bone crunching felids at the end of the pleistocene in fuego-patagonia, chile

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Martín

Article JTa072. All rights reserved. *E-mail: [email protected]

2008 Journal of Taphonomy

PROMETHEUS PRESS/PALAEONTOLOGICAL NETWORK FOUNDATION (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. Martín*

Fundación 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

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Taphonomy of Panthera onca

Materials Information recovered at several paleontological and archaeological caves is revised in this paper. Lago Sofía 4 cave (CLS4) is a dark cave, at Ultima Esperanza, Chile (51º 31’ 54’’ S, 72º 34’ 12’’W) excavated by Alfredo Prieto and Pedro Cárdenas. 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 19th century (Roth, 1899; Nordenskjöld, 1996 [1900]). The finding of large carnivores like bears (Arctotherium tarijense) and panthers (Panthera onca mesembrina) (Roth, 1899, 1904; Smith Woodward, 1900; Nordenskjöld, 1996 [1900]) in late Pleistocene contexts at Milodón 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 Milodón cave bone assemblage described marks on Mylodon bones -most of which today we assign to extinct cats (Martín, 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 19th century or early in the 20th 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.

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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.

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Table 2. Selected taxon dates for carnivores and herbivores of Fuego-Patagonia.

Site T

axon E

lement

Age (B

P) L

ab Source

Sofía 4 cave M

ylodon V

ertebra 11.590 ± 100

PITT-0940 Prieto, 1991

Sofía 4 cave M

ylodon D

ermal ossicle

13.400 ± 90 A

A-11498

Borrero et al., 1997

Sofía 4 cave Panthera onca m

esembrina

Mandible

10.840 ± 60 G

X 31643

Patricio Moreno, pers. com

m.

Milodón cave

S. populator R

adius 11.265 ± 45

Oxa-13717

Barnett et al., 2005

Milodón cave

S. populator R

adius 11.420 ± 50

Oxa-14457

Barnett et al., 2005

Milodón cave

Mylodontinae

Dung

10.200 ± 400 Sa-49

Emperaire &

Laming, 1954

Milodón cave

Mylodontinae

Bone

13.630 ± 50 B

eta-164895 B

orrero, pers. comm

. M

edio cave Panthera onca m

esembrina

Mandible

11.410 ± 80 U

a-24687 This paper

Dos H

erraduras 3 M

ylodontinae R

ib 12.825 ± 110

AA

-12574 B

orrero & M

assone, 1994 D

os Herraduras 3

Mylodontinae

Rib

11.380 ± 150 LP-421

Borrero et al., 1991

Fell cave M

ylodontinae cf. Mylodon

Pelvis 10.295 ± 65

Ua-34249

This paper C

hingues cave M

ylondontinae D

ermal ossicle

12.165 ± 80 U

a-32861 This paper

Chingues cave

Hippidion saldiasi

Hum

erus 11.990 ± 90

Ua-24685

This paper C

hingues cave R

heidae Tibia-tarso

10.165 ± 70 U

a-24684 This paper

Chingues cave

Hippidion saldiasi

Phalanx 11.210 ± 50

Beta-147744

San Rom

án et al., 2000 Pum

a cave A

rctotherium tarijense

Femur

10.345 ± 75 U

a-21033 M

artín et al., 2004 Pum

a cave Lam

a sp. R

adio-ulna 11.575 ± 80

Ua-21035

Martín et al., 2004

Tres Arroyos 1

Panthera onca mesem

brina M

etatarso 11.085 ± 70

Oxa-9248

Massone &

Prieto, 2004 Tres A

rroyos 1 H

ippidion saldiasi B

one 12.540 ± 70

Beta-123152

Borrero, 2003

Tres Arroyos 1

Hippidion saldiasi

Bone

10.685 ± 70 O

XA

-9247 M

assone & Prieto, 2004

Tres Arroyos 1

Dusicyon avus

Mandible

10.575 ± 65 O

XA

-9245 M

assone & Prieto, 2004

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Martín

extinction of the Pleistocene fauna (Borrero, 1997). Milodón cave is a huge cave of about 150 m by 180 m located on the Cerro Benitez flanks (51º 33’ 54’’S; 72º 37’ 13’’W). 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 Milodón cave (Nami, 1987; Nami & Menegaz, 1991, Nami & Nakamura, 1995). We already mentioned that carnivore bones were identified at Milodón cave before the end of the 19th century, as part of the rich bone assemblage excavated by Hauthal and Nordenskjöld (Hauthal, 1899; Roth, 1899, 1904; Smith Woodward, 1900; Nordenskjöld, 1996 [1900]). There were new excavations throughout the 20th century that produced abundant bones. The recorded species are Mylodon darwini, Hippidion saldiasi, Panthera onca mesembrina,

an excavated sample of 3 m2. 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 Milodón 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 Sofía 4 cave.

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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° 44’31’’W). 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 Román 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 Milodón cave. Judging from the available written sources, Medio cave can be suggested as the origin of that collection (Martín, 2007). In this paper I will present evidence obtained in a restudy of the Hauthal collection (1899-1900) stored at the Departamento Científico Paleontología 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 Milódon cave (51º 33’ 36’’ S; 72º 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 m2, 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.

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Martín

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 (Martín, 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 m2 and Junius Bird in 1969-1970 from an unspecified area. In addition, we have studied some bones recovered by Pedro Cárdenas 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 (Martín, 2007) (Table 4).

Tres Arroyos is a small site of about 6 m by 4 m located at Cerro de los Onas (53º 23’ S, 68º 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 Román 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 m2 is presented here. All the sediments were screened. Puma cave is a large and complex site located at the Estancia “Brazo Norte”, (52º 1’ 12,6’’ S, 69º 58’ 41,2’’ W), about 2 km from the Chico River (Martín 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

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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 Goñalons, 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 m2 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

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Martín

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 Fernández-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 & Fernández-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 (Domínguez-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 (Domínguez-Rodrigo & Piqueras, 2003; Domínguez-Rodrigo & Barba, 2006). Comparative evidence was gathered from modern carnivores who existed in Patagonia at the end of the Pleistocene (e.g., puma [Martín & Borrero, 1997; Martín, 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; Domínguez-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 element’s 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 & Fernández-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

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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; Nordenskjöld, 1996 [1900]; Esteban, 1996; Latorre, 1998; S. Vizcaíno and S. Bargo, pers. comm.), Camelidae (Roth, 1899, 1904; Nami & Menegaz, 1991; Prieto & Canto, 1997; Latorre, 1998), Macrauchenia patachonica (Nordenskjöld, 1996 [1900]), Panthera onca mesembrina (Roth, 1899, 1904; Cabrera, 1934; Nami & Menegaz, 1991; Prevosti & Martín, 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 (Vizcaíno & 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 tortoise’s 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 & Antón, 1997; Schiaffino et al., 2002). The description of this strategy indicates that “it bites through the animal’s skull between the ears or horns. The thick skulls of horses and cattle

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Martín

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; Martín, 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 (Fariña 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) (Martín, 2007). Marks are also important on horse bones, an animal of about 200 kg of body mass (Fariña 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 Sofía 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).

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Table 8. Sofía 4 cave. Carnivore marks on Mylodontinae specimens.

Table 7. Sofía 4 cave. Mylodontinae. Number of identified specimens (NISP) for each skeletal element.

Table 6. Sofía 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. Sofía 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

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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.

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produced today by jaguars hunting large prey (Martín, 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 19th 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 (Martín, 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 (Martín & Borrero, 1997; Martín, 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. Milodón 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

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Table 10. Milodón cave. Hippidion saldiasi. Number of identified specimens (NISP) for each skeletal element.

Table 9. Milodón 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

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Taxon

Specimens

Punctures Furrow

s Pits

Scoop-ing-out

Scores C

renulated

Mylodon darw

ini Skull, posterior part

1 1

1

Mylodon darw

ini Skull, base

1

Mylodon darw

ini Skull, supraoccipital *

1 1

1

Mylodon darw

ini Skull, tem

poral + petrosum *

1

Mylodon darw

ini M

andible, left 1

1 M

ylodon darwini

Mandible, right

1

1

Mylodon darw

ini M

axilla, frag.

1

M

ylodontinae cf. Mylodon

Stylohyale, frag.

1

1

Mylodontinae cf. M

ylodon A

xis, complete

1

Mylodontinae cf. M

ylodon V

ertebra, neural arc 1

1

1

1

Mylodontinae cf. M

ylodon R

ib, fragment

1

Mylodontinae cf. M

ylodon R

ib, fragment

1

Mylodontinae cf. M

ylodon Scapula, glenoid cavity

1

1

Mylodontinae cf. M

ylodon Tibia, com

plete 1

1

Mylodontinae cf. M

ylodon Tibia, com

plete

1 1

1

Mylodontinae cf. M

ylodon Tibia, prox. shaft

1

1

1

Mam

malia cf. M

ylodontinae Skull?, fragm

ent

1

1

Mam

malia cf. M

ylodontinae V

ertebra, frag.

1

1

Mam

malia cf. M

ylodontinae U

ndetermined

4

14

6 2

Total

14

5 25

2 9

7

Table 11. Milodón cave. C

arnivore marks on ground sloth bones. The num

ber of marks is larger than the num

ber of marked bones.

* Same elem

ent.

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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.

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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.

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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 Milodón 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. Milodón 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

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Table 14. Dos Herraduras Rockshelter. Mylodontinae. Number of identified specimens (NISP) for each skeletal element.. Dermal ossicles were not quantified.

Table 13. Milodón 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 & Martín, 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 (Martín, 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; Martín & Borrero, 1997; Domínguez-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

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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.

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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 (Martín, 2007). There is a MNI of three for Mylodon, including two young, one of them probably a newborn (Sergio Vizcaíno, 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 14C 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

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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.

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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

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Table 19. Fell cave. Mylodontinae. Number of identified specimens (NISP) for each skeletal element. Samples obtained by John Fell (1952–1959), Junius Bird (1969-1970) and Pedro Cárdenas (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

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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 Goñalons (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

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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

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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 Milodón 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.

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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 Milodón cave, but many individuals were probably hunted at this huge herbivore den, where they were at least partially processed (Martín, 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 (Fariña 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 - Milodón 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; Domínguez-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 Milodón 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

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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 Milodón 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 (Martín, 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 & Martín, 1996). San Román 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 (Martín, 2007). Direct late Pleistocene radiocarbon dates of Panthera onca mesembrina are available for Sofía 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).

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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 Milodón 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 Domínguez-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 Hereñú 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, Milodón cave,

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