Canadian Journal of Earth Sciences
Pleistocene Mammals From Extinction Cave, Belize
Journal: Canadian Journal of Earth Sciences
Manuscript ID cjes-2018-0178.R3
Manuscript Type: Article
Date Submitted by the 04-May-2019 Author:
Complete List of Authors: Churcher, C.S.; University of Toronto, Zoology
Central America, Pleistocene, Fauna, Vertebrate Palaeontology, Keyword: Limestone cave
Is the invited manuscript for consideration in a Special Not applicableDraft (regular submission) Issue? :
https://mc06.manuscriptcentral.com/cjes-pubs Page 1 of 43 Canadian Journal of Earth Sciences
1
1 PLEISTOCENE MAMMALS FROM EXTINCTION CAVE, BELIZE
2 by C.S. CHURCHER1
Draft
1Department of Zoology, University of Toronto, Toronto, Ontario Canada M5S 2C6 and
322-240 Dallas Rd., Victoria, British Columbia, Canada V8V 4X9 (corresponding address):
e-mail [email protected]
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 2 of 43
2
4
5 ABSTRACT. A small mammalian fauna is recorded from Extinction Cave (also called Sibun
6 Cave), east of Belmopan, on the Sibun River, Belize, Central America. The animals recognized
7 are armadillo (†Dasypus bellus), American lion (†Panthera atrox), jaguar (P. onca), puma or
8 mountain lion (Puma concolor), Florida spectacled bear (†Tremarctos floridanus), javelina or
9 collared peccary (Pecari tajacu), llama (Camelidae indet., ?†Palaeolama mirifica), red brocket
10 deer (Mazama americana), bison (Bison sp.) and Mexican half-ass (†Equus conversidens), and
11 sabre-tooth cat († Smilodon fatalis) may also be represented (‘†’ indicates an extinct taxon).
12 Bear and bison are absent from the region today. The bison record is one of the more southernly 13 known. The bear record is almost the mostDraft westerly known and a first for Central America. 14 KEYWORDS. Fauna, Rancholabrean, Wisconsinian, Dasypus bellus, Homo sapiens, Panthera
15 atrox, P. concolor, P. onca, Smilodon, Tremarctos floridanus, Pecari tajacu, Palaeolama
16 mirifica, Mazama americana, Bison sp., Equus conversidens, ecology.
17 Introduction. Extinction Cave is the first Rancholabrean faunal deposit to be reported
18 in Belize. It links the Belizean Rancholabrean mammalian fauna to that of Mexico and
19 Guatemala, and shows that it was part of the midcontinental Nearctic fauna and fitted into the
20 continental species cline of distribution. It demonstrates that Belize’s extensive limestone
21 exposures carried a fauna similar to that on the Mexican Yucatán to the west or the Guatemalan
22 highlands and coastal plain to the south.
23 Extinction Cave is located in a small isolated limestone hill about 50 m south of a logging
24 road that gives access to the area. It is the lowest and only fossiliferous member of a trio of caves
25 and is entered from the north side of the hill through a wide entrance in a cliff overhang. Logan
https://mc06.manuscriptcentral.com/cjes-pubs Page 3 of 43 Canadian Journal of Earth Sciences
3
26 McNatt of the Belize Department of Archareology describes it as a narrow passage that is about
27 70 m long, and passes through the hill to a second opening on the south or Ventura Creek side of
28 the hill. In the middle of the passage is a low silt-filled hollow which lies some 2.5 m lower than
29 the main passage floor. McNatt records ‘We collected a small amount of animal bones and teeth
30 from this area’ (pers. comm, 2000; Field Notes, AC 62 #100, June 21, 1984). McNatt notes it as
31 Cave A and indicates that the cave was once a water conduit within the limestone. An upper
32 cave, Cave B, about halfway up the side of the hill, has two or three small chambers and contains
33 some remnant Mayan rockwalling. A third shelter, Cave C. is located on the north side of the
34 hill with a wide, pillared, walk-in entrance, but becomes low at the back. Potsherds were 35 scattered throughout this cave; 21 were Draftcollected, together with a ‘chert biface’ and an 36 unidentified canine tooth (these specimens were not included in the faunal sample) (data from
37 ‘Report of Visit to new site: Caves at Tiger Sandy Bay Outcrops - West’, 4 May, 1984' by L.
38 McNatt, 21 June, 1984).
39 The faunal sample was collected by Logan McNatt, then an Archaeological Officer with
40 the Belize Department of Archaeology, Belmopan, on May 4th, 1984. Caves in Tiger Sandy Bay
41 Limestone Formation outcrops had been reported by Diane Weyer of Mile 30, Western
42 Highway, about April 27th., to Joriel Boriek, Carol Pepe, George Vani and Logan McNatt. On
43 May 4th these five visited the site, which is located east of Belmopan about 28 km (16.5 miles)
44 east of the settlement of Churchyard and about 3.5 km (2.2 miles) across the Sibun River on its
45 south or right bank (Fig.1). The cave lies on the north slope of the valley of Ventura Creek at
46 approximately Lat. 17̊ 20.0' N, Long. 88̊ 31.5', and an elevation of less than 40 m amsl, in the
47 middle flood plain of the Sibun River. The nearest town is La Democracia on the Western
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 4 of 43
4
48 Highway some 5.5 km (3.5 miles) due northwest.
49 The sample of mammalian bones and teeth, calcareous cave earth and bone breccia, in ten
50 packets, was handed by McNatt to Maryijka Mychajlowycz on May 5th, 1984, for identification.
51 She was then on leave from working as my laboratory assistant in the Department of Zoology,
52 University of Toronto, and had access to the requisite comparative materials in the Royal Ontario
53 Museum (ROM). We prepared the specimens, performed preliminary identifications, composed
54 a faunal list, and estimated the faunal age as Wisconsinan. In October, 1999, I refined our
55 examinations, took measurements, made reference searches and comparisons, and used the
56 collections of the Departments of Mammalogy and Palaeobiology (Vertebrate Palaeontology) of 57 the Royal Ontario Museum, Toronto, toDraft confirm our earlier diagnoses. 58 The specimens came into my care without any agreements as to their ownership or rights
59 of collection or transport, and have been deposited in the Royal Ontario Museum, Toronto, for
60 safe keeping.
61 Provenance and Geology. Extinction Cave is an isolated dry karst solution conduit in
62 the eastern lobe of the Vaca Plateau (Tiger Sandy Bay Formation) limestones that form the
63 bedrock of the western half of southern Belize District, west central Belize (Fig. 1). It lies on the
64 Sibun River valley bottom in bush covered rolling topography where the limestone valley walls
65 become submerged in the alluvium of the river plain. The limestones are honeycombed by
66 solution chambers, pipes and conduits (e.g., Chiquibul Sinks in the Vaca Plateau limestones to
67 the southwest).
68 The Extinction Cave fossils comprise a single assemblage within a uniform cave earth
69 matrix on the evidence of bones still with matrix (77579 & 77580) and the matrix sample itself.
https://mc06.manuscriptcentral.com/cjes-pubs Page 5 of 43 Canadian Journal of Earth Sciences
5
70 The fossils were limited to a small pocket within the cave, fide McNatt’s and Mychajlowycz’s
71 May, 1984, field notes. Samples such as this are usually random remnant agglomerations of
72 specimens from various ages and may represent a broad temporal sample.
73 The deposit that contained the bones and teeth is a typical brownish-buff, fine-grained
74 powdery limestone cave earth lacking any stratification, sand grains or pebbles. It is the residue
75 of the dissolved limestone bedrock in which the cave system formed. Accumulation of such
76 earths occur in any solution cavity that is available and is not scoured by floods. The occurrence
77 of a mass of small pieces of bone intermingled with the earth suggests that there was minimal
78 scouring of the deposition site. The fossils may have accumulated on a dry cave floor by the 79 mixing of small bone fragments derivedDraft from carnivore chewing and faeces, mixed with 80 uningested larger bone fragments and teeth from animals that died in or were brought in as prey
81 into the cave, and cave earth stirred by animal feet in a dry floor deposit. No microvertebrate or
82 invertebrate fossils were recovered from matrix dissolved in dilute (10% by volume) glacial
83 acetic acid in tap water. I interpret the age of this deposit as Late Pleistocene (Rancholabrean)
84 based on the faunal composition and the presence of Bison. Radiometric dating based on faunal
85 evidence is open to major errors when early representatives or lingering descendants of groups
86 are sampled and considered to represent animals from the main time range for the deposit. The
87 presence of the extinct species Tremarctos floridanus places part of the fauna earlier than ca.
88 10,000 BP, and thus probably not Holocene. All the other mammals could have been
89 contemporaneous with the extinct bear. However, a small cave deposit that is from a small
90 pocket, as this is, is notoriously hard to date (e.g., Swartkrans ‘Hanging Remnant’, Brain 1985)
91 and this sample might have taken considerable time to have accumulated. The absence of bison
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 6 of 43
6
92 and black bear in the region today may reflect mankind’s hunting presence but might instead
93 reflect climatic change and the withdrawal of some forms into more boreal regions. This would
94 suggest that an earlier age for the fauna might explain the differences between the cave fauna and
95 the modern fauna of the region. A faunal age is therefore used.
96 Preparation of Specimens and Measurements. The specimens were preserved within a
97 porous calcareous cave breccia matrix. Preparation of most specimens was mechanical to free
98 them from the cave earth or neighbouring bone fragments. Prior to photographing, a few
99 specimens were treated briefly in 10% dilute acetic acid in tap water to clear the finer details.
100 The two equid milk teeth visible in matrix blocks (77579 & 77580) were prepared by dissolving 101 the blocks in changes of dilute acetic acid.Draft Specimens were coated with two applications of a 102 dilute solution of polyvinyl acetate in acetone, with the field specimen numbers emplaced
103 between coats. The Royal Ontario Museum, Department of Vertebrate Palaeontology accession
104 numbers were later applied over white background patches. The specimens were then
105 photographed. The residue remaining after the acid treatment of ca. 15 litres of matrix was
106 examined for microfossils (e.g., rodent or bat teeth or bones) but none was found.
107 The specimens are identified based on comparisons with museum specimens and from
108 published descriptions of Pleistocene materials. Measurements (in millimetres) of the significant
109 specimens were taken with dial calipers.
110 The specimens are accessioned in the Collection of Vertebrate Fossils, Department of
111 Palaeobiology, Royal Ontario Museum, Toronto, Numbers 77555-77583 & 69680. Each number
112 is prefaced by ‘ROM.VP’, which is omitted here.
113 A remnant matrix sample with a partial lower milk premolar is conserved as 69680.
https://mc06.manuscriptcentral.com/cjes-pubs Page 7 of 43 Canadian Journal of Earth Sciences
7
114 Uncatalogued Materials in bags: S3 - Bag of nine small bone fragments. Unidentified.
115 S6 - Bagged archaeological artifacts (see Mychajlowicz’s field notes). a. Two large chert flakes,
116 struck from a core; b. Two sherds, plain, red pottery ware. S9 - Bag of matrix sample, either
117 ‘cave dust’ or debris from acid preparation.
118 Systematic Palaeontology
119 .
120 Dasypus bellus, extinct armadillo (Xenarthra, Dasypodidae)
121 Three osteoderms from the anterodorsal area of a pectoral shield (77582a, b & c: Figs
122 2A, 2B & 2C). Measurements: diagonals across hexagons: a – 13.5, 12.7, 11.6; b – 18.8, 18.0,
123 16.0, & c – 10.2, 10.0, 9.9. Draft
124
125 Panthera atrox (Felis atrox), extinct American lion (Carnivora, Felidae)
126 Midshaft fragment of a robust ?left ulna (77569; Fig. 3A, section), rounded triangular in
127 section, ends rounded by removal of small fragments, weathered. Probably chewed by carnivore
128 lacking crushing molars, i.e., not a canid. Measurements: length ca. 84: greater and lesser
129 diameters 29.5 & 20.5; wall thickness 4.5 - 6.5.
130 Midshaft fragment of ?left radius (77570; Figs 3B, section, & 4A), rounded semi-circular
131 in section and ends rounded. Both 77569 & 77570 weathered by flaking and spalling.
132 Specimens 77569 and 77570 are possibly from the same individual. Measurements: length ca.
133 64; greater and lesser diameters 30.5 & 22.5; wall thickness 4.0 - 5.0.
134 The American lion (Panthera atrox) was widespread but is a relatively infrequent fossil
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 8 of 43
8
135 usually represented by isolated or fragmentary bones or teeth, as are most felines. The full
136 skeleton is well-known from Rancho La Brea, California (Merriam and Stock, 1932). Bravo-
137 Cuevas et al.’s (2016) report of P. atrox from the Mexican state of Hildalgo, west of Belize, is
138 based on an isolated canine.
139 Specimens 77569 & 77570 possibly from same individual.
140 Puma concolor, puma or cougar (Carnivora, Felidae)
141 Proximal phalanx (77555), possibly from right digit III or IV. Large carnivore, probably
142 P. concolor (Figs 4E & F). Measurements: length 40.2; proximal width 15.3; proximal depth
143 12.3; midshaft width 9.4; midshaft depth 8.9; distal width 11.8; distal depth 8.8. 144 Proximal end of right femur (77557Draft), with the trochanteric fossa, eroded on the greater 145 trochanter, and showing punctates from carnivore teeth on the compact bone below the fossa
146 (Fig. 4K). This specimen appears small for P. atrox and large for recent P. onca. Due to its
147 worn and partial nature, it is tentatively assigned to a feline, either P. concolor augusta or P.
148 onca. Measurements (either worn or taken over insignificant markers) indicate its intermediate
149 size: transverse width 47.4 (lateral surface of trochanter to base of collis caput femoris);
150 anteroposterior width 20.0 over lesser trochanter; anteroposterior & proximodistal diameters of
151 collis capitis femoris 18.0 & 21.4 respectively; shaft width 20.5e (estimated as it is worn); shaft
152 depth 23.5.
153 The puma or mountain lion (Puma concolor) is present in Central America today; it
154 varies in size over its distribution from Alaska or Quebec to Chile and Argentina making
155 distinction of Panthera or Puma phalanges difficult.
156 Panthera onca (= Felis onca), jaguar (Carnivora, Felidae)
https://mc06.manuscriptcentral.com/cjes-pubs Page 9 of 43 Canadian Journal of Earth Sciences
9
157 Right lower fourth premolar, RP4 (77556), worn on cusps to expose dentine, in dentary
158 fragment (Figs 3D and E). Measurements of enamel crown: mesiodistal length 19.5; height of
159 main cusp 11.4. worn (13.5e unworn); buccolingual widths - mesial 8.6; midcrown 9.4; distal
160 9.8.
161 Damaged metapodial shaft remnant (77577, Fig. 4I), from a carnivore as large as P. onca.
162 Gnawed by rodents on one face. Measurement: shaft diameter 12.1.
163 Comments. 77555 & 77556 are identified as ‘carnivore’ based on their morphology;
164 77556 matches the fourth lower permanent premolar of P. onca, and 77556 & 77557 are similar
165 to the appropriate skeletal elements of P. onca though 77557 is somewhat small. Specimens 166 77569 & 77570 are large enough to deriveDraft from a large jaguar. 77569 is sufficiently massive 167 possibly to derive from a small lion (Panthera atrox) or sabretooth (Smilodon fatalis), but both
168 show straight shafts and none of the curvature or torsion typical of Smilodon. Smilodon sp. is
169 reported from the Late Pleistocene Hormiguero Local Fauna of El Salvador (Stirton and Gealey
170 1949; Webb and Perrigo 1984), and both Smilodon and P. onca from the Rancholabrean Cedazo
171 Fauna, Aguascalientes, Mexico (Mooser and Dalquest 1975b).
172 The gnawed metapodial shaft 77577 appears feline and is assigned to P. onca, but may
173 derive from the spectacled bear (Tremarctos floridanus, vide infra).
174 The jaguar is a member of the Recent Central American mammal fauna (Hall and Kelson,
175 1959) and is widely recorded throughout the Late Pleistocene of the United States (FAUNMAP,
176 1994, 376-379), and Central and South America. The Wisconsin form was 15-20% larger than
177 the extant form (Kurtén and Anderson, 1980, 192; Seymour, 1993), probably as a response to the
178 cooler ice age environment.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 10 of 43
10
179 Tremarctos floridanus, Florida spectacled bear (Carnivora, Ursidae)
180 The isolated lower left second molar, LM2 (77573), with unworn wrinkled enamel, has
181 fine wrinkled crown enamel similar to teeth of the extinct Florida spectacled bear (Tremarctos
182 floridanus) (Figs 2F & 4G). It is similar in size to the corresponding tooth in both the North
183 American black bear (Ursus americanus) and the extinct short-faced bear (Arctodus pristinus)
184 (Tabs 1 & 2). Means of lengths and widths of M2's of Recent U. americanus (19.98 & 12.02,
185 respectively; N = 25-30 individuals; Kurtén (1963, tab. 2)) compare well with those for the
186 Extinction Cave molar. Comparisons with M2's of T. floridanus (Kurtén, 1963, 1966) and A.
187 pristinus show 77573 to resemble those of T. floridanus in both morphology and size, although
188 its crown is narrower occlusally than theDraft T. floridanus M2's illustrated or measured by Kurtén
189 (1966, table 1). Table 1 gives means and ranges of M2's of these species from Kurtén (1963).
190 The Extinction Cave tooth 77573 best resembles in size an isolated tooth from Seminole, Florida,
191 or one in a jaw from San Josecito Cave, Mexico (Stock, 1950).
192 Kurtén (1966, tab. 1) gives measurements for M2's of T. ornatus and T. floridanus (Table
193 2). The Recent South American species T. ornatus is smaller in all dimensions but overlaps the
194 observed ranges of the fossil North American species T. floridanus (Table. 2). The Extinction
195 Cave molar’s measurements lie within the ranges for the fossil T. floridanus tooth and near the
196 means for dimensions that do not suffer change due to wear, but near the maximum size for
197 specimens of Recent T. ornatus. Thus the metrical data confirm the observation that the
198 Extinction Cave molar best resembles one illustrated by Kurtén (1966, figs 4C & D). This tooth
199 is assigned to Tremarctos floridanus on the fine morphology of the crown enamel and its size,
200 which match teeth for T. floridanus reported in the literature.
https://mc06.manuscriptcentral.com/cjes-pubs Page 11 of 43 Canadian Journal of Earth Sciences
11
201 The bear is a Tremarctos species. It is small for Arctodus, even for A. pristinus, which is
202 only known from the east coast of North America and east of the Mississippi River (FAUNMAP
203 1994, 412-413; Richards et al., 1996). Rancholabrean Tremarctos floridanus is known from
204 Florida and coastal Gulf of Mexico at Ingleside, Texas (Lundelius, 1972) and San Josecito Cave,
205 Nuevo Leon, Mexico, where it is associated with the black bear Ursus americanus (Stock, 1950).
206 A nearly complete skeleton of a juvenile and probably female T. floridanus was found on the
207 floor of Cebada Cave within the Chiquibul cavern system (Miller, 2000, 65; H.G. McDonald,
208 pers. comm., 1999). The genus is extinct in North America post-Wisconsin but persists in South
209 America as the smaller spectacled bear, T. ornatus, which ranges into eastern Panama (Nowak, 210 1991; Peyton, 1980). The Belizean recordDraft suggests a continuous Pleistocene Texas- to-South 211 America distribution when the Gulf of Mexico coastal plain connecting Florida to Yucatan was
212 exposed by a 26-31 m lower sea level than present today (Watts and Hansen, 1988).
213 An adult right iliac fragment (77571), showing some sacroiliac suture (auricular surface),
214 and chewed by carnivores and/or rodents, is too small for jaguar or horse, appears similar to ilia
215 of some carnivores, and thus may derive from T. floridanus.
216
217 Equus conversidens, Mexican half-ass (Perissodactyla, Equidae)
218 A left upper cheektooth, probably M1, less likely, P4 (77558; Fig. 4D), is the only whole
219 mature cheektooth with a worn occlusal surface showing the enamel pattern. Crown enamel
220 pattern details (Fig. 2D): ectostyles rounded; protocone oblong, flat medial face; ectostyle
221 valleys smoothly concave; no pli caballin; 1 pli protoloph; 1 bifurcated pli protoconule; 2 plis
222 prefossettes; 3 (1 robust) plis postfossettes, no pli hypoloph; enamel of fossette walls not
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 12 of 43
12
223 strongly sinuous nor crenulate.
224 Measurements: mesiodistal diameter of occlusal surface 24.6 (24.9); buccolingual
225 diameter of occlusal surface 24.5 (22.6); protocone length on occlusal surface 12.4 (11.8),
226 preserved crown height from roots over protocone 69.0 & over ectoloph (81.0); (measurements
227 in parentheses are averages for the two P4's of the type palate no. 403; Hibbard, 1955, tab. I).
228 The worn first incisor, I1 (77559) preserves no infundibulum or mark and derives from an
229 aged individual. Measurements: mesiodistal diameter 11.6; buccolingual diameter 14.6, on worn
230 enamel margins of occlusal surface.
231 A lower molar fragment, (77560), possibly from a RM1, preserves the entoflexid area and 232 is from a mature adult. Measurement: preservedDraft crown height 81.0 (cf. 77558 above).
233 Two right upper milk premolars, ?p3 (77561, Fig. 3C), lightly worn on ectoloph crests
234 and major cusps, and ?p4 (77562) lacking protocone, worn on ectoloph crest and beginning wear
235 on lingual cusps, both unrooted, from foals. A right upper milk premolar, ?p2 (77579), newly
236 worn on ectoloph and beginning wear on lingual cusps, has cement lingually, mesially and
237 distally; buccal face of metacone missing, unrooted. Developed from matrix block (Fig. 4B),
238 77579 is slightly larger than 61 but at same stage of wear. (Comparative measurements in
239 Tables 5 & 6.)
240 A left upper milk premolar, ?Lp4 (77563), unworn and unrooted, from a foal, matches
241 77561 and is probably from the same individual as 77561 & 62.
242 Two isolated lower right milk premolars, p3 or p4 (77564 & 77580), lacking cement,
243 unworn and unrooted, are from a young foal (Measurements in Tab. 4). 77580 was developed
https://mc06.manuscriptcentral.com/cjes-pubs Page 13 of 43 Canadian Journal of Earth Sciences
13
244 from matrix block (Fig. 4C).
245 Four milk premolar scraps (77565), comprising a worn lower milk molar entoflexid area,
246 two fragments of milk upper molars, one unworn, and a worn lower molar fragment with
247 entoflexid and metastylid (77566), showing adult entoflexid and compressed and elongated
248 metastylid, are also recognized together with a distal fragment of a splint metapodial (77583).
249 The small New World Mexican half-ass, Equus conversidens, was common during the
250 Pleistocene in North America and ranged from southern Alberta and Saskatchewan to the U.S.-
251 Mexican border states (Churcher 1968, 1969; FAUNMAP 1994, 427) and south into Mexico and
252 Central America. It was first described from the Valley of Mexico by Owen (1869) and later 253 recorded from Loltún Cave, Yucatán, northDraft of Belize, by Alvarez (1982) where it was the most 254 numerous taxon recovered. Azzaroli (1998) considered E. conversidens Owen 1869 the senior
255 synonym for E. barcenai Cope 1884 and possibly also for Onager zoyatalis Mooser 1958. Apart
256 from noting significant occurrences of this equid in the United States, he notes the sample of
257 skeletal elements, including two skulls, several jaws, and a large number of postcranial bones
258 from San Josecito Cave, Nuevo Leon (Stock, 1953), Barranca del Muerto, Tequixquiac, and
259 Tosopoco, Sonora, all in highland Mexico. The upper molar (777558) has the typically fine
260 enamel ridges and the lower molar fragment (777566) the compressed metastylid characteristic
261 of equines that are asses or half-asses and not true horses. The enamel sinuosity here may
262 confirm it as E. conversidens.
263 Comments: The molar specimens 77558-77564 all derive from a small equid, which is probably
264 the Mexican half-ass, Equus conversidens. 77558, 77560 & 77566 could derive from a single
265 prime adult, 77559 from an aged adult, and 77561-77564 from a new foal on the minimal wear
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 14 of 43
14
266 on the milk cheekteeth. Two other milk cheek teeth (upper 77579 & lower 77580) also derive
267 from foals and 77579 is probably from a third foal. These specimens represent at least two adult
268 and two young animals. The crown patterns of molar 77558 and fragments 77560 & 77566 are
269 consistent with details of adult teeth of this equid (Churcher, 1968; Mooser and Dalquest 1975b)
270 and measurements of 77558 fall within ranges for the dimensions of M1's for post-Wisconsin E.
271 conversidens from Medicine Hat (mesiodistal length 25.0-19.5, buccolingual width 25.9-21.2,
272 and protocone length 12.8-10.5; Churcher, 1969, 252). The crowns of the milk premolars are
273 insufficiently worn and show no enamel characters. Milk premolar 77579 is slightly larger and
274 may represent a second, slightly larger foal. All measurements lie within the variation observed 275 in E. conversidens milk teeth from the MedicineDraft Hat Buried Valley deposits, Alberta (Churcher,
276 1969). The lower right milk premolar, p3 or p4 (77580), matches in size two milk premolars
277 from Medicine Hat (Churcher, 1969; lengths without hypoconulid 32.7 & 31.9, buccolingual
278 width without cement 11.0).
279 Equid materials are also reported from 14 localities in Costa Rica, but most of these
280 occurrences ‘lack stratigraphic documentation and/or are from superficial (upper Quaternary)
281 deposits’ (Lucas et al., 1997, 417). Equus fraternus was identified by J.W. Gidley in 1905 on an
3 282 upper M and lower P2 and P4 (figured by Alfaro, 1911) from Cedral de Aserrí (Loc 28 of Lucas
283 et al, 1997), but Lucas et al. only refer these isolated teeth to ‘Equus sp.’ Laurito et al. (1993)
284 reported an equid incisor from Barra Honda, Río Nacaome (Loc. 5 of Lucas et al., 1997) which,
285 unless it is Amerhippus s.l., is best referred to Equus s.l. Lucas et al. (1997, tab. 1) list nine
286 occurrences of Pleistocene equids in Costa Rica, five as ‘Equus sp.’ and four as ‘horse’; seven of
287 these samples are lost, including that identified by Gidley, which reduces the usefulness of these
https://mc06.manuscriptcentral.com/cjes-pubs Page 15 of 43 Canadian Journal of Earth Sciences
15
288 records.
289 The teeth from Aserrí illustrated by Alfaro (1911) have a morphology of ‘slender,
290 elongated protocone, complex folded enamel’. The M3 is ‘24 mm long over the ectoloph’
291 [mesiodistal?] ‘and 24 mm wide at right angles to the ectoloph’ (Lucas et al., 1997, 422). These
292 characters and dimensions are possible for E. conversidens, as observed in the Medicine Hat,
293 Alberta population (Churcher, 1969), and do not disagree with the characters and size observed
294 for the adult cheek tooth and fragments available (77558, 60 & 65). Previously E. fraternus was
295 distinguished from E. complicatus on dental characters that have been shown to be individually
296 highly variable. Identification of equid species on isolated teeth is considered unreliable and 297 more an art than a science (Cooke 1950,Draft Wells 1959, Churcher 1970). Azzaroli (1998) 298 considered E. fraternus and E. conversidens closely related.
299 Equus sp. is recorded for the Hormiguero Local Fauna, El Salvador by Webb and Porrigo
300 (1984) as a newly recognised taxon for the locality, but no description or discussion is given.
301 Olson and McGrew (1942) noted Equus sp. from the Yeroconte Local Fauna, Departamento de
302 Copan, Honduras and Webb and Porrigo (1984) record it in the Orillas del Humuya Local Fauna
303 (cutbank of the Río Humuya), Honduras. None of these equids are identifiable to species.
304 Silva-Bárcenas (1993) described a subspecies, Equus conversidens leoni, in the Museum of
305 Geology, National Autonomous University of Mexico, but gave no provenance or details.
306 Pecari (= Dicotyles) tajacu, collared peccary or javelina (Artiodactyla, Tayassuidae)
307 An incomplete worn right upper tooth row, RP3-4- M1-2 (77572: Fig. 2G) in maxillary
308 fragment. The teeth, with alveoli only of P2 and M3, and a worn M1 in which the protocone and
309 hypocone are abraded, show this to be from an aged individual. The bilophodont cheekteeth and
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 16 of 43
16
310 size (Table 3) match those of the collared peccary or javelina (Pecari tajacu).
311
312 A maxillary facial fragment (77568) shows the nasal contact and the internal surface of
313 the nasal passage. It is from a mature individual and may be from the same individual as 77572.
314 The collared peccary, Pecari tajacu, is recognized on a molar row (77572) and a rostral
315 fragment (77568). Kurtén and Anderson (1980) considered it unknown in Rancholabrean faunas
316 in North America, but it is now known from three localities in Florida (Peace River 5A, six local
317 faunas in De Soto County, and the Suwanee River in north Florida) (Hurlburt et al., 2001).
318 Today it is extant over much of Central America (Hall and Kelson, 1959; Nowak, 1991). The 319 Peace River (Florida) 5A peccary mandibleDraft was associated with a fauna of 16 species, including 320 Platygonus compressus, indicating a Rancholabrean age, but its restricted Floridan distribution
321 suggests it represents a brief colonisation episode. Alvarez (1982) records peccary from the
322 limestone cavern of Loltún, Yucatán. Peccaries occupy a great variety of habitats and ‘limestone
323 caves ... serve as winter quarters in some areas’ (Nowak, 1991, 1346). Woodburne (1969)
324 described a full tooth row in a maxilla collected by Barnum Brown from the ‘bed and associated
325 playas of the Río de la Pasión and adjacent Río Santa Amelia, [near] the town of Santa Amelia’
326 from the late Pleistocene of Guatemala. Its age, though uncertain, is considered late or latest
327 Pleistocene (Woodburne, 1969).
328 Stinnesbeck et al. (2017) describe Mukhalia minima n. gen. et n. sp. from a drowned
329 karst cave in Quintana Roo, Mexico. This is dated as probably from the Latest Pleistocene (? =
330 Wisconsinian) and is distinguished by mandibular characters that cannot be compared with those
331 in 77572. M. minima is geographically close and may be somehow related to the Extinction
https://mc06.manuscriptcentral.com/cjes-pubs Page 17 of 43 Canadian Journal of Earth Sciences
17
332 Cave peccary.
333 Camelidae indet., extinct llama, ?Palaeolama mirifica (Artiodactyla, Camelidae)
334 A left radial carpal or scaphoid (77576; proximo-medial carpal) of robust size may
335 represent the medium-sized llama, Palaeolama mirifica, best known in North America from
336 Florida but also from the Texas and Mexican Gulf coast (FAUNMAP 1994, 440-441), or less
337 likely, the widespread North American Pleistocene Western camel, Camelops hesternus, known
338 from the Great Plains region of North America. Identification to species or even genus on this
339 element is unreliable as juvenile carpals may appear adult, and size and morphology are too
340 variable. The element is well ossified and appears adult. The suggested camelid identity of the 341 scaphoid is based on size, the known RancholabreanDraft distribution of llamas, and the coastal Gulf 342 distribution of P. mirifica. Measurements: anteroposterior diameter 42.9; transverse diameter
343 25.9; proximodistal diameter 33.0.
344 The llama scaphoid sets a limital record for any North American camel. Camelops cf.
345 hesternus and C. traviswhitei are described from the Rancholabrean Cedazo Fauna,
346 Aguascalientes (Mooser and Dalquest, 1975a), C. cf. hesternus from Tequixquiac, Mexico
347 (Hibbard, 1955), a large species of Tanupolama (= Hemiauchenia) from the Cedazo Fauna,
348 Aguascalientes by Mooser and Dalquest (1975a), and Camelops sp. and Tanupolama (=
349 Hemiauchenia) mirifica from the Late Wisconsinan of Ingleside, Texas (Lundelius, 1972) and
350 Emery Borrow Pit, California (Kurtén and Anderson, 1980). Alvarez (1982, 30) reported a
351 single proximal phalanx of ‘Tanupolama’ from the cave of Loltún, Yucatán, identified it as too
352 small (‘es pequeña para’) for Camelops, and assigned it to Tanupolama. P. mirifica is best
353 known in North America from Florida but is sparse on the Texas and Mexican Gulf coast
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 18 of 43
18
354 (FAUNMAP 1994, 440-441). Isolated cameline carpals are not specifically identifiable, so this
355 bone may represent either a southern limital C. hesternus or a Caribbean coastal record of P.
356 mirifica (Webb, 1974). C. hesternus is not known from southern Mexico and this carpal may
357 represent P. mirifica, though absence of evidence is no evidence of absence. Camelops is
358 reported from Honduras (Alvarez, 1982) and Camelidae indet. from Guatamala, Honduras, El
359 Salvador and Nicaragua. The nearest record of P. mirifica is in Mexico (Bravo and Jimenez,
360 2015).
361 Mazama americana, brocket deer (Artiodactyla, Cervidae)
362 A mature worn right lower M2-3 series (77575, Figs 3G & 4H), in a dentary fragment. 363 Three small ectostylid pillars are presentDraft between the buccal lophs and an oblique crista is
364 present on the mesiobuccal face of the protoconid of M2. This represents a small cervid,
365 identified as the Nearctic red brocket, Mazama americana (Table 4). The identification as
366 Mazama americana rests on its small size, smooth dental enamel, and simple tooth pattern,
367 which precludes a small individual of Odocoileus and matches Mazama. Two cervid metapodial
368 shaft fragments (77567 & 81) match in size and cross-section those of M. americana.
369 The red brocket deer, Mazama americana, is extant in Belize. Mazama is also reported
370 from the cavern of Loltún, Yucatán by Alvarez (1982): he gives the modern local species as M.
371 gouazoubira, contra Nowak (1991) who uses M. americana and limits the former to Panama and
372 further south and east. Mazama is also known from Guatemala.
373 Bison sp., cf. B. bison, New World bison (Artiodactyla, Bovidae)
374 An isolated left lower third molar, LM3 (77574, Fig. 3F), in any early stage of wear and
375 lacking roots, is indistinguishable from modern Bison bison. However, identification of Bison to
https://mc06.manuscriptcentral.com/cjes-pubs Page 19 of 43 Canadian Journal of Earth Sciences
19
376 species on single teeth is unreliable. A fragment of the medial surface of an unworn left lower
377 premolar, P3 (77578), best matches that tooth of Bison bison.
378 Measurements: mesiodistal lengths, overall 45.1, and over paraconid-metaconid 36.0;
379 buccolingual widths at midcrown height, 17.0 over protoconid-paraconid, 17.5 over hypoconid-
380 metaconid; crown height of enamel above cingulum on metaconid, 66.1. Ectostylid pillar
381 present between protoconid and hypoconid only.
382 Mooser and Dalquest (1975b) give the alveolar (mesiodistal) length of M3 of their new
383 species Bison aguascalientes, from the Cedazo Fauna, Aguascalientes, Mexico as 49.6 mm,
384 which compares reasonably with 45.1 for the unworn Extinction Cave tooth. 385 Wisconsinan Bison is known fromDraft the greater part of North America south of the last ice 386 sheets (Kurtén and Anderson, 1980) including Mexico and the United States west of the
387 Mississippi River (FAUNMAP 1994, 460-461). Bison chaneyi and B. occidentalis are reported
388 from Tequixquiac, Mexico (Hibbard, 1955) and Bison aguascalientes from the Cedazo Fauna,
389 Aguascalientes, Mexico (Mooser and Dalquest, 1975b). Mooser and Dalquest (1975b) gave the
390 alveolar (mesiodistal) length of M3 of B. aguascalientes as 49.6, which compares reasonably
391 with 45.1 for the unworn Extinction Cave tooth. Alvarez (1982) reported a partial Bison
392 humerus from the limestone cavern of Loltún, Yucatán, Mexico. Lucas et al. (1997) record its
393 presence in Honduras and El Salvador. While Bison was present in the Central American fauna,
394 it is considered to have been a minor member in that it did not form massive herds as the
395 environment was rough bush rather than grassy plains.
396 Ecological Interpretation. The fauna includes animals not usually found in caves, such as
397 horse, llama, and bison: ungulates that are often considered as plains adapted but may be found
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 20 of 43
20
398 in broken wooded country. Their presence in a cave may be attributed to their occurrence as
399 prey items brought in by carnivores. Remains of animals such as lion, jaguar, puma, bear and
400 peccary are often found in caves: all five of them use caves as lairs, wintering dens for
401 hibernation, or transient resting places. The fragments of carnivores suggest that these animals
402 alternated in their occupations as cohabitation seems unlikely. Any of the carnivores could have
403 been the agent that brought parts of other animals into the cave as food. The presence of milk
404 teeth of equids and the unworn condition of the bison cheektooth show that immature animals
405 comprised a major part of the prey spectrum. The aged peccary may show that old animals were
406 taken, or that it retired to the cave to die. No remains of cave dwelling rodents, bats, birds, 407 small carnivores, reptiles or amphibiansDraft were found. 408 The presence of multiple carnivore species in such a small faunal sample is unusual, and
409 may be attributed to the assembling propensities of caverns for fossils. Morgan and Seymour
410 (1997) record co-occurrences of P. concolor and Miracinonyx inexpectatus in Florida and other
411 taxa found with either one or both. They list 14 sites in five of which Smilodon fatalis (Seymour
412 et al., 2018) and Tremarctos floridanus co-occur with P. concolor, and usually with Canis dirus
413 (dire wolf). In only one deposit is a third felid known (Panthera atrox in the Cutler Hammock
414 carnivore den, Florida), which suggests that S. fatalis did not compete directly with P. concolor
415 for prey or shelter. Emslie and Morgan (1995) consider the taphonomy of the Cutler Hammock
416 den. They date it on its taxa as late Pleistocene Rancholabrean, an age that matches the probable
417 age of the Extinction Cave fauna. Both caves are karstic solution cavities in limestones, and lie
418 at similar heights above sea level. Cutler Hammock and Extinction Cave share Tremarctos
419 floridanus, Panthera onca, P. atrox and Puma concolor. All four carnivores probably used
https://mc06.manuscriptcentral.com/cjes-pubs Page 21 of 43 Canadian Journal of Earth Sciences
21
420 caves for dens or shelter, but probably separately. Any of them might bring prey items into the
421 cave and thus contribute to the variety of herbivore taxa identified. Emslie and Morgan remark
422 that Cutler Hammock is unusual in that it yielded four large felids (Puma concolor, Panthera
423 onca, P. atrox, and Smilodon fatalis). (Smilodon may be present in the Extinction Cave fauna as
424 some specimens are difficult to identify.) Nonetheless, Extinction Cave is remarkable in
425 yielding three large felids and a bear.
426 The small horse is represented by the largest number of recovered elements (11 complete
427 teeth and other fragments) and individuals (at least 3, possibly 5), the jaguar is the next with 5
428 elements recovered, but perhaps from only one individual, the lion and bison are represented by 429 two specimens but probably only one individualDraft of each, and the other taxa are represented by 430 single individuals and few specimens. The lion, jaguar and puma are strictly carnivorous and
431 may have preyed on members of the other taxa, or scavenged stray carcasses. Bear and peccary
432 are essentially omnivorous and will eat a wide variety of foods, although modern Peruvian
433 Tremarctos ornatus is largely vegetarian (Peyton, 1980). Horse and bison are mainly grazers,
434 the small deer has a mixed diet of browse and forbs, and most New World Pleistocene camels
435 were mixed browsers that did not ingest large amounts of grass or ground-level vegetation
436 (Dompierre and Churcher, 1996). Thus, the diversity is great considering the restricted sample
437 size and the few specimens. Comparisons with the fauna of Cutler Hammock show a similarity
438 in the more numerous representations of immature equid remains (Cutler Hammock - 2 adults,
439 17 juveniles: Extinction Cave - 1 prime and 1 aged adult, and 2 or 3 juveniles) and both caves
440 appear to owe most of the larger fossil assemblage to the actions of their carnivore residents.
441 Tooth marks on the bones comprise punctates, heavy scorings and rodent and other
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 22 of 43
22
442 gnawings (77557 & 77571). Large cats such as jaguars do not typically gnaw bones and there is
443 no direct evidence for a gnawing and crushing carnivore such as a coyote (Canis latrans) or wolf
444 (C. lupus or extinct C. dirus) from the bone debris in the cave breccia. There is no fossil
445 evidence of rodents but only incisive tooth marks ca. 2.5 mm across (77571 & 77577) which
446 suggests a rat-sized animal such as packrats (Neotoma) or cotton rats (Sigmodon). These gnaw
447 marks are too small for Mexican porcupines (Coendou mexicanus) and too large for voles, e.g.,
448 Microtus.
449 The environments inhabited by modern members of the cave species differ only slightly
450 from that of the area today. Jaguar, puma and collared peccary are comfortable in broken, well 451 wooded country, and find caves convenientDraft for denning or resting. Peccary and brocket inhabit 452 bushy woodland and thickets as these give good cover and a wide range of foodstuffs. Bears use
453 caves for denning and sleeping, although the modern T. ornatus generally does not do so
454 (Nowak, 1991). However, only jaguar and puma carry their kills to a cave for consumption.
455 Colts of the half-ass would be easy to carry because of their small size, a dead brocket weighs
456 less than a young equid, and a peccary might not weigh much more than a colt. Pieces of llama
457 and bison may have been scavenged and carried in the manner that canids will carry bones long
458 distances to a den. The fragments of lion, jaguar, puma and peccary may represent ones that died
459 in the cave, and were scavenged by later occupants. Llama, horse and bison require reasonable
460 grazing in nearby areas, and perhaps signal that the limestone upland strata where the caves
461 occur would have supported a mixed grassland with small bushes and trees or a scrub savanna.
462 The lack of evidence for canids is unexpected.
463 It is noteworthy that only the armadillo represents a true South American mammal, and
https://mc06.manuscriptcentral.com/cjes-pubs Page 23 of 43 Canadian Journal of Earth Sciences
23
464 that a single llama represents a possible South American derivative from an invading North
465 American stock, while the remainder of the taxa are of Nearctic origin.
466 Few pieces of bone from the matrix blocks in samples 77579 and 77580 are large and not
467 rounded or abraded, suggesting not having passed through a digestive tract. Analysis of the
468 solution fine residue yielded no reptile, bird, bat, insectivore, or rodent bones or teeth. The
469 samples received include pottery and flint tools brought in by man.
470
471 Conclusions. Ten or 11 mammals are recognised from Extinction Cave († = extinct taxon):
472 Xenarthra: armadillo (†Dasypus bellus): Carnivora: puma or mountain lion (Puma concolor), 473 jaguar (Panthera onca), American lion (†DraftPanthera atrox) and Florida bear (†Tremarctos 474 floridanus) with also the possibility of sabre-tooth cat (†Smilodon fatalis ); Perissodactyla;
475 American horse or half-ass (†Equus conversidens); Artiodactyla; collared peccary or javelina
476 (Pecari tajacu), red brocket deer (Mazama americana), South American llama (?†Palaeolama
477 mirifica), bison (Bison cf. bison). These identifications are based on cheek teeth or skeletal
478 fragments, some of which have been chewed by rodents.
479 The Extinction Cave fauna is remarkable for a high proportion of carnivores, and the
480 absence of micromammals, birds, reptiles and amphibians. In terrestrial mammalian
481 palaeofaunas carnivores usually comprise ca. 4% of the taxa but in Extinction Cave they are ca.
482 33% (4/11 or 12 taxa). No Canidae or Procyonidae are represented and it is surprising that these
483 two scavenging families are absent as they are usually present in Nearctic palaeofaunas. Cats are
484 generally predatory and usually avoid scavenging. Bears are more omnivorous and often accept
485 small prey, or others’ kills.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 24 of 43
24
486 The cave was used as a den by the carnivores and perhaps as a shelter by the peccary and
487 armadillo. It is a remnant karstic water conduit in a limestone hill that is isolated with two
488 openings, and provides good shelter for those animals that require a quick exit.
489 Humans were present in the caves as pottery shards and tools attest, but these may
490 represent Holocene visits rather than Late Pleistocene/Holocene or Rancholabrean incidents.
491 None of the herbivores are large, even when adult. Juvenile half-asses and brocket deer
492 are easy prey for either the puma or jaguar to bring into the cave. The American lion could kill
493 larger prey such as adult half-asses, peccary, llama or bison. Only tooth marks on bone
494 fragments attest rodents, and these are not identifiable to species. 495 The environment suggested by theDraft fauna is a mixed woodland with grazers (half-ass, 496 bison) and browsers (brocket deer, llama), and with peccaries rooting for forbs and tubers. The
497 large cats may have ambushed the herbivores, both juvenile and aged adult, such as half-ass and
498 peccary.
499
https://mc06.manuscriptcentral.com/cjes-pubs Page 25 of 43 Canadian Journal of Earth Sciences
25
501 Acknowledgements. I wish to thank Logan McNatt, of the Department of Archaeology,
502 Ministry of Natural Resources, Belmopan, Belize for collecting the fossils from Extinction Cave,
503 for allowing Ms. Maryijka Mychajlowycz to carry them to Toronto for our eventual study, and
504 for access to his field notes on the cave and the fossils’ collection. I thank Ms. Mychajlowycz
505 for bringing the collection to Toronto, for cleaning the specimens and for performing the
506 preliminary identifications. I also wholeheartedly acknowledge Ms. Mychajlowycz’s help as my
507 laboratory assistant when I was active in the Dept. Zoology, Univ. Toronto and Research
508 Associate, in the Dept. Palaeobiology, Royal Ontario Museum, Toronto, Canada. Dr. H.
509 Gregory McDonald of the U.S. Bureau of Land Management, Salt Lake City, Utah offered 510 valuable advice over the identification ofDraft the Tremarctos floridanus molar. Dr. Thomas E. 511 Miller of the Dept. Geology, University of Puerto Rico, Mayagüaz, gave generously of his
512 advice and caving experience in Belize. Dr. Kevin Seymour of the Dept. Palaeobiology, Royal
513 Ontario Museum, Toronto allowed me access to his departmental modern osteological collection
514 and Dr. Judith Eger also of the Dept. Mammalogy, Royal Ontario Museum, Toronto allowed me
515 similar access to her departmental osteological collection. I thank my wife, Bee, for
516 photographing the fossils and for the half-tone views of the illustrated specimens, and my
517 daughter Jaclyn for assistance with computer matters.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 26 of 43
26
References.
Alfaro, A. 1911. Comprobaciones geológicas. Boletín del Fomento (San José, Costa Rica), 1:
123-131.
Alvarez, T. 1982. Restos de mamíferos recientes y pleistocénicos procedentes de las grutas de
Loltún, Yucatán, México. In Restos de moluscos y mamíferos cuaternarios procedentes de
Loltún, Yucatán. Edited by T. Alvarez and O.J. Polaco. Cuaderno de Trabajo, Departamento de
Prehistoria, Instituto Nacional de Antropologia e Historia, 26: 7-31
Azzaroli, A. 1998. The genus Equus in North America - The Pleistocene species.
Palaeontographia Italica, 85: 1-60. Brain, C.K. 1985. New insights into earlyDraft hominid environments from the Swartkrans Cave, 325-343. In L’Environnement des Homindés au Plio-Pléistocène. Edited by Y. Coppens. Paris,
Masson.
Bravo, V., and E. Jimenez. 2015. First reported occurrence of Palaeolama mirifica (Camelidae,
Lamini) from the Late Pleistocene (Rancholabrean) of Puebla, central Mexico. Boletin de la
Sociedad Geologica Mexicana 67: 13-20.
Bravo-Cuevas, V.M., M.A. Cabral-Perdumo and M.A. Pineda-Maldonaldo. 2016. First
Occurrences of Panthera atrox (Felidae, Pantherinae) in the Mexican state of Hildalgo and a review of the record of felids from the Pleisrocene of Mexico. Fossil Record 16, 131-141.
Churcher, C.S. 1968. Pleistocene ungulates from the Bow River gravels at Cochrane, Alberta.
Canadian Journal of Earth Sciences, 5: 1467-1488.
Churcher, C.S. 1969. A Fourth Report on Some Pleistocene Localities in Southern Alberta and
Saskatchewan and their Vertebrate Fossil Faunas. Unpublished Report to The Department of
https://mc06.manuscriptcentral.com/cjes-pubs Page 27 of 43 Canadian Journal of Earth Sciences
27
Energy, Mines and Resources, Geological Survey of Canada, Ottawa.
Churcher, C.S. 1970. The fossil Equidae from the Krugersdorp Caves. Annals of the Transvaal
Museum, 26: 145-168.
Cooke, H.B.S. 1950. A critical revision of Quaternary Perissodactyla of southern Africa.
Annals of the South African Museum, 32: 393-479.
Cope, E.D. 1884. The extinct Mammalia of the Valley of Mexico. Proceedings of the
American Philosophical Society, Philadelphia, 22: 1-21.
Dompierre, H., and C.S. Churcher. 1996. Premaxillary shape as an indicator of the diet of seven
extinct Late Cenozoic New World camels. Journal of Vertebrate Paleontology, 16: 141-148. Emslie, S.D. and G.S. Morgan. 1995. TaphonomyDraft of a Late Pleistocene Carnivore Den, Dade County, Florida, 65-83. In Late Quaternary Environments and Deep History: A tribute to Paul S.
Martin. Edited by D.W. Steadman and J.I. Mead, The Mammoth Site of Hot Springs, South
Dakota, Inc., Scientific Papers, Vol. 3.
FAUNMAP. 1994. A Database Documenting Late Quaternary Distributions of Mammal
Species in the United States. FAUNMAP Working Group, Graham, R.W. and Lundelius, E.L.,
Jr., and Collaborators. Illinois State Museum, Scientific Papers, 25: 1-690.
Hall, E.R., and K.R. Kelson. 1959. The Mammals of North America. Ronald Press, NewYork.
Hibbard, C.W. 1955. Pleistocene vertebrates from the Upper Becerra (Becerra Superior)
Formation, Valley of Tequixquiac, Mexico, with notes on other Pleistocene forms.
Contributions from the Museum of Paleontology, University of Michigan, 12: 47-96.
Hurlburt, R.C., Jr., G.S. Morgan and A. Kerner. 2009. Collared peccary (Mammalia,
Artiodactyla, Tayassuidae, Pecari) from the late Pleistocene of Florida. Museum of Northern
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 28 of 43
28
Arizona Bulletin, 15: 543-555.
Kurtén, B. 1963. Fossil bears from Texas, The Pearce-Sellards Series No. 1, 1-15. Texas
Memorial Museum, University of Texas.
Kurtén, B. 1966. Pleistocene bears of North America. 1. Genus Tremarctos, Spectacled Bears.
Acta Zoologica Fennica, 15: 1-120.
Kurtén, B. and E. Anderson. 1980. Pleistocene Mammals of North America. New York;
Columbia University Press.
Laurito, C.A., Valerio, W. and Vega, E. 1993. Nuevos hallazgos paleovertebradológicos en la
Península de Nicoya: implicaciones paleoambientales y culturales de la fauna de Nacaome. Revista Geológica de América Central, Draft16: 113-115. Lucas, S.G., Alvarado, G.E. and Vega, E. 1997. The Pleistocene mammals of Costa Rica.
Journal of Vertebrate Paleontology, 17: 413-427.
Lundelius, E.L. 1972. Fossil vertebrates from the late Pleistocene Ingleside fauna, San Patricio
County, Texas. Reports of Investigations, Bureau Economic geology, University of Texas, 77:
1-74.
Miller, T. 2000. Chiquibul Cave. National Geographic, 197: 54-71.
Merriam, C.H., and C. Stock. 1932. The Felidae of Rancho La Brea. Publications of the
Carnegie. Institute of Washington, no. 422.
Mooser, O. 1958. La fauna “Cedazo” del Pleistocenico en Aguacalientes. Anales del Instituto de Biologia de la Universidad Nacional Autonoma de Mexico, 29: 409-452.
Mooser, O. and W.W. Dalquest. 1`975a. A new species of camel (genus Camelops) from the
Pleistocene of Aguascalientes, Mexico. Southwestern Naturalist, 19: 341-345.
https://mc06.manuscriptcentral.com/cjes-pubs Page 29 of 43 Canadian Journal of Earth Sciences
29
Mooser, O. and W.W. Dalquest. 1975b. Pleistocene mammals from Aguascalientes, central
Mexico. Journal of Mammalogy, 56: 781-820.
Morgan, G.S. and K.L. Seymour. 1997. Fossil History of the panther (Puma concolor) and the
cheetah-like cat (Miracinonyx inexpectatus) in Florida. Bulletin of the Florida Museum of
Natural History, 40: 177-219.
Nowak, R.M. 1991. Walker’s Mammals of the World, edit. 5. John Hopkins University Press.
Olson, E.C. and P.O. McGrew. 1942. Field Museum Paleontological Expedition to Honduras.
Science, 96: 85.
Owen, R. 1869. On the fossil teeth of equines from Central and South America. Proceedings of the Royal Society, London, 17: 267-268.Draft Owen R. 1870. On fossil remains of equines from Central and South America referable to
Equus conversidens Ow., Equus tau Ow., and Equus arcidens Ow. Philosophical Transactions
of the Royal Society, London, 159: 559-573.
Peyton, B. 1980. Ecology, distribution, and food habits of spectacled bears, Tremarctos
ornatus, in Peru. Journal of Mammalogy, 61: 639-52.
Richards, R.L., Churcher, C.S. and Turnbull, W.D. 1996. Distribution and size variation in
North American short-faced bears, Arctodus simus, 191-246. In Palaeoecology and
Palaeoenvironments of Late Cenozoic Mammals: Tributes to the career of C.S. (Rufus)
Churcher. Edited by K.M. Stewart and K.L. Seymour. University of Toronto Press.
Seymour, K.L. 1993. Size change in North American Quaternary jaguars, 343-372. In
Morphological Change in Quaternary mammals of North America. Edited by R.A. Martin and
A.D. Barnosky. Cambridge University, Cambridge, UK.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 30 of 43
30
Seymour, K. L., A. R. Reynolds and C.S. Churcher. In press (2018). Smilodon fatalis from
Talara, Peru: Sex, Age, Mass, and Histology: 30-52. In Smilodon: The Iconic Sabertooth.
Edited by L. Werdelin. H.G. McDonald, and C.A. Shaw. Johns Hopkins University Press,
Baltimore, Maryland, USA.
Silva-Bárcenas, Á. 1993. Vertebrados fósiles del México de Geologia de la UNAM
(Universidad Nacional Autonomia do Mexico). Revista de la Societad Mexicana do
Paleontologia 6: 71-103.
Stinnesbeck, S.R., E. Frey, W. Stinnesbeck, J.A. Olguin, P. Zell, A.T. Mata, M.B. Sanvincente,
A.G. Gonzàlez, C.R. Sandoval and E.A. Nuñez. 2017. A new fossil peccary from the Pleistocene-Holocene boundary of the YukatanDraft Peninsula, Mexico. Journal of South American Earth Sciences, 77: 341-349.
Stirton, R.A. and W.K. Gealey. 1949. Recent geology and vertebrate paleontology in El
Salvador. Geological Society of America Bulletin, 60: 1731-1754.
Stock, C. 1950. Bears from the Pleistocene Cave of San Josecito, Nuevo Leon, Mexico.
Journal of the Washington Academy of Science. 40: 317-321.
Stock, C. 1953. El caballo pleistocénico (Equus conversidens leoni subsp. nov.) de la Cueva de
San Josecito, Aramberri, Nuevo Léon. Memorial del Congresso Cientifico de Mexico, 3: 170-
171.
Watts, W.A. and B.C.S. Hansen. 1988. Environments of Florida in the late Wisconsin and
Holocene, 307-323. In Wet Site Archaeology. Edited by B.A. Purdy. Telford Press, Caldwell,
New Jersey.
Webb, S.D. 1974. Pleistocene llamas of Florida, with a brief review of the Lamini, 170-213. In
https://mc06.manuscriptcentral.com/cjes-pubs Page 31 of 43 Canadian Journal of Earth Sciences
31
Pleistocene Mammals of Florida. Edited by S.D. Webb. Gainesville; University of Florida
Presses.
Webb, S.D. and Porrigo, S.C. 1984. Late Cenozoic vertebrates from Honduras and El Salvador.
Journal of Vertebrate Paleontology, 4: 237-254.
Wells, L.H. 1959. The nomenclature of South African fossil equids. South African Journal of
Science, 55: 64-66.
Woodburne, M.O. 1969. A late Pleistocene occurrence of the collared peccary, Dicotyles
tajacu, in Guatemala. Journal of Mammalogy, 50: 121-125.
518 Draft
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 32 of 43
32
Table 1.
Dimension Extinction T. floridanus (N = 8) A. pristinus (N = 6)
Cave (77573) Max. - Min. Max. - Min.
Mesiodistal length 21.4 22.1 24.0 - 21.5 28.9 29.7 - 28.51
25.6 27.2 - 23.62
Buccolingual width 12.4 12.7 13.2 - 12.0 19.7 20.0 - 19.21
16.6 18.7 - 15.22
Table 1. Measurements of TremarctosDraft lower second molar (M2) from Extinction Cave, Belize; compared with data from samples from Seminole, Florida; Ingleside and Friesenhahn
Cave, Texas; San Josecito Cave, Mexico; and Potter Creek Cave, California. Data are from
Kurtén (1963, tab. 41) and from Richards et al. (1996), Appendix 2, southeast sample from
Florida (2). ‘’ = Mean value.
https://mc06.manuscriptcentral.com/cjes-pubs Page 33 of 43 Canadian Journal of Earth Sciences
33
Table 2.
Dimension Extinction T. floridanus T. ornatus
Cave (77573) Max. - Min. SD Max. - Min. SD
Mesiodistal length 21.4 21.86 24.0 - 19.0 1.06 18.80 21.0 - 16.5 1.26
Buccolingual width, mesial 11.4 13.07 14.6 - 11.8 0.66 11.22 12.6 - 9.9 0.72
.. distal 12.4 12.91 14.5 - 11.6 0.67 11.09 12.4 - 10.0 0.73
crown height, protoconid 8.2 8.04 8.8 - 7.3 0.46 7.38 8.1 - 6.7 0.16
.. metaconid 6.7 7.18 8.2 - 6.6 0.50 6.76 7.5 - 6.2 0.44 .. hypoconid 7.2 Draft 7.41 8.5 - 6.9 0.44 6.70 7.3 - 5.8 0.58 Table 2. Comparative measurements of the Extinction Cave Tremarctos lower second
molar (M2) and those of T. floridanus and T. ornatus. Data are from Kurtén (1966, tab. 1).
‘’ = Mean value; ‘SD’ = Standard Deviation. Sample sizes 23 – 28 for lengths and widths, and
17 - 18 for crown heights.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 34 of 43
34
Table 3
Dimension Extinction Cave toothrow P3-M2 (77572)
Mesiodistal row lengths P3-M2 44.9; P3-P4 19.0; M1-M2 24.8.
Tooth P3 P4 M1 M2 M3
Mesiodistal length on buccal face 18.6 19.7 10.8 13.8 14.0e
Buccolingual width mesially 20.019.5 11.1 11.6 13.8e
Santa Amelia toothrow P2-M3 (AMNH 48799), selected teeth Mesiodistal row lengths Draft P2-M3 63.15 P3-M2 42.4e Tooth P3 P4 M1 M2 M3
Mesiodistal length on buccal face 18.70 19.95 10.95 12.60 13.35
Buccolingual width mesially 19.25 10.55 10.75 12.40
Table 3. Comparative measurements of upper cheekteeth, P3-M2 of Pecari tajacu from
Extinction Cave, Belize and those of a toothrow from Santa Amelia, Guatemala (Woodburne,
1969). ‘e’ = estimated measurement.
https://mc06.manuscriptcentral.com/cjes-pubs Page 35 of 43 Canadian Journal of Earth Sciences
35
Table 4
Dimension 77575
Mesiodistal length over M2-3 23.7
Tooth M2 M3
Mesiodistal length on buccal face 11.0 14.1 (9.1 less hypoconulid)
Mesial buccolingual width (protoconid-paraconid) 7.9 7.2
Distal buccolingual width (hypoconid-metaconid) 7.4 7.0
Buccolingual width of hypoconulid/talon - 3.8 Mesial crown heights - buccallyDraft 4.2 5.2 - lingually 5.5 6.2
Depth of dentary between M2 and M3 on lingual face 20.7
- distal to M3 on lingual face 21.4
Table 4. Measurements of right lower molars, RM2-3 (77575) of Mazama americana
from Extinction Cave, Belize.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 36 of 43
36
Table 5
Dimension 77561 77562 77563 77579
Mesiodistal diameter 31.1 30.3 31.0 33.6
Buccolingual diameter 19.9 - 20.0 25e
Protocone length 21.1 17.2 21.2 10.6
Crown height 30.1 25.8 30e 32.6
Table 5. Measurements of upper milk premolars of Equus cf. conversidens from Extinction Cave, Belize. ‘e’ = estimatedDraft measurement.
https://mc06.manuscriptcentral.com/cjes-pubs Page 37 of 43 Canadian Journal of Earth Sciences
37
Table 6
Dimension77564 77580
Mesiodistal diameter over hypoconulid 32.2 32.7
.. protoconid to hypoconid 20.5 13.4
Buccolingual diameter 10.6 -
Crown height 30.6 30+
Table 6. Measurements of lower milk premolars of Equus cf. E. conversidens from Extinction Cave, Belize. ‘+’ = minimalDraft measurement.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 38 of 43
38
Legends for Figures
Figure 1. Location Map of Extinction Cave, Belize, Central America. Extinction Cave
● ; Settlements ■ ; Roads = heavy lines. Oriented to geographic grid north. Drawn from DGS
958, Edition 4, Government of the United Kingdom, Directorate of Overseas Surveys, 1981.
Figure 2. 77582 A, B & C - Osteoderms from the pectoral shield of Dasypus bellus. 2 A
& 2C – Osteoderms from the centre , and 2B – Osteoderm from near the margin of the shield.
Scale in millimetres. 2D - Crown enamel pattern of equid left upper cheektooth, probably LM1
(77558) of Equus cf. conversidens. 2E - Enamel crown patterns of molars RM (77575) of Draft 2-3 Mazama americana. 2F - Enamel crown pattern of lower left second molar, LM2 (77573), of
Tremarctos floridanus, showing unworn wrinkled enamel. 2G - Enamel crown patterns of an
incomplete worn right upper cheektooth row P3-4- M1-2, with alveoli of P2 and M3 outlined, of
Pecari tajacu. Scales in millimetres.
Figure 3. 3A - Panthera atrox. 77569. Cross-section of ulna. 3B - Panthera atrox.
77570, Cross-section of radius. 3C - Equus cf. conversidens. 77561 - Right milk p3, occlusal view, early wear. 3D & E - Panthera onca. 77556. Right P4, occlusal and buccal views. 3F -
Bison sp. 77574. Left M3, buccal view; note cement remnants in interlophar re-entrants. 3G -
Mazama americana. 77575. Right M2-3, in buccal view. Dentine exposed by wear in black.
Arrows indicate mesial/anterior. For 3A, B & E: d = dorsal/anterior; v = ventral/volar; m = medial, and l = lateral. Scale - All figures to scale except 3F, which is half scale.
Figure 4. 4A -77570. ?Panthera onca, ?Left radius, midshaft portion, ends rounded by
https://mc06.manuscriptcentral.com/cjes-pubs Page 39 of 43 Canadian Journal of Earth Sciences
39
gnawing, antero-lateral aspect. Line indicates locus of section (Fig. 3B). 4B -77559. Equus, cf.
E. conversidens. Right upper milk premolar p4; paracone, protocone, protoconule newly in
occlusion; hypocone area damaged post mortem. 4C - 77580. Equus cf. E. conversidens. Left
lower milk premolar (?p)4 unerupted, occlusal aspect. 4D - 77578. Equus, cf. E. conversidens,
Left P3 or P4, mature adult, mesial aspect, occlusal table to right. NB. Cementum spalled off.
4E -77555. ?Puma concolor, proximal (?right manual) phalanx, abplantar aspect. 4F - 77555.
?Puma concolor, ibid super, lateral aspect (distal to right). 4G - 77573. Tremarctos floridanus.
Left M2, lingual view. 4H - 77575. Mazama americana . Right M2-3, worn, in dentary
fragment, buccal aspect. 4I - 77577. ?Panthera sp., metapodial shaft fragment, plantar aspect,
with rodent tooth marks. 4J - 77556. PantheraDraft onca. Right P4, slightly worn, in dentary fragment, lingual aspect. 4K - 77557. ?Panthera onca or P. concolor, Right femur, proximal
end, gnawed and weathered (half size of other figures).
Scale in millimetres for Figs 4A-J: 4K is half scale.
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 40 of 43
Draft
Location Map of Extinction Cave, Belize, Central America. Extinction Cave ● ; Settlements ■ ; Roads = heavy lines. Oriented to geographic grid north. Drawn from DGS 958, Edition 4, Government of the United Kingdom, Directorate of Overseas Surveys, 1981.
220x193mm (300 x 300 DPI)
https://mc06.manuscriptcentral.com/cjes-pubs Page 41 of 43 Canadian Journal of Earth Sciences
Draft
Specimens referenced in text.
213x252mm (300 x 300 DPI)
https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 42 of 43
Draft
Bone sections and teeth from Exctinction Cave
214x279mm (300 x 300 DPI)
https://mc06.manuscriptcentral.com/cjes-pubs Page 43 of 43 Canadian Journal of Earth Sciences
Draft
Specimens referenced in text
https://mc06.manuscriptcentral.com/cjes-pubs