Canadian Journal of Earth Sciences
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;Draft Independent Kooyman, Brian; University of Calgary , Department of Anthropology and Archaeology
Is the invited manuscript for consideration in a Special N/A Issue? :
Homotherium serum , Homotheriini, Dentition, Wally's Beach, Keyword: Pleistocene
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1 Scimitar Cat ( Homotherium serum Cope) from 2 Southwestern Alberta, Canada
3 Tatyanna Ewald, L.V. Hills*, Shayne Tolman, and Brian Kooyman
4 *Deceased
5 T. Ewald. (email: [email protected]) Department of Anthropology and Archaeology, 6 University of Calgary, 2500 University Dr. NW, Calgary, AB T2N 1N4, Canada.
7 L.V Hills. * Department of Geoscience, University of Calgary , 2500 University Dr. N.W., 8 Calgary, AB T2N 1N4, Canada
9 S. Tolman. (email: [email protected]) Box 1146, Cardston, AB TOK 0Y0, Canada.
10 B. Kooyman. (email: [email protected]) Department of Anthropology and Archaeology, 11 University of Calgary, 2500 University Dr. NW, Calgary, AB T2N 1N4, Canada.
12 Corresponding author: Draft
13 Tatyanna Ewald 14 Email: [email protected] 15 Phone: 1 403 827 2245 16 Fax: 1 403 284 5467 17 Address: Department of Anthropology and Archaeology 18 2500 University Drive NW 19 T2N 1N4 20 Canada
21 *Deceased.
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22 Abstract 23 Skull and tooth fragments of Homotherium serum recently recovered from the Wally’s Beach
24 site (DhPg 8) in southwestern Alberta provide the first indications that scimitar cat populated the
25 area of the St. Mary Reservoir. AMS radiocarbon dating provides a 2 sigma range calibrated age
26 of Cal BP 12 715 to 12 655. This is the fourth known occurrence of the species in Canada, the
27 first outside of Yukon, and is currently the youngest precisely dated occurrence of the species in
28 North America. Well preserved dentition combined with the temporal and geographic context
29 allows the sample to be identified as Homotherium serum . The specimen is significant as it
30 represents an extension of the geographic and chronological range of the species.
31 Keywords Draft 32 Homotherium serum , Homotheriini, Dentition, Wally’s Beach, Pleistocene
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33 Introduction
34 The Wally’s Beach site, near Cardston in southwestern Alberta, consists of a series of
35 Late Pleistocene paleontological and archaeological remains which were exposed by erosion
36 associated with the operation of the St. Mary Reservoir (Hills et al. 2014; Kooyman et al. 2012;
37 Waters et al. 2015). Species present include Equus conversidens , Camelops hesternus ,
38 Bootherium bombifrons , Rangifer tarandus , and Bison antiquus among others. A number of
39 species are also represented by preserved tracks, including Mammuthus (McNeil et al.
40 2005:1255). Recently, skull and tooth fragments of an individual scimitar cat, Homotherium
41 serum, were discovered. The purpose of this paper is to describe and illustrate this specimen, and 42 to discuss its location and date in relationDraft to other known occurrences of the species.
43 Geological Setting and Context
44 The Wally’s Beach site (also known by its Borden designation DhPg 8) is located in
45 southwestern Alberta near the town of Cardston (Fig. 1), in the drawdown zone of the St. Mary
46 Reservoir (Hills et al. 2014). While the H. serum remains are paleontological, the Wally’s Beach
47 site is predominately an archaeological site and is therefore defined by the Borden system. The
48 Borden system divides Canada into a series of small blocks inside larger blocks based on latitude
49 and longitude. The capital letters in the Borden designation identify the large block, while the
50 lower case letters denote the small block to which the site belongs. Each archaeological site is
51 then designated sequentially within these blocks (Borden 1954).
52 Cranial remains of H. serum were discovered in situ, within a 16 18cm radius. The
53 location consists of Late Pleistocene loess and eolian sands, upon which a Holocene paleosol
54 developed (Hills et al. 2014:25) The specimen was recovered from the very top portion of the
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55 Pleistocene sediments, embedded in the carbonate horizon that resulted from the development of
56 the overlying paleosol. The enamel layer of one of the canines shows extensive fibrous root
57 etchings on the surface (Fig. 2). This suggests that this tooth was incorporated in the soil horizon
58 while it was actively forming, before the soil’s final burial.
59 A bone sample from the skull provides an AMS (Beta 440230) 2 sigma range calibrated
60 age of Cal BP 12 715 to 12 655 (conventional age of BP 10 740 ±40). This date was calibrated
61 using IntCal13. The CN (carbon:nitrogen) ratio of +3.3 is within the normal range expected for
62 live protein, therefore the collagen was well preserved (the δ13C value of 19.3 o/oo and δ15N
63 value of +6.6 o/oo are also both within the normal range expected for live protein). This date
64 places the specimen in the Late Pleistocene and makes it the youngest dated specimen of
65 Homotherium serum in Canada. Draft
66 Materials and Methods
67 The following description is based on fragments of five incisors, two upper canines, the
68 left lower canine, the upper fourth premolar, the lower first molar, and an unidentifiable root
69 (Fig. 3). Preservation is largely restricted to the enamel layer of each tooth. The canine fragments
70 were measured using electronic calipers providing digital readouts to the nearest millimetre (see
71 Dental Remains Recovered). Several small skull fragments (DhPg 8 3847) are also present
72 within the assemblage, but were not considered in this study. The skull fragments are too small
73 and fragmented for partial skull reconstruction, therefore the teeth were the focus of this analysis.
74 The identification of the tooth fragments was reliant predominately upon Rawn
75 Schatzinger (1983; 1992), although several other sources (Antón et al. 2014; Biknevicius et al.
76 1996; Cope 1893; Martin et al. 2011a; Martin et al. 2011b; Martin et al. 2011c; Martin et al.
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77 2011d; Wheeler 2011) and some comparisons to modern Felidae taxa (Felis concolor, Lynx
78 canadensis , and Leopardus pardalis , Department of Anthropology and Archaeology, University
79 of Calgary) were also utilized.
80 Results
81 Systematic Paleontology
82 We follow the systematics proposed by Antón et al. (2014).
83 Mammalia Linnaeus, 1758
84 Order Carnivora Bowdich, 1821 85 Suborder Feliformia Kretzoi, 1945 Draft 86 Family Felidae Fischer von Waldheim, 1817
87 Subfamily Machairodontinae Gill, 1872
88 Tribe Homotheriini Kurtén, 1962
89 Genus Homotherium Fabrini, 1890
90 Homotherium serum (Cope 1893)
91 Referred Specimens
92 The material used in this study will be stored in the Archaeology collections at the Royal
93 Alberta Museum: DhPg 8 3836, crown of the upper left canine; DhPg 8 3837, crown of the
94 upper right canine; DhPg 8 3838, fragment of the crown from an upper first or second incisor;
95 DhPg 8 3839, fragment of the crown from the lower right second incisor; DhPg 8 3840,
96 fragment of the crown from the lower left second incisor; DhPg 8 3841, fragment of the crown
97 from an upper incisor; DhPg 8 3842, fragment of the crown from an unidentified incisor; DhPg
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98 8 3843, crown of the lower left canine; DhPg 8 3844, metacone of the upper left fourth
99 premolar; DhPg 8 3845, paracone of the lower right first molar; DhPg 8 3846, partial root;
100 DhPg 8 3847, several small skull fragments. Cataloguing of these materials followed the Borden
101 system.
102 Dental Remains Recovered
103 The Homotherium genus is particularly characterized and separated from other saber
104 tooth cats by its dentition, and this is particularly useful in distinguishing Homotherium from its
105 close Smilodon relatives, also present in North America during the Pleistocene. A total of 11
106 tooth fragments identified as belonging to Homotherium serum were recovered at Wally’s 107 Beach. The teeth recovered indicate thatDraft only a single individual is represented; no elements are 108 duplicated and the size and wear on the teeth are consistent for a single individual. In addition,
109 the remains were discovered in situ. Distinct, fine serrations are present along the cutting edge of
110 each tooth and are characteristic of H. serum , as serrations are present on all teeth of both
111 juvenile and adult specimens (Rawn Schatzinger 1983). “The term “finely serrate” as used by
112 Cope (1983) and Churcher (1966) refers to the consistent and regular serration of H. serum ’s
113 teeth,” (Rawn Schatzinger 1992:4). Within this specimen, serrations are most clearly pronounced
114 on the upper canines (Fig. 4), upon which they are present along the entire length of the anterior
115 and posterior edge (Rawn Schatzinger 1983; 1992; Wheeler 2011:28).
116 The most distinctive of the teeth recovered from Wally’s Beach are the upper canines
117 (Fig. 5), of which only the enamel caps are present. As noted by Rawn Schatzinger (1983:53),
118 "the permanent canine has one half of its length covered with enamel.” As such, it is reasonable
119 to assume that these enamel caps represent approximately one half of the original length of each
120 tooth. The fragment of the left tooth (DhPg 8 3836) is 51.8 mm in length and 20.1 mm wide
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121 antero posteriorly at its widest aspect, while the right (DhPg 8 3837) is 48.2 mm long and 18.7
122 mm wide. Medio laterally, the left tooth is 8.3 mm wide and the right is 8.4 mm, once again at
123 their widest aspects. This corresponds with descriptions by Rawn Schatzinger (1983:52; 1992:3)
124 of H. serum upper canines which are laterally compressed or flat as compared to conical tooth
125 cats. Also unique to Homotherium, and exhibited in this specimen, is that the canines are
126 relatively short and broad in comparison to other saber tooths (Martin et al. 2011a:6; Martin et
127 al. 2011d:208; Wheeler 2011:27) and that they are more complex on their lingual surface (Rawn
128 Schatzinger 1983:52). Both teeth are relatively straight with a very slight posterior curvature.
129 The convex nature of the posterior as opposed to the anterior face of the upper canines, as in this
130 specimen, was noted by Cope (1893:897). Both canines exhibit heavy post depositional root
131 etching. Draft
132 Several incisor fragments were also identified. All display a slight posterior convex
133 curvature and all are procumbent and rather robust, as is characteristic of Homotherium (Martin
134 et al. 2011a:8; Wheeler 2011:28). The first fragment (Fig. 6A; DhPg 8 3838) is comprised of the
135 anterior portion of the enamel cap. Partial cuspules are present on both lateral sides of this
136 fragment, allowing it to be identified as either the upper first or second incisor (Rawn
137 Schatzinger 1983:53), although the side is indeterminable in this state. An additional incisor
138 fragment (Fig. 6B; DhPg 8 3839) consists predominately of the posterior and medial portion of
139 the enamel cap. Basal cuspules are present on both lateral sides of the fragment, one partial and
140 the other complete. This tooth has therefore been identified as the lower second incisor (Rawn
141 Schatzinger 1983:53). The medial cuspule is noticeably higher on the tooth than the lateral,
142 indicating that this is the right second incisor (Martin et al. 2011b: Figure 4.8). Yet another
143 incisor fragment, consisting predominately of the posterior and lateral portion of the enamel cap,
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144 exhibits mirrored identical features (Fig. 6C; DhPg 8 3840). This tooth has been identified as the
145 lower left second incisor.
146 Yet another obviously incisiform fragment consists of a lateral portion of the enamel cap,
147 and terminates just below a basal cuspule (Fig. 6D; DhPg 8 3841). This tooth has been identified
148 as part of the upper incisor dentition; however further identification is not possible. It is larger
149 than the lower second incisors already identified for this specimen and is therefore not a lower
150 first incisor, as the lateral incisors are typically larger and more robust in both saber tooth cats
151 and modern carnivores (Biknevicius et al. 1996). The obvious cuspule present on this fragment
152 also excludes it from being identified as the lower third incisor, as this tooth lacks basal cuspules 153 in H. serum (Rawn Schatzinger 1983:52).Draft This incisor is therefore part of the upper dentition, 154 however is not identifiable within a sequence. A fifth incisor fragment (DhPg 8 3842) is poorly
155 preserved and further identification is not possible.
156 Also present within this sample is a lower canine (Fig. 7; DhPg 8 3843), of which the
157 entire crown appears to be present. As noted by Rawn Schatzinger (1983:53), enamel covers
158 approximately one third of this tooth, which indicates that this fragment is likely representative
159 of one third of the original length of the tooth. Although the lower canines of Homotherium are
160 diagnostically reduced and incisor like (Biknevicius et al. 1996:518; Wheeler 2011:32), at 19.6
161 mm long this tooth fragment is noticeably larger than all the incisor fragments present in this
162 sample, which is typical in H. serum (Rawn Schatzinger 1983:50). In their description of
163 Homotherium serum , Antón et al. (2014:263) state that in the occlusal view the lower canine has
164 “a strong lateral flattening, and a small size, just slightly larger than the i3.” This lateral
165 compression is also exhibited in this specimen and is also characteristic of H. serum (Rawn
166 Schatzinger 1983:53). It is also much smaller than the upper canines, which would allow the cat
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167 more clearance when biting (Wheeler 2011:29). A small basal cuspule located on its medial
168 aspect (Rawn Schatzinger 1983:53) indicates that this tooth is the left. This specimen also
169 exhibits a distinct convex posterior curvature which is more pronounced than in the upper
170 canines.
171 A single cusp of the upper fourth premolar, or the upper carnassial (Fig. 8; DhPg 8 3844)
172 is also present. This cusp is identifiable as such because the edge is nearly straight, allowing it to
173 function in its role as a specialized cutting tooth, a characteristic unique to Homotherium (Rawn
174 Schatzinger 1983:52; Rawn Schatzinger 1992:3). The shape of this specimen also indicates that
175 it is the metacone, which is the only cusp within H. serum which is distinctively flat (Rawn
176 Schatzinger 1983). The main ridge of the cusp leans slightly medially and two slight bulges are
177 present on the buccal aspect of the tooth,Draft with a slight vertical groove between them (Cope
178 1893:897). Combined with indications of post mortem breakage on the anterior portion of the
179 cusp and a lack of such evidence on the posterior portion, this fragment has been identified as the
180 most posterior cusp (metacone) of the left upper carnassial (Rawn Schatzinger 1983:54). The
181 identification of this tooth was corroborated by comparisons to modern Felidae samples ( Felis
182 concolor, Lynx canadensis , and Leopardus pardalis , Department of Anthropology and
183 Archaeology, University of Calgary).
184 A cusp with a pointed and rather symmetrical apex was also recovered (Fig. 9; DhPg 8
185 3845). The lower carnassial, or lower first molar, of Homotherium is distinguished by its two
186 blade like cusps (Rawn Schatzinger 1983:54). The paracone in particular is both large and
187 vertical (Martin et al. 2011c:193), indicating that this fragment is the paracone of the lower
188 carnassial. The presence of two vertical concavities on the lingual aspect of the cusp, as well as
189 indications of post mortem breakage on the posterior but not anterior aspect of the cusp, further
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190 indicates that this is a fragment of the right carnassial. Comparisons with modern Felidae
191 samples ( Felis concolor, Lynx canadensis , and Leopardus pardalis , Department of Anthropology
192 and Archaeology, University of Calgary) corroborated this identification. Shearing wear is also
193 present on both carnassial fragments; this is particularly well preserved on the fragment of the
194 lower carnassial (see Fig. 9A).
195 The final fragment is a partial root (Fig. 10; DhPg 8 3846). Three separate roots are
196 identifiable on this fragment, the middle of which projects outwards from the other two. Based
197 on comparisons with modern Felidae samples ( Felis concolor, Lynx canadensis , and Leopardus
198 pardalis, Department of Anthropology and Archaeology, University of Calgary), it is likely that 199 this belongs to a molar or premolar of HomotheriumDraft serum . As this fragment is not diagnostic, 200 further evaluation of the tooth roots in Homotherium is beyond the scope of this study.
201 In the present material, the upper canines are lenticular in cross section with serrations
202 present on both the anterior and posterior edges. The canines are blunted with numerous flake
203 scars and the serrations are polished and worn (Fig. 11), indicative of a mature individual. Rawn
204 Schatzinger (1983) subdivided the scimitar cat into an eight stage age progression ranging from
205 Stage I, deciduous teeth just erupting beyond the alveolus, to Stage VIII, with extensive wear and
206 teeth blunted by wear polish. These last features are visible on this specimen; hence it is
207 classified as a Stage VIII adult. Rawn Schatzinger (1983) did not provide an age for Stage VIII
208 adults, however that for Stage VII is two or more years, which equates to the approximate age
209 for complete functional permanent dentition. No teeth in this specimen are in an extreme state of
210 wear evidenced by broken teeth or teeth worn to the pulp cavity (Rawn Schatzinger 1983:52).
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211 Discussion
212 The Wally’s Beach scimitar cat specimen provides a significant new addition to the Late
213 Pleistocene paleofauna of southwestern Alberta, Canada. It is the fourth known occurrence of H.
214 serum in Canada (Debicki 1983:24; Harington 1977:526 528, 529 531; Harington 1989:95;
215 Harington 1997; Harington 2003:385; Rawn Schatzinger and Collins 1981:18) and the first in
216 Alberta. The calibrated age of Cal BP 12 715 to 12 655 is corroborated by the specimen’s
217 stratigraphic context and makes this the youngest precisely dated occurrence of this species in
218 North America.
219 Due to the fragmented nature of the skull fragments recovered, an analysis of the
220 specimen recently recovered from Wally’sDraft Beach was entirely dependent upon the partial
221 dentition. While the distinctive tooth morphology (e.g., fine serrations present along the cutting
222 edge of all teeth) exhibited here was the basis for identifying this specimen, the geographic and
223 temporal context of the finds supports this identification.
224 The saber tooth cats were common primarily during the Pliocene and Pleistocene, periods
225 correspondingly characterized by large prey species which these cats were well adapted to hunt.
226 Homotherium is recognized as a wide ranging genus from throughout Europe, Asia, Africa, and
227 North America (Rawn Schatzinger 1992), and as the most abundant scimitar cats from northern
228 Eurasia and North America (Martin et al. 2011d). Recently, a specimen from Venezuela
229 (Homotherium venezuelensis ) has also been described (Rincón et al. 2011), adding South
230 America to the range of Homotherium and making it the most geographically dispersed genus of
231 the saber tooth cats (Antón et al. 2014). Homotherium was the only saber tooth cat that achieved
232 such a wide range (Martin et al. 2011a).
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233 The genus Homotherium is the last of a long line of machairodont Homotheriini, ranging
234 in age between four million and 10,000 years (Antón et al. 2014). Their extinction correlated
235 with megafauna extinctions as well as climatic fluctuations (Martin et al. 2011a). Diagnosis of
236 species within Homotherium remains a debated issue among researchers. While a single species
237 tends to be recognized for Europe and Asia ( Homotherium latidens ), there are several
238 classifications of North American species (Antón et al. 2014). Homotherium ischyrus is often
239 described as an earlier North American species recovered most often from Pliocene and early
240 Pleistocene contexts, while Homotherium serum characterizes the Late Pleistocene of North
241 America (Martin et al. 2011c). Homotherium serum, while abundant primarily during the later
242 stages of the Pleistocene, was not uncommon earlier in this epoch (Antón et al. 2014). Draft 243 While both H. ischyrus and H. serum occupied North America, H. serum characterized
244 the Late Pleistocene and may have been the only scimitar species on the continent at the time
245 (Antón et al. 2014). The distribution of Homotherium in North America is predominately within
246 the Camelops Faunal Province, “which was considered to be composed mostly of pine parkland
247 and characterized by the most cursorially adapted large mammals,” (Martin et al. 2011d:206).
248 Homotherium serum is known from over 30 sites in North America which are concentrated in the
249 United States (see Fig. 1; Table 1). Only three sites within Canada (not including Wally’s
250 Beach) are currently known to contain H. serum remains (Debicki 1983:24; Harington 1977:526
251 528, 529 531; Harington 1989:95; Harington 1997; Harington 2003:385; Rawn Schatzinger and
252 Collins 1981:18). As the only known specimen in Alberta, the remains from Wally’s Beach
253 represent an extension of the range of H. serum . Furthermore, as the youngest occurrence in
254 Canada by a significant margin, this specimen may represent a Late Pleistocene extension of the
255 species northward into modern Canada from the United States.
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256 The calibrated age of Cal BP 12 715 to 12 655 for this specimen corroborates its
257 identification as H. serum, which characterized the Late Pleistocene of North America, rather
258 than the much older H. ischyrus (Antón et al. 2014). More significantly, this specimen is the
259 youngest known precisely dated occurrence of the species in North America. In Canada, three
260 other specimens have been identified from Yukon, however these are significantly older than that
261 recovered from Wally’s Beach. These Yukon specimens date to over 40,000 years BP at
262 Sixtymile Loc. 3 (the only Yukon specimen which has been precisely dated) (Harington 1997;
263 Harington 2003:385), over 25,000 years BP at Old Crow Loc. 21 (Harington 1977:526 528;
264 Harington 1989:95), and 39,900 to 22,200 years BP at Dawson Loc. 9 (Debicki 1983:24;
265 Harington 1977:529 531; Rawn Schatzinger and Collins 1981:18). In North America more
266 broadly, most specimens date to an earlierDraft Pleistocene context (see Table 1). Only one site
267 contains specimens which have been precisely radiocarbon dated to the last 15,000 years:
268 Laubach Cave No. 2, Texas (13,970 years BP; Graham and Lundelius 2010). The Wally’s Beach
269 specimen is therefore currently the most recent precisely dated specimen in North America and
270 may be a useful indicator of the survival of the species into the Younger Dryas (as defined in
271 Widga et al. 2017).
272 Conclusions
273 The scimitar cat Homotherium serum was a specialized predator which lived during the
274 Late Pleistocene and was widely dispersed across North America (Martin et al. 2011c; Martin et
275 al. 2011d). Homotherium serum is most notably identified by its dentition. The serrations present
276 on all teeth allowed for a blade like function (Rawn Schatzinger 1983) while the robustness of
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277 the dentition, compared to its sister taxa, allowed for the capture and consumption of large
278 bodied prey (Rawn Schatzinger 1992).
279 The dental materials recovered from Wally’s Beach permit identification of this specimen
280 as H. serum based primarily on the presence of serrations on all teeth in addition to other
281 diagnostic aspects of the morphology. Individual teeth can be identified based on distinguishing
282 characteristics outlined by previous researchers, and by comparisons to modern felid taxa. The
283 upper canines are particularly diagnostic due to their laterally compressed form, the serrations
284 present on the entire posterior and anterior edges, and their smaller size and slight curvature
285 which differs from Smilodon relatives. The assignment of this specimen to H. serum is 286 corroborated based on geographic and temporalDraft criteria. The context of the find is important as it 287 expands the known distribution of the species, while the age of the specimen, at Cal BP 12 715
288 to 12 655, is currently the youngest precisely dated occurrence of the species in North America
289 and further extends its range temporally. This specimen is significant as it represents the fourth
290 record of H. serum in Canada and the first in Alberta, contributing to our knowledge of the
291 distribution of the species in North America, and provides an indicator of the survival of the
292 species into the Younger Dryas (Widga et al. 2017).
293 Acknowledgments
294 Grants from the Alberta Historical Resources Foundation, Alberta Transportation,
295 Natural Sciences and Engineering Research Council of Canada (NSERC) (L.V.H.), and
296 Westwinds School Division (S.T.) are gratefully acknowledged. The assistance of Tracy Wyman
297 with ArcGIS to produce Figure 1 is greatly appreciated. We are also thankful to the reviewers
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298 and editorial staff of the Canadian Journal of Earth Sciences for their suggestions for improving
299 the manuscript.
Draft
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345 (Camelops hesternus ) hunting in southwestern Canada. American Antiquity, 77 : 115–124.
346 Kretzoi, M. 1945. Bemerkungen über das Raubtiersystem. Annales historico naturales Musei
347 nationalis hungarici, 38 : 59–83.
348 Kurtén, B. 1962. The sabre toothed cat Megantereon from the Pleistocene of Java. Zoologiscge
349 Mededelingen Leiden, 38 : 101–104.
350 Linnaeus, C. 1758. Systema naturae perDraft regna tria naturae, secundum classes, ordines, genera,
351 species, cum characteribus, differentiis, synonymis, locis. (tenth edition), Laurentius Salvius,
352 Stockholm.
353 Martin, L.D., Babiarz, J.P., and Naples, V.L. 2011a. Introduction. In The other saber tooths:
354 scimitar tooth cats of the Western Hemisphere. Edited by V.L. Naples, L.D. Martin, and J.P.
355 Babiarz. The John Hopkins University Press, Baltimore, Md. pp. 3–18.
356 Martin, L.D., Babiarz, J.P., and Naples, V.L. 2011b. The osteology of a cookie cutter cat,
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358 Edited by V.L. Naples, L.D. Martin, and J.P. Babiarz. The John Hopkins University Press,
359 Baltimore, Md. pp. 43–98.
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360 Martin, L.D., Naples, V.L, and Babiarz, J.P. 2011c. Revision of the New World Homotheriini. In
361 The other saber tooths: scimitar tooth cats of the Western Hemisphere. Edited by V.L. Naples,
362 L.D. Martin, and J.P. Babiarz. The John Hopkins University Press, Baltimore, Md. pp. 185–194.
363 Martin, L.D., Babiarz, J.P., and Naples, V.L. 2011d. A framework for the North American
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366 Md. pp. 201–209.
367 McNeil, P.L., Hills, L., Kooyman, B., and Tolman, S.M. 2005. Mammoth tracks indicate a
368 declining Late Pleistocene population in southwestern Alberta, Canada. Quaternary Science 369 Reviews, 24 : 1253–1259. Draft 370 Rawn Schatzinger, V. 1983. Development and eruption sequence of deciduous and permanent
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372 49–57.
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374 morphology, and predatory behaviour. Illinois State Museum, Springfield, Ill.
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376 Gassaway Fissure, Cannon County, Tennessee and the North American distribution of
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381 Smith, K.S., and Cifelli, R.L. 2000. A synopsis of the Pleistocene vertebrates of Oklahoma.
382 Oklahoma Geological Survey Bulletin 147, The University of Oklahoma, Norman, Okla.
383 Waters, M.R., Stafford, Jr. T.W., Kooyman, B., and Hills, L.V. 2015. Late Pleistocene horse and
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390 Widga, C., Lengyel, S.N., Saunders, J.,Draft Hodgins, G., Walker, J.D., and Wanamaker, A.D. 2017. 391 Late Pleistocene proboscidean population dynamics in the North American Midcontinent.
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398 Tables
399 Table 1. Localities containing Homotherium serum in Canada and the United States.
Site State/Province Geologic Age
Fairbanks Alaska Rancholabrean 1
Wally’s Beach Alberta Late Wisconsinan/Younger Dryas
111 Ranch Local Fauna Arizona Blancan 1
Coyote Local Fauna California Irvingtonian 1
Fairmead Landfill Locality California Middle Irvingtonian 2
Vallecito Creek Local Fauna California Irvingtonian1 Bass Point Waterway I Draft Florida Blancan 1 Inglis IA Florida Irvingtonian 1
Leisey Shell Pits Florida Irvingtonian 1
Reddick 1A Florida Wisconsinan 1
Santa Fe River 1 (includes 1A and 1B) Florida Blancan 1
American Falls Idaho Early Wisconsinan; Rancholabrean 1
Birch Creek Idaho Blancan 1
Dam Local Fauna Idaho Middle Wisconsinan 1
Duck Point Idaho Wisconsinan 1
Anderson Gravel Pit Local Faunule Kansas Rancholabrean 1
Filmore County Minnesota Late Wisconsinan 3
Merrell Montana Middle Wisconsinan 1
Quinlan Local Fauna Oklahoma Irvingtonian 4
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Delmont Local Fauna South Dakota Blancan 1
Gassaway Fissure Tennessee Wisconsinan 1
Blanco Local Fauna Texas Blancan 1
Friesenhahn Cave Texas Late Wisconsinan 5
Gilliland Local Fauna Texas Irvingtonian 1
Laubach Cave No. 2 Texas Late Wisconsinan 1
Marmot Quarry Texas Blancan 1
Slaton Quarry Local Fauna Texas Rancholabrean 1
Dawson Loc. 9 Yukon Middle to Late Wisconsinan 6–8
Old Crow Loc. 21 Yukon Middle Wisconsinan 7, 9
Sixtymile Loc. 3 Draft Yukon Middle Wisconsinan 10, 11
400 1Graham and Lundelius 2010
401 2Woodruff 2014
402 3Widga et al. 2012
403 4Smith and Cifelli 2000
404 5Graham 1987
405 6Debicki 1983
406 7Harington 1977
407 8Rawn Schatzinger and Collins 1981
408 9Harington 1989
409 10 Harington 1997
410 11 Harington 2003
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411 Figure Captions
412 Figure 1. Distribution of Homotherium serum in Canada and the United States (for locality
413 information see Table 1).
414 Figure 2. Root etching on the upper right canine, lateral aspect. Scale bar = 10mm.
415 Figure 3. Position of tooth fragments within the dentition of H. serum (shaded area represents
416 Wally’s Beach specimen).
417 Figure 4. Serrations on the posterior aspect of the left upper canine. Scale bar = 10mm.
418 Figure 5. Upper canines, medial aspect. A, left canine (DhPg 8 3836). B, right canine (DhPg 8
419 3837). Scale bars = 10mm. 420 Figure 6. Identifiable incisor fragments.Draft A, upper first or second incisor, anterior aspect (DhPg 421 8 3838). B, lower right second incisor, medial/lingual aspect (DhPg 8 3839). C, lower left
422 second incisor, lateral/lingual aspect (DhPg 8 3840). D, upper incisor, lateral or medial aspect
423 (DhPg 8 3841). Scale bars = 10mm.
424 Figure 7. Lower left canine (DhPg 8 3843). A, anterior aspect. B, posterior aspect. Scale bars =
425 10mm.
426 Figure 8. Metacone of the upper left fourth premolar (DhPg 8 3844). A, lingual aspect. B,
427 buccal aspect. Scale bars = 10mm.
428 Figure 9. Paracone of the lower right first molar (DhPg 8 3845). A, lingual aspect. B, buccal
429 aspect. Scale bars = 10mm.
430 Figure 10. Root fragment, possibly from a molar or premolar (DhPg8 3846). Scale bar = 10mm.
431 Figure 11. Polish on the anterior aspect of the right upper canine. Scale bar = 10mm.
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432 Figures
433 Figure 1
Draft
434
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435 Figure 2
436
Draft
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437 Figure 3
Draft
438
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439 Figure 4
440
Draft
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441 Figure 5
442 443
Draft
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444 Figure 6
Draft 445
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446 Figure 7
447
Draft
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448 Figure 8
449
Draft
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450 Figure 9
451
A Draft B
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452 Figure 10
Draft
453
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454 Figure 11
455
Draft
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