHS paleohuman localities. Ohio outline in white.

177 158 Legend Per km² 50 km Figure 51 Kernel density map created from all paleohuman localities with radiocarbon dates. Ohio outline in white. Legend Per km² 50 km Figure 52 Kernel density map created from paleohuman localities from the 15,460-14,030 cal yr interval. Ohio outline in white.

178 159 Legend Per km² 50 km Figure 53 Kernel density map created from paleohuman localities from the 13,840-12,950 cal yr interval. Ohio outline in white. Legend Per km² 50 km Figure 54 Kernel density map created from paleohuman localities from the 12,900-12,000 cal yr interval. Ohio outline in white.

179 160 Legend Per km² 50 km Figure 55 Kernel density map created from paleohuman localities from the 11,255-10,245 cal yr interval. Ohio outline in white. Legend Per 100 km² 50 km Figure 56 Kernel density map for all Proboscidea localities in the OPMDB. Ohio outline in white.

180 161 Legend Per 100 km² 50 km Figure 57 Kernel density map for the Mastodon Fauna. Ohio outline in white. Legend Per 100 km² 50 km Figure 58 Kernel density map for the Mammoth Fauna. Ohio outline in white.

181 162 Legend Per 100 km² 50 km Figure 59 Kernel density map for all hoofed mammals. Ohio outline in white Figure 60 Dendrogram of taxa using full dataset. Analysis clusters groups of taxa according to how similar their localities are, using the Raup- Crick similarity index. Cophenetic correlation coefficient is Six large clusters are defined here: 1) Groups , 2) Group 304, 3) Groups 305 and 308, 4) Groups 306, 307, and 309, 5) Group 310, and 6) Group 311. Table 10 Taxonomic composition of each group defined by cluster analysis on localities. Table values are percentage contribution of that taxon to that dendrogram group. See Figure 60 for dendrogram. Cluster Number Group Number Carnivora Bison sp Castoroides ohioensis Cervalces scotti Cervus elaphus Equus sp. Mammut americanum Mammuthus primigenius Megalonyx jeffersonii Megatherium sp. Mylohyus nasutus Platygonus compressus Ovibos moschatus Proboscidea Rangifer tarandus Symbos cavifrons Tapirus sp.

182 km Figure 61 Directional deviation ellipses for Cluster 1. Included taxa are M. jeffersonii (gray), S. cavifrons (red), R. tarandus and Tapirus sp. (pink), and P. compressus (orange). Notice considerable overlap of ellipses, suggesting an ecological association among these mammals. Ohio outline is in black.

183 km Figure 62 Directional deviation ellipse for Cluster 2. This cluster contains just one taxon, M. americanum (yellow). Ohio outline is in black. 50 km Figure 63 Directional deviation ellipses for Cluster 3. Included taxa are C. ohioensis (green), and Equus spp. and O. moschatus (lt. green). Notice considerable overlap of ellipses, suggesting an ecological association among these mammals. Ohio outline is in black.

184 km Figure 64 Directional deviation ellipses for Cluster 4. Included taxa are Bison spp., Megatherium sp., and M. nasutus (blue), C. scotti (turquoise), and unidentified Proboscidea (lt. blue). Notice considerable overlap of ellipses, suggesting an ecological association among these mammals. Ohio outline is in black. 50 km Figure 65 Directional deviation ellipse for Cluster 5. This cluster contains just one taxon, M. primigenius. Ohio outline is in black.

185 km Figure 66 Directional deviation ellipse for Cluster 6. Included taxa are Family Carnivora and C. elaphus. Ohio outline is in black Figure 67 Dendrogram of taxa using simplified site and taxon data. Analysis clusters groups of taxa according to how similar their localities are, using the Raup-Crick similarity index. Cophenetic correlation coefficient is Six large clusters can be defined: 1) Groups , 2) Groups , 3) Groups 102, 103, 107, 112, 4) Groups , 5) Group 110, 6) Group 111.

186 167 Table 11 Taxonomic composition of each group defined by cluster analysis on localities using simplified dataset. Table values are percentage contribution of that taxon to that dendrogram group. See Figure 67 for dendrogram. Cluster Number Group Number Carnivora Bison sp. C. ohioensis C. scotti C. elaphus Equus spp. M. americanum M. primigenius M. jeffersonii Megatherium sp. Tayassuidae O. virginianus O. moschatus Proboscidea R. tarandus S. cavifrons Tapirus sp Figure 68 Dendrogram of taxa excluding M. americanum. Analysis clusters groups of taxa according to how similar their localities are, using the Raup-Crick similarity index. Cophenetic correlation coefficient is Four large clusters can be defined: 1) Groups , 2) Groups , 611, 3) Group 610, 4) Group 609.

187 168 Table 12 Taxonomic contents and taxon percentage of each group for Figure 68 dendrogram. Cluster Number Group Number A. simus B. bison B. latifrons C. ohioensis C. scotti C. elaphus Equus spp. M. primigenius M. jeffersonii Megatherium sp. M. nasutus O. virginianus O. moschatus P. compressus Proboscidae R. tarandus S. cavifrons Tapirus sp. U. americanus km Figure 69 M. americanum kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

188 km Figure 70 Mastodon Fauna kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white. 50 km Figure 71 M. primigenius kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

189 km Figure 72 Mammoth Fauna kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white. 50 km Figure 73 Megalonyx jeffersonii kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

190 km Figure 74 Castoroides ohioensis kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white. 50 km Figure 75 Cervalces scotti kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

191 km Figure 76 Equus spp. kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white. 50 km Figure 77 Bison bison kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

192 km Figure 78 Platygonus compressus kernel density map overlain by paleohuman occupation ellipse. Ohio outline in white.

193 174 S.cavifrons 0.44% R. tarandus 1.32% U. americanus 0.88% A. simus 1.76% B. bison 1.76% B. bombifrons 0.88% O. virginianus 0.44% Proboscidea 10.57% C. ohioensis 3.52% C. scotti 5.73% M. nasutus 0.44% P. compressus 10.13% C. elaphus 1.32% Equus spp. 3.52% M. jeffersonii 1.32% M. primigenius 10.13% M. americanum 45.81% Figure 79 Relative frequency of taxa found within the human occupation ellipse.

194 175 P. compressus 2% R. tarandus 2% B. bombifrons 2% C. ohioensis 4% Proboscidea 23% C. scotti 23% M. jeffersonii 2% Proboscidea 10% C. scotti 6% M. primigenius 19% M. americanum 52% M. americanum 54% A B Figure 80 Relative frequency of taxa found within the human occupation ellipse for sites dated to 15,460-14,030 cal yr (A) and 13,840-12,950 cal yr (B). R. tarandus 2% P. compressus 4% O. virginianus 1% M. nasutus 1% U. americanus 1% S. cavifrons 1% Proboscidea 11% A. simus 2% B. bison 2% B. bombifrons 1% C. ohioensis 5% C. scotti 7% C. elaphus 2% R. tarandus 1% Proboscidea 10% B. bombifrons 1% C. scotti 6% C. ohioensis 2% Equus spp. 4% M. jeffersonii 2% Equus spp. 1% P. compressus 17% M. primigenius 6% M. primigenius 9% M. americanum 50% M. americanum 51% A B Figure 81 Relative frequency of taxa found within the human occupation ellipse for sites dated to 12,900-12,000 cal yr (A) and 11,255-10,245 cal yr (B).

195 176 Table 13 Correlation table between kernel density rasters of paleohumans and relevant prey mammals. Mammal groups are listed on the left starting with the highest correlation with paleohumans. Mammal group Paleohuman Mastodon Fauna Mammoth Fauna Caprinae R. tarandus P. compressus M. nasutus M. jeffersonii M. americanum M. primigenius Equus spp. C. elaphus C. scotti C. ohioensis B. bison Paleohuman Mastodon Fauna M. americanum Mammoth Fauna B. bison M. primigenius M. jeffersonii P. compressus R. tarandus C. scotti Equus spp C. ohioensis Caprinae C. elaphus M. nasutus Table 14 Correlation table between kernel density rasters of paleohumans dated 15,460-14,030 cal yr and relevant prey mammals. Mammal groups are listed on the left starting with the highest correlation with paleohumans. Mammal group Paleohuman Mastodon Fauna Mammoth Fauna Caprinae R. tarandus P. compressus M. nasutus M. jeffersonii M. americanum M. primigenius Equus spp. C. elaphus C. scotti C. ohioensis B. bison Paleohuman C. scotti Mastodon Fauna M. americanum M. jeffersonii M. primigenius Mammoth Fauna R. tarandus C. elaphus C. ohioensis M. nasutus P. compressus B. bison Caprinae

196 177 Equus spp Table 15 Correlation table between kernel density rasters of paleohumans dated 13,840-12,950 cal yr and relevant prey mammals. Mammal groups are listed on the left starting with the highest correlation with paleohumans. Mammal group Paleohuman Mastodon Fauna Mammoth Fauna Caprinae R. tarandus P. compressus M. nasutus M. jeffersonii M. americanum M. primigenius Equus spp. C. elaphus C. scotti C. ohioensis B. bison Paleohuman C. scotti Mastodon Fauna M. primigenius M. americanum Mammoth Fauna C. ohioensis P. compressus M. jeffersonii Caprinae Equus spp R. tarandus C. elaphus M. nasutus B. bison

197 178 Table 16 Correlation table between kernel density rasters of paleohumans dated 12,900-12,000 cal yr and relevant prey. Mammal groups are listed on the left starting with the highest correlation with paleohumans. Mammal group Paleohuman Mastodon Fauna Mammoth Fauna Caprinae R. tarandus P. compressus M. nasutus M. jeffersonii M. americanum M. primigenius Equus spp. C. elaphus C. scotti C. ohioensis B. bison Paleohuman C. elaphus Mastodon Fauna M. americanum M. jeffersonii C. scotti C. ohioensis M. primigenius Mammoth Fauna M. nasutus R. tarandus B. bison Equus spp Caprinae P. compressus Table 17 Correlation table between kernel density rasters of paleohumans dated 11,255-10,245 cal yr and relevant prey mammals. Mammal groups are listed on the left starting with the highest correlation with paleohumans. Mammal group Paleohuman Mastodon Fauna Mammoth Fauna Caprinae R. tarandus P. compressus M. nasutus M. jeffersonii M. americanum M. primigenius Equus spp. C. elaphus C. scotti C. ohioensis B. bison Paleohuman Mastodon Fauna M. americanum C. scotti Caprinae Equus spp Mammoth Fauna M. primigenius R. tarandus C. elaphus M. nasutus C. ohioensis P. compressus M. jeffersonii

198 179 B. bison Table 18 Taxon occurrences in each sedimentological category. Category Mixed Peat Clay Gravel Sand Sand Mixed Silt Uncategorized Carnivora Bison spp Caprinae C. ohioensis Cervidae Equus spp M. americanum M. primigenius Pilosa Tayassuidae Unidentified Proboscidae R. tarandus Tapirus sp Total taxa per group

199 180 Unidentified Proboscidea 12% Caprinae 8% C. ohioensis 4% Equus spp. 2% Unidentified Proboscidea 9% Equus spp. 18% M. primigenius 16% Tayassuidae 36% M. americanum 37% M. americanum 58% A B Unidentified Proboscidea 5% Bison spp. 5% Caprinae 5% Equus spp. 5% M. americanum 9% Tayassuidae 25% C. ohioensis 25% Tayassuidae 57% M. primigenius 14% M. primigenius 13% M. americanum 37% C D Figure 82 Relative frequency of taxon occurrences in gravel (A), sand (B), sand-mixed (C), and silt (D) settings.

200 181 Unidentified Proboscidea 3% Order Pilosa 3% M. primigenius 6% R. tarandus 3% Tapirus sp. 3% Cervidae 3% Unidentified Proboscidea 9% Order Pilosa 7% M. primigenius 4% R. tarandus 2% Carnivora 2% Caprinae 2% Cervidae 11% C. ohioensis 9% Equus spp. 2% M. americanum 79% M. americanum 52% A B Tayassuidae 7% R. tarandus 7% Order Pilosa 7% Carnivora 33% M. primigenius 7% M. americanum 6% C. ohioensis 6% Cervidae 27% C Figure 83 Relative frequency of taxon occurrences in clay (A), peat (B), and mixed (C) settings.

201 182 Skeleton 3% Partial skeleton 2% Cranium-upper palate 8% Mandible 1% Unidentified 8% Unidentified 11% Mandible 17% Hip girdle 2% Vertebrae 2% Rib 2% Long bone 8% Tooth 28% Skeleton 33% Tooth 17% Horn, horn core, & antler 6% Tusk 27% Tusk 25% A B Unidentified 6% Cranium-upper palate 8% Tooth 8% Mandible 6% Unidentified 20% Tooth 7% Cranium_uppe r palate 6% Mandible 6% Skeleton 33% Tusk 8% Partial skeleton 13% Tusk 20% Misc. 3% Long bone 6% Hip girdle 3% Vertebrae 8% Rib 8% Horn, horn core, & antler 3% Long bone 7% Vertebrae 7% Rib 7% Shoulder girdle 7% C D Figure 84 Relative frequency of skeletal elements preserved in gravel (A), sand (B), sand-mixed (C), and silt (D) settings.

202 183 Partial skeleton 4% Unidentified 9% Cranium-upper palate 1% Mandible 7% Unidentified 10% Cranium-upper palate 3% Mandible 3% Misc. 1% Skeleton 9% Manus/pes 9% Long bone 8% Hip girdle 3% Vertebrae 10% Rib 9% Tooth 12% Tusk 14% Horn, horn core, & antler 1% Shoulder girdle 3% Misc. 3% Manus/pes 4% Skeleton 8% Partial skeleton 10% Long bone 13% Hip girdle 4% Vertebrae 6% Rib 7% Tooth 19% Tusk 6% Horn, horn core, & antler 1% Shoulder girdle 3% A B Tooth 23% Unidentified 55% Rib 9% Tusk 4% Vertebrae 4% Manus/pes 5% C Figure 85 Relative frequency of skeletal elements preserved in clay (A), peat (B), and mixed (C) settings.

203 184. Figure 86 Cluster analyses of skeletal elements. Top left, cluster diagram grouping skeletal elements by Voorhies group. Cophenetic correlation coefficient is Bottom left, cluster diagram grouping skeletal elements by taxon. Cophenetic correlation coefficient is Bottom right, cluster diagram grouping skeletal elements by sedimentology. Cophenetic correlation coefficient is See Table 19 for composition of groups. Table 19 Key to groups shown in Figure 86. Dendrogram/color Gray Pink Brown Tan Green Blue Purple Skeletal elements by Voorhies group Crania Mandible Misc. Shoulder girdle Long bone Manus/pes Hip girdle Vertebrae Rib --- Skeletal element by taxon Horn-horn core Crania Tusk Rib Shoulder girdle Hip girdle Vertebrae Long bone Manus/pes Tooth Mandible --- Misc. Skeletal element by sedimentology Horn-horn core Crania Tusk Hip girdle Shoulder girdle Vertebrae Manus/pes Rib Long bone Mandible Misc. Tooth

204 Figure 1 Simplified taphofacies dendrogram. Cluster analysis using Euclidean distances produced the dendrogram showing skeletal elements grouped by locality. The initial dendrogram was produced from a trimmed OPMDB dataset using all localities that have skeletal elements and sedimentary data. The lower row of pie charts indicates pooled skeletal element data for each branch of related localities. These are then compared to occurrences by sedimentological class (top row of graphs). Three clusters were denoted from skeletal element data and sedimentology categories that correspond to glaciofacies as defined in text (Miller, 2009). Cluster A is Ice Contact Deposition. Cluster B is Late and Post Glacial Deposition. Cluster C is Proglacial Lake Deposition. Key totaphofacies dendrogram, lower skeletal pie charts (left) and upper sedimentology pie charts (right). 185

205 186 Manus/pes 1% Vertebrae 3% Long bone 15% Mandible 8% peat 24% clay 9% silt 1% sand 8% Rib 2% sand mixed 8% Tooth 42% mixed 8% Tusk 29% gravel 42% A B Cranium_uppe r palate 16% peat 16% clay 22% Tooth 9% Unidentified 51% Tusk 9% mixed 24% silt 8% Horn_horn core_antler 10% sand mixed 6% sand 2% gravel 22% C Hip girdle 1% Vertebrae 4% D Figure 88 Cluster A Glacio-facies One-Ice Contact Deposition (A & B) and Cluster B Glacio-facies Three-Late and Post Glacial Deposition (C & D). Cluster A Glacio-facies One-Ice Contact Deposition relative frequency of skeletal elements (A) and sedimentology (B). Cluster B Glaciofacies Three-Late and Post Glacial Deposition relative frequency of skeletal elements (C) and sedimentology (D).

206 187 Unidentified 6% Cranium-upper palate 1% Manus/pes 10% Misc 4% Mandible 7% Tooth 11% peat 20% Long bone 10% Tusk 10% mixed 10% clay 45% Hip girdle 6% Shoulder girdle 6% gravel 5% Vertebrae 12% Rib 17% Horn, horn core, & antler 0% sand mixed 15% silt 5% A B sand 0% Figure 89 Cluster C Glacio-facies Two Proglacial Lake Deposition. Relative frequency of skeletal elements (A) and sedimentology (B).

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225 206 APPENDIX A. ADDITIONAL BACKGROUND TEXT AND FIGURES Faunal species synopses Bison spp. Bison are believed to have emigrated into North American via Beringia from Asia in the middle Pleistocene, about ka (Jiménez-Hidalgo et al., 2013; Scott, 2010; Shapiro et al., 2004). They moved southward into central North America around ka and then across the United States (Jiménez-Hidalgo et al., 2013; Shapiro et al., 2004). Pleistocene Bison spp. fossils are abundant throughout Beringia (Shapiro et al., 2004). By the late Pleistocene and early Holocene, bison were widely distributed across North America and inhabited most regions not covered by glacial ice (Jiménez-Hidalgo et al., 2013; Rivals et al., 2007; Scott, 2010). Bison are so prevalent at this time they are the mammalian index fossil of the Rancholabrean North American Land Mammal Age below the 55 latitude (Jiménez-Hidalgo et al., 2013; Scott, 2010). Currently, six species of Bison are recognized of Quaternary age in North America: Bison latifrons, Bison antiquus, Bison occidentalis, Bison alaskensis, Bison priscus, and Bison bison (also historically termed Bison sylvestris) (Jiménez-Hidalgo et al., 2013; Scott, 2010). All but the last are extinct members of the group. Only B. bison and B. latifrons are relevant to this study. An early species, B. latifrons were generally not as geographically widespread as the other bison species. During the Illinoian glacial stage, B. latifrons probably had a relatively larger range due to the extensive woodland and forest habitats preferred by the species (Scott, 2010). Bison bison is recognized as containing at least two sub-species, B. bison bison (the plains bison) and B. bison athabascae (wood bison) (Jiménez-Hidalgo et al., 2013).

226 207 Post glacially, B. bison would begin to dominate up through central Alberta, Canada. Paleontological and genetic data explain this as a recolonization from the south through (Jass et al., 2011). Data presents a picture of these large herbivores flourishing and becoming dominant in the late Pleistocene over areas previously held by Equus. Bison continue to thrive in areas of North America (Jass et al., 2011; Zazula et al., 2009). Modern bison are large, herding ungulates often considered a keystone species, i.e. species that have a dominant impact upon the ecosystem relative to other cohabiting species. It is presumed that paleobison may have served a similar role in ecosystems (Scott, 2010). However, other evidence suggested that Pleistocene species may have behaved dissimilar to the modern. Analyses of fossil teeth suggest at least some paleobison species utilized more browse than modern bison (Feranec et al., 2009; Jiménez-Hidalgo et al., 2013; Rivals et al., 2007; Scott, 2010). Modern wood bison have differing mesowear than modern plains bison and one which mimics closely that of many fossil species (Fenerac et al., 2009; Rivals et al., 2007). Isotopic and mesowear analyses imply fossil bison species, except B. latifrons, were perhaps more flexible in their dietary regimes than the modern plains bison (Fenerac et al., 2009; Jiménez-Hidalgo et al., 2013). Herding behavior does not seem to be consistent between all bison species. B. latifrons has been inferred to be less social and more solitary than other species because the frequency of individual finds within the fossil record. Other extinct species have been presumed to follow the pattern of modern bison, though perhaps not in such vast herds as seen historically (Scott, 2010). As a result of their wide distribution and primarily herding nature, bison are well represented within relevant mammal fossil assemblages. Most bone elements are found in quantity within the

227 208 fossil record. Teeth are most common but other cranial and post cranial material are preserved (Burns, 2010; Czaplewski, 2012; Jiménez-Hidalgo et al., 2013; Harington, 2011; Rivals et al., 2007; Scott, 2010; Velivetskaya et al., 2011; Zazula et al., 2009). The muscle, skin, and hair of Steppe Bison (B. priscus) have even been discovered (Harington, 2011). Fossil specimens have been commonly recovered from silts (Harington, 2011; Jiménez- Hidalgo et al., 2013), in gravel bars and pits (Burns, 2010; van Kolfschoten et al., 2011), in sands and beach deposits (Jiménez-Hidalgo et al., 2013; Zazula et al., 2009), and in clays (Czaplewski, 2012; Jiménez-Hidalgo et al., 2013). Caprinae Musk-ox is a common term for some ovibovoid mammals (Campos et al., 2010). Although represented by a single taxon modernly, musk-oxen were more diverse during the Pleistocene and earliest Holocene (Boeskrorov, 2006; Campos et al., 2010; MacPhee et al., 2005). The Helmeted Musk-ox (Bootherium) was the most widespread and dominant in the Western Hemisphere. The genus ranged throughout all of North America. The genus Bootherium first appeared in eastern Beringia, within the broad, intermontane valleys, and later dispersing to the lower latitude plains during the Illinoian and Wisconsinian glaciations, ka. The latest known specimens, about 12,900 cal yr, are from Michigan and southern Alberta (Campos et al., 2010). There are questions about the relationship between the two genera Bootherium and Symbos that arose soon after the description of the two genera as to whether they represent distinct genera or are simply divided upon characteristics of sexual dimorphism. Fossil examples of both often occur within the same localities. Similarly, the elements physically resemble the

228 209 males and females of the extant Ovibos and bone ratios are consistent with sexual dimorphism. As a result, some researchers consider Symbos to be the male and Bootherium the female of the genus Bootherium (Campos et al., 2010). The genus Ovibos is a more cosmopolitan member of Caprinae as their geographic distribution was not limited to the Americas (Campos et al., 2010). Fossil evidence indicates Ovibos had Eurasian origin. Data suggests all three genera entered North America around the same time (MacPhee et al., 2005). During the Pleistocene, Ovibos had a periglacial distribution. Modernly, it is limited to Greenland and the Arctic Archipelago (Campos et al., 2010; MacPhee et al., 2005). This restriction appears to be a rather recent one as Ovibos were common in Alaska during the earlier Pleistocene but relatively rare by 40 ka (MacPhee et al., 2005). Few Ovibos fossils are found prior to the Wisconsinan in Greenland and Canada after glacial retreat, the fossil record increased noticeably, most especially during the mid-holocene (MacPhee et al., 2005). This genus does not seem to have been limited to a tundra existence and some researchers believe this flexibility may be the key to its success in surviving the end Pleistocene (Campos et al., 2010). Data suggests these mammals preferred grasslands, alpine meadows, and woodlands (Campos et al., 2010). Diet of ovibovoids varied, though fossil dung evidence suggests they browsed more than grazed (Campos et al., 2010). Modern O. moschatus is a browser of a differential diet depending upon season (Barboza & Reynolds, 2004; Kazmin & Abaturov, 2011). Preferred food sources are leaves and shoots of willows (Salicaceae) (Barboza & Reynolds, 2004; Lawler & White, 2006; Kazmin & Abaturov, 2011; Kazmin, et al 2011). However, legumes (Fabaceae), sedges and rushes, as well as some alpine and meadow grasses compose a considerable portion of their diet (Barboza & Reynolds, 2004; Kazmin & Abaturov,

229 ; Kazmin et al., 2011). Requirements are very similar to that of caribou (Rangifer taradus) and the two groups are often paired within arctic research (Barboza & Reynolds, 2004; Kazmin & Abaturov, 2011; Kazmin et al., 2011). Peculiarly although ovibovoids were widely distributed during the Quaternary, they are uncommon as fossils (MacPhee et al., 2005). Fossils are most often found in silts and gravels (Campos et al., 2010; Jass et al., 2011). Common elements found are crania, cranial fragments, and horn cores (Jass et al., 2011; Ray, 1966; Zazula et al., 2009). Some post-cranial material, such as vertebrae and vertebral fragments as well as few metatarsals, have been discovered (Jass et al., 2011). Carnivora Recreating predator-prey relationships in the past is a particularly frustrating endeavor as predators are naturally underrepresented within the fossil assemblage (Burns, 2010; Jass et al., 2011; McHorse et al., 2012; Muñoz-Durán & van Valkenburgh, 2006). The only exceptions to this generalization apply to what many researchers denote as predator traps, such as caves and asphalt seeps (McHorse et al., 2012; Muñoz-Durán & van Valkenburgh, 2006). In these cases, it is often that a few carnivores become predominantly overrepresented. A good example of this might be the Rancho La Brea fauna which has a unusually large population of Smilodon sp. and Dirus canis represented (Springer et al., 2010). Another good example are the cave systems of Natural Trap Cave and Little Box Elder Cave, Wyoming which are dominated by canids, felids, and ursids (Kohn & McKay, 2012). Ohio does not possess active asphalt seeps, though it does possess karst regions which might represent predator traps. One such area is in Wyandot county at the Sheriden and Indian Trails cave systems and there are a few fossils from large carnivores here. However overall,

230 211 Ohio has an unusual paucity of large predators within the Pleistocene fossil assemblage. This is most especially apparent when the fossil records from surrounding states are compared to the Ohio record. Some generalities may be made for carnivores, irrespective of their size. Predators tend to be larger than their prey (Brose et al., 2006; Shultz & Finlayson, 2010). Predator size and prey size are positively correlated as when one increases, the other does as well (Brose et al., 2006). Research of extant predators also suggests differences in habitat breadth are disconnected from the incidence of fossil appearance in assemblages. However, there is a considerable, positive correlation between size and frequency of fossil representation where larger carnivores are better represented than smaller-bodied ones (Muñoz-Durán & van Valkenburgh, 2006). Social predators tend to exploit larger prey sizes as opposed to lone hunters, although groups do not show significant biases towards the largest sizes available (Shultz & Finlayson, 2010). Terrestrial carnivores commonly are present in low densities, have large home ranges upon which they move, and feed as opportunity presents itself. These factors make discrimination of habitats and behaviors somewhat difficult to puzzle out. In modern carnivore studies, the traditional methods of observation, direct and indirect, are utilized to determine diet, home range, and behaviors. Even with this, it is still problematic as many predators rely on being unseen to successfully hunt prey. With these facts in mind, discovering the habits of extinct predators can be even more challenging as direct observation is impossible and modern environmental analogs are often unavailable. The major large predators for the Pleistocene Midwest would be from three main families, the Felidae, the Ursusae, and the Canidae. Felids tend to be underrepresented within the fossil record, though there are exceptions (Sommers & Nadachowski, 2006). The largest among cat groups were those of the saber-toothed

231 212 cat (Smilodon fatalis) and the American lion (Panthera atrox). These cats were members who went extinct at the end Pleistocene though the reason is not yet clear (Barnett et al., 2009; DeSantis et al., 2012). Large and morphologically diverse varieties of lion were scattered across the Northern Hemisphere. The American lion is generally thought to be a close relative of the cave or steppe lion (Panthera leo spelaea) present throughout Europe and Asia (Barnett et al., 2009; Deidrich, 2011). Studies suggest that an ancestral Beringian population of P. leo spelaea migrated into North America and developed into P. atrox between ka before moving into central North America by ka and persist until the end of the Pleistocene, about 11 ka (Barnett et al., 2009). Social behavior in large extinct felids may be inferred but not proven. Usually, evidence relies upon obvious sexually dimorphic anatomical features, such as a long mane or significantly larger body size. Generally, P. atrox is thought to have been social. Saber-toothed felids, Machairodontinae, were extensive and prevalent group of large predators often proposed to be social (Christiansen & Harris, 2012, Dundas, 1999; McHorse et al., 2012). The difficulty rests in determining the difference between species specific sexual dimorphism versus characters which denote differences between distinct species of Machairodontinae (Christiansen & Harris, 2012; Dundas, 1999). The Pleistocene jaguar (Panthera onca augusta) generally ranged through central and southern North America. Harrodsburg Crevice, a Pleistocene site in southern Indiana, has the northernmost known site for this felid (Smith & Polly, 2013). Modernly, the jaguar (Panthera onca) is restricted to mostly Central and South America, with a few incursions into the southern United States. However, this appears to be of historic origin as individuals had been found east

232 213 of the Mississippi River prior to the early 19 th century. Verifiable kills of jaguars were reported in Kentucky, northern Ohio, and western Pennsylvania (Smith & Polly, 2013). Habitat information about North American felids is sparse. The Machairodontinae and P. atrox probably preferred open, grassy areas with scattered tree stands, much like the modern P. leo. P. onca augusta was interpreted to have a range of tropical habitat similar to that of the modern P. onca. However, P. onca has historically been less particular about habitat than it is at present (Smith & Polly, 2013). Other than those of certain Machairodontinae, few felid fossils are found. Large cats are frequent members of predator trap assemblages and appear in crevice and cave deposits as well as asphalt traps (MacFadden et. al., 2012; Rincón et al., 2011; Smith & Polly, 2013). As yet, no large felid has been found in Ohio, though they have in the Midwest. Similodons are common in the fossil record and nearly the entire skeleton has been represented within all age cohorts (Barnett et al., 2009; Christiansen & Harris, 2012). Generally other felid finds include crania, mandibles, and teeth with less postcranial material available (Czaplewski, 2012; Diedrich, 2011; Rincón et al., 2011; Smith & Polly, 2013). Ursidae are one of the most studied families within the Carnivora. Yet, plylogenetic relationships between groups are still controversial. DNA sequences with radiocarbon, stable isotope, and paleoclimate data combined to depict the phylogeographic change of bear lineages in the late Pleistocene. Studies reveal these alterations predate the Last Glacial Maximum, as well as human entry into North America and the late Pleistocene extinction (Barnes et al., 2002). Which groups are related more closely seems to depend entirely upon the method of analysis (Krause et al., 2008).Genetic evidence points towards a rapid radiation of bears at the Miocene-Pliocene boundary. As the radiation is also common in other groups of mammals, it has

233 214 been implied climate change played a large role (Krause et al., 2008). Ursids exist upon nearly every continent and environment from arctic to rainforest. Many members are omnivorous and feed upon varied diets. They subsist upon plants, insects and other invertebrates, as well as meat (Barnes et al. 2002; McHorse et al., 2012). For the purpose of this research, only three members of this family are of importance, the brown bear (Ursus arctos), the American short-faced bear (Arctodus simus), and the black bear (Ursus americanus). U. arctos are extant and have extensive modern distribution across Europe, Asia, and North America (Barnes et al., 2002; Boeskrorov, 2006). Evidence suggests this species migrated from Asia, across Beringia about ka (Burns, 2010; Matheus et al., 2004) Radiocarbon dates depict that U. arctos reached areas south of Beringia before the coalescence of the Laurentide and Cordilleran glaciers (Burns, 2010; Matheus et al., 2004). Permafrost remains from the area which was eastern Beringia allows for genetic studies of the past 60,000 BP and shows interesting patterns of localized extinction, emigration, and possibly, interspecies competition (Barnes et al., 2002). Genetic studies seem to indicate that U. arctos in the Pleistocene were more diverse than modernly (Barnes et al., 2002; Matheus et al., 2004). A. simus diverged earlier than either U. arctos or U. americanus. It is believed this the short-faced bear is more basal and branched off with their sister clade, spectacled bears, around 5.7 Ma (Krause et al., 2008). Stable isotope evidence indicates A. simus as more of a carnivore than that of U. arctos. It has been suggest that some of the patterns of local extinction and recolonization of U. arctos seen in parts of Beringia could be competition from the larger, more carnivorous A. simus (Barnes et al., 2002). U. americanus is an extant omnivore species which has been present within North America for at least 3 Ma and has largely dispersed across the continent within favorable habitat

234 215 for much of its history. Currently black bears range from coast to coast, as far north as Alaska and as far south as Mexico. The Pleistocene forest distribution is considered one of the most important influences to current U. americanus genetic diversity. Refugia created by Pleistocene events created extant diversity patterns. Modernly, forested areas extend across much of North America but Pleistocene forest dispersion was patchier and created isolates of bear populations (Wooding & Ward, 1997). U. americanus fossils are rare, perhaps because they are extant and up until recently, considered a pest species which was frequently hunted down and exterminated. Pleistocene ursids have been found in Ohio. Like many other terrestrial predators, Ursid fossil material is rare in the record. Typical elements found are crania, mandible, and teeth (Barnes, 2010; Harington, 2011; Jass et al., 2010).Unlike felids, ursids are not well represented predator trap situations such as asphalt seeps. Though, U. arctos is more commonly found in seeps than U. americanus (McHorse et al., 2012). Samples have been found in permafrost layers of North America (Barnes et al., 2002), in gravels of fluvial systems (Burns, 2010; MacFadden et. al., 2012), and marls and sands (MacFadden et. al., 2012). The family Canidae is better represented than most carnivores within the North American fossil assemblages. In the past, canids were widespread throughout North and South America. Late Pleistocene sites for C. dirus are concentrated in the United States, especially in California, Texas, Missouri, and Florida. The dire wolf (Canus dirus) occurs within many Pleistocene fauna assemblages. As there are no Holocene records of C. dirus, it has been presumed the taxa went extinct throughout its range at the end Pleistocene extinctions (Dundas, 1999). A North American origin is favored for C. dirus for a number of reasons. Among the most evident is the substantially better fossil distribution of C. dirus within North America than in South America, the probable ancestors of C. dirus existed within the middle Pleistocene North

235 216 America, and the initial appearance of C. dirus occured earlier in the North American fossil record. The oldest record currently known is that of Salamander Cave in the Black Hills of South Dakota and is approximately 252 ka based upon uranium series dating (Dundas, 1999). There are also many sites throughout the United States which are of Sangamonian Age (Dundas, 1999). At that time, C. dirus ranged from Alberta, Canada to Texas and from the Atlantic to Pacific coasts. In the Wisconsinan, their range expanded south into much of Mexico. Wisconsinan sites are more common than those of Sangamonian so the seemingly expanded range may just be an artifact of collection bias. C. dirus finds appear more rarely in South America and are probably Lujanian in age (Dundas, 1999). There are also C. dirus sites in Kentucky, Pennsylvania, Indiana, Illinois, Tennessee, West Virginia, Virginia, and Wisconsin but surprisingly, none have yet been found in Ohio. The last appearance for the species remains undetermined despite the vast number of late Pleistocene sites. The Harrodsburg Crevice, Indiana site has the presence of C. dirus at least to 29,700 cal yr (Smith & Polly, 2013). The problem remains that most of the sites have not been adequately dated to determine the extent and timing of the extinction. Thus far the latest known specimens are questionable radiocarbon dates of about 11,240 cal at La Brea, C.A., 12,790 cal yr at La Brea, C.A, and 16, 020 cal yr at Powder Mill Creek Cave, M.O.(Dundas, 1999). Morphologic studies have suggested the dire wolf was a hypercarnivore. Robust skulls and tooth breakage indicate they likely were more adapted for larger prey, whether through active hunting or scavenging is not as clear. Isotopic studies suggest they were not species specific predators (Fox-Dobbs et al., 2007). Dire wolves were not however, considered to have the bone-crushing abilities of carnivores like the spotted hyena (Crocuta crocuta) (Dundas, 1999). Fossils of C. dirus are common. As these mammals have often been found preserved

236 217 together in quantity, it is presumed C. dirus were gregarious in nature. Large modern canids are social so it is probable the dire wolf was as well (McHorse et al., 2012). Most sites are of one or a few individuals but some have many more. La Brea, California has thousands (Dundas, 1999; McHorse et al., 2012). Sites in Talara, Peru, Cutler Hammock, Florida, and in San Josecito Cave, Mexico have dozens of finds each (Dundas, 1999). Conversely in the Pleistocene, the gray wolf (Canis lupus) are much less common, especially in the United States (Czaplewski, 2012). However, C. lupus is much less robust than their C. dirus cousins (Fox-Dobbs et al., 2007). C. lupus originated in Europe during the earlymiddle Pleistocene and travelled into North America during the late Irvingtonian via the Bering Land bridge (Dundas, 1999; Randi, 2010). Genetically, North American C. lupus populations are closely related remnants of a previously geographically widespread and abundant species (Fox- Dobbs et al., 2007). C. lupus and C. dirus were temporally co-occurrent species. It may be that C. lupus was displaced from areas where C. dirus were most common, or perhaps the larger dire wolf preyed upon different species. It is unknown to what extent these canids interacted (McHorse et al., 2012). Modernly, C. lupus separate into a number of distinct groups divided by habitat, preypreference, and geographic distribution. It is not known how many wolf-like sub-species and species exist and they readily hybridize with other canids such as coyotes (Canis latrans) (Randi, 2010). C. dirus lived in a variety of habitats from forested areas to open plains (Dundas, 1999; Rincón et al., 2011). They resided from sea level to 2255 m elevation (Dundas, 1999). It has been purported that C. dirus migrated southward into South America via the Andean Corridor, following cool, dry, open habitats (Dundas, 1999). C. lupis lives in similar intermixed forests and

237 218 open-grass lands modernly and was likely to have done so in the past as well (Dundas, 1999; MacFadden et. al., 2012; Rincón et al., 2011). C. dirus is a common Pleistocene carnivore fossil species, C. lupus is not. Evidence suggests dire wolves were frequent members of predator trap situations and gray wolves were not (Dundas, 1999; MacFadden et. al., 2012; Rincón et al., 2011). Dire wolf bone elements have occurred in asphalt seeps (Dundas, 1999; MacFadden et. al., 2012; Rincón et al., 2011; Smith & Polly, 2013), within cave deposits (McHorse et al., 2012), and in gravels and sands (MacFadden et. al., 2012). Since C. dirus are common so crania and post-cranial material are wellrepresented. C. lupus are rarely found within fossil assemblages though sparse elements of post cranial, such as pelvic and tarsal bones, and some cranial material are found (Czaplewski, 2012). Castoroides ohioensis Castoroides ohioensis, or giant beaver, were known throughout North America in the Plio-Pleistocene and in some locations, into the Holocene (Burns, 2010; Harington, 2011). C. ohioensis was the largest ice age rodent, measuring up to 2.5 meters in length and a estimated weight of 200 kg (Miller, 2000). Changing conditions of the Late Pleistocene restricted the giant beaver to the Eastern United States (McDonald & Bryson, 2010). Earlier studies of giant beaver suggested their habits were much like those of their smaller and co-occurring cousin C. canadensis, the modern beaver, as they share similar crania characteristics (Miller, 2000; Swinehart & Richards, 2001). Miller (2000) noted there is little concrete evidence of tree-felling, dam building, or den construction behaviors except for a record in the early 1900s of a possible lodge from Knoxville, Ohio where a portion of a giant beaver skull and den constructed of saplings 7.2 centimeters in diameter which measured 1.2 meters in height and 2.4 meters in diameter (Miller, 2000). Morphological studies depict a lifestyle

238 219 consistent with Ondatra zibethicus (muskrat) rather than modern beavers (Swinehart & Richards, 2001). Conditions generally accepted as C. ohioensis habitats are those of wet meadows bordering lakes and ponds with emergent forested areas of white spruce, tamarack, balsam fir as well as some aspen and birch (McDonald & Bryson, 2010; Swinehart & Richards, 2001). Fossils are usually located in sediments indicative of boggy settings, such as peat, loam, and marls (Lanken, 1993; McDonald & Bryson, 2010; Miller, 2000; Swinehart & Richards, 2001). Incisors seem to be the most common bone element found within the fossil record (Lanken, 1993; Laub, 2003; Miller, 2000). Occasionally crania and mandibles are found (Lanken, 1993; Miller, 2000). Much more rarely, limb bones are found (Lanken, 1993; Swinehart 2001). Cervalces scotti No evidence currently exists of paleohumans caused the decline of Cervalces scotti (elk moose). Paleohuman tools have been found in association, such as at Sheriden Cave, Ohio. However, there is no conclusive evidence Cervalces were butchered or scavenged. It has been proposed that these mammals were out competed by other artiodactyls such as bison moving in from the south and west (Long & Yahnke, 2011). At the time of the late Pleistocene, C. scotti had two distinctive and geographically separate populations within North America. One population resided in Beringia and the other lay immediately south of the Laurentide ice sheet (Schubert et al., 2004). Elk-moose are often associated with the ice margin and related tiaga, bogs, and marshes, similar to that of Rangifer tarandus. The changes from this vegetative habitat to that of grasslands, oak forests, and other composites likely initiated a faunal turnover from elk-moose and caribou to that of moose, elk, and deer in the Pleistocene-Holocene transition (Long & Yahnke, 2011). C. scotti has also been

239 220 associated with Megalonyx jeffersonii, as well as C. ohioensis, B. bombifrons, and Mammut americanum (McDonald et al., 2000; Schubert et al., 2004). C. scotti likely inhabited an environment which does not have a modern analog, intermediary between midlatitude tundra to mixed conifer and deciduous woodlands (Long & Yahnke, 2011; Schubert et al., 2004). Evidence suggests that while Rangifer taradus and elkmoose shared a similar cool temperature regime, C. scotti seems to have preferred a wetter periglacial climate with heavier winter precipitation (Long & Yahnke, 2011; Schubert et al., 2004). Their range appears to follow this condition eastward and southward regardless of time and relative distance from glacial fronts. Dietary requirements are not yet known for elk-moose but it has been suggested that they fed upon sedges and willows typically found within the environs they inhabited (Long & Yahnke, 2011). Elk-moose fossils are not typical finds. C. scotti fossils are found within peat and associated marls (Long & Yahnke, 2011; Robinson et al., 2005; Schubert et al., 2004) and within marls and sands do occur (Long & Yahnke, 2011). Antler beams and fragments as well as crania and cranial fragments are the most common elements found (Laub, 2003; Long & Yahnke, 2011; Schubert et al., 2004). Post cranial material is rare (Robinson et al., 2005; Schubert et al., 2004). Cervus elaphus Researching Cervus elaphus presents a difficulty. Among European researchers, C. elaphas refers to the red deer common through Europe and Asia during the Pleistocene. To the American researcher, C. elaphus refers to the Rocky Mountain or Canadian elk (C. elaphus = C. Canadensis) (Sanders, 2002). As a result, it is often unclear to which animal C. elaphus of literature refers. For the purpose of this study, C. elaphus will refer to the elk of the Americas.

240 221 Of the North American ungulate taxa, the elk are among the most geographically widespread and studied (Sawyer et al., 2007). Elk are known from Irvingtonian Cape Deceit fauna of Alaska and persist to the present day (Sanders, 2002). Although Pleistocene elk resemble modern elk, they seem to differ in that they were considerably larger. This has been suggested to result from the greater abundance of quality forage available at the terminal Pleistocene (Lyman, 2010). Before European colonization of North America, C. elaphus inhabited all of North America. The last known elk from eastern North America was purportedly killed in eastern Tennessee in 1849 (Kindall et al., 2011). Large cervids such as elk, deer, and caribou were habitually exploited by human groups from paleo to historic times, although associations of artifacts and fossil remains regarding early human hunting situations are rare (Jackson, 1987). Fossil records indicate C. elaphas populated both glacial steppe and interglacial woodland habitats, indicating environmental flexibility within the species (Bender et al., 2012; Sommer et al., 2008). Modernly, elk usually inhabit rugged elevations of mountains and canyons with forests and grasslands. They are also capable of living within arid environs and are known to migrate seasonally between winter and summer ranges (Bender et al., 2012; Cooper et al., 2003; Sawyer et al., 2007). Preferred areas are forest proximal habitats were quality cover and good forage are both accessible (Sawyer et al., 2007). C. elaphus tends to prefer forest plant species and forest habitat (Bender et al., 2012; Kindall et al., 2011). However, they will adapt to open areas (Bender et al., 2012; Sawyer et al., 2007). C. elaphus is a browse-dominated mixed feeder (Rivals et al., 2010). Grasses and forbs are consumed in some quantities as is woody vegetation (Conard & Gipson, 2012; Rivals et al.,

241 ; Sommer et al., 2008). Diet depends largely upon available seasonal flora and regionality (Conard & Gipson, 2012). C. elaphus fossil materials tend to be rare and little sedimentological data about finds is available. Some elements have been reported in conjunction with open water settings, such as shoreline deposits (Van Kolfschoten et al., 2011) and within silts interpreted as an incipient paleosol (Lyman, 2010). Skeletal elements found are often fragmentary. Sometimes teeth and vertebrae are discovered (Sanders, 2002). More rarely, a complete skeleton may be discovered (Jackson, 1987; Lyman, 2010). Equus spp. Evolution of the horse occurred in North America. The family emigrated from the continent, across the Beringian land bridge, and into Eurasia before becoming extinct from the continent of its origin (Boeskorov, 2006; MacPhee et al., 2005). Equus sp. of the Pleistocene were typical members of the Mammoth fauna and share the same open grazing requirements of many large herbivores from the time period. In Alberta, Canada, Equus was the most abundant megaherbivore before being replaced by Bison by the latest Pleistocene. This condition was common and persistent within northern North America, although it occurred at different rates spatially and temporally (Jass et al., 2011). Zazula et al. (2009) observed from their research that aridity, rather than diet alone was indicative of horse-dominated assemblages. Modern horse diet is generally dominated by grasses (C 4 plants). There is evidence of a more flexible diet within earlier horse populations from the Rancha Le Brea fauna of California. Additionally, data records that up to half of the modern North American wild (feral) horse diet is habitat dependent and up to 50% of populations consume browse (Fenarec, 2009). McDonald and Pelikan (2006) concluded that paleo Equus tooth enamel held similar isotopic signatures to

242 223 two other sympatric grazers, Bison and Mammuthus, who were determined to be C 4 grazer/mixed feeders. Horses are frequent taxa within Pleistocene fossil assemblages. Equus fossil elements are commonly found in fluvial deposition environments with sand, interbedded sands and silts, such as those found in marginal beach environs (Harington, 2011; MacFadden et. al., 2012; Zazula 2009) and from mixed sand and gravel situations (MacFadden et. al., 2012; Van Kolfschoten et al., 2011), Common bone elements found in the fossil record include cranial and post cranial material (Burns 2010; Czaplewski, 2012; Harington, 2011). Mammut americanum Ancestors of all proboscideans in North America originated in Eurasia and crossed the Bering land bridge from Siberia to Alaska (Yansa & Adams, 2012). Proboscideans once thrived throughout Africa, Eurasia, and the Americas. Living elephants are the last remnants of a previously more diverse order. The proboscideans have been extensively researched as of late using modern and ancient mitochondrial DNA of order members determine phylogenetic relationships (Rohland et al., 2007). There are many reasons for this interest though the two which likely most affect this most are the availability of samples from the Pleistocene preserved intact in permafrost, particularly Siberia but in other locales as well, and the very recent, geologically, extinction of many members of this order (Rohland et al., 2007). It is generally believed the early ancestor Archidiskodon meridionalis migrated across the Beringia land bridge approximately 1.8 Ma. Mastodons, Mammut americanum, were the first to differentiate within these proboscidean groups, separating from Elephanidae about Ma. The Elephanidae, which include the extinct mammoths and modern elephants, do not begin to differentiate until Miocene-Pliocene boundary (Rohland et al., 2007).

243 224 In the context of this research, the mastodon referred to is the American mastodon (Mammut americanum) and the mammoth will be the woolly mammoth (Mammuthus primigenius). There were more mammoth species found within North America, and possibly Ohio as well. However, those which have been positively identified within the literature examined were all those of the woolly mammoth. It is for that reason all others have been excluded. In the Great Lakes region, mastodon and mammoth bones are more common than in anywhere else in North America. Areas such as Ontario, Canada, as well as Illinois, Indiana, Ohio, Michigan, Minnesota, and Wisconsin of the United States all have numerous bones and teeth found and recorded through history (Yansa & Adams, 2012). New York also has a large collection of proboscidean fossils and they represent the most abundant megafaunal taxa post- Late Glacial Maximum (Feranec & Kozlowski, 2012; Teale & Miller, 2012). Mastodons were very abundant compared to mammoths, 4 to 1 more abundant in Michigan and Ohio (Yansa & Adams, 2012). There are physical and ecological differences between mastodons and mammoths. Generally, mastodons are believed to be browsers of shrubs and trees while their cousin mammoths are believed to be grazers of grass and grass-like plants (Teale & Miller 2012; Yansa & Adams, 2012). However, they often appear contemporaneously in many areas around the Great Lakes region presumably because of the heterogeneous habitats present at this point within the Pleistocene (Feranec & Kozlowski, 2012; Yansa & Adams, 2012). Yansa and Adams (2012) purpose this close association within the Great Lakes region promoted interspecies competition for waning resources within the time period leading up to 13,000 cal yr as the latest mastodons and mammoths display fossil evidence of distress.

244 225 The M. americanum and M. primigenius are very different animals (Figure A). Mastodon have a more robust, stocky build with equidistant limb proportions in fore and hind legs. The skull is flatter and their tusks grew horizontally out from the jaws. The tooth, with two parallel rows of cusps running the length, is what granted the mastodon its distinctive name breast tooth. (Figure A). Early mastodonts arrived in North America around Ma and developed within this continent into M. americanum (Rohland et al., 2007; Teale & Miller, 2012; Yansa & Adams, 2012). The species would roam from the ice margins in the north southward into Florida, the Great Plains, as well as portions of the southwestern states and Mexico wherever suitable habitat was found (Teale & Miller, 2012; Yansa & Adams, 2012). The mastodon has been proposed to be victim of Clovis culture or the Younger Dryas event(s). However curiously enough, there are radiocarbon dates which suggest they survived well up to the end of the Pleistocene, most especially within the Great Lakes region (Laub, 2003; Teale & Miller, 2012; Yansa & Adams, 2012). A northern Indiana mastodon has yielded dates of about 11,615 cal yr and 11,600 cal yr indicating this mammal persisted through both the advent of Clovis and into the late Younger Dryas within the mid-continent (Woodman & Athfield, 2009). Mastodon remains are often recovered from areas with sediments rich in plant macrofossils and pollen indicative of open forest environment (Harington, 2011; Schubert et al., 2004; Teale & Miller, 2012; Yansa & Adams, 2012). In the western part of the United States, distribution of mastodons closely follows that of available forest and consequently, occurs at higher elevations (McDonald & Pelikan, 2006). The mastodon as a browser is well-established but dietary components are less certain as new data suggests the mastodon menu was more complex than initially supposed. It is generally

245 226 presumed that mastodons consumed mainly leaves and branches of spruce and other trees within a signature spruce parkland-sedge wetland environment before the habitat later became spruce dominated (Teale & Miller, 2012; Yansa & Adams, 2012). Typical conditions indicated are those of open white spruce (Picea glauca) forest with tamarack (Larix laricina), balsam fir (Abies balsamea), and pine (Pinus). Paper birch (Betula papyrifera), balsam poplar (Populus balsamifera), and willow (Salix) are also often present in quantity (Teale & Miller, 2012). Previous studies of stomach contents and dung support such a conclusion as spruce needles and twigs are frequent components, as are tamarack (Laub, 2003; Teale & Miller, 2012; Yansa & Adams, 2012). However, this may not be as straightforward as it appears since a growing body of evidence points to a more varied diet containing broadleaf and herbaceous plants as well (Teale & Miller, 2012). Stomach contents of mastodons from Florida contained deciduous tree and shrub materials, as well as fruits such as wild grape (Vitis) and gourd (Cucurbita pepo). Teeth from mastodons within the same region show enamel scarring consistent with eating bark and fruits. Stomach contents from an Ohio mastodon living within the spruce habitat contained aquatic species and pigweed (Amaranthus) but no conifer twigs or needles (Teale & Miller, 2012). Perhaps mastodons have more flexible dietary needs then first supposed, or one which was at least controlled more by seasonal variability and regionality than suspected. There is also initial evidence in isotopic analysis for mastodon migration, at least in the southeastern United States (Teale & Miller, 2012). Additional data supports the likelihood that mastodons, as with elephants and mammoths, utilized salt licks, seeps, and springs (Teale & Miller, 2012; Yansa & Adams, 2012). These appear in great quantities throughout the Great Lakes and the Upper South of the United States.

246 227 Analysis of mastodon dung revealed they ate mineral-rich clay. Southern Michigan had many of these mineral-rich deposits available and a considerable amount of fossil material has been found in association with numerous salt seeps and shallow saline springs (Yansa & Adams, 2012). Ohio also has mastodon fossils in association with salt springs, especially from the Scioto Saline Works. Fossils are often found in marls, clays, clay mixtures, peat and other organic sediments as these sediments are often co-occurrent (Feranec & Kozlowski, 2012; Harrington, 2011; Robinson et al., 2005; Teale & Miller, 2012; Whittecar et al., 2007; Woodman & Athfield, 2009), sands, and gravels (Harrington, 2011; MacFadden et. al., 2012; Robinson et al., 2005; Zazula et al., 2009). All portions of the skeleton are well-represented within the fossil record (Feranec & Kozlowski, 2012; Harington, 2011; Laub, 2003; Robinson et al., 2005; Teale & Miller, 2012; Woodman & Athfield 2009). Mammuthus primigenius The first appearance of mammoth currently defines the beginning of the Irvingtonian North American Land Mammal Age at about 1.4 Ma. By the late Pleistocene, mammoths become common and geographically extensive through North America, so much so that the mammoth is one of the primary index terrestrial fossils for the epoch (MacFadden & Hulbert, 2009). The genus Mammuthus originated in Africa during the Pliocene, then traveled northward, through Asia and across Beringia to North America (MacFadden & Hulbert, 2009) M. trogontherii from Asia developed into the Columbian mammoth (Mammuthus columbi) in North America around Ma (Debruyne et al., 2008; Yansa and Adams, 2012). M. columbi occupied the western United States and down into Nicaragua (McDonald & Pelikan, 2006; MacFadden & Hulbert, 2009; Yansa & Adams, 2012). There exists an east-west

247 228 gradient in North America between Columbian mammoths and American mastodons whereby mastodon density are highest in the eastern, more forested areas and decreases westward across the continent. The opposite occurs in mammoth density which are highest in the west and decreases eastward across the continent (McDonald & Pelikan, 2006). The second Beringian immigration came about 100 ka when the woolly mammoth (Mammuthus primigenius) arrived in Alaska from across the Bering land bridge (Debruyne et al., 2008; Elias & Crocker, 2008; MacFadden & Hulbert, 2009; Yansa & Adams, 2012). M. primigenius descended from M. trogontherii which remained in Siberia (Yansa & Adams, 2012). The woolly mammoth ranged from the southern border of the ice sheets to the middle of the United States and eastward to the Atlantic coast (Álvarez-Lao et al., 2009; McDonald & Pelikan, 2006; MacFadden & Hulbert, 2009; Yansa & Adams, 2012). Woolly mammoth are also very common in the Great Lakes region (Yansa & Adam, 2012). The youngest mammoths from the Great Lakes region are of 13,500 to 13,000 cal yr (Yansa &Adams, 2012). Intriguingly while the woolly mammoth would go extinct on mainland North America and Eurasia at the end of the Pleistocene, the species would survive on proximal islands for thousands more years (Boeskrorov, 2006; MacPhee et al., 2006). In fact, M. primigenius remains, literally numbering into hundreds of thousands, are one of the most common fossil finds from the northern and middle latitudes of North America and Eurasia as well (Álvarez-Lao et al., 2009). At some point in the Middle Pleistocene, the Jefferson mammoth (Mammuthus jeffersonii) appeared in the Midwest United States. However, this designation is uncertain as some researchers maintain the Jefferson mammoth is only a variant of the Columbian mammoth, or a hybrid of the Columbian mammoth and woolly mammoth while others insist it is a distinct species (Yansa & Adams, 2012). For the purpose of this research, only the woolly mammoth will

248 229 be considered as the Columbian mammoth did not appear in the Great Lakes region and documented occurrences of the Jefferson mammoth in Ohio are inconclusive. M. primigenius are considered to be closely related to living elephants (McNeil et al., 2005; Yansa & Adams, 2012). They were similar physically in that they had longer front legs than hind legs which created a sloping back with a hump behind the neck and over the shoulders. The skull was domed and tusks exited the mouth downward before curling up and outward (Figure A1). Teeth are similar to modern elephants with the occlusal surface composed of a series of tightly packed, transverse enamel plates (Figure A2). Mammoth were similar in size, though generally larger, to the African elephant (L. Africana) but their bones were more robust so mammoths may likely have been more massive (McNeil et al., 2005). Studies have indicated that despite this similarity of appearance, mammoths are likely more closely related to Asian elephants than African ones (Rohland et al. 2007). Most mammoth sites, about 90%, are located at higher elevation, often along lake shorelines (Yansa & Adams, 2012). Many of the attendant microfauna species found at mammoth sites modernly live within higher, and consequently cooler, latitudes and elevations suggesting that mammoths also must have preferred cooler climates at the minimum (Holen, 2006). During the Late Pleistocene, northern North America, middle to northern Europe and northern Asia where places which contained favored mammoth habitat of open steppe environment (Boeskrorov, 2006; Rivals, 2010). However, the Mammoth Steppe differs from the modern Asian significantly. Modernly, a steppe is defined as dry, cold grassland. It may be semi-desert, covered in grasses, or a combination of shrubs and grasses. The arid climate is too dry to support substantial forested areas but does not qualify as desert. Mammoth Steppe is extinct, its collection of fauna

249 230 and flora no longer globally exist anywhere. Some animal and plant species are extant but they no longer appear in the same distributions or combinations with each other as they did before the Pleistocene-Holocene transition. Instead, the Mammoth Steppe was replaced by modern tundra, tiaga, and steppe regions of North America and Eurasia (Rivals et al., 2010). Mammoth habitat was a montage with bits of tundra, open boreal woodlands, mixed conifer-deciduous forest and pine woodlands. There was greater diversity of flora with fewer trees just south of the ice sheets, and fostering an abundance of favored mammoth foods, grasses and sedges (Yansa & Adams, 2012). Mammoths have been interpreted as principally grazing animals both due to study of carcasses and associated faunal assemblages (McDonald & Pelikan, 2006; Yansa & Adams, 2012). Examinations of frozen mammoths divulge a diet largely, 90%, of grasses and grass-like sedges with smaller amounts of leaves and twigs of birch, alder, willow, and tamarack needles (Yansa & Adams, 2012). Other research depicts diets heavily concentrated in grasses and sedges, up to 99% (McDonald & Pelikan, 2006). Mammoth fossils are common in North America. They are found within fine-grained alluvial sediments such as sand and silt (Feranec &Kozlowski, 2012; Harington, 2011; Holen, 2006; MacFadden, 2012; Whittecar et al., 2007), clay deposits associated with open-water situations (Czaplewski, 2012; Feranec & Kozlowski, 2012; MacFadden et al., 2012; Van Kolfschoten et al., 2011), in coarser sediments, such as gravels, generally associated with active waterways such as banks, bars, and beach debris (MacFadden et al., 2012; Zazula et al., 2009), and in peat (Álvarez-Lao et al., 2009). Entire or near complete skeletons of mammoth are commonly found within North America (Álvarez-Lao et al., 2009; Harington, 2011; Holen, 2006; Teale & Miller, 2012,).

250 231 Disassociated cranial and post cranial material is relatively common (Álvarez-Lao et al., 2009; Czaplewski, 2012; Feranec & Kozlowski, 2012; Velivetskaya et al., 2011; Zazula et al., 2009). Pilosa Pilosa originate in South America and later colonized North America during the Great American Biotic Interchange. Pilosa included members which are small such as Megatherium altiplanicum, medium such as Megatherium sundti, and large Megatherium americanum. (De Iuliis et al., 2009). The endemic North American giant ground sloth (Megalonyx jeffersonii) ranged in the Pleistocene throughout the continent and even southward into southern Mexico (Hoganson & McDonald, 2007; McDonald et al., 2000; Schubert et al., 2004). This bear sized sloth ranged from coast to coast and an ungual from North Dakota dated from about 13,770 cal yr indicates it was still widely distributed even into the Late Pleistocene North America, including higher latitudes (Hoganson & McDonald, 2007). These sloths were large mammals with long unretractable claws indicating they were not fast moving individuals. They are frequently imaged as sitting bipedally, propped up by their large, stout tail and using their claws to pull down vegetation. M. jeffersonii would not have been easy prey for any predator. The large claws would have been formidable weapons and their hide was thick, much like that of the modern elephant (McDonald & Pelikan, 2006). Furthermore, ground sloth hides contained thousands of osteoderms, creating a sort of dermal armor (Borrero & Martin, 2012; McDonald & Pelikan, 2006). If M. jeffersonii resembles the Patagonian ground sloth Mylodon, the skeleton would have been difficult to disarticulate. The forelimbs of Mylodon are strongly articulated and its spine has additional articulations cognizant to its digging activities as the last few thoracic vertebrae and sacrum are firmly attached forming a more rigid skeletal element (Borrero & Martin, 2012).

251 232 It is generally accepted that giant ground sloths were solitary animals who probably came together only for the purpose of breeding (McDonald & Pelikan, 2006). Ground sloth finds are single individuals with no evidence of large family groups like mammoth, mastodons, and other herding mammals (McDonald & Pelikan, 2006). There is evidence of cave dwelling in the Patagonian ground sloth Mylodon but it is not yet known if these were living spaces in a traditional sense, maternal dens, or seasonally utilized for some other purpose (Borrero & Martin, 2012). There is also evidence for at least sporadic utilization of caves by M. jeffersonii in the United States (Schubert et al., 2004). The dissemination of M. jeffersonii at higher latitudes in the western part of its range resembles that of the elk-moose, giant beaver, and helmeted musk ox (McDonald et al., 2000). Generally, M. jeffersonii finds with M. americanum are also co-occurent with M. primigenius and rarely with only mastodon alone (McDonald & Pelikan, 2006). These taxa are also linked in the Pleistocene faunas from the eastern United States. This close association implies they shared similar ecological or environmental requirements despite dissimilar appearances and habits (McDonald et al., 2000). Evidence from related ground sloths (Mylodon darwini) in Fuego- Patagonia, South America suggests they occupied open habitats and had diets based upon grasses, sedges, and similar vegetation also akin to that favored by mammoths (McDonald & Pelikan, 2006). Preferred climate has been inferred to be cool and moist, as much of the vegetation associated with M. jeffersonii thrives within areas of high soil moisture (Borrero & Martin, 2012; Hoganson & McDonald, 2007; McDonald & Pelikan, 2006; Schubert et al., 2004). Hoganson (2007) reveals evidence of M. jeffersonii as a member of spruce-dominated forest in a riparian setting, browsing upon forest vegetation and likely using river valleys as migratory routes

252 233 (Hoganson & McDonald, 2007). However, it seems difficult to determine exactly what the preferred habitat for M. jeffersonii may be, as it was so geographically widely distributed and fossils are present in diverse depositional environments (Schubert et al., 2004). Ground sloth fossil elements are found in a variety of sedimentological situations such as gravels (Brandoni, 2011; Hoganson & McDonald, 2007; Schubert et al., 2004), beach sands and associated marls (MacFadden et. al., 2012; Schubert et al., 2004), peat, clays, silts, cave deposits, and tar pits (Schubert et al., 2004). As a result, ground sloth skeletal elements seem well represented with both cranial and post cranial material (Borrero & Martin, 2012; Brandoni, 2011; De Iuliis et al., 2009; Hoganson & McDonald, 2007; McDonald et al., 2000; Schubert et al., 2004). The hide and dung of giant ground sloths has even be found, though certainly less commonly than other mineralized parts (Borrero & Martin, 2012). Rangifer tarandus Extant Rangifer tarandus, caribou, persist throughout the Arctic Circle. Their range during the Pleistocene was considerably more expansive as within North America, R. tarandus fossils are known from as far south and east as Alabama by around 11.8 ka (Burns, 2010). Caribou are a frequent member of Mammoth fauna (van Kolfschoten et al., 2011). Subspecies of caribou have consistently been exploited through time to some degree by humans in the Northern hemisphere. From the Paleolithic period to modern day, caribou bones are common elements within archaeological sites where sufficient habitat supports, or supported, a population (Puputti & Niskanen, 2008; Robinson et al., 2005).The extent of R. taradus of the past seemed limited only by conditions permitting taiga forest habitat of tundra and conifer-based woods (Long & Yahnke, 2011; Schubert et al., 2004).

253 234 Pollen sequences from central Indiana bogs indicate the dominate habitats utilized by caribou ranged from those of spruce parklands to mixed conifer forests (Schubert et al., 2004). Replacement of these taiga-type forests to more open prairie conditions would have pushed out the caribou. Caribou migrated northward as the Wisconsinan glacier retreated, following their preferred boreal habitats (Long & Yahnke, 2011). Modernly, caribou feed upon leaves and shoots of willows (Salicaceae), sedges, rushes, and legumes. Lichens and mosses may also be consumed but do not seem to be preferred forage if given other selections (Kazmin & Abaturov, 2011; Kazmin et al., 2011). Caribou fossil material is rare and generally fragmentary in nature. Many records have not been adequately recorded in scientific literature. In fact, van Kolfschoten, et al. (2011) reported even the complete picture of body size variation within Middle Pleistocene R. tarandus remains unknown. R. taradus has been found in bog deposits and sandy tills (Long & Yahnke, 2011), Sands and gravels (van Kolfschoten et al., 2011), and from cave deposits (Robinson et al., 2005). Common fossil elements found are antler beams and fragments (Harrington, 2011; Laub, 2003; Long & Yahnke, 2011; R. Miller, 2000; van Kolfschoten et al., 2011; Zazula et al., 2009). Post cranial representatives are rare (van Kolfschoten et al., 2011; Zazula et al., 2009). To give some idea of the uniqueness of R. taradus finds, van Kolfschoten, et al. (2011) reported a collection from Woerden, the Netherlands, of antler fragments, three mandibles with some dentition, isolated premolars, several vertebrae, and a number of post cranial bones as an unusually rich record. The researchers also remark this was one of the few instances were R. taradus remains were a significant component of the find (van Kolfchoten et al., 2011). Tapirus spp.

254 235 Tapiroids likely originated within the Eocene of Asia and North America, later giving rise to the Tapirids (Eberle, 2005; Scherler et al., 2011). The oldest and most northernmost tapir lineage fossil, Thuliadanta mayri, hales from Ellesmere Island in the Canadian Arctic leaving the possibility of a North American origin. This suggests that tapirs both began in North America and at high latitudes before dispersing into Asia and throughout the mid-latitudes of the Americas (Eberle, 2005). Diversity of the lineage decreased through the Oligocene. There are only four extant species of the genus Tapiridae which persist in Central and South America and southeast Asia (Hollanda et al., 2011; Scherler et al., 2011). T. terrestris is widely distributed through the tropical habitats of Brazil, Argentina, Paraguay, Bolivia, Peru, Ecuador, Columbia, the Guianas, and Venezuela. T. bairdii extends from southern Mexico and Central America to Columbia and Ecuador. T. pinchaque lives within the Andes of Columbia, Ecuador, and Peru. Only T. indicus is found in southeast Asia (Holanda et al., 2011). Fossil tapirs are rare but may be found throughout North, Central, and South America as well as Asia (Eberle, 2005). Extant tapirs are solitary creatures living primarily within forested areas in warm, humid climates proximal to water sources. Dentition and a short proboscis promote a herbivorous, browsing form. As fossil tapirids share these same characteristics, it is presumed they share this condition (Scherler et al., 2011). Additionally, the lithology of many fossil localities seems to indicate extinct tapirs inhabited lushly vegetated environs close to water (DeSantis, 2008; Eberle, 2005; Scherler et al., 2011). As a result, tapir fossils are frequently found within marls, coquinas, conglomerates, and sand (Hollanda et al., 2011; Hollanda & Ferrero, 2013; MacFadden et al., 2012; Scherler et al., 2011). Generally, the tapir fossil record is sparse. Common elements found are mandible and teeth (Eberle, 2005; Hollanda et al., 2011; Hollanda & Ferrero, 2013; Scherler

255 236 et al., 2011). Occasionally, crania and postcranial bones are found (Hollanda et al., 2011; Hollanda & Ferrero, 2013; Scherler et al., 2011). Tayassuidae Tayassuidae were one of the first immigrants into South America during the Great American Biotic Interchange, though the exact timing the arrival is controversial at this time (Gasparini, 2013). Modernly, Tayssuidae are distributed throughout the southwestern United States and down into north-central Argentina and represented by two genera and three species. In the past, taxonomic diversity and geographic distribution were greater as there are records of them in Asia, Europe, Africa, as well as throughout the Americas (Gasparini et al., 2010). Frailey and Campbell, Jr. (2012) suggests the genera Pecari and Tayssu developed within tropical forests instead of temperate deciduous forests, or subtropical savannahs based upon their skull morphology. The short, robust skull would have been more beneficial in rooting through leaf litter and heavier clayey soils for nuts, fruits, and tubers. There are distinct differences in craniomandibular features between rainforest and desert populations of Pecari which have been related to the more resistant plant foods available in tropical rainforests (Frailey & Campbell, Jr., 2012). Modern tayassuines occupy a diversity of environments, from open range lands to forested areas with a resulting diverse array of food stuffs utilized. Dental microwear and isotopic analysis evidence suggests members of the family Tayassuidae lived on an herbivorous diet with probable foraging habits, such as that of a C 3 mixed feeder (Gasparini & Ubilla, 2011). The most commonly occurring late Pleistocene age fossil peccaries in North America are those of the Flat-headed (Platygonus compressus), and Long-nosed (Mylohyus nastus) peccaries. The extant White-collared (Pecari tajacu) peccary is only known from Holocene deposits (Czaplewski, 2012).

256 237 M. nastus was primarily an eastern North American species with only a few specimens as far west as Texas and Kansas and into the south and central Great Plains (Czaplewski, 2012). Research suggests P. compressus are adapted to drier and more open, grassy habitats with scattered trees (Gasparini, 2013; Gasparini & Ubilla, 2011). Open and arid environments which developed during glacial cycles would have allowed latitudinal expansion of P. compressus. Catagonus wagneri replaced P. compressus around the middle Pleistocene as open environments to which P. compressus were better adapted shrank (Gasparini, 2013). Sedimentary data available for fossil peccaries suggests they frequented coastal settings such as mangrove, marsh, fluvial, woodlands, and arid tropical forested habitats (MacFadden, et. al., 2010). Other sedimentary evidence suggests at least some Tayassuidae species utilized dense canopy, temperate oak-hickory forests (DeSantis 2008 and MacFadden, et. al., 2010). Peccaries are also often rarer elements of fossil localities, though there are exceptions to this (MacFadden et. al., 2010). Peccary fossils have been found in riverine lag deposits, beaches, creek beds, and coastal deposits from deltaic to nearshore (Czaplewski, 2012; DeSantis, 2008; Frailey & Campbell, 2012; MacFadden, et. al., 2010; MacFadden, et. al., 2012), relatively common as karst deposits, along with larger carnivores, suggesting they are often subjects of predator traps (Czaplewski, 2012; DeSantis 2008; Gasparini 2012; Smith and Polly 2013), and conglomerates from fluvial and volcanoclastic situations (Frailey & Campbell, 2012; MacFadden, et. al., 2010). Most common fossil elements are crania, teeth (including tusks), and mandibles (Czaplewski, 2012; Frailey & Campbell, 2012; Gasparini et al., 2010; Gasparini & Ubilla, 2011; Gasparini, 2013; MacFadden et. al., 2012). Some post cranial skeletal elements are found

257 238 (Czaplewski, 2012; Frailey & Campbell, 2012; Gasparini, 2013). Occasionally, partial skeletons are found (Gasparini et al., 2010).

258 239 Figure A1 Comparison of mammoth (left) and mastodon (right). Image was created in Photoshop by Dantheman9758. Image available as opensource illustration on the internet. Figure A2 Comparison of mastodon molar (left) and mammoth molar (right). Photos available as open-source images on the internet.