Submitted : May 2 0th , 2020 – Accepted : July 30 th , 2020 – Posted online : August 3 rd , 2020
To link and cite this article:
doi: 10.5710/AMGH.30.07.2020.3368
1 Paleontological Notes
2 ‘A VERY LAZY DEER’: REVISION OF THE COTYPE OF NOTHROPUS
3 CARCARANENSIS (MAMMALIA, XENARTHRA)
4 RAÚL IGNACIO VEZZOSI 1,2,3, ALLINE ROTTI4,5 and LEONARDO DOS SANTOS
5 AVILLA4,5
6
7 1Laboratorio de Paleontología de Vertebrados, Centro de Investigación Científica y de
8 Transferencia Tecnológica a la Producción (CONICET, Gob. E.R., UADER), España
9 149, Diamante E3105BWA, Argentina. E-mail: [email protected]
10 2 Facultad de Ciencia y Tecnología, Universidad Autónoma de Entre Ríos, Ruta
11 Provincial 11 km 10,5, Oro Verde E3100XAD, Entre Ríos, Argentina.
12 3Programa de Desarrollo de las Ciencias Básicas (PEDECIBA – UDELAR) and Museo
13 Nacional de Historia Natural de Montevideo, Uruguay.
14 4Laboratório de Mastozoologia, Instituto de Biociências, Universidade Federal do
15 Estado do Rio de Janeiro. Avenida Pasteur, 458, sala 501, Urca. 22290-240, Rio de
16 Janeiro, RJ, Brasil. E-mails: [email protected], [email protected]
17 5Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva (PPGBBE),
18 Universidade Federal do Rio de Janeiro, Centro de Ciências da Saúde, Prédio das Pós-
19 graduações do Instituto de Biologia, Interbloco B/C, Cidade Universitária. 21941-902,
20 Rio de Janeiro, RJ, Brasil.
21
22
23 Total number of pages (14), figures (3)
24 Proposed header. VEZZOSI ET AL.: REVISION OF NOTHROPUS CARCARANENSIS
25 BORDAS, 1942.
1 26 Corresponding author: [email protected]
27 Short Description. The cotype of the sloth Nothropus carcaranensis is reviewed and
28 systematically assigned to the extinct Cervidae Morenelaphus.
29
30 Key words. Anatomy. Systematic. Cervidae. Morenelaphus. Braincase
31 Palabras clave. Anatomía. Sistemática. Cervidae. Morenelaphus. Caja craneana.
2 32 THE HOLOTYPE of the nothrotheriid Nothropus carcaranensis Bordas, 1942 is a partial
33 dentary from the Pleistocene of Santa Fe Province (Argentina) associated to other
34 isolated bones (see Bordas, 1942; Brandoni & McDonald, 2015). Among these remains,
35 an isolated braincase had been considered the cotype of N. carcaranensis in the original
36 publication of Bordas (1942:174–175).
37 This contribution provide for the first time detailed anatomical description and
38 analysis of this braincase assigned to N. carcaranensis, and to discuss its original
39 assignment.
40 Anatomical abbreviations. Acr, articular crest; af, articular fossa; ali,
41 alisphenoid; atu, articular tuberosity; bs, basisphenoid; bo, basioccipital; bu, bumps;
42 eam, external acoustic meatus; eop, external occipital process; fm, foramen magnum;
43 fo, foramen ovale; for, foramen orbitorotundum; fr, frontal; hf, hypoglossal foramen;
44 jf, jugular foramen; lf, lacerum foramen; mp, mastoid process; ms, mesiodistal sulcus;
45 mt, meatus temporalis; oc, occipital condyle; orb, orbital; p, pedicle; pal, palatine; pm,
46 procesus muscularis; par, parietal; ppa, paroccipital process; pop, postglenoid process;
47 pos, parieto-occipital suture; ptub, posterior tuberosity; pte, pterygoid; sc, sagittal
48 crest; sf, stylohyoid fossa; so, supraoccipital; sq, squamosal; tc, temporal crest; th,
49 tympanohyal recess; tm, tuberculum musculare; tyb, tympanic bulla; zp, zygomatic
50 process of squamosal.
51 MATERIALS AND METHODS
52 A direct comparison of the cotype of Nothropus carcaranensis with the skulls of
53 nothrotheres (e.g., Mionothorpus cartellei De Iuliis, Gaudin, Vicars [2011],
54 Nothrotherium maquinense [Lund, 1839], Pronothrotherium mirabilis [Kraglievich,
55 1925], Thalassocnus spp. McDonald & Muizon, 2012) and extant and extinct deer (see
56 Appendix 1) was made. The cranial nomenclatural of mammals and deer follows
3 57 Cabrera (1941), NAV (2005), and Heckeberg (2020). Comparisons to some taxa, such
58 as Megaloceros giganteus Blumenbach, 1799, Navahocerus fricki (Schultz & Howard,
59 1935) were based on Webb (1992) and Vislobokoba (2013). A list of the analyzed
60 specimens and their institutional collections are present in Appendix 1 (Supplementary
61 Online Information).
62 GEOGRAPHIC AND STRATIGRAPHIC PROVENANCE OF THE FOSSIL
63 DEER
64 The cotype of Nothropus carcaranensis (MACN-Pv 12630, Fig. 1) was
65 recovered by Osvaldo Coronel between 1930 to 1940 from the Carcarañá beds at the
66 National Road number 11, near Villa La Rivera, Santa Fe Province, Argentina
67 (32°38'23.03''S; 60°48'54.16''W). The fossiliferous strata can be correlated to the Upper
68 Pleistocene sequence (Cohen et al., 2013; ICC, 2018), which is part of the Timbúes
69 Formation (Iriondo & Kröhling, 2009; Vezzosi et al., 2019a, 2019b).
70 [Figure 1 insert here]
71
72 SYSTEMATIC PALEONTOLOGY
73
74 Order CETARTIODACTYLA Montgelard, Catzeflis & Douzery, 1997
75 Suborder RUMINANTIA Scopoli, 1777
76 Infraorder PECORA Linnaeus, 1758
77 Family CERVIDAE Goldfuss, 1820:374
78 Genus Morenelaphus Carette, 1922
79 Type species. Morenelaphus brachyceros (Gervais & Ameghino, 1880)
80 Morenelaphus sp.
81 Figures 1–3
4 82
83 Referred material. MACN-Pv 12630 posterior portion of the braincase of a young-adult
84 deer (Fig. 1), originally considered as cotype of Nothropus carcaranensis by Bordas
85 (1942:174–175).
86 Locality and age. Late Pleistocene levels of La Rivera town (Carcarañá River between
87 Timbúes and Oliveros, 32°38'23.03"S – 60°48'54.16"W), Santa Fe Province, Argentina.
88
89 Description and comparisons. MACN-Pv 12630 does not resemble another species of
90 Nothrotheriidae, and its general morphology is clearly different of any sloth braincases
91 (Fig. 1–3). In fact, it lacks features clearly recognized in best-known nothrotheriid
92 species like Mionothorpus cartellei, Nothrotherium maquinense, Pronothrotherium
93 mirabilis, even in Thalassocnus spp., including: circular and flat ectotympanic bone
94 (Fig. 2.3–2.4); flat basioccipital-basiesphenoid bones without posterior tuberosities (Fig.
95 2.5–2.6); absence of tympanic bullae and deep tympanohyal recess (Fig. 2.3–2.6); large
96 sagittal crest (Fig. 2.8); and large occipital condyles, very conspicuous and with oval
97 section; (Fig. 2.9); amongst others (Fig. 2). Moreover, the braincase of MACN-Pv
98 12630 is very similar to a young-adult deer in several aspects (Fig. 1), which include a
99 weak sagittal crest and basioccipital-basisphenoid bones fused (Fig. 3.3). The nuchal
100 plane of the occipital is composed by fully fused bones, but keeping a parieto-occipital
101 suture (Fig. 3.4). This suture is positioned dorsally near the parietal bone and disto-
102 laterally from the dorso-posterior area of squamosal. The mastoid process extends
103 ventro-laterally to the posterior edge of squamosal. Posteriorly, some specimens of
104 Morenelaphus (e.g., MFA-G-Pv 1311) lack a lateral constriction near the mastoid
105 process and the nuchal area of the occipital has a subcircular shape (Fig. 3.5). The
106 mastoid in Morenelaphus is a vertical and wide plate, which is compressed between the
5 107 posterior surface of the squamosal and the anterolateral edge of the infraoccipital; the
108 entire occipital area is subcircular in a posterior view (Fig. 3.6). Such as in Blastocerus
109 dichotomus males (Fig. 3.1), the posterior edge of the braincase, in lateral view, is
110 extremely concave and both the supraoccipital region and the occipital condyles are
111 projected much more posteriorly than other extant Cervidae (Fig. 3.3–3.4).
112 Nevertheless, senile individuals of Odocoileus virginianus have a very prominent edge
113 along the lambdoid suture (Cabrera, 1941), but never at the level recognized for B.
114 dichotomus and MACN-Pv 12630 (Fig. 3). Although both paroccipital processes are
115 slightly fragmented, they are ventrally evident such as in Hippocamelus spp. Leuckart,
116 1816; B. dichotomus; Ozotocerus bezoarticus (Linnaeus, 1758); Mazama spp.
117 Rafinesque, 1817; Cervus elaphus Linnaeus, 1758 and Rangifer tarandus Linnaeus,
118 1758 (Fig. 3). The occipital condyles are comparatively large and well-developed as in
119 Navahocerus fricki (Webb, 1992), much more than in all extant South American
120 Cervidae (SAC). MACN–Pv 12630 and Morenelaphus have similar large size and
121 reniform occipital condyles (Fig. 2.10–3). Adult individuals of Morenelaphus that bear
122 large antlers (MFA-G-Pv 1312, MFA-G-Pv 1722; Fig. 3.5) have one or two
123 conspicuous bumps dorsally to the foramen magnum. MACN–Pv 12630 lacks this
124 feature (Fig. 3.6), probably because it is a young-adult. Both petrosals in MACN–Pv
125 12630 bear a similar morphology to Morenelaphus (see Orcesi et al., 2019), in which
126 the promontorium is short and ovoid, the epitympanic wings are short and have a flat
127 surface and the tegmen tympani is concave, with an oval transverse section and with less
128 conspicuous anterior process (see Vezzosi, 2015, fig. II.102). In contrast with the
129 Nothrotheriidae, the basioccipital-basisphenoid have a triangular section in ventral
130 view, with two posterior tuberosities, which are distally project to both sides of
131 paroccipital processes (Fig. 3.8). These tuberosities exhibit a similar development in B.
6 132 dichotomus (MACN-Ma 13-1), O. bezoarticus (MACN-Ma 49183), and O. virginianus
133 (AMNH 100289); while in Hippocamelus spp., Mazama spp., and Pudu puda Molina,
134 1782, these are proportionally less developed. As in Morenelaphus, the mesiodistal
135 sulcus continues in a convex crest, limited by two conspicuous tuberculum musculare
136 with variable size (Fig. 3.7–3.8). Although MACN–Pv 12630 has the basisphenoid
137 preserved only until the level of its convex crest, it stills very similar to the one in
138 Morenelaphus (Fig. 3.7); showing a basicranium with small jugular foramen and
139 associated to the lacerum foramen, circular foramen orbitorotundum and similar in size
140 to the foramen ovale (Fig. 3.8). The concave articular fossa and the postglenoid process
141 are transversely positioned, but this latter is crosswise to foramen ovale. Behind this
142 foramen, the tympanic bulla is less inflated compared to Mazama spp., O. virginianus,
143 P. puda, and R. tarandus, and almost flattened such as in B. dichotomus, C. elaphus, O.
144 bezoarticus, P. mephistophiles (de Winton, 1896) and Hippocamelus spp. The
145 depression to tympanohyal recess is very deep in Morenelaphus (MNHNM 689), a
146 well-defined cavity, while in MACN-Pv 12630, which have only preserved the
147 tympanohyal, this is circular and lesser deep (Fig. 3.8).
148 [Figure 2 insert here]
149 DISCUSSION
150 Antlers are the most SAC common fossil, but skulls are extremely rare.
151 Therefore, the taxonomy of extinct SAC is traditionally based on antler morphology,
152 and not on braincases or postcranial remains (Cabrera, 1929; Kraglievich, 1932;
153 Churcher, 1966; Alcaraz, 2010; Rotti et al., 2018). However, anatomical characters
154 present in the braincase are very useful in mammals to evaluate some taxonomic issues.
155 Several fossils collected by Osvaldo Coronel during 1930 and 1940 come from
156 Carcarañá river at the Northern Pampa region and are housed at MACN and were
7 157 identified as Nothrotherinae indet., Nothropus carcaranensis, and Nothrotheriops sp.
158 (Brandoni & McDonald, 2015; Brandoni & Vezzosi, 2019). Among these remains,
159 Bordas (1942) recognizes the basicranium study here as part of the syntype (MACN-Pv
160 12630, cotype) of N. carcaranensis, and is pointed out substantial differences with
161 Nothrotherium maquinense (e.g., sharp external occipital process, slightly evident
162 paroccipital processes, acoustic meatus near the paroccipital process and posterior to the
163 postglenoid process). These morphological differences are present in deer (Cabrera,
164 1941; Vezzosi, 2015), while others (e.g., large intercondylar area, tympanohyal in a
165 large and concave surface, conspicuous entotympanic and superficial petrosal bones)
166 are clearly recognized in Nothrotheriinae and Thalassocninae sloths (De Iuliis et al.,
167 2011; McDonald & Muizon, 2012).
168 Furthermore, the morphological comparisons between MACN-Pv 12630 and
169 extant and extinct American deers allowed to recognize that the braincase previous
170 attributed to a Nothrotheriidae shares several characters with Morenelaphus (e.g., large
171 and posteriorly projected reniform condyles and tuberculae musculari).
172 Additionally, Morenelaphus have several other features especially in the
173 basicranium that are distinct from all other deer (i.e., conspicuous bumps, small jugular
174 foramen and associated to the lacerum foramen, circular orbitorotundum foramen with
175 similar size to foramen ovale). Also, considering the petrosal bones described by Orcesi
176 et al. (2019), it is possible to observe a short and ovoid promontorium; short and flat
177 epitympanic wing; tegmen tympani concave transversely convex, with a less
178 conspicuous anterior process. Thus, this unique combination of features in the braincase
179 confirms without any doubt the assignment of MACN-Pv 12630 to a young-adult
180 Morenelaphus Cervidae and not a sloth.
181 [Figure 3 insert here]
8 182 ACKNOWLEDGMENTS
183 We thank to the curators of CFA, CML, MACN, MDPA, MFA, MLP, MMP,
184 MNHN, MUFYCA; for facilitating access to collections of their institutions. Special
185 thanks to Guillermo Cassini (MACN) for facilitating the use of photos and information
186 of living deer from the collection of the AMNH, and to Andrés Rinderknecht (MNHN)
187 and Diego Brandoni (CICYTTP) for improving a previous version and providing useful
188 comments. We thank the reviewers François Pujos (IANIGLA) and H. Gregory
189 McDonald (BLM, USA) for providing useful reviews. This work was funded by PICT-
190 ANPCyT 2017-0954 (Argentina) and PEDECIBA-UDELAR (Uruguay).
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295 Figure captions
296 Figure 1. Braincase of MACN-Pv 12630 assigned to Morenelaphus in; 1, Left lateral;
297 2, Posterior; 3, Ventral; 4, Dorsal views. Scale 5 cm.
298 Figure 2. 1, Skull anatomy of cervids sensu Heckeberg (2020); 2, MACN-Pv 12360 in
299 lateral view; 3, Draw of the holotype of Mionothropus cartellei (modified from
300 De Iuliis et al., 2011); 4, Left lateral view of the braincase of Pronothrotherium
301 mirabilis (FC-DPV 271); 5, Draw of the basicranium of Mionothorpus cartellei
13 302 in ventral view (modified from De Iuliis et al., 2011); 6, Brasicranium of P.
303 mirabilis (FC-DPV 271) in ventral view; 7, Basicranium of MACN-Pv 12360 in
304 ventral view; 8-9, Dorsal and posterior views of P. mirabilis (FC-DPV 271); 10-
305 11, Posterior and dorsal views of MACN-Pv 12360. For anatomical
306 abbreviations, see material and methods. Scale 1 cm.
307 Figure 3. Braincases of living and fossil SAC. 1, Male of Blastocerus dichotomus
308 (MLP 22-II-99-1) in left lateral view; 2, Female of B. dichotomus (MLP 9-XI-
309 01-1) in left lateral view; 3, Male of Morenelaphus in left lateral view; 4,
310 Young-adult specimen of Morenelaphus (MACN-Pv 12630) in left lateral view;
311 5, Morenelaphus (MFA-G-Pv 1312) in posterior view; 6, Morenelaphus
312 (MACN-Pv 12630) in posterior view; 7, Morenelaphus (MFA-G-Pv 1312) in
313 ventral view; 8, Morenelaphus (MACN-Pv 12630) in ventral view. For
314 anatomical abbreviations, see material and methods. Scale 5 cm.
14
Paleontological Notes
‘A VERY LAZY DEER’: REVISION OF THE COTYPE OF NOTHROPUS
CARCARANENSIS (MAMMALIA, XENARTHRA)
RAÚL IGNACIO VEZZOSI 1,2,3, ALLINE ROTTI4,5 and LEONARDO DOS SANTOS
AVILLA4,5
1Laboratorio de Paleontología de Vertebrados, Centro de Investigación Científica y de Transferencia
Tecnológica a la Producción (CONICET, Gob. E.R., UADER), España 149, Diamante E3105BWA,
Argentina. E-mail: [email protected]
2 Facultad de Ciencia y Tecnología, Universidad Autónoma de Entre Ríos, Ruta Provincial 11 km 10,5,
Oro Verde E3100XAD, Entre Ríos, Argentina.
3Programa de Desarrollo de las Ciencias Básicas (PEDECIBA – UDELAR) and Museo Nacional de
Historia Natural de Montevideo, Uruguay.
4Laboratório de Mastozoologia, Instituto de Biociências, Universidade Federal do Estado do Rio de
Janeiro. Avenida Pasteur, 458, sala 501, Urca. 22290-240, Rio de Janeiro, RJ, Brasil. E-mails: [email protected], [email protected]
5Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva (PPGBBE), Universidade Federal do Rio de Janeiro, Centro de Ciências da Saúde, Prédio das Pós-graduações do Instituto de Biologia,
Interbloco B/C, Cidade Universitária. 21941-902, Rio de Janeiro, RJ, Brasil.
SUPPLEMENTARY ONLINE INFORMATION
Institutional abbreviations. AMNH, American Museum of Natural History, New
York, EEUU; CFA-Ma, Colección de Mastozoología, Fundación de Historia Natural
‘Félix de Azara’, Buenos Aires, Argentina; CML, Colección Mamíferos Lillo,
Universidad Nacional de Tucumán-Fundación Miguel Lillo, Tucumán, Argentina;
LACM, Natural History Museum of Los Angeles County, Los Angeles, EEUU; MACN-A, Colección Nacional Ameghino, Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia”, Buenos Aires, Argentina; MACN-Ma, Sección
Mastozoología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”,
Buenos Aires, Argentina; MACN-Pv, Sección Paleontología de Vertebrados, Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina;
MDPA, Museo Municipal Darwin de Punta Alta, Bahía Blanca, Argentina; MFA-G-
Pv, Área Paleontología de Vertebrados, Museo Provincial de Ciencias Naturales
“Florentino Ameghino”, Santa Fe, Argentina; MFA-Zv, Área Zoología de Vertebrados,
Museo Provincial de Ciencias Naturales “Florentino Ameghino”, Santa Fe, Argentina;
MLP, Museo de La Plata, Buenos Aires, Argentina; MMP, Museo Municipal de Mar del Plata “Lorenzo Scaglia”, Buenos Aires, Argentina; MNHNM, Museo Nacional de
Historia Natural de Montevideo, Uruguay; MNHN SAS, Muséum National d'Historie
Naturelle, Paris, France; MUFYCA, Museo Universitario Florentino y Carlos
Ameghino, Rosario, Santa Fe, Argentina.
Appendix 1. Taxonomic list of specimen of Morenelaphus and living deer used.
Morenelaphus, MACN-Pv 11408, MACN-Pv 11479, MACN-Pv 11564, MACN-Pv
11722, MACN-Pv 11730, MACN-Pv 12082, MACN-Pv 15697, MACN-Pv 16578,
MACN-Pv 16580; MLP 9-10, MLP 51-IV-11-69, MLP 63-V-31-1; MPA-85-189-I-D,
MFA-G-Pv 1311, MFA-G-Pv 1312, MFA-G-Pv 1681, MFA-G-Pv 1722, MFA-G-Pv
1723, MFA-G-Pv 1724, MFA-G-Pv 1725, MFA-G-Pv 1726; MUFYCA 1027,
MUFYCA 1432, MNHN 689. Hippocamelus sulcatus, MLP 9-11 (type of Furcifer seleniticus), MLP 9-23 (plastotipo de Furcifer sulcatus). Morenelaphus brachyceros,
MLP 9-15 (type of Morenelaphus lydekkeri); MLP 9-21 (type of Morenelaphus rothi);
MLPM 249 (calcotype of Cervus brachyceros), MLPM–250 (calcotype of Cervus palaeoplatensis), MLP 91-1. Morenelaphus lujanensis, MLP 9-10 (type of M. pseudoplatensis); MLP 9-17 (type of Blastocerus azpeitianus); MLP 9-26 (type of
Pampaeocervus platensis), MLP 63-V-31-1. Blastocerus dichotomus, CFA–Ma 6805,
CML 3738, MACN–ZV 13.1, MACN–Zv 13.2, MACN–Ma 45-8, MACN–Ma 5172,
MACN–Ma 13437, MACN–Ma 16269, MACN–Ma 21645, MACN–Ma 21646
MACN–Ma 21647, MACN–Ma 26601, MACN–Ma 31231, MACN–Ma 42.320, MFA–
Zv 1264, MFA–Zv 179, MFA–Zv 180, MFA–Zv 181, MFA–Zv 1263, MLP 763, MLP
1523, MLP 1524, MLP 1687, MLP 1724, MLP 9-XI-01-2, MLP 9-XI-01-3, MLP 9-XI-
01-4, MLP 11.X.46.8, MLP 11.X.46.14, MLP 11.X.46.15, MLP 11.X.46.17, MLP
12.XI.01.1, MLP 12.XI.01.3, MLP 12-XI-01-7, MLP 12-XI-01-9, MLP 12-XI-01-13,
MLP 12-XI-10-14, MLP 22.II.99.I. Axis axis, MACN–Ma 44.2, MACN–Ma 47216.
Cervus elaphus, MACN–Ma 4338, MACN–Ma s/n. Dama dama, MACN–Ma 4397,
MACN–Ma 25959. Hippocamelus antisensis, AMNH 217149, CML 3737, CML 3740,
CM; 3742, CML 3745, CML 3777, CML 3779, MCN-UNSa 003583, MACN–Ma
5362, MCN-UNSa 01, MCN-UNSa 03, MCNUNSa 01/m, MCN-UNSa 02/m, MCN-
UNSa 03/m, MCN-UNSa 04, MCN-UNSa 05, MACN–Ma 53.60, MACN–Ma 53.62,
SGO.Ma 567, SGO.Ma 1023. Hippocamelus bisulcus, CML 3276, CML 3739, CML
3743, CML 3744, CML 3776, MACN–Ma 2.16, MACN–Ma 4.30, MACN–Ma 11.8,
MACN–Ma 44.28, MACN–Ma 15665 MACN–Ma 38261, MLP 397, MLP 423, MLP
516, MLP 1346, MLP 1350, MLP 1352, MLP 1353, MLP 1354, MLP 1359, MLP 1360,
MLP 1361, MLP 1362, MLP 1363, MLP 1364, MLP 1522, MLP 10-VIII-00-30, MLP
19-X-00-11, SGO.Ma 1024. Mazama americana, CFA-Ma 03911, CFA-Ma 03983.
Mazama gouazoupira, CFA-Ma 04369, CFA-Ma 06345, CFA-Ma 10793, CFA-Ma
11044, MACN–Ma 3632, MFA–Zv 40, MFA–Zv 106, MFA–Zv 125, MFA–Zv 126,
MFA–Zv 127, MFA–Zv 128, MFA–Zv 156, MFA–Zv 157, MFA–Zv 194, MFA–Zv 937, MFA–Zv 753, MFA–Zv 760, MFA–Zv 782, MFA–Zv 783, MFA–Zv 784, MFA–
Zv 785, MLP 713, MLP 1-IX-00-4, MLP18-X-02-5. Mazama nana, CFA-Ma 05719,
CFA-Ma 0263, MACN–Ma 24688, MACN–Ma 24691. Mazama sp., MLP 735, MLP
736, MLP 1343, MLP 1344, MLP 1525. Moschus moschiferus, MACN–Ma 35211.
Odocoileus virginianus, AMNH 100289. Ozotoceros bezoarticus, MACN–Ma 4.14,
MACN–Ma 18.14, MACN–Ma 24507, MACN–Ma 24523, MACN–Ma 24749,
MACN–Ma 4297, MACN–Ma 31231, MACN–Ma 49182, MACN–Ma 49183, MACN–
Ma 49240, MFA–Zv 147, MFA–Zv 438, MFA–Zv 870, MFA–Zv 880, MFA–Zv 881,
MFA–Zv 882, MFA–Zv 883, MFA–Zv 915, MFA–Zv 1000, MFA–Zv 1182, MFA–Zv
1183, MFA–Zv 1184, MFA–Zv 1185, MFA–Zv 1186, MFA–Zv 1187, MFA–Zv 1188,
MFA–Zv 1189, MFA–Zv 1190, MFA–Zv 1191, MFA–Zv 1192, MFA–Zv 1195, MFA–
Zv 1196, MFA–Zv 1197, MFA–ZV 1198, MLP 692, MLP 1337, MLP 1340, MLP
1341, MLP 1-XII-00-2, MLP 5-VI-97-3, MLP 6-VIII-98-1, MLP 14-VI-00-2, MLP 18-
VIII-92-1, MLP 18-VIII-92-2, MLP 18.VIII.92.7, MLP 18.VIII.92.9, MLP
18.VIII.92.15, MLP 4-14. Pudu mephistophiles, AMNH 181505, AMNH 181506,
AMNH 181504. Pudu pudu, MACN–Ma 17809, MACN–Ma 22368, MACN–Ma
47219, SGO.Ma 480, SGO.MA 559, AMNH 93321. Rangifer tarandus, MACN–Ma
6.3, MACN–Ma 8.24, MACN–Ma 4331. Taxonomic list of nothrotheres used.
Mionothropus cartellei, LACM 4609 ⁄ 117533; Nothrotheriops shastensis, LACM 208,
166, 632,633; Nothrotherium, MUN STRI 12924; Pronothrotherium mirabilis, FC-
DPV 271; Thalassocnus natans, MNHN SAS 734; T. littoralis, MNHN SAS 1615.