Scimitar Cat ( Homotherium serum Cope) from · 2017. 11. 13. · 113 teeth,” (Rawn-Schatzinger...

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Scimitar Cat (Homotherium serum Cope) from

Southwestern Alberta, Canada

Journal: Canadian Journal of Earth Sciences

Manuscript ID cjes-2017-0130.R1

Manuscript Type: Article

Date Submitted by the Author: 10-Aug-2017

Complete List of Authors: Ewald, Tatyanna; University of Calgary, Department of Anthropology and Archaeology Hills, Len; The University of Calgary, Department of Geoscience Tolman, Shayne; Independent Kooyman, Brian; University of Calgary , Department of Anthropology and Archaeology

Is the invited manuscript for consideration in a Special

Issue? : N/A

Keyword: <i>Homotherium serum </i>, Homotheriini, Dentition, Wally's Beach, Pleistocene

https://mc06.manuscriptcentral.com/cjes-pubs

Canadian Journal of Earth Sciences

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Scimitar Cat (Homotherium serum Cope) from 1

Southwestern Alberta, Canada 2

Tatyanna Ewald, L.V. Hills*, Shayne Tolman, and Brian Kooyman 3

*Deceased 4

T. Ewald. (email: [email protected]) Department of Anthropology and Archaeology, 5

University of Calgary, 2500 University Dr. NW, Calgary, AB T2N 1N4, Canada. 6

L.V Hills. * Department of Geoscience, University of Calgary, 2500 University Dr. N.W., 7

Calgary, AB T2N 1N4, Canada 8

S. Tolman. (email: [email protected]) Box 1146, Cardston, AB TOK 0Y0, Canada. 9

B. Kooyman. (email: [email protected]) Department of Anthropology and Archaeology, 10

University of Calgary, 2500 University Dr. NW, Calgary, AB T2N 1N4, Canada. 11

Corresponding author: 12

Tatyanna Ewald 13

Email: [email protected] 14

Phone: 1-403-827-2245 15

Fax: 1-403-284-5467 16

Address: Department of Anthropology and Archaeology 17

2500 University Drive NW 18

T2N 1N4 19

Canada 20

*Deceased. 21

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Abstract 22

Skull and tooth fragments of Homotherium serum recently recovered from the Wally’s Beach 23

site (DhPg-8) in southwestern Alberta provide the first indications that scimitar cat populated the 24

area of the St. Mary Reservoir. AMS radiocarbon dating provides a 2 sigma range calibrated age 25

of Cal BP 12 715 to 12 655. This is the fourth known occurrence of the species in Canada, the 26

first outside of Yukon, and is currently the youngest precisely dated occurrence of the species in 27

North America. Well-preserved dentition combined with the temporal and geographic context 28

allows the sample to be identified as Homotherium serum. The specimen is significant as it 29

represents an extension of the geographic and chronological range of the species. 30

Keywords 31

Homotherium serum, Homotheriini, Dentition, Wally’s Beach, Pleistocene 32

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Introduction 33

The Wally’s Beach site, near Cardston in southwestern Alberta, consists of a series of 34

Late Pleistocene paleontological and archaeological remains which were exposed by erosion 35

associated with the operation of the St. Mary Reservoir (Hills et al. 2014; Kooyman et al. 2012; 36

Waters et al. 2015). Species present include Equus conversidens, Camelops hesternus, 37

Bootherium bombifrons, Rangifer tarandus, and Bison antiquus among others. A number of 38

species are also represented by preserved tracks, including Mammuthus (McNeil et al. 39

2005:1255). Recently, skull and tooth fragments of an individual scimitar cat, Homotherium 40

serum, were discovered. The purpose of this paper is to describe and illustrate this specimen, and 41

to discuss its location and date in relation to other known occurrences of the species. 42

Geological Setting and Context 43

The Wally’s Beach site (also known by its Borden designation DhPg-8) is located in 44

southwestern Alberta near the town of Cardston (Fig. 1), in the drawdown zone of the St. Mary 45

Reservoir (Hills et al. 2014). While the H. serum remains are paleontological, the Wally’s Beach 46

site is predominately an archaeological site and is therefore defined by the Borden system. The 47

Borden system divides Canada into a series of small blocks inside larger blocks based on latitude 48

and longitude. The capital letters in the Borden designation identify the large block, while the 49

lower-case letters denote the small block to which the site belongs. Each archaeological site is 50

then designated sequentially within these blocks (Borden 1954). 51

Cranial remains of H. serum were discovered in-situ, within a 16-18cm radius. The 52

location consists of Late Pleistocene loess and eolian sands, upon which a Holocene paleosol 53

developed (Hills et al. 2014:25) The specimen was recovered from the very top portion of the 54

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Pleistocene sediments, embedded in the carbonate horizon that resulted from the development of 55

the overlying paleosol. The enamel layer of one of the canines shows extensive fibrous root 56

etchings on the surface (Fig. 2). This suggests that this tooth was incorporated in the soil horizon 57

while it was actively forming, before the soil’s final burial. 58

A bone sample from the skull provides an AMS (Beta 440230) 2 sigma range calibrated 59

age of Cal BP 12 715 to 12 655 (conventional age of BP 10 740 ±40). This date was calibrated 60

using IntCal13. The CN (carbon:nitrogen) ratio of +3.3 is within the normal range expected for 61

live protein, therefore the collagen was well-preserved (the δ13C value of -19.3 o/oo and δ15N 62

value of +6.6 o/oo are also both within the normal range expected for live protein). This date 63

places the specimen in the Late Pleistocene and makes it the youngest dated specimen of 64

Homotherium serum in Canada. 65

Materials and Methods 66

The following description is based on fragments of five incisors, two upper canines, the 67

left lower canine, the upper fourth premolar, the lower first molar, and an unidentifiable root 68

(Fig. 3). Preservation is largely restricted to the enamel layer of each tooth. The canine fragments 69

were measured using electronic calipers providing digital readouts to the nearest millimetre (see 70

Dental Remains Recovered). Several small skull fragments (DhPg-8-3847) are also present 71

within the assemblage, but were not considered in this study. The skull fragments are too small 72

and fragmented for partial skull reconstruction, therefore the teeth were the focus of this analysis. 73

The identification of the tooth fragments was reliant predominately upon Rawn-74

Schatzinger (1983; 1992), although several other sources (Antón et al. 2014; Biknevicius et al. 75

1996; Cope 1893; Martin et al. 2011a; Martin et al. 2011b; Martin et al. 2011c; Martin et al. 76

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2011d; Wheeler 2011) and some comparisons to modern Felidae taxa (Felis concolor, Lynx 77

canadensis, and Leopardus pardalis, Department of Anthropology and Archaeology, University 78

of Calgary) were also utilized. 79

Results 80

Systematic Paleontology 81

We follow the systematics proposed by Antón et al. (2014). 82

Mammalia Linnaeus, 1758 83

Order Carnivora Bowdich, 1821 84

Suborder Feliformia Kretzoi, 1945 85

Family Felidae Fischer von Waldheim, 1817 86

Subfamily Machairodontinae Gill, 1872 87

Tribe Homotheriini Kurtén, 1962 88

Genus Homotherium Fabrini, 1890 89

Homotherium serum (Cope 1893) 90

Referred Specimens 91

The material used in this study will be stored in the Archaeology collections at the Royal 92

Alberta Museum: DhPg-8-3836, crown of the upper left canine; DhPg-8-3837, crown of the 93

upper right canine; DhPg-8-3838, fragment of the crown from an upper first or second incisor; 94

DhPg-8-3839, fragment of the crown from the lower right second incisor; DhPg-8-3840, 95

fragment of the crown from the lower left second incisor; DhPg-8-3841, fragment of the crown 96

from an upper incisor; DhPg-8-3842, fragment of the crown from an unidentified incisor; DhPg-97

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8-3843, crown of the lower left canine; DhPg-8-3844, metacone of the upper left fourth 98

premolar; DhPg-8-3845, paracone of the lower right first molar; DhPg-8-3846, partial root; 99

DhPg-8-3847, several small skull fragments. Cataloguing of these materials followed the Borden 100

system. 101

Dental Remains Recovered 102

The Homotherium genus is particularly characterized and separated from other saber-103

tooth cats by its dentition, and this is particularly useful in distinguishing Homotherium from its 104

close Smilodon relatives, also present in North America during the Pleistocene. A total of 11 105

tooth fragments identified as belonging to Homotherium serum were recovered at Wally’s 106

Beach. The teeth recovered indicate that only a single individual is represented; no elements are 107

duplicated and the size and wear on the teeth are consistent for a single individual. In addition, 108

the remains were discovered in-situ. Distinct, fine serrations are present along the cutting edge of 109

each tooth and are characteristic of H. serum, as serrations are present on all teeth of both 110

juvenile and adult specimens (Rawn-Schatzinger 1983). “The term “finely serrate” as used by 111

Cope (1983) and Churcher (1966) refers to the consistent and regular serration of H. serum’s 112

teeth,” (Rawn-Schatzinger 1992:4). Within this specimen, serrations are most clearly pronounced 113

on the upper canines (Fig. 4), upon which they are present along the entire length of the anterior 114

and posterior edge (Rawn-Schatzinger 1983; 1992; Wheeler 2011:28). 115

The most distinctive of the teeth recovered from Wally’s Beach are the upper canines 116

(Fig. 5), of which only the enamel caps are present. As noted by Rawn-Schatzinger (1983:53), 117

"the permanent canine has one-half of its length covered with enamel.” As such, it is reasonable 118

to assume that these enamel caps represent approximately one-half of the original length of each 119

tooth. The fragment of the left tooth (DhPg-8-3836) is 51.8 mm in length and 20.1 mm wide 120

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antero-posteriorly at its widest aspect, while the right (DhPg-8-3837) is 48.2 mm long and 18.7 121

mm wide. Medio-laterally, the left tooth is 8.3 mm wide and the right is 8.4 mm, once again at 122

their widest aspects. This corresponds with descriptions by Rawn-Schatzinger (1983:52; 1992:3) 123

of H. serum upper canines which are laterally compressed or flat as compared to conical-tooth 124

cats. Also unique to Homotherium, and exhibited in this specimen, is that the canines are 125

relatively short and broad in comparison to other saber-tooths (Martin et al. 2011a:6; Martin et 126

al. 2011d:208; Wheeler 2011:27) and that they are more complex on their lingual surface (Rawn-127

Schatzinger 1983:52). Both teeth are relatively straight with a very slight posterior curvature. 128

The convex nature of the posterior as opposed to the anterior face of the upper canines, as in this 129

specimen, was noted by Cope (1893:897). Both canines exhibit heavy post-depositional root 130

etching. 131

Several incisor fragments were also identified. All display a slight posterior convex 132

curvature and all are procumbent and rather robust, as is characteristic of Homotherium (Martin 133

et al. 2011a:8; Wheeler 2011:28). The first fragment (Fig. 6A; DhPg-8-3838) is comprised of the 134

anterior portion of the enamel cap. Partial cuspules are present on both lateral sides of this 135

fragment, allowing it to be identified as either the upper first or second incisor (Rawn-136

Schatzinger 1983:53), although the side is indeterminable in this state. An additional incisor 137

fragment (Fig. 6B; DhPg-8-3839) consists predominately of the posterior and medial portion of 138

the enamel cap. Basal cuspules are present on both lateral sides of the fragment, one partial and 139

the other complete. This tooth has therefore been identified as the lower second incisor (Rawn-140

Schatzinger 1983:53). The medial cuspule is noticeably higher on the tooth than the lateral, 141

indicating that this is the right second incisor (Martin et al. 2011b: Figure 4.8). Yet another 142

incisor fragment, consisting predominately of the posterior and lateral portion of the enamel cap, 143

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exhibits mirrored identical features (Fig. 6C; DhPg-8-3840). This tooth has been identified as the 144

lower left second incisor. 145

Yet another obviously incisiform fragment consists of a lateral portion of the enamel cap, 146

and terminates just below a basal cuspule (Fig. 6D; DhPg-8-3841). This tooth has been identified 147

as part of the upper incisor dentition; however further identification is not possible. It is larger 148

than the lower second incisors already identified for this specimen and is therefore not a lower 149

first incisor, as the lateral incisors are typically larger and more robust in both saber-tooth cats 150

and modern carnivores (Biknevicius et al. 1996). The obvious cuspule present on this fragment 151

also excludes it from being identified as the lower third incisor, as this tooth lacks basal cuspules 152

in H. serum (Rawn-Schatzinger 1983:52). This incisor is therefore part of the upper dentition, 153

however is not identifiable within a sequence. A fifth incisor fragment (DhPg-8-3842) is poorly 154

preserved and further identification is not possible. 155

Also present within this sample is a lower canine (Fig. 7; DhPg-8-3843), of which the 156

entire crown appears to be present. As noted by Rawn-Schatzinger (1983:53), enamel covers 157

approximately one-third of this tooth, which indicates that this fragment is likely representative 158

of one-third of the original length of the tooth. Although the lower canines of Homotherium are 159

diagnostically reduced and incisor-like (Biknevicius et al. 1996:518; Wheeler 2011:32), at 19.6 160

mm long this tooth fragment is noticeably larger than all the incisor fragments present in this 161

sample, which is typical in H. serum (Rawn-Schatzinger 1983:50). In their description of 162

Homotherium serum, Antón et al. (2014:263) state that in the occlusal view the lower canine has 163

“a strong lateral flattening, and a small size, just slightly larger than the i3.” This lateral 164

compression is also exhibited in this specimen and is also characteristic of H. serum (Rawn-165

Schatzinger 1983:53). It is also much smaller than the upper canines, which would allow the cat 166

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more clearance when biting (Wheeler 2011:29). A small basal cuspule located on its medial 167

aspect (Rawn-Schatzinger 1983:53) indicates that this tooth is the left. This specimen also 168

exhibits a distinct convex posterior curvature which is more pronounced than in the upper 169

canines. 170

A single cusp of the upper fourth premolar, or the upper carnassial (Fig. 8; DhPg-8-3844) 171

is also present. This cusp is identifiable as such because the edge is nearly straight, allowing it to 172

function in its role as a specialized cutting tooth, a characteristic unique to Homotherium (Rawn-173

Schatzinger 1983:52; Rawn-Schatzinger 1992:3). The shape of this specimen also indicates that 174

it is the metacone, which is the only cusp within H. serum which is distinctively flat (Rawn-175

Schatzinger 1983). The main ridge of the cusp leans slightly medially and two slight bulges are 176

present on the buccal aspect of the tooth, with a slight vertical groove between them (Cope 177

1893:897). Combined with indications of post-mortem breakage on the anterior portion of the 178

cusp and a lack of such evidence on the posterior portion, this fragment has been identified as the 179

most posterior cusp (metacone) of the left upper carnassial (Rawn-Schatzinger 1983:54). The 180

identification of this tooth was corroborated by comparisons to modern Felidae samples (Felis 181

concolor, Lynx canadensis, and Leopardus pardalis, Department of Anthropology and 182

Archaeology, University of Calgary). 183

A cusp with a pointed and rather symmetrical apex was also recovered (Fig. 9; DhPg-8-184

3845). The lower carnassial, or lower first molar, of Homotherium is distinguished by its two 185

blade-like cusps (Rawn-Schatzinger 1983:54). The paracone in particular is both large and 186

vertical (Martin et al. 2011c:193), indicating that this fragment is the paracone of the lower 187

carnassial. The presence of two vertical concavities on the lingual aspect of the cusp, as well as 188

indications of post-mortem breakage on the posterior but not anterior aspect of the cusp, further 189

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indicates that this is a fragment of the right carnassial. Comparisons with modern Felidae 190

samples (Felis concolor, Lynx canadensis, and Leopardus pardalis, Department of Anthropology 191

and Archaeology, University of Calgary) corroborated this identification. Shearing wear is also 192

present on both carnassial fragments; this is particularly well-preserved on the fragment of the 193

lower carnassial (see Fig. 9A). 194

The final fragment is a partial root (Fig. 10; DhPg-8-3846). Three separate roots are 195

identifiable on this fragment, the middle of which projects outwards from the other two. Based 196

on comparisons with modern Felidae samples (Felis concolor, Lynx canadensis, and Leopardus 197

pardalis, Department of Anthropology and Archaeology, University of Calgary), it is likely that 198

this belongs to a molar or premolar of Homotherium serum. As this fragment is not diagnostic, 199

further evaluation of the tooth roots in Homotherium is beyond the scope of this study. 200

In the present material, the upper canines are lenticular in cross section with serrations 201

present on both the anterior and posterior edges. The canines are blunted with numerous flake 202

scars and the serrations are polished and worn (Fig. 11), indicative of a mature individual. Rawn-203

Schatzinger (1983) subdivided the scimitar cat into an eight-stage age progression ranging from 204

Stage I, deciduous teeth just erupting beyond the alveolus, to Stage VIII, with extensive wear and 205

teeth blunted by wear polish. These last features are visible on this specimen; hence it is 206

classified as a Stage VIII adult. Rawn-Schatzinger (1983) did not provide an age for Stage VIII 207

adults, however that for Stage VII is two or more years, which equates to the approximate age 208

for complete functional permanent dentition. No teeth in this specimen are in an extreme state of 209

wear evidenced by broken teeth or teeth worn to the pulp cavity (Rawn-Schatzinger 1983:52).210

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Discussion 211

The Wally’s Beach scimitar cat specimen provides a significant new addition to the Late 212

Pleistocene paleofauna of southwestern Alberta, Canada. It is the fourth known occurrence of H. 213

serum in Canada (Debicki 1983:24; Harington 1977:526-528, 529-531; Harington 1989:95; 214

Harington 1997; Harington 2003:385; Rawn-Schatzinger and Collins 1981:18) and the first in 215

Alberta. The calibrated age of Cal BP 12 715 to 12 655 is corroborated by the specimen’s 216

stratigraphic context and makes this the youngest precisely dated occurrence of this species in 217

North America. 218

Due to the fragmented nature of the skull fragments recovered, an analysis of the 219

specimen recently recovered from Wally’s Beach was entirely dependent upon the partial 220

dentition. While the distinctive tooth morphology (e.g., fine serrations present along the cutting 221

edge of all teeth) exhibited here was the basis for identifying this specimen, the geographic and 222

temporal context of the finds supports this identification. 223

The saber-tooth cats were common primarily during the Pliocene and Pleistocene, periods 224

correspondingly characterized by large prey species which these cats were well adapted to hunt. 225

Homotherium is recognized as a wide-ranging genus from throughout Europe, Asia, Africa, and 226

North America (Rawn-Schatzinger 1992), and as the most abundant scimitar cats from northern 227

Eurasia and North America (Martin et al. 2011d). Recently, a specimen from Venezuela 228

(Homotherium venezuelensis) has also been described (Rincón et al. 2011), adding South 229

America to the range of Homotherium and making it the most geographically dispersed genus of 230

the saber-tooth cats (Antón et al. 2014). Homotherium was the only saber-tooth cat that achieved 231

such a wide range (Martin et al. 2011a). 232

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The genus Homotherium is the last of a long line of machairodont Homotheriini, ranging 233

in age between four million and 10,000 years (Antón et al. 2014). Their extinction correlated 234

with megafauna extinctions as well as climatic fluctuations (Martin et al. 2011a). Diagnosis of 235

species within Homotherium remains a debated issue among researchers. While a single species 236

tends to be recognized for Europe and Asia (Homotherium latidens), there are several 237

classifications of North American species (Antón et al. 2014). Homotherium ischyrus is often 238

described as an earlier North American species recovered most often from Pliocene and early 239

Pleistocene contexts, while Homotherium serum characterizes the Late Pleistocene of North 240

America (Martin et al. 2011c). Homotherium serum, while abundant primarily during the later 241

stages of the Pleistocene, was not uncommon earlier in this epoch (Antón et al. 2014). 242

While both H. ischyrus and H. serum occupied North America, H. serum characterized 243

the Late Pleistocene and may have been the only scimitar species on the continent at the time 244

(Antón et al. 2014). The distribution of Homotherium in North America is predominately within 245

the Camelops Faunal Province, “which was considered to be composed mostly of pine parkland 246

and characterized by the most cursorially adapted large mammals,” (Martin et al. 2011d:206). 247

Homotherium serum is known from over 30 sites in North America which are concentrated in the 248

United States (see Fig. 1; Table 1). Only three sites within Canada (not including Wally’s 249

Beach) are currently known to contain H. serum remains (Debicki 1983:24; Harington 1977:526-250

528, 529-531; Harington 1989:95; Harington 1997; Harington 2003:385; Rawn-Schatzinger and 251

Collins 1981:18). As the only known specimen in Alberta, the remains from Wally’s Beach 252

represent an extension of the range of H. serum. Furthermore, as the youngest occurrence in 253

Canada by a significant margin, this specimen may represent a Late Pleistocene extension of the 254

species northward into modern Canada from the United States. 255

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The calibrated age of Cal BP 12 715 to 12 655 for this specimen corroborates its 256

identification as H. serum, which characterized the Late Pleistocene of North America, rather 257

than the much older H. ischyrus (Antón et al. 2014). More significantly, this specimen is the 258

youngest known precisely dated occurrence of the species in North America. In Canada, three 259

other specimens have been identified from Yukon, however these are significantly older than that 260

recovered from Wally’s Beach. These Yukon specimens date to over 40,000 years BP at 261

Sixtymile Loc. 3 (the only Yukon specimen which has been precisely dated) (Harington 1997; 262

Harington 2003:385), over 25,000 years BP at Old Crow Loc. 21 (Harington 1977:526-528; 263

Harington 1989:95), and 39,900 to 22,200 years BP at Dawson Loc. 9 (Debicki 1983:24; 264

Harington 1977:529-531; Rawn-Schatzinger and Collins 1981:18). In North America more 265

broadly, most specimens date to an earlier Pleistocene context (see Table 1). Only one site 266

contains specimens which have been precisely radiocarbon dated to the last 15,000 years: 267

Laubach Cave No. 2, Texas (13,970 years BP; Graham and Lundelius 2010). The Wally’s Beach 268

specimen is therefore currently the most recent precisely dated specimen in North America and 269

may be a useful indicator of the survival of the species into the Younger Dryas (as defined in 270

Widga et al. 2017). 271

Conclusions 272

The scimitar cat Homotherium serum was a specialized predator which lived during the 273

Late Pleistocene and was widely dispersed across North America (Martin et al. 2011c; Martin et 274

al. 2011d). Homotherium serum is most notably identified by its dentition. The serrations present 275

on all teeth allowed for a blade-like function (Rawn-Schatzinger 1983) while the robustness of 276

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the dentition, compared to its sister taxa, allowed for the capture and consumption of large-277

bodied prey (Rawn-Schatzinger 1992). 278

The dental materials recovered from Wally’s Beach permit identification of this specimen 279

as H. serum based primarily on the presence of serrations on all teeth in addition to other 280

diagnostic aspects of the morphology. Individual teeth can be identified based on distinguishing 281

characteristics outlined by previous researchers, and by comparisons to modern felid taxa. The 282

upper canines are particularly diagnostic due to their laterally compressed form, the serrations 283

present on the entire posterior and anterior edges, and their smaller size and slight curvature 284

which differs from Smilodon relatives. The assignment of this specimen to H. serum is 285

corroborated based on geographic and temporal criteria. The context of the find is important as it 286

expands the known distribution of the species, while the age of the specimen, at Cal BP 12 715 287

to 12 655, is currently the youngest precisely dated occurrence of the species in North America 288

and further extends its range temporally. This specimen is significant as it represents the fourth 289

record of H. serum in Canada and the first in Alberta, contributing to our knowledge of the 290

distribution of the species in North America, and provides an indicator of the survival of the 291

species into the Younger Dryas (Widga et al. 2017). 292

Acknowledgments 293

Grants from the Alberta Historical Resources Foundation, Alberta Transportation, 294

Natural Sciences and Engineering Research Council of Canada (NSERC) (L.V.H.), and 295

Westwinds School Division (S.T.) are gratefully acknowledged. The assistance of Tracy Wyman 296

with ArcGIS to produce Figure 1 is greatly appreciated. We are also thankful to the reviewers 297

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and editorial staff of the Canadian Journal of Earth Sciences for their suggestions for improving 298

the manuscript. 299

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Kooyman, B., Hills, L.V., Tolman, S.M., and McNeil, P.E. 2012. Late Pleistocene western camel 344

(Camelops hesternus) hunting in southwestern Canada. American Antiquity, 77: 115–124. 345

Kretzoi, M. 1945. Bemerkungen über das Raubtiersystem. Annales historico-naturales Musei 346

nationalis hungarici, 38: 59–83. 347

Kurtén, B. 1962. The sabre-toothed cat Megantereon from the Pleistocene of Java. Zoologiscge 348

Mededelingen Leiden, 38: 101–104. 349

Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, 350

species, cum characteribus, differentiis, synonymis, locis. (tenth edition), Laurentius Salvius, 351

Stockholm. 352

Martin, L.D., Babiarz, J.P., and Naples, V.L. 2011a. Introduction. In The other saber-tooths: 353

scimitar-tooth cats of the Western Hemisphere. Edited by V.L. Naples, L.D. Martin, and J.P. 354

Babiarz. The John Hopkins University Press, Baltimore, Md. pp. 3–18. 355

Martin, L.D., Babiarz, J.P., and Naples, V.L. 2011b. The osteology of a cookie-cutter cat, 356

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declining Late Pleistocene population in southwestern Alberta, Canada. Quaternary Science 368

Reviews, 24: 1253–1259. 369

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teeth in the saber-tooth cat Homotherium serum Cope. Journal of Vertebrate Paleontology, 3: 371

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morphology, and predatory behaviour. Illinois State Museum, Springfield, Ill. 374

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Gassaway Fissure, Cannon County, Tennessee and the North American distribution of 376

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camel hunting at the southern margin of the ice-free corridor: reassessing the age of Wally’s 384

Beach, Canada. Proceedings of the National Academy of Sciences of the United States of 385

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http://cenozoiclife.blogspot.ca/2014/02/fairmead-landfill-california.html [cited 28 July 2017]. 397

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Tables 398

Table 1. Localities containing Homotherium serum in Canada and the United States. 399

Site State/Province Geologic Age

Fairbanks Alaska Rancholabrean1

Wally’s Beach Alberta Late Wisconsinan/Younger Dryas

111 Ranch Local Fauna Arizona Blancan1

Coyote Local Fauna California Irvingtonian1

Fairmead Landfill Locality California Middle Irvingtonian2

Vallecito Creek Local Fauna California Irvingtonian1

Bass Point Waterway I Florida Blancan1

Inglis IA Florida Irvingtonian1

Leisey Shell Pits Florida Irvingtonian1

Reddick 1A Florida Wisconsinan1

Santa Fe River 1 (includes 1A and 1B) Florida Blancan1

American Falls Idaho Early Wisconsinan; Rancholabrean1

Birch Creek Idaho Blancan1

Dam Local Fauna Idaho Middle Wisconsinan1

Duck Point Idaho Wisconsinan1

Anderson Gravel Pit Local Faunule Kansas Rancholabrean1

Filmore County Minnesota Late Wisconsinan3

Merrell Montana Middle Wisconsinan1

Quinlan Local Fauna Oklahoma Irvingtonian4

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Delmont Local Fauna South Dakota Blancan1

Gassaway Fissure Tennessee Wisconsinan1

Blanco Local Fauna Texas Blancan1

Friesenhahn Cave Texas Late Wisconsinan5

Gilliland Local Fauna Texas Irvingtonian1

Laubach Cave No. 2 Texas Late Wisconsinan1

Marmot Quarry Texas Blancan1

Slaton Quarry Local Fauna Texas Rancholabrean1

Dawson Loc. 9 Yukon Middle to Late Wisconsinan6–8

Old Crow Loc. 21 Yukon Middle Wisconsinan7, 9

Sixtymile Loc. 3 Yukon Middle Wisconsinan10, 11

1Graham and Lundelius 2010 400

2Woodruff 2014 401

3Widga et al. 2012 402

4Smith and Cifelli 2000 403

5Graham 1987 404

6Debicki 1983 405

7Harington 1977 406

8Rawn-Schatzinger and Collins 1981 407

9Harington 1989 408

10Harington 1997 409

11Harington 2003 410

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Figure Captions 411

Figure 1. Distribution of Homotherium serum in Canada and the United States (for locality 412

information see Table 1). 413

Figure 2. Root etching on the upper right canine, lateral aspect. Scale bar = 10mm. 414

Figure 3. Position of tooth fragments within the dentition of H. serum (shaded area represents 415

Wally’s Beach specimen). 416

Figure 4. Serrations on the posterior aspect of the left upper canine. Scale bar = 10mm. 417

Figure 5. Upper canines, medial aspect. A, left canine (DhPg-8-3836). B, right canine (DhPg-8-418

3837). Scale bars = 10mm. 419

Figure 6. Identifiable incisor fragments. A, upper first or second incisor, anterior aspect (DhPg-420

8-3838). B, lower right second incisor, medial/lingual aspect (DhPg-8-3839). C, lower left 421

second incisor, lateral/lingual aspect (DhPg-8-3840). D, upper incisor, lateral or medial aspect 422

(DhPg-8-3841). Scale bars = 10mm. 423

Figure 7. Lower left canine (DhPg-8-3843). A, anterior aspect. B, posterior aspect. Scale bars = 424

10mm. 425

Figure 8. Metacone of the upper left fourth premolar (DhPg-8-3844). A, lingual aspect. B, 426

buccal aspect. Scale bars = 10mm. 427

Figure 9. Paracone of the lower right first molar (DhPg-8-3845). A, lingual aspect. B, buccal 428

aspect. Scale bars = 10mm. 429

Figure 10. Root fragment, possibly from a molar or premolar (DhPg8-3846). Scale bar = 10mm. 430

Figure 11. Polish on the anterior aspect of the right upper canine. Scale bar = 10mm. 431

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Figures 432

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