Get 20M+ Full-Text Papers For Less Than $1.50/day. Start a 14-Day Trial for You or Your Team.

Learn More →

Use of meat resources in the Early Pleistocene assemblages from Fuente Nueva 3 (Orce, Granada, Spain)

Use of meat resources in the Early Pleistocene assemblages from Fuente Nueva 3 (Orce, Granada,... Over the last few decades, several types of evidence such as presence of hominin remains, lithic assemblages, and bones with anthropogenic surface modifications have demonstrated that early human communities inhabited the European sub - continent prior to the Jaramillo Subchron (1.07–0.98 Ma). While most studies have focused primarily on early European lithic technologies and raw material management, relatively little is known about food procurement strategies. While there is some evidence showing access to meat and other animal-based food resources, their mode of acquisition and associated butchery processes are still poorly understood. This paper presents a taphonomic and zooarchaeological analysis of the Fuente Nueva-3 (FN3) (Guadix-Baza, Spain) faunal assemblage, providing a more in-depth understanding of early hominin subsistence strategies in Europe. The present results show that hominins had access to the meat and marrow of a wide range of animal taxa, including elephants, hippopotami, and small- and medium-sized animals. At the same time, evidence of carnivore activity at the site suggests that these communities likely faced some degree of competition from large predators when acquiring and processing carcasses. Keywords Early Pleistocene · Taphonomy · Hominin-carnivore interactions · Zooarchaeology · Cut marks · Palaeolithic archaeology * José Yravedra Department d’Historia i Historia de l’Art, Universitat Rovira joyravedra@hotmail.com I Virgili (URV), Tarragona, Spain Institut Català de Paleoecologia Humana I Evolució Social, Department of Prehistory, Ancient History and Archaeology, Tarragona, Spain Complutense University of Madrid, Madrid, Spain Prehistory Area, Universitat Rovira I Virgili, Tarragona, C.A.I. Archaeometry and Archaeological Analysis, Spain Complutense University, Madrid, Spain Museum of the First Settlers of Europe “Josep Gibert”, Orce, Department of Prehistory and Archaeology, University Granada, Spain of Sevilla, Sevilla, Spain Archaeological and Ethnological Museum of Granada, Department of Cartographic and Land Engineering, Higher Granada, Spain Polytechnic School of Avila, University of Salamanca, Avila, Spain Department of Geology, Unitat d’Estratigrafia, University of Barcelona, Barcelona, Spain Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Department of Prehistory and Archaeology, University of Granada, Granada, Spain School of Archaeology, St. Hugh’s College, University of Oxford, Oxford, UK Institute of Peace and Conflict Research, University of Granada, Granada, Spain History and Arts Doctoral Program, University of Granada, Granada, Spain GEOREC Master Program, University of Granada, Granada, Spain Vol.:(0123456789) 1 3 213 Page 2 of 23 Archaeol Anthropol Sci (2021) 13:213 scavenging, as proposed by Bunn & Ezzo 1993); (3) when Introduction hominins encountered an animal carcass dead from natu- ral causes. The first scenario entails a series of important Recent publications have amply demonstrated that hom- behavioural capacities among early hominins, since hunt- inins inhabited Europe prior to the Jaramillo Subchron ing involves some degree of planning, cooperation, and (1.07–0.98 Ma). Thus, sites such as Barranco León, Fuente the ability to kill prey. Scenarios 2–3 also entail some Nueva 3 (FN3) (Barsky et al., 2010, 2015; Titton et al., degree of landscape knowledge through the monitoring 2019, 2020; Toro-Moyano et  al., 2009, 2010a, 2010b, of suitable procurement spots (Clark and Linares-Matás 2011, 2013), and Sima del Elefante in Spain (Parés et al., 2021), although some authors have highlighted a series 2006; Carbonell et al 2008), Le Vallonnet (Michel et al., of issues associated with them. For example, Treves and 2017), Bois-de-Riquet, and Pont-de-Lavaud in France Treves (1999) and Domínguez-Rodrigo (2002) have noted (Bourguignon et al., 2015; Despriée et al., 2018; Lozano- how scenario 2 is dangerous and unlikely, while the third Fernández et al., 2019), and Kozarnika cave in Bulgaria scenario is highly reliant on trophic pressures and sea- (Sirakov et al., 2010) as well as Pirro Nord in Italy (Arza- sonality (Blumenschine, 1986, 1989; Tappen, 1995; Clark rello et al., 2016; Pavia et al., 2012) have provided exten- and Linarés-Matás, 2021), which renders recurrent and sive evidence of hominin activities older than 1 Ma. The reliable access to carcasses rather unlikely (Domínguez- studies carried out so far in most of these early sites have Rodrigo (2002; Yravedra 2006: 162) beyond opportunis- focused on improving our understanding of the technologi- tic exploitation of catastrophic carrion pulses. From the cal behaviours reflected in the lithic assemblages, docu- 1970s and 1980s, Binford (1981) challenged the “Man the menting the use of Oldowan technocomplexes (Mode 1) Hunter” paradigm of human subsistence by proposing that at most of these sites (Carbonell et al., 2008; Barsky et al., hominins were scavengers, kickstarting an intense “hunt- 2010, 2015; Sirakov et  al., 2010; Toro-Moyano et  al., ing versus scavenging” debate that has generated a vast 2011, 2013; de Lombera-Hermida et al. 2015; Arzarello academic literature involving several hypotheses and the et al. 2016; Bourguignon et al 2015). Nevertheless, com- elaboration of models derived from actualistic reference paratively little is still known about their food procurement frameworks (see Yravedra 2006; Parkinson 2018). After strategies, which included both plant matter and animal more than 30 years of debates, there is an emerging con- resources. In addition to the material traces of their activi- sensus that hominins had primary access to animal car- ties, the presence of these early human communities in casses of different sizes at Early Stone Age sites in Africa the European subcontinent has been directly demonstrated such as FLK-Zinj and BK (Olduvai Gorge, Tanzania), St through the discovery of hominin remains at sites such as (Peninj, Tanzania), FwJj14A, FwJj14B, and GaJi14 (Koobi Barranco León (Toro-Moyano et al. 2013) and Sima del Fora, Kenya), KJS (Kanjera, Kenya), and Swartkrans Elefante (Carbonell et al., 2008). (South Africa) (Monahan, 1996, Domínguez-Rodrigo With regards to the exploitation of animal resources, (2002; Domínguez-Rodrigo et al. 2002, 2009; Pickering some evidence is known of cut-marked bones, providing et al., 2008; Pobiner et al., 2008; Bunn & Pickering, 2010; direct evidence of hominin access to meat resources. In Ferraro et al., 2013; Parkinson, 2018; Oliver et al., 2019; some cases, such as Kozarnika cave (Sirakov et al., 2010), Clark & Linares-Matás 2021). Trlica (Vislobokova et al., 2020), or Pirro Nord (Cheheb In Europe, Sima del Elefante (Atapuerca, Spain) is the et al., 2019), sample sizes are small and present ambiguous only site older than 1 Ma with conclusive evidence of early distribution patterns. This unfortunately limits the extent human access to animal carcasses (Huguet et  al., 2013, to which inferences can be inferred about procurement 2017). At this site, 5% of cervid, bovid, and equid remains modalities and the order of which hominins had access from different anatomical parts possess cut marks, evidenc- to these carcasses (primary or secondary). The timing of ing butchery activities such as skinning, dismembering, and hominin access to carcasses is nonetheless a fundamen- defleshing (Huguet et al., 2017). Moreover, the presence of tal issue for understanding the subsistence strategies of percussion marks in this highly fragmented assemblage evi- Early Pleistocene populations. When conceptualising dences the exploitation of bone marrow. Alongside anthro- carcass acquisition strategies, primary access implies that pogenic evidence, the Sima del Elefante faunal assemblage hominins processed the remains before any other predator, also has some taphonomic evidence of carnivore activity while secondary access implies that hominin consumption (5%), resulting from the action of medium-sized carnivores, of animal resources took place after other predators had i.e. wolves or hyenas, according to Huguet et al. (2017). In already been feeding upon the carcass. contrast with the distribution of cut marks, most tooth marks Primary access is only possible in either of these sce- are found on fat-rich elements, such as vertebrae, ribs, and narios: (1) when hominins hunted prey; (2) when hominins epiphyses/metadiaphyses, a pattern generally associated with dispossessed another carnivore of its prey (confrontational secondary access, according to Huguet et al. (2017). 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 3 of 23 213 At Orce, a model of hominin-carnivore competition has provide a more developed understanding of the subsistence been proposed for both Barranco León and FN3 (Rodríguez- strategies employed by the Early Pleistocene populations Gómez et al., 2016; Espigares et al., 2019). At FN3, the at FN3. discovery of an elephant carcass with nearby lithic imple- ments and giant hyena (Pachycrocuta brevirostris) copro- lites was interpreted as a place of encounter and competition between both agents (Espigares et al., 2013). The study of The FN3 archaeological site the bone assemblages from Barranco León and FN3 in Espi- gares et al. (2019) shows the remains from multiple animals FN3 is an Early Pleistocene archaeological site located modified by humans and carnivores, interpreted as homi- 7 km west of the town of Orce (Granada, southern Spain), nins having secondary access to the carcasses. However, situated in the northeasternmost part of the Guadix-Baza the results of Espigares et al. (2019) have been challenged Basin (Toro-Moyano et al., 2010a, 2010b) (Fig. 1). This by Domínguez-Rodrigo et al. (2020), who reassessed some intermontane basin was formed in the upper-middle Mio- of the evidence. In any case, it is unexpected for Espigares cene after the closure of one of the Betic corridors between et al. (2019) to claim that hominins had secondary access the Mediterranean and the Atlantic (Hüsing et al. 2010). to carcasses when they are documenting evisceration cut From the Upper Miocene to the Middle-Upper Pleistocene, marks on the ventral side of the ribs and vertebrae, as well this area has been interpreted as an endorheic continen- as defleshing cut marks on long bone diaphyses (Espigares tal basin, that is, a closed drainage basin, which eventu- et al., 2019, SF Table S12). Furthermore, carnivore tooth ally formed the saline Baza lake (Anadón et  al., 1994; marks are relatively infrequent in these assemblages. As Anadón and Gabàs, 2009), around which various archaeo- such, there is scope for putting forward alternative hypoth- palaeontological localities such as Barranco León, Fuente eses to those presented by Espigares et al. (2019) and other Nueva, or Venta Micena have been observed (Fig. 1). The authors (e.g. Martínez-Navarro et al. 2003, 2010; Palmqvist waters of this endorheic system were drained by the chang- et al. 2005, 2011; Rodríguez-Gómez et al., 2016). ing course of the Guadalquivir River and its tributaries. Through the present zooarchaeological and taphonomic Today, the basin preserves a succession of alluvial (Gua- study, which analyses the faunal remains excavated during dix) and colluvial deposits (Baza: lacustrine clays, silts, the 2017–2020 fieldwork seasons at FN3, we expand the and sands with evaporitic limestone crusting), reaching FN3 assemblage and assess the hypotheses put forward Espi- up to 100 m thick and covering a timescale ranging from gares et al. (2013, 2019). The new data may enable us to the Upper Miocene to the Upper Pleistocene. Over the Fig. 1 Location of the FN3 archaeological site 1 3 213 Page 4 of 23 Archaeol Anthropol Sci (2021) 13:213 last half-century, numerous fossiliferous and archaeologi- Since the discovery of FN in 1991 by Alain Bocquet, cal localities have been documented (Maldonado-Garrido and until 2016, several research projects have demonstrated et al., 2017). important associations between the faunal remains and the FN3 is located near the margins of the lake (Fig.  1). lithic industries from this locality (Gibert et al., 1992, 1998; The age of FN3 has been determined using both relative Roe, 1995; Tixier et al., 1995; Turq et al., 1996; Martínez- and absolute dating methods, employing both micro- and Navarro et  al., 1997, 2010; Toro-Moyano et  al., 2003). macro-vertebrate biochronological calibrations with mag- Since 2017, a new research project under the direction of neto-stratigraphical data (Agustí et al., 1987, 1996, 2007, the University of Granada has undertaken fieldwork at the 2010; Martinez-Navarro et  al., 1997, 2003, 2010; Oms site. The present study explores the zooarchaeological and et al., 1999, 2000a, 2000b; Agustí and Madurell, 2003; Scott taphonomic insights from the faunal assemblage retrieved et  al., 2007; Lozano-Fernández et al., 2015a), as well as from field campaigns carried out between 2017 and 2020. combined U-series/ESR dating of quartz grains and tooth The taxonomy of the faunal assemblages represented in enamel (Duval et al., 2011, 2012a, 2012b). The stratigraphi- the archaeo-palaeontological sites of Orce has undergone cal sequence at the site (Fig.  2) correlates to the Matuy- numerous changes over the last five decades (see Luzón ama Chron, situated between the Olduvai and Jaramillo et al., 2021, Supplementary Notes S2 for a further discus- subchrons (1.78–1.48 Ma 1.07–0.98 Ma, Gradstein et al., sion). Nevertheless, we present an updated list of the site 2005). In particular, an age of around 1.2 Ma is accepted for of FN3. These include carnivores, such as the bear Ursus the site of FN3, on the basis of the derived features of the etruscus (Medin et al., 2017), the canids Canis mosbachensis rodents Allophaiomys aff. lavocati (Agustí and Madurell, (Luzón et al., 2021, Supplementary Notes S2), Xenocyon 2003; Lozano-Fernández et al., 2015b). (Lycaon) lycaonoides (Luzón et al., 2021, Supplementary Fig. 2 Stratigraphic sequence of FN3 (after Oms et al., 2011; Reinoso-Gordo et al., 2020) 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 5 of 23 213 Notes S2), and Vulpes alopecoides (Bartolini-Lucenti & The lithic industry of FN3 has been studied in detail by Madurell-Malapiera, 2020), the hyena Pachycrocuta brevi- Toro-Moyano et al. (2003, 2010a, 2010b, 2011, 2013) and rostris (Martínez-Navarro et al., 2010), Felidae indet. (Mar- Barsky et al. (2010, 2013, 2015), presenting typical Old- tínez-Navarro et al., 2010), the small felid Lynx cf. pardinus owan (Mode 1) artefacts. The lithic industry recovered in (Boscaini et al., 2015), the badger Meles meles (Madurell- FN3 level 5 represents 5% of the archaeological material Malapeira et al., 2011), and the mustelid Martellictis ardea recovered at the site, compared to the fauna that comprises (Ros-Montoya et al., 2021); and large herbivores, such as 95%. Flint is the predominant raw material, followed by the proboscidean Mammuthus meridionalis (Ros-Montoya, limestone, both of which were obtained from local sources 2010), the rhino Stephanorhinus cf. etruscus (Luzón et al., (Barsky et al., 2010, 2015). The predominant knapping tech- 2021, Supplementary Notes 2), two horses Equus altidens niques include bipolar knapping on anvil, as well as direct and E. sussenbornesis (Alberdi, 2010), the hippo Hippo- percussion knapping, used to produce small-sized flakes and potamus antiquus (Martínez-Navarro et  al., 2010), the choppers (Zaidner, 2013; Barsky et al., 2010, 2015). Lime- bovid Bison sp. (Martínez-Navarro et al., 2010), the caprids stone blocks and some cobbles were mainly used as heavy- Ammotragus europaeus (Martínez-Navarro et  al., 2010) duty tools. Although both flint and limestone were used to and Capra cf. alba (van der Made et al., 2008), and the knap small flakes, the abundance of limestone percussion cervids Praemegaceros cf. verticornis and Metacervocerus tools indicates a clear, task-related differentiation between rhenanus (Abbazzi, 2010). From a paleoecological point the two materials (Toro-Moyano et al., 2010a; Barsky et al., of view, two variables have been calculated both based on 2015). As in other Oldowan assemblages, the toolkit is non- dental ecometrics in large herbivorous mammal guilds of standardized and contains few retouched items. In addition, FN3 and Barranco León: Mean Annual Precipitation (MAP) some heavy-duty scrapers have been identified in the macro- and Net Primary Production (NPP). Results have provided toolkit (Barsky et al., 2018). Stone tool production was car- equal values for MAP (602 mm) and NPP (914 g(C)/m /a) ried out with hard hammers, using both bipolar-on-anvil and in both sites (Saarinen et al., 2021). It is worth noting that free-hand percussion methods. the MAP calculation on dental ecometrics is slightly lower than such calculated by Mutual Ecogeographic Range (FN3 MAP: 738 mm; Blain et al., 2016). In addition, drier climatic Materials and methods conditions (MAP: 449 mm) and lesser productivity (NPP: 747 g(C)/m /a) have been calculated from the dental features The materials analysed for this study were retrieved from of the Venta Micena herbivore community (Saarinen et al., levels 1–6 of FN3 during the 2017–2020 field seasons, with 2021). most remains coming from level 5 (Table 1). This faunal The micromammals from FN3 include the insectivores assemblage encompasses the fossils bones obtained during Erinacenae indet., Crocidura sp., Sorex minutus, Sorex sp., the new field seasons from 2017 to 2020, retrieved from 2 2 Asoriculus gibberodon, and Galemys sp. (Agustí et  al., 44 m in level 5, and 51 m in all other levels. This mate- 2010); and the rodents, Allophaiomys aff. lavocati, Allo- rial expands the faunal assemblages discussed by previous phaiomys sp., Mimomys savini, Castillomys crusafonti, research. Apodemus aff. mystacinus, and Hystrix sp. (Agustí and Faunal remains were quantified by the number of iden - Madurell, 2003; Agustí et al., 2007, 2010). tifiable specimens (NISP), minimun number of elements The herpetofaunal list of FN3 is somewhat poorer (13 (MNE), and minimum number of individuals (MNI). MNI taxa in total) than the nearby site of Barranco León with estimates considered element side and ontogenetic age four anurans (Discoglossus cf. jeanneae, Pelobates cultripes, (Brain, 1969). Mortality profiles were developed by assign- Bufo bufo s.l., and Pelophylax cf. perezi), four lizards (Chal- ing relevant elements/individuals to one of four categories: cides cf. bedriagai, Timon sp., indeterminate small lacertids, infant, juvenile, prime adult-adult, and senile, based on tooth and cf. Ophisaurus), and five snakes (Coronella girondica, eruption and crown wear. Natrix maura, Natrix natrix s.l., Zamenis scalaris, and Skeletal part profiles were based on a division of the skel- Malpolon monspessulanus) (Blain, 2005, 2009; Blain and eton into the following anatomical regions: cranial (i.e. horn, Bailon, 2010; Blain et al., 2011, 2016; Sánchez-Bandera cranium, mandible, and teeth); axial (vertebrae, ribs, pel- et al., 2020). Some fish and chelonian remains are also pre- ves, and scapulae, according to Yravedra and Domínguez- sent in the faunal assemblage. Rodrigo (2009)); upper appendicular elements (humerii, The stratigraphic sequence of FN3 was first described by femora); intermediate appendicular limbs (radii, tibiae, Turq et al. (1996). Nevertheless, it has since been redefined patellae, ulnae); and lower appendicular elements (metapo- by Oms et al. (2010), recognizing 12 stratigraphic levels dials, carpals, tarsals, phalanges, and sesamoids). Long limb (Fig. 2), among which level 5 stands out as it contains most bones were further divided into anterior elements (scapu- of the macro-vertebrate and lithic remains. lae, humerii, radii, ulnae, carpals, and metacarpals), as well 1 3 213 Page 6 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 1 Taxonomic NISP MNI representation in the zooarchaeological assemblages Layer 2 3 5 6 Total 2 3 5 S/A/J/I 6 Total of FN3, based on the faunal Mammuthus meridionalis 1 389 3 393 1 4 0/2/1/1 1 6 remains retrieved from the 2017–2020 field seasons, Stephanorhinus etruscus 1 18 1 20 1 1 0/1/0/0 1 3 quantified in terms of NISP Equus altidens 5 5 1 0/1/0/0 1 and MNI. Age classes: S: Equus sp. 2 1 27 30 1 1 2 0/2/0/0 4 seniles; A: adults; J: juveniles; Hippopotamus antiquus 53 53 3 0/1/1/1 3 I: infants. The Bold entries reflected the total Bison sp. 8 8 2 0/2/0/0 2 Bovidae 3 1 18 22 1 1 2 0/2/0/0 4 Praeovibos sp. 1 1 1 0/1/0/0 1 Capra alba 6 6 1 0/1/0/0 1 Sorgelia minor 1 1 1 0/1/0/0 1 Cervidae 1 36 37 1 3 0/2/1/0 4 Metacervocerus rhenanus 1 2 3 1 1 0/1/0/0 2 Praemegaceros cf. verticornis 1 5 6 1 1 0/1/0/0 2 Artiodactyla size 2 1 3 4 Artiodactyla size 3 1 1 Artiodactyla size 3b 6 6 Oryctolagus lacosti 6 6 1 0/1/0/0 1 Ursus etruscus 3 3 1 0/1/0/0 1 Carnivore indet 3 3 1 1 Carnivore size 2 1 1 1 0/1/0/0 1 Chelonia 35 35 1 0/1/0/0 Ave 1 1 1 Mammalia indet size 0 4 4 Mammalia indet size 1 6 6 Mammalia indet size 2 6 59 65 Mammalia indet size 3 1 6 47 1 55 Mammalia indet size 3a 1 1 14 16 Mammalia indet size 3b 2 3 118 123 Mammalia indet size 4 1 9 10 Mammalia indet size 5 5 2 99 106 Indet 84 81 3567 39 3771 Total 100 107 4550 44 4801 as posterior elements (pelves, femora, tibiae, patellae, tar- carnivores (e.g. wolves); and large carnivores (e.g. lions, sals, and metatarsals). Herbivore and carnivore specimens hyenas), following Espigares et al. (2019). that could not be identified at a species level, but were not Several procedures were followed to reconstruct site entirely considered indeterminable, were assigned a weight/ formation processes, assessing and evaluating both site size class. The categories used for herbivores follow the integrity and the contribution of various biological agents adaptation made by Espigares et al. (2019) of the classifi- to the faunal assemblage. Bone fragmentation was ana- cation system developed by Bunn (1982): microfaunal (0), lysed according to three variables; first, bones were divided including species weighing less than 25 kg; very small size into several categories according to their length: < 3 cm, (1), including macro-vertebrates species weighing 25–50 kg; 3.1–5.0 cm, 5.1–10 cm, and > 10 cm; secondly, bones were small size (2), including species weighing 50–125 kg; inter- classified based on the nature of their breakage planes, i.e. mediate size (3), including species weighing 125–500 kg, green or dry fractures, following Villa and Mahieu (1991). with an additional division between 3a (125–250 kg) and Criteria used to diagnose dry breaks included the existence 3b (250–500 kg); large size (4), including species weigh- of abundant breaks that are longitudinal and/or transverse ing 500–1000 kg; and very large size (5) for species weigh- to the axis of the bone, as well as breakage planes that are ing > 1000 kg. Carnivores were classified according to three uneven, rough, and in possession micro-step fractures. size classes: small carnivores (e.g. foxes); intermediate Dry breaks are further characterized by cortical medullary 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 7 of 23 213 surface angles that are close to 90°. In contrast, specimens rhinoceroses, are well represented, alongside animals better broken when fresh frequently have smoother surfaces and adapted to wooded environments (such as cervids), general- more abundant oblique breakage planes. Lastly, bone dia- ist animals adapted to rocky environments (such as caprids), physis preservation was recorded according to the relative and animals that tend to rely on watercourses (such as hippo- length of their fragments, as well as circumference types; potami). In relation to the latter, other non-herbivore species where type 1 refers to specimens with < 25% of the shaft cir- such as turtles are also present. Mortality patterns show a cumference intact; type 2 refers to specimens with 25–50% predominance of adults in all levels across all taxa, with only of the shaft circumference intact; and type 3 refers to speci- level 5 yielding infant or juvenile individuals, found among mens with > 50% shaft circumference. proboscideans, hippopotami, and cervids (Table 1). The impact of u fl vial alterations was estimated with frag - The skeletal profiles analysed for levels 2, 3, and 6 are ment size distributions, as well as the presence of abrasion, not very representative due to their small sample size (see polishing, and rounding on bone surfaces. Rounding and Suppl. File). The level 5 sample is larger, but with the excep- abrasion were additionally classified into different stages; tion of size 3 animals, most size classes are generally poorly light, intermediate, and intense. Weathering intensity was represented, and tend to show a predominance of cranial ele- analysed following Behrensmeyer (1978). Bone surface ments due to the overrepresentation of heavily fragmented modification analyses were carried out using hand-held teeth and ivory tusks. Small animals (sizes 0–2) also show lenses at 10–40 × magnification. Tooth marks were classified some presence of axial and appendicular elements. Among as pits, scores, or punctures, while furrowing was also ana- medium-sized animals (size 3), all the skeletal portions are lysed following the criteria established by multiple authors represented including the axial and appendicular elements, (Binford, 1981; Blumenschine, 1995; Blumenschine et al., even if cranial remains are clearly overrepresented (see 1996). Cut and percussion marks were classified according Tables 2–3 and Suppl. File). Among the appendicular ele- to Binford (1981), Blumenschine and Salvaggio (1988), and ments, we observe that both forelimbs and hindlimbs have Blumenschine et al. (1996). Modifications were quantified a fairly similar representation. Very few remains could be for specimens with well-preserved bone surfaces in terms of attributed to large animals (size 4), while very large animals NISP values. Other processes, such as manganese staining (size 5) are also represented mainly by cranial elements (see or biochemical alterations, were recorded using the criteria Tables 2–3 and Suppl. File). outlined by Fernández Jalvo and Andrews (2017). Taphonomic analyses Results The faunal assemblage shows a very high degree of frag- mentation across all levels, with a predominance of Zooarchaeological analyses bones < 3  cm, rendering more than 86% of the analysed sample indeterminable (Table 4). Among long limb bones, The sample analysed comprises 4801 remains, with level the high level of fragmentation has considerably influenced 5 being the one that has provided the largest volume of their circumference index and relative length values. In this remains (n = 4550), of which 14% were considered determi- regard, more than 90% of long bones from level 5 have less nable remains (Table 1). The other levels have only provided than 25% of their shaft circumference and less than 25% of a relatively small sample; in levels 2 and 3, hardly more than their relative length (Table 4). In levels 3 and 6, the sam- 100 remains per level have been found and only 6% of them ple is very small but also shows high fragmentation rates. could be determined. In level 6, the sample is even more lim- This high degree of fragmentation is likely due to diverse ited, with only 44 remains documented. Remains identified biostratinomic processes that have induced green fractures at the site belong to several herbivore and carnivore taxa. in > 40% of the level 5 bone assemblage, and in > 50% of the Among the herbivores, Mammuthus meridionalis stands bone samples from levels 2, 3, and 6 (Table 4). out with 62% of the determinable remains, and 14% of the Although bone fragmentation is high, cortical surface MNI. The second best-represented taxa is Hippopotamus preservation in the assemblage is very good (> 75% of the antiquus with 8.5% of the remains and 10.7% of the indi- bones are in a good state of preservation), and only level 2 viduals. Artiodactyla (cervids, caprids, and bovids) are bones are poorly preserved (Table 5). This good state of cor- better represented than equids. Finally, there are several tical preservation is likely due to the rapid burial rates that carnivore remains, of which only an ursid (Ursus etruscus) the assemblage experienced, as indicated by the very low could be identified at a species level. From a palaeoeco- incidence of weathering alterations: over 99% of the sam- logical perspective, the species represented belong to a wide ple exhibit slight or no weathering (Table 3). Furthermore, range of different ecological settings. Species associated waterborne alterations (abrasion, polishing, or rounding) with open environments, such as proboscidians, equids, and are not particularly common, with only 13% of bones from 1 3 213 Page 8 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 2 Skeletal profiles Species size level 5 according to NISP of the level 5 of FN3 assemblage from the NISP 0 1 2 3 3a 3b 4 5 Carniv 1 Carniv 3 Indet Total 2017–2020 field seasons. See Ivory 228 228 complete information in SF. The Bold entries reflected the total Cranial 1 1 2 1 3 8 Mandible 1 3 1 5 Tooth 2 31 18 1 55 202 3 120 432 Vertebrae 4 3 8 4 19 Rib 4 14 16 3 33 4 32 1 122 229 Scapule 1 1 1 1 4 Humerus 3 1 3 7 14 Radius 1 2 3 Ulna 1 1 Carpal 1 1 8 4 14 Metacarpal 3 3 Pelvis 3 1 4 Femur 1 2 1 1 6 3 14 Tibia 1 1 6 1 10 19 Metatarsal 1 1 4 1 7 Metapodial 2 1 6 1 7 1 18 Tarsal 1 1 2 Phalange 1 5 2 8 Indet 5 1 21 11 4 42 4 69 3302 3459 Placa 35 35 Total 11 6 85 65 16 196 9 548 2 3 3585 4526 Table 3 Skeletal profiles according to MNE of the level 5 of FN3 assemblage from the 2017–2020 field seasons. See complete information in SF. The Bold entries reflected the total Species size level 5 MNE 0 1 2 3 3a 3b Total size 3 % with tooth % no tooth 4 5 Carniv 1 Carniv 3 Ivory 5 Cranial 1 1 1 2 1.6 2.7 1 1 Mandible 1 2 2 1.6 2.7 1 Tooth 2 20 12 1 39 52 40.9 23 3 Vertebrae 2 1 3 4 3.1 5.3 Rib 1 3 3 1 10 14 11.0 18.7 2 8 1 Scapule 1 1 2 1.6 2.7 1 Humerus 1 1 1 2 4 3.1 5.3 Radius 1 1 1 0.8 1.3 Ulna 0.0 0.0 1 Carpal 1 1 8 10 7.9 13.3 4 Metacarpal 3 3 2.4 4.0 Pelvis 1 0.0 0.0 1 Femur 1 1 1 1 4 6 4.7 8.0 1 Tibia 1 1 5 1 7 13 10.2 17.3 Metatarsal 1 1 4 5 3.9 6.7 1 Metapodial 1 1 2 1 2 5 3.9 6.7 Tarsal 1 0.0 0.0 1 Phalange 1 4 4 3.1 5.3 2 Total 5 2 35 28 8 91 127 100.0 169.3 3 49 2 3 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 9 of 23 213 Table 4 Fragmentation patterns Taphonomic features Lev-2 Lev-3 Lev-5 Lev-6 of the FN3 bone assemblage (levels 2, 3, 5, and 6). The Bold Fragmentation Sample NISP 100 107 4550 44 entires reflected the total of Bones < 3 cm 82 87 3843 40 NISP of sample Bones 3.1–5 cm 12 10 416 3 Bones > 5.1–9.9 cm 6 10 218 1 Bones > 10 cm 73 % long bone with % long bone with green fracture 60 50 41 100 % Long bone with dry fracture 40 50 18.4 % long bone with indet fracture 40.6 % degree of long bone shaft circumference < 25% 75 91.1 100 26–50% 12.5 5.48 0 > 51% 12.5 3.42 0 % shaft length of long bones < 25% 87.5 95.9 100 26–50% 12.5 2.72 0 > 51% 1.4 levels 3 and 5 showing evidence of these alterations, always large species, and ribs, vertebrae, and shafts of intermediate of low or moderate intensity (Table 5). These alterations size species. Furthermore, the presence of percussion marks indicate that the assemblage did not experience considerable on long bones indicates that these bones were intention- water transport and that these different waterborne altera- ally fractured in order to access energy-dense within-bone tions were produced by small sedimentary particles sliding resources, such as marrow (Table 5). over the bone surface during processes of site formation. Very few specimens show calcitic concretions, and surface staining by manganese or oxides, which affects around 15% of the assemblage although it has not impacted bone pres- Discussion and conclusions ervation (Table 5). Biochemical alterations, such as root- etching or rodent damage, were relatively unimportant in The evidence reported in this study resembles the previous the FN3 assemblage. analyses of FN3 presented by Espigares et al. (2019). Both Carnivore activity has been recorded for levels 2, 3, and studies show similarly diverse taxonomic profiles with spe- 5, and if we exclude bone fragments < 2 cm, tooth-marked cies from different ecological settings, with a predominance bones can reach up to 11% in the smaller samples (levels in the MNI of very large animals, such as elephants and hip- 2–3). Nevertheless, overall tooth mark frequencies are lower popotami. The mortality patterns are also consistent, high- than 3%, with very few digested bones or instances of fur- lighting that the mortality profiles of sizes 1, 2, and 3 show rowing are present. Similarly, only one bone exhibited 5 or a predominance of adult individuals, whereas larger animals more tooth marks, suggesting that the incidence of carnivore present a higher percentage of infants and juveniles. Skeletal activity at the site is low. Tooth marks are usually 2–3 mm in part profiles also show a similar scarcity of axial elements diameter (with only one instance reaching 5 mm), and these and an overrepresentation of cranial elements, with teeth traces of carnivore activity are mostly found on the edges playing a major role in this regard. The presence of bones of ribs and long bones of small-, medium-, and large-sized from all skeletal portions suggests that some carcasses may animals (see Suppl. File). have arrived complete at the site. Hominins also contributed to the accumulation and On the basis of the presence and distribution of cut marks, modification of the faunal assemblage, as evidenced by the percussion marks, and tooth marks, both studies also agree presence of cut and percussion marks on bone elements. that humans and carnivores had access to the meat and Cut mark frequencies are lower than 1% (Table 5), although within-bone nutrients of several animal species at FN3. The they are found across all carcass sizes, including very large identification of cut marks on small-, medium-, and very animals, such as hippopotami or elephantids (Table 6). The large–sized animals such as elephantids or hippopotami distribution patterns of cut marks show that these anthropo- also indicate that hominins exploited multiple animal taxa. genic alterations appear on ribs, diaphysis of femur of very Carnivores have also left feeding traces on a wide range of 1 3 213 Page 10 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 5 Taphonomical alterations for FN 3 Taphonomic characteristics Lev-2 Lev-3 Lev-5 Lev-6 % lev-2 % lev-3 % lev-5 % lev-6 Sample NISP 100 107 455 44 Sample excluding teeth and Ivory 94 97 3821 37 Bone surfaces Samples with bad preservation 68 16 1101 7 68.0 15.0 24.2 15.9 Samples with good preservation 26 81 272 30 32.0 85.0 75.8 84.1 Weathering stage 0 97 105 4378 44 97.0 98.1 96.2 100 Weathering Weathering stage 1–2 2 2 162 0 2.0 1.9 3.6 0.0 Weathering stage 3–4 1 10 1.0 0.0 0.2 0.0 Hydrolic alteration 4 14 605 4 4.0 13.1 13.3 9.1 Water alteration Abrasion 4 10 293 2 4.0 9.3 6.4 4.5 Light stage abrasion 2 5 222 2 50.0 50.0 75.8 100 Intermediate stage abrasion 2 3 54 50.0 60.0 18.4 Intense stage abrasion 16 0.0 0.0 5.5 Polishing 3 5 154 3.0 4.7 3.4 0 Rounding 2 5 379 3 2.0 4.7 8.3 6.8 Light stage rounding 1 3 270 1 50.0 60.0 58.2 33.3 Intermediate stage rounding 1 1 116 1 50.0 20.0 25.0 33.3 Intense stage rounding 1 78 1 0.0 20.0 16.8 33.3 Calcitic concretions 1 14 1.0 0.0 0.3 0.0 Oxides (MN, FE) 8 18 681 13 8.0 16.8 15.0 29.5 Trampling alterations 2 2 99 1 7.7 2.5 3.6 3.3 Biochemical and root alterations 6 12 183 1 6.0 11.2 4.0 2.3 Bones with rodent tooth marks 1 0.9 Carnivore activity Bones with carnivore tooth marks 1 2 21 0 3.8 2.5 0.8 0 Bones with carnivore tooth marks 1 2 21 0 11.1 11.1 2.6 0 (excluded bone fragments < 2 cm) Bones with tooth pits 1 1 15 100 100 71.4 Bones with tooth scores 10 47.6 Bones with punctures 1 4.8 Bones with both pits and scores 1 3 50 14.3 Digestive alterations 1 Human activity Bones with cut marks 1 2 22 3.8 2.5 0.8 Bones with cut marks (excluded 1 2 22 11.0 11.0 2.7 bone fragments < 2 cm) Bones with peeling 1 Bones with percussion marks 1 14 1 0.0 1.2 0.5 3.3 carcass sizes, from smaller animals such as rabbits to very documented at FN3 with the results of several actualistic large animals, including elephants. frameworks. The butchery activities inferred from the distribution pat- These actualistic reference frameworks were developed terns of cut marks (Table 6) appear to be related to eviscera- by different authors (see SF 2) with the aim of establish- tion and defleshing activities, which, alongside the evidence ing empirically the order in which hominins and carnivores for percussion marks, suggests that hominins were accessing accessed animal carcasses. Carcasses of different sizes were the meat and marrow from different carcass sizes. However, made available to different human and carnivore agents in the presence of tooth marks also indicates that carnivores different experimental settings, in order to quantify the fre- played a role in the alteration of the faunal assemblage at quencies of taphonomic alterations that they generated. It the site. These taphonomic data are all consistent with the was noted that the frequency and distribution of cut marks, results described by Espigares et  al. (2019). In order to percussion marks, and tooth marks varied depending on ascertain the temporality of carcass access, we have sub- which agent(s) accessed the carcass first, which allowed sequently compared the cut and tooth mark frequencies researchers to empirically establish the relative temporality 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 11 of 23 213 Table 6 Cut-marked bones Taxon Species size Skeletal bone Function identified among the FN3 assemblage from the 2017–2020 Bovidae 3b Rib Disarticulation field seasons Mammalia indet 1 Rib Defleshing Mammalia indet 5 Rib Defleshing Mammalia indet 5 Rib Defleshing Hippopotamus antiquus 5 Femur diaphysis Defleshing Mammalia indet 3b Femur diaphysis Defleshing Mammalia indet 3 Long bone diaphysis Defleshing Mammalia indet 5 Long bone Indet Mammalia indet 3b Long bone Indet Mammalia indet 3b Long bone diaphysis Defleshing Mammalia indet 3b Long bone diaphysis Defleshing Mammalia indet Indet Indet Defleshing Mammalia indet 2 Indet Defleshing Mammalia indet 3a Indet Defleshing Mammalia indet Indet Indet Indet Indet Indet Indet Indet Mammalia indet 2 Pelvis Defleshing Mammalia indet 5 Pelvis Evisceration Mammalia indet 3b Tibia diaphysis Indet Mammalia indet 3 Tibia diaphysis Defleshing Mammalia indet 3b Tibia diaphysis Defleshing Mammalia indet 3b Vertebrae Defleshing Mammalia indet 3b Vertebrae Defleshing of animal resource acquisition. This approach has gener- happens when analysing the frequency of tooth marks on ally yielded positive results in taphonomic studies of Early limb bone diaphyses: the results obtained are well below the Pleistocene sites in Africa (Domínguez-Rodrigo, 2002; outcomes of actualistic observations of carnivore-first mod- Domínguez-Rodrigo et  al. 2002, 2009; Pickering et  al., els. Instead, the low tooth mark frequencies fit much better 2008; Pobiner, 2007; Pobiner et al., 2008; Pante, 2013; Par- the patterns left by carnivores when they engage with car- kinson, 2018; Yravedra et al., 2020). casses after human intervention, including those instances Nevertheless, it is worth noting that the comparative when carcasses were initially modified by vultures (Fig.  5). assessment of bone surface modifications for interpreting While tooth mark data seems to indicate that carnivores the order of carcass intervention at FN3 using experimental were not the primary agent responsible for the accumula- or actualistic frameworks is hindered by small sample sizes tion of the faunal assemblage at FN3, it is still necessary to (Figs. 3–7). conclusively prove the degree of human intervention through The very low frequencies of carnivore tooth marks docu- the analysis of the cut mark and percussion mark frequen- mented at FN3 are not consistent with the percentage ranges cies at the site. generated by carnivore-hominin models (Fig. 3), with the When the frequencies of percussion marks left by humans exception of cheetah or leopard accumulations of size 1–2 are compared with the actualistic framework, the very low animals. However, the FN3 assemblage has a considerable percentages found at FN3 do not readily fit any of the pro- representation of animals > 150 kg, a size range that neither posed models (Fig. 6), leading to a somewhat inconclusive cheetahs nor leopards target, which suggests that carnivores result. were not the main accumulation agents at FN3 (Fig. 3). Neither do the very low cut mark frequencies seem to Across all three categories of appendicular elements, correspond to primary anthropogenic accumulations either tooth mark frequencies are considerably lower than those (Fig. 7), resembling more closely the profiles that result from that would be expected in episodes of primary access by secondary hominin access to carcasses previously consumed carnivores (Fig. 4). ULB and ILB in particular have very by carnivores. Breaking down cut mark frequencies by low tooth mark frequencies, a pattern inconsistent with the upper, intermediate, and lower appendicular elements, ULB observations of carnivore feeding when they have access frequencies would match those associated with secondary to the most nutritious portions of the carcasses. The same access, while cut mark frequencies on diaphyses/bone shafts, 1 3 213 Page 12 of 23 Archaeol Anthropol Sci (2021) 13:213 Fig. 3 Tooth mark frequencies on appendicular elements from FN3 yellow box shows the tooth mark frequencies resulting from a model by size classes (1–2, 3, 4–5) in relation to the actualistic framework in which vultures accessed first the carcasses, followed by humans derived from carnivore feeding behaviours documented in different and then carnivores, while the red box documents the main range of contexts. The blue box highlights the range of tooth mark frequen- tooth mark frequencies associated with primary carnivore accumula- cies that carnivores tend to leave on bones during secondary scav- tions. See Suppl. file 2 for the full bibliographic details of the relevant enging activities after human abandonment of the carcasses. The sources Fig. 4 Tooth mark frequencies on appendicular remains from FN3 such as humerii and femoral remains, ILB refers to intermediate on animal size classes 1–2, 3, and 4–5, in relation to the actualistic limbs (tibiae, radii), while LLB stands for lower limb bones (i.e. framework derived from carnivore feeding behaviours documented in metapodials). See suppl. file 2 for more information on the compara- different contexts. ULB refers to tooth marks on upper limb bones, tive samples and their full bibliographic details intermediate, and lower elements are inconclusive since they available does not allow us to determine with certainty the overlap with models of both primary and secondary accesses temporality of hominin access to animal carcasses at FN3, (Fig. 8). Consequently, the taphonomic evidence currently 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 13 of 23 213 Fig. 5 Tooth mark frequencies on limb bone diaphyses from FN3 yellow box shows the tooth mark frequencies resulting from a model on animal size classes 1–2, 3, and 4–5, in relation to the actualistic in which vultures accessed first the carcasses, followed by humans framework derived from carnivore feeding behaviours documented in and then carnivores, while the red box documents the main range of different contexts. The blue box highlights the range of tooth mark tooth mark frequencies associated with primary carnivore accumula- frequencies that carnivores tend to leave on bones during secondary tions. See the Suppl. file 2 for the full bibliographic details of relevant scavenging activities after human abandonment of the carcasses. The sources Fig. 6 Percussion mark frequen- cies on appendicular elements from FN3 on the basis of carcass size (1–2, 3, and 4–5) in relation to a comparative actualistic framework. The blue box represents the percussion mark frequencies generated by humans when they have early access to carcasses, and the red box encompasses the percussion mark frequencies in contexts of secondary access. See Suppl. File 2 for bibliographic details of the reference samples employed a problem of ambiguity further compounded by the small while the low frequency of tooth marks does not match the sample sizes of bone surface modifications. expectations of primary carnivore access (Figs. 3–5). With respect to the temporality of carcass access, the When analysing the distribution and inferred functional- comparative assessment of cut mark, percussion mark, and ity of cut marks (Table 6, Fig. 8), we note their presence tooth mark frequencies in the FN level 5 bone assemblage on the diaphyses of long bones such as the femur or tibia with actualistic framework returned ambiguous and incon- (Figs. 9, 10, 11), as well as on axial elements such as ribs or clusive results. The low frequency of cut marks (Fig. 7 and the pelvis (Fig. 11). This pattern would be consistent with 8) resemble more a pattern of secondary hominin access, defleshing and evisceration activities on carcass elements 1 3 213 Page 14 of 23 Archaeol Anthropol Sci (2021) 13:213 Fig. 7 Cut mark frequencies on appendicular elements from FN3 (level 5) on the basis of carcass size (1–2, 3, and 4–5) in relation to the comparative framework generated by several experimental and actualistic studies. The blue box represents the cut mark frequencies gener- ated by humans when they have early access to carcasses, the yellow box corresponds to cut mark frequencies when human access followed the presence of vultures but preceded carni- vore access, while the red box encompasses cut mark frequen- cies from contexts of secondary access. See suppl. File 2 for the full bibliographic details of the comparative samples employed Fig. 8 Cut mark frequencies on appendicular remains from FN3 frequencies left by humans in early carcass access contexts, while the (level 5) on animal size classes 1–2, 3, and 4–5. ULB refers to tooth red box shows cut mark frequencies in contexts of secondary access. marks on upper limb bones, such as humerii and femoral remains, Cut mark frequencies on diaphyses are shown on the right hand side ILB refers to intermediate limbs (tibiae, radii), while LLB stands for of the graph. See suppl. file 2 for more information on the compara- lower limb bones (i.e. metapodials). The blue box represents cut mark tive samples and their full bibliographic details with high nutritional values, which may indicate early access At the same time, the low frequencies of anthropogenic to at least some of the carcasses, rather than secondary scav- traces do not allow to conclusively establish the nature of enging. This is especially relevant when considering actu- carcass acquisition strategies. The sample analysed in the alistic reports of carnivore feeding patterns to first consume present study thus experiences similar limitations than the the visceral packages. This pattern is documented among work conducted by Espigares et al. (2019): both have few a wide range of wild carnivores, such as felines (Schaller, bones with anthropogenic and carnivore alterations, and thus 1972; Blumenschine, 1986; Blumenschine & Cavallo, neither can determine in which order did humans and carni- 1992), canids (Mech, 1970; Stahler et al, 2006; Yravedra vores access prey carcasses at the site. As such, there is little et al., 2011), and hyenas (Kruuk, 1972; Blumenschine, 1995; evidence to support previous models of hominin acquisi- Faith, 2007). tion of animal resources at Orce consisting in the secondary 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 15 of 23 213 Fig. 9 Shaft of femur of hippopotamus with cut marks in FN3 level 5 Fig. 10 Shaft with cut marks and 3D modelled of FN3 level 5 1 3 213 Page 16 of 23 Archaeol Anthropol Sci (2021) 13:213 2016, according to Van Valkenburgh and Ruff, 1987). This circumstance would provide hominins with opportunities to scavenge sizeable chunks of muscle tissue before giant hyena individuals accessed the carcasses. This passive scav- enging scenario might help explain the cut mark distribution patterns and the scarcity of tooth marks that is observed by Espigares et al. (2019) documented at FN3. However, there are multiple lines of evidence that prompt a reconsideration of the interpretations proposed by Mar- tínez-Navarro and Palmqvist (1995), Palmqvist et al. (1996, 1999, 2005, 2011), Arribas and Palmqvist (1999, 2002), Espigares et al. (2013, 2019), and Rodríguez-Gómez et al. (2016) regarding the Megantereon-Hominins-Pachycrocuta model of carcass acquisition at FN3. Several authors have already shown in different publica- tions that skeletal parts and mortality profiles are generally inconclusive when modelling the behaviour of humans and carnivores, since the resulting outcomes can experience issues of equifinality (see discussion in Domínguez-Rodrigo (2002)). From a taphonomic perspective, it is worth bearing in mind that there are no empirical studies of the feeding behaviour of M. whitei, which precludes reliable interpreta- tions and inferences regarding their role in the alteration of fossil assemblages. The only available taphonomic study of an accumulation altered by machairodonts is described by Marean and Ehrhardt (1995). While the assemblage corre- sponds to Homotherium rather than M. whitei—which may have generated slightly different patterns—the high fre- quencies of bone damage, particularly in relation to those documented at FN3, are noteworthy, indicating that some sabretooth felids were capable of inflicting considerable damage to the bones of their prey. Moreover, Harstone-Rose (2008, 2011) has argued that Megantereon would have had carcass processing behaviours no more hypercarnivorous Fig. 11 Several cut marks of FN3 level 5 on animal size level 3, pel- than extant felids, which have been shown to generate con- vis of animal size level 5, and indeterminate fragment of level 5 siderable tooth mark damage (see e.g. Rodríguez-Alba et al. 2019 for jaguars; Brain 1969 and Arriaza et al. 2019 for leopards; or Arriaza et al. 2016, Haynes & Klimowicz 2015, exploitation of carcasses left by large felids, and prior to the and Haynes & Hutson 2020 for lions). Furthermore, bio- intervention of the giant hyena. mechanical studies of sabretooth dentition suggest that these Several authors, including Martínez-Navarro and carnivores would have had no issue making contact with Palmqvist (1995), Palmqvist et al. (1996, 1999, 2005, 2011), bone surfaces when they were feeding on animal carcasses Arribas and Palmqvist (1999, 2002), Espigares et al. (2013, (Bryant et al., 1995; Harstone-Rose, 2008; 2011; Desantis 2019), and Rodríguez-Gómez et  al. (2016), suggest, on et al. 2012). Therefore, the teeth of Megantereon whitei were the basis of skeletal part profiles, mortality patterns, and neither as fragile nor as inefficient as Van Valkenburgh and taphonomic evidence, that large felids, such as Megantereon Ruff (1987) and Palmqvist et al. (1996, 1999, 2005, 2011) whitei, would have had primary access to animal carcasses have proposed. at FN3. They assume that M. whitei individuals would have Regarding the action of the giant hyena, the low tooth only accessed part of their prey’s visceral package, since the mark frequencies documented at FN3 are in stark contrast size and fragility of their canines may have prevented them with the damage generated by hyenas when they are the pri- from consuming much meat (Palmqvist et al., 1996, 1999, mary agent of carcass modification documented by different 2005, 2011; Espigares et al., 2013; Rodríguez-Gómez et al., authors (Kruuk 1972; Blumenschine, 1986; Villa & Bartram, 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 17 of 23 213 1996; Villa & Soresi, 1998; Faith, 2007; Domínguez-Rod- marks derived from marrow extraction (Table 5). These pat- rigo et al., 2015; Haynes & Klimowicz, 2015; Fernández- terns suggest that hominins had recurrent access to meat and Jalvo & Andrews, 2017; Haynes & Hutson 2020). This marrow resources at FN3. divergence suggests that the action of giant hyena at FN3 The lithic evidence described by Toro-Moyano et  al. was likely more limited than previously envisioned. (2003, 2009, 2011, 2013) and Barsky et al. (2010, 2013, Espigares et al. (2013) describe a M. meridionalis carcass 2015) is also in support of these interpretations. The lithic surrounded by lithic industry and giant hyena coprolites at analyses at FN 3 show a predominance of small-sized single FN3, which indicate the presence of these carnivores at the flakes of flint and limestone obtained from local sources, site. Nevertheless, no tooth marks or anthropogenic marks and some choppers (Toro-Moyano et al., (2003, 2009, 2011, were observed on the elephantid bones, thus not conclu- 2013; Barsky et al. 2010, 2013, 2015). These heavy-duty sively proving the interaction of humans or carnivores with tools may be associated with percussive activities oriented the proboscidean carcass. in part to the fragmentation of bones, as suggested by the Cut marks demonstrating direct hominin engagement existence of bones with green fractures (Table 4) and per- with the remains have been found on Middle Pleistocene cussion marks (Table 5). Furthermore, use-wear analyses elephant carcasses (e.g. Yravedra et  al. 2010), and this of FN3 flakes conducted by Toro-Moyano et  al. ( 2003) absence of taphonomic alterations is also in stark contrast documented their use in butchery activities. Recent exper- with actualistic reports of carnivore engagement with ele- imental studies have shown that small flakes are suitable phant carcasses, which tend to present a high number of for defleshing activities, even on very large carcasses, such tooth marks (Haynes & Klimowicz, 2015; Haynes & Hutson as proboscideans (Starkovich et al., 2021; Marinellu et al., 2020). Therefore, this spatial association of lithic finds and 2021). Lower Palaeolithic sites with small, simple flake coprolites in association with these elephant carcasses may lithic assemblages associated with cut-marked remains are have been fortuitous, resulting from independent episodes indeed very common (see Marinellu et al., 2021 for a more that coalesced into a palimpsest through complex site for- in-depth discussion). mation processes. Such coincidental associations of stone Based on these taphonomic arguments, the possibility tools and faunal remains are not infrequent in the Palaeo- that hominins at FN3 had early access to meat from fresh lithic record. At Bois Roche, lithic artefacts were found carcasses prior to carnivore engagement with the remains inside a hyena den, product of fluvial transportation (Villa cannot be dismissed. However, low cut mark frequencies in & Bartram, 1996; Villa & Soresi, 1998). Likewise, associa- relation to actualistic models (Figs. 6–8) hinder the robustic- tions of lithic artefacts with proboscidean remains are not ity of the conclusions that can be established regarding the uncommon (Gaudzinski et al 2005). Nevertheless, many of timing of hominin carcass access at FN3. It is also worth these associations have been shown to be the result of spatial considering the possibility that hominins may have followed processes unrelated to human subsistence strategies (Yrave- different carcass procurement strategies on the basis of ani- dra et al., 2014; Haynes et al 2020). mal size (small, medium, large, or very large), given the On the basis of the evidence documented at FN3, there diverse range of animal carcass sizes at FN3 level 5 present- are several important inferences that can be made regarding ing evidence of anthropogenic alterations. the role of hominins and carnivores at the site. The low tooth A convergence of different processes at the site is also mark frequencies at the site (Table 5, Figs. 3–5) suggest that plausible, with some animal deaths by natural causes, sev- carnivore impact on the FN3 bone assemblage was relatively eral episodes of primary and/or secondary hominin access to unimportant. The scarcity of digested bones, the absence of prey, and sporadic carnivore feeding bouts. This hypothetical diaphysiary cylinders, bones with pit tooth marks > 5 mm, scenario would explain the low cut mark frequencies while or individual bones with more than 5 tooth marks all sug- also accounting for the small number of tooth marks docu- gest that the role of the giant hyena was rather limited in mented in the faunal assemblage. the formation of the FN3 faunal assemblage. In the future, Nevertheless, until the limited evidence is complemented a geometric morphometric assessment of tooth marks (cf. by a revision of the materials from previous excavations Aramendi et al., 2017; Courtenay et al., 2021) could provide and sample sizes are enlarged with further newly excavated valuable insights into the specific carnivore(s) that played a remains, it will not be possible to conclusively define what role in the accumulation and/or alteration of the bone assem- role did humans play in the accumulation of the faunal blage at the site. assemblage at FN3. Likewise, more advanced statistical Regarding the role played by hominins, it is worth high- techniques may be able to shine new light on the nature lighting the considerable degree of bone sample fragmen- of this assemblage. Additionally, the incorporation of new tation (Table 4) and the presence of cut marks associated technologies applied to the study of cut marks could help us with butchery practices such as disarticulation, defleshing, develop new approaches. For example, recent approaches and evisceration (Table 6, Figs. 9–11), as well as percussion may allow the identification of which raw materials and 1 3 213 Page 18 of 23 Archaeol Anthropol Sci (2021) 13:213 the article's Creative Commons licence and your intended use is not types of lithic implements were employed in the butchery permitted by statutory regulation or exceeds the permitted use, you will activities documented at the site (Courtenay et al., 2019; need to obtain permission directly from the copyright holder. To view a Linares-Matás et  al., 2019; Maté-González et  al., 2019; copy of this licence, visit http://cr eativ ecommons. or g/licen ses/ b y/4.0/ . Yravedra et al., 2017, 2019). Lastly, the current study provides further zooarchaeologi- cal data on hominin subsistence strategies in the Early Pleis- References tocene settlement of the European subcontinent. The present study demonstrates the complexities and issues inherent to Abbazzi L (2010) La fauna de cérvidos de Barranco León y Fuente the study and interpretation of Early-Middle Pleistocene Nueva 3. In: Toro I, Martínez-Navarro B, Agustí J (Eds.), Ocupa- ciones Humanas en el Pleistoceno Inferior y Medio de la Cuenca assemblages, given the limitations imposed by the repre- de Guadix-Baza. Junta de Andalucía, Consejería de Cultura, sentativity of faunal samples. Nevertheless, FN3 joins other Sevilla, 273–290 Early Pleistocene sites older than 1 Ma across Europe where Agustí J, Madurell J (2003) Los arvicólidos (Muroidea, Rodentia, hominins had access to animal carcasses, such as Kozarnika Mammalia) del Pleistoceno inferior de Barranco León y Fuente Nueva 3 (Orce, Granada). Datos preliminares. In: Toro-Moyano, (Sirakov et al., 2010), Trilika (Vislobokova et al., 2020), or I., Agustí, J., Martínez-Navarro, B. (Eds.), El Pleistoceno infe- Pirro Nord (Cheheb et al., 2019), although there are still rior de Barranco León y Fuente Nueva 3, Orce (Granada). Junta some unresolved questions related to the temporality of car- de Andalucía. Consejería de Cultura. E.P.G.P.C. Arqueología cass access or whether hominins exhibited different acquisi- Monografías, Sevilla, pp. 137–147. Memoria Científica cam- pañas 1999–2002 tion strategies on the basis of carcass size. Agustí J, Arbiol S, Martin-Suarez E (1987) Roedores y lagomorfos (Mammalia) del Pleistoceno inferior de Venta Micena (Depresión Supplementary Information The online version contains supplemen- de Guadix-Baza, Granada). Paleontologia i Evolució, Memoria tary material available at https://doi. or g/10. 1007/ s12520- 021- 01461-7 . Especial 1:95–107 Agustí J, Oms O, Garcés M, Parés JM (1996) Calibration of the Late Acknowledgements We thank the researchers and students involved Pliocene-Early Pleistocene transition in the continental beds in the excavations, recovery, and preparation of the archeo-paleonto- of the Guadix-Baza Basin (southeastern Spain). Quatern Int logical record from the FN3. Juan Manuel Jiménez Arenas belongs to 40:93–100 the Excellence Unit “Archaeometrical Studies. Inside the artefacts & Agustí J, Oms O, Parés JM (2007) Biostratigraphy, paleomagnetism ecofacts” (University of Granada) and the Junta de Andalucía Research and geology of the Orce ravine (Southern Spain) Comment on Project “HUM-607”. Last but not least, we would like to thank the the paper by Gibert et al. (2006). Quaternary Science Reviews reviewers of this paper for their fruitful comments that have greatly 26:568–572 improved the first version of this paper. Agustí J, Blain HA, Furió M, De Marfá R, Santos-Cubedo A (2010) The Early Pleistocene small vertebrate succession from the Orce Funding Open Access funding provided thanks to the CRUE-CSIC region (Guadix-Baza Basin, SE Spain) and its bearing on the agreement with Springer Nature. This research was funded by the first human occupation of Europe. Quatern Int 223–224:162–169 Junta de Andalucía, Consejería de Educación, Cultura y Deporte: Alberdi MT (2010) Estudio de los caballos de los yacimientos de Orce Research Project “Primeras ocupaciones humanas y contexto Fuente Nueva-3 y Barranco León-5 (Granada). In: Toro I, paleoecológico a partir de los depósitos pliopleistocenos de la cuenca Martínez-Navarro B, Agustí J (eds) Ocupaciones Humanas en Guadix-Baza: zona arqueológica de la Cuenca de Orce (Granada, el Pleistoceno Inferior y Medio de la Cuenca de Guadix-Baza España), 2017–2020 (Ref. BC.03.032/17)”. We also received sup- Junta de Andalucía. Consejería de Cultura, Sevilla, pp 291–306 port from the PALARQ Foundation with the convocatory of Analitics Anadón P, Gabàs M (2009) Paleoenvironmental evolution of the Early 2019: “Identificando Carnívoros a partir de análisis Tafonómicos de Pleistocene lacustrine sequence at Barranco León archeological última generación aplicando Fotogrametría y Morfometría Geométrica site (Orce, Baza Basin, Southern Spain) from stable isotopes de las Marcas de Diente. Aplicación a Yacimientos del Pleistoceno and Sr and Mg chemistry of ostracod shells. J Paleolimnol Inferior Ibérico: FN3, Venta Micena 3 y 4 (Granada), Pontón de la 42:261–279 Oliva (Patones, Madrid)”. Lloyd Austin Courtenay is also funded by Anadón P, Utrilla R, Julià R (1994) Palaeoenvironmental reconstruc- the Spanish Ministry of Science, Innovation and Universities with an tion of a Pleistocene lacustrine sequence from faunal assem- FPI Predoctoral Grant (Ref. PRE2019-089411) associated to project blages and ostracode shell geochemistry, Baza Basin, SE Spain. RTI2018-099850-B-I00 and the University of Salamanca. The Institut Palaeogeogr Palaeoclimatol Palaeoecol 111:191–205 Català de Paleoecologia Humana I Evolució Social (IPHES-CERCA) Aramendi J, Maté-González MA, Yravedra J, Ortega MC, Arriaza has received financial support from the Spanish Ministry of Science MC, González-Aguilera D, Baquedano E, Domínguez-Rodrigo and Innovation through the “María de Maeztu” program for Units of M (2017) Discerning carnivore agency through the three-dimen- Excellence (CEX2019-000945-M). sional study of tooth pits: revisiting crocodile feeding behaviour at FLK- Zinj and FLK NN3 (Olduvai Gorge, Tanzania). Palaeo- Open Access This article is licensed under a Creative Commons Attri- geogr Palaeoclimatol Palaeoecol 488:93–102 bution 4.0 International License, which permits use, sharing, adapta- Arriaza MC, Domínguez-Rodrigo M, Yravedra J, Baquedano E (2016) tion, distribution and reproduction in any medium or format, as long Lions as bone accumulators? Paleontological and ecological as you give appropriate credit to the original author(s) and the source, implications of a modern bone assemblage from Olduvai Gorge. provide a link to the Creative Commons licence, and indicate if changes PLoS ONE 11(5):e0153797. https:// doi. or g/ 10. 1371/ jour n al. were made. The images or other third party material in this article are pone. 01537 97 included in the article's Creative Commons licence, unless indicated Arriaza MC, Aramendi J, Maté-González MÁ, Yravedra J, Stratford otherwise in a credit line to the material. If material is not included in D (2019) Characterizing leopard as taphonomic agent through 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 19 of 23 213 the use of micro-photogrammetric reconstruction of tooth marks Blumenschine RJ (1989) A landscape taphonomic model of the scale and pit to score ratio. Hist Biol 33:176–185 of prehistoric scavenging opportunities. J Hum Evol 18:345–371 Arribas A, Palmqvist P (1999) On the ecological connection between Blumenschine RJ, Cavallo JA (1992) Carroñeo y evolución humana. In: sabre-tooths and hominids: faunal dispersal events in the lower Libros de Investigación y Ciencia. Orígenes del Hombre Mod- Pleistocene and a review of the evidence for the first human erno, pp 90–97 arrival in Europe. J Archaeol Sci 26:571–585 Blumenschine RJ (1995) Percussion marks, tooth marks, and experi- Arribas A, Palmqvist P (2002) The first human dispersal to Europe: mental determination of the timing of hominid and carnivore remarks on the archaeological and palaeoanthropological record access to long bones at FLK Zinjanthropus, Olduvai Gorge, Tan- from Orce (Guadix-Baza basin, southeastern Spain). Hum Evol zania. J Hum Evol 29:21–51 17:55–78 Blumenschine RJ, Salvaggio MM (1988) Percussion marks on bone Arzarello M, De Weyer L, Peretto C (2016) The first European peo- surfaces as a new diagnostic of hominid behaviour. Nature pling and the Italian case: peculiarities and “opportunism.” Qua- 333:763–765 ternay International 393:41–50 Blumenshchine RJ, Marean CW, Capaldo SD (1996) Blind test of inter- Barsky D, Celiberti V, Cauche D, Grégoire S, Lebègue F, Lum- analyst correspondence and accuracy in the identification of cut ley H, Toro Moyano I (2010) Raw material discernment marks, percussion marks, and carnivore tooth marks on bone and technological aspects of the Barranco León and Fuente surfaces. J Archaeol Sci 23:493–507 Nueva 3 stone assemblages (Orce southern Spain). Quatern Boscaini A, Madurell-Malapiera J, Llenas M, Martínez-Navarro B Int 223–224:201–219 (2015) The origin of the critically endangered Iberian lynx: spe- Barsky D, Garcia J, Martínez K, Sala R, Zaidner Y, Carbonell ciation, diet and adaptive changes. Quatern Sci Rev 123:247–253 E, Toro-Moyano I (2013) Flake modification in European Bourguignon L, Crochet JY, Capdevila R, Ivorra J, Antoine PO, Agustí Early-Middle Pleisocne Stone tool assemblages. Quatern Int J, Barsky D, Blain HA, Boulbes N, Bruxelles L, Claude J, 316:140–154 Cochard D, de Weyer L, Filoux A, Firmat C, Lozano-Fernández Barsky D, Sala R, Menéndez L, Toro-Moyano I (2015) Use and re-use: I, Magniez P, Pelletier M, Rios J, Testu A, Valensi P (2015) re-knapped a fl kes from the Mode 1 site of Fuente Nueva 3 (Orce, Bois-de-Riquet (Lézignan-la-Cèbe, l’Hérault): a late Early Pleis- Andalucía, Spain). Quatern Int 361:21–33 tocene archeological occurrence in southern France. Quatern Int Barsky D, Vergès JM, Titton S, Guardiola M, Sala R, Toro-Moyano I 393:24–40 (2018) The emergence and significance of heavy-duty scrapers Brain CK (1969) The probable role of leopards as predators of the in ancient stone toolkits. CR Palevol 17:201–219 Swartkrans Australopithecines. South African Archaeological Bartolini-Lucenti SB, Madurell-Malapiera J (2020) Unraveling the Bulletin 24:170–171 fossil record of foxes: an updated review on the Plio-Pleisto- Bryant HN, Russell AP, Thomason JJ (1995) Carnassial functioning cene Vulpes spp. from Europe. Quaternary Science Reviews in nimravid and felid sabertooths: theoretical basis and robust- 236:106296 ness of inferences. In: Thomason JJ (ed) Functional morphology Behrensmeyer AK (1978) Taphonomic and ecologic information from in vertebrate paleontology. Cambridge University Press, Cam- bone weathering. Paleobiology 4:150–162 bridge, pp 116–135 Binford LR (1981) Bones: ancient men and modern myths. Academic Bunn HT (1982) Meat eating and human evolution: studies on the diet Press, New York and subsistence patterns of plio-pleistocene hominids in East Blain H-A (2005) Contribution de la paléoherpétofaune (Amphibia Africa. Universidad de California, Berkeley, PhD & Squamata) à la connaissance de l’évolution du climat et du Bunn HT, Ezzo JA (1993) Hunting and scavenging by Plio-Pleistocene paysage du Pliocène supérieur au Pléistocène moyen d’Espagne. Hominids: nutritional constraints, archaeological patterns, and Doctorat du Muséum National d’Histoire Naturelle de Paris. behavioural implications. J Archaeol Sci 20:365–398. https:// Département de Préhistoire. Institut de Paléontologie humainedoi. org/ 10. 1006/ jasc. 1993. 1023 Blain H-A (2009) Contribution de la paléoherpétofaune (Amphibia Bunn HT, Pickering TR (2010) Bovid mortality profiles in paleoeco- & Squamata) à la connaissance de l’évolution du climat et du logical context falsify hypotheses of endurance running–hunting paysage du Pliocène supérieur au Pléistocène moyen d’Espagne. and passive scavenging by Early Pleistocene hominins. Quatern Treballs Del Museu Geologic De Barcelona 16:39–170 Res 74:395–404. https:// doi. org/ 10. 1016/j. yqres. 2010. 07. 012 Blain H-A, Bailon S (2010) Anfibios y escamosos del Pleistoceno infe- Carbonell E, Bermúdez de Castro JM, Parés JM, Pérez-González A, rior de Barranco León y de Fuente Nueva 3 (Orce, Andalucía, Cuenca- Bescós G, Ollé A, Mosquera M, Huguet R, Van der España). In: Toro, I., Martínez-Navarro, B., Agustí, J. (Eds.), Made J, Rosas A, Sala R, Vallverdú J, García N, Granger DJ, Ocupaciones humanas en el Pleistoceno inferior y medio en la Martinón-Torres M, Rodríguez XP, Stock GM, Vergés JM, Allué Cuenca de Guadix-Baza. Arqueología Monografías, Sevilla, pp. E, Burjachs F, Cáceres I, Canals A, Benito A, Díez C, Lozano M, 165–183 Mateos A, Navazo M, Rodríguez J, Rosell J, Arsuaga JL (2008) Blain H-A, Bailon S, Agustí J, Martínez-Navarro B, Toro I (2011) The first hominin of Europe. Nature 452:465–469 Paleoenvironmental and paleoclimatic proxies to the Early Cheheb RC, Arzarello M, Arnaud J, Berto C, Cáceres I, Caracausi Pleistocene hominids of Barranco León D and Fuente Nueva 3 S, Colopi F, Daffara S, Canini GM, Huguet R, Karambatsou T, (Granada, Spain) by means of their amphibian and reptile assem- Sala B, Zambaldi M, Berruti GLF (2019) Human behavior and blages. Quatern Int 243:44–53 Homo-mammal interactions at the first European peopling: new Blain H-A, Lozano-Fernández I, Agustí J, Bailon S, Menèndez L, Espí- evidence from the Pirro Nord site (Apricena, Southern Italy). The gares Ortiz MP, Ros-Montoya S, Jiménez Arenas JM, Toro I, Science of Nature 106:16 Martínez-Navarro B, Sala R (2016) Refining upon the climatic Clark J, Linares-Matas G (2021) Seasonality and Oldowan behavioral background of the Early Pleistocene hominid settlement in West- variability in East Africa. J Hum Evol, in press. https:// doi. org/ ern Europe: Barranco León and FN3 (Guadix-Baza basin, SE 10. 1016/j. jhevol. 2021. 103070 Spain). Quatern Sci Rev 144:132–144 Courtenay LA, Yravedra J, Mate-González MÁ, Aramendi J, González- Blumenschine RJ (1986) Early hominid scavenging opportunites. Aguilera D (2019) 3D analysis of cut marks using a new geomet- Implications of carcass availability in the Serengeti and ric morphometric methodological approach. Archaeol Anthropol Ngorongoro ecosystems. Oxford: BAR International Series 283 Sci 11:651–665 1 3 213 Page 20 of 23 Archaeol Anthropol Sci (2021) 13:213 Courtenay LA, Herranz-Rodrigo D, González-Aguilera D et al (2021) Fernández Jalvo Y, Andrews P (2017) Atlas of taphonomic identifica- Developments in data science solutions for carnivore tooth tion. Springer. Ed. pit classification. Sci Rep 11:10209. https:// doi. org/ 10. 1038/ Ferraro JV, Plummer TW, Pobiner BL, Oliver JS, Bishop LC, Braun s41598- 021- 89518-4 DR, Ditchfield PW, Seaman JW III, Binetti KM, Seaman JW de Lombera-Hermida A, Bargallo A, Terradillos-Bernal M, Huguet R, Jr, Hertel F, Potts R (2013) Earliest archaeological evidence of Vallverdú J, García-Anton M-D, Mosquera M, Ollé A, Sala R, persistent hominin carnivory. PLoS ONE 8:e62174. https:// doi. Carbonell E, Rodríguez-Alvarez XP (2015) The lithic industry org/ 10. 1371/ journ al. pone. 00621 74 of Sima del Elefante (Ata- puerca, Burgos, Spain) in the context Gaudzinski S, Turner E, Anzidei AP, Àlvarez-Fernández E, Arroyo- of Early and Middle Pleistocene technology in Europe. J Hum Cabrales J, Cinq-Mars J, Dobosi VT, Hannus A, Johnson E, Evol 82:95–106 Münzel Ss C, Scheer A, Villa P (2005) The use of proboscidean Desantis LRG, Schubert BW, Scott JR, Ungar PS (2012) Implications remains in every-day Palaeolithic life. Quaternary International of diet for the extinction of saber-toothed cats and American 126–128:179–194. https://doi. or g/10. 1016/j. q uaint.2004. 04. 022 lions. PLoS ONE 7:e52453. https:// doi. or g/ 10. 1371/ jour n al. Gibert J, Iglesias A, Maillo A, Gibert L (1992) Industrias líticas en el pone. 00524 53 Pleistoceno inferior de la región de Orce. Projecto Orce-Cueva Despriée J, Moncel MH, Arzarello M, Courcimault G, Voinchet P, Victoria 1988e1992. In: Gibert, J. (Ed.), Museo de Prehisto- Bahain JJ, Falguéres C (2018) The 1-million-year-old quartz ria: Presencia humana en el Pleistoceno inferior de Granada assemblage from Pont-de-Lavaud (Centre, France) in the Euro- y Murcia. Ediciones del Museo de Prehistoria, Ayuntamiento pean context. J Quat Sci 33:639–661 de Orce, 219–282 Domínguez-Rodrigo M (2002) Hunting and scavenging by early Gibert J, Gibert L, Iglesias A, Maestro E (1998) Two ‘Oldowan’ humans: the state of debate, Journal of World Prehistory 16(1), assemblages in the Plio-Pleistocene deposits of the Orce Heidelberg, 1–54 region, southeast Spain. Antiquity 72:17–25 Domínguez-Rodrigo M et al (2002) The ST site complex at Peninj, Gradstein FM, Ogg JG, Smith AG (2005) A geologic time scale. West Lake Natron, Tanzania: implications for early hominid Cambridge University Press, Cambridge behavioural models. J Archaeol Sci 29, Oxford, 639–665 Harstone-Rose A (2008) Evaluating the hominin scavenging niche Domínguez-Rodrigo M, Mabulla A, Bunn HT, Barba R, Diez-Martín F, through analysis of the carcass-processing abilities of the car- Egeland CP, Espílez E, Egeland A, Yravedra J, Sánchez P (2009) nivore guild. PhD of Philosophy in the Department of Bio- Unraveling hominin behavior at another anthropogenic site from logical Anthropology and Anatomy in the Graduate School of Olduvai Gorge (Tanzania): new archaeological and taphonomic Duke University research at BK, Upper Bed II. J Hum Evol 57(3):260–283. Hartstone-Rose A (2011) Reconstructing the diets of extinct South https:// doi. org/ 10. 1016/j. jhevol. 2009. 04. 006 African carnivorans from premolar “intercuspid notch” mor- Domínguez-Rodrigo M, Yravedra Y, Organista E, Gidna A, Fourvel phology. J Zool 285:119–127 J-B, Baquedano E (2015) A new methodological approach to the Haynes G, Klimowicz J (2015) Recent elephant-carcass utilization taphonomic study of paleontological and archaeological faunal as a basis for interpreting mammoth exploitation. Quatern Int assemblages: a preliminary case study from Olduvai Gorge (Tan- 359–360:19–37 zania). J Archaeol Sci 59:35–53 Haynes G, Krasinski K, Wojtal P (2020) A study of fractured pro- Domínguez-Rodrigo M, Cifuentes G, Jiménez B, Abellán N, Pizarro boscidean bones in Recent and Fossil Assemblages. J Archaeol M, Organista E, Baquedano E (2020) Artificial intelligence pro- Method Theory. https:// doi. org/ 10. 1007/ s10816- 020- 09486-3 vides greater accuracy in the classification of modern and ancient Haynes G, Hutson J (2020) African elephant bones modified by bone surface modifications. Sci Rep 10:18862 carnivores: Implications for interpreting fossil proboscidean Duval M, Aubert M, Hellstrom J, Grün R (2011) High resolution LA- assemblages. J Archaeol Sci Rep 34:102596. https:// doi. org/ ICP-MS mapping of U and Th isotopes in an Early Pleistocene 10. 1016/j. jasrep. 2020. 102596 equid tooth from FN3 (Orce, Andalusia, Spain). Quat Geochro- Huguet R, Saladié P, Cáceres I, Díez C, Rosell J, Bennasar M, Blasco nol 6:458–467. https:// doi. org/ 10. 1016/j. quageo. 2011. 04. 002 R, Esteban-Nadal M, Gabucio J, Rodríguez-Hidalgo A, Car- Duval M, Falguères C, Bahain JJ (2012a) Age of the oldest hominin bonell E (2013) Successful subsistence strategies of the first settlements in Spain: contribution of the combined U-series/ humans in south-western Europe. Quatern Int 295:168–182 ESR dating method applied to fossil teeth. Quat Geochronol Huguet R, Vallverdú J, Rodríguez-Álvarez XP, Terradillos-Bernal M, 10:412–417 Bargalló A, Lombera-Hermida A, Menéndez L, Modesto-Mata Duval M, Falguères C, Bahain J-J, Grün R, Shao Q, Aubert M, Dolo M, Van der Made J, Soto M, Blain H-A, García N, Cuenca- JM, Agustí J, Martínez-Navarro B, Palmqvist P, Toro-Moyano I Bescós G, Gómez-Merino G, Pérez-Martínez R, Expósito I, (2012b) On the limits of using combined U-series/ESR method Allué E, Rofes J, Burjachs F, Canals A, Bennàsar M, Nuñez- to date fossil teeth from two Early Pleistocene archaeological Lahuerta C, Bermúdez de Castro JM, Carbonell E (2017) Level sites of the Orce area (Guadix-Baza basin, Spain). Quatern Res TE9c of Sima del Elefante (Sierra de Atapuerca, Spain): a 77:482–491 comprehensive approach. Quatern Int 433:278–295 Espigares MP, Martinez-Navarro B, Palmqvist P, Ros-Montoya S, Toro Kruuk H (1972) The sppoted hyena: a study of predation and social I, Agustí J, Sala R (2013) Homo vs. Pachycrocuta: earliest evi- behavior. University of Chicago Press, Chicago dence of competition for an elephant carcass between scavengers Linares-Matás G, Yravedra J, Maté-González J, Aramendi J, Cour- at FN3 (Orce, Spain). Quatern Int 295:113–125 tenay Ll, Cuartero F, González-Aguilera D (2019) A geomet- Espigares MP, Palmqvist P, Ros-Montoya G-M, S., García-Aguilar, ric-morphometric assessment of three-dimensional models J.M. Rodríguez-Gómez, G., Serrano, F., Martínez-Navarro, B. of experimental cut-marks generated using flint and quartzite (2019) The earliest cut marks of Europe: a discussion on hominin flakes and handaxes. Quatern Int 517:45–54 subsistence patterns in the Orce sites (Baza basin, SE Spain). Lozano-Fernandez I, Blain HA, Agusti J, Piñero P, Barsky D, Ivorra Sci Rep 9:15408 J, Bourguignon L (2019) New clues about the late Early Pleis- Faith JT (2007) Sources of variation in carnivore tooth-mark frequen- tocene peopling of Western Europe: small vertebrates from the cies in a modern spotted hyena (Crocuta crocuta) den assem- Bois-de-Riquet archeopaleontological site (Lézignan-La-Cèbe, blage, Amboseli Park, Kenya. J Archaeol Sci 34:1601–1609 southern France). Quatern Sci Rev 219:187–203 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 21 of 23 213 Lozano-Fernandez I, Blain HA, López-García JM, Agustí J (2015a) Michel V, Shen CC, Woodhead J, Hu HM, Wu CC, Moullé PÉ, … De Biochronology of the first hominid remains in Europe using Lumley H (2017). New dating evidence of the early presence of the vole Mimomys savini: Fuente Nueva 3 and Barranco hominins in Southern Europe. Scientific Reports 7, 10074 León D, Guadix-Baza Basin, south-eastern Spain. Hist Biol Monahan CM (1996) New zooarchaeological data from Bed II, Olduvai 27:1021–1028 Gorge, Tanzania: implications for hominid behavior in the Early Lozano-Fernandez S, Mateos A, Rodríguez J (2015b) Exploring Pleistocene. J Hum Evol 31:93–128 paleo food-webs in the European Early and Middle Pleisto- Oliver JS, Plummer TW, Hertel F, Bishop LC (2019) Bovid mortality cene: a Network Analysis. Quatern Int 413:44–54 patterns from Kanjera South, Homa Peninsula, Kenya and FLK- Luzón C, Courtenay LA, Yravedra J, Saarinen J, Blain H-A, Zinj, Olduvai Gorge, Tanzania: evidence for habitat mediated DeMiguel D, Viranta S, Azanza B, Rodríguez-Alba JJ, Her- variability in Oldowan hominin hunting and scavenging behavior. ranz-Rodrigo D, Solano JA, Oms O, Agustí J, Fortelius M, J Hum Evol 131:61–75. https://doi. or g/10. 1016/j. jhe vol.2019. 03. Jiménez Arenas JM (2021) Taphonomic and spatial analyses 009 from the Early Pleistocene site of Venta Micena 4 (Orce, Gua- Oms O, Dinares-Turell J, Agustí J, Parés JM (1999) Refinements of dix-Baza Basin, southern Spain). Sci Rep 11:13977 the European mammal chronology from the magnetic polarity Madurell-Malapeira J, Martínez-Navarro B, Ros-Montoya S, Espi- record of the Plio-Pleistocene Zújar section, Guadix-Baza Basin, gares MP, Toro I, Palmqvist P (2011) The earliest European SE Spain. Quatern Res 51:94–103 badger (Meles meles), from the Late Villafranchian site of Oms O, Agustí J, Gabàs M, Anadón P (2000a) Lithostratigraphical cor- Fuente Nueva 3 (Orce, Granada, SE Iberian Peninsula). CR relation of micromammal sites and biostratigraphy of the Upper Palevol 10:609–615 Pliocene to Lower Pleistocene in the Northeast Guadix-Baza Maldonado-Garrido E, Piñero P, Agustí J (2017) A catalogue of Basin. J Quat Sci 15:43–50 the vertebrate fossil record from the Guadix-Baza Basin (SE Oms O, Parés JM, Martínez-Navarro B, Agustí J, Toro-Moyano I, Mar- Spain). Spanish Journal of Palaeontology 32:207–236 tínez Fernández G, Turq A (2000b) Early human occupation of Marean CW, Ehrhardt CL (1995) Paleoanthropological and paleoeco- Western Europe: paleomagnetic dates for two Paleolithic sites in logical implications of the taphonomy of a sabertooth’s den. J Spain. Proceedings of the National Academy of Sciences USA Hum Evol 29:515–547 97:10666–10670 Marinellu F, Lemorini C, Barkai R (2021) Lower palaeolithic small Oms O, Agustí J, Parés JM (2010) Litoestratigrafía, magnetoes- flakes and megafauna: the contribution of experimental approach tratigrafía y bioestratigrafía de los yacimientos de Barranco and use-wear analysis to reveal the link. In Konidaris, GE Barkai, León 5 y Fuente Nueva 3 (Cuenca, Guadix-Baza). En B. Mar- R Tourloukis, V Harvati, K (eds) Human-elephant interactions: tínez-Navarro, J. Agustí, I. Toro Moyano (coords.), Ocupaciones from past to present. Tübingen University Press, Tübingen, p humanas en el Pleistoceno Inferior y Medio de la cuenca de 237–260. https:// doi. org/ 10. 15496/ publi kation- 55604 Guadix-Baza. Junta de Andalucía, Consejería de Cultura. Sevilla, Martinez-Navarro B, Palmqvist P (1995) Presence of the African 107–120 machairodont Megantereon whitei (Broom, 1937) (Felidae, Oms O, Anadón P, Agustí J, Julià R (2011) Geology and chronology Carnivora, Mammalia) in the Lower Pleistocene site of Venta of the continental Pleistocene archeological and mammal sites Micena (Orce, Granada, Spain), with some considerations on of the Orce Area (Baza Basin, Spain). Quatern Int 243:33–43 the origin, evolution and dispersal of the genus. J Archaeol Sci Palmqvist P, Martínez-Navarro B, Arribas A (1996) Prey selection 22:569–582 by terrestrial carnivores in a lower Pleistocene paleocommunity. Martinez-Navarro B, Turq A, Agustí J, Oms O (1997) FN3 (Orce, Paleobiology 22:514–534 Granada, Spain) and the first human occupation of Europe. J Palmqvist P, Arribas A, Martínez-Navarro B (1999) Ecomorphological Hum Evol 33:611–620 study of large canids from southeastern Spain. Lethaia 32:75–88 Martínez-Navarro B, Espigares MP, Ros-Montoya S, (2003) Estudio Palmqvist P, Martínez-Navarro B, Toro I, Espigares MP, Ros-Montoya preliminar de las asociaciones de grandes mamíferos de Fuente S, Torregrosa V, Pérez-Claros JA (2005) A re-evaluation of the Nueva 3 y Barranco León(Orce, Granada, España) (Informe de evidence of human presence during Early Pleistocene times in las campañas de 1999e2002). In: Toro-Moyano, I., Agustí, J., southeastern Spain. L’anthropologie 109:411–450 Martínez-Navarro, B. (Eds.), El Pleistoceno inferior de Barranco Palmqvist P, Martínez-Navarro B, Pérez-Claros JA, Torregrosa V, León y Fuente Nueva 3, Orce (Granada). Junta de Andalucía. Figueirido B, Jiménez-Arenas JM, Patrocinio-Espigares M, Ros- Consejería de Cultura. E.P.G.P.C. Arqueología Monografías, Montoya S, De Renzi M (2011) The giant hyena Pachycrocuta Sevilla, 115–137. Memoria Científica campañas 1999–2002 brevirostris: modelling the bone-cracking behavior of an extinct Martínez-Navarro B, Palmqvist P, Madurell-Malapeira J, Ros-Montoya carnivore. Quatern Int 243:61–79 S, Espigares M-P, Torregrosa V, Pérez-Claros JA (2010) La fauna Pares JM, Pérez-González A, Rosas A, Benito A, Bermúdez de Castro de grandes mamíferos de FN3 y Barranco León-5: Estado de JM, Carbonell E, Huguet R (2006) Matuyama-age lithic tools la cuestión. In: Toro, I., Martínez-Navarro B, Agustí J (Eds.), from the Sima del Elefante site, Atapuerca (northern Spain). J Ocupaciones Humanas en el Pleistoceno Inferior y Medio de Hum Evol 50:163–169 la Cuenca de Guadix-Baza. Junta de Andalucía, Consejería de Parkinson JA (2018) Revisiting the hunting-versus-scavenging debate Cultura, Sevilla, 197–236 at FLK Zinj: a GIS spatial analysis of bone surface modifications Maté-González MA, González-Aguilera D, Linares-Matás G, Yrave- produced by hominins and carnivores in the FLK 22 assemblage, dra J (2019) New technologies applied to modelling taphonomic Olduvai Gorge, Tanzania. Palaeogeogr Palaeoclimatol Palaeo- alterations. Quatern Int 517:4–15 ecol 511:29–51. https:// doi. org/ 10. 1016/j. palaeo. 2018. 06. 044 Mech LD (1970) The wolf: the ecology and behavior of an Endargered Pante MC (2013) The larger mammal fossil assemblage from JK2, Species. The Natural History Press, Garden City Bed III, Olduvai Gorge, Tanzania: implications for the feeding Medin T, Martínez-Navarro B, Rivals F, Madurell-Malapiera J, Ros- behavior of Homo erectus. J Hum Evol 64:68–82 Montoya S, Espigares MP, Figueirido B, Rook L, Palmqvist P Pavia M, Zunino M, Coltorti M, Angelone C, Arzarello M, Bagnus C, (2017) Late Villafranchian Ursus etruscus and other large car- Bellucci L, Colombero S, Marcolini F, Peretto C, Petronio C, nivorans from the Orce sites (Guadix-Baza basin, Andalusia, Petrucci M, Pieruccini P, Sardella R, Tema E, Villier B, Pavia G (2012) Stratigraphical and palaeontological data from the Early southern Spain): taxonomy, biochronology, paleobiology, and ecogeographical context. Quatern Int 431:20–41 1 3 213 Page 22 of 23 Archaeol Anthropol Sci (2021) 13:213 Pleistocene Pirro 10 site of Pirro Nord (Puglia, south eastern Stahler DR, Smith DW, Guernsey DS (2006) Foraging and feeding Italy). Quatern Int 267:40–55 ecology of the gray wolf (Canis lupus): lessons from Yellowstone Pickering TR, Egeland CP, Dominguez-Rodrigo M, Brain CK, Schnell National Park, Wyoming, USA. J Nutr 136(7 Suppl.):1923–1926 AG (2008) Testing the “shift in the balance of power” hypoth- Starkovich BM et al (2021) Minimal tools, maximum meat: a pilot esis at Swartkrans, South Africa: hominid cave use and subsist- experiment to butcher an elephant foot and make elephant bone ence behavior in the Early Pleistocene. J Anthropol Archaeol tools using Lower Paleolithic stone tool technology. Ethnoar- 27:30–45 chaeology 12:118–147 Pobiner, B., (2007) Hominid-carnivore interactions: evidence from Tappen M (1995) Savanna ecology and natural bone deposition: impli- modern carnivore bone modification and Early Pleistocene cations for early hominid site formation, hunting and scavenging. archaeofaunas (Koobi Fora, Kenya; Olduvai Gorge, Tanzania). Curr Anthropol 36:223–260 Ph.D. dissertation. Department of Anthropology, Rutgers Uni- Titton S, Barsky D, Bargalló A, Serrano-Ramos A, Vergès JM, versity, New Brunswick. Toro-Moyano I, Sala R, García-Solano J, Jiménez-Arenas JM Pobiner BL, Rogers MJ, Monahan CM, Harris JWK (2008) New (2020) Subspheroids in the lithic assemblage of Barranco León evidence for hominin carcass processing strategies at 1.5 Ma, (Spain): recognizing the late Oldowan in Europe. PLoS One Koobi Fora, Kenya. Journal of Human Evolution 55:103–130 15(3):e0231036 Reinoso-Gordo JF, Barsky D, Serrano-Ramos A, Solano-García Titton S, Barsky D, Bargalló D, Vergès JM, Guardiola M, García- JA, León-Robles CA, Luzón-González C, Titton S, Jiménez- Solano J, Jiménez-Arenas JM, Toro-Moyano I, Sala R (2019) Arenas JM (2020) Walking among mammoths. Remote sensing Active percussion tools from the Oldowan site of Barranco León and virtual reality supporting the study and dissemination of (Orce, Andalousie, Spain): the fundamental role of pounding Pleistocene archaeological sites: the case of Fuente Nueva 3 activities in hominin lifeways. J Archaeol Sci 96:131–147 in Orce, Spain. Sustainability 12:4785 Tixier PJ, y Roche, H. (1995) Polyedre, sub-spheroïde, spheroïde et Rodríguez J, Rodríguez-Gómez G, Martín-González JA, Goikoetxea bola: des segments plus ou moins longs d’une même chaîne I, Mateos A (2012) Predator-prey relationships and the role operatoire. Cahier Noir 7:31–40 of Homo in Early Pleistocene food webs in Southern Europe. Thompson CEL, Ball S, Thompson TJU, Gowland R (2011) The abra- Palaeogeogr Palaeoclimatol Palaeoecol 365–366:99–114 sion of modern and archaeological bones by mobile sediments: Rodríguez-Gómez G, Palmqvist P, Rodríguez J, Mateos A, Martín- the importance of transport modes. J Archaeol Sci 38:784–793 González JA, Espigares MP, Ros-Montoya S, Martínez-Nav- Toro-Moyano I, Lumley H de, Barsky D, Cauche D, Celiberti, V. y arro B (2016) On the ecological context of the earliest human Moncel M.-H. (2003) Las industrias líticas de Barranco León y settlements in Europe: resource availability and competition Fuente Nueva 3. Estudio técnico y tipológico. Las cadenas opera- intensity in the carnivore guild of Barranco León-D and FN3 tivas. Análisis traceológico. Resultados preliminares. Memoria (Orce, Baza Basin, SE Spain). Quatern Sci Rev 134:69–83 Científica campañas 1999–2002. In : Toro, I., Agustí, J., Mar - Rodríguez-Alba J-J, Linares-Matás G, Yravedra J (2019) First assess- tínez-Navarro, B. (Editores), El Pleistoceno inferior de Barranco ments of the taphonomic behaviour of jaguar (Panthera onca). León y FN3, Orce (Granada). Junta de Andalucía. Consejería de Quatern Int 517:88–96 Cultura, Arqueología Monografías, Sevilla, 173–183 Roe DA (1995) The Orce Basin (Andalucia, Spain) and the initial Toro-Moyano I, de Lumley H, Fajardo B, Barsky D, Celiberti V, Gré- Palaeolithic of Europe. Oxf J Archaeol 14:1–12 goire S, Martínez-Navarro B, Espigares MP, y Ros-Montoya, S., Ros-Montoya S (2010) Los Proboscídeos del Plio-Pleistoceno de las (2009) L’industrie lithique des gisements du pleistocène infé- cuencas de Guadix-Baza y Granada. Editorial de la Universi- rieur de Barranco León et Fuente Nueva 3, Granade, Espagne. dad de Granada, Granada PhD L’anthropologie 113:111–124 Ros-Montoya S, Bartolini-Lucenti S, Espigares MP, Palmqvist P, Mar- Toro-Moyano I, Lumley H de, Barrier P, Barsky D, Cauche D, Celiberti tínez-Navarro B (2021) First review of Lyncodontini material V, Grégoire S, Lebègue F, Mestour B. y Moncel MH (2010a) Les (Mustelidae, Carnivora, Mammalia) from the Lower Pleistocene industries lithiques archaïques du Barranco León et de Fuente archaeo-palaeontological sites of Orce (Southeastern Spain). Nueva 3, Orce, basin du Guadix-Baza, Andalousian. En : Mon- Rivista Italiana De Paleontologia e Stratigrafia 127:33–47 ography, CNRS Éditions Saarinen J, Oksanen O, Žliobaitė I, Fortelius M, DeMiguel D, Azanza Toro-Moyano I, Martínez-Navarro B, Agustí J, (2010b) Ocupaciones B, Bocherens H, Luzón C, Solano-García J, Yravedra J, Cour- Humanas en el Pleistoceno inferior y medio de la cuenca de tenay LA, Blain H-A, Sánchez-Bandera C, Serrano-Ramos A, Guadix-Baza. Memoria Científica. Junta de Andalucía, Conse- Rodriguez-Alba JJ, Viranta S, Barsky B, Tallavaara M, Oms O, jería de Cultura, EPG Arqueología Monografias Agustí J, Ochando J, Carrión JS, Jiménez-Arenas JM (2021) Plio- Toro-Moyano I, Barsky D, Cauche D, Celiberti V, Grégoire S, Leb- cene to Middle Pleistocene climate history in the Guadix-Baza egue F, Moncel MH, de Lumley H (2011) The archaic stone-tool Basin, and the environmental conditions of early Homo dispersal industry from Barranco León and Fuente Nueva 3, (Orce, Spain): in Europe. Quaternary Science Reviews 268:107132 evidence of the earliest hominin presence in southern Europe. Schaller GB (1972) The Serengeti lion: a study of predator-prey rela- Quatern Int 243:80–91 tions. University of Chicago Press, Chicago Toro-Moyano I, Martínez-Navarro B, Agustí J, Souday C, Bermúdez Scott G, Gibert L, Gibert J (2007) Magnetostratigraphy of the Orce de Castro JM, Martinón-Torres M, Fajardo B, Duval M, Fal- region (Baza basin), SE Spain: new chronologies for Early Pleis- guères C, Oms O, Parés JM, Anadón P, Julià R, García-Aguilar tocene faunas and hominid occupation sites. Quatern Sci Rev JM, Moigne AM, Espigares MP, Ros-Montoya S, Palmqvist P 26:415–435 (2013) The oldest human fossil in Europe, from Orce (Spain). J Sirakov N, Guadelli J-L, Ivanova S, Sirakova S, Boudadi-Maligne M, Hum Evol 65:1–9 Dimitrova I, Fernandez P, Ferrier C, Guadelli A, Iordanova D, Treves A, Treves HL (1999) Risk and oportunity for humans coexisting Iordanova N, Kovatcheva M, Krumov I, Leblanc JC, Miteva V, with large carnivores. J Hum Evol 336:275–282 Popov R, Spassov R, Taneva S, Tsanova T (2010) An ancient Turq A, Martínez-Navarro B, Palmqvist P, Arribas A, Agustí J, Rod- continuous human presence in the Balkans and the beginnings of ríguez Vidal J (1996) Le Plio-Pléistocène de la région d’Orce, human settlement in western Eurasia: a Lower Pleistocene exam- province de Grenade, Espagne : bilan et perspectives de recherche. Paléo 8:161–204 ple of the Lower Palaeolithic levels in Kozarnika cave (North- western Bulgaria). Quatern Int 223–224:94–106 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 23 of 23 213 Van der Made J, Carlos Calero JA, Mancheño MA (2008) New material Yravedra J, Lagos L, Bárcena F (2011) A taphonomic study of wild of the goat Capra? alba from the Lower Pleistocene of Quibas wolf (Canis lupus) modification of horse bones in Northwestern (Spain); notes on sexual dimorphism, stratigraphic distribution Spain. Journal of Taphonomy 9:37–65 and systematics. Bollettino Della Società Paleontologica Italiana Yravedra J, Maté-González MA, Palomeque-González JF, Aramendi 47:13–23 J, Estaca-Gómez V, San Juan Blazquez M, García Vargas E, Van Valkenburgh B, Ruff CB (1987) Canine tooth strength and killing Organista E, González-Aguilera D, Arriaza MC, Cobo-Sánchez behaviour in large carnivores. J Zool 212:379–397 L, Gidna A, Uribelarrea Del Val D, Baquedano E, Mabulla A, Villa P, Mahieu E (1991) Breakage patterns of human long bones. J Domínguez-Rodrigo M (2017) A new approach to raw mate- Hum Evol 21:27–48 rial use in the exploitation of animal carcasses at BK (Upper Villa P, Bartram L (1996) Flaked bone from a hyaena den. Paléo Bed II, Olduvai Gorge, Tanzania): a micro-photogrammetric and 8:143–159 geometric morphometric analysis of fossil cut marks. Boreas Villa P, Soresi M (1998) Stones tools in carnivore sites. The case of 46:860–873. https:// doi. org/ 10. 1111/ bor. 12224 Bois Roche. J Archaeol Res 56:187–215 Yravedra J, Maté-González MA, Courtenay L, López-Cisneros P, Est- Vislobokova IA, Agadzhanyan AK, Lopatin AV (2020) The case of aca-Gómez V, Aramendi J, de Andrés-Herrero M, Linares-Matás Trlica TRL11–10 (Montenegro): implications for possible early G, González-Aguilera D, Álvarez-Alonso D (2019) Approaching hominin dispersals into the Balkans in the middle of the Early raw material functionality in the Upper Magdalenian of Coímbre Pleistocene. Quatern Int 554:15–35 cave (Asturias, Spain) through geometric morphometrics. Qua- Yravedra J (2006) Tafonomía aplicada a Zooarqueología. Aula Abierta, tern Int 517:97–106 UNED, Madrid Yravedra J, Rubio-Jara S, Courtenay Ll, A., Martos, J.A. (2020) Mam- Yravedra J, Domínguez-Rodrigo M (2009) The shaft-based methodo- mal butchery by Homo erectus at the Lower Pleistocene acheu- logical approach to the quantification of long limb bones and its lean site of Juma’s korongo 2 (JK2), bed III, Olduvai Gorge. relevance to understanding hominin subsistence in the Pleisto- Tanzania Quaternary Science Reviews 249(1):106612. https:// cene: application to four Paleolithic sites. J Quat Sci 24:85–96doi. org/ 10. 1016/j. quasc irev. 2020. 106612 Yravedra J, Domínguez-Rodrigo M, Santonja M, Pérez-González A, Zaidner Y (2013) Adaptive flexibility of Oldowan hominins: secondary Panera J, Rubio-Jara S, Baquedano E (2010) Cut marks on the use of flakes at Bizat Ruhama. Israel. PLoS One 8(6):e66851 Middle Pleistocene elephant carcass of Áridos 2 (Madrid, Spain). J Archaeol Sci 37:2469–2476 Publisher's note Springer Nature remains neutral with regard to Yravedra J, Panera J, Rubio-Jara S, Manzano I, Expósito A, Pérez- jurisdictional claims in published maps and institutional affiliations. González A, Soto E, López-Recio M (2014) Neanderthal and Mammuthus interactions at EDAR Culebro 1 (Madrid, Spain). J Archaeol Sci 42:500–508 1 3 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Archaeological and Anthropological Sciences Springer Journals

Loading next page...
 
/lp/springer-journals/use-of-meat-resources-in-the-early-pleistocene-assemblages-from-fuente-2fQMcquCx2

References (195)

Publisher
Springer Journals
Copyright
Copyright © The Author(s) 2021
ISSN
1866-9557
eISSN
1866-9565
DOI
10.1007/s12520-021-01461-7
Publisher site
See Article on Publisher Site

Abstract

Over the last few decades, several types of evidence such as presence of hominin remains, lithic assemblages, and bones with anthropogenic surface modifications have demonstrated that early human communities inhabited the European sub - continent prior to the Jaramillo Subchron (1.07–0.98 Ma). While most studies have focused primarily on early European lithic technologies and raw material management, relatively little is known about food procurement strategies. While there is some evidence showing access to meat and other animal-based food resources, their mode of acquisition and associated butchery processes are still poorly understood. This paper presents a taphonomic and zooarchaeological analysis of the Fuente Nueva-3 (FN3) (Guadix-Baza, Spain) faunal assemblage, providing a more in-depth understanding of early hominin subsistence strategies in Europe. The present results show that hominins had access to the meat and marrow of a wide range of animal taxa, including elephants, hippopotami, and small- and medium-sized animals. At the same time, evidence of carnivore activity at the site suggests that these communities likely faced some degree of competition from large predators when acquiring and processing carcasses. Keywords Early Pleistocene · Taphonomy · Hominin-carnivore interactions · Zooarchaeology · Cut marks · Palaeolithic archaeology * José Yravedra Department d’Historia i Historia de l’Art, Universitat Rovira joyravedra@hotmail.com I Virgili (URV), Tarragona, Spain Institut Català de Paleoecologia Humana I Evolució Social, Department of Prehistory, Ancient History and Archaeology, Tarragona, Spain Complutense University of Madrid, Madrid, Spain Prehistory Area, Universitat Rovira I Virgili, Tarragona, C.A.I. Archaeometry and Archaeological Analysis, Spain Complutense University, Madrid, Spain Museum of the First Settlers of Europe “Josep Gibert”, Orce, Department of Prehistory and Archaeology, University Granada, Spain of Sevilla, Sevilla, Spain Archaeological and Ethnological Museum of Granada, Department of Cartographic and Land Engineering, Higher Granada, Spain Polytechnic School of Avila, University of Salamanca, Avila, Spain Department of Geology, Unitat d’Estratigrafia, University of Barcelona, Barcelona, Spain Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Department of Prehistory and Archaeology, University of Granada, Granada, Spain School of Archaeology, St. Hugh’s College, University of Oxford, Oxford, UK Institute of Peace and Conflict Research, University of Granada, Granada, Spain History and Arts Doctoral Program, University of Granada, Granada, Spain GEOREC Master Program, University of Granada, Granada, Spain Vol.:(0123456789) 1 3 213 Page 2 of 23 Archaeol Anthropol Sci (2021) 13:213 scavenging, as proposed by Bunn & Ezzo 1993); (3) when Introduction hominins encountered an animal carcass dead from natu- ral causes. The first scenario entails a series of important Recent publications have amply demonstrated that hom- behavioural capacities among early hominins, since hunt- inins inhabited Europe prior to the Jaramillo Subchron ing involves some degree of planning, cooperation, and (1.07–0.98 Ma). Thus, sites such as Barranco León, Fuente the ability to kill prey. Scenarios 2–3 also entail some Nueva 3 (FN3) (Barsky et al., 2010, 2015; Titton et al., degree of landscape knowledge through the monitoring 2019, 2020; Toro-Moyano et  al., 2009, 2010a, 2010b, of suitable procurement spots (Clark and Linares-Matás 2011, 2013), and Sima del Elefante in Spain (Parés et al., 2021), although some authors have highlighted a series 2006; Carbonell et al 2008), Le Vallonnet (Michel et al., of issues associated with them. For example, Treves and 2017), Bois-de-Riquet, and Pont-de-Lavaud in France Treves (1999) and Domínguez-Rodrigo (2002) have noted (Bourguignon et al., 2015; Despriée et al., 2018; Lozano- how scenario 2 is dangerous and unlikely, while the third Fernández et al., 2019), and Kozarnika cave in Bulgaria scenario is highly reliant on trophic pressures and sea- (Sirakov et al., 2010) as well as Pirro Nord in Italy (Arza- sonality (Blumenschine, 1986, 1989; Tappen, 1995; Clark rello et al., 2016; Pavia et al., 2012) have provided exten- and Linarés-Matás, 2021), which renders recurrent and sive evidence of hominin activities older than 1 Ma. The reliable access to carcasses rather unlikely (Domínguez- studies carried out so far in most of these early sites have Rodrigo (2002; Yravedra 2006: 162) beyond opportunis- focused on improving our understanding of the technologi- tic exploitation of catastrophic carrion pulses. From the cal behaviours reflected in the lithic assemblages, docu- 1970s and 1980s, Binford (1981) challenged the “Man the menting the use of Oldowan technocomplexes (Mode 1) Hunter” paradigm of human subsistence by proposing that at most of these sites (Carbonell et al., 2008; Barsky et al., hominins were scavengers, kickstarting an intense “hunt- 2010, 2015; Sirakov et  al., 2010; Toro-Moyano et  al., ing versus scavenging” debate that has generated a vast 2011, 2013; de Lombera-Hermida et al. 2015; Arzarello academic literature involving several hypotheses and the et al. 2016; Bourguignon et al 2015). Nevertheless, com- elaboration of models derived from actualistic reference paratively little is still known about their food procurement frameworks (see Yravedra 2006; Parkinson 2018). After strategies, which included both plant matter and animal more than 30 years of debates, there is an emerging con- resources. In addition to the material traces of their activi- sensus that hominins had primary access to animal car- ties, the presence of these early human communities in casses of different sizes at Early Stone Age sites in Africa the European subcontinent has been directly demonstrated such as FLK-Zinj and BK (Olduvai Gorge, Tanzania), St through the discovery of hominin remains at sites such as (Peninj, Tanzania), FwJj14A, FwJj14B, and GaJi14 (Koobi Barranco León (Toro-Moyano et al. 2013) and Sima del Fora, Kenya), KJS (Kanjera, Kenya), and Swartkrans Elefante (Carbonell et al., 2008). (South Africa) (Monahan, 1996, Domínguez-Rodrigo With regards to the exploitation of animal resources, (2002; Domínguez-Rodrigo et al. 2002, 2009; Pickering some evidence is known of cut-marked bones, providing et al., 2008; Pobiner et al., 2008; Bunn & Pickering, 2010; direct evidence of hominin access to meat resources. In Ferraro et al., 2013; Parkinson, 2018; Oliver et al., 2019; some cases, such as Kozarnika cave (Sirakov et al., 2010), Clark & Linares-Matás 2021). Trlica (Vislobokova et al., 2020), or Pirro Nord (Cheheb In Europe, Sima del Elefante (Atapuerca, Spain) is the et al., 2019), sample sizes are small and present ambiguous only site older than 1 Ma with conclusive evidence of early distribution patterns. This unfortunately limits the extent human access to animal carcasses (Huguet et  al., 2013, to which inferences can be inferred about procurement 2017). At this site, 5% of cervid, bovid, and equid remains modalities and the order of which hominins had access from different anatomical parts possess cut marks, evidenc- to these carcasses (primary or secondary). The timing of ing butchery activities such as skinning, dismembering, and hominin access to carcasses is nonetheless a fundamen- defleshing (Huguet et al., 2017). Moreover, the presence of tal issue for understanding the subsistence strategies of percussion marks in this highly fragmented assemblage evi- Early Pleistocene populations. When conceptualising dences the exploitation of bone marrow. Alongside anthro- carcass acquisition strategies, primary access implies that pogenic evidence, the Sima del Elefante faunal assemblage hominins processed the remains before any other predator, also has some taphonomic evidence of carnivore activity while secondary access implies that hominin consumption (5%), resulting from the action of medium-sized carnivores, of animal resources took place after other predators had i.e. wolves or hyenas, according to Huguet et al. (2017). In already been feeding upon the carcass. contrast with the distribution of cut marks, most tooth marks Primary access is only possible in either of these sce- are found on fat-rich elements, such as vertebrae, ribs, and narios: (1) when hominins hunted prey; (2) when hominins epiphyses/metadiaphyses, a pattern generally associated with dispossessed another carnivore of its prey (confrontational secondary access, according to Huguet et al. (2017). 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 3 of 23 213 At Orce, a model of hominin-carnivore competition has provide a more developed understanding of the subsistence been proposed for both Barranco León and FN3 (Rodríguez- strategies employed by the Early Pleistocene populations Gómez et al., 2016; Espigares et al., 2019). At FN3, the at FN3. discovery of an elephant carcass with nearby lithic imple- ments and giant hyena (Pachycrocuta brevirostris) copro- lites was interpreted as a place of encounter and competition between both agents (Espigares et al., 2013). The study of The FN3 archaeological site the bone assemblages from Barranco León and FN3 in Espi- gares et al. (2019) shows the remains from multiple animals FN3 is an Early Pleistocene archaeological site located modified by humans and carnivores, interpreted as homi- 7 km west of the town of Orce (Granada, southern Spain), nins having secondary access to the carcasses. However, situated in the northeasternmost part of the Guadix-Baza the results of Espigares et al. (2019) have been challenged Basin (Toro-Moyano et al., 2010a, 2010b) (Fig. 1). This by Domínguez-Rodrigo et al. (2020), who reassessed some intermontane basin was formed in the upper-middle Mio- of the evidence. In any case, it is unexpected for Espigares cene after the closure of one of the Betic corridors between et al. (2019) to claim that hominins had secondary access the Mediterranean and the Atlantic (Hüsing et al. 2010). to carcasses when they are documenting evisceration cut From the Upper Miocene to the Middle-Upper Pleistocene, marks on the ventral side of the ribs and vertebrae, as well this area has been interpreted as an endorheic continen- as defleshing cut marks on long bone diaphyses (Espigares tal basin, that is, a closed drainage basin, which eventu- et al., 2019, SF Table S12). Furthermore, carnivore tooth ally formed the saline Baza lake (Anadón et  al., 1994; marks are relatively infrequent in these assemblages. As Anadón and Gabàs, 2009), around which various archaeo- such, there is scope for putting forward alternative hypoth- palaeontological localities such as Barranco León, Fuente eses to those presented by Espigares et al. (2019) and other Nueva, or Venta Micena have been observed (Fig. 1). The authors (e.g. Martínez-Navarro et al. 2003, 2010; Palmqvist waters of this endorheic system were drained by the chang- et al. 2005, 2011; Rodríguez-Gómez et al., 2016). ing course of the Guadalquivir River and its tributaries. Through the present zooarchaeological and taphonomic Today, the basin preserves a succession of alluvial (Gua- study, which analyses the faunal remains excavated during dix) and colluvial deposits (Baza: lacustrine clays, silts, the 2017–2020 fieldwork seasons at FN3, we expand the and sands with evaporitic limestone crusting), reaching FN3 assemblage and assess the hypotheses put forward Espi- up to 100 m thick and covering a timescale ranging from gares et al. (2013, 2019). The new data may enable us to the Upper Miocene to the Upper Pleistocene. Over the Fig. 1 Location of the FN3 archaeological site 1 3 213 Page 4 of 23 Archaeol Anthropol Sci (2021) 13:213 last half-century, numerous fossiliferous and archaeologi- Since the discovery of FN in 1991 by Alain Bocquet, cal localities have been documented (Maldonado-Garrido and until 2016, several research projects have demonstrated et al., 2017). important associations between the faunal remains and the FN3 is located near the margins of the lake (Fig.  1). lithic industries from this locality (Gibert et al., 1992, 1998; The age of FN3 has been determined using both relative Roe, 1995; Tixier et al., 1995; Turq et al., 1996; Martínez- and absolute dating methods, employing both micro- and Navarro et  al., 1997, 2010; Toro-Moyano et  al., 2003). macro-vertebrate biochronological calibrations with mag- Since 2017, a new research project under the direction of neto-stratigraphical data (Agustí et al., 1987, 1996, 2007, the University of Granada has undertaken fieldwork at the 2010; Martinez-Navarro et  al., 1997, 2003, 2010; Oms site. The present study explores the zooarchaeological and et al., 1999, 2000a, 2000b; Agustí and Madurell, 2003; Scott taphonomic insights from the faunal assemblage retrieved et  al., 2007; Lozano-Fernández et al., 2015a), as well as from field campaigns carried out between 2017 and 2020. combined U-series/ESR dating of quartz grains and tooth The taxonomy of the faunal assemblages represented in enamel (Duval et al., 2011, 2012a, 2012b). The stratigraphi- the archaeo-palaeontological sites of Orce has undergone cal sequence at the site (Fig.  2) correlates to the Matuy- numerous changes over the last five decades (see Luzón ama Chron, situated between the Olduvai and Jaramillo et al., 2021, Supplementary Notes S2 for a further discus- subchrons (1.78–1.48 Ma 1.07–0.98 Ma, Gradstein et al., sion). Nevertheless, we present an updated list of the site 2005). In particular, an age of around 1.2 Ma is accepted for of FN3. These include carnivores, such as the bear Ursus the site of FN3, on the basis of the derived features of the etruscus (Medin et al., 2017), the canids Canis mosbachensis rodents Allophaiomys aff. lavocati (Agustí and Madurell, (Luzón et al., 2021, Supplementary Notes S2), Xenocyon 2003; Lozano-Fernández et al., 2015b). (Lycaon) lycaonoides (Luzón et al., 2021, Supplementary Fig. 2 Stratigraphic sequence of FN3 (after Oms et al., 2011; Reinoso-Gordo et al., 2020) 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 5 of 23 213 Notes S2), and Vulpes alopecoides (Bartolini-Lucenti & The lithic industry of FN3 has been studied in detail by Madurell-Malapiera, 2020), the hyena Pachycrocuta brevi- Toro-Moyano et al. (2003, 2010a, 2010b, 2011, 2013) and rostris (Martínez-Navarro et al., 2010), Felidae indet. (Mar- Barsky et al. (2010, 2013, 2015), presenting typical Old- tínez-Navarro et al., 2010), the small felid Lynx cf. pardinus owan (Mode 1) artefacts. The lithic industry recovered in (Boscaini et al., 2015), the badger Meles meles (Madurell- FN3 level 5 represents 5% of the archaeological material Malapeira et al., 2011), and the mustelid Martellictis ardea recovered at the site, compared to the fauna that comprises (Ros-Montoya et al., 2021); and large herbivores, such as 95%. Flint is the predominant raw material, followed by the proboscidean Mammuthus meridionalis (Ros-Montoya, limestone, both of which were obtained from local sources 2010), the rhino Stephanorhinus cf. etruscus (Luzón et al., (Barsky et al., 2010, 2015). The predominant knapping tech- 2021, Supplementary Notes 2), two horses Equus altidens niques include bipolar knapping on anvil, as well as direct and E. sussenbornesis (Alberdi, 2010), the hippo Hippo- percussion knapping, used to produce small-sized flakes and potamus antiquus (Martínez-Navarro et  al., 2010), the choppers (Zaidner, 2013; Barsky et al., 2010, 2015). Lime- bovid Bison sp. (Martínez-Navarro et al., 2010), the caprids stone blocks and some cobbles were mainly used as heavy- Ammotragus europaeus (Martínez-Navarro et  al., 2010) duty tools. Although both flint and limestone were used to and Capra cf. alba (van der Made et al., 2008), and the knap small flakes, the abundance of limestone percussion cervids Praemegaceros cf. verticornis and Metacervocerus tools indicates a clear, task-related differentiation between rhenanus (Abbazzi, 2010). From a paleoecological point the two materials (Toro-Moyano et al., 2010a; Barsky et al., of view, two variables have been calculated both based on 2015). As in other Oldowan assemblages, the toolkit is non- dental ecometrics in large herbivorous mammal guilds of standardized and contains few retouched items. In addition, FN3 and Barranco León: Mean Annual Precipitation (MAP) some heavy-duty scrapers have been identified in the macro- and Net Primary Production (NPP). Results have provided toolkit (Barsky et al., 2018). Stone tool production was car- equal values for MAP (602 mm) and NPP (914 g(C)/m /a) ried out with hard hammers, using both bipolar-on-anvil and in both sites (Saarinen et al., 2021). It is worth noting that free-hand percussion methods. the MAP calculation on dental ecometrics is slightly lower than such calculated by Mutual Ecogeographic Range (FN3 MAP: 738 mm; Blain et al., 2016). In addition, drier climatic Materials and methods conditions (MAP: 449 mm) and lesser productivity (NPP: 747 g(C)/m /a) have been calculated from the dental features The materials analysed for this study were retrieved from of the Venta Micena herbivore community (Saarinen et al., levels 1–6 of FN3 during the 2017–2020 field seasons, with 2021). most remains coming from level 5 (Table 1). This faunal The micromammals from FN3 include the insectivores assemblage encompasses the fossils bones obtained during Erinacenae indet., Crocidura sp., Sorex minutus, Sorex sp., the new field seasons from 2017 to 2020, retrieved from 2 2 Asoriculus gibberodon, and Galemys sp. (Agustí et  al., 44 m in level 5, and 51 m in all other levels. This mate- 2010); and the rodents, Allophaiomys aff. lavocati, Allo- rial expands the faunal assemblages discussed by previous phaiomys sp., Mimomys savini, Castillomys crusafonti, research. Apodemus aff. mystacinus, and Hystrix sp. (Agustí and Faunal remains were quantified by the number of iden - Madurell, 2003; Agustí et al., 2007, 2010). tifiable specimens (NISP), minimun number of elements The herpetofaunal list of FN3 is somewhat poorer (13 (MNE), and minimum number of individuals (MNI). MNI taxa in total) than the nearby site of Barranco León with estimates considered element side and ontogenetic age four anurans (Discoglossus cf. jeanneae, Pelobates cultripes, (Brain, 1969). Mortality profiles were developed by assign- Bufo bufo s.l., and Pelophylax cf. perezi), four lizards (Chal- ing relevant elements/individuals to one of four categories: cides cf. bedriagai, Timon sp., indeterminate small lacertids, infant, juvenile, prime adult-adult, and senile, based on tooth and cf. Ophisaurus), and five snakes (Coronella girondica, eruption and crown wear. Natrix maura, Natrix natrix s.l., Zamenis scalaris, and Skeletal part profiles were based on a division of the skel- Malpolon monspessulanus) (Blain, 2005, 2009; Blain and eton into the following anatomical regions: cranial (i.e. horn, Bailon, 2010; Blain et al., 2011, 2016; Sánchez-Bandera cranium, mandible, and teeth); axial (vertebrae, ribs, pel- et al., 2020). Some fish and chelonian remains are also pre- ves, and scapulae, according to Yravedra and Domínguez- sent in the faunal assemblage. Rodrigo (2009)); upper appendicular elements (humerii, The stratigraphic sequence of FN3 was first described by femora); intermediate appendicular limbs (radii, tibiae, Turq et al. (1996). Nevertheless, it has since been redefined patellae, ulnae); and lower appendicular elements (metapo- by Oms et al. (2010), recognizing 12 stratigraphic levels dials, carpals, tarsals, phalanges, and sesamoids). Long limb (Fig. 2), among which level 5 stands out as it contains most bones were further divided into anterior elements (scapu- of the macro-vertebrate and lithic remains. lae, humerii, radii, ulnae, carpals, and metacarpals), as well 1 3 213 Page 6 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 1 Taxonomic NISP MNI representation in the zooarchaeological assemblages Layer 2 3 5 6 Total 2 3 5 S/A/J/I 6 Total of FN3, based on the faunal Mammuthus meridionalis 1 389 3 393 1 4 0/2/1/1 1 6 remains retrieved from the 2017–2020 field seasons, Stephanorhinus etruscus 1 18 1 20 1 1 0/1/0/0 1 3 quantified in terms of NISP Equus altidens 5 5 1 0/1/0/0 1 and MNI. Age classes: S: Equus sp. 2 1 27 30 1 1 2 0/2/0/0 4 seniles; A: adults; J: juveniles; Hippopotamus antiquus 53 53 3 0/1/1/1 3 I: infants. The Bold entries reflected the total Bison sp. 8 8 2 0/2/0/0 2 Bovidae 3 1 18 22 1 1 2 0/2/0/0 4 Praeovibos sp. 1 1 1 0/1/0/0 1 Capra alba 6 6 1 0/1/0/0 1 Sorgelia minor 1 1 1 0/1/0/0 1 Cervidae 1 36 37 1 3 0/2/1/0 4 Metacervocerus rhenanus 1 2 3 1 1 0/1/0/0 2 Praemegaceros cf. verticornis 1 5 6 1 1 0/1/0/0 2 Artiodactyla size 2 1 3 4 Artiodactyla size 3 1 1 Artiodactyla size 3b 6 6 Oryctolagus lacosti 6 6 1 0/1/0/0 1 Ursus etruscus 3 3 1 0/1/0/0 1 Carnivore indet 3 3 1 1 Carnivore size 2 1 1 1 0/1/0/0 1 Chelonia 35 35 1 0/1/0/0 Ave 1 1 1 Mammalia indet size 0 4 4 Mammalia indet size 1 6 6 Mammalia indet size 2 6 59 65 Mammalia indet size 3 1 6 47 1 55 Mammalia indet size 3a 1 1 14 16 Mammalia indet size 3b 2 3 118 123 Mammalia indet size 4 1 9 10 Mammalia indet size 5 5 2 99 106 Indet 84 81 3567 39 3771 Total 100 107 4550 44 4801 as posterior elements (pelves, femora, tibiae, patellae, tar- carnivores (e.g. wolves); and large carnivores (e.g. lions, sals, and metatarsals). Herbivore and carnivore specimens hyenas), following Espigares et al. (2019). that could not be identified at a species level, but were not Several procedures were followed to reconstruct site entirely considered indeterminable, were assigned a weight/ formation processes, assessing and evaluating both site size class. The categories used for herbivores follow the integrity and the contribution of various biological agents adaptation made by Espigares et al. (2019) of the classifi- to the faunal assemblage. Bone fragmentation was ana- cation system developed by Bunn (1982): microfaunal (0), lysed according to three variables; first, bones were divided including species weighing less than 25 kg; very small size into several categories according to their length: < 3 cm, (1), including macro-vertebrates species weighing 25–50 kg; 3.1–5.0 cm, 5.1–10 cm, and > 10 cm; secondly, bones were small size (2), including species weighing 50–125 kg; inter- classified based on the nature of their breakage planes, i.e. mediate size (3), including species weighing 125–500 kg, green or dry fractures, following Villa and Mahieu (1991). with an additional division between 3a (125–250 kg) and Criteria used to diagnose dry breaks included the existence 3b (250–500 kg); large size (4), including species weigh- of abundant breaks that are longitudinal and/or transverse ing 500–1000 kg; and very large size (5) for species weigh- to the axis of the bone, as well as breakage planes that are ing > 1000 kg. Carnivores were classified according to three uneven, rough, and in possession micro-step fractures. size classes: small carnivores (e.g. foxes); intermediate Dry breaks are further characterized by cortical medullary 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 7 of 23 213 surface angles that are close to 90°. In contrast, specimens rhinoceroses, are well represented, alongside animals better broken when fresh frequently have smoother surfaces and adapted to wooded environments (such as cervids), general- more abundant oblique breakage planes. Lastly, bone dia- ist animals adapted to rocky environments (such as caprids), physis preservation was recorded according to the relative and animals that tend to rely on watercourses (such as hippo- length of their fragments, as well as circumference types; potami). In relation to the latter, other non-herbivore species where type 1 refers to specimens with < 25% of the shaft cir- such as turtles are also present. Mortality patterns show a cumference intact; type 2 refers to specimens with 25–50% predominance of adults in all levels across all taxa, with only of the shaft circumference intact; and type 3 refers to speci- level 5 yielding infant or juvenile individuals, found among mens with > 50% shaft circumference. proboscideans, hippopotami, and cervids (Table 1). The impact of u fl vial alterations was estimated with frag - The skeletal profiles analysed for levels 2, 3, and 6 are ment size distributions, as well as the presence of abrasion, not very representative due to their small sample size (see polishing, and rounding on bone surfaces. Rounding and Suppl. File). The level 5 sample is larger, but with the excep- abrasion were additionally classified into different stages; tion of size 3 animals, most size classes are generally poorly light, intermediate, and intense. Weathering intensity was represented, and tend to show a predominance of cranial ele- analysed following Behrensmeyer (1978). Bone surface ments due to the overrepresentation of heavily fragmented modification analyses were carried out using hand-held teeth and ivory tusks. Small animals (sizes 0–2) also show lenses at 10–40 × magnification. Tooth marks were classified some presence of axial and appendicular elements. Among as pits, scores, or punctures, while furrowing was also ana- medium-sized animals (size 3), all the skeletal portions are lysed following the criteria established by multiple authors represented including the axial and appendicular elements, (Binford, 1981; Blumenschine, 1995; Blumenschine et al., even if cranial remains are clearly overrepresented (see 1996). Cut and percussion marks were classified according Tables 2–3 and Suppl. File). Among the appendicular ele- to Binford (1981), Blumenschine and Salvaggio (1988), and ments, we observe that both forelimbs and hindlimbs have Blumenschine et al. (1996). Modifications were quantified a fairly similar representation. Very few remains could be for specimens with well-preserved bone surfaces in terms of attributed to large animals (size 4), while very large animals NISP values. Other processes, such as manganese staining (size 5) are also represented mainly by cranial elements (see or biochemical alterations, were recorded using the criteria Tables 2–3 and Suppl. File). outlined by Fernández Jalvo and Andrews (2017). Taphonomic analyses Results The faunal assemblage shows a very high degree of frag- mentation across all levels, with a predominance of Zooarchaeological analyses bones < 3  cm, rendering more than 86% of the analysed sample indeterminable (Table 4). Among long limb bones, The sample analysed comprises 4801 remains, with level the high level of fragmentation has considerably influenced 5 being the one that has provided the largest volume of their circumference index and relative length values. In this remains (n = 4550), of which 14% were considered determi- regard, more than 90% of long bones from level 5 have less nable remains (Table 1). The other levels have only provided than 25% of their shaft circumference and less than 25% of a relatively small sample; in levels 2 and 3, hardly more than their relative length (Table 4). In levels 3 and 6, the sam- 100 remains per level have been found and only 6% of them ple is very small but also shows high fragmentation rates. could be determined. In level 6, the sample is even more lim- This high degree of fragmentation is likely due to diverse ited, with only 44 remains documented. Remains identified biostratinomic processes that have induced green fractures at the site belong to several herbivore and carnivore taxa. in > 40% of the level 5 bone assemblage, and in > 50% of the Among the herbivores, Mammuthus meridionalis stands bone samples from levels 2, 3, and 6 (Table 4). out with 62% of the determinable remains, and 14% of the Although bone fragmentation is high, cortical surface MNI. The second best-represented taxa is Hippopotamus preservation in the assemblage is very good (> 75% of the antiquus with 8.5% of the remains and 10.7% of the indi- bones are in a good state of preservation), and only level 2 viduals. Artiodactyla (cervids, caprids, and bovids) are bones are poorly preserved (Table 5). This good state of cor- better represented than equids. Finally, there are several tical preservation is likely due to the rapid burial rates that carnivore remains, of which only an ursid (Ursus etruscus) the assemblage experienced, as indicated by the very low could be identified at a species level. From a palaeoeco- incidence of weathering alterations: over 99% of the sam- logical perspective, the species represented belong to a wide ple exhibit slight or no weathering (Table 3). Furthermore, range of different ecological settings. Species associated waterborne alterations (abrasion, polishing, or rounding) with open environments, such as proboscidians, equids, and are not particularly common, with only 13% of bones from 1 3 213 Page 8 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 2 Skeletal profiles Species size level 5 according to NISP of the level 5 of FN3 assemblage from the NISP 0 1 2 3 3a 3b 4 5 Carniv 1 Carniv 3 Indet Total 2017–2020 field seasons. See Ivory 228 228 complete information in SF. The Bold entries reflected the total Cranial 1 1 2 1 3 8 Mandible 1 3 1 5 Tooth 2 31 18 1 55 202 3 120 432 Vertebrae 4 3 8 4 19 Rib 4 14 16 3 33 4 32 1 122 229 Scapule 1 1 1 1 4 Humerus 3 1 3 7 14 Radius 1 2 3 Ulna 1 1 Carpal 1 1 8 4 14 Metacarpal 3 3 Pelvis 3 1 4 Femur 1 2 1 1 6 3 14 Tibia 1 1 6 1 10 19 Metatarsal 1 1 4 1 7 Metapodial 2 1 6 1 7 1 18 Tarsal 1 1 2 Phalange 1 5 2 8 Indet 5 1 21 11 4 42 4 69 3302 3459 Placa 35 35 Total 11 6 85 65 16 196 9 548 2 3 3585 4526 Table 3 Skeletal profiles according to MNE of the level 5 of FN3 assemblage from the 2017–2020 field seasons. See complete information in SF. The Bold entries reflected the total Species size level 5 MNE 0 1 2 3 3a 3b Total size 3 % with tooth % no tooth 4 5 Carniv 1 Carniv 3 Ivory 5 Cranial 1 1 1 2 1.6 2.7 1 1 Mandible 1 2 2 1.6 2.7 1 Tooth 2 20 12 1 39 52 40.9 23 3 Vertebrae 2 1 3 4 3.1 5.3 Rib 1 3 3 1 10 14 11.0 18.7 2 8 1 Scapule 1 1 2 1.6 2.7 1 Humerus 1 1 1 2 4 3.1 5.3 Radius 1 1 1 0.8 1.3 Ulna 0.0 0.0 1 Carpal 1 1 8 10 7.9 13.3 4 Metacarpal 3 3 2.4 4.0 Pelvis 1 0.0 0.0 1 Femur 1 1 1 1 4 6 4.7 8.0 1 Tibia 1 1 5 1 7 13 10.2 17.3 Metatarsal 1 1 4 5 3.9 6.7 1 Metapodial 1 1 2 1 2 5 3.9 6.7 Tarsal 1 0.0 0.0 1 Phalange 1 4 4 3.1 5.3 2 Total 5 2 35 28 8 91 127 100.0 169.3 3 49 2 3 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 9 of 23 213 Table 4 Fragmentation patterns Taphonomic features Lev-2 Lev-3 Lev-5 Lev-6 of the FN3 bone assemblage (levels 2, 3, 5, and 6). The Bold Fragmentation Sample NISP 100 107 4550 44 entires reflected the total of Bones < 3 cm 82 87 3843 40 NISP of sample Bones 3.1–5 cm 12 10 416 3 Bones > 5.1–9.9 cm 6 10 218 1 Bones > 10 cm 73 % long bone with % long bone with green fracture 60 50 41 100 % Long bone with dry fracture 40 50 18.4 % long bone with indet fracture 40.6 % degree of long bone shaft circumference < 25% 75 91.1 100 26–50% 12.5 5.48 0 > 51% 12.5 3.42 0 % shaft length of long bones < 25% 87.5 95.9 100 26–50% 12.5 2.72 0 > 51% 1.4 levels 3 and 5 showing evidence of these alterations, always large species, and ribs, vertebrae, and shafts of intermediate of low or moderate intensity (Table 5). These alterations size species. Furthermore, the presence of percussion marks indicate that the assemblage did not experience considerable on long bones indicates that these bones were intention- water transport and that these different waterborne altera- ally fractured in order to access energy-dense within-bone tions were produced by small sedimentary particles sliding resources, such as marrow (Table 5). over the bone surface during processes of site formation. Very few specimens show calcitic concretions, and surface staining by manganese or oxides, which affects around 15% of the assemblage although it has not impacted bone pres- Discussion and conclusions ervation (Table 5). Biochemical alterations, such as root- etching or rodent damage, were relatively unimportant in The evidence reported in this study resembles the previous the FN3 assemblage. analyses of FN3 presented by Espigares et al. (2019). Both Carnivore activity has been recorded for levels 2, 3, and studies show similarly diverse taxonomic profiles with spe- 5, and if we exclude bone fragments < 2 cm, tooth-marked cies from different ecological settings, with a predominance bones can reach up to 11% in the smaller samples (levels in the MNI of very large animals, such as elephants and hip- 2–3). Nevertheless, overall tooth mark frequencies are lower popotami. The mortality patterns are also consistent, high- than 3%, with very few digested bones or instances of fur- lighting that the mortality profiles of sizes 1, 2, and 3 show rowing are present. Similarly, only one bone exhibited 5 or a predominance of adult individuals, whereas larger animals more tooth marks, suggesting that the incidence of carnivore present a higher percentage of infants and juveniles. Skeletal activity at the site is low. Tooth marks are usually 2–3 mm in part profiles also show a similar scarcity of axial elements diameter (with only one instance reaching 5 mm), and these and an overrepresentation of cranial elements, with teeth traces of carnivore activity are mostly found on the edges playing a major role in this regard. The presence of bones of ribs and long bones of small-, medium-, and large-sized from all skeletal portions suggests that some carcasses may animals (see Suppl. File). have arrived complete at the site. Hominins also contributed to the accumulation and On the basis of the presence and distribution of cut marks, modification of the faunal assemblage, as evidenced by the percussion marks, and tooth marks, both studies also agree presence of cut and percussion marks on bone elements. that humans and carnivores had access to the meat and Cut mark frequencies are lower than 1% (Table 5), although within-bone nutrients of several animal species at FN3. The they are found across all carcass sizes, including very large identification of cut marks on small-, medium-, and very animals, such as hippopotami or elephantids (Table 6). The large–sized animals such as elephantids or hippopotami distribution patterns of cut marks show that these anthropo- also indicate that hominins exploited multiple animal taxa. genic alterations appear on ribs, diaphysis of femur of very Carnivores have also left feeding traces on a wide range of 1 3 213 Page 10 of 23 Archaeol Anthropol Sci (2021) 13:213 Table 5 Taphonomical alterations for FN 3 Taphonomic characteristics Lev-2 Lev-3 Lev-5 Lev-6 % lev-2 % lev-3 % lev-5 % lev-6 Sample NISP 100 107 455 44 Sample excluding teeth and Ivory 94 97 3821 37 Bone surfaces Samples with bad preservation 68 16 1101 7 68.0 15.0 24.2 15.9 Samples with good preservation 26 81 272 30 32.0 85.0 75.8 84.1 Weathering stage 0 97 105 4378 44 97.0 98.1 96.2 100 Weathering Weathering stage 1–2 2 2 162 0 2.0 1.9 3.6 0.0 Weathering stage 3–4 1 10 1.0 0.0 0.2 0.0 Hydrolic alteration 4 14 605 4 4.0 13.1 13.3 9.1 Water alteration Abrasion 4 10 293 2 4.0 9.3 6.4 4.5 Light stage abrasion 2 5 222 2 50.0 50.0 75.8 100 Intermediate stage abrasion 2 3 54 50.0 60.0 18.4 Intense stage abrasion 16 0.0 0.0 5.5 Polishing 3 5 154 3.0 4.7 3.4 0 Rounding 2 5 379 3 2.0 4.7 8.3 6.8 Light stage rounding 1 3 270 1 50.0 60.0 58.2 33.3 Intermediate stage rounding 1 1 116 1 50.0 20.0 25.0 33.3 Intense stage rounding 1 78 1 0.0 20.0 16.8 33.3 Calcitic concretions 1 14 1.0 0.0 0.3 0.0 Oxides (MN, FE) 8 18 681 13 8.0 16.8 15.0 29.5 Trampling alterations 2 2 99 1 7.7 2.5 3.6 3.3 Biochemical and root alterations 6 12 183 1 6.0 11.2 4.0 2.3 Bones with rodent tooth marks 1 0.9 Carnivore activity Bones with carnivore tooth marks 1 2 21 0 3.8 2.5 0.8 0 Bones with carnivore tooth marks 1 2 21 0 11.1 11.1 2.6 0 (excluded bone fragments < 2 cm) Bones with tooth pits 1 1 15 100 100 71.4 Bones with tooth scores 10 47.6 Bones with punctures 1 4.8 Bones with both pits and scores 1 3 50 14.3 Digestive alterations 1 Human activity Bones with cut marks 1 2 22 3.8 2.5 0.8 Bones with cut marks (excluded 1 2 22 11.0 11.0 2.7 bone fragments < 2 cm) Bones with peeling 1 Bones with percussion marks 1 14 1 0.0 1.2 0.5 3.3 carcass sizes, from smaller animals such as rabbits to very documented at FN3 with the results of several actualistic large animals, including elephants. frameworks. The butchery activities inferred from the distribution pat- These actualistic reference frameworks were developed terns of cut marks (Table 6) appear to be related to eviscera- by different authors (see SF 2) with the aim of establish- tion and defleshing activities, which, alongside the evidence ing empirically the order in which hominins and carnivores for percussion marks, suggests that hominins were accessing accessed animal carcasses. Carcasses of different sizes were the meat and marrow from different carcass sizes. However, made available to different human and carnivore agents in the presence of tooth marks also indicates that carnivores different experimental settings, in order to quantify the fre- played a role in the alteration of the faunal assemblage at quencies of taphonomic alterations that they generated. It the site. These taphonomic data are all consistent with the was noted that the frequency and distribution of cut marks, results described by Espigares et  al. (2019). In order to percussion marks, and tooth marks varied depending on ascertain the temporality of carcass access, we have sub- which agent(s) accessed the carcass first, which allowed sequently compared the cut and tooth mark frequencies researchers to empirically establish the relative temporality 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 11 of 23 213 Table 6 Cut-marked bones Taxon Species size Skeletal bone Function identified among the FN3 assemblage from the 2017–2020 Bovidae 3b Rib Disarticulation field seasons Mammalia indet 1 Rib Defleshing Mammalia indet 5 Rib Defleshing Mammalia indet 5 Rib Defleshing Hippopotamus antiquus 5 Femur diaphysis Defleshing Mammalia indet 3b Femur diaphysis Defleshing Mammalia indet 3 Long bone diaphysis Defleshing Mammalia indet 5 Long bone Indet Mammalia indet 3b Long bone Indet Mammalia indet 3b Long bone diaphysis Defleshing Mammalia indet 3b Long bone diaphysis Defleshing Mammalia indet Indet Indet Defleshing Mammalia indet 2 Indet Defleshing Mammalia indet 3a Indet Defleshing Mammalia indet Indet Indet Indet Indet Indet Indet Indet Mammalia indet 2 Pelvis Defleshing Mammalia indet 5 Pelvis Evisceration Mammalia indet 3b Tibia diaphysis Indet Mammalia indet 3 Tibia diaphysis Defleshing Mammalia indet 3b Tibia diaphysis Defleshing Mammalia indet 3b Vertebrae Defleshing Mammalia indet 3b Vertebrae Defleshing of animal resource acquisition. This approach has gener- happens when analysing the frequency of tooth marks on ally yielded positive results in taphonomic studies of Early limb bone diaphyses: the results obtained are well below the Pleistocene sites in Africa (Domínguez-Rodrigo, 2002; outcomes of actualistic observations of carnivore-first mod- Domínguez-Rodrigo et  al. 2002, 2009; Pickering et  al., els. Instead, the low tooth mark frequencies fit much better 2008; Pobiner, 2007; Pobiner et al., 2008; Pante, 2013; Par- the patterns left by carnivores when they engage with car- kinson, 2018; Yravedra et al., 2020). casses after human intervention, including those instances Nevertheless, it is worth noting that the comparative when carcasses were initially modified by vultures (Fig.  5). assessment of bone surface modifications for interpreting While tooth mark data seems to indicate that carnivores the order of carcass intervention at FN3 using experimental were not the primary agent responsible for the accumula- or actualistic frameworks is hindered by small sample sizes tion of the faunal assemblage at FN3, it is still necessary to (Figs. 3–7). conclusively prove the degree of human intervention through The very low frequencies of carnivore tooth marks docu- the analysis of the cut mark and percussion mark frequen- mented at FN3 are not consistent with the percentage ranges cies at the site. generated by carnivore-hominin models (Fig. 3), with the When the frequencies of percussion marks left by humans exception of cheetah or leopard accumulations of size 1–2 are compared with the actualistic framework, the very low animals. However, the FN3 assemblage has a considerable percentages found at FN3 do not readily fit any of the pro- representation of animals > 150 kg, a size range that neither posed models (Fig. 6), leading to a somewhat inconclusive cheetahs nor leopards target, which suggests that carnivores result. were not the main accumulation agents at FN3 (Fig. 3). Neither do the very low cut mark frequencies seem to Across all three categories of appendicular elements, correspond to primary anthropogenic accumulations either tooth mark frequencies are considerably lower than those (Fig. 7), resembling more closely the profiles that result from that would be expected in episodes of primary access by secondary hominin access to carcasses previously consumed carnivores (Fig. 4). ULB and ILB in particular have very by carnivores. Breaking down cut mark frequencies by low tooth mark frequencies, a pattern inconsistent with the upper, intermediate, and lower appendicular elements, ULB observations of carnivore feeding when they have access frequencies would match those associated with secondary to the most nutritious portions of the carcasses. The same access, while cut mark frequencies on diaphyses/bone shafts, 1 3 213 Page 12 of 23 Archaeol Anthropol Sci (2021) 13:213 Fig. 3 Tooth mark frequencies on appendicular elements from FN3 yellow box shows the tooth mark frequencies resulting from a model by size classes (1–2, 3, 4–5) in relation to the actualistic framework in which vultures accessed first the carcasses, followed by humans derived from carnivore feeding behaviours documented in different and then carnivores, while the red box documents the main range of contexts. The blue box highlights the range of tooth mark frequen- tooth mark frequencies associated with primary carnivore accumula- cies that carnivores tend to leave on bones during secondary scav- tions. See Suppl. file 2 for the full bibliographic details of the relevant enging activities after human abandonment of the carcasses. The sources Fig. 4 Tooth mark frequencies on appendicular remains from FN3 such as humerii and femoral remains, ILB refers to intermediate on animal size classes 1–2, 3, and 4–5, in relation to the actualistic limbs (tibiae, radii), while LLB stands for lower limb bones (i.e. framework derived from carnivore feeding behaviours documented in metapodials). See suppl. file 2 for more information on the compara- different contexts. ULB refers to tooth marks on upper limb bones, tive samples and their full bibliographic details intermediate, and lower elements are inconclusive since they available does not allow us to determine with certainty the overlap with models of both primary and secondary accesses temporality of hominin access to animal carcasses at FN3, (Fig. 8). Consequently, the taphonomic evidence currently 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 13 of 23 213 Fig. 5 Tooth mark frequencies on limb bone diaphyses from FN3 yellow box shows the tooth mark frequencies resulting from a model on animal size classes 1–2, 3, and 4–5, in relation to the actualistic in which vultures accessed first the carcasses, followed by humans framework derived from carnivore feeding behaviours documented in and then carnivores, while the red box documents the main range of different contexts. The blue box highlights the range of tooth mark tooth mark frequencies associated with primary carnivore accumula- frequencies that carnivores tend to leave on bones during secondary tions. See the Suppl. file 2 for the full bibliographic details of relevant scavenging activities after human abandonment of the carcasses. The sources Fig. 6 Percussion mark frequen- cies on appendicular elements from FN3 on the basis of carcass size (1–2, 3, and 4–5) in relation to a comparative actualistic framework. The blue box represents the percussion mark frequencies generated by humans when they have early access to carcasses, and the red box encompasses the percussion mark frequencies in contexts of secondary access. See Suppl. File 2 for bibliographic details of the reference samples employed a problem of ambiguity further compounded by the small while the low frequency of tooth marks does not match the sample sizes of bone surface modifications. expectations of primary carnivore access (Figs. 3–5). With respect to the temporality of carcass access, the When analysing the distribution and inferred functional- comparative assessment of cut mark, percussion mark, and ity of cut marks (Table 6, Fig. 8), we note their presence tooth mark frequencies in the FN level 5 bone assemblage on the diaphyses of long bones such as the femur or tibia with actualistic framework returned ambiguous and incon- (Figs. 9, 10, 11), as well as on axial elements such as ribs or clusive results. The low frequency of cut marks (Fig. 7 and the pelvis (Fig. 11). This pattern would be consistent with 8) resemble more a pattern of secondary hominin access, defleshing and evisceration activities on carcass elements 1 3 213 Page 14 of 23 Archaeol Anthropol Sci (2021) 13:213 Fig. 7 Cut mark frequencies on appendicular elements from FN3 (level 5) on the basis of carcass size (1–2, 3, and 4–5) in relation to the comparative framework generated by several experimental and actualistic studies. The blue box represents the cut mark frequencies gener- ated by humans when they have early access to carcasses, the yellow box corresponds to cut mark frequencies when human access followed the presence of vultures but preceded carni- vore access, while the red box encompasses cut mark frequen- cies from contexts of secondary access. See suppl. File 2 for the full bibliographic details of the comparative samples employed Fig. 8 Cut mark frequencies on appendicular remains from FN3 frequencies left by humans in early carcass access contexts, while the (level 5) on animal size classes 1–2, 3, and 4–5. ULB refers to tooth red box shows cut mark frequencies in contexts of secondary access. marks on upper limb bones, such as humerii and femoral remains, Cut mark frequencies on diaphyses are shown on the right hand side ILB refers to intermediate limbs (tibiae, radii), while LLB stands for of the graph. See suppl. file 2 for more information on the compara- lower limb bones (i.e. metapodials). The blue box represents cut mark tive samples and their full bibliographic details with high nutritional values, which may indicate early access At the same time, the low frequencies of anthropogenic to at least some of the carcasses, rather than secondary scav- traces do not allow to conclusively establish the nature of enging. This is especially relevant when considering actu- carcass acquisition strategies. The sample analysed in the alistic reports of carnivore feeding patterns to first consume present study thus experiences similar limitations than the the visceral packages. This pattern is documented among work conducted by Espigares et al. (2019): both have few a wide range of wild carnivores, such as felines (Schaller, bones with anthropogenic and carnivore alterations, and thus 1972; Blumenschine, 1986; Blumenschine & Cavallo, neither can determine in which order did humans and carni- 1992), canids (Mech, 1970; Stahler et al, 2006; Yravedra vores access prey carcasses at the site. As such, there is little et al., 2011), and hyenas (Kruuk, 1972; Blumenschine, 1995; evidence to support previous models of hominin acquisi- Faith, 2007). tion of animal resources at Orce consisting in the secondary 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 15 of 23 213 Fig. 9 Shaft of femur of hippopotamus with cut marks in FN3 level 5 Fig. 10 Shaft with cut marks and 3D modelled of FN3 level 5 1 3 213 Page 16 of 23 Archaeol Anthropol Sci (2021) 13:213 2016, according to Van Valkenburgh and Ruff, 1987). This circumstance would provide hominins with opportunities to scavenge sizeable chunks of muscle tissue before giant hyena individuals accessed the carcasses. This passive scav- enging scenario might help explain the cut mark distribution patterns and the scarcity of tooth marks that is observed by Espigares et al. (2019) documented at FN3. However, there are multiple lines of evidence that prompt a reconsideration of the interpretations proposed by Mar- tínez-Navarro and Palmqvist (1995), Palmqvist et al. (1996, 1999, 2005, 2011), Arribas and Palmqvist (1999, 2002), Espigares et al. (2013, 2019), and Rodríguez-Gómez et al. (2016) regarding the Megantereon-Hominins-Pachycrocuta model of carcass acquisition at FN3. Several authors have already shown in different publica- tions that skeletal parts and mortality profiles are generally inconclusive when modelling the behaviour of humans and carnivores, since the resulting outcomes can experience issues of equifinality (see discussion in Domínguez-Rodrigo (2002)). From a taphonomic perspective, it is worth bearing in mind that there are no empirical studies of the feeding behaviour of M. whitei, which precludes reliable interpreta- tions and inferences regarding their role in the alteration of fossil assemblages. The only available taphonomic study of an accumulation altered by machairodonts is described by Marean and Ehrhardt (1995). While the assemblage corre- sponds to Homotherium rather than M. whitei—which may have generated slightly different patterns—the high fre- quencies of bone damage, particularly in relation to those documented at FN3, are noteworthy, indicating that some sabretooth felids were capable of inflicting considerable damage to the bones of their prey. Moreover, Harstone-Rose (2008, 2011) has argued that Megantereon would have had carcass processing behaviours no more hypercarnivorous Fig. 11 Several cut marks of FN3 level 5 on animal size level 3, pel- than extant felids, which have been shown to generate con- vis of animal size level 5, and indeterminate fragment of level 5 siderable tooth mark damage (see e.g. Rodríguez-Alba et al. 2019 for jaguars; Brain 1969 and Arriaza et al. 2019 for leopards; or Arriaza et al. 2016, Haynes & Klimowicz 2015, exploitation of carcasses left by large felids, and prior to the and Haynes & Hutson 2020 for lions). Furthermore, bio- intervention of the giant hyena. mechanical studies of sabretooth dentition suggest that these Several authors, including Martínez-Navarro and carnivores would have had no issue making contact with Palmqvist (1995), Palmqvist et al. (1996, 1999, 2005, 2011), bone surfaces when they were feeding on animal carcasses Arribas and Palmqvist (1999, 2002), Espigares et al. (2013, (Bryant et al., 1995; Harstone-Rose, 2008; 2011; Desantis 2019), and Rodríguez-Gómez et  al. (2016), suggest, on et al. 2012). Therefore, the teeth of Megantereon whitei were the basis of skeletal part profiles, mortality patterns, and neither as fragile nor as inefficient as Van Valkenburgh and taphonomic evidence, that large felids, such as Megantereon Ruff (1987) and Palmqvist et al. (1996, 1999, 2005, 2011) whitei, would have had primary access to animal carcasses have proposed. at FN3. They assume that M. whitei individuals would have Regarding the action of the giant hyena, the low tooth only accessed part of their prey’s visceral package, since the mark frequencies documented at FN3 are in stark contrast size and fragility of their canines may have prevented them with the damage generated by hyenas when they are the pri- from consuming much meat (Palmqvist et al., 1996, 1999, mary agent of carcass modification documented by different 2005, 2011; Espigares et al., 2013; Rodríguez-Gómez et al., authors (Kruuk 1972; Blumenschine, 1986; Villa & Bartram, 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 17 of 23 213 1996; Villa & Soresi, 1998; Faith, 2007; Domínguez-Rod- marks derived from marrow extraction (Table 5). These pat- rigo et al., 2015; Haynes & Klimowicz, 2015; Fernández- terns suggest that hominins had recurrent access to meat and Jalvo & Andrews, 2017; Haynes & Hutson 2020). This marrow resources at FN3. divergence suggests that the action of giant hyena at FN3 The lithic evidence described by Toro-Moyano et  al. was likely more limited than previously envisioned. (2003, 2009, 2011, 2013) and Barsky et al. (2010, 2013, Espigares et al. (2013) describe a M. meridionalis carcass 2015) is also in support of these interpretations. The lithic surrounded by lithic industry and giant hyena coprolites at analyses at FN 3 show a predominance of small-sized single FN3, which indicate the presence of these carnivores at the flakes of flint and limestone obtained from local sources, site. Nevertheless, no tooth marks or anthropogenic marks and some choppers (Toro-Moyano et al., (2003, 2009, 2011, were observed on the elephantid bones, thus not conclu- 2013; Barsky et al. 2010, 2013, 2015). These heavy-duty sively proving the interaction of humans or carnivores with tools may be associated with percussive activities oriented the proboscidean carcass. in part to the fragmentation of bones, as suggested by the Cut marks demonstrating direct hominin engagement existence of bones with green fractures (Table 4) and per- with the remains have been found on Middle Pleistocene cussion marks (Table 5). Furthermore, use-wear analyses elephant carcasses (e.g. Yravedra et  al. 2010), and this of FN3 flakes conducted by Toro-Moyano et  al. ( 2003) absence of taphonomic alterations is also in stark contrast documented their use in butchery activities. Recent exper- with actualistic reports of carnivore engagement with ele- imental studies have shown that small flakes are suitable phant carcasses, which tend to present a high number of for defleshing activities, even on very large carcasses, such tooth marks (Haynes & Klimowicz, 2015; Haynes & Hutson as proboscideans (Starkovich et al., 2021; Marinellu et al., 2020). Therefore, this spatial association of lithic finds and 2021). Lower Palaeolithic sites with small, simple flake coprolites in association with these elephant carcasses may lithic assemblages associated with cut-marked remains are have been fortuitous, resulting from independent episodes indeed very common (see Marinellu et al., 2021 for a more that coalesced into a palimpsest through complex site for- in-depth discussion). mation processes. Such coincidental associations of stone Based on these taphonomic arguments, the possibility tools and faunal remains are not infrequent in the Palaeo- that hominins at FN3 had early access to meat from fresh lithic record. At Bois Roche, lithic artefacts were found carcasses prior to carnivore engagement with the remains inside a hyena den, product of fluvial transportation (Villa cannot be dismissed. However, low cut mark frequencies in & Bartram, 1996; Villa & Soresi, 1998). Likewise, associa- relation to actualistic models (Figs. 6–8) hinder the robustic- tions of lithic artefacts with proboscidean remains are not ity of the conclusions that can be established regarding the uncommon (Gaudzinski et al 2005). Nevertheless, many of timing of hominin carcass access at FN3. It is also worth these associations have been shown to be the result of spatial considering the possibility that hominins may have followed processes unrelated to human subsistence strategies (Yrave- different carcass procurement strategies on the basis of ani- dra et al., 2014; Haynes et al 2020). mal size (small, medium, large, or very large), given the On the basis of the evidence documented at FN3, there diverse range of animal carcass sizes at FN3 level 5 present- are several important inferences that can be made regarding ing evidence of anthropogenic alterations. the role of hominins and carnivores at the site. The low tooth A convergence of different processes at the site is also mark frequencies at the site (Table 5, Figs. 3–5) suggest that plausible, with some animal deaths by natural causes, sev- carnivore impact on the FN3 bone assemblage was relatively eral episodes of primary and/or secondary hominin access to unimportant. The scarcity of digested bones, the absence of prey, and sporadic carnivore feeding bouts. This hypothetical diaphysiary cylinders, bones with pit tooth marks > 5 mm, scenario would explain the low cut mark frequencies while or individual bones with more than 5 tooth marks all sug- also accounting for the small number of tooth marks docu- gest that the role of the giant hyena was rather limited in mented in the faunal assemblage. the formation of the FN3 faunal assemblage. In the future, Nevertheless, until the limited evidence is complemented a geometric morphometric assessment of tooth marks (cf. by a revision of the materials from previous excavations Aramendi et al., 2017; Courtenay et al., 2021) could provide and sample sizes are enlarged with further newly excavated valuable insights into the specific carnivore(s) that played a remains, it will not be possible to conclusively define what role in the accumulation and/or alteration of the bone assem- role did humans play in the accumulation of the faunal blage at the site. assemblage at FN3. Likewise, more advanced statistical Regarding the role played by hominins, it is worth high- techniques may be able to shine new light on the nature lighting the considerable degree of bone sample fragmen- of this assemblage. Additionally, the incorporation of new tation (Table 4) and the presence of cut marks associated technologies applied to the study of cut marks could help us with butchery practices such as disarticulation, defleshing, develop new approaches. For example, recent approaches and evisceration (Table 6, Figs. 9–11), as well as percussion may allow the identification of which raw materials and 1 3 213 Page 18 of 23 Archaeol Anthropol Sci (2021) 13:213 the article's Creative Commons licence and your intended use is not types of lithic implements were employed in the butchery permitted by statutory regulation or exceeds the permitted use, you will activities documented at the site (Courtenay et al., 2019; need to obtain permission directly from the copyright holder. To view a Linares-Matás et  al., 2019; Maté-González et  al., 2019; copy of this licence, visit http://cr eativ ecommons. or g/licen ses/ b y/4.0/ . Yravedra et al., 2017, 2019). Lastly, the current study provides further zooarchaeologi- cal data on hominin subsistence strategies in the Early Pleis- References tocene settlement of the European subcontinent. The present study demonstrates the complexities and issues inherent to Abbazzi L (2010) La fauna de cérvidos de Barranco León y Fuente the study and interpretation of Early-Middle Pleistocene Nueva 3. In: Toro I, Martínez-Navarro B, Agustí J (Eds.), Ocupa- ciones Humanas en el Pleistoceno Inferior y Medio de la Cuenca assemblages, given the limitations imposed by the repre- de Guadix-Baza. Junta de Andalucía, Consejería de Cultura, sentativity of faunal samples. Nevertheless, FN3 joins other Sevilla, 273–290 Early Pleistocene sites older than 1 Ma across Europe where Agustí J, Madurell J (2003) Los arvicólidos (Muroidea, Rodentia, hominins had access to animal carcasses, such as Kozarnika Mammalia) del Pleistoceno inferior de Barranco León y Fuente Nueva 3 (Orce, Granada). Datos preliminares. In: Toro-Moyano, (Sirakov et al., 2010), Trilika (Vislobokova et al., 2020), or I., Agustí, J., Martínez-Navarro, B. (Eds.), El Pleistoceno infe- Pirro Nord (Cheheb et al., 2019), although there are still rior de Barranco León y Fuente Nueva 3, Orce (Granada). Junta some unresolved questions related to the temporality of car- de Andalucía. Consejería de Cultura. E.P.G.P.C. Arqueología cass access or whether hominins exhibited different acquisi- Monografías, Sevilla, pp. 137–147. Memoria Científica cam- pañas 1999–2002 tion strategies on the basis of carcass size. Agustí J, Arbiol S, Martin-Suarez E (1987) Roedores y lagomorfos (Mammalia) del Pleistoceno inferior de Venta Micena (Depresión Supplementary Information The online version contains supplemen- de Guadix-Baza, Granada). Paleontologia i Evolució, Memoria tary material available at https://doi. or g/10. 1007/ s12520- 021- 01461-7 . Especial 1:95–107 Agustí J, Oms O, Garcés M, Parés JM (1996) Calibration of the Late Acknowledgements We thank the researchers and students involved Pliocene-Early Pleistocene transition in the continental beds in the excavations, recovery, and preparation of the archeo-paleonto- of the Guadix-Baza Basin (southeastern Spain). Quatern Int logical record from the FN3. Juan Manuel Jiménez Arenas belongs to 40:93–100 the Excellence Unit “Archaeometrical Studies. Inside the artefacts & Agustí J, Oms O, Parés JM (2007) Biostratigraphy, paleomagnetism ecofacts” (University of Granada) and the Junta de Andalucía Research and geology of the Orce ravine (Southern Spain) Comment on Project “HUM-607”. Last but not least, we would like to thank the the paper by Gibert et al. (2006). Quaternary Science Reviews reviewers of this paper for their fruitful comments that have greatly 26:568–572 improved the first version of this paper. Agustí J, Blain HA, Furió M, De Marfá R, Santos-Cubedo A (2010) The Early Pleistocene small vertebrate succession from the Orce Funding Open Access funding provided thanks to the CRUE-CSIC region (Guadix-Baza Basin, SE Spain) and its bearing on the agreement with Springer Nature. This research was funded by the first human occupation of Europe. Quatern Int 223–224:162–169 Junta de Andalucía, Consejería de Educación, Cultura y Deporte: Alberdi MT (2010) Estudio de los caballos de los yacimientos de Orce Research Project “Primeras ocupaciones humanas y contexto Fuente Nueva-3 y Barranco León-5 (Granada). In: Toro I, paleoecológico a partir de los depósitos pliopleistocenos de la cuenca Martínez-Navarro B, Agustí J (eds) Ocupaciones Humanas en Guadix-Baza: zona arqueológica de la Cuenca de Orce (Granada, el Pleistoceno Inferior y Medio de la Cuenca de Guadix-Baza España), 2017–2020 (Ref. BC.03.032/17)”. We also received sup- Junta de Andalucía. Consejería de Cultura, Sevilla, pp 291–306 port from the PALARQ Foundation with the convocatory of Analitics Anadón P, Gabàs M (2009) Paleoenvironmental evolution of the Early 2019: “Identificando Carnívoros a partir de análisis Tafonómicos de Pleistocene lacustrine sequence at Barranco León archeological última generación aplicando Fotogrametría y Morfometría Geométrica site (Orce, Baza Basin, Southern Spain) from stable isotopes de las Marcas de Diente. Aplicación a Yacimientos del Pleistoceno and Sr and Mg chemistry of ostracod shells. J Paleolimnol Inferior Ibérico: FN3, Venta Micena 3 y 4 (Granada), Pontón de la 42:261–279 Oliva (Patones, Madrid)”. Lloyd Austin Courtenay is also funded by Anadón P, Utrilla R, Julià R (1994) Palaeoenvironmental reconstruc- the Spanish Ministry of Science, Innovation and Universities with an tion of a Pleistocene lacustrine sequence from faunal assem- FPI Predoctoral Grant (Ref. PRE2019-089411) associated to project blages and ostracode shell geochemistry, Baza Basin, SE Spain. RTI2018-099850-B-I00 and the University of Salamanca. The Institut Palaeogeogr Palaeoclimatol Palaeoecol 111:191–205 Català de Paleoecologia Humana I Evolució Social (IPHES-CERCA) Aramendi J, Maté-González MA, Yravedra J, Ortega MC, Arriaza has received financial support from the Spanish Ministry of Science MC, González-Aguilera D, Baquedano E, Domínguez-Rodrigo and Innovation through the “María de Maeztu” program for Units of M (2017) Discerning carnivore agency through the three-dimen- Excellence (CEX2019-000945-M). sional study of tooth pits: revisiting crocodile feeding behaviour at FLK- Zinj and FLK NN3 (Olduvai Gorge, Tanzania). Palaeo- Open Access This article is licensed under a Creative Commons Attri- geogr Palaeoclimatol Palaeoecol 488:93–102 bution 4.0 International License, which permits use, sharing, adapta- Arriaza MC, Domínguez-Rodrigo M, Yravedra J, Baquedano E (2016) tion, distribution and reproduction in any medium or format, as long Lions as bone accumulators? Paleontological and ecological as you give appropriate credit to the original author(s) and the source, implications of a modern bone assemblage from Olduvai Gorge. provide a link to the Creative Commons licence, and indicate if changes PLoS ONE 11(5):e0153797. https:// doi. or g/ 10. 1371/ jour n al. were made. The images or other third party material in this article are pone. 01537 97 included in the article's Creative Commons licence, unless indicated Arriaza MC, Aramendi J, Maté-González MÁ, Yravedra J, Stratford otherwise in a credit line to the material. If material is not included in D (2019) Characterizing leopard as taphonomic agent through 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 19 of 23 213 the use of micro-photogrammetric reconstruction of tooth marks Blumenschine RJ (1989) A landscape taphonomic model of the scale and pit to score ratio. Hist Biol 33:176–185 of prehistoric scavenging opportunities. J Hum Evol 18:345–371 Arribas A, Palmqvist P (1999) On the ecological connection between Blumenschine RJ, Cavallo JA (1992) Carroñeo y evolución humana. In: sabre-tooths and hominids: faunal dispersal events in the lower Libros de Investigación y Ciencia. Orígenes del Hombre Mod- Pleistocene and a review of the evidence for the first human erno, pp 90–97 arrival in Europe. J Archaeol Sci 26:571–585 Blumenschine RJ (1995) Percussion marks, tooth marks, and experi- Arribas A, Palmqvist P (2002) The first human dispersal to Europe: mental determination of the timing of hominid and carnivore remarks on the archaeological and palaeoanthropological record access to long bones at FLK Zinjanthropus, Olduvai Gorge, Tan- from Orce (Guadix-Baza basin, southeastern Spain). Hum Evol zania. J Hum Evol 29:21–51 17:55–78 Blumenschine RJ, Salvaggio MM (1988) Percussion marks on bone Arzarello M, De Weyer L, Peretto C (2016) The first European peo- surfaces as a new diagnostic of hominid behaviour. Nature pling and the Italian case: peculiarities and “opportunism.” Qua- 333:763–765 ternay International 393:41–50 Blumenshchine RJ, Marean CW, Capaldo SD (1996) Blind test of inter- Barsky D, Celiberti V, Cauche D, Grégoire S, Lebègue F, Lum- analyst correspondence and accuracy in the identification of cut ley H, Toro Moyano I (2010) Raw material discernment marks, percussion marks, and carnivore tooth marks on bone and technological aspects of the Barranco León and Fuente surfaces. J Archaeol Sci 23:493–507 Nueva 3 stone assemblages (Orce southern Spain). Quatern Boscaini A, Madurell-Malapiera J, Llenas M, Martínez-Navarro B Int 223–224:201–219 (2015) The origin of the critically endangered Iberian lynx: spe- Barsky D, Garcia J, Martínez K, Sala R, Zaidner Y, Carbonell ciation, diet and adaptive changes. Quatern Sci Rev 123:247–253 E, Toro-Moyano I (2013) Flake modification in European Bourguignon L, Crochet JY, Capdevila R, Ivorra J, Antoine PO, Agustí Early-Middle Pleisocne Stone tool assemblages. Quatern Int J, Barsky D, Blain HA, Boulbes N, Bruxelles L, Claude J, 316:140–154 Cochard D, de Weyer L, Filoux A, Firmat C, Lozano-Fernández Barsky D, Sala R, Menéndez L, Toro-Moyano I (2015) Use and re-use: I, Magniez P, Pelletier M, Rios J, Testu A, Valensi P (2015) re-knapped a fl kes from the Mode 1 site of Fuente Nueva 3 (Orce, Bois-de-Riquet (Lézignan-la-Cèbe, l’Hérault): a late Early Pleis- Andalucía, Spain). Quatern Int 361:21–33 tocene archeological occurrence in southern France. Quatern Int Barsky D, Vergès JM, Titton S, Guardiola M, Sala R, Toro-Moyano I 393:24–40 (2018) The emergence and significance of heavy-duty scrapers Brain CK (1969) The probable role of leopards as predators of the in ancient stone toolkits. CR Palevol 17:201–219 Swartkrans Australopithecines. South African Archaeological Bartolini-Lucenti SB, Madurell-Malapiera J (2020) Unraveling the Bulletin 24:170–171 fossil record of foxes: an updated review on the Plio-Pleisto- Bryant HN, Russell AP, Thomason JJ (1995) Carnassial functioning cene Vulpes spp. from Europe. Quaternary Science Reviews in nimravid and felid sabertooths: theoretical basis and robust- 236:106296 ness of inferences. In: Thomason JJ (ed) Functional morphology Behrensmeyer AK (1978) Taphonomic and ecologic information from in vertebrate paleontology. Cambridge University Press, Cam- bone weathering. Paleobiology 4:150–162 bridge, pp 116–135 Binford LR (1981) Bones: ancient men and modern myths. Academic Bunn HT (1982) Meat eating and human evolution: studies on the diet Press, New York and subsistence patterns of plio-pleistocene hominids in East Blain H-A (2005) Contribution de la paléoherpétofaune (Amphibia Africa. Universidad de California, Berkeley, PhD & Squamata) à la connaissance de l’évolution du climat et du Bunn HT, Ezzo JA (1993) Hunting and scavenging by Plio-Pleistocene paysage du Pliocène supérieur au Pléistocène moyen d’Espagne. Hominids: nutritional constraints, archaeological patterns, and Doctorat du Muséum National d’Histoire Naturelle de Paris. behavioural implications. J Archaeol Sci 20:365–398. https:// Département de Préhistoire. Institut de Paléontologie humainedoi. org/ 10. 1006/ jasc. 1993. 1023 Blain H-A (2009) Contribution de la paléoherpétofaune (Amphibia Bunn HT, Pickering TR (2010) Bovid mortality profiles in paleoeco- & Squamata) à la connaissance de l’évolution du climat et du logical context falsify hypotheses of endurance running–hunting paysage du Pliocène supérieur au Pléistocène moyen d’Espagne. and passive scavenging by Early Pleistocene hominins. Quatern Treballs Del Museu Geologic De Barcelona 16:39–170 Res 74:395–404. https:// doi. org/ 10. 1016/j. yqres. 2010. 07. 012 Blain H-A, Bailon S (2010) Anfibios y escamosos del Pleistoceno infe- Carbonell E, Bermúdez de Castro JM, Parés JM, Pérez-González A, rior de Barranco León y de Fuente Nueva 3 (Orce, Andalucía, Cuenca- Bescós G, Ollé A, Mosquera M, Huguet R, Van der España). In: Toro, I., Martínez-Navarro, B., Agustí, J. (Eds.), Made J, Rosas A, Sala R, Vallverdú J, García N, Granger DJ, Ocupaciones humanas en el Pleistoceno inferior y medio en la Martinón-Torres M, Rodríguez XP, Stock GM, Vergés JM, Allué Cuenca de Guadix-Baza. Arqueología Monografías, Sevilla, pp. E, Burjachs F, Cáceres I, Canals A, Benito A, Díez C, Lozano M, 165–183 Mateos A, Navazo M, Rodríguez J, Rosell J, Arsuaga JL (2008) Blain H-A, Bailon S, Agustí J, Martínez-Navarro B, Toro I (2011) The first hominin of Europe. Nature 452:465–469 Paleoenvironmental and paleoclimatic proxies to the Early Cheheb RC, Arzarello M, Arnaud J, Berto C, Cáceres I, Caracausi Pleistocene hominids of Barranco León D and Fuente Nueva 3 S, Colopi F, Daffara S, Canini GM, Huguet R, Karambatsou T, (Granada, Spain) by means of their amphibian and reptile assem- Sala B, Zambaldi M, Berruti GLF (2019) Human behavior and blages. Quatern Int 243:44–53 Homo-mammal interactions at the first European peopling: new Blain H-A, Lozano-Fernández I, Agustí J, Bailon S, Menèndez L, Espí- evidence from the Pirro Nord site (Apricena, Southern Italy). The gares Ortiz MP, Ros-Montoya S, Jiménez Arenas JM, Toro I, Science of Nature 106:16 Martínez-Navarro B, Sala R (2016) Refining upon the climatic Clark J, Linares-Matas G (2021) Seasonality and Oldowan behavioral background of the Early Pleistocene hominid settlement in West- variability in East Africa. J Hum Evol, in press. https:// doi. org/ ern Europe: Barranco León and FN3 (Guadix-Baza basin, SE 10. 1016/j. jhevol. 2021. 103070 Spain). Quatern Sci Rev 144:132–144 Courtenay LA, Yravedra J, Mate-González MÁ, Aramendi J, González- Blumenschine RJ (1986) Early hominid scavenging opportunites. Aguilera D (2019) 3D analysis of cut marks using a new geomet- Implications of carcass availability in the Serengeti and ric morphometric methodological approach. Archaeol Anthropol Ngorongoro ecosystems. Oxford: BAR International Series 283 Sci 11:651–665 1 3 213 Page 20 of 23 Archaeol Anthropol Sci (2021) 13:213 Courtenay LA, Herranz-Rodrigo D, González-Aguilera D et al (2021) Fernández Jalvo Y, Andrews P (2017) Atlas of taphonomic identifica- Developments in data science solutions for carnivore tooth tion. Springer. Ed. pit classification. Sci Rep 11:10209. https:// doi. org/ 10. 1038/ Ferraro JV, Plummer TW, Pobiner BL, Oliver JS, Bishop LC, Braun s41598- 021- 89518-4 DR, Ditchfield PW, Seaman JW III, Binetti KM, Seaman JW de Lombera-Hermida A, Bargallo A, Terradillos-Bernal M, Huguet R, Jr, Hertel F, Potts R (2013) Earliest archaeological evidence of Vallverdú J, García-Anton M-D, Mosquera M, Ollé A, Sala R, persistent hominin carnivory. PLoS ONE 8:e62174. https:// doi. Carbonell E, Rodríguez-Alvarez XP (2015) The lithic industry org/ 10. 1371/ journ al. pone. 00621 74 of Sima del Elefante (Ata- puerca, Burgos, Spain) in the context Gaudzinski S, Turner E, Anzidei AP, Àlvarez-Fernández E, Arroyo- of Early and Middle Pleistocene technology in Europe. J Hum Cabrales J, Cinq-Mars J, Dobosi VT, Hannus A, Johnson E, Evol 82:95–106 Münzel Ss C, Scheer A, Villa P (2005) The use of proboscidean Desantis LRG, Schubert BW, Scott JR, Ungar PS (2012) Implications remains in every-day Palaeolithic life. Quaternary International of diet for the extinction of saber-toothed cats and American 126–128:179–194. https://doi. or g/10. 1016/j. q uaint.2004. 04. 022 lions. PLoS ONE 7:e52453. https:// doi. or g/ 10. 1371/ jour n al. Gibert J, Iglesias A, Maillo A, Gibert L (1992) Industrias líticas en el pone. 00524 53 Pleistoceno inferior de la región de Orce. Projecto Orce-Cueva Despriée J, Moncel MH, Arzarello M, Courcimault G, Voinchet P, Victoria 1988e1992. In: Gibert, J. (Ed.), Museo de Prehisto- Bahain JJ, Falguéres C (2018) The 1-million-year-old quartz ria: Presencia humana en el Pleistoceno inferior de Granada assemblage from Pont-de-Lavaud (Centre, France) in the Euro- y Murcia. Ediciones del Museo de Prehistoria, Ayuntamiento pean context. J Quat Sci 33:639–661 de Orce, 219–282 Domínguez-Rodrigo M (2002) Hunting and scavenging by early Gibert J, Gibert L, Iglesias A, Maestro E (1998) Two ‘Oldowan’ humans: the state of debate, Journal of World Prehistory 16(1), assemblages in the Plio-Pleistocene deposits of the Orce Heidelberg, 1–54 region, southeast Spain. Antiquity 72:17–25 Domínguez-Rodrigo M et al (2002) The ST site complex at Peninj, Gradstein FM, Ogg JG, Smith AG (2005) A geologic time scale. West Lake Natron, Tanzania: implications for early hominid Cambridge University Press, Cambridge behavioural models. J Archaeol Sci 29, Oxford, 639–665 Harstone-Rose A (2008) Evaluating the hominin scavenging niche Domínguez-Rodrigo M, Mabulla A, Bunn HT, Barba R, Diez-Martín F, through analysis of the carcass-processing abilities of the car- Egeland CP, Espílez E, Egeland A, Yravedra J, Sánchez P (2009) nivore guild. PhD of Philosophy in the Department of Bio- Unraveling hominin behavior at another anthropogenic site from logical Anthropology and Anatomy in the Graduate School of Olduvai Gorge (Tanzania): new archaeological and taphonomic Duke University research at BK, Upper Bed II. J Hum Evol 57(3):260–283. Hartstone-Rose A (2011) Reconstructing the diets of extinct South https:// doi. org/ 10. 1016/j. jhevol. 2009. 04. 006 African carnivorans from premolar “intercuspid notch” mor- Domínguez-Rodrigo M, Yravedra Y, Organista E, Gidna A, Fourvel phology. J Zool 285:119–127 J-B, Baquedano E (2015) A new methodological approach to the Haynes G, Klimowicz J (2015) Recent elephant-carcass utilization taphonomic study of paleontological and archaeological faunal as a basis for interpreting mammoth exploitation. Quatern Int assemblages: a preliminary case study from Olduvai Gorge (Tan- 359–360:19–37 zania). J Archaeol Sci 59:35–53 Haynes G, Krasinski K, Wojtal P (2020) A study of fractured pro- Domínguez-Rodrigo M, Cifuentes G, Jiménez B, Abellán N, Pizarro boscidean bones in Recent and Fossil Assemblages. J Archaeol M, Organista E, Baquedano E (2020) Artificial intelligence pro- Method Theory. https:// doi. org/ 10. 1007/ s10816- 020- 09486-3 vides greater accuracy in the classification of modern and ancient Haynes G, Hutson J (2020) African elephant bones modified by bone surface modifications. Sci Rep 10:18862 carnivores: Implications for interpreting fossil proboscidean Duval M, Aubert M, Hellstrom J, Grün R (2011) High resolution LA- assemblages. J Archaeol Sci Rep 34:102596. https:// doi. org/ ICP-MS mapping of U and Th isotopes in an Early Pleistocene 10. 1016/j. jasrep. 2020. 102596 equid tooth from FN3 (Orce, Andalusia, Spain). Quat Geochro- Huguet R, Saladié P, Cáceres I, Díez C, Rosell J, Bennasar M, Blasco nol 6:458–467. https:// doi. org/ 10. 1016/j. quageo. 2011. 04. 002 R, Esteban-Nadal M, Gabucio J, Rodríguez-Hidalgo A, Car- Duval M, Falguères C, Bahain JJ (2012a) Age of the oldest hominin bonell E (2013) Successful subsistence strategies of the first settlements in Spain: contribution of the combined U-series/ humans in south-western Europe. Quatern Int 295:168–182 ESR dating method applied to fossil teeth. Quat Geochronol Huguet R, Vallverdú J, Rodríguez-Álvarez XP, Terradillos-Bernal M, 10:412–417 Bargalló A, Lombera-Hermida A, Menéndez L, Modesto-Mata Duval M, Falguères C, Bahain J-J, Grün R, Shao Q, Aubert M, Dolo M, Van der Made J, Soto M, Blain H-A, García N, Cuenca- JM, Agustí J, Martínez-Navarro B, Palmqvist P, Toro-Moyano I Bescós G, Gómez-Merino G, Pérez-Martínez R, Expósito I, (2012b) On the limits of using combined U-series/ESR method Allué E, Rofes J, Burjachs F, Canals A, Bennàsar M, Nuñez- to date fossil teeth from two Early Pleistocene archaeological Lahuerta C, Bermúdez de Castro JM, Carbonell E (2017) Level sites of the Orce area (Guadix-Baza basin, Spain). Quatern Res TE9c of Sima del Elefante (Sierra de Atapuerca, Spain): a 77:482–491 comprehensive approach. Quatern Int 433:278–295 Espigares MP, Martinez-Navarro B, Palmqvist P, Ros-Montoya S, Toro Kruuk H (1972) The sppoted hyena: a study of predation and social I, Agustí J, Sala R (2013) Homo vs. Pachycrocuta: earliest evi- behavior. University of Chicago Press, Chicago dence of competition for an elephant carcass between scavengers Linares-Matás G, Yravedra J, Maté-González J, Aramendi J, Cour- at FN3 (Orce, Spain). Quatern Int 295:113–125 tenay Ll, Cuartero F, González-Aguilera D (2019) A geomet- Espigares MP, Palmqvist P, Ros-Montoya G-M, S., García-Aguilar, ric-morphometric assessment of three-dimensional models J.M. Rodríguez-Gómez, G., Serrano, F., Martínez-Navarro, B. of experimental cut-marks generated using flint and quartzite (2019) The earliest cut marks of Europe: a discussion on hominin flakes and handaxes. Quatern Int 517:45–54 subsistence patterns in the Orce sites (Baza basin, SE Spain). Lozano-Fernandez I, Blain HA, Agusti J, Piñero P, Barsky D, Ivorra Sci Rep 9:15408 J, Bourguignon L (2019) New clues about the late Early Pleis- Faith JT (2007) Sources of variation in carnivore tooth-mark frequen- tocene peopling of Western Europe: small vertebrates from the cies in a modern spotted hyena (Crocuta crocuta) den assem- Bois-de-Riquet archeopaleontological site (Lézignan-La-Cèbe, blage, Amboseli Park, Kenya. J Archaeol Sci 34:1601–1609 southern France). Quatern Sci Rev 219:187–203 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 21 of 23 213 Lozano-Fernandez I, Blain HA, López-García JM, Agustí J (2015a) Michel V, Shen CC, Woodhead J, Hu HM, Wu CC, Moullé PÉ, … De Biochronology of the first hominid remains in Europe using Lumley H (2017). New dating evidence of the early presence of the vole Mimomys savini: Fuente Nueva 3 and Barranco hominins in Southern Europe. Scientific Reports 7, 10074 León D, Guadix-Baza Basin, south-eastern Spain. Hist Biol Monahan CM (1996) New zooarchaeological data from Bed II, Olduvai 27:1021–1028 Gorge, Tanzania: implications for hominid behavior in the Early Lozano-Fernandez S, Mateos A, Rodríguez J (2015b) Exploring Pleistocene. J Hum Evol 31:93–128 paleo food-webs in the European Early and Middle Pleisto- Oliver JS, Plummer TW, Hertel F, Bishop LC (2019) Bovid mortality cene: a Network Analysis. Quatern Int 413:44–54 patterns from Kanjera South, Homa Peninsula, Kenya and FLK- Luzón C, Courtenay LA, Yravedra J, Saarinen J, Blain H-A, Zinj, Olduvai Gorge, Tanzania: evidence for habitat mediated DeMiguel D, Viranta S, Azanza B, Rodríguez-Alba JJ, Her- variability in Oldowan hominin hunting and scavenging behavior. ranz-Rodrigo D, Solano JA, Oms O, Agustí J, Fortelius M, J Hum Evol 131:61–75. https://doi. or g/10. 1016/j. jhe vol.2019. 03. Jiménez Arenas JM (2021) Taphonomic and spatial analyses 009 from the Early Pleistocene site of Venta Micena 4 (Orce, Gua- Oms O, Dinares-Turell J, Agustí J, Parés JM (1999) Refinements of dix-Baza Basin, southern Spain). Sci Rep 11:13977 the European mammal chronology from the magnetic polarity Madurell-Malapeira J, Martínez-Navarro B, Ros-Montoya S, Espi- record of the Plio-Pleistocene Zújar section, Guadix-Baza Basin, gares MP, Toro I, Palmqvist P (2011) The earliest European SE Spain. Quatern Res 51:94–103 badger (Meles meles), from the Late Villafranchian site of Oms O, Agustí J, Gabàs M, Anadón P (2000a) Lithostratigraphical cor- Fuente Nueva 3 (Orce, Granada, SE Iberian Peninsula). CR relation of micromammal sites and biostratigraphy of the Upper Palevol 10:609–615 Pliocene to Lower Pleistocene in the Northeast Guadix-Baza Maldonado-Garrido E, Piñero P, Agustí J (2017) A catalogue of Basin. J Quat Sci 15:43–50 the vertebrate fossil record from the Guadix-Baza Basin (SE Oms O, Parés JM, Martínez-Navarro B, Agustí J, Toro-Moyano I, Mar- Spain). Spanish Journal of Palaeontology 32:207–236 tínez Fernández G, Turq A (2000b) Early human occupation of Marean CW, Ehrhardt CL (1995) Paleoanthropological and paleoeco- Western Europe: paleomagnetic dates for two Paleolithic sites in logical implications of the taphonomy of a sabertooth’s den. J Spain. Proceedings of the National Academy of Sciences USA Hum Evol 29:515–547 97:10666–10670 Marinellu F, Lemorini C, Barkai R (2021) Lower palaeolithic small Oms O, Agustí J, Parés JM (2010) Litoestratigrafía, magnetoes- flakes and megafauna: the contribution of experimental approach tratigrafía y bioestratigrafía de los yacimientos de Barranco and use-wear analysis to reveal the link. In Konidaris, GE Barkai, León 5 y Fuente Nueva 3 (Cuenca, Guadix-Baza). En B. Mar- R Tourloukis, V Harvati, K (eds) Human-elephant interactions: tínez-Navarro, J. Agustí, I. Toro Moyano (coords.), Ocupaciones from past to present. Tübingen University Press, Tübingen, p humanas en el Pleistoceno Inferior y Medio de la cuenca de 237–260. https:// doi. org/ 10. 15496/ publi kation- 55604 Guadix-Baza. Junta de Andalucía, Consejería de Cultura. Sevilla, Martinez-Navarro B, Palmqvist P (1995) Presence of the African 107–120 machairodont Megantereon whitei (Broom, 1937) (Felidae, Oms O, Anadón P, Agustí J, Julià R (2011) Geology and chronology Carnivora, Mammalia) in the Lower Pleistocene site of Venta of the continental Pleistocene archeological and mammal sites Micena (Orce, Granada, Spain), with some considerations on of the Orce Area (Baza Basin, Spain). Quatern Int 243:33–43 the origin, evolution and dispersal of the genus. J Archaeol Sci Palmqvist P, Martínez-Navarro B, Arribas A (1996) Prey selection 22:569–582 by terrestrial carnivores in a lower Pleistocene paleocommunity. Martinez-Navarro B, Turq A, Agustí J, Oms O (1997) FN3 (Orce, Paleobiology 22:514–534 Granada, Spain) and the first human occupation of Europe. J Palmqvist P, Arribas A, Martínez-Navarro B (1999) Ecomorphological Hum Evol 33:611–620 study of large canids from southeastern Spain. Lethaia 32:75–88 Martínez-Navarro B, Espigares MP, Ros-Montoya S, (2003) Estudio Palmqvist P, Martínez-Navarro B, Toro I, Espigares MP, Ros-Montoya preliminar de las asociaciones de grandes mamíferos de Fuente S, Torregrosa V, Pérez-Claros JA (2005) A re-evaluation of the Nueva 3 y Barranco León(Orce, Granada, España) (Informe de evidence of human presence during Early Pleistocene times in las campañas de 1999e2002). In: Toro-Moyano, I., Agustí, J., southeastern Spain. L’anthropologie 109:411–450 Martínez-Navarro, B. (Eds.), El Pleistoceno inferior de Barranco Palmqvist P, Martínez-Navarro B, Pérez-Claros JA, Torregrosa V, León y Fuente Nueva 3, Orce (Granada). Junta de Andalucía. Figueirido B, Jiménez-Arenas JM, Patrocinio-Espigares M, Ros- Consejería de Cultura. E.P.G.P.C. Arqueología Monografías, Montoya S, De Renzi M (2011) The giant hyena Pachycrocuta Sevilla, 115–137. Memoria Científica campañas 1999–2002 brevirostris: modelling the bone-cracking behavior of an extinct Martínez-Navarro B, Palmqvist P, Madurell-Malapeira J, Ros-Montoya carnivore. Quatern Int 243:61–79 S, Espigares M-P, Torregrosa V, Pérez-Claros JA (2010) La fauna Pares JM, Pérez-González A, Rosas A, Benito A, Bermúdez de Castro de grandes mamíferos de FN3 y Barranco León-5: Estado de JM, Carbonell E, Huguet R (2006) Matuyama-age lithic tools la cuestión. In: Toro, I., Martínez-Navarro B, Agustí J (Eds.), from the Sima del Elefante site, Atapuerca (northern Spain). J Ocupaciones Humanas en el Pleistoceno Inferior y Medio de Hum Evol 50:163–169 la Cuenca de Guadix-Baza. Junta de Andalucía, Consejería de Parkinson JA (2018) Revisiting the hunting-versus-scavenging debate Cultura, Sevilla, 197–236 at FLK Zinj: a GIS spatial analysis of bone surface modifications Maté-González MA, González-Aguilera D, Linares-Matás G, Yrave- produced by hominins and carnivores in the FLK 22 assemblage, dra J (2019) New technologies applied to modelling taphonomic Olduvai Gorge, Tanzania. Palaeogeogr Palaeoclimatol Palaeo- alterations. Quatern Int 517:4–15 ecol 511:29–51. https:// doi. org/ 10. 1016/j. palaeo. 2018. 06. 044 Mech LD (1970) The wolf: the ecology and behavior of an Endargered Pante MC (2013) The larger mammal fossil assemblage from JK2, Species. The Natural History Press, Garden City Bed III, Olduvai Gorge, Tanzania: implications for the feeding Medin T, Martínez-Navarro B, Rivals F, Madurell-Malapiera J, Ros- behavior of Homo erectus. J Hum Evol 64:68–82 Montoya S, Espigares MP, Figueirido B, Rook L, Palmqvist P Pavia M, Zunino M, Coltorti M, Angelone C, Arzarello M, Bagnus C, (2017) Late Villafranchian Ursus etruscus and other large car- Bellucci L, Colombero S, Marcolini F, Peretto C, Petronio C, nivorans from the Orce sites (Guadix-Baza basin, Andalusia, Petrucci M, Pieruccini P, Sardella R, Tema E, Villier B, Pavia G (2012) Stratigraphical and palaeontological data from the Early southern Spain): taxonomy, biochronology, paleobiology, and ecogeographical context. Quatern Int 431:20–41 1 3 213 Page 22 of 23 Archaeol Anthropol Sci (2021) 13:213 Pleistocene Pirro 10 site of Pirro Nord (Puglia, south eastern Stahler DR, Smith DW, Guernsey DS (2006) Foraging and feeding Italy). Quatern Int 267:40–55 ecology of the gray wolf (Canis lupus): lessons from Yellowstone Pickering TR, Egeland CP, Dominguez-Rodrigo M, Brain CK, Schnell National Park, Wyoming, USA. J Nutr 136(7 Suppl.):1923–1926 AG (2008) Testing the “shift in the balance of power” hypoth- Starkovich BM et al (2021) Minimal tools, maximum meat: a pilot esis at Swartkrans, South Africa: hominid cave use and subsist- experiment to butcher an elephant foot and make elephant bone ence behavior in the Early Pleistocene. J Anthropol Archaeol tools using Lower Paleolithic stone tool technology. Ethnoar- 27:30–45 chaeology 12:118–147 Pobiner, B., (2007) Hominid-carnivore interactions: evidence from Tappen M (1995) Savanna ecology and natural bone deposition: impli- modern carnivore bone modification and Early Pleistocene cations for early hominid site formation, hunting and scavenging. archaeofaunas (Koobi Fora, Kenya; Olduvai Gorge, Tanzania). Curr Anthropol 36:223–260 Ph.D. dissertation. Department of Anthropology, Rutgers Uni- Titton S, Barsky D, Bargalló A, Serrano-Ramos A, Vergès JM, versity, New Brunswick. Toro-Moyano I, Sala R, García-Solano J, Jiménez-Arenas JM Pobiner BL, Rogers MJ, Monahan CM, Harris JWK (2008) New (2020) Subspheroids in the lithic assemblage of Barranco León evidence for hominin carcass processing strategies at 1.5 Ma, (Spain): recognizing the late Oldowan in Europe. PLoS One Koobi Fora, Kenya. Journal of Human Evolution 55:103–130 15(3):e0231036 Reinoso-Gordo JF, Barsky D, Serrano-Ramos A, Solano-García Titton S, Barsky D, Bargalló D, Vergès JM, Guardiola M, García- JA, León-Robles CA, Luzón-González C, Titton S, Jiménez- Solano J, Jiménez-Arenas JM, Toro-Moyano I, Sala R (2019) Arenas JM (2020) Walking among mammoths. Remote sensing Active percussion tools from the Oldowan site of Barranco León and virtual reality supporting the study and dissemination of (Orce, Andalousie, Spain): the fundamental role of pounding Pleistocene archaeological sites: the case of Fuente Nueva 3 activities in hominin lifeways. J Archaeol Sci 96:131–147 in Orce, Spain. Sustainability 12:4785 Tixier PJ, y Roche, H. (1995) Polyedre, sub-spheroïde, spheroïde et Rodríguez J, Rodríguez-Gómez G, Martín-González JA, Goikoetxea bola: des segments plus ou moins longs d’une même chaîne I, Mateos A (2012) Predator-prey relationships and the role operatoire. Cahier Noir 7:31–40 of Homo in Early Pleistocene food webs in Southern Europe. Thompson CEL, Ball S, Thompson TJU, Gowland R (2011) The abra- Palaeogeogr Palaeoclimatol Palaeoecol 365–366:99–114 sion of modern and archaeological bones by mobile sediments: Rodríguez-Gómez G, Palmqvist P, Rodríguez J, Mateos A, Martín- the importance of transport modes. J Archaeol Sci 38:784–793 González JA, Espigares MP, Ros-Montoya S, Martínez-Nav- Toro-Moyano I, Lumley H de, Barsky D, Cauche D, Celiberti, V. y arro B (2016) On the ecological context of the earliest human Moncel M.-H. (2003) Las industrias líticas de Barranco León y settlements in Europe: resource availability and competition Fuente Nueva 3. Estudio técnico y tipológico. Las cadenas opera- intensity in the carnivore guild of Barranco León-D and FN3 tivas. Análisis traceológico. Resultados preliminares. Memoria (Orce, Baza Basin, SE Spain). Quatern Sci Rev 134:69–83 Científica campañas 1999–2002. In : Toro, I., Agustí, J., Mar - Rodríguez-Alba J-J, Linares-Matás G, Yravedra J (2019) First assess- tínez-Navarro, B. (Editores), El Pleistoceno inferior de Barranco ments of the taphonomic behaviour of jaguar (Panthera onca). León y FN3, Orce (Granada). Junta de Andalucía. Consejería de Quatern Int 517:88–96 Cultura, Arqueología Monografías, Sevilla, 173–183 Roe DA (1995) The Orce Basin (Andalucia, Spain) and the initial Toro-Moyano I, de Lumley H, Fajardo B, Barsky D, Celiberti V, Gré- Palaeolithic of Europe. Oxf J Archaeol 14:1–12 goire S, Martínez-Navarro B, Espigares MP, y Ros-Montoya, S., Ros-Montoya S (2010) Los Proboscídeos del Plio-Pleistoceno de las (2009) L’industrie lithique des gisements du pleistocène infé- cuencas de Guadix-Baza y Granada. Editorial de la Universi- rieur de Barranco León et Fuente Nueva 3, Granade, Espagne. dad de Granada, Granada PhD L’anthropologie 113:111–124 Ros-Montoya S, Bartolini-Lucenti S, Espigares MP, Palmqvist P, Mar- Toro-Moyano I, Lumley H de, Barrier P, Barsky D, Cauche D, Celiberti tínez-Navarro B (2021) First review of Lyncodontini material V, Grégoire S, Lebègue F, Mestour B. y Moncel MH (2010a) Les (Mustelidae, Carnivora, Mammalia) from the Lower Pleistocene industries lithiques archaïques du Barranco León et de Fuente archaeo-palaeontological sites of Orce (Southeastern Spain). Nueva 3, Orce, basin du Guadix-Baza, Andalousian. En : Mon- Rivista Italiana De Paleontologia e Stratigrafia 127:33–47 ography, CNRS Éditions Saarinen J, Oksanen O, Žliobaitė I, Fortelius M, DeMiguel D, Azanza Toro-Moyano I, Martínez-Navarro B, Agustí J, (2010b) Ocupaciones B, Bocherens H, Luzón C, Solano-García J, Yravedra J, Cour- Humanas en el Pleistoceno inferior y medio de la cuenca de tenay LA, Blain H-A, Sánchez-Bandera C, Serrano-Ramos A, Guadix-Baza. Memoria Científica. Junta de Andalucía, Conse- Rodriguez-Alba JJ, Viranta S, Barsky B, Tallavaara M, Oms O, jería de Cultura, EPG Arqueología Monografias Agustí J, Ochando J, Carrión JS, Jiménez-Arenas JM (2021) Plio- Toro-Moyano I, Barsky D, Cauche D, Celiberti V, Grégoire S, Leb- cene to Middle Pleistocene climate history in the Guadix-Baza egue F, Moncel MH, de Lumley H (2011) The archaic stone-tool Basin, and the environmental conditions of early Homo dispersal industry from Barranco León and Fuente Nueva 3, (Orce, Spain): in Europe. Quaternary Science Reviews 268:107132 evidence of the earliest hominin presence in southern Europe. Schaller GB (1972) The Serengeti lion: a study of predator-prey rela- Quatern Int 243:80–91 tions. University of Chicago Press, Chicago Toro-Moyano I, Martínez-Navarro B, Agustí J, Souday C, Bermúdez Scott G, Gibert L, Gibert J (2007) Magnetostratigraphy of the Orce de Castro JM, Martinón-Torres M, Fajardo B, Duval M, Fal- region (Baza basin), SE Spain: new chronologies for Early Pleis- guères C, Oms O, Parés JM, Anadón P, Julià R, García-Aguilar tocene faunas and hominid occupation sites. Quatern Sci Rev JM, Moigne AM, Espigares MP, Ros-Montoya S, Palmqvist P 26:415–435 (2013) The oldest human fossil in Europe, from Orce (Spain). J Sirakov N, Guadelli J-L, Ivanova S, Sirakova S, Boudadi-Maligne M, Hum Evol 65:1–9 Dimitrova I, Fernandez P, Ferrier C, Guadelli A, Iordanova D, Treves A, Treves HL (1999) Risk and oportunity for humans coexisting Iordanova N, Kovatcheva M, Krumov I, Leblanc JC, Miteva V, with large carnivores. J Hum Evol 336:275–282 Popov R, Spassov R, Taneva S, Tsanova T (2010) An ancient Turq A, Martínez-Navarro B, Palmqvist P, Arribas A, Agustí J, Rod- continuous human presence in the Balkans and the beginnings of ríguez Vidal J (1996) Le Plio-Pléistocène de la région d’Orce, human settlement in western Eurasia: a Lower Pleistocene exam- province de Grenade, Espagne : bilan et perspectives de recherche. Paléo 8:161–204 ple of the Lower Palaeolithic levels in Kozarnika cave (North- western Bulgaria). Quatern Int 223–224:94–106 1 3 Archaeol Anthropol Sci (2021) 13:213 Page 23 of 23 213 Van der Made J, Carlos Calero JA, Mancheño MA (2008) New material Yravedra J, Lagos L, Bárcena F (2011) A taphonomic study of wild of the goat Capra? alba from the Lower Pleistocene of Quibas wolf (Canis lupus) modification of horse bones in Northwestern (Spain); notes on sexual dimorphism, stratigraphic distribution Spain. Journal of Taphonomy 9:37–65 and systematics. Bollettino Della Società Paleontologica Italiana Yravedra J, Maté-González MA, Palomeque-González JF, Aramendi 47:13–23 J, Estaca-Gómez V, San Juan Blazquez M, García Vargas E, Van Valkenburgh B, Ruff CB (1987) Canine tooth strength and killing Organista E, González-Aguilera D, Arriaza MC, Cobo-Sánchez behaviour in large carnivores. J Zool 212:379–397 L, Gidna A, Uribelarrea Del Val D, Baquedano E, Mabulla A, Villa P, Mahieu E (1991) Breakage patterns of human long bones. J Domínguez-Rodrigo M (2017) A new approach to raw mate- Hum Evol 21:27–48 rial use in the exploitation of animal carcasses at BK (Upper Villa P, Bartram L (1996) Flaked bone from a hyaena den. Paléo Bed II, Olduvai Gorge, Tanzania): a micro-photogrammetric and 8:143–159 geometric morphometric analysis of fossil cut marks. Boreas Villa P, Soresi M (1998) Stones tools in carnivore sites. The case of 46:860–873. https:// doi. org/ 10. 1111/ bor. 12224 Bois Roche. J Archaeol Res 56:187–215 Yravedra J, Maté-González MA, Courtenay L, López-Cisneros P, Est- Vislobokova IA, Agadzhanyan AK, Lopatin AV (2020) The case of aca-Gómez V, Aramendi J, de Andrés-Herrero M, Linares-Matás Trlica TRL11–10 (Montenegro): implications for possible early G, González-Aguilera D, Álvarez-Alonso D (2019) Approaching hominin dispersals into the Balkans in the middle of the Early raw material functionality in the Upper Magdalenian of Coímbre Pleistocene. Quatern Int 554:15–35 cave (Asturias, Spain) through geometric morphometrics. Qua- Yravedra J (2006) Tafonomía aplicada a Zooarqueología. Aula Abierta, tern Int 517:97–106 UNED, Madrid Yravedra J, Rubio-Jara S, Courtenay Ll, A., Martos, J.A. (2020) Mam- Yravedra J, Domínguez-Rodrigo M (2009) The shaft-based methodo- mal butchery by Homo erectus at the Lower Pleistocene acheu- logical approach to the quantification of long limb bones and its lean site of Juma’s korongo 2 (JK2), bed III, Olduvai Gorge. relevance to understanding hominin subsistence in the Pleisto- Tanzania Quaternary Science Reviews 249(1):106612. https:// cene: application to four Paleolithic sites. J Quat Sci 24:85–96doi. org/ 10. 1016/j. quasc irev. 2020. 106612 Yravedra J, Domínguez-Rodrigo M, Santonja M, Pérez-González A, Zaidner Y (2013) Adaptive flexibility of Oldowan hominins: secondary Panera J, Rubio-Jara S, Baquedano E (2010) Cut marks on the use of flakes at Bizat Ruhama. Israel. PLoS One 8(6):e66851 Middle Pleistocene elephant carcass of Áridos 2 (Madrid, Spain). J Archaeol Sci 37:2469–2476 Publisher's note Springer Nature remains neutral with regard to Yravedra J, Panera J, Rubio-Jara S, Manzano I, Expósito A, Pérez- jurisdictional claims in published maps and institutional affiliations. González A, Soto E, López-Recio M (2014) Neanderthal and Mammuthus interactions at EDAR Culebro 1 (Madrid, Spain). J Archaeol Sci 42:500–508 1 3

Journal

Archaeological and Anthropological SciencesSpringer Journals

Published: Dec 1, 2021

Keywords: Early Pleistocene; Taphonomy; Hominin-carnivore interactions; Zooarchaeology; Cut marks; Palaeolithic archaeology

There are no references for this article.