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2017 Late of the flat-headed on the Ozark Plateau: Paleozoological insights from Peccary Cave, Kurt M. Wilson Iowa State University

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Recommended Citation Wilson, Kurt M., "Late Pleistocene extinction of the flat-headed peccary on the Ozark Plateau: Paleozoological insights from Peccary Cave, Arkansas" (2017). Graduate Theses and Dissertations. 15461. https://lib.dr.iastate.edu/etd/15461

This Thesis is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Graduate Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Late Pleistocene extinction of the flat-headed peccary on the Ozark Plateau: Paleozoological insights from Peccary Cave, Arkansas

by

Kurt M. Wilson

A thesis submitted to the graduate faculty

in partial fulfillment of the requirements for the degree of

MASTER OF ARTS

Major: Anthropology

Program of Study Committee: Matthew G. Hill, Major Professor Alan D. Wanamaker Jill D. Pruetz

The student author and the program of study committee are solely responsible for the content of this thesis. The Graduate College will ensure this thesis is globally accessible and will not permit alterations after a degree is conferred.

Iowa State University

Ames, Iowa

2017

Copyright © Kurt M. Wilson, 2017. All rights reserved.

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TABLE OF CONTENTS

Page

LIST OF FIGURES ...... iv LIST OF TABLES ...... v ABSTRACT ...... vi ACKNOWLEDGMENTS ...... vii

CHAPTER 1. INTRODUCTION 1

CHAPTER 2. LATE PLEISTOCENE 3 Overkill and Human Causation ...... 3 Climate Change and Non-Human Causation ...... 5

CHAPTER 3. FLAT-HEADED PECCARY 7 General Characteristics ...... 7 Geographic Distribution ...... 8 Temporal Distribution ...... 9 Behavior ...... 10 Diet...... 11 Archaeological Associations ...... 11

CHAPTER 4. PECCARY CAVE 12 Geologic and Sedimentary Setting ...... 12 Excavations ...... 13 Previous Research on the Faunal Remains ...... 14 Environmental Context ...... 15 Age of the Faunal Remains ...... 19

CHAPTER 5. PALEOZOOLOGY 20 General Assemblage Characteristics ...... 20 Skeletal Element Representation ...... 21 Minimum Number of Elements ...... 22 Minimum Number of Individuals ...... 23 Assemblage Age Composition ...... 24 Carnivore Damage ...... 25 Density-Mediated Attrition ...... 25 ...... 26 Discussion ...... 27 Summary ...... 29

CHAPTER 6. STABLE ISOTOPES 31 Methods ...... 33 Results ...... 34 Discussion ...... 36 Summary ...... 39

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CHAPTER 7. CONCLUSIONS 41 The Uncommon and Unadaptable Flat-Headed Peccary ...... 41 Modern Peccary Reproductive Biology ...... 44 Extinction of Flat-Headed Peccary on the Ozark Plateau ...... 45

REFERENCES CITED ...... 64

APPENDIX A. RECORDS OF FLAT-HEADED PECCARY ...... 88 APPENDIX B. SCAN SITE MNES AND VOLUME DENSITIES ...... 95 APPENDIX C. RAW STABLE ISOTOPES DATA ...... 96

iv

LIST OF FIGURES

Page

Figure 1. Location of Peccary Cave and other selected places on and around the Ozark Plateau ...... 49

Figure 2. Location of published records of peccary in the contiguous ...... 50

Figure 3. General character of peccary assemblages ...... 51

Figure 4. Radiocarbon ages for peccary ...... 51

Figure 5. MNI estimates for peccary assemblages ...... 52

Figure 6. Location of Peccary Cave at the confluence of Ben’s Branch and Cave Creek ...... 53

Figure 7. Plan map of Peccary Cave, including excavation trenches and sections ...... 53

Figure 8. AMS radiocarbon ages for selected taxa on the Ozark Plateau ...... 54

Figure 9. AMS radiocarbon ages for peccary and dire wolf ...... 54

Figure 10. Box-and-whisker plot of mandibular third molar length ...... 55

Figure 11. Examples of breccia coverage ...... 55

Figure 12. Skeletal element profiles for sub/adult and fetal/neonate peccary ...... 56

Figure 13. Distribution of selected classes of faunal and archaeological material ...... 56

Figure 14. Stable isotope values for peccary and dire wolf ...... 57

Figure 15. Intratooth δ13C and δ18O profiles for peccary mandibular canines ...... 57

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LIST OF TABLES

Page

Table 1. Direct dates for peccary ...... 58

Table 2. Ozark Plateau environment from Pre-LGM through the Oldest Dryas ...... 59

Table 3. Radiocarbon AMS measurements and δ13C results on peccary and dire wolf ...... 59

Table 4. Summary of rodent modification ...... 60

Table 5. Summary of breccia coverage ...... 60

Table 6. Summary of peccary remains ...... 61

Table 7. Summary of sub/adult peccary teeth ...... 62

Table 8. Summary of proximal epiphyseal union for sub/adult peccary calcanea ...... 62

Table 9. Spearman rank-order correlations (rs) between scan site MNE and volume density ...... 63

Table 10. Records of litter size in modern ...... 63

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ABSTRACT

Paleozoological research on flat-headed peccary ( compressus) and dire wolf (Canis dirus) remains from Peccary Cave, Arkansas, provides information on the ecology and extinction of these taxa. The peccary assemblage is large, totaling nearly 4,000 specimens, and includes all skeletal elements, while the dire wolf assemblage includes only a handful of specimens. A minimum of 61 sub/adult peccaries, 15 fetal/neonatal peccaries, and 3 dire wolves are represented in the collection. Peccaries died naturally in the cave. The dire wolf remains were transported into the cave by colluvial or fluvial processes. Five direct AMS ages on peccary and three direct AMS ages on dire wolf bone range between approximately 25,500 cal B.P. and 21,300 cal B.P., and indicate both assemblages are time-averaged accumulations. Dire wolves did not den in the cave or transport peccary carcasses there, however bulk stable isotopes indicate that dire wolves likely preyed on peccaries regularly. Serial isotopic sampling of two peccary canine teeth paired with stable isotope values from bone indicates the foraged primarily on C3 plants year-round, mostly legumes. Early Late Glacial Maximum conditions on the Ozark

Plateau were warming and drying, which contributed to deficits in peccary reproduction and, ultimately, to the regional extinction of the taxon, and by extension, dire wolf.

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ACKNOWLEDGMENTS

Many people contributed to this project and deserve acknowledgement for helping me see it through to completion. Mary Suter (University of Arkansas Museum) arranged the loan that made the research possible, including providing copies of the documentation associated with the extant collection.

Leo Carson Davis (Magnolia, Arkansas) and Holmes Semken (University of Iowa) shared their knowledge of the field investigations, the extant collections, and previous research on material from the cave. Iowa State University undergraduate students, Natalie Hassebrock, Madison Pullis, and Daniel

Dalmas, clocked many volunteer hours getting the collection in good working order for analysis. John

Lambert, University of (Davis) (and a former Iowa State University undergraduate student), spent several days in Ames helping analyze the chipped stone artifacts. Jim Theler, University of

Wisconsin (La Crosse), and Laura Halverson-Monahan, University of Wisconsin Zoological Museum

(Madison), provided comparative skeletal material that was invaluable. The isotope work would not have been possible without the assistance of Chris Widga (East Tennessee State University), Al Wanamaker,

Suzy Ankjerstine, and Betsy Swanner (all Iowa State University), who were incredibly generous in sharing their time, knowledge, experience, and laboratory resources. My committee, Drs. Matthew G.

Hill, Jill D. Pruetz, and Alan D. Wanamaker, was instrumental in all stages of the research, for which I am most grateful. Generous financial support was provided by a Miller Fellowship, the Graduate College, the College of Liberal Arts and Sciences, and the Department of Anthropology. 1

CHAPTER 1. INTRODUCTION

Global mass extinction of occurred during the late Pleistocene. In , 35 genera disappeared around or just prior to the time the first arrival of people on the landscape, which spawned debate and much research on the cause of extinctions (Martin and Klein 1984). For North

America, the prevailing explanations revolve around climate change (Graham and Lundelius 1984;

Graham and Mead 1987; Grayson 2007), though some researchers attribute the extinctions to “overkill” by human hunters (Fiedel and Haynes 2004; Lyons et al. 2004; Martin 1967, 1984). The timing of the extinctions is also debated, with some scholars arguing for a synchronous extinction event, while others contend that they occurred over the course of tens of thousands of years (Grayson 2007).

Much research has focused on extinction event generally or on several ‘spectacular’ megafauna, for example, and mastodon (Widga et al. 2017), with relatively little attention given to some other taxa, for example, small- and medium-sized . This is not an ideal situation because, as Grayson notes, arguably the most effective method for understanding late Pleistocene extinctions as a whole involves first “reconstructing the histories of individual taxa, making no assumptions about the chronology of extinction” (2007:198). One animal that has received limited attention is the flat-headed peccary (), which is somewhat surprising since it has been described as “probably the most numerous medium-sized during the late Pleistocene”

(Kurten and Anderson 1980:295). At first blush, the animal does appear to have been relatively common, however, close inspection of the evidence reveals that this may not be the case. Save for several large accumulations of peccary remains from caves in the eastern United States, most records for the taxon consist of isolated specimens or a small handful of remains. Moreover, the distribution of the animal in space and, especially, in time, is not well documented. To date, 25 direct dates on peccary bone from 15 localities are available for assessment, including 5 new assays reported here. These dates are from the eastern United States, except for one from New and one from . 2

Another matter is that peccary has been recovered from late Pleistocene archaeological contexts at several locations. While it would seem that the animal would have been easy prey for prehistoric hunters, direct evidence for exploitation occurs at only Sheriden Cave, (Redmond and Tankersley

2005:524). This raises the possibility that humans may have somehow contributed to their extinction.

Alternatively, as Guilday et al. (1971) noted years ago, given their inferred numbers on the landscape, their inferred widespread distribution, and the fact that modern peccaries occupy a variety of habitats, flat-headed peccary ought to have been relatively resistant to extinction triggered by climate change. The question of why these animals went extinct is thus central to understanding the bigger picture of late

Pleistocene extinctions.

Provided this background, the objective of this research is to shed fresh light on the ecology and extinction of flat-headed peccary through comprehensive analysis of the large and previously unreported assemblage of flat-headed peccary remains from Peccary Cave in northeast Arkansas (Davis 1973; Quinn

1972; Semken 1985) (Figure 1). A handful of dire wolf (Canis dirus) remains from the cave also figure into to the research. Multiple lines of evidence are used to explore these topics. Skeletal element frequencies supply information on taphonomic history of the material, including, in particular, how the cave was used by peccaries and dire wolves. Accelerator mass spectrometry (AMS) radiocarbon ages on peccary and dire wolf bone provide a temporal context for the accumulation and the taxa. Stable carbon, oxygen, and nitrogen isotopes reveal dietary patterns and trophic relationships for peccaries and dire wolves. Review of aspects of modern peccary ecology provides a context for understanding how climate change may have contributed to extinction of flat-headed peccary.

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CHAPTER 2. LATE PLEISTOCENE EXTINCTIONS

Since the discovery of extinct megafauna in North America, the question of what caused their extinction has garnered significant attention. Over the past 50 years two primary theories on extinction have emerged, presenting humans and climate change as the active agents driving loss of fauna (Martin

1967; Graham 1985a). Though at least one other large-scale hypothesis has been posited, a meteorite/comet impact (Firestone et al. 2007), most hypotheses revolve around overkill, climate or environmental change, or some combination of the two. Despite years of debate, neither overkill apologists nor climate change proponents have managed to provide indisputable proof they are correct

(Haynes 2009; Grayson 2016). Due to the ongoing debate over the cause of extinction it is necessary to provide a brief overview of the main arguments to situate this research.

Overkill and Human Causation

The most well-known of the overkill arguments is the “blitzkrieg” model proposed by Paul

Martin (Martin 1967, 1973, 1984; Martin and Steadman 1999). Martin’s arguments hinge on naïve fauna in a landscape previously untouched by humans. As hunters entered North America from they encountered fauna which had no knowledge or concept of humans and the danger they could present. As a result these naïve fauna had no defensive mechanisms and were ripe for easy killing. According to this model, humans entering North America engaged on a killing spree of these naïve fauna, slaughtering their way rapidly from west to east across North America, wiping out most megafauna in their path. The model was developed on the premise of human caused extinctions on islands and viewing the North American continent as one large island. Arguing North America as an island presents problems in that islands have limited space and ecosystems which is not true of a continent. Further problems emerge as proving

“blitzkrieg” false is impossible because the argument claims no evidence would be left as mass kills on the surface would not have been preserved (Haynes 2009). Since it cannot be falsified it functionally cannot be proved.

Despite the arguments against the “blitzkrieg” overkill model it remains a popular explanation for

Pleistocene faunal extinction. Burney and Flannery (2005) proposed overkill could account for 4

extinctions due to temporally stepped megafaunal collapses which appear to closely correlate with the first human appearances in landscapes around the world. In their view, climate change does not coincide with all extinctions and overkill thus presents a better model. Lyons et al. (2004) argued human was more impactful on extinction than climate change based on the size of creatures that do not go extinct. This is partially based on modern extinctions, where hunters tend to threaten large game much more so than small game. Alroy (2001) also argues for overkill, using computer simulations incorporating numerous factors to determine that over hundreds of years thousands of human hunters could have killed most of the extinct in North America. Some of the key factors in the model are increasing human population and improving efficiency which would, over hundreds of years, overtake megafauna reproductive capabilities and eventually drive them to extinction.

Several other models have been generated focusing on human causation but not directly tied to human predation on herbivores. Haynes (2002, 2006) proposed a mixed model wherein killing during first human contact combined with climate stress caused by human actions in the environment, such as burning, would drive extinction. Another argument, based on human arrival on Pacific Islands, posits the loss of many species of birds on Pacific Islands was due only in part to overkill and driven equally as strongly by human depletion of habitat through land clearance and predation by rats and dogs which were introduced by humans (Steadman 2007; Steadman et al. 2002). Finally, Whitney-Smith (2004) provided a second-order predation model where humans entering the North American landscape killed off apex predators allowing herbivore populations to boom. These booms resulted in herbivore populations exceeding environmental holding capacities resulting in mass die-offs and extinctions.

As Grayson and Meltzer (2002, 2003, 2004, 2015) point out there are problems with the overkill hypotheses and models. Primarily, there is a lack of evidence for human predation on many of the extinct animals, with only 15 sites providing compelling evidence for human use of fauna, the majority of which are mammoth sites (Grayson and Meltzer 2002, 2015). Further issues arise in dating the fauna as, despite arguments put forth by Fiedel (2009), current quality dates for extinction show some fauna go extinct before humans enter the landscape (Grayson and Meltzer 2015). 5

Climate Change and Non-Human Causation

The alternate argument to extinction caused by humans tends to be extinction by late Pleistocene climate change. Russell W. Graham and Ernest L. Lundelius have been key proponents of this argument

(Graham 1985a, 1985b, 1986; Graham and Lundelius 1984; Graham and Mead 1987; Lundelius 1988,

1989). Their argument is based around changes in wetness, temperature, and seasonality. During the

Pleistocene the climate was undoubtedly cooler than the succeeding but was also wetter due to lower temperatures. The climate was more equable with smaller seasonal temperature shifts. Near the end of the Pleistocene this changed with the climate becoming warmer and dryer with greater seasonal temperature extremes. Such changes led to a decrease in plant community diversity which could no longer support diverse mammalian fauna leading to extinction (Graham and Lundelius 1984).

More recently Graham (2001, 2003) showed ecosystems were continually stressed and impacted during each glacial and interglacial period. Repeated stress to the environment eventually fragmented floral and faunal communities, impacting the ability to maintain viable populations and resulting in ecosystem collapse and extinction.

Guthrie (1982, 1984) provides similar arguments showing that a changing climate would have changed growing seasons. Pleistocene growing seasons were lengthy and allowed fauna to accumulate large amounts of nutrients. As the climate changed the growing season shortened and the heterogeneous plant landscape of the Pleistocene gave way to a more homogenous plant community. Thus less time was available for nutrient accumulation and greater homogeneity prevented some fauna from finding preferred food sources, the stress of which could have caused extinction for taxa that could not adapt. An example of this was shown by Grayson and Delpech (2005) who demonstrated warming during the end of the

Pleistocene may have caused the extirpation of in southern France as increased summer temperatures impacted forage and had a direct negative effect on reindeer and caribou populations.

Not all arguments for non-human caused extinction explicitly claim climate change for causation.

Owen-Smith (1987) postulated key species like mammoth died off, cause unknown. Their death could have caused environmental changes on a mass scale as may have served as environmental 6

engineers, and their absence enabled environmental change leading other fauna to extinction. Widga et al.

(2017) showed mammoth and mastodon extinction in the midcontinent may be related to trophic control failure. The loss of apex predators, whose extinction is not well-understood, resulted in boom-bust population cycles in proboscideans. Such boom-bust cycles caused the populations to become less resilient to climate and ecological changes. Climate change may have driven the final extinction but the loss of trophic controls initiated the process (Widga et al. 2017).

Like overkill, climate change hypotheses have weaknesses. Primarily, a lack of well constrained regional extinction dates means it is not possible to fully determine terminal dates for most species. This dearth of dates leaves open the plausibility of extinction occurring rapidly during human occupation or staggered, with some extinctions before and others after arrival of humans. Further, shrinking and fracturing range arguments make for compelling extinction possibilities but some evidence exists for range expansion for creatures such as mammoths immediately prior to their extinction (Agenbroad 2005).

In general, a lack of well-developed regional environmental reconstructions combined with quality radiocarbon dates holds back climate change arguments.

As the primary cause of megafauna extinction at the end of the Pleistocene remains unknown it is necessary to continue to investigate the extinction of individual species (Grayson 2007) to determine regional patterns. Such knowledge will contribute to filling the gaps that persist in our understanding of extinction causes for many Late Glacial Maximum (LGM) and Late Pleistocene species.

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CHAPTER 3. FLAT-HEADED PECCARY

General Characteristics

The flat-headed peccary (Mammalia – Artiodactyla – Tayassuidae – Platygonus compressus) was first reported by LeConte in 1848 (LeConte 1848) and, several years later, described in detail by Leidy

(1853). Indications are it was “about the size of a European (Sus scrofa)” (Kurten and Anderson

1980:300), though somewhat taller and leggier. It is related to extant New World peccaries (Tayassuidae), most closely to the (Catagonus wagneri) (Mayer and Wetzel 1986). Several distinctive features include large canine teeth, fused radius-ulna, and a relatively wide, short, and flat cranium set over a relatively flat, robust mandible (LeConte 1848; Leidy 1853, 1889). Sexual dimorphism has proven to be difficult to detect, and like modern peccaries, males and females were approximately the same size

(Sowls 1997; Woodruff 2016; Wright 1993).

The flat-headed peccary (and the Chacoan peccary) possess only two hind toes, relatively higher set eyes, and longer limbs compared to other extant peccary (Guilday et al. 1971; Mayer and Wetzel

1986). Guilday et al. (1971) report that the eyes of the flat-headed peccary faced outwards and upwards and remained more horizontal during browsing, suggesting that the animal relied heavily on vision for survival. The eye placement further suggests a preference for open environments. Relatively short proximal forelimbs and relatively long distal limbs also point toward adaptation to open environments and fleetness.

The nasal cavity is also highly complex and well developed (Leidy 1853), and suggests that, in addition to excellent vision, the flat-headed peccary likely possessed excellent olfactory capabilities, perhaps for locating food and mates as well as monitoring their surroundings for predators. Although the animal was undoubtedly omnivorous like its extant relatives (Bodmer 1989a), the teeth and chewing apparatus suggest it was largely a browser that foraged primarily on tough, fibrous foods (Guilday et al.

1971).

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Geographic Distribution

Two online databases – FAUNMAP and NEOTOMA – were used as starting point to document the geographic distribution of flat-headed peccary in the contiguous United States. The primary literature associated with each record of Platygonus sp. was checked to ‘scrub’ the data as well as to collect basic quantitative and qualitative information that is not included in either database. The effort revealed minor problems with both databases, ranging from duplicate entries (resulting from use of synonyms for the same locality) to inconsistencies in /species designations between the primary literature and that reported in either FAUNMAP and/or NEOTOMA (excluding taxonomic synonyms). Several records were not able to be verified, and several published records were missing, which is not surprising given that they lurk in obscure places. A handful of records post-date compilation of NEOTOMA and are incorporated here. All told, there are 91 verifiable records of flat-headed peccary (P. compressus or

Platygonus cf. compressus) from 88 localities (Appendix A). The number drops to 83 records from 81 localities if only material classified as P. compressus is tallied.

Several patterns are apparent in the distribution of the records (Figure 2). First, they generally occur south of a diagonal line that stretches from New York and to . The finds in southeast Idaho and in eastern Wyoming/western are noteworthy, northerly exceptions to this general pattern. Second, the majority of records occur: 1) on and around the Ozark Plateau, which centers roughly on southern ; 2) the Driftless Area of southwestern Wisconsin and northwestern

Illinois; 3) the Interior Low Plateau in Tennessee and as well as the Allegheny Plateau in Ohio; and 4) the Appalachian Mountains. Third, the Southeast is virtually devoid of records, except for about a dozen localities in . Fourth, a handful of widely scattered records occur in the Southwest and southern California.

The general nature of these records also deserves comment. Nearly half of them are represented by ten or fewer identified specimens (NISP) (n = 43) (Figure 3). Seventeen records (and 17 localities) are represented by only one specimen, and another 10 records (and 10 localities) are represented by only two specimens. Three assemblages contain between 100 and 500 specimens, while three assemblages include 9

between 501 and 1,000 specimens. Eight assemblages contain more than 1,000 specimens, for example,

Guy Wilson Cave, Welsh Cave, Zoo Cave, Cherokee Cave, and Peccary Cave. Several localities are represented by what are loosely described as ‘herds’ that contain the remains of three or more individuals.

Since the minimum number of individuals (MNI) represented in assemblage is derived from the pool of identified specimens (Lyman 2008), there is also strong correlation between the assemblage size and the number of peccaries represented therein. The vast majority of the assemblages include the remains of two or fewer individuals (n = 49) (Figure 4). It seems probable that the most of the assemblages that lack MNI data (n = 19) also contain very few animals; if this supposition is accurate, then roughly three-quarters (68 of 91) of the assemblages are contain approximately 1-2 individuals. Most of the balance of the sample is comprised of assemblages with MNIs between 5 and 50. Three assemblages include the remains of between 51 and 100 individuals, these being Peccary Cave (MNI =

61), Bat Cave (MNI = 70), and Zoo Cave (MNI = 81). The greatest number of individuals is found at

Megenity Peccary Cave, which contains the remains of over 500 peccaries.

Temporal Distribution

Absolute dates for flat-headed peccary are far between and few in number. Currently, there are

25 radiometric assays from 14 localities on peccary bone, including the five AMS radiocarbon ages reported here from Peccary Cave. Conventional and AMS radiocarbon ages are represented. These data are summarized in Table 1 and displayed in Figure 5. All of the dates are from the eastern United States, except for one result from Oven, . The current terminal age for the taxon is 10,750 ± 50 B.P. on a specimen from near Gainesville, New York (Feranec and Kozlowski 2010). The result for Fremont,

Ohio, is highly suspect and is not considered to be reliable (Hoare 1964). The relatively young results for

Brynjulfson Cave, Missouri, and Baker Bluff, Tennessee, are also suspect. Not only are these conventional radiocarbon ages but they are also considerably younger than the (more reliable) AMS ages for Peccary Cave and ecological inferences drawn from patterning in AMS assays on other taxa from

Ozark Caves (this is discussed in greater detail below). The most recent and reliable indirect age is from 10

Agate Basin, Wyoming, where a mandibular condyle was recovered from a Folsom archaeological component (Walker 1982) dated to 10,430 ± 25 B.P. (Surovell et al. 2016:84-85).

Among the oldest absolute dates for the taxon are four conventional assays with very large standard errors from Guy Wilson Cave, Baker Bluff, Patton Cave, and Brynjulfson Cave. The youngest date among these sites is a 19,000 B.P. result from Guy Wilson Cave, while the oldest is a 34,600 B.P. result from Brynjulfson Cave. Five AMS ages from Peccary Cave range between roughly 20,600 B.P. and

18,000 B.P., and are regarded her to represent the oldest reliable ages for flat-headed peccary.

It currently difficult to ascertain much in the way of spatio-temporal patterning in the radiocarbon ages. The most that can be mustered is that the oldest dates occur at Peccary Cave, and dates in the ballpark of 11,000 B.P. occur at three, widely separated places: Butterworth, Idaho; Sheriden Cave,

Ohio; and Gainesville, New York. Clearly, additional radiocarbon dates are needed to elucidate direction(s) and rate(s) of extinction across the range (Figure 2).

Behavior

Observations of the behavior of modern peccaries and inferences drawn from study of flat- headed peccary remains and localities provides some insight into aspects of the latter’s behavior. Modern peccaries are gregarious animals that routinely travel in herds of 6-15 individuals in open environments

(Sowls 1997:143-145). Evidence for similar behavior in flat-headed peccaries comes from several remarkable discoveries where small groups of animals apparently died together (Figure 3) (Finch et al.

1972; Hoare et al. 1964; Klippart 1875; Wagner 1903; Williston 1894). At Chickasaw Bluffs, for example, it appears as though the largest animal led the group, followed by three subadults, which, in turn, were trailed by another adult (Finch et al. 1972), similar to that reported in modern peccaries when protecting vulnerable individuals (Sowls 1997).

Modern peccaries also regularly use natural topography for protection against extreme temperatures (Sowls 1984), particularly caves and fissures to escape cold nighttime temperatures. Such locations were also used regularly by flat-headed peccaries. While large carnivores likely contributed peccary remains to some caves and fissures, the large number of peccaries at places like Megenity 11

Peccary Cave and Peccary Cave (Figure 4), for example, leave little doubt that these locations were used regularly and that individuals periodically died in them.

Diet

LeConte’s (1848) early observation that flat-headed peccary possessed teeth used primarily to handle coarse, fibrous food is supported by more recent tooth microwear and functional morphology research suggesting the animal was predominantly a horizontal browser (Graham and Lundelius 1984;

Guilday et al. 1971; Schmidt 2008). Although the dentition clearly belongs to an , and modern peccaries in the rain forest consume an astounding range of food (Bodmer 1989a), there is no firm evidence that other foods were a regular part of flat-headed peccary diet. In fact, Guilday et al. (1971:311) reasoned that the animal was primarily an herbivore despite their tooth morphology and omnivorous capability. Recent isotopic work supports this inference and points to a diet focused on C3 browse or

CAM-using plants (Giesler 2002; MacFadden and Cerling 1996; Yann and DeSantis 2014).

Archaeological Associations

Associations of archaeological material and the remains of extinct animals central to assessing the role that people may have played in the loss of these animals (Grayson and Meltzer 2002). To date, remains of flat-headed peccary occur in association with archaeological material at several places. Several of these association are easily dismissed because the archaeological material is much too recent to be tied to peccary exploitation by human predators (e.g., Guilday et al. 1971; Richards and Munson 1988).

However, there are three co-occurrences that deserve brief mention. One case is Agate Basin, Wyoming, mentioned above, another is Lubbock Lake, , where a mandibular premolar was recovered from the

Clovis component. Perhaps the most compelling evidence for possible human exploitation of flat-headed peccary comes from Sheriden Cave, Ohio, where remains of the animal were recovered in close association with Clovis artifacts in a sediment package dated between 11,130 ± 60 B.P. and 11,060 ± 60

B.P. (Tankersley 1997:718, 1999:Table 9.4). However, is no direct evidence (e.g., cut marks) that Clovis hunters exploited these animals (Waters et al. 2009:110).

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CHAPTER 4. PECCARY CAVE

Peccary Cave is located in northwest Arkansas on the southern Ozark Plateau (Figures 1 and 6).

It was discovered in January, 1965 by Jack McCutcheon, who lowered himself into a chimney and encountered numerous faunal remains (Davis 1969; Quinn 1972). The cave, which totals about 1,000 ft. of passages, and its deposits were subsequently investigated between September 1967 and August 1969 by a team of researchers led by James H. Quinn and L. Carson Davis. Additional work at the site was done in the summers of 1971 and 1972 (Semken 1984). The combined work yielded a large faunal collection that likely totals over 10,000 specimens and includes a diversity of micro-, small-, medium-, and large-sized taxa, the most visible of which are the peccary remains, which totals over 4,000 specimens. A small collection of chipped stone artifacts was also recovered. It includes 318 pieces of debitage and 107 formal and informal tools. Several temporally diagnostic projectile points (Dickson

Cluster) suggests most of the archaeological material is Woodland in age (Justice 1987). The human occupation of the cave is beyond the scope of this research.

Geologic and Sedimentary Setting

The cave is situated in limestone on the north-facing hillside near the confluence of

Ben’s Branch and Cave Creek, a tributary to the Buffalo River (Ball and Davis 1993; Quinn 1972). The original entrance is estimated to have been about 9 m across and 2 m high (Figure 7), and was sealed with alluvium around 2,200 B.P. Beneath a 2 m open chimney is an 8 m-high talus cone. Most of the debris entering via the chimney was contained in the immediate alcove into which it fell. Other interior portions of the cave have experienced roof-rock falls rather than surface deposition (Davis 1969).

The sediment within the cave is a combination of clay, silt, sand, pebbles, rock fragments, and angular chert, though very little is pebble-size material (Quinn 1972). One portion of the cave floor near an internal drain is clay, which settled in water, and rests above a sandy layer. Though the cave is 27 feet above Ben’s Branch, the alluvium in the cave matches the alluvium present around the stream. Alluvial benches in the valley indicate alluvium previously rested near the same level as the cave opening (Quinn

1972). 13

Davis (1973) identified four lithologic units within the cave. The oldest unit, Unit D, is yellow sand and gravel resting on the bedrock floor, lacking . Above this lay Unit C, a fossiliferous light reddish-brown clay which may indicate a change in moisture. The next layer, Unit B, is containing dark red-brown clay suggestive of previously extant pools of water. Unit B also carries evidence of limestone roof-falls. The youngest unit, Unit A, is comprised of angular chert pebbles, light brown friable material, fallen stalactites, and feces. The chert pebbles likely moved into the cave via slope wasting from the Boone Formation that closed off the entrance. Most of the faunal remains are derived from Unit B.

The horizons between units D and C and units B and A are not conformal, presenting evidence of mixing, while the horizon between C and B is conformal.

Excavations

Excavations were laid out with a trench, section format in which a trench would be marked off and broken into sub-sections which were used for provenience (Figure 7). Trenches were designed to be roughly three feet wide with sections roughly three feet square. Depth of excavations ranged from three to five feet.

Excavation depth was recorded by level, with level depth of six or twelve inches (Davis 1973).

Trenches 4, 8, 13, and 18 were excavated using six inch levels while all other trenches used twelve inch levels. Matrix was removed from the cave in wheelbarrows and drums to be screened (Quinn 1972).

Undried matrix was water-screened via submersion for four hours in screen-bottomed boxes. After this matrix was dried and passed through nested screens (1/2-in, 1/4-in, and 1/8-in). Dry screening of unwashed material occurred using the same meshes (Davis 1973). In clay and wet soil areas screening was not performed and recovery was dependent on excavators seeing specimens while excavating with hammer and trowel. The majority of faunal remains were recovered in the top 20 inches of sediment.

During excavations researchers noted strong evidence of animal bones, snails, , and other organic material washing into the cave from various surface entrances or flooding of Ben’s Branch (Davis

1973; Quinn 1972). Identified fauna from the cave include , , dire wolf, flat-headed peccary, giant armadillo, mammoth or mastodon, a large , , and many micromammals (Quinn 1972). 14

Radiocarbon dates from several items were obtained during initial excavation research. A piece of charcoal near a suspected latrine area was dated to 2,980 ± 180 14C years B.P. (Quinn 1972).

Additional charcoal was dated to 4,290 ± 110 and 2230 ± 120 14C years B.P. to go along with a date of

9,510 ± 140 on a snail and a composite date of 16,700 ± 250 (Quinn 1972:95).

In 1992 excavations were resumed by Kenneth Ball and L. Carson Davis. During this period the sinkhole within Peccary Cave was excavated in 15cm levels to a depth of 2.28 m (Ball and Davis

1993). Sediments were removed and dried on screens as most of the matrix in the sinkhole was clay. Once dried, sediments were water screened using a stacked set of screens with mesh ranging from 2.2 cm to 0.2 cm. All zoological and archaeological specimens were then removed from the screens (Ball and Davis

1993).

Previous Research on the Faunal Remains

Peccary Cave has been described as “the most intensely analyzed Pleistocene vertebrate site in

Arkansas” (Ball and Davis 1993:123). To be sure, the herpetofauna (Davis1969) and microfauna microfauna (Guilday and Parmalee 1972; Hallberg et al. 1974; Semken 1984; Semken et al. 2010) are very well studied. (Heretofore, the same could not be said about the medium- and large-size animals.)

Guilday and Parmalee (1972) used the presence of the vole Phenacomys intermedius to show the locale, during the late Pleistocene, was spruce forest. Hallberg et al. (1974), focusing on Microtus xanthognathus, inferred that the area around the cave was likely an open forest by 11,000 B.P. Utilizing co-occurrence of three species of Blarina (short-tailed shrew), Graham and Semken (1976) interpreted the local, Pleistocene environment as a more equable climate than present. And Stafford and Semken (1990) pushed this inference further in demonstrating the existence of a disharmonious, non-analog fauna.

Davis (1973) found evidence of a warming and drying trend throughout the biostratigraphy. In particular, based on presence of Crotaphytus collaris (common collared ), the climate dried out near the end of the Wisconsinan (Davis 1973:77). Ball and Davis (1993) added further evidence for this interpretation through their later sinkhole excavations in which they found toads significantly outnumbered frogs, representing a drier environment. 15

Expanding on previous work, and utilizing a greater diversity of micromammals, Semken (1984) investigated the shifting climate around Peccary Cave from the Late Wisconsinan period into the

Holocene. He found evidence of individualistic and progressive faunal change rather than catastrophic overhaul of the fauna. These progressive faunal changes reflected environmental changes with the local landscape shifting from cool steppe with patches of coniferous forest toward deciduous forest and temperate prairie.

More recently, Semken et al. (2010) dated 38 micromammal remains from the site and showed that the environment was non-analogous, bearing microfauna from boreal, tundra, neotropical, steppe, and deciduous forest environs over time. Despite this variance, for significant periods of time during the

Pleistocene the environment around Peccary Cave was relatively open (Semken et al. 2010). The non- analog fauna of Peccary Cave likely represents chronological and geographical variation in the environments of the southeastern United States during the LGM. Semken et al. (2010:Table 3) did find remains of beautiful armadillo (Dasypus bellus), a cold-intolerant species (Klippel and Parmalee 1984;

Schubert and Graham 2000), in the cave dated to 16,380 ± 70 B.P. potentially indicating the environment in the area at that time was relatively warm. Wide seasonal variation is believed to be detrimental to beautiful armadillo (Schubert and Graham 2000) and the absence of younger examples of the taxa in the cave may represent increasing seasonality.

Environmental Context

Environmental reconstructions of the Ozark Plateau are limited in number, but an overview of the existing literature is presented here to describe the environment as best as possible (Table 2). In general, the overall climate of North America during the Pleistocene was cool but more equable than

Holocene conditions (Axelrod 1967; Graham 1976a, 1985a; Graham and Grimm 1990; Graham and Mead

1987; Guilday et al. 1978; Hibbard 1960; Slaughter 1967). Many microfaunal assemblages from the LGM are non-analog, representing the more equable nature of the Pleistocene climate. Loss of such assemblages at the end of the Pleistocene is interpreted as a change to a more variable climate (Graham

1985b, 1986; Graham and Lundelius 1984; Graham and Mead 1987; Lundelius 1988, 1989). 16

In eastern North America the LGM environment was cold with mixed forests and steppe landscapes (Jackson et al. 2000). The mean January temperatures were 15 to 20oC colder than modern eastern North America with mean July temperatures 10oC colder than now (Jackson et al. 2000:500). By

21,000 years ago the region had significantly less precipitation than today which would have reduced the moisture available for plant and animal life (Bartlein et al. 2011).

In the LGM the CO2 levels across North America were low, an environmental factor that would have reduced the growth of C3 plants, providing an advantage to plants utilizing C4 photosynthetic pathways (Harrison and Prentice 2003; Prentice and Harrison 2009). However, the colder temperatures of the region negated this effect, enabling growth of C3 vegetation (Ward et al. 2008). As the LGM came to an end, significant climate change impacted eastern North America. By 16,000 years before present wide seasonal temperature variations occurred (Williams et al. 2001) as did rapid vegetation community change (Williams et al. 2004).

Prior to the LGM (before 22,200 ka) the Ozark Plateau climate was relatively stable. Delcourt and Delcourt (1996:229) showed from 34,000 to 23,000 years B.P. the Ozark Plateau biome would have been majority open pine parkland. Using speleothems from Crevice Cave, Missouri Dorale et al. (1998) demonstrated the environment was likely forest from 55,000 to 25,000 thousand years ago. King’s (1973) pollen analyses of several springs in the showed an open pine-parkland from 40,000 to 25,000 thousand years ago. Cupola Pond, MO was pine dominated parkland around 25,000 ka (Cushing

1975:63). Pollen from 34,000 to 24,000 years ago in the deposits from Trolinger Spring in the Missouri

Ozarks reveal a pine-dominated parkland in the region as well (King and Lindsay 1976:75). Haynes

(1985) reported the period from 26,000 to 22,000 ka around Boney Springs, Missouri was stable. At

Peccary Cave, only Neotoma floridana (eastern woodrat), C. dirus, and P. compressus are known to have been present prior to the LGM (Figure 8). The eastern woodrat is typically a deciduous forest inhabitant reliant on cover in its environment that prefers woody, wet conditions (Rainey 1956), consistent with the reconstructions. 17

Climate within the Ozark Plateau began shifting during the early LGM (22,200 to 20,000 ka).

From 25,000 to 20,000 ka the open pine parkland King (1973) described in the pre-LGM changed to a cool, wetter forest. The area around Powers Fort Swale in the western lowlands of Southeastern Missouri was cold, wet spruce swamp by 20,000 years cal B.P. (Royall et al. 1991:167). By 23,000 ka climate change resulted in the Trolinger Spring area shifting from open pine parkland to a spruce dominated landscape as well (King and Lindsay 1976:75). Delcourt and Delcourt (1996:230) showed from 23,000 to

16,500 ka vegetation shifted from open parkland to closed forest. Haynes (1985) also noted climate changed in the Boney Springs area after 22,000 ka. Such changes appear reflected in the micromammal component at Peccary Cave with the appearance of four additional species (Table 2), several of which are now found only in the arctic. Short-tailed shrew in particular, which prefer cool, wet conditions in deciduous forests (Pruitt 1953) show up in this period at Peccary Cave.

By the late LGM (16,800 to 20,000 ka) in the Ozarks the environment changed further. While

Cupola Pond was still dominated by cold tolerant spruce around 19,000 ka (Smith 1984:44) the dominant pattern is of warming and drying. King (1973) found a shift to a warmer clime, evidenced by the arrival of more thermophile plants, began slightly after 20,000 ka. Beautiful armadillo arrived in Peccary Cave around 19,700 cal B.P. (Semken et al. 2010), coinciding with the arrival of more thermophile plants.

Along with beautiful armadillo, four micromammal taxa appear for the first time in Peccary Cave, potentially indicative of the shifting climate (Semken et al. 2010). Cool, spruce preferring Blarina carolinensis, along with P. compressus and C. dirus no longer appear in the Peccary Cave faunal record by the late LGM (Table 2). Other wet and cold adapted micrommamals such as taiga vole (Swanson

1996), eastern mole (Arlton 1936), and northern bog lemming (Hafner et al. 1998) also fail to persist past this period (Figure 8). Unpublished data from speleothem research at Crevice Cave suggest the climate became dry, with speleothems experiencing significantly reduction in, or complete termination of, growth rates (Dorale 2017, personal communication). This restriction occurred between 25,000 and 18,000 years ago. 18

Within the Ozark Plateau, an increase in faunal diversity is seen within the early LGM (Figure

8). Prior to the LGM only the eastern woodrat, beautiful armadillo, flat-headed peccary, and dire wolves have been dated. As the climate shifted from open pine parkland to wetter spruce forest more microfauna appear in the dated record (Semken et al. 2010). Abruptly between 20,000 and 19,000 ka termination of faunal use of the Ozark Plateau occurs (Figure 8). Termination appears to coincide with the onset of warming and drying evidenced in the late LGM environmental recreations discussed above. Species reliant on cooler or moist conditions such as taiga vole, eastern mole, northern bog lemming, and short- tailed shrew drop out of the record. Around 7,000 years later eastern mole and taiga vole reappear briefly, while the other taxa have not been dated in the Ozark Plateau again. Even the eastern woodrat, a micromammal with relatively strong environmental plasticity (Rainey 1956), that still lives in the region today, disappears from the Ozark record for around 9,000 years. The pattern suggests the Ozark Plateau may have become inhospitable from roughly 19,000 ka until 13,000 ka. The single dated specimen within this timeframe, a plains pocket gopher, seems anomalous.

Overall, the general picture of the environment of the pre-LGM Pleistocene in the Ozarks is one of a relatively equable, cool climate dominated by open pine parkland. By the early LGM vegetation shifted to a wetter, spruce dominated environment. Around 20,000 years ago, the climate began warming and drying out, coinciding with the arrival of more thermophile plants and the disappearance of fauna for

6,000 years. However, pollen record recreations may not capture all landscape vegetation. Legumes

(Fabaceae) are common plant forms but drastically underrepresented in pollen rain, and thus pollen records, due to their mode of pollination (Grimm 1988). Studies of modern eastern woodrat have shown leguminous plants comprise nearly 20% of dietary material (Post et al. 1993; Wagle and Feldhammer

1997) and, as demonstrated later in this work, leguminous plants appear to be a primary food source for flat-headed peccary. It is likely leguminous vegetation was present in the Ozarks but is not recorded in current reconstructions.

19

Age of the Faunal Remains

The faunal assemblage from Peccary Cave is very well dated. Semken et al. (2010:Table 3) report 38 AMS ages on microfauna (and a beautiful armadillo) that range from 34,400 ± 790 B.P. to 3,400

± 70 B.P. As part of this study, 5 AMS ages on peccary bone and 3 AMS ages on dire wolf (Canis dirus) bone were obtained to situate these animals in a temporal context. These data are presented in Table 3.

The five peccary dates range between 20,610 ± 140 B.P. and 17,990 ± 100 B.P. As noted previously, these are the oldest, secure ages for flat-headed peccary in the contiguous United States. Dates for dire wolf range between 20,850 ± 80 B.P. and 17,880 ± 100 B.P. These dates not only bookend the dates for peccary but also demonstrate spatial and temporal overlap of the taxa (Figure 9). The dates are also considerably older than the 13,170 ± 600 B.P. date from Powder Mill Creek Cave, which is located in

Owls Bend, Missouri (Krueger and Weeks (1965:48).

20

CHAPTER 5. PALEOZOOLOGY

Most prior research on faunal material from Peccary Cave has focused on the large and diverse micro- and herpetofaunas, which include 35 and 17 taxa, respectively (Davis 1973; Semken 1984). Quinn

(1972) reports the assemblage also includes a number of other small, medium, large, and very large animals that are either extinct or no longer present in the region. In addition to flat-headed peccary and dire wolf, the extinct fauna includes muskox (Symbos sp.) elk- (Cervalces sp.), mammoth

(Mammthus sp.), mastodon (Mammut sp.), and stilt-legged deer (Sangamona sp.). Extant taxa that no longer occupy the region include the Andean tapir (Tapirus sp.), porcupine (Erethizon sp.), and

( sp.). The peccary assemblage is by far the largest, totaling several thousand identified specimens and representing at least 61 sub/adults and 15 fetal/neonates. The sample of dire wolf remains is not large, totaling about 20 identified specimens and representing 3 individuals. The other taxa are represented by isolated specimens or a handful of specimens.

The objective of this chapter is to elucidate the taphonomic history of peccary and dire wolf assemblages. Most attention is given to the peccary collection due to its size. To this end, skeletal element frequencies and bone modifications are used primarily to infer the mode of accumulation and how carnivores, rodents, or sediment chemistry may have shaped the general character of the assemblages

(Brain 1981; Lyman 1994).

General Assemblage Characteristics

The peccary assemblage includes 2,704 sub/adult specimens and 1,192 fetal/neonatal specimens.

Another 922 specimens are peccary-size but cannot be identified to a specific element. Although a systematic taxonomic assessment of the peccary material has not been conducted, maximum length measurements analyzed via a One-Way ANOVA test of the mandibular third molar demonstrate that the teeth of Peccary Cave peccaries are statistically indistinguishable from those at Sheriden Cave and Bat

Cave (Figure 10) F(3, 97) = 16.505, p = <.001. Use of Tukey post-hoc comparison reveals specimens from Laubach Cave are statistically distinct. Why these teeth are statistically different from those at

Laubach Cave is a question that deserves attention. 21

Generally speaking, the remains are relatively good physical condition although many specimens have suffered extreme gnawing from rodents, possibly by the Eastern woodrat, as woodrats are known for gnawing bone (Poole 1940). Approximately 20% of the peccary remains and 35% of the dire wolf remains have been gnawed (Table 4). These numbers likely underestimate the incidence of rodent gnawing because breccia variably covers many specimens (Table 5) (Figure 11). Interestingly, however, gnawing of fetal/neonatal remains is not common; although speculative, this may be because rodents ignored them due to their soft, porous nature.

Skeletal Element Representation

Figure 12 profiles the skeletal element frequencies for sub/adult and fetal/neonatal peccary remains (Table 6). For sub/adults material, small specimens such as carpals, tarsals, patellae, and phalanges are not common, while cranial and mandibular specimens are well represented. All other skeletal elements are fairly evenly represented. Calcanea and astragali, elements often indicative of complete limb deposition, are represented in similar numbers to most other appendicular elements. Lack of small skeletal elements, and overrepresentation of dental specimens, may represent recovery biases.

General presence of most skeletal elements among flat-headed peccary remains demonstrates natural death of peccary within the cave.

A slightly different pattern exists for the fetal/neonatal material (Figure 12). Apart from an abudance of ribs (probably because ‘concentrations’ of fetal/neonatal material were collected en masse), most elements are evenly represented. The absence of vertebrae, carpals, tarsals, and patella is simply due to the fact that these specimens can only identified generally to element given their size. Most vertebral bodies and carpals are small nubbins of bone. Similarly, the near-absence of metacarpals and metatarsals is due to the inability to differentiate between the two elements in fetal specimens. Even distribution in the element profile of fetal remains demonstrates near complete preservation and recovery of these fetal peccaries. Complete deposition of fetal remains implies fetuses were deposited in the cave through death of pregnant sows.

22

Minimum Number of Elements

Skeletal element abundance is a useful analytical tool for paleozoological research (Lyman

1994; 2008). Minimum number of element (MNE) data reported here uses the comprehensive MNE as established by Bunn (1986). Despite the evidence for lack of carpals and tarsals (Figure 12), reviewing the summary data demonstrates most appendicular elements are fairly evenly represented. Axial elements vary in their abundances, though some of this is due to the nature of the elements themselves. Crania of flat-headed peccary are thin and easily broken, impairing the ability to identify specific cranial elements.

Similarly, the atlas and axis vertebra are relatively fragile and can easily become unrecognizable. Other axial elements however are more evenly represented.

Examination of the fetal peccary remains reveals clear patterning as well. Among fetal remains long bones, inonimate and ribs, and calcaneii and astragalii provide the highest MNE counts. Carpals, tarsals, vertebrae, and phalanges may appear absent but this is not true. As mentioned, many carpals, tarsals and vertebrae were present but due to lack of morphological definition are unidentifiable beyond the level of vertebra or carpal/tarsal. This is different from mature peccaries where carpals and tarsals are absent. Long bone element MNEs show consistent recovery of fetal remains. Recovery of calcaneii and astragali from fetal peccary provide evidence of strong preservation of fetal material. This presence may indicate a lack of density mediated destruction. Further, relatively equal distribution of fetal elements implies fetal remains came to reside in the cave through death of the mother.

MNE counts on teeth reveal slightly different patterns. Tooth size generally increases from the incisors to premolars to molars to canines. Peccary possess deciduous incisors, canines, and premolars in both mandibular and maxillary dentition (Kirkpatrick and Sowls 1962). Reviewing the summary data for teeth (Table 7) reveals a bias toward recovery of mandibular canines. Despite a complete dentition possessing only four canines, as compared to eight incisors and twelve premolars and molars, mandibular canines NISP drastically outnumbers the next most frequent tooth, mandibular first molar. This overabundance is most likely due to preferential recovery of these distinctive specimens during excavation. Canine MNE values are much larger than the nearest MNE values from appendicular 23

elements. Molars are recovered in similar relative abundances to the appendicular skeletal elements while premolars and incisors are lacking.

Clear size-influenced recovery bias for dental elements exists (Table 7). A pattern of increasing presence with increasing overal size is apparent with mandibular and maxillary MNEt values of incisors, premolars, molars, and canines. Though 42 more molar than canine elements are present this is offset due to typical dentition carrying more molars than canines. Analysis of dental presence by NISP shows canines as the most recovered element, followed by molars, premolars, and incisors. The overabundance of canines, and drastic variation in recovery of teeth by size, is not due to natural dentition but indicative of either differential preservation or recovery. It is plausible smaller teeth were washed away during periodic flooding but they may also have fallen through screen mesh or gone unnoticed during recovery from matrix.

Since the underrepresnted and overrepresented peccary elements may be explained through excavator bias, the relatively even representation of axial, appendicular, and molar elements provides evidence for natural death. Predation, by humans or carnivores, should result in differential preservation of elements (Binford 1981; Todd and Rapson 1988).

Minimum Number of Individuals

Utilizing the MNE values by side allows for determination of minimum number of individual

(MNI) estimates following the method set forth by White (1953). Results reveal at least 61 sub/adult mature peccaries are represented in the assemblage along with at least 15 fetal peccaries and 3 dire wolves.

The MNI for sub/adults is based on left mandibular canines (Table 7). This leads to an element

MNI that is disproportionately large compared to other elements because of the excavation bias in recovering peccary canines. Appendicular elemental MNIs cluster in the upper 30s. Identification of specimens impacted by breccia formation, rodent modification, or other post-depositional damage may help explain the discrepancy in MNI estimates between dental and other skeletal elements, though recovery preference toward canines appears the primary cause. 24

Deciduous canines (mandibular MNE = 44) in the assemblage confirm the presence in the cave, either through behavioral use or natural death, of juvenile peccary. At least 12 individuals are represented by deciduous canines, though the number is low as 21 deciduous canines unable to be sided are present

(Table 7). Since peccaries possess deciduous and permanent canines, losing their deciduous teeth as the permanent canines erupt (Kirkpatrick and Sowls 1962), it is plausible the deciduous canines were deposited by peccaries that later died in the cave, leaving behind mature canines as well. Thus only mature canines are included in MNI count. Eight mandibular canines were unable to be identified as either deciduous or permanent and have not been included in the MNI count.

Fetal MNI is based on femora, with at least 15 individuals represented (Table 5). This is closely followed by other appendicular elements providing MNI estimates between 9 and 12. Comparisons between MNI estimates on teeth are not possible for fetal peccary. Premolars and first and second incisors erupt after birth and therefore a fetal MNI based on those dental remains is not possible to ascertain. At birth modern peccary possess all four canines and IC3 (Sowls 1997), a tooth P. compressus lacks. Even though the deciduous canines are present at birth it is not possible to use them to estimate fetal numbers either as they are replaced between 29 and 41 weeks in modern peccaries (Kirkpatrick and Sowls 1962).

Assemblage Age Composition

Using age-mortality profiles of sites is a standard practice for archaeological investigations of faunal remains (Klein 1982). While many profiles are based on dental wear it is also possible to obtain basic age structure from epiphyseal fusion of elements. At Peccary Cave, general age structure is determined by epiphyseal fusion of the proximal epiphysis of calcanei. Calcanei were chosen due to their relative abundance (MNE = 73) and their overall preservation of epiphyseal ends compared to other elements with epiphyseal fusion.

The assemblage is comprised of at least 14 sub-adult and 12 fully mature peccaries (Table 8).

While not representative of the entire assemblage, from the available specimens it is apparent both skeletally mature and immature peccary died in the cave. Though the age structure (MNI = 15 fetal, 14 sub-adult, 12 mature) is reminiscent of a catastrophic mortality profile (Klein 1982) the assemblage was 25

not created via a catastrophic event based on radiocarbon dates. Therefore, at various times, pregnant females, sub-adults, and mature adults died in the cave.

Carnivore Damage

The assemblage from Peccary Cave, though relatively well preserved, is not in pristine condition. As noted above, many specimens were modified by rodents and/or coated in breccia. Another modification that needs to be examined is damage to the remains by carnivores. Several recent studies have shown spatial overlap of dire wolves and peccaries in caves tied to carnivore modification of peccary remains (Nye 2007; Woodruff 2016). It should be noted that Hawksley et al. (1973) concluded that there was little carnivore damage on peccary remains from Bat Cave, although more recently,

Woodruff (2016) reports extensive carnivore damage. These findings of apparent carnivore usage are not replicated at Peccary Cave. While carnivore modification is present, only 10 specimens possess definitive evidence of carnivore modification. Seven of these specimens show evidence of carnivore pitting while three show evidence of furrowing. Additionally, carnivore modification typically results in the loss of epiphyses of long bones (Binford 1981; Todd and Rapson 1988). The relatively common presence of proximal and distal epiphyses on long bone elements in Peccary Cave indicates a lack of carnivore modification. The majority of alteration comes from rodent modification and breccia formation.

Density-Mediated Attrition

Due to the presence of rodent modification and breccia formation, the overall impression of the

Peccary Cave flat-headed peccary assemblage is that it has undergone remodeling due to density- mediated attrition. Following the procedure outlined by Morlan (1994), Spearman’s rank-order correlations (rs) between scan site MNE and scan site bone volume density (Table 9) were employed to test the overall impression. The correlations are based on scan site MNEs summarized in Appendix B.

Since peccary bone volume density values are unknown, bone volume density values of (Ioannidou

2003) were utilized.

Typically volume density values for all elements are analyzed to determine assemblage level patterns of destruction. With collection and identification biases impacting the Peccary Cave assemblage, 26

only proximal long bones, calcanei, and astragali were incorporated in the analysis due to relative abundances indicating they were subjected to less recovery bias. Scan site locations are based on Lyman

(1984) and Ioannidou (2003).

For the flat-headed peccary assemblage, significant correlations exist between volume density and preservation for humeri (p = <.001) and the assemblage as a whole (p =.006) (Table 9). Less dense bone suffered greater loss than denser bone overall. This result supports the impression that density- mediated attrition impacted the overall assemblage. The reason for the pattern may be less clear.

Lack of significant correlations for the remaining elements, radius-ulna, femur, tibia, calcanea, and astragali reveals potentially strong preservation of specimens. Carnivore modification and soil chemistry induced destruction disproportionately destroy proximal and distal epiphyses of long bones over long bone shafts (Binford 1981; Todd and Rapson 1988). Many proximal and distal ends were preserved suggesting the greatest impacts on assemblage frequency patterning may not have been natural phenomenon. The presence of numerous fetal remains supports this suggestion.

Were all elements to be included it is anticipated volume density would be highly correlated with survivorship of specimens. Excavation actions likely resulted in some of the patterning. It is possible more broken epiphyses went unrecovered during excavation due to their smaller size whereas long bone shaft pieces tended to be larger. Rodent modification may also have differentially impacted identification of long bone epiphyses, contributing to density-mediated attrition. Despite significant assemblage level patterns among the proximal long bones, calcanea, and astragali the presence of fetal remains and lack of significant correlation among proximal long bones apart from humeri suggest overall preservation was strong and support the conclusion of natural deaths by peccary within Peccary Cave.

Dire Wolf

Remains of dire wolf are not common at Peccary Cave. The sample includes 14 specimens, mostly distal limb elements: 12 metapodials plus one calcaneus and one humerus. The drastic difference in skeletal element representation, overrepresentation of metapodial elements, provides evidence for accumulation of dire wolf remains via a different taphonomic pathway than peccaries. Natural death 27

would leave numerous appendicular and axial elements in relatively even distribution, as seen with the peccaries. The skeletal element profile for dire wolf remains presents no such pattern, nearly completely consisting of metapodials. Initial excavators believed fluvial deposition of material occurred in Peccary

Cave (Quinn 1972; Davis 1973). It is plausible, perhaps likely, the dire wolf remains were deposited through either stream flooding or other fluvial or alluvial movement.

Despite possessing a small assemblage, at least three dire wolves are represented by right fifth metacarpals. The large difference between minimum numbers of peccary and dire wolf point to different use of the cave by these animals. The relative absence of dire wolf remains coupled with the virtual absence of carnviore damage on the peccary remains strongly suggests dire wolves rarely if ever used the cave.

Discussion

The paleozoology of Peccary Cave presents insights into the creation of the faunal assemblage.

Peccaries were dying naturally within the cave system. Dire wolves were not and likely entered through fluvial or alluvial movement bringing skeletal elements from outside the cave.

Consistent evidence exists from skeletal element frequencies to demonstrate peccaries were dying in the cave. Most elements appear in relatively even distribution, while those overrepresented or underrepresented are explainable through excavation induced biases. Modern peccaries will bed down in caves to escape temperature extremes (Neal 1959; Sowls 1997) and it is plausible such use was occurring at Peccary Cave. The recovery of peccary remains from other cave sites (Appendix A) supports the interpretation of peccaries using caves behaviorally.

Though skeletal element frequencies show peccaries were utilizing and dying in the cave, evidence does not exist for mass death. Radiocarbon dates (Table 3) indicate a time averaged accumulation generated the assemblage. At minimum a 3,000 year time span is represented. With 61 peccaries, this represents roughly one peccary dying in the cave every 50 years.

Fetal peccary remains further these interpretations. Their presence, with relatively little loss of material, means pregnant females were dying in the cave. Lack of destruction of fetal remains reveals a 28

lack of carnivore presence, as consumption of pregnant females should include consumption of fetal remains resulting in loss of the material. Since this did not happen it is likely the pregnant females died without predator involvement.

Modern peccaries average litter size is two (Table 10). Thus it is reasonable to assume, with an

MNI of 13 fetal peccaries, six or seven pregnant females died in the cave. As the assemblage is known to be time averaged this represents one pregnant peccary dying every 430 years over the minimum 3,000 year use of Peccary Cave. Most fetal remains came from only two excavation sections (Figure 13) which may be the result of excavator action rather than true spatial spread. The spatial overlap of fetal remains need not imply contemporaneous death of pregnant females as it is unlikely six or seven pregnant females, dying around roughly the same time, would die directly on top of each other, within a 3 by 3 foot square. The spatial overlap of remains may result from deliberate recovery of fetal material from the specific units or from another process, such as fluvial action pushing small, fetal remains into compact zones.

Spatial distribution of remains also supports the interpretation of peccaries utilizing Peccary

Cave behaviorally (Figure 13). Non-fetal peccary remains were recovered from most excavated locations in the cave, though concentrated in the more northerly sections. The relative uniformity in spread of remains would not be expected if significant fluvial action had impacted the assemblage nor would this be expected if the peccaries were dragged into the cave by carnivores. The spatial spread aligns with other evidence of use and natural death. Interestingly, there is a visual pattern of increased chipped stone density in areas of high peccary remain density. Since temporal periods for peccaries and human use are separated by nearly 20,000 years this may represent excavation bias wherein a focus on faunal remains resulted in recovery of chipped stone from those excavation sections while chipped stone only sections were less thoroughly excavated or not excavated at all.

A lack of carnivore modification combined with the paucity of dire wolf material indicates

Peccary Cave was not a carnivore den. It is unlikely dire wolves used the cave despite their presence on the landscape at the same time as the peccaries. If three wolves had died in the cave there should have 29

been many more dire wolf specimens, particularly as metapodials are not the densest or largest, and therefore not the most easily preserved specimens. If dire wolves were not using the cave it lends further credence to the interpretation of behavioral use by Pleistocene peccaries.

Excavation bias is clearly present in the faunal assemblage. The evidence is strong for natural death of peccaries in the cave, and the lack of significance in long bone density-mediated attrition may rule out significant chemical damage. Element frequencies show relatively even distribution in recovery of most elements apart from small, but dense, elements like carpals and tarsals (Figure 12). Excavation of

Peccary Cave was coarse grained, particularly in areas comprised largely of clay, which likely resulted in the loss of smaller skeletal elements. The overabundance of canines and molars versus scarcity of incisors and premolars (Table 7) also shows excavation bias toward recovery of larger specimens. The relative overrepresentation of canines compared to other skeletal elements likely reflects excavator preference for recovery of the distinctive canines.

As a natural death location for flat-headed peccary, Peccary Cave is a unique setting from which to examine further behavior and possible extinction causes. While other cave assemblages may have been carnivore dens, or carnivore mediated (Nye 2007; Woodruff 2016), Peccary Cave provides behavioral insight that has previously been unknown for these extinct creatures. Lack of density mediated destruction on the assemblage makes the peccaries themselves useful study specimens for understanding behavioral ecology of peccaries on a larger scale. Furthermore, though some material of the cave suffered rodent modification and breccia formation, the material overall is well preserved. Fetal remains show little damage and evidence strong preservation, as does the relative completeness of the assemblage.

Summary

1. Peccary Cave represents the natural death location for at least 61 adult flat-headed peccary and 15

fetal peccary. Natural death of pregnant females, along with other peccaries, may imply hardship for

during the time period of use of the cave. 30

2. Peccary Cave assemblage is time averaged accumulation. Deaths occurred over at least 3,000 years

which equates to one peccary every 50 years, one pregnant peccary every 430 years, and one dire

wolf every 1,000 years.

3. Post-depositional taphonomic process affected the assemblage. Rodent modification and breccia

formation limited identification of certain features and elements possibly lowering the amount of

identified specimens and carnivore modification occurrences. Regardless, minimal carnivore

modification occurred and density mediated attrition does not appear to have impacted the

assemblage significantly.

4. Overall preservation of the assemblage is strong. Fetal remains and skeletal element frequencies show

the presence of most elements in even representation demonstrating the skeletal assemblage is well

preserved.

5. Future studies investigating density mediated attrition should consider using bone-density values

from and pigs as analogs for peccary. values should be avoided.

6. Collection bias is evidenced in element frequencies in appendicular and dental elements. Differential

recovery of larger teeth, particularly canines, is apparent. Lack of small elements like carpals and

tarsals may be due to coarse recovery methodology.

7. Peccary Cave is not a dire wolf den. Element frequencies and carnivore modification demonstrate dire

wolves did not use the cave and their remains were likely introduced through post-depositional

processes. 31

CHAPTER 6. STABLE ISOTOPES

The use of stable isotope biogeochemistry for determining ecological relationships and diet for extinct and extant taxa is well established, and aids in understanding late Pleistocene extinctions when climate change caused shifts in vegetation types, abundances, productivity, and moisture (Graham 1985a,

1986; Kutzbach et al. 1998; Prentice et al. 1991; Webb et al. 2004; Williams et al. 2001).

Stable isotope values from mammalian teeth samples accurately record δ13C and δ15N per mil values of isotopic composition of diet (DeNiro and Epstein 1981; Cerling et al. 1997; Lee-Thorp and van der Merwe 1987). For herbivores, δ13C values from teeth reveal dietary preference for certain plant types,

13 based on photosynthetic pathways known as C3 and C4 plants, whose δ C values vary greatly (Bender

13 1971). In C3 plants δ C values range from -35‰ to -22‰ whereas for C4 plants the range is -20‰ to -

8‰. Despite the large range, most C3 plants cluster between -28‰ to -23‰ with most C4 plants between -

13‰ to -10‰ (Bender 1971; Cerling et al. 1997). A fractionation occurs between dental enamel and diet

13 12 13 δ C/ δ C ratios, therefore, in teeth, Cerling et al. (1997) showed enamel δ C for animals with C3 dominant diets range between -13‰ to -8.6‰ while C4 dominated diets range from -2‰ to 4‰.

Previously Cerling and MacFadden (1996:Table 1) found enamel ranges of -19‰ to -9‰ δ13C indicated

13 C3 dietary patterns. As with enamel, dentine and dentine-enamel mixes from teeth record dietary δ C

(Passey et al. 2005; Zazzo et al. 2006).

δ18O values may also be obtained from dental samples. The relative ratio of δ18O to δ16O of precipitation varies in predictable patterns over landscapes based on temperature, source, and distance from source, known as Rayleigh distribution (Gat 1996). Due to the impact temperature has on fractionation of oxygen, tracing the δ18O value change over time in a set location can record change in seasons. This will be complicated by variation in source and amount of precipitation. Despite complicating factors, seasonal variation has been recorded in mammalian teeth enamel and dentine

(Fricke and O’Neil 1996; Fricke et al. 1998; Gadbury et al. 2000; Higgins and MacFadden 2003; Hoppe

2006; Kirsanow et al. 2008; Koch et al. 1989; Kohn et al. 1998). Pairing δ18O values from serially 32

sampled teeth with δ13C values from the same sample locations may reveal change in diet or behavior by season (Cerling et al. 2009; Chisholm et al. 1986; Laffoon et al. 2014; Widga et al. 2010).

Another stable isotope, δ15N, is commonly used in archaeological and zoological studies to identify trophic levels and food sources. Heavy elemental isotopes are incorporated preferentially from food sources, resulting in an increase of ~3 – 4‰ δ15N and ~1 - 2‰ δ13C with each trophic level (DeNiro and Epstein 1981; Minagawa and Wada 1984). Comparisons of δ15N with δ13C values from different species thus possess potential to reveal predation pathways. Variations in nitrogen values reflect diet sources within species as well (DeNiro 1987; DeNiro and Epstein 1981). Broadly, differences in plant nitrogen intake enables use of δ15N values to separate plants into nitrate fixing, legumes, and non- legumes, or non-nitrate fixing (Burns and Hardy 1975), with legumes possessing lower δ15N values than other plants (Delwiche and Steyn 1970; Delwiche et al. 1979; DeNiro 1987; DeNiro and Hastorf 1985).

Research shows ranges from ~0 - 4‰ for legumes with non-legumes varying between ~5 – 12‰ δ15N

(DeNiro 1987:Figure 3; DeNiro and Hastorf 1985). Due to these variations, values of δ15N combined with

13 δ C reveal dietary reliance on C3 legumes, C3 non-legumes, or C4 plants.

To date, few reports of isotopic data from flat-headed peccary for diet exist. Therefore this research utilized isotopic investigation to further study of peccary for preferred diet. Composite lifetime

δ13C and δ15N values were obtained during radiocarbon dating (Table 3). Serial samples of two peccary canines were analyzed to obtain paired δ13C and δ18O values to investigate diet by season. Serial intra- tooth samples are essential tools for understanding herbivore diets (Balasse 2002; Britton et al. 2009;

Feranec 2007; Feranec et al. 2009; Gadbury et al. 2000; Larson et al. 2001). Variability by season may indicate a capacity for shifting food sources, potentially limiting damage brought by climate change on a population. Conversely, lack of seasonal diet change may indicate strong reliance upon a particular diet source, increasing the possibility climate change may negatively impact species through vegetation change.

33

Methods

Bone Samples

Five peccary and three dire wolf bones were sampled for radiocarbon dating and sent to the

UC-Irvine Keck Carbon Cycle AMS Facility. During radiocarbon dating stable isotope values for δ13C and δ15N were obtained. Measurements were performed on extracted collagen. δ13C was measured on prepared graphite using a Fisons NA 1500NC elemental analyzer/Finnigan Delta Plus isotope ratio mass spectrometer. δ13C and δ15N values were measured to a precision of <0.1‰ and <0.2‰, respectively, on aliquots of ultrafiltered collagen. All results were corrected for isotopic fractionation according to the conventions of Stuiver and Polach (1977).

Serial Samples

Two unbroken, mature peccary tusks were selected for stable isotope analysis. Using a Dremel drill bit, 18 powdered samples were generated at roughly 2mm intervals on specimen 67-300-14. 17 samples at 2mm intervals were taken from specimen 65-158-18. Based on modern peccary tusk growth rates of approximately 1.5 mm per month (Sowls 1984:20) 2 mm intervals achieved as near a month by month sample as was possible with the given tools. Due to the nature of the drill bit used, and the thinness of the enamel on peccary canines, samples were a mix of enamel and dentine. Though enamel is preferred over dentine for isotopic studies (Lee-Thorp and van der Merwe 1991; Shahack-Gross et al. 1999) the difference in isotopic value between enamel and dentine is not well understood yet and both structures have demonstrated accurate records of isotopic information (Passey et al. 2005; Kirsanow et al. 2008;

Koch et al. 1989; Uno 2012; Zazzo et al. 2006). After obtaining a mixed enamel-dentine powder sample from a location a dentine only powder sample was taken.

Due to a lack of material the first two samples from specimen 67-300-14 failed to provide measurements resulting in 16 samples of enamel-dentine and dentine each for analysis. All 17 sample locations for both enamel-dentine and dentine were analyzed on specimen 65-158-18.

All samples were separated into individual vials upon drilling. Samples were then weighed using a microbalance to obtain 4 to 6 mg from each sample location for mixed enamel-dentine and 34

dentine only. No procedures for separating collagen from apatite within the samples were performed, and all samples therefore are bulk structural carbonate which formed within the enamel and dentine of the peccary tusks. After samples were weighed, pretreatment based on the methods specified in Rountrey

(2009) was performed. Each sample was soaked in 200mL of sodium hypochlorite 2.5% NaOCl for 24 hours. Samples then underwent five rinses with pure water. After pure water rinsing each sample was soaked in acetic acid 0.100 Normal for 4 hours. Once soaking in acid was complete another series of five rinses with pure water was utilized. Samples were then taken for isotopic analysis.

Isotopic measurements were performed via a Finnigan MAT Delta Plus XL mass spectrometer in continuous flow mode connected to a Gas Bench with a CombiPAL autosampler at Iowa State

University. The isotope ratio measurements are calibrated based on reference standards NBS-18 and

NBS-19 with at least one reference standard used every five samples. The combined uncertainty

(analytical uncertainty and average correction factor) for all samples from specimen 65-158-18 and the mixed enamel-dentine series of specimen 67-300-14 is ± 0.09‰ (VPDB) for δ13C and is ± 0.16‰

(VPDB) for δ18O. Error was determined via regression-based error analysis. Due to device/software problems the samples for specimens 67-300-14 and 65-158-18 were run 48-72 hours after acidification.

The dentine samples from specimen 67-300-14 were run prior to machine errors and possess a combined uncertainty of ± 0.07‰ (VPDB) for δ13C and ± 0.12‰ (VPDB) for δ18O.

Results

13 δ Ccoll. values ranged from -20.5 ± 0.1 to -21.3 ± 0.1‰ on dated peccary bone samples (Table

15 3). On the same specimens, δ Ncoll. values ranged from 3.7 ± 0.2 to 4.8 ± 0.2‰. These results demonstrate likely dietary habits focused on plants and place peccaries in the second trophic level. Dire

13 15 wolf remains ranged in δ Ccoll. from -18.4 ± 0.1 to -18.9 ± 0.1‰ and δ Ncoll. from 6.1 ± 0.2 to 8.0 ±

0.2‰ (Table 3). The carbon and nitrogen values fall in the range of the third trophic level, one above the peccaries (Figure 14).

Due to the known issue of a mixed enamel and dentine signal, sample series for both enamel- dentine and dentine only were run for specimens 67-300-14 and 65-158-18. The raw results of the 35

enamel-dentine and dentine only samples are shown in Appendix C. While the results are not identical,

13 the average differences in δ Ctusk of enamel-dentine minus dentine only are -1.69‰ for specimen 67-300-

18 14 and 0.21‰ for specimen 65-158-18. Average differences in δ Otusk for enamel-dentine minus dentine only are -1.26‰ and 0.33‰ for specimens 67-300-14 and 65-158-18 respectively. The relative similarity in results implies samples mixing enamel with dentine, or simply using dentine, may provide accurate results. Despite the relative similarity, only the mixed enamel-dentine results are interpreted below due to prior work favoring use of enamel (Lee-Thorp and van der Merwe 1991; Shahack-Gross et al. 1999). The similarity in values between the series may indicate the dentine signature from enamel-dentine mixed samples overrode the enamel signature.

13 Average δ Ctusk values were -13.34‰ and -13.50‰ for samples 67-300-14 and 65-158-18 respectively, a C3 signature based on previous studies showing fractionation occurs between diet and enamel, or enamel-dentine mixes, of herbivorous animals with heavier isotopes preferentially incorporated in teeth (Cerling and Harris 1999; Passey et al. 2005; Zazzo et al. 2006). On teeth particularly, Passey et al. (2005) have shown dental values of δ13C are 13.3‰ higher than diet. Pig and peccary teeth are quite similar (Hillson 1986) and transforming dental values from Peccary Cave by subtracting 13.3‰ would show a dietary value of roughly -26‰. Sullivan and Krueger (1984) showed collagen δ13C averages 5‰ higher than diet. Transforming collagen values from Peccary Cave by subtracting 5‰ would also show a dietary δ13C value of roughly -26‰.

13 A roughly 3‰ variance over the lifespan in δ Ctusk values is present in both specimens (Figure

15). Specimen 67-300-14 carries a maximum variation of 3.21‰ based on the mixed enamel-dentine sample series between values of -12.00‰ at 86 mm from the enamel junction and -15.21‰ at 56 mm

13 from the junction. Maximum lifetime variance in δ Ctusk values for specimen 65-158-18 is 3.03‰ with a

13 value of -15.26‰ at 71 mm and -12.23‰ at 57 mm. Lifetime variance in δ Cdentine was slightly smaller, with maximum variation in specimen 67-300-14 of 2.43‰ and specimen 65-158-18 of 2.73‰. Despite

13 variance over time, the highest δ Ctusk values were -12.00‰ and -12.23‰ for specimens 67-300-14 and

13 13 65-158-18 respectively. A 1.11‰ higher maximum δ C value, -10.89‰, was recorded on δ Cdentine than 36

13 13 13 δ Ctusk for specimen 67-300-14. The highest δ C value for specimen 65-158-18 was from δ Ctusk at -

12.23‰.

13 18 13 Figure 15 shows the change in δ Ctusk and δ Otusk via down-tooth sampling. Over time, δ Ctusk

18 18 and δ Otusk generally move synchronously. δ Otusk lows at 56 mm, 39 mm, and 23 mm on specimen 67-

300-14 likely indicate seasons as roughly a year, based on tooth growth projections, separates each low point. It seems probable that peccary at Peccary Cave would take drinking water from Ben’s Branch, potentially a relatively unbuffered small stream. Observed annual δ18O lows may indicate cold seasons

18 18 due to greater δ O fractionation with colder temperatures (Gat 1996). The δ Otusk value movements between sample points at 56 mm, 39 mm, and 23 mm on specimen 67-300-14 mimic expected meteoric water line patterns for seasonal change.

18 The δ Otusk signature from specimen 65-158-18 is less clear, with fewer blatant peaks and dips.

Fluctuations between measurements at 62 mm and 44 mm may capture a single season. δ18O values may have been impacted due to precipitation source or peccary obtainment of water as modern peccaries intake some water from plants rather than obligate drinking (Sowls 1997). Additionally, modern δ18O value movements from Arkansas rivers near the Ozark Plateau do not precisely follow seasons (Coplen and Kendall 2000:51, 55). Modern patterns reflect amount and source effects which likely influenced

δ18O values during the Pleistocene as well.

Discussion

The isotopic data provided here aligns with prior isotopic work and may grant new insights into flat-headed peccary relationship with dire wolves. Though limited in number of samples, the results provide strong implications for predation on peccary by dire wolves, a peccary reliance on C3 leguminous vegetation, and a lack of seasonal variance in peccary diet.

Three prior studies reported flat-headed peccary isotope average values from Florida and

Tennessee. MacFadden and Cerling (1996:111) found an average enamel δ13C value of -8.8‰ from peccaries in Florida dated to around 100,000 years ago, which they interpreted as indicative of mixed

13 C3/C4 or CAM feeding. Yann and DeSantis (2014:Table 6) reported an average of -10.7‰ δ C on enamel 37

from four peccary samples of early Rancholabrean age at Haile 8A, Florida, interpreted as a signature of a

13 C3 diet. Finally, Giesler (2002) provided an average value of -12.7‰ δ C from seven peccary dental samples from Guy Wilson Cave, Tennessee. This average also was seen as a C3 signature. The average

13 dental value for δ Ctusk from both Peccary Cave specimens combined is -13.42‰, also a C3 signature. All

13 four studies show δ C values indicate C3 or mixed C3/C4 diets. In particular, the two more northerly sites strongly suggest reliance on C3 plant life for diet. One further study, on Neogene peccary, showed a lack of seasonal variation in diet (DeSantis and Wallace 2008). This may indicate reliance on single vegetation types for ancient Tayassuidae. Their data correlates well with the samples analyzed here.

Bone Samples

Bone collagen isotope analysis of dated flat-headed peccary strongly suggests peccary relied on

13 C3 legumes year round (Figure 14). Bone values present lifetime averages, with the δ Ccoll. signature of the peccaries at roughly -21‰ falling within the range of C3 plants (Bender 1971; Cerling et al. 1997).

15 13 15 Pairing δ Ncoll. values with those from δ Ccoll. strongly suggests peccary ate C3 vegetation with δ N values between 0 – 3‰.

Prior research has shown animals are expected to increase in δ15N values by 3 to 4‰ with a single trophic level increase (DeNiro and Epstein 1981; Minagawa and Wada 1984: Schoeninger and

15 DeNiro 1984). Flat-headed peccary values from Peccary Cave ranging from 3.9 to 4.8‰ δ Ncoll. (Table 3) demonstrate a dietary source with δ15N values between 0 and 2‰, values matching the proposed δ15N for prehistoric legumes (DeNiro and Hastorf 1985). Current pollen reconstruction for the Ozark Plateau reports presence of leguminous plants at both Boney and Koch Springs in the LGM and late Pleistocene, though neither are key contributors to the pollen record (King 1973). Such lack of legume pollen does not mean leguminous plants were not present, nor does it mean they were rare, as legume pollination relies on insects leading to drastic underrepresentation of legume pollen in pollen rain in lake and wetland

15 13 sediments (Grimm 1988). Thus combining δ Ncoll. with δ Ccoll. from Peccary Cave specimens suggests flat-headed peccary were heavily reliant on legumes for diet over their lifetime (Figure 14). 38

15 δ Ncoll. values also demonstrate a likely predator-prey relationship at Peccary Cave. Dire wolf

15 δ Ncoll. values range from 6.1 to 8.0‰ (Table 3). These are between 2 to 4‰ higher than peccary values

15 for δ Ncoll. (Figure 14), representing the traditional expectations for a single trophic level increase

13 (DeNiro and Epstein 1981; Minagawa and Wada 1984). However, the dire wolf δ Ccoll. values are between 2 and 3‰ higher than peccary, slightly above the expected increases. Fox-Dobbs et al. (2007) showed enrichment for δ13C and δ15N of wolves from prey to be 1.3 ± 0.6‰ and 4.6 ± 0.7‰ respectively with a fully constrained diet. As the modern wolf δ15N enrichment is slightly beyond the average

13 differences reported from Peccary Cave specimens, and the δ Ccoll. values of dire wolves are between 2 and 3‰ higher, rather than 1 to 2‰, it appears the dire wolves were preying on multiple species, not

15 solely peccary. However, as the δ Ncoll. values of peccary and dire wolf fit the expected values for a one

13 trophic level increase, and the δ Ccoll. values are only marginally higher than expected, the values reported here strongly suggest that near Peccary Cave flat-headed peccary formed a part of the dire wolf diet.

This isotopic predator-prey relationship is supported by the spatial and temporal overlap of both taxa on the LGM landscape at Peccary Cave (Figure 9). Review of other Ozark Plateau Pleistocene faunal assemblages (Hawksley 1986; Hawksley et al. 1973; Hood and Hawksley 1975; Parmalee and Oesch

1972; Schubert 2003; Simpson 1946, 1949) reveals few other mid-sized herbivores that might have served as key prey for dire wolves. Lacking many other likely prey suspects provides intriguing evidence that flat-headed peccary may have been a key prey species for dire wolves on the Ozark Plateau. If peccary were a key prey species for dire wolves it is possible peccary extirpation could have impacted dire wolf populations, contributing to their disappearance as well.

Serial Samples

13 13 All washed δ Ctusk values on enamel-dentine samples were lower than -12.00‰ δ C. Serial sampling here recorded years 1 – 3 in life of the peccaries based on replacement age of deciduous canines with permanent teeth in modern peccary (Kirkpatrick and Sowls 1962). Put together, this data reveals that flat-headed peccaries relied on C3 vegetation for their diet, at least from 1 – 3 years of age. This could 39

have been advantageous at the time due to low temperatures which enabled C3 plant life to effectively

15 compete with C4 plants (Harrison and Prentice 2003; Prentice and Harrison 2009). Moreover, δ Ncoll.

13 values combined with the dental δ Ctusk data shows peccaries likely relied on leguminous plants over C3 browse.

δ18O values from the serially sampled tusks provide evidence that seasonality is likely recorded in the tusks of flat-headed peccary (Figure 15). While seasonal shifts in δ18O value were recorded

13 13 coinciding with shifts in δ C, the shifts in δ C fail to rise into ranges of C4 plants, or even mixed signature values suggested by prior research (Cerling et al. 1997; Cerling and MacFadden 1996). It may be that during warmer seasons different types of C3 vegetation were preferred than during cold seasons or

13 18 that seasonal temperature impacted δ C values of the plants similarly to δ O. Incorporation of small amounts of C4 plant material during warmer seasons may also be indicated by the synchronous pattern

13 (Sharp and Cerling 1998). Despite the potential for mixing, δ Ctusk values throughout the sampled

13 lifespan of both individuals remain firmly within C3 plant δ C value ranges. This suggests peccaries restricted their diet to C3 plant material, likely legumes, regardless of season.

Although these peccaries may have possessed omnivorous capability, isotopically they are herbivorous. Yann and DeSantis (2014) noted flat-headed peccary may have been able to shift dietary preference for C3 or C4 vegetation over large timescales but determining this will require more samples across time and space paired with isotopic data than are currently available. Though Yann and DeSantis

(2014) reported peccaries could fluctuate in diet, they concluded peccaries likely relied more on C3 than

C4 plants near the end of the Pleistocene. Such reliance is supported in the serial and bone samples from

13 Peccary Cave. C3 dietary preference is congruous with other known flat-headed peccary δ C values.

13 15 More samples are needed to confirm this pattern and pairing δ C with δ N for additional specimens may help reinforce or undermine the apparent reliance on leguminous vegetation.

Summary

13 1. Seasonality is likely captured in peccary tusks. While seasonality is present, and some δ Ctusk

variation occurs, isotopic evidence suggests peccaries relied on C3 plant life for diet in all seasons. 40

15 13 2. δ Ncoll. combined with δ Ccoll. support the interpretation of reliance on C3 plant life for diet,

particularly suggestive of a strict focus on legumes.

13 15 3. Isotopic values for δ Ccoll. and δ Ncoll. from peccary and dire wolves at Peccary Cave strongly imply

dire wolves ate peccaries. Radiocarbon dates confirm the possibility of a predator prey relationship

due to overlap temporally and spatially on the landscape.

4. Flat-headed peccary isotopes from Peccary Cave align well with other known peccary δ13C values,

evidencing a species-wide adaptation toward C3 vegetation. 41

CHAPTER 7. CONCLUSION

Peccaries’ extinction at the end of the Pleistocene has presented a problem for extinction theories. Previous work has highlighted their geographic range, supposed flexible dietary capacities

(Graham and Lundelius 1984; Guilday et al. 1971; Schmidt 2008), and populous numbers (Kurtén and

Anderson 1980) implying a species which should resist extinction. Minimal evidence for human predation however suggests overkill is not a likely explanation for their disappearance. This leaves the possibility that climate change impacted these creatures in a manner unanticipated based on the initial interpretations of the species. The research conducted here suggests the latter may be true. Peccaries may not have been highly adaptable or as common as assumed. Combined with their dietary reliance on C3 plants, low population numbers may have made them very susceptible to changing environments.

The Uncommon and Unadaptable Flat-Headed Peccary

The argument for a large population appears uncited in the summary of flat-headed peccaries provided in Kurtén and Anderson (1980). It seems reasonable to assume their interpretation derives from the presence of several large peccary assemblages reported in the late 1960s and 1970s (Appendix A).

However, dated remains from large assemblages are rare (Table 1), leaving unknown the accumulation period for the assemblages.

For the three sites with large MNIs (>4) and more than one date (Peccary Cave, Patton Cave, and Brynjulfson Cave) large date ranges exist (Figure 5). At Peccary Cave, the site with the largest MNI

(61) and smallest range of dates (min. 3000 cal B.P. years) of the three sites, the assemblage could be accumulated with one peccary death every 50 years; hardly evidence of mass death or large populations.

At Patton Cave, two dates represent a minimum date range of 10,000 years cal B.P. With an MNI of 22

(Grady 1988:13), the assemblage could be generated with one peccary death every 450 years. For

Brynjulfson Cave, the minimum calibrated date range of 22,000 years combines with a peccary MNI of

15 (Parmalee and Oesch 1972:37) meaning one peccary would need to die every 1,450 years to create the assemblage. All three of these caves present evidence of time-averaged accumulation. Large peccary assemblages are not indicative of large population numbers at these three locations. 42

While more dates from additional sites are needed to confirm the pattern, if other large assemblages of peccary remains are also time accumulated averages little evidence then exists for the commonality of peccaries on the landscape. Supporting this possibility, most reported finds of peccary

(49 of 96) possess only one or two peccaries (Figure 4). All of the largest assemblages derive from caves, likely representing time-averaged accumulations (Appendix A). While synchronous death locations do exist, these exclusively demonstrate small herds, similar to the behavior of modern peccaries (Sowls

1997). Evidence does not exist to show great numbers of small herds roamed the Pleistocene landscape, though it is plausible increased numbers of dates from peccary sites could provide such evidence. At the moment, all available data point to large assemblages as time-averaged accumulations and few assemblages demonstrating large populations.

Low population numbers are supported by modern peccary reproductive data. Average litter sizes for modern peccary are two infants (Table 10). Once born, greater than 50% infant mortality is seen in modern peccaries (Bissonette 1982:49; Ellisor and Harwell 1979:15). With small litters and high infant mortality it becomes difficult to see how peccaries would have come to dominate the landscape. A loss of apex predators near the end of the Pleistocene resulted in proboscidean population booms (Widga et al.

2017) and it is possible peccary could have experienced similar booms, generating large populations.

However peccaries are not mega-herbivores on the scale of mammoths and mastodons and conceivably would have remained prey for smaller predators incapable of killing proboscideans. Without precise and accurate constraint of regional dating on peccaries a boom-bust series of events cannot be identified. Thus interpretation of peccaries as the most common mid-sized mammal of the Pleistocene appears unsubstantiated and incorrect. More likely they had small populations spread out among relatively small herds.

The adaptability of flat-headed peccary appears to be another assumption lacking strong evidence. Despite wide geographic spread in sites, there is little evidence of widespread population numbers. Large MNI assemblages only emerge in the eastern United States. The largest peccary assemblage west of Missouri is 9 at Goodland, KS followed by an MNI of 5 at Denver, CO (Appendix 43

A). Non-cave sites in the eastern United States also tend toward small MNIs, indicative of few numbers of peccaries. Their presence across the geographic range of North America may represent some adaptability but their greater numbers in the eastern United States than the west shows a potential preference for the region.

Further, from the peccary assemblages associated with environmental reconstructions it appears they preferred cool, seasonally muted, open landscapes (Guilday and Parmalee 1972; Guilday et al. 1971;

Finch et al. 1972). Though caves are time-averaged accumulation, the presence of peccaries in numerous cave assemblages, and modern peccary use of caves, may indicate an adaptive preference to landscapes which were open in terms of vegetation but possessed cave systems. The combination of preferences may have limited the preferred environmental ranges to certain zones to which peccary were specially adapted.

The geographic range of peccary may not indicate adaptability to numerous environments but populations seeking specific environmental niches. Further research on environmental reconstruction from peccary assemblages will help confirm or refute this possibility. Additionally, as most peccary sites remain undated, it is not possible to constrain when peccary used certain landscapes and regions. With such information it may be possible to show a very specific environmental preference or adaptability. At the present though, extant information suggests a specialized environmental niche for flat-headed peccary rather than widespread adaptation to many different environments and locations.

Peccary diet al.so appears inflexible, despite the potential for dietary diversity (Schmidt 2008).

Review of isotopic data demonstrates P. compressus may have specially adapted to a diet based on C3 legumes (Figure 14) regardless of season (Figure 15). Prior reports of peccary dental isotopes (Giesler

2002; MacFadden and Cerling 1996; Yann and DeSantis 2014) show strong reliance on C3 vegetation but

15 do not report δ N values from which to evaluate legume versus non-legume C3 preference. Lack of dietary flexibility throughout a year, or over a lifetime, demonstrates a clear focus on C3 plant life and a lack of adaptation to other food sources.

The typical story of flat-headed peccary has been one of large populations of adaptable creatures roaming the continental United States (Kurtén and Anderson 1980). Based on a more thorough 44

investigation of peccary this interpretation appears faulty. Peccary may not have been overly common and lacked the facilities for rapidly repopulating an environment. Their diet may have relied almost exclusively on C3 legumes and they possessed specific environmental preferences which would have made them susceptible to environmental change brought on by changing climate.

Modern Peccary Reproductive Biology

Lacking commonality and showing evidence of adaptation to a particular food source and region made the flat-head peccary more susceptible to climate change extinction than may have previously been thought. Combining what is known about flat-headed peccary with information on modern peccary biology strengthens this possibility. Flat-headed peccary are inferred to most closely related to the Chacoan peccary (Mayer and Wetzel 1986), however little is known about Chacoan peccary reproduction so information on all three extant peccary species are included here as analogs.

As enumerated in Table 10, average peccary litter size is two. Mortality rate typically is greater than 50% (Bissonette 1982:49; Ellisor and Harwell 1979:15). Gestation is an average of 145 days in (Lochmiller et al. 1984; Sowls 1961, 1966, 1984) and 158 days in white-lipped peccary

( 1966). Modern peccaries are capable of producing one litter per year. While captive peccaries whose infant young are immediately removed after birth may, in a small percentage of instances, become pregnant again straightaway, thus bearing two litters in a year, this has not been shown to happen in wild peccaries (Sowls 1966, 1984). Therefore, it appears modern peccary typically produce one small litter bearing two young per year with only one infant surviving.

Beyond small litter sizes, modern peccary breeding is quite susceptible to environmental conditions. The Chacoan peccary tends to have increased litter size with increased rainfall and reduced litter size in times of dryness (Yahnke et al. 1997). Low (1970:84) reported wild peccaries conceived few to no litters during drought conditions. Sowls (1966, 1968) found during dry conditions with lower quality forage fewer young are born and survive. Drought induced nutritional stress may depress male peccary libido and reduce male breeding activity (Hellgren 1984; Lochmiller et al. 1985:56) while similar poor nutrition results in a reduction, or cessation of, reproductive activity in female peccary (Lochmiller 45

et al. 1986:299). Drought and diet induced stress not only impacts reproductive behavior but reduces the survival of young during dry periods (Ellisor and Harwell 1979; Sowls 1966). While Hellgren et al.

(1985:358,366) found peccaries could still reproduce when facing some diet stress, negative physiological impacts occurred and peccaries eating lower nutrition food took longer to return to estrus and normal ovarian function.

Peccaries are capable breeders during strong environmental conditions but suffer when experiencing drought or a reduction in nutritional value of preferred forage. While typical peccary reproduction during good conditions results in a single litter of two infants with one surviving beyond the first year, reproduction during poor conditions often results in few to no infants. If flat-headed peccary reproduction was similar to the modern analogs, climate change resulting in drying of the landscape could have produced drastic negative impacts on reproduction and survivorship of young.

Extinction of Flat-Headed Peccary on the Ozark Plateau

The flat-headed peccary, as a C3 legume browser lacking large litter sizes and vulnerable to drought-like conditions provides compelling evidence for climate change as the driver of extinction.

Climate change at the end of the Pleistocene would have disrupted herbivore feeding strategies (Graham and Lundelius 1984) and reduced effective nutritional value of vegetation (Guthrie 1982). Similar changes today would have very detrimental impacts on modern peccaries and it appears they likely had a drastic impact on those living in the Pleistocene.

In food-rich environments the evolution of novelty, or specialization, is well served (Geist

1998, 1999). Specialization allows a creature to possess a defined food-source to which it may be the only, or one of a few, users. Based on the dietary information available for peccaries via stable isotope analysis, Pleistocene peccaries were highly C3 legume adapted with little to no seasonal variation, evidence of specialization. When food becomes limited, efficiency phenotypes such as the ability to be a dietary generalist or produce large numbers of offspring, become favored (Geist 1998). Based on modern peccary reproduction, peccaries could not produce large numbers of offspring, and despite the potential for dietary plasticity no evidence of this exists. When environments shift and food-rich locations become 46

food-limited ecosystems specialization puts mega-herbivores and other specialized fauna at greater risk for extinction than animals with efficiency phenotypes (Geist 1998, 1999). The environment during the

Pleistocene was food rich (Huston 1993; Huston and Wolverton 2009) which would have enabled specialized adaptation but changing climate could have made the succeeding landscape food poor for peccaries.

Peccaries fit the pattern of dispersal phenotype adaptation as described by Wolverton et al.

(2009). They utilized herd behavior to protect young, possessed relatively small litter sizes, were larger than present peccaries, and developed during a time of food-richness. C3 specialization was viable during the Pleistocene for peccaries, as low temperatures enabled growth of their preferred dietary vegetation.

Climate warming as the Pleistocene ended meant C4 plants began to vastly outperform C3. A C3 specialization would have then shifted from a boon to a detriment for the flat-headed peccary.

As shown by Wolverton et al. (2009:39) when environmental disturbances increase they disproportionately impact fauna which have developed specializations and novelty over generalization and plasticity. In the LGM of the southeastern United States a changing environment existed.

Environmental disturbances were commonplace, demonstrated by biomass, wetness, and temperature changes shown in studies of Ozark Pleistocene environments (Table 2). Changes in the terminal

Pleistocene to intra-annual growing seasons (Kiltie 1984) furthered environmental disturbances.

Johnson (2002) found mammals with relatively low reproductive rates were prone to extinction during the late Pleistocene. The species with low reproductive rates that survived were adapted to closed forest environments rather than open steppe. Cardillo (2003) correlated litter size strongly with extinction, showing smaller litter size animals possessed greater risk of extinction. Peccaries, though possessing a relatively short gestation, had small litter sizes. Further, inhibition of reproduction would have occurred during drying conditions. As has been shown, environmental conditions as the Pleistocene came to an end were not ideal for flat-headed peccary. Warming and drying of the environment occurred (Table 2) around the same time peccary use of the cave ceased. 47

Using the flat-headed peccary population from Peccary Cave as a case example, climate change explains their disappearance. The peccaries recovered from the cave were specialized in their diet, focused on C3 legumes year-round. Based on the recovery of fetal and mature peccary skeletal element frequencies, pregnant mothers were dying naturally in Peccary Cave. This may be expected as modern peccaries respond poorly to dry conditions that reduce nutritional diet. Near the end of the peccary use of

Peccary Cave the environment in the Ozark region was warming and drying out. Such drying, along with environmental shifts, would have disrupted the ecosystem for peccaries, resulting in vegetation change and dietary stress. The environment would have become untenable due to negative impacts on reproduction from drying conditions, forcing either extinction or migration.

Coinciding with peccary and dire wolf extirpation at Peccary Cave, at least six taxa of microfauna went regionally extinct and six additional taxa ceased use of the Ozark Plateau for around

6,000 years (Figure 8). Even the most adaptable of microfauna present in the Ozarks, the eastern woodrat, disappeared from the landscape for thousands of years. Their adaptability should have enabled them to tolerate increased seasonality and warmer and drier conditions (Rainey 1953). However, one of the food sources for eastern woodrat are legumes (Jones et al. 1985:224). It is possible climate change resulted in a loss of leguminous forage in the Ozark Plateau between 19,000 and 13,000 ka, explaining the loss of eastern woodrat and flat-headed peccary. Even if leguminous forage was still present, the stark pattern of temporary abandonment of the Ozark Plateau by fauna around 19,000 ka strongly suggests the area became inhospitable. Peccaries and dire wolves were unable to maintain their presence in the region during this change.

Only near the end of the Allerød and beginning of the Younger Dryas periods do fauna reappear in the dated record on the Ozark Plateau. Most of the returning fauna are warm tolerant, such as the beautiful armadillo, suggesting by the time the region became inhabitable again the area was also warming. At this time long-horned bison appeared in the Ozark Plateau (Lyman and Bassett 2004) for the first time. Bison are known for their ability to subsist on numerous food sources, so their appearance only at a time when other fauna also reappear on the Ozark Plateau lends support to the idea the area was 48

uninhabitable between ~19,000 and 13,000 ka. While more radiocarbon dates from Ozark Plateau

Pleistocene faunal assemblages are required to test this possibility, extant data suggests the climate became intolerable for fauna around ~19,000 ka, coinciding with the regional extirpation of flat-headed peccary and dire wolf at Peccary Cave.

Extinction of peccaries may have impacted survival of dire wolves as well. As seen in the remains from Peccary Cave, dire wolves likely were preying on fauna with similar isotopic signatures to peccaries. Dating from Peccary Cave specimens demonstrate dire wolves and peccaries overlapped spatially and temporally. Though dire wolf dates outside of Rancho la Brea are poorly constrained the disappearance of dire wolves from Peccary Cave appears to coincide with the loss of the peccaries amid changing climate. Dire wolves do not appear to persist past ~13,000 14C years in eastern North America

(Krueger and Weeks 1965). While their disappearance may predate peccaries as a whole, minimal peccary populations may have reduced encounter rates for dire wolves and caused, along with other mammalian extinction, a dramatic reduction in available, obtainable prey. Much more research is needed to investigate this possibility but such a pattern is plausible from review of data from Peccary Cave.

Warming and drying, associated with a decrease in C3 and increase in C4 plant life in the

Ozarks explains regional peccary extirpation at Peccary Cave. The pattern and data available from skeletal element frequencies, radiocarbon dates, and stable isotopes of the Peccary Cave assemblage demonstrates climate change would have stressed peccary populations immensely. With small litter sizes and reproductive cycles likely to produce only a single litter per year, peccaries were not poised to compensate for population losses. Their specialization in C3 legumes and reproductive response to dry conditions made the animal vulnerable to climate change, patterns of which are present during the later

Pleistocene. Peccaries appear not to go extinct due to kill off by humans. The most likely cause of extinction of flat-headed peccary was climate change.

49

Figure 1. Location of Peccary Cave and other selected places on and around the Ozark Plateau.

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Figure 2. Location of published records of peccary (P. compressus and Platygonus cf. compressus) in the contiguous United States (n = 88) (Appendix A): 1) McKittrick, CA; 2) Rancho La Brea, CA; 3) Diamond Valley Lake, CA; 4) Butterworth, ID; 5) Oven, NM; 6) Sheep Camp Shelter, NM; 7) Pit Stop Quarry, AZ; 8) Papago Springs Cave, AZ; 9) Agate Basin, WY; 10) Hay Springs, NE; 11) Smith Falls, NE; 12) Denver Juvenile Hall, CO; 13) Selby, CO; 14) Dutton, CO; 15) Gillen Pits, NE; 16) Davidson Pits, NE; 17) Goodland, KS; 18) Cragin Quarry, KS; 19) Jones Ranch, KS; 20) Canyon Basin, TX; 21) Rock Creek, TX; 22) Lubbock Lake, TX; 23) Yukon, OK; 24) Rush Creek, OK; 25) Moore Pit, TX; 26) Zesch Cave, TX; 27) Laubach Cave, TX; 28) Hall’s Cave, TX; 29) Friesenhahn Cave, TX; 30) Ingleside, TX; 31) Blue Mounds, WI; 32) Dubuque, IA; 33) Galena, IL; 34) Burlington, IA; 35) Brynjulfson Caves, MO; 36) Benton Co., MO; 37) Elisha Estes Cave, MO; 38) Jacob’s Cave, MO; 39) Bat Cave, MO; 40) Great Spirit Cave, MO; 41) Roubidoux Cave, MO; 42) Little Beaver Cave, MO; 43) Ozark Underground Laboratory, MO; 44) Zoo Cave, MO; 45) Peccary Cave, AR; 46) Alton, IL; 47) Cherokee Cave, MO; 48) Crevice Cave, MO; 49) Chickasaw Bluffs, KY; 50) Belding, MI; 51) Ann Arbor, MI; 52) Fremont, OH; 53) Sheriden Cave, OH; 54) Laketon, IN; 55) Marion Co., IN; 56) Columbus, OH; 57) Indun Rockshelter, IN; 58) Reading, OH; 59) Megenity Peccary Cave, IN; 60) Welsh Cave, KY; 61) Lone Star Peccary Cave, KY; 62) Cave of the Skulls, TN; 63) Rochester, NY; 64) Gainesville, NY; 65) Mosherville, PA; 66) Dutchess Quarry Caves, NY; 67) Hot Run, VA; 68) New Trout Cave, WV; 69) Augusta Co., VA; 70) Patton Cave, WV; 71) Rogersville, TN; 72) Baker Bluff Cave, TN; 73) Guy Wilson Cave, TN; 74) Yarbrough Cave, GA; 75) Ladds Quarry, GA; 76) Haile, FL; 77) Arredondo, FL; 78) Reddick, FL; 79) Devil’s Den, FL; 80) Waccasassa River, FL; 81) Rock Springs, FL; 82) Melbourne, FL; 83) Seminole Field, FL; 84) Pool Branch, FL; 85) Sebastian Canal, FL; 86) Vero, FL; 87) Cutler Hammock, FL; 88) Monkey Jungle Hammock, FL.

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Figure 3. General character of peccary assemblages (n = 91) (Appendix A).

Figure 4. MNI estimates for peccary assemblages (n = 91) (Appendix A).

52

Figure 5. Radiocarbon ages for peccary (P. compressus) (see also Table 1). Each assay is either a conventional or AMS radiocarbon age on bone, and is calibrated using OxCal 4.2 (Bronk Ramsey 2009) and IntCal09 (Reimer et al. 2009).

53

Figure 6. Location of Peccary Cave at the confluence of Ben’s Branch and Cave Creek. Inset shows the location in northeast Arkansas.

Figure 7. Plan map of Peccary Cave, including excavation trenches and sections (after Davis 1973:Figure 1).

54

Figure 8. AMS radiocarbon ages for selected taxa on the Ozark Plateau. Ages from microfauna and beautiful armadillo from Semken et al. (2010:Table 3), long-horned bison from Lyman and Bassett (2004), and short-faced bear from Schubert (2004). Each assay is a direct age on bone, and is calibrated using OxCal 4.2 (Bronk Ramsey 2009) and IntCal09 (Reimer et al. 2009).

Figure 9. AMS radiocarbon ages for peccary and dire wolf. Each assay is a direct age on bone, and is calibrated using OxCal 4.2 (Bronk Ramsey 2009) and IntCal09 (Reimer et al. 2009).

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Figure 10. Box-and-whisker plot of mandibular third molar length in several peccary assemblages. Peccary Cave (PC), Bat Cave (BC, Woodruff 2012:Appendix D), Sheriden Cave (SC, Gobetz 1998:Appendix B), and Laubach Cave (LC, Slaughter 1966a:Table 2). The number of measured specimens in each sample is indicated in the respective box.

Figure 11. Examples of breccia coverage on peccary metapodials. From left to right are percent coverages: 0, 1-25, 26-50, 51-75, and 76-100.

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Figure 12. Skeletal element profiles for sub/adult and fetal/neonate peccary.

Figure 13. Distribution of selected classes of faunal and archaeological material.

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Figure 14. Stable isotope values for peccary and dire wolf (Table 3). Legume data from DeNiro and Hastorf (1985:Table 3).

Figure 15. Intratooth δ13C and δ18O profiles for peccary mandibular canines. 58

Table 1. Direct dates for peccary. Locality 14C Age (B.P.) Reference Comments Fremont, OH 4,290 ± 150 Hoare 1964:427 Brynjulfson Cave #1, MO 9,440 ± 760 Coleman 1973:84 mixed species Baker Bluff, TN 10,560 ± 220 Guilday et al. 1978 Ann Arbor, MI 10,790 ± 246 Eshelman et al. 1972:246 Gainesville, NY 10,750 ± 50 Feranec and Kozlowski 2010:206 AMS Sheriden Cave, OH 11,060 ± 60 Tankersley 1997:718 AMS Sheriden Cave, OH 11,130 ± 60 Tankersley 1999:Table 9.4 AMS Butterworth, ID 11,340 ± 50 McDonald 2002:142 AMS Baker Bluff, TN 11,640 ± 250 Guilday et al. 1978 Mosherville, PA 11,900 ± 750 Ray et al. 1970:84 Oven, NM 12,060 ± 1,780 Lucas and Smartt 1995:293 Dutchess Quarry Caves, NY 12,160 ± 80 Steadman et al. 1997:Table 3 AMS Dutchess Quarry Caves, NY 12,220 ± 60 Steadman et al. 1997:Table 3 AMS Dutchess Quarry Caves, NY 12,430 ± 70 Steadman et al. 1997:Table 3 AMS Welsh Cave, KY 12,950 ± 550 Buckley and Willis 1972:116 Patton Cave, WV 13,350 ± 120 Grady 1988:10, 15 Guy Wilson Cave, TN 19,700 ± 600 Buckley and Willis 1972:115 Peccary Cave, AR 17,990 ± 100 this study, Table 3 AMS Baker Bluff, TN 19,100 ± 850 Guilday et al. 1978 Peccary Cave, AR 19,460 ± 130 this study, Table 3 AMS Peccary Cave, AR 19,600 ± 70 this study, Table 3 AMS Peccary Cave, AR 20,500 ± 80 this study, Table 3 AMS Peccary Cave, AR 20,610 ± 140 this study, Table 3 AMS Patton Cave, WV 22,620 ± 240 Grady 1988:10, 15 Brynjulfson Cave #1, MO 34,600 ± 2,100 Coleman and Liu 1975:172 Note: all dates are conventional radiocarbon ages unless noted in the comments.

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Table 2. Ozark Plateau environment from Pre-LGM through the Oldest Dryas. Period ka Peccary Cave Taxona Environment References Pre-LGM >22.2 Canis dirus Open pine parkland characterized by cool, Delcourt and Delcourt (1996); Neotoma floridana equable seasons. Dorale et al. (1998); King (1973); Platygonus compressus King and Lindsay (1976) Early LGM 20 – 22.2 Blarina carolinensis Cool, wet spruce dominated forest. Loss of Haynes (1985); King and Lindsay C. dirus climate stability beginning. (1976); Royall et al. (1991); Clethrionomys gapperi Semken et al. (2010) Geomys bursarius N. floridana P. compressus Late LGM 16.8 – 20 C. gapperi Landscape warming and drying. King (1973); Semken (1984); Dasypus bellus Thermophile plants emerge and greater Semken et al. (2010) Microtus xanthognathus seasonal variation occurs. N. floridana Phenacomys intermedius Scalopus aquaticus

Synaptomys borealis 5

Oldest Dryas 14.6 – 16.8 No dated specimens Steppe environment with coniferous forest Semken (1984) 9 patches and mixed forest parkland. aPeccary Cave taxon data from Semken et al. (2010) and this volume.

Table 3. Radiocarbon AMS measurements and δ13C results on peccary and dire wolf. Lab. no. KWNO. Cat. no. Taxon Element Provenience (UCI-AMS) 930 67-300-31 P. compressus right HM T3-S4-L1 165399 923 65-158-12 P. compressus right HM T2-S6-L2 165398 916 67-300-4 P. compressus right HM T2-S7-L3 175968 931 67-300-254 P. compressus right HM T21-S1-L4 175967 918 67-300-51 P. compressus right HM T11-S3-L2 165400 2964 67-300-239 C. dirus left CL T15-S5-L1 165393 2958 67-300-254 C. dirus right MCV T21-S1-L4 175965 2959 67-300-51 C. dirus right MCV T11-S3-L2 175966 Note: OxCal 4.2 (Bronk Ramsey 2009) and IntCal09 (Reimer et al. 2009) used to calibrate measured radiocarbon ages.

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Table 4. Summary of rodent modification. Type Sub/Adult Fetal/Neonate dire wolf peccary-size No modification 1,928 1,184 12 705 1 surface 208 4 0 63 2 surfaces 74 1 0 18 3 surfaces 27 0 0 9 4+ surfaces 95 0 2 30 Medullary cavity 49 0 0 3 exposed Gnawing on proximal 25 1 0 0 end of specimen Gnawing on distal 65 0 0 2 end of specimen Gnawing on proximal 33 0 0 8 and distal ends of specimen Gnawing on lateral 3 0 0 0 portion of specimen Gnawing on medial 3 0 0 0 portion of specimen Gnawing on medial 5 1 0 0 and lateral portions of specimen Indeterminate 189 1 0 85 NISPt 2,704 1,192 14 923

Table 5. Summary of breccia coverage. % Coverage Taxon NISP 0 1-25 26-50 51-75 76-100 sub/adult peccary 2,704 1,853 416 157 87 191 fetal/neonate peccary 1,192 1,036 90 52 13 1 dire wolf 14 9 4 1 0 0 61

Table 6. Summary of peccary remains. Sub/Adult Fetal/Neonate Element NISP MNE MNI NISP MNE MNI cranium 283 45 45 57 8 5 mandible 569 61 61 32 10 5 stylohyoid 0 0 - 0 0 - atlas vertebra 21 11 - 0 0 - axis vertebra 12 12 - 0 0 - cervical vertebra (3-7) 59 48 - 0 0 - thoracic vertebra (1-14) 131 103 - 25 19 - rib 42 42 - 367 130 - costal cartilage 3 3 - 0 0 - manubrium 1 1 - 0 0 - lumbar vertebra (1-5) 58 50 - 0 0 - sacrum 29 12 12 3 3 3 caudal vertebra 0 0 - 0 0 - scapula 99 44 23 24 18 9 humerus 192 83 41 23 22 12 radius-ulna 134 55 31 42 20 12 cuneiform 3 3 2 0 0 0 lunar 4 4 2 0 0 0 scaphoid 11 11 7 0 0 0 magnum 10 10 5 0 0 0 unciform 5 5 3 0 0 0 metacarpal III-IV 85 41 24 1 1 1 innominate 80 38 22 33 21 11 femur 181 73 30 28 25 15 patella 18 18 11 1 1 1 tibia 131 60 32 23 22 12 fibula 26 10 - 23 16 - astragalus 68 68 34 19 19 2 calcaneus 73 73 38 27 26 13 first cuneiform 3 3 2 0 0 0 cuboid 6 6 4 0 0 0 navicular 17 17 10 0 0 0 metatarsal III-IV 74 53 30 1 1 1 metapodial 109 - - 120 - - first phalanx 81 76 - 60 60 - second phalanx 52 50 - 40 40 - third phalanx 31 30 - 11 11 - proximal sesamoid 3 3 - 0 0 - distal sesamoid 0 0 - 0 0 -

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Table 7. Summary of sub/adult peccary teeth, expressed as MNE (isolated and emplaced combined). Mandibular Maxillary Tooth left right n left right n deciduous canine 12 11 21 2 2 0 canine 61 50 26 45 34 2 deciduous incisors (1-2) 1 1 0 0 0 0 incisors (1-2) 6 7 4 1 1 0 deciduous 2nd premolar 1 0 0 1 0 0 deciduous 3rd premolar 3 7 0 0 0 0 deciduous 4th premolar 18 17 0 0 0 0 2nd premolar 9 8 0 5 5 1 3rd premolar 15 8 0 8 6 0 4th premolar 16 10 0 6 4 1 1st molar 47 51 0 29 27 1 2nd molar 43 36 0 23 17 0 3rd molar 23 23 0 8 16 1

Table 8. Summary of proximal epiphyseal union for sub/adult peccary calcanea from Peccary Cave. MNE Side Proximal Left Right MNI unfusedUnion 14 10 14 partially 0 0 0 fused line 1 3 3 completelyvisible 12 12 12 indeterminatefused 8 13 13

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Table 9. Spearman rank-order correlations (rs) between scan site MNE and scan site volume density for sub/adult peccary.

Element N r p s humerus 5 1.00 <.001**

radius-ulna 7 .536 .108 femur 6 .600 .104 calcaneus 4 .800 .100 astragalus 2 - - tibia 5 .300 .312 assemblagea 22 .527 .006** Note: N (number of ranks); *p <.05, >.01; **p <.01. aFour proximal long bones plus calcaneus and astragalus.

Table 10. Records of litter size in modern peccaries. Litter Size (number of Taxon Context Recorded individuals) Reference P. tajacu captive live 1.88 ± 0.13 Hellgren et al. 1985:360 P. tajacu captive in utero 2 Smith and Sowls 1975:620 P. tajacu captive live 1.8 ± 0.1 Lochmiller et al. 1984:146 P. tajacu captive live 2 Sowls 1984:70 P. tajacu wild live 2.2 Sowls 1984:70 P. tajacu wild and captive in utero 1.98 Sowls 1984:70 P. tajacu captive live 2.17 Sowls 1966:163 P. tajacu wild in utero 2.1 Sowls 1966:163 P. tajacu wild in utero 1.28 Jennings and Harris 1953:21 P. tajacu wild in utero 1.81 Low 1970:90 P. tajacu wild live 1.44 Knipe 1957:31 P. tajacu wild live 1.81 Neal 1959:179 P. tajacu wild live 1.5 Ellisor and Harwell 1979:15 P. tajacu wild live 1.3 Bissonette 1982:49 P. tajacu wild in utero 1.7 Bodmer 1989b:Table 4 C. wagneri wild in utero 2.72 Mayer and Brandt 1982:443 C. wagneri wild live 2.46 Mayer and Brandt 1982:443 C. wagneri captive live 2.4 ± 0.8 Yahnke et al. 1997:303 C. wagneri wild live 1.7 Taber et al. 1993:450 C. wagneri captive live 2.23 Brooks 1992:316 T. wild in utero 1.63 Bodmer 1989b:Table 4 T. pecari wild in utero 2 Leopold 1959:498 T. pecari captive live 2 Roots 1966:199 T. pecari/P. tajacu hybrid captive live 2 Zuckerman 1953:906

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APPENDIX A. INFORMATION FOR LOCALITIES WITH REPORTED REMAINS OF FLAT-HEADED PECCARY (P. COMPRESSUS AND PLATYGONUS CF. COMPRESSUS).1

igure 2 igure Locality F MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments 44WR61, VA - - - 8 1 1 - - Haynes 1978 no mention of peccary misc. elements (NISP ≥1, NISP Agate Basin, WY 9 1 1 1 1 1 ≤ 5) mandibular condyle Walker 1982:296 isolated tooth, teeth, or tooth "Platygonus Alton, IL 46 1 1 2 - - row (NISP ≥ 1, NISP ≤ 10) mandibular canine Hay 1920:111 compressus?" American Falls, ID - - - 8 1 1 - - Gazin 1935; Hopkins et al. 1969 no mention of peccary "the three skeletons were assumed to be in an articulated condition" Ann Arbor, MI 51 3 251 1 1 1 "herd" (MNI ≥ 3) (Eshelman et al. 1972:250) Eshelman et al. 1972 Arredondo IIA, FL 77 ? ? 1 1 1 unspecified unspecified Webb and Wilkins 1984:Table 1 "primarily of foot and lower

leg bones (astragalus, 8 8

metapodials, phalanges)" misc. elements (NISP ≥ 20, (Yates and Lundelius Aubrey, TX - ? 23 3 1 1 NISP ≤ 30) 2001:108) Yates and Lundelius 2001:Table 8.1, 108 isolated tooth, teeth, or tooth Augusta Co., VA 69 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) dp4 Leidy 1860:100 isolated tooth, teeth, or tooth Baker Bluff Cave, TN 72 2 2 1 1 1 row (NISP ≥ 1, NISP ≤ 10) maxillary molar and dp2 Guilday et al. 1978:Table 1 NISP includes 34 fetal remains; NISP does not include dental rows with emplaced teeth because unable to determine if each Hawksley et al. 1973:Table 1; Woodruff row include more than Bat Cave, MO 39 70 >2,680 1 1 1 NISP ≥ 1,000 multiple 2016:62, 69, Appendix A one tooth "five specimens [i.e., skeletons] are represented in the collection" (Wagner Belding, MI 50 5 ? 1 1 - "herd" (MNI ≥ 3) 1903:777) Finch et al. 1972:20; Wagner 1903:777 isolated tooth, teeth, or tooth mandibular canine attached Benton Co., MO 36 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) to fragmentary symphysis LeConte 1852:5; Leidy 1860:100

89

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments "several fragments of jaws and some teeth, with a variety of other bones of the teeth plus other (NISP ≥1, peccary…" (Whitney Blue Mounds, WI 31 ? ? 1 - - NISP ≤ 10) 1862:135) Whitney 1862:135 Brynjulfson Cave #1, MO 35 15 475 1 1 1 NISP ≥ 100, NISP ≤ 500 multiple Parmalee and Oesch 1972:Table 3 isolated tooth, teeth, or tooth Brynjulfson Cave #2, MO 35 1 2 1 1 1 row (NISP ≥ 1, NISP ≤ 10) two isolated cheek teeth Parmalee and Oesch 1972:Table 4 Burlington, IA 34 ? ? 1 - - unspecified unspecified Whitney 1862:135 "almost complete skeleton" Butterworth, ID 4 1 54 1 1 - "skeleton" (MNI ≥ 1, MNI ≤ 2) (McDonald 2002:141) McDonald 2002; Stokes 1953 "skeletons of an adult and a

peccary baby" (Dalquest and 8

Canyon Basin, TX 20 2 ? 1 - - "skeleton" (MNI ≥ 1, MNI ≤ 2) Schultz 1992:53) Dalquest and Schultz 1992:53 9

isolated tooth, teeth, or tooth identified by Clayton Castle Rock Cave, WI - 1 1 4 1 1 row (NISP ≥ 1, NISP ≤ 10) canine Palmer 1974:219 E. Ray

Cave of the Skulls, TN 62 12 ? 1 1 - NISP ≥ 501, NISP ≤ 1,000 unspecified Corgan and Breitburg 1996:143 MNI = 21 (43 m3s divided by 2, and rounding down) (Simpson 1949:Table 4); NISP from Cherokee Cave, MO 47 21 2-3,000 1 1 1 NISP ≥ 1,000 multiple Simpson 1946, 1949 Simpson (1946) "a herd of five adult peccaries" (Finch et al. Chickasaw Bluffs, KY 49 5 ? 1 1 1 "herd" (MNI ≥ 3) 1972:1) Finch et al. 1972 "twelve nearly complete skeletons…" (Newberry 1875:vi); two "nests" bones Columbus, OH 56 12 ? 1 - - "herd" (MNI ≥ 3) (Klippart 1875:1) Klippart 1875; Newberry 1875:vi isolated tooth, teeth, or tooth maxillary tooth row with M2- Cragin Quarry (Loc. 1), KS 18 1 1 2 2 - row (NISP ≥ 1, NISP ≤ 10) 3 Hibbard and Taylor 1960:183-184 Crevice Cave, MO 48 2 17 1 1 1 NISP ≥ 11, NISP ≤ 20 multiple Hawksley 1986:23

90

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments

Cutler Hammock, FL 87 ? ? 1 1 1 unspecified unspecified Emslie and Morgan 1995:Table 1 isolated tooth, teeth, or tooth Davidson Pits (Rw-102), NE 16 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) partial maxillary tooth row Corner 1977:84 "the five individuals were found pointing toward…" Denver Juvenile Hall, CO 12 5 ? 1 1 - "herd" (MNI ≥ 3) (Lewis 1970:B140) Lewis 1970:B139-140 Devil's Den, FL 79 ? 10 1 1 1 NISP ≥ 11, NISP ≤ 20 multiple Martin and Webb 1974:132 Diamond Valley Lake, CA 3 1 1 1 - - unspecified unspecified Springer et al. 2009:Table 1, Appendix "A number of teeth and teeth plus other (NISP ≥1, bones of [peccary].." Dubuque, IA 32 ? ? 1 - - NISP ≤ 10) (Whitney 1862:135) Whitney 1862:135

isolated tooth, teeth, or tooth 9

Dutchess Quarry Caves, NY 66 1 2 1 - - row (NISP ≥ 1, NISP ≤ 10) two mandibular canines Steadman et al. 1997:Table 2, 112 0 teeth plus other (NISP ≥1, mandible fragments, various Dutton, CO 14 2 >8 1 1 1 NISP ≤ 10) teeth, and first phalange Graham 1981:44 Eshelman and Grady (1986:Table 3) report P. compressus, which Early's Cave, VA - - - 5 - - - - Hay 1923:221 appears to be an error. isolated tooth, teeth, or tooth Elisha Estes Cave, MO 37 1 1 1 1 1 row (NISP ≥ 1, NISP ≤ 10) mandibular canine Hawksley 1986:4 "The numbers of various bones present…point out the presence of at least four individual peccaries" (Hoare Finch et al. 1972:19; Hoare 1964; Hoare et Fremont, OH 52 4 432 1 1 1 "herd" (MNI ≥ 3) et al. 1964:212). al. 1964:207-212, Table 8 teeth plus other (NISP ≥1, Friesenhahn Cave, TX 29 1 2 1 1 1 NISP ≤ 10) m3 and metatarsal Graham 1976b:132 "The material…belongs to two individuals…" (Clarke Clarke 1916:34; Feranec and Kozlowski Gainesville, NY 64 2 21 1 - - "skeleton" (MNI ≥ 1, MNI ≤ 2) 1916:34) 2010:206; Hartnagel and Bishop 1922:85

Galena, IL 33 ? 15 1 1 1 NISP ≥ 11, NISP ≤ 20 multiple LeConte 1848 isolated tooth, teeth, or tooth partial maxillary tooth row Gillen Pits (Rw-101), NE 15 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) with left/right P3-4, M2-3 Corner 1977:84

91

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments "In all, nine animals have been uncovered, all lying P. leptorhinus (= close together…"( Williston Wagner 1903:708; Williston 1894:23, compressus) (Wagner Goodland, KS 17 9 ? 1 1 1 "herd" (MNI ≥ 3) 1894:23) 1897:303 1903:780) teeth plus other (NISP ≥1, Great Spirit Cave, MO 40 1 2 1 1 1 NISP ≤ 10) canine and tibia Hawksley 1986:40 Guy Wilson Cave, TN 73 16 1,086 1 1 1 NISP ≥ 1,000 multiple Nye 2007:Table 7, Appendix B Haile 7A, FL 76 ? ? 1 1 1 unspecified unspecified Webb 1974:Table 2.1 isolated tooth, teeth, or tooth Yann and DeSantis 2014:Supplementary Haile 8A, FL 76 1 4 1 1 1 row (NISP ≥ 1, NISP ≤ 10) teeth Table 1 teeth plus other (NISP ≥1, mandible, two isolated teeth, Hall's Cave, TX 28 1 4 1 1 1 NISP ≤ 10) and a third phalange Toomey 1993:414

isolated tooth, teeth, or tooth "upper and lower jaws" 9 Hay Springs, NE 10 ? ? 1 - - row (NISP ≥ 1, NISP ≤ 10) (Matthew 1902:318) Matthew 1902:318 1

Hill Peccary, NY - - - 8 1 - - - teeth plus other (NISP ≥1, mandibular canine plus three Hot Run, VA 67 1 4 1 1 1 NISP ≤ 10) appendicular elements Ott and Weems 1986:130 isolated tooth, teeth, or tooth deciduous incisors and Indun Rockshelter, IN 57 1 9 2 2 - row (NISP ≥ 1, NISP ≤ 10) premolars only Richards and Munson 1988:Table 1 teeth plus other (NISP ≥1, 7 isolated teeth, 1 premaxilla, Ingleside, TX 30 1 9 1 1 - NISP ≤ 10) and 1 metatarsal Lundelius 1972:44 Jacob's Cave, MO 38 7 221 1 1 1 NISP ≥ 100, NISP ≤ 500 multiple Hawksley 1986:Table 1 isolated tooth, teeth, or tooth Davis 1987:101; Frye and Hibbard Jones Ranch, KS 19 1 2 1 1 1 row (NISP ≥ 1, NISP ≤ 10) two fragmentary molars 1941:419; Hibbard 1958:21 isolated tooth, teeth, or tooth Ladds Quarry, GA 75 1 1 1 1 1 row (NISP ≥ 1, NISP ≤ 10) left m3 Ray 1967:147 isolated tooth, teeth, or tooth Laketon, IN 54 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) mandible with p2-m2 Cope and Wortman 1884:20 MNI = 9 (19 m2s divided by 2, and rounding down) (Slaughter 1966:Table Laubach Cave, TX 27 9 ? 1 1 1 NISP ≥ 501, NISP ≤ 1,000 multiple Slaughter 1966 3) unerupted maxillary premolar isolated tooth, teeth, or tooth and fragmentary, unerupted Little Beaver Cave, MO 42 1 2 1 1 - row (NISP ≥ 1, NISP ≤ 10) cheeth tooth Schubert 2003:189

92

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments "partial skeleton from a young adult flat-headed peccary" (Wilson et al. Lone Star Peccary Cave, KY 61 1 25 1 1 1 "skeleton" (MNI ≥ 1, MNI ≤ 2) 1975:83) Wilson et al. 1975:83 isolated tooth, teeth, or tooth from water screen in Lubbock Lake, TX 22 1 1 1 1 1 row (NISP ≥ 1, NISP ≤ 10) pm2 Johnson 1987:82 Clovis component "The five peccaries…died simultaneously, and their proximity argues that this was a herd" (Munson Marion Co., IN 55 5 ? 1 - - "herd" (MNI ≥ 3) 1991:207) Munson 1991 mandibular canine, mandible teeth plus other (NISP ≥1, fragment with dp4, p3-m3,

McKittrick, CA 1 1 3 2 6 - NISP ≤ 10) and metacarpal Schultz 1937:150; Stock 1928:25

9 2

Megenity Peccary Cave, IN 59 >500 ? 1 1 1 NISP ≥ 1,000 multiple Richards 1988; Schmidt 2008:98 isolated tooth, teeth, or tooth Ray 1958:442-445; Simpson 1929:253; Melbourne, FL 82 1 5 1 1 1 row (NISP ≥ 1, NISP ≤ 10) teeth Webb 1974:Table 2.1 Monkey Jungle Hammock, FL 88 ? ? 1 1 1 unspecified unspecified Morgan 2002:Table 1 isolated tooth, teeth, or tooth Moore Pit, TX 25 1 1 2 2 - row (NISP ≥ 1, NISP ≤ 10) M2 Slaughter 1966:81 "major part of a skeleton" Mosherville, PA 65 1 ? 1 - - "skeleton" (MNI ≥ 1, MNI ≤ 2) (Ray et al. 1970:78) Ray et al. 1970:78 isolated tooth, teeth, or tooth New Trout Cave, WV 68 ? ? 1 1 1 row (NISP ≥ 1, NISP ≤ 10) teeth Grady and Garton 1982:67 "partial peccary skeleton" (Lucas and Smartt 1995:293) and a fragmentary canine, vertebrae, and other Oven, NM 5 2 ? 1 - - "skeleton" (MNI ≥ 1, MNI ≤ 2) fragments Lucas and Smartt 1995:293 Ozark Underground teeth plus other (NISP ≥1, mandibular canine and distal Laboratory, MO 43 1 2 1 1 1 NISP ≤ 10) femur Hawksley 1986:52 P. alemanii (= compressus) Papago Springs Cave, AZ 8 4 >15 1 - - NISP ≥ 11, NISP ≤ 20 multiple Skinner 1942:172-173; Slaughter 1966:488 (Slaughter 1966:488)

Patton Cave (lower), WV 70 22 644 1 1 1 NISP ≥ 501, NISP ≤ 1,000 multiple Grady 1988:13 Patton Cave (upper), WV 70 6 108 1 1 1 NISP ≥ 100, NISP ≤ 500 multiple Grady 1988:13

93

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments MNI and NISP = Peccary Cave, AR 45 61 2,704 1 1 1 NISP ≥ 1,000 multiple this study sub/adult isolated tooth, teeth, or tooth Pit Stop Quarry, AZ 7 1 1 1 - 1 row (NISP ≥ 1, NISP ≤ 10) complete mandible Murray et al. 2005:366-367 Pool Branch, FL 84 ? ? 1 1 - unspecified unspecified Webb 1974:Table 2.1 misc. elements (NISP ≥1, NISP partial cranium plus three Merriam and Stock 1921; Stock 1929:Figure Rancho La Brea, CA 2 2 4 1 1 1 ≤ 5) appendicular elements 3 "one peccary skeleton..and part of a second…. more than 1½ skeletons may have been present " (Finch et al. Reading, OH 58 2 ? 1 3 3 "skeleton" (MNI ≥ 1, MNI ≤ 2) 1972:19) Finch et al. 1972:19 classification ambiguous:

"one peccary tooth" Platygonus cf. 9

isolated tooth, teeth, or tooth (Dalquest and Schultz compressus or 3 Red Springs, TX - 1 1 3 2 - row (NISP ≥ 1, NISP ≤ 10) 1992:59) Dalquest and Schultz 1992:59-60 Platygonus sp. Reddick (IA, IB, and 2C), Gut and Ray 1964:326; Martin 1974:84; FL 78 ? ? 1 1 1 unspecified unspecified Webb 1974:Table 2.1 "two adult individuals…. Of one individual is the greater part of the skeleton…. Of the second individual there is a less perfect skull with the Finch et al. 1972:20; Hartnagel and Bishop Rochester, NY 63 2 ? 1 - - "skeleton" (MNI ≥ 1, MNI ≤ 2) upper teeth" (Leidy 1889:41) 1922:85; Leidy 1889:41 mandibular tooth row with isolated tooth, teeth, or tooth p4-m2? and isolated Rock Creek, TX 21 1 2 1 7 - row (NISP ≥ 1, NISP ≤ 10) mandibular canine Troxell 1915:635 Rock Springs, FL 81 ? ? 1 - - unspecified unspecified Webb 1974:Table 2.1 teeth plus other (NISP ≥1, mandibular canine plus three P. setiger (= Rogersville, TN 71 1 4 1 1 1 NISP ≤ 10) appendicular elements Hay 1920:84-85 compressus) misc. elements (NISP ≥1, NISP astragalus and second Roubidoux Cave, MO 41 1 2 1 1 1 ≤ 5) phalange Hawksley 1986:42 isolated tooth, teeth, or tooth Rush Creek, OK 24 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) mandible with p2-4 Czaplewski 2012:113 Santa Fe (IA), FL - - - 8 1 1 - - Bullen et al. 1970; Dunbar 1991:Table 1 no mention of peccary

Sebastian Canal, FL 85 ? ? 1 1 1 unspecified unspecified Webb 1974:Table 2.1

94

APPENDIX A, CONTINUED

Locality MAP MNI NISP Taxon FAUNMAP NEOTOMA Find Type Specimens Primary References Comments isolated tooth, teeth, or tooth Selby, CO 13 1 >1 1 1 1 row (NISP ≥ 1, NISP ≤ 10) isolated teeth Graham 1981:44 isolated tooth, teeth, or tooth Simpson 1929:563, 591; Webb 1974:Table Seminole Field, FL 83 ? ? 1 1 1 row (NISP ≥ 1, NISP ≤ 10) isolated teeth 2.1 Sheep Camp Shelter, NM 6 ? ? 2 - - unspecified unspecified Gillespie 1985:22 NISP and MNI combined: CMNH sample (NISP = 1,685, MNI = 36) + KSU sample (NISP = 534, Sheriden Cave, OH 53 41 2,219 1 1 1 NISP ≥ 1,000 multiple Gobetz 1988:23, 26 MNI = 5)

Smith Falls, NE 11 1 ? 2 2 - unspecified unspecified Voorhies and Corner 1985:Table 1 isolated tooth, teeth, or tooth Hay 1917:48-50, 1923:222; Webb Vero, FL 86 1 ? 1 1 1 row (NISP ≥ 1, NISP ≤ 10) teeth 1974:Table 2.1; Weigel 1962:41

Waccasassa River (IIB and 9 4

III), FL 80 ? ? 1 1 1 unspecified unspecified Webb 1974:Table 2.1 Welsh Cave, KY 60 31 1,183 1 1 1 NISP ≥ 1,000 multiple Guilday et al. 1971:Table 1 Yarbrough Cave, GA 74 ? ? 1 1 - unspecified unspecified Martin 2001:Table 1 isolated tooth, teeth, or tooth Yukon, OK 23 1 1 1 - - row (NISP ≥ 1, NISP ≤ 10) mandible with p3-m3 Czaplewski 2012:114 misc. elements (NISP ≥1, NISP Zesch Cave, TX 26 1 1 2 2 3 ≤ 5) sacrum Lundelius 1967:Table 1; Sagebiel 2010:169 Zoo Cave, MO 44 81 2,456 1 1 1 NISP ≥ 1,000 multiple Hood and Hawksley 1975:Table 2 1Codes for Taxon, FAUNMAP, and NEOTOMA columns: 1) P. compressus; 2) Platygonus cf. compressus; 3) Platygonus sp.; 4) Platygonus cf. cumberlandensis; 5) nasutus; 6) Platygonus af. compressus); 7) P. vetus; 8) unable to verify. 95

APPENDIX B. SCAN SITE MNES AND VOLUME DENSITIES

Element Scan Site MNE Volume Density Humerus 1 34 0.21 Humerus 2 42 0.22 Humerus 3 83 0.43 Humerus 4 81 0.41 Humerus 5 62 0.39 Radius 1 57 0.45 Radius 2 52 0.54 Radius 3 32 0.56 Radius 4 37 0.42 Radius 5 26 0.4 Ulna 6 16 0.33 Ulna 7 55 0.43 Femur 1 47 0.34 Femur 2 20 0.4 Femur 3 39 0.39 Femur 4 73 0.51 Femur 5 48 0.41 Femur 6 33 0.23 Calcaneus 1 29 0.42 Calcaneus 2 68 0.64 Calcaneus 3 73 0.60 Calcaneus 4 43 0.56 Astragalus 1 68 0.35 Astragalus 2 68 0.47 Tibia 1 29 0.3 Tibia 2 49 0.29 Tibia 3 52 0.5 Tibia 4 60 0.39 Tibia 5 40 0.48 Volume density values and scan sites from Ioannidou (2003). Scan sites 6 and 7 for ulna correspond to Ioannidou (2003) scan sites 1 and 2. 96

APPENDIX C. RAW STABLE ISOTOPE DATA

δ13C and δ18O from Mixed Enamel and Dentin Samples Specimen Distance From δ13C ± 0.09‰ δ18O ± 0.16‰ ERJ (mm) 67-300-14 87-85 -12.00513 -3.31264 67-300-14 82-80 -14.19287 -4.03240 67-300-14 78-76 -13.62105 -4.98954 67-300-14 74-72 -13.15530 -5.00707 67-300-14 70-68 -13.53591 -4.79551 67-300-14 65-63 -14.03564 -4.61870 67-300-14 61-59 -13.42211 -5.15006 67-300-14 57-55 -15.21486 -6.97794 67-300-14 53-51 -13.07437 -3.41032 67-300-14 49-47 -12.70472 -3.10764 67-300-14 45-43 -12.41268 -3.20686 67-300-14 40-38 -14.38589 -6.11345 67-300-14 37-35 -12.99093 -4.35558 67-300-14 33-31 -12.51522 -3.76046 67-300-14 29-27 -13.07437 -4.02871 67-300-14 24-22 -13.16616 -5.38827 Specimen Distance From δ13C ± 0.09‰ δ18O ± 0.16‰ ERJ (mm) 65-158-18 *92-90 -14.35130 -1.25926 65-158-18 88-86 -13.79527 -3.16350 65-158-18 84-82 -14.02981 -2.99469 65-158-18 81-79 -13.72098 -3.55997 65-158-18 77-75 -14.02971 -3.70716 65-158-18 72-70 -15.26311 -3.07064 65-158-18 68-66 -14.59981 -3.35856 65-158-18 63-61 -13.36249 -3.28937 65-158-18 58-56 -12.22630 -1.24604 65-158-18 54-52 -12.39459 -1.41075 65-158-18 49-47 -12.55755 -2.46579 65-158-18 45-43 -12.51321 -2.94262 65-158-18 41-39 -13.09961 -3.06849 65-158-18 37-35 -13.85780 -3.14065 65-158-18 33-31 -13.38521 -2.31685 65-158-18 29-27 -13.18837 -3.79993 65-158-18 25-23 -13.14625 -2.97818 *Sample was below ideal detection window and had higher possible errors.

97

APPENDIX C, CONTINUED

δ13C and δ18O from Dentin Only Samples Specimen Distance From δ13C ± 0.07‰ δ18O ± 0.12‰ ERJ (mm) 67-300-14 87-85 -11.78660 -4.30215 67-300-14 82-80 -13.31958 -4.51512 67-300-14 78-76 -12.91470 -3.85574 67-300-14 74-72 -11.34903 -2.95119 67-300-14 70-68 -11.61038 -2.73198 67-300-14 65-63 -11.24942 -2.67663 67-300-14 61-59 -11.94713 -4.28958 67-300-14 57-55 -11.49731 -2.65171 67-300-14 53-51 -11.49691 -3.00425 67-300-14 49-47 -11.39761 -3.00560 67-300-14 45-43 -11.19881 -2.83644 67-300-14 40-38 -11.69125 -2.49284 67-300-14 37-35 -11.54823 -3.51743 67-300-14 33-31 -10.89181 -2.63146 67-300-14 29-27 -11.06105 -2.56054 67-300-14 24-22 - - Specimen Distance From δ13C ± 0.09‰ δ18O ± 0.16‰ ERJ (mm) 65-158-18 *92-90 -13.75335 -1.25926 65-158-18 88-86 -13.84001 -3.16350 65-158-18 84-82 -13.64186 -2.99469 65-158-18 81-79 -13.84986 -3.55997 65-158-18 **77-75 - - 65-158-18 72-70 -15.30684 -3.70716 65-158-18 68-66 -14.36347 -3.07064 65-158-18 63-61 -13.26417 -3.35856 65-158-18 58-56 -13.07890 -3.28937 65-158-18 54-52 -12.72925 -1.24604 65-158-18 49-47 -12.58037 -1.41075 65-158-18 45-43 -12.88045 -2.46579 65-158-18 41-39 -13.77557 -2.94262 65-158-18 37-35 -14.16975 -3.06849 65-158-18 33-31 -14.41474 -3.14065 65-158-18 29-27 -14.06158 -2.31685 65-158-18 25-23 -13.08503 -3.79993 *Sample was below ideal detection window and had higher possible errors. **Sample material lost during washing process.