at SciVerse ScienceDirect

Journal of Human Evolution 63 (2012) 568e575

Contents lists available

Journal of Human Evolution

journal homepage: www.elsevier .com/locate/ jhevol

News and views

Defending the early human occupation of Vallparadís (Barcelona, IberianPeninsula): A reply to Madurell-Malapeira et al. (2012)

Joan Garcia a,b,*, Kenneth Martínez a,b, Eudald Carbonell a,b,c, Jordi Agustí a,b,d, Francesc Burjachs a,b,d

a IPHES, Institut Català de Paleoecologia Humana i Evolució Social (IPHES), C/Escorxador s/n, 43005 Tarragona, SpainbURV, Universitat Rovira i Virgili, Àrea de Prehistòria, Avinguda de Catalunya 35, 43002 Tarragona, Spainc Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP), Chinad ICREA, Institut Català de Paleoecologia Humana i Evolució Social (IPHES), Universitat Rovira i Virgili, C/Escorxador s/n, 43005 Tarragona, Spain

a r t i c l e i n f o

Article history:Received 21 February 2012Accepted 1 June 2012Available online 28 June 2012

Keywords:Late Early PleistoceneLithic industryCut marksHuman settlement

* Corresponding author.E-mail address: jgarcia@prehistoria.urv.cat (J. Garc

0047-2484/$ e see front matter � 2012 Elsevier Ltd.http://dx.doi.org/10.1016/j.jhevol.2012.06.001

differentiated accumulations of material, which suggests therecould have been several episodes of sedimentation and which,

Introduction

In the last decade, the debate regarding the early settlement ofEurope at the end of the Early Pleistocene has been resolved. Themature Europe hypothesis, which asserts that hominins have beenpresent on the continent for more than a million years, is no longerquestioned. Vallparadís has helped to close this debate (Martínezet al., 2010) by contributing new lines of interpretation andpointing to the continued occupation by hominins of the IberianPeninsula from the end of the Early Pleistocene (1.4e1.2 Ma[millions of years ago]) up to the MatuyamaeBrunhes boundary(0.78 Ma) (Garcia and Martínez, 2011; Garcia et al., 2011a, b). In thepresent article, we refute the opinions offered recently by the teamof paleontologists (Madurell-Malapeira et al., 2012) with whomweexcavated Vallparadís (Martínez et al., 2010). In their article, theyquestion the human occupation of Vallparadís during the EarlyPleistocene. As we have shown in previous papers (Martínez et al.,2010; Duval et al., 2011; Garcia and Martínez, 2011; Garcia et al.,2011a, b): 1) a lithic set attributed to the Mode 1 has been foundat the site; 2) associated with this set are bone remains with cut-marks and anthropogenic fractures; 3) the vertical distribution ofthe fossil record at EVT7 (levels 10 and 10c) presents three

ia).

All rights reserved.

along with the identification of four direct lithic refittings, mayindicate that part of the record was in primary position; 4) thepresence of lithic industry at different levels throughout thestratigraphic sequence suggests continued human occupation; and5) the radiometric (ESR-U/Series) and paleomagnetic datings, andthe biochronological data place the sequence and the differentlevels with lithic industry at between 1.07 (lower limit of the Jar-amillo subchron) and 0.6 Ma (Madurell-Malapeira et al., 2010;Martínez et al., 2010; Duval et al., 2011) (Fig. 1).

Archaeological record and depositional environment

Madurell-Malapeira et al. (2012) support their assertion that thelithic industry is ‘doubtful’ based on attributes of the depositionalenvironment. Just because part of the lithic record is in secondaryposition or presents rounded edges mainly by sediment abrasionand to a lesser extent by transport in water, does not rule out theexistence of lithic industry. What turns a lithic object into anarchaeological one is the presence of technical features that origi-nate during the knapping process. There is a large record of lithicindustry at Vallparadís made from local stones (quartz, flint, lydite,quartzite and sandstone) using a chaîne opératoire that is shortboth in time and space, and that involves minimum interventionand exploitation of the objects. The following knapping methodshave been identified among the cores: 1) bipolar on anvil, gener-ating matrices with volumetric structures that are usually cubic orpolyhedral; 2) unifacial centripetal with plano-convex structures;and 3) bifacial centripetal with discoidal features. The first system isthe most common, most probably because it was related to thetechnological tradition of these human groups and not merelybecause it allowed small cobbles to be more easily knapped(Martínez et al., 2010). For their part, although the unifacials andbifacials centripetal cores are clearly secondary, they can be inter-preted as a sign of archaic technology given that they are found atsites such as Dmanisi in Georgia (de Lumley et al., 2005), andUbeidiya in Israel (Bar-Yosef and Goren-Inbar, 1993).

The flakes generated by these knapping methods, particularlythose obtained from the bipolar on anvil method, present

Figure 1. Geological context: summary of the lithostratigraphy and chronology of the Quaternary sedimentation at the Vallparadís site. The synthetic column shows the differentstratigraphic units to the right and the magnetostratigraphy column to the left. (1) Organic material and fossil wood remains, (2) root marks, (3) gastropod remains, (4) CaCO3

remains, (5) units bearing archaeological remains, (6) cross-lamination, (7) Upper Pleistocene terrace, (8) clays and muds with gastropods, (9) unit 5, (10) red clays and muds, (11)unit 7, (12) brown clays and muds, (13) conglomerates, and (14) paleo-floor.

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575 569

quadrangular morphologies, although there are also rectangularflakes with long cutting dihedrals of up to 50 mm. Many of theflakes show cortical surfaces both on the dorsal face and on thestriking platform, without prior preparation of the percussionplatforms, and with straight or irregular ventral surfaces. We havecarried out extensive experiments to better differentiate the tech-nical features of these types of objects. These experiments looked atthe use of quartz as the rawmaterial, the knapping method and thesedimentary context of the archaeological level. This revealed thatthe flakes and cores obtained from bipolar knapping on an anvilpresent some technical features to take into account. Thus, forexample, flakes that present a double striking platform are habit-ually attributed to these methods, but during the experimentsthese only represented 30% of the end product. Likewise, theexperiments revealed a high degree of fracturing of the strikingplatform resulting from the blow by the percussor and the coun-terblow from the anvil. For the same reason, the flakes show highpercentages of transversal factures. Moreover, the intense

exploitation of the cores using this method completely eliminatesthe morpho-technical characteristics that would allow us torecognize them.

Themost common retouched artefacts are denticulates on smallpebbles, occasionally associated with notches and segments withscraper retouches. The most common retouched flakes are alsodenticulate and are usually made of short segments with unifacialretouches. The notches tend to be retouched, unifacial, deep andlocated on the same edge, although tools with associated doublenotches have been documented. There are few side-scrapers onwhich series of continuous or regular retouches have been carriedout around the perimeter. A certain technical specialization can befound among the retouched tools in that there are various technicalmodels or standardized morphotypes. One of the most interestingis the bec, made by means of either a double notch or two abruptand continuous retouched segments that converge on a dihedral ortrihedral convex area. Another model presents an abrupt denticu-late lateral associated with a distal notch. This morphotype is

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575570

present in sites such as Gran Dolina TD6 (Carbonell et al., 1999) andPont de Lavaud in France (Despriée et al., 2006). In general, there isa high percentage of retouched tools, which is in contrast to otherEuropean sites with Mode 1 industry, such as Fuente Nueva 3(Martínez-Navarro et al., 1997), Barranco León 5 (Toro-Moyanoet al., 2009), Gran Dolina TD6 (Carbonell et al., 1995), Sima delElefante (Carbonell et al., 2008), Dmanisi, (de Lumley et al., 2005)and Ubeidiya (Bar-Yosef and Goren-Inbar, 1993). This highpercentage could be because there is an indeterminate number offlakes generated by the bipolar method that do not conserve theirtechnical characteristics and that have therefore been classified asfragments (Fig. 2 and Fig. 3).

The characteristics of the lithic industry in units 3 (EVT3) and 4(EVT4) suggest a technological rupture at the MatuyamaeBrunhesboundary, because they are larger in size and show longer chaîneopératoire than in level 10 (Fig. 2, num. 23 and 24). Madurell-Malapeira et al. (2012:169) state that the lithic industry recoveredfrom Middle Pleistocene units 3 and 4 is “out of context” and usethis as an argument against continued human occupationthroughout the sequence. In fact, because of the archaeologicalrichness of level 10, we excavated the entirety of it in accordancewith archaeological methodology. The archaeological record in theremaining levels was much less dense. Therefore, we excavatedthese levels with the paleontological team when we encounteredaccumulations of lithic and faunal remains, and grouped the arte-facts according to the level in which they were found. Therefore, allof the archaeological material found at Vallparadís is placed incontext and thus lends itself to diachronic interpretationsregarding human settlements (Martínez et al., 2010; Garcia andMartínez, 2011; Garcia et al., 2011a, b).

Moreover, there is further evidence of a human presence atlevel 10 of Vallparadís in the form of the anthropic modificationsmade to the faunal remains (e.g., cut-marks and fractures) (Fig. 4).These have already been described in other papers (Martínezet al., 2010; Cáceres et al., 2011), but are not mentioned byMadurell-Malapeira et al. (2012). Although the definitive tapho-nomic study has yet to be published, these preliminary dataalready mean that Vallparadís has more documented remainswith anthropic marks than any of the other open air EuropeanEarly Pleistocene sites such as Orce (Huguet, 2007; Espigares,2010), Ubeidiya (Gaudzinski, 2004) and Dmanisi (Lordkipanidzeet al., 2007). For our part, we have not denied that the boneremains recovered at Vallparadís were principally accumulatedand modified by Pachycrocuta brevirostris. Nevertheless, thisscavenger causes a bias in the record by eliminating remainsdepending on the animal size and structure, mineral density,marrow content and fat of the bones.

Madurell-Malapeira et al. (2012: 172), refer to Huguet (2007),when they assert that hominins would have had “occasionalprimary access to carcasses.” However, if we take into account thelow density of human populations during the Early Pleistocene andthe evidence from all Early Pleistocene sites (Dmanisi, Orce, Ubei-diya, Sima del Elefante, Gran Dolina TD6 and Vallparadís) of cut-marks and anthropic fracturing, it may be said that the activitiesof the scavengers have obscured any evidence that humans regu-larly had primary access to carcasses (Faith et al., 2007). Therefore,we propose that hominins were just one of several contemporaryEuropean predators that were all in direct competition with eachother. The Gran Dolina TD6 site presents both the greatest evidenceof anthropic intervention in fossil remains and the lowestpercentage of tooth-marks. These cut-marks occur in the context ofa cave, in contrast with the other Early Pleistocene sites in theIberian Peninsula. We therefore conclude that anthropic activity atVallparadís could be underrepresented because of the activities ofcarnivores and scavengers at open air sites.

Furthermore, the sediment at the different levels of the strati-graphic sequence has been described and presented in previouspapers (Martínez et al., 2010; Duval et al., 2011; Garcia andMartínez, 2011; Garcia et al., 2011a, b), and coincides in thisrespect with the interpretation put forward by Madurell-Malapeiraet al. (2012). EVT7 is made up of two archaeological levels: theupper level 10 (with a maximum thickness of 2 m) and the lowerlevel 10c (up to 40 cm thick). The sediment of level 10 is formed bygrey-green clays and mud with angular and subangular clasts ofquartz, lydite and metamorphic rock ranging in size from 1 mm to1 cm in a matrix supported fabric in which abundant root markscan be found. Level 10c is made up of gravel and conglomeratesdominated by large-scale rounded blocks in a matrix supportedfabric. This level was probably formed by the accumulation oftransported massive debris-flow and to a lesser extent by sheetflows with erosive capabilities. Level 10 was formed when clay wasdeposited as a result of dense mudflows moving in an NEeSWdirection. These mudflows were caused by successive seasonalriver level rises and were rapidly colonized by vegetation. Fieldobservations in EVT7, which were confirmed by vertical projectionsof fossil material, show that the record is distributed between threeaccumulations separated by 20e30 cm intervals (Fig. 5 and Fig. 6).The first accumulation is the least dense and is situated in the upperpart of the level among mudstone with root marks (contactbetween levels 9 and 10). Of particular note are proboscideandiaphyseal elements with sub-aerial alteration, although most ofthe remains come from medium sized fauna (e.g., Cervidae andEquidae). The main accumulation (level 10) appears at mid heightand contains singular elements that are absent from the previousaccumulation, such as hyena coprolites and deer horns completewith tooth marks, in addition to a large number of appendicularelements fromHippopotamidae and Rhinocerotidae that have beensignificantly modified by carnivores (e.g., P. brevirostris, Pantheragombaszoegensis and Felidae indet.). The last accumulation (level10c) is situated at the bottom of the unit EVT7 in gravel andcontains isolated dental remains of hippopotamus and abundantpolyspecific concentrations. These remains and concentrationscontain examples of anatomically connected material and thatgenerally show less interference from carnivores. The lithic arte-facts were recovered from all three accumulations.

This vertical distribution of fossil material, the different degreeof post-depositional alteration of bones and lithic industry, and theidentification of four direct refittings of lithic objects (one of them ismade of two quartz flakes that together form an exploitationsequence, Fig. 2, num. 22) suggest that EVT7 (levels 10 and 10c)could not have been formed during a single catastrophic episode(contra Madurell-Malapeira et al., 2012). Therefore, we believe thatthe sedimentary history of level 10 is more complex than thatproposed by Madurell-Malapeira et al. (2012) in that there werevarious episodes that were subsequently covered by successivemudslides, which would allow that part of the record was inprimary position (Figs. 5 and 6).

Chronological and paleoenvironmental dates

Madurell-Malapeira et al. (2012) argue that the lithic industryfrom units 3 and 4 is clearly out of context, when in reality it wasfound in association with faunal remains. ESR-U/Series analysisobtained from equine teeth and quartz grains yielded ages of0.83 � 0.07 Ma (Martínez et al., 2010) and 0.83 � 0.13 Ma (Duvalet al., 2011) for level 10. Paleomagnetic analysis placed level 10directly above the Jaramillo subchron and the entire stratigraphicsequence between the Jaramillo subchron and Early-Middle Pleis-tocene (Madurell-Malapeira et al., 2010; Martínez et al., 2010;Minwer-Barakat et al., 2011). Biostratigraphic analysis of the

Figure 2. Lithic tools recovered at Vallparadís. Level 10: (1) quartzite anvil, (2) cubic flint core knapped on an anvil bipolar technique, (3) freehand centripetal unifacial flint core, (4)bifacial centripetal quartz core, (5e12) jasper (5), quartzite (6), quartz (7e9) and flint (10e11) denticulate tools, (12 and 13) retouched quartz notches, (14 and 15) flint side-scrapers,(16) quartz distal trihedral or bec, (17e19) distal notches associated with lateral retouches on quartz (17 and 18) and flint (19), (20 and 21) quartz flakes, and (22) direct refitting onquartz flakes. Unit 4: (23 and 24) quartz denticulates.

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575 571

Figure 3. Close-ups of two denticulates on flint recovered from level 10.

Figure 5. Key archaeological section of EVT7 at Vallparadís.

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575572

micrommamals dates level 10 closer to the Sima del Elefante(Martínez et al., 2010) than TD6. According to these data, anapproximate age would be ca. 0.85e0.9 Ma for level 10 and for thewhole stratigraphic sequence an age between 1 Ma and 0.6 Ma.Given that lithic industry has been found in association with faunathroughout the sequence, there is evidence of human occupationthroughout the time period of the site (Martínez et al., 2010; Garciaand Martínez, 2011; Garcia et al., 2011a, b), a finding that is highlysignificant in the context of early continental colonization.

At a biochronological level, the statement by Madurell-Malapeira et al. (2012: 171) that “Iberomys huescarensis (.) hasnever been reported” at Sima del Elefante is incorrect. Cuenca-Bescós et al. (2001) assigned the microtine from Sima del Elefanteto I. huescarensis. The material published from Sima del Elefantematches I. huescarensis samples from the type-locality Huéscar 1and Vallparadís (Martínez et al., 2010). Recently, Cuenca-Bescóset al. (2010) have modified their taxonomic assessment, butwithout providing additional information to support such a change.In any case, it seems clear that regardless of the final taxonomicstatus of the secondmicrotine species at Sima del Elefante, it is very

Figure 4. Tibiae of medium-sized herbivores with

close to the beginning of the Iberomys lineage in the IberianPeninsula and so does not change the biochronological relationshipwith Huéscar 1 and Vallparadís. The lower molars of I. huescarensisfrom Vallparadís show less derived morphology than those fromGran Dolina. This and the absence of other advanced microtine

oblique defleshing cut-marks from level 10.

Figure 6. Vertical distribution of the fossil remains of levels 10 and 10c at Vallparadís. Above: antlers, coprolites, teeth and bones in column V (NeS); middle: coprolites in columnV; below: antlers, coprolites, teeth and bones in line 27 (EeW).

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575 573

species indicate that Vallparadís is older than the Gran Dolina site(Martínez et al., 2010).

Madurell-Malapeira et al. (2012: 172) state that “contrary to thepaleoenvironmental inferences drawn by Agustí et al. (2009), theeffects of the glacial stage 22 (.) were milder in the lower latitudes(.) than in Central Europe.” However, Madurell-Malapeira et al.(2012) seem to have misunderstood the Mutual Climatic Rangemethod (MCR), as presented by Blain et al. (2009) and Agustí et al.(2009). This method compares basic climatic data (Mean AnnualTemperature, Temperature of the Coldest Month, Temperature ofthe Warmest Month, Rainfall) within the same area, and, therefore,detects climatic changes in a restricted geographic zone, not inrelationship with other distant areas. The question as to whetherthe climate conditions were milder in Southern Europe than in

Central Europe is irrelevant here because the MCR method showswhether local conditions were milder or harsher than the presentconditions at exactly the same point. In this way, Agustí et al. (2009)were able to identify a significant climatic deterioration in a sectionof Cal Guardiola, irrespective of what had happened in other areasof Europe. In any case, the climatic deterioration detected by Agustíet al. (2009) at Cal Guardiola is older than the levels with lithicindustry in the Vallparadís section.

Conclusions

Madurell-Malapeira et al. (2012) omit evidence that demon-strates the human occupation of Vallparadís, such as the conclu-sive presence of cut-marks in level 10 (Fig. 4). Furthermore, on the

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575574

basis of interpretations regarding the processes by which thedeposit was formed, they place in doubt the anthropic origin ofthe lithic industry, without paying attention to the technicalattributes that show it to be so. The only lithic artefacts thatMadurell-Malapeira et al. (2012) do not question are thoserecovered from units 3 and 4. They argue that these objects are outof context because they were not recovered during a systematicarchaeological excavation. However, these lithic objects wererecovered from the Middle Pleistocene levels using the samemethodology as that used by the team of paleontologists to extractfossil remains. All of this leads Madurell-Malapeira and colleaguesto conclude that “the currently available data from Vallparadís donot support the presence of early human populations” (Madurell-Malapeira et al., 2012: 172). However, no consistent data havebeen provided that actually refute our hypothesis (Martínez et al.,2010; Garcia et al., 2011a).

On the basis of the diachronic interpretation of the archaeo-logical record at Vallparadís, we believe that the ways of life andtechnology of the first European populations documented at all theSpanish Early Pleistocene sites (e.g., Orce, Sima del Elefante, GranDolina TD6 and Vallparadís) were maintained up to the Matuya-maeBrunhes limit. Or as Madurell-Malapeira et al. (2012: 172) putit: “significant changes in human behavior and paleoecology didnot occur until slightly later in time, probably around the Early-Middle Pleistocene boundary.”

Furthermore, the geochronological and biochronological datafrom level 10 at Vallparadís situate it close to the Jaramillo subchron(0.98 Ma; Martínez et al., 2010), whilst the Iberian sites of Orce,Barranco León and Fuente Nueva 3 are dated to 1.4e1.2 Ma(Martínez-Navarro et al., 1997; Oms et al., 2000; Agustí andMadurell, 2003; Duval et al., 2012), and the Atapuerca sites ofSima del Elefante and Gran Dolina TD6 are dated to 1.22 � 0.16 Maand 0.88e0.78 Ma, respectively (Carbonell et al., 1995, 2008;Moreno-García, 2011). On the basis of this we can therefore statethat these sites in Western Mediterranean Europe were repeatedlyand perhaps continually occupied from 1.4 to 1.2 Ma until the Early-Middle Pleistocene (0.78 Ma), thus closing the chronological gap inthe Late Early Pleistocene of the Iberian Peninsula (Garcia et al.,2011a; contra; Madurell-Malapeira et al., 2012; Duval et al., inpress). However, this hypothesis is controversial because otherauthors maintain that the colonization of Eurasia was discontin-uous and probably interrupted by climatic fluctuations (Dennell,2003, 2010; Agustí et al., 2009; Blain et al., 2009; Moncel, 2010;Martinón-Torres et al., 2011; Bermúdez de Castro and Martinón-Torres, in press). The data available regarding the climatic condi-tions throughout this long period of time (Almogi-Labin, 2011)show certain contradictions in the north-eastern Iberian Peninsulabecause while Vallparadís and Cal Guardiola show constant flux(Agustí et al., 2009), Bòbila Ordis, 100 km to the north and at thesimilar altitude, shows no significant changes in vegetation (Sucand Popescu, 2005).

Given the archaeological similarities between the Spanish sitesfrom the period, we propose that hominins could have continu-ously occupied the Iberian Peninsula throughout the Early Pleis-tocene (Garcia et al., 2011a; Mosquera et al., in press). This ability toadapt to Iberian ecosystems was based on efficient lithic tech-nology and social cohesion, which allowed the hominins to colo-nize the territory and to act as a predator (Carbonell et al., 2010;Rolland, 2010; Saladié et al., 2011).

Acknowledgments

The fieldwork in Vallparadís was financed by the Generalitat deCatalunya (Catalan Government), under the supervision of theDepartment of Culture. This paper has been developed in the

framework of the Research Projects CGL2009-7896 (MICINN) andSGR-324 (Generalitat de Catalunya). We would like to thank Ter-rassa City Council (Ajuntament de Terrassa), the Rovira i VirgiliUniversity Foundation and the archaeological field-work team fortheir collaboration.We are grateful to I. Cáceres, R. Huguet, J. Vilalta,J. Mestre, M. Rovira, and S. Olivares for the stone tool and faunalremains pictures. We also thank the editor (David Begun), theassociate editor and three reviewers for their helpful commentsand help in improving this paper.

References

Agustí, J., Blain, H.-A., Cuenca-Bescós, G., Bailón, S., 2009. Climate forcing the firsthominid dispersal in Western Europe. J. Hum. Evol. 57, 815e821.

Agustí, J., Madurell, J., 2003. Los arvicólidos (Muroidea, Rodentia, Mammalia) delPleistoceno inferior de Barranco León y Fuente Nueva 3 (Orce, Granada).Arqueol. Monogr. 17, 105e114.

Almogi-Labin, A., 2011. The paleoclimate of the Eastern Mediterranean during thetransition from Early to Mid Pleistocene (900 to 700 ka) based on marine andnon-marine records: an integrated overview. J. Hum. Evol. 60, 428e436.

Bar-Yosef, O., Goren-Inbar, N., 1993. The Lithic Assemblages of ‘Ubeidiya: A LowerPalaeolithic Site in the Jordan Valley. Qedem 34, Monographs of the Institute ofArchaeology. The Hebrew University of Jerusalem, Jerusalem.

Bermúdez de Castro, J.M., Martinón-Torres, M. A new model for the evolution of thehuman Pleistocene populations of Europe. Quatern. Int., in press, http://dx.doi.org/10.1016/j.quaint.2012.02.036.

Blain, H.-A., Bailon, S., Cuenca-Bescós, G., Arsuaga, J.L., Bermúdez de Castro, J.M.,Carbonell, E., 2009. Long-term climate record inferred from earlyemiddlePleistocene amphibian and squamate reptile assemblages at the Gran DolinaCave, Atapuerca, Spain. J. Hum. Evol. 56, 55e65.

Cáceres, I., Huguet, R., Bennàsar, M., Vilalta, J., Garcia, J., Martínez, K., Carbonell, E.,2011. Carnivores and Hominins in Early Pleistocene of Vallparadís (Terrassa,Spain). Poster of the Hominid-Carnivore Interactions International Congress,Salou, October 25e28, 2011.

Carbonell, E., Bermúdez de Castro, J.M., Arsuaga, J.L., Díez, J.C., Rosas, A., Cuenca-Bescós, G., Sala, R., Mosquera, M., Rodríguez, X.P., 1995. Lower Pleistocenehominins and artifacts from Atapuerca-TD6 (Spain). Science 269, 826e830.

Carbonell, E., Bermúdez de Castro, J.M., Parés, J.M., Pérez, G.A., Cuenca-Bescós, G.,Ollé, A., Mosquera, M., Huguet, R., van der Made, J., Rosas, A., Sala, R.,Vallverdú, J., García, N., Granger, D.E., Martinón-Torres, M., Rodríguez, X.P.,Stock, G.M., Vergès, J.M., Allué, E., Burjachs, F., Cáceres, I., Canals, A., Benito, A.,Díez, C., Lozano, M., Mateos, A., Navazo, M., Rodríguez, J., Rosell, J., Arsuaga, J.L.,2008. The first hominin of Europe. Nature 452, 465e469.

Carbonell, E., García-Antón, M.D., Mallol, C., Mosquera, M., Ollé, A., Rodríguez, X.P.,Sahnouni, M., Sala, R., Vergès, J.M., 1999. The TD6 level lithic industry from GranDolina, Atapuerca (Burgos, Spain): production and use. J. Hum. Evol. 37,653e693.

Carbonell, E., Sala, R., Rodríguez, X.P., Mosquera, M., Ollé, A., Vergès, J.M., Martínez-Navarro, B., Bermúdez de Castro, J.M., 2010. Early hominid dispersals: a tech-nological hypothesis for ‘out of Africa’. Quatern. Int. 223e224, 36e44.

Cuenca-Bescós, G., Canudo, J.I., Laplana, C., 2001. La séquence des rongeurs(Mammalia) des sites du Pléistocène inférieur et moyen d’Atapuerca (Burgos,Espagne). L’Anthropologie 105, 115e130.

Cuenca-Bescós, G., Rofes, J., López-García, J.M., Blain, H.-A., de Marfá, J.R., Galindo-Pellicena, M.A., Bennàsar-Serra, M.L., Melero-Rubio, M., Arsuaga, J.L., Bermúdezde Castro, J.M., Carbonell, E., 2010. Biochronology of Spanish Quaternary smallvertebrate faunas. Quatern. Int. 212, 109e119.

de Lumley, H., Nioradzé, M., Barsky, D., Cauche, D., Celiberti, V., Nioradzé, G.,Notter, O., Zvania, D., Lordkipanidze, D., 2005. Les industries lithiques pré-oldowayennes du début du Pléistocène inférieur du site de Dmanissi enGéorgie. L’Anthropologie 109, 1e182.

Dennell, R.W., 2003. Dispersal and colonisation, long and short chronologies: howcontinuous is the Early Pleistocene record for hominins outside East Africa?J. Hum. Evol. 45, 421e440.

Dennell, R.W., 2010. The colonization of ‘Savannahstan’: issues of timing(s) andpatterns of dispersal across Asia in the Late Pliocene and Early Pleistocene. In:Norton, C.J., Braun, D.R. (Eds.), Asian Paleoanthropology: From Africa to Chinaand Beyond. Vertebrate Paleobiology and Paleoanthropology. Springer, Dor-drecht, pp. 7e30.

Despriée, J., Gageonnet, R., Voinchet, P., Bahain, J.-J., Falguères, C., Varache, F.,Courcimault, G., Dolo, J.M., 2006. Une occupation humaine au Pléistocèneinférieur sur la bordure nord du Massif central, vol. 5. C.R. Palevol, pp. 821e828.

Duval, M., Falguères, C., Bahain, J.-J. Age of the oldest hominin settlements in Spain:contribution of the combined U-series/ESR dating method applied to fossilteeth. Quatern. Geochron., in press, http://dx.doi.org/10.1016/j.quageo.2012.02.025.

Duval, M., Falguères, C., Bahain, J.-J., Grün, R., Shao, Q., Aubert, M., Dolo, J.-M.,Agustí, J., Martínez-Navarro, B., Palmqvist, P., Toro-Moyano, I., 2012. On thelimits of using combined U-series/ESR method to date fossil teeth from twoEarly Pleistocene archaeological sites of the Orce area (Guadix-Baza basin,Spain). Quatern. Res. 77, 482e491.

J. Garcia et al. / Journal of Human Evolution 63 (2012) 568e575 575

Duval, M., Moreno, D., Shao, Q., Voinchet, P., Falguères, C., Bahain, J.-J., García, T.,Garcia, J., Martínez, K., 2011. Datación por ESR del yacimiento arqueológico delPleistoceno inferior de Vallparadís (Terrassa, Cataluña, España). Trab. Preh. 68,7e24.

Espigares, M.P., 2010. Análisis y modelización del contexto sedimentario y losatributos tafonómicos de los yacimientos pleistocénicos del borde Nororientalde la Cuenca de Guádix-Baza. Ph.D. Dissertation, Universidad de Granada.

Faith, J.T., Marean, C.W., Behrensmeyer, A.K., 2007. Carnivore competition, bonedestruction, and bone density. J. Archaeol. Sci. 34, 2025e2034.

Garcia, J., Martínez, K., 2011. Lower Pleistocene hominid settlement in northeasternSpain: Vallparadís (Barcelona). Abstracts of the Paleoanthropology Society 2011Meeting, Minneapolis, April 12e13, 2011. PaleoAnthropology, p. A12, http://dx.doi.org/10.4207/PA.2011.ABS9.

Garcia, J., Martínez, K., Carbonell, E., 2011a. Continuity of the first human occupationin the Iberian Peninsula: Closing the archaeological gap. C.R. Palevol. 10,279e284.

Garcia, J., Martínez, K., Carbonell, E., 2011b. Human occupation and paleoecologyduring late Lower Pleistocene in north-eastern Spain: the key sites of Vallpar-adís (Barcelona) and Bòbila Ordis (Girona). 1st Annual Meeting for the EuropeanSociety for the Study of Human Evolution, Leipzig, September 23e24, 2011.Abstracts ESHE, p. 40.

Gaudzinski, S., 2004. Subsistence patterns of Early Pleistocene hominins in theLevant: taphonomic evidence from the Ubeidiya Formation (Israel). J. Archaeol.Sci. 31, 65e75.

Huguet, R., 2007. Primeras ocupaciones humanas en la Península Ibérica: paleo-economía en la Sierra de Atapuerca (Burgos) y en la Cuenca de Guadix-Baza(Granada) durante el Pleistoceno Inferior. Ph.D. Dissertation, UniversitatRovira i Virgili.

Lordkipanidze, D., Jashashvili, T., Vekua, A., Ponce de León, M.S., Zollikofer, Ch.P.E.,Rightmire, G.Ph., Pontzer, H., Ferring, R., Oms, O., Tappen, M., Bukhsianidze, M.,Agustí, J., Kahlke, R., Kiladze, G., Martínez-Navarro, B., Mouskhelishvili, A.,Nioradze, M., Rook, L., 2007. Postcranial evidence from early Homo fromDmanisi, Georgia. Nature 449, 305e310.

Madurell-Malapeira, J., Alba, D.M., Minewr-Barakat, R., Aurell-Garrido, J., 2012. Earlyhuman dispersals into the Iberian Peninsula: a comment on Martínez et al.(2010) and Garcia, et al. (2011). J. Hum. Evol. 62, 169e173.

Madurell-Malapeira, J., Minwer-Barakat, R., Alba, D.M., Garcés, M., Gómez, M.,Aurell-Garrido, J., Ros-Montoya, S., Moyà-Solà, S., Berástegui, X., 2010. TheVallparadís section (Terrassa, Iberian Peninsula) and the latest Villafranchianfaunas of Europe. Quatern. Sci. Rev. 29, 2972e2982.

Martínez, K., Garcia, J., Carbonell, E., Agustí, J., Bahain, J.-J., Blain, H.-A., Burjachs, F.,Cáceres, I., Duval, M., Falguères, C., Gómez, M., Huguet, R., 2010. A new lowerPleistocene archeological site in Europe (Vallparadís, Barcelona, Spain). Proc.Natl. Acad. Sci. 107, 5762e5767.

Martínez-Navarro, B., Turq, A., Agustí, J., Oms, O., 1997. Fuente Nueva-3 (Orce,Granada, Spain) and the first human occupation of Europe. J. Hum. Evol. 33,611e620.

Martinón-Torres, M., Martín-Francés, L., Gracia, A., Olejniczak, A., Prado-Simón, L.,Gómez-Robles, A., Lapresa,M., Carbonell, E., Arsuaga, J.L., Bermúdez de Castro, J.M.,2011. Early Pleistocene human mandible from Sima del Elefante (TE) cave site inSierra de Atapuerca (Spain): a palaeopathological study. J. Hum. Evol. 61, 1e11.

Minwer-Barakat, R., Madurell-Malapeira, J., Alba, D.M., Aurell-Garrido, J., Moyà-Solà, S., 2011. Pleistocene rodents from the Torrent de Vallparadís section(Northeastern Spain): biochronological implications. J. Vert. Paleontol. 31,849e865.

Moncel, M.-H., 2010. Oldest human expansions in Eurasia: favouring and limitingfactors. Quatern. Int. 223e224, 1e9.

Moreno-García, D., 2011. Datation par ESR de quartz optiquement blanchis (ESR-OB)de la région de Atapuerca (Burgos, Espagne). Application au site préhistoriquede Gran Dolina (contexte karstique) et aux systèmes fluviatiles Quaternaires del’Arlanzón et l’Arlanza. Ph.D. Dissertation, Universitat Rovira i Virgili.

Mosquera, M., Ollé, A., Rodríguez, X.P. From Atapuerca to Europe: tracing theearliest peopling of Europe. Quatern. Int., in press, http://dx.doi.org/10.1016/j.quaint.2012.01.031.

Oms, O., Parés, J.M., Martínez-Navarro, B., Agustí, J., Toro, I., Martínez-Fernández, G.,Turq, A., 2000. Early human occupation of western Europe: paleomagneticdates for two Paleolithic sites in Spain. Proc. Natl. Acad. Sci. 97, 10666e10670.

Rolland, N., 2010. The earliest hominid dispersals beyond Subsaharan Africa:a survey of underlying causes. Quatern. Int. 223e224, 54e64.

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 deci-sions in Homo antecessor subsistence strategies. J. Hum. Evol. 61, 425e446.

Suc, J.P., Popescu, S.M., 2005. Pollen records and climatic cycles in the NorthMediterranean region since 2.7 Ma. In: Head, M.J., Gibbard, M. (Eds.), Early-Middle Pleistocene Transitions: The Land-Ocean Evidence. Geological Society,London, p. 247.

Toro-Moyano, I., de Lumley, H., Fajardo, B., Barsky, D., Cauche, D., Celiberti, V.,Grégoire, S., Martínez-Navarro, B., Espigares, M.P., Ros-Montoya, S., 2009. L’in-dustrie lithique des gisements du Pléistocène inférieur de Barranco León etFuente Nueva 3 à Orce, Grenade, Espagne. L’Anthropologie 113, 111e124.