Bone modification and destruction patterns of leporid carcasses by Geoffroy’s cat (Leopardus geoffroyi): An experimental study

Quaternary International 278 (2012) 71e80

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Bone modification and destruction patterns of leporid carcasses by Geoffroy’s cat (Leopardus geoffroyi): An experimental study María C. Álvarez a, *, Cristian A. Kaufmann a, Agustina Massigoge a, María A. Gutiérrez a, Daniel J. Rafuse b, Nahuel A. Scheifler b, Mariela E. González a a

CONICET, INCUAPA, Facultad de Ciencias Sociales, Universidad Nacional del Centro de la Provincia de Buenos Aires, Avenida del Valle 5737, B7400JWI Olavarría, Buenos Aires, Argentina INCUAPA, Facultad de Ciencias Sociales, Universidad Nacional del Centro de la Provincia de Buenos Aires, Avenida del Valle 5737, B7400JWI Olavarría, Buenos Aires, Argentina

b

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 2 December 2011

This paper characterizes the bone modification patterns on leporid carcasses generated by captive Geoffroy’s cats (Leopardus geoffroyi). The modification pattern from non-ingested remains is described in terms of anatomical representation, breakage, and tooth marks. Results from this experimental study suggest that Geoffroy’s cat tends to mostly destroy ribs and vertebrae. A high proportion of fractures were registered in the scapulas, cranium, and the epiphysis of long bones. The innominate, scapula, mandible, and long bones showed high frequencies of tooth marks, dominated by pits and punctures. If the activity of this particular predator is identified in fossil assemblages, the differential inter- and intrabone survival produced by this agent on small mammal prey should be taken into consideration when discussing anatomical part representation. Ó 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction In the pampean region of Argentina, small mammal (<10 kg) remains in archaeological sites have commonly served as indicators in the reconstruction of the paleoclimate. The presence and association of some small mammal species constitutes good indicators of the climate, as well inferences of the biotic communities and their relation to past environments (Pardiñas, 1991; Tonni, 1992; Pearson and Pearson, 1993; Vizcaíno et al., 1995). However, until the second half of the 1990s, small vertebrates had received little attention in the discussion of hunter-gatherer diet as greater emphasis was put on the role of medium and large sized prey such as the guanaco (Lama guanicoe), pampas deer (Ozotoceros bezoarticus), greater rhea (Rhea americana), and sea mammals (Arctocephalus australis and Otaria flavescens). Smaller mammals were generally considered as marginal or complementary resources. Different factors may have proportioned the lack of attention given to small fauna, including the difficulty in identifying anthropic modifications in smaller species or biases during excavation and taxonomic identification (Behrensmeyer, 1978; Stahl, 1996; Sunquist and Sunquist, 2001). However, in accordance with worldwide tendencies (Hudson, 1991; Stiner et al., 2000; Lupo and

* Corresponding author. E-mail address: [email protected] (M.C. Álvarez). 1040-6182/$ e see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.11.029

Schmitt, 2002; 2005; Schmitt et al., 2004), this situation has changed, in the last decades small sized fauna have received special interest in pampean archaeology, particularly when discussing issues such as the implication of diversification and intensification strategies in the exploitation of faunal resources during the Late Holocene (González de Bonaveri, 1997; Quintana et al., 2002). The development of actualistic models of distinct taphonomic agents that help to evaluate the origin of small vertebrates in archaeological sites is fundamental not only to making paleoenviromental interpretations but also human subsistence strategies. In Europe, the growing interest in these problems led to the development of methods and criteria for the identification of different taphonomic agents involved in the formation of microvertebrate fossil records (Andrews, 1990; Fernández-Jalvo and Andrews, 1992; Bochenski, 2005; Lloveras et al., 2008a). Nevertheless, in South America, the importance of these kinds of studies has only recently been established. Carnivores are considered one of the primary taphonomic agents that can accumulate and modify bones (Cruz-Uribe, 1991; Marean and Spencer, 1991; Capaldo and Blumenschine, 1994; Faith et al., 2007). The most common taphonomic effects of carnivore activity on bone remains in archaeological sites are differential anatomical representation, breakage, tooth marks, and digestive attrition; affecting in this mode, the anatomical and taxonomic representation of elements in the archaeological site (Binford, 1981; Cruz-Uribe, 1991; Marean and Spencer, 1991). The

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M.C. Álvarez et al. / Quaternary International 278 (2012) 71e80

Small felids from South America are diverse with eight known species. The present geographical distribution and fossil record suggest that at least five or six of these species evolved in South America, including Geoffroy’s cat, which last shared a common ancestor with the kodkod (Leopardus guigna) around 2.3 Ma (Nowell and Jackson, 1996; Johnson et al., 1999; Prevosti, 2006). Small cat remains have been found in both archaeological and paleontological sites from the Pampas to Patagonia (Mazzanti and Quintana, 2001; Prado et al., 2001; Martínez and Gutierrez, 2004; Zubimendi et al., 2010). However, compared to larger felids such as the puma (Puma concolor) or other carnivores from South America, the current fossil record of this species is scarce. Currently, Geoffroy’s cat habitat distribution includes Bolivia and southern Brazil, Argentina, and Chile (Johnson and Franklin, 1991; Lucherini et al., 2006). This carnivore is principally nocturnal and its range includes both dry and semiarid environments, although they utilize a variety of habitats such as grasslands, open forests, and wetlands. Geoffroy’s cat is an opportunistic solitary predator, and their current diet consists primarily of lagomorphs, small rodents, and aquatic birds (Ximénez, 1975; Manfredi et al., 2004; 2011; Perovic and Pereira, 2006). Although their weight varies geographically, the average adult male is around 4.9 kg, while females are on the order of 4.2 kg. While the females maintain territories of 2e6 km2, males are known to maintain areas up to 12 km2 (Cabrera and Yepes, 1960; Johnson and Franklin, 1991; Oliveira, 1994; Lucherini et al., 2006).

Notwithstanding, this field of study has focused in large part in the identification of modification patterns of digested remains from scats and pellets (Andrews and Evans, 1983; Andrews, 1990; Stallibrass, 1990; Schmitt and Juell, 1994; Gómez, 2007; Montalvo et al., 2007; Lloveras et al., 2008b). In consequence, there have been few taphonomic investigations which provide information regarding non-ingested remains by small carnivores (Lloveras et al., 2011; Montalvo et al., 2011). In South America, actualistic investigations of mammalian carnivore taphonomy has focused on the modifications of medium and large ungulates (e.g., Ovis aries, Lama sp.) by small foxes (Pseudalopex culpaeus, P. griseus and P. gymnocercus) and puma (Puma concolor). In general, these investigations have examined digested and non-ingested remains found in kill-sites, dens, and latrines; and have focused in the identification of bone damage and transport patterns produced by these carnivores (Borrero, 1988; 1990; Mondini, 1995; 2000; 2001; 2004; Borrero and Martín, 1996; Martín and Borrero, 1997; Nasti, 2000; Borrero et al., 2005; Montalvo et al., 2007; Mondini and Muñoz, 2008; Muñoz et al., 2008). Among the few experimental studies of carnivore mammals in South America is the work of Elkin and Mondini (2001), who evaluated the modifications produced on sheep by captive pampas fox (P. gymnocercus). In regards to small felids, some experimental studies have been developed, such as the analysis of gnawing damage by domestic cats on sheep bones realized by Moran and O’Connor (1992). These authors offered 13 roosted and defleshed lamb bones to domestic cat, however limiting their results to superficial bone modifications produced on the scapulas and humerus. In other studies, DelaneyRivera et al. (2009) conducted feeding experiments directed specifically to study tooth mark dimensions produced by several omnivore and carnivore species, among which included Leopardus pardalis, Lynx rufus, and Leiptailurus serval. In South America, the only experimental study of small felids is from the work of Gómez (2007) who analyzed the digestive effects on house mouse (Mus musculus) by distinct predators which currently inhabit the Pampas. In addition, this author studied the digestive effects produced by the jaguarundi (Herpailurus yaguarondi) in natural settings. However, the sample size collected by Gómez was small. Particularly significant to this work are the results from the studies of leporid bones recovered from modern Iberian lynx (Lynx pardinus) scats by Lloveras et al. (2008b) and of both scats and noningested domesticated rabbit from the Red fox (Vulpes vulpes) by Lloveras et al. (2011). Although their analysis comes from a different geographical region, the results provide a good example of how distinguishing taphonomic signatures of leporid remains accumulated by small mammalian carnivores can help understand past human subsistence and ecology.

3. Background

4. Materials and methods

From the beginning of the 1960s, with the pioneer work of Brain (1967); (1969); (1981), actualistic taphonomy studies of carnivore bone modification and destruction have focused on the scavenging and/or predation action of medium and large sized predators (>10 kg) on ungulates (e.g., Binford and Bertram, 1977; Binford, 1981; Haynes, 1983; Blumenschine, 1988; Marean and Spencer, 1991; Marean et al., 1992; Capaldo and Blumenschine, 1994; Selvaggio, 1994; Capaldo, 1995; Selvaggio and Wilder, 2001; Domínguez-Rodrigo and Piqueras, 2003). Advances in microvertebrate taphonomy prompted actualistic investigations of small predator carnivores (terrestrial mammals as well as raptor birds), emphasizing in predation studies such as the cause of death, and the accumulation of their remains in archaeological sites (Andrews, 1990; Fernández-Jalvo and Andrews, 1992; Stahl, 1996).

The specimens analyzed in this study correspond to the noningested remains of 10 European domestic rabbits (Oryctolagus cuniculus) by two Geoffroy’s cats in captivity sharing the same enclosure at a local zoo (Parque Zoológico La Máxima, Olavarría, Argentina). The rabbits chosen for this analysis were sub adults and adults, with an average weight of approximately 3 kg. The ten rabbits were offered to the cats during the months of November 2010 to July 2011. In each feeding, one rabbit was placed in the enclosure for three days, during which time the cats were not given other food. After three days, the enclosure was cleaned and all noningested and scats were collected. European domestic rabbit was chosen in this experiment because its size and bone structure are similar to many native small mammals that inhabited the pampean region during pre-historic

macroscopic characteristics of superficial damage produced by carnivores on bone permits the analysist to identify the participation of this agent in the formation of the assemblage (Binford, 1981; Johnson, 1985; Blumenschine and Selvaggio, 1991; Capaldo and Blumenschine, 1994). The study of multiple attributes (e.g., localization and morphology of the marks) further helps to identify the type of carnivore responsible of the bone modifications. The objective of this paper is to characterize the bone modification patterns generated by Geoffroy’s cat (Leopardus geoffroyi; D’Orbigny and Gervais, 1844), an extant felid from South America known to prey on small sized vertebrates commonly found in archaeological sites. This work focuses on the non-ingested remains and describes the modification pattern in terms of anatomical representation, breakage, and tooth marks. These results will be used to generate a model of the destructive behavior of this small feline, which can later be utilized as a means to evaluate the integrity of the archaeofaunal assemblages of sites located in regions where this predator or other small native felines inhabit or inhabited. 2. Geoffroy’s cat

M.C. Álvarez et al. / Quaternary International 278 (2012) 71e80

times, including the plains vizcacha (ca. 4 kg in adult), juvenile coypu (Myocastor coypus, ca. 7.5 kg in adult), and juvenile Patagonian mara (Dolichotis patagonum, ca. 8 kg in adult). Although it would be preferable to experiment with these indigenous prey or other South American small mammals (e.g., Sylvilagus brasiliensi), they are not locally accessible for repetitive experimentation or are protected species (D. patagonum). Characterization of the bone modifications produced by Geoffroy’s cat considered the anatomical representation, breakage, and tooth marks.

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complete, differentiating between isolated specimens and in situ specimens (Fernández-Jalvo and Andrews, 1992). 4.3. Tooth marks To analyze the tooth mark damage, each specimen was examined under a microscope (Motic ST-39) at 20 and 40 magnification. The marks were categorized using the types defined by Binford (1981): pits, punctures, crenulated edges, scores, and furrowing.

4.1. Anatomical representation 5. Results The minimum number of skeletal elements (MNE) and the relative abundance of the distinct elements that make up the skeleton (Ri) (Andrews, 1990) were calculated from the proportional relationship between the number of times that a determined element was represented in the study sample, and the expected number of this determined element (Ei) in relation to the minimum number of individuals (MNI): [Ri ¼ (MNE/MNI)  Ei]  100 (Andrews, 1990). The MNI was calculated from the most frequent skeletal element (Andrews, 1990). To evaluate if the destruction pattern generated by Geoffroy’s cat could be conditioned by bone mineral density (BMD), the relative frequency of parts (Ri%) was compared with the BMD index of Pavao and Stahl (1999) for domestic rabbit (O. cuniculus). To evaluate the representation of determined groups of elements, the following indexes were utilized: A) Measurement of the proportion of cranial elements in relation to postcranial elements used the indexes PCRT/CR: [(femur þ humerus)/(mandible þ maxilla)]  100 (Andrews, 1990), and PCRAP/CR: [(humerus þ radius þ ulna þ femur þ tibia)  26 (mandible þ maxilla þ molars)  10]  100. This last index was modified from Fernández-Jalvo and Andrews (1992), considering the dental formula of the rabbit. B) Measurement of the proportion of distal element ends in relation to proximal element ends used the index Z/E: [(tibia þ ulna)/(femur þ humerus)]  100 (Andrews, 1990). C) Measurement of the proportion of elements from the anterior limbs and posterior limbs used the index AN/PO: [(humerus þ radius þ ulna þ metacarpus)/(femur þ tibia þ metatarsus)]  100 (Lloveras et al., 2008a). D) To measure the relation between the postcranial axial skeleton and the appendicular skeleton, the index AX/AP was developed: [(vertebras þ ribs)  10/(humerus þ ulna þ radius þ femur þ tibia)  66]  100. E) Measurement of the relative proportion of isolated molars and incisors used two indexes: Isolated molars ¼ [(isolated molars)/ (empty mandible alveoli þ empty maxilla alveoli)]  100 (Andrews, 1990), and Isolated incisors ¼ [(isolated incisors)/ (empty mandible alveoli þ empty maxilla alveoli)]  100 (Andrews, 1990). 4.2. Breakage To evaluate the intensity of breakage, the integrity of each specimen was registered using the breakage categories proposed by Lloveras et al. (2008a). In addition, to assess the breakage of the cranial skeleton, the loss of molar and incisors was calculated based on the indexes used by Fernández-Jalvo and Andrews (1992): (empty mandible molar alveoli/expected mandible molars in situ)  100; (empty maxilla molar alveoli/expected maxilla molars in situ)  100; (empty maxilla incisor alveoli/expected maxilla incisors in situ)  100. Evaluation of the damage to molars and incisors took into account if the teeth were fragmented or

5.1. Anatomical representation A total of 793 bone specimens were recovered from the feeding experiments, of which 439 (55.4%) were identified anatomically and a MNI of 8 was calculated from the pelvis. In all 10 individuals there was a complete absence of ribs and sternebrae, as well as a low survival of vertebrae and scapula. The anterior limb survival was variable, while the posterior limb elements and the innominate survived in numerous individuals (Fig. 1). Table 1 and Fig. 2 show the anatomical composition of the total bone sample. The average relative abundance was calculated at 40.2%, which indicates a significant loss of bone. The results demonstrate that the mandible, maxilla, innominate, femur, tibia, and metatarsus present the highest average relative abundance (>60%), while the humerus, radius, calcaneum, astragalus, and carpal/tarsal present moderate representation (>30%). The isolated molars and incisors, scapula, patellae, phalanges 1/2 and phalanges 3 presented low representation (between 20% and 30%), while the vertebrae (4.9%) and metacarpus (10%) were scarce. The correlation between the MAU% and BMD was positive and significant (rs ¼ 0.468; p ¼ 0.0002), indicating that the anatomical profile observed in the non-ingested sample is conditioned by this property (Fig. 2). Table 2 presents the relative proportion of the skeletal element groups. The results indicate the following: The indexes that compare the cranial with the postcranial elements (PCRT/CR; PCRAP/CR) show distinct results. While the index that evaluates the relation between the proximal appendicular elements with the mandible and maxilla (PCRT/CR) shows a greater representation of cranial elements; the index that compares the proportion of appendicular elements with the maxilla, mandible and molars (PCRAP/CR) indicates a greater representation of postcranial elements. In reference to the index that evaluates the proportion of distal element ends in relation to proximal element ends (Z/E), data shows that both groups of elements are represented in similar proportions. The index AN/PO revels that there is a greater representation of posterior limb elements in relation to anterior limb bones, and the index AX/AP indicates a significant loss of the postcranial axial skeleton compared to the appendicular skeleton. Lastly, the indexes that evaluate the abundance of isolated teeth show that the relation between isolated molars, mandible and maxilla is almost proportional, while the isolated incisors are well represented. 5.2. Breakage From the number of identified bone specimens, the percentage of complete elements was 88.8% (390/439) (Table 2). The values varied in relation to the size of the bones, as observed here, the smaller sized skeletal elements present 100% completeness (tarsal/ carpal, phalanges 3, metatarsus, patellae, molars, and incisors). On the other hand, only 45.5% of the long bones were complete. Table 3

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M.C. Álvarez et al. / Quaternary International 278 (2012) 71e80

Fig. 1. Schematic representation of the relative frequency of parts (Ri%) and the minimum number of skeletal elements (MNE).

presents the number and percentage of skeletal parts included in each breakage category. The results from the breakage analysis suggest the following: The cranium presents significant breakage, and was identified through parts of the maxillary, incisors and maxilla, zygomatic arches and neurocranium. There was a low completeness of mandibles, represented by the mandible body and incisive parts, and the condylar process. From the few vertebrae recovered in the sample, a high percentage was complete, while fragments were represented by the spinous process and vertebral body. There was a low completeness of innominate, in the majority of the occasions it was identified by the acetabulum, ischium and ilium, as well as fragments of the acetabulum and ischium. All scapula were incomplete and identified in every case by the glenoid cavity and neck. With the exception of the metacarpus and metatarsus, which were found complete between 80% and 100% of the time, long bone fragments included the integrated shaft and distal epiphysis, distal epiphysis, and the proximal epiphysis and shaft. The indexes used to evaluate the loss of teeth indicate that for the mandible there was a significant loss of molars and their alveoli (86.1%), while

similar results were also found in the maxilla (87.9%). In reference to the incisors, the indexes indicate that both the mandible (50%) and the maxilla (36.4%) received moderate losses. Finally, although the fracture types were not quantified in this analysis, it’s important to mention that various shapes were recognized including irregular, transverse, and oblique fractures. 5.3. Tooth marks From the total number of recovered bones, 19.8% (n ¼ 81) contained tooth marks. The types of marks registered with greater frequency were pits (59%) and punctures (49%), followed by crenulated edges (37%), and scoring (26%). Furrows were registered in only 2% of the sample. Totals registered were 278 pits, 76 punctures, and 125 scores (Table 4). In reference to the frequency of marks, the phalanges 3, metatarsus, astragalus, and tarsal/carpal did not present any tooth marks, while the patellae and calcaneum presented low frequencies. The remaining of the elements (with the exception of the ribs and sternebrae) presented a considerable variety of tooth marks

M.C. Álvarez et al. / Quaternary International 278 (2012) 71e80 Table 1 Non-ingested rabbit skeletal parts.

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Table 2 Anatomical representation indexes.

Skeletal element

NR

MNE

Ri %

Indexes

%

Cranium (maxilla) Mandible Upper molars Lower molars Incisors Vertebrae Ribs Scapula Humerus Radius Ulna Innominate Femur Tibia Patellae Calcaneum Astragalus Carpal/tarsal Metacarpus Metatarsus Phalanges 1/2 Phalanges 3 Total Average relative abundance

15 14 31 23 14 18 0 4 5 10 9 17 20 16 4 7 6 65 8 40 77 35 339

11 12 31 23 14 17 0 4 5 7 7 13 14 12 4 7 6 65 8 40 76 35 411

68.7 75 25.8 28.7 29.2 4.9 0 25 31.2 43.7 43.7 81.2 87.5 75 25 43.7 37.5 33.8 10 62.5 27.9 24.3 884.3 40.2

PCRT/CR PCRAP/CR Z/E AN/PO AX/AP Isolated molars Isolated incisors Complete elements

82.6 151.9 100 30.7 5.7 90 140 88.8

Abbreviations; NR: number of remains; MNE: minimum number of skeletal elements; Ri%: relative frequency of parts.

and frequencies. The more affected elements by pits, punctures, and scoring were the innominate and scapula, followed by the humerus, femur, mandible, tibia, and ulna. Crenulated edges were found in greater percentages in the innominate, scapula, and mandible (Fig. 3). 6. Discussion 6.1. Bone modification pattern of Geoffroy’s cat In regards to the bone modification patterns generated by Geoffroy’s cat, some general tendencies were found. First, the relative abundance of the remains that survived is 40%, which indicates that an important percentage of remains from the original prey carcasses were lost. The elements that tended to be mostly

Abbreviations are provided in the methods section.

destroyed by this carnivore were ribs (absent in all individuals), followed by vertebrae, metacarpus, scapula, patellae, and phalanges (with a <30% representation). This bone modification pattern is also seen through some of the indexes, which indicate a difference in the anterior limbs in relation to the posterior limbs (AN/PO), and the postcranial axial elements with respect to the appendicular elements (AX/AP). In total, the skeletal part representation generated by Geoffroy’s cat indicates that the most represented elements tend to be those of the posterior limbs together with the innominate and cranium. As for the relationship between the cranial and postcranial elements, the results from the two indexes are dissimilar, with cranial elements dominating in the PCRT/CR index, and the postcranial in the PCRAP/CR. It is important to mention here the results from a preliminary analysis of the scats recovered from the Geoffroy’s cat enclosure. To date, a total of 294 specimens recovered from the scats, including bone and teeth remains, have been analyzed. From this total, 43% were anatomically identifiable. As anticipated, some of the absent or scarce elements from the non-ingested sample, such as phalanges were well represented in the scats samples. Likewise, some elements with greater abundance in the non-ingested sample, such as the mandible, innominate, and tibia were minimal in the scats. Other general tendencies observed in the scats sample are the greater representation of anterior limbs with respect to the posterior limbs, and the postcranial axial skeleton with respect to the appendicular skeleton. In reference to the breakage in the non-ingested sample, the percentage of complete identifiable elements was around 89%. While this is a high percentage, most of the elements correspond to

Fig. 2. Relative abundance of different skeletal parts, and correlation between the relative frequency of parts (Ri%) and the bone mineral density (BMD) of rabbit. Abbreviations; man: mandible, max: maxilla, inc: incisors, u mol: upper molars, I mol: lower molars, hum: humerus, rad: radius, uln: ulna, fem: femur, tib: tibia, pat: patella, sc: scapula, inn: innominate, mtc: metacarpus, mts: metatarsus, phal 1/2: phalanges 1/2, phal 3: phalanges 3, cal: calcaneum, ast: astragalus, c/t: carpal/tarsal, ver: vertebrae, rib: ribs.

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Table 3 Number and percentage of skeletal parts included in each breakage category. Long bones and metapodial

C N

%

PE N

%

PES N

%

S N

%

SDE N

%

DE N

%

Humerus Radius Ulna Femur Tibia Metacarpus Metatarsus

2 0 2 3 2 32 8

40 0 22.2 15 12.5 80 100

0 1 2 4 0 0 0

0 10 22.2 20 0 0 0

0 5 3 4 3 8 0

0 50 33.3 20 18.7 20 0

0 0 0 7 4 0 0

0 0 0 35 25 0 0

2 2 0 2 5 0 0

40 20 0 10 31.2 0 0

1 2 2 0 2 0 0

20 20 22.2 0 12.5 0 0

Mandible

N

%

Cranium

N

%

Innominate

N

%

Scapula

N

%

C IP MBI MB MBB CP

3 0 8 1 0 2

21.4 0 57.1 7.1 0 14.3

C IB IBM M ZA NC

2 1 2 6 3 2

12.5 6.25 12.5 37.5 18.7 12.5

C A AIS AISIL AIL IS IL

2 2 1 9 0 1 1

12.5 12.5 6.2 56.2 0 6.2 6.2

C GC GCN NF F

0 0 4 0 0

0 0 100 0 0

Vertebrae

N

%

Phalanges 1/2

N

%

Phalanges 3

N

%

C VB VE SP

13 1 0 4

72.2 5.5 0 22.2

C P D F

69 5 3 0

89.6 6.5 3.9 0

C P D F

35 0 0 0

100 0 0 0

Patella

N

%

Carpal/trasal

N

%

Calcaneum

N

%

Astragalus

N

%

Ribs

N

%

C F

4 0

100 0

C F

65 0

100 0

C F

7 0

100 0

C F

6 0

100 0

C F

0 0

0 0

Teeth

in-situ

isolated

Incisors

C F

Upper molars

Lower molars

Incisors

Upper molars

Lower molars

N

%

N

%

N

%

N

%

N

%

N

%

11 0

100 0

28 2

93.3 6.7

28 0

100 0

14 0

100 0

31 0

100 0

23 0

100 0

References: Long bones, metacarpal and metatarsal bones were classified as: complete (C), proximal epiphysis (PE), proximal epiphysis þ shaft (PES), shaft (S), shaft þ distal epiphysis (SDE) and distal epiphysis (DE). Mandible as: complete (C), incisive part (IP), mandible body þ incisive part (MBI), mandible body (MB), mandible body þ branch (MBB) and condylary process (CP). Cranium as: complete (C), incisive bone (IB), incisive bone þ maxilla (IBM), maxilla (M), zygomatic arch (ZA) and neurocranium (NC). Innominate as: complete (C), acetabulum (A), acetabulum þ ischium (AIS), acetabulum þ ischium þ illium (AISIL), acetabulum þ illium (AIL), ischium (IS) and illium (IL). Scapula as: complete (C), glenoid cavity (GC), glenoid cavity þ neck (GCN), neck þ fossa (NF) and fossa (F). Vertebrae as: complete (C), vertebral body (VB), vertebral epiphysis (VE) and spinous process (SP). Phalanges as: complete (C), proximal fragment, (P), distal fragment (D) and fragment (F). Patella, carpal/tarsal, calcaneum, astragalus, ribs and teeth as: complete (C) and fragment (F).

small articulating bones such as tarsal/carpal. On the other hand, a high proportion of fractures were registered in the scapula and cranium, and more than 50% of the long bones. In general, long bone breakage was more frequent in the epiphysis. In cases in

which the shaft was attached to an epiphysis, this tended to be the denser in the element (e.g., proximal humerus and proximal tibia). The positive and significant result obtained in the correlation between Ri% and BMD suggests that the differential inter-bone and

Table 4 Frequency of tooth marks. Skeletal Element

Pits FS

%

FM

FS

Punctures %

FM

FS

Scores %

FM

FS

%

FS

%

Maxilla Mandible Vertebrae Scapula Humerus Radius Ulna Innominate Femur Tibia Patellae Calcaneum Astragalus Carpal/tarsal Metarcapus Metatarsus Phalanges 1/2 Phalanges 3 Total

0 2 1 3 4 4 4 9 5 5 1 1 0 0 0 3 6 0 48

0 16.6 5.9 75 80 57.1 57.1 69.2 35.7 41.7 25 14.3 0 0 0 7.5 7.9 0

0 2 7 23 24 22 37 30 14 42 1 1 0 0 0 38 37 0 278

2 5 5 3 2 0 4 9 6 3 0 0 0 0 0 1 0 0 40

18.2 41.7 29.4 75 40 0 57.1 69.2 42.8 25 0 0 0 0 0 2.5 0 0

4 9 10 4 3 0 8 27 6 4 0 0 0 0 0 1 0 0 76

0 3 1 0 3 0 1 6 8 7 0 0 0 0 0 0 2 0 21

0 25 5.9 0 60 0 14.3 46.1 57.1 58.3 0 0 0 0 0 0 2.6 0

0 6 3 0 9 0 5 16 50 32 0 0 0 0 0 0 4 0 125

2 7 0 4 0 1 0 13 3 0 0 0 0 0 0 0 0 0 30

18.2 50 0 80 0 14.3 0 100 21.4 0 0 0 0 0 0 0 0 0

0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2

0 0 0 0 40 0 0 0 0 0 0 0 0 0 0 0 0 0

Abbreviations; FS: frequency of specimens with tooth marks, FM: frequency of marks.

Crenulated edges

Furrows

M.C. Álvarez et al. / Quaternary International 278 (2012) 71e80

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Fig. 3. Different rabbit bones with surface damage. (A) radius shaft with multiple pitting; (B) scapula blade with crenulated edge and puncture; (C) tibia distal with shaft fracture; (D) innominate with puncture; (E) humerus distal epiphysis with furrowing; (F) femur shaft with scoring.

intra-bone survival was mediated by bone density. However, other factors could have also conditioned the skeletal profile, including the mode of access to the carcass, hunger, carnivore group size, amongst others. Finally, around 20% of the non-ingested bones presented tooth marks. The pits and punctures dominated the sample, although other modifications were also registered. It is important to mention that the large majority of the anatomical units, with the exception of some carpal/tarsal, metacarpus, and maxilla presented tooth marks. The innominate and scapula, as well as the long bones and mandible were the elements that contained higher frequencies of modifications.

6.2. Comparison with other mammalian carnivores As mentioned earlier, there are few actualistic taphonomic studies that address the modifications of non-ingested remains by small mammalian carnivores. An exception to this is the work of Lloveras et al. (2011) study of scats and non-ingested domesticated rabbit from the Red fox (Vulpes vulpes). Table 5 compares the bone modification of Geoffroy’s cat with those of Red fox presented by these authors. Comparing samples of non-ingested bones from these two predators, the canid generates more destruction than the feline (Table 5). Evidence of this is revealed by the average relative

Table 5 Anatomical representation, breakage and tooth mark comparisons on leporid bone remains modified by small terrestrial carnivores. Vulpes vulpes Type of sample

Scat

Reference

Lloveras et al. (2011)

N* Mean Ri% Ri% > values Ri% < values PCRAP/CR** Z/E AN/PO Complete element % Mean value long bones Mean value total Tooth marks %

265 26.6 long bones-sc-cra mtc-c/t- inn ¼ þ proximal ¼ 0 12 3

Leopardus geoffroyi Non-ingested

Scat

Non-ingested

Unpublished personal data

Present study

639 25.6 mts-cal-ast-tib cra-sc-rib-inn-ver þ poscraneal þ distal þ hindlimb

294 34 cra-phal-cal-ast man-hum-tib-pat-inn ¼ ¼ þ forelimb

439 40.2 fem-inn-tib-man-max-mts ver-rib-mtc þ poscraneal ¼ þ hindlimb

5.4 89.4 9.5

0 15 1.6

15 88.8 19.8

Abbreviations; cra: cranium, man: mandible, max: maxilla, hum: humerus, rad: radius, uln: ulna, fem: femur, tib: tibia, pat: patella, sc: scapula, inn: innominate, mtc: metacarpus, mts: metatarsus, phal: phalanges, cal: calcaneum, ast: astragalus, c/t: carpal/tarsal, ver: vertebrae, rib: ribs. References: *Anatomically identified specimens. **Corresponds to the index PCRLB/CR in Lloveras et al. (2011).

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abundance, which in the Red fox is 25.6% and 40.2% in Geoffroy’s cat. Both predators completely destroy the ribs and in large part the vertebrae, which is a pattern also observed in larger sized carnivores (see discussion in Cleghorn and Marean, 2007). As seen in Table 5, there is greater survival of the posterior limbs in relation to the anterior limbs. In regards to the differences in bone modification between these two predators, in the case of the Red fox there is more destruction of the cranium, mandible, innominate, scapula, proximal limb elements, patella, and metacarpus; while in Geoffroy’s cat there is a significant percentage of bone survival of the cranium, mandible, innominate, and femur. Although the Red fox presented greater destruction of long bone elements, complete long bones that survived in both species are minimal. On the other hand, the overall average of complete bone is high and very similar in both cases (ca. 89%); although, it is important to mention that the majority of the complete elements correspond to small articulating bones such phalanges and tarsal/carpal. The results from the scats analysis of both carnivores corresponds with the expectations as previously described for bone destruction. In this sense, remains of ribs and vertebrae were identified, although in low percentages. Also abundant are fragments of scapula and long bones from the anterior limbs. The phalanges, carpal/tarsal, calcaneum, and astragalus were also well represented, which could be due in part to its minimal fragmentation, a condition that later favors their identification. The average percentage of complete bones was low in both carnivore scat samples (12% Red fox, 15% Geoffroy’s cat); while most of these elements correspond to carpal/tarsal and phalanges. In neither of the scat samples were any long bones found complete, an expected result given the size relationship between predator and prey. In the non-ingested remains, the percentage of elements with tooth marks is 9.5% for the Red fox and 19.8% for Geoffroy’s cat. The greater frequency of tooth marks registered in Geoffroy’s cat sample could be related in part to the greater survival of long bones, innominate, and scapula. On the other hand, the percentage of bones with tooth marks in the scats from both species is low (3% Red fox, 1.6% Geoffroy’s cat). It is possible, as mentioned by Lloveras et al. (2011), that this characteristic is conditioned by the degree of fragmentation and digestion. 6.3. Implications for the study of archaeofaunal assemblages The potential contribution of Geoffroy’s cat must be considered when studying the formation of faunal assemblages from archaeological sites in South America, particularly those sites which contain small sized prey with similar or smaller body mass as the rabbit. In the southern portion of the continent, the zooarchaeological record suggests that small mammal prey played an important role in Late Holocene hunter-gatherer subsistence, and many authors relate this fact to an intensification process (González de Bonaveri, 1997; Quintana et al., 2002; Escosteguy, 2011; Quintana and Mazzanti, 2011; among others). Given the importance that the intensification process may have had for hunter-gatherer societies, it is necessary to evaluate in depth the participation of Geoffroy’s cat (and other small carnivores) prey in the formation of the archaeofaunal assemblages. If the activity of this predator is identified, the differential inter- and intra-bone survival produced by this agent on small mammal prey should be taken into consideration when discussing anatomical part representation. Faunal exploitation models for the Late Holocene in the Pampas suggest an intensification and diversification in the exploitation of resources (see Martínez and Gutierrez, 2004). In this sense, the presence of rodents of similar size or slightly larger than rabbits (less than 10 kg body weight), such as the coypu in the Salado Depression (Escosteguy, 2011) or the plains vizcacha in the sierras

and inter-sierras of the Pampas (Martínez et al., 2001; Quintana and Mazzanti, 2011; among others) are common in archaeological sites, and there is clear evidence of processing and consumption of these mammals, which appear to have been exploited in an integrated manner. The anatomic relative abundance data from plains vizcacha in sites from the sierras show a medium to low representation of ribs and vertebrae (Quintana and Mazzanti, 2011), while the percentage of the number of individual specimens (NISP%) from sites in the Salado Depression region and the northeast of the province of Buenos Aires also show low values in these elements (Escosteguy, 2011). This anatomical pattern registered in sites where there is abundant evidence of anthropic activity of small mammals coincides with the results of the skeletal part profiles produced by Geoffroy’s cat. Thus, when evaluating the origin of the archaeofaunal assemblages, the skeletal part profiles are not a diagnostic criterion for determining the causal agent. It is necessary to consider multiple variables, such as the type and characteristics of the surface modifications (tooth marks, cut marks), as well as the breakage and types of fractures in the bones. Although the anatomical pattern cannot be considered a diagnostic criteria for identifying the involvement of this agent in the formation of the assemblage, the information obtained in this experimental study on differential anatomical representation is important when evaluating the skeletal part representation in the archaeofaunal assemblage, particularly where there is clear evidence of the participation of small carnivores (e.g., tooth marks). In this regard, the possibility for the inclusion of dissimilar proportions of distinct skeletal elements should be considered. 7. Conclusion This work demonstrates how Geoffroy’s cat can have a high impact on small mammal bones, thus conditioning the anatomical profile. This agent is capable of completely destroying prey elements of similar size to domesticated rabbit (ca. 3 kg). Ecological studies in the pampean grassland suggest that hares (Lepus europaeus, body mass ca. 4.5 kg) are especially important in the diet of modern Geoffroy’s cat (Manfredi et al., 2011: 317). Therefore, in the wild, bone destruction of ribs and vertebrae of rabbit sized prey is expected from this feline. Though the extent of destruction likely varies as a function of prey availability, cats hunger, and carnivore competition, the bone-crushing abilities of this species is a significant factor in prey damage. The results suggest that Geoffroy’s cat is physically capable of destroying bones from prey weighing up to ca. 60% of its own body mass. In reference to the inter-bone survival, this study suggests that Geoffroy’s cat selectively destroys elements. Particularly relevant was the complete absence of ribs and low frequencies of vertebrae. The results also support the premise proposed by other researchers (Brain, 1967; 1969; Marean and Spencer, 1991; Blumenschine and Marean, 1993; Cleghorn and Marean, 2007), that carnivores selectively destroy those portions with lesser bone mineral density and with greater bone fat (e.g. axial elements and long bone epiphysis). This study provides an initial baseline for the recognition of the taphonomic effects of Geoffroy’s cat on small mammal assemblages, which should be called to attention when interpreting archaeofaunal remains. Faunal analysts working in regions which contain small sized prey should be aware of small felids as agents in the accumulation and destruction of faunal remains. Additionally, the results of this article are particularly relevant to researchers who work with archaeological leporid remains in areas where small felids with similar behavior and ecology to Geoffroy’s cat are or were present (e.g., European wild cat; Felis silvestris). Detailed analysis of the tooth marks in the experimental sample as well as

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complementary naturalistic studies of Geoffroy’s cat will contribute to strengthen this preliminary bone modification model for this carnivore. These studies will help determine the participation of small felids in bone accumulations and thus strengthen the foundations upon which archaeological and paleoecological interpretations are made. Acknowledgements Financial support for this research was provided by CONICET (grant PIP 112-200801-00291) and ANPCyT (grant PICT 08-814). This paper was developed within the research program INCUAPA (Facultad de Ciencias Sociales, UNCPBA). Special thanks to the Parque Zoológico La Máxima (Municipalidad de Olavarría), Sandra S. Botasi, and Horacio M. Grand. We also want to thank Agustin Venzi and Juan Rodríguez for helping to process the samples. We thank the anonymous reviewers who helped improve the previous version of this paper. Interpretations and any errors, however, are those of the authors. References Andrews, P., 1990. Owls, Caves and Fossils. University of Chicago Press, Chicago. Andrews, P., Evans, E.M., 1983. Small mammal bone accumulations produced by mammalian carnivores. Paleobiology 9, 289e307. Behrensmeyer, A.K., 1978. Taphonomic and ecological information from bone weathering. Paleobiology 4, 150e162. Binford, L.R., 1981. Bones: Ancient Men and Modern Myths. Academic Press, New York. Binford, L.R., Bertram, J.B., 1977. Bone frequencies-and attritional processes. In: Binford, L.R. (Ed.), For Theory Building in Archaeology. Academic Press, New York, pp. 77e153. Blumenschine, R.J., 1988. An experimental model of the timing of hominid and carnivore influence on archaeological bone assemblages. Journal of Archaeological Science 15, 483e502. Blumenschine, R.J., Marean, C.W., 1993. A carnivore’s view of archaeological bone assemblage. In: Hudson, J. (Ed.), From Bones to Behavior. Center for Archaeological Investigation, Southern Illinois University, Carbondale, pp. 273e300. Blumenschine, R.J., Selvaggio, M., 1991. On the marks of marrow bone processing by hammerstones and hyaenas: their anatomical patterning and archaeological implications. In: Clark, J.D. (Ed.), Cultural beginnings: approaches to understanding early hominid life-ways in the African savanna. Romisch-Germanisches Zentralmuseum, Forschunginstitut fur Vor- und Fruhgeschichte, Monographien 19, Bonn, pp. 17e31. Bochenski, Z.M., 2005. Owls, diurnal raptors and humans: signature on avian bones. In: O’Connor, T. (Ed.), Biosphere and Lithosphere. New Studies in Vertebrate Taphonomy. Oxbow Books, Oxford, pp. 31e45. Borrero, L.A., 1988. Estudios tafonómicos en Tierra del Fuego: su relevancia para entender procesos de formación del registro arqueológico. In: Yacobaccio, H. (Ed.), Arqueología Contemporánea Argentina: actualidad y perspectivas. Ediciones Búsqueda, Buenos Aires, pp. 13e32. Borrero, L.A., 1990. Taphonomy of guanaco bones in Tierra del Fuego. Quaternary Research 34, 361e371. Borrero, L.A., Martín, F.M., 1996. Tafonomía de carnívoros: un enfoque regional. In: Gómez Otero, J. (Ed.), Arqueología solo Patagonia. CENPAT-CONICET, Puerto Madryn, pp. 189e198. Borrero, L.A., Martín, F.M., Vargas, J., 2005. Tafonomía de la interacción entre pumas y guanacos en el Parque Nacional Torres del Paine, Chile. Magallania 33 (1), 95e114. Brain, C.K., 1967. Hottentot food remains and their bearing on the interpretation of fossil bone assemblages. Scientific Papers of Namib Desert Research Station 32, 1e11. Brain, C.K., 1969. The probable role of leopards as predators of the Swartkrans australopithecines. South African Archaeological Bulletin 24, 127e143. Brain, C.K., 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. The University of Chicago Press, Chicago. Cabrera, A., Yepes, J., 1960. Mamíferos Sudamericanos. Tomo I. EDIAR, Buenos Aires. Capaldo, S.D., 1995. Inferring hominid and carnivore behavior from dual-patterned archaeological assemblages. Ph.D. Dissertation, Rutgers University, New Brunswick. Capaldo, S.D., Blumenschine, R.J., 1994. A quantitative diagnosis of notches made by hammerstone percussion and carnivore gnawing on bovid long bones. American Antiquity 59, 724e748. Cleghorn, N., Marean, C.W., 2007. The destruction of human-discarded bone by carnivores: the growth of a general model for bone survival and destruction in zooarchaeological assemblages. In: Pickering, T.R., Toth, N., Schick, K. (Eds.), African Taphonomy: a Tribute to the Career of C. K. "Bob" Brain. Stone Age Press, Bloomington, pp. 13e42. Cruz-Uribe, K., 1991. Distinguishing hyena from hominid bone accumulations. Journal of Field Archaeology 18, 467e486.

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