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Home , Albuquerque Basin, Amphicyon, Arctoidea, Barbourofelis, Bassariscus, Bear dog

UNDERSTANDING CARNIVORAN ECOMORPHOLOGY THROUGH DEEP TIME, WITH A CASE STUDY DURING THE CAT-GAP OF FLORIDA

SHARON ELIZABETH HOLTE

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2018

To Dr. Larry, thank you

ACKNOWLEDGMENTS

I would like to thank my family for encouraging me to pursue my interests. They have always believed in me and never doubted that I would reach my goals. I am eternally grateful to my mentors, Dr. Jim Mead and the late Dr. Larry Agenbroad, who have shaped me as a paleontologist and have provided me to the strength and knowledge to continue to grow as a scientist. I would like to thank my colleagues from the Florida Museum of Natural History who provided insight and open discussion on my research. In particular, I would like to thank Dr. Aldo Rincon for his help in researching procyonids. I am so grateful to Dr. Anne-Claire Fabre; without her understanding of R and knowledge of 3D morphometrics this project would have been an immense struggle. I would also to thank Rachel Short for the late-night work sessions and discussions. I am extremely grateful to my advisor Dr. David Steadman for his comments, feedback, and guidance through my time here at the University of Florida. I also thank my committee, Dr. Bruce MacFadden, Dr. Jon Bloch, Dr. Elizabeth Screaton, for their feedback and encouragement.

I am grateful to the geosciences department at East Tennessee State University, the American Museum of Natural History, and the Museum of Comparative Zoology at

Harvard for the loans of specimens. This research is based upon work supported by the

National Science Foundation Grant No. 1701587. Other financial support for this research came from the Grinter Fellowship and CLAS Graduate Travel Award from the

University of Florida College of Liberal Arts and Sciences as well as the Miss Lucy

Dickinson Fellowship and Gary S. Morgan Award from the Florida Museum of Natural

History and the Mitchell Hope Scholarship from the Southwest Florida Fossil Society.

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 7

LIST OF FIGURES ...... 14

LIST OF ABBREVIATIONS ...... 17

ABSTRACT ...... 18

CHAPTER

1 INTRODUCTION ...... 20

Functional Ecomorphology ...... 21 Thomas Farm Fossil Locality ...... 23 Cat-Gap ...... 30 3D Morphometric Analyses ...... 31

2 NEW PROCYONIDS FROM THE EARLY HEMINGFORDIAN (EARLY MIOCENE) THOMAS FARM FOSSIL SITE, FLORIDA ...... 37

Introductory Remarks...... 37 Descriptions and Comparative Morphology ...... 40 Systematic Paleontology: Thomas Farm potosine ...... 40 Systematic Paleontology: Thomas Farm cf. Probassariscus ...... 41 Phylogenetic Analysis ...... 47 Discussion ...... 50

3 3D GEOMETRIC MORPHOMETRICS ON COMPLETE VS. PARTIAL FORELIMB ELEMENTS OF CARNIVORANS ...... 63

Introductory Remarks...... 63 Materials and Methods...... 64 Materials ...... 64 Methods...... 64 Phylogeny ...... 68 Phylogenetic Signal ...... 69 MANOVA and Phylogenetic MANOVA ...... 69 Results ...... 69 Geometric Morphometrics ...... 69 Family-level PCAs ...... 69 Locomotion PCAs ...... 73 Hunting strategy PCAs ...... 76

Phylogenetic Signal Results ...... 78 MANOVA and Phylogenetic MANOVA ...... 79 Linear Discriminant Analyses ...... 79 k-NN Analyses ...... 81 Discussion ...... 82

4 ECOMORPHOLOGY OF CAT-GAP CANIDS FROM THOMAS FARM FOSSIL SITE, (EARLY MIOCENE) FLORIDA...... 109

Introductory Remarks...... 109 Materials and Methods...... 110 Materials ...... 110 Thomas Farm Canids ...... 111 Geometric Morphometrics ...... 111 k-NN Analysis and Predictions ...... 112 Results ...... 113 Family-Level PCAs ...... 113 Locomotion PCAs ...... 117 Hunting Strategy PCAs...... 121 k-NN and Predictions ...... 125 Discussion ...... 126

5 CONCLUSIONS ...... 145

APPENDIX

A UPPER P4 MEASUREMENTS ...... 148

B LOWER m1 MEASUREMENTS ...... 155

C LIST OF CHARACTERS EDITED FROM BASKIN, 2004 ...... 162

D EXTANT TAXA SCANNED ...... 165

E SUPPLEMENTAL INFORMATION FOR CHAPTER 3...... 171

F SUPPLEMENTAL INFORMATION FOR CHAPTER 4...... 254

LIST OF TABLES

Table page

1-1 Thomas Farm (TF) Carnivorans ...... 36

3-1 Samuels et al., 2013 locomotor classification...... 106

3-2 Van Valkenburgh, 1985 hunting strategy classification...... 106

3-3 LDA Rate of Reclassification ...... 107

3-4 k-NN Rate of Reclassification...... 108

E-1 Definition of the landmarks of the humerus used in the analysis ...... 171

E-2 Definition of the landmarks of the radius used in the analysis ...... 172

E-3 Definition of the landmarks of the ulna used in the analysis ...... 173

E-4 MAOVAs and Phylogenetic MAOVAs ...... 174

E-5 Calculation of Phylogenetic Signal...... 176

E-6 Complete Humerus Family-level LDA ...... 177

E-7 Complete Humerus Locomotion LDA ...... 178

E-8 Complete Humerus Hunting Strategy LDA ...... 179

E-9 Complete Humerus Activity Pattern LDA ...... 180

E-10 Complete Humerus Diet LDA ...... 180

E-11 Complete Humerus Habitat LDA ...... 181

E-12 Complete Humerus Social Behavior LDA ...... 181

E-13 Proximal Humerus Family-level LDA ...... 182

E-14 Proximal Humerus Locomotion LDA ...... 183

E-15 Proximal Humerus Hunting Strategy LDA ...... 184

E-16 Proximal Humerus Activity Pattern LDA ...... 185

E-17 Proximal Humerus Diet LDA ...... 185

E-18 Proximal Humerus Habitat LDA ...... 186

E-19 Proximal Humerus Social Behavior LDA ...... 186

E-20 Distal Humerus Family-level LDA ...... 187

E-21 Distal Humerus Locomotion LDA ...... 188

E-22 Distal Humerus Hunting Strategy LDA ...... 189

E-23 Distal Humerus Activity Pattern LDA ...... 190

E-24 Distal Humerus Diet LDA ...... 190

E-25 Distal Humerus Habitat LDA ...... 191

E-26 Distal Humerus Social Behavior LDA ...... 191

E-27 Complete Radius Family-level LDA ...... 192

E-28 Complete Radius Locomotion LDA ...... 193

E-29 Complete Radius Hunting Strategy LDA ...... 194

E-30 Complete Radius Activity Pattern LDA ...... 195

E-31 Complete Radius Diet LDA ...... 195

E-32 Complete Radius Habitat LDA ...... 196

E-33 Complete Radius Social Behavior LDA ...... 196

E-34 Proximal Radius Family-level LDA ...... 197

E-35 Proximal Radius Locomotion LDA ...... 198

E-36 Proximal Radius Hunting Strategy LDA ...... 199

E-37 Proximal Radius Activity Pattern LDA ...... 200

E-38 Proximal Radius Diet LDA ...... 200

E-39 Proximal Radius Habitat LDA ...... 201

E-40 Proximal Radius Social Behavior LDA ...... 201

E-41 Distal Radius Family-level LDA ...... 202

E-42 Distal Radius Locomotion LDA ...... 203

E-43 Distal Radius Hunting Strategy LDA ...... 204

E-44 Distal Radius Activity Pattern LDA ...... 205

E-45 Distal Radius Diet LDA ...... 205

E-46 Distal Radius Habitat LDA ...... 206

E-47 Distal Radius Social Behavior LDA ...... 206

E-48 Complete Ulna Family-level LDA ...... 207

E-49 Complete Ulna Locomotion LDA ...... 208

E-50 Complete Ulna Hunting Strategy LDA ...... 209

E-51 Complete Ulna Activity Pattern LDA ...... 210

E-52 Complete Ulna Diet LDA ...... 210

E-53 Complete Ulna Habitat LDA...... 211

E-54 Complete Ulna Social Behavior LDA ...... 211

E-55 Proximal Ulna Family-level LDA ...... 212

E-56 Proximal Ulna Locomotion LDA ...... 213

E-57 Proximal Ulna Hunting Strategy LDA ...... 214

E-58 Distal Humerus Activity Pattern LDA ...... 215

E-59 Proximal Ulna Diet LDA ...... 215

E-60 Proximal Ulna Habitat LDA ...... 216

E-61 Proximal Ulna Social Behavior LDA ...... 216

E-62 K values for k-NN analysis...... 217

E-63 Complete Humerus Family-level k-NN ...... 218

E-64 Complete Humerus Locomotion k-NN ...... 218

E-65 Complete Humerus Hunting Strategy k-NN ...... 219

E-66 Complete Humerus Activity Pattern k-NN ...... 219

E-67 Complete Humerus Diet k-NN ...... 219

E-68 Complete Humerus Habitat k-NN ...... 220

E-69 Complete Humerus Social Behavior k-NN ...... 220

E-70 Proximal Humerus Family-level k-NN ...... 221

E-71 Proximal Humerus Locomotion k-NN ...... 221

E-72 Proximal Humerus Hunting Strategy k-NN ...... 222

E-73 Proximal Humerus Activity Pattern k-NN ...... 222

E-74 Proximal Humerus Diet k-NN ...... 222

E-75 Proximal Humerus Habitat k-NN ...... 223

E-76 Proximal Humerus Social Behavior k-NN ...... 223

E-77 Distal Humerus Family-level k-NN ...... 224

E-78 Distal Humerus Locomotion k-NN ...... 224

E-79 Distal Humerus Hunting Strategy k-NN ...... 225

E-80 Proximal Humerus Activity Pattern k-NN ...... 225

E-81 Distal Humerus Diet k-NN...... 225

E-82 Distal Humerus Habitat k-NN ...... 226

E-83 Distal Humerus Social Behavior k-NN ...... 226

E-84 Complete Radius Family-level k-NN ...... 227

E-85 Complete Radius Locomotion k-NN ...... 227

E-86 Complete Radius Hunting Strategy k-NN ...... 228

E-87 Complete Radius Activity Pattern k-NN ...... 228

E-88 Complete Radius Diet k-NN ...... 228

E-89 Complete Radius Habitat k-NN...... 229

E-90 Complete Radius Social Behavior k-NN ...... 229

E-91 Proximal Radius Family-level k-NN ...... 230

E-92 Proximal Radius Locomotion k-NN ...... 230

E-93 Proximal Radius Hunting Strategy k-NN ...... 231

E-94 Proximal Radius Activity Pattern k-NN ...... 231

E-95 Proximal Radius Diet k-NN ...... 231

E-96 Proximal Radius Habitat k-NN ...... 232

E-97 Proximal Radius Social Behavior k-NN...... 232

E-98 Distal Radius Family-level k-NN...... 233

E-99 Distal Radius Locomotion k-NN ...... 233

E-100 Distal Radius Hunting Strategy k-NN ...... 234

E-101 Proximal Radius Activity Pattern k-NN ...... 234

E-102 Distal Radius Diet k-NN ...... 234

E-103 Distal Radius Habitat k-NN ...... 235

E-104 Distal Radius Social Behavior k-NN ...... 235

E-105 Complete Ulna Family-level k-NN ...... 236

E-106 Complete Ulna Locomotion k-NN ...... 236

E-107 Complete Ulna Hunting Strategy k-NN ...... 237

E-108 Complete Ulna Activity Pattern k-NN ...... 237

E-109 Complete Ulna Diet k-NN ...... 237

E-110 Complete Ulna Habitat k-NN ...... 238

E-111 Complete Ulna Social Behavior k-NN ...... 238

E-112 Proximal Ulna Family-level k-NN ...... 239

E-113 Proximal Ulna Locomotion k-NN ...... 239

E-114 Proximal Ulna Hunting Strategy k-NN ...... 240

E-115 Proximal Ulna Activity Pattern k-NN ...... 240

E-116 Proximal Ulna Diet k-NN ...... 240

E-117 Proximal Ulna Habitat k-NN ...... 241

E-118 Proximal Ulna Social Behavior k-NN ...... 241

F-1 Chapter 4 groupings ...... 254

F-2 Best k-values for analyses ...... 254

F-3 Humerus “All” Family-level k-NN ...... 255

F-4 Humerus TFcafemusthypro Family-level k-NN ...... 255

F-5 Humerus TFcafemusthy Family-level k-NN ...... 255

F-6 Humerus TFcafe Family-level k-NN ...... 256

F-7 Radius “All” Family-level k-NN ...... 256

F-8 Radius TFcafemusthy Family-level k-NN ...... 256

F-9 Radius TFcafe Family-level k-NN ...... 256

F-10 Ulna “All” Family-level k-NN...... 257

F-11 Ulna TFcafemusthy Family-level k-NN ...... 257

F-12 Ulna TFcafe Family-level k-NN ...... 257

F-13 Humerus “All” Hunting Strategy k-NN ...... 258

F-14 Humerus TFcafemusthypro Hunting Stratgey k-NN ...... 258

F-15 Humerus TFcafemusthy Hunting Strategy k-NN ...... 258

F-16 Humerus TFcafe Hunting Strategy k-NN ...... 259

F-17 Radius “All” Hunting Strategy k-NN ...... 259

F-18 Radius TFcafemusthy Hunting Strategy k-NN ...... 259

F-19 Radius TFcafe Hunting Strategy k-NN ...... 259

F-20 Ulna “All” Hunting Strategy k-NN ...... 260

F-21 Ulna TFcafemusthy Hunting Strategy k-NN ...... 260

F-22 Ulna TFcafe Hunting Strategy k-NN ...... 260

F-23 Humerus “All” Locomotion k-NN ...... 261

F-24 Humerus TFcafemusthypro Locomotion k-NN...... 261

F-25 Humerus TFcafemusthy Locomotion k-NN ...... 261

F-26 Humerus TFcafe Locomotion k-NN ...... 262

F-27 Radius “All” Locomotion k-NN ...... 262

F-28 Radius TFcafemusthy Locomotion k-NN ...... 262

F-29 Radius TFcafe Locomotion k-NN ...... 263

F-30 Ulna “All” Locomotion k-NN ...... 263

F-31 Ulna TFcafemusthy Locomotion k-NN ...... 263

F-32 Ulna TFcafe Locomotion k-NN ...... 264

LIST OF FIGURES

Figure page

1-1 Phylogeny of the Order Carnivora ...... 33

1-2 Timeline depicting arrival and disappearance of nimravids, barbourofelids, and felids in North America ...... 34

1-3 A comparison of the nimravid Dinictis felina (left), the barbourofelid Barbourofelis loveorum (middle), to the true felid Nimravides catocopis (right) .. 35

2-1 Distribution of fossil procyonids from the Oligocene through the Miocene ...... 52

2-2 TF potosine, fossil, lower m1: UF 406549. cf. Probassariscus, fossil, lower m1s:...... 53

2-3 Lower m1 box plots of the Natural log of APL...... 54

2-4 cf. Probassariscus, fossil, upper P4s and upper M1...... 55

2-5 cf. Probassariscus, fossil, UF 276543, left dentary ...... 56

2-6 cf. Probassariscus, fossil, UF 181215, right partial dentary ...... 57

2-7 Bassariscus astutus, modern, UF 11933 ...... 58

2-8 Upper P4 box plots of the Natural log of APL ...... 59

2-9 Bassariscus astutus, modern, UF11933, left dentary ...... 60

2-10 Possible phylogenetic relationships of TF cf. Probassariscus, fossil taxa ...... 61

2-11 Character matrix from Baskin, 2004 with Thomas Farm cf. Probassariscus scored...... 62

3-1 Landmark placement of the humerus ...... 85

3-2 Landmark placement of the radius ...... 86

3-3 Landmark placement of the ulna ...... 87

3-4 Landmark placement of the proximal humerus ...... 88

3-5 Landmark placement of the distal humerus ...... 89

3-6 Landmark placement of the proximal radius ...... 90

3-7 Landmark placement of the distal radius ...... 91

3-8 Landmark placement of the proximal ulna ...... 92

3-9 Character matrix for individual extant species...... 93

3-1 Carnivoran phylogeny of the extant 81 species included in dataset, modified from Nyakatura and Bininda-Emonds (2012)...... 96

3-11 Family phylomorphospaces for the humerus ...... 97

3-12 Family phylomorphospaces for the radius ...... 98

3-13 Family phylomorphospaces for the ulna ...... 99

3-14 Locomotion PCAs for the humerus ...... 100

3-15 Locomotion PCAs for the radius ...... 101

3-16 Locomotion PCAs for the ulna ...... 102

3-17 Hunting strategy PCAs for the humerus ...... 103

3-18 Hunting strategy PCAs for the radius ...... 104

3-19 Hunting strategy PCAs for the ulna ...... 105

4-1 Morphology of the distal humerus in a typical extant canid (Canis latrans 13412), extant felid (Puma concolor UF 25908), and an extinct TF canid (UF 1485)...... 128

4-2 Morphology of the radius in a typical extant canid (Canis latrans, UF 13412), extant felid (Puma concolor, UF 25908), and an extinct TF canid (UF 60541). 129

4-3 Morphology of the proximal ulna in a typical extant canid (Canis latrans, UF 13412), extant felid (Puma concolor, UF 25908), and an extinct TF canid (UF 267416)...... 130

4-4 Family PCAs for the distal humerus ...... 131

4-5 Family PCAs for the radius ...... 132

4-6 Family PCAs for the proximal ulna ...... 133

4-7 Locomotion PCAs for the distal humerus ...... 134

4-8 Locomotion PCAs for the radius...... 135

4-9 Locomotion PCAs for the proximal ulna ...... 136

4-10 Hunting strategy PCAs for the distal humerus ...... 137

4-11 Hunting strategy PCAs for the radius ...... 138

4-12 Family PCAs for the proximal ulna ...... 139

4-13 Thomas Farm humeri predictions...... 140

4-14 Thomas Farm radii predictions...... 141

4-15 Thomas Farm ulnae predictions...... 142

4-17 Summary figure of the most common characteristic of TF canids...... 144

E-1 Activity pattern PCAs for the humerus ...... 242

E-2 Activity pattern PCAs for the radius ...... 243

E-3 Activity pattern PCAs for the ulna ...... 244

E-4 Diet PCAs for the humerus ...... 245

E-5 Diet PCAs for the radius ...... 246

E-6 Diet PCAs for the ulna ...... 247

E-7 Social behavior PCAs for the humerus ...... 248

E-8 Social behavior PCAs for the radius...... 249

E-9 Social behavior PCAs for the ulna ...... 250

E-10 Habitat PCAs for the humerus ...... 251

E-11 Habitat PCAs for the radius ...... 252

E-12 Habitat PCAs for the ulna ...... 253

LIST OF ABBREVIATIONS

3D 3 dimensional

AMNH American Museum of Natural History

APL Anterior-posterior length

CI Consistency index

CM Carnegie Museum

CSSL Curve sliding semi-landmarks

FAM American Museum of Natural History specimen

HI Homoplasy index k-NN k-nearest neighbors analysis

KUM Kansas University Natural History Museum

LDA Linear discriminant analysis

MCZ Museum of Comparative Zoology, Harvard University

MPT Most parsimonious tree

NA North America

NALMA North American Land Mammal Age

PC Principal component

PCA Principal component analysis

RI Retention index

SSSL Surface sliding semi-landmarks

TF Thomas Farm fossil site

TW Transverse width

UF University of Florida

Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

UNDERSTANDING CARNIVORAN ECOMORPHOLOGY THROUGH DEEP TIME, WITH A CASE STUDY DURING THE CAT-GAP OF FLORIDA

Sharon Elizabeth Holte

May 2018

Chair: David Steadman Major: Zoology

The early Miocene is an important time in carnivoran evolution. Felids had not dispersed into North America where canids and procyonids already existed. A new taxa of the bassariscine procyonid cf. Probassariscus from the Thomas Farm fossil locality helps to bridge the phylogenetic gap bewteen Bassariscus and Probassariscus. A second procyonid (from the subfamily Potosininae) was also present at Thomas Farm; these two taxa represent the first two procyonids described at this fossil site.

Interpreting the behavior of extinct animals can be difficult, especially when examining partial fossil specimens. This research demonstrates the feasibility of incorporating incomplete specimens into datasets using 3D (3 dimensional) morphometrics. A 3D database of 81 species of modern carnivorans was compiled for comparison with early Miocene canids. These analyses allow for a better comparison of functional ecomorphology among modern taxa and provides a backdrop with which to compare extinct taxa. K-nearest neighbors (k-NN) analyses and principal component analyses can be used to interpret behavior and morphological relationships fairly accurately. Such statistical tools show that focusing research on the weight bearing

epiphyses of forelimbs may provide a higher rate of accurate classification than complete elements.

Early Hemingfordian (early Miocene) canids have been described as having more mobility in their forelimbs than modern canids. This research examines how canids from the early Miocene Thomas Farm locality relate in their forelimb movement to modern carnivorans, in order to better understand their paleoecomorphology. The function of the forelimb provides insight on how these canids may have adapted to open felid niches during the Cat-Gap, a nearly 5-million-year time span (~21-16 Ma) in which there were no felids in North America. 3D morphometric analyses and k-NN predictions have suggested that these fossil canids were most likely scansorial and ambush hunted their prey similar to extant felids. These results indicate that the locomotor and hunting strategy behavior of canids is different in the past than today.

CHAPTER 1 INTRODUCTION

Research Objectives

This dissertation seeks to better understand carnivorans from the Thomas Farm fossil locality (Gilchrist County, Florida) during the early Hemingfordian North American

Land Mammal Age (NALMA).

The research will result in three papers for publication. The first manuscript

(Chapter 2) reports the morphology and systematics of procyonid fossils from Thomas

Farm (TF). This paper is written for submission to the Bulletin of the Florida Museum of

Natural History with collaborator Aldo Rincón. Rincón created the map of Figure 2-1, provided support for the methodology, assisted in compiling and describing morphological characters, and edited the document. The other two research papers are based on the ecomorphology of carnivorans. The first (Chapter 3) examines the usefulness of 3D morphometric software on partial/incomplete specimens of the forelimb for determining ecomorphology. The second paper (Chapter 4) compares the ecomorphology of canids at Thomas Farm to that of extant carnivorans including musteloids, canids, and felids. This issue is important because TF lies within the “Cat-

Gap,” an interval of time (~21 - 16 Ma) when no felid fossils are known in North

America.

The aim of my research is to answer the following questions:

• What characters define the forelimb niche of felids, musteloids and canids? • How can these defining characters be refined with 3D morphometric software using a database built of modern taxa? • How complete do forelimb elements need to be in order to determine phylogenetic and/or functional ecomorphology using 3D morphometric software? • How do early Miocene Thomas Farm canids fall into the defined forelimb niches?

• How does this research affect our overall understanding of carnivoran ecomorphology during the Cat-Gap?

Functional Ecomorphology

Carnivorans display many different modes of locomotion such as cursorial

(running), scansorial/arboreal (climbing), fossorial (digging), and natatorial (swimming)

(Van Valkenburgh, 1985; Polly and MacLeod, 2008; Samuels et al, 2013). Some carnivorans employ more than one of these behaviors but typically spend more time doing one than the other. An animal’s locomotory mode is reflected in the morphology of its appendicular skeleton (Van Valkenburgh, 1985; Polly, 2007). The overall shape and orientation of each tubercle or condyle of an element can indicate the animal’s locomotory style as well as its overall range of limb motion (Samuels and Van

Valkenburgh, 2008; Samuels et al, 2013).

Mammals that are adapted for speed are known as cursorial. A cursor’s gait (trot, pace, run, half-bound or gallop) depends on its morphology and size (Taylor, 1989).

Cursorial mammals typically have relatively elongated limbs and distal elements, either an unguligrade or digitigrade stance, and long, narrow scapulae to increase stride length (Polly, 2007). Cursorial carnivorans tend to have a humerus with a relatively large proximally protruding greater tuberosity, a reduced bicipital notch, and a proximally located bicipital tuberosity of the radius (Taylor, 1989). They also are restricted in their ability to supinate the forearm for joint stabilization while running

(Polly, 2007).

The terms scansorial and arboreal are sometimes used interchangeably in the literature. Scansorial refers to taxa that spend some time in the trees whereas arboreal means that an animal spends most of its time in trees. Scansorial mammals tend to

climb trees when they need to escape; arboreal mammals are rarely on the ground and they shelter and forage in the trees (Van Valkenburgh, 1987). According to Taylor

(1989), arboreal animals have three primary strategies to obtain food: 1) they must be fast either in the trees or on the ground; 2) they must be stealthy (e.g., many cats); or 3) they are slow-moving herbivores such as tree sloths and lack the need for quick movements. MacLeod and Rose (1993) demonstrated that the outline of the radial head can help to discriminate scansorial from other locomotor behaviors. Scansorial carnivorans are likely to have these characters: an enlarged groove between the head of the humerus and the greater tuberosity for the biceps tendon, allowing for greater flexibility of the forearm; radial notch of the radius laterally oriented and a prominent bicipital tuberosity; and an olecranon process of the ulna angled slightly anteriorly and not elongate (Taylor, 1989).

Carnivorans that are truly fossorial (spend most of time underground) can be divided into two groups: those that excavate their own burrows and those that modify preexisting ones; semi-fossorial mammals also dig for both shelter and food, although they spend most of their time above ground (Van Valkenburgh, 1987; Taylor, 1989).

Fossorial and semi-fossorial carnivorans are likely to have strong forearm musculature with pronounced areas for muscle attachment resulting in an elongated olecranon process of the ulna and relatively short and robust diaphyses of the ulna and radius

(Taylor, 1989). Fossorial (and natatorial) carnivorans also tend to have a triangular- shaped scapula (Polly, 2007).

Natatorial (swimming) carnivorans are adapted to spending a large amount of time in the water and only some time on land (Taylor, 1989). Forelimbs of natatorial

carnivorans tend to be similar to those that are semifossorial, such as a shortened humerus and an elongated olecranon process of the ulna, because aquatic carnivorans utilize their forelimbs to help them swim (Polly, 2007).

Thomas Farm Fossil Locality

The Thomas Farm Fossil Site is located in Gilchrist County, Florida about 70 km northwest of the University of Florida in Gainesville. The site was discovered in 1931 by

Clarence Simpson of the Florida Geological Survey. The site was deeded to the

University of Florida in the late 1930s. Thomas Farm is a sinkhole deposit with an associated cave system (Pratt, 1989). During the early Miocene, the sinkhole acted as a

“natural trap” into which animals fell or carcasses washed in. The site dates biochronologically to the early Hemingfordian North American Land Mammal Age

(NALMA) of the early Miocene (~18 Ma) based on the immigration of the amphicyonid genus Amphicyon, and the presence of the ursid Phoberocyon, the mustelid Leptarctus, the rhinoceros Floridaceras, and the canid Metatomarctus (Tedford et al., 2004).

Numerous localities in the western United States have been suggested to be of comparable biochronological age to Thomas Farm including the early Hemingfordian

Runningwater Formation in Nebraska (Tedford and Frailey, 1976; Woodburne and

Robinson, 1977; Wang, 2003; Morgan and Lucas, 2009). There is also a correlation between the carnivoran fossils at Thomas Farm and those from the early Burdigalian age (~ 20.44 Ma) of Eurasia (Tedford and Frailey, 1976; Hunt, 1998b). For example, a group of mustelids called neomustelids originated in Europe in the early Miocene and immigrated to North America resulting in a turnover in the types of mustelids living in

North America during the Hemingfordian (Baskin, 1998b).

At the end of the early Miocene, climatic conditions were changing, with the ice sheets that developed during the late Oligocene (27 - 26 Ma) beginning to melt. By the middle Miocene (~15 Ma), global ice volume remained low (Zachos et al., 2001). This warming trend led to changes in habitats: the forested areas of Florida that restricted the need for highly cursorial carnivorans shrank as grasslands began to spread (Wang and Tedford, 2008). The largest predators of the early Hemingfordian, the amphicyonids, may have been outcompeted by the more cursorial borophagine canids

(Wang and Tedford, 2008). Rising temperatures peaked in the Barstovian, known as the

Miocene climatic optimum (17 - 15 Ma), and were followed by a gradual cooling (Zachos et al., 2001).

Viranta (2003) examined the geographic and temporal ranges of carnivorans from the middle and late Miocene of North America and confirmed the hypothesis proposed by Janis (1993) that Eurasian faunas had a higher degree of endemism. For example, canids are understood to be primarily endemic to North America until the late

Miocene when they made their way into western Eurasia where hyaenids had been thriving. This suggests that prior to the arrival of canids in Eurasia, the canid niche there was occupied by hyaenids, ursids, and amphicyonids. Amphicyonids (Cynelos,

Amphicyon) and ursids (Phoberocyon (Hemicyon), Ursavus) occurred in both Eurasia and North America during the early and middle Miocene, probably because of multiple dispersal events (Hunt, 1996).

Thomas Farm – Hemingfordian Carnivorans

Carnivorans are distinguished from other mammals by dental characteristics such as modified carnassial teeth in the upper fourth premolars and first lower molars

(Janis et al., 1998). The Order Carnivora includes Nimravidae (extinct cat-like animals),

Barbourofelidae (extinct cat-like animals), Felidae (cats), Viverridae (meerkats and civets), Hyaenidae (hyenas), Herpestidae (mongooses), Eupleridae (fossa), Canidae

(dogs), Amphicyonidae (extinct bear-dogs), Ursidae (bears), Phocidae (true seals),

Otariidae (fur seals and sea lions), Odobenidae (walruses), Ailuridae (red pandas),

Mephitidae (skunks), Procyonidae (raccoons), and Mustelidae (weasels, badgers, etc.;

Figure 1-1).

The currently identified carnivoran fossils at Thomas Farm represent the

Amphicyonidae, Canidae, Ursidae, Mustelidae, and Procyonidae (Table 1-1). No felids, nimravids, or barbourofelids have been found at Thomas Farm. A 5-million-year interval

(known as the Cat-Gap, discussed in detail later) exists in North America between the earliest Miocene, when nimravids become extinct (~21 Ma), until the late early Miocene

(~16 Ma) when felids first appear (Figure 1-2) (Martin, 1998). This time gap would be compromised with a discovery of nimravids or felids at the early Miocene Thomas Farm site.

Temperatures during the early Miocene were milder than today (Wang and

Tedford, 2008). During this time, central peninsular Florida, being above sea level, sustained diverse mammalian faunas (Hine, 2013). The faunas from the early-late

Arikareean through the early Hemingfordian are generally considered the “Cat-Gap” assemblage, which also is known as the Runningwater chronofauna (Hulbert, 2001).

Faunas during this time lacked nimravids, barbourofelids, and felids with amphicyonids being the top predators, whereas the canid, hyaenid, and ursid niches were filled by diverse canids (Janis et al., 1998). By the early-late Hemingfordian, the top predators

were large wolf-like amphicyonids and hemicyonids and a variety of medium to small canids; this is known as the amphicyonid-hemicyonid assemblage (Hulbert, 2001).

Amphicyonidae is a family of extinct mammals known as bear-dogs because they have characters resembling those of both bears and dogs (Tomiya and Tseng, 2016).

These animals ranged in size from small dog-like species to larger hyena-like animals.

The genus Amphicyon first appears in North America during the early Miocene in the

Runningwater Formation of western Nebraska and the Troublesome Formation in north- central Colorado (Hunt, 2003). These first New World amphicyonids may have had a similar ecological role to the modern lion, Panthera leo (Hunt, 2003). Amphicyon and

Cynelos are the two genera of amphicyonids known from Thomas Farm. Amphicyon longiramus was a large amphicyonid that may have acted as sort of a grizzly bear with its plantigrade stance, broad limbs, and ability to crush bone (Hunt, 1998a).

Another amphicyonid predator in Florida during the Hemingfordian was Cynelos caraniavorus, known only from Thomas Farm (Hunt, 1998a). The genus Cynelos first appears in the late Oligocene of Europe at Pech Desse, Quercy, at ~25 Ma (Zhanxiang,

2003). The first record North American record of Cynelos is from the early Miocene upper Harrison Formation, Nebraska, at 19.2 Ma (Hunt, 1998b). Cynelos caraniavorus was a small bear-dog close in size to the Eurasian bear-dog C. schlasseri of the

Burdigalian (Tedford and Frailey, 1976).

Another carnivoran that defines the Hemingfordian NALMA is the hemicyonid

Phoberocyon. Hemicyonidae is a member of the superfamily Ursoidea which also contains the modern family Ursidae. Thomas Farm has the only record of the genus

Phoberocyon in North America, whereas in Eurasia the genus is known from the

Burdigalian through the Tortonian ages (~20.4 - 7.2 Ma) (Tedford and Frailey, 1976;

Hulbert, 2001). Phoberocyon johnhenryi (sometimes referred to as Hemicyon johnhenryi) was digitigrade and most likely long-footed, so it may have been more cursorial than the large amphicyonids (Hunt, 1998b).

Two out of the three known Hemingfordian subfamilies of canids occurred in

Florida, the hesperocyonines and the borophagines. Osbornodon iamonensis, a hesperocyonine distinguished from the borophagines by the lack of a transverse crest on the lower m1 connecting the hypoconid and the entoconid, was thought to have been a relatively good climber and omnivorous (Hulbert, 2001). Osbornodon iamonensis was a coyote-like canid that was similar in size to another canid, Desmocyon matthewi, which is known from the Miller Local Fauna, Florida (early Hemingfordian, ~19 Ma) and the Runningwater Formation (also early Hemingfordian, ~18.8 Ma) (Wang, 2003).

Desmocyon matthewi eventually may be discovered at Thomas Farm since it has been found at fossil sites that are only slightly older. Osbornodon iamonensis has been recorded at both Thomas Farm and in the Runningwater Formation (Wang, 1994).

Euoplocyon spissidens was a small borophagine canid found at Thomas Farm. It was first assigned to the genus Aelurocyon by White (1947), who also established the species name. The species was classified as Enhydrocyon spissidens by Olsen (1958).

Tedford and Frailey (1976), however, referred it to the genus Euoplocyon based on dental characters. The two species of Euoplocyon in North America are E. spissidens from Thomas Farm and E. praedator from the Lower Snake Creek Fauna (Tedford and

Frailey, 1976). The amount of material referred to E. spissidens is limited (Wang et al.,

1999).

Metatomarctus canavus was a small canid also known from the correlative

Runningwater Formation in Nebraska (~18.8 Ma; Wang et al., 1999). Metatomarctus canavus is thought to have been a generalist with the possibility of scavenging as well

(Hulbert, 2001).

Phlaocyon was one of many canids that developed an omnivorous diet and dental specializations similar to those of procyonids (Wang and Tedford, 2008). The genus ranged from the middle Oligocene to the middle Miocene with about nine small to medium sized species described (Wang and Tedford, 2008). Phlaocyon was originally described by Wortman and Matthew (1899), who noted postcranial features similar to those of the canid Hesperocyon, yet its dental affinities more resemble those of the procyonid Bassariscus (Munthe, 1998). Morphology of the auditory region (Hough,

1948) indicated that Phlaocyon was indeed a canid, and Romer (1966) formally described it as such. Phlaocyon was further classified as a borophagine by Tedford

(1978) and placed within the group cynarctines with other hypocarnivorous canids (diet less than 30% meat such as the modern black bear, Ursus americanus; Munthe,1998).

Some of the isolated carnivoran teeth from Thomas Farm were originally referred to

Phlaocyon, a borophagine canid with hypocarnivorous dental characters thought to be convergent with procyonids. The Thomas Farm specimens are now identified as the procyonid cf. Probassariscus (Holte and Rincon, in review).

Mustelidae is a family of generally small carnivorans such as weasels, ferrets, martens, fishers, badgers, and wolverines. They tend to be highly sexually dimorphic in size in both extant and extinct species (Bever and Zakrzewski, 2009). Mustelid paleobiology is poorly understood because of the limited amount of cranial and

postcranial material (Baskin, 1998b). Nevertheless, mustelid identifications are frequently used to distinguish North American Land Mammal Ages based on proposed immigration events in the late Arikareean, Hemingfordian, late Hemphillian, and the

Blancan (Tedford et al., 2004).

Leptarctus ancipidens was a small badger-like carnivore known from Thomas

Farm. This species first was identified based only on dentition (Olsen, 1957a). A lower jaw, discovered around the same time, was assigned to Mephititaxus ancipidens

(Olsen, 1957b). Olsen (1958) reexamined the jaw and synonymized the genus

Mephititaxus with Leptarctus. Baskin (2005) suggested that Leptarctus ancipidens is the oldest known species of Leptarctus and possibly the least derived.

Miomustela(?) sp. is a small weasel discovered at Thomas Farm, originally identified by Olsen (1956). This designation was later discredited by Tedford and Frailey

(1976) who believed that the same material was more comparable to the genus

“Plesictis.” Later, the Thomas Farm material that was described as Miomustela was regarded as cf. Stenogale, a primitative aeluroid (Hunt, 1989; Baskin, 1998b).

Oligobunis floridanus was a large badger-like mustelid described by White

(1941). This designation was questioned by Hochstein (2007), who stated that the badly worn dentition of the Thomas Farm material provided no diagnostic features to attribute it to the Olgiobuninae.

Zodiolestes freundi was a medium sized mustelid (paleo-wolverine) that may have been semifossorial because of cranial similarities with the extant wolverine, Gulo gulo (Hochstein, 2007).

The early Hemingfordian carnivorans from Thomas Farm have been identified mainly from cranial and dental elements. Most postcranial carnivoran elements have not been identified beyond the family level. My research will provide characters to group specimens together of the fossil species already described from Thomas Farm. There is also the potential of identifying undescribed or unrecognized species from Thomas

Farm. For these reasons, the canids from Thomas Farm provide an ideal assemblage to study.

Cat-Gap

Multiple nimravid genera occurred in North America from the middle Eocene (~40

Ma) to the early Miocene (~21 Ma; Anton, 2013). During the Oligocene, nimravids were one of the top predators in North America and were thought to have filled what is viewed today as the felid niche; they were adapted to the edges of forests, parklands, and open spaces and are thought to have been highly skilled climbers (Martin, 1998).

Even though nimravids, barbourofelids, and felids evolved independently of each other, they are very similar in morphology and inferred behavioral characteristics (Anton, 2013;

Figure 1-3). Compared to other carnivorans, the nimravids and barbourofelids had shorter rostra, sectorial dentition, reduced molars and retractable claws, all of which are characters attributed to felids today (Martin, 1998).

The first true felids appear in North America at ~16 Ma. The span of roughly five million years (~21 - 16 Ma) when no felids or felid-like carnivorans (nimravids, barbourofelids) occur in North America is known as the Cat-Gap (Turner, 1997; Figure

1-2 herein).

Currently, the genus Proailurus is described as the first true felid in North

America, although Hunt (1998b) mentions another true felid skull (FAM 61847) from the

Ginn Quarry of Nebraska that he believes to be from the late Hemingfordian NALMA.

This skull was found in a fluvial channel fill possibly from the Box Butte or Sheep Creek formations, both of which overlie the early Hemingfordian Runningwater Formation. By the late middle Miocene (~13 Ma), members of the family Barbourofelidae had made their way into North America (Barbourofelis) but were extinct globally by the early late

Miocene (~11 Ma; Turner, 1997; Anton, 2013). A factor contributing to the extinction of barbourofelids may have been the reduction of forested regions during the late Miocene

(Martin, 1998). The Cat-Gap is based on negative evidence; a felid, nimravid, or barbourofelid may still be found within this time span, although, after nearly two centuries of excavation across North America, numerous early Hemingfordian canids, amphicyonids, and other carnivorans have been discovered but no felids, nimravids, or barbourofelids.

3D Morphometric Analyses

Many different methods have been used over the years to interpret the shapes of bones. One traditional 2D method used in paleontology is to place reference points (X-,

Y- coordinates) known as landmarks on the study object (Reyment, 2010). Difficulties with the landmark method are that equivalent points must be identified by the researcher and be readily repeated on each specimen in the study. Another method of landmark placement is an eigenshape analysis that places the landmarks around the shape of the object at equidistant spaces, traditionally done in 2D. This 2D eigenshape analysis method can be problematic because it may not cover the necessary morphological structures the researcher wants to analyze (Reyment, 2010). This error is easier to avoid by using 3D eigenshape morphometrics, which allows the researcher to

analyze the entire element at one time instead of sectioning the element into pieces and potentially losing some data.

There are various methods of using 3D morphometrics. For example, Polly and

MacLeod (2008) interpreted locomotor function based solely on the calcaneum through an eigenshape analysis by superimposing an equidistant landmark point grid onto their

3D scans. They then defined the shape variation by performing principal component analyses, where each axis depicted the amount of shape variation between the specimens.

Gunz and Mitteroecker (2013) demonstrated that equidistant grid semilandmarks can introduce unnecessary noise such as visualization and statistical artifacts into the data, so they developed a technique where sliding-landmarks can be placed around curves and other features deemed potentially important by the researcher. Since surface semilandmarks are more difficult to place on homologous locations on different specimens, Gunz and Mitteroecker (2013) recommended using a template specimen and then projecting its landmarks using a ThinPlate Spline interpolation function on to the specimens in the dataset. Fabre et al. (2013a, 2013b) followed their techniques to interpret the effect of body mass on the morphology of musteloid forelimbs (humerus, radius, and ulna) as well as to understand the evolution of grasping in musteloids.

Analyses by Cornette et al. (2015) demonstrated that partial specimens can be accurately classified using both 2D traditional landmarks and sliding semilandmarks, although sliding semilandmarks produced better results. Gunz and Mitteroecker (2013) discussed the feasibility of using semilandmarks to estimate the location of the missing

landmarks, although this method is currently viable only in specimens that are bilaterally symmetrical.

Therefore, by understanding the feasibility of 3D morphometric analyses on partial specimens, one can accurately describe the characteristics of fossil species.

That is why Chapter 3 explores this very concept, and Chapter 4 implements it. Chapter

4 defines the ecomorphology of canids during the Cat-Gap from the Thomas Farm fossil site by comparing their ecomorphology to that of modern carnivorans. Partial specimens of the humerus and ulna, as well as the complete radius, are used in the 3D morphometric analysis to interpret the modern family-level guild, locomotor type, and potential hunting behavior of these Thomas Farm carnivorans.

Figure 1-1. Phylogeny of the Order Carnivora (from Anton, 2013). *extinct family

Figure 1-2. Timeline depicting arrival and disappearance of nimravids, barbourofelids, and felids in North America. Geologic timescale modified from Janis et al., 1998.

Figure 1-3. A comparison of the nimravid Dinictis felina (left), the barbourofelid Barbourofelis loveorum (middle), to the true felid Nimravides catocopis (right) not to scale. Sketches modified from (Anton, 2013 pages 92, 107, and 123).

Table 1-1. Thomas Farm (TF) Carnivorans Family Subfamily Genus Species Pertinent Information Ampicyonidae Amphicyoninae Amphicyon longiramus Top predator of the time Cynelos caraniavorus Known only from TF Ursidae/Hemicyonidae Phoberocyon/Hemicyon johnhenryi TF earliest and only NA occurrence Canidae Hesperocyonine Osbornodon iamonensis TF and Runningwater Fm, NE Borophagine Euoplocyon spissidens One of two species within NA Metatomarctus canavus TF and Runningwater Fm, NE Mustelidae Leptarctinae Leptarctus ancipidens Oldest known species of Leptarctus Unnamed Group Miomustela(?) TF Miomustela may be Stenogale Oligobuninae Oligobunis floridanus Not enough material to validate Oligobuninae Zodiolestes freundi May be semifossorial Indeterminate Stenogale(?) Procyonidae Procyoninae cf. Probassariscus Subject of Chapter 2 potosine Subject of Chapter 2

CHAPTER 2 NEW PROCYONIDS FROM THE EARLY HEMINGFORDIAN (EARLY MIOCENE) THOMAS FARM FOSSIL SITE, FLORIDA

Introductory Remarks

The extant Superfamily Musteloidea includes Mustelidae, Ailuridae, and

Procyonidae (Tedford, 1976; Baskin, 1998a). Morphological evidence suggests a divergence of procyonids from mustelids at a minimum of 27.6 Ma, in close agreement with molecular estimates of the divergence ~29 Ma in Europe (Arnason et al., 2007;

Yonezawa et al., 2007). Characters in the auditory region, such as having a well- developed suprameatal fossa outlined by Hough (1944; 1948), define the family

Procyonidae. Early Miocene stem procyonids of Europe (Plesictis of France, Broiliana and Stomeriella of southern Germany) sustain the procyonid auditory arrangement

(Beaumont, 1968). While stem procyonids appeared in the Old World in the Oligocene, they probably did not disperse to North America until the earliest Miocene (Hunt, 1996;

Tedford et al., 2004).

The oldest stem procyonids in North America are Amphictis sp. from the early

Miocene [Hemingfordian North American Land Mammal Age (NALMA)] Anderson

Ranch Formation, Nebraska (Baskin, 2004) and an isolated hypocarnivorous carnivore similar to Bassaricyonoides from the late Arikareean in Panama, Central America (Bloch et al., 2016). Moreover, the beginning of the early Miocene (early Hemingfordian)

Runningwater chronofauna is marked by the occurrence of the procyonids Edaphocyon and Amphictis (?) in Nebraska (Webb and Opdyke, 1995). These endemic New World procyonids had a more inflated auditory bulla as well as a relatively short and narrow

 Submitted to the Bulletin of the Florida Museum of Natural History for publishing.

paraoccipital process (Baskin, 2004). Although the affinities of the oldest New World procyonids are still debated, some of the stem procyonids that arrived in North America during the early Miocene might have been more similar in dental morphology to the hypocarnivorous potosines, the morphologically less-derived tribe of procyonids

(Baskin, 2004). Most fossil and recent procyonines have hypocarnivorous dentitions that reflect either omnivorous or frugivorous diets with the mesocarnivorous Bassariscus and

Probassariscus as the only exception (Baskin, 1982). Extant procyonids are well-known for their scansorial to arboreal behavior and by their small slender bodies with distinctive long tails. Except for Bassariscus, derived procyonids are omnivorous or frugivorous with hypocarnivorous dentition (Baskin, 2003, 2004). The occurrence of the canid

Phlaocyon in the late Arikareean (Buda Local Fauna) in Florida suggests that no procyonids with remarkable hypocarnivory inhabited the eastern coast of the Mississippi

River during the late Paleogene. Furthermore, all late Oligocene carnivorans present in the subtropical fossil record are either borophagine canids; however, by the earliest

Hemingfordian (He1), basal potosines and procyonines inhabited a wide area in North

America ranging from the Gulf Coast region (from Florida to Texas), Panama (Bloch et al., 2016), and the even reaching in Nebraska (Figure 2-1). The occurrence of representatives of the Tribe Procyonini (Edaphocyon) in early Miocene (earliest

Hemingfordian) sequences of southern North America suggest a tropical “ghost” diversification of stem procyonids occurred near the earliest Miocene just after the arrival of stem procyonids to North America. Furthermore, the presence of procyonines and potosines in the earliest Hemingfordian Miller Site (ca. 19 Ma) suggests the

morphological divergence between the tribes potosini and procyonini occurred during the earliest Miocene (Baskin, 2004).

The formal definition of the early Hemingfordian NALMA incorporates the oldest occurrence of the amphicyonid Amphicyon, the canids Phoberocyon and Leptarctus, and the rhinoceros Floridaceras, all of which are present at Thomas Farm in Florida

(Tedford et al., 2004). Moreover, early Hemingfordian diagnostic taxa such as the canid

Metatomarctus, and the floridatraguline camelid Floridatragulus, are also present in association with the rhinoceros Menoceras at this site. Excavations at TF have been ongoing since the early 1930s yet the increase of screen washing in the last few years has uncovered a wealth of small-sized fossil specimens, including many isolated mesocarnivoran teeth (extant mesocarnivorans have a diet of 50-70% meat;

VanValkenburgh, 2007). Some of these teeth were originally referred to Phlaocyon, a borophagine canid with dental morphologies thought to be convergent with procyonids due to their very similar mesocarnivorous dentition (Baskin, 2003). These similarities are so strong that Phlaocyon was originally attributed to the Subfamily Procyoninae

(McGrew, 1938). Further studies on the basicranium placed this taxon the Canidae

(Hough, 1948) on the basis of having a cyonoid septum formed by the posterior inflation of the ventral section of the caudal-entotympanic (Ivanoff, 2001).

Herein, fossils of two early Miocene procyonids from Thomas Farm (TF) are described. These fossil specimens fill a phylogenetic and temporal gap in subtropical latitudes during a time of procyonid diversification leading to the appearance of extant genera during the late Neogene (Simpson, 1945; Fulton and Strobeck, 2006; Baskin,

1989, 2003, 2004).

Comparative extant specimens are from the Florida Museum of Natural History,

University of Florida (UF) Mammalogy collection. Fossil specimens are from the UF

Vertebrate Paleontology collection, Carnegie Museum (CM), Kansas University Natural

History Museum (KUM), and the Museum of Comparative Zoology, Harvard University

(MCZ). Dental terminology follows that proposed by Van Valen (1966) for describing dental material. Measurements (in mm) include anterior-posterior length (APL), transverse width (TW), and were measured at the base of the crown near the enamel dentine junction. All measurements were taken with electronic digital calipers Storm-

3C301 (Appendix A and B).

Descriptions and Comparative Morphology

Systematic Paleontology: Thomas Farm potosine

Order: Carnivora (Bowdich, 1821)

Family: Procyonidae (Gray, 1825)

Tribe: Potosini (Trouessart, 1904)

Referred material: UF 406549, right m1

Horizon and locality: Early Miocene, He1, Thomas Farm fossil site, Gilchrist

County, Florida.

Age: Early Hemingfordian, He1, NALMA (ca. 19 Ma).

Description: The trigonid basin of UF 406549 is missing the summit of the protoconid and the metaconid (Figures 2-2 A, B, C), although there is a relatively distinct anteromedial paraconid. The talonid basin is shallow, representing only the 47% of the APL. The hypoconid is robust and slightly shorter than the paraconid. Anterior to the hypoconid is a highly reduced mesoconid. The hypoconulid and entoconid are

subequal in size and height, with the hypoconid having a slight secondary cusp. A labial cingulid extends from the protoconid to the hypoconid.

Comparison: The trigonid/talonid APL ratio is greater than 1.0, consistent with the

APL ratios of fossil potosines such as B. phyllismillerae from Florida (Baskin, 2004).

Furthermore, the presence of a relatively lingually open trigonid and the location of the paraconid apex along the midline of the tooth further support the identification. The lower m1 is approximately 64% larger than that of the living Bassariscus astutus

(Appendix A). UF 406549 lacks the open trigonid and relatively narrow talonid present in borophagine canids from the same fossil assemblage. Although broken, it has a distinct and relatively close trigonid on the m1. The talonid is basined, differing remarkably from that of borophagine canids and mustelids in having a functional hypoconulid. Although this specimen is relatively large relative to Bassaricyonoides, when comparing the natural log of APL and TW of the talonid it appears within the normal range of size variation for other procyonids from TF included in our sample (Figure 2-3). It differs further from any other previously described carnivore from TF in having a relatively wider talonid, having a remnant of a lingual cingulid on the labial part of protoconid and hypoconid, a nearly closed trigonid with an open trigonid, and larger size (Figure 2-3)..

Systematic Paleontology: Thomas Farm cf. Probassariscus

Order: Carnivora (Bowdich, 1821)

Family: Procyonidae (Gray, 1825)

Tribe: Procyonini (Gray, 1825)

cf. Probassariscus

Referred Material: MCZ 21446, right P4; UF 406542, left P4; UF 1406541 right

P4; UF 259985, left M1; UF 276543, partial left dentary with p2-p3 and a partial p4;

UF181315, right dentary with p4; UF 209500, right m1; UF 225330, left m1; UF 406540, left m1.

Horizon and locality: Early Miocene, Thomas Farm Fossil Site, Gilchrist County,

Florida.

Age: Early Hemingfordian, He1, NALMA (ca. 19 Ma).

Description: The P4 (MCZ 21446) has a short metacone blade and the paracone is enlarged with slight wear on the occlusal surface (Figures 2-4 A, B, C). The protocone extends anteriorly and posteriorly into short pre- and postprotocristae. The paraconule is distinct but not as enlarged as the hypocone. UF 1406541 exhibits considerable wear on the cusps, with a slight longitudinal crack that runs up to the paracone (Figures 2-4

G, H, I). This specimen differs from MCZ 21446 and UF 406542 (Figures 2-4 D, E, F) by having a paraconule larger than the protocone. UF 406542 has a crack that runs through the carnassial notch, while the paraconule is more reduced than in UF

1406541. Among the sample reported from TF, the variation in the position of the paraconule varies from a small but distinct paraconule posterior to the protocone in UF

1406541 to a relatively more reduced cusp on UF 406542 and MCZ 21446. An MNI based on the presence of P4 suggests at least three different individuals present in TF.

The left M1 UF 259985 (Figures 2-4 J, K, L) has a reduced metaconule and lacks the paraconule. It has an enlarged protocone that is subequal in height to the metacone and paracone. A robust postprotocrista connects the highly reduced hypocone with the metacone. The left dentary UF 276543 (Figure 2-5) does not preserve the ascending ramus and the morphology of the anterior part of the p1 alveolus; it retains a complete p2-p4 series along with the mandibular condyle and angular process below the tooth

row, yet there is no m3 alveoli. A complete p4 is preserved in UF 181315, a partial right dentary with the alveoli of the p2 and p3 (Figure 2-6). The m1 (UF 209500) has a highly reduced anteriolabial cingulum, a prominent hypoconid, and a slightly posterointernal placed hypoconulid (Figures 2-2 D, E, F). The mesoconid is in contact with the hypoconulid and it has a barely developed entoconid. The protoconid and metaconid are nearly subequal in height while the paraconid is distinct. UF 225330 has a slightly shorter paraconid relative to UF 209500 and UF 406540 (Figures 2-2 G, H, I). The trigonid is lingually closed (formed by the close proximity of the metaconid and paraconid) in UF 225330 (Figures 2-2 J, K, L) and UF 406540, but not reaching the extent observed in UF 209500.

Comparisons: The TF cf. Probassariscus specimens represent procyonids because they have a closed trigonid on the m1 and lack of m3. Associated isolated M1s lack the distinct metaconule of borophagine canids. The presence of a distinct hypocone on the P4 and a relatively narrow talonid on the m1 are also indicative of New

World procyonines (Baskin, 2004). Furthermore, the well-developed parastyle on the P4 and the more posteriorly situated hypocone on the M1 suggest that the TF specimen belongs to the subfamily Procyoninae. The P4 has a distinct parastyle that projects about 10% of the total length of the tooth with a metacone blade that is shorter, and a protocone that is more posterior than in the stem-procyonid Broiliana (Figure 2-4). The ratio of the width of the trigonid to the talonid of the m1 places the TF specimens in the

Tribe Procyonini, whereas UF 406549 is regarded as a potosine (Baskin, 2004).

In comparison to Phlaocyon, the P4 of the TF procyonid has a more labially oriented paraconule but lacks the distinctive prominent external cingulum of Phlaocyon

(Wang et al., 1999). Compared to the P4 of Broiliana nobilis (Sammlung München 1937

II 13524), the metacone blade of the TF taxon is more reduced, the protocone is located more posterior, and the parastyle is more developed (Dehm, 1950). The metacone blade is relatively shorter than that of Phlaocyon, and the paraconule is located more labially than in Phlaocyon, Osbornodon, or Metatomarctus and lacks the external cingulum of Phlaocyon. Finally, the hypocone is distinct in the TF taxon, Bassariscus astutus, and Phlaocyon willistoni, whereas it is faint in Phlaocyon annectens and absent in Phlaocyon minor; this may be a derived character. In contrast to that of P4 (UF

1406541), the hypocone of Phlaocyon is more lingually placed. Furthermore, the M1 from TF lacks the very prominent paraconule present in Phlaocyon.

Even though the TF M1 has a reduced metaconule and lacks a paraconule, the protocone is enlarged and subequal in height to the metacone and paracone. A robust postprotocrista connects the highly reduced hypocone with the metacone. A small cuspule is present lingual to the protocone in B. astutus yet is not present in the TF specimen (Figures 2-4, 2-7). The parastylar shelf is labially expanded in relation to

Phlaocyon but to the extent in Bassariscus astutus. The M1 from TF also has a more labially expanded parastylar shelf; the m1 has a relatively shorter trigonid which opens lingually. The paraconid is the shortest cuspid while the metaconid and protoconid are relatively sub-equal in height; in Phlaocyon, the protoconid is the tallest cuspid of the trigonid. It differs from Phlaocyon in lacking a distinctive paraconule on the P4, and in having an m1 with more anteroposteriorly elongated talonid with a distinct hypoconulid, a narrower and closed trigonid on the m1, a barely distinct metaconid on the M1, and retaining the m3. The P4 seems to be convergent with that of Phlaocyon. It has a slight

enlargement of the cingulum; but does not have the distinct paraconule present in the

Hemingfordian Phlaocyon minor (Wood and Wood, 1937). Bassariscus has a distinct hypocone as do the TF specimens. The hypocone is placed more lingually in Phlaocyon but is not as distinct as in P4s from TF.

Procyonids differ from mustelids by having a P4 with a hypocone, a posterointernal cingulum, and a protocone that is slightly more posterior (Baskin, 1982).

The M2/m2 are enlarged, while the m2 has an elongated talonid (Baskin, 1989). The TF procyonid differs from the mustelid Zodiolestes from the same fossil assemblage in lacking an m3, and in having an m2 with no external cingulid, a P4 with a discontinuous protoconal shelf, and a distinct hypocone (Figures 2-4, 2-5). It differs from

Potamotherium in having a P4 with a more anteriorly reduced metacone shelf, an M1 with a posterior margin wider than the anterior margin, presence of asymmetric lingual cingula on M1, and more reduced metaconal shelf. The m1 has a relatively shallow talonid basin and a closed trigonid. The left dentary (UF 276543) differs from that of

Miomustela in having a p3 lacking that cusp and premolars (p2-p4) with distinct anteriolingual cingulum.

Compared with the M1 of the potosine Bassaricyonoides from the slightly older earliest Hemingfordian Miller site in Florida, the M1 has a distinct internal cingulum and the m1 lacks a hypocone. The m1 is more reduced anteromedially and the talonid is slightly basined with poorly developed cuspids (Baskin, 2003). In contrast, procyonines have a P4 has a more distinct parastyle, the M1 has a more posteriorly situated hypocone, and similar to extant procyonids, the bulla has a more inflated entotympanic

(Baskin, 2004).

Cf. Probassariscus differs from all other procyonid genera in having a closed talonid on the m1, a distinct hypocone on P4, and retaining the m3. None of the fossil species referred to Bassariscus (e.g., B. antiquus and B. parvus) or Probassariscus matthewi have a closed talonid. Probassariscus matthewi is from the early Barstovian

Virgin Valley Formation in Nevada and Bassariscus parvus is from the early late

Barstovian Stewart Spring Fauna in Nevada. The TF procyonine taxa differs from B. parvus in lacking a fourth cusp on the P4 and having an m1 similar to that of B. astutus.

The variation in the dental dimensions for APL and TW of both the P4 and m1 is within the normal range of modern procyonids (Figure 2-8). The TF specimens differ from Bassariscus astutus in having a relatively wider talon on M1 with a less distinct metaconule and reduced postprotocrista. The M1 metaconule is relatively smaller than that of B. astutus and with a more distinct postprotocrista. Similar to that of B. astutus, the paracone is the largest cusp on the trigon, differing only in having a protocone with equivalent height to that seen on the metacone and the paracone. The parastylar shelf, although labially expanded, does not reach the extent of B. astutus, but it is more similar to that of Probassariscus (Baskin, 2004;Figure 2-6C). The Thomas Farm procyonids are also similar to Probassariscus with a tiny paraconule in the preprotocrista, a morphology not present in Bassariscus. However, TF M1 differs from P. matthewi in having a relatively square shape and a more pronounced metaconule.

Similar to some amphicynodontines, the lower subequal premolars (p2 and p3) have poorly developed cuspids that are also poorly developed in Bassariscus and in UF

276543 therefore suggesting that it might be a primitive character of the procyonidae

(Baskin, 1989; Clark and Guensburg, 1972). The m1 hypoconulid is located lower in the

crown, but more distinct and in a wider talonid, than in B. astutus. This specimen is similar to B. astutus in the relative height of the trigonid cuspids. The protoconid is the largest, the paraconid is the smallest, and the metaconid is intermediate in size. The TF specimens are similar to B. astutus and B. antiquus in lacking anterolabial or external cingulids on m1. The mesoconid of the TF specimen is in contact with the hypoconid and both are located on the lingual side. In comparison to Phlaocyon, the relatively shorter trigonid basin opens lingually with a paraconid being the shortest cuspid while the metaconid and protoconid are relatively subequal in height. The mesoconid is more distinct in the TF specimens than in B. astutus (Figures 2-5, 2-9). The talonid basin is wider and the trigonid basin is less expanded posteriorly than in B. astutus.

Phylogenetic Analysis

To evaluate the phylogenetic relationships of the TF procyonine taxa, a phylogenetic analysis including 18 genera and 40 morphological characters was performed, 26 of which were scored (see Figure 2-11, Appendix C). The TF specimen

UF 406549, referred to as a postone, will not be classified due to the low number of characters to be scored following Baskin (2004) character matrix. Data were scored in

Mesquite version 2.72 and then analyzed using PAUP version 4.0b10 branch and bound algorithm under parsimony; four trees were produced (Figure 2-10, Figure 2-11,

Appendix C) The trees have a length of 104 steps, a consistency index (CI) of 0.596, a retention index (RI) of 0.753, and a homoplasy index (HI) of 0.404.

Old world procyonids exhibit a double-rooted P1/p1 [11(2)] not seen in New

World procyonids excluding Paranausa and Nasua (Baskin, 2004). Western

Hemisphere procyonids have an inflated auditory bulla [3(1/2)], a paraoccipital process that is narrower in cross section and relatively shorter [6(10)], and a knob-like protocone

on the P4 [13(1)]. Among these fossil carnivorans, the subfamily Procyoninae has a deep suprameatal fossa [1(1)], a P4 with an internal shelf [15(1)], an M2 with a separated paracone and metacone [27(1)], and an m2 with an enlarged hypoconid

[38(1/2)], and lacks an alisphenoid canal [2(1)]. The Tribe Potosini sensu lato has a reduced to absent P1/p1, a morphology also present in derived procyonines such as

Cynonasua, Amphinasua, and Brachynasaua) [11(1)]. The upper molars are triangular and have rounded cusps [19(1)], and have an m1 with a basined talonid [36(0)].

Furthermore, the Tribe Procyonini has a mastoid process that is either at a 45-degree angle (a synapomorphy only present in Bassariscus) or vertical (all other procyonines)

[5(1/2)], and an enlarged M1 parastyle [21(1)].

The TF specimen’s relationship with both Bassariscus and Probassariscus is supported by having an P4 protocone that is relatively not as anterior as the potosines, while in all other procyonines it is more medially located [14(2)]. The P4 also has a somewhat reduced metacone blade [17(1)] and an M1 with a hypocone while most other procyonids have a reduced (procyonini) or greatly reduced to absent (potosini) hypocone [22(2)]. TF specimens differ from Bassariscus and Probassaricus by having a

P4 with a weaker parastyle [18(1)], a shared character with Broiliana, Bassaricyonoides,

Bassaricyon, and Potos. The m1 trigonid of TF has a paraconid, protoconid, and metaconid that is more or less equally spaced [28(0)], while the paraconid and metaconid are close together in Bassariscus and Probassariscus. Finally, TF specimens have an m1 entoconulid, which is a shared with Arctonasua, Cyonasua, Amphinasua, and Champalmalania [33(1)] but absent in Bassariscus and Probassariscus.

Surprisingly, in all of our MPT’s (most parsimonious trees), taken together, the

TF specimens share a common ancestor with Bassariscus, and more interestingly, the topology suggests that a procyonid dentally similar to Bassariscus might have been the ancestor to an early Miocene Bassariscus-like mesocarnivoran. For instance, MPT A, places the TF specimens as sister taxa to Bassariscus having an M1 hypocone that is expanded posterointernally [23(2)] and an m1 with a hypoconulid [34(1)]. In MPT B, TF is outside of Bassariscus and Probassariscus and shares a common ancestor to all other procyonines. This is supported by having an m1 entoconulid [33(1)], a shared morphology with Arctonasua, Cyonasua, Amphinasua, and Champalmalania. In MPT C, the TF specimens fall in a polytomy with Probassariscus. In contrast to Bassariscus,

Probassariscus and the TF specimens have a metaconule on the M1 a cusp that is prominent or secondarily reduced to absent in most other procyonines [24(1)]. Similarly, to Edaphocyon, Arctonasua, and Paranasua, the P4 of TF (and Probassariscus) have a small hypocone [16(1)]. Finally, the TF specimens represent an intermediate procyonid with an m1 with equal talonid and trigonid width [35(1)] like in Edaphocyon, the potosines Parapotos and Potos, and the procyonines Probassariscus, Arctonasua,

Cynonasua and Chapalmalania in MPT D. One should note that the TF procyonine is closely related to Bassariscus and Probassariscus. More complete fossils perhaps could be better classified as a basal species of Bassariscus, a derived species of

Probassariscus, or a new genus. These fossils are tentatively attributed to

Probassariscus because they differ in only five characters versus six characters with

Bassariscus. In assembling the character matrix, it was noticed that character 18 has

only two-character states and therefore the data set was ran with Potos scored as 1 for character 32 and only having 2 character states (Appendix C).

Discussion

A procyonid, likely representing a potosini based on the morphology of the talonid of an isolated m1 (Figures 2-2 J, K, L) is reported from the TF fossil site.

Although basal potosines have a distinctive upper P4 with an anterior-posteriorly elongated protocone lingual to the paracone (Baskin, 2017) with no hypocone on the internal cingulum, the morphology preserved in UF 406549 is more similar to that preserved in similarly sized isolated teeth likely representing a potosines similar to

Bassaricyonoides phyllismillerae from the Miller Site Fauna also in Florida (Baskin,

2004).

A second procyonid species is represented by an isolated partial mandible (UF

276543) that preserves the following synapomorphies for Procyonidae identified by

Baskin (2004): 1) the m3 is absent; 2) lower m1 with distinct hypoconulid, and 3) the anterior premolars (p2-p4) are relatively wider than those from any other hypocarnivorous taxon present in TF (e.g., the mustelid Zodiolestes, the borophagine canid Osbornodon). The isolated partial dentitions from TF is referred to the Subfamily

Procyoninae based on: 1) the presence of a knoblike protocone on the P4 with a better developed parastyle, 2) having a more posteriorly placed protocone, and 3) having a lower m1 with a distinct entoconid and a shorter metacone blade (Baskin, 2004).

Furthermore, this new procyonid taxon from TF has a distinct mesocarnivorous dentition that is not present in hypocarnivorous potosines from older assemblages in Florida. The

P4 protocone is more posteriorly and more medially located than in postosines, the P4

also has a reduced metacone blade, and the M1 exhibits a hypocone similar to other procyonines (Baskin, 2004).

The TF taxon has an M1 with a prominent hypocone that is located more posterior to the protocone than in other procyonids, but also characteristic of

Bassariscus (Baskin, 2004). The morphologies present in bassariscine procyonids

(Bassariscus and Probassariscus) are not present in the canid Phlaocyon also from

Florida. These TF partial isolated procyonid teeth are more similar to those of

Probassariscus than Bassariscus astutus, and therefore they are now tentatively referred to cf. Probassariscus. Furthermore, the presence of a reduced to absent hypocone on the M1 confirms that this is a primitive procyonid morphology, which seems appropriate for one of the earliest occurrences of procyonines in North America

(Tedford et al., 2004). Although based mainly on few synapomorphies preserved in the partially preserved fossils, not enough material is available to fully define a new species of Probassariscus.

Both procyonines (cf. Probassariscus) and potosines inhabited Florida during the earliest Hemingfordian. The occurrence of bats, rodents, and other small carnivorans suggests that the karstic habitat in TF (see Pratt, 1989) was diverse enough to sustain scansorial and arboreal carnivores. Independent of the ancestral relationships of extant potosines, our result confirms the presence of meso and hypocarnivorous in Florida during the earliest Hemingfordian. More interestingly, our results also support the monophyly of the Tribe Potosini, which includes the extant Bassaricyon and Potos, had diverged from the Procyonini by the early Miocene in North America (Baskin, 2004).

Figure 2-1. Distribution of fossil procyonids from the Oligocene through the Miocene.

Figure 2-2. TF potosine, fossil, lower m1: UF 406549 right. A. labial, B. lingual, C. occlusal views. cf. Probassariscus, fossil, lower m1: UF 209500, left, D. labial, E. lingual, F. occlusal views; UF 406540, left, G. labial, H. lingual, I. occlusal views; UF 225330, right, J. labial, K. lingual, L. occlusal views.

Figure 2-3. Lower m1 box plot of the Natural log of APL (blue) and PMW (gray) with number of individuals measured. A. Potos flavus, B. Bassaricyonoides phyllismillerae, C. TF potosine, D. Nasua narica, E. Procyon cancrivorus, F. Procyon lotor, G. Bassariscus astutus, H. Bassariscus ogallala, I. TF cf. Probassariscus, J. Phlaocyon sp., K. Phlaocyon ansechens, L. Pachycynodon dobius, M. Phycynodon tenuis, N. Leptarctus ancipidens, O. Miomustela sp., P. Oligobunis sp.

Figure 2-4. cf. Probassariscus, fossil, Upper P4: MCZ 21446, right, A. labial, B. lingual, C. occlusal views; UF 406542, left, D. labial, E. lingual, F. occlusal views; UF 1406541, right, G. labial, H. lingual, I. occlusal views; cf. Probassariscus, fossil, Upper M1: UF 259985, left, J. labial, K. lingual, L. occlusal views.

Figure 2-5. cf. Probassariscus, fossil, UF 276543, left dentary with p2-p4, A. labial, B. lingual, C. occlusal views.

Figure 2-6. cf. Probassariscus, fossil, UF 181215, right partial dentary with lower p4 A. labial, B. lingual, C. occlusal views.

Figure 2-7. Bassariscus astutus, modern, UF 11933, A. left labial, B. lingual left cross section, C. occlusal view of left side.

Figure 2-8. Upper P4 box plots of the Natural log of APL (blue) and TW (gray) with number of individuals measured. A. Potos flavus, B. Nasua narica, C. Procyon cancrivorus, D. Procyon lotor, E. Bassariscus astutus, F. TF cf. Probassariscus, G. Phalocyon (Notocyon) sp., H. Phalocyon sp., I. Phalocyon willistoni, J. Leptarctus ancipidens, K. Zodiolestes sp., L. Parictis sp.

Figure 2-9. Bassariscus astutus, modern, UF11933, left dentary A. labial, B. lingual, C. occlusal views.

TF cf. TF cf.

A. .

TF cf. TF cf.

Figure 2-10. Possible phylogenetic relationships of TF cf. Probassariscus, fossil taxa. A. Tree 1 TF specimen is closely related to Bassariscus, B. Tree 2 TF specimen more closely related to Probassariscus, C. Tree 3 TF specimen in a polytomy with Probassariscus, D. Tree 4 TF specimen shares a common ancestor with Probassariscus. † depicts an extinct genus.

Figure 2-11. Character matrix from Baskin, 2004 with Thomas Farm cf. Probassariscus scored.

CHAPTER 3 3D GEOMETRIC MORPHOMETRICS ON COMPLETE VS. PARTIAL FORELIMB ELEMENTS OF CARNIVORANS

Introductory Remarks

A major challenge in paleontology is to obtain statistically significant sample sizes. Complete specimens often are rare in collections because of taphonomic processes. Incomplete specimens can be difficult to classify or to make interpretations of behavioral characteristics. Geometric morphometrics has become the favored methodology for interpreting shape variation (Rohlf and Marcus, 1993; Adams et al.,

2004, 2013). Cornette et al. (2015) demonstrated that partial specimens can be accurately classified using both 2D traditional landmarks and sliding semi-landmarks, although sliding semi-landmarks produced better results. A more recent analysis by

Cardini (2016) looked at the accuracy of using only one side of bilaterally symmetrical specimens and demonstrated that congruency was relatively high on a large scale but is much lower on a microevolutionary scale.

Can non-bilaterally symmetrical partial specimens (i.e., the distal end of the humerus) be used in morphometric analyses that seek ecomorphological interpretations? 2D morphological studies on the humeral epiphyseal shapes in species of Felidae found a higher classification accuracy for locomotion on the distal end than the proximal end of the humerus (Walmsley et al., 2012). Therefore, it is hypothesized that using 3D morphometrics and expanding beyond the epiphysis will provide more accurate rates of classification for the humerus and that this can also be applied to the radius and ulna. It is also hypothesized that using k-Nearest neighbors analyses will provide a higher rate of classification in comparison to leave-one-out linear discriminant

analyses. Through these techniques one can make morphological interpretations beyond phylogeny and locomotion, such as hunting strategy and potentially diet.

Materials and Methods

Materials

Between one and five individuals from each of 81 species representing 57 genera from 10 Carnivoran families were included: Canidae (11 species), Felidae (18 species), Eupleridae (1 species), Hyaenidae (2 species), Mustelidae (22 species),

Mephitidae (4 species), Procyonidae (8 species), Ailuridae (1 species), Ursidae (8 species), and Viverridae (6 species). Marine pinnipeds (families Phocidae, Odobenidae, and Otariidae) were excluded from the study because their forelimbs are highly modified for swimming and thus are not comparable to the other terrestrial carnivorans. When possible, both males and female adults were analyzed for each species. The specimens are from the Florida Museum of Natural History, Gainesville, Florida, USA; East

Tennessee State University, Johnson City, Tennessee, USA; and the Muséum National d’Histoire Naturelle, Paris, France. Specimens were digitized using a reuckmann 3D surface scanner (camera resolution of 1.4 megapixels), a NextEngine 3D surface scanner, or by NanoCT imaging. The specimen list is found in Appendix D.

Methods

To accurately evaluate the nature of forelimb functional morphology, 3D sliding semi-landmarks were chosen as the preferred method to analyze the complexity of the elbow articulation. Methods follow the protocol outlined by Gunz and Mitteroecker

(2013). Homologous anatomical landmark coordinates (traditional landmarks) were first placed on the elements resulting in 30 traditional landmarks for the humerus, 22 for the radius, and 29 for the ulna (based on Fabre et al. 2013b) (Appendix E). Curve sliding

semi-landmarks (CSSL) were placed along the articulation surfaces, muscle attachments, and any other important morphological features present on the elements that traditional landmarks would not measure (Figures 3-1, 3-2, 3-3). These CSSL both started and ended at a traditional landmark to ensure consistency. Surface sliding semilandmarks (SSSL) are more difficult to place on homologous locations consistently. To reduce error of SSSL placement, a template specimen was created for the complete and partial humeri, radii, and ulnae. Each template is marked with both traditional landmarks and CSSL, after which a mesh of SSSL was hand-placed on each template.

In the case of the ulna, a second set of CSSL was placed along the entire length of the diaphysis to allow for a more accurate project of the template mesh due to the high variability and slender morphology of the ulna. A thin-plate spline interpolation function was computed from the traditional landmarks and CSSL to extrapolate the SSSL from the template onto each specimen. Spline relaxation was performed to allow for the

CSSL and SSSL to slide along curves and tangents to take the shape of the actual specimen (Fabre et al., 2013b; Gunz and Mitteroecker, 2013). Numbers of SSSL differed from template to template. A separate template was created for each proximal, distal, and complete analysis for each element.

To analyze the proximal and distal ends of the bones, a new template and therefore a new set of SSSL were created for each (Figures 3-4, 3-5, 3-6, 3-7, 3-8). A sub-set of the traditional landmarks and CSSL was selected to analyze each end. In some cases, additional CSSL were added to enclose the curves; this was vital for an accurately placed projection of SSSL. A new mesh was created for an extrapolation onto the enclosed aspects of the CSSL, such as the lunar notch on the proximal end of

the ulna (Figure 3-8). Without an enclosure of CSSL, the SSSL would not be contained and therefore would not be accurately projected onto the elements. For accurate comparisons between the complete, proximal, and distal aspects of an element, all traditional and CSSL coordinates were consistent. The distal end of the ulna was excluded from this analysis due to its simplistic morphology and the difficulty to place the CSSL to provide for a large enough surface area for the projection of the SSSL.

A Procrustes superimposition (Rohlf and Slice, 1990) was performed using the package Rmorph1 in R2. The Procrustes superimposition rotates, translates, and equally scales all the specimens to remove size as a possible variable. Principal components analyses (PCA) was performed on the data to determine how specimens are distributed within morphospace. Because animals with a similar evolutionary background are expected to exhibit similar morphologies, a phylogenetic MANOVA was performed on the PCAs (Fabre et al., 2013b; Garland et al., 1993) as well as a multivariate K statistic to determine phylogenetic signal relative to feasibility under Brownian motion (Adams,

2014) based on the univariate K statistic described by Blomberg et al. (2003).

The F-values from the complete elements were compared against the F-values of the distal and proximal ends to find reliability. Any phylogenetically significant F-value analyses were noted with all components subjected to a discriminant analysis. The discriminant determined how accurately this analysis can predict an unknown’s characteristics, i.e., if the unknown is cursorial or arboreal.

1 Baylac, M. 2012. Rmorph: a R geometric and multivariate morphometrics library. Available from the author: baylac@ mnhn.fr via personal communication with A.-C. Fabre

2 R Development Core Team. 2014.R: a language and environment for statistical computing. Vienna: R Foundation for Statistical Computing, ISBN 3-900051-07-0. Available at: http://www.R-project.org

The characters for locomotion, hunting strategy, activity pattern, diet, vegetation/habitat, and social behavior were collected from the literature to build a matrix for analysis (Figure 3-9). Body mass was also collected, which will be an important factor for Chapter 4. Locomotion categories (Samuels et al., 2013) include: terrestrial, cursorial, scansorial, arboreal, semifossorial, and semiaquatic (Table 3-1).

Hunting strategy definitions (from Van Valkenburgh, 1985) include: ambush, pursuit and pounce, pursuit, and occasional. I added two other variables: various (for animals that implemented more than one strategy) and aquatic (Table 3-2). Activity pattern, diet, and vegetation categories were obtained from Gittleman (1985, 1989): activity pattern

(nocturnal (N), diurnal (D), crepuscular (C), nocturnal and crepuscular (O), and arrhythmic (A)), diet (defined as constituting at least 60% of the animal’s diet – carnivorous (M), insectivorous/worm eating (I), folivorous/frugivorous (V), piscivorous

(F), or omnivorous (O)), vegetation which herein I call habitat (open grassland and forest (N), open grassland (O), forest (F), open grassland and woodland (S), dense brush and scrub (D), desert (T), woodland (W), aquatic (Q)). Social behavior is categorized as solitary, pairs, groups, and variable groups (Ortolani and Caro, 1996).

To determine the reliability of the elements and the partial elements for predicting unknowns, a leave-one-out cross-validated linear discriminant analysis (LDA) was performed on each characteristic via the ‘MASS’ package in R. The LDA assumes equal covariance matrices among groups and works by assigning a new observation to an element via seeking the closest mean vector. The LDA method predicts a classification per individual by comparing it to all the remaining specimen designations (Fabre et al.,

2015b). After each individual prediction is made, the percent of classification accuracy

for each designation (i.e., hunting behavior) is calculated. LDAs were run on the means for each species.

ecause LDA’s focus is on maximizing the differences between the groups, which can sometimes force the data and skew results, a k- nearest neighbors test (k-

NN) was also preformed via the ‘CLASS’ package in R. A k-NN focuses on the differences between individuals rather than the differences between groups, and therefore may provide a more reliable result. Protocol for the k-NN analyses is similar to that in Hanot et al. (2017). The amount of variation included in the analyses was ~90%, so the number of principal components varied from analysis to analysis. Also, the number of Nearest-Neighbors for each analysis varied from k=1 to k=5. Only the k values with the highest percent of correct reclassification were included in the discussion.

This research project included running the PCAs, MANOVAs, phylogenetic

MANOVAs, LDAs, and k-NN analyses on family-level, activity pattern, diet, locomotion, hunting strategy, social behavior, and habitat for the 81 extant carnivoran species. This paper focuses on which of these aspects are the most useful for the interpretation of fossils, which will be the focus of Chapter 4.

Phylogeny

The phylogenetic tree used in this analysis is from Nyakatura and Bininda-

Emonds (2012), whose Carnivora supertree was shortened to include only the 81 species of carnivorans in the dataset (Figure 3-10). The phylogenetic positions of the fossa and the red panda are highly disputed; in Nyakatura and Bininda-Emonds (2012) the fossa (Eupleridae) lies just outside of Hyaenidae and the red panda (Ailuridae) near the procyonids, mustelids, and mephitids.

Phylogenetic Signal

To determine the strength of the phylogenetic signal, a multivariate K-statistic was calculated (Adams, 2014) on datasets of the whole and partial specimens. To do this the ‘GEOMORPH’ library (Adams and Otarola-Castillo, 2013) in R was used. A strong phylogenetic signal is represented by a higher K-value, therefore a K-value of greater than 1 suggests a strong phylogenetic affinity. On the other hand, a weak K- value such as those close to 0 suggests a strong degree of morphological convergence.

A phylogeny of our dataset was mapped using the polymorphospace function in R, which was implemented in the ‘PHYTOOLS’ library (Revell, 2012).

MANOVA and Phylogenetic MANOVA

A multivariate analysis of variance (MANOVA) is a test to see if there are mean differences in the shape of the element (i.e., humerus, radius, or ulna). This statistical analysis allows for more than one dependent variable to be analyzed at a time. The phylogenetic MANOVA analyses evaluates how the phylogeny affects the data and therefore, the strength of the phylogenetic signal in the element’s morphology.

Results

Geometric Morphometrics

Family-level PCAs

Humerus Whole. Over 75% of the overall shape variation is accounted for in the first three axes. When examining the morphospace defined by PC1 (55.4% total shape variation) and PC2 (12.3% total shape variation), the family Ursidae is well separated from the other extant families (Figure 3-11A). Ailuropoda melanoleuca, the giant panda, is outside most other members of Ursidae and falls closer to some species of

Mustelidae. Mustelidae is the least separated family, which is not surprising due to its

wide variation in morphology. Mephitidae overlaps with Mustelidae and is separated by the second axis. Viverridae is also separated by the second axis and overlaps slightly in morphospace with Procyonidae. The first axis separates Nasua narica, N. nasua, and

Potos flavus from the other procyonids in this analysis. Ailurus fulgens, the red panda and only extant member of Ailuridae, falls near members of Viverridae, Procyonidae, and Ursidae. Felid morphospace overlaps with members of Viverridae, Mustelidae,

Procyonidae, and Canidae. It is the least well defined in this PCA. Canid morphospace is nicely separated by the first axis. It overlaps slightly with Acinonyx jubatus (cheetah),

Felis chaus, and F. margarita. Interestingly, Spethos venaticus (bush dog) falls within the morphospace of Hyaenidae, which is well separated by both axes. The only member of Eupleridae in this analysis is near the origin of the axes.

Humerus Proximal. In comparison to the whole humerus, only 47% of the overall shape variation is accounted for in the first three axes of the proximal humerus analysis (Figure 3-11B). The first axis represents 27.3% of the total shape variation while axis two represents only 11.3%. The family Ursidae continues to be well separated from other carnivoran families. It is more separated by the second axis than the first axis. Mustelidae is well separated by the first axis and overlaps mainly with procyonids and mephitids. Mephitidae is well separated by both axes and is more constrained than in the complete humerus PCA. Viverridae is the least constrained and overlaps with Felidae morphospace. All members are separated by the second axis except for the large Indian civet, Viverra zibetha. Procyonidae is well separated by the first axis and overlaps with some mustelids. Compared to the complete humerus PCA, felids are more constrained and well separated by the first axis. Canids are also well

separated by the first axis with the proximal morphology of Spethos venaticus falling within viverrids and outside of the hyaenid morphospace. Hyaenidae is well separated by both axes and Cryptoproctra ferox (Eupleridae) is near the origin as in the complete humerus PCA.

Humerus Distal. The distal humerus has 57% of the overall shape variation accounted for in the first three axes, which is more than the proximal humerus (47%) but not as much as the whole humerus at 75% (Figure 3-11C). Axis one accounts for

26% of the total shape variation, while the second axis accounts for 21.8%. In this PCA,

Ursidae is not as well defined as in the complete humerus and proximal humerus PCAs.

Its morphospace spans the first axis and is separated by the second. The giant panda,

Ailuropoda melanoleuca, falls within the procyonid morphospace. Mustelidae is not separated well by either axis although most members are near the origin. Mephitids are relatively constrained and are separated by both axes as is the mustelid Taxidea taxus, the American badger, within its morphospace. Viverridae is again the least constrained family of carnivorans, ranging across the PCA’s morphospace. Procyonidae is separated by the first axis and encompasses the euplerid morphospace. Most felids are separated by both axes, although members of Panthera have a more positive shape variation along the second axis.

Radius Whole. Over 71% of the shape variation is accounted for in the first three axes of the whole radius. The first PC accounts for 32.6% of the total variation while the second PC accounts for 15.9% (Figure 3-12A). In contrast to PCAs of the humerus, the fossa (Eupleridae) is well separated in each radius PCA from other carnivorans. PC1 separates the ursids, hyaenids, and canids (excluding Spethos venaticus, the bush

dog). Interestingly, Spethos venaticus falls near the mustelids Mustela lutreola and M. putoris. Most mustelids are also separated by the PC1 axis. Aside from being separated by PC1, all ursids except Melursus ursinus and Ursus arctos, fall on the negative end of

PC2. Hyaenidae is well separated by both axes and forms a non-overlapping morphospace. Felidae is well separated by PC2.

Radius Proximal. Though not as great as the complete radius, the proximal radius’ first three axes account for nearly 62% of the total shape variation (Figure 3-

12B). This large percentage consists mostly of the first two axes with axis 1 accounting for 32% and PC2 for ~13% of the shape variation. As with the complete radius, PC1 is effective at separating the canids and ursids, whereas PC2 separates the felids and hyaenids excluding the cheetah Acinonyx jubatus, which falls within the canid morphospace. Most mustelids (except Melogale moschata and Lyncodon patagonicus) are also separated by the first principal component.

Radius Distal. The first three PCs for the distal radius account for about 49% of the total shape variation. The first axis accounts for almost 24% percent of the variation while the second axis is only ~3% (Figure 3-12C). Felidae has a distinct morphospace in the distal radius PCA, being well separated by both PC1 and PC2 as well as from other carnivoran families. Hyaenids are also well separated by both axes and fall near the felid morphospace. There is some overlap on PC1 for canids, although PC2 does separate them except for Vulpes velox which falls near the ursid morphospace. As with the complete radius PCA, the mephitid morphospace and mustelid morphospace overlap. The fossa is well separated from all other carnivorans which is also seen in the complete radius PCA.

Ulna Whole. The first three axes of the complete ulna account for the highest percentage of shape variation in this analysis at over 77%. PC1 had over 55% of the total shape variation while PC2 had 17% (Figure 3-13A). The first axis separates the

Canidae, Hyaenidae, Ailuridae, and most of Ursidae (except the sloth bear, Melursus ursinus). As with most of the PCA analyses, the viverride morphospace is widely distributed over both axes and, unlike the radii PCAs, the euplerid is more centrally located and falls near mustelids and felids.

Ulna Proximal. The proximal ulna has a relatively high percentage of shape variation for the first three PCs at almost 59%, consisting of mostly PC1 at ~36% and

PC2 at 13% (Figure 3-13B). Ursidae separates out nicely in its own morphospace by the first axis outside of all other carnivorans. There is much more overlap of the canid morphospace when looking only at the proximal ulna versus the other described PCAs.

Felids are well separated by the second axis as well as hyaenids, viverrids, the euplerid, and mephitids. Most mustelids are also well separated by PC2 and overlap some with canids.

Locomotion PCAs

The six defined locomotion categories were overlaid onto the PCAs. For the whole humerus, there is some overlap of the locomotor morphospaces (Figure 3-14A).

The first axis separates the cursorial morphospace from the other modes of locomotion, making this clearly defined. The semiaquatic and semifossorial locomotor behaviors have a few individuals not separated by PC1. The same is also true for the terrestrial morphospace. The second axis separates most of the scansorial species, whereas the morphospace for arboreal carnivorans overlaps most locomotor categories for the complete humerus PCA.

In comparison to the complete humerus PCA, the proximal humerus PCA’s arboreal morphospace is more clearly defined and is mostly separated by the first axis

(Figure 3-14B). The first axis, PC1, also is effective at separating out the semiaquatic morphospace with only one species on the other side of the axis. The cursorial morphospace is even more clearly defined by PC1 in the proximal humerus PCA than in the complete humerus PCA. The second axis also separates out the arboreal morphospace with only one species on the negative side. Like the complete humerus

PCA, the proximal humerus PCA has overlapping semifossorial, scansorial, and terrestrial morphospaces.

In contrast to the complete humerus and proximal humerus PCAs, the distal humerus PCA is the most effective at defining the arboreal morphospace via the first axis (Figure 3-14C). The distal humerus PCA also clearly defines the cursorial morphospace. Most of the semiaquatic morphospace is found on the positive side of the second axis with only three species on the negative side. Overall, there is overlap of terrestrial, scansorial, and semifossorial morphospaces which is consistent with all humeri PCAs.

Similar to the humerus PCAs, the complete radius PCA clearly defines cursorial locomotion morphospace and in this case by PC1 (Figure 3-15A). Due to the high amount of total shape variation across the first axis, it acts as a good separator for different methods of locomotion. The first axis also defines the semiaquatic morphospace and most of the arboreal species. On the other hand, the second axis nearly defines arboreal morphospace and semifossorial, although with some overlap

with a few species on either side. Scansorial and terrestrial morphospaces have more overlap and are not as well defined by these axes.

The same is true for these locomotor morphospaces in the proximal radius. The cursorial morphospace is well defined and separated by the first axis (Figure 3-15B).

Most of the overlap of the cursorial morphospace is with the terrestrial which is not well defined by either axis. Semiaquatic and scansorial are also poorly defined by these axes, although the arboreal morphospace is defined by both axes with only one species outside of each axis.

The distal radius analysis for locomotion shows more morphological overlap than in the proximal or complete radius (Figure 3-15C). Similarly, the cursorial morphospace is the most well defined and in this case separated solely by the second axis. Neither axis is effective in defining the other locomotor categories for the distal radius.

In comparison to the radii locomotor PCAs, the complete ulna PCAs have more clearly defined locomotor morphospaces across the first two axes (Figure 3-16A). The first axis fully separates the cursorial morphospace and it also separates most of the semiaquatic morphospaces. The semiaquatic morphospace is also largely separated by the second axis. PC2 separates more species with terrestrial locomotion on the negative side. Scansorial, semifossorial, and arboreal fall largely near the origin and do not separate as well via the first two PCs.

In the case of the proximal ulna, the scansorial and semifossorial morphospaces are greatly expanded across the first axis (Figure 3-16B). The first PC, however, does separate the arboreal morphospace and largely the semiaquatic. The second axis is more effective in separating the scansorial and semifossorial morphospaces than in the

complete ulna PCA. Unlike the complete ulna, the proximal ulna’s first two PCs do not separate the cursorial morphospace.

Hunting strategy PCAs

When describing the hunting strategy PCA of the complete humerus, PC1 is relatively effective at separating out different hunting techniques such as pursuit and pounce morphospace (excluding one species), the pursuit morphospace, and most of the aquatic morphospace (Figure 3-17A). The majority of the aquatic morphospace is also separated PC2, which distinguishes the majority of semifossorial hunters, pursuit hunters, and various strategy hunters. Occasional hunting morphospace and ambush hunting morphospaces tend to overlap.

In comparison to the complete humerus hunting strategy PCA, the proximal humerus separates out the occasional hunters across the first axis (Figure 3-17B). The first axis can be used to distinguish the aquatic hunters as well as the various strategy hunters and pursuit. All but five species of the ambush hunters fall on the positive side of the second axis. Various hunters have a very clear defined morphospace on the positive side of axis two, but that could also be due to the low number of species.

In the case of the distal humerus, the pursuit morphospace is distinct and separated out from the pursuit and pounce morphospace. It can be defined by the first axis, as can the various hunting strategy morphospace. The semifossorial morphospace and aquatic morphospaces have only one species that falls across the second axis

(Figure 3-17C). While there is still overlap between and over the point of origin, occasional hunting and ambush hunting have a more distinct morphospace than in other humeri PCAs.

The complete radius hunting strategy PCA clearly defines the aquatic and pursuit hunting strategies across PC1 (Figure 3-18A). All but one species of semifossorial hunters can also be separated by this first axis. Similarly, the second axis clearly separates pursuit carnivorans, semifossorial carnivorans, and most ambush predators.

The occasional hunters fall over the point of origin and like the PCAs of the humerus, they overlap with many of the different hunting strategies’ morphospaces.

In comparison to the complete radius, the morphospace of the pursuit and pounce hunters is greatly expanded across the second axis with some overlap over the first axis as well in the proximal radius PCA (Figure 3-18B). The morphospace of semifossorial hunting strategy is also greatly expanded but can be separated by the second axis. The morphospace of pursuit predators and ambush predators overlaps with some of the semifossorial and pursuit and pounce strategies. The aquatic morphospace falls squarely within the occasional hunting morphospace and is clearly defined by the first axis and nearly defined by the second axis with one species on PC2 and another on the negative side. Most ambush predators fall on the positive side of

PC1, as do with most of the occasional hunters.

The distal radius PC1 separates the aquatic, and various hunting strategy morphospaces. Pursuit predators are separated by the positive side of the second axis where most of the pursuit and pounce and the ambush carnivorans fall (Figure 3-18C).

The morphospace of the various hunting strategies and those of the pursuit and pounce strategy are widely spaced but not to the extent as those in the proximal radius PCA.

Most of the species that occasionally hunt fall on the negative side of PC2, while most of the ambush hunters are on the positive side.

Aquatic and semifossorial hunting strategy morphospaces are clearly defined by the first component axis in the PCA of the complete ulna (Figure 3-19A). Unsurprisingly, the pursuit carnivoran morphospace overlaps somewhat with the pursuit and pounce morphospace and with the ambush predators. This morphospace is clearly separated by both axes. The majority of the ambush and pursuit and pounce predators fall on the negative side of PC2. The occasional hunters, as with most of the PCAs, fall near the origin and overlap with most of the hunting techniques.

In comparison to the complete ulna, the morphospace of the occasional hunters is more spread out across the first axis (Figure 3-19B). Also, widely distributed across the first and second axis are the ambush hunters, although most of these species fall on the negative side of PC2. Aquatic hunters are separated by most axes though not as clearly as in the complete ulna PCA. There is also more overlap of the semifossorial and pursuit and pounce hunters.

Not as significant as the family, locomotion, or hunting strategy, the PCAs for diet and social behavior are included as appendices. Habitat and activity pattern PCAs are available upon request and not included as appendices because they lack interpretable information.

Phylogenetic Signal Results

The multivariate K-statistic results indicate that the shape data for each element of the forelimb is significant (Appendix E). They also show that each partial element

(e.g., the distal humerus) is also significant when looking at family level, although the signal strength varies. For example, the signal is slightly stronger in the whole humerus than in the distal end, suggesting more morphological convergence in the proximal end.

The complete radius has a higher K value (0.31) than the proximal and distal radii (0.24

and 0.23, respectively). The K-statistic values of the ulnar analyses show a different pattern with the proximal ulna (0.34) being just slightly higher than in the complete ulna

(0.33).

MANOVA and Phylogenetic MANOVA

The MANOVAs at 90% and 95% of the overall variance for the complete humerus indicated a significant P-value for all character criteria except the activity pattern at 95% variance. The phylogenetic MANOVAS portray the same pattern

(Appendix E). This pattern is not seen, however, in the proximal and distal humerus phylogenetic MANOVAs, where the characters of locomotion, diet, and social behavior are typically phylogenetically significant. The MANOVAs of the radius show a significant

P-value for all characters excluding activity pattern, although the phylogenetic MANOVA

P-values are typically not statistically significant when examining just the proximal and distal ends. The phylogenetic MANOVA of the complete radius is similar to that of the complete humerus. The MANOVAs of the ulnar analyses have significant P-values, while the P-values for locomotion are the most significant when examining the phylogenetic MANOVAs.

Linear Discriminant Analyses

The LDAs typically have higher rates of reclassification on complete specimens than on just the proximal or distal ends (Table 3-3 a,b). Family-level designation was best classified overall with a minimum of 67% accuracy for all levels of element completeness. Therefore, due to the lower levels of classification for other categories, family-level is the only LDAs described here; all other LDA analyses are available upon request.

The complete humerus had an 89% rate of reclassification for family level designation (Table 3-3b). Classification rates for the proximal and distal humeri were lower at 72% and 81%, respectively. Both Canidae and Felidae were well classified on the complete humerus with only one felid being classified as a viverrid (Appendix E).

Mustelids, being one of the most morphological diverse families of carnivorans, were classified as mustelids, procyonids, ailurids, or felids based on the complete humerus.

The proximal humerus also is an effective way of classifying canids and felids, although two species of felids were interpreted as viverrids (Appendix E). Compared to the complete humerus and distal humerus, the proximal humerus classification of mustelids much more varied and separated out into 6 different families. The distal humerus saw better classification for the felids, while one of the canids was classified as a hyaenid, most likely being the bush dog (Appendix E). A higher rate of classification is also seen in six mustelids, with only two species being classified as ailurids. Overall, the distal humerus had an overall lower rate of classification compared to the complete.

Just as in the LDA humeri analyses, the LDA radii analyses have a higher rate of classification of the complete element (Table 3-3b). Canids, felids, viverrids, hyaenids, and ursids are well classified using the complete radius (Appendix E). The rate of classification is almost as high for the complete radius as for the complete humerus,

87% and 89%, respectively. Once again, mustelids are the most difficult to classify correctly with some species categorized as ailurids, mephitids, or procyonids. When examining just the proximal end of the radius, the classification rates go down by ~20% with certain canids now classified as mephitids or mustelids, and some felids classified as mustelids or viverrids (Appendix E). Accuracy rates increase somewhat when

focusing on just the distal end compared to the proximal end. Canids and felids were more accurately classified using the distal end than the proximal end (Appendix E).

In comparison the LDAs of the radii, the ulnae had relatively higher classification equivalent to that of the LDAs of the humerus (Table 3-3b). Canids and felids were well classified with only one species of felid classified as a canid (Appendix E) Mustelids were relatively well categorized as well with only three species falling within

Procyonidae. The proximal ulna also is useful to classify canids and felids, with one canid classified as a hyaenid and two felids as a hyaenid and procyonid (Appendix E). k-NN Analyses

The k value for each analysis can be found in Appendix E. In most of the cases, a k value of 1 for both analysis with and without centroid size as a factor was found to have the highest percent of correctly classified specimens. Overall the k-NN analyses provided a higher rate of reclassification when compared to the LDA analyses. Table 3-

4 shows the highest rate of classification (with or without centroid size) with varying k values. Family level and hunting strategies were the most likely to be accurately classified. In the humerus and the radius, the distal end provided higher rates of classification than the proximal end or even the complete element in all cases. The patterns in the ulna were similar, although the proximal end provides for a better rate classification than the distal.

In the case of the ulna k-NN analyses, just focusing on the proximal end provided the highest rate classification of all k-NN analyses (92%), yet the complete ulna provided one of the lowest rates (76%) for family-level classification.

Discussion

The results of these analyses show that complete and partial specimens can be used to classify the characteristics of family-level designation, locomotor style, hunting strategy, diet, and social behavior with varying levels of accuracy. These analyses can also be used to classify albeit with less accuracy, activity pattern and habitat. The phylogenetic signals for each analysis indicated that the morphology of the elements are to some extent constrained by the animal’s phylogeny.

The PCAs reflect the variation of locomotor styles and hunting strategies within the families but also so indicate distinct morphospaces that can be used for interpretation of element morphologies. Family-level morphospaces are the most distinct and are in some cases clearer (e.g., the proximal humerus) when looking at partial elements. Examining only the proximal or distal ends of an element helps to eliminate some of the noise in the data and allows for more distinct morphospaces that can be separated out by the axes. The complete elements first axes consistently show higher levels of variation. For example, the complete humerus first axis has a percentage of total variation of 55.4% while the proximal humerus has 27.3% and the distal humerus has 26%. This makes sense because there is more morphological variation across the entire element than just looking at a partial element.

Nevertheless, one must take into consideration the importance of the elbow joint in phylogenetic shape variation. The distal humerus had more shape variation in the first two axes than the proximal humerus, the proximal radius had more shape variation in the first two axes than the distal radius, and the proximal ulna had still higher shape variation than the distal humerus. This suggests that the elbow joint plays a significant role in phylogenetic morphology across Carnivora.

The results of the LDA classifications indicate that most specimens can be accurately classified to family level, but less accurately to the other criteria. A complete element gives a higher rate of classification than examining partial specimens for all criteria in the LDA analyses. The distal humerus provides for a better classification of family, locomotion, hunting strategy, diet, and habitat than proximal humerus. The distal radius LDA analyses better classify family, hunting strategy, activity pattern, habitat, and social behavior than the proximal radius. The proximal ulna LDA was better at classifying the hunting strategy of the carnivorans than partial specimens of the humerus and radius.

While classification of specimens was relatively low for the LDA analyses, the k-

NN analyses were all over 50%. Thus, a k-NN would be a better statistical tool to classify an unknown specimen. One reason for this is that a k-NN analysis focuses on the individual specimens rather than the average within a species and therefore, allows for more variation within said species. Based on these results, if one wanted to classify an unknown (fossil) specimen, it would be better to look at the partial specimen rather than the complete to eliminate the noise. Overall, the distal humerus provided a higher rate of classification than the proximal or complete humerus. This pattern is seen again with the distal radius, which has a higher rate of classification than the proximal or complete. With the ulna, the proximal ulna provided a considerably higher rate of classification than the complete ulna and, in most cases, higher than the distal humerus or distal radius.

Therefore, the data show that partial specimens can be used to interpret family level, locomotion, and hunting strategy and that their morphospaces are relatively

distinct and clear in PCAs. It also shows how these analyses are phylogenetically significant and that phylogeny does play a role in carnivoran forelimb morphology. LDA are not as accurate as k-NN for classifying unknowns in carnivoran forelimbs and that by eliminating some noise and focusing on partial specimens may help provide a more accurate classification. This will give researchers more confidence in classifying partial

specimens from the fossil record.

A. . E. F.

Figure 3-1. Landmark placement of the humerus. A. anterior view. B. posterior view. C. proximal view. D. distal view. Landmark placement on template. E. anterior view. F. posterior view.

G. E.

H. A. B. C. D.

Figure 3-2. Landmark placement of the radius. A. anterior view. B. medial view. C. posterior view. D. lateral view. E. proximal view. F. distal view. Landmark placement on template. G. anterior view. H. distal view.

A. . C. D. E. F.

Figure 3-3. Landmark placement of the ulna. A. anterior view. B. medial view. C. posterior view. D. lateral view. Landmark placement on template. E. anterior view. F. posterior view.

A. . C. D.

F. G. H. I.

Figure 3-4. Landmark placement of the proximal humerus. A. anterior view. B. medial view. C. posterior view. D. lateral view. E. proximal view. Landmark placement on template. F. proximal view. G. anterior view. H. medial view. I. posterior view.

A. . D. E.

Figure 3-5. Landmark placement of the distal humerus. A. anterior view. B. posterior view. C. distal view. Landmark placement on template. D. anterior view. E. posterior view.

. C. D.

Figure 3-6. Landmark placement of the proximal radius. A. proximal view. B. anterior view. C. medial view. D. posterior view. Landmark placement on template. E. proximal view.

. C. D.

Figure 3-7. Landmark placement of the distal radius. A. distal view. B. anterior view. C. medial view. D. posterior view. Landmark placement on template. E. distal view.

A. . C.

E. F. G.

Figure 3-8. Landmark placement of the proximal ulna. A. medial view. B. anterior view. C. lateral view. D. proximal view. Landmark placement on template. E. medial view. F. anterior view. G. lateral view.

Figure 3-9. Character matrix for individual extant species.

Figure 3-9. Continued.

Figure 3-10. Carnivoran phylogeny of the extant 81 species included in dataset, modified from Nyakatura and Bininda- Emonds (2012).

Figure 3-11. Family phylomorphospaces for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

Figure 3-12. Family phylomorphospaces for the radius. A. complete radius. B. proximal radius. C. distal radius.

Figure 3-13. Family phylomorphospaces for the ulna. A. complete ulna. B. proximal ulna.

Figure 3-14. Locomotion PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

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Figure 3-15. Locomotion PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

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Figure 3-16. Locomotion PCAs for the ulna. A. complete ulna. B. proximal ulna.

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Figure 3-17. Hunting strategy PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

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Figure 3-18. Hunting strategy PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

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Figure 3-19. Hunting strategy PCAs for the ulna. A. complete ulna. B. proximal ulna.

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Table 3-1. Samuels et al., 2013 locomotor classification. Locomotion Description Rarely swims or climbs, may dig to make a burrow (but not Terrestrial extensively) (e.g., ermine, grisons). Regularly displays rapid locomotion with bounding characterized by Cursorial unsupported intervals (e.g., cheetah, gray wolf). Capable of climbing, usually for escape, does not forage in trees Scansorial (e.g., pumas, black bears). Capable of and regularly seen climbing for escape, shelter; forages actively in trees, and may rarely come down to the ground (e.g., red Arboreal panda, palm civets). Regularly digs to build burrows for shelter, may dig to forage Semifossorial underground (e.g., badgers). Regularly swims for dispersal, escape, or foraging (e.g., river otters, Semiaquatic American mink).

Table 3-2. Van Valkenburgh, 1985 hunting strategy classification. Hunting Strategy Description Short distance (<500 m) rush frequently preceded by a stalk (e.g. Ambush felids); forelimbs (i.e. grappling) often used in kill A moving search which ends in either a pounce or chase (e.g., Pounce/pursuit foxes); grappling with prey infrequent A chase, typically long distance (>500 km), rarely preceded by a Pursuit stalk (e.g., wolves); no grappling with prey Occasional Rarely hunts at all (e.g., most bears) Semifossorial Frequently digs for prey (e.g., badgers) Employs multiple hunting techniques such as pouncing in trees Various as well as diving (e.g., bearcat) Typically captures prey with mouth in aquatic environments (e.g., Aquatic otters) may consume inverts

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Table 3-3. LDA Rate of Reclassification: a) Discriminant Analysis. b) Cross Validation.

A. Complete Proximal Distal Complete Proximal Distal Complete Proximal Humerus Humerus Humerus Radius Radius Radius Ulna Ulna Family 100% 100% 100% 100% 100% 100% 100% 100% Locomotion 93% 98% 100% 90% 94% 94% 93% 98% Hunting 96% 100% 100% 95% 100% 99% 96% 99% Activity Pattern 90% 94% 89% 83% 88% 86% 90% 91% Diet 93% 94% 100% 91% 98% 93% 93% 88% Habitat 85% 96% 99% 91% 98% 94% 85% 100% Social Behavior 91% 98% 98% 94% 95% 98% 91% 91%

B. Complete Proximal Distal Complete Proximal Distal Complete Proximal Humerus Humerus Humerus Radius Radius Radius Ulna Ulna Family 89% 72% 81% 87% 67% 71% 89% 72% Locomotion 44% 38% 52% 48% 38% 38% 44% 32% Hunting 60% 42% 54% 65% 48% 53% 60% 60% Activity Pattern 47% 33% 28% 46% 25% 27% 47% 38% Diet 42% 49% 53% 54% 54% 43% 42% 33% Habitat 36% 21% 37% 37% 27% 32% 36% 35% Social Behavior 60% 57% 54% 57% 42% 47% 60% 47%

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Table 3-4. k-NN Rate of Reclassification. Complete Proximal Distal Complete Proximal Distal Complete Proximal Humerus Humerus Humerus Radius Radius Radius Ulna Ulna Family 86% 82% 90% 78% 77% 84% 77% 92% Locomotion 71% 57% 70% 61% 57% 64% 64% 71% Hunting 75% 69% 80% 71% 72% 76% 66% 83% Activity Pattern 61% 55% 62% 61% 59% 67% 56% 71% Diet 73% 64% 73% 65% 69% 71% 69% 79% Habitat 56% 53% 67% 52% 54% 59% 53% 62% Social Behavior 80% 73% 78% 73% 71% 76% 73% 81%

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CHAPTER 4 ECOMORPHOLOGY OF CAT-GAP CANIDS FROM THOMAS FARM FOSSIL SITE, (EARLY MIOCENE) FLORIDA

Introductory Remarks

Interpretations on fossil carnivoran behavior can be challenging due to incomplete specimens and small sample sizes. Chapter 3 demonstrated that partial specimens may provide more accurate classifications of behavior than complete ones. It is fairly common for paleontologists to interpret an extinct carnivoran’s behavior based on morphological similarities to modern taxa. These interpretations help to understand the evolutionary history of a species and to interpret changing community dynamics through time.

The forelimb can be a good indicator of ecomorphology because the morphology of its long bones has a strong functional signal (Van Valkenburgh, 1985, 1987, 1988;

Samuels and Van Valkenburgh, 2008; Walmsley et al., 2012; Samuels et al., 2013;

Fabre et al., 2013b; Fabre et al., 2015a). In particular, the well-studied carnivoran elbow joint complex has been shown to be indicative of locomotor and feeding strategies

(Gonyea, 1978; Andersson and Werdelin, 2003). It provides stability through ligaments and the humeroulnar and humeroradial articulations while maintaining mobility and transferring loads between the brachium and antebrachium (Andersson, 2004; Fabre et al., 2013a). The olecranon process of the ulna produces an outforce to the distal radius that allows for motion to be transferred down the arm; therefore, the direction or the angle at which the olecranon process is projected affects locomotory behavior

(Christiansen, 2002; Ercoli et al., 2012). Researchers have also shown that the carnivoran elbow joint complex has evolved in two directions: retention of their ability to supinate (to rotate forelimb so palm side is up) or loss of that ability and adaptation to

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cursoriality (Andersson and Werdelin, 2003). Fabre et al. (2015a) stressed the importance of studying the forelimb as a whole versus looking at each element independently. They demonstrated that the morphology of forearm long bones in musteloids is influenced by body mass, phylogeny, and lifestyle (Andersson, 2003;

Fabre et al., 2013a; Fabre et al., 2015b). Chapter 3 demonstrates that these characters apply to most extant carnivorans and therefore should be applicable in fossil carnivorans.

Materials and Methods

Materials

The humerus, radius, and ulna were scanned from 81 species of extant carnivorans, as well as fossil canids, felids, nimravids, and barbourofelids. The number of modern specimens from each species ranged from one to five individuals and included the same extant specimens as studied in Chapter 3 (Figure 3-9). When possible, both male and females were analyzed for each species. Specimens are from the Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA

(UF), East Tennessee State University, Johnson City, Tennessee, USA (ETSU), and the Museum National d’Histoire Naturelle, Paris, France (MNHN). Specimens were digitized using a Breuckmann 3D surface scanner, (camera resolution of 1.4 megapixels), a NextEngine 3D surface scanner or by NanoCT imaging. Only fossil specimens complete enough to be analyzed were chosen to be scanned. Fossil specimens are from UF, Museum of Comparative Zoology, Harvard University (MCZ),

American Museum of Natural History (AMNH), and were digitized by a NextEngine 3D surface scanner or by NanoCT imaging.

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Thomas Farm Canids

As mentioned in Chapter 1, three species of canids occur at TF: a hesperocyonine Osbornodon iamonensis and two borophagines Euoplocyon spissidens and Metatomarctus canavus (Tedford and Frailey, 1976; Wang, 1994). Osbornodon iamonensis is the largest and most abundant species of canid at TF; it and

Metatomarctus canavus are both index species for the He1 of the Hemingfordian

(Tedford et al., 2004). Euoplocyon spissidens was hypercarnivorous and was a primitive species of Euoplocyon (Wang et al., 1999).

Geometric Morphometrics

Methodology of the landmark placements and sub-selections follow the protocol outlined in Methodology in Chapter 3 and include the same coordinates. As described in

Chapter 3, incomplete elements can be used to determine some criteria such as family level, hunting strategy, and locomotion with relatively good accuracy. Therefore, it was deemed reasonable to examine distal humeri and proximal ulnae from Thomas Farm.

One difference in methodology from Chapter 3 is that the PCAs of Chapter 4 are on individuals within species instead of the mean morphology of a species.

Groupings and criteria. The modern specimens were split into three different groupings for ecomorphological comparisons. In the first grouping, the TF fossil specimens were compared against the 81extant species outlined in Chapter 3 as well as with an extinct nimravid (Hoplophoneus insolens and Hoplophoneus sp. (35-29 Mya), barbourofelid (Barbourofelis loveorum (~9.5 Ma)), felid (Nimravides galiani (~9.5 Ma)), and canids from before (Hesperocyon gregarious (~32 Ma), Mesocyon temnodon (~33-

23? Ma), Mesocyon coryphaeus (~32 Ma)), during (Osbornodon fricki (~15 Ma),

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Osbornodon iamonensis (21-18 Ma), and after the Cat-Gap (Tomarctus thompsoni

(~16-14 Ma)), Epicyon sp. (~9.5 Ma)) (see Appendix D).

The second set of analyses examined only extant felis, canids, mustelids, hyaenids, and procyonids for the humerus alone. This analysis was not run on the radius and ulna because there was not enough of a signal to consider procyonids further. The third set of analyses involved reducing the “noise” by examining only extant felids, canids, mustelids, and hyaenids (totaling 51 species) and was run on all three elements. This subset was chosen due to morphological similarities of the TF canids to extant canids, felids, and mustelids. The fourth set of analyses, also conducted on all three elements, reduced further “noise” by comparing the TF specimens to only felids and canids, to evaluate the hypothesis that TF specimens are more felid-like in their morphology even though they are identified as fossil canids.

Based on the research from Chapter 3, family-level, locomotion, and hunting strategy were considered to be the best criteria to examine the ecomorphology of fossil canids. Chapter 3 showed that these qualities can be classified relatively accurately through PCAs and k-NN analyses. k-NN Analysis and Predictions

Instead of using a Leave-one-out cross-validated linear discriminant analysis

(LDA), a k-nearest neighbors algorithm was used to determine the unknown classifications of the fossil elements. This was based on the comparison of the LDA to the k-NN analyses found in Chapter 3. The best k was determined for each element

(between 1 and 5) and each category analyzed (family, hunting strategy, and locomotion) based on accuracy of the classification and then reclassified to include the centroid size. Centroid size allows us to see if a more accurate classification is possible

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when taking the size into account. Through the k-NN analyses one can label specimens as unknowns and have software make predictions based on each individual unknown’s nearest neighbor.

Results

Like in most fossil sites, the Thomas Farm fossils are typically incomplete with a number of the fossil canid forelimb elements missing one epiphysis or the other.

Morphology of the TF distal humeri is similar to that of modern canids by having a deep olecranon fossa, a large radial fossa, and a sharp lateral edge to the capitulum. The TF humeri also have an entepicondylar foramen and lack a perforated radial/olecranon fossa similar to extant felids (Figure 4-1). The TF radii; the TF canid radii characteristics of the shape and level of medial inflection of the radial head, the prominence of the bicipital tuberosity, the curvature of the diaphysis, and the degree of mediolateral expansion of the distal end among others are in-between that of extant canids and felids

(Figure 4-2). The TF proximal ulnae have a triangular radial notch and a proximodistally expanded lunar notch as in extant canids (Figure 4-3). These morphological features are typically strong indicators of behavior. Because the TF fossil site falls within the Cat-

Gap, this research seeks to understand the behavior of the TF canids with the hypothesis that they may have occupied an open felid predator guild.

Family-Level PCAs

The first two axes of the first distal humerus PCA account for almost 43% of the total variation (Figure 4-4A). This PCA examines the 81 extant species plus all the fossil specimens including the nimravids, barbourofelids, fossil canids, and fossil felids. The extant canid, hyaenid, procyonid, mephitid and most of the extant felid, ursid, and mustelid morphospaces are well separated by the first axis. The second axis separates

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the hyaenid, ursid, and mephitid morphospaces as well as most of the extant felid and canid morphospaces. The barbourofelid surprisingly falls within the mustelid and mephitid morphospace, while the nimravid is directly within the felid morphospace. The fossil canids Epicyon, Tomarctos thompsoni, Osbornodon, and Hesperocyon gregarius are on the outer edge of the extant felid morphospace closer to the extant canids, whereas the pre-Cat-Gap canid Mesocyon falls within the ailurid and mustelid morphospace. The TF specimens fall within the extant felid morphospace and outside of the extant canid morphospace.

Figure 4-4B examines the distal humerus morphospaces of the extant felids, canids, mustelids, hyaenids, and procyonids. The first axis accounts for 28.5% of the total variation while the second axis accounts for just over 15%. The first axis separates the canid, hyaenid, and procyonid morphospaces as well as most of the extant felid and mustelid morphospaces. The second axis distinctly separates out the hyaenid and mustelid morphospaces as well as most of the morphospaces of the remaining extant families. The majority of TF canids fall within the extant felid morphospace with a few specimens falling in the morphospace between extant felids and canids with most being closest to extant felids.

The distal humerus morphospaces become more distinct when comparing just the extant felids, canids, mustelids, and hyaenids (Figure 4-4C). Most species within a family are well separated by both axes. The first axis covers ~28% of the total variation while the second axis has ~16% of the total variation. The hyaenid and canid morphospaces fall on the negative side of PC1 and are separated from each other via

PC2. The extant canid, Spethos venaticus falls within the morphospace of the hyaenids

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and outside that of modern canids. The felid and mustelid morphospaces overlap slightly but are mostly distinguished from each other via PC2. The TF canid specimens clearly fall on the edge of the felid morphospace, well away from the mustelid, hyaenid, and extant canid morphospaces.

The extant canids and felids have very distinct morphospaces that are well separated by the first axis (Figure 4-4D). The first two axes account for over 48% of the total variation across the felids, canids, and TF taxa. The second axis separates out all of the TF taxa while the edge of the TF morphospace overlaps with the edge of the extant felid morphospace; the extant canid morphospace is outside of both.

The first two axes of complete radius PCA with all 81 extant species and the fossil specimens account for 60.5% of the total variation (Figure 4-5A). The first PC separates most of the extant canids, hyaenids, and ursids. The second axis separates the mephitids and hyaenids as well as most of the TF taxa and felids. The nimravid falls within the outer limits of the felid morphospace that is overlapped by the mustelids and mephitids, whereas the barbourofelid morphospace falls within the mustelids and in close proximity to the ursids. The fossil canids Osbornodon and Mesocyon fall within the overlapping ursid and felid morphospace, whereas Hesperocyon gregarius and Epicyon are closer to the TF taxa. The TF specimens are mostly on the positive side of PC2 and fall within the overlapping sections of the felid, mephitid, mustelid, and procyonid morphospaces. All fossil canids and TF taxa are outside of the extant canid morphospace, with S. venaticus again being an outlier.

Figure 4-5B examines the complete radius with just the extant felids, canids, mustelids, hyaenids, and the TF carnivoran taxa. Almost 66% of the total variation is

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within the first two axes, much higher than in Figure 4-5A. The first axis separates out the hyaenids and the majority of the canids and mustelids, whereas the second axis separates the hyaenids and most felids and canids. There is some overlap of morphospaces in all the extant families. Mustelids have the largest morphospace due to the morphological variation in the radius for this family. The TF taxa fall outside of the hyaenid morphospace and on the edge of the mustelid morphospace. Some TF specimens fall within felid morphospace as well. If one were to exclude the extant canid

S. venaticus from the analysis, all the TF canid specimens would be outside of the extant canid morphospace.

When comparing the complete radius of TF taxa to exclusively extant canids and felids, the morphospace becomes more distinct (Figure 4-5C). The TF taxa are well separated by the second axis which accounts for ~19% of the total variation. As with

Figure 4-5B, excluding S. venaticus, the TF taxa would be outside of the extant canid morphospace. The extant canid and felid morphospaces overlap along the first axis, which accounts for over 50% of the total variation.

The proximal ulna PCA with all carnivorans shows over 40% of the total variation with the first two axes (Figure 4-6A). The first axis separates out the hyaenids, ursids, procyonids, and mephitids, as well as most of the mustelids. The second axis separates most of the mustelids, ursids, canids, felids, procyonids, viverrids, and mephitids. The ursid morphospace is the most distinct with most species falling on the negative side of axis one and the positive side of axis two. The felid morphospace is also relatively distinct and overlaps slightly with the viverrid and canid morphospaces. The barbourofelids and nimravid fall within the felid morphospace. The fossil canids,

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Meoscyon and Hesperocyon sp., also fall within the felid morphospace, whereas,

Epicyon and Tomarctos are within the extant canid morphospace. The TF specimens are more widely dispersed than in the radius and humerus PCAs. They can be separated into three groupings, one that falls clearly within extant felid morphospace, another within extant canid morphospace, and a third within the procyonid morphospace.

This same pattern is somewhat present in the analysis with only extant canids, felids, mustelids, and hyaenids (Figure 4-6B). Four individuals fall clearly within felid morphospace, a larger grouping within canid morphospace, and another cluster in the mustelid morphospace that also overlaps with extant canids and felids. The first two axes of this PCA accounts for ~48% of the total variation. The first axis separates the hyaenids and most of the felids and mustelids. The second axis separates most of the individuals in each of the families.

In Figure 4-6C, the canid and felid morphospaces are separated by the first axis, which accounts for 23.3% of the total variation. Three TF specimens fall squarely in the felid morphospace, while a majority falls in or near the extant canid morphospace.

Some TF specimens are outside both canid and felid morphospace, often falling between the two.

Locomotion PCAs

The locomotor habits of extant carnivorans is described in 6 categories (Chapter

3). When looking at the 81 species of carnivorans for the distal humerus, there is much overlap in the locomotor morphospaces (Figure 4-7A). The first axis separates the arboreal morphospace as well as most of the semifossorial, cursorial, and scansorial.

The second axis separates most of the semifossorial and semiaquatic locomotor

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morphospaces. The barbourofelid is within the overlapping morphospace of terrestrial, scansorial, and semifossorial locomotor abilities. The nimravid falls within the overlapping scansorial and terrestrial morphospace. The fossil canid Mesocyon lies within the arboreal morphospace, whereas Osbornodon is within the scansorial morphospace. Hesperocyon gregarius is on the outer edge of the cursorial morphospace, whereas the remaining fossil canids fall within the TF specimen morphospace, which are within the overlapping cursorial and scansorial morphospaces.

Figure 4-7B depicts the relationships of locomotor morphospace for extant felids, canids, mustelids, hyaenids, and procyonids. The cursorial morphospace is dominated by hyaenids and canids and therefore it is well separated by the first axis. The scansorial morphospace which is dominated by felids and the arboreal (dominated by procyonids) is also well separated by the first axis. The second axis separates most of the arboreal and semiquatic morphospaces. The TF canids fall in the overlapping scansorial, terrestrial, and cursorial morphospaces.

When simplifying the analysis to only include felids, canids, mustelids, and hyaenids, the morphospaces shift (Figure 4-7C). The first axis separates the arboreal and most of the scansorial, cursorial, and semifossorial. The second axis separates the arboreal and the majority of the semiaquatic, cursorial, and semifossorial. The TF specimens are all within the overlapping morphospace of the cursorial, scansorial, and terrestrial locomotor abilities, although they are closest to the scansorial felid species.

The locomotor morphospaces become more distinct with less locomotor categories in the analysis of extant felid and canid species than the previous two distal humeri analyses (Figure 4-7D). The first axis separates the arboreal, as well as most of

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the scansorial and cursorial species. The second axis separates arboreal and semifossorial as well as most of the cursorial and terrestrial species. The TF specimens fall within or slightly outside of the scansorial morphospace.

The locomotion PCA for the complete radius of 81 species of carnivorans shows more morphospatial overlap with each other than that of the distal humerus (Figure 4-

8A). The first axis separates most of the arboreal, cursorial, semiaquatic, and semifossorial morphospaces, whereas the second axis separates most of the cursorial species. The barbourofelids fall on the outer edge of the scansorial morphospace and within the semiaquatic (although it seems unlikely that they were semiaquatic), whereas the nimravid falls closest to a scansorial species. The fossil canids Mesocyon,

Tomarctos, and Osbornodon fall within the overlapping semifossorial, scansorial, and terrestrial morphospaces. Hesperocyon gregarius is on the outer limits of the terrestrial morphospace and falls within the overlapping scansorial and semifossorial, whereas the

Epicyon individuals fall closest to scansorial and cursorial individuals. The TF specimens have two outliers along the positive side of axis one; these individuals fall near the barbourofelids. The remaining TF specimens overlap with modern individuals that are scansorial, terrestrial, arboreal, or cursorial.

There is less overlap in the locomotor morphospaces of the complete radius when only looking at the mustelids, canids, felids, and hyaenids (Figure 4-8B). The first axis separates the semiaquatic, cursorial, and arboreal as well as most of the semifossorial locomotor habits. The second axis only separates most of the semifossorial species. One individual fossil TF carnivoran is an outlier from the others and is on the positive side of axis one and the negative side of axis two. This individual

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falls squarely in the felid morphospace and is within the arboreal, terrestrial, and scansorial morphospaces. The remaining TF carnivorans are on the positive side of axis two. Three individuals are in the semifossorial morphospace while half of the remaining are on the overlapping semifossorial and terrestrial morphospaces. The six remaining

TF carnivoran individuals are in the overlapping semifossorial, terrestrial, and scansorial morphospaces.

When comparing the complete radius TF specimens to only extant felids and canids, the first axis separates the arboreal, semifossorial, and cursorial as well as most of the terrestrial (Figure 4-8C). The second axis separates the arboreal and most of the scansorial, cursorial, terrestrial, and semifossorial species. The TF specimens fall within their own morphospace with only some overlap with the scansorial morphospace.

The first two axes of the PCA for the proximal ulna, including all 81 species, account for ~40% of the total variation (Figure 4-9A). The morphospace for scansorial is the most widespread. It is overlapped in part by each other form of locomotion. The first axis separates most of the arboreal, cursorial, and semifossorial morphospaces whereas the second axis does not. Nimravids and barbourofelids fall on the negative side of axis two, with barbourofelids being closest to a scansorial and a cursorial species and the nimravids being closest to a scansorial species. The fossil canids

Hesperocyon sp. and Mesocyon fall on the positive side of axis one and the negative side of axis two, both of which are closest to scansorial species. Epicyon, Tomarctos, and Osbornodon all fall on the positive side of axis two. Epicyon and Osbornodon fall closest to individuals that are defined as terrestrial whereas Tomarctos fall next to a

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scansorial species. The TF specimens are congregated near the center of the two axes and more often associate with individuals that are scansorial.

When looking at the proximal ulna and restricting the families to canids, hyaenids, mustelids, and felids, the morphospaces become easier to interpret (Figure 4-

9B). The first two axes account for ~39% of the total variation with the first axis separating most of the semifossorial and semiaquatic species and the second axis separating most of the arboreal, cursorial, and scansorial individuals. The TF specimens again fall near the center of the two axes. Most of the specimens are closest to either scansorial, cursorial, or terrestrial individuals.

In Figure 4-9C, the first axis separates the arboreal as well as most of the scansorial, terrestrial, and cursorial individuals whereas the second axis separates most of the cursorial individuals. The TF specimens fall in three different groups: one group is in the negative side of both axes and in the overlapping morphospace of scansorial and cursorial; the second group falls near the point of origin and within overlapping cursorial, scansorial, and terrestrial morphospaces; and the third group falls on the positive side of both axes but also within the overlapping cursorial, scansorial, and terrestrial morphospaces.

Hunting Strategy PCAs

The PCA for the distal humerus in extant species and fossil individuals does a relatively good job of separating out the different hunting strategy morphospaces

(Figure 4-10A). The first axis distinguishes out the pursuit predators as well as most of the semifossorial, occasional, aquatic, ambush, and pursuit and pounce categories. The second axis separates the pursuit and semifossorial as well as most of the aquatic, occasional, ambush, and pursuit and pounce. The nimravid and barbourofelids fall

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within the ambush predator morphospace. Epicyon, Osbornodon, Tomarctos, and

Hesperocyon gregarius fall within the ambush morphospace whereas Mesocyon is in the overlap of ambush and occasional hunting morphospace. The TF carnivorans are within the ambush morphospace dominated by the felids.

Figure 4-10B depicts the hunting strategy morphospaces for extant felids, canids, mustelids, hyaenids, and procyonids. The first axis separates out the pursuit, occoasional and most of the pursuit and pounce, ambush, and semifossorial morphospaces. The second axis separates the pursuit, semifossorial, and aquatic morphospaces as well as most of the occasional hunting individuals. Most of the TF canids fall within the ambush hunting morphospace with a few individuals falling inbetween the ambush and the pursuit and pounce morphospaces dominated by felids and canids respectively.

Looking at the hunting strategies of the distal humerus in just felids, canids, mustelids, and hyaenids, the first axis separates the pursuit predators as well as most of the ambush, semifossorial, aquatic, and pursuit and pounce (Figure 4-10C). The second axis separates the aquatic and semifossorial as well as most of the pursuit and pounce canids. The TF specimens clearly fall within the ambush predator morphospace.

When just looking at canid and felid distal humeri, the first axis separates all three hunting strategies (ambush, pursuit and pounce, and ambush), whereas the second axis separates only the pursuit predators (Figure 4-10D). Most of the TF carnivorans fall within their own morphospace that overlaps only slightly with the ambush predators (felids) and is well outside of the pursuit and pursuit and pounce morphospaces.

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The hunting strategy PCA for the complete radius in extant species and fossil individuals does a relatively good job of characterizing the different hunting strategy morphospaces (Figure 4-11A). The first axis separates the aquatic as well as most of the semifossorial and pursuit and pounce individuals. Semifossorial hunting strategy is well separated by the second axis. The barbourofelids fall within the aquatic morphospace and just outside of the ambush hunting morphospace, whereas the nimravid is within the ambush morphospace. The fossil canids Hesperocyon gregarius and Epicyon fall within the overlapping pursuit and pounce and semifossorial morphospace. Mesocyon, Osbornodon, and Tomarctos fall closest to ambush individuals. Most of the TF specimens are within the overlapping ambush and pursuit and pounce morphospaces.

Figure 4-11B is the PCA for the complete radius with just the mustelids, felids, canids, and hyaenids. The first axis separates semifossorial, aquatic, pursuit, and pursuit and pounce whereas the second axis separates most of the ambush and semifossorial individuals. The aquatic morphospace overlaps the edge of the semifossorial and ambush categories. The semifossorial morphospace overlaps mostly with the ambush morphospace. The TF specimens mostly fall within the ambush morphospace and the overlapping ambush and semifossorial morphospace.

When looking at only the felids and canids of the complete radius, the morphospaces are dominated by pursuit and pounce and ambush predators (Figure 4-

11C). The first axis separates the pursuit canids and most of the pursuit and pounce predators whereas the second axis separates most of the ambush predators. The TF specimens lie outside of the extant morphospace, within the pursuit and pounce

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morphospace, and in some cases within the overlapping pursuit and pounce and ambush morphospace.

The hunting PCA with all extant species and fossil taxa of the proximal ulna separates most of the semifossorial, aquatic, various, and pursuit predators across the first axis (Figure 4-12A). The second axis separates most of the ambush, various, pursuit, and occasional predators. The fossil felids, nimravids, and barbourofelids fall within the ambush hunting strategy morphospace. The fossil canids Mesocyon and

Hesperocyon sp. fall within the ambush morphospace whereas Epicyon, Osbornodon, and Tomarctos fall within the pursuit and pounce morphospace. The TF specimens seem to separate into three groups, one within the ambush morphospace, a second in the overlapping ambush and pursuit and pounce morphospace, and another within the pursuit and pounce morphospace.

The three groupings of TF specimens are also present in Figure 4-12B, the proximal ulna PCA of extant felids, canids, mustelids, and hyaenids. The first grouping of TF specimens falls within the ambush morphospace, the second in the overlapping ambush and pursuit and pounce morphospaces, and the third in the overlapping pursuit and pounce and semifossorial morphospaces. The first axis separates the pursuit, semifossorial, and most of the aquatic. The second axis separates most of the pursuit and the pursuit and pounce individuals.

The hunting strategies for the proximal ulna of extant felids and canids, have a very distinct separation of morphospaces (Figure 4-12C). The felids and canids and therefore the ambush, pursuit and pounce, and pursuit hunting strategies are well separated by the first axis. The second axis works to separate the pursuit predators.

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The TF specimens fall either in the ambush morphospace, the pursuit and pounce morphospace, or in between. k-NN and Predictions

All the predictions made in this section are based on the k-NN analyses with the best k-value (Appendix F). Tables of the best k-NN analyses for each category (distal humerus, complete radius, and proximal) and for each grouping (explained in methods) can be found in Appendix F. When looking at the distal humerus compared to the 81 extant species, the average prediction for the TF specimens was an scansorial procyonid that is an ambush hunter (Table 4-1). The distal humerus TF specimens’ predictions in the second grouping suggest a scansorial felid that digs for its prey whereas the third grouping of extant canids and felids generally predicts a scansorial felid with ambush hunting techniques.

The predictions for the complete radius are a little different (Table 4-2). When compared to the extant species, they are more likely to be classified as a terrestrial canid that has a pursuit and pounce hunting strategy. The grouping of modern canids, felids, mustelids, and hyaenids suggests a scansorial mustelid that also has a pursuit and pounce hunting strategy. When compared only to modern felids and canids, the analyses suggest a scansorial felid with ambush hunting strategy.

The predictions of the proximal ulna, when compared to extant species, were more variable across the categories than in the distal humerus or complete radius

(Table 4-3). The top two family-level predictions were felids and canids, whereas the analyses predicted a scansorial locomotion and an ambush hunting strategy. In the second grouping, the analyses predict a scansorial felid with an ambush hunting

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strategy. Nevertheless, when looking at just felids and canids, the analyses predicts a scansorial canid with ambush hunting strategy.

Discussion

One interesting finding of this study involves the locomotor and hunting strategy behavior of the non-Thomas Farm taxa (Table 4-4). Barbourofelis loveorum was predicted to be scansorial with an ambush hunting strategy, which is unsurprising given its short, well-muscled limbs. The nimravid Hoplophoneus was indicated to be more terrestrial and was also likely to be an ambush predator.

As predicted, the results of the PCAs indicate that the morphology of the forelimb of the TF canids typically falls within the felid morphospace or just outside of it in their own morphospace. The distal humeri PCAs suggest a more similar morphology to felids, with scansorial locomotion, and an ambush-like hunting strategy whereas the complete radii and proximal ulna PCAs suggest a morphospace between extant felids and canids. Such a locomotion would have the ability to be scansorial or semifossorial, and a hunting strategy that varied between ambush and pursuit and pounce.

The results of the k-NN analyses are very similar to that of the PCAs. The average distal TF humerus has a morphology resembling that in procyonids and felids, suggesting that they were scansorial and were most likely ambush hunters (Figure 4-

17). However, the average TF complete radius is more mustelid and felid-like, also characteristic of a scansorial locomotion, but differing in the possibility that they were pursuit and pounce predators. The average TF ulna suggests a scansorial ambush hunting as in a felid-like canid.

All these analyses suggest that the TF canids were most likely able to supinate their forelimbs, climb, and hunt using similar techniques to modern felids. The canids

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before the Cat-Gap varied in their niches. Mesocyon was more arboreal to scansorial, with an ambush hunting strategy, whereas Hesperocyon was more terrestrial to cursorial and was more likely a pursuit and pounce predator more similar to modern canids (Table 4-4). Non-TF Osbornodon, near the end of the Cat-Gap, was unsurprisingly similar to TF canids; they were most likely ambush hunters that were scansorial yet had a more cursorial ulna. Tomarctos and Epicyon were more terrestrial to cursorial but may have retained the ability to climb and preferred the pursuit and pounce hunting technique of modern canids.

Overall, my data suggest that prior to the Cat-Gap (> 21 Ma) canids such as

Mesocyon were more arboreal and would ambush their prey, whereas Hesperocyon was terrestrial and used the pursuit and pounced hunting technique. During the Cat-

Gap (~21 - 16 Ma), canids behaved more similar to modern felids by maintaining the ability to climb (scansorial) and ambush their prey. Just after the Cat-Gap (16 Ma), potentially related to the arrival of felids in North America, canids such as Tomarctos were less scansorial and more terrestrial, spent less time in the trees, and used a variety of techniques to hunt prey such as ambushing and pursuing. The extinct canids

Epicyon and modern canids tend to be more cursorial and pursue then pounce their prey. Therefore, we see a shift in locomotor and hunting strategy behavior of canids that corresponded with the presence vs. absence of nimravids, barbourofelids, and felids in

North America.

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Figure 4-1. Morphology of the distal humerus in a typical extant canid (Canis latrans 13412), extant felid (Puma concolor UF 25908), and an extinct TF canid (UF 1485).

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Figure 4-2. Morphology of the radius in a typical extant canid (Canis latrans, UF 13412), extant felid (Puma concolor, UF 25908), and an extinct TF canid (UF 60541).

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Figure 4-3. Morphology of the proximal ulna in a typical extant canid (Canis latrans, UF 13412), extant felid (Puma concolor, UF 25908), and an extinct TF canid (UF 267416).

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0 0

. 0

0 0

1 1

. 0

0.20 0.1 0.10 0.0 0.00 0.0 0.10 A. . 0.20 0.1 0. 10 0.0 0.00 0.0 0.10 PC1 2 .2% of Total ariance PC1 2 . % of Total ariance

0.1 0.10 0.0 0.00 0.0 0.10 0.1 0.10 0.0 0.00 0.0 0.10 C. PC1 2 . % of Total ariance D. PC1 3 . % of Total ariance

Thomas Farm Canid Thomas Farm Fossil Canidae iverridae Fossil Felidae Mephitidae Canidae Ursidae Felidae Ailuridae Mustelidae Eupleridae Hyaenidae arbourofelidae Procyonidae Nimravidae

Figure 4-4. Family PCAs for the distal humerus. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, procyonids, and TF fossils. C. extant mustelids, hyaenids, felids, canids, and TF fossils. D. extant felids, canids, and TF fossils.

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A. 0.10 0.0 0.00 0.0 0.10 0.1 . 0.10 0.0 0.00 0.0 0.10 0.1 PC1 .1% of Total ariance PC1 1. % of Total ariance

Thomas Farm Fossil Canidae Fossil Felidae Canidae Felidae Mustelidae Hyaenidae Procyonidae iverridae Mephitidae Ursidae Ailuridae Eupleridae arbourofelidae 0.0 0.00 0.0 Nimravidae C. PC1 0.6% of Total ariance

Figure 4-5. Family PCAs for the radius. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Figure 4-6. Family PCAs for the proximal ulna. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Figure 4-7. Locomotion PCAs for the distal humerus. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, procyonids, and TF fossils. C. extant mustelids, hyaenids, felids, canids, and TF fossils. D. extant felids, canids, and TF fossils.

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Figure 4-8. Locomotion PCAs for the radius. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Figure 4-9. Locomotion PCAs for the proximal ulna. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Figure 4-10. Hunting strategy PCAs for the distal humerus. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, procyonids, and TF fossils. C. extant mustelids, hyaenids, felids, canids, and TF fossils. D. extant felids, canids, and TF fossils.

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Figure 4-11. Hunting strategy PCAs for the radius. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Figure 4-12. Family PCAs for the proximal ulna. A. 81 extant species and fossil taxa. B. extant mustelids, hyaenids, felids, canids, and TF fossils. C. extant felids, canids, and TF fossils.

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Humerus All HumerusTFcafemusthypro HumerusTFcafemusthy Humerus Tfcafe Family Locomotion Hunting Family LocomotionHunting Family Locomotion Hunting Family Locomotion Hunting MC HA 20 Procyonidae Scansorial Ambush Mustelidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Semiaquatic Ambush MC HC 20 Procyonidae Terrestrial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush MC HD 20 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Terrestrial Ambush UF001 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Terrestrial Ambush UF01 6 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Scansorial Ambush UF01 66 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF01 66 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF01 3 Procyonidae Scansorial Ambush Mustelidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Scansorial Ambush UF0606 Procyonidae Terrestrial Occasional Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF0 3 20 Procyonidae Terrestrial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF0 3 21 Procyonidae Terrestrial Ambush Felidae Arboreal Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF0 3 36 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF0 3 3 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF0 3 3 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF1 1 Procyonidae Terrestrial Ambush Procyonidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF1 1 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF16 1 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF16 3 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF1 26 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Terrestrial Ambush UF21 Procyonidae Terrestrial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF21 3 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2 1 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2 3 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2 Procyonidae Scansorial Ambush Felidae Terrestrial Semifossorial Felidae Terrestrial Semifossorial Felidae Terrestrial Ambush UF261133 Procyonidae Terrestrial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF262 2 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2660 2 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF26 2 6 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF26 1 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF26 3 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Terrestrial Semifossorial Felidae Terrestrial Ambush UF2 221 Procyonidae Terrestrial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2 2 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF2 0 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF31 16 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush UF 01 6 Procyonidae Scansorial Ambush Felidae Scansorial Semifossorial Felidae Scansorial Semifossorial Felidae Scansorial Ambush

Figure 4-13. Thomas Farm humeri predictions.

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Figure 4-14. Thomas Farm radii predictions.

141

Figure 4-15. Thomas Farm ulnae predictions.

142

Figure 4-16. Non-Thomas Farm fossil predictions.

143

Figure 4-17. Summary figure of the most common characteristic of TF canids.

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CHAPTER 5 CONCLUSIONS

The Thomas Farm fossil locality in Florida is world renowned. Specimens have been studied here since the 1930s and the carnivorans have been no exception.

Nevertheless, with close examination of existing vertebrate paleontology collections and ongoing fieldwork, new Early Miocene taxa are still being discovered in this sinkhole deposit.

The identification of procyonids from Thomas Farm has provided us with new insight to their diversity and distribution during a time of great importance in their North

American diversification. At this time, it is difficult to determine if the first procyonid taxa from Thomas Farm belongs to the living genus Bassariscus or the extinct

Probassariscus. These genera are very similar, with few synapomorphies to distinguish them. The specimens in hand have slightly more characteristics in common with

Probassariscus than Bassariscus and therefore, are designated as cf. Probassariscus.

With more material, the Thomas Farm bassariscine procyonid may prove to be a more derived Probassariscus or the oldest known Bassariscus species. Future examination of procyonid postcranial material may also provide more synapomorphies for distinguishing these genera. A second, slightly larger procyonid taxa also occurs at

Thomas Farm. This procyonid is believed to be a potosine but more material needs to be recovered and identified.

Postcranial elements are found frequently at Thomas Farm, providing opportunities to look at ecomorphology of early Hemingfordian carnivorans. The material collected, as at most fossil localities, consists mainly of incomplete elements.

Typically, the ends of long bones tend to survive the taphonomic process more

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frequently, although most 3D morphometric research is done on complete modern specimens.

Chapter 3 demonstrates the practicality of using the distal and proximal ends of the humerus and radius, and the proximal end of the ulna, in leu of complete elements in 3D morphometric analyses. This is highly beneficial to paleontologists, who often are limited by sample size. By focusing on the weight bearing aspects of forelimbs and allowing for partial specimens to be scanned, the sample sizes can be greatly increased and a fuller range of morphological diversity can be analyzed.

Chapter 3 also discusses the viability of distinguishing phylogenetic and behavioral characteristics of species based on these forelimb elements. Principal component analyses are a strong statistical method for making morphological interpretations on the family-level differences as well as behavioral differences. PCAs coupled with k-NN analyses can help to make well supported interpretations of the behavior of extinct species. The research shows that the distal humerus, distal radius, and the proximal ulna in most cases provide higher accuracy in classifying locomotion, hunting strategy, and family-level than by looking at the complete elements.

Therefore, Chapter 4 uses PCAs and k-NN analyses on partial fossil specimens, to interpret Cat-Gap (~21 - 16 Ma) canid taxa from the Thomas Farm fossil site. 3D scans of the distal humerus, complete radius, and proximal ulna of TF canids were compared to those of 81 extant carnivoran species to evaluate extant family-level designations, locomotor behavior, and hunting style. Based on these analyses, the typical TF canid was very felid-like, was scansorial, and most likely hunted by ambush, all characteristics similar to those of average extant felid species. These canids would

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have occupied the open felid predator guild of Florida during 5 million years when no felids, nimravids, or barbourofelids lived in North America.

Future directions for this research would include adding more canid species before and after the Cat-Gap to get a fuller understanding of the evolution of canid morphology across time and through lineages. For wider geographical approach, canids across North America should also be included. In conjunction with this, a better understanding of the musculature of canids can provide a stronger framework for making behavioral interpretations based on morphology.

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APPENDIX A UPPER P4 MEASUREMENTS

APL APL TW TW Species Catalogue # Side Sex (mm) NatLog (mm) NatLog Potos flavus UF 20619 R 4.31 1.46 6.3 1.84 Unknown UF 20619 L 4.35 1.47 6.25 1.83 Unknown UF 20620 R 4.32 1.46 5.8 1.76 Unknown UF 20620 L 4.17 1.43 5.94 1.78 Unknown Nasua narica UF 6858 R 8.01 2.08 7.64 2.03 Male UF 6858 L 7.58 2.03 7.22 1.98 Male UF 12303 R 7 1.95 7.38 2 Unknown UF 33184 R 6.75 1.91 6.26 1.83 Female UF 33184 L 6.97 1.94 7.01 1.95 Female UF 33185 R 6.74 1.91 5.61 1.72 Male UF 33185 L 7.51 2.02 6.12 1.81 Male UF 33190 R 7.02 1.95 7.22 1.98 Female UF 33190 L 7.28 1.99 6.83 1.92 Female Procyon UF 13457 R 12.41 2.52 12.71 2.54 Unknown cancrivorus UF 13457 L 12.55 2.53 12.64 2.54 Unknown UF 33182 R 9.71 2.27 11.4 2.43 Male UF 33182 L 9.67 2.27 11.39 2.43 Male UF 40 R 8.39 2.13 8.35 2.12 Male Procyon lotor UF 40 L 8.28 2.11 8.34 2.12 Male UF 361 R 7.69 2.04 7.7 2.04 Male UF 361 L 7.71 2.04 7.71 2.04 Male UF 362 R 7.53 2.02 7.77 2.05 Female UF 362 L 7.5 2.01 7.54 2.02 Female UF 363 R 8.71 2.16 8.05 2.09 Male UF 363 L 8.05 2.09 8.33 2.12 Male UF 364 R 8.02 2.08 8.23 2.11 Male UF 364 L 8.1 2.09 8.29 2.12 Male UF 367 R 7.69 2.04 8.28 2.11 Unknown UF 367 L 7.6 2.03 7.9 2.07 Unknown UF 368 R 9.45 2.25 7.58 2.03 Unknown UF 372 R 8.14 2.1 8.13 2.1 Male UF 372 L 8.21 2.11 8.14 2.1 Male UF 602 R 7.77 2.05 7.68 2.04 Male UF 602 L 7.77 2.05 7.67 2.04 Male UF 605 R 7.82 2.06 7.99 2.08 Female UF 605 L 7.79 2.05 8.05 2.09 Female UF 606 R 8.02 2.08 7.55 2.02 Male UF 606 L 8.05 2.09 7.56 2.02 Male

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UF 607 R 7.55 2.02 7.86 2.06 Male UF 607 L 7.59 2.03 7.84 2.06 Male Procyon lotor UF 665 R 6.6 1.89 7.23 1.98 Male UF 665 L 6.49 1.87 7.17 1.97 Male UF 666 R 7.31 1.99 7.62 2.03 Male UF 666 L 7.35 1.99 7.67 2.04 Male UF 836 R 8.23 2.11 7.89 2.07 Male UF 836 L 8.08 2.09 8.01 2.08 Male UF 837 R 7.8 2.05 7.97 2.08 Male UF 837 L 8 2.08 8.03 2.08 Male UF 838 R 8.2 2.1 7.83 2.06 Female UF 838 L 8.06 2.09 7.99 2.08 Female UF 841 L 7.66 2.04 7.39 2 Unknown UF 842 R 7.96 2.07 7.77 2.05 Male UF 843 L 7.62 2.03 7.66 2.04 Unknown UF 1005 R 7.44 2.01 7.79 2.05 Female UF 1005 L 7.61 2.03 7.89 2.07 Female UF 1006 L 7.38 2 7.33 1.99 Male UF 1007 R 7.67 2.04 6.98 1.94 Male UF 1007 L 7.34 1.99 6.71 1.9 Male UF 1049 R 8.18 2.1 7.82 2.06 Male UF 1049 L 8.12 2.09 7.93 2.07 Male UF 1069 R 7.01 1.95 7.26 1.98 Male UF 1107 R 8.2 2.1 7.52 2.02 Female UF 1107 L 8.34 2.12 7.65 2.03 Female UF 1216 R 7.15 1.97 7.4 2 Male UF 1216 L 7.02 1.95 7.3 1.99 Male UF 1231 R 7.9 2.07 8.02 2.08 Female UF 1231 L 8.05 2.09 8.04 2.08 Female UF 1409 R 8.17 2.1 8.18 2.1 Unknown UF 1409 L 8.1 2.09 8.14 2.1 Unknown UF 1592 R 7.28 1.99 7.51 2.02 Male UF 1592 L 7.3 1.99 7.41 2 Male UF 1593 L 6.25 1.83 7.23 1.98 Male UF 1594 R 7.26 1.98 7.64 2.03 Male UF 1594 L 7.3 1.99 7.5 2.01 Male UF 1596 R 7.95 2.07 7.56 2.02 Female UF 1596 L 7.96 2.07 7.6 2.03 Female UF 1597 R 7.74 2.05 7.73 2.05 Male UF 1597 L 7.81 2.06 7.69 2.04 Male UF 1598 R 8.5 2.14 8.45 2.13 Male UF 1598 L 8.54 2.14 8.39 2.13 Male UF 1637 R 7.86 2.06 8.06 2.09 Unknown

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UF 1637 L 7.96 2.07 7.89 2.07 Unknown UF 1875 R 7.66 2.04 8.07 2.09 Female UF 1875 L 7.62 2.03 7.9 2.07 Female UF 1914 R 6.91 1.93 7.25 1.98 Male Procyon lotor UF 1914 L 7.01 1.95 7.19 1.97 Male UF 1915 R 8.57 2.15 8.63 2.16 Male UF 1915 L 8.29 2.12 8.19 2.1 Male UF 1916 R 8.67 2.16 8.04 2.08 Male UF 1916 L 8.08 2.09 7.94 2.07 Male UF 1917 R 8.11 2.09 7.95 2.07 Male UF 1917 L 8.2 2.1 7.9 2.07 Male UF 1940 R 7.59 2.03 7.66 2.04 Female UF 1940 L 7.55 2.02 7.62 2.03 Female UF 2103 R 8.09 2.09 7.84 2.06 Unknown UF 2103 L 8.17 2.1 7.87 2.06 Unknown UF 2105 R 7.58 2.03 7.77 2.05 Unknown UF 2105 L 7.58 2.03 7.65 2.03 Unknown UF 2107 R 8.3 2.12 8.08 2.09 Unknown UF 2107 L 8.44 2.13 8.02 2.08 Unknown UF 2108 R 8.18 2.1 7.84 2.06 Unknown UF 2108 L 7.76 2.05 7.88 2.06 Unknown UF 3836 L 7.33 1.99 7.42 2 Female UF 3977 R 8.22 2.11 8.36 2.12 Female UF 3977 L 7.95 2.07 8.28 2.11 Female UF 3978 R 7.68 2.04 7.86 2.06 Male UF 3978 L 7.76 2.05 7.9 2.07 Male UF 3979 R 7.39 2 7.52 2.02 Unknown UF 3979 L 7.49 2.01 7.44 2.01 Unknown UF 3980 R 7.72 2.04 7.7 2.04 Unknown UF 3980 L 7.68 2.04 7.72 2.04 Unknown UF 4547 L 7.49 2.01 7.66 2.04 Male UF 4571 R 7.5 2.01 7.48 2.01 Unknown UF 4571 L 7.47 2.01 7.41 2 Unknown UF 4572 R 8.79 2.17 8.33 2.12 Unknown UF 4572 L 8.37 2.12 8.19 2.1 Unknown UF 4591 R 8.05 2.09 7.71 2.04 Male UF 4591 L 8.12 2.09 7.61 2.03 Male UF 4609 R 7.9 2.07 8.54 2.14 Unknown UF 4609 L 8.11 2.09 8.55 2.15 Unknown UF 4610 R 7.73 2.05 7.5 2.01 Unknown UF 4610 L 7.74 2.05 7.49 2.01 Unknown UF 4614 R 8.5 2.14 8.4 2.13 Unknown UF 4615 R 7.8 2.05 7.78 2.05 Unknown

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UF 4615 L 8.06 2.09 7.75 2.05 Unknown UF 4695 R 8.16 2.1 7.85 2.06 Male UF 4695 L 7.94 2.07 7.46 2.01 Male UF 4696 R 7.83 2.06 8.28 2.11 Unknown UF 4696 L 8.04 2.08 8.08 2.09 Unknown Procyon lotor UF 4925 R 7.67 2.04 7.86 2.06 Male UF 4925 L 7.35 1.99 7.49 2.01 Male UF 4926 R 7.95 2.07 7.64 2.03 Male UF 4926 L 8.01 2.08 7.53 2.02 Male UF 5106 L 8.04 2.08 8.21 2.11 Unknown UF 5584 R 7.79 2.05 8 2.08 Female UF 5584 L 8.35 2.12 8.03 2.08 Female UF 5681 R 8.44 2.13 8.49 2.14 Unknown UF 5681 L 8.53 2.14 8.23 2.11 Unknown UF 5826 R 7.95 2.07 8.41 2.13 Male UF 5826 L 8.19 2.1 8.76 2.17 Male UF 5828 L 7.72 2.04 7.26 1.98 Unknown UF 5829 L 8.09 2.09 8.11 2.09 Male UF 5830 L 7.36 2 7.34 1.99 Female UF 5867 R 9.16 2.21 8.55 2.15 Unknown UF 5867 L 9.05 2.2 8.57 2.15 Unknown UF 5937 L 7.81 2.06 7.32 1.99 Male UF 6653 R 7.85 2.06 7.72 2.04 Female UF 6653 L 7.56 2.02 7.7 2.04 Female UF 6677 R 7.58 2.03 7.4 2 Female UF 6677 L 7.58 2.03 7.72 2.04 Female UF 6731 R 7.66 2.04 7.65 2.03 Unknown UF 6731 L 7.59 2.03 7.97 2.08 Unknown UF 6732 R 8.01 2.08 7.86 2.06 Unknown UF 6732 L 8.06 2.09 7.46 2.01 Unknown UF 6733 L 7.25 1.98 7.39 2 Female UF 7167 L 7.51 2.02 7.15 1.97 Unknown UF 7169 R 8.23 2.11 7.74 2.05 Unknown UF 7169 L 8.11 2.09 7.66 2.04 Unknown UF 7751 R 7.58 2.03 7.05 1.95 Unknown UF 7751 L 7.6 2.03 6.93 1.94 Unknown UF 7850 R 7.4 2 7.38 2 Unknown UF 7850 L 7.11 1.96 7.28 1.99 Unknown UF 7851 R 7.19 1.97 7.46 2.01 Unknown UF 8072 R 8.9 2.19 8.92 2.19 Unknown UF 8072 L 8.93 2.19 8.92 2.19 Unknown UF 8073 R 7.67 2.04 7.8 2.05 Unknown UF 8073 L 7.63 2.03 7.76 2.05 Unknown

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UF 8074 R 8.93 2.19 8.98 2.19 Unknown UF 8074 L 9.11 2.21 9.01 2.2 Unknown UF 8075 L 7.63 2.03 7.37 2 Unknown UF 8077 R 7.74 2.05 8.38 2.13 Unknown UF 8077 L 7.96 2.07 8.23 2.11 Unknown UF 8078 R 8.77 2.17 8.62 2.15 Unknown UF 8078 L 8.39 2.13 8.64 2.16 Unknown Procyon lotor UF 8124 R 7.61 2.03 6.79 1.92 Unknown UF 9205 L 8.07 2.09 7.91 2.07 Male UF 9225 L 7.97 2.08 7.78 2.05 Female UF 9867 R 8.05 2.09 7.46 2.01 Unknown UF 9990 R 7.8 2.05 7.62 2.03 Unknown UF 9990 L 7.81 2.06 7.62 2.03 Unknown UF 9998 L 7.75 2.05 7.39 2 Unknown UF 10008 R 7.87 2.06 7.96 2.07 Male UF 10008 L 7.89 2.07 8.09 2.09 Male UF 10089 R 8.04 2.08 8.01 2.08 Male UF 10089 L 8.11 2.09 7.93 2.07 Male UF 10095 L 7.56 2.02 7.46 2.01 Male UF 10098 L 7.68 2.04 7.4 2 Male UF 10125 R 8.5 2.14 8.26 2.11 Male UF 10125 L 8.49 2.14 8.17 2.1 Male UF 10400 L 7.42 2 7.73 2.05 Unknown UF 11947 R 8.36 2.12 8.38 2.13 Unknown UF 11947 L 8.49 2.14 8.57 2.15 Unknown UF 12349 R 8.03 2.08 7.48 2.01 Unknown UF 12349 L 7.88 2.06 7.42 2 Unknown UF 12509 L 8.06 2.09 8.02 2.08 Male UF 12651 R 7.77 2.05 7.69 2.04 Unknown UF 12651 L 7.76 2.05 7.95 2.07 Unknown UF 12655 R 7.33 1.99 7.78 2.05 Female UF 12655 L 7.38 2 7.83 2.06 Female UF 12657 R 8.32 2.12 7.62 2.03 Unknown UF 12657 L 8.06 2.09 7.62 2.03 Unknown UF 12800 R 6.96 1.94 7.37 2 Male UF 12800 L 7.13 1.96 7.37 2 Male UF 13259 R 7.99 2.08 7.57 2.02 Unknown UF 13259 L 7.96 2.07 7.48 2.01 Unknown UF 13282 R 8.31 2.12 7.87 2.06 Male UF 13282 L 8.2 2.1 7.76 2.05 Male UF 14663 R 7.57 2.02 7.5 2.01 Unknown UF 14663 L 7.65 2.03 7.59 2.03 Unknown UF 14680 R 7.76 2.05 8.2 2.1 Unknown

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UF 14680 L 7.73 2.05 8.03 2.08 Unknown UF 20618 R 4.25 1.45 5.34 1.68 Unknown UF 20618 L 4.28 1.45 5.82 1.76 Unknown UF 21685 R 7.97 2.08 7.6 2.03 Unknown UF 21685 L 8.09 2.09 7.54 2.02 Unknown UF 21706 R 7.84 2.06 7.57 2.02 Male UF 21706 L 7.89 2.07 7.57 2.02 Male UF 22129 R 7.04 1.95 7.21 1.98 Female UF 22129 L 7.25 1.98 7.23 1.98 Female Procyon lotor UF 22130 R 7.66 2.04 7.8 2.05 Male UF 22130 L 7.66 2.04 8.01 2.08 Male UF 22420 R 7.71 2.04 8.15 2.1 Male UF 22420 L 9.34 2.23 9.36 2.24 Male UF 22569 L 8.28 2.11 7.81 2.06 Male UF 24012 R 7.59 2.03 7.73 2.05 Male UF 24012 L 7.44 2.01 7.54 2.02 Male UF 25921 R 7.87 2.06 8.19 2.1 Male UF 25921 L 7.8 2.05 8.28 2.11 Male UF 26007 R 7.34 1.99 7.21 1.98 Male UF 26007 L 7.19 1.97 7.21 1.98 Male UF 26010 R 8.34 2.12 7.75 2.05 Unknown UF 26010 L 8.34 2.12 7.73 2.05 Unknown UF 26011 R 7.65 2.03 7.52 2.02 Unknown UF 26011 L 7.77 2.05 7.59 2.03 Unknown UF 29873 R 7 1.95 7.18 1.97 Unknown UF 29873 L 7.05 1.95 7.03 1.95 Unknown UF 32659 L 7.6 2.03 7.73 2.05 Unknown UF 33183 R 8.53 2.14 8.42 2.13 Male UF 33183 L 8.86 2.18 8.12 2.09 Male UF 33546 L 7.67 2.04 8.21 2.11 Male Bassariscus UF 7192 R 7.57 2.02 5.6 1.72 Male astutus UF 7192 L 7.8 2.05 5.59 1.72 Male UF 7257 R 7.36 2 4.94 1.6 Unknown UF 7257 L 7.55 2.02 4.84 1.58 Unknown UF 7865 R 7.02 1.95 4.51 1.51 Female UF 7865 L 6.93 1.94 4.71 1.55 Female UF 8474 R 7.41 2 5.05 1.62 Unknown UF 8474 L 7.34 1.99 4.98 1.61 Unknown UF 11933 R 7.44 2.01 5.11 1.63 Unknown UF 11933 L 6.01 1.79 7.8 2.05 Unknown UF 13741 R 6.95 1.94 4.39 1.48 Female UF 13741 L 6.99 1.94 4.55 1.52 Female Thomas Farm UF 163893 R 6.59 1.89 5.5 1.7 Unknown

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UF 163894 R 6.29 1.84 4.93 1.6 Unknown Thomas Farm UF 406541 R 6.73 1.91 5.45 1.7 Unknown UF 406542 L 6.09 1.81 5.05 1.62 Unknown UF 406543 L 6.5 1.87 4.6 1.53 Unknown Phlaocyon (Notocyon) CM 1602 L 8.59 2.15 4.84 1.58 Unknown sp. Phlaocyon CM 11332 L 7.66 2.04 5.6 1.72 Unknown (Brookesville) UF 163533 R 5.72 1.74 3.06 1.12 Unknown Phlaocyon UF 163534 R 5.86 1.77 3.71 1.31 Unknown willistoni Leptarctus cast MCZ L 10.42 2.34 7.62 2.03 Unknown ancipidens 3658 MCZ 6223 L 10.88 2.39 7.7 2.04 Unknown Zodiolestes UF 206245 R 15.96 2.77 11.12 2.41 Unknown Parictis sp. MCZ 8441 L 8.02 2.08 5.85 1.77 Unknown

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APPENDIX B LOWER m1 MEASUREMENTS

Catalogue APL APL PMW PMW Species Side Sex # (mm) NatLog (mm) NatLog Potos flavus UF 20618 R 5.4 1.69 3.95 1.37 Unknown UF 20618 L 5.44 1.69 3.92 1.37 Unknown UF 20619 R 5.59 1.72 3.97 1.38 Unknown UF 20619 L 5.4 1.69 4.02 1.39 Unknown UF 20620 R 5.47 1.7 4.24 1.44 Unknown UF 20620 L 5.41 1.69 4.12 1.42 Unknown Nasua narica UF 6858 R 8.62 2.15 5.09 1.63 Male UF 6858 L 8.55 2.15 5.34 1.68 Male UF 33184 R 8.46 2.14 5.35 1.68 Female UF 33184 L 8.5 2.14 5.25 1.66 Female UF 33185 R 8.06 2.09 4.97 1.6 Male UF 33189 R 8.77 2.17 5.86 1.77 Unknown UF 33189 L 9.22 2.22 5.9 1.77 Unknown UF 33192 R 7.89 2.07 5.29 1.67 Female UF 33192 L 7.89 2.07 5.29 1.67 Female Procyon UF 13457 R 13.29 2.59 9.78 2.28 Unknown cancrivorus UF 13457 L 12.81 2.55 9.65 2.27 Unknown UF 33182 R 12.87 2.55 8.62 2.15 Male UF 33182 L 13.12 2.57 8.65 2.16 Male Procyon lotor UF 37 R 8.45 2.13 5.86 1.77 Female UF 37 L 5.52 1.71 5.86 1.77 Female UF 40 R 10.33 2.34 6.44 1.86 Male UF 40 L 10.36 2.34 6.41 1.86 Male UF 361 R 9.13 2.21 6.09 1.81 Male UF 361 L 9.03 2.2 6.03 1.8 Male UF 362 R 9.49 2.25 6.43 1.86 Female UF 362 L 9.62 2.26 6.26 1.83 Female UF 363 R 10.03 2.31 6.44 1.86 Male UF 363 L 9.88 2.29 6.3 1.84 Male UF 364 R 9.72 2.27 6.62 1.89 Male UF 364 L 9.8 2.28 6.09 1.81 Male UF 370 R 9.82 2.28 5.78 1.75 Unknown UF 370 L 9.8 2.28 5.89 1.77 Unknown UF 371 R 9.65 2.27 6.24 1.83 Unknown UF 602 R 9.99 2.3 6.15 1.82 Male UF 602 L 9.87 2.29 6.59 1.89 Male UF 603 R 9.25 2.22 6.01 1.79 Male UF 603 L 9.28 2.23 5.79 1.76 Male UF 604 R 9.34 2.23 6.13 1.81 Male

155

UF 604 L 9.37 2.24 6.27 1.84 Male Procyon lotor UF 605 R 10.28 2.33 6.7 1.9 Female UF 606 R 9.85 2.29 6.22 1.83 Male UF 606 L 9.83 2.29 6.04 1.8 Male UF 607 R 9.55 2.26 6.25 1.83 Male UF 607 L 9.6 2.26 6.22 1.83 Male UF 665 R 8.58 2.15 5.86 1.77 Male UF 665 L 8.72 2.17 5.83 1.76 Male UF 666 R 9.22 2.22 6.08 1.81 Male UF 667 R 9.34 2.23 6.49 1.87 Female UF 667 L 9.46 2.25 6.51 1.87 Female UF 836 L 10.27 2.33 6.44 1.86 Male UF 837 R 9.98 2.3 6.3 1.84 Male UF 837 L 9.87 2.29 6.13 1.81 Male UF 838 R 10.13 2.32 6.7 1.9 Female UF 838 L 10.17 2.32 6.68 1.9 Female UF 841 R 9.81 2.28 6.08 1.81 Unknown UF 841 L 9.89 2.29 6.15 1.82 Unknown UF 842 R 9.29 2.23 6.22 1.83 Male UF 842 L 9.41 2.24 6.35 1.85 Male UF 844 R 9.78 2.28 6.6 1.89 Unknown UF 844 L 9.81 2.28 6.68 1.9 Unknown UF 1005 R 10.1 2.31 6.41 1.86 Female UF 1005 L 9.54 2.26 6.31 1.84 Female UF 1006 R 9.48 2.25 5.61 1.72 Male UF 1006 L 9.36 2.24 5.83 1.76 Male UF 1007 R 9.05 2.2 5.14 1.64 Male UF 1007 L 8.84 2.18 5.12 1.63 Male UF 1049 R 9.61 2.26 6.05 1.8 Male UF 1049 L 9.61 2.26 6.03 1.8 Male UF 1069 R 9.46 2.25 5.85 1.77 Male UF 1069 L 9.58 2.26 5.82 1.76 Male UF 1107 R 9.68 2.27 6.38 1.85 Female UF 1107 L 9.63 2.26 6.2 1.82 Female UF 1216 R 9.42 2.24 5.95 1.78 Male UF 1216 L 9.58 2.26 5.77 1.75 Male UF 1231 R 10.75 2.37 6.74 1.91 Female UF 1231 L 10.65 2.37 6.62 1.89 Female UF 1592 R 9.31 2.23 5.95 1.78 Male UF 1592 L 9.36 2.24 5.9 1.77 Male UF 1593 R 8.99 2.2 5.84 1.76 Male UF 1593 L 8.67 2.16 5.77 1.75 Male UF 1594 R 9.32 2.23 5.83 1.76 Male UF 1594 L 9.34 2.23 5.88 1.77 Male UF 1596 R 9.67 2.27 6.03 1.8 Female

156

UF 1596 L 9.84 2.29 5.96 1.79 Female Procyon lotor UF 1597 R 9.73 2.28 6.33 1.85 Male UF 1597 L 9.33 2.23 6.4 1.86 Male UF 1598 R 10.46 2.35 6.86 1.93 Male UF 1598 L 10.41 2.34 6.67 1.9 Male UF 1637 R 9.98 2.3 6.55 1.88 Unknown UF 1637 L 9.73 2.28 6.53 1.88 Unknown UF 1875 R 10.15 2.32 6.43 1.86 Female UF 1875 L 10.01 2.3 6.33 1.85 Female UF 1914 R 8.69 2.16 5.84 1.76 Male UF 1914 L 8.6 2.15 5.94 1.78 Male UF 1915 R 10.45 2.35 6.87 1.93 Male UF 1915 L 10.07 2.31 6.69 1.9 Male UF 1916 R 9.76 2.28 6.11 1.81 Male UF 1917 R 10.45 2.35 6.56 1.88 Male UF 1917 L 10.15 2.32 6.44 1.86 Male UF 1940 R 10.17 2.32 6.17 1.82 Female UF 1940 L 10.02 2.3 6.11 1.81 Female UF 3836 R 9.53 2.25 6.17 1.82 Female UF 3836 L 10.21 2.32 6.07 1.8 Female UF 3977 R 9.62 2.26 6.19 1.82 Female UF 3977 L 9.84 2.29 6.22 1.83 Female UF 3978 R 9.56 2.26 6.31 1.84 Male UF 3978 L 9.37 2.24 6.17 1.82 Male UF 3979 R 9.6 2.26 6.09 1.81 Unknown UF 3979 L 9.53 2.25 6.03 1.8 Unknown UF 3980 R 9.77 2.28 6.53 1.88 Unknown UF 3980 L 9.78 2.28 6.48 1.87 Unknown UF 4574 R 9.57 2.26 6.28 1.84 Male UF 4574 L 9.28 2.23 6.26 1.83 Unknown UF 4695 R 9.86 2.29 6.21 1.83 Male UF 4695 L 10.07 2.31 6.3 1.84 Male UF 4696 R 11.04 2.4 7.12 1.96 Unknown UF 4696 L 11 2.4 7.07 1.96 Unknown UF 4925 R 10.01 2.3 6.19 1.82 Male UF 4925 L 10.01 2.3 6.17 1.82 Male UF 4926 R 9.85 2.29 6.41 1.86 Male UF 4926 L 10.04 2.31 6.45 1.86 Male UF 4951 R 9.74 2.28 6.31 1.84 Male UF 4951 L 9.63 2.26 6.21 1.83 Male UF 5104 R 9.77 2.28 6.49 1.87 Male UF 5104 L 10.07 2.31 6.59 1.89 Male UF 5106 R 9.62 2.26 6.39 1.85 Unknown UF 5584 R 9.94 2.3 6.18 1.82 Female UF 5584 L 9.93 2.3 6.08 1.81 Female

157

UF 5681 R 10.8 2.38 6.38 1.85 Unknown Procyon lotor UF 5681 L 10.42 2.34 6.57 1.88 Unknown UF 5826 R 11.18 2.41 6.91 1.93 Male UF 5826 L 10.89 2.39 7.37 2 Female UF 5828 R 9.93 2.3 6.12 1.81 Unknown UF 5828 L 10.03 2.31 6.07 1.8 Unknown UF 5829 R 9.72 2.27 6.53 1.88 Male UF 5829 L 9.53 2.25 6.64 1.89 Male UF 5830 R 9.32 2.23 5.87 1.77 Female UF 5830 L 9.05 2.2 5.98 1.79 Female UF 5867 R 11.19 2.42 7.01 1.95 Unknown UF 5937 R 9.27 2.23 6.3 1.84 Male UF 5937 L 9.74 2.28 6.57 1.88 Male UF 6653 R 10.29 2.33 6.92 1.93 Female UF 6653 L 10.5 2.35 6.45 1.86 Female UF 6677 R 9.82 2.28 5.9 1.77 Female UF 6677 L 9.78 2.28 5.96 1.79 Female UF 6731 L 10.47 2.35 6.64 1.89 Unknown UF 6733 R 8.46 2.14 5.48 1.7 Female UF 6733 L 8.7 2.16 5.43 1.69 Female UF 6768 R 9.74 2.28 6.35 1.85 Male UF 6768 L 10.04 2.31 6.18 1.82 Male UF 7167 R 9.68 2.27 5.74 1.75 Unknown UF 7167 L 9.74 2.28 5.84 1.76 Unknown UF 7169 R 10.63 2.36 6.41 1.86 Unknown UF 7169 L 10.42 2.34 6.4 1.86 Unknown UF 7751 R 9.4 2.24 5.8 1.76 Unknown UF 7751 L 9.48 2.25 5.76 1.75 Unknown UF 7850 R 9.05 2.2 5.92 1.78 Unknown UF 7850 L 9.04 2.2 6.12 1.81 Unknown UF 7851 R 9.67 2.27 5.58 1.72 Unknown UF 7851 L 9.66 2.27 5.67 1.74 Unknown UF 8072 R 11.23 2.42 7.32 1.99 Unknown UF 8072 L 10.34 2.34 7.42 2 Unknown UF 8073 R 9.7 2.27 6.4 1.86 Unknown UF 8073 L 9.72 2.27 6.45 1.86 Unknown UF 8074 R 10.1 2.31 7.09 1.96 Unknown UF 8074 L 10.04 2.31 7.02 1.95 Unknown UF 8075 R 9.72 2.27 6.07 1.8 Unknown UF 8075 L 9.88 2.29 6.04 1.8 Unknown UF 8085 R 9.65 2.27 6.34 1.85 Unknown UF 8085 L 9.27 2.23 6.37 1.85 Unknown UF 8124 L 10.02 2.3 6.4 1.86 Unknown UF 9205 L 9.54 2.26 6.44 1.86 Male UF 9225 R 9.26 2.23 5.97 1.79 Female

158

UF 9225 L 9.26 2.23 6.07 1.8 Female Procyon lotor UF 9867 R 10.49 2.35 6.32 1.84 Unknown UF 9867 L 10.33 2.34 6.34 1.85 Unknown UF 9990 R 10.09 2.31 6.5 1.87 Unknown UF 9990 L 10.13 2.32 6.54 1.88 Unknown UF 9998 R 9.71 2.27 6 1.79 Female UF 9998 L 9.65 2.27 6.32 1.84 Female UF 10008 R 9.6 2.26 5.88 1.77 Male UF 10008 L 9.85 2.29 5.89 1.77 Male UF 10089 R 10.02 2.3 6.08 1.81 Male UF 10089 L 9.95 2.3 6.19 1.82 Male UF 10095 R 9.9 2.29 5.74 1.75 Male UF 10095 L 9.91 2.29 5.84 1.76 Male UF 10098 R 10.27 2.33 6.12 1.81 Male UF 10098 L 10.49 2.35 5.99 1.79 Male UF 10125 R 9.85 2.29 6.71 1.9 Male UF 10125 L 10.69 2.37 6.77 1.91 Male UF 10189 R 9.55 2.26 5.66 1.73 Male UF 10189 L 9.48 2.25 5.54 1.71 Male UF 10400 R 9.72 2.27 6.44 1.86 Unknown UF 10400 L 9.6 2.26 6.45 1.86 Unknown UF 11947 R 10.35 2.34 6.85 1.92 Unknown UF 11947 L 10.18 2.32 6.89 1.93 Unknown UF 12349 L 10.39 2.34 6.63 1.89 Unknown UF 12509 R 9.76 2.28 6.14 1.81 Male UF 12509 L 9.37 2.24 6.21 1.83 Male UF 12651 R 9.1 2.21 5.84 1.76 Unknown UF 12651 L 9.18 2.22 5.62 1.73 Unknown UF 12655 R 10.09 2.31 6.31 1.84 Female UF 12655 L 10.14 2.32 6.27 1.84 Female UF 12656 R 10.35 2.34 6.45 1.86 Female UF 12656 L 10.54 2.36 6.67 1.9 Female UF 12657 R 10.16 2.32 6.1 1.81 Unknown UF 12657 L 10.04 2.31 6.03 1.8 Unknown UF 12800 R 9.71 2.27 5.99 1.79 Female UF 12800 L 9.55 2.26 6.01 1.79 Female UF 13259 R 9.7 2.27 6.3 1.84 Unknown UF 13259 L 9.85 2.29 6.51 1.87 Unknown UF 13282 R 10.44 2.35 6.4 1.86 Male UF 13282 L 10.16 2.32 6.56 1.88 Male UF 21685 R 9.56 2.26 5.95 1.78 Unknown UF 21685 L 9.84 2.29 5.8 1.76 Unknown UF 21706 L 9.77 2.28 5.93 1.78 Male UF 22129 R 9.16 2.21 5.7 1.74 Female UF 22129 L 9.32 2.23 5.65 1.73 Female

159

UF 22130 R 10.36 2.34 6.62 1.89 Male Procyon lotor UF 22130 L 10.11 2.31 6.58 1.88 Male UF 22420 R 10.68 2.37 6.85 1.92 Male UF 22420 L 10.51 2.35 6.69 1.9 Female UF 22569 R 10.91 2.39 6.48 1.87 Male UF 22569 L 10.58 2.36 6.6 1.89 Male UF 24012 R 9.13 2.21 5.88 1.77 Male UF 24012 L 9.11 2.21 6 1.79 Male UF 24554 R 9.97 2.3 5.93 1.78 Female UF 24554 L 9.56 2.26 6.12 1.81 Female UF 25921 R 10.02 2.3 6.44 1.86 Male UF 25921 L 10.11 2.31 6.55 1.88 Male UF 26007 R 9.88 2.29 6.18 1.82 Male UF 26007 L 9.79 2.28 5.28 1.66 Male UF 26010 R 10.2 2.32 6.21 1.83 Unknown UF 26010 L 10.05 2.31 5.97 1.79 Unknown UF 32659 R 10.06 2.31 5.79 1.76 Unknown UF 32659 L 9.88 2.29 5.74 1.75 Unknown UF 33183 R 10.64 2.36 6.57 1.88 Male UF 33183 L 10.39 2.34 6.74 1.91 Male Bassariscus UF 7192 R 7.45 2.01 3.32 1.2 Male astutus UF 7192 L 7.44 2.01 3.23 1.17 Male UF 7257 R 7.54 2.02 3.34 1.21 Unknown UF 7257 L 7.54 2.02 3.22 1.17 Unknown UF 7865 R 7.38 2 3.35 1.21 Female UF 7865 L 7.29 1.99 3.32 1.2 Female UF 8474 R 7.45 2.01 3.33 1.2 Unknown UF 8474 L 7.94 2.07 3.23 1.17 Unknown UF 11933 R 7.4 2 2.92 1.07 Unknown UF 11933 L 7.38 2 3.36 1.21 Unknown UF 13741 R 7.46 2.01 3.05 1.12 Female UF 13741 L 7.54 2.02 3.13 1.14 Female Bassariscus KUM 3749 X 7.53 2.02 3.42 1.23 Unknown ogallalae Thomas Farm UF 209500 R 9.17 5.09 4.95 1.6 Unknown UF 225330 L 6.73 3.83 3.46 1.24 Unknown UF 406540 L 7.01 3.92 3.74 1.32 Unknown UF V 6371 R 6.69 3.36 3.24 1.17 Unknown Phlaocyon UF 163503 R 6.82 1.92 3.22 1.17 Unknown (Brooksville) UF 163512 L 6.49 1.87 3.09 1.13 Unknown Phlaocyon CM 1602 R 9.91 2.29 4.22 1.44 Unknown ansectens CM 1602 L 10.04 2.31 3.83 1.34 Unknown Pachycynodon MCZ 8913 L 9.95 2.3 4.17 1.43 Unknown dobius MCZ 8926 L 8.12 2.09 3.63 1.29 Unknown

160

Phycynodon MCZ 914 L 8.05 2.09 4.25 1.45 Unknown tenuis Leptarctus cast MCZ X 9.14 2.21 9.31 2.23 Unknown ancipidens 3658 Miomustela MCZ 7016 L 6.12 1.81 1.95 0.67 Unknown sp. Oligobunis cast of L 16.31 2.79 5.91 1.78 Unknown floridanus MCZ 4064

161

APPENDIX C LIST OF CHARACTERS EDITED FROM BASKIN, 2004

1. Suprameatal fossa: shallow (0); deep (1).

2. Alisphenoid canal: present (0); absent (1).

3. Auditory bulla: uninflated (0); somewhat inflated (1); inflated (2).

4. Mastoid process: unreduced (0); reduced (1).

5. Mastoid process: horizontal (0); at a 45 degree angle (1); vertical (2).

6. Paroccipital process: large and transversely widened (0); narrower in cross-section

and shorter (1).

7. Palate extends: to level of M2 (0); just posterior of M2 (1); well posterior of M2 (2).

8. Rostrum: normal (0); elongate (1); shortened (2).

9. Frontal sinuses: normal (0); enlarged (1).

10. Angular process of dentary: below level of tooth row (0); above (1).

11. P1/pl: single rooted (0); reduced to absent (1); double rooted (2).

12. P-3/p2-3: slender (0); anteroposteriorly compressed/transversely broadened (1).

13. P4 protocone: low (0); knoblike (1).

14. P4 protocone: anterior (0); not as anterior (1); medial (2).

15. P4 internal shelf: weakly developed (0); present (1).

16. P4 hypocone: absent (0); small (1); large (2).

17. P4 metacone blade: elongate (0); somewhat reduced (1); greatly reduced (2);

absent (3).

18. P4 parastyle: absent (0); present (1); enlarged (2).

19. Upper molars: triangular with distinct cusps (0); rounded, with reduced cusps (1).

20. Upper molars with external cingulum: present (0); reduced to absent (1).

162

21. M1 parastyle: small (0); enlarged (1).

22. M1 hypocone: present (0); reduced (1); greatly reduced to absent (2).

23. M1 hypocone (or internal cingulum if hypocone is absent): internal (0); posterior to

protocone (1); expanded posterointernally (2).

24. M1 metaconule: absent (0); present (1); prominent (2); secondarily reduced to

absent (3).

25. M1 metaconule: posteroexternal to protocone (0); posterior to protocone (1).

26. M1 metacone: not connected to metaconule (0); connected by crest (1).

27. M2 paracone and metacone: close together (0); separated (1).

28. ml trigonid: with paraconid, protoconid, and metaconid more or less equally spaced

(0); paraconid and metaconid close together (1); paraconid and metaconid adjacent

(2); paraconid blade-like, anteriorly placed (3).

29. ml protoconid: taller than p4 (0); equal (1); lower (2).

30. ml paraconid: single cusped (0); bifid (1); with internal ridge to metaconid (2).

31. ml paraconid: unreduced (0); reduced (1); very reduced to absent (2).

32. ml entoconid: ridge like (0); as cusp (1).

33. ml entoconulid: absent (0); present (1).

34. ml hypoconulid: absent (0); present (1).

35. ml talonid: narrower than trigonid (0); equal (1); wider (2).

36. ml talonid: basined (0); unbasined (1).

37. m2 paraconid: present (0); absent (1).

38. m2 hypoconid: small (0); enlarged (1); larger than protoconid (2).

39. m2 hypoconulid: posteroexternal (0); posterioror posterointernal (1).

163

40. m2 entoconid: absent (0); present (1); fused with reduced hypoconulid (2).

164

APPENDIX D EXTANT TAXA SCANNED

Family Genus Species Common Name Sex Humerus Radius Ulna U UF008079 UF008079 UF008079 M UF014707 UF014707 UF014707 Acinonyx jubatus Cheetah F UF014386 UF014386 UF014386 F UF023688 UF023688 UF023688 F UF026520 UF026520 UF026520 F UF031220 UF031220 UF031220 Puma concolor Puma M UF025908 UF025908 UF025908 M UF031296 UF031296 UF031296 F UF026773 UF026773 UF026773 Puma yagouaroundi Jaguarundi M UF024159 UF024159 UF024159 M 0 UF024622 UF024622 F UF031973 UF031973 UF031973 Lynx lynx Eurasian Lynx M UF031972 UF031972 UF031972 U UF006293 UF006293 0

Felidae F UF024547 UF024547 UF024547 F UF024022 UF024022 UF024022 Lynx rufus Bobcat M UF024548 UF024548 UF024548 M UF024023 UF024023 UF024023 Felis catus Domestic Cat F UF008614 UF008614 UF008614 U 2014_566 0 0 Felis margarita Sand Cat U UF031984 UF031984 UF031984 Felis chaus Jungle Cat U ETVP03013 ETVP03013 ETVP03013 Leopardus colocolo Pampas Cat M UF015218 UF015218 UF015218 F UFZ006780 UFZ006780 UFZ006780 M UFZ006781 UFZ006781 UFZ006781 Leopardus pardalis Ocelot M UF004644 UF004644 UF004644 U UFZ011121 UFZ011121 UFZ011121

165

Catopuma Asian Golden temminckii M UF013415 UF013415 UF013415 (Pardofelis) Cat F UF032134 UF032134 UF032134 M UF032618 UF032618 UF032618 Caracal caracal Caracal U UF019097 UF019097 UF019097 U 1919_25 0 0 U 1936_75 0 0 Leptailurus serval Serval U ETVP07255 ETVP07255 ETVP07255 (Caracal) M UF011932 UF011932 UF011932 F UF010912 UF010912 UF010912 Clouded M UF010771 UF010771 UF010771 Neofelis nebulosa Leopard M UF026153 UF026153 UF026153 U UF031967 UF031967 UF031967 U UF019187 UF019187 UF019187 Panthera leo Lion M UF010643 UF010643 UF010643 U UF008428 UF008428 UF008428 F UF011858 UF011858 UF011858 Panthera pardus Leopard M UF031971 UF031971 UF031971 F UF011928 UF011928 UF011928 Panthera onca Jaguar M UF023685 UF023685 UF023685 M UF032010 UF032010 UF032010 F UF031965 UF031965 UF031965 Panthera tigris Tiger M UF031966 UF031966 UF031966 M UF029264 UF029264 UF029264 Bearcat/ Arctictis binturong M CG1990_88 CG1990_88 CG1990_88 Binturong

Small-toothed Arctogalidia trivirgata M CG2001_495 CG2001_495 CG2001_495 Palm Civet Large Indian Viverra zibetha U A3493 A3493 A3493 Civet

Viverridae M CG1884-1810 CG1884-1810 0 Viverra tangalunga Malayan Civet A12953 (left and U A12953 A12953 right)

166

Small Indian Viverricula indica U CG1951_4 CG1951_4 CG1951_4 Civet Genetta servalina Servaline Genet U CG1983_15 CG1983_15 CG1983_15 Eupleridae Cryptoprocta ferox Fossa U ETVP5468 ETVP5468 ETVP5468 M ETVP07232 ETVP07232 ETVP07232 MNHN_enfants MNHN_enfants MNHN_enfants Crocuta crocuta Spotted Hyena U

perdus perdus perdus U 1936_656 0 0 MNHN_CG1906_1 MNHN_CG1906_ MNHN_CG1906 M 0 10 _10

Hyaenidae MNHN_CG1929_1 MNHN_CG1929_ MNHN_CG1929 Hyaena hyaena Striped Hyena U 734 1734 _1734 MNHN_CG1930_2 MNHN_CG1930_ MNHN_CG1930 U 20 220 _220 Ailuropoda melanoleuca Panda U 2000_363 2000_363 2000_363 M UF0014206 0 UF0014206 Helarctos malayanus Sun Bear U 1881_586 1881_586 1881_586 U 0 1913_505 1913_505 Melursus ursinus Sloth Bear U 1979_307 1979_307 1979_307 Tremarctos ornatus Spectacled Bear U 1992_1469 1992_1469 1992_1469 American Black Ursidae Ursus americanus U 1902_1415 1902_1415 1902_1415 Bear F 0 UF006388 0 Ursus arctos Brown Bear U 1945_11 1945_11 1945_11 U A79655 A79655 A79655 Ursus maritimus Polar Bear U 1912_103 1912_103 1912_103 Asian Black Ursus thibetanus U 1903_151 1903_151 1903_151 Bear F 1960_84 1960_84 1960_84 Ailuridae Ailurus fulgens Red Panda M 1960_85 1960_85 1960_85

European/Com Lutra lutra U 1996_2466 1996_2466 1996_2466 mon Otter North American M UF006676 UF006676 UF006676 Lontra canadensis Otter F 0 0 UF013254 Mustelidae Lontra felina Marine Otter F 1995_185 1995_185 1995_185

167

U 1935_124 1935_124 1935_124 Enhydra lutris Sea Otter U A12503 A12503 A12503 Galictis vittata Greater Grison M CG_2001_1971 CG_2001_1971 CG_2001_1971 African Striped Poecilogale albinucha U 1934_107 1934_107 1934_107 Weasel Pteronura brasiliensis Giant Otter U A1918 A1918 A1918 Neovison vison American Mink M 1958_165 1958_165 1958_165 Mustela eversmanii Steppe Polecat M 2005_668 2005_668 2005_668 European F 1991_605 1991_605 1991_605 Mustela putorius Polecat U 2004_639 2004_639 2004_639 U 1962_332 1962_332 1962_332 Mustela lutreola European Mink M 1987_177 1987_177 1987_177 Ictonyx striatus Striped Polecat M 2001_2185 2001_2185 2001_2185 M 8548 8548 8548

Vormela peregusna Marbled Polecat F 8554 8554 8554 Chinese Ferret- Melogale moschata F 1892_1015 1892_1015 1892_1015 Badger

Mustelidae Eira barbara Tayra M 2000_646 2000_646 2000_646 F 1983_946 1983_946 1983_946 Gulo gulo Wolverine U 1995_1208 1995_1208 1995_1208 M 1998_187 unknown 1998_187 Martes foina Beech Marten M 1994_808 1994_808 1994_808 M 1994_806 1994_806 1994_806 Martes martes Pine Marten M 2005_232 2005_232 2005_232 European F 1917_131 1917_131 1917_131 Meles meles Badger U 1987_28 1987_28 1987_28 American F 9186 9186 9186 Taxidea taxus Badger M 9783 9783 9783 Patagonian Lyncodon patagonicus U 1897_422 1897_422 1897_422 Weasel U 1928_61 1928_61 1928_61 Mellivora capensis Honey Badger M 2004_453 2004_453 2004_453

168

Eastern Lowland F MCZ_37922 MCZ_37922 MCZ_37922 Bassaricyon alleni Olingo M MCZ_37923 MCZ_37923 MCZ_37923 U 395837 0 395837 Bassaricyon gabbii Northern Olingo F 0 305748 305748 U 0 1983_945 0 U 1911_768 1911_768 1911_768 Bassariscus astutus Ring-tailed Cat M 1854_128 1854_128 1854_128 U 0 1888_846 1888_846

Nasua narica Coatimundi U 1894_204 1894_204 1894_204

F 1918_10 0 0 U 0 1968_784 1968_784 South American Nasua nasua M 2010_640 0 0 Coati

Procyonidae F CG_1974_108 CG_1974_108 CG_1974_108 Crab-Eating M 2000_364 2000_364 2000_364 Procyon cancrivorus Raccoon U CG_1932_2235 CG_1932_2235 CG_1932_2235 U 1934_556 1934_556 1934_556 Procyon lotor Raccoon U 0 1948_512 0 U 6038 0 6038 U 0 1889_292 1889_292 U 0 1970_366 1970_366 Potos flavus Kinkajou F 1995_956 0 0 M 1995_957 0 0 Molina's Hog- Conepatus chinga U 1897_432 1897_432 1897_432 nosed Skunk M USNM_087236 USNM_087236 USNM_087236

Mephitis mephitis Striped Skunk F 0 564277 564277 F USNM_260921 0 0 M 564281 564281 564281 Eastern Spotted Spilogale putorius F USNM_564280 0 USNM_564280 Mephitidae Skunk U 0 589252 0 Sunda Stink U 8300 8300 8300 Mydaus javanensis Badger U A3404 A3404 A3404

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U 0 0 UFZ002003 Falkland Islands Dusicyon australis U UF012022 UF012022 UF012022 Wolf U UF012024 UF012024 0 Chrysocyon brachyurus Maned Wolf U UF020778 UF020778 UF020778 F UF027710 UF027710 UF027710 F UF023848 0 UF023848 Canis latrans Coyote M UF013628 UF013628 UF013628 M UF013412 UF013412 UF013412 M UF024340 UF024340 UF024340 U 1973_3 1973_3 1973_3 Canis lupus Grey Wolf U 1984_036 1984_036 1984_036 U ETVP05936 ETVP05936 ETVP05936

Speothos venaticus Bush Dog F UF019126 UF019126 UF019126 F UF013551 UF013551 UF013551 M UF014189 UF014189 UF014189

Canidae Vulpes vulpes Red Fox M UF025919 UF025919 UF025919 U UF030873 UF030873 UF030873 Vulpes (Alopex) lagopus Arctic fox F ETVP14285 ETVP14285 ETVP14285 U UFZ03170A UFZ03170A UFZ03170A Vulpes macrotis Kit Fox U 0 UFZ03170B UFZ03170B U UFZ003172 0 0 Vulpes velox Swift Fox M UF031214 UF031214 UF031214 U ETVP02082 ETVP02082 ETVP02082 Vulpes zerda Fennec Fox U ETVP05558 ETVP05558 ETVP05558 F UF023691 UF023691 UF023691 cinereoargente F UF024270 UF024270 UF024270 Urocyon Grey Fox us M 0 UF019000 UF019000 M UF019130 UF019130 UF019130

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APPENDIX E SUPPLEMENTAL INFORMATION FOR CHAPTER 3

Table E-1. Definition of the landmarks of the humerus used in the analysis. Definitions in quotes are from Fabre et al., 2013b. Landmark Definition 1 Most proximo-lateral point on anterior side of capitulum Insertion point for the ulnaris lateralis muscle. The lateral side of the 2 capitulum centered within concavity. 3 “Most latero-proximal point of the anterior side of the capitulum” 4 Most medio-proximal point of the anterior side of the capitulum 5 “Most medio-proximal point of the anterior side of the trochlea” 6 “Point of maximum curvature of the radial fossa” 7 “Point of maximum curvature of the coronoid fossa” Most latero-proximal point of the medial epicondylar crest where it meets 8 the diaphysis Most latero-distal point of the medial epicondylar crest where it meets the 9 medial epicondyle 10 Most lateral point of the medial epicondyle 11 “Most distal point of the medial epicondyle” 12 Most antero-distal point where the medial epicondyle meets the trochlea 13 Most distal point of where the trochlea meets the lateral epicondyle “Point of maximum concavity of the caudo-medio-distal part of the 14 trochlea” 15 “Most medio-proximal point of the trochlea on the posterior side” “Most proximal point of the olecranon fossa/point of maximum curvature 16 of the olecranon fossa” Most lateral point of the olecranon fossa where it meets with the 17 capitulum Most proximal point where the capitulum and trochlea meet on the 18 posterior side 19 “Point of maximum convexity of the lateral epicondylar crest” 20 Point where the lateral condylar crest meets the diaphysis 21 “Most distal point of the deltopectoral crest” 22 Proximal point of the greater tuberosity on anterior side 23 Point of deepest distal concavity of the bicipital groove 24 “Most proximal tip of the lesser tuberosity” 25 Most disto-medial point of the lesser tuberosity Most proximo-medial point the distal edge of the head where it meets the 26 lesser tuberosity 27 Most distal edge of the posterior side of the head Most proximo-lateral point of the distal edge of the head where it meets 28 the greater tuberosity 29 Most latero-distal point of the greater tuberosity 30 Most proximal point of the greater tuberosity on the posterior side

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Table E-2. Definition of the landmarks of the radius used in the analysis. Definitions in quotes are from Fabre et al., 2013b. Landmark Definition 1 “Most disto-lateral point of anterior side of the ulnar facet” 2 Most proximal point of medial side 3 “Most proximo-lateral point of anterior side of the ulnar facet” 4 “Point of maximum of concavity of the anterior part of the fovea” 5 Most proximal point of the fovea 6 Most proximal point of lateral side 7 Most disto-lateral point of articular circumference 8 Most proximal point of bicipital tuberosity 9 Most lateral point of bicipital tuberosity 10 Most distal point of bicipital tuberosity 11 “Most proximal point of curvature of the distal articular facet with the ulna” 12 “Most disto-medial point of the distal articular facet with the ulna” 13 “Most disto-lateral point of the distal articular facet with the ulna” 14 Most postero-lateral point of the dorsal articular facet 15 Most posterior point of dorsal side of the radius 16 Point where styloid process meets the diaphysis 17 “Medial tip of styloid process” 18 “Most proximal point of groove for extensor carpi radialis longus and brevis” 19 Most distal projection for the groove 20 “Distal tip of the styloid process” 21 “Most disto-lateral point of the dorsal side of the radius” 22 “Most proximal point of the groove for extensor digitorium and extensor indicis”

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Table E-3. Definition of the landmarks of the ulna used in the analysis. Definitions in quotes are from Fabre et al., 2013b. Landmark Definition 1 “Most palmar-medial point of olecranon process” 2 Most distal point of the olecranon in anterior view 3 “Most palmar-lateral point of olecranon process” 4 Narrowest portion of olecranon on lateral side in anterior view 5 Most proximal point of trochlear notch on lateral side 6 “Most proximo-medial point of the incisure of the trochlear notch” 7 “Point of maximum concavity of the proximal part of the trochlear notch” 8 “Most proximo-lateral point of the incisure of the trochlear notch” 9 Most narrow point of the incisure of the trochlear notch along lateral side 10 Most proximo-lateral point of the incisure of the radial notch 11 Most lateral point of the incisure of the radial notch 12 “Point of maximum concavity between the radial notch and the trochlear notch” 13 Most distal point of the radial notch 14 Point of maximum concavity of the proximal part of the coronoid process? 15 Most proximal point of the coronoid process 16 Most medial point of the trochlear notch 17 Most narrow point of the trochlear notch on medial side 18 “Most dorsal-medial point of the olecranon process” 19 “Most dorsal-lateral point of the olecranon process” 20 “Most proximal point of insertion of the medial epicondylar crest on the diaphysis” 21 “Most distal point of insertion of the medial epicondylar crest on the diaphysis” 22 Point where the posterior edge of the distal diaphysis begins to expand 23 Most postero-lateral point of the distal diaphysis 24 The point where the diaphysis expands to form the articular facet that articulates with the radius 25 “Most proximal point of the articular facet that articulates with the radius” 26 “Most distal point of the articular facet that articulates with the radius” 27 Point of greatest concavity between the articular facet that articulates with the radius and the styloid process 28 Most distal point of the styloid process 29 Attachment location for the Pronator quadratus

173

Table E-4. MAOVAs and Phylogenetic MAOVAs for 90% of overall variance and 95% of overall variance MANOVA Phylogenetic MANOVA MANOVA Phylogenetic MANOVA 90% of overall variance 95% of overall variance Character F P-value P-value Df Residuals Wilks F P-value P-value Df Residuals Wilks

Family 10.43 0.001 N/A 9 71 0.001 7.70 0.001 N/A 9 71 0.001 Locomotion 5.11 0.001 0.001 5 75 0.08 3.77 0.001 0.001 5 75 0.03 Hunting 7.14 0.001 0.001 6 74 0.02 5.26 0.001 0.001 6 74 0.004 Activity 1.55 0.030 0.190 4 76 0.48 1.10 0.300 0.771 4 76 0.34 Diet 2.95 0.001 0.001 6 74 0.14 2.62 0.001 0.001 6 74 0.04 Habitat 3.14 0.001 0.001 7 73 0.09 2.29 0.001 0.001 7 73 0.03 CompleteHumerus Social Behavior 2.42 0.000 0.010 3 77 0.44 2.04 0.001 0.006 3 77 0.26

Family 5.78 0.001 N/A 9 71 0.001 5.45 0.001 N/A 9 71 0.001 Locomotion 3.14 0.001 0.002 5 75 0.04 2.58 0.001 0.003 5 75 0.02 Hunting 3.69 0.001 0.391 6 74 0.01 3.34 0.001 0.274 6 74 0.004 Activity 1.08 0.328 0.820 4 76 0.35 1.01 0.471 0.879 4 76 0.23 Diet 3.37 0.001 0.001 4 76 0.07 2.73 0.001 0.001 4 76 0.04 Habitat 1.68 0.001 0.124 7 73 0.06 1.50 0.001 0.271 7 73 0.03 Proximal Proximal Humerus Social Behavior 1.86 0.002 0.040 3 77 0.29 1.74 0.002 0.036 3 77 0.18

Family 7.30 0.001 N/A 9 71 0.001 6.17 0.001 N/A 9 71 0.001

Locomotion 3.00 0.001 0.001 5 75 0.05 2.87 0.001 0.001 5 75 0.01 Hunting 3.49 0.001 0.631 6 74 0.02 3.06 0.001 0.509 6 74 0.004 Activity 1.34 0.057 0.291 4 76 0.28 1.44 0.015 0.084 4 76 0.13 Diet 2.68 0.001 0.004 4 76 0.11 2.45 0.001 0.011 4 76 0.05

DistalHumerus Habitat 1.72 0.001 0.096 7 73 0.06 1.63 0.001 0.059 7 73 0.02 Social Behavior 2.06 0.003 0.007 3 77 0.26 1.69 0.003 0.047 3 77 0.17

Family 6.69 0.001 N/A 9 71 0.003 6.05 0.001 N/A 9 71 0.001 Locomotion 5.08 0.001 0.001 5 75 0.06 4.36 0.001 0.001 5 75 0.02 Hunting 4.79 0.001 0.193 6 74 0.04 4.80 0.001 0.002 6 74 0.01 Activity 1.20 0.201 0.671 4 76 0.52 1.02 0.448 0.865 4 76 0.37 Diet 3.02 0.001 0.003 4 76 0.23 2.92 0.001 0.002 4 76 0.09

Habitat 2.12 0.001 0.013 7 73 0.15 1.95 0.001 0.008 7 73 0.04 CompleteRadius Social Behavior 2.34 0.001 0.006 3 77 0.41 2.13 0.001 0.005 3 77 0.25

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Table E-4. Continued. MANOVA Phylogenetic MANOVA MANOVA Phylogenetic MANOVA 90% of overall variance 95% of overall variance Character F P-value P-value Df Residuals Wilks F P-value P-value Df Residuals Wilks

Family 5.99 0.001 N/A 9 71 0.001 4.97 0.001 N/A 9 71 0.001

Locomotion 2.61 0.001 0.042 5 75 0.10 2.12 0.001 0.193 5 75 0.05 Hunting 3.05 0.001 0.981 6 74 0.05 2.55 0.001 0.994 6 74 0.02 Activity 1.05 0.393 0.846 4 76 0.44 1.00 0.480 0.906 4 76 0.28 Diet 1.99 0.001 0.300 4 76 0.24 1.83 0.000 0.372 4 76 0.13

Habitat 2.16 0.001 0.003 7 73 0.06 2.02 0.001 0.001 7 73 0.02 Proximal Proximal Radius Social Behavior 1.52 0.032 0.323 3 77 0.43 1.29 0.102 0.527 3 77 0.31 Family 7.87 0.001 N/A 9 71 0.001 6.79 0.001 N/A 9 71 0.001

Locomotion 2.64 0.001 0.017 5 75 0.09 2.39 0.001 0.024 5 75 0.04 Hunting 3.28 0.001 0.855 6 74 0.03 2.86 0.001 0.939 6 74 0.01 Activity 1.10 0.307 0.780 4 76 0.39 1.05 0.387 0.845 4 76 0.27 Diet 2.40 0.001 0.038 4 76 0.16 2.20 0.001 0.055 4 76 0.09

DistalRadius Habitat 1.68 0.001 0.172 7 73 0.09 1.63 0.001 0.117 7 73 0.03 Social Behavior 1.63 0.013 0.164 3 77 0.38 1.37 0.058 0.399 3 77 0.30 Family 8.39 0.001 N/A 9 71 0.01 6.67 0.001 N/A 9 71 0.001 Locomotion 4.85 0.001 0.001 5 75 0.15 3.35 0.001 0.001 5 75 0.06 Hunting 5.30 0.001 0.227 6 74 0.09 3.85 0.001 0.500 6 74 0.03 Activity 1.06 0.393 0.860 4 76 0.67 1.35 0.066 0.322 4 76 0.36 Diet 3.65 0.001 0.002 4 76 0.30 3.11 0.001 0.001 4 76 0.13

CompleteUlna Habitat 2.41 0.001 0.007 7 73 0.22 1.74 0.000 0.138 7 73 0.10 Social Behavior 1.94 0.010 0.152 3 77 0.59 1.77 0.005 0.092 3 77 0.38 Family 8.53 0.001 N/A 9 71 0.001 7.41 0.001 N/A 9 71 0.001 Locomotion 2.91 0.001 0.005 5 75 0.71 2.29 0.001 0.034 5 75 0.03

Ulna Hunting 3.96 0.001 0.306 6 74 0.02 3.31 0.001 0.340 6 74 0.01

Activity 1.38 0.047 0.265 4 76 0.32 1.36 0.035 0.211 4 76 0.16 Diet 2.63 0.001 0.012 4 76 0.14 2.31 0.001 0.014 4 76 0.67

Proximal Habitat 1.77 0.001 0.076 7 73 0.08 1.66 0.001 0.067 7 73 0.02 Social Behavior 1.51 0.030 0.291 3 77 0.40 1.25 0.127 0.593 3 77 0.29

175

Table E-5. Calculation of Phylogenetic Signal. K P Complete Humerus 0.38 0.001 Proximal Humerus 0.25 0.001 Distal Humerus 0.32 0.001 Complete Radius 0.31 0.001 Proximal Radius 0.24 0.001 Distal Radius 0.23 0.001 Complete Ulna 0.33 0.001 Proximal Ulna 0.34 0.001

176

Table E-6. Complete Humerus Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC42 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

B. Cross Validation Predicted Group sum=79 Rate of reclassification= 82% Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 17 0 0 0 0 0 1 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 1 0 0 0 3 0 0 0 0 Mustelidae 3 0 0 1 0 0 17 1 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 1 0 1 6 0 Viverridae 0 0 0 2 0 0 0 3 0 1

177

Table E-7. Complete Humerus Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC42 Locomotion Predicted Group sum=81 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 0 0 21 0 0 4 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 1 1 0 9 1 Terrestrial 0 0 1 0 0 16

B. Cross Validation Predicted Group sum=81 Rate of reclassification = 41% Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 4 0 4 1 0 1 Cursorial 0 5 0 0 0 3 Scansorial 2 2 13 1 1 6 Semiaquatic 2 0 0 4 1 2 Semifossorial 1 2 3 1 4 1 Terrestrial 1 2 7 1 3 3

178

Table E-8. Complete Humerus Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation. A. Discriminant Analysis Hunting Strategy Predicted Group sum=81 Actual Group Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 24 0 0 1 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 0 0 19 0 0 0 0 Pounce/pursuit 1 0 0 15 1 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

B. Cross Validation Predicted Group sum=81 Rate of reclassification = 58% Actual Group Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 18 0 3 1 0 2 1 Aquatic 0 3 2 1 0 1 0 Occasional 4 0 13 1 0 1 0 Pounce/pursuit 3 2 1 10 1 0 0 Pursuit 0 0 0 2 0 0 0 Semifossorial 2 0 1 0 0 3 3 Various 1 0 0 0 0 1 0

179

Table E-9. Complete Humerus Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Rate of reclassification = Activity Pattern Predicted Group sum=81 Predicted Group 33% Actual Group A C D N O Actual Group A C D N O A 13 1 0 1 0 A 8 1 1 4 1 C 0 3 0 0 0 C 1 0 0 1 1 D 0 0 7 2 0 D 2 1 1 3 2 N 2 0 0 37 1 N 6 6 5 16 7 O 1 0 0 3 10 O 2 3 2 5 2

Table E-10. Complete Humerus Diet LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Diet Predicted Group sum=7 Predicted Group Rate of reclassification = 51% Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 0 2 1 0 0 I 0 7 0 0 0 I 0 2 2 2 1 M 0 0 40 0 0 M 0 5 24 6 5 O 0 0 4 23 0 O 1 2 10 13 1 V 0 0 0 0 4 V 0 0 1 1 2

180

Table E-11. Complete Humerus Habitat LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Habitat Predicted Group sum=81 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 1 2 1 0 0 0 0 0 F 0 27 0 0 0 0 1 1 F 2 15 3 2 0 2 3 3 N 0 2 14 0 0 0 0 0 N 0 5 4 3 0 1 2 2 O 0 0 0 9 0 0 1 0 O 0 1 4 2 0 0 0 1 Q 0 0 0 0 6 0 0 0 Q 1 0 0 1 2 0 1 0 S 0 0 0 0 0 7 0 0 S 0 1 2 0 0 2 3 1 T 0 0 0 0 0 0 2 0 T 0 1 0 1 0 0 0 0 W 0 1 0 0 0 0 0 6 W 0 5 0 0 0 0 0 2

Table E-12. Complete Humerus Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Rate of reclassification = Social Behavior Predicted Group sum=81 Predicted Group 62% variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 5 1 0 0 group 1 1 2 2 Pairs 0 6 1 0 pairs 0 2 2 3 Solitary 0 0 52 0 solitary 3 2 40 7 Variable group 0 0 2 14 variable group 1 5 3 7

181

Table E-13. Proximal Humerus Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC45 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

B. Cross Validation Predicted Group Rate of reclassification = Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 16 0 0 0 0 0 2 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 2 1 0 1 0 Mustelidae 1 0 1 1 0 1 14 4 0 0 Procyonidae 0 0 0 0 0 0 3 5 0 0 Ursidae 1 0 0 0 0 0 0 0 7 0 Viverridae 1 0 0 4 0 0 0 1 0 0

182

Table E-14. Proximal Humerus Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC45 Locomotion Predicted Group sum=79 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 1 0 24 0 0 0 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 0 0 12 0 Terrestrial 0 0 1 0 0 16

B. Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 3 1 2 2 1 1 Cursorial 0 5 1 0 1 1 Scansorial 3 2 13 2 0 5 Semiaquatic 0 0 1 4 2 2 Semifossorial 0 2 3 0 4 3 Terrestrial 1 2 5 3 4 2

183

Table E-15. Proximal Humerus Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis Hunting Strategy Predicted Group sum=81 Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 25 0 0 0 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 0 0 19 0 0 0 0 Pounce Pursuit 0 0 0 17 0 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

B. Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 15 1 4 3 0 2 0 Aquatic 1 2 1 1 0 2 0 Occasional 3 2 9 2 0 3 0 Pounce Pursuit 4 1 1 5 3 3 0 Pursuit 0 0 0 2 0 0 0 Semifossorial 1 1 0 4 0 3 0 Various 2 0 0 0 0 0 0

184

Table E-16. Proximal Humerus Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=77 Predicted Group Actual Group A C D N O Actual Group A C D N O A 13 1 0 1 0 A 6 1 1 5 2 C 0 3 0 0 0 C 1 0 0 1 1 D 0 0 9 0 0 D 2 1 0 3 3 N 2 0 0 38 0 N 9 5 6 16 4 O 1 0 0 0 14 O 2 1 3 3 5

Table E-17. Proximal Humerus Diet LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Diet Predicted Group sum=76 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 1 0 1 0 1 I 0 7 0 0 0 I 0 2 3 2 0 M 0 0 38 2 0 M 0 5 23 11 1 O 0 0 3 24 0 O 1 2 12 12 0 V 0 0 0 0 4 V 1 0 1 0 2

185

Table E-18. Proximal Humerus Habitat LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Habitat Predicted Group sum=77 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 0 1 1 0 0 1 0 1 F 1 28 0 0 0 0 0 0 F 2 13 5 2 1 3 2 1 N 0 1 14 0 0 0 0 0 N 3 2 1 5 0 4 0 1 O 0 0 0 10 0 0 0 0 O 2 0 2 1 0 1 3 1 Q 0 0 0 0 6 0 0 0 Q 0 1 2 1 1 0 0 1 S 0 0 0 0 0 6 0 1 S 2 1 3 0 0 0 0 1 T 0 0 0 0 0 0 2 0 T 0 0 1 1 0 0 0 0 W 0 0 0 0 0 0 0 7 W 1 1 1 1 0 2 0 1

Table E-19. Proximal Humerus Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=79 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 6 0 0 0 Group 3 1 0 2 Pairs 0 6 1 0 Pairs 1 5 1 0 Solitary 0 0 52 0 Solitary 5 4 32 11 Variable group 0 0 1 15 Variable group 3 2 5 6

186

Table E-20. Distal Humerus Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC46 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

B. Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 10 0 0 1 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 3 0 0 0 1 Mustelidae 2 0 0 0 0 0 20 0 0 0 Procyonidae 2 0 0 0 0 0 0 4 0 2 Ursidae 0 0 0 0 0 0 0 1 6 1 Viverridae 0 0 0 1 0 1 0 3 0 1

187

Table E-21. Distal Humerus Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC46 Locomotion Predicted Group sum=81 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 0 0 25 0 0 0 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 0 0 12 0 Terrestrial 0 0 0 0 0 17

B. Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 5 0 2 1 0 2 Cursorial 0 8 0 0 0 0 Scansorial 0 2 12 2 2 7 Semiaquatic 1 0 1 4 1 2 Semifossorial 0 0 2 1 7 2 Terrestrial 3 1 4 2 1 6

188

Table E-22. Distal Humerus Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation.

B. Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 18 1 3 1 0 2 0 Aquatic 0 5 1 1 0 0 0 Occasional 4 1 11 2 0 1 0 Pounce Pursuit 2 0 1 9 2 3 0 Pursuit 0 0 0 2 0 0 0 Semifossorial 3 1 1 3 0 1 0 Various 1 0 0 0 0 1 0

189

Table E-23. Distal Humerus Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=72 Predicted Group Actual Group A C D N O Actual Group A C D N O A 14 0 0 0 0 A 5 0 2 3 5 C 0 3 0 0 1 C 0 0 1 1 1 D 0 0 8 1 0 D 1 2 2 2 2 N 2 0 0 37 1 N 8 3 5 15 9 O 1 0 0 3 10 O 4 2 1 6 1

Table E-24. Distal Humerus Diet LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Diet Predicted Group sum=81 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 1 0 1 1 0 I 0 7 0 0 0 I 1 0 3 2 1 M 0 0 40 0 0 M 2 5 27 5 1 O 0 0 0 27 0 O 2 2 8 13 2 V 0 0 0 0 4 V 0 0 1 1 2

190

Table E-25. Distal Humerus Habitat LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Habitat Predicted Group sum=80 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 1 0 1 0 0 0 0 2 F 0 29 0 0 0 0 0 0 F 3 13 3 5 0 3 1 1 N 0 0 16 0 0 0 0 0 N 2 2 6 4 0 1 0 1 O 0 1 0 9 0 0 0 0 O 0 2 4 2 0 1 1 0 Q 0 0 0 0 6 0 0 0 Q 0 0 0 2 4 0 0 0 S 0 0 0 0 0 7 0 0 S 0 1 2 2 0 1 0 1 T 0 0 0 0 0 0 2 0 T 0 1 1 0 0 0 0 0 W 0 0 0 0 0 0 0 7 W 0 3 1 0 0 0 0 3

Table E-26. Distal Humerus Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=79 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 6 0 0 0 Group 2 1 1 2 Pairs 0 7 0 0 Pairs 1 1 1 4 Solitary 0 0 51 1 Solitary 3 4 38 7 Variable group 0 0 1 15 Variable group 2 4 7 3

191

Table E-27. Complete Radius Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC40 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

B. Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 2 0 0 0 2 0 0 Mustelidae 1 0 0 0 0 1 17 3 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 1 0 0 0 7 0 Viverridae 0 0 0 0 0 0 0 0 0 6

192

Table E-28. Complete Radius Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC40 Locomotion Predicted Group sum=73 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 7 0 0 0 1 Scansorial 0 0 22 0 0 3 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 1 0 11 0 Terrestrial 0 1 2 0 0 14

B. Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 6 0 2 0 1 1 Cursorial 0 5 0 0 0 3 Scansorial 0 3 13 0 2 7 Semiaquatic 0 0 0 7 1 1 Semifossorial 1 0 2 2 5 2 Terrestrial 1 3 7 1 2 3

193

Table E-29. Complete Radius Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis Hunting Strategy Predicted Group sum=77 Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 24 0 0 1 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 0 0 18 1 0 0 0 Pounce Pursuit 0 0 1 15 1 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

B. Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 20 0 2 1 0 2 0 Aquatic 1 6 0 0 0 0 0 Occasional 2 0 12 2 2 0 1 Pounce Pursuit 2 0 3 9 2 1 0 Pursuit 0 0 0 2 0 0 0 Semifossorial 0 0 2 0 1 6 0 Various 0 0 1 1 0 0 0

194

Table E-30. Complete Radius Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=67 Predicted Group Actual Group A C D N O Actual Group A C D N O A 12 0 0 3 0 A 7 0 3 5 0 C 0 2 0 0 1 C 0 0 0 2 1 D 0 0 8 1 0 D 1 1 5 2 0 N 2 0 0 36 2 N 6 2 2 21 9 O 0 0 0 5 9 O 2 1 0 7 4

Table E-31. Complete Radius Diet LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Diet Predicted Group sum=74 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 2 0 0 0 1 I 0 7 0 0 0 I 1 4 1 1 0 M 0 0 38 2 0 M 0 4 23 10 3 O 0 0 5 22 0 O 1 0 12 13 1 V 0 0 0 0 4 V 0 1 0 1 2

195

Table E-32. Complete Radius Habitat LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Habitat Predicted Group sum=74 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 0 1 2 0 0 0 0 1 F 1 27 0 0 0 1 0 0 F 3 12 2 0 1 3 1 7 N 1 1 14 0 0 0 0 0 N 3 4 7 0 0 1 0 1 O 0 1 0 9 0 0 0 0 O 3 2 1 3 0 1 0 0 Q 0 0 0 0 6 0 0 0 Q 0 1 0 1 2 0 0 2 S 0 0 0 0 0 7 0 0 S 0 1 1 0 0 4 0 1 T 0 0 0 0 0 0 1 1 T 0 0 0 0 0 0 0 2 W 0 1 0 0 0 0 0 6 W 0 3 0 1 0 1 0 2

Table E-33. Complete Radius Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=76 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 4 0 1 1 Group 0 1 3 2 Pairs 0 5 1 1 Pairs 1 2 3 1 Solitary 0 0 52 0 Solitary 3 5 36 8 Variable group 0 0 1 15 Variable group 3 2 3 8

196

Table E-34. Proximal Radius Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC44 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

B. Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 8 0 0 0 2 1 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 14 0 0 3 0 0 1 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 2 0 1 0 1 Mustelidae 2 0 1 1 0 1 13 2 0 2 Procyonidae 0 0 0 0 0 0 1 7 0 0 Ursidae 0 0 1 1 0 1 0 0 5 0 Viverridae 0 0 0 0 0 0 1 3 0 2

197

Table E-35. Proximal Radius Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC44 Locomotion Predicted Group sum=76 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 9 0 1 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 0 0 24 0 1 0 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 1 0 11 0 Terrestrial 0 1 1 0 0 15

B. Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 3 0 4 1 1 1 Cursorial 0 5 1 2 0 0 Scansorial 3 2 8 3 4 5 Semiaquatic 0 1 3 4 1 0 Semifossorial 1 0 4 1 4 2 Terrestrial 2 2 3 2 1 7

198

Table E-36. Proximal Radius Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation.

B. Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 13 2 4 4 0 0 2 Aquatic 2 3 0 1 0 1 0 Occasional 3 0 12 3 0 0 1 Pounce Pursuit 1 2 1 5 3 3 2 Pursuit 0 0 0 2 0 0 0 Semifossorial 0 1 0 2 0 6 0 Various 0 0 1 1 0 0 0

199

Table E-37. Proximal Radius Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=71 Predicted Group Actual Group A C D N O Actual Group A C D N O A 15 0 0 0 0 A 15 0 0 0 0 C 0 3 0 0 0 C 0 3 0 0 0 D 0 0 9 0 0 D 0 0 9 0 0 N 1 0 0 35 4 N 1 0 0 35 4 O 0 0 0 5 9 O 0 0 0 5 9

Table E-38. Proximal Radius Diet LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Diet Predicted Group sum=79 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 0 1 1 1 0 I 0 7 0 0 0 I 0 3 2 1 1 M 0 0 39 1 0 M 0 6 27 5 2 O 0 0 1 26 0 O 0 1 8 13 5 V 0 0 0 0 4 V 0 0 1 2 1

200

Table E-39. Proximal Radius Habitat LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Habitat Predicted Group sum=79 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 0 1 1 0 1 0 0 1 F 0 29 0 0 0 0 0 0 F 3 9 3 6 0 5 0 3 N 0 1 15 0 0 0 0 0 N 3 0 6 4 2 0 0 1 O 0 1 0 9 0 0 0 0 O 1 3 1 2 2 1 0 0 Q 0 0 0 0 6 0 0 0 Q 1 2 1 0 2 0 0 0 S 0 0 0 0 0 7 0 0 S 0 2 0 0 0 1 0 4 T 0 0 0 0 0 0 2 0 T 0 0 1 1 0 0 0 0 W 0 0 0 0 0 0 0 7 W 0 2 0 0 0 3 0 2

Table E-40. Proximal Radius Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation.

A. B. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=77 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 6 0 0 0 Group 0 1 2 3 Pairs 0 6 1 0 Pairs 0 1 2 4 Solitary 0 0 51 1 Solitary 5 5 31 11 Variable group 0 0 2 14 Variable group 4 1 9 2

201

Table E-41. Distal Radius Family-level LDA: a) Discriminant Analysis. b) Cross Validation.

B. Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 10 0 0 1 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 17 1 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 1 0 0 0 2 0 1 0 Mustelidae 0 0 0 0 1 2 13 4 1 1 Procyonidae 1 0 0 0 0 1 2 3 0 1 Ursidae 1 0 0 0 0 0 1 0 6 0 Viverridae 1 0 0 0 0 0 0 0 0 5

202

Table E-42. Distal Radius Locomotion LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis PC40 Locomotion Predicted Group sum=76 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 0 0 24 0 0 1 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 0 0 11 1 Terrestrial 0 2 1 0 0 14

B. Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 5 0 2 2 0 1 Cursorial 0 3 3 0 1 1 Scansorial 0 2 14 2 3 4 Semiaquatic 3 1 1 1 0 3 Semifossorial 2 1 1 1 4 3 Terrestrial 1 2 4 6 0 4

203

Table E-43. Distal Radius Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation.

A. Discriminant Analysis Hunting Strategy Predicted Group sum=80 Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 24 0 0 1 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 0 0 19 0 0 0 0 Pounce Pursuit 0 0 0 17 0 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

B. Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 15 3 5 2 0 0 0 Aquatic 1 3 0 1 1 1 0 Occasional 2 0 13 1 1 1 1 Pounce Pursuit 1 2 1 8 4 0 1 Pursuit 0 0 0 2 0 0 0 Semifossorial 0 1 1 2 0 4 1 Various 0 0 1 1 0 0 0

204

Table E-44. Distal Radius Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation.

Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=70 Predicted Group Actual Group A C D N O Actual Group A C D N O A 12 0 0 2 1 A 7 0 2 3 3 C 0 3 0 0 0 C 0 0 1 2 0 D 0 0 9 0 0 D 2 1 1 5 0 N 1 0 1 36 2 N 7 4 7 12 10 O 0 0 0 4 10 O 3 1 2 6 2

Table E-45. Distal Radius Diet LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Diet Predicted Group sum=75 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 1 0 2 0 0 I 0 6 1 0 0 I 1 1 3 2 0 M 0 1 37 2 0 M 2 4 19 12 3 O 0 0 2 25 0 O 0 3 10 11 3 V 0 0 0 0 4 V 0 0 1 0 3

205

Table E-46. Distal Radius Habitat LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Habitat Predicted Group sum=76 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 1 1 1 0 1 0 0 0 F 0 28 0 0 0 0 0 1 F 0 13 5 2 2 2 1 4 N 0 1 15 0 0 0 0 0 N 0 1 8 1 2 4 0 0 O 0 0 1 9 0 0 0 0 O 1 1 3 2 0 0 0 3 Q 0 0 0 0 6 0 0 0 Q 1 2 0 0 2 1 0 0 S 0 1 1 0 0 5 0 0 S 0 1 3 1 0 0 0 2 T 0 0 0 0 0 0 2 0 T 1 0 0 0 0 1 0 0 W 0 0 0 0 0 0 0 7 W 0 2 1 1 1 2 0 0

Table E-47. Distal Radius Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=79 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 6 0 0 0 Group 0 0 2 4 Pairs 0 6 1 0 Pairs 1 0 5 1 Solitary 0 0 52 0 Solitary 1 8 34 9 Variable group 0 0 1 15 Variable group 7 0 5 4

206

Table E-48. Complete Ulna Family-level LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis PC37 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 1 0 17 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 1 3 0 0 0 0 Mustelidae 0 0 0 0 0 0 19 3 0 0 Procyonidae 0 0 0 0 0 0 1 7 0 0 Ursidae 0 0 0 0 0 0 1 0 7 0 Viverridae 0 0 0 0 0 0 0 2 0 4

207

Table E-49. Complete Ulna Locomotion LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis PC37 Locomotion Predicted Group sum=75 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 7 1 0 0 0 Scansorial 0 1 24 0 0 0 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 2 0 10 0 Terrestrial 0 1 1 0 0 15

Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 6 0 3 0 1 0 Cursorial 0 4 2 0 0 2 Scansorial 2 1 14 1 2 5 Semiaquatic 2 0 1 4 1 1 Semifossorial 0 1 3 2 2 4 Terrestrial 1 3 3 0 4 6

208

Table E-50. Complete Ulna Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Hunting Strategy Predicted Group sum=78 Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 24 0 0 1 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 1 0 18 0 0 0 0 Pounce Pursuit 0 0 0 16 1 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce pursuit Pursuit Semifossorial Various Ambush 20 0 3 2 0 0 0 Aquatic 0 4 0 2 0 1 0 Occasional 3 1 11 1 1 1 1 Pounce Pursuit 2 0 3 11 1 0 0 Pursuit 1 0 0 1 0 0 0 Semifossorial 0 1 1 0 1 3 3 Various 1 0 1 0 0 0 0

209

Table E-51. Complete Ulna Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=73 Predicted Group Actual Group A C D N O Actual Group A C D N O A 14 0 0 1 0 A 9 1 0 4 1 C 0 2 0 1 0 C 0 0 1 2 0 D 0 0 8 0 1 D 2 1 4 0 2 N 2 0 0 37 1 N 7 6 1 18 8 O 0 0 0 2 12 O 1 0 0 6 7

Table E-52. Complete Ulna Diet LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Diet Predicted Group sum=75 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 0 1 1 1 0 I 0 7 0 0 0 I 0 2 3 2 0 M 0 0 39 1 0 M 1 5 23 9 2 O 0 0 4 22 1 O 1 2 12 8 4 V 0 0 0 0 4 V 0 0 1 2 1

210

Table E-53. Complete Ulna Habitat LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Habitat Predicted Group sum=69 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 3 1 0 0 0 0 0 0 D 2 1 0 1 0 0 0 0 F 0 24 1 0 0 0 1 3 F 1 11 3 3 0 2 3 6 N 0 2 14 0 0 0 0 0 N 0 2 9 1 0 3 0 1 O 1 0 0 9 0 0 0 0 O 2 1 2 2 0 3 0 0 Q 0 0 0 0 6 0 0 0 Q 0 1 0 0 3 1 1 0 S 0 1 0 0 0 6 0 0 S 0 1 0 0 1 0 2 2 T 0 0 0 0 0 0 2 0 T 0 1 0 0 0 1 0 0 W 0 2 0 0 0 0 0 5 W 1 4 0 0 0 0 0 2

Table E-54. Complete Ulna Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=74 Predicted Group variable variable Actual Group group pairs solitary group Actual Group group pairs solitary group Group 4 0 1 1 Group 1 1 3 1 Pairs 0 6 1 0 Pairs 2 2 2 0 Solitary 0 0 50 2 Solitary 4 7 36 5 Variable group 1 0 1 14 Variable group 1 2 4 9

211

Table E-55. Proximal Ulna Family-level LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis PC43 Family Predicted Group sum=81 Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 11 0 0 0 0 0 0 0 0 Eupleridae 0 0 1 0 0 0 0 0 0 0 Felidae 0 0 0 18 0 0 0 0 0 0 Hyaenidae 0 0 0 0 2 0 0 0 0 0 Mephitidae 0 0 0 0 0 4 0 0 0 0 Mustelidae 0 0 0 0 0 0 22 0 0 0 Procyonidae 0 0 0 0 0 0 0 8 0 0 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 0 0 0 0 0 0 0 0 6

Cross Validation Predicted Group Actual Group Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 10 0 0 1 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 16 1 0 0 1 0 0 Hyaenidae 0 0 0 1 0 0 0 0 0 1 Mephitidae 0 0 0 0 0 2 1 0 0 1 Mustelidae 1 0 0 4 0 3 15 3 0 0 Procyonidae 0 0 0 0 0 0 1 6 0 1 Ursidae 1 0 0 0 0 0 0 0 7 0 Viverridae 0 0 0 0 1 1 0 0 0 4

212

Table E-56. Proximal Ulna Locomotion LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis PC43 Locomotion Predicted Group sum=79 Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 0 0 0 0 Cursorial 0 8 0 0 0 0 Scansorial 0 0 25 0 0 0 Semiaquatic 0 0 0 9 0 0 Semifossorial 0 0 0 0 12 0 Terrestrial 0 1 0 0 1 15

Cross Validation Predicted Group Actual Group Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 4 1 3 1 1 0 Cursorial 2 1 1 0 2 2 Scansorial 3 1 12 1 2 6 Semiaquatic 1 0 3 4 1 0 Semifossorial 1 3 3 0 2 3 Terrestrial 1 4 5 0 4 3

213

Table E-57. Proximal Ulna Hunting Strategy LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Hunting Strategy Predicted Group sum=80 Actual Group Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 24 0 1 0 0 0 0 Aquatic 0 7 0 0 0 0 0 Occasional 0 0 19 0 0 0 0 Pounce/pursuit 0 0 0 17 0 0 0 Pursuit 0 0 0 0 2 0 0 Semifossorial 0 0 0 0 0 9 0 Various 0 0 0 0 0 0 2

Cross Validation Predicted Group Actual Group Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 16 0 3 1 1 4 0 Aquatic 0 6 0 0 0 1 0 Occasional 3 0 12 3 1 0 0 Pounce/pursuit 0 0 4 11 0 2 0 Pursuit 0 0 0 1 0 1 0 Semifossorial 2 0 1 2 0 4 0 Various 0 0 1 1 0 0 0

214

Table E-58. Distal Humerus Activity Pattern LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Activity Pattern Predicted Group sum=74 Predicted Group Actual Group A C D N O Actual Group A C D N O A 13 0 0 2 0 A 8 2 0 2 3 C 0 2 0 1 0 C 0 0 1 2 0 D 0 0 8 1 0 D 1 1 4 2 1 N 0 0 0 39 1 N 5 5 3 15 12 O 1 0 0 1 12 O 3 0 2 5 4

Table E-59. Proximal Ulna Diet LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Diet Predicted Group sum=71 Predicted Group Actual Group F I M O V Actual Group F I M O V F 3 0 0 0 0 F 0 2 1 0 0 I 0 7 0 0 0 I 1 1 2 3 0 M 0 0 36 4 0 M 1 7 17 14 1 O 0 0 6 21 0 O 0 2 13 8 4 V 0 0 0 0 4 V 0 0 1 2 1

215

Table E-60. Proximal Ulna Habitat LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Habitat Predicted Group sum=81 Predicted Group Actual Group D F N 0 Q S T W Actual Group D F N O Q S T W D 4 0 0 0 0 0 0 0 D 3 0 1 0 0 0 0 0 F 0 29 0 0 0 0 0 0 F 1 9 7 4 0 1 1 6 N 0 0 16 0 0 0 0 0 N 0 2 9 2 0 2 1 0 O 0 0 0 10 0 0 0 0 O 1 1 1 0 0 2 2 3 Q 0 0 0 0 6 0 0 0 Q 0 0 0 0 4 1 0 1 S 0 0 0 0 0 7 0 0 S 0 1 1 2 0 2 0 1 T 0 0 0 0 0 0 2 0 T 0 1 0 0 0 0 0 1 W 0 0 0 0 0 0 0 7 W 0 5 0 1 0 0 0 1

Table E-61. Proximal Ulna Social Behavior LDA: a) Discriminant Analysis. b) Cross Validation. Discriminant Analysis Cross Validation Social Behavior Predicted Group sum=74 Predicted Group Variable Variable Actual Group Group Pairs Solitary group Actual Group Group Pairs Solitary group group 6 0 0 0 group 0 0 1 5 pairs 0 5 2 0 pairs 1 1 5 0 solitary 0 1 49 2 solitary 1 10 33 8 variable group 1 0 1 14 variable group 4 3 5 4

216

Table E-62. K values for k-NN analysis without centroid size included and with centroid size included. Bolded values had highest percentage of accuracy. Proximal Complete Humerus Humerus Distal Humerus Complete Radius Proximal Radius

no no no no no centroid centroid centroid centroid centroid centroid centroid centroid centroid centroid Family-level 1 1 3 1 3 1 1 1 5 1 Locomotion 1 1 1 1 1 1 1 1 5 1 Hunting 1 1 1 1 1 1 4 1 2 1 Activity Pattern 1 1 1 1 5 1 1 1 5 4 Diet 1 1 1 1 1 1 1 1 4 3 Habitat 1 1 1 1 1 1 1 1 5 1 Social Behavior 1 1 4 1 1 1 5 1 3 4

Distal Radius Complete Ulna Proximal Ulna no no no centroid centroid centroid centroid centroid centroid Family-level 1 1 5 1 1 1 Locomotion 1 1 3 1 1 1 Hunting 1 1 3 1 1 1 Activity Pattern 1 1 1 1 1 1 Diet 3 1 1 1 1 2 Habitat 1 1 1 1 1 1 Social Behavior 1 1 1 1 1 1

217

Table E-63. Complete Humerus Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 0 0 Canidae 0 23 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 1 0 0 0 0 0 0 Felidae 0 0 1 48 0 0 0 2 0 1 Hyaenidae 0 1 0 0 6 0 0 0 0 0 Mephitidae 0 0 0 0 0 6 1 0 0 0 Mustelidae 0 0 0 0 0 1 27 3 0 1 Procyonidae 0 0 0 0 0 0 2 10 1 2 Ursidae 0 0 0 0 0 0 0 0 9 0 Viverridae 1 1 0 0 0 0 2 0 0 3

Table E-64. Complete Humerus Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 9 0 5 0 0 1 Cursorial 0 23 3 1 1 0 Scansorial 4 1 42 0 6 4 Semiaquatic 0 0 0 6 1 2 Semifossorial 0 2 3 2 13 2 Terrestrial 4 0 1 2 0 16

218

Table E-65. Complete Humerus Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 52 0 7 3 1 0 1 Aquatic 1 6 0 1 0 2 0 Occasional 7 0 21 3 0 1 0 Pounce/pursuit 0 1 2 23 2 0 1 Pursuit 0 0 0 1 3 0 0 Semifossorial 0 2 0 2 0 11 0 Various 0 0 0 0 0 0 0

Table E-66. Complete Humerus Activity Pattern k-NN. A C D N O A 17 2 1 10 3 C 3 3 1 4 1 D 2 0 12 1 2 N 7 4 0 52 9 O 4 0 2 4 10

Table E-67. Complete Humerus Diet k-NN. F I M O V F 2 1 0 0 0 I 1 6 2 1 0 M 1 2 72 15 3 O 0 3 12 30 1 V 0 0 0 0 2

219

Table E-68. Complete Humerus Habitat k-NN. D F N O Q S T W D 6 0 0 1 0 0 0 0 F 1 33 7 0 0 6 1 7 N 0 5 13 2 1 1 0 1 O 0 1 2 13 1 1 0 0 Q 0 0 2 0 4 0 0 0 S 1 11 4 1 1 8 1 0 T 0 1 1 0 0 0 2 0 W 0 4 1 0 0 2 0 7

Table E-69. Complete Humerus Social Behavior k-NN. Group Pairs Solitary Variable group Group 4 1 0 2 Pairs 2 8 1 3 Solitary 1 3 85 4 Variable group 6 3 5 26

220

Table E-70. Proximal Humerus Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 21 0 0 0 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 4 1 49 0 0 0 2 1 3 Hyaenidae 0 0 0 0 6 0 0 0 0 0 Mephitidae 0 0 0 0 0 3 0 1 0 0 Mustelidae 0 0 0 0 0 0 27 2 0 0 Procyonidae 2 0 0 0 0 4 5 10 0 2 Ursidae 0 0 0 0 0 0 0 0 9 0 Viverridae 0 0 0 0 0 0 0 0 0 2

Table E-71. Proximal Humerus Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 6 0 1 0 0 1 Cursorial 0 23 1 0 1 3 Scansorial 11 3 44 1 11 13 Semiaquatic 0 0 1 5 3 1 Semifossorial 0 0 5 2 6 3 Terrestrial 0 0 2 3 0 4

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Table E-72. Proximal Humerus Hunting Strategy k-NN. . Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 53 1 7 4 0 1 2 Aquatic 1 5 1 0 0 1 0 Occasional 2 1 18 3 0 3 0 Pounce/pursuit 4 2 0 20 3 1 0 Pursuit 0 0 0 3 3 0 0 Semifossorial 0 0 4 3 0 8 0 Various 0 0 0 0 0 0 0

Table E-73. Proximal Humerus Activity Pattern k-NN. A C D N O A 22 1 3 11 4 C 1 5 1 3 0 D 2 1 6 2 2 N 5 2 3 42 10 O 3 0 3 13 9

Table E-74. Proximal Humerus Diet k-NN. F I M O V F 2 0 0 1 0 I 0 1 1 1 1 M 2 6 68 19 2 O 0 5 16 25 0 V 0 0 1 0 3

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Table E-75. Proximal Humerus Habitat k-NN. D F N O Q S T W D 4 2 1 2 0 0 0 1 F 1 34 6 2 0 8 2 0 N 0 5 15 1 3 3 0 4 O 1 2 0 10 1 0 0 1 Q 0 0 2 1 2 0 0 0 S 0 7 3 0 0 6 0 0 T 2 3 1 0 0 0 2 0 W 0 2 2 1 1 1 0 9

Table E-76. Proximal Humerus Social Behavior k-NN. Group Pairs Solitary Variable group Group 3 0 1 5 Pairs 0 6 1 1 Solitary 4 6 86 11 Variable group 6 3 3 18

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Table E-77. Distal Humerus Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 24 0 0 0 0 0 0 0 1 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 0 48 0 0 0 2 1 2 Hyaenidae 0 0 0 0 6 0 0 0 0 0 Mephitidae 0 0 0 0 0 6 0 0 0 0 Mustelidae 2 0 0 0 0 0 32 1 0 0 Procyonidae 0 0 0 0 0 0 0 12 1 2 Ursidae 0 0 0 0 0 0 0 0 8 0 Viverridae 0 1 1 1 0 1 0 0 0 2

Table E-78. Distal Humerus Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 12 0 2 0 1 0 Cursorial 0 19 0 0 1 2 Scansorial 3 1 44 0 1 3 Semiaquatic 0 0 3 6 1 1 Semifossorial 1 5 0 1 13 5 Terrestrial 1 1 5 4 4 14

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Table E-79. Distal Humerus Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 53 0 5 1 0 3 1 Aquatic 0 6 0 3 0 0 0 Occasional 4 1 24 0 0 1 0 Pounce/pursuit 0 1 0 25 1 0 1 Pursuit 0 0 1 2 5 0 0 Semifossorial 2 1 0 1 0 10 0 Various 1 0 0 1 0 0 0

Table E-80. Proximal Humerus Activity Pattern k-NN. A C D N O A 18 1 1 8 6 C 1 7 1 1 0 D 2 1 11 1 1 N 7 0 2 50 8 O 5 0 1 11 10

Table E-81. Distal Humerus Diet k-NN. F I M O V F 2 0 0 0 0 I 1 7 2 1 0 M 0 2 72 15 1 O 1 3 11 29 2 V 0 0 1 1 3

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Table E-82. Distal Humerus Habitat k-NN.

D F N O Q S T W D 6 2 1 1 0 0 0 0 F 1 41 7 1 0 5 2 4 N 1 3 18 2 1 2 0 0 O 0 1 2 12 1 0 0 0 Q 0 1 0 1 4 0 0 0 S 0 4 2 0 0 10 0 1 T 0 0 0 0 0 0 2 0 W 0 3 0 0 1 1 0 10

Table E-83. Distal Humerus Social Behavior k-NN. Group Pairs Solitary Variable group Group 9 0 3 1 Pairs 1 8 4 4 Solitary 1 6 77 4 Variable group 2 1 7 26

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Table E-84. Complete Radius Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 1 1 0 0 Canidae 0 22 0 0 1 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 2 0 44 0 0 0 0 3 1 Hyaenidae 0 0 0 0 3 2 0 0 0 0 Mephitidae 0 0 0 0 0 3 2 0 0 0 Mustelidae 0 0 0 1 0 1 26 1 0 1 Procyonidae 1 0 0 0 0 1 1 12 1 2 Ursidae 0 0 0 1 0 0 1 0 6 0 Viverridae 1 1 1 1 1 0 1 2 1 3

Table E-85. Complete Radius Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 11 0 4 1 1 1 Cursorial 0 18 4 0 2 3 Scansorial 4 2 37 1 4 5 Semiaquatic 0 0 1 8 1 1 Semifossorial 0 0 2 1 8 2 Terrestrial 3 4 7 0 5 12

227

Table E-86. Complete Radius Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 50 2 5 6 0 1 0 Aquatic 0 6 0 1 0 0 0 Occasional 5 1 25 2 0 5 1 Pounce/pursuit 1 0 0 20 4 1 1 Pursuit 0 0 0 2 1 0 0 Semifossorial 2 0 2 2 0 7 0 Various 0 0 0 0 0 0 0

Table E-87. Complete Radius Activity Pattern k-NN. A C D N O A 23 0 0 6 2 C 2 2 1 1 1 D 0 1 11 8 0 N 7 5 4 48 14 O 1 1 0 6 9

Table E-88. Complete Radius Diet k-NN. F I M O V F 2 0 2 1 0 I 0 3 0 1 0 M 0 6 64 17 1 O 2 3 16 25 1 V 0 0 0 4 5

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Table E-89. Complete Radius Habitat k-NN. D F N O Q S T W D 5 0 2 0 0 1 1 0 F 1 29 6 5 0 8 1 5 N 2 5 14 2 0 1 0 1 O 0 2 3 9 1 0 0 0 Q 0 2 0 0 6 0 0 0 S 0 9 4 0 0 7 1 0 T 1 1 0 1 0 0 0 0 W 0 7 0 0 0 0 0 10

Table E-90. Complete Radius Social Behavior k-NN. Group Pairs Solitary Variable group Group 0 0 0 1 Pairs 1 5 3 0 Solitary 5 8 85 11 Variable group 6 3 3 22

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Table E-91. Proximal Radius Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 1 1 0 0 Canidae 0 20 0 0 2 0 1 1 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 1 44 1 0 0 0 0 1 Hyaenidae 0 3 0 0 2 0 0 0 0 0 Mephitidae 0 1 0 0 0 2 0 0 0 2 Mustelidae 1 1 0 0 0 2 24 1 0 0 Procyonidae 1 0 0 2 0 0 4 13 0 2 Ursidae 0 0 0 0 0 0 0 0 11 0 Viverridae 0 0 0 1 0 3 2 0 0 2

Table E-92. Proximal Radius Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 11 0 8 2 0 2 Cursorial 0 19 4 1 0 3 Scansorial 4 4 35 2 5 8 Semiaquatic 1 0 0 4 1 2 Semifossorial 0 1 2 1 12 3 Terrestrial 2 0 6 1 3 6

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Table E-93. Proximal Radius Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 47 0 1 4 0 2 1 Aquatic 1 4 1 1 0 1 0 Occasional 6 3 29 0 0 0 0 Pounce/pursuit 3 1 1 21 3 3 0 Pursuit 0 0 0 2 2 0 0 Semifossorial 0 0 0 5 0 7 1 Various 1 1 0 0 0 1 0

Table E-94. Proximal Radius Activity Pattern k-NN. A C D N O A 17 0 0 8 4 C 1 6 1 3 0 D 1 0 9 4 0 N 13 3 4 46 10 O 1 0 2 8 12

Table E-95. Proximal Radius Diet k-NN. F I M O V F 2 1 1 0 0 I 0 5 1 2 0 M 2 2 65 13 1 O 0 4 15 30 2 V 0 0 0 3 4

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Table E-96. Proximal Radius Habitat k-NN. D F N O Q S T W D 6 3 0 1 0 0 0 0 F 2 37 7 3 3 3 0 5 N 1 5 11 3 0 5 3 0 O 0 3 3 8 1 0 0 0 Q 0 1 2 0 2 0 0 0 S 0 3 4 2 1 8 0 1 T 0 0 1 0 0 0 0 0 W 0 3 1 0 0 1 0 10

Table E-97. Proximal Radius Social Behavior k-NN. Group Pairs Solitary Variable group Group 4 0 3 3 Pairs 0 7 1 3 Solitary 3 5 79 10 Variable group 5 4 8 18

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Table E-98. Distal Radius Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 1 0 0 0 0 0 0 0 1 0 Canidae 0 23 0 0 0 0 1 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 1 0 45 0 0 0 0 0 1 Hyaenidae 0 0 0 2 5 0 0 0 0 0 Mephitidae 0 0 0 0 0 5 3 3 0 0 Mustelidae 0 0 1 0 0 1 25 3 0 1 Procyonidae 0 0 0 0 0 1 2 10 0 0 Ursidae 1 0 0 0 0 0 1 0 10 0 Viverridae 0 1 0 0 0 0 0 0 0 5

Table E-99. Distal Radius Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 8 0 3 0 1 4 Cursorial 0 19 4 1 0 3 Scansorial 6 2 40 0 4 6 Semiaquatic 0 0 1 7 0 0 Semifossorial 0 1 1 3 14 1 Terrestrial 4 2 6 0 2 10

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Table E-100. Distal Radius Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 49 2 0 2 0 2 1 Aquatic 2 4 0 0 0 2 0 Occasional 1 1 27 1 0 1 0 Pounce/pursuit 3 1 3 26 3 0 1 Pursuit 1 0 0 1 2 0 0 Semifossorial 2 1 2 1 0 9 0 Various 0 0 0 2 0 0 0

Table E-101. Proximal Radius Activity Pattern k-NN. A C D N O A 23 0 0 9 0 C 2 4 0 3 1 D 1 0 8 2 4 N 6 4 4 51 5 O 1 1 4 4 16

Table E-102. Distal Radius Diet k-NN. F I M O V F 1 0 0 0 0 I 0 3 6 0 1 M 1 3 68 12 1 O 2 5 8 32 1 V 0 1 0 4 4

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Table E-103. Distal Radius Habitat k-NN. D F N O Q S T W D 5 0 1 1 1 0 0 0 F 2 37 7 7 0 5 1 7 N 0 5 17 0 1 0 0 1 O 0 4 1 9 1 2 0 0 Q 1 0 0 0 4 0 0 0 S 0 5 2 0 0 10 0 1 T 0 0 0 0 0 0 2 0 W 1 4 1 0 0 0 0 7

Table E-104. Distal Radius Social Behavior k-NN. Group Pairs Solitary Variable group Group 3 0 4 1 Pairs 1 8 4 2 Solitary 4 3 78 4 Variable group 4 5 5 27

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Table E-105. Complete Ulna Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 Canidae 0 23 0 4 2 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 2 2 1 42 1 0 1 5 1 3 Hyaenidae 0 0 0 0 1 1 0 0 0 0 Mephitidae 0 0 0 0 0 0 0 0 0 0 Mustelidae 0 1 0 0 0 6 30 1 0 1 Procyonidae 0 0 0 0 0 0 2 10 0 1 Ursidae 0 0 0 0 0 0 0 0 10 0 Viverridae 0 0 0 0 1 0 0 0 0 2

Table E-106. Complete Ulna Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 11 0 6 0 2 0 Cursorial 0 19 2 0 1 3 Scansorial 6 4 42 2 3 9 Semiaquatic 0 0 1 6 1 1 Semifossorial 0 0 2 3 12 2 Terrestrial 1 2 2 1 1 9

236

Table E-107. Complete Ulna Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 43 1 10 2 1 0 1 Aquatic 0 4 0 2 0 2 0 Occasional 3 0 21 4 0 1 1 Pounce/pursuit 10 3 0 22 1 2 0 Pursuit 0 0 0 2 3 0 0 Semifossorial 1 2 0 2 0 9 0 Various 0 0 1 0 0 0 0

Table E-108. Complete Ulna Activity Pattern k-NN. A C D N O A 25 0 0 5 3 C 1 5 3 2 1 D 1 1 2 5 0 N 4 3 8 43 11 O 1 0 4 15 11

Table E-109. Complete Ulna Diet k-NN. F I M O V F 0 0 1 0 0 I 2 6 3 0 0 M 2 5 67 18 1 O 0 1 12 29 1 V 0 0 0 1 5

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Table E-110. Complete Ulna Habitat k-NN. D F N O Q S T W D 1 0 2 2 0 0 0 0 F 2 41 5 3 0 8 0 6 N 2 3 14 2 2 2 0 0 O 1 2 3 7 2 0 0 0 Q 0 0 1 0 3 1 0 0 S 0 3 4 1 1 5 1 1 T 2 0 1 0 0 1 2 0 W 1 6 0 1 0 0 0 9

Table E-111. Complete Ulna Social Behavior k-NN. Variable Group Pairs Solitary group Group 7 0 6 0 Pairs 1 9 2 4 Solitary 4 1 76 12 Variable group 0 6 6 20

238

Table E-112. Proximal Ulna Family-level k-NN. Ailuridae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Procyonidae Ursidae Viverridae Ailuridae 2 0 0 0 0 0 0 0 0 0 Canidae 0 25 0 0 1 0 1 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 Felidae 0 1 1 46 1 0 0 0 0 1 Hyaenidae 0 0 0 0 3 0 0 0 0 1 Mephitidae 0 0 0 0 0 6 1 1 0 1 Mustelidae 0 0 0 0 0 0 31 0 0 0 Procyonidae 0 0 0 0 0 1 0 15 0 1 Ursidae 0 0 0 0 0 0 0 0 11 0 Viverridae 0 0 0 0 0 0 0 0 0 3

Table E-113. Proximal Ulna Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 12 0 3 0 0 0 Cursorial 1 20 1 1 0 2 Scansorial 3 4 46 2 4 6 Semiaquatic 0 0 1 7 1 2 Semifossorial 2 0 3 1 13 3 Terrestrial 0 1 1 1 2 11

239

Table E-114. Proximal Ulna Hunting Strategy k-NN. Ambush Aquatic Occasional Pounce/pursuit Pursuit Semifossorial Various Ambush 52 0 1 3 1 2 1 Aquatic 1 7 0 1 0 1 0 Occasional 1 1 29 1 0 1 0 Pounce/pursuit 1 1 1 28 2 0 0 Pursuit 0 1 0 0 2 0 1 Semifossorial 2 0 1 1 0 10 0 Various 0 0 0 0 0 0 0

Table E-115. Proximal Ulna Activity Pattern k-NN. A C D N O A 27 0 1 5 2 C 1 6 0 2 0 D 0 1 12 3 0 N 3 1 4 49 8 O 1 1 0 11 16

Table E-116. Proximal Ulna Diet k-NN. F I M O V F 2 0 0 0 0 I 0 9 1 3 0 M 2 1 74 11 1 O 0 2 8 31 0 V 0 0 0 3 6

240

Table E-117. Proximal Ulna Habitat k-NN. D F N O Q S T W D 4 4 2 3 0 0 0 0 F 3 40 6 1 2 1 0 4 N 0 1 18 2 1 2 0 0 O 2 3 0 7 1 0 0 0 Q 0 0 0 1 4 0 0 0 S 0 2 4 0 0 10 1 1 T 0 0 0 0 0 1 2 0 W 0 5 0 2 0 3 0 11

Table E-118. Proximal Ulna Social Behavior k-NN. Group Pairs Solitary Variable group Group 7 0 1 0 Pairs 0 10 3 6 Solitary 3 3 80 2 Variable group 2 3 6 28

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Figure E-1. Activity pattern PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

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Figure E-2. Activity pattern PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

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Figure E-3. Activity pattern PCAs for the ulna. A. complete ulna. B. proximal ulna.

244

Figure E-4. Diet PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

245

Figure E-5. Diet PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

246

Figure E-6. Diet PCAs for the ulna. A. complete ulna. B. proximal ulna.

247

Figure E-7. Social behavior PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

248

Figure E-8. Social behavior PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

249

Figure E-9. Social behavior PCAs for the ulna. A. complete ulna. B. proximal ulna.

250

Figure E-10. Habitat PCAs for the humerus. A. complete humerus. B. proximal humerus. C. distal humerus.

251

Figure E-11. Habitat PCAs for the radius. A. complete radius. B. proximal radius. C. distal radius.

252

Figure E-12. Habitat PCAs for the ulna. A. complete ulna. B. proximal ulna.

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APPENDIX F SUPPLEMENTAL INFORMATION FOR CHAPTER 4

Table F-1. Chapter 4 groupings. Abbreviations All 81 extant species and all fossil species TFcafemusthypro Thomas Farm canids and extant canids, felids, mustelids, hyaenids, and procyonids TFcafemusthy Thomas Farm canids and extant canids, felids, mustelids, and hyaenids TFcafe Thomas Farm canids and extant canids and felids

Table F-2. Best k-values for analyses

Best k values for analyses k values - no centroid k values - with centroid Family- Family-level Hunting Locomotion level Hunting Locomotion Humerus All 4 5 1 1 1 1 Humerus TFcafemusthy 5 1 1 1 1 2 Humerus TFcafe 1 2 3 1 3 4 Radius All 1 1 1 1 1 1 Radius TFcafemusthy 1 1 1 3 1 1 Radius TFcafe 3 1 4 1 1 1 Ulna All 1 1 1 1 1 1 UlnaTFcafemusthy 1 1 1 1 1 1 Ulna TFcafe 1 1 1 1 1 1

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Table F-3. Humerus “All” Family-level k-NN.

Ailuridae Barbourofelidae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Nimravidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 0 0 0 0 0 Barbourofelidae 0 0 0 0 0 0 0 0 0 0 0 0 Canidae 0 0 27 0 1 0 0 0 0 0 0 2 Eupleridae 0 0 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 2 1 49 0 2 0 1 2 0 2 Hyaenidae 0 0 1 0 0 6 0 0 0 0 0 0 Mephitidae 0 0 0 0 0 0 5 0 0 0 0 0 Mustelidae 2 0 0 0 0 0 0 32 0 0 0 0 Nimravidae 0 0 0 0 0 0 0 0 0 0 0 0 Procyonidae 0 1 0 0 0 0 0 0 0 13 1 2 Ursidae 0 0 0 0 0 0 0 0 0 0 9 0 Viverridae 0 0 1 0 0 0 0 0 0 0 0 1

Table F-4. Humerus TFcafemusthypro Family-level k-NN. Canidae Felidae Hyaenidae Mustelidae Procyonidae Canidae 25 0 0 0 0 Felidae 0 49 0 0 1 Hyaenidae 0 0 6 0 0 Mustelidae 0 0 0 32 0 Procyonidae 0 0 0 0 14

Table F-5. Humerus TFcafemusthy Family-level k-NN. Canidae Felidae Hyaenidae Mustelidae Canidae 25 0 0 0 Felidae 0 49 0 0 Hyaenidae 0 0 6 0 Mustelidae 0 0 0 32

255

Table F-6. Humerus TFcafe Family-level k-NN. Canidae Felidae Canidae 25 0 Felidae 0 49

Table F- . Radius “All” Family-level k-NN.

Ailuridae Barbourofelidae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Nimravidae Procyonidae Ursidae Viverridae Ailuridae 0 0 0 0 0 0 0 1 0 1 0 0 Barbourofelidae 0 2 0 0 0 0 0 1 0 0 1 0 Canidae 0 0 25 0 1 2 1 0 0 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 4 0 45 0 0 0 0 0 2 1 Hyaenidae 0 0 1 0 0 3 1 0 0 0 0 0 Mephitidae 0 0 0 0 0 0 3 2 0 0 0 0 Mustelidae 0 0 0 0 1 0 1 25 1 1 1 0 Nimravidae 0 0 0 0 0 0 0 0 0 0 0 0 Procyonidae 1 0 0 0 0 0 1 2 0 12 1 2 Ursidae 0 0 1 0 0 0 0 0 0 0 6 0 Viverridae 1 0 0 1 2 0 0 1 0 2 0 4

Table F-8. Radius TFcafemusthy Family-level k-NN. Canidae Felidae Hyaenidae Mustelidae Canidae 22 1 1 0 Felidae 3 45 0 0 Hyaenidae 0 0 4 0 Mustelidae 0 1 0 32

Table F-9. Radius TFcafe Family-level k-NN. Canidae Felidae Canidae 24 1 Felidae 1 46

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Table F-10. Ulna “All” Family-level k-NN.

Ailuridae Barbourofelidae Canidae Eupleridae Felidae Hyaenidae Mephitidae Mustelidae Nimravidae Procyonidae Ursidae Viverridae Ailuridae 2 0 0 0 0 0 0 0 0 0 0 0 Barbourofelidae 0 2 0 0 0 1 0 0 0 0 0 0 Canidae 0 0 27 0 0 1 0 1 1 0 0 0 Eupleridae 0 0 0 0 0 0 0 0 0 0 0 0 Felidae 0 0 2 1 48 1 0 0 0 0 0 1 Hyaenidae 0 0 0 0 0 2 0 0 0 0 0 1 Mephitidae 0 0 0 0 0 0 6 1 0 0 0 0 Mustelidae 0 0 0 0 0 0 0 31 0 0 0 0 Nimravidae 0 0 1 0 0 0 0 0 0 0 0 0 Procyonidae 0 0 1 0 0 0 1 0 0 16 0 2 Ursidae 0 0 0 0 0 0 0 0 0 0 11 0 Viverridae 0 0 0 0 0 0 0 0 0 0 0 3

Table F-11. Ulna TFcafemusthy Family-level k-NN. Canidae Felidae Hyaenidae Mustelidae Canidae 25 0 1 1 Felidae 1 45 1 0 Hyaenidae 0 1 3 0 Mustelidae 0 0 0 32

Table F-12. Ulna TFcafe Family-level k-NN. Canidae Felidae Canidae 25 0 Felidae 1 46

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Table F-13. Humerus “All” Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Various Ambush 54 0 5 2 0 3 1 Aquatic 0 6 0 3 0 0 0 Occasional 4 1 25 0 0 1 0 Pursuit & Pounce 0 1 0 25 1 0 1 Pursuit 0 0 0 2 5 0 0 Semifossorial 2 1 0 0 0 10 0 Various 0 0 0 1 0 0 0

Table F-14. Humerus TFcafemusthypro Hunting Stratgey k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Ambush 52 0 3 0 0 2 Aquatic 0 6 0 3 0 0 Occasional 3 1 13 0 0 0 Pursuit & Pounce 0 1 0 25 1 0 Pursuit 0 0 0 2 5 0 Semifossorial 2 1 0 0 0 7

Table F-15 Humerus TFcafemusthy Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Ambush 55 0 0 0 0 2 Aquatic 0 7 1 3 0 0 Occasional 0 0 0 0 0 0 Pursuit & Pounce 0 1 0 25 1 0 Pursuit 0 0 0 2 5 0 Semifossorial 2 1 0 0 0 7

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Table F-16. Humerus TFcafe Hunting Strategy k-NN. Ambush Pursuit & Pounce Pursuit Ambush 49 0 0 Pursuit & Pounce 0 21 1 Pursuit 0 1 2

Table F-1 . Radius “All” Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Various Ambush 49 0 6 4 0 1 0 Aquatic 0 8 0 1 0 1 0 Occasional 4 0 22 4 0 3 1 Pursuit & Pounce 5 1 3 23 4 3 1 Pursuit 0 0 0 0 1 1 0 Semifossorial 0 0 1 1 0 5 0 Various 0 0 0 0 0 0 0

Table F-18. Radius TFcafemusthy Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Ambush 51 0 0 5 0 1 Aquatic 0 8 0 1 0 1 Occasional 0 0 0 1 0 0 Pursuit & Pounce 4 1 1 21 4 2 Pursuit 0 0 0 1 1 0 Semifossorial 0 0 0 1 0 5

Table F-19. Radius TFcafe Hunting Strategy k-NN. Ambush Pursuit & Pounce Pursuit Ambush 47 2 0 Pursuit & Pounce 0 20 2 Pursuit 0 0 1

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Table F-20. Ulna “All” Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Various Ambush 52 0 3 3 1 1 1 Aquatic 1 7 0 0 0 2 0 Occasional 1 1 28 1 0 1 0 Pursuit & Pounce 1 2 1 29 2 0 0 Pursuit 0 0 0 0 2 0 1 Semifossorial 2 0 0 1 0 10 0 Various 0 0 0 0 0 0 0

Table F-21. Ulna TFcafemusthy Hunting Strategy k-NN. Ambush Aquatic Occasional Pursuit & Pounce Pursuit Semifossorial Ambush 49 0 0 2 1 0 Aquatic 1 8 0 1 0 0 Occasional 0 0 0 0 0 0 Pursuit & Pounce 1 2 1 27 2 0 Pursuit 1 0 0 1 2 0 Semifossorial 2 0 0 0 0 9

Table F-22. Ulna TFcafe Hunting Strategy k-NN. Ambush Pursuit & Pounce Pursuit Ambush 46 0 0 Pursuit & Pounce 0 21 1 Pursuit 0 2 2

260

Table F-23. Humerus “All” Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 13 0 3 0 1 0 Cursorial 0 18 0 0 1 2 Scansorial 3 1 43 0 1 4 Semiaquatic 0 0 2 5 1 1 Semifossorial 1 6 1 1 11 4 Terrestrial 0 1 5 5 6 14

Table F-24. Humerus TFcafemusthypro Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 2 0 0 0 Cursorial 0 18 0 1 1 1 Scansorial 1 0 39 0 0 4 Semiaquatic 0 0 0 7 0 1 Semifossorial 0 7 0 1 8 4 Terrestrial 1 1 5 1 3 11

Table F-25. Humerus TFcafemusthy Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 4 0 0 1 0 0 Cursorial 0 18 0 1 1 2 Scansorial 1 1 33 0 0 4 Semiaquatic 0 0 0 7 0 1 Semifossorial 0 6 0 0 7 3 Terrestrial 0 1 5 1 4 11

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Table F-26. Humerus TFcafe Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 4 0 0 0 0 0 Cursorial 0 18 1 0 1 3 Scansorial 0 1 30 0 1 3 Semiaquatic 0 0 0 0 0 0 Semifossorial 0 0 0 0 0 1 Terrestrial 0 1 2 1 1 6

Table F-2 . Radius “All” Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 10 0 5 1 0 1 Cursorial 0 19 4 0 2 3 Scansorial 6 1 35 0 5 3 Semiaquatic 0 0 1 8 1 1 Semifossorial 0 0 4 1 7 2 Terrestrial 2 4 6 1 6 14

Table F-28. Radius TFcafemusthy Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 1 0 2 0 0 0 Cursorial 0 18 4 0 0 3 Scansorial 3 2 30 1 1 6 Semiaquatic 0 0 0 8 1 1 Semifossorial 0 0 0 0 4 1 Terrestrial 1 4 3 1 5 9

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Table F-29. Radius TFcafe Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 1 0 1 0 0 0 Cursorial 0 15 3 0 0 3 Scansorial 3 3 29 1 2 6 Semiaquatic 0 0 0 0 0 0 Semifossorial 0 0 0 0 0 0 Terrestrial 0 1 1 0 0 3

Table F-30. Ulna “All” Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 14 0 2 0 0 0 Cursorial 0 20 3 1 0 4 Scansorial 3 4 48 1 2 5 Semiaquatic 1 0 0 8 2 1 Semifossorial 0 0 1 1 12 6 Terrestrial 0 1 1 1 4 8

Table F-31. Ulna TFcafemusthy Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 3 0 0 0 0 0 Cursorial 0 20 3 1 0 4 Scansorial 1 4 32 0 0 4 Semiaquatic 0 0 0 9 0 1 Semifossorial 0 0 0 0 7 6 Terrestrial 1 1 3 1 4 5

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Table F-32. Ulna TFcafe Locomotion k-NN. Arboreal Cursorial Scansorial Semiaquatic Semifossorial Terrestrial Arboreal 3 0 1 0 0 0 Cursorial 0 17 1 0 0 2 Scansorial 1 3 29 1 1 3 Semiaquatic 0 0 0 0 0 0 Semifossorial 0 0 1 0 0 1 Terrestrial 0 0 1 0 1 6

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BIOGRAPHICAL SKETCH

Sharon Holte has a PhD from the University of Florida (2018), a Master of

Science from East Tennessee State University (2012), and a Bachelor of Science from

South Dakota School of Mines and Technology (2009). She was a guest editor on a

Quaternary International volume entitled: Quaternary Proboscideans: Papers in Memory of Larry D. Agenbroad. She was nominated and elected as a Science Associate of The

Mammmoth Site of Hot Springs, South Dakota. Science Associates are scientific professionals that acts as an advisory committee for researchers at The Mammoth Site.

Sharon received an NSF Doctoral Dissertation Improvement Grant for her work on early Miocene carnivorans. She was also an NSF GK-12 Fellow for the academic years of 2009-2010 and 2010-2011. In her first year as a Fellow, she worked with

Prekindergarten and Kindergarten teachers. During the second year, Sharon worked with third grade teachers at the Title 1 North Side Elementary School in Johnson City,

TN. She developed and taught state standard based science lessons weekly in three classrooms and posted them on a GK-12/Teacher online resource center. Through these two years, Sharon developed and taught weekly science lesson plans for the media center that were adaptable to all grade levels at the elementary school. One of

Sharon’s passions is public outreach and education; through this interest, she has been invited to provide multiple presentations to public events and fossil clubs.

Sharon’s first experience excavating at a fossil site was as a volunteer, so she understands the importance that volunteers provide for research. Through her multiple years of field experience, she has worked with many different people from a wide range of ages and backgrounds. These volunteers and citizen scientists have helped her to learn how to communicate science affectively to the general public. Some of her field

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experience has included among others: Montbrook Fossil Site, North central FL; 2016-

2017 (Excavator); Persistence Cave, Wind Cave National Park, SD, June 2015 (Site

Co-PI); Thomas Farm Fossil Site, Northern FL, Spring 2013 (Site Foreman); Panama

Canal, Panama, Fall 2012 (Excavator); Gray Fossil Site, Gray TN, 2009 -2012

(Excavator); Saltville, VA, Summer 2009 (Excavator), Fossil Lake, OR, Summer 2008

(Excavator/prospector); Geology Field Camp in Turkey, SDSMT, Summer 2008; Wind

Cave National Park, SD, Fall 2007 (Park Service Volunteer). She plans to continue doing education and outreach through paleontology as an educator at The Mammoth

Site of Hot Springs, South Dakota.

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