Neanderthal and Homo sapiens subsistence strategies in the Cantabrian region of northern Spain

1 23

Archaeological and AnthropologicalSciences ISSN 1866-9557 Archaeol Anthropol SciDOI 10.1007/s12520-015-0253-4

Neanderthal and Homo sapienssubsistence strategies in the Cantabrianregion of northern Spain

José Yravedra-Sainz de los Terreros,Alberto Gómez-Castanedo, JuliaAramendi-Picado, Ramón Montes-Barquín & Juan Sanguino-González

1 23

Your article is protected by copyright and

all rights are held exclusively by Springer-

Verlag Berlin Heidelberg. This e-offprint is

for personal use only and shall not be self-

archived in electronic repositories. If you wish

to self-archive your article, please use the

accepted manuscript version for posting on

your own website. You may further deposit

the accepted manuscript version in any

repository, provided it is only made publicly

available 12 months after official publication

or later and provided acknowledgement is

given to the original source of publication

and a link is inserted to the published article

on Springer's website. The link must be

accompanied by the following text: "The final

publication is available at link.springer.com”.

ORIGINAL PAPER

Neanderthal and Homo sapiens subsistencestrategies in the Cantabrian region of northern Spain

José Yravedra-Sainz de los Terreros1 & Alberto Gómez-Castanedo2 &

Julia Aramendi-Picado1 & Ramón Montes-Barquín3&

Juan Sanguino-González4

Received: 11 November 2014 /Accepted: 4 June 2015# Springer-Verlag Berlin Heidelberg 2015

Abstract The Iberian Peninsula is key for the study of thetransition from the Middle to the Upper Palaeolithic inEurope, as well as for the replacement of Neanderthals by an-atomically modern humans (AMH). On this subject, the mostwidespread misconception assumed that both human speciescoexisted during a certain period of time, after which Homosapiens imposed on Neanderthals who finally got extinct.However, recent proposals based on improved dating methods,discuss this possibility, arguing that the arrival of AMH wasmarked by the complete absence of Homo neanderthalensis inthis territory. In that way, new theories deny the possibility ofcoexistence and the disappearance of Neanderthals by culturaldisplacement. Covalejos Cave (Velo, Pielagos, Cantabria), oneof the few settlements in the northern Peninsula with Final

Mousterian and Early Aurignacian levels, supports this hypoth-esis. Nevertheless, in this paper, we try to avoid a direct discus-sion about this question in order to centre our analysis on iden-tifying possible different subsistence strategies betweenH. neanderthalensis and anatomically modern humans in thenorth of the Iberian Peninsula. Our zooarchaeological and taph-onomic studies reflect that Neanderthals and anatomically mod-ern humans exploited the same faunal species, pointing out thatthere does not seem to be significant differences in their behav-iour in Covalejos Cave.

Keywords Homo neanderthalensis . Anatomicallymodernhumans .Mousterian .Aurignacian .Subsistence .Cantabrianregion . Northern Iberian Peninsula

Introduction

The north of the Iberian Peninsula, especially the Cantabrianregion, has become a reference point for the study of the so-called transition which includes the replacement of Neanderthaltechno-complex by the ones produced by anatomically modernhumans (AMH) and, consequently, the displacement and laterextinction of Homo neanderthalensis. New explanatory frame-works led to the rise of the Ebro Frontier hypothesis (Zilhão2000, 2006; Zilhão and Trinkaus 2002). This proposal exposedthe existence of a Bborder^ along the Ebro river valley, separat-ing Neanderthals and AMH during the Early UpperPalaeolithic. EarlyHomo sapienswould have established northof the border, thanks to a better adaptive capacity to the ecolog-ical and environmental conditions of this area, and in so doing,they would have developed the Aurignacian technology. On theother side of the valley, Neanderthals would have shown pref-erence for the southern environment characterised by more be-nign weather conditions where they had worked on the

Electronic supplementary material The online version of this article(doi:10.1007/s12520-015-0253-4) contains supplementary material,which is available to authorized users.

* José Yravedra-Sainz de los [email protected]

Alberto Gó[email protected]

Ramón Montes-Barquí[email protected]

Juan Sanguino-Gonzá[email protected]

1 Department of Prehistory, Universidad Complutense de Madrid,Ciudad Universitaria s/n, 28040 Madrid, Spain

2 Department of Historical Sciences, University of Cantabria, EdificioInterfacultativo, Avda. Los Castros s/n, 39005 Santander, Cantabria,Spain

3 Itinerario Cultural del Consejo de Europa Prehistoric Rock Art,Madrid, Spain

4 Consejería de Educación, Comunidad de Madrid, Madrid, Spain

Archaeol Anthropol SciDOI 10.1007/s12520-015-0253-4

Author's personal copy

http://dx.doi.org/10.1007/s12520-015-0253-4

http://crossmark.crossref.org/dialog/?doi=10.1007/s12520-015-0253-4&domain=pdf

Mousterian technology. Nevertheless, Neanderthals could alsohave used some shelters in the North until very recently (Baenaet al. 2005, 2012; Garriga et al. 2012).

Other authors have focused on demonstrating the existenceof a great stratigraphic discontinuity (Mallol et al. 2012)among sites with Mousterian and Upper Palaeolithic levels.According to them, from a geological point of view, no clearcontinuous geomorphologic episode can be seen at the sites.Besides, in each of the aforementioned cases, the separationbetween the Mousterian and Upper Palaeolithic levels ischaracterised by erosive events or stratigraphic breaks.However, this work does not aim at establishing a chrono-stratigraphic interrelation of different sites that may indicateepisodes of contemporaneity between Mousterian,Aurignacian or Châtelperronien deposits.

In the same vein, but from a cultural point of view, someresearchers (Martínez-Moreno et al. 2010) observed an episodeof rupture between the Mousterian and Aurignacian levels ofCova Gran (Catalunya, Spain), indicating the existence of differ-ent cultures developed by different hominins. Nevertheless, theydo not dismiss the contemporaneity of both techno-complexes inother regions, thus leaving open the possibility of Neanderthalsand AMH occupying nearby spaces at the same time.

In parallel, a different theory holds that regardless of a possiblecontemporaneity of Neanderthals and AMH, the former got ex-tinct as a result of a less cultural efficiency in comparison withH. sapiens (Binford 1968, 1984; Gamble 1986; Mellars 1989,1996, 2005, 2006; Stiner 1994; Klein 2008; Brown et al. 2009;Shea 2009). However, this theory has come in for considerablecriticism since the existence of major cultural differences, whichcould have led to the disappearance of a certain species and thesuccess of another, is still to be proven (Villa and Roebroeks2014). Precisely, in this respect, many studies have revealed thecomplexity and efficiency of Neanderthals subsistence strategies,which include the exploitation of a wide range of animals andplants that indicate indeed their adaptation to different environ-ments (Scott 1986; Madella et al. 2002; Delpech and Grayson2007; Stringer et al. 2008; Blasco and Fernández Peris 2009;Hardy and Moncel 2011; Yravedra et al. 2015a).

Although Neanderthal hunting efficiency has become in-creasingly accepted, it is still believed that AMHs showedgreater exploitation versatility, including a wider array of re-sources ranging from ungulates to small mammals, bivalves,crustaceans, birds and all kinds of living beings includingextra large animals such as proboscidea (Bailey 1983;Altuna 1990; Straus 1992; Gaudzinski et al. 2005; Surovelland Waguespack 2008; Adán et al. 2009; Hockett and Haws2009; Hoffecker 2009; Richards and Trinkaus 2009; Álvarez-Fernández 2011; Fa et al. 2013).

Nevertheless, recently some studies have highlighted howNeanderthals did also exploit birds (Blasco and FernándezPeris 2009, 2012; Peresania et al. 2011; Finlayson et al.2012; Blasco et al. 2013), small animals like rabbits

(Sanchis and Fernández Peris 2008; Blasco and FernándezPeris 2012; Cochard et al. 2012; Sanchis 2012; Blasco et al.2013), reptiles (Díez et al. 1998; Speth and Tchernov 2002;Blasco 2008; Morales and Sanchis 2009; Blasco et al. 2013);as well as fish and marine resources like bivalve and molluscs(McBurney 1967; Álvarez-Fernández 2005; Bicho and Haws2008; Cortés-Sánchez et al. 2008, 2011; Fa 2008; Stringeret al. 2008; Stiner 2009; Zilhão et al. 2010; Brown et al.2011), large animals like elephants (Yravedra et al. 2012;Smith 2015) and even plant resources (Hardy et al. 2001;Madella et al. 2002; El Zaatari et al. 2011; Salazar-Garcíaet al. 2013).

In recent years, the perception of this transition has sub-stantially changed in Europe and in the Iberian Peninsula.Nowadays, all these models have to face chronological ques-tions (Callaway 2012; De la Rasilla and Santamaría 2013;Higham et al. 2014). Technological improvements of radio-carbon dating depict a much more complex situation: theEuropean territory would have been occupied by differentpopulations, probably even presenting scenarios characterisedby the overlap of Neanderthals andH. sapiens occupations, asit might have been the case in the eastern Mediterranean.Therefore, the Iberian Peninsula has become a particular pointof discussion. Several authors claim that the Spanish mainlandwas one of the last Neanderthals shelters, which have hadenabled the simultaneous presence of both taxa in the samespaces (Vega 1988; Zilhão 1993, 2006; Baena et al. 2005,2012; Finlayson et al. 2006, 2008; Martínez-Moreno et al.2010; Garriga et al. 2012; Straus 2013; Higham et al. 2014).Meanwhile, there is a new model, which revolves around thedisappearance of Neanderthals happening concurrently to thearrival of the first AMHs in the Iberian Peninsula (Jöris et al.2003; Wood et al. 2013; Santamaría and de la Rasilla 2013).This hypothesis is based on the climatic deterioration occurredduring the ISO 3 that could have generated a significant re-duction of the Neanderthal population (D’Errico and SánchezGoñi 2003; Finlayson et al. 2006, 2007; Finlayson andCarrión 2007; Fedele et al. 2008).

Following on from that, in our opinion, the study ofsamples formed by transitional layers covering from theMiddle to the Upper Palaeolithic would facilitate theunderstanding of this transition. In other words, the ex-position of these deposits to the same topographic con-ditions would enable to carry out a comparison on equalterms. Covalejos Cave (Velo, Piélagos, Cantabria), withits different strata dating from isotope stages 3 to 5 andits remains ofMousterian and Aurignacian techno-complexes,meets these requirements. Such chronological and culturaldiversity provides us with the necessary information to ana-lyse on the one hand how Neanderthal subsistence strategiesevolved over time and, on the other hand, to compare them tothe patterns of the firstH. sapienswho lived in the North of theIberian Peninsula.

Archaeol Anthropol Sci


Materials and methods

The analysis of subsistence strategies involves different disci-plines that address diverse aspects such as exploitation pat-terns and the use of rawmaterials, the type of tools used for theacquisition of resources or the analysis of palaeovegetables. Inthis paper, we have focused our attention on the study ofhunted animal resources recovered in Covalejos Cave. Weadopted a taphonomic and zooarchaeological perspective tostudy the 49.799 skeletal remains recovered from levels A, B,C, D, E, H, I, J, K, L, M, O and Q, although in our study wewill refer only to levels B, C, D, E, H, I, J, K and Q becausethey are the most representative. The materials that we haveanalysed include all identifiable and unidentifiable fragments,except for levels J and K whose samples are so numerous andfragmented that we decided to focus on the identifiable andunidentifiable bone fragments >4 cm and to conduct a randomsampling among those fragments <4 cm, selecting 25 % ofthese remains. However, the fragmentation of the assemblageof certain levels is extremely high. For example, as it will bediscussed below, 100 % of the remains of level K are <5 cm,as well as 91.6 % of those found in level J.

As already mentioned, the methods used for the analysis ofsubsistence strategies in Covalejos have a taphonomic back-ground. This is due to previous studies (Yravedra 2006,2010a, b, 2013) where we have shown that taphonomy is aprerequisite for a proper interpretation. Remaining resolute inthis criterion, we will present the results of our taxonomical,mortality patterns and skeletal profile results.

The taxonomical identification is based on reference mate-rial. In the cases where the exact taxonomical determinationwas not feasible, epiphyses and shaft fragments were assignedto approximate animal weight/size classes. In our analysis, wefollowed Uerpmann’s (1973) guideline where small refers toBunn’s (1982) sizes 1 and 2 (animals <250 kg, e.g. Rupicapraor Capra), medium refers to Bunn’s (1982) size 3 (animals250–750 kg, e.g. deer and horse) and large refers to Bunn’s(1982) sizes 4–6 (>700 Kg, e.g. Bison).

The estimation of the number of identified specimens(NISP) and minimum number of individuals (MNI) is usedto quantify the remains in order to determine the most appro-priate features for the description of the faunal taxonomicrepresentation. NISP determination follows Lyman’s (1994)synthesis, whereas MNI is based on Brain’s (1969) model,which includes bone laterality and animal age. Furthermore,skeletal profiles and MNI reconstruction consider shaft thick-ness, section shape and medullar surface properties (Barbaand Domínguez-Rodrigo 2005). In this way, bones are dividedinto four anatomical regions: cranial (i.e. horn, cranium, man-dible and teeth), axial (vertebrae, ribs, pelvis and scapula,sensu Yravedra and Domínguez-Rodrigo 2009), upper appen-dicular limb bones (humerus, radius, ulna, femur, patella andtibia) and lower appendicular limb bones (metapodials,

carpals, tarsals, phalanges and sesamoideal). Long limb boneswere also divided into upper (humerus and femur), intermedi-ate (radius and tibia) and lower (metapodials) limb bones(Domínguez-Rodrigo 1997).

Estimates of the minimum number of elements (MNE) atarchaeological sites often differ substantially depending onwhether epiphyses or shafts are used for element identificationor not (Pickering et al. 2003; Cleghorn and Marean 2004;Domínguez-Rodrigo et al. 2007). Therefore, some researchershave used a GIS-based method to calculate MNE (Mareanet al. 2001). However, we feel more confident in elementestimation if overlap among specimens is documented byhand. Thus, we employed an integrative approach using thebone section division proposed by Patou-Mathis (1984,1985), Münzel (1988) and Delpech and Villa (1993), as de-scribed in detail in Yravedra and Domínguez-Rodrigo (2009).Following Delpech and Villa (1993) and Münzel (1988),shafts were divided in equally sized parts, irrespective of areasof muscular insertion. These sectors (upper shaft, mid-shaft,lower shaft) can be easily differentiated and oriented (cranial,caudal, lateral, medial). Yravedra and Domínguez-Rodrigo(2009) described the criteria used for the division of each shaftsection, taking into account the orientation of each specimen.We also used the criteria of Barba and Domínguez-Rodrigo(2005) for long limb element identification, which is based onshaft thickness, section shape and medullar surface properties.After having identified the element and shaft section, MNEwas quantified by laying out together all specimens from thesame element and size group. This procedure enabled us toobserve all the criteria used in our comprehensive analysis(Lyman 1994), such as element size, side, age and biometrics.

To estimate the MNE of cranial elements such as the upperand lower jaws, we chose the most representative tooth inorder to not give an over-represented quantification. For in-stance, if we have 4 lower jaws, 5 lower M1 and 9 lower M2,the MNE would be 9.

Mortality patterns are divided into several categories: juve-nile, juvenile-adult and adult. Age profiles were estimated fromtooth crown wear and the emergence of the teeth. We based ouranalysis on Guadelli (1998) for the horses and on the formulaproposed by Steele (2002)) for deer, using the lower molars.

We also employed a wide number of methods to reconstructsite formation processes, assess site integrity and evaluate thecontributions of various biogenic agents to the faunal assemblage.

This was done at three levels. First of all, we carried out asize-sorting examination among all fragments. Regardingbone fragmentation indexes, bones were classified into twocategories: <5 or >5.1 cm. Secondly, only long bone frag-ments were considered, as cancellous axial bones undergodifferent fragmentation patterns than do denser limb bones(Domínguez-Rodrigo and Martinez-Navarro 2012). Shaft cir-cumference was also taken into account, since as Bunn (1982)demonstrated, anthropogenic bone concentrations show



greater fragmentation than carnivore ones. Bunn proposesthree categories for the determination of shaft circumferencewhere (1) stands for shaft circumference <25%, (2) covers therange between 25 and 75 % and (3) refers to shaft circumfer-ence >75 %. Thirdly, only those long bone fragmentsdisplaying green breakage were considered. This distinctionis made because diagenetic (dry) breakage is relatively com-mon in archaeological assemblages, and thus, the specimensize distribution in the recovered assemblages may be quitedifferent from that of the original deposited accumulations.

Signs of polishing, rounding or abrasion were recorded as afinal appraisal of water presence at the site. These signs maybe found in transported and non-transported assemblageswhen exposed to moving water and sediments, such as thosefound in sand strata (Thompson et al. 2011). Several bonesshowed cracks and diagenetic breakage planes that causedfurther fragmentation during excavation. Identification ofbreakage planes was carried out following Villa andMahieu’s (1991) guidelines. According to them, dry breakageplanes tend to be longitudinal and/or transverse to the longaxis of the bone; they possess a nearly 90° angle betweenthe cortical and medullar surfaces and show an uneven break-age plane surface with micro-step fractures and a rough tex-ture. On the other hand, green breakage planes arecharacterised by smoother surfaces and are more likely to beoriented obliquely to the long axis of the bone. Breakage pat-tern analysis also followed the methods outlined byDomínguez-Rodrigo et al. (2007), but specifically for thestudy of breakage angles, we based on Alcántara et al. (2006).

Additionally, a systematic examination of bone surfacemodifications such as cut, percussion and tooth marks wascarried out with ×10–×20 hand lenses and different lighting(Blumenschine 1988, 1995). The diagnostic criteria definedby Bunn (1982), Potts and Shipman (1981) and DomínguezRodrigo et al. (2009) guided our identification of cut marks,whereas tooth marks were recorded following Binford (1981)and Blumenschine (1988, 1995). Finally, the identification ofpercussion marks followed Blumenschine and Selvaggio(1988) and Blumenschine (1995). For comparative purposes,surface modifications include the evaluation of epiphysis andshafts (Blumenschine 1988, 1995). Modifications were alsoquantified by element type and bone section (Domínguez-Rodrigo 1997; Domínguez-Rodrigo and Barba 2005) basedon NISP values (Bunn 1982). The presence of tooth, percus-sion and cut marks was recorded for the entirety of remains,whereas estimated percentages include only well-preservedbone surfaces.

Covalejos Cave

Covalejos Cave (Velo, Pielagos, Cantabria) is situated close tothe mouth of the river Pas in the Bay of Santander, 48m above

sea level. The surrounding environment is characterised by agentle landscape with low hills below 250 m (Sanguino andMontes 2005) where other sites with a similar chronologysuch as El Pendo, El Ruso or Santián are to be found (Fig. 1).

Eduardo de la Pedraja discovered this cave in 1872. Sincethem, until 1879, several excavation works were carried outthere. Later, in 1968, Alfonso Moure Romanillo undertook thecleaning of the profiles (Sanguino andMontes 2005). And final-ly, during the threshold from the twentieth to the twenty-firstcentury, R. Montes and J. Sanguino directed four excavationcampaigns (1997–1999 and 2002). Those last years of researchexposed the faunal collection that we present in this paper.

The excavated space of the upper levels comprises an areaof 10 m2 (Fig. 2). The stratigraphic sequence includes somelevels covering isotope stages 3–5 with Mousterian andAurignacians occupations. The most recent levels (B and C)are Early Aurignacian; for the Mousterian period, severallayers have been found (D, H, I, J, K andM); and finally, levelQ might be Acheulean or archaic Mousterian (Sanguino andMontes 2005). This stratigraphic sequence stretching over al-most 4 m (380 cm) is characterised by a gentle dip and ahorizontal layout, where many levels are affected by erosionprocesses (Fig. 3; for further details, see Sanguino andMontes2005). Thus, level Q is covered by an erosive event that sep-arates it from level P. Similarly, levels M and L are isolated bysuch a process. Level K, with its 40 cm thickness, shows a net-plane, contacting level J (60 cm thick). The contact with levelI (20 cm thick) is also flat, but the connection surface withlevel H is erosive. Level E also shows a flat contact with levelD. The latter presents an abrupt contact with the UpperPalaeolithic level C, exposing a stratigraphic disconformityor local redeposition of soliflucted sediment (Sanguino andMontes 2005).

The chronological dating provided by this cavity issummarised in Table 1; this dating supplies a fairly continuoussequence covering the ISO 3–5. Particularly noteworthy arelevels C and D, which synthesise the entire period of thetransition, comprising a time frame around 40,000 BP for boththe Middle and the Upper Palaeolithic.

Palynological (Ruiz-Zapata and Gil 2005) andanthracological analyses (Uzquiano 2005) refer only to theupper levels. The palaeoenvironment seems to have beendominated by a wooded landscape with different taxa depend-ing on weather conditions. Levels H–I show an overall trendtowards a temperate environment with a variety of tree spe-cies, being the main ones birch, pine and Quercus. But fromlevel D to B, a cooling process that leads to lower plant covercan be appreciated. Such an event would have resulted in theemergence of mixed forests with a higher importance ofsteppe and shrub species such as Ericaceae and Juniperus(Ruiz-Zapata and Gil 2005).

The faunal remains cannot be considered a goodpalaeoenvironmental marker, but in the case of Covalejos,



Fig. 2 Covalejos excavated area(1997–2002) from Sanguino andMontes (2005)

Fig. 1 Location of Covalejos



some evidence enables us to infer certain ecological and envi-ronmental conditions. The abundance of deer along the entiresequence highlights the presence of a forested landscapearoundCovalejos. However, in certain situations, the incidenceof reindeer and horse may suggest environmental changes.Following the results published by Castaños (2005), level Jreflects a reduction in deer representation, an increase in horseremains and the presence of reindeer. This would indicate the

existence of a cold climate event that would have led to adecrease of the forested areas while open spaces would havegained importance, certifying a greater environmental dryness.This would have been followed by a forest recovery as shownin the next levels and by a final return of cold conditions inlevel B, as stressed by Ruiz-Zapata and Gil (2005).

Among the micromammals identified by Sesé (2005), thereis a wide variety of species including Pliomys lenki, Galemyspyrenaicus, Microtus oeconommus, Microtus arvalis,Arvicola terrestris, Terricula lusitanicus, Sorex araneus, Glisglis, Clethrionomys glareolus, Apodemus sylvaticus,Chimomys nivalis, Erineaceus europeus and Talpa europeaa.Also the marmot and the rabbit must be added to the group.These mammals suggest the presence of open landscapes withneighbouring forest and shrub areas. They also indicate mildclimatic periods, being the presence of marmot in levelJ and of Microcebus in levels B and J the only evidenceof a possible slight cooling. The scarcity of lagomorphs’remains precludes us from establishing any possible linkbetween them and humans.

Results

Faunal representation and mortality patterns

Covalejos Cave contained remains of Bos-Bison, Canis lupus,Capra pyrenaica, Capreolus capreolus, Cervus elaphus,Crocuta crocuta, Dama sp., Equus caballus, Felix silvestris,Equus hydruntinus, Megaloceros sp., Orictolagus cuniculus,Rangifer tarandus, Dicerorhinus sp., Rupicapra rupicapra,Sus scrofa, Ursus speleus and Vulpes vulpes (Tables 2 and 3).

The entire sequence shows a rather homogeneous faunalrepresentation. Deer is the most represented species in alllevels except for level Q. The second most important specieschanges among levels are as follows: in some cases, it is theroe (level B); in others, the Bos/Bison (levels C, H, I, J) or the

Fig. 3 Covalejos stratigraphic sequence. Figure modified from Sanguinoand Montes (2005)

Table 1 Dates of Covalejos Cave. See dating discussion in Maroto et al. (2012)

Culture Level Date Reference

Achel-Moust? Covalejos Q 101,000 BP (U/TH) Sanguino and Montes (2005)

Estalactitic costra Covalejos N 91,857+4000–4000 (U/TH) Sanguino and Montes (2005)

Mousterian Quina Covalejos J >45,000 BP (GrA-33812) Maroto et al. (2012)

Mousterian Discoide Covalejos D 43,050+750–550 BP (GrA-33811) Maroto et al. (2012)

Mousterian Discode Covalejos D 41,640+650–650 BP Sanguino and Montes (2005)

Mousterian Discoide Covalejos D 40,650+2300–2300 BP Sanguino and Montes (2005)

Mousterian Covalejos H 38,344+3560–3900 BP (TL) Sanguino and Montes (2005)

Archaic Aurignacian Covalejos C 37,940+400–350 BP (GrA-33877) Maroto et al. (2012)

Archaic Aurignacian Covalejos C 32,840+280–250 BP (GrA-24200) Sanguino and Montes (2005)

Mousterian Quina Covalejos I 30,860+340–300 BP (GrA-33822) Maroto et al. (2012)

Aurignacian Covalejos B 30,380+250–250 BP (GrA-22443) Sanguino and Montes (2005)



wild boar (level K). Meanwhile, deer maintains high fluctuat-ing percentages: MNI figures are around 30–85 %, and theirNISP are always over 75 % of the total, with the exception oflevels B and Q. The frequency of species is also constantthroughout the sequence, except for the less representativelevels (E, L–Q) and level J. The latter is in fact the most repre-sentative level of the entire sequence, showing the greatest tax-onomic variability with 15 species (Tables 2 and 3).

Middle and Upper Palaeolithic levels exhibit the same tax-onomic representation, being deer the main species (Fig. 4),especially in level B. This could be related to a mild environ-mental episode. A further difference is the important presenceof carnivores in theMousterian levels. However, regarding theproportion of deer, levels C and D display similar frequencies(Fig. 4; Supplementary File 1).

The diversity of macromammals found in this cave high-lights different climatic and environmental conditions. Taxasuch as reindeer reflect cold conditions in levels B, C, I, J andK. This fact is also corroborated by the presence of chamois inthose levels (except level K). The finding of chamois remainscan only be explained as the result of seasonal movement(Crampe et al. 2007). So during winters, chamois would haveleft the mountains and descended to the woods to seek shelter.Only in very cold or harsh moments chamois could have

moved to the forests of the coastal plain; Covalejos is a goodexample of this type of situation. Moreover, environmentals t r ingency is also supported by the presence ofE. hydruntinus and horse. These species suggest the presenceof dry environments that enable the proliferation of openspaces. However, the finding of deer, roe deer, wild boar andcarnivores—such as bobcat—in these levels, points out that,close to these open spaces, there were also wooded areas.

Finally, the presence of animals like fallow deer in level Dreveals more temperate and humid conditions, a fact that isalso corroborated by the absence of E. hydruntinus, the scar-city of horse and the increase in deer individuals.

Mortality patterns show a predominance of adult individ-uals among all levels and taxa (Table 3). The incidence ofinfantile or juvenile specimens is very scarce and only somenon-adults are to be found in most of the levels.

Skeletal bone profiles

Covalejos skeletal profiles reflect a rather biased sample com-posed of a high number of cranial bones as reflected by theabundance of teeth and a scarce number of axial elements(see tables of NISP in Supplementary File 3). Deer is the bestrepresented animal; all parts of the skeleton have been

Table 2 Taxonomic patterns according to NISP of Covalejos

NISP A B B–C C D E F–G H I J K L M O Q Total

Dicerorhinus 1 5 6

Bos-Bison 3 11 46 10 12 20 60 120 33 52 367

Equus 5 26 57 2 2 9 32 153 54 340

Megaloceros 1 2 1 4

E. hydruntinus 3 1 7 11

Cervus 50 162 66 689 252 178 1 131 517 1198 776 1 7 4 41 4073

Dama 1 1

Rangifer 2 1 2 1 1 7

Size: large-intermediate 47 278 153 45 31 112 2133 1323 5 127 11 273 4538

Rupicapra 2 1 1 2 1 7

Capra 3 4 1 7 20 2 8 28 32 105

Capreolus 6 22 36 5 2 1 7 41 43 37 200

Sus 4 1 3 2 1 4 1 7 23

Size: small indet. 28 713 51 18 17 81 282 552 2 3 47 1794

Crocuta 1 1

Ursus 1 1 2

Canis 2 5 1 2 10

Vulpes 1 2 1 3 2 9

Felix 7 2 1 2 12

Mustela 3 1 4

Carnivore indet. 4 4

Orictolagus 1 2 1 2 3 9

Unidentified 793 9850 452 4964 3341 560 111 801 2285 6836 7377 117 22 762 38,271

Total 864 10,167 520 6803 3845 820 113 1020 3144 10,826 10,198 125 156 18 1180 49,799



identified. On the contrary, the rest of the species are worseportrayed by their skeletal remains, being in some cases onlyidentified by the presence of their teeth (Supplementary File 3).

Bone bias can also be seen in the disparity between theMNI and MNE (Table 4). For example, in levels J and K,more than 10 specimens could be found per individual, whilein other levels, the MNE is usually <10 per individual. Furtherevidence of the bias in Covalejos can be observed whenanalysing the representation of long bone sections. Here, thereis a high disproportion between diaphysis and epiphysis inci-dence, being the former much better represented than the latter(Fig. 5). Hence, estimation of long bonesMNE has beenmadebased on diaphysis fragments; only for levels B and E,

epiphysis of the metatarsals and, for levels J and K, epiphysisof the metacarpals have been used (Fig. 5).

After computing the MNE and grouping the different ana-tomical elements in categories (cranial bones, upper and lowerlimb bones and axial) per animal size—small (Fig. 6), medium(Fig. 7) and large (Fig. 8)—it could be observed that someportions are better represented than we first thought. In thatway, we could verify that each group was made up of bones ofall anatomical parts, suggesting that each and every anatomi-cal section could have been introduced into the site. But mostsignificant results were related to levels B, E, J and K wheresmall-sized animals show a good representation of axial ele-ments and to levels C, H and I where all skeletal portions

Table 3 MNI of the mostimportant levels of Covalejos,where A adults, J juvenile, Iinfantile. See Supplementary File2

MNI B C D E H I J K Q

Ages A/J/I A/J/I A/J/I A/J/I A/J/I A/J/I A/J/I A/J/I A/J/I

Dicerorhinus 1/0/0 1/0/0

Bos-Bison 2/0/0 3/0/0 1/0/0 1/0/0 2/1/0 5/1/0 4/1/0 2/0/0 2/1/0

Equus 2/0/0 1/0/0 1/0/0 1/0/0 1/0/0 2/0/0 5/0/0 1/1/0

Megaloceros 1/0/0 1/0/0 1/0/0

E. hydruntinus 1/0/0 1/0/0 2/0/0

Cervus 5/2/2 16/6/2 9/1/2 22/1/0 8/1/0 34/2/3 42/2/3 5/1/0 2/0/0

Dama 1/0/0

Rangifer 1/0/0 1/0/0 1/0/0 1/0/0 1/0/0

Rupicapra 1/0/0 1/0/0 1/0/0 1/0/0 1/0/0

Capra 1/0/0 1/0/0 1/0/0 1/0/0 1/0/0 1/1/0 1/0/0

Capreolus 4/1/0 2/0/0 1/0/0 1/0/0 1/0/0 4/0/0 2/0/1 1/0/0

Sus 1/0/0 1/0/0 1/0/0 1/0/0 1/0/0 1/0/0 2/1/0

Crocuta 1/0/0

Ursus 1/0/0 1/0/0

Canis 1/0/0 1/0/0 1/0/0 1/0/0

Vulpes 1/0/0 1/0/0 1/0/0 1/0/0

Felix 1/0/0 1/0/0 1/0/0 1/0/0

Mustela 1/0/0 1/0/0

Carnivore indet. 1/0/0

Total 23 29 21 26 18 51 75 20 6

Fig. 4 Taxonomic representation of species introduced by humans in Covalejos. NISP on the left; MNI on the right



appear to be equally represented. Just level D shows a poorlevel representation of the axial skeleton (Fig. 6). This obser-vation contradicts the comments we made at the beginning ofthis section. However, it has to be taken into consideration thatsmall animals show poor significant skeletal profiles with fewspecimens. Only levels C, J and K rely on samples >50 spec-imens; indeed, level K contains more than 100 items, showinga ratio of 15 specimens per individual (Table 4).

Among medium-sized animals, the above-described osteo-logical bias can be observed. Lower appendicular elements arebest represented in levels B, C, D, H and J and cranial ele-ments in E and I, being the axial skeleton predominant only inlevel K (Fig. 7). Upper appendicular bones and axial speci-mens are often poorly represented among medium size spe-cies; there is no level (except K) where both together exceed30 % of the total skeletal representation (Fig. 7).

Large animals seem to be only well represented in levels Jand K with more than 100 specimens, which also means that>16 elements per individual could be recognised (Table 4). Inlevel J, a balanced incidence of all portions could be identi-fied, while level K shows a predominance of axial bones(Fig. 8), similar to that documented among medium- andsmall-sized animals.

In sum, Covalejos includes elements of all bone sections.Nevertheless, some evidence such as the disproportion of el-ements in relation to the MNI (Table 4), the low number ofappendicular elements identified from epiphysis remains(Fig. 5) and the scarcity of axial elements among the bestrepresented animals (Fig. 7 and Supplementary File 3) suggestthat bone sample was exposed to a relative anatomical bias.

Bone brakeage and bone surface modifications

Covalejos faunal remains show a high fragmentation degree ineach and every level that resulted in bone fragments <5 cm.Indeed, 98 % of the total remains are <5 cm and only levels Eand J show a bit lower percentage, with 96.3 and 91.6 %,respectively (Table 5).

The high fragmentation can also be observed when study-ing the long bones: 100% of the shafts correspond to category1 of the index circumference described by Bunn (1982). Thismeans that no shaft exceeds 25 % of its total circumference.

The causes of this may be different, but the abundance oflong bones with green fractures and longitudinal and obliquefracture patterns seems to point out that human and carnivoreactivity could have been the main reason, rather than diage-netic factors.

The preservation of bone surfaces differs among levels.Levels B, C or D, for example, show a poor preservation,while other layers, such as I, J and K contain much well-preserved remains. The cause of poor preservation appearsto be related to different hydraulic disorders. The incidenceof other processes such as weathering is virtually nonexistent.Instead, we have mostly documented other alterationslike abrasion, polishing and rounding (Table 6). Onlylevel K reflects a low incidence of this type of processwith few bones affected by water alteration. By contrast,in the other levels, the percentage of rounded, polishedor abraded bones make up more than 20 %, indicatingthat the materials were exposed to water flows(Table 6). However, these alterations are mainly super-ficial (Supplementary File 4), suggesting that movingwater was not the cause of the sample contribution.The water may rather have flowed on the deposit, erod-ing the bone surfaces without displacing or fracturingthem, as it has been experimentally demonstrated byThompson et al. (2011). This explains why no dry frac-tures have been identified. In fact, most fractures onlong bones seem to be related to processes generatedby carnivores and humans. Despite of its intensity, water flowsmay have caused the disappearance of axial bones, which dueto their buoyancy may have suffered some kind of movement(Voorhies 1969). This would enlighten the low representationof the axial skeleton in the entire sequence, except for level K,which is characterised by a lower incidence of water alter-ation. Nevertheless, the action of other agents such as carni-vores could also elucidate the lack of such elements.

The impact of biological agents on Covalejos faunal re-mains is important. The traces generated by carnivores andhumans insinuate that they were primarily responsible forthe fracturing of the bones and possibly the osteological biasdescribed above.

The action of carnivores has been observed in each andevery level of the cave; tooth marks on animal bones of large,medium and small size were identified (Fig. 9). Levels H and Ishow the highest frequencies of tooth-marked bones with per-centages ranging between 25 and 35 % for medium- andlarge-sized animals. The rest of the layers provide us withlower tooth mark frequencies that do not exceed 15% for eachsize group. The Upper Palaeolithic levels B and C also showsimilar frequencies (Fig. 9).

Table 4 Proportion of elements for each individual

Levels Large Intermediate Small

MNE/MNI N MNE/MNI N MNE/MNI N

B 20/5 4 64/11 5.8 35/9 3.8

C 27/5 5.4 129/25 5.2 52/7 7.4

D 14/2 7 77/13 5.9 24/7 3.5

E 20/3 6.7 66/23 2.9 15/2 7.5

H 15/4 3.8 45/10 4.5 12/4 3

I 35/8 4.4 112/40 2.8 35/7 5

J 182/11 16.5 384/51 7.5 57/13 4.4

K 103/4 25.8 205/7 29.3 120/8 15



Fig. 5 Representation of the epiphysis and diaphysis of limb bones in Covalejos for large- and medium-sized animals



Carnivore activity has affected all skeleton sections, beingthe axial and appendicular bones the most harmed ones(Supplementary File 4). In some levels, only elements relatedto those portions display tooth marks: medium-sized animalsfrom levels B, C, D, E and H do not have marks on their

cranial elements, as well as large-sized species recovered fromlevels B and H. Concerning compact bones of medium-sizedanimals, no tooth marks have been observed in level I, neitheron large and small species from levels B, C, D, E, H, I and K(Supplementary File 4).

Fig. 6 Skeletal profiles of small-sized animals



According to this evidence, we conclude that carnivorescould have been the cause of the registered anatomical bias(Supplementary File 3 and Figs. 6, 7, 8 and 9). But in terms ofactivity, evidence tends to indicate that they were not theagents responsible for the carcasses contribution to the site,but they seem rather to have acted as secondary agents, alter-ing previously introduced bones.

The high fragmentation of the remains reflected in Table 5 andthe absence of cylinders do not correspond to the typical patternof accumulations left by carnivores (Bunn 1982). The absence of

deciduous teeth of carnivores, as well as the lack of coprolites,digested bones and the scarcity of remains of carnivores couldalso be used as arguments to propose that Covalejoswas not usedas a den site. Furthermore, the size of the tooth marks point outthat carnivores ravaging in Covalejos were small. The pits wemeasured do not exceed 4 mm; predominantly, marks are 2×1 mm or 3×2 mm (Supplementary File 4: levels J and K).

According to previous studies that analyse the activity ofcarnivores based on the size of the tooth marks, only foxes,wolves, leopards and lynx could have been responsible for

Fig. 7 Skeletal profiles of medium-sized animals



the faunal assemblage of Covalejos. The dimensions of the pitsare smaller than those left by hyenas, lions and bears(Dominguez-Rodrigo and Piqueras 2003; Selvaggio andWilder 2001; Pinto-Llona et al. 2005; Andrés et al. 2012).The measurements only fit those produced by foxes(Yravedra et al. 2015b), wolves (Yravedra et al. 2011; Andrés

et al. 2012) and medium-sized cats such as leopards (Selvaggioand Wilder 2001; Dominguez-Rodrigo and Piqueras 2003;Domínguez-Rodrigo et al. 2007; Andrés et al. 2012).

Figures 11 and 12 highlight that the frequency of toothmarks observed in Covalejos is also lower than the one expect-ed when carnivores have primary access to the preys. Foxes

Fig. 8 Skeletal profiles of large-sized animals



tend to generate marks on over 40 % of the bones of smallanimals (Yravedra et al. 2015b); hyenas’ rates on small preysare quite similar (>35 %) (Bunn 1982) and they can even leavefrequencies over 50 % on medium- to large-sized animals asshown by Blumenschine (1995) and Capaldo (1997). Wolvescan also generate mark frequencies exceeding 30% onmediumto large animals (Yravedra et al. 2011). But there is no level inCovalejos where these frequencies exceed 25 %. Even level H,which is the most affected by carnivores, shows a total percent-age of 22 % (Fig. 10). Also tooth mark frequencies on limbbones (see Figs. 13 and 14) present lower percentages as ex-pected from a carnivore-created assemblage. Only the leopardgenerates tooth mark frequencies (Domínguez-Rodrigo et al.2007) that could be related to the observed frequencies inCovalejos (Figs. 10, 11, 12, 13 and 14). Moreover, the skeletalprofiles (Supplementary File 3 and Figs. 7, 8 and 9), as well asthe epiphysis (Fig. 5) and axial element bias (SupplementaryFile 3) and the low representation of elements in proportion tothe MNI (Table 4), do not fit the patterns normally created bycats in their accumulations (Brain 1981; Ruiter and Berger2000). Hence, it does not seem likely that this carnivore couldhave been primarily responsible for the carcasses deposit.

Based on all these arguments—high fragmentation, ab-sence of epiphysis, scarcity of axial elements, lack of

deciduous teeth and coprolites, low frequency of tooth marksand small dimension of pits—we propose that carnivores wereinvolved as secondary agents, ravaging assemblages that hadbeen previously transported by humans. Evidence for this hy-pothesis was identified in the form of anthropogenic marks onbones (Fig. 9) and their distribution, functionality and fre-quencies (see bellow), and further supported by the overlapof tooth marks on cut marks observed in levels I, J, K and Q.In addition to that, those bones presenting anthropogenic evis-ceration or fleshing traces that do also show tooth marks sug-gest a carnivore post-anthropic access to the carcasses, too.

Apart from the overlaying of tooth and cut marks, there areother arguments that support the hypothesis that humans wereresponsible for the faunal accumulation in Covalejos.

The anatomical distribution of cut marks and their highfrequency indicate an anthropogenic early access to meat re-sources. The frequencies of cut marks obtained in Covalejosfit the experimental results showing human primary access tomeat on large, medium and small animals (Domínguez-Rodrigo 1997; Lupo and O’Connell 2002; Barba andDomínguez-Rodrigo 2005) (Figs. 14 and 15). The surface ofupper limb bones—femur and humerus—shows frequencies>20 % for large (Fig. 14) and small animals (Fig. 15) whichmatch the results obtained by Domínguez-Rodrigo (1997).The low frequency of cut marks on metapodials at Covalejosis also similar to the one observed on frameworks for primaryanthropogenic access (Figs. 14 and 15). Consequently, bothNeanderthals and H. sapiens would have been responsible forthe faunal accumulation of Covalejos.

Moreover, the distribution of the marks shows varioustypes of activities, such as skinning, inferred from cut markson phalanges and cranial elements; evisceration, marks on theventral side of ribs; fleshing, traces on axial such external sideof ribs and appendicular elements; and disarticulation, markson various articular elements, metadiaphysis and condyles,such as the mandibular condyle (Supplementary File 4,Fig. 16).

The presence of percussion marks reveals that after meatconsumption, the exploitation of the carcasses continued withthe acquisition of the bone marrow. This intensive use of thecarcasses explains the high fragmentation of the faunal assem-blage in Covalejos and acts, meanwhile, as evidence for thehuman authorship, since high fracturing is a well-known fea-ture of anthropic accumulations (Bunn 1982).

In this sense, bones presenting thermal alterationswork as further evidence for human activity inCovalejos. Levels concerning the Upper as well as theMiddle Palaeolithic contain burnt bones, with levels B,C and D having a higher frequency (Supplementary File4). However, despite the presence of burnt bones, weare not able to link them with a possible use as fuel, aspreviously noted in other Upper and Middle Palaeolithicsites in Cantabria (Yravedra et al. 2005; Yravedra and

Table 5 Covalejos fragmentation patterns. The NISP includes teeth

NR % Bones <5 cm % Shafts <25 % O

B 10,167 98 100

C 6803 99.1 100

D 3845 99.9 100

E 820 96.3 100

H 912 100 100

I 3165 99 100

J 10,826 91.6 100

K 10,198 100 100

Q 1180 99 100

Table 6 Preservation of bone surface and water alteration. The NISPconsidered here does not include teeth

NISP Bad preservation Rounded Polished Abrasion

B 10,010 7367 (73.6) 2066 (20.6) 2066 (20.6) 2031 (20)

C 6232 4306 (69.1) 5115 (82.1) 4951 (79.4) 5106 (81.9)

D 3677 2175 (59.2) 2748 (74.7) 2748 (74.7) 2762 (75.1)

E 697 36 (5.2) 199 (28.6) 199 (28.6) 193 (27.7)

H 912 570 (62.5) 100 (11) 47 (5.2) 130 (4.3)

I 2668 194 (7.3) 770 (28.9) 735 (23.2) 390 (14.6)

J 10,149 2859 (28) 4071 (40) 4107 (40) 3285 (32)

K 10,101 2575 (25.5) 1036 (10.3) 379 (3.8) 911 (9)

Q 1154 476 (41.3) 861 (74.6) 636 (55.1) 834 (72.3)



Uzquiano 2013). Not even level B, where 85.6 % of thetotal burnt remains were found, suggests this type ofuse; the burnt bones only prove low degrees ofwarming, far from giving evidence of carbonization orcalcination processes.

Discussion and conclusion

The study of the transition from Middle to UpperPalaeolithic does not only involve the analysis ofchanging techno-complexes or biological substitutions.

Fig. 9 Frequency of tooth, cut and percussion marks among large-, medium- and small-sized animal groups in the most representative levels



Other issues that include similarities and notable dis-crepancies need to be addressed in order to fully under-stand the significance of this period. A comparativestudy regarding subsistence behaviour of Neanderthalsand H. sapiens would be useful, providing a corpus ofdata that facilitates the comprehension of the phenome-non as a whole.

Covalejos Cave is an archaeological site that has provided awide stratigraphic sequence, ranging from isotope stages 3 to5. Therefore, this place is of great importance for the under-standing of the evolution of Neanderthal way of life over along period of time during the Middle Palaeolithic. On top ofthat, the higher levels of Covalejos belong to the UpperPalaeolithic and are therefore linked to H. sapiens; this means

Fig. 10 Tooth mark frequencies on large-sized carcasses in CovalejosCave compared to the patterns generated by wolves, where Wolf* refersto infant carcasses of horse which were hunted and consumed by wolvesin only one visit; Wolf**: carcasses of adult horse scavenged by wolvesin many events; Wolf***: carcasses of infant animals scavenged bywolves inmany visits (Yravedra et al. 2011); Vulpes*: carcasses modifiedby foxes in Ayllón; Vulpes**: carcasses modified by foxes in Ourtiaga(Yravedra et al. 2015b); carcasses modified by Leopards and Cheetah

(Domínguez-Rodrigo et al. 2007); Hy1-2: hyena tooth mark frequen-cies on size 1-2 carcasses (sensu Bunn 1982); Hy3-4: hyena toothmark frequencies on size 3-4 carcass, where hyena is the first andhumans the secondary consumer (c-h) and vice versa (H-C)(Blumenschine 1995); Hy H-C1: hyena tooth mark frequencies wherehyena is the first and human the subsequent consumer; and Hy H-C2: hyena tooth mark frequencies where hyena is the first and hu-man the second consumer (Capaldo 1997)

Fig. 11 Tooth mark frequencies on Small-sized carcasses in CovalejosCave compared to the patterns generated by wolves, where Wolf* refersto infant carcasses of horse which were hunted and consumed by wolvesin only one visit; Wolf**: carcasses of adult horse scavenged by wolvesin many events; Wolf***: carcasses of infant animals scavenged bywolves inmany visits (Yravedra et al. 2011); Vulpes*: carcasses modifiedby foxes in Ayllón; Vulpes**: carcasses modified by foxes in Ourtiaga(Yravedra et al. 2015b); carcasses modified by Leopards and Cheetah

(Domínguez-Rodrigo et al. 2007); Hy1-2: hyena tooth mark frequen-cies on size 1-2 carcasses (sensu Bunn 1982); Hy3-4: hyena toothmark frequencies on size 3-4 carcass, where hyena is the first andhumans the secondary consumer (c-h) and vice versa (H-C)(Blumenschine 1995); Hy H-C1: hyena tooth mark frequencies wherehyena is the first and human the subsequent consumer; and Hy H-C2: hyena tooth mark frequencies where hyena is the first and hu-man the second consumer (Capaldo 1997)



that this cave is key for the comparative analysis of the sub-sistence strategies of the last Neanderthals and the first AMHs.

The aim of this work revolved around the analysis of sub-sistence strategies throughout the occupation time of the cav-ity, trying to determine the evolution of Neanderthal behav-iour and, in so doing, to establish differences and simi-larities in relation to H. sapiens. Such a study is not asimple task, since it implies carrying out a holistic workthat involves different disciplines. Our work has ad-dressed these phenomena by analysing the fossil evi-dence from two different and specialised perspectives:zooarchaeology and taphonomy.

One of the most significant findings is that from theMiddlePalaeolithic level K to the Upper Palaeolithic level B, deer hasremained the most important taxon throughout the sequence.Its abundance is such that, in many levels, like the UpperPalaeolithic level C and the Mousterian levels E, I and J, itexceeds 65 % of the MNI, and in level D, it represents >55 %of the total faunal assemblage.

This could be a reflection of hunting strategies with a highdegree of specialisation and deer as the most commonlyhunted species. This finding adds to the observations madeby numerous authors who have already recognised huntingspecialised behaviours among Neanderthals contradicting

Fig. 13 Toothmark frequencies onappendicular bones of small-sizedcarcasses in Covalejos Cavecompared to the patterns generatedby wolves, where Wolf* refers toinfant carcasses which were huntedand consumed by wolves in onlyone visit; Wolf** carcasses adultanimals scavenged by wolves inmany events; Wolf***: are infantanimals scavenged by wolves inmany visits (Yravedra et al. 2011);-Vulpes*: carcasses modified byfoxes inAyllón. Vuleps**: carcassesmodified by foxes in Ourtiaga(Yravedra et al. 2015b) and car-casses modified by leopards andcheetah (Domínguez-Rodrigoet al. 2007)

Fig. 12 Tooth mark frequencies onappendicular bones of large-sizedcarcasses in Covalejos Cavecompared to the patterns generatedby wolves, where Wolf* refers toinfant carcasses which were huntedand consumed by wolves in onlyone visit; Wolf** carcasses adultanimals scavenged by wolves inmany events; Wolf***: are infantanimals scavenged by wolves inmany visits (Yravedra et al. 2011);-Vulpes *: carcasses modified byfoxes in Ayllón. Vuleps**: carcassesmodified by foxes in Ourtiaga(Yravedra et al. 2015b) and car-casses modified by leopards andcheetah (Domínguez-Rodrigoet al. 2007)



the traditional interpretation that advocates that such habits arecultural developments of the Upper Palaeolithic (Binford andBinford 1966; Mellars 1989, 1996, 2005; Ready 2010). ManyMousterian sites have been found to reflect specialised behav-iours or the predominance of one species (Burke 2000;Gaudzinski and Roebroeks 2000; Gaudzinski 2006; Nivenet al. 2012; Rendu et al. 2012).

Taxonomic patterns also highlight that Neanderthal hunt-ing strategies have not varied greatly over time. Thus, there isnot only a significant predominance of deer catches, but alsovery similar patterns of mortality—with a preponderance of

adult individuals in all taxa—can be appreciated throughoutthe sequence (Table 3 and Supplementary File 1).

However, regardless of occupational recurrence of the cav-ity during the Middle and Upper Palaeolithic, evidence sug-gests that the occupations did not last long. Such data wouldbe, for example, the existence of significant hydraulic alter-ations on many bones found in different levels, which meansthat at times the cave must have been flooded, complicatingthereby the habitability of the place. Similarly, bone destruc-tion caused by carnivores has been observed in each and everylevel. This implies carnivore ravaging of bones previously

Fig. 14 Cut mark frequencies onappendicular elements of large-sized animals in Covalejos com-pared to the models for early hu-man access to large-sized carcassesof Hadza (Lupo and O'Connell2002), early human access tolarge-sized carcasses (A)(Domínguez-Rodrigo 1997), earlyhuman access to the small-sizedcarcasses (B) (Barba andDomínguez-Rodrigo 2005) andearly carnivore access (C)(Domínguez-Rodrigo 1997). Darkboxes represent the range of valuesfor early carnivore access (C)

Fig. 15 Cut mark frequencies onappendicular elements of small-sized animals in Covalejos com-pared to the models for early hu-man access to large-sized carcassesof Hadza (Lupo and O’Connell2002), early human access tolarge-sized carcasses (A)(Domínguez-Rodrigo 1997), earlyhuman access to the small-sizedcarcasses (B) (Barba andDomínguez-Rodrigo 2005) andearly carnivore access (C)(Domínguez-Rodrigo 1997). Darkboxes represent the range of valuesfor early carnivore access (C)



discarded by humans when the cave had been abandoned bythe latter.

Carnivore activity is a fairly common phenomenon inarchaeological sites. Previous works (see Straus 1982 orLindly 1988) showed how their activity is relativelycommon in many Middle and Upper PalaeolithicIberian caves. In other cases, an alternation of human

and carnivore occupation over time could be detected.In sites such as Cave Amalda (Yravedra 2007, 2010a),El Ruso (Yravedra et al. 2010) or Hornos de la Peña(Yravedra 2010b), anthropogenic contributions duringthe Upper and Middle Palaeolithic could be distin-guished from those made by carnivores, since both fo-cused on different taxa.

Fig. 16 Anatomical distribution of cut marks on large-, medium- and small-sized animals in Covalejos levels J andK. Supplementary File 4 provides thefrequencies obtained



In connection with this issue, we could discuss how impor-tant the activity of humans and carnivores was in Covalejos;however, as noted in the BResults^ section, evidence showsthat different carnivores acted after Neanderthals and AMHshad left the site. Those conclusions focus on the high frag-mentation of the bone assemblage, the absence of cylinders,the frequencies of tooth marks, the overlapping of tooth markson previous cut marks and on the anatomical distribution ofmarks and their frequencies. All these together compose asolid argumentation to support the anthropic activity as maincause for the accumulation in Covalejos. Regarding the activ-ity of carnivores, the small size of the tooth marks, the scarcityof axial elements and the absence of epiphysis would suggestthat small canids like the fox could be responsible for the post-human damage. Nevertheless, these tooth marks could also berelated to the action of leopards and lynx, but the lack ofepiphysis and the skeletal profiles do not fit the patterns ofthese carnivores. Moreover, cats primarily rely on meat re-sources and Covalejos cut mark frequencies on the upper limbbones suggest that humans had already processed the majormuscle packets before carnivores could intervene. In sum, itseems likely that small canids would have been the perpetra-tors of carnivore damage on the assemblage. This fact is con-firmed by the size of the tooth marks (always <5 mm pits).

Synthesising, we could conclude that there were two mainactors in Covalejos: Neanderthals for the Mousterian levelsand H. sapiens for the Aurignacians levels.

Regarding the transport mechanisms of the carcasses to thesite, there has been a major skeletal bias and intense fracturingcaused by carnivores. These two factors have hindered thecomplete reconstruction of the skeletal profiles. However,the representation of all anatomical sections for large, mediumand small animals suggests that the entire body of the preyswould have been transported to the site. So far, this is only ahypothesis; future excavations at the site will allow us to bettersubstantiate this proposal.

Finally, we will address the subsistence strategies of the lastNeanderthals and the first AMHs in northern Iberia. As notedabove,we defend the need to carry out studies on siteswith levelsof both periods in order to reliably assess whether there wererelevant behavioural differences or similarities betweenH. neanderthalensis and H. sapiens. Concerning Covalejos, wehave documented evidence for very similar behaviour patterns.

Nearby sites such as El Ruso or El Pendo show similarpatterns to the one identified in Covalejos, with a clear pre-dominance of deer remains. The former contains Mousterianand Aurignacian levels where the frequency of deer is lowerthan in Covalejos: <50 % of the MNI but >70 % of NISP(Yravedra et al. 2010; Yravedra 2013). The latter suffers fromstratigraphic problems that have led to question the impor-tance of the stratigraphic package (Montes et al. 2005).Nevertheless, zooarchaeological studies showed a predomi-nance of deer throughout the El Pendo stratigraphic sequence

(Fuentes 1980), and seasonality analyses reflect similarities inthe occupation patterns during the Upper and MiddlePalaeolithic (Pike-Tay et al. 1999).

AMHs getting access to a wider range of resources couldsuggest an important difference in comparison to the subsis-tence strategies of Neanderthals. Thus, some authors haveproposed that AMH exploited a wider range of resources in-cluding birds, reptiles, bivalves, fish and other marine re-sources, as well as small mammals (Bailey 1983; Altuna1990; Straus 1992; Adán et al. 2009; Hockett and Haws2009; Álvarez-Fernández 2011; Fa et al. 2013). Also isotopicanalyses have shown that AMHs had a more varied and rich inaquatic resources diet than Neanderthals who focused on theconsumption of large preys (Hoffecker 2009; Richards andTrinkaus 2009).

However, these findings present some problems and thedebate about subsistence strategies among AMHs andNeanderthals is more complex than it may seem.

Although it is possible that AMH got access to a greaterrange of resources, studies show that the wide range of exploi-tation patterns developed by AMH did not become general-ised until the ISO 2 at the end of the Gravettian or EarlySolutrean or even until the Magdalenian (Straus 1992; Adánet al. 2009; Álvarez-Fernández 2005, 2011; Hockett andHaws2009; Gutierrez et al. 2013). Gutierrez et al. (2013) andÁlvarez-Fernández (2005, 2011) have demonstrated that fish-ing and the exploitation of marine resources had been sporad-ically carried out in the North of the Iberian Peninsula sincethe Mousterian. Also during the Aurignacian, this type ofpractice has been recognised, yet rare. Adán et al. (2009))pointed out that fishing in the Cantabrian coast can only beexceptionally documented before the Solutrean andMagdalenian, with evidence during the Aurignacian andGravettian being very scarce. Only a greater increase in theuse of resources and shellfish gathering can be appreciatedsince the end of the Upper Palaeolithic and especially fromthe Magdalenian period.

In the Portuguese area, Hockett and Haws (2009) observedthat the exploitation of large and small game, birds or aquaticresources is known from the Middle Palaeolithic, but its gen-eralisation took place from the Gravettian onwards, especiallyduring the Solutrean. In the Italian area, Phoca-Cosmetatou(2009) showed that the generalisation of small prey exploita-tion occurred in the Magdalenian.

The same process is observed in the Iberian Mediterraneanregion. However, since the Middle Palaeolithic sites with ev-idence of small prey exploitation are relatively frequent: rab-bits are often found in Bolomor, Les Canalettes or Quebrada(Sanchis and Fernández Peris 2008; Blasco and FernándezPeris 2012; Cochard et al. 2012; Sanchis 2012; Blasco et al.2013; Salazar-García et al. 2013), birds in Bolomor andGorham’s cave (Blasco and Fernández Peris 2009, 2012;Finlayson et al. 2012; Blasco et al. 2013), reptiles (Blasco



2008; Morales and Sanchis 2012; Blasco et al. 2013) and fishand marine resources in Andalucia, Murcia and Gibraltar(Cortés-Sánchez et al. 2008, 2011; Fa 2008; Stringer et al.2008; Zilhão et al. 2010; Brown et al. 2011).

Although it seems that AMHswere exploiting a wide rangeof different resources, this only started to become widespreadin the Late Upper Palaeolithic. Therefore, and becauseNeanderthals were also exploiting these kinds of resourcesin various sites such as Bolomor, the Quebrada, Gorham’sCave, Los Aviones Cave, as well as in several other sites,we can only conclude that the first AMHs arriving to south-western Europe had no different behaviour from that foundamong Neanderthals. Therefore, it seems likely that subsis-tence strategies of Neanderthals and H. sapiens were not sig-nificantly different.

By contrast, isotopic analyses may lead to different conclu-sions. Some isotopic studies have shown that Neanderthals’consumption was based on the ingest of large animals unlikethat of AMHs who had a more varied diet including marineresources (Hoffecker 2009; Richards and Trinkaus 2009).However, these findings are debatable. First, these analysesare based on few sites and few individuals. Second, most ofthe studied deposits linked to AMH are sites from the ISO 2,and in some cases such as Paviland, Arene Candide orRochette, the selected sites are close to the coast or largerivers. Moreover, some of the Neanderthal sites, e.g. LesRochers-de-Villeneuve 1, present isotopic evidence of marineresources consumption. Thus, only the analysis of new sam-ples aimed at comparing the remains of Neanderthals andH. sapiens during the ISO 3 living in nearby areas will allowto establish more reliable comparative patterns.

Regarding the exploitation of ungulates, other sites in theNorth of the Iberian Peninsula like Hornos de la Peña(Yravedra 2010b), Morín (Yravedra and Gómez-Castanedo2010), El Castillo (Dari 2003; Landry and Burke 2006) orLezetxiki (Altuna 1972) also point out some continuity insubsistence and hunting patterns. At all these sites, the samespecies were hunted during the Mousterian and the UpperPalaeolithic. This characteristic can also be recognised at siteslike Amalda with Mousterian and Gravettian levels (Yravedra2010a). In all these cases, the most represented taxon is thedeer. Indeed, only in certain mountainous or rocky environ-ments, hunting strategies focused on other resources such asthe Iberian ibex, as observed at the Mousterian site ofEsquilleu (Yravedra et al. 2015a).

Future zooarchaeological and taphonomic studies at transi-tional sites with Upper and Middle Palaeolithic layers like LaGüelga, El Mirón, Arrillor or La Viña will extend the refer-ences on the subsistence strategies of Neanderthals andH. sapiens in northern Iberia. But based on the current dataavailable for this area, we conclude that no significant differ-ences were recorded among the subsistence behaviour of bothhominids, and we could rather speak of certain continuity and

the hunting differences cannot be used as an explanatory ar-gument for the extinction of Neanderthals with respect toH. sapiens.

Acknowledgments We thank the Museum of Prehistory and Archae-ology of Cantabria (MUPAC) and especially Raul Gutiérrez and PedroFernández Vega for their attention and help in 2007 while we conductedthe study of the materials presented in this work. We also appreciate thesuggestions and comments of the reviewers.

References

Adán G, Álvarez-Lao D, Turrero P, Arbizu M, García-Vázquez E (2009)Fish as diet resource in North Spain during the Upper Paleolithic. JArchaeol Sci 36:895–899

Alcántara V, Barba R, Barral Del Pino JM, Crespo AB, Vidal A, FalquinaA, Herrero S, Ibarra A, Megías M, Pérez M, Pérez V, Rolland J,Yravedra J, Vidal A, Domínguez-Rodrigo M (2006) Determinaciónde procesos de fractura sobre huesos frescos: un sistema de análisisde los ángulos de los planos de fracturación como discriminador deagentes bióticos. Trab Prehist 63:37–45

Altuna J (1972) Fauna de Mamíferos de los Yacimiento Prehistórico deGuipuzcua. Munibe XXIV:1–404

Altuna J (1990) La caza de herbívoros durante el paleolítico y mesolíticodel P.Vasco. Munibe 42:229–240

Álvarez-Fernández E (2005) La explotación de los moluscos marinosdurante el Paleolítico superior y el Mesolítico en la RegiónCantábrica y en el Valle del Ebro: pasado y presente de lainvestigación. Munibe 57:359–368

Álvarez-Fernández E (2011) Humans and marine resource interactionreappraised: archaeofauna remains during the late Pleistocene andHolocene in Cantabrian Spain. J Anthropol Archaeol 30:327–343

Andrés M, Gidna AO, Yravedra J, Domínguez-Rodrigo M (2012) Astudy of dimensional differences of tooth marks (pits and scores)on bones modified by small and large carnivores. ArchaeolAnthropol Sci 4:209–219

Baena J, Carrión E, Ruiz B, Ellwood B, Sesé C, Yravedra J, Jordá J,Uzquiano P, Velázquez R, Manzano I, Sánchez A, Hernández F(2005) Paleoecología y comportamiento humano durante elPleistoceno Superior en la comarca de Liébana: la secuencia de laCueva del Esquilleu, Occidente de Cantabria, España. In: LasherasJA,Montes R (eds) Neandertales cantábricos. Estado de la Cuestión.Monografías Museo de Altamira 461–487

Baena J, Carrión E, Cuartero F, Fluck H (2012) A chronicle of crisis: thelate Mousterian in northern Iberia (Cueva del Esquilleu, Cantabria,Spain). Quat Int 247:199–211

Bailey G (1983) Economic change in late Pleistocene Cantabria. In:Bailey G (ed) Hunter-gatherer economy in prehistory: a Europeanperspective. 149–165

Barba R, Domínguez-RodrigoM (2005) The taphonomic relevance of theanalysis of bovid long limb bone shaft features and their applicationto element identification. Study of bone thickness and morphologyof the medullar cavity. J Taphonomy 3:29–42

Bicho N, Haws J (2008) At the land’s end: marine resources and theimportance of fluctuations in the coastline in the prehistoric hunt-er–gatherer economy of Portugal. Quat Sci Rev 27:2166–2175

Binford L (1968) Early Upper Pleistocene adaptations in the Levant. AmAnthropol 70:707–717

Binford LR (1981) Bones: ancient men and modern myths. Academic,New York

Binford LR (1984) Faunal remains fromKlasies RiverMouth. Academic,New York



Binford L, Binford A (1966) The predatory revolution: a consideration ofthe evidence for a new subsistence level. AmAnthropol 68:508–512

Blasco R (2008) Human consumption of tortoises at level IVof BolomorCave (Valencia, Spain). J Archaeol Sci 35:2839–2848

Blasco R, Fernández Peris J (2009) Middle Pleistocene bird consumptionat level XI of Bolomor Cave (Valencia, Spain). J Archaeol Sci 36:2213–2223

Blasco R, Fernández Peris J (2012) Small and large game: human use ofdiverse faunal resources al level IV of Bolomor Cave (Valencia,Spain). Comptes Rendus Palevol 11:265–282

Blasco R, Rosell J, Fernández Peris J, Arsuaga JL, Bermúdez de CastroJM, Carbonell E (2013) Environmental availability, behavioural di-versity and diet: a zooarchaeological approach from the TD10–1sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain) andBolomor Cave (Valencia, Spain). Quat Sci Rev 70:124144

Blumenschine R (1988) An experimental model of the timing of hominidand carnivore influence on archaeological bone assemblages. JArchaeol Sci 15:483–502

Blumenschine R (1995) Percussion marks, tooth marks and the experi-mental determinations of the timing of hominid and carnivore accessto long bones at FLK Zinjanthropus, Olduvai Gorge, Tanzania. JHum Evol 29:21–51

Blumenschine R, Selvaggio MM (1988) Percussion marks on bone sur-faces as a new diagnostic of hominid behaviour. Nature 333:763–765

Brain CK (1969) The contribution of Namib Desert Hottentot to under-standing of Australopithecus bone accumulations. Sci PapNamibian Desert Res Station 32:1–11

Brain CK (1981) The hunters or the hunted? University of Chicago Press,Chicago

Brown KS et al (2009) Fire as an engineering tool of early modernhumans. Science 325:859–862

Brown K, Fa DA, Finlayson G, Finlayson C (2011) Small game andmarine resource exploitation by Neanderthals: the evidence fromGibraltar. In: Bicho NF et al. (eds) Trekking the shore: changingcoastlines and the antiquity of coastal settlement, interdisciplinarycontributions to archaeology. Springer 247–272

Bunn HT (1982) Meat-eating and human evolution: studies on the dietand subsistence patterns of Plio-Pleistocene hominins in East Africa.Ph. Dissertation. University of California, Berkeley

Burke A (2000) The view from Starosole: faunal exploitation at a middlePalaeolithic site inWestern Crimea. Int J Osteoarchaeol 10:325–335

Callaway E (2012) Date with history. Nature 485:27–29Capaldo SD (1997) Experimental determinations of carcass processing

by Plio-Pleistocene hominids and carnivores at FLK 22(Zinjanthropus), Olduvai Gorge, Tanzania. J Hum Evol 33:555–598

Castaños P (2005) Revisión actualizada de las faunas demacromamíferosdel Würn antiguo en la Región cantábrica. In: Neandertalescantábricos. Estado de la cuestión. Museo Nacional y Centro deInvestigación de Altamira 20: 201–207

Cleghorn N,Marean CW (2004) Distinguishing selective transport and insitu attrition: a critical review of analytical approaches. JTaphonomy 2:43–67

Cochard D, Brugal JP, Morin E,Meignen L (2012) Evidence of small fastgame exploitation in the middle Paleolithic of Les Canalettes(Aveyron, France). Quat Int 264:32–51

Cortés-Sánchez M et al (2008) Palaeoenvironmental and cultural dynam-ics of the coast of Málaga (Andalusia, Spain) during the UpperPleistocene and early Holocene. Quat Sci Rev 27:2176–2193

Cortés-SánchezM et al. (2011) Earliest known use ofmarine resources byNeanderthals. PLoS One 6–24026

Crampe JP, Bon R, Gerard JF, Serrano E, Caens P, Florence E, GonzalezG (2007) Site fidelity, migratory behaviour, and spatial organizationof female sards (Rupicapra pyrenaica) in the Pyrenees NationalPark, France. Can J Zool 85:16–25

D’Errico F, Sánchez Goñi MF (2003) Neanderthal extinction and themillennial scale climatic variability of OIS 3. Quat Sci Rev 22:769–788

Dari A (2003) Comportement de subsistence pendant le transitionpaaleolithique moyen-Paleolithique superieur en Cantabria à partirde l'etude archaeozooologique des restes osseaux des grandsmamiferes de la Grotte d'el Castillo (Espagne) Museun NationaleD’histoire Naturalle IPH MNHN

De la Rasilla M, Santamaría D (2013) El Paleolítico medio en Asturias.Mainake 23:23–62

Delpech F, Grayson D (2007) Chasse et subsistance aux temps deNeandertal. Biologie et cultures Vandermersch B; Maurelle B. andCopens Y. Documents prehistoriques

Delpech EP, Villa P (1993) Activités de chasse et boucherie dans la grottedes Eglises. In Exploitation des animaux sauvages a travers detemps. IV, In: Desse J and Audoin-Rouzeau F (eds), ColloqueInternational de l'Homme et l'Animal. Editions APDCA: 79–102

Díez C, Arribas Herrera A, Jordá JF (1998) Torrejones (TamajónGuadalajara). A hyaena den on human occupation. En economicprehistorique: Les compertements de subssistence auPaleolithique. En XVIII Rencontres internationale d'Archeologie etd’histoire d’Antibes- Edit. APDECA. Sophia Antipolis

Domínguez Rodrigo M, De Juana S, Galán AB, Rodríguez M (2009) Anew protocol to differentiate trampling marks from butchery cutmarks. J Archaeol Sci 36:2643–2654

Domínguez-Rodrigo M (1997) Meat eating by early homids at FLK Zinj22 site, Olduvay Gorge Tanzania: an experimental approach usingcut-mark data. J Hum Evol 33:669–690

Domínguez-Rodrigo M, Barba R (2005) A study of cut marks on small-sized carcasses and its application to the study of cut marked bonesfrom small mammals at the FLK Zinj site. J Taphon 3:121–134

Domínguez-Rodrigo M, Martinez-Navarro B (2012) Taphonomic analy-sis of the early Pleistocene (2.4 Ma) faunal assemblage from A.L.894 (Hadar, Ethiopia). J Hum Evol 62:315–327

Dominguez-Rodrigo M, Piqueras A (2003) The use of tooth pits to iden-tify carnivore taxa in tooth-marked archaeofaunas and their rele-vance to reconstruct hominid carcass processing behaviours. JArchaeol Sci 30:1385–1391

Domínguez-Rodrigo M, Barba M, Egeland CP (2007) DeconstructingOlduvai: a taphonomic study of the bed I sites. Springer, Dordrecht

El Zaatari SE, Grinec FE, Ungar PS, Hublin JJ (2011) Ecogeographicvariation in Neandertal dietary habits: evidence from occlusal molarmicrowear texture analysis. J Hum Evol 61:411–424

Fa DA (2008) Effects of tidal amplitude on intertidal resource availabilityand dispersal pressure in prehistoric human coastal populations: theMediterranean–Atlantic transition. Quat Sci Rev 27:2194–2209

Fa JE, Stewart JR, Lloveras L, Vargas JM (2013) Rabbits and homininsurvival in Iberia. J Hum Evol 64:233–241

Fedele FG, Giaccio B, Hajdas I (2008) Timescale and cultural process at40,000 BP in the light at the Campanian Ignimbrite eruption,Western Eurasia. J Hum Evol 55:834–857

Finlayson C, Carrión JC (2007) Rapid ecological turnover and its impacton Neanderthal and other human population. Trends Ecol Evol 22:213–222

Finlayson C et al (2006) Late survival of Neanderthals at the southern-most extreme of Europe. Nature 443:850–853

Finlayson C et al (2007) Caves as archives of ecological and climaticchanges in the Pleistocene—the case of Gorham’s Cave, Gibraltar.Quat Int 181:55–63

Finlayson C et al (2008) Gorham’s cave, Gibraltar—the persistence of aNeanderthal population. Quat Int 181:64–71

Finlayson C et al (2012) Birds of a feather: Neanderthal exploitation ofraptors and corvids. PLoS One 7:e45927. doi:10.1371/journal.pone.0045927



http://dx.doi.org/10.1371/journal.pone.0045927

http://dx.doi.org/10.1371/journal.pone.0045927

Fuentes C (1980) Estudio de la Fauna del Pendo en González Echegaray,El Yacimiento de la Cueva del Pendo exc. 1953.57. Bibl PrehistóricaHisp 17:215–238

Gamble C (1986) The paleolithic settlement of Europe. CambridgeUniversity Press, Cambridge

Garriga J, Martínez K, Baena J (2012) Neanderthal survival in the northof the Iberian Peninsula? Reflections from a Catalan and Cantabrianperspective. J World Prehist 25:81–121

Gaudzinski S (2006) Monospecific or species-dominated faunal assem-blages during the Middle Paleolithic of Europe. In: Hovers E, KuhnS (eds) Transitions before the transition. Springer, New York, pp137–147

Gaudzinski S, Roebroeks W (2000) Adults only, reindeer hunting at theMiddle Palaeolithic site Salzgitter Lebenstedt, northern Germany. JHum Evol 38:497–521

Gaudzinski S, Turner E, Anzidei AP, Álvarez-Fernández E, Arroyo-Cabrales J, Cinq-Mars J, Dobosi VT, Hannus A, Johnson E,Münzel SC, Scheer A, Villa P (2005) The use of Proboscideanremains in every-day Palaeolithic life. Quat Int 126–128:179–194

Guadelli JL (1998) Détermination de l’age des caveaux fossiles etétablissement des chasses d’age. Paléo 10:87–93

Gutierrez I, Cuenca-Solana D, Rasines del Río P, Muñoz E, Santamaría S,Morlote JM (2013) The role of shellfish in hunter–gatherer societiesduring the Early Upper Palaeolithic: a view from El Cucorockshelter, northern Spain. J Anthropol Archaeol 32:242–256

Hardy BL, Moncel MH (2011) Neanderthal use of fish, mammals, birds,starchy plants and wood 125–250,000 years ago. PLoS ONE 6(8):23–68

Hardy BL, Kay M, Marks AE, Monigal K (2001) Stone tool function atthe paleolithic sites of Starosele and Buran Kanya III, Crimea: be-havioral implications. Proc Natl Acad Sci U S A 98:10972–10977

Higham T et al (2014) The timing and spatiotemporal patterning ofNeanderthal disappearance. Nature 512:306–309

Hockett B, Haws J (2009) Continuity in animal resource diversity in thelate Pleistocene human diet of Central Portugal. Before Farming2009(2):1–14

Hoffecker JH (2009) Neanderthal and modern human diet in easternEurope. In: Hublin JJ, Richards MP (eds) The evolution ofHominin diets: integrating approaches to the study of Palaeolithicsubsistence, Springer Sci 87–98

Jöris O, Alvarez Fernandez E, Weninger B (2003) Radiocarbon evidenceof the Middle to Upper Palaeolithic transition in SouthwesternEurope. Trab Prehist 60:15–38

Klein RG (2008) Out of Africa and the evolution of human behavior.Evol Anthropol 17:267–281

Landry G, Burke A (2006) El Castillo: the Obermaier faunal collection.Zona Arqueológica. Homenaje 7:104

Lindly J (1988) Hominid and carnivore activity at Middle and UpperPaleolithic cave sites in eastern Spain. Munibe 40:45–70

Lupo KD, O’Connell JF (2002) Cut and tooth mark distributions on largeanimal bones: ethnoarchaeological data from the Hadza and theirimplications for current ideas about early human carnivory. JArchaeol Sci 29:85–109

Lyman RL (1994) Relative abundance of skeletal specimens and tapho-nomic analysis of vertebrate remains. Palaios 9:288–298

Madella M, Jones MK, Goldberg P, Goren Y (2002) The exploitation ofplant resources byNeanderthals in AmudCave (Israel): the evidencefrom phytolith studies. J Archaeol Sci 29:703–719

Mallol C, Hernández C, Machado J (2012) The significance of strati-graphic discontinuities in IberianMiddle-to-Upper Palaeolithic tran-sitional sites. Quat Int 275:4–13

Marean CW, Abe Y, Nilssen P, Stone E (2001) Estimating the minimumnumber of skeletal elements (MNE) in zooarchaeology: a reviewand a new image-analysis GIS approach. Am Antiq 66:333–348

Maroto J, Vaquero M, Arrizabalaga A, Baena J, Baquedano E,Jordá J, Juliá R, Montes R, Van Der Plicht J, Rasines P,

Wood R (2012) Current issues in late Middle Palaeolithic chronolo-gy: New assessments from Northern Iberia. Quaternary International247:15–25

Martínez-Moreno J, Mora R, De la Torre I (2010) The Middle-to-UpperPalaeolithic transition in Cova Gran (Catalunya, Spain) and the ex-tinction of Neanderthals in the Iberian Peninsula. J Hum Evol 58:211–226

McBurney CBM (1967) The Haua Fteah (Cyrenaica) and the Stone Ageof the southeast Mediterranean. Cambridge University Press,Cambridge

Mellars P (1989) Major issues in the emergence of modern humans. CurrAnthropol 30:349–385

Mellars P (1996) The Neanderthal legacy. Princeton University Press,Princeton

Mellars P (2005) The impossible coincidence. A single-species model forthe origins of modern human behaviour in Europe. Evol Anthropol14:12–27

Mellars P (2006) Archeology and the dispersal of modern humans inEurope: deconstructing the BAurignacian^. Evol Anthropol 15:167–182

Montes R, Sanguino J, Martín P, Gómez AJ, Morcillo C (2005). Lasecuencia estratigráfica de la Cueva del Pendo (Escobedo deCamargo, Cantabria): Problemas geoarqueológicos de un referentecronocultural. In: Geoarqueología y patrimonio en la PenínsulaIbérica y el entorno mediterráneo. Santonja M, Pérez González A,and Machado MJ (ed) 139–159

Morales JV, Sanchis A (2009) The quaternary fossil record of the genustestudo in the Iberian Peninsula. Archaeological implications anddiacronic distribution in the western Mediterranean. J Archaeol Sci36:1152–1162

Münzel SC (1988) Quantitative analysis and archaeological site interpre-tation. Archaeozoologia 2:93–110

Niven L, Steele MT, Rendu W, Mallye JB, McPherron SP, Soressi M,Jaubert J, Hublin JJ (2012) Neandertal mobility and large-gamehunting: the exploitation of reindeer during the Quina Mousterianat Chez-Pinaud Jonzac (Charente-Maritime, France). J Hum Evol63:624–635

Patou-Mathis ME (1984). Contribution a l’étude des mammifères descouches supérieures de la Grotte du Lazaret. M. A. Dissertation.Université de la Sorbonne, Paris (unpublished)

Patou-Mathis ME (1985) La fracturation des os longs de grandsmammifères: élaboration d’un lexique et d’une fiche type.Outillage peu élabore en os et en bois de cervidés. Artefacts 1:11–22

Peresania M, Fiore I, Gala M, Romandini M, Tagliacozzo A (2011) LateNeandertals and the intentional removal of feathers as evidencedfrom bird bone taphonomy at Fumane Cave 44 ky B.P., Italy. ProcNatl Acad Sci 108:3888–3893

Phoca-Cosmetatou N (2009) Specialisation & diversification: a tale oftwo subsistence strategies from Late Glacial Italy. Before farming2009/3. 1–29

Pickering TR, Marean C, Domínguez-Rodrigo M (2003) Importance oflimb bone shaft fragments in zooarchaeology: a response to Bon insitu attrition and vertebrate body part profiles^ (2002), by M.C.Stiner. J Archaeol Sci 30:1469–1482

Pike-Tay A, Cabrera V, Bernaldo de Quirós F (1999) Seasonal variationof the Middle-Upper Paleolithic transition at El Castillo, CuevaMorin and El Pendo (Cantabria Spain). J Hum Evol 36:283–317

Pinto-Llona AC, Andrews PJ, Etxebarría F (2005) Tafonomía yPaleobiología de úrsidos cuaternarios cantábricos. Fundación osode Asturias, Proaza

Potts R, Shipman P (1981) Cutmarks made by stone tools from OlduvaiGorge, Tanzania. Nature 291:577–580

Ready E (2010) Neandertal man the hunter: a history of Neandertal sub-sistence. Explor Anthropol 10:58–80



Rendu W, Costamango S, Meignen L, Soulier MC (2012) Monospecificfaunal spectra inMousterian contexts: implications for social behav-ior. Quat Int 247:50–58

Richards M, Trinkaus E (2009) Isotopic evidence for the diets ofEuropean Neanderthals and early modern humans. PNAS 106(38):16034–16039

Ruiter JD, Berger LR (2000) Leopard as taphonomic agents in dolomiticcaves—implications for bone accumulations in the hominid-bearingdeposits of South Africa. J Archaeol Sci 27:665–684

Ruiz-Zapata MB, Gil MJ (2005) Los neandertales cántabros su paisajevegetal. In Neandertales cantábricos. Estado de la cuestión. MuseonNac Cent Invest Altamira 20:275–284

Salazar-García DC, Power RC, Sanchis A, Villaverde V, Walker J, HenryAG (2013) Neanderthal diets in central and southeasternMediterranean Iberia. Quat Int 318:3–18

Sanchis A (2012) Los lagomorfos del Paleolítico medio en la vertientemediterránea ibérica. In: Humanos y otros predadores como agentesde aporte y alteración de los restos óseos en yacimientosarqueológicos. Serie Trabajos Varios 115. Servicio deInvestigación Prehistórica de la Diputación de Valencia, Valencia

Sanchis A, Fernández Peris J (2008) Procesado y consumo antrópico deconejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia).El nivel XVIIc (ca 350 ka). Complutum 19:25–46

Sanguino J, Montes R (2005) Nuevos datos para el conocimiento delPaleolítico Medio en el centro de la región cantábrica: La cueva deCovalejos. In: Neandertales cantábricos. Estado de la cuestión.Museon Nacional y Centro de Investigación de Altamira 20:443–460

Santamaría D, de la Rasilla M (2013) Datando el final del Paleolíticomedio en la Península Ibérica. Problemas metodológicos y límitesde la interpretación. Trab Prehist 70:241–263

Scott K (1986) The bone assemblages of layers 3 and 6. In: Callow P,Cornford JM (eds) La Cotte de St. Brelade, 1961–1978. Geo Books,Norwich, pp 159–183

Selvaggio MM, Wilder J (2001) Identifying the involvement of multiplecarnivore taxon with archaeological bone assemblages. J ArchaeolSci 28:465–470

Sesé C (2005) Aportación de los micromamíferos al conocimientopaleoambiental del Pleistoceno Superior de la Región Cantábrica:Nuevos datos y síntesis. In: Montes Barquín, R. y LasherasCorruchaga, JA (eds): Neandertales Cantábricos, estado de lacuestión. Monografías del Museo Nacional y Centro deInvestigación de Altamira, 20: 167–200

Shea JJ (2009) The ecological impact of projectile weaponry in LatePleistocene human evolution. In: Hublin J-J, Richards MP (eds)The evolution of hominin diets: integrating approaches to the studyof Palaeolithic subsistence. Springer, Leipzig, pp 189–199

Smith GM (2015) Neanderthal megafaunal exploitation in westernEurope and its dietary implications: a contextual reassessment ofLa Cotte de St Brelade (Jersey). J Hum Evol 78:181–201

Speth JD, Tchernov E (2002) Middle Paleolithic tortoise use at KebaraCave (Israel). J Archaeol Sci 29:471–483

Steele MTE (2002) Red deer: their ecology and how they were hunted bylate Pleistocene Hominids in western Europe. Department ofAnthropological sciences and the Committee on Graduate Studiesof Stanford University in partial fulfillment of the requirements forthe degree of Doctor of Philosophy

Stiner M (1994) Honor among thieves: a zooarchaeological study ofNeanderthal ecology. Princeton University Press, Princeton

Stiner MC (2009) Prey choice, site occupation intensity & economicdiversity in the Middle–early Upper Palaeolithic at the ÜçağizliCaves, Turkey. Before Farming 3:1–20

Straus LG (1982) Carnivores and cave sites in Cantabrian Spain. JAntropol Res 1:75–96

Straus LG (1992) Iberia before the Iberians. The Stone Age prehistory ofCantabrian Spain. University of New México Press, Albuquerque

Straus LG (2013) Iberian archaeofaunas and hominin subsistence duringmarine isotope stages 4 and 3. In: Clark J, Speth JD (eds)Zooarchaeology and modern human origins: human hunting behav-ior during the Later Pleistocene. Vertebrate Paleobiology andPaleoanthropology, DOI: 10.1007/978-94-007-6766-96 Springer97–128

Stringer C, Finlayson C, Barton RNE, Fernández-Jalvo Y, Cáceres I,Sabin RC, Rhodes EJ, Currant AP, Rodríguez-Vidal J, Giles-Pacheco F, Riquelme JA (2008) Neanderthal exploitation of marinemammals in Gibraltar. PNAS 105(38):14319–14324

Surovell TA, Waguespack NN (2008) How many elephant kills are 14?Quat Int 191:82–97

Thompson CEL, Ball S, Thompson TY, Gowland R (2011) The abrasionof modern and archaeological bones by mobile sediments: the im-portance of transport modes. J Archaeol Sci 38:784–793

Uerpmann HP (1973) Animal bone finds and economic archaeology: acritical study of Bosteoarchaeological^ method. World Archaeol 4:307–322

Uzquiano P (2005) El registro antracológico durante la transiciónMusteriense-Paleolítico superior inicial en la Región Cantábrica.Vegetción, paleoambiente y modos de vida anrrededor del fuego.In: Neandertales cantábricos. Estado de la cuestión. MuseonNacional y Centro de Investigación de Altamira. 20:255–274

Vega G (1988) El Paleolítico Medio del Suroeste español y AndalucíaOriental. PHD Universidad Complutense Madrid

Villa P, Mahieu E (1991) Breakage patterns of human long bones. J HumEvol 20:1–22

Villa P, Roebroeks W (2014) Neandertal demise: an archaeological anal-ysis of the modern human superiority complex. PLoS ONE 9(4):96424. doi:10.1371/journal.pone.0096424http://www.plosone.org/article/info:doi/10.1371/journal.pone.0096424

Voorhies M (1969) Taphonomy and population dynamics of an earlyPliocene vertebrate fauna, Knox country, Nebraska, special paperno. 1. University of Wyoming Press

Wood RE et al (2013) Radiocarbon dating casts doubt on the late chro-nology of the Middle to Upper Palaeolithic transition in southernIberia. Proc Natl Acad Sci U S A 110:2781–2786

Yravedra J (2006) Acumulaciones biológicas en yacimientosarqueológicos: Amalda VII y Esquilleu III-IV. Trab Prehist62:55–78

Yravedra J (2007) Nuevas contribuciones en el comportamientocinegético de la Cueva de Amalda. Munibe 58:43–88

Yravedra J (2010a) A taphonomic perspective on the origins of the faunalremains from Amalda Cave (Spain). J of tapho 8:301–334

Yravedra J (2010b) Zooarqueología y tafonomía del yacimiento deHornos de la Peña (San Felices de Buelna, Cantabria). Complutum21:69–86

Yravedra J (2013) New contributions in the subsistence of Middle-UpperPalaeolithic in northern of Spain. In: Clark J, Speth JD (eds)Zooarchaeology and modern human origins: human hunting behav-ior during the Later Pleistocene. Vertebrate Paleobiology andPaleoanthropology, doi: 10.1007/978-94-007-6766-96 Springer77–95

Yravedra J, Domínguez-Rodrigo M (2009) The shaft-based methodolog-ical approach to the quantification of long limb bones and its rele-vance to understanding hominid subsistence in the Pleistocene: ap-plication to four Palaeolithic sites. J Quat Sci 24:85–96

Yravedra J, Gómez-Castanedo A (2010) Tafonomía en Cueva Morín.Resultados preliminares de un estudio necesario. Zephirus LXVII:69–90

Yravedra J, Uzquiano P (2013) Burnt bone assemblages from ElEsquilleu cave (Cantabria, Northern Spain): deliberate use for fuelor systematic disposal of organic waste? Quat Sci Rev 68:175–190

Yravedra J, Baena J, Arrizabalaga A, Iriarte MJ (2005) El empleo dematerial óseo como combustible durante el Paleolítico Medio ySuperior en el Cantábrico. Observaciones experimentales. In:



http://dx.doi.org/10.1007/978-94-007-6766-96

http://dx.doi.org/10.1371/journal.pone.0096424http://www.plosone.org/article/info:doi/10.1371/journal.pone.0096424

http://dx.doi.org/10.1371/journal.pone.0096424http://www.plosone.org/article/info:doi/10.1371/journal.pone.0096424

http://dx.doi.org/10.1007/978-94-007-6766-96

Neandertales cantábricos estado de la cuestión (ed) Mesa deTrabajo. El Paleolítico Medio cantábrico Museo de Altamira,Cantabria, pp 369–383

Yravedra J, Muñoz E, Gómez-Castanedo A (2010) Estrategias deSubsistencia en el yacimiento del Ruso (Igollo, Camargo,Cantabria España). Espacio, Tiempo y Forma 3: 39–58

Yravedra J, Lagos L, Bárcena F (2011) A taphonomic study of wild wolf(Canis lupus) modification of horse bones in northwestern Spain. JTaphon 9:37–67

Yravedra J, Rubio Jara S, Panera J, Uribelarrea D, Pérez-González A(2012) Elephants and subsistence. Evidence of the human exploita-tion of extremely large mammal bones from the Middle Palaeolithicsite of PRERESA (Madrid, Spain). J Archaeol Sci 39:1063–1071

Yravedra J, Gómez-Castanedo A, Aramendi J, Baena J (2015a)Neanderthal hunting specialisation on Iberian Ibex during theMiddle Palaeolithic at El Esquilleu. Antiquity 88:1035–1049

Yravedra J, Fosse P, Andrés M, Besson JP (2015b) Taphonomic analysisof small ungulates modified by fox (Vulpes vulpes) in southwesternEurope. J Taphonomy (in press)

Zilhão J (1993) Le passage du Paléolithique moyen au PalèolithiqueSupérieur dans le Portugal. In: Cabrera V (ed) El origen del hombremoderno en el Suroeste de Europa. UNED 127–145

Zilhão J (2000) The Ebro frontier: a model for the late extinction ofIberian Neanderthals. In: Stringer CB, Barton RNE, Finlayson JC(eds) Neanderthal on the edge. Oxbow Books, Oxford, pp 111–121

Zilhão J (2006) Neandertals and moderns mixed, and it matters. EvolAnthropol 15:183–195

Zilhão J, Trinkaus E (2002) Portrait of the artist as a child. The gravettianhuman skeleton from the Abrigo do Lagar Velho and its archaeo-logical context. In: Trabalhos deArqueologia 22. Instituto Portuguêsde Arqueologia, Lisboa

Zilhão J, Angelucci DE, Badal-García E, d’Errico F, Daniel F, Dayet L,Douka K, Higham TFG, Martínez-Sánchez MJ, Montes-Bernárdez R, Murcia-Mascarós S, Pérez-Sirvent C, Roldán-García C, Vanhaeren M, Villaverde V, Wood R, Zapata J(2010) Symbolic use of marine shells and mineral pigmentsby Iberian Neandertals. Proceedings of the National Academyof Sciences Early Edition 1–6



Neanderthal and Homo sapiens subsistence strategies in the Cantabrian region of northern Spain

Documents

Transcript of Neanderthal and Homo sapiens subsistence strategies in the Cantabrian region of northern Spain