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Article JTa147. All rights reserved. *E-mail: [email protected]

2012

Journal of Taphonomy PROMETHEUS PRESS/PALAEONTOLOGICAL NETWORK FOUNDATION (TERUEL)

VOLUME 10 (ISSUE 3-4)

Available online at www.journaltaphonomy.com

The Búho and the Zarzamora caves (Segovia, Spain) are two small karstic cavities in the North of the Central System Cretaceous limestones, in the transitional region between the Sierra de Guadarrama Mountains and the Castilian Plateau. The infilling sediment was excavated during two periods, from 1988-1990 and from 2008-actuality, and subsequently has been assigned to the Late Pleistocene. The aim of this study is the taphonomical analysis of the macrofaunal remains from the old and the new excavation campaigns.

A Taphonomic study of the Búho and Zarzamora caves.

Hyenas and Humans in the Iberian Plateau (Segovia, Spain) during the Late Pleistocene

M.T. Nohemi Sala* Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos

C/ Monforte de Lemos 5, 28029 Madrid

Milagros Algaba Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos

C/ Monforte de Lemos 5, 28029 Madrid, Spain

Juan Luis Arsuaga Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos

C/ Monforte de Lemos 5, 28029 Madrid Departamento Paleontología, Facultad de Ciencias Geológicas

Universidad Complutense de Madrid, Spain

Arantza Aranburu DepartamentoMineralogía y Petrología, Facultad de Ciencia y Tecnología

UPV/EHU, Bilbao, Spain

Ana Pantoja Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos

C/ Monforte de Lemos 5, 28029 Madrid Departamento Paleontología, Facultad de Ciencias Geológicas

Universidad Complutense de Madrid, Spain

Journal of Taphonomy 10 (3-4) (2012), 477-497. Manuscript received 15 March 2012, revised manuscript accepted 15 November 2012.

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A taphonomic study of Búho and Zarzamora caves

2008 and 2013 and the data collected have led to the publishing of new articles (Sala et al., 2009, 2010, 2011). The systematic study of the macrofauna remains has been employed to elaborate the following taxonomical list, which comprises all the taxa present throughout the entire sequence: Carnivora (Crocuta crocuta, Lynx sp., cf. Panthera sp., Canis lupus, Meles meles); Perisodactyla (Equus ferus, Equus hydruntinus, Stephanorhinus hemitoechus); Artiodactyla (Sus scrofa, Cervus elaphus, Bos primigenius, Bison priscus) (Sala et al., 2009, 2010, 2011). None of the carnivores identified suggests either a maximum cold period or a typical forest environment. This absence suggests that the association found at the Zarzamora Cave could easily belong to a mild temperature period in an open environment inhabited by such species as hyena and lynx, which need open spaces and herbaceous cover (Sala et al., 2011). The presence of Bison priscus in association with the steppe rhinoceros (Stephanorhinus hemitoechus), as well as the presence of grazing species such as Equus ferus and Equus hydruntinus also support this interpretation of an open habitat in this area during the Late Pleistocene (Sala et al., 2010, 2011). The microfauna fossil assemblage, with a high-prevalence of Microtus cabrerae and M. duodecimcostatus, is also indicative of temperate climate. The set of rodent species

Introduction The Búho and Zarzamora caves are a paleontological sites located in central Iberia, which includes rich and diversified fauna and flora from the MIS 3 period. This site is a Late Pleistocene cave deposit in a limestone outcrop that extends from the crystalline piedmont of the Sierra de Guadarrama to the Tertiary Castilian Plateau. The small cave is located in Perogordo (40º 55.906´N, 04º 08.220´W), near the city of Segovia. The cave is situated at the edge of a small stream, the “Arroyo Tejadilla”, a tributary of the Eresma River (Figure 1). Previous works The Late Pleistocene site of the Búho Cave was discovered in the late eighties. Excavations were done at the site from 1988 until 1990. During an earlier phase of excavation undertaken during the 1980s, some papers were published that dealt with taxonomical aspects (Molero et al., 1989; Íñigo, 1995; Maldonado, 1996; Íñigo et al., 1998). Additionally a hyena den origin was suggested for the accumulation at the site (Íñigo et al., 1996, 1998). In 2008 a multidisciplinary research team started excavating the site again. These new excavations were undertaken between

The taxonomical list includes: Carnivora (Crocuta crocuta, cf. Panthera sp., Lynx sp., Canis lupus, Vulpes vulpes and Meles meles), Perissodactyla (Equus ferus, Equus hydruntinus and Stephanorhinus hemitoechus) and Artiodactyla (Sus scrofa, Cervus elaphus, Bison priscus and Bos primigenius). The abundance of hyena juveniles and coprolites, as well as carnivore tooth marks and digested bones suggest that the Búho and Zarzamora caves worked as a spotted hyena den during the Late Pleistocene. Nevertheless some human activity is also present in the Zarzamora cave site with evidences of cut marks in carnivore remains (Lynx sp.). The macro faunal association suggests an open environment where equids were the most abundant herbivores. Keywords: HYENA DEN, ZARZAMORA CAVE, KARSTIC SITE, TAPHONOMY

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forests during this period in this area where the oak and the holm oak, together with the juniper and other mesophilous plants, are the dominant taxa (Sala et al., 2011). The present work describes a complete taphonomical study of the Búho and Zarzamora caves site. The fossil material studied comes from both the initial and more recent excavation periods.

found at the site, where small moles are the dominant group, suggests an environment dominated by open areas but also featuring some spots of woody vegetation which acted as refugia for mice and dormice (Sala et al., 2011). The analysis of the pollen gathered from the hyena coprolites also supports the existence of an open environment of steppes, grasslands, and small and cleared

Figure 1. Geographical location of the Zarzamora and Búho cave sites in relation with the Sistema Central Mountains and the city of Segovia. Aerial view of the Tejadilla Valley, home to the Búho and Zarzamora caves. Detail of the different sectors of the site (Búho Cave, Zarzamora Cave and Cata Exterior).

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that is associated with this lamination. The clays have an identical stratigraphic position and a similar mineralogical composition, with a consequent interpretation that they belong to the same stratigraphic level. The clay minerals that comprise this deposit are Illite, Kaolinite and Smectite. The argillaceous level of the base could be due to water transport (Figure 2). Middle unit: The next stratigraphic level of the Cata Exterior (level II) is a deposit of macrovertebrate pebbles and fossils cemented in a carbonated matrix. In the Zarzamora Cave, this level is present in relict patches affixed to the wall that fill the irregularities of the paleorelief in the chamber (level 3).The top of this level on the Cata Exterior matches the height of the inside of the Zarzamora Cave. Nonetheless, the inside of the cave was some 10 cm thick, while the thickness of the Cata Exterior reached 40 cm. The fossils are floating in a matrix of equigranular idiomorph dolomite crystals with a greyish orange 10YR7/4 colour. The appearance of the level is homogeneous, without any observable sedimentary structures, and contains 0.5-2.0 cm pebbles of altered dolostone. This level is interpreted as dolostone arenisation, where the blocks are part of the same rock that forms the cave. Applying ultrafiltration pre-treatment protocols, radiocarbon dating places fossils at this level at over 44,400 years BP (OxA-24566, Research Laboratory for Archaeology and the History of Art, University of Oxford) (Figure 2). Upper unit: Level 1-2 inside the Zarzamora Cave and Level III of the Cata Exterior contain similar mineralogical ratios, and are the only levels with plagioclase and potassium feldspar. This fact lets both levels be correlated (Figure 2). Detritic material practically reaches the top, which caused the cavity to fill up inside Zarzamora Cave.

The site The Zarzamora Cave is a small gallery with a NE-SW direction that is approximately 1 m high, 2 m wide and some 7 m of depth with accessible development. The total length is still unknown due to the sediment infilling. The “keyhole-shaped” section suggests that the karstic system evolved from phreatic to vadose conditions. The sedimentary infilling is around 90 cm thick and three stratigraphic units have been identified. The upper two units are made up of a crystalline dolomite matrix with macro and microvertebrate fossils. The lower unit is formed of plastic clay with no paleontological evidence. The entrance to the Zarzamora Cave is a few meters away from the entrance to the Búho Cave. It was during the excavation of the latter in the late 1980s that the entrance to the Zarzamora Cave was discovered. Then, the site was divided into three sectors: the Búho Cave, the Zarzamora Cave and Cata Exterior (Figure 1). Stratigraphy The deposits consist of three major stratigraphic units correlated in two sectors of the site (Zarzamora and Cata Exterior). The stratigraphy was established and described by Sala et al. (2009, 2011) (Figure 2). Lower unit: The first deposit of sedimentary fill both inside the Zarzamora Cave (level 4) and at the Cata Exterior (level I) are comprised of plastic clays that are homogenous and sterile of paleontological content. Following colour charts, these clays are ‘yellowish brown 10YR 5/4’.This level reveals parallel low energy lamination. It contains a fine patina of manganese oxide

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Material and Methods This study has collected data from at least 2033 macrovertebrate fossil remains from the three sectors, belonging to both the excavation campaigns back in the 1980s and more recent ones. A Nikon SMZ800 (stereoscopic zoom microscope) has been used to examine surface modification on all bone fragments (NR=1219). In order to evaluate the skeletal element represented in the macrofaunal assemblage, the Number of Remains (NR), Number of Identified Specimens (NISP),

The matrix is comprised of equigranular idiomorph dolomite crystals with a moderate yellowish brown 10YR 5/4 colour, rich in organic matter and with some small scattered charcoal, as well as allochthonous contributions of potassium feldspar and plagioclase. In addition to dolomite pebbles, this matrix includes macrovertebrate fossils and hyena coprolites. This level presents intense bioturbation produced by small mammals. Radiocarbon dating of a fossil at this level is 32,510 ± 240 years BP (Beta‐

252209, Beta Analytic Laboratory, Florida) (Sala et al., 2010).

Figure 2. Stratigraphic section of Zarzamora cave (right) and Cata Exterior (left).

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determine which ones were significantly different. With the aim of identifying the breakage process (green or dry bone fragmentation), all long bone fragments were analysed following the criteria developed by Villa & Mahieu (1991), Bunn (1983), Haynes (1983b) and Lyman (1993, 1994). This method considers: the fracture location following Lyman (1993, 1994); the fracture outline (longitudinal, transverse and V-shaped); the angle formed by the fracture surface and the bone’s cortical surface (oblique, right or mixed); the characteristics of the fracture edges (jagged or smooth); shaft fragment length (1: shafts that are less than one-fourth the original length; 2: length comprised between one-fourth and one-half; 3: between one-half and three-fourths; 4: is more than three-fourths, essentially a complete or almost complete shaft); and shaft circumference (1: bone circumference is less than half of the original; 2: circumference is more than half in at least a portion of the bone length and 3: complete circumference in at least a portion of the bone length). Results

Faunal analysis from the site involves 2033 remains. Out of these, 895 were attributed to the categories established by weight sizes and 828 have been identified to the taxonomical level. As one can observe in Table 1, equids are the predominant ungulates in the faunal association and hyenas are the most abundant carnivores. These proportions are upheld at both stratigraphic levels. In addition to hyena fossil remains, 26 coprolites have been recovered. The features of these coprolites suggest that they belong to spotted hyena. These characteristics are:

the Minimum Number of Elements (MNE), the Minimum Number of Individuals (MNI) and the Skeletal survival rate (% Survi.=MNEi x 100 / number of elements i in the animal skeleton x MNI) were calculated (Brain, 1969, 1981; Klein & Cruz-Uribe, 1984; Lyman, 1994). The age of death of the Crocuta crocuta remains have been calculated following the criteria of Stiner (1994). Carnivore damage on bone surfaces is classified into pits, punctures, and scores. The crenulated edges, furrowing and evidence of bone corrosion caused by stomach acid is attributed to carnivore modification (Haynes, 1980, 1983a; Maguire et al., 1980; Binford, 1981). Tooth-mark distribution on bone portions was done by following the methods of (Selvaggio & Wilder, 2001; Domínguez-Rodrigo & Piqueras, 2003; Delaney-Rivera et al., 2009). Conspicuous marks were measured in length and breadth (Figure 3) and measurements were taken with an digital calliper. For the identification of the carnivore species that acted on the assemblage, tooth marks have been measured and then compared to data published by Selvaggio & Wilder (2001), Domínguez Rodrigo & Piqueras (2003), Saladié et al. (2013) and Delaney-Rivera et al. (2009). Furthermore, metric data have been compared to experimental data obtained from living carnivores (P. leo, P. pardus, C. lupus, U. arctos, P. concolor, P. onca and C. crocuta) (Sala, 2012; Sala & Arsuaga, 2013; Sala et al., in press). For this comparison, the location of tooth marks on dense and cancellous or cortical bone was taken into account. For the univariate analysis, we performed a Kruskal Wallis test to compare the differences between the Búho and Zarzamora and living carnivore’s samples. When a significant difference (p < 0.05) was found in a variable, we performed a Mann Whitney (Mann & Whitney, 1947) test between all possible pairs of samples to

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and radii. There is little or no presence of axial skeletons. A very similar pattern is also seen with regard to medium-size ungulates (Figure 4), namely, a high representation of the inferior appendicular skeleton (metapodial and tarsal bones), as well as a high representation of dental remains. Of the superior appendicular skeleton, only humeri have been conserved and there are no remains from the axial skeleton. This bias against the axial skeleton and superior appendicular skeleton could be interpreted as owing to the higher bone density of autopods, which make them more resistant to destruction by both carnivores and geological processes. To be able to relate these two factors (bone density with respect to the high ratio of anatomical representation), a correlation analysis was performed between them. Bone density data were taken from Lyman (1984, 1994). There is a relationship between these two factors, whose correlation coefficient is 0.83. Of the remains analysed, 156 (8.8% of the sample) were complete bones, with

They show a spherical morphology with their typical central depression in some cases (Carrión et al., 2001; Larkin et al., 2000); their large size, between 3 and 6 cm, is consistent with the measures taken in different European Pleistocene sites (Carrión et al., 2001; Kruuk, 1972); bone digested fragments are found inside these coprolites as documented by various authors (Carrión et al., 2007; Kruuk, 1972; Larkin et al., 2000); their internal morphology show numerous vesicles like those described in the A Valiña cave and other sites (Fernández Rodríguez et al., 1995). During the early campaigns some spotted hyena coprolites were also recovered and published (Ínigo et al. 1998), which have been revised and incorporated into the present study. For carnivores, dental remains are dominant in almost all taxa. The spotted hyena is represented at the site by adult and immature individuals. The remains of the juveniles consist of milk teeth and tooth germs, while the adult group is represented by permanent teeth showing several stages of wear. We estimate that at least 4 juveniles, 3 primes and two old Crocuta crocuta individuals are represented in the Zarzamora cave sample. Two phalanges assigned to hyena have also been recovered. A few dental remains and one postcranial remain of Canis lupus have been recovered. A single remain (a phalange) of a medium-sized felid has been attributed to cf. Panthera pardus (Sala et al., 2011), and a single dental remain to Vulpes vulpes. A few postcranial remains, all of the mautopods, of Lynx sp. have also been recovered in both the upper and middle units. With respect to skeletal remains of the ungulates, we observe that the anatomical parts of Bos/Bison, E. ferus and S. hemitoechus that are best represented are carpal, tarsal and metapodial bones, followed by tibias

Figure 3. Examples of SEM images of tooth marks (a: scores; b: punctures and c-d: pits) showing the measurements considered in this study.

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no fractures. Thus, over 91% of the bone samples had fractures. Of the fractured remains, 1304 (73.8%) corresponded to unidentifiable bone splinter fragments. For bone fragments that could be identified, at least for the size of the ungulate, the fracturing characteristics were established as depicted in Table 2. As can be seen in Table 2, the transverse orientation of the fractures is dominant in both weight sizes, where longitudinal and oblique fractures are similarly represented. With regard to the fracture angle, there is a predominance of fractures with oblique and mixed angles. According to Villa & Mahieu (1991), this

pattern indicates fresh bone breakage. The properties of the fracture edges in the majority of cases are characterised by being smooth, with a high percentage of crenulated edges in the sample. Regarding post-depositional breakage, transverse outlines with right angles have been identified rarely. When quantifying the fracturing patterns of the long bones for diaphyseal lengths and circumferences, several samples were taken into account. First, the complete sample was considered, namely, identifiable and unidentifiable elements. Fracturing for the complete sample reveals a predominance of fragments whose circumference is less than half the total and where diaphyseal fragments

UPPER UNIT MIDDLE UNIT NISP MNE MNI NISP MNE MNI Equus ferus 189 184 8 79 79 4 Equus hydruntinus 43 42 4 9 8 1 Equus sp. 131 - - 40 - - Bos/Bison 47 - - 14 - - Bos primigenius 10 10 2 1 1 1 Bison priscus 6 6 2 1 1 1 Cervus elaphus 28 25 3 14 13 2 Sus scrofa 3 2 1 8 7 2 Stephanorhinus hemitoechus 13 13 2 1 1 1 Crocuta crocuta 50 44 7 20 17 4 Lynx sp. 3 3 1 3 3 1 cf. Panthera pardus 1 1 1 0 0 0 Canis lupus 5 5 1 1 1 1 Vulpes vulpes 0 0 0 1 1 1 Meles meles 6 6 1 0 0 0 Large sized ungulate 50 - - 8 - - Medium sized ungulate 3 - - 0 - - Small sized ungulate 3 - - 0 - - Indet (NR) 532 - - 710 - -

TOTAL 1123 910

Table 1. NR, NISP, MNE and MNI by taxa and size categories from the upper and middle unit’s faunal assemblage of Zarzamora and Búho Caves.

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cylinders, with general diaphysis lengths less than half of the total. Figure 5 depicts the results for medium-size ungulates. This figure reveals that the majority of the sample is represented by diaphyses with complete circumferences and incomplete diaphyseal lengths. The sampling size for these ungulates was low. This is because the large part of the remains for this ungulate size are unidentifiable splinters. Due to this, these values do not match the real

are small (< ¼ of the total), so that the majority are small bone splinter (Figure 5). Given that the number of splinters is so high, it was decided to also perform the same analysis on the remains identified -at least anatomically- at a level of the ungulate size. Figure 5 shows that more than 28% of large ungulates are represented by fragments of long-bone splinters. Some 61% of the sample is characterised by having complete circumferences, by way of diaphyseal

Figure 4. % of survival of the different skeletal portions in large size (superior image) and medium size ungulates (inferior image).

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bottom units are scores and pitting. In general, there are very few punctures, and furrowing is well represented, particularly in the epiphyses of long and large-sized bones, compact bones and flat bones. Metric data on the tooth marks are summarised in Table 4. Another modification typical of carnivores is evidence of disintegration due to gastric acids. At the Búho/Zarzamora Cave site, 20.54% of the total sample analysed have been found with clear evidence that they were digested. The mean length of the fragments with digestion evidence is 31.6 mm. The bone fragments inside of the coprolites are much smaller, so the digestion evidence of the bone splinters are likely due to the regurgitation but not to the faeces disintegration. These data are compatible with the results obtained by Horwitz (1990). No evidence has been found of burned bones among the collection of fossil remains discovered at the site. No stone tools or signs of lithic industry have been found either, with the exception of three small pieces, one hewn from quartzite and the other two, quartz. Their typology is uncertain due to their small size. Neither was any signs found on the bones that are typical of human use, such as evidence of cut marks or fracturing by percussion. There is only one exception. Lynx sp. humerus PG-2010/01/50 has a series of incisions perpendicular to the long axis of the diaphysis. The incisions have a V-shaped section and display internal microstriation, which have been interpreted as cut marks and differ of those scores caused by carnivore activity (Figure 6). However, there is a tooth pit superimposed over these cut marks that were made by a carnivore, as depicted in Figure 6. Table 5 shows the percentages of others bone surface modifications.

sample, as only remains with enough conservation to lend themselves to identification have been considered. After conducting the taphonomic study on the bone collection at the deposit, marks from carnivore activity were estimated to be the most abundant alterations both in the upper and middle units. Thus, 38.8% of the material analysed from the upper unit shows modifications allocated to carnivore activity, and for the middle unit this frequency is 43.96% (Table 3). As can be seen in Table 3, the most characteristic marks both in the upper and

LSU MSU

Fracture outline

Longitudinal 26.27 20.00

Transversal 51.90 55.29

Curved 21.84 24.71

Fracture angle Oblique 50.65 54.35

Right 22.08 28.26

Mixed 27.27 17.39

Edges Smooth 43.12 66.67

Jagged 15.60 10.42

Crenulated 20.64 22.92

Shaft circunference*

< 1/2 27.27 11.76

> 1/2 10.74 2.94

Complete 61.98 85.29

Shaft fragment*

<1/4 50.00 45.71

1/4-1/2 30.33 28.57

1/2-3/4 11.48 22.86

>3/4 8.20 2.86

* Long bones only

Table 2. Fracture properties in Zarzamora and Búho Caves assemblage.

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Nonetheless and in low percentages, the upper unit does include minerals that are not present in the insoluble residue of the dolomite (potassium feldspar and plagioclase), although its composition is fundamentally dolomitic (Sala et al., 2011). This could indicate slight allochthonous contributions to the upper unit and, therefore, facies at the cave entrance. Furthermore, the analysis of

Discussion

Fossilipherous sedimentary packages from the three Zarzamora Cave sections are primarily autochthonous, meaning that sediment proceeds from the arenisation of the cave rock, which thus means that it was not responsible for the incorporation of skeletons inside the karstic system.

Figure 5. Three-dimensional bar diagrams showing relative frequencies of shaft length by shaft circumference in large size (superior image) and medium size ungulates (inferior image). Shaft length categories are: 1: <1/4 of the original length; 2: 1/4 to 1/2; 3: 1/2 to 3/4 and 4: >3/4 or complete. Shaft circumference categories are: 1: bone circumference is less than half of the original; 2: circumference is more than half, and 3: complete circumference in at least a portion of the bone length.

Large size ungulates

Middle size ungulates

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UPPER UNIT

Large sized ungulates

Long bones Flat bones Articular bones

Prox. ep. Diaphysis Dist. ep. Cortical Cancellous Compact bone

Pits 0.00 10.61 1.52 20.69 3.45 14.67

Scores 2.27 11.36 5.30 17.24 6.90 14.67

Punctures 0.76 0.00 0.00 0.00 0.00 0.00

Furrowing 6.82 7.58 15.91 37.93 10.34 30.67

Medium sized ungulates



Pits 0.00 7.14 0.00 50.00 0.00 6.25

Scores 0.00 28.57 0.00 25.00 0.00 6.25

Punctures 0.00 0.00 0.00 0.00 0.00 0.00

Furrowing 0.00 14.29 7.14 25.00 0.00 18.75

MIDDLE UNIT

Large sized ungulates



Pits 0.00 4.35 0.00 16.67 0.00 3.85

Scores 0.00 4.35 0.00 0.00 0.00 11.54

Punctures 0.00 0.00 0.00 16.67 0.00 7.69

Furrowing 0.00 4.35 4.35 50.00 0.00 30.77

Medium sized ungulates Long bones Flat bones Articular bones


Pits 0.00 0.00 0.00 0.00 0.00 0.00

Scores 0.00 0.00 0.00 0.00 0.00 22.22

Punctures 0.00 0.00 0.00 0.00 0.00 11.11

Furrowing 0.00 0.00 0.00 0.00 50.00 22.22

Table 3. Tooth mark frequencies of the Zarzamora and Búho Caves’ assemblage according to the distribution in the bone portions, geological unit and ungulate size.

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carnivores after humans. Thus, accumulation by humans at the Búho/Zarzamora Cave seems highly unlikely. There are several Iberian Pleistocene sites where carnivore remains do have anthropic markings. In the case of the Lynx, cases of using this felid have been documented at sites such as Peña de Estebanvela (Yravedra, 2005). They have found Lynx pardinus remains at this Magdalenian site in Segovia with taphonomic man-made alterations that seem to point to using this animal’s meat at the very end of the Late Pleistocene epoch (Yravedra, 2005). Cases of larger felids have been documented such as the level TD-10-1in Gran Dolina (Atapuerca, Burgos) (Blasco et al., 2010). Felid remains with cut marks have also been

weathering marks on the bones shows that the remains were not exposed to atmospheric agents prior to their burial. No evidence of water transport was found either, such as rounding or polishing of the bone remains. Thus, we can rule out this hypothesis and attend to geological and taphonomic factors. The hypothesis has been discussed that the skeletons may have been transported to the cave by a biological agent (carnivores, humans or both). Evidence is scarce or null of anthropic activity at this site. After the detailed taphonomic study of all bone remains at the site, only one case was logged of anthropic marks from removal of flesh. This remain also contains superimposed marks by carnivores over the cut marks, which proves access by

n Mean Min. Max. St. Desv. CI -95% CI + 95%

PUNCTURES

Length Cancellous 5 5.64 4.32 7.60 1.50 3.77 7.50

Thin cortical 5 5.59 3.95 6.92 1.17 4.14 7.04

Width Cancellous 3 5.29 4.51 5.95 0.73 3.48 7.10

Thin cortical 3 4.34 2.90 5.94 1.53 0.54 8.13

PITTING

Length

Cancellous 7 3.04 2.60 3.50 0.33 2.74 3.34

Thin cortical 8 2.50 1.13 3.40 0.73 1.88 3.11

Cortical 17 2.42 1.21 4.10 0.82 2.00 2.84

Width

Cancellous 3 2.07 1.72 2.48 0.38 1.12 3.02

Thin cortical 5 2.65 1.60 5.21 1.47 0.83 4.48

Cortical 11 2.04 0.84 3.27 0.72 1.56 2.53

SCORES Max. Width

Cancellous 26 1.78 0.83 3.73 0.57 1.55 2.01

Cortical 31 1.53 0.60 2.90 0.47 1.35 1.70

Thin cortical 10 1.61 0.53 3.12 0.84 1.01 2.21

Table 4. Statistical values of pits, punctures and scores analyzed in the sample of the Zarzamora and Búho Caves. The data displayed in the tables how length and width (n: number of cases; Max: maximum value; Min: minimum value; C.I.: Confidence Interval for mean and St. Desv.: Standard Deviation).

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Table 5. Bone surface modification frequencies of the Zarzamora and Búho Caves’ assemblage.

Figure 6. Lynx sp. humeral remain (PG-2010/01/50) showing cut marks (white arrows) with a tooth pit superimposed (black arrow).

% Surface modifications

Weathering, following Behrensmeyer (1978)

0 1 2 3 4 5 Upper unit 75.12 7.58 1.42 1.66 1.66 0.47 Middle unit 83.52 8.79 0.00 0.00 0.00 1.10

Concretion

0 1 2 3 Upper unit 32.46 31.99 18.01 15.40 Middle unit 53.85 29.67 9.89 1.10

Trampling (%) Upper unit 1.18 Middle unit 0.00

Root etching (%) Upper unit 12.80 Middle unit 17.58

Rodents activity (%)

Upper unit 16.35

Middle unit 13.19

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to destruction, both by carnivores and by geological processes. In addition to their density, the skeletal elements from the inferior appendicular skeleton belong to regions of the body with lower flesh mass which are thus less likely to be consumed by carnivores. Therefore, accumulation agents would not necessarily have had given preference to some parts of carrions over others in transporting them to the cave. For Cruz-Uribe (1991), the ratio of cranial/post-cranial elements tends to decrease with the size of the ungulates in hyena-induced associations. Stated differently, the post-cranial skeleton is better represented in large than small ungulates. The post-cranial skeleton tends to be well represented for large ungulates (ratio=1.16), whereas medium-sized ungulates are better represented by their dentition than by the post-cranial skeleton (ratio=0.47). For small ungulates, no post-cranial bone has been identified, only one small dental piece (ratio=0). This is related to hyenas’ capacity to reduce the carcasses of small ungulates to unidentifiable bone splinters. Fracturing pattern results show that there is a predominance of fractures with oblique and mixed angles with respect to the fracture angle. The properties of the fracture edges in the majority of cases are characterised by being smooth, with a high percentage of crenulated edges in the sample. These data suggest that the majority of the sample was fractured during a biostratinomic stage and that carnivores played a role in the fracturing processes (Villa & Mahieu, 1991). Furthermore, the extremely-high degree of fracturing in the sample suggests that the fracturing agent was very effective. Large ungulates reveal high fracturing levels with complete diaphyseal circumferences. As the size of the ungulate decreases, the quantity of fractures in long bones increases, until becoming unidentifiable

registered at the Foradada Cave site (Pantoja et al., 2011). Traces of human presence have also been recorded in other carnivore dens of the Iberian Peninsula, such as Cueva del Camino (Arsuaga et al., 2012) and Cueva de la Buena Pinta (Huguet et al., 2010), or other sites such as Wezmeh Cave (Iran) (Mashkour et al., 2009), Geula Cave (Israel) (Monchot, 2005), Les Auzières 2 and Bois Roche (France) (Marchal et al., 2009, Villa et al., 2010) and the Zourah Cave (Morocco) (Monchot & Aouraghe, 2009). Moreover, there is abundant evidence of carnivorous activity in the fossilipherous association at this site. However, it is not also simple to assign a taxon responsible for modifying and accruing these bones. The association of macromammals at the site include seven species of ungulates and six of carnivores. Spotted hyenas are the predominant carnivores in the association, where both adult and young specimens were found (Sala et al., 2011). For Cruz-Uribe (1991), the ratio of carnivore MNI/carnivore + ungulate MNI in faunal associations could be employed as a discernment criterion between man-made accumulations versus hyena-made accumulations. Thus, in accumulations made by hyenas, this ratio reaches values above 20%, whereas human accumulations are below 13%. At Zarzamora Cave, this ratio is 37.5%. This datum may be useful for discerning carnivore-made accumulations, albeit not necessarily produced by hyenas. Dentition, followed by the inferior appendicular skeleton (metapodial, carpal, tarsal and phalanges), have the best representation in the site’s faunal association. Based on these results, the skeletal representations found are believed to be related to the bone density of autopod elements, which make them more resistant

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reviewing all these criteria, Kuhn et al. (2010) suggested that the best discriminatory criteria for differentiating hyena-made accumulations are the presence of its coprolites and the presence of the remains of the species’ young. More than 26 coprolites assigned to the spotted hyena have been recovered from the site, both from the insides of the Zarzamora Cave and Búho Cave and at the Cata Exterior. Moreover, Crocuta crocuta juvenile individuals have been factored in, which entail almost half of the specimens represented. Therefore and according to these criteria, Zarzamora/Búho Cave site were probably used as a spotted hyena den during the Late Pleistocene epoch. Differences have been observed in post-depositional appearances between the upper and middle units. Firstly, there are in situ post-depositional fractures in the middle level and we can therefore conclude that it is a primary deposit without transport. Conversely, the upper level reveals greater concretion of fossil remain, a higher rate of rodent activity and, although in low proportions, evidence of trampling. Moreover, no post-depositional fractures were logged without transport. This fact may be explained by the abundant bioturbation that this level contains or by small-scale sedimentary transport at this level from other points of the karstic system. Conclusion No taphonomic differences were observed in the biostratinomic stage between macromammal associations at different site levels. Thus, the origin of accumulations in the different levels seems to be of the same nature. The high number of equids merits mention, both with respect to the number of remains and the

bone splinters. For Cruz-Uribe (1991), hyenas tend to consume the long bones from the epiphyses, generating diaphyseal cylinders. Conversely, humans tend to break the long bones from the centre of the diaphysis to access the marrow. It follows that in man-made accumulations, there are an abundance of complete epiphyses, while in hyena accumulations, there are many long bones consumed from the epiphyses. For Kuhn et al. (2010), this criterion is valid for discriminating between accumulations made by humans or carnivores, although not specifically by hyenas. Some of the fractures whose characteristics are assigned to post-depositional processes where the bones are dry have been logged onsite at the middle unit of the site without moving the different fractured fragments. Tooth-marks metric data have been compared at this site with those published earlier (Figure 7). Table 6 summarises the statistical values comparing the dimensions taken from site samples and measurements taken during experiments with carnivores today. With regard to the maximum measures taken in the cortical, the fossil sample is statistically different from the data obtained from bears and lions. The measurements of the Búho/Zarzamora Cave cannot be metrically differentiated with respect to wolves and hyenas. This may be due to the modification of the skeletal remains by juvenile hyenas. However, if we look at the ratio between the maximum and minimum dimensions taken in the cortical bone, we can see that data from the Búho/ Zarzamora Cave fall within the range of variation of C. crocuta (Figure 8). Recognition criteria for different bone accumulations at archaeological-paleontological sites have been debated for decades (Cruz-Uribe, 1991; Kuhn et al., 2010; Pickering, 2002; Stiner, 1991). After

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has little representation in any size. Skeletal representations are related to bone density and the associated flesh mass and, therefore, the accumulation agent or agents would not necessarily have given preference to some parts of carrions over others in transporting them to the cave. Evidence is scarce or null of anthropic activity at this site. Only one case

number of individuals represented. Spotted hyenas are the predominant carnivores in the association, where both adult and young individuals were found. Dentition followed by the appendicular skeleton are the best-represented elements in the site’s faunal association. The postcranial skeleton is better represented in the larger size ungulate. The axial skeleton

Figure 7. Mean ± 1 Standard Deviation of carnivore tooth pit sizes according to bone type and length/breadth recovered from Zarzamora site. A: Length in cancellous bone; B: breadth in cancellous bone; C: Length in dense cortical; D: breadth in dense cortical. Legend data from: *Domínguez-Rodrigo & Piqueras, 2003; ^Delaney-Rivera et al., 2009; ´´Saladié et al., 2013 and without symbol means this study together with Sala, 2012, Sala & Arsuaga, 2013 and Sala et al., in press.

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Búho/Zarzamora Cave site is interpreted as a hyena den based on the following criteria: the extremely high degree of fracturing in the sample suggests that the fracturing agent was very effective, given that the majority of fractures are biostratinomic fractures. The percentage of bone splinters is predominant, and grows higher the smaller the ungulate. Almost half of bone matter originating at the site shows some type of mark allocated to carnivore activity. Metric tooth-mark aspects allow discrimination

of man-made marks of removing flesh was recorded. This remain also contains superimposed tooth marks by carnivores over the anthropic marks, which proves later access by carnivores after humans. Thus, accumulation by humans at the Búho/Zarzamora Cave seems highly unlikely. Study data on fracturing patterns suggest that the majority of the sample was fractured during a biostratinomic stage and that carnivores played a primordial role in the fracturing processes.

Búho / Zarzamora C. lupus C. crocuta P. leo U. arctos

n U p n U p n U p n U p Length cortical (n=30)

274 3795,00 0,49 213 3184,00 0,98 134 869,5* 0,00* 15

89,5* 0,00*

Table 6. Results of the Mann Whitney U-test for the differences between the means in the Zarzamora and Búho samples and those from experimental data with living carnivores. Values with *are significant at p<0.05.

Figure 8. Relationship between maximum length and breadth of pits/punctures in the Zarzamora sample in relation with other carnivores (experimental own data).

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between hyenas and other carnivores analysed in experiments. The spotted hyena is the dominant carnivore in the fossil association. Bone remains both of adult and young individuals were found, as well as numerous coprolites assigned to this species. Acknowledgments The following organizations have funded and supported the excavation and analysis of the site: Caja Segovia, Junta de Castilla y León, Ministerio de Educación, cultura y deporte (FPU grants AP2006-04737 and AP2009-4096 to Nohemi Sala and Ana Pantoja), Fundación Atapuerca grants and Fundación Ancestros. Thanks to the Ministerio de Economía y Competitividad (CGL2009-12703-C03-03 project). We are grateful to the excavation and research team: Francisco Gracia, Adrián Pablos, Jorge Rodríguez, Eva Poza, Alejandro Bonmatí, Ignacio Martínez, Nuria García and Mª Cruz Ortega. We are indebted to many people that have allowed us access to important skeletal collections under their care and provided their kindly help: Gary Haynes, Curtis Marean and Mary Stiner. Thanks to Silvia Menéndez (Museo Geominero de Madrid) and Paloma Sevilla (UCM) for allowing us access to the material of the first excavation periods. Thanks also to the BBP group for their support and discussions. Thanks to the hominid-carnivore interaction organisation people. References Arsuaga, J.L., Baquedano, E., Pérez-González, A.,

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Sala, N., Arsuaga, J.L. & Haynes, G. (in press). Taphonomic comparison of bone modifications caused by wild and captives wolves (Canis lupus). Quaternary International, DOI: 10.1016/j.quaint.2013.08.017.

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