Going beyond the potential equifinality problems: A response to Saladié and Rodríguez-Hidalgo (2019)

Journal Pre-proof Going beyond the potential equifinality problems: A response to Saladié and Rodríguez-Hidalgo (2019) Jordi Rosell, Ruth Blasco, Maite Arilla, Yolanda Fernández-Jalvo PII:

S1040-6182(19)30876-6

DOI:

https://doi.org/10.1016/j.quaint.2019.11.031

Reference:

JQI 8064

To appear in:

Quaternary International

Please cite this article as: Rosell, J., Arilla, M., Blasco, R., Fernández-Jalvo, Y., Going beyond the potential equifinality problems: a response to Saladié and Rodríguez-Hidalgo (2019), Quaternary International, https://doi.org/10.1016/j.quaint.2019.11.031. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier Ltd.

Going beyond the potential equifinality problems: a response to Saladié and Rodríguez-Hidalgo (2019) Jordi Rosell1,2,*, Ruth Blasco3, Maite Arilla1,2, Yolanda Fernández-Jalvo4

1

Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya, 35, 43002 Tarragona, Spain. E-mail: [email protected] (J.R.); [email protected] (M.A.) 2 IPHES; InstitutCatalà de Paleoecologia Humana i Evolució Social, Zona educacional 4, Campus Sescelades URV (Edifici W3), 43007 Tarragona, Spain. 3 Centro Nacional de Investigación sobre la Evolución Humana (CENIEH), Paseo Sierra de Atapuerca 3, 09002 Burgos, Spain. Email: [email protected]; [email protected] 4 Museo Nacional de Ciencias Naturales (CSIC), José Gutiérrez Abascal, 2, 28906, Madrid, Spain. Email: [email protected] *Corresponding author: [email protected] Abstract Actualistic studies have been commonly used as valid analogies in taphonomic research and, as the growing body of data demonstrate, have proved to be highly informative to explain the formation of terrestrial vertebrate fossil faunas. In Rosell et al. (2019), we conducted an experimental study with free-ranging brown bears (Ursus arctos arctos) with the aim of modeling their behavior and characterizing the bone damage caused on large, medium and small-sized ungulate carcasses. The purpose of the study was to highlight the equifinality processes observed experimentally based on the assumption that “some carnivores show physical and dental characteristics that could lead to bone modifications potentially like those generated by humans” (Rosell et al. 2019, p.67). In the case of bears, their “bunodont dentition and plantigrade locomotion –the latter allows them to frequently release and use their claws as ‘hands’”– have led to the production of peeling and tooth marks that show important similarities with those generated during the feeding activities of humans and chimpanzees (Pan troglodytes), although anecdotally also made by other taphonomic agents. Saladié and Rodríguez-Hidalgo (2019) interpret our study as an attempt to invalidate their inferences about human tooth marks from the TD6.2 level of Gran Dolina (Sierra de Atapuerca, Burgos, Spain), even though we do not make any archeological application. We also clearly maintain that ours is an initial and merely descriptive study that aims to raise awareness of the existence of taphonomic equifinality phenomena between bears and humans. The present work intends, therefore, to respond to their criticisms about the contexts in which humans and bears produce peeling as well as about the methodology used for assessing the tooth mark measurements. We have tried to read positively the Saladié and Rodríguez-Hidalgo’s (2019) paper in order to make progress in the main challenge of finding elements and features that allow us to discriminate bone alterations potentially attributable to more than one taphonomic agent. Keywords: Equifinality; Neo-taphonomy; Bears; Humans; Peeling; Tooth marks

1. Introduction Actualistic studies rely on direct observations in the modern world in order to document the chain of causal relationships between taphonomic actors (in this case, bears), their actions (feeding), effectors (teeth), and effects (damage to bones) (see, e.g., Gifford-Gonzalez, 1991). The results of actualistic studies (of known origin) can then be productively applied to interpret paleontological or archaeological faunas of unknown origins. However, researchers sometimes face damage that can potentially be attributed to more than one taphonomic agent (e.g., Lyman, 1985, 1987, 2004; Gifford-Gonzalez, 1991). This is the case for our study in Rosell et al. (2019), in which we documented equifinality phenomena with the appearance of peeling in actualist bone assemblages modified by Ursus arctos arctos and recorded striking similarities in some bear tooth marks resembling those experimentally attributed to humans or chimpanzees (Pan troglodytes) (e.g., Brain, 1981; Pobiner et al., 2007; Fernández-Jalvo and Andrews, 2011, 2016; Saladié et al., 2013a, 2014). Among axial bones, the main alterations we identified were transversal fractures, crushing, and different types of tooth marks (pits, punctures and scores), as well as crenulated edges (mainly on vertebrae). Classic, general, and incipient peeling (bent ends, fraying) were also documented on different portions of the ribs and on the apophyses of the vertebrae, which also showed occasional furrowing on the bodies. An element to highlight is the frequent co-occurrence of these alterations, mainly tooth marks with transverse fractures and peeling. The analysis of the images derived from the photo- and video-trap monitoring also allowed us to relate evidence-based activity and resulting bone damage on the carcasses. Most peeling alterations result from the movements of the bear around the carcass during viscera removal. In the course of this process, bears put pressure with their anterior paws on the ribs and expand the rib cage from the sternum with the claws and later with the help of the teeth. Once this process concludes, bears also tend to show interest in the marrow located inside the spine. In such cases, bears hold the spine with one of the front paws and use the mouth as a “hand” to bend the spine with strong head movements (see detailed descriptions in Arilla et al., 2014; Rosell et al., 2019). Regardless of the phase or stage of consumption, we observed specific characteristics in the tooth marks that closely resembled the human ones. For instance, some scores showed flaked surfaces, both on the shoulder and on the bottom (16.7%). All of them were located on axial bones, and three of them showed internal microstriation (4.2%) such as those described by Saladié and colleagues (2013a) for human scores. Among the pits, crescent-shaped morphologies were also documented on limb and axial bones as well as punctures with incomplete contours on axial bones (see Figures 1 and 2 from Rosell et al., 2019). Double-arch morphologies were also observed (3%), although only one of them was located on a crenulated edge, as described by Fernández-Jalvo and Andrews (2011) (see Figure 3 from Rosell et al., 2019). All these characteristics were identified as a result of the descriptive analysis of the tooth marks, which were produced not only in a stage of initial consumption but also when the carcass was practically depleted in external resources and certain skeletal regions were already disarticulated. As a reaction to the documentation of these alterations in non-anthropogenic contexts, Saladié and Rodríguez-Hidalgo (2019) elaborate an article in which they try to justify their assignment

of human tooth marks in TD6.2 of the Gran Dolina, Sierra de Atapuerca, Burgos, Spain (Saladié et al., 2014), expressing doubts about the similarities and context as well as the methodology used in our study. Although Saladié and Rodríguez-Hidalgo (2019, p.) write, “Rosell et al. (2019) proposed possible problems in our assignation of producers of tooth marks, especially those of the early Pleistocene assemblage of TD6.2 of the Gran Dolina […]”, the point is that we never dealt with any archaeological application or a critical review of this assemblage. In fact, two of us (JR and YFJ) proposed human cannibalism in TD6.2 more than a decade ago (Fernández-Jalvo et al., 1996; 1999; Díez et al., 1999) as well as to subsequently suggest the presence of human tooth marks in the assemblage (Fernández-Jalvo and Andrews, 2011). In fact, the only mention of TD6.2 throughout our text is in Figure 3, in which the image of a tooth mark interpreted as human by Fernández-Jalvo and Andrews (2011) is used as a model to illustrate such similarities. Either way, we will try to address here all the points that are highlighted as problematic by Saladié and Rodríguez-Hidalgo (2019) in order to contribute to the evolving research program on potential equifinality problems, which are vital for understanding the formation of terrestrial vertebrate fossil faunas suspected to have been formed and/or modified by bears and/or humans. 2. Human tooth marks and peeling Bending fractures and co-related modifications as a result of a single activity or several overlapping actions on the same bone have attracted attention from the scientific community since White (1992) first described the so-called peeling. Saladié and Rodríguez-Hidalgo (2019) dedicate a section to this damage and its combination with other alterations, offering a good review of the definitions and their variants, which we would like to clarify based on the proposals made by Pickering et al. (2013). As mentioned, classic peeling was first described by White (1992, p.140) "when freshbone is fractured and peeled apart similar to bending a small fresh twig from a tree branch between two hands’’. More detail description has been published by Pickering et al. (2013, p.1299): “with classic peeling, layer(s) of lamella(e) is/are missing in strip(s) from the rib’s dorsal, ventral or both cortices […] Classic peeling can occur at various points along a rib’s length but is commonly observed at the sternal terminations of short fragments of the vertebral ends of ribs”. “With general peeling, an area of the whole dorsal or ventral cortex of a rib is peeled backed for some length, revealing the internal trabeculae of the rib” (Pickering et al., 2013, p.1299). Incipient peeling is observed on the “bent rib ends” chewed by humans and described by Fernández-Jalvo and Andrews (2011, p.121) as result of bending the ends of these thin skeletal elements "pushing up or down on the ends of the bone, using the hands and holding the ends between the upper and lower cheek teeth". This has also been observed on bones chewed by chimpanzees (Pickering and Wallis, 1997, Plummer and Standford, 2000), and Pobiner et al. (2007) and the latest named it as fraying. It is “a type of peeling where a strip(s) of lamaella(e) is/are only partially peeled back against the rib shaft, not fully removed from the specimen” (Pickering et al., 2013, p.1300). Pickering et al. (2013) demonstrated that, unlike

classic and general peeling, incipient peeling is more a consequence of the intrinsic physical properties of an affected rib than of any taphonomic force(s) applied to it. For this reason, we give greater weight to classic and general peeling, although we also find it convenient to record that incipient peeling also occurs extensively in our study. This same type, together with classic and general peeling, also appears combined with other alterations (n=39; 29.32%), such as crenulated edges on the distal shaft of ribs (n=2); transversal fractures on mid-shafts, distal ends and, to a lesser extent, necks of ribs (n=7); transversal fractures on the spinous process of thoracic vertebrae (n=1) and on the spinous, transverse and mammillary processes of lumbar vertebrae (n=12); crushing on the head and distal shafts of ribs (n=4) and on the spinous processes of lumbar vertebrae (n=2); longitudinal cracks on distal shafts of ribs (n=3); and furrowing on the body of one thoracic vertebra. Saladié and Rodríguez-Hidalgo (2019) point out that the association between peeling, gnawing marks, and cut marks for the attribution of tooth marks to an anthropogenic origin is a valid option as long as the characteristics of the archaeological site are taken into account. We agree that no direct relationship can be made between effector (e.g., teeth) and actor (e.g., human) in these contexts given precisely the problem of equifinality that we openly manifest in our study. Therefore, we strongly emphasize the need to characterize in detail the damages and their combinations following the lines of argumentation in other works (e.g., Lyman, 1987; Gifford-González, 1991) and showing special concern for the genesis of those assemblages in which bears and humans could have played a part. In fact, peeling as an individual damage is difficult to ascribe to one agent or another in the current state of the investigation, since the alteration is morphologically similar and is also recorded in the same anatomical areas. Making clear the necessity of determining the general characteristics and combination of modifications, we must bear in mind that jumping to the “archeological domain” entails greater complexity. Archaeological assemblages are often the result of numerous depositional events and overlapping occupations produced by different actors, which are normally accompanied by disruptive processes that can alter the preexisting sets by biasing them and/or adding different nature remains (e.g., Bailey, 2007; Bailey and Galadinou, 2009; Brochier, 1999; Malinsky-Buller et al., 2011; Vaquero, 2008; Vaquero et al., 2012). The results are palimpsests with a disordered appearance that can confuse the archaeological interpretations, not only in terms of the sets’ spatial organization but in terms of the taxonomic and anatomical profile and/or the bone surface modifications themselves –see, for example, Behrensmeyer et al. (1986), who showed the strong similarities between cut marks and trampling marks and is still the subject of debate among specialists (e.g., Olsen and Shipman, 1988; Domínguez-Rodrigo et al., 2009; Pineda et al., 2014; Fernández-Jalvo and Andrews, 2016). This cumulative character has important consequences for the interpretation of archaeological assemblages, especially “when attempting to apply ethnographically derived middle-range theories constructed in high resolution temporal contexts” (Vaquero et al., 2012, p.2785). Therefore, the taphonomic analysis of an archaeological site constitutes an interpretative challenge in which elements other than the general dynamic can explain an individual alteration. 3. Human tooth marks in TD6.2 level

Saladié and Rodríguez-Hidalgo (2019, p.) mention that “the context in which [they] found TD6.2 assemblage was decisive in the evaluation of the tooth marks documented”. TD6 is a ~3m thick lithostratigraphic unit situated in the middle of the 25m sedimentary infill of the Gran Dolina site and divided into three sub-units (TD6.3, TD6.2 and TD6.1, from bottom to top). TD6.2 is a subunit that presents great stratigraphic complexity, displaying up to five different strata (Bermúdez de Castro et al., 2008; Campaña et al., 2016). As a result of this complexity, it is assumed that different processes and agents could have been involved with higher or lower incidence in their formation. Proof of this is that although there is a general dynamic common to all strata based on the presence of Homo antecessor remains (e.g., Carbonell et al., 2010; Mosquera et al., 2018), the action of other actors on the bones has also been identified as it has on the skeletal remains of different carnivores (Crocuta crocuta, Ursus dolinensis, Canis mosbachensis, Vulpes praeglaciaris, Lynx sp.; Saladié et al., 2011, 2012, 2014) and coprolites in TD6.2.0 (n=6), TD6.2.2/3 (n=11) y TD6.4 (n=16) strata. Saladié et al. (2014) showed that 8.9% of mid-shafts exhibited carnivore tooth marks when they analyzed TD6.2 as a whole. As the human presence runs through all TD6.2 strata, the identification of peeling (30.2%) and human tooth marks (2.6% of the total) is not strange in the assemblage and completes, in some way, the rest of the taphonomic butchery signals, such as cut marks or diagnostic elements of intentional fracturing to access the bone marrow (e.g., Fernández-Jalvo et al., 1996; 1999; Díez et al., 1999; Carbonell et al., 2010; Fernández-Jalvo and Andrews, 2011; Saladié et al., 2011). We do not question the validity of the experimental data collected in Saladié et al. (2013a, 2014). In fact, the experimental actualistic (EA) approaches allow undoubtedly relating effectors (teeth) and effects (damage to bones). Consequently, their experimental data of human tooth marks are used as valid to compare them with those produced by bears in our naturalistic actualistic (NA) approaches in an attempt to establish diagnostic elements of discrimination (Arilla et al., 2014; Rosell et al., 2019). However, Saladié and Rodríguez-Hidalgo (2019) rule out our neotaphonomic data (especially the classic peeling) as valid for TD6.2 because they consider the context to be different. We agree that TD6.2 is a cave context, which shows significant human activity, and the interpretation of cannibalism and transport of ungulate carcasses together with Homo antecessor support a main anthropic origin (e.g., Fernández-Jalvo et al., 1996; 1999; Díez et al., 1999; Carbonell et al., 2010; Fernández-Jalvo and Andrews, 2011; Saladié et al., 2011). Nevertheless, as the researchers themselves point out, the activity of carnivores is also present during the formation of the assemblage; a phenomenon that is, on the other hand, widely recorded in many Pleistocene sites with similar characteristics (e.g., Binford, 1981, 1988; Stiner, 1994; Selvaggio and Wilder 2001; Faith et al. 2007; Domínguez-Rodrigo et al. 2010). According to Saladié et al. (2014), the role of these predators (including bears) would be mainly considered as secondary accesses to the remains abandoned by hominins based on the rate of carnivores’ activity. Conversely, our data come from complete, fresh carcasses of large-, medium- and small-sized ungulates as primary access and, therefore, a context opposite to that interpreted in TD6.2. So, we agree that the context is different. In fact, at no time were we thinking of recreating a hypothetical scenario similar to that of TD6.2, but despite this, the authors insist on linking our experimental work with their studied archaeological assemblage (Saladié and Rodríguez-Hidalgo, 2019, p.): “Rosell et al. (2019) suggested a series of possible

errors that could be incurred by not considering bears as scavengers in the cave and as the potential causes of these damages”. This leads them to invalidate our experimental observations recalling the validity principles of scientific models based on realism, precision, and generality (Levins, 1968) to interpret the assemblage. Indeed, the scenario that we tried to reproduce has nothing to do with the scavenging of bears in anthropogenic assemblages and cannibalistic practices, but it must be remembered that this was not the objective of the work. Our goal, we emphasize again, was simply to show descriptively the similarities between specific taphonomic modifications made by bears and humans. At this point we wonder if the authors have made a bad reading of our article. In any case, our long-term purpose is to try to reproduce all possible scenarios (including the scavenging of bears on assemblages previously modified by humans). Actually, we are currently working on this research line, monitoring carnivorous activities on remains left by humans through modeling and idealized reproductions (Arilla et al., submitted). Despite this, we would like to express a careful thought. Saladié et al. (2011, 2012, 2013a, 2014) propose that Homo antecessor carcasses are probably introduced whole to the cave, and therefore the nutritional exploitation is done in an integral and intensive way in the TD6.2 habitat place. In our NA approaches, we also offer complete and fresh carcasses to bears in a first stage of the experimental development of each observation (OB). Disregarding the cultural component and the anatomical and physiological distances between small/mediumsized ungulates (e.g., <300 kg cervids) and Homo antecessor (previously compared in Saladié et al. [2012]), we would have, on the one hand, an intact carcass scenario exposed to humans and, on the other, an intact carcass scenario exposed to bears. The initial stage in both scenarios could be considered similar and, therefore, the alterations inflicted by both agents – those, of course, that do not involve the use of tools– could be studied from the point of view of equifinality (i.e., different agents producing a similar modification on whole carcasses). We would also like to clarify that the experimental exposure time took more than a monthand-a-half in some observations (e.g., OB1). During this process the carcass underwent several phases or stages. The latest stage implies the complete absence of soft tissues, an evident limb disarticulation, and axial skeleton fragmentation. As mentioned in Arilla et al. (2014) and Rosell et al (2019), most of the peeling comes from the early stages (viscera removal and extraction of marrow located inside the spine) but not exclusively. Peeling is also identified in the final stages of consumption when bears carry on with their movements around the carcass using both their claws and their teeth in an attempt to continue taking advantage of nutrients. An incomplete carcass and the existence of fragmented skeletal elements together with the lack of soft tissues would perhaps be the scenario most similar to scavenging activities or late access. In any case, the similarity of peeling in vertebrae and ribs, as well as some tooth marks, is in our opinion relevant (see, for example, Figures 1-2 and 5 in Rosell et al., 2019), and following the objective of our study, we think it is worth a thorough investigation. 4. Human tooth marks in other archaeological contexts

Human tooth marks have been identified in several archaeological sites, linked not only to cannibalism but to the routine processes of human consumption (e.g., Cáceres et al. 2007; Bello et al., 2015; Fernández-Jalvo and Andrews, 2016). As we have mentioned repeatedly in the text, we have never questioned these results; all the contrary, we used them as valid and reference evidence in an attempt to establish diagnostic criteria of discrimination with those derived from NA assemblages modified by bears. The criteria proposed by Saladié et al. (2013a) have, therefore, been used by many of us as a valid complement to previous studies that pointed in the same direction (e.g., Landt, 2004, 2007; Fernández-Jalvo and Andrews, 2011). In this sense, we do not understand why Saladié and Rodríguez-Hidalgo (2019) are surprised when some of us have used them in other works, especially those focused on small prey, including leporids, quelonids, and birds (e.g., Blasco, 2008; Blasco and Fernández Peris, 2009; Blasco et al., 2013, 2014, 2016; Blasco et al., 2019). The small prey size favors the use of hands and teeth for consumption and, therefore, the need for processing tools is smaller. Thus, the probability of observing human tooth marks on these animals is proportionally higher than observing them in animals of larger dimensions (e.g., Landt, 2007). In addition, there is more extensive and increasing literature about tooth marks on small prey than on larger carcasses (e.g., Laroulandie, 2000, 2001, 2005; Cochard, 2004; Pérez Ripoll, 2005; Landt, 2004, 2007; Laroulandie et al., 2008; Martínez, 2009; Sanchis Serra, 2010; Sanchis Serra and Fernández Peris, 2008; Fernández-Jalvo and Andrews, 2011, 2016; Romero et al., 2016; Morin et al., 2019; Pelletier et al., 2019). All these studies agree on the combination of alterations together with the identification of patterns to establish an appropriate assignment in direct human consumption. Some of these works were already described in detail in Rosell et al. (2019) with the objective of collecting those taphonomic referents that would be used in our study. The only issue that was not addressed is the presence of diaphyseal cylinders (or tubes) in small prey bones, because our bear-induced assemblages did not include this type of damage or this category of carcass. Despite this, Saladié and Rodríguez-Hidalgo (2019) mention bone fragmentation traits (including the presence of distal ends of long bones and crenulated edges) and their possible equifinality problem. We are convinced that many of the efforts should also be aimed at solving these phenomena in the case of small game given that fragmentation is often present in assemblages produced by cats (with values <10% for long bone elements; e.g., Andrews and Evans, 1983; Lloveras et al., 2008; Marin-Monfort et al., 2019) and can also be produced by other pre- and post-burial processes (e.g., sediment pressure, trampling; Cochard, 2004). Many researchers have been very careful when establishing the attribution of these modifications to an anthropogenic origin (e.g., Cochard et al., 2012; Blasco et al., 2013; Martínez-Valle et al., 2016; Morín et al., 2019; Pelletier et al., 2019). Most authors use a combination of damages, such as 1) the percentage of whole long bones; 2) break type pattern by skeletal element (following Lloveras et al., 2008); 3) the percentage of shaft cylinders and their maximum length, assuming that shaft cylinders are usually short in natural deposits (e.g., Hockett, 1993; Brugal, 2006); 4) the abundance of dry vs. green-bone fractures (distinguishing natural, carnivore, and human origin; see, for example, Figure 7 in Cochard et al., 2012; see also an application to small mammals in Armstrong, 2016a, 2016b); and 5) the presence of patterns on the olecranon region of the ulna in combination with a breakage near the radial

fossa of the humerus (as a result of the overextension of the elbow during disarticulation (e.g., Laroulandie et al., 2008; Cochard et al., 2012; Blasco et al., 2016). The results are also combined with other anthropic-inflicted damage such as cut marks to establish whether humans have been the main agent acting on the small prey in the assemblage. In the specific case of the lynxes, we must mention that the authors themselves register a very scarce fragmentation, with 94% of the rabbit specimens complete (Rodríguez-Hidalgo et al., 2015), something that would already be far from the archaeological sites in which an anthropogenic origin has been determined. In addition, the non-ingested accumulations published by Rodríguez-Hidalgo et al. (2016) for red-legged partridge remains involves the Iberian lynx under captive breeding programs, and this circumstance should be considered when assessing the results. This is also applicable to Rodríguez-Hidalgo et al. (2013) where the lynxes are affected by a chronic disease in a population. Several studies have previously demonstrated that animals in captivity alter bones differently and generate a higher intensity of damage than wild animals (e.g., Gidna et al., 2013; Sala and Arsuaga, 2013; Arilla et al., 2014). With this in mind, we agree with the authors that more analysis should be done to resolve any doubt about the equifinality problems regarding damage inflicted on small prey in future research. 5. Human and bear tooth marks: metric problems In relation to the metric problems of the tooth marks, Saladié and Rodríguez-Hidalgo (2019, p.) state “[...] they [we] cannot compare their metric results with those of Saladié et al. (2013a), since they [we] do not consider the stratification of bone tissues in the same way [...]”. The classification of the bones carried out by Selvaggio (1994), and later adopted by other authors (e.g., Domínguez-Rodrigo and Piqueras, 2003; Delaney-Rivera et al., 2009; Andrés et al., 2012; Saladié et al., 2013a), focuses mainly on long bones and is based on criteria of bone thickness and cortical density in each portion of the bone: thick cortical or mid-shaft, thin cortical or metaphysis, and cancellous tissue or epiphyses. This division is due to the fact that each of these surfaces absorbs the bites differently, which generates significant differences in the measurements of the generated tooth marks. Subsequently, Andrés et al. (2012, p.211) propose to suppress the thin cortical category (metaphysis) because “[...] their thickness varied according to element and these sections do not systematically sample bone thickness intermediate between end and midshafts”. Saladié and Rodríguez-Hidalgo (2019) seem to concur with this proposal. However, we consider it a mistake not taking into account the thin cortical, since most of the modifications of our NA approaches come from flat bones of the axial skeleton (vertebrae and ribs), followed by girdles (see, for example, Figure 4 in Arilla et al., 2014). These types of bones are not described by the preceding authors in terms of cortical type. In our opinion, and using the same criteria described by Selvaggio (1994), these are bones with lower thicknesses and densities than the mid-shaft diaphysis of the long bones. Therefore, most portions of these bones should be grouped within the thin cortical, except the rib head, glenoid of the scapula, and vertebral bodies, which should be considered cancellous tissue. At this point, we do not understand why Saladié and Rodríguez-Hidalgo (2019) say we do not consider the stratification of the bones, as data of bear tooth marks both in this work and in Arilla et al. (2014) are presented following this classification (see methods section and boxplots of Figure 4 and Table 4 in Rosell et al., 2019).

It should also be mentioned that not all authors who perform experimental reproductions with carnivores do it in the same way. Apart from the differences in the environmental condition (captivity vs. freedom) already mentioned above, there are works in which they also select specific skeletal elements previously disarticulated and defleshed prior to being subjected to carnivorous activity (e.g., Saladié et al. 2013b; Delaney-Rivera et al., 2009). This should also be taken into account when making comparisons, since we do not know to what degree the amount of meat on the bones could affect tooth mark dimensions. Saladié and Rodríguez-Hidalgo (2019, p.) also criticize how the metric parameters of tooth marks were used in our study, suggesting that we obviate the most recent methodological proposals for a more precise evaluation of the results. We do not know exactly what they are referring to. We assume that they are perhaps referring to the micro-photogrammetry and/or geometric morphometric technique as analysis method (e.g., Arriaza et al., 2017, 2018, 2019; Aramendi et al., 2017; Maté-González et al., 2019; Yravedra et al., 2017c). The application of these analysis systems for the reconstruction of tooth mark morphology has recently provided satisfactory results in differentiating among carnivore taxa with similar body size (e.g., Arriaza et al., 2019). In Rosell et al. (2019), we made a first descriptive approximation of the damages using a light stereomicroscope with a magnification of up to 120x, a KH-8700 3D digital microscope, and an analytical FEI QUANTA 600 environmental scanning electron microscope (ESEM) operated in low vacuum mode with a magnification of up to 300x. The images used were taken by means of these devices in order to compare them with those previously published in other works on human tooth marks in which these systems were also used (Fernández-Jalvo et al., 2011; Saladié et al., 2013a; Saladié et al., 2014). The authors themselves use these same systems to present their results in all the articles cited. However, we have no doubt that the use of computer vision systems will significantly improve our ability to document and present highly precise and accurate digital models of the damage. We agree with Saladié and Rodríguez-Hidalgo (2019) that it is important to consider the statistical results of the populations studied and not only the ranges of measures. Figure 4 in Rosell et al. (2019) shows the data of the tooth marks in different boxplots (n˃33) according to the tissue type in which they are found (thin cortical and cancellous tissue). The work of Arilla et al. (2014) also provides important statistical deployment in this regard. However, it is surprising that, following this comment, Saladié and Rodríguez-Hidalgo (2019) attach two figures using only the maximum and minimum ranges of the tooth marks with the mean (see Figures 1-2 in Saladié and Rodríguez-Hidalgo, 2019) and without taking into account the outliers. In a graphic such as the one presented by them, outliers should always be considered, since they always increase the maximum and minimum ranges. Otherwise, they should specify that outliers in the other works have also been discarded or included. In any case, Table 4 and Figure 4 of our work show the superposition of the means between bears and humans, which was surprising even to us. Data obtained on thin cortical are even more evident. Following this line, they say that the tooth marks sample recovered by Arilla et al. (2014) on cancellous tissue is very small and cannot be used in comparisons. Table S1 of that article shows the quantitative data, where the number of pits on cancellous tissue amounts to 49. It is

a sufficient number to be considered, especially taking into account that it is a study with wild animals (principle of realism). Obviously, this number is lower than those obtained by Saladié et al. (2013a, b) for both bears and humans, not to mention that the former are carried out in contexts of captivity with the alteration in intensity and frequency that these conditions entail and the latter correspond to experiments in which individuals are induced to produce marks. Therefore, we consider that their arguments are not sufficient to invalidate our data with wild animals, and phrases such as “[…] the only available data are those of Saladié et al. (2013b)” (Saladié and Rodríguez-Hidalgo, 2019, p.) seem to us excessively categorical and even pretentious. It is possible that subjectivity in the perception of the modifications’ characteristics may influence the methods of analysis, especially when the origin is unknown (Domínguez-Rodrigo et al., 2017). However, we have tried to overcome this handicap with a descriptive analysis by offering images taken not only by us but by other researchers for comparison (see Figures 1-3 in Rosell et al., 2019) as well as by presenting peeling images, both classic and incipient, so that the reader can see the diagnostic characteristics (see Figure 5 in Rosell et al., 2019). Obviously, applications to archaeological contexts entail complexity and always carry a risk of interpretation, as already mentioned above. An interpretative generalization of human activities in an assemblage cannot lead to an assumption that a specific incidence of bone damage (or its co-occurrence with other alterations on the same bone) is produced solely and exclusively by the same collecting agent. Besides the necessity to develop better statistical methods and new analysis techniques, only the realization of appropriate experimental protocols in NA approaches and common methodologies will enable the observation of bone damage and its accurate comparison. We consider our work to be a starting point in this line. 6. Conclusions It is evident that our study and the comments made by Saladié and Rodríguez-Hidalgo (2019) show the need to continue working on the characterization of predators that intervene in faunal assemblages based on the analysis of their taphonomic signals and, especially, tooth marks or damage related to bone flexion. These works are essential to identify the different agents and sequence them in time, which is highly informative for understanding the processes of origin and formation of archaeological sites. Such works become more relevant when they refer to human tooth marks since their identification entails important connotations that can influence subsequent interpretations of the socio-economic behavior of human groups. Therefore, the identification of these modifications must always be doubly accurate and, from this point of view, any new technique/method and discussion that help to move in this direction are always welcome. We think that with this work we clarify the doubts set out by Saladié and Rodríguez-Hidalgo (2019) about the similarities, context, and methodology used in our study, always making a positive reading of the comments. The existence of these controversies encourages the setting up and discussion of common and uniform action protocols in order to truly make progress in clarifying comparisons and discrimination of elements in the specific case of equifinality problems. Acknowledgements

This study is part of the Spanish MINECO/FEDER projects CGL2015-65387-C3-1-P (JR), CGL2016-80000-P (JR) and CGL2015-68604-P (RB), the Generalitat de Catalunya-AGAUR projects 2017 SGR 836 and CLT009/18/00055 (JR, RB, MA). MA is the beneficiary of a research fellowship (FI) from AGAUR (2017FI-B-00096). Special thanks to Thijs van Kolfschoten for allowing us to reply and participate in this debate within the journal. Referencias Andrés, M., Yravedra, J., Domínguez-Rodrigo, M., 2012. A study of dimensional differences of tooth marks (pits and scores) on bones modified by small and large carnivores. Archaeological and Anthropological Sciences 4, 209-219. Andrews, P., Evans, E. M. N., 1983. Small mammals bone accumulations produced by mammalian carnivores. Paleobiology 9, 289-307. Aramendi, J., Maté-González, D., Yravedra, J., Ortega, M. C., Arriaza, M. C., González-Aguilera, D., Baquedano, E., Domínguez Rodrigo, M., 2017. Discerning carnivore agency through the three-dimensional study of tooth pits: revisiting crocodile feeding behaviour at FLK- Zinj and FLK NN3 (Olduvai Gorge, Tanzania). Palaeogeography, Palaeoclimatology, Palaeoecology 488, 93-102. Arilla, M., Rosell, J., Blasco, R., Domínguez-Rodrigo, M., Pickering, T. R., 2014. The ‘‘bear’’ essentials: actualistic research on Ursus arctos arctos in the Spanish Pyrenees and its implications for Paleontology and Archaeology. Plos One 9 (7), e102457. Arilla, M., Rosell, J., Blasco, R. (submitted). The campsites’ looters: an experimental (neotaphonomic) approach with free-ranging wild carnivores to biogenic destruction/”production” of hominin campsites. Scientific Reports. Armstrong, A., 2016a. Eagles, owls, and coyotes (oh my!): Taphonomic analysis of rabbits and guinea pigs fed to captive raptors and coyotes. Journal of Archaeological Science: Reports 5, 135-155. Armstrong, A., 2016b. Small mammal utilization by Middle Stone Age humans at Die Kelders Cave 1 and Pinnacle Point Site 5-6, Western Cape Province, South Africa. Journal of Human Evolution 101, 14-44. Arriaza, M., Organista, E., Yravedra, J., Santonja, M., Baquedano, E., Domínguez-Rodrigo, M., 2018. Striped hyenas as bone modifiers in dual human-to-carnivore experimental models. Archaeological and Anthropological Sciences https://doi.org/10.1007/s12520-018-0747-y. Arriaza, M. C., Aramendi, J., Maté-González, D., Yravedra, J., Stratford, D., 2019. Characterising leopard as taphonomic agent through the use of micro-photogrammetric reconstruction of tooth marks and pit to score ratio. Historical Biology https://doi.org/10.1080/08912963.2019.1598401. Arriaza, M. C., Yravedra, J., Domínguez-Rodrigo, M., Mate-González, M. A., García Vargas, E., Palomeque-González, J. F., Aramendi, J., González-Aguilera, D., Baquedano, E., 2017. On applications of micro-photogrammetry and geometric morphometrics to studies of tooth mark morphology: the modern Olduvai Carnivore Site (Tanzania). Palaeogeography, Palaeoclimatology, Palaeoecology 488, 103-112.

Bailey, G., 2007. Time perspectives, palimpsests and the archaeology of time. Journal of Anthropological Archaeology 26, 198-223. Bailey, G., Galadinou, N., 2009. Caves, palimpsests and dwelling spaces: examples from the Upper Palaeolithic of south-east Europe. World Archaeology 41 (2), 215-241. Binford, L. R., 1981. Bones: Ancient Men and Modern Myths. New York, Academic Press. Binford, L. R., 1988. Etudetaphonomique des restes fauniques de la GrotteVaufrey, Couche VIII. Rigaud, J. P. (Ed.). La GrotteVaufrey: Paléoenvironnement, chronologie, activitéshumaines, Mémoires de la Société PréhistoriqueFrançaise. 19, 535-564. Blasco, R., 2008. Human consumption of tortoises at Level IV of Bolomor Cave (Valencia, Spain). Journal of Archaeological Science 35, 2839-2848. Blasco, R., Fernández Peris, J., 2009. Middle Pleistocene bird consumption at Level XI of Bolomor Cave (Valencia, Spain). Journal of Archaeological Science 36, 2213-2223. Blasco, R., Finlayson, C., Rosell, J., Sánchez Marco, A., Finlayson, S., Finlayson, G., Negro, J. J., Giles Pacheco, F., Rodríguez-Vidal, J., 2014. The earliest pigeon fanciers. Scientific Reports 4, 5971. Blasco, R., Rosell, J., Fernández Peris, J., Arsuaga, J. L., Bermúdez de Castro, J. M., Carbonell, E., 2013. Environmental availability, behavioural diversity and diet: a zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain). Quaternary Science Reviews 70, 124-144. Blasco, R., Rosell, J., Rufà, A., Sánchez Marco, A., Finlayson, C., 2016. Pigeons and choughs, a usual resource for the Neanderthals in Gibraltar. Quaternary International 421, 62-77. Blasco, R., Rosell, J., Sánchez-Marco, A., Gopher, A., Barkai, R., 2019. Feathers and food: Human-bird interactions at Middle Pleistocene Qesem Cave, Israel. Journalof Human Evolution 136, 102653. Brain, C. K., 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago, Universityof Chicago Press. Behrensmeyer, A. K., Gordon, K. D., Yanagi, G. T., 1986. Trampling as a cause of bone surface damage and pseudocutmarks. Nature 319, 768-771. Bello, S. M., Saladié, P., Cáceres, I., Rodríguez-Hidalgo, A., Parfitt, S. A., 2015. Upper Palaeolithic ritualistic cannibalism at Gough's Cave (Somerset, UK): The human remains from head to toe. Journalof Human Evolution 82, 170-189. Bermúdez de Castro, J. M., Pérez-González, A., Martinón-Torres, M., Gómez-Robles, A., Rosell, J., Prado, L., Sarmiento, S., Carbonell, E., 2008. A new early Pleistocene hominin mandible from Atapuerca-TD6, Spain. Journal of Human Evolution 55, 329-335. Brochier, J. E., 1999. Couche archéologique, sol archéologique et distributions spatiales: quelques réflexions (géo)archéologiques sur un vieux problème. Geoarqueologia i Quaternari litoral. Memorial M.P. Fumanal. València, Universitat de València, 91-95.

Brugal J.Ph. 2006. Petit gibier et fonction de sites au Paléolithique supérieur. Paleo 18, 45-68. Cáceres, I., Lozano, M., Saladié, P., 2007. Evidence for Bronze Age Cannibalism in El Mirador Cave (Sierra de Atapuerca, Burgos, Spain). American Journal of Physical Anthropology 133, 119. Campaña, I., Pérez-González, A., Benito-Calvo, A., Rosell, J., Blasco, R., Bermúdez de Castro, J. M., Carbonell, E., Arsuaga, J. L., 2016. New interpretation of the Gran Dolina-TD6 bearing Homo antecessor deposits through sedimentological analysis. ScientificReports DOI: 10.1038/srep34799. Carbonell, E., Cáceres, I., Lozano, M., Saladié, P., Rosell, J., Lorenzo, C., Vallverdú, J., Huguet, R., Canals, A., Bermúdez de Castro, J. M., 2010. Cultural cannibalism as a paleoeconomic system in the European Lower Pleistocene. Current Anthropology 51 (4), 539-549. Cochard, D., 2004. Étude taphonomique des léporidés d’unetanière de renard actuelle: apport d’un référentiel à la reconnaisance des accumulations anthropiques. Revue de Paléobiologie 23 (2), 659-673. Cochard, D., Brugal, J. P., Morin, E., Meignen, L., 2012. Evidence of small fast game exploitation in the Middle Paleolithic of Les Canalettes, Aveyron, France. Quaternary International 264, 3251. Delaney-Rivera, C., Plummer, T. W., Hodgson, J. A., Forrest, F., Hertel, F., Oliver, J. S., 2009. Pits and pitfalls: taxonomic variability and patterning in tooth mark dimensions. Journal of Archaeological Science 36, 2597-2608. Díez, J. C., Fernández-Jalvo, Y., Rosell, J., Cáceres, I., 1999. Zooarchaeology and taphonomy of Aurora Stratum (Gran Dolina, Sierra de Atapuerca, Spain). Journal of Human Evolution 37 (3/4), 623-652. Domínguez-Rodrigo, M., Piqueras, A., 2003. The use of tooth pits to identify carnivore taxa in tooth-marked archeofaunas and their relevance to reconstruct hominid carcass processing behaviours. Journal of Archaeological Science 30, 1385-1391. Domínguez-Rodrigo, M., Juana, S. d., Galán, A. B., Rodríguez, M., 2009. A new protocol to differentiate trampling marks from butchery cut marks. Journal of Archaeological Science 36, 2643–2654. Domínguez-Rodrigo, M., Bunn, H. T., Mabulla, A. Z. P., Ashley, G. M., Diez-Martin, F., Barboni, D., Prendergast, M. E., Yravedra, J., Barba, R., Sánchez, A., Baquedano, E., Pickering, T. R., 2010. New excavations at the FLK Zinjanthropus site and its surrounding landscape and their behavioral implications. Quaternary Research 4 (3), 315-332. Domínguez Rodrigo, M., Saladié, P., Cáceres, I., Huguet, R., Yravedra, J., Rodríguez -Hidalgo, A., Martín, P., Pineda, A., Marín, J., Gené, C., Aramendi, J., Cobo-Sánchez, L., 2017. Use and abuse of cut mark analyses: The Rorschach effect. Journal of Archaeological Science 86, 14-23. Faith, J. T., Marean, C. W., Behrensmeyer, A. K., 2007. Carnivore competition, bone destruction, and bone density. Journal of Archaeological Science 34, 2025-2034.

Fernández-Jalvo, Y., Díez, J. C., Bermúdez de Castro, J. M., Carbonell, E., Arsuaga, J. L., 1996. Evidence of early cannibalism. Science 271, 277-278. Fernández-Jalvo, Y., Díez, J. C., Cáceres, I., Rosell, J., 1999. Human cannibalism in the Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain). Journal of Human Evolution 37 (3/4), 591-622. Fernández-Jalvo, Y., Andrews, P., 2011. When humans chew bones. Journal of Human Evolution 60, 117-123. Fernández-Jalvo, Y., Andrews, P., 2016. Atlas of Taphonomic Identification. Dordrecht, Heidelberg, New York, London, Springer. Gidna, A., Yravedra, J., Domínguez-Rodrigo, M., 2013. A cautionary note on the use of captive carnivores to model wild predator behaviour: a comparison of bone modification patterns on long bones by captive and wild lions. Journal of Archaeological Science 40, 1903-1910. Gifford-Gonzalez, D., 1991. Bones are not enough: analogues, knowledge, and interpretative strategies in zooarchaeology. Journal of Anthropological Archaeology 10, 215-254. Hockett, B. S., 1993. Taphonomy of the leporid bones from Hogup Cave, Utah: Implications for cultural continuity in the Eastern Great Basin. .Reno, University of Reno. Landt, M. J., 2004. Investigations of human gnawing on small mammal bones: among contemporary Bofi foragers of the Central African Republic. Department of Anthropology, Washington State University. Ph. D. Disertation, 178. Landt, M. J., 2007. Tooth marks and human consumption: ethnoarchaeological mastication research among foragers of the Central African Republic. Journal of Archaeological Science 34, 1629-1640. Laroulandie, V., 2000. Taphonomie et archéozoologie des oiseaux en grotte: applications aux sites paléolithiques du Bois Ragot (Vienne), de Combe Saunière (Dordogne) et de La Vache (Ariège). Bordeaux, Université de Bordeaux-I. Laroulandie, V., 2001. Les traces liées à la boucherie, à la cuisson et à la consommation d’oiseaux: apport de l’expérimentation. Bourguignon, L., Ortega, I. and Frère-Sautot, M.-C. (Ed.). Préhistoire et approche expérimentale. Montagnac, Monique Mergoual, Collection préhistoire. 5, 97-108. Laroulandie, V., 2005. Bird explotaition pattern: the case of Ptarningan Lagopus sp. in the Upper Magdalenian site of la Vache (Ardiège, France). Grupe, G. and Peters, J. (Ed.). Feathers, Grid and Symbolism. Birds and Humans in the Ancient Old and New Worlds. Munich, Proceedings of the 5th. Meeting of the ICAZ. Bird Working Group, 165-178. Laroulandie, V., Costamagno, S., Cochard, D., Mallye, J.-B., Beauval, C., Castel, J. C., Ferriè, J. G., Gourichon, L., Rendu, W., 2008. Quand désarticulerais des traces: le cas de l'hyperextension du coude. Annales de Paléontologie 94, 287-302. Levins, R., 1968. Evolution in Changing Environments. Some Theoretical Explorations. Princenton 8 (Mpb-2), Princenton University Press.

Lloveras, L., Moreno-Garcia, M., Nadal, J., 2008. Taphonomic analysis of leporid remains obtained from modern Iberian lynx (Lynx pardinus) scats. Journal of Archaeological Science 35, 1-13 Lyman, R. L., 1985. Bone frecuencies: differential survivorship of fossil classes. Journal of Archaeological Science 12, 221-236. Lyman, R. L., 1987. Archaeofaunas and butchery studies: a taphonomic perspective. Schiffer, M. B. (Ed.). Advances in Archaeological Method and Theory. New York, Academic Press. 10, 249-337. Lyman, R. J., 2004. The concept of equifinality in Taphonomy. Journal of Taphonomy 2 (1), 1526. Malinsky-Buller, A., Hovers, E., Marder, O., 2011. Making time: ‘Living floors’, ‘palimpsests’ and site formation processes. A perspective from the open-air Lower Paleolithic site of Revadim Quarry, Israel. Journal of Anthropological Archaeology 30, 89-101. Marin-Monfort, M.D., García-Morato, S., Olucha, R. Yravedra, J. Piñeiro, A., Barja, I, Andrews, P., Fernández-Jalvo, Y. 2019. Wildcat scats: Taphonomy of the predator and its micromammal prey. Quaternary Science Reviews 225 https://doi.org/10.1016/j.quascirev.2019.106024 Martínez, G., 2009. Human chewing bone surface modification and processing of small and medium prey amongst the Nukak (forager of the Colombian Amazon). Journal of Taphonomy 7 (1), 1-20. Martínez Valle, R., Guillem, P. M., Villaverde, V., 2016. Bird consumption in the final stage of Cova Negra (Xàtiva, Valencia). Quaternary International 421, 85-102. Maté-González, D., Courtenay, L. A., Aramendi, J., Yravedra, J., Mora, R., González-Aguilera, D., Domínguez Rodrigo, M., 2019. Application of geometric morphometrics to the analysis of cut mark morphology on different bones of differently sized animals. Doessizereallymatter? Quaternary International 517, 33-44. Morin, E., Meier, J., El Guennouni, K., Moigne, A. M., Lebreton, L., Rusch, L., Valensi, P., Conolly, J., Cochard, D., 2019. New evidence of broader diets for archaic Homo populations in the northwestern Mediterranean. Science Advances 5, eaav9106. Mosquera, M., Ollé, A., Rodríguez-Álvarez, X. P., Carbonell, E., 2018. Shedding light on the Early Pleistocene of TD6 (Gran Dolina, Atapuerca, Spain): the technological sequence and occupational inferences. PlosOne 13 (1), e0190889. Olsen, S. L., Shipman, P., 1988. Surface modification on bone: trampling versus butchery. Journal of Archaeological Science 15, 535-553. Pelletier, M., Desclaux, E., Brugal, J. Ph., Texier, P.-J., 2019. The exploitation of rabbits for food and pelts by last interglacial Neandertals. Quaternary Science Reviews 224, 105972. Pérez Ripoll, M., 2005. Caracterización de las fracturas antrópicas y sus tipologías en huesos de conejo procedentes de los niveles gravetienses de la Cova de les Cendres (Alicante). Munibe 57, 239-254.

Pickering, T. R., Domínguez-Rodrigo, M., Heaton, J. L., Yravedra, J., Barba, R., Bunn, H. T., Musiba, C., Baquedano, E., Díez-Martín, F., Mabulla, A., Brain, C. K., 2013. Taphonomy of ungulate ribs and the consumption of meat and bone by 1.2-million-year-old hominins at Olduvai Gorge, Tanzania. Journal of Archaeological Science 40, 1295-1309. Pineda, A., Saladié, P., Vergès, J. M., Huguet, R., Cáceres, I., Vallverdú, J., 2014. Trampling versus cut marks on chemically altered surfaces: an experimental approach and archaeological application at the Barranc de la Boella site (la Canonja, Tarragona, Spain). Journal of Archaeological Science 50, 84-93. Pobiner, B. L., De Silva, J., Sanders, W. J., Mitani, J. C., 2007. Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda. Journal of Human Evolution 52, 614-636. Rodríguez -Hidalgo, A., Saladié, P., Marín, J., Canals, A., 2015. Expansion of the referential framework for the rabbit fossil accumulations generated by Iberian lynx. Palaeogeography, Palaeoclimatology, Palaeoecology 418, 1-11. Rodríguez -Hidalgo, A., Saladié, P., Marín, J., Canals, A., 2016. Bird-bone modifications by Iberian lynx: A taphonomic analysis of non-ingested red-legged partridge remains. Quaternary International 421, 228-238. Romero, A. J., Díez, C., Saladié, P., 2016. Mammal bone surface alteration during human consumption: an experimental approach. Journal of Archaeological Science: Reports 8, 82-89. Rosell, J., Blasco, R., Arilla, M., Fernández-Jalvo, Y., 2019. Very human bears: wild brown bear neo-taphonomic signature and its equifinality problems in archaeological contexts. Quaternary International 517, 67–78. Sala, N., Arsuaga, J. L., 2013. Taphonomic studies with wild brown bears (Ursus arctos) in the mountains of northern Spain. Journal of Archaeological Science 40, 1389-1396. Saladié, P., Huguet, R., Díez, C., Rodríguez-Hidalgo, A., Cáceres, I., Vallverdú, J., Rosell, J., Bermúdez de Castro, J. M., Carbonell, E., 2011. Carcass transport decisions in Homo antecessor subsistence strategies. Journal of Human Evolution 61 (4), 425-446. Saladié, P., Huguet, R., Rodríguez -Hidalgo, A., Cáceres, I., Esteban-Nadal, M., Arsuaga, J. L., Bermúdez de Castro, J. M., Carbonell, E., 2012. Intergroup cannibalism in the European Early Pleistocene: The range expansion and imbalance of power hypotheses. Journal of Human Evolution 63 (5), 425-446. Saladié, P., Rodríguez -Hidalgo, A., Díez, C., Martín-Rodríguez, P., Carbonell, E., 2013a. Range of bone modifications by human chewing. Journal of Archaeological Science 40, 380-397. Saladié, P., Huguet, R., Díez, C., Rodríguez-Hidalgo, A., Carbonell, E., 2013b. Taphonomic modifications produced by modern brown bears (Ursus arctos). International Journal of Osteoarchaeology 23, 13-33. Saladié, P., Rodríguez -Hidalgo, A., Huguet, R., Cáceres, I., Díez, C., Vallverdú, J., Canals, A., Soto, M., Santander, B., Bermúdez de Castro, J. M., Arsuaga, J. L., Carbonell, E., 2014. The role of carnivores and their relationship to hominin settlements in the TD6-2 level from Gran Dolina (Sierra de Atapuerca, Spain). Quaternary Science Reviews 93, 47-66.

Saladié, P., Rodríguez-Hidalgo, A. 2019. Reply to “Very human bears” by Rosell et al. 2019 Quaternary International, this volume. Sachis Serra, A. 2010. Los lagomorfos del Paleolítico medio de la región central y sudoriental del mediterráneo ibérico: caracterización tafonómica y taxonómica. PhD University of Valencia. Sanchis Serra, A., Fernández Peris, 2008. Procesado y consumo antro’ pico de conejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia). Complutum19 (1), 25–46. Selvaggio, M. M., 1994. Carnivore tooth marks and stone tool butchery marks on scavenged bones: archaeological implications. Journal of Human Evolution 27, 215-228. Selvaggio, M. M., Wilder, J., 2001. Identifying the involvement of multiple carnivore taxa with archaeological bone assemblages. J Journal of Archaeological Science 28, 465-470. Stiner, M. C., 1994. Honor among thieves: A zooarchaeological study of Neandertal ecology. Princeton, Princeton University Press. Vaquero, M., 2008. The history of stones: behavioural inferences and temporal resolution of an archaeological assemblage from the Middle Palaeolithic. Journal of Archaeological Science 35, 3178-3185. Vaquero, M., Chacón, M. G., García-Antón, M. D., Gómez de Soler, B., Martínez, K., Cuartero, F., 2012. Time and space in the formation of lithic assemblages: the example of Abric Romaní Level J. Quaternary International 247, 162-181. White, T. D., 1992. Prehistoric Cannibalism at Mancos 5MTURM-2346. Princeton: Princeton University Press. Yravedra, J., García-Vargas, S., Maté-González, D., Aramendi, J., Palomeque-González, J. F., Vallés-Iriso, J., Matasanz-Vicente, J., González-Aguilera, D., Domínguez Rodrigo, M., 2017. The use of Micro-Photogrammetry and Geometric Morphometrics for identifying carnivore agency in bone assemblage. Journal of Archaeological Science: Reports 14, 106-115.