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E N. Jb. Geol. Paläont. Abh. 289/2 (2018), 177–187Article
Stuttgart, August 2018

Panthera onca (Carnivora, Felidae) in the late Pleistocene-early

Holocene of northern Argentina
Sergio Gabriel Rodriguez, Cecilia Méndez, Esteban Soibelzon, Leopoldo Héctor Soibelzon,
Silvina Contreras, Juan Friedrichs, Carlos Luna, and Alfredo Eduardo Zurita
With 2 figures and 1 table

Abstract: The most northern fossil record of Panthera onca (Linnaeus, 1758) from Argentina is
here reported. The specimen, PVE-F 130, represented by an articulated and well preserved cranium
and mandible, atlas and a fragment of left humerus, was exhumed from levels assigned to the Río
Bermejo Formation (late Pleistocene-early Holocene; ca. 12.0-9.7 ka), near Villa Escolar, Formosa
Province. The associated paleofauna, comprised mostly of large and megafaunal mammals, has a clear
taxonomic similarity to that of the Pampean region of Argentina, and suggests open arid to semiarid
environments. Indeed, previous analyses of plant remains associated with these vertebrates support
the predominance of mega/mesothermal grasslands characterized by C3/C4 grass species.

Key words: Felidae, Quaternary, Formosa, Eastern Chaco, South America, Río Bermejo Formation.

1. Introduction recently the southernmost record of the species were

the fossils recovered from the “Cueva del Milodon”
Panthera onca (Linnaeus, 1758) is among the largest in southern Chile (see Cabrera 1934), dated between
felids and the single living representative of the ge- 13560 ± 180 and 10200 ± 400 ry BP (Tonni et al. 2013).
nus in the Americas, whereas numerous species are However, recently Chimento & Agnolin (2017) indi-
currently distributed throughout the Old World. Up to cated that these materials actually correspond to the
sixteen subspecies of P. onca have been described pri- extinct American lion (Panthera atrox).
marily on the basis of cranial characteristics (Nelson The oldest records of P. onca are those from the
& Goldman 1933), though other authors regard them Irvingtonian (early to middle Pleistocene) of North
as variations within populations (Larson 1997; Eizirik America (Schultz et al. 1985). Based on mitochondrial
et al. 2001). DNA, Eizirik et al. (2001) stated that P. onca origina-
From an historical viewpoint, P. onca was wide- ted between 280 ka and 510 ka BP, whereas the fossil
ly distributed in the Americas, from southern North record suggests this event was probably older, around
America to Rio Negro province in southern Argentina 850 ka BP (Eizirik et al. 2001).
(Seymour 1989; Nowak & Paradiso 1991). However, Felidae arrived in South America during the Great
this species currently occupies less than 50% of that American Biotic Interchange (GABI) after the arrival
territory and it is classified as Near Threatened by the of other carnivorans such as Procyonidae (Huayquerian;
IUCN (Caso et al. 2008). It should be noted that until late Miocene), Galictinae, and Canidae (Marplatan; late

©2018 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/njgpa/2018/0758 0077-7749/2018/0758 $ 2.75

eschweizerbart_xxx
178 S.G. Rodriguez et al.

Fig. 1. A – General outline of the megafan of the Bermejo River showing the location of the fossiliferous locality; B – Geo-
logical section exposed in the margins of the Bermejo River at the Villa Escolar, Formosa Province, with the provenance
of Panthera onca remains (PVE-F 130).

Pliocene) (Cione et al. 2015). The Felidae were one of The large Felidae are represented in the fossil re-
the most successful of the placental carnivorans that dis- cord of the current territory of Argentina by Smilodon
persed from Central and North America during GABI, populator Lund, 1842 and Puma concolor (Linnae-
with at least seven invasions to South America (Sunquist us, 1771) since the early Pleistocene (Soibelzon et al.
& Sunquist 2002; Johnson et al. 2006; Prevosti 2006; 2008; Chimento & Dondas 2017), and by Panthera
Soibelzon & Prevosti 2007; Prevosti & Soibelzon onca (Linnaeus, 1758) since the late Pleistocene, accor-
2012). The oldest South American records of Felidae are ding to Seymour (1983) and Arroyo-Cabrales (2002).
those of the Pleistocene, together with Lutrinae, Mephi- However, other authors (e.g., Van Valkenburgh 1991);
tidae and Ursidae (Berta & Marshall 1978; Soibelzon Berman 1994; Cione et al. 1999; Soibelzon et al. 2008)
& Prevosti 2007; Prevosti & Soibelzon 2012). stated that P. onca is recorded since the early Pleisto-

eschweizerbart_xxx
 Panthera onca (Carnivora, Felidae) in the late Pleistocene-early Holocene of northern Argentina 179

cene. During the late Pleistocene and early Holocene, that according to Iriondo (2010) represent swamp depo-
P. onca is recorded in Argentina, Bolivia, Brazil, Chile, sits from the overflows produced by the aforementioned
Ecuador, Peru, Uruguay, and Venezuela (Winge 1895; river during the Last Glacial Maximum (LGM). These
Hoffstetter 1952; Berta & Marshall 1978; Ochseni- swamp deposits levels are massive, formed predomi-
us 1980; Berman 1994; Cione et al. 1999; San Roman nantly by clay sediments, bioturbated, with the pres-
et al. 2000; Ubilla & Perea 1999; Ubilla et al. 2004; ence of rhizoliths and crotovines and abundant remains
Labarca & Lopez 2006; Soibelzon & Prevosti 2013). of vertebrates (Zurita et al. 2009, 2014; Méndez et al.
In Argentina, fossil records of P. onca are present in the 2017). A radiocarbon dating on bivalves from this sec-
provinces of Córdoba, Entre Ríos, Corrientes, Tucumán tion of the Río Bermejo Formation yielded 9570 ± 90
and numerous localities of Buenos Aires (Bonaparte ry BP (Zurita et al. 2011) (Fig. 1B), whereas two others
& Bobonicov 1974; Berta & Marshall 1978; Berman from the type locality of this unit (Las Lomitas) yiel-
1994; Scillato-Yané et al. 1998; Cione et al. 1999; Fer- ded ca. 12,400 and 11,800 ry BP (Kruck et al. 2011).
rero 2008; Cruz et al. 2012; Francia 2014; Zurita et In accordance with these dates, the fossils collected in
al. 2014; among others). this unit are typically considered a late Pleistocene-
An associated cranium and mandible, together with early Holocene faunal assemblage (Zurita et al. 2009,
post-cranial remains, are here assigned to Panthera 2014). Overlying this unit is the Fidelidad Formation,
onca and were found in outcrops of the Río Bermejo which represents the paleo-channel of the Bermejo Ri-
Formation (late Pleistocene-Holocene) on the banks ver, accumulated by avulsion from the Holocene to the
of the Bermejo River (Formosa Province) in an excel- present. These are formed by intercalations of clay-silty
lent state of preservation. The main objective of this levels with abundant bioturbation and the presence of
contribution is to report this specimen (PVE-F 130), stem and leaf impressions intermixed with mollusks
representing the most northern fossil record of P. onca and ostracods (Contreras 2010; Zamudio 2013; Con-
for Argentina (Fig. 1A, B) and discuss it paleoenviron- treras & Lutz 2014), which would correspond to
mental framework. floodplain deposits, while sandy silt levels finely lami-
nates would correspond to deposits of channels.

2. Geological setting
3. Material and methods
The material analyzed in this contribution (PVE-F 130;
Fig. 2) comes from the margins of the Bermejo River, The specimen described was thoroughly compared
near to Villa Escolar (26°37’ S; 58°40’ W), Formosa with modern and fossil specimens of both Panthera
Province (Argentina) of the Chaco Region. The Cha- onca and Puma concolor. This is because the latter
co Region is a plain that corresponds to an extremely is the only other sympatric felid of comparable size
large sedimentary basin limited by the Andes and the and similar morphology. Smilodon, the other large felid
Brazilian Region. The Paraguay and Paraná rivers flow recorded in the area, was ruled out as an option based
along the eastern boundary, in association with four on the morphology and development of the upper ca-
other principal rivers (Parapetí, Pilcomayo, Bermejo, nines, the lateral compression of the braincase, and the
and Juramento-Salado) that cross the Chaco plain from particular morphology of the zygomatic arches, among
northwest to southeast (Iriondo 1987, 2010); there, per- others. For other hand, the recently published large fe-
manent and temporary swamps are crossed by ancient lid from southern Chile (Panthera atrox, Chimento &
fluvial channels (Iriondo 1984, 2010; Orfeo & Iriondo Agnolin, 2017) duplicates in size fossils and current P.
2012). onca (Prevosti & Martin 2014; Chimento & Agnolin
The eastern part of this region belongs to the Ea- 2017); for this reason we do not take in account for the
stern Chaco Ecoregion of the Chacoan phytogeogra- comparisons.
phical Province (Cabrera 1971; Morrone 2014). This Skull, dental and humeral measurements were taken
includes the east of Chaco and Formosa provinces, nor- with a digital caliper with an accuracy of 0.01 mm.
thwest of Corrientes and north of Santa Fe provinces. Measurements follow those proposed by Morales &
In particular, the fossiliferous site is located within the Giannini (2010), Martin et al. (2011), and Ruiz-García
distal sector of the Río Bermejo megafan (Fig. 1A). & Payan (2013).
From a stratigraphic viewpoint, the material comes The Multidimensional Test of the XLSTAT
from the levels of the Río Bermejo Formation (Fig. 1B) software was used. The tests are used to compare

eschweizerbart_xxx
180 S.G. Rodriguez et al.

samples described by several variables. Instead of 4. Systematic paleontology

comparing the average of two samples as with the
Student t test, we compare here simultaneously for
the same samples averages measured for several Order Carnivora Bowdich, 1821
variables. Compared to a procedure that would Family Felidae Fischer v. Waldheim, 1817
involve as many Student t tests as there are variables, Subfamily Pantherinae Pocock, 1917
the method proposed here has the advantage of using Genus Panthera Oken, 1816
the structure of covariance of the variables and of
obtaining an overall conclusion, reducing significantly Panthera onca Linnaeus, 1758
the probability of commit a Type I error. Here the Fig. 2
significance was analyzed by the Wilks’ Lambda Test
Referred material: PVE-F 130. Articulated cranium and
(through Rao’s approximation). mandible, associated with a fragment of cervical vertebra
(atlas), and a large portion of the diaphysis with complete
Institutional abbreviations: MACN: Museo Argentino de distal epiphysis of the left humerus (Fig. 2). The specimen
Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, was found in sediments of the Río Bermejo Formation, Villa
Argentina. MLP-Ma: Museo de La Plata, División Zoología Escolar (26°39´39.3” S 58°38´19.6” W), Formosa Province,
de Vertebrados, Sección Mastozoología, Facultad de Cien- Argentina (Fig. 1A). Lujanian Stage/Age (late Pleistocene-
cias Naturales y Museo, Universidad Nacional de La Plata, early Holocene).
Buenos Aires, Argentina. LHS: Colección personal L. H.
Soibelzon. PVE-F: Museo Provincial de Ciencias Naturales, Remarks: PVE-F 130 is among the best preserved and most
Villa Escolar, Formosa, Argentina. PVL: Colección de Pa- complete specimens known. This preservation suggests scar-
leontología de Vertebrados del Museo de Ciencias Naturales ce or negligible post mortem transport, little exposure to
“Miguel Lillo”, Tucumán, Argentina. weathering, and rapid burial. The closure of cranial sutures
and that of the humeral epiphysis, coupled with the strong
Dental abbreviations: Upper: I1, first incisor; I2, second wear of the apex and lingual side of the lower canines, sug-
incisor; I3, third incisor; C, canine; PM2, second premolar; gest that it corresponds to an adult individual.
PM3, third premolar; PM4, fourth premolar. Lower: i1, first
incisor; i2, second incisor; i3, third incisor; c, canine; pm3, Description and comparison: The cranium is very well pre-
third premolar; pm4, fourth premolar; m1, first molar. served on the right side, and the left is somewhat weathered,
lacking most of the zygomatic arch and basicranium. The left
Measurements (see Table 1): Skull: TL, total length; POW, hemimandible has no angular apophysis, and both tympanic
postorbital width; IOW, interorbital width; UCL, upper ca- bullae and nasals are missing. As for the upper teeth, the
nine length. Dental: MWUC, mesiodistal width of upper right I1-3, C, and PM 2-4 as well as the left C and PM2-4 are
canine; MWUPM3, mesiodistal width of upper third premo- preserved. In the mandible, the right c, pm4 and m1, and the
lar; MWUPM4, mesiodistal width of upper fourth premo- left i1-2, c pm3-4 and m1 are preserved. The humerus pre-
lar; UCPM4L, upper canine-fourth premolar length; LCL, serves the distal epiphysis and a large part of the diaphysis,
lower canine length; MWLC, mesiodistal width of lower missing only the center portion of the bridge that defines the
canine; MWLPM3, mesiodistal width of lower third premo- entepicondylar foramen.
lar; MWLPM4, mesiodistal width of lower fourth premolar; The cranium of PVE-F 130 differs from that of Puma
LPM3PM4L, lower third premolar-fourth premolar length; concolor because it is anteroposteriorly elongated (as in
MTL, mandible total length. Humerus: MWD, mesiodistal Panthera onca) and not globose; the dorso-posterior end of
width of diaphysis; AWD, anteroposterior width of diaphysis; the skull (where the sagittal and lambdoidal crests merge)
DEW, distal epiphysis width. is dorsoposteriorly oriented as in P. onca, and not postero-
ventrally as observed in P. concolor. The posterior angle of
Reference material: Panthera onca: MLP-Ma 24-IX-01-6, the zygomatic arch is well defined (as in P. onca), whereas
MLP-Ma 24-IX-01-7, MLP-Ma 24-IX-01-8, MLP-Ma 119, the zygomatic arch of P. concolor describes almost a cir-
MLP-Ma 294, MLP-Ma 456, MACN 7.7, MACN 7.8, MACN cumference arch. In addition, the cranium length of PVE-F
8-42, MACN 8.43, MACN 17.1, MACN 17.2, MACN 253, 130 (271.00 mm) falls within the range of Panthera onca
MACN 25.66, MACN 992, MACN 21622, MACN 25778, following ranges established by Chimento & Dondas (2017)
MACN 26.217, MACN 33.168, MACN 38.265, MACN (P. onca, 253.00-300.00 mm; Puma concolor, 171.00-228.81
51.147, LHS s/n. Puma concolor: MLP-Ma 1633, MLP-Ma mm). The mandible of PVE-F 130 is similar to that of P.
1306, MLP-Ma 292, MLP-Ma 1312, MLP-Ma 455, MLP- onca, which clearly differs from that of P. concolor due to
Ma 552, MLP-Ma 1309, MLP-Ma 14-IV-48-2, MLP-Ma the greater relative development of the ventral edge of the
14-IV-48-3, MLP-Ma 14-IV-48-4, MLP-Ma 1770, MLP-Ma masseteric fossa, morphology that is notoriously laterally
1310, MLP-Ma 9-X-92.2, MLP-Ma 2-VI-60.4, MLP-Ma expanded in the former species.
694, MLP-Ma 31.VIII-98-2, MLP-Ma 3.IX.01.15, MLP-Ma The atlas of PVE-F 130 preserves only a portion of the
3.IX.01.18, MLP-Ma 3-IV.01.16, MLP-Ma 13.IV.99.45, MLP- body, featuring the dorsal and ventral arches, the right po-
Ma 16.XII.98.18. sterior articular facet, and a small part of the right transverse

eschweizerbart_xxx
 Panthera onca (Carnivora, Felidae) in the late Pleistocene-early Holocene of northern Argentina 181

Fig. 2. A-C – Skull of PVE-F 130 in lateral, dorsal and anterior views, respectively; D-E – left humerus of PVE-F 130 in
posterior and anterior views, respectively; F, G – atlas of PVE-F 130 in anterior and dorsal views, respectively. Scale equals
1 cm.

eschweizerbart_xxx
182 S.G. Rodriguez et al.

apophysis. Due to the fragmentary state of the material it temperate/warm-temperate and semiarid/subhumid cli-
cannot reliably be compared with P. concolor. mates with seasonal conditions and open environments
The condyle of the humerus of PVE-F 130 is more me-
dominated by mixed grasslands, suitable for the large
mammals (Contreras & Zucol 2018).
diolaterally expanded than in Puma concolor, as observed in
Panthera onca. The capitulum is relatively more developed
than the trochlea, as in P. onca, whereas these structures are The presence of P. onca together with the taxa listed
subequal in size in P. concolor. In P. concolor the trochlea above is further evidence of the diversity of habitats to
extends more distally respect to the capitulum than in PVE-F which this taxon is adapted. Labarca & López (2006)
130 and P. onca. The entepicondyle of PVE-F 130, as well as stated that it is difficult to infer paleoenvironmental con-
that of P. onca, is a little more developed than in P. concolor.
The lateral crest is more laterally developed in PVE-F 130 ditions based solely on the presence of P. onca due to the
and in P. onca compared to P. concolor. high adaptive capacity of this felid. The species inhabits
When comparing the measurements of PVE-F 130 with myriad environments including rain forests with tropical
those of several specimens of Panthera onca (Table 1), six vegetation, areas with ephemeral marshes, grasslands,
fall within the range of this species, three are below the lower shrubby regions, and dry deciduous forests (Nowell &
Jackson 1996). In modern ecosystems, P. onca depends
limit, and eight exceed the upper limit.
Based on the above descriptive and quantitative mor-
phological comparisons, PVE-F 130 is here assigned to the large on water resources and has a marked preference
species Panthera onca. In agreement with this interpretation, for rivers and lagoons (Mondolfi & Hoogesteijn 1986).
the Wilks Lambda Test results suggested that fossil PVE-F Hence, the tolerance for different climatic-environmental
130 and modern specimens of Panthera onca are not signi- conditions by P. onca is evident. Noteworthy, carnivo-
ficantly different (P= 0, 8).
rans exhumed at levels assignable to the Río Bermejo
Formation include the procyonid Procyon cancrivorus
(Soibelzon et al. 2010), and the canid Protocyon cf. tro-
5. Paleoenvironment
glodytes (Prevosti et al. 2005).
The mammal assemblage of the late Pleistocene-ear- From a paleoecological perspective, the recorded
ly Holocene of the Río Bermejo Formation is mostly taxa were likely adapted to diverse climatic and envi-
“Pampean-Patagonian” in origin, and is characterized ronmental conditions during the Pleistocene-Holocene.
by, among others, the Artiodactyla Hemiauchenia pa- In this scenario, short humid and warm pulses may
radoxa and Morenelaphus lujanensis, the Xenarthra have stimulated the presence of taxa with intertropical
Megatherium sp., Glyptodon sp., Neosclerocalyptus affinities, whereas arid and cold pulses would have fa-
paskoensis, Pampatherium typum and the Notoungula- vored the presence of taxa of Pampean “origin” (Car-
ta Toxodon sp. (Zurita et al. 2009; Alcaraz & Ferrero lini et al. 2008, 2004). Such a combination of species is
2011). To date, the single taxon with tropical affinities comparable to the fossil record of Dique “Los Quiroga”
recorded in this unit is Holmesina paulacoutoi (Pam- in the Santiago del Estero Province where a “mix” of
patheriidae, Cingulata). typically Pampean and Brazilian taxa is recorded (Chi-
Zurita et al. (2014) stated that, from a paleoenviron- mento & Agnolin 2011). Similar situation was obser-
mental point of view, most taxa found in Formosa Pro- ved throughout the Mesopotamian region of Argentina.
vince indicate the development of open environments, In this area, some taxa suggest open grasslands with
suitable for the settlement of large mammals, with an cold and arid/semiarid climate (e.g., Neosclerocalyptus
arid to semiarid climate. This interpretation is congruent paskoensis, Glyptodon reticulatus), while others sug-
with palaeoclimatic inferences established for the late gest warm and humid environments with tropical to
Pleistocene-early Holocene (Tonni et al. 1999; Cione et subtropical climate (e.g., Holmesina paulacoutoi, Boa
al. 2007; Tonni 2009). In addition, phytolitic analysis constrictor, Tayassu pecari, and Tapirus sp.) (Zurita
of the strata associated with these vertebrates showed et al. 2014).
mega/mesothermal mesophytic grasslands (especially
Panicoideae (C3/C4) and Chloriroideae (C4) subfamilies);
with some herb and shrub or wood dicot species (Con-
6. Conclusions
treras & Zucol 2018). C4 grass species tend to occupy
dryer niches than C3 grasses (Taylor et al. 2011). The The most northern record of Panthera onca from the
presence of Podostemaceae and sponge spicules sug- late Pleistocene-early Holocene of Argentina is re-
gests that a floodplain with water bodies such as rivers ported. PVE-F 130 is a very well preserved specimen
or streams were also present. In additions, during this found in the Formosa Province. Panthera onca repre-
period, the Chaco environment was characterized by sents a new taxon added to the list of fossil fauna of the

eschweizerbart_xxx
 Panthera onca (Carnivora, Felidae) in the late Pleistocene-early Holocene of northern Argentina 183

Eastern Chaco, and joins the small number of species Bogan, S., Zurita, A., Miño-Boilini, A.R., Suárez, P., Frie-
not typically considered Pampean, such as Holmesina drichs, J., Lutz, A. & Friedrichs, J. (2012): Primer reg-

paulacoutoi and Procyon cancrivorus, present in this istro de Synbranchiformes (Teleostei) para el Holoceno
temprano de Argentina. – Comunicaciones del Museo
region. Panthera onca is recorded in a wide diversity Argentino de Ciencias Naturales “Bernardino Rivada-
of habitats, suggesting a high degree of environmental via”, 14 (1): 41-45.
plasticity for this taxon. Bowdich, T.E. (1821): An Analysis of the Natural Classifica-
The excellent state of preservation of this specimen tions of Mammalia for the Use of Students and Travelers.
will allow future contributions, including systematic – 115 pp.; Paris (J. Smith).
Cabrera, A. (1934): Los yaguares vivientes y extinguidos
review of the taxon as well as geomorphometric and de la América austral. – Notas del Museo de La Plata,
phylogenetically-informed analyses to determine and 2: 34-50.
quantify intraspecific variation within fossil and mo- Cabrera, A. (1971): Fitogeografía de la República Argentina.
dern specimens of P. onca. – Boletín de la Sociedad Argentina de Botánica, 14: 1-42.
Cabrera, A.L. & Willink, A. (1973): Biogeografía de Amé-
rica Latina. Monografía 13, Serie de Biología, OEA,
Washington D.C.: 120 pp.
Carlini, A.A., Zurita, A.E., Gasparini, G.M. & Noriega J.I.
Acknowledgements (2004): Los mamíferos del Pleistoceno de la Mesopota-
To the staff of the institutions that allowed access to the mia argentina y su relación tanto con aquéllos del Centro
collections: Marcelo Reguero, Martín de los Reyes, Itatí Norte de la Argentina, Paraguay y Sur de Bolivia, como
Olivares, Diego H. Verzi (Museo de La Plata), Alejandro con los del Sur de Brasil y Oeste de Uruguay: Paleobio-
K ramarz, Laura Chornogubsky, Sergio Lucero, Pablo geografía y Paleoambientes. – In: Aceñolaza, F.G. (Ed.):
Teta (Museo Argentino de Ciencias Naturales “Bernardino Temas de la Biodiversidad del Litoral Fluvial Argentino:
Rivadavia”), José Friedrichs (Museo Provincial de Ciencias Tucumán, Universidad Nacional de Tucumán. – INSUG-
Naturales, Formosa), Pablo Ortiz (Museo de Ciencias Na- EO, Miscelánea, 12: 83-90.
turales “Miguel Lillo”). We also want to thank Eric Lynch Carlini, A.A., Zurita, A.E. & Miño-Boilini, A.R. (2008):
(East Tennessee State University) for the correctness of the Reseña paleobiogeográfica de los Xenarthra (Mamma-
language and his comments that improved the manuscript. lia) del Pleistoceno tardío de la región Mesopotámica
The comments of Nicolás Chimento and another anonymous (Argentina). – INSUGEO, Miscelánea, 17 (2): 259-270.
reviewer allowed us to improve the final version of the ma- Caso, A., Lopez-Gonzalez, C., Payan, E., Eizirik, E., de
nuscript. This contribution was partially funded by projects Oliveira, T., Leite-Pitman, R., K elly, M. & Valder-
PICT 2016/0609, PICT 2017/0765 and PIQ 002/17. rama, C. (2008): Panthera onca. – The IUCN Red
List of Threatened Species 2008: e.T15953A5327466.
http://dx.doi.org/10.2305/IUCN.UK.2008.RLTS.
T15953A5327466.
Chimento, N.R. & Agnolin, F.L. (2011). Mamíferos del
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186 S.G. Rodriguez et al.

Manuscript received: April 27th, 2018. Esteban Soibelzon, Leopoldo Héctor Soibelzon, División
Revised version accepted by the Stuttgart editor: July 2nd, Paleontología Vertebrados, Museo de La Plata, Facultad de
2018. Ciencias Naturales y Museo, Universidad Nacional de La
Plata (CONICET), Paseo del Bosque s/n, 1900 Buenos Aires,
Argentina.
Addresses of the authors: Juan Friedrichs, Museo Provincial de Ciencias Naturales de
Sergio Gabriel Rodriguez, Facultad de Ciencias Naturales
Villa Escolar. Formosa, Argentina.
y Museo (UNLP), 122 y 60, La Plata, Buenos Aires, Ar-
gentina;
e-mail: [email protected]
Cecilia Méndez, Silvina Contreras, Carlos Luna, Alfredo
Eduardo Zurita, Centro de Ecología Aplicada del Litoral
(CECOAL-CONICET-UNNE). Ruta 5, km 2.5. W3400 Cor-
rientes, Argentina.

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 Panthera onca (Carnivora, Felidae) in the late Pleistocene-early Holocene of northern Argentina 187

Table 1. Comparative measurements of specimens of Panthera onca and PVE-F 130 (see Materials and Methods). Measu-
rements in mm.
MLP-Ma 24- MLP-Ma 24- MLP-Ma 24-
MLP-Ma 119 MLP-Ma 456 MLP-Ma 294 LHS s/n MACN 992 MACN 8-42 MACN 25778
IX-01-7 IX-1-8 IX-01-6

TL 272 282 291 302 271 260 253 279 285 295

APO 49.8 51.9 62.35 66.8 47.2 51.5 57 68.6 53.2 63.4

AIO 51.53 50.1 57.72 58.5 49.7 46.1 51 54.9 50.1 57

LCS 21.68 24.5 23.58 25.9 21.9 22.3 20.2 22.1 23.7 24

ACS 16.9 19.6 20.02 23.1 16.7 17.4 17 17.4 20.2 20.5

APM3 19.83 20.09 20.8 20.6 19.9 19.2 19.4 20.8 20.6 19.5

APM4 30.1 29.8 31.07 29.9 29.7 27.5 29.2 31.2 29.9 29.7

LCPM4 81.49 84.1 86.9 87.55 82.8 80.8 81.48 88.4 87.8 89.2

LCI 22.58 22.4 23.1 x 21.1 21.3 20.5 19.3 23.4 23

ACI 14.83 16.1 15.9 x 18.6 15.7 14.9 16.8 17.3 16.8

APM3I 15.25 16.99 15.86 x 15.3 15.03 14.5 16.7 16.6 15.8

APM4I 20.45 22 22.46 x 18.8 20.1 21.8 22.6 22.5 20.8

LPM3PM4I 35.7 37.9 37.85 x 34.6 35.4 37.2 38.3 42.1 35

LTM 183 193 187 x 174 172 165 190 186 188

ADH 22.36 x x x x x x x x x

AEH 63.1 x x x x x x x x x

AAPH x x x x x x x x x x

Table 1. (continued)
MACN MACN MACN MACN MACN MACN MACN MACN MACN MACN MACN MACN PVE-F
Range
253 17.1 17.2 8.43 25.66 7.7 21622 51.147 38.265 33.168 26.217 7.8 130

247 266 270 241 246 266 244 269 229 265 263 231 229-302 271

46.88 50.4 46.3 45.9 45.1 47.9 47.3 x 48.6 46.6 55.2 48.4 45.1-68.6 42.1

46.6 51.7 52.5 47.6 43.2 46.2 40.4 49.3 43.3 49.9 56.6 43.5 40.4-58.5 46.8

20.76 22.7 24.8 26.2 24.3 24.8 16.2 18.9 18.9 20 22.4 19 16.2-26.2 21.3

16.3 18 18.1 19.8 18.1 18.2 12.7 x 14 15.7 16.5 14.8 12.7-23.1 18.2

19.7 18.8 20.2 19.7 21.1 19.9 16.7 x 19 20.5 21.2 18.7 16.7-21.2 21.5

27.9 26.8 30.8 29.3 29.2 30 25.3 28.4 25.7 29.3 29.2 29 25.3-31.2 33.7

75.9 77.8 82 82.2 82 83.3 74.6 x 75.1 81 81.3 75.9 74.6-89.2 93.2

21.4 23.7 22.9 20.8 22.5 24 17.7 20.7 17.5 19.8 21.5 19.8 17.5-23.7 16.8

15.9 15.8 18.7 14.7 16.3 17.8 11.7 14.4 13.1 15.6 14.8 15.8 11.7-18.7 15.2

16.1 14.3 16.5 16.9 17 17.3 12.7 15.3 15.2 15.4 16.4 14.9 12.7-17.3 19.6

22 19 21.6 22.3 21.2 22.3 18.6 20.7 19.4 20.6 22.4 20.2 18.6-22.6 22.8

40.6 33.3 41.7 42 39.8 42 34.8 36.7 36.7 40.8 40.9 38.8 33.3-42.1 44.3

161.5 169 162 154 162 176 154 166 148 167 176 155 148-193 187

x x x x x x x 22.4 x x x x 22.36-22.4 22.2

x x x x x x x 63.7 x x x x 63.1-63.7 65.7

x x x x x x x 34 x x x x 34 38.2

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