337

Martn
Article JTa072. All rights reserved.
*E-mail: fabiana1917@yahoo.com.ar
2008

Journal of Taphonomy
PROMETHEUS PRESS/ PALAEONTOLOGI CAL NETWORK FOUNDATI ON (TERUEL)

VOLUME 6 (ISSUE 3-4)


Available online at www.journaltaphonomy.com

The fragmented bone remains of extinct mammals recovered at several late Pleistocene sites in Fuego-
Patagonia are analyzed. Indications of human involvement with the bones are not abundant and some of
the sites are purely paleontological. However, all of them preserve large carnivore tooth-marks. Some of
the sites can be explained as accumulations produced by extinct felids.

Keywords: FELIDS, MYLODONTINAE, HORSES, FUEGO, PATAGONIA, PLEISTOCENE.


Bone Crunching felids at the End of the
Pleistocene in Fuego-Patagonia, Chile

Fabiana M. Martn*
Fundacin CEQUA, Punta Arenas, Chile


Journal of Taphonomy 6 (3-4) (2008), 337-372.
Manuscript received 4 January 2008, revised manuscript accepted 3 April 2008.
Introduction

Fuego-Patagonia is an extensive landmass
located in the southern tip of South America.
It is characterized by a variety of open
environments (Moore, 1978). Three of these
areas, Pali Aike, Ultima Esperanza and
North of Isla Grande of Tierra del Fuego
(Figure 1), have yielded important late
Pleistocene bone assemblages. Paleoclimatic
and palaeocological data based mainly on
pollen columns indicate that steppe or
Patagonian tundra environments have
dominated the region during post-glacial times
(ca. 14,000 BP) (Markgraf, 1988; Clapperton,
1993; Heusser, 2003; Zolitschka et al., 2004;
McCulloch et al., 2005; Villa-Martinez &
Moreno, 2007). Only near the Cordillera has
a Northofagus forest been present since
10,000 BP (Moore, 1978; Heusser, in Prieto
1991; Heusser et al., 1992; Markgraf, 1993;
Heusser, 2003).
The geography of Fuego-Patagonia at
the end of the Pleistocene was quite different
from today. Ecosystems were unstable after
the retreat of the ice masses (Pisano, 1975),
as what is today the island of Tierra del
Fuego was still connected to the continent
(McCulloch et al., 1997). In addition, the
faunal community was more diverse than today
(Miotti & Salemme, 1999) and included
many extinct species, among them several
338
Taphonomy of Panthera onca

Materials

Information recovered at several paleontological
and archaeological caves is revised in this
paper. Lago Sofa 4 cave (CLS4) is a dark
cave, at Ultima Esperanza, Chile (51 31
54 S, 72 34 12W) excavated by Alfredo
Prieto and Pedro Crdenas. Hundreds of
remains from both extinct and extant fauna
were found in the cave, which measures ca.
6.0 m by 2.5 m with a maximum height of
about 1.5 m. There is no evidence of human
presence or activities at this cave. The site
was dated between 13.4 and 10.8 thousand
years (ka) BP (Table 2) and was partially
synchronic with the human occupations
discovered at nearby CLS1, which is dated
between 11.5 and 10.1 ka BP (Prieto, 1991;
Massone & Prieto, 2004). The absence of
humans at CLS4 could be related with the
morphology, darkness, and difficulties of
access of the cave, which is not adequate for
human habitation. The presence of carnivore
bones and the study of the marks found on
the bone assemblage prompted an interpretation
of the site as a Panthera onca mesembrina
den, a study that demonstrated the importance
of Pleistocene carnivores as bone accumulators
and destructors (Prieto, 1991; Borrero et al.,
1997). As we will see below, this interpretation
is not shared by Tonni et al. (2003). The
study of the bone assemblage produced
evidence for the presence of Smilodon sp.
(Canto, 1991); however, the marks and
fragmentation were attributed to panthers
(Borrero et al., 1997). The study of the marks
was difficult due to the regular preservation
of the bones. A recent reanalysis of the
carnivore bones by Francisco Prevosti
identified the presence of three panthers at
the site, one of them a cub with milk teeth
(Figure 2). We present the taphonomic
analysis of the bone assemblage obtained in
carnivores. Table 1 includes all herbivores
and carnivores for which there is good
information for each of the three areas. It
must be noted that the carnivore guild
included large felids with bone destruction
capabilities.
Despite several important paleontological
discoveries, the presence of bone-crunching
carnivores in Fuego-Patagonia was not
recognized until recently. The remains of
large extinct cats and other carnivores were
known at least since the end of the 19
th

century (Roth, 1899; Nordenskjld, 1996
[1900]). The finding of large carnivores
like bears (Arctotherium tarijense) and panthers
(Panthera onca mesembrina) (Roth, 1899,
1904; Smith Woodward, 1900; Nordenskjld,
1996 [1900]) in late Pleistocene contexts at
Milodn cave, Ultima Esperanza, Chile, was
the first indication. However, their presence
was not accompanied by the recognition of
their marks on bones. Thus, Lehmann-Nitsche
(1899) in his study of the Milodn cave
bone assemblage described marks on Mylodon
bones -most of which today we assign to
extinct cats (Martn, 2007)- as inflicted by
humans. The paradigm of the time was to
look for the early presence of humans, a
search that at least partially inhibited the
exploration of alternatives.
In this paper we present the results
of taphonomic studies of several bone
assemblages, some of them recovered in the
19
th
century or early in the 20
th
century. For
that reason the quality of the basic information,
especially that related with the recovery
techniques of the samples, is variable. However,
we believe that it is valuable to use these
collections, since combined with newly acquired
chronological information this taphonomic
study helps to understand the role of large
carnivores at the end of the Pleistocene in
Fuego-Patagonia.
339

Martn
Table 1. Taxa identified in the three study regions.
Taxon Ultima Esperanza Pali-Aike Tierra del Fuego
Hippidion saldiasi X X X
Mylodon sp. X X X
Lama guanicoe X X X
Lama morfotipo owenii X - -
Vicugna sp. X - X
Macrauchenia patachonica X - -
Rheidae - X -
Panthera onca mesembrina X X X
Smilodon populator X - -
Puma concolor X - -
Arctotherium tarijense X X -
Dusicyon avus X X X
Dusicyon griseus X X X
Dusicyon culpaeus X X X
Figure 1. Map showing the location of archaeological and paleontological sites in the three study areas.
340
Taphonomy of Panthera onca

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341

Martn
extinction of the Pleistocene fauna
(Borrero, 1997).
Milodn cave is a huge cave of about
150 m by 180 m located on the Cerro Benitez
flanks (51 33 54S; 72 37 13W). Its
late Pleistocene component is dated by several
radiocarbon dates between 13.6 and 10.2 ka
BP (Borrero, 1997; Borrero, pers. comm.). It
must be noted that Medio cave, with an
important penecontemporaneous archaeological
assemblage dated 11.0 and 9.5 ka BP, is
located about 1000 m east of Milodn cave
(Nami, 1987; Nami & Menegaz, 1991, Nami
& Nakamura, 1995). We already mentioned
that carnivore bones were identified at
Milodn cave before the end of the 19
th

century, as part of the rich bone assemblage
excavated by Hauthal and Nordenskjld
(Hauthal, 1899; Roth, 1899, 1904; Smith
Woodward, 1900; Nordenskjld, 1996 [1900]).
There were new excavations throughout the
20
th
century that produced abundant bones.
The recorded species are Mylodon darwini,
Hippidion saldiasi, Panthera onca mesembrina,
an excavated sample of 3 m
2
. All the sediments
were screened.
Beyond CLS4, several bone assemblages
can be used to discuss the importance of
carnivore activities at the end of the Pleistocene,
among them Milodn cave and the Dos
Herraduras rockshelters, also located at Ultima
Esperanza, and Chingues cave (CDLCH),
Puma cave, and Fell cave at the Pali Aike
Volcanic Field. Also, the Tres Arroyos
rockshelter, located in the island of Tierra
del Fuego (Figure 1), provides a relevant
bone assemblage. Bones of several species
recovered at these sites display large
carnivore marks, although in this paper we
will concentrate on Mylodon and Hippidion
saldiasi, since these are the larger animals
present in those assemblages. These bone
assemblages are all dated between ca. 13.8
and 10.0 ka BP and will be succinctly
presented here (Table 2). This is a time
period at which two main proceses took
place, the human colonization of the
southern tip of the Americas and the
Figure 2. Mandible of a Panthera
onca mesembrina cub from
Lago Sofa 4 cave.
342
Taphonomy of Panthera onca

Table 3. Chingues cave. Hippidion saldiasi
specimens with cutmarks.
regionally dated to about 12.6

ka BP (Stern,
1992; Borrero & Massone, 1994; McCulloch
et al., 2005). This tephra rests above a
geological deposit formed at the shores of
an ancient lake. A complete femur preserves
large carnivore marks (Favier Dubois &
Borrero, 1997). When this discovery was
published this element was anomalous, but
after the publication of CLS4 and evidence
found at other sites it can be considered as
part of the taphonomic background noise for
southern Patagonia. In other words, bones
with felid marks larger than those produced
by living carnivores are present at most of the
known late Pleistocene bone assemblages.
Chingues cave is a relatively small
site-a lava tube of approximately 10 m by 3
m-with a somewhat restricted access that is
located within a maar (52 05 37 S; 69
4431W). It contains two chambers, one of
them dark and only accessible through a narrow
passage. Two components were identified,
one Pleistocene and one Holocene. The
Pleistocene component was considered
purely paleontological, while the Holocene
presents alternate use by humans and
carnivores (San Romn et al., 2000). The
Pleistocene context is dated between 12.1 and
10.1 ka BP (Table 2) and was interpreted
as a large felid den. This restudy of the
late Pleistocene bone assemblage suggests
that there is also evidence for ephemeral
human use of the cave at that time. The
evidence for late Pleistocene human
presence at the site consists of four horse
bones with cutmarks (Table 3). In addition,
three beads made on Mylodontinae dermal
ossicles were found. However, the latter
could also be the result of early Holocene
humans-whose presence is well attested at
the cave-scavenging the dermal ossicles.
The faunal remains at the Pleistocene
component includes bear (Prevosti et al.,
Arctotherium tarijense, Dusicyon avus,
Macrauchenia patachonica and Camelidae
(Roth, 1899; Smith-Woodward, 1900;
Emperaire & Laming, 1954; Saxon, 1979;
Bird, 1988; Borrero et al., 1991). According
to Mol et al. (2003), Smilodon sp. is also
present at the cave with two bone samples
dated around 11.4 and 11.2 ka BP (Barnett
et al., 2005) (Table 2). These samples are
stored at the Zoological Museum of Amsterdam
University, The Netherlands, and its exact
provenience is not necessarily Milodn
cave. Judging from the available written
sources, Medio cave can be suggested as the
origin of that collection (Martn, 2007). In
this paper I will present evidence obtained
in a restudy of the Hauthal collection (1899-
1900) stored at the Departamento Cientfico
Paleontologa de Vertebrados, Museo de La
Plata, Argentina. A small sample stored at the
Museo Argentino de Ciencias Naturales
Bernardino Rivadavia (MACN) will be also
described.
The Dos Herraduras rockshelters are
located at cerro Benitez, 500 m north of
Mildon cave (51 33 36 S; 72 37 22).
It is a purely paleontological context dated
between 12.8 and 11.3 ka BP (Table 2)
which was recorded at rockshelters 2 and 3.
Mylodontinae bones were found partially
embedded within a tephra layer at rockshelter
3 in an excavated area of 10 m
2
, practically
the whole roofed space. This tephra was
fingerprinted to the Reclus volcano and was

Portion
Scapula dist.
Radius dist.
Carpal complete
Metapodial III dist.
343

Martn
by different teams produced a long sequence
of human occupations beginning at the end
of the Pleistocene. The remains of extinct fauna
together with human artifacts were recovered
in the lower layers, which are dated between
11.0 and 10.0 ka BP (Bird, 1988). Extinct
faunal remains included Mylodon darwini
(Martn, 2007), Mylodontinae cf. Mylodon,
Hippidion saldiasi (Alberdi & Prieto, 2000),
and Dusicyon avus (Caviglia, 1985-1986).
However, modern fauna, especially guanaco
(Lama guanicoe), dominates the bone assemblage.
The site was excavated in the
1930s and it was originally concluded that
all the bones resulted from human predation
activities (Bird, 1988). Additional work by
Emperaire et al. (1963) suggested that the
ground sloth bones were deposited before
the arrival of humans. However, most
modern sources indicate that ground sloths
constituted human prey. Physically associated
with the bone assemblage were found several
projectile points of the so-called fishtail
model, retouched lithic tools, flakes, debitage,
bone tools and several hearths. In this paper we
present the results of a study of a previously
unstudied collection obtained by John Fell in
1952-1959 from an area of about 11 m
2
and
Junius Bird in 1969-1970 from an
unspecified area. In addition, we have
studied some bones recovered by Pedro
Crdenas on the surface of the shelter near
the only exposed profile in 1999. The study
of these samples demostrated the presence of
cutmarks on Hippidion saldiasi and
Mylodontinae bones (Martn, 2007) (Table
4).
Tres Arroyos is a small site of
about 6 m by 4 m located at Cerro de los
Onas (53 23 S, 68 47.W), Tierra del
Fuego (Massone, 1987, 2004; Borrero, 2003).
An archaeological bone assemblage dated
between 10.6 and 10.1 ka BP was recovered
2003; Soibelzon et al., 2005), camelids,
Rheidae, Mylodontinae (San Romn et al.,
2000), Panthera onca mesembrina (F. Prevosti,
pers. comm.), and Hippidion saldiasi (Alberdi
& Prieto, 2000). The study of a sample of
the bone assemblage recovered in an excavated
area of 6 m
2
is presented here. All the sediments
were screened.
Puma cave is a large and complex
site located at the Estancia Brazo Norte,
(52 1 12,6 S, 69 58 41,2 W), about 2
km from the Chico River (Martn et al.,
2004). Its small entry opens toward the north
on the external wall of an extinct crater.
Puma cave has a maximum depth of 49 m
with a variable width. Several lateral chambers
of varied shape and size define a complex
tunnel system. A 16-meter-long passage,
almost completely filled with sediments,
leads to a large dark chamber. The topography
of this chamber is irregular and its height is
variable, reaching a maximum of about 6 m.
A large bone assemblage was discovered
on the surface and in stratigraphy at different
chambers of the cave. The faunal list for
extinct animals includes Mylodon sp.,
Hippidion saldiasi, Panthera onca mesembrina,
Arctotherium tarijense, Dusicyon avus, and
Camelidae. Preliminary studies of this bone
assemblage preserve carnivore marks. The
only evidence of human activity so far
discovered at Puma cave is a single sidescraper
found on the surface of one of the lateral
chambers. However, no evidence in the
form of cutmarks was found on the bones.
All the evidence suggests it is basically a
paleontological accumulation deposited
between 11.5 and 10.3 ka BP (Table 2).
Fell cave is a small site that measures
about 11 m by 8.5 m and opens to the valley
of the Chico river (52 02' 40'' S, 70 03'
23'' W) and was formed by river erosion
(Bird, 1988). Excavations through the years
344
Taphonomy of Panthera onca

Table 4. Fell cave. Mylodontinae cf. Mylodon and Hippidion saldiasi specimens with cutmarks.
reevaluation of the history of the peopling
of the region (Borrero, 2003). By considering
the existence of non-anthropic elements in the
assemblages, as well as by giving importance
to paleontological sites, a more balanced
panorama of the ecology of the end of the
Pleistocene was possible. The consideration
of large carnivores and their palaeoecology
made it possible to discuss the relationships
of these animals with their prey, and to
begin to understand their interaction with
humans. The study of the bone surface
modifications proceeded by the identification
of marks produced by carnivores. In order
to do that a reference collection of bones
marked by pumas (Puma concolor) obtained
in a taphonomic naturalistic study conducted
at Torres del Paine National Park, Chile
(Borrero et al., 2005) was used, as well as
published evidence for modern analogs for
extinct species.
Fossil Panthera onca in the Pampas
was a large animal, with a body mass of
about 95-137 kg (Prevosti & Vizcaino,
2006:409). The patagonian panthers were
larger (F. Prevosti pers. comm.) and were
compared in size with the African lion
(Cabrera, 1934). Their presence at many late
Pleistocene bone assemblages, sometimes
associated with tooth-marked bones, suggest
that they were important agents of bone
in several excavations (Massone et al., 1998;
Massone & Prieto, 2004).
Tres Arroyos is the only important
rockshelter in the north of the island. The
archaeological assemblage is characterized by
five discrete hearths (Massone, 2004), fragments
of at least two projectile points -probably
belonging to the fishtail model (Jackson,
2002)- retouched lithic tools, debitage, and
bone tools. The faunal remains are heavily
fragmented, and some bones preserve cutmarks,
including a horse right tibia shaft (Mengoni
Goalons, 1987). Below the archaeological
layers there is a paleontological bone
accumulation dated between 12.5 and 11.0
ka BP (Table 2), basically constituted by a
few ground sloth and horse specimens with
the presence of Panthera onca mesembrina.
The distinction between layers V and VI is
not clear cut, and for that reason the
discussion will not make that difference. A
bone assemblage recovered in an excavated
area of 9 m
2
obtained at both layers was
studied. The sediments were screened.


Methodology

The introduction of taphonomic analysis to the
study of Fuego-Patagonian bone assemblages
was an important component of the
Taxon Element Portion
Mylodontinae cf. Mylodon Pelvis cotiloid cavity
Mylodontinae cf. Mylodon Ulna complete
Hippidion saldiasi Humerus dist. + shaft
Hippidion saldiasi Metacarpal III y II prox. + shaft
Hippidion saldiasi Sesamoid dist. complete
Hippidion saldiasi Metapodial III shaft
Hippidion saldiasi? Femur? shaft
345

Martn
known that bears are solitary hunters or
scavengers (Frison, 2004; Pinto Llona et al.,
2005), they do not usually accumulate
bones (Pinto Llona et al., 2005:83) and only
rarely inflict the full range of damage of
which they are capable (Haynes, 1983:169).
However, they can damage compact bone of big
ungulates, even producing some fragmentation
(Haynes, 1980a:348; Pinto Llona et al.,
2005:74). Bears can produce scores that
appear similar to rodent gnaw marks: short
and parallel, shallowly etched, straight score
lines (Haynes, 1983:169) and modify bones
creating nearly flat-bottomed holes and sets
of parallel furrows on the crest of long
bones (Haynes, 1983:169; Pinto Llona et
al., 2005). The use of their cheek teeth to
grind off the lateral and medial edges of the
proximal epiphyses was recorded (Haynes,
1980a:347). Andrews and Fernndez-Jalvo
recorded tooth-marks-pits and punctures up
to 10.4 mm diameter-very likely made by
bears on bear fossil bones at Sima de los
Huesos, Spain. They found that marks are
present on the shaft surfaces, as well as on
articular surfaces and on intact bone edges
(Andrews & Fernndez-Jalvo, 1997: 212).
Summing up the bone surface
modifications produced by bears, Haynes
mentions low frequencies of tooth-marking,
grinding bone prominences with teeth-leaving
smooth stumps-production of square or
rectangular tooth impressions in trabecular
bone, and short scratches on shafts similar
to those produced by rodents; i.e., short,
straight, parallel and shallowly etched lines
(Haynes, 1983:169).
The percentage of marked bones as
well as the location of the marks were also
considered. Pits alone are not good enough
to identify specific taxa (Domnguez-Rodrigo
& Piqueras, 2003). However, it is possible
to make a distinction between pits produced
accumulation and destruction. The study of the
marks of fossil felids is not easy. As indicated
by Haynes, the marks produced by large
felids are wide, deep, and countable (Haynes
1983:169). Another criterium is the size and
shape of the pits and scores (Domnguez-
Rodrigo & Piqueras, 2003; Domnguez-Rodrigo
& Barba, 2006). Comparative evidence was
gathered from modern carnivores who
existed in Patagonia at the end of the
Pleistocene (e.g., puma [Martn & Borrero,
1997; Martn, 2007]), modern carnivores
that can be considered analogs of extinct
animals (e.g., jaguars, Panthera onca
[Hoogesteijn & Mondolfi, 1992; Sunquist &
Sunquist, 2002] and leopards, Panthera
pardus [Brain, 1981; de Ruiter & Berger, 2000;
Domnguez-Rodrigo & Piqueras, 2003]),
and extinct carnivores (e.g., fossil evidences
attributed to Smilodon fatalis [Van Valkenburgh
& Hertel, 1993) and Homotherium serum
(Marean & Ehrhardt, 1995]). Felids are
accurately classified as flesh specialists, and
for that reason initial consumers (Selvaggio
& Wilder, 2001). Pickering et al. (2004:601)
indicate that leopards do, in fact, leave
tooth-marks on limb bones midshafts (see
also Selvaggio, 1994). Tooth scores made
by felids are perpendicular to the elements
long axis, and shallow but rather sharply
incised (Haynes, 1983:169). A common
pattern in leopards is to produce bone
assemblages characterized by leg bones,
fewer foot bones and skulls, and fewer still
vertebrae and ribs (Andrews & Fernndez-
Jalvo, 1997: 213). Felids can also destroy bones
when the right ecological conditions are met
(Brain, 1981). All considered, a certain degree
of bone breakage is expected from the
activities of felids.
The available evidence of marks
produced by bears was also used (e.g.,
Haynes, 1983; Pinto Llona et al., 2005). It is
346
Taphonomy of Panthera onca

killed by jaguars often have one or two holes
punched through the temporal bone (Sunquist
& Sunquist, 2002:309). Given that the jaguar
can break large mammal bones, we can
assume that the panther was able to break
and consume megamammal bones.
Beyond the marks in the braincase
associated with the classic method of jaguar
hunting, sometimes the prey ends violently
falling to the ground, causing the breakage
of the neck at the first and second cervical
vertebrae. In at least one case the occipital
was also broken (Hoogesteijn & Mondolfi,
1992:69). It is known that occasionally jaguar
bites do not puncture the bones (Schaller &
Vasconcelos, 1978), but crush them to the
point that they looked as if they had been
sawn (Hoogesteijn & Mondolfi, 1992:58).
It is a common pattern of consumption
of large prey to feed only on part of the
carcass, with the rear quarters and vertebral
column many times left abandoned and
unconsumed (Crawshaw & Quigley, 2002;
Sunquist & Sunquist, 2002).
The taxonomic identification is
derived from specific studies of horses
(Alberdi & Prieto, 2000; Alberdi & Prado,
2004; Weinstock et al., 2005), ground sloth
(Roth, 1899, 1904; Nordenskjld, 1996
[1900]; Esteban, 1996; Latorre, 1998; S.
Vizcano and S. Bargo, pers. comm.),
Camelidae (Roth, 1899, 1904; Nami & Menegaz,
1991; Prieto & Canto, 1997; Latorre, 1998),
Macrauchenia patachonica (Nordenskjld,
1996 [1900]), Panthera onca mesembrina
(Roth, 1899, 1904; Cabrera, 1934; Nami &
Menegaz, 1991; Prevosti & Martn, 2008;
A. Currant, pers. comm.), Smilodon populator
(Canto, 1991; Mol et al., 2003; Barnett et
al., 2005), Arctotherium tarijense (Smith-
Woodward, 1899, Prevosti et al., 2003;
Soibelzon et al., 2005; F. Prevosti, pers.
comm.), and Dusicyon sp. (Roth, 1899,
by large and small carnivores. Selvaggio
(1998:192) concluded that [t]he evidence
of defleshing tooth-marks and butchery marks
on midshafts is not significantly altered by a
final bout of feeding by hyaenas. For that
reason, the incidence of felids feeding on
bones should be no problem for recognizing
previous butchery marks.
The more important analog for the
patagonian panther is the jaguar. This is
based on phylogeny (Vizcano & De Iuliis,
2003). Tonni et al. (2003:610) indicate that
patagonian panthers were between 5% and
30% larger than living jaguars. Even though
the jaguar is smaller than the Patagonian
panther, it is the living felid whose canine
teeth are more robust and have a more
powerful bite than those of the other big
cats (Sunquist & Sunquist, 2002:306). The
hunting capabilities of the jaguar are well
recorded. Jaguars can break open the tortoises
tough shell with its strong teeth (Hoogesteijn
& Mondolfi, 1992: 54). Predation on the
Orinoco crocodile (Crocodilus intermedius),
caiman (Caiman crocodilus), pecari (Tayassu
tajacu), white-tailed deer (Odocoileus
virginianus), giant anteater (Myrmecophaga
tridactyla), tapir (Tapirus terrestris), and a
variety of large domestic animals is well
attested (Hoogesteijn & Mondolfi, 1992:55-59;
Kuroiwa & Ascorra, 2002; Scognamillo et
al., 2002; Sunquist & Sunquist, 2002). In
many cases the prey bones are reported as
highly fragmented or crushed. A particularly
important pattern which is characteristic of
jaguars killing their prey is breaking the
skull with a powerful bite, specifically at the
base or the dorsal portion of the neck
(Emmons, 1992; Turner & Antn, 1997;
Schiaffino et al., 2002). The description of
this strategy indicates that it bites through
the animals skull between the ears or
horns. The thick skulls of horses and cattle
347

Martn
The study by Borrero and coauthors
(1997) found that camelids are well
represented by a variety of specimens, while
horses were not well represented, even
though their bones are more resistant to
destruction (also Alberdi & Prieto, 2000;
Martn, 2007). The Lama sp. specimens
include low density bones, which are highly
fragmented and marked by carnivores. The
paucity of horse specimens probably indicates
the selective introduction of its parts (Table 5).
Some horse specimens preserve carnivore
marks (Table 6). It must be stated that the
smaller elements, like carpals, tarsals, and
phalanges are corroded by gastric acids,
suggesting their introduction in carnivore
scats. At least one Mylodon skull is present,
as well as postcranial bones and thousands
dermal ossicles (Table 7). Carnivore marks
are preserved on some specimens (Table 8).
This indicates that large carnivores were
able to transport large bones to restricted
and protected places.
At CLS4 several Mylodon postcranial
bones are recorded, for example an
incomplete shaft of a left humerus, probably
a juvenile (Figure 3). This bone preserves
carnivore marks on the proximal and distal
ends. The distal end is crenulated while the
proximal end displays hinge and spiral
fractures. There is a large pit above the
nutrient foramen. It must be stated that the
reduction of the element to a shaft appears to
be the result of the carnivore actions.
Evidence of carnivore activities was
also found on an undetermined bone assigned
to Mammalia cf. Mylodontinae that exhibits
tooth notches and conchoidal flake scars
(Figure 4).
In general, the number of
carnivore-marked bones found at CLS4 is
not high, but affects elements of different
species. The size of the marks is larger
1904; Caviglia, 1985-1986; Clutton-Brock,
1988; Arroyo, 1998; Latorre, 1998; F. Prevosti,
pers. comm.). A 10-power hand lens, as well
as a 60-power binocular microscope, were
used in the study of the bone surface
modifications.


Results

Among the large herbivores of Pleistocene
Patagonia, Mylodon darwini was the
largest, with an estimated body mass of
1000 kg (Faria et al., 1998). A taphonomic
restudy of the Mylodontinae found at
several sites produced clear evidence of
carnivore marks whose size was much
larger than those produced by pumas
(Puma concolor) (Martn, 2007). Marks
are also important on horse bones, an
animal of about 200 kg of body mass
(Faria et al., 1998), and on camelids of
different sizes.
The basic evidence for the presence
of felid activities at bone assemblages in
Fuego-Patagonia will be introduced here.
The description of selected marked bones,
especially of Mylodon and horse will be the
focus of this presentation.


Lago Sofa 4 cave

The faunal remains obtained at CLS4
belong were attributed to Mammalia,
Camelidae, Vicugna sp., Lama guanicoe,
Mylodon darwini, Hippidion saldiasi,
Smilodon populator, Panthera onca
mesembrina, Hippocamelus bisulcus,
Hippocamelus bisulcus, Lagidium viscacia,
Carnivora, Felidae, Canidae, Rodentia,
Aves, and undetermined (Borrero et al.,
1997).
348
Taphonomy of Panthera onca

Table 8. Sofa 4 cave. Carnivore marks on Mylodontinae specimens.
Table 7. Sofa 4 cave. Mylodontinae. Number of identified specimens (NISP) for each skeletal element.
Table 6. Sofa 4 cave. Carnivore marks on horse specimens.

Punctures Pits Furrowing
Phalanx 1, prox. + shaft

1

Phalanx 1, prox

1

Phalanx 2, complete 1

1
Phalanx 3, prox.

1

Total 1 3 1
Table 5. Sofa 4 cave. Hippidion saldiasi. Number of identified specimens (NISP) for each skeletal element.
Total
Carpals 3
Metatarsal 1
Metapodial 2
Distal sesamoid 1
Phalanges 15
Total 22

Mylodon
darwini
Mylodontinae cf.
Mylodon
Mammalia cf.
Mylodontinae Total
Skull 1 1
Teeth 3 3
Vertebra 1 5 6
Ribs 2 2
Pelvis 2 1 3
Scapula 1 1
Humerus 1 1
Phalanges 4 4
Dermal ossicle 2685 2685
Undetermined 2 9 11
Total 3 2697 17 2717

Punctures Pits Scores Crenulated Furrowing
Skull, yugal

1

Vertebra, body + arc

1

1

Vertebra, body

1
Humerus, shaft

1

1

Undetermined, frag. 1

Undetermined, frag.

1

1

Total 1 3 1 3 1
349

Martn
Figure 3. Shaft of a Mylodon humerus.
Figure 4. Undetermined fragment assigned to Mammalia
cf. Mylodontinae with notches and flake scars on the
internal view of the rim.
Figure 5. Medial view of Mylodon darwini left mandible
with punctures and associated damages.
Figure 6. Punctures and associated fracture line. Note the
crenulated rim with in situ flake scars.
350
Taphonomy of Panthera onca

produced today by jaguars hunting large
prey (Martn, 2007). As already mentioned
this is a typical pattern of modern jaguars.
This evidence points to fossil panthers as
the killing agents. Some bones identified as
Mylodontinae, including dermal ossicles,
preserve gastric corrosion. This evidence is
in line with the existence of carnivore scats
recovered at the cave at the end of the 19
th

century that include fragments of Mylodon
skin with dermal ossicles and hair. Ten
panther bones were also recovered (Table
12), two of which preserve carnivore marks
(Table 13).
One of the marked Mylodon
darwini bones is an adult left mandible with
four teeth (Figure 5) and well-preserved soft
tissues (Lehmann-Nitsche, 1899; Roth, 1899).
Marks on this bone were described in great
detail by Lehmann-Nitsche (1899). There are
large and shallow punctures of about 12 mm in
diameter. Also, there is a break line starting
from the two main punctures. One of them
cross-cuts the mandible toward the gnawed
inferior rim (Figure 6), while the other joins
one of the punctures with the crushed zone
below the third tooth (Figure 7).
Figure 8 presents a nearly complete
and very well-preserved adult right tibia of a
Mylodontinae cf. Mylodon with abundant
soft tissues (Roth, 1899:432). The medial
condyle of the proximal tibia presents scooping
out, furrows and pits (Figure 9), while the
articular face presents several pits on bone
and cartilage (Figure 10). There is also one
score over the cartilage. The size of the pits
varies between 4.34 mm and 9.05 mm, denoting
carnivores larger than pumas (Martn, 2007).
The marks just presented, as well
as many others, were described by
Lehmann-Nitsche (1899) as the result of
human activities. Using the criteria presented
above, I attribute these marks to the actions of
than those produced by living pumas on
guanaco or horse bones (Martn & Borrero,
1997; Martn, 2007). Together with the limited
number of punctures and bone fragmentation,
this evidence points to fossil panthers as the
destructive agents. A conclusion that is
reinforced by the presence of three panthers
at the site, including one juvenile and one cub.


Milodn cave

Several Mylodontinae and Hippidion saldiasi
skeletal parts are present in the Hauthal
collection (Tables 9, 10). Previous studies
of the bone collections produced by the
different expeditions rarely mentioned the
presence of carnivore marks (Lehmann-
Nitsche, 1904). We recently reanalyzed the
sample obtained by Hauthal at this site
(Hauthal, 1899; Roth, 1899, 1904) and found
that large carnivore marks are present on
many of the herbivore and panther bones
and in one horse bone, a maxilla. Thirty-six
Mylodontinae bones display large carnivore
marks, including skulls, scapulae, tibiae,
vertebrae, ribs, and several undetermined
bones (Table 11). Pits predominate, followed
by punctures and scores, which are less
abundant. The punctures are frequently located
on the rim of the bone, producing notches.
There are cases at which the strength of the
bite collapsed the bone, creating attached
small flakes. There are several bone flakes
and bone fragments displaying negatives of
hinge fractures resulting from the action of
the carnivores. The shape of the punctures
and the low frequency of scores and furrows
indicate a felid pattern (Haynes, 1983). The
size of punctures and pits is larger than that
of living pumas. The skulls and mandibles,
attributed to Mylodon darwini (Esteban, 1996),
preserve marks which are similar to those
351

Martn
Table 10. Milodn cave. Hippidion saldiasi. Number of
identified specimens (NISP) for each skeletal element.
Table 9. Milodn cave. Mylodontinae. Number of identified specimens (NISP) for each skeletal element.
Mylodon darwini
Mylodontinae cf.
Mylodon
Mammalia cf.
Mylodontinae Total
Skull 1 20 10 31
Stylohyale 3 3
Maxilla 5 5
Mandible 4 1 5
Teeth 5 2 7
Vertebra 6 5 11
Ribs 12 2 14
Clavicle 1 1
Scapula 9 3 12
Humerus 1 1
Carpals-tarsals 3 3
Tibia 3 3
Patella 1 1
Sesamoid 1 1
Phalanges 6 6
Claw 6 2 8
Dermal ossicle 166 166
Undetermined 3 391 394
Total 15 244 413 672
Total
Skull 1
Mandible 1
Teeth 3
Vertebra 1
Carpal 3
Metacarpal 2
Metatarsal 1
Phalanges 6
Hooves 1
Total 19
352
Taphonomy of Panthera onca

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353

Martn
Figure 7. Punctures and associated fracture line connecting
with crushed area. Note the presence of the well-preserved
gum.
Figure 8. Anterior view of Mylodontinae cf. Mylodon
right tibia.
Figure 9. Medial view of Mylodontinae cf. Mylodon
right tibia with scooping out. The white line bordering
the damaged area was probably painted by Lehmann-
Nitsche (1899) when describing the material.
Figure 10. Mylodontinae cf. Mylodon proximal right
tibia with pits and score on cartilage and bone. The white
line bordering the damaged area was probably painted by
Lehmann-Nitsche (1899) when describing the material.
354
Taphonomy of Panthera onca

Figure 11. Left femur and left tibia with scooping out.
Figure 12. Antero-lateral view of left femur
with scooping out and two large punctures.
Figure 13. Lateral view of left tibia with
scooping out and puncture on proximal end.
Figure 14. Medial view of left tibia with
detail of puncture.
355

Martn
hinge fractures. Pits and a large puncture are
present in the lateral view of the condyle
(Figure 12). In the posterior view there is
scooping out associated with large pits as
well as a large notch and hinge fractures on
the rims.
The tibia is also incomplete and
has the proximal end removed. Evidence of
scooping out are present on the anterior,
lateral, and posterior views (Figure 13). The
anterior view also presents pits and scores
on the compact bone near the removed bone
area, while in the postero-medial view
below the superior fibular articular facet
there is a pit and a large puncture (Figure
14). Both bones, recovered at a place
interpreted as a carnivore hunting site, may
reflect conditions under which felids produced
higher numbers of marks.


Dos Herraduras rockshelters

The Mylodontinae skeletal parts include one
femur, several ribs, and hundreds of dermal
ossicles (Table 14). The complete left femur
of Mylodontinae is very well-preserved
(Figure 15). Six punctures were identified
around the head of the proximal end, below
carnivores. Together with the presence of
large felid scats, the marks on the skulls and
the presence of panthers-including a young
individual-I suggest that these marks were
produced by fossil panthers.
The intensity of gnawing damage
recorded on two bones from Milodn cave
that are stored at the MACN under catalog
number 6866 is larger than that observed for
most of the analyzed bones. For that reason
I will describe those bones here. They are
one left femur and one left tibia, probably
from the same individual, as can be judged
from their size, anatomical coherence and
the location of the carnivore marks (Figure
11). The femur is incomplete, comprising
the distal end plus most of the shaft. The
proximal end appears to be removed by
carnivore gnawing (Figure 11). The rim is
irregular and highly polished. In the anterior
view there are a few pits, while in the
posterior and lateral views pits and scores
are abundant. On the distal end there are
missing portions, and scooping out is
present above the lateral condyle on the
anterior, posterior, and medial views. In the
anterior view of the compact bone there are
pits, scores, and one large puncture (Figure
12), while the rim exhibits angular and
Table 12. Milodn cave. Panther. Number of identified specimens (NISP) for each skeletal element.

P. onca
mesembrina
Felidae cf. P. onca
mesembrina Total
Teeth 2 2
Mandible 2 2
Pelvis + sacrum 1 1
Humerus 1 1
Metatarsal 1 1
Metapodial 1 1
Phalanges 2 2
Total 7 3 10
356
Taphonomy of Panthera onca

Table 14. Dos Herraduras Rockshelter.
Mylodontinae. Number of identified specimens
(NISP) for each skeletal element.. Dermal ossicles
were not quantified.
Table 13. Milodn cave. Panthera onca mesembrina specimens with tooth marks.
puncture 9.00 mm by 7.60 mm, a proximal
rib with a puncture 8.13 mm by 4.79 mm
(Figures 17 and 18), a proximal radius
epiphysis with large carnivore pits and
scores (Figure 19), and a femur medial
condyle with pits. The rest of the tooth-
marked bones include small elements like
carpals and tarsals. A fragment of a right
mandible of a large feline and a fragment of
a left mandible of a young P. o. mesembrina
were also found (Prevosti & Martn, 2008).
The latter preserves a large puncture (4.9 x
4.2 mm) attributable to the action of a large
carnivore. The presence of a young panther
is concordant with the interpretation of the
site as a panther den (Martn, 2007). Even
when bear specimens were found at the site
(two teeth, one atlas), the frequency, shape,
and size of marks are similar to those
produced by felids.
Summing up, human utilization of
the cave at the end of the Pleistocene was
very sporadic and there is no record of any
interaction with carnivores.

the fusion line (Figure 16). None of the
punctures was located on the epiphysis
itself. There are two groups of two punctures
and two isolated punctures. Collapsed bone
is present and two of the punctures indicate
an oblique angle of entry. The size of the
punctures as well as the size of the femur on
which they were found implicate a large
felid. The pattern of the punctures is the
same produced by pumas on the proximal
end of guanaco (Lama guanicoe) femora
(Borrero et al., 2005) and similar to damage
produced by felids on proximal femora of
different species (Haynes, 1983; Martn &
Borrero, 1997; Domnguez-Rodrigo, 1999).
The observed damage on this femur is
consistent with the interpretation of the
bone accumulation resulting from an adult
individual that was partially consumed by a
large extinct felid near the shores of a lake
(Borrero, 2001).


Chingues cave

At Chingues cave, 752 Mylodontinae dermal
ossicles were found together with a single
phalanx (Table 15). Only four of the
ossicles preserve gastric corrosion. The horse
bones are more abundant in comparison
with other sites (a total of 285 specimens,
Table 16) and some of them display
carnivore marks (Table 17). Eleven of these
bones are marked by large carnivores. The
sample includes a distal metapodium with a
Portion Punctures
Pits Scores
Furrows
Humerus dist. + shaft 1
1 1

Pelvis + sacrum incomplete 1
1

1
Total 1 1 1 1

Total
Femur 1
Ribs 4
Total 5
357

Martn
Figure 15. Anterior view of left femur with large
puncture in the head.

Figure 16. Posterior view of left femur with
large puncture indicated by arrow.
Figure 17. Horse rib with a large carnivore puncture. Figure 18. Close up view of horse rib with a large carnivore
puncture.
358
Taphonomy of Panthera onca

Pleistocene bones. The morphology of the
marks and the presence of bones of panther
are concordant with this interpretation.


Fell cave

The list of taxonomic identifications in the
analyzed sample is presented in Table 18
(see also Tables 19 and 20). A recent review
of a previously unstudied bone collection
from the site shows that Mylodon was not
only processed by humans but also by large
carnivores (Martn, 2007). There is a MNI of
three for Mylodon, including two young, one
of them probably a newborn (Sergio Vizcano,
pers. comm.), and an adult. The number of
bone specimens with marks is low (Table
Puma cave

The bone assemblage is characterized by the
presence of carnivore fibres and scats.
However, preservation is not always good,
especially in some chambers where the
bones were exposed to circulating water and
other processes.
Puma cave is one of the few sites,
together with CLS4, where well-preserved
bones from extinct fauna were recovered mixed
with modern fauna on the surface. The temporal
range recorded on the surface of the cave is
about 10,300
14
C years, constituting an extreme
example of averaged faunas on a surface. All
the bones of extinct fauna were found isolated,
suggesting that the carcasses were not
transported complete to the cave. The size
of the passage, about 60 cm in height, prevents
adult medium-sized mammals from finding
their way to the interior.
The bone assemblage is only
preliminarily studied, but at least a
Camelidae radio-ulna presents pits that can
be attributed to the activities of a large
carnivore. Bones with marks of smaller
carnivores were also identified. The cave
can be interpreted as a den that was used by
more than one carnivore species, but it is
not yet clear what specific carnivore agents
caused the bone accumulations. The panther
is a good candidate to explain most of the
Table 15. Chingues cave. Mylodontinae.
Number of identified specimens (NISP) for
each skeletal element.
Total
Phalanx
1
Dermal ossicle
752
Total
753
Table 16. Chingues cave. Hippidion saldiasi and
Mammalia cf. H. saldiasi. Number of identified
specimens (NISP) for each skeletal element.
Total
Skull 22
Teeth 38
Mandible 8
Vertebra 29
Pelvis 1
Ribs 3
Scapula 1
Humerus 5
Radius-ulna 2
Radius 4
Carpals-tarsals 50
Femur 14
Tibia 8
Fibula 7
Metatarsal 3
Metapodial 16
Patella 3
Sesamoids 38
Phalanges 33
Total 285
359

Martn
Figure 19. Horse radius epiphysis with pits and scores.
Figure 20. Mylodontinae cf. Mylodon fragment of pelvis
with crenulated rim, pits and scores.
Figure 21. Close up view of Mylodontinae cf. Mylodon
fragment of pelvis with pits, scores and polished rim.
360
Taphonomy of Panthera onca

Table 17. Chingues cave. Carnivore marks on horse specimens.
while carnivore tooth-marks are not abundant.
The sequence of production of the tooth-
and cutmarks is not known.


Tres Arroyos rockshelter

The list of animals found at this shelter
includes Mylodontinae, Hippidion saldiasi,
Panthera onca mesembrina, Dusicyon avus,
D. culpaeus, Camelidae including Vicugna
sp., and Lama guanicoe (Prieto & Canto,
1997; Latorre, 1998; Alberdi & Prieto,
2000; Massone, 2004) (Table 22). It is
possible that Lynchailurus colocolo was
21), but constitute good evidence for the
activities of a large carnivore at Fell cave. A
fragment of Mylodontinae cf. Mylodon pelvis
found at Fell cave presents marks (pits and
scores on compact bone). Smooth crenulated
marks are present on the edges of the bone,
and furrows on the trabecular bone. The
glenoid cavity was removed and there is a
hinge fracture (Figures 20 and 21). The size
of the marks as well as its paucity were
interpreted as indicating the activities of a
felid.
The evidence for human use of Fell
cave at the end of the Pleistocene is
substantial, including cutmarks (Table 4),

Punctures Pits Scores

Crenulated Furrowing
Atlas, frag.

1
Rib, frag. 1

Vertebra, apophysis 1

Scapula, frag.

1
Radius, epiphysis prox.

1 1

1
Radius, dist. 1

1

Carpal, complete

1 1

Carpal, complete 1 1 1

Pelvis, cotiloid cavity

1 1

Femur, dist.

1

Patella, complete 1

Tibia, shaft

1 1

Metapodial, prox. shaft 1

Metapodial, dist. + shaft 1

1

Proximal sesamoid, complete

1

Proximal sesamoid, complete

1

Phalanx 1, dist + shaft

1

Phalanx 2, complete

1 1

Total 7 9 8 1 3
361

Martn
Table 19. Fell cave. Mylodontinae. Number of identified specimens (NISP) for each skeletal element. Samples
obtained by John Fell (19521959), Junius Bird (1969-1970) and Pedro Crdenas (1999).
Table 18. Fell cave. Number of identified specimens (NISP) and percentage for each taxon.
Taxon NISP %
Camelidae 3 3,1
Lama guanicoe 1 1
Hippidion saldiasi 50 51,5
Hippidion saldiasi? 2 2
Mylodon darwini 1 1
Mylodontinae cf. Mylodon 18 18,5
Mylodontinae? 1 1
Mammalia 19 19,6
Mammalia cf. Mylodontinae 2 2
Total 97 100

Mylodon darwini Mylodontinae cf.
Mylodon
Mammalia cf.
Mylodontinae
Mylodontinae?
Skull

1

Maxilla 1

Mandible

1

Vertebra

5

Ribs

1 2

Pelvis

3

Ulna

1

Femur

1

Tarsals

2

1
Dermal ossicle

3

Subtotal 1 18 2 1
Total 22
362
Taphonomy of Panthera onca

Table 21. Fell Cave. Carnivore marks on Mylodontinae and horse specimens.
parts (Table 23). Carnivore marks were
identified on horse bones (Table 24). A
complete Hippidion unfused thoracic vertebra
preserves abundant carnivore marks. The
neural apophysis is heavily gnawed, with
several overlapping small pits, similar to
those produced by foxes. Polish on the
epiphysis is another result of the carnivore
activities. There is a large puncture below
the neural apophysis. Three punctures and
furrows are present on the transverse
apophysis. The size and location of the
marks indicate that the animal was initially
consumed by a large carnivore and later
scavenged by canids.
On the other hand, a partial unfused
proximal epyphisis of a right horse humerus
presents large carnivore marks, which were
initially recognized by Mengoni Goalons
(1987). They are located mainly on the
posterior view of the head, and include pits
and wide parallel scores (Figure 22). Pits
and scores are also found on the articular
face of the humerus with the scapula.
There are furrows in the posterior view of
the medial tuberosity. A Panthera onca
mesembrina metatarsal presents small
superimposed pits and scores. They are
similar to those produced by small canids. It
must be remembered that D. avus and D.
culpaeus are present in the bone assemblage.
There is no basis on which to claim
interaction between large carnivores and
also present in the Pleistocene, although its
stratigraphic position does not support this
(Prevosti, 2006).
The dermal ossicles and highly
fragmented bones attributable to Mylodontinae
do not present carnivore marks. Horse is
represented by a large variety of skeletal
Table 20. Fell cave. Hippidion saldiasi. Number
of identified specimens (NISP) for each skeletal
element. Samples obtained by John Fell (1952
1959) and Junius Bird (1969-1970).
* Hippidion saldiasi?

Total
Skull 2
Teeth 16
Mandible 5
Vertebra 4
Rib 1
Humerus 1
Radius 1
Radius 1
Metacarpals 3
Carpals-tarsals 4
Femur?* 1
Tibia 4
Patella 1
Metapodial 1
Sesamoids 5
Phalanges 2
Total 52
Taxon Element Pits Scores Crenulated
Mylodontinae cf. Mylodon Vertebra, complete 1 1

Mylodontinae cf. Mylodon Pelvis, frag. 1 1 1
Hippidion saldiasi Skull, occipital condyle 1 1

Hippidion saldiasi Radius, ep. dist 1 1

Total

4 4 1
363

Martn
Table 24. Tres Arroyos. Carnivore marks on horse specimens. * Mammalia cf. Hippidion saldiasi.
Taxon NISP
layer V
% NISP
layer VI
% NISP
total
Hippidion saldiasi 16 34,0 3 33,3 19
Mammalia cf H. saldiasi 6 12,8 1 11,1 7
Panthera onca mesembrina 1 2,1 -

1
Mammalia cf. Mylodontinae 2 4,2 - - 2
Mammalia 22 46,8 5 55,5 27
Total 47 100 9 100 56
Total
Skull 1
Teeth 3
Vertebra 5
Ribs 4
Humerus 1
Carpals-tarsals 3
Tibia 1
Patella 1
Sesamoids 5
Phalanges 2
Total 26
Table 23. Tres Arroyos 1. Hippidion saldiasi and Mammalia cf. H. saldiasi, Layers V - VI. Number of identified
specimens (NISP) for each skeletal element.
Table 22. Tres Arroyos 1, Layers V-VI. Number of identified specimens (NISP) and percentage for each taxon.

Punctures Pits Scores Furrows
Thoracic, complete 1 1 1 1
Rib, frag.*

1

Humerus, epiphysis

1 1 1
Tarsal, frag.

1

Total 1 3 3 2
364
Taphonomy of Panthera onca

patagonian panthers had important destruction
capabilities.
It is also necessary to consider bears
as potential contributors to the production
of marks on the bones. Scavenging by bears
should have produced more marks in the
bones. The recorded punctures in general
are not square, rectangular or flat-bottomed
as bear marks are. Scores on shafts rarely
are short and they usually are not parallel.
In fact, most of the analyzed bones display
less conspicuous marks. Then, bears do not
appear as important accumulators. It is a
fact that the range of gnawing damage that
can be attributed to fossil felids is not well
known. Using different sources (Brain, 1981;
Marean, 1989), it was previously argued that
Smilodon populator did not mark or
fragment the bones of their prey (Borrero et
al., 1997). However, the bones collected by
Homotherium serum at Friesenham cave,
Texas, USA, displayed carnivore marks
humans at Tres Arroyos. The human imprint
is very strong at this site, where hearths,
lithics, and bone tools were found, while the
large carnivore activity is restricted to some
horse bones.


Discussion

The list of tooth-marked bones from each of
the mentioned sites is not high. However,
there is enough evidence to sustain that
carnivores capable of breaking megamammal
bones, probably felids, existed at the end of
the Pleistocene in Fuego-Patagonia. Some
cases, like the pelvis recovered at Fell cave
and the tibia and the femur from Milodn
cave stored at the MACN (#6866) present
more damage than that usually associated with
felid activities. However, the mentioned
tooth-marked bones at several sites indicate
that there is good reason to sustain that the
Figure 22. Furrows and scores on a horse epiphysis of a humeus.
365

Martn
components. Borrero et al. (1997) found no
evidence of gastric corrosion in the 4246
dermal ossicles, suggesting that they did not
reach the cave in carnivore excrements. The
transport of Mylodon bone segments that
included parts of the hide is a more
parsimonious explanation. Access to the
cave was a limitant for the introduction of
complete animals. I suggest that the carcass
was processed and disarticulated by carnivores,
and some segments were later selected for
transport to the cave. This is probably the
case also for the dermal ossicles at CDLCH,
for which the evidence indicates that only a
limited number reached the cave through
now-desintegrated carnivore scats. The presence
of Mylodon bones at Puma cave also imply
transport by carnivores. The size of the
access to the dark chamber where these
bones were found is too small to allow these
large animals to find their way inside.
Mylodon remains, both with and without
carnivore marks, are abundant at Milodn
cave, but many individuals were probably
hunted at this huge herbivore den, where
they were at least partially processed
(Martn, 2007). The Mylodon bones found
at the Dos Herraduras rockshelters can also
be considered a residual assemblage at a
place at which one individual was probably
hunted and processed by a large felid.
We must remember that bones of
horse and camelids, smaller animals of about
200 and 100 kg of body mass (Faria et al.,
1998), are usually highly fragmented. They also
display carnivore marks that appear to be
the result of large felids. In at least two of the
sites - Milodn cave (Table 13) and CDLCH -
there are carnivore marks on panther bones.
Generally speaking, the number of carnivore
marks at each of the sites - including those
postulated as dens - is not abundant, which
is precisely one of the properties of felid
(Marean & Ehrhardt, 1995), a fact that suggests
that sabertooth cats can also be candidates
to produce marks on the patagonian bones.
However, Marean and Ehrhardt (1995:528)
utilize the little damage recorded on the
bones to argue that Homoherium was the
main gnawing agent, instead of dire wolf
(Canis dirus). They report that tooth-marks
rarely penetrated the bone cortical surface
or resulted in bone fracture (Marean &
Ehrhardt, 1995:529). In addition, the presence
of sabertooth cats at patagonian caves is far
from abundant. This combined information
suggests that sabertooth cats were not important
accumulators of these bone assemblages.
Several experiments with African
carnivores indicate that low percentages of
tooth-marked bones are associated with early
access of humans to a carcass (Blumenschine
1988, 1995). However, variation across
experimental, ethnoarchaeological and
archaeological samples indicates that this
pattern can be complicated (Capaldo, 1998;
Selvaggio, 1998; Domnguez-Rodrigo &
Barba, 2006; Faith, 2007). It must be
accepted that the degree of utilization of a
carcass or a bone should be variable in
accordance with ecological conditions (Haynes,
1980b; Borrero et al., 2005), an issue that
still needs to be fully investigated in South
America.
With the exception of Fell cave
(where the deposition of small-sized juvenile
Mylodon bones was mediated by humans)
and Milodn cave, at most of the known
sites Mylodon is represented by small
elements (phalanges, carpals, tarsals, dermal
ossicles). The presence of a relatively
complete humerus, a scapula, a pelvis,
vertebrae, ribs, and a skull together with a
high number of dermal ossicles at CLS4
inform on a difference in comparison with
other caves characterized by important den
366
Taphonomy of Panthera onca

As for the use of caves by modern jaguars it
must be noted that these shelters are not
abundant at most of their current distribution.
However, at places where caves are available
they make use of them and transport their prey
(Schaller & Vasconcelos, 1978; Hoogesteijn
& Mondolfi, 1992). As for the size and
behaviour of Mylodon sp., two points must
be kept in mind. First, the study of the
skulls of three individuals of Mylodon
darwini recovered at Milodn cave display
marks which not only parallel the shape and
size of those that are attributed to panthers,
but also preserve these marks in the same
pattern produced by jaguars today when
they hunt large prey (Martn, 2007). Second,
it has been shown that a limited number of
Mylodontinae bones find their way into
CLS4, and CDLCH and Puma cave for that
matter, and were recovered in dark chambers
whose access was not possible for live
ground sloths. Moreover, studies by Paul
Martin suggest that ground sloths were not
necessarily a difficult prey for human
hunters (Martin, 2005:139). Indeed, the case
for carnivore hunters remains open, but
Prevosti & Vizcaino (2006:416) indicated
that the diet of P. onca would include large
mammals of up to 600 kg. Given the larger
size of the fossil jaguar ... this species could
have fed on preys that were somewhat larger
than those hunted by living individuals. It is
also highly probable that it preyed upon
juvenile megamammals. The sum of evidences
is at odds with the interpretation of Tonni et
al. (2003) of CLS4. Even when the bone
sample obtained at CLS4 is large, the remains
of megamammals are relatively scarce, and
the weathering -which is not present at the
sample- cannot be used to explain their
absence. The representation of a limited
selection of bones at the cave, plus the fact
that the herbivores could not have entered
activities (Haynes, 1983; de Ruiter & Berger,
2000). Another pattern is that spiral and
hinge fractures are frequently associated with
carnivore marks.
Summing up, the selective presence
of Mylodontinae bones at several sites
with dark chambers and access difficulties
is best explained as the result of transport.
The majority of the marks on ground sloths
are on dense cortical bone. The degree of
bone fragmentation as well as these marks
indicate the activities of large felids. The
panther is the only felid which is well
represented at many of the sites, and can
be considered the transporting agent.
It must be remembered that the
faunal evidence from two sites at the Pali-
Aike Lava Field was used to infer the
presence of felids in the region before any
large carnivore bone was found. Effectively,
Fell cave included Hippidion saldiasi bones
with marks that were attributed to large
felids (Borrero & Martn, 1996). San Romn
et al. (2000) also inferred large felid activities
at Chingues cave. At both sites the evidence
was indirect - only the marks on the bones - but
recent research demonstrated the presence
of Panthera onca at two sites at Pali-Aike,
Puma cave, and Chingues cave, thus supporting
those inferences (Martn, 2007). Direct late
Pleistocene radiocarbon dates of Panthera
onca mesembrina are available for Sofa 4 cave,
Medio cave, and Tres Arroyos rockshelter
(Table 2).
The interpretation of CLS4 as a
panther den was criticized by Tonni et al.
(2003) on the basis of: a) Panthera onca
does not transport prey to caves today, and
b) the size and behaviour of Mylodon sp.
makes it a difficult prey for a solitary
carnivore. They suggest that the presence of
Mylodon sp. at the site (CLS4) is a result of
its use of the cave (Tonni et al., 2003:611).
367

Martn
appears to have been panther hunting
grounds, while the evidence at Fell cave,
Tres Arroyos rockshelter, or Medio cave is
clearly different. In the latter three sites there
is archaeological evidence that indicates
that humans were the main occupants, in
contrast to the cases of CDLCH and Milodn
cave with only sporadic human use, and
CLS4, Dos Herraduras and Puma cave which
appear to be purely paleontological. Then,
interaction between humans and carnivores
appears to have been minimal in Fuego-
Patagonia.
In conclusion, the study of faunal
assemblages in Fuego Patagonian sites should
take into account the existence of bone-
crunching felids. The presented evidence indicates
the possibility of interactions between the
first human settlers and large carnivores.
This discussion shows that these carnivores
not only accumulate bones at sites which
before or after where used by humans, but
that they also damage the bones. The felids from
Pleistocene Fuego-Patagonia were perfectly
capable of fracturing, marking, and consuming
large mammal and megamammal bones.


Acknowledgements

To Manuel Domnguez-Rodrigo for his
invitation to participate in this issue of the
Journal of Taphonomy. To Luis A. Borrero
and Francisco Prevosti for their critical
comments on this paper and support. To
Marcelo Reguero and Lucas Pommi, curators
of the MLP collections and to Alejandro
Kramarz, curator of the MACN collections,
for their support. To Patricio Moreno who
share with me his unpublished radiocarbon
date for Panthera onca mesembrina. To
Ramiro Barberena, Daniel Here and
Azrahel Martinez and for their help with the
the cave on their own constitute good reasons
to suggest that this is a case of selective
transport by carnivores (de Ruiter &
Berger, 2000).

Conclusion

Bears and sabertooth cats are also members
of the carnivore guild of Fuego-Patagonia,
and thus can be implicated in the
accumulation and processing of at least part
of the bone assemblages examined here.
However, Panthera onca mesembrina appears
as one of the more ubiquitous large carnivores
during the end of the Pleistocene, as well
as the only carnivore represented by several
age groups. Its presence at most of the sites
indicates its frequent use of caves and, judging
from the number of young individuals, part
of this use was related to maternal/birth
denning. The presence of bears and sabertooth
cats is, on the other hand, more restricted
in number of sites and number of bones.
Moreover, the high fragmentation of the
bone assemblage as well as the recorded
marks point towards panthers as the main
agent of accumulation.
I believe that the available evidence
is inclined towards the interpretation of
CLS4 as a den. A similar pattern was found
at Puma cave and Chingues cave, where
megamammals are less abundant and
dominated by dermal ossicles. This evidence,
together with the fact that jaguars transport
bones to caves (Hoogesteijn & Mondolfi,
1992), and the presence of young panthers, can
be used to sustain that the best explanation
for sites like CLS4, CDLCH and Puma cave
is that they were panther dens, places where
this carnivore accumulated prey bones.
Activities of panthers are recorded at other
sites as well. One of them, Milodn cave,
368
Taphonomy of Panthera onca

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To Mauricio Massone, Alfredo Prieto, Pedro
Crdenas, Manuel San Romn and Flavia
Morello who provided information on the
sites that they have excavated. To Mauricio
Quercia for facilitating access to the Fell
cave collection stored at the Museo Regional
of Punta Arenas. I acknowledge to the reviews
by S. Capaldo, C. Egeland and M. Domngez-
Rodrigo who helped to improve the paper. I
finally thank the CEQUA Foundation and
Proyecto FONDECYT 1070709 for its support
for my research.


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