Content uploaded by Patricio López Mendoza
Author content
All content in this area was uploaded by Patricio López Mendoza on May 10, 2019
Content may be subject to copyright.
Estudios Geológicos
julio-diciembre 2015, 71(2), e041
ISSN-L: 0367-0449
doi: http://dx.doi.org/10.3989/egeol.42011.357
Presence of Arctotherium (Carnivora, Ursidae, Tremarctinae)
in a pre-cultural level of Baño Nuevo-1 cave
(Central Patagonia, Chile)
Presencia de Arctotherium (Carnivora, Ursidae, Tremarctinae)
en un nivel pre-cultural de cueva Baño Nuevo-1 (Patagonia
Central, Chile)
P. López Mendoza1, F. Mena Larraín2, E. Bostelmann3
1 Departamento de Antropología, Facultad de Ciencias Sociales, Universidad de Chile and ARQMAR-Centre for
Maritime Archaeology Research of the Southeastern Pacic, Ignacio Carrera Pinto N° 1045, Ñuñoa, Santiago, Chile.
Email: patriciolopezmend@gmail.com
2 Centro de Investigación en Ecosistemas de la Patagonia (CIEP; CONICYT-Regional R10C 1003), Simpson 471, Coyhaique,
Chile. Email: francisco.mena@ciep.cl
3 Instituto de Ciencias de la Tierra, Facultad de Ciencias, Universidad Austral de Chile. Isla Teja s/n, Valdivia, Chile.
E-mail: ebostel@yahoo.com
ABSTRACT
The description of an I3 assigned to Arctotherium sp. obtained from the Baño Nuevo-1 site (Central Patagonia,
Chile) is presented. The finding was recovered from Layer 5 and it is associated to Macrauchenia sp., Lama guani-
coe, Felidae, Camelidae, Equidae and Mylodontidae, within a sterile deposit of cultural material, dated between ca.
13.500 and 11.200 BP. Despite the fact that it is only a single specimen, such finding extends the known distribution
for the genus in Chile.
Keywords: Ursidae; Arctotherium; Late Pleistocene; Central Patagonia.
RESUMEN
Se presenta la descripción de un I3 asignado a Arctotherium sp. proveniente del sitio Baño Nuevo-1 (Patagonia
Central, Chile). El hallazgo fue realizado en la Capa 5 y está asociado a restos de Macrauchenia sp., Lama guani-
coe, Felidae, Camelidae, Equidae y Mylodontidae dentro de un depósito estéril de material cultural, datado entre
los ca. 13.500 y 11.200 años AP. Aunque se trata de un único espécimen, amplía el rango de distribución conocido
para este género en Chile.
Palabras clave: Ursidae; Arctotherium; Pleistoceno final; Patagonia Central.
Recibido el 7 de enero de 2015 / Aceptado el 23 de septiembre de 2015 / Publicado online el 20 de noviembre de 2015
Citation / Cómo citar este artículo: P. López Mendoza, et al. (2015). Presence of Arctotherium (Carnivora, Ursidae, Tremarctinae)
in a pre-cultural level of Baño Nuevo-1 cave (Central Patagonia, Chile). Estudios Geológicos 71(2): e041. http://dx.doi.org/10.3989/
egeol.42011.357.
Copyright: © 2015 CSIC. This is an open-access article distributed under the terms of the Creative Commons Attribution-Non
Commercial (by-nc) Spain 3.0 License.
2 P. López Mendoza et al.
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
Introduction
Recent evaluations of the Baño Nuevo-1 collections
have led to the identification of an Arctotherium sp.
(Carnivora, Ursidae, Tremarctinae) tooth fragment
from the upper sections of Layer 5 (Late Pleistocene),
associated to Macrauchenia sp., Equidae, Felidae,
Lama guanicoe, Camelidae and Mylodontidae re -
mains (mainly dermal bones). On the immediately
overlaying Layer 4B, Diabolotherium cf. nordenski-
oldi, Lama guanicoe, Macrauchenia sp., Equidae,
Camelidae, Felidae and Mylodontidae remains have
also been recorded (López, 2009; Bostelmann, et al.,
2011; López & Mena, 2011).
This sample (one specimen) opens new issues for
the osseous record of the final Pleistocene at the site,
as another possible taphonomic agent has to be con-
sidered, while discussing the faunal remains founded
on rockselters from this time period at the Chilean
central Patagonia in general. Beyond description and
taxonomic identification, it also opens new grounds
for paleoecological studies, such as those already
developed in southern Patagonia (Martín, 2008; 2013;
Martín & San Román, 2010).
The finding of an Arctotherium remain at Baño
Nuevo-1 has been barely mentioned in previous
studies (see López & Mena, 2011). Given its impor-
tance, however, this brief note aims to provide valu-
able information on its stratigraphical setting and
chronological context and a detailed description as
well as a preliminary discussion on the origins of the
fossil.
The nding in context
Baño Nuevo-1 is located ca. 80 km, NE of
Coyhaique (45° 17 S-71° 32′W) (Fig. 1). The cave
is located on an Aptian volcanic complex, locally
known as “Cerro Grande del Campo Seis” (Large
Hill on Section 6). With a depth of 20 m and an aver-
age width of 4 m, the cave nowadays has a relatively
restricted access due to the entry of coluvial debris
fallen through a lateral shaft (Velásquez & Mena,
2006). The stratigraphic record has been previously
presented in several articles (see Mena et al., 2000,
2003; Núñez et al., 2005; Mena & Stafford, 2006;
Velásquez & Mena, 2006; Mena, 2009; López &
Mena, 2011; Trejo & Jackson 1998).
The tooth specimen presented, was recovered
from Layer 5, characterized by clay and organic
sand, dated between ca. 13.500 and 11.200 cal 14C
B.P. (see Fig. 2). Just like Layer 4A, this layer pres-
ents abundant fossil remains and organic sediments
that could be assigned to Mylodontidae dung (Mena
& Stafford, 2006; Núñez et al., 2005). This layer
overlies Layer 6, composed by fine microlaminated
sands deposited by a large Pleistocene proglacial
lake devoid of any faunal remains (Mena & Stafford,
2006; Núñez et al., 2005).
Material and methods
The material corresponds to a left I3. Currently,
it is stored at the Facultad de Ciencias Sociales,
Universidad de Chile, under the code Baño Nuevo-1/
Unidad 5C/Capa 5/N°F171. The specimen was com-
pared with collections of Arctotherium vetustum at the
Museo Argentino de Ciencias Naturales Bernardino
Rivadavia, Lycalopex culpaeus, Otaria sp. and Puma
concolor from the Laboratorio de Zooarqueología of
the Universidad de Chile, and Panthera onca from a
private collection. Since part of the cusp is broken,
mesio-distal and lingual-labial measurements had
to be taken only at the crown base. The occlusal
surface was observed through Scanning Electron
Microscopy (SEM) (Zeiss, model EVO MA10) at the
Departamento de Antropología of the Universidad
de Chile.
Figure 1.—A. Location of the Baño Nuevo-1 site; B. Plan drawing
showing recovery location of the Arctotherium sp. sample.
Presence of Arctotherium in a pre-cultural level of Baño Nuevo-1 cave 3
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
Institutional abbreviations: MACN, Museo Argen-
tino de Ciencias Naturales Bernardino Rivadavia.
Systematic Paleontology
Carnivora Bowdich, 1821
Ursidae Gray, 1825
Tremarctinae Merriam and Stock, 1925
Arctotherium Burmeister, 1879
Arctotherium sp.
Material: Baño Nuevo-1/Unidad 5C/Capa 5/N°
F171, an isolated I3 (Fig. 3A, B, C and D).
Geographical occurrence: Baño Nuevo-1 cave,
Coyhaique, Central Patagonia, Chile. UTM: 0301821
E-4981638 N (Fig. 1A-B).
Stratigraphical occurrence: Layer 5, Late Pleisto-
cene dated between ca. 13.500 and 11.200 cal 14C
B.P. (Fig. 2).
Description and comparisons
The small and fragmentary nature of the sample
precludes a clear-cut, absolute identification, yet both
its size and shape allows us to assign it to Ursidae
and, moreover, to Arctotherium Burmeister, 1879.
The finding was done on unit 5C and it corresponds to
a left I3 from an adult. The incisor presents a fracture
on its oclussal surface allowing us to see the pulp cav-
ity of the tooth. This surface exhibits a subtriangular
shape, while on the lingual facies we can see an enamel
layer crossing the surface. The root also presents a
subtriangular section being compressed on its messio-
distal direction with two grooves reaching the dental
crown and parallel edges. At the basis of the crown in
the mesial level, the root rises to form an inverted V,
while on the distal surface we observe a well devel-
oped marginal lobe at the point where the crown
meets the root, that is fractured on its occlusal surface
(Fig. 3A, B, C and D).
The incisor from Baño Nuevo-1 was found to be
much larger than those from canids, felids and ota-
rids used as comparative references. It also pres-
ents a marked development of the marginal distal
lobe and grooves on the root. None of these traits
were observed on canids, felids or otarids. The
specimen also presents a thick enamel layer on its
labial face, being much thinner on the lingual one.
Morphologically the Baño Nuevo-1 incisor is similar
to Arctotherium vetustum (MACN 1201), although
larger, with 8.5 mm mesio-distal measurement in the
base and 12.6 mm labial-lingual measurement in the
base.
Biogeographic panorama
Nowadays the Tremarctinae Subfamily can be found
exclusively on the Americas (Soibelzon, 2004).
Four genera are recognized: Plionarctos Frick,
1926 and Arctodus Leidy, 1854 in North America,
Arctotherium Burmeister, 1879 in South America
Figure 2.—Stratigraphic cross section of the Baño Nuevo-1 site.
4 P. López Mendoza et al.
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
and Tremarctos Gervais, 1855 with an extinct species
from North America and a living one in South America
(Soibelzon, 2004).
According to the latest reviews (Soibelzon, 2004;
Soibelzon et al., 2005) the South American extinct
bears have been assigned to the genera Arctotherium,
being recorded in Venezuela, Brazil, Bolivia, Uruguay,
Argentina and Chile from the Ensenadan (Upper
Pliocene to Middle Pleistocene) to the Lujanian
(Upper Pleistocene-Early Holocene). Five species
are recognized within this genera: Arctotherium
vetustum Ameghino, 1885 from Bonaerian levels
(Middle Pleistocene) at the Argentine provinces of
Buenos Aires and Entre Ríos and one possible finding
Figure 3.—Views from the de Arctotherium sp. I3 from Baño Nuevo-1: A. Oclusal; B. Labial; C. Lateral; D. Lingual; E. Drawings
of I3 of Arctotherium sp. of the Baño Nuevo-1 site indicating the principal features mentioned in the text; F. Arctotherium vetustum
(MACN 1201) mandible with left I3 in situ. Abbreviations: ml. marginal lobe; pc. pulp cavity; rc. root canal.
Presence of Arctotherium in a pre-cultural level of Baño Nuevo-1 cave 5
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
from Brazil; Arctotherium wingei Ameghino, 1902
from Brazilian and Venezuelan Lujanian levels
and Pleistocene deposits in Bolivia; Arctotherium
bonariense (Gervais 1848–1852) from Bonaerian
and Lujanian levels of the Buenos Aires province
(Argentina); Arctotherium angustidens Gervais &
Ameghino, 1880 from Ensenadan levels at the
same province and at Tarija, Bolivia; and finally
Arctotherium tarijense Ameghino, 1902 from Bonae-
rian and Lujanian levels at this Bolivian locality,
Argentine provinces of Santa Fe and Buenos Aires,
Uruguay and at the XII Region of Magallanes, Chile
(Soibelzon, 2004).
In Chile, the record of Ursidae is restricted to cen-
tral and southern Patagonia. It is composed by a molar
from the Pilauco site (40° S) assigned to Ursidae
(Pino et al., 2013). Another finding corresponds to
a femur from Cueva del Milodón (51°S), originally
assigned to Arctotherium sp. and later on reassigned
to Pararctotherium pamparun by Oliver-Schneider
(1935). At Cueva de Los Chingues (52° S) an I2 was
recorded. Originally assigned to Pararctotherium sp.
by Prevosti et al. (2003), it was later identified as
Arctotherium tarijense by Soibelzon (2004) and has
recently been reassigned to Arctotherium sp. by Prevosti
& Martín (2013). Finally, there must be mentioned
another femur fragment collected from the surface of
Cueva del Puma (52° S) and identified as Arctotherium
tarijense with a date of 10.345±75 B.P. (Martín et al.,
2004, Martín, 2013). Thus, the evidence now reported
from Baño Nuevo-1 fills the intermediate gap between
the areas from where it was known before.
The Arctotherium sp. remain from Layer 5 at
Baño Nuevo-1 cave is associated to Mylodontinae,
Felidae (aff. Panthera onca mesembrina), Equidae,
Lama guanicoe and Macrauchenia sp. remains with
no traces of human presence (López & Mena, 2011).
Paleoenvironmental studies based on pollen records
from neighboring areas (High Cisnes River, De
Porras et al., 2012; High Simpson river, Markgraf
et al., 2007; Middle Chacabuco river, Villa-Martínez
et al., 2011) reveal an open steppe environment.
Discussion and conclusions
One of the questions triggered by the finding
of Arctotherium at Baño Nuevo-1 is related to the
taphonomy of the fossil assemblage at the cave.
Studies about the role of Arctotherium as a potential
agent on the formation and transformation of bone
assemblages are almost nonexistent, unlike the Old
World Ursidae situation, whose taphonomic effects
have been subject of intensive research (Gargett,
1996; DʼErrico et al., 1998; Stiner et al., 1998;
Wolverton, 2001; Quilès et al., 2006; Rabal-Garcés
& Cuenca-Bescós, 2009; Arilla et al., 2014). One
of the few American studies, specifically geared
to the case of Arctotherium angustidens, reveals a
high proportion of broken teeth, as likely result of
chewing on hard materials such as bones (Figueirido
& Soibelzon, 2010). On the other hand, studies by
Haynes (1983) have revealed that bears (Ursus arc-
tos and Ursus americanus in particular) leave traces
like those made by rodents, that is short and paral-
lel with punctures related to parallel grooves on the
crest of long bones. Andrews and Fernández-Jalvo
(1997) found punctures of up to 10.4 mm in diam-
eter, attributed to the action of bears. These marks
were recorded both on the shaft and articular sur-
faces, just like those findings done by DʼErrico and
collaborators (1998). The only puncture recorded on
remains from Layer 5 at Baño Nuevo-1 was found
on a Lama sp. patella and both its shape and size are
consistent with those produced by canids.
However, with this finding, Arctotherium joins the
Felidae (aff. Panthera onca mesembrina) and Dusicyon
avus previously known from the site as potential
agents in the formation of the bone assemblages from
the layers of the Final Pleistocene at the site (Trejo
& Jackson, 1998). As said before, the wear patterns
and dental pathologies observed in different samples
indicate an omnivorous diet with non- negligible meat
consumption, including bones (Prevosti & Vizcaíno,
2006; Figueirido & Soibelzon, 2010; Martín, 2013).
In fact, most of the known Arctotherium angustidens
remains have broken teeth’s, most likely due to chew-
ing on hard bones (Soibelzon et al., 2009; Figueirido
& Soibelzon, 2010).
In the occlusal surface of the I3 of Baño Nuevo-1, part
of the cusp is fragmented and it presents a caries that
could be interpreted as a result of eating carbohydrate-
rich foods, such as fruit or honey (Ferigolo, 1992;
Soibelzon & Prevosti, 2007; Soibelzon et al., 2014).
Though a microwear analysis is not applicable to
South American giant short-faced bear, because in spe-
cies with omnivore diet the interpretation of hard plant
6 P. López Mendoza et al.
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
versus bone consumption is problematic (Soibelzon
et al., 2014:1241), we performed a SEM observa-
tion to see microwear that could reveal chewing use
after cusp breakage. We looked the labial and lingual
faces of the occlusal surface, finding striations and
pits that point to a chewing action on the lingual face,
while no such traits were observed on the labial face
(Fig. 4A–B). Such evidences points to a postdeposi-
tional fracture of a portion of the cusp, or an immedi-
ate loss of it together with the complete incisive from
its alveolar cavity.
Given the lack of direct evidence of damage
due to bear action, the origin of Arctotherium at
Baño Nuevo-1 has to be questioned. Soibelzon
and collaborators (2009) point to two possible sce-
narios for the presence of bears in caves: the use
for hibernating, and the sporadic search of carrion
inside the caves. Hibernation has been mentioned
in other works (Martín, 2013:28), but it is not clear
if it can be applied to Tremarctinae. Accordingly,
the sporadic use of Baño Nuevo-1 cave for both
food and shelter purposes, is the most plausible
scenario, taking into account the number and sort
of evidence recovered, along with no presence of
taphonomic traces attributable to Arctotherium. In
any case, this very low density of Arctotherium
material is consistent with the record from other
sites in Patagonia, such as Milodón, Los Chinges
or El Puma caves, as it is the almost null evidence
of taphonomic traces left by this taxon (Martín,
2008:364).
The record of Arctotherium at Baño Nuevo-1
raises several questions that must be dealt with in the
future, such as the competition with other animals
to occupy the cave or its role in the formation and
transformation of the deposits.
ACKNOWLEDGMENTS
To Dr. Francisco Prevosti for his help identifying the sample,
and his valuable comments to an earlier version of the manuscript.
Thanks to the Museo Argentino de Ciencias Naturales Bernardino
Rivadavia for granting the access to their collections. We also
want to thank the anonymous reviewers and Elvira Latorre B. for
her drawings.
References
Andrews, P. & Fernández-Jalvo, Y. (1997). Surface modi-
fications of the Sima de los Huesos fossil humans.
Journal of Human Evolution, 33: 191–217. http://
dx.doi.org/10.1006/jhev.1997.0137
Ameghinio, C. (1885). Nuevos restos de mamíferos
fósiles oligocenos, recogidos por el profesor Pedro
Scalabrini y pertenecientes al Museo provincial de
la ciudad de Paraná. Boletín de la Academia de
Ciencias de Córdoba, 8: 3–207.
Ameghinio, C. (1902). Notas sobre algunos mamíferos
fósiles nuevos o poco conocidos del valle de Tarija.
Anales del Museo Nacional de Buenos Aires, 3:
225–261.
Arilla, M.; Rosell, J.; Blasco, R.; Domínguez-Rodrigo,
M.; Pickering, T. (2014). The “bear” essentials: actu-
alistic research on Ursus arctos arctos in the span-
ish pyrenees and its implications for Paleontology
and Archaeology. PLoS ONE, 9 (7): e102457. http://
dx.doi.org/10.1371/journal.pone.0102457
Bostelmann, E.; López, P.; Salas-Gismondi, R. & Mena,
F. (2011). First record of Diabolotherium cf. nor-
deskioldi, Kraglievich 1926, (Mammalia, Tardigrada,
Megalonychidae), from the Late Pleistocene of Chile.
Ameghiniana, 48 (4): 146.
Bowdich, T.E. (1821). An analysis of the natural classifica-
tons of Mammalia for the use of students and travellers.
J. Smith, Paris. 115 pp.
Burmeister, H. (1879). Description physique de la Répub-
lique Argentine dʼaprès des observations personnelles
et étrangéres. Traduit de lʼallemand avec le concourse
de E. Daireaux. Tome troisiém. Animaux vertébrés.
Premiére partie: Mammiféres vivants et éteints, 3 (1):
1–556 with Atlas.
DʼErrico, F.; Villa, P.; Pinto Llona, A. & Ruíz Idarraga,
R. (1998). A Middle Palaeolithic origin of music?
Using cave-bear bone accumulations to assess the
Divje Babe I bone ‘flute’. Antiquity, 72: 65–79.
http://dx.doi.org/10.1017/S0003598X00086282
De Porras, M.E.; Maldonado, A.; Abarzúa, A.M.;
Cárdenas, M.L.; Francois, J.P.; Martel-Cea, A.; Stern,
Ch.; Méndez, C. & Reyes, O. (2012). Postglacial veg-
etation, fire and climate dynamics at Central Chilean
Patagonia (Lake Shaman, 44° S). Quaternary Science
Reviews, 50: 71–85. http://dx.doi.org/10.1016/j.
quascirev.2012.06.015
Figure 4.—Occlusal view of the I3 of Baño Nuevo-1 showing:
A. SEM photography of the area of fracture; B. SEM photography
and schematic drawing indicating tracks of strias and pits product
of the use as masticatory surface.
Presence of Arctotherium in a pre-cultural level of Baño Nuevo-1 cave 7
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
Ferigolo, J. (1992). Non-human vertebrate paleopathology
of some Brazilian Pleistocene mammals. In: Paleopa-
tologia and Paleoepidemiologia-Estudos Multidisci-
plinares (Araujo, A. & Ferreira, L., Eds.), Panorama/
Escola Nacional de Saude Publica, Rio de Janeiro,
213–234.
Figueirido, B. & Soibelzon, L. (2010). Inferring palaeo-
ecology in extinct tremarctine bears (Carnivora,
Ursidae) using geometric morphometrics. Lethaia,
43 (2): 209–222. http://dx.doi.org/10.1111/j.1502-
3931.2009.00184.x
Frick, C. (1926). The Hemocyoninae and an American
Tertiary Bear. American Museum of Ntural History
Bulletin, 56: 1–110. http://hdl.handle.net/2246/1321
Gargett, R. (1996). Cave bears and modern human origins.
University Press of America, Lanham, Maryland.
288 pp.
Gervais, P. (1855). Recherches sur les mammiféres
fossiles de lʼAmérique du Sud. In: Expédition
dans les parties centrales de lʼAmérique du Sud:
De Rio de Janeiro á Lima, et de Lima au Para
(Castelnau, F., dir.). Vol. 7, Zoologuie. Bertrand,
Paris, 116 pp.
Gervais, P. (1848–1852). Zoologie et Paléontologie Fran-
caises (animaux vertébrés) ou Nouvelles Recherches
sur les Animaux Vivantes et Fossiles de la France.
A. Bertrand, Paris, 271 pp.
Gervais, H. & Ameghino, F. (1880). Los Mamíferos Fósiles
de la América Meridional. F. Savy, Paris, 225 pp.
Gray, J.E. (1825). Outline of an attempt at the disposition
of the Mammalia into tribes and families with a list of
genera apparently appertaining to each tribe. Annals
of Philosophy, 10: 337–344.
Haynes, G. (1983). A guide for differentiating Mamma-
lian carnivore taxa responsible for gnaw damage to
herbivore limb bones. Paleobiology, 9 (2): 164–172.
Leidy, J. (1854). Remarks on Sus americanus, or Harlanus
americanus and other extinct mammals. Proceedings
of the Academy of Natural Sciences of Philadelphia,
7: 89–90.
López, P. (2009). El mundo perdido de Patagonia Central:
una aproximación tafonómica al estudio de los mamífe-
ros extintos del sitio Baño Nuevo-1 (XI Región-Chile).
In: Zooarqueología y Tafonomía en el Confín del
Mundo (López, P.; Cartajena, I.; García, Ch. & Mena,
F., Eds.), Ediciones Universidad Internacional SEK-
Chile, Santiago, 181–198.
López, P. & Mena, F. (2011). Extinct ground sloth dermal
bones and their role in the taphonomic research of
caves: the case of Baño Nuevo-1 (Andean Central
Patagonia, Chile). Revista Mexicana de Ciencias
Geológicas, 28 (3): 519–532.
Markgraf, V.; Whitlock, C. & Haverle, S. (2007). Veg-
etation and fire history during the last 19,000 cal
yr B.P. in southern Patagonia: Mallín, Pollux, Coy-
haique, Province Aisén (45° 41′ 30″, 71° 50′ 30″ W,
640 m elevation). Paleogeography, Paleoclimatology,
Palaeoecology, 254: 492–507. http://dx.doi.org/10. 1016/
j.palaeo.2007.07.008
Martín, F. (2008). Bone Crunching felids at the end of the
Pleistocene in Fuego-Patagonia, Chile. Journal of
Taphonomy, 6 (3–4): 337–372.
Martín, F.M. & San Román, M. (2010). Explorando la
variabilidad del registro arqueológico y tafonómico
en Pali-Aike (Chile) a través de la búsqueda de
registros pleistocenos a cielo abierto. Magallania,
38 (1): 199–214. http://dx.doi.org/10.4067/S0718-
22442010000100012
Martín, F. (2013). Tafonomía y paleocología de la tran-
sición Pleistoceno-Holoceno en Fuego Patagonia.
Interacción entre humanos y carnívoros y su impor-
tancia como agentes en la formación del registro
fósil. Ediciones de la Universidad de Magallanes,
Punta Arenas. 406 pp.
Martín, F.; Prieto, A.; San Román, M.; Morello, F.;
Prevosti, F.; Cárdenas, P. & Borrero, L.A. (2004).
Late-Pleistocene megafauna at Cueva del Puma,
Pali-Aike Lava Field, Chile. Current Research in the
Pleistocene, 21: 101–103.
Mena, F. (2009). Aves en Cueva Baño Nuevo. In:
Zooarqueología y Tafonomía en el Confín del Mundo
(López, P.; Cartajena, I.; García, Ch. & Mena, F.,
Eds.), Ediciones Universidad Internacional SEK-
Chile, Santiago, 59–71.
Mena, F., & Stafford, T. (2006). Contexto estratigráfico y
fechación directa de esqueletos humanos del Holo-
ceno Temprano en Cueva Baño Nuevo (Patagonia
Centra, Chile). In: Segundo Simposio Internacional
el Hombre Temprano en América (Jiménez, J., Ed.),
Instituto Nacional de Antropología e Historia, México,
139–154.
Mena, F.; Lucero, V.; Reyes, O.; Trejo, V. & Velásquez, H.
(2000). Cazadores tempranos y tardíos en la Cueva
Baño Nuevo-1, margen occidental de la estepa cen-
tropatagónica (XI Región de Aysén; Chile). Anales
del Instituto de la Patagonia, 28: 173–195.
Mena, F.; Reyes, O.; Stafford, T. & Southon, J. (2003).
Early human remains from Baño Nuevo-1 cave,
Central Patagonian Andes, Chile. Quaternary Interna-
cional, 109–110: 113–121. http://dx.doi.org/10.1016/
S1040-6182(02)00207-0
Merriam, J. & Stock, C. (1925). Relationships and struc-
ture of the short-faced bear, Arctotherium, from the
Pleistocene of California. Carnegie Institution of
Washington Publication, 347: 1–35.
Núñez, H., Stafford, T. & Frassinetti, D. (2005). Primer
registro de fósiles de Liolaemus en Chile (Reptilia,
Sauria). Noticiario Mensual del Museo de Historia
Natural, 356: 3–7.
Oliver-Schneider, C. (1935). Mamíferos fósiles de Chile:
Adiciones y correcciones a la lista preliminar. Revista
Chilena de Historia Natural, 39: 297–304.
Pino, M., Chávez-Hoffmeister, M., Navarro-Harris, X. &
Labarca, R. (2013). The late Pleistocene Pilauco site,
8 P. López Mendoza et al.
Estudios Geológicos, 71(2), julio-diciembre 2015, e041, ISSN-L: 0367-0449. doi: http://dx.doi.org/10.3989/egeol.42011.357
Osorno, south-central Chile. Quaternary Interna-
tional, 299: 3–12. http://dx.doi.org/10.1016/j.quaint.
2012.05.001
Prevosti, F. & Vizcaíno, S. (2006). Paleoecology of the large
carnivore guild from the late Pleistocene of Argentina.
Acta Palaeontologica Polonica, 51 (3): 407–422.
Prevosti, F. & Martín, F. (2013). Paleoecology of the mam-
malian predator guild of Southern Patagonia during
the latest Pleistocene: Ecomorphology, stable iso-
topes, and Taphonomy. Quaternary International, 305:
74–84. http://dx.doi.org/10.1016/j.quaint.2012.12.039
Prevosti, F.; Soibelzon, L.; Prieto, A.; San Román, M. &
Morello, F. (2003). The southernmost bear Pararctoth-
erium (Carnivora: Ursidae: TreMartínae) in the latest
Pleistocene of southern Patagonia, Chile. Journal of
Vertebrate Paleontology, 23: 709–712. http://dx.doi.
org/10.1671/0272-4634(2003)023[0709:TSBPCU]
2.0.CO;2
Quilès, J.; Petrea, C.; Moldovan, O.; Zilhão, J.; Rodrigo,
R.; Rougierf, H.; Constantin, S; Milota, S.; Gherase,
M.; Sarcină, L.; & Trinkaus, E. (2006). Cave bears
(Ursus spelaeus) from the Peştera cu Oase (Banat,
Romania). Comptes Rendus Palevol, 5: 927–934.
http://dx.doi.org/10.1016/j.crpv.2006.09.005
Rabal-Garcés, R. & Cuenca-Bescós, R. (2009). Tafonomía
del yacimiento de osos de las cavernas de Coro Tracito
(Tella, Huesca, España). Paleolusitana, 1: 397–402.
Soibelzon, L. (2004). Revisión sistemática de los Tremarc-
tinae (Carnivora: Ursidae) fósiles de América del
Sur. Revista Museo Argentino de Ciencias Naturales,
6: 107–133.
Soibelzon, L. & Prevosti, F. (2007). Los carnívoros
(Carnivora, Mammalia) terrestres del Cuaternario
de América del Sur. In: Geomorfologia Litoral i
Quaternari. Homenatge a Joan Cuerda Barceló (Pons,
G. & Vicens, D., Eds.), Monografies de la Societat
dʼHistoria Natural de les Balears, Mallorca, 49–68.
Soibelzon, L.; Tonni, E. & Bond, M. (2005). The fos-
sil of South American short-faced bears (Ursidae,
Tremarctinae). Journal of South American Earth
Sciences, 20: 105–113. http://dx.doi.org/10.1016/j.
jsames.2005.07.005
Soibelzon, L.; Pomi, L.; Tonni, E.; Rodríguez, S. &
Dondas, A. (2009). First report of a South American
short-faced bears’ den (Arctotherium angustidens):
palaeobiological and palaeoecological implications.
Alcheringa, 33 (3): 211–222. http://dx.doi.org/10.
1080/03115510902844418
Soibelzon, L.; Grinspan, G.; Bocherens, H.; Acosta, W.;
Jones, W.; Blanco, E.; Prevosti, F. (2014). South
American giant short-faced bear (Arctotherium
angustidens) diet: evidence from pathology, mor-
phology, stable isotopes and biomechanics. Journal
of Paleontology, 88 (6): 1240–1250. http://dx.doi.
org/10.1666/13-143
Stiner, M.; Achyuthan, H.; Arsebük, G.; Clark Howell, F.;
Josephson, S.; Juell, K.; Pigati, J. & Quade, J. (1998).
Reconstructing cave bear paleoecology from skeletons:
a cross-disciplinary study of middle Pleistocene bears
from Yarimburgaz Cave, Turkey. Paleobiology, 24 (1):
74–98.
Trejo, V. & Jackson, D. (1998). Cánidos patagónicos: iden-
tificación taxonómica de mandíbulas y molares del
sitio arqueológico cueva Baño Nevo-1 (Alto Ñirehuao,
XI Región). Anales del Instituto de la Patagonia 26:
181–194.
Velásquez, H. & Mena, F. (2006). Distribuciones óseas de
ungulados en la Cueva Baño Nuevo-1 /XI Región, Chile):
un primer acercamiento. Magallania, 34 (2): 91–106.
http://dx.doi.org/10.4067/S0718-22442006000200009
Villa-Martínez, R., Moreno, P. & Valenzuela, M.A.
(2012). Deglacial and postglacial vegetation changes
on the Eastern slopes of the central Patagonian Andes
(47° S). Quaternary Science Reviews, 32: 86–99.
http://dx.doi.org/10.1016/j.quascirev.2011.11.008
Wolverton, S. (2001). Caves, ursids, and artifacts: a
natural- trap hypothesis. Journal of Ethnobiology,
21 (2): 55–72.
