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Symbolic use of marine shells and mineral pigmentsby Iberian NeandertalsJoo Zilhoa,1, Diego E. Angeluccib, Ernestina Badal-Garcac, Francesco dErricod,e, Floral Danielf, Laure Dayetf,Katerina Doukag, Thomas F. G. Highamg, Mara Jos Martnez-Snchezh, Ricardo Montes-Bernrdezi,Sonia Murcia-Mascarsj, Carmen Prez-Sirventh, Clodoaldo Roldn-Garcaj, Marian Vanhaerenk,

Valentn Villaverdec

, Rachel Woodg

, and Josefi

na Zapatal

aUniversity of Bristol, Department of Archaeology and Anthropology, Bristol BS8 1UU, United Kingdom; bUniversit degli Studi di Trento, Laboratorio diPreistoria B. Bagolini, Dipartimento di Filosofia, Storia e Beni Culturali, 38122 Trento, Italy; cUniversidad de Valencia, Departamento de Prehistoria yArqueologa, 46010 Valencia, Spain; dCentre National de la Recherche Scientifique, Unit Mixte de Recherche 5199, De la Prhistoire lActuel: Culture,Environnement et Anthropologie, 33405 Talence, France; eUniversity of the Witwatersrand, Institute for Human Evolution, Johannesburg, 2050 Wits, SouthAfrica; fUniversit de Bordeaux 3, Centre National de la Recherche Scientifique, Unit Mixte de Recherche 5060, Institut de Recherche sur lesArchomatriaux, Centre de recherche en physique applique l archologie, 33607 Pessac, France; gUniversity of Oxford, Research Laboratory forArchaeology and the History of Art, Dyson Perrins Building, Oxford OX1 3QY, United Kingdom; hUniversidad de Murcia, Departamento de Qumica Agrcola,Geologa y Edafologa, Facultad de Qumica, Campus de Espinardo, 30100 Murcia, Spain; iFundacin de Estudios Murcianos Marqus de Corvera, 30566 LasTorres de Cotillas (Murcia), Spain; jUniversidad de Valencia, Instituto de Ciencia de los Materiales, 46071 Valencia, Spain; kCentre National de la RechercheScientifique, Unit Mixte de Recherche 7041, Archologies et Sciences de l Antiquit, 92023 Nanterre, France; and lUniversidad de Murcia, rea deAntropologa Fsica, Facultad de Biologa, Campus de Espinardo, 30100 Murcia, Spain

Communicated by Erik Trinkaus, Washington University, St. Louis, MO, December 5, 2009 (received for review October 16, 2009)

Two sites of the Neandertal-associated Middle Paleolithic of Iberia,

dated to as early as approximately 50,000 years ago, yielded

perforated and pigment-stained marine shells. At Cueva de losAviones, three umbo-perforated valves of Acanthocardia and Gly-

cymeriswere found alongside lumps of yellow and red colorants,

and residues preserved inside a Spondylus shell consist of a red

lepidocrocite base mixed with ground, dark red-to-black fragments

of hematite and pyrite. A perforated Pecten shell, painted on itsexternal, white side with an orange mix of goethite and hematite,

wasabandonedafter breakage at Cueva Antn,60 km inland. Com-

parable early modern human-associated material from Africa and

the Near East is widely accepted as evidence for body ornamenta-

tion, implying behavioral modernity. The Iberian finds show that

European Neandertalswere no differentfrom coevalAfricansin thisregard, countering genetic/cognitive explanations for the emer-

gence of symbolism and strengthening demographic/social ones.

behavioral modernity | Iberia | Middle Paleolithic | shell ornaments |symbolism

Items of body decorationperforated and pigment-stainedshells of marine mollusksare a feature of the Middle StoneAge (MSA) of southern Africa and the Middle Paleolithic (MP)of the Maghreb and Near East (1, 2). Dated to 70120 ka cal-endar (cal) B.P., these finds are widely accepted as evidence ofsymbolic thinking among the earliest anatomically modernhumans of Africa and adjacent regions of Southwest Asia (3).

An absence of similar finds from the coeval archeologicalrecord of Europe has supported the notion that Neandertals weredevoid of symbolism, their cognitive inferiority explaining whymodern humans dispersing out of Africa eventuallyreplaced them

across the entire continent approximately 40 ka cal B.P. (4, 5). Inthis view, claimed instances of Neandertal symbolism are dis-regarded on different groundse.g., as coming from old exca-

vations and too ill-recorded to be certain or of ambiguousinterpretation and in any case too scant to be meaningful. A casein point is the Mousterian cemetery of La Ferrassie (6), for whichregional culture-stratigraphy indicates an age of approximately6570 ka cal B.P. (7) and where the burial pit of an adult malecontained a bone fragment decorated with four sets of parallelincisions, whereas that of a 3- to 5-year-old child was covered by acupule-decorated limestone slab.

In this context, the presence of body ornaments (namely,pierced and grooved animal teeth) in Neandertal-associatedarcheological cultures (such as the Chtelperronian of France)has been variously explained by stratigraphic mixing, accultur-

ation, imitation without understanding, or independent Nean-dertal innovation (814). As this evidence comes from near the

time of contact with modern humans in Europe, unresolved issuesof dating and taphonomy impacting the broader paleoanthropo-logical problem explain the persistence of the debate (1518).

Here,wereportsecureevidencethat,approximately50kacalB.P., 10 millennia before modern humans are first recorded inEurope, the behavior of Neandertals was symbolically organizedand continued to be so until the very end of their evolutionarytrajectory. This conclusion is based on the same types offinds andcriteria used to assess the African and Near Eastern evidencepigment-stained, perforated marine shells and the associatedprocurement, processing, and application of colorants for bodyornamentation. The evidence comes from two sites in the Murciaprovince of southeast Spain, Cueva de los Aviones (37357.30N,0598.66W) and Cueva Antn (38351.84N; 12947.20W) (SIAppendix, Sections I and II).

Results

The Sites. Aviones is a large cave whose fill has for the most partbeen eroded away by postglacial marine erosion (SI Appendix,Section I, Figs. S1S4). At the time of occupation, the Medi-terranean was 5090 m lower (19), implying a distance of 1.57.0km between site and shore (ormore, because the Murcia coast is asubsidence area). A brecciated baulk preserved against thenorthwest wall of the cave was excavated in 1985 (20, 21),revealing an MP stratigraphy resting on an Eemian rock beachand made up of slope deposits of continental origin interspersed

with flowstones and carbonate crusts (SI Appendix, Section I,Tables S1 and S2). The archeological levels (IV), separated fromthe rock beach by a thick, sterile deposit (level VI), date to the

approximately 45

50 ka cal B.P. interval (SI Appendix, Section III,Table S4, Fig. S13). They yielded abundant lithic assemblagesprimarily made of quartz but with a significant component offlintsidescrapers and points (SI Appendix, Section I, Fig. S5)in

Author contributions: J. Zilho and J. Zapata designed research; J. Zilho, D.E.A., E.B.-G.,F.d., F.D., L.D., K.D., T.F.G.H., M.J.M.-S., R.M.-B., S.M.-M., C.P.-S., C.R.-G., M.V., V.V., R.W.,and J. Zapata performed research; J. Zilho, D.E.A., E.B.-G., F.d., F.D., L.D., K.D., T.F.G.H.,M.J.M.-S., S.M.-M., C.P.-S., C.R.-G., M.V., V.V., R.W., and J. Zapata analyzed data; andJ. Zilho, D.E.A., E.B.-G., F.d., F.D., L.D., K.D., T.F.G.H., M.J.M.-S., S.M.-M., C.P.-S., C.R.-G.,M.V., V.V., R.W., and J. Zapata wrote the paper.

The authors declare no conflict of interest.

1To whom correspondence should be addressed. E-mail: [email protected].

This article contains supporting information online at www.pnas.org/cgi/content/full/0914088107/DCSupplemental.

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association with marine mollusk shells and bone remains of horse,deer, ibex, rabbit, and tortoise.

Antn (22) is a large rockshelter excavated by the Mula river,which runs through an interior basin located approximately 60 kmfrom the present shore line. The MP occupations are preserved ina more than 4-m-thick sequence of low-energy flood depositsoverlain by silts accumulated during recent episodes of inundationby a dam reservoir (SI Appendix, Section II, Table S3, Figs. S11and S12). The levels are artifact-rich andorganized around hearth

features in thelower part of thesequence, but occupation remainsare scarce toward the top, reflecting sporadic and ephemeralincursions. The uppermost fertile units are levels II-l and I-k, for

which the ages indicated by radiocarbon dating are, respectively,greater than 43.5 and approximately 37.4 ka cal B.P. (SI Appendix,Section III, Table S5, Figs. S13 and S14).

The Aviones Finds. Three categories of species are represented inthe Aviones marine shell assemblage (Table 1). Most (95.7%) areof edible intertidal genera (Cerastoderma, Monodonta, Mytilus,and Patella) and have well preserved surfaces, indicating collec-tion of living animals. A residual proportion (0.2%) are acci-dentally introducedNassarius incrassatus and Gibbula sp.As thesetaxa live or feed on algae, their presence is likely to reflectpackaging of the edible species in water-soaked algae, a tradi-tional, preindustrial technique of maintaining harvested mollusksalive (they rot very rapidly and must be eaten or cooked extremelyfresh) during transportation from the point of collection to thepoint of consumption. Indeed, remains of Jania rubens var. cor-niculata, an intertidal epiphytic alga whose calcareous articulatedfrond allows preservation and recognition in carbonated cave fills,

were recovered in levels I and II (20, 21).The remaining 4.0% are infra- and circalittoral genera with

variably abraded and bioeroded surfaces, reflecting beach collec-tionoftheshellsofdeadanimals( SI Appendix, Section I, Figs. S8S10). The Chama, Laevicardium, Charonia, and Thais material istoo broken to assess original condition, and the one Truncular-iopsis trunculus is a complete specimen with an irregular, probablypostdepositional perforation of the body whorl in an area where

the shell is very thin.Of the three Acanthocardia tuberculata, one is complete,another bears an excavation break with loss of the umbo area,the third is a conjoin of two fragments with an ancient, post-depositional break, and all must have been collected and intro-duced to the site as whole valves. The smaller, from level II, isumbo-perforated (Fig. 1, 1).

Of the Glycymeris insubrica, four are complete, ten are ventralmargin fragments, and four are small fragments of the middle ofthe shell; although excavation breaks are apparent in a few, mostfeature ancient breaks only. Two complete specimens from levelII are umbo-perforated (Fig. 1, 23), and residues of a red col-orant, identified as hematite (SI Appendix, Section IV Figs. S15and S16), were found while cleaning the carbonate coatingaround the perforation of the larger shell. As argued for the

comparable modern human-associated material from NearEastern sites of the MP and early Upper Paleolithic (UP)(Qafzeh, Ksar Akil, aizli), the parsimonious interpretationofGlycymeris shells, even in the absence of pigment residues andirrespective of the origin of the perforation, is that they arepersonal ornaments (23, 24).

The three Spondylus gaederopus shells were originally complete.The inner side of a level II upper valve (Fig. 2) bears residues of apigmentatious mass composed of a red lepidocrocite base mixed

with ground particles of charcoal, dolomite, hematite, and pyrite(SI Appendix, Section IV, Table S6, Figs. S17S19), suggesting useas a container for the storage of colorants or as a kind of paint cupfor their preparation. Although examination of their surfaces forpigment residues is hindered by extensive carbonate coating, asimilar function can be envisaged forthe other valves ofSpondylus

as well as forthose of taxa with a similar concave morphologyanexcavation-broken Callista chione and two lower valve fragmentsofPecten maximus (SI Appendix, Section I, Figs. S9 and S10).

Orange pigment was also observed on the tip of an unmodifiedancillary metatarsal ofEquus sp. fromlevel III, but its compositioncould not be established, due to the extensive carbonate coatingand attendant high fluorescence of the specimen (Fig. 3; SIAppendix, Section IV, Fig. S20). This naturally pointed bone mayhave been used as a stiletto for the preparation or application ofmineral dyes or as a pin or awl to perforate soft materials (e.g.,hides) that were themselves colored with such dyes.

Lumps of red and yellow colorants were noted and sampled at

thetimeof excavation (25) (SI Appendix, Section V, Table S7, Figs.S21S23). X-ray diffraction showed the reddish material to besiderite, goethite, hematite, and nontronite, and the yellow to besiderite and natrojarosite. These pigments can only be manuportsbecause both thecave and the catchment of its sedimentaryfill arein Mesozoic limestone and dolomite bedrock, where the identi-fied minerals either do not exist or do not occur as masses ofmacroscopic size (in the region, such masses formed as a result of

volcanism and hydrothermalism processes of Neogene age; SIAppendix, Section V).

Sources for the red colorants can be found 35 km to thenorthwest, in the mining district of La Unin, exploited for gold,silver and base metals since antiquity. For the yellow natrojar-osite, the purity of the level III samplea large (23 g) and

homogeneous accumulation of prisms, granules, and powder,probably the contents of a small purse made of perishable mate-rialconstrains potential sources to a limited number of local-ities, theclosest lying approximately 7 km to theeast (SI Appendix,Section V, Figs. S22 and S24).

The Antn Finds. Level I-k of Cueva Antn, which caps the MParcheological succession and is directly overlain by the reservoirinundation silts, yielded half of an upper, flat valve ofP. maximusfeaturing a circular perforation approximately 6 mm in diameter(Fig. 4). The shells numerous sponge holes indicate that it wasbeached after laying on the sea bottom for a significant amount oftime. Whether the circular perforation relates to the same bio-erosion processes (i.e., is natural) or was made after collection (i.e., is anthropogenic) cannot be assessed due to postdepositional

Table 1. Cueva de los Aviones mollusks: taxa and provenience

I II III IV V Total

Food taxa

Cerastoderma edule 8 1 5 14Monodonta turbinata 12 14 234 151 16 427Mytilus edulis 36 11 31 30 108Patella sp.* 10 61 96 69 236

Nonfood taxa

Acanthocardia tuberculata 1 1 1 3Callista chione 1 1Chama gryphoides 1 1Charonia lampas 1 1Gibbula cf varia 1 1Glycymeris insubrica 10 6 2 18Laevicardium oblongum 1 1Nassarius incrassatus 1 1Pecten maximus 2 2Spondylus gaederopus 1 1 1 3Thais hemastoma 1 1Trunculariopsis trunculus 1 1

Unclassified bivalve 1 1Total 72 103 368 260 17 820

*Mostly P. ferruginea; a few are P. aspera and P. lusitanica

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degradation of the holes margins (e.g., by exfoliation and rubbingagainst the angular clast matrix), which caused the extant sharp-nessof its paper-thin edges. But we can exclude a postdepositionalorigin for the perforation itself because the presence of orangepigment residues on the interior surfaces of the exposed structuralhollows of the shell implies that the perforation preexisted theapplication of the colorant.

The pigmenta mix of yellow goethite and red hematite forwhich potential sources exist within approximately 5 km from thesiteis ubiquitously preserved but on the shell s external, dis-colored side only, suggesting that it may have been deliberatelypainted, either to regain the original appearance or to make it thesame color as the internal side, which remained its natural red (SIAppendix, Section VI, Figs. S25 and S26). Theephemeral nature ofthe occupation of level I-k argues against on-site tool productionor tool maintenance tasks and strengthens the case for theinterpretation of this shell as an item of body decoration. The

alternativesuse as a container or palettecould hold for alower valve color-stained on the inner, concave surface or if theperforation was postdepositional, but neither is the case.

Comparative Study. It has been argued that symbolic inter-pretations of the African and Near Eastern MSA/MP marine shellbeads are questionable because the anthropogenic nature of theirperforations is uncertain, whereas those in the marine shell beadsfound among modern human-associated cultures of the AfricanLate Stone Age (LSA) and the European UP are clearly man-

made (26). By extension, this argument applies to the materialpresented here too, but is it valid? To assess this issue, we col-lected from regional beach thanatocenoses a modern referencesample of shells of the bivalve genera that are most common inornamental assemblages of prehistoric IberiaAcanthocardia,Cerastoderma, and Glycymeris (SI Dataset and SI Appendix, Sec-tion VII, Tables S8S10, Figs. S27S30). We then compared the

Fig. 1. The perforated shells from level II of Cueva de los Aviones (after cleaning): (1) Acanthocardia tuberculata; (23). Glycymeris insubrica (maximuminternal diameter of the perforations: 4.2, 9.5, and 6.8 mm, respectively). See also SI Appendix, Sections I and IV, Figs. S6, S7, S15, and S16.

Fig. 2. (Left) Excavation-broken upper valve of Spondylus gaederopus from Cueva de los Aviones (the pigment residue is indicated); (Right) close-up view ofthe pigment residue. See also SI Appendix, Section IV, Figs. S17S19.

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Murcia MP specimens with that collection and with (i) the earlymodern human-associated MP material from the Near East, and(ii) the relevant components of ornament assemblages from theUP and the Neolithic of Mediterranean Spain (SI Appendix,Section VIII, Table S11, Figs. S31S42).

In the natural collection, umbo-perforated shells occur fre-quently (45.4%) in the case ofGlycymeris, less so in the case ofAcanthocardia (2.7%). Both the occurrence and the size of theperforations correlate positively with discrete indicators of

weathering such as decalcification/exfoliation, sponge holes, andabrasion (Table 2). Thus, in the absence of production tool-marks, perforations can be presumed anthropogenic only if theumbo holes are large but the shell is fresh or only lightly

weathered. Conversely, as umbo perforations occur in 90% ofthe modern Glycymeris that feature advanced weathering, anatural origin must be presumed for archeological Glycymeris insimilar condition. Based on these uniformitarian criteria, theperforations in the MP Glycymeris from Qafzeh and Aviones, as

well as in those from the UP and the Early Neolithic of Medi-terranean Spain that we examined, are natural, and so too arethose in all of the Acanthocardia and Cerastoderma shells fromthe same sites.

The archeological Glycymeris, however, were not gatheredcompletely at random. When our three collections (MP, UP, andNeolithic) are considered together, a pattern of selection basedon perforation size emergesirrespective of height, shells withholes in the 4.56.5 mm interval were targeted (SI Appendix,Section VIII, Fig. S31). These observations suggest that the pat-tern is cross-cultural and determined by threading constraints,

with color, shine, and overall surface appearance being of sec-ondary importanceexplaining why all of this material featuresadvanced weathering, despite lightly weathered shells (that pre-

serve more of theoriginal color butlack a perforation or only havea very small one) being rather easy to find in nature too.Although the perforations on the Nassarius kraussianus from

theMSA of Blombos are anthropogenic (1), only oneout of the41Nassarius gibbosulus beads from Skhul and the North African MPsites featured striationsmade by a stone toolmostare weatheredspecimens with types of holes whose size and placement on theshell can also be found in natural thanatocenoses, albeit in dif-ferent frequencies, indicating selection but not necessarily mod-ification. The use wear, however, is distinctive, and shows they

were used as beads even when their perforations are natural (2).In the Murcia MP shellsas is the case, due to the high water

content of depositional environments, in most ornamental shellassemblages of the European Paleolithic (27)chemical micro-

weathering (SI Appendix,Section I, Fig.S7)wouldhavelongerased

any perforation-associated use wear evidence. Shallow notchesobserved on the internal contour of the perforations of two Gly-cymeris from Qafzeh, where they were interpreted as caused by

friction from a string (23), are also apparent in the Avionesspecimens (Fig. 1 and SI Appendix, Section I, Fig. S7). However,because similar notches exist in naturally umbo-perforated mate-rial (SI Appendix, Section VII, Fig. S28), this evidence is incon-clusive. Such irregular internal contours may be caused bycontinued abrasion in sandy, high-energy burial environments,eventually eliminating the shell margin side of the contour andcreating an open hole perforation. Although not uncommon inshells from natural assemblages (SI Appendix, Section VII, TableS10), we found no such open holes among the archeologicalspecimensas should be expected, given their unsuitability forsuspension, and strengthening a link between perforation andsuspension even when preservation factors prevent assessment ofuse wear.

The origin of perforation argument is therefore not valid.

Our findings suggest that naturally perforated marine shells wereextensively used at the time of emergence of the European MPand African MP/MSA traditions of body decoration and con-tinued to be very much in fashion even as the repertoire ofornament types expanded to include more elaborated items.

Discussion

Natural surface processes and animal behavior can explain inlandmarine shell accumulations but,as discussed for the South Africanand Near Eastern beads (2,23), distance to shore and difference inelevation preclude involvement of such agents in the Murcia sites.The use of the robustshellofC. chione as a replacement for flint isdocumented in the coastal MP of Italy (28), and the taxon is rep-resented at Aviones by one specimen (SI Appendix, Section I, Fig.

S10). Conceivably, this sites thicker ventral marginandmiddle-of-the-shell Glycymeris fragments (SI Appendix, Section I, Fig. S8)

could represent broken-and-discarded instances of such tools andthe complete specimens unmodified blanks for their manufacture.However, neitheruse wear nor any kind of retouch, even incipient,

were observed in the specimens intact edges, so the parsimoniousinterpretation of this Glycymeris material is that it reflects post-depositional breakage or on-site discard of broken shells that werebeach-collected and used as complete valvesfor ornamentation,

when perforated, perhaps as containers, when not.Although, at Aviones, mineral colorants were present in the

deposits, the residues observed on the finds discussed here are notincidental or postdepositional contamination. If so, one wouldexpect pigment to be randomly distributed across the differentfind categories. This expectation was not met for a sample of 50

Fig. 3. Different views of an ancillary metatarsal of horse with an excavation break from Cueva de los Aviones (Left) and binocular microscope close-ups of itspigment-dotted tip (Centerand Right). See also SI Appendix, Section IV, Fig. S20.

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shells of different species examined with a 10 hand magnifierafter cleaning in an ultrasound vat (examination of the totality ofthe assemblage was not possible due to the extensive carbonatecoating of the surfaces): no pigment was found on the food shells,and, of the nonfood ones, only the one Spondylus discussed above(in addition to one of the two perforated Glycymeris) had any.Moreover, all of the lumps of red pigment sampled during exca-

vation feature in their composition siderite, a mineral that was notidentified in the pigment residues of either the Spondylus or theGlycymeris (SI Appendix, Sections IV and V, Tables S6 and S7,Figs. S16S19, S22, and S23); therefore, these residues cannot beaccidentally derived from pigments present in the shells imme-diate burial environment.

Ithasbeenarguedthatochredoesnot necessarily equate withsymbolism, as its manipulation may relate to technology, e.g., thepreparationof mastics for hafting (29), and we cannot excludethatthe Aviones pigments are in part related to such tasks. However,no pigmentresidues were observed in the Avionesstone tools (20),andno tool production or tool maintenancetasks were carried outat Antn, so suchactivities cannot provide a functional context forthe pigment-stained shells found at both sites.

Moreover, the limited capacity (approximately 5 cm3) ofS. gae-deropus upper valves argues against their use for the processing orstorageofmaterialrelatedtoroutinetoolingactivities.Theadditionto the pigmentatious mass of ground bits of hematite and pyrite(which, when fresh, have a brilliant black, reflective appearance) isalso inconsistent with that function,and suggests instead thekind ofinclusion for effect that one would expect in a cosmetic prepa-ration.TheuseofSpondylus shellsforsuchaparticularpurposemayrelate to their attention-grabbing crimson, red, or violet color andexuberant sculpture, which have led to their symbolic- or ritual-related collection in a variety of archeological contexts worldwide,namely Neolithic Europe and pre-Columbian America (30, 31).

These inferences are consistent with thefact that, in therecordedhistory of pigments, all known uses (in Ancient Egypt) of yellowminerals of the jarosite family are in cosmetics or painting (namely,for the rendition of female skin) (32). Decorationof body, cloth-ing, living space, or artistic representations thereof (33)is also theonly archeologically and ethnographically documented function ofcolored,umbo-perforated Glycymeris andAcanthocardia shells.The

well known use ofP. maximus valves underlying the species com-monname (pilgrim shell) is the same, andthe Antn specimenmaybe exceptional for the MP only in its degree of preservation, as P.maximus fragments were also found in level III of Aviones (Table 1)andlevel V of Higueralde Valleja (Cdiz) (34), an Andaluciancavelocated almost as far inland (more than 50 km) as Antn.

Conclusion

The Pecten from Antn and Higueral de Valleja relate to the veryend of the period of late persistence of Neandertals in Iberianregions to the south of the Ebro drainage (in the Andalucian site,the find comes from undated deposits capping a thick MPsequence, a chronostratigraphic position akin to Antns andsuggesting a similar age) (35). Therefore, they cannot be disen-tangled from the controversies surrounding the interpretation ofcontact period ornaments found elsewhere in Europe (818).The Aviones material, however, comes from the approximately 50ka cal B.P. brecciated remnant of a site fill entirely of MP age and

where the notion of later intrusions is an oxymoronthe associ-ation of this material with the Neandertals is, literally, rock-solid.

Thesymbolicimplicationsofbodypaintingandoftheornamentaluse of pigment-stained and perforated marine shells are uncon-

troversial in UP and later prehistoric contexts but, as shown by theevidence from Africa, the Near East andnowIberia, both behaviorsfirst occur in the MP/MSA. Their emergence in two continents,among two different lineages and, in the time scale of human evo-lution, at about the sametime, is inconsistentwithcognitive-geneticexplanations and implies that these innovations were fulfilling aneedaidinginthepersonalorsocialidentificationofpeoplethatdid not exist in the preceding two million years of human evolution.

Our findings therefore support models of the emergence ofbehavioral modernity as caused by technological progress, dem-ographic increase,and social complexification and show that thereis no biunivocal correlation between modern anatomy andmodern behavior (13, 3638). This much is also implied by theNorth African beads, which are from a time when the region wasinhabited by the Dar-es-Soltan people, who, anatomically, were

Fig. 4. K19-3, a perforated upper half-valve of Pecten maximus from level I-k at Cueva Antn (height: 120 mm). (Upper) The internal, naturally red side(Left) and the external, whitish side that was painted with an orange colo-rant made of goethite and hematite (Right). (Lower) binocular microscopeclose-ups on pigment masses and pigment stains around the perforation. Seealso SI Appendix, Section VI, Figs. S25 and S26.

Table 2. Modern reference collection: perforation and

weathering

Perforated Nonperforated

Cardiidae (n = 55) (n = 770)Exfoliation 50.9% 9.7%Sponge through-holes 3.6% 0.6%

Glycymeris (n = 132) (n = 150)Weathering stages 23 99.2% 50.7%

Zilho et al. PNAS Early Edition | 5 of 6

ANTHROPOLO

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nonmodern (39, 40). Where the personal ornaments of the Ch-telperronian and coeval transitional cultures of Europe areconcerned, a final corollary of our results is that Neandertalauthorship is the null hypothesis.

Materials and Methods

Archeological excavation was by natural stratigraphic units and used 3D piece-plotting.Descriptionofthereferenceprofilesfollowedgeoarcheologicalcriteria,accounting for stratigraphic, sedimentary, and diagenetic features to differ-

entiate units and group them, on the basis of the observed discontinuities, intocomplexes (formally, allostratigraphic units). Dating was carried out by theradiocarbon method, on samples of shells, at Aviones, and charcoal, using boththeABA andABOx-SC methods, at Antn.Nondestructiveelementalanalysis ofpigment residues onshells from both sites wascarriedout by X-Ray Fluorescenceand Energy Dispersive Spectroscopy and compound analysis by micro-Ramanspectroscopy. The mineralogical composition of the colorant lumps from

Aviones was established by X-Ray Diffraction. Assessment of the origin andnature of perforations in the bivalve shells from Aviones and later prehistoricsitesoftheregionwasbasedonvisualandmicroscopicinspectionofthesurfaces,with reference to systematically collected modernbeach thanatocenoses of thesamespecies.For additionaldetails on materials and methods, see SI Appendix.

ACKNOWLEDGMENTS. PedroCallapez,Ivan Mulero, Arnaldo Marn,and MarioSnchez advised on the taxonomy and ecology of marine mollusks and recentregional tectonics. We are indebted to the directors and curators of the Carta-gena andValenciamuseumsfor facilitatingaccess totheircollections, andto the

Spectroscopy of Solids Group of the Materials Science Institute (University ofValencia), especially Nuria Garro, for access to their equipment. F. dE. was sup-ported by the Origin of Man, Language and Languages Programme of theEuropean Science Foundation, the Origin II project of the Aquitaine Region,and the CHIMARC Research Group. Fieldwork was supported by grants to J.Zapata from the Seneca Foundation, the University of Murcia, and the Munici-pality of Mula and dating by ORADS-NERC Grant 2006/1/4.

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the Iberian Peninsula. Pyrenae 37:784.36. Gilman A (1984) Marxist Perspectives in Archaeology, ed Spriggs M (University Press,

Cambridge, UK), pp 115126.37. ZilhoJ (2001)AnatomicallyArchaic,Behaviorally Modern:The LastNeanderthalsand Their

Destiny (StichtingNederlands Museum voorAnthropologieen Praehistoriae,Amsterdam).38. Powell A, Shennan S, Thomas MG (2009) Late Pleistocene demography and the

appearance of modern human behavior. Science 324:12981301.39. Klein R (1992) The archaeology of modern human origins. Evol Anthropol 1:514.40. Trinkaus E (2005) Early modern humans. Annu Rev Anthropol34:207230.

http://www.pnas.org/cgi/data/0914088107/DCSupplemental/Supplemental_PDF#nameddest=STXThttp://www.pnas.org/cgi/doi/10.1073/pnas.0914088107http://www.pnas.org/cgi/doi/10.1073/pnas.0914088107http://www.pnas.org/cgi/data/0914088107/DCSupplemental/Supplemental_PDF#nameddest=STXT


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Symbolic Use of Marine Shells and

Mineral Pigments by Iberian Neandertals

Joo ZILHO, Diego E. ANGELUCCI, Ernestina BADAL-GARCA, Francesco dERRICO,Floral DANIEL, Laure DAYET, Katerina DOUKA, Thomas F. G. HIGHAM, Mara Jos

MARTNEZ-SNCHEZ, Ricardo MONTES-BERNRDEZ, Sonia MURCIA-MASCARS,Carmen PREZ-SIRVENT, Clodoaldo ROLDN-GARCA, Marian VANHAEREN, Valentn

VILLAVERDE, Rachel WOOD, Josefina ZAPATA

Supporting Information Appendix

Supporting Information I:The site of Cueva de los Aviones

Supporting Information II:The site of Cueva Antn

Support ing Information III:Radiocarbon dating

Supporting Information IV:Analysis of mineral p igments in f inds from Cueva de los Aviones

Supporting Information V:Analysis of mineral pigment samples from Cueva de los Aviones

Supporting Information VI:Analysis of mineral p igments in a Cueva Antn Pecten

Support ing Information VII:Reference collection ofAcanthocardia, Cerastoderma and

Glycymeris shells from modern beaches of Murcia and Alicante

Support ing Information VIII:Perforated bivalves from prehistor ic si tes of Mediterranean Spain

Supporting Information IX:

References


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Supporting Information I:The site of Cueva de los Aviones

Cueva de los Aviones is a sea cave located at the base of a natural promontory protectingthe SW entrance to the Cartagena harbor (SI Figs. 1-2). Local bedrock is made up of carbonaterocks belonging to the Lower Alpujrride unit, of Middle-Upper Triassic age (1-2). The cave

walls are composed of moderately deformed light grey and grey limestone (sometimescontaining small chert nodules) and dark grey dolomite with intercalations of calcareous

breccia and fine layers of yellow sandstone and clay. Locally, bedding of bedrock dips W/20.The Lower Alpujrride structural unit also contains lithotypes such as slate, quartzite anddiabase, all of which outcrop at a short distance from the site. The preservation of brecciateddeposits and flowstones well outside the roofed area indicates that the cave was much larger inthe Pleistocene, extending several meters E/SE of the present drip-line, whose retreat is relatedto coastal erosion triggered by sea level rise during the Late Glacial and the Holocene.

These erosive processes almost entirely removed the original fill, except for a brecciatedremnant preserved against the northwestern wall of the cave, ~4 m of which were excavated in

1985 in a two week-long (September 11-26) salvage operation (3-4) (SI Figs. 2-3). Theexcavation was carried out in 5 cm spits that followed the natural dip of the stratification andwith three-dimensional plotting of stone tool and animal bone finds. The extreme cementationof the deposit (explaining its preservation against millennia of sea erosion and repeated impactof strong waves) made it necessary to proceed with chisel and hammer most of the time,inevitably entailing a significant incidence of excavation breakage among the most fragilecategory of finds, the shells.

The extant succession, described below following criteria outlined elsewhere (5), iscomposed of slope sediments accumulated in two distinct cycles and resting on a possiblyEemian rock-beach. From top to bottom, three stratigraphic complexes have been recognized:(1) US (Upper Slope Sediment), which survives in the upper part of the 1985 profile and ascemented remnants hanging from the cave walls; (2) LS (Lower Slope Sediment), whichcorresponds to the excavated archeological sequence; (3) CC (Cemented Cobbles), whichoutcrops slightly above modern sea level on the NE side of the current cave entrance and iscomposed of cemented cobbles and pebbles. Correlation with the units recognized during the1985 excavation is given in SI Table 1, and a stratigraphic column is presented in SI Fig. 4.

Complex US is made of diamict-like limestone and dolomite forming a heterogeneouscemented breccia that contains rare fragments of bones and lithic artifacts with randomorientation. It rests on an erosive surface with evidence of post-depositional deformation, andis separated from underlying complex LS by a discontinuously preserved carbonate flowstone.Complex LS is composed, on average, of coarse material (mainly fragments of carbonate

rocks) with a variable degree of post-depositional carbonation (usually intense) and poorlyrecognizable bedding. It dips SE-E with an inclination of 20-25 and has an erosive base, withangular unconformity. Over a thickness of ~3 m, this complex, originally subdivided in sixarcheostratigraphic levels, comprises ten different units, all of which contain a variable sandfraction formed of angular and subangular grains of the same lithology as the coarser material,with minor amounts of silt and clay (SI Table 2). Complex CC underlies the archeologicalsequence, and is shaped as a ridge made up of cemented, clast-supported, rounded cobbles ofdolomite and limestone, and occasional quartz pebbles.

As the lithology of the fill is identical to that of the cave, a mainly local source is inferredfor the bulk of the US and LS complexes. The geometry of these deposits and of the

discontinuous flowstones found along the walls indicates that a several meter-thick seaward-sloping talus cone originally filled the entrance to the cave. The apex of that cone would havebeen located at the base of the passage that connects the rear part of the site with the inner

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karst, whence came the geological components of the deposit (complemented by direct fallsfrom the caves roof and walls). The sedimentary dynamics involved consist mainly of theaccumulation of single fragments (scree-like), although mechanisms related to the action ofsurface-running waters, such as overland-flow and grain-flow, are also responsible for theformation of some of the observed facies (e.g., units LS04 and LS09).

Despite the abundant archeological remains, neitherin situ features nor anthropic

microfacies were observed in the extant profiles, although a concentration of burnt bones andcharcoal (identified as holm oak by J.-L. Vernet, University of Montpellier) is reported fromlevel IV (3) and may have been part of a disturbed combustion feature. The orientation of theartifacts and bones in the different layers is consistent with the orientation pattern of the rockfragments, indicating that all accumulated together with the sediment and by means of thesame sedimentary dynamics. As a result of these processes, stone tools and animal remainsunderwent lateral displacement, and their horizontal redistribution across the entire surface ofthe cave means that their place of excavation does not necessarily coincide with the actual areaof use and discard. Given the morphology of the cave and the overall geometry of the deposit,human occupation is likely to have taken place towards the inner part of the talus cone, whencethe remains recovered in the area of the 1985 excavation were eventually displaced by suchlow-energy, syndepositional geological processes as discussed above.

Although original site reports (3-4) suggested that a transgressive marine level could beobserved in level V (=unit LS061), the entire LS complex is in fact of continental origin.Moreover, there is no evidence that any of its units could correspond, even if only in part, tothe redeposition of pre-Tyrrhenian marine deposits once extant in the cliff face at a higherelevation. Therefore, the marine shells found in the archeological sequence are non-geogenic,and their association with abundant stone tools and mammal bones (of horse, red deer, ibex,rabbit and tortoise) indicates an anthropic origin.

Five Patella shells from levels I-IV of the 1985 excavation (levels V-VI were sterile) have

been radiocarbon dated (Supporting Information III). The results for levels I-III (=units LS02-LS05) are in stratigraphic order, indicate accumulation during the ~45-50 ka cal BP (calendaryears before present) interval, and suggest correlation of the LS01 flowstone with GreenlandInterstadial 12. The fact that, at ~45 ka 14C BP, the results for levels II and III are statisticallythe same is consistent with the fact that, in our reading of the stratigraphy (SI Figs. 3-4), the

base of level II groups with level III to form a single geological unit, LS05. One of the tworesults for level IV (=unit LS06u), OxA-19312, is not inconsistent with this chronology and, ifaccepted as valid, leads to a Bayesian model (SI Fig. 13) that constrains the deposition of thearcheological sequence to the ~45-48 ka cal BP interval, broadly coinciding with the very coldepisode Heinrich Event 5. However, the mid-point of the uncertainty interval of this level IVresult is younger than that for levels II and III, which, given the challenging nature of

radiocarbon dating in this time range, could reflect undetected, residual contamination. In thiscase, OxA-19312 would be a minimum age only. In fact, the deposition of level IV (=upperpart of unit LS06) may well significantly predate that of level III, from which it is separated bya noticeable discontinuity (the base of the overlying LS05 unit features common carbonateconcretions and has a clear lower boundary).

The other sample from level IV (OxA-20906; ~42.5 ka cal BP) is statistically younger.However, this sample came from square C2, in the seaward row of the excavation grid, wherethe in situ deposits are covered by a sheet of reworked material derived from the erosion of

both the upper part of the LS complex and the lower part of the US complex (SI Fig. 3). As itwent unrecognized at the time of excavation, this sheet must have contributed to the collections

provenanced to the different archeological levels. Bearing in mind that the OxA-20906 result isyounger than the uppermost unit of the LS complex (level I), the simplest explanation for theanomaly is that this C2 sample was in derived position and originally belonged in the UScomplex. If so, it would place the deposition of the US complex ~42.5 ka cal BP, in the cold

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stadial comprised between Greenland Interstadials 11 and 12 (SI Fig. 13). Given thetopographic location of the US remnants preserved towards the rear of the cave (high along thecave walls), a corollary of this interpretation is that the site would have become entirely filledup by the end of the Middle Paleolithic, in agreement with the fact that no Upper Paleolithicitems were observed among the thousands of lithic artifacts recovered in the 1985 excavations.

In all levels, the stone tool assemblages are mostly quartz, suggesting that flint is non-local,

but the Levallois method was used in the production of blanks made on both kinds of raw-materials. The absence of flint cores and the high values of the Sidescraper Index (a toolcategory for which flint was preferentially used) suggest long-distance transport and curationof the finer raw-materials (3-4).

The two perforated Glycymeris shells were found in close association with diagnosticlithics, as noted in the excavation diary, which records their discovery on August 16 andAugust 17, 1985, during the excavation of layer II of square B1 and in the vicinity of two flinttools, a Levallois point and a Mousterian point (SI Fig. 5). No excavation record exists of the

provenience of the perforatedAcanthocardia, but in the collections of the Museum ofCartagena it was clearly labeled as coming from level II, in agreement with the published

information (4). In the framework of this study, we partially cleaned these shells in order toaccess the carbonate-coated surfaces around the umbo, for use-wear analysis; they areillustrated in their original condition in SI Figs. 6-7, together with close-up views of thecorresponding umbo holes before and after cleaning.

Besides these three previously reported perforated shells, and a Spondylus upper valve withpigment residues identified among the unclassified material in the Museum of Cartagena(Fig. 2), levels I-IV yielded other beach-collected shells of non-food gastropods and bivalves,for the most part coming from levels I and II. The majority are G. insubrica whole valves orfragments thereof (SI Fig. 8), but there are also complete or excavation-broken shells of

A. tuberculata and S. gaederopus(SI Fig. 9). The other taxa areillustrated in SI Fig. 10:

Callista chione (no. 1, dorsal fragment with umbo); Chama gryphoides (no. 4, dorsal fragmentwith umbo); Charonia lampas (no. 2, body whorl fragment);Laevicardium oblongum (no. 7,ventral fragment); Pecten maximus (two fragments of a large right valveno. 3 is a posteriorauricle and no. 6 is part of the anterior ventral edge); Thais haemastoma (no. 5, body whorlfragment); and Trunculariopsis trunculus (no. 8, whole shell with a perforation, in alllikelihood post-depositional, in the thinner part of the body whorl). This range of taxa includes(and in fact is wider than) that associated with the perforatedNassarius gibbosulus from theearly modern human-associated Middle Paleolithic levels of the Near Eastern site of Skhul,which also featuredAcanthocardia deshayesii,Laevicardium crassum and Pecten jacobaeus(6).

In terms of ornamental shell find densities, the Aviones values are of the same order of

magnitude as those observed in the African MP/MSA sites that yielded significant amounts ofNassariusbeads. Considering only the perforated material, the round figures are: for the M1phase of Blombos, 39 shells over 20 m and an average thickness of 50 cm (7), i.e., 2/m or4/m; for layer E of Taforalt, 19 shells over 26 m and an average thickness of 40 cm (8), i.e.,1/m and 2/m; for unit II of Aviones, 3 shells over 3 m and 1 m, i.e., 1/m and 3/m (andthese are minima only, given the many Glycymeris fragments where the umbo area is missingand that originally may have been perforated too).

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SI Table 1. Synopsis of Cueva de los Aviones succession (the thicker dotted lines indicatediscontinuities).

Unit Short description Correlation with Montes 1987, 1991 Age

US slope sediment above flowstone OIS-3

LS01 flowstone

LS02slope sediment, random orientationpattern

LS03 slope, parallel orientation pattern

Iconglomerado muy cementado de cantos

dolomticos y tierra negra estructura de greze

lite

LS04 slope, random orientation pattern

slope, random orientation patternII conglomerado dolomtico y tierra rojiza

LS05 slope sediment, with a discontinuouscarbonate crust at the base (LS05k)

III caliche

slope sediment , with few largestones (LS06u)

IV microconglomerado areniscoso y tierra rojiza

LS06 slope sediment, with common largestones (LS06l)

V (no description)

OIS-3

LS07 slope sediment, strongly cemented

LS08slope sediment, some parallelorientation pattern

LS09 discontinuous sand intercalation

LS10slope sediment, no orientationpattern

VI tierra rojiza con algunos cantos dolomticosOIS-3orOIS-4

CC rock-beach Eemian

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SI Table 2. Description of the stratigraphic units in complex LS.

Unit Description

LS01 Carbonate flowstone with laminar structure formed of well-crystallised calcite crystals, probablymicrosparite, strongly cemented. This flowstone is visible at several places along the cave wall, alwaysdisplaying laminar structure. Its thickness is varied, reaching some 15 cm along the N wall.

LS02 Heterogeneous clast-supported breccia, locally open-work, formed of subangular (with rare subroundedand angular) fragments with random orientation, sometimes parallel to the lower boundary, with clastsfrom 2 mm to 15 cm (clasts ranging 2-4 cm are the most represented); clasts are composed of white andlight gray dolomite, white and light gray limestone, fine calcareous breccia, and yellow or pinkweathered sandstone; the matrix is 7.5YR4/2.5 silty-sandy-loam; the sand fraction is fine and formed ofangular and subangular grains of dolomite; the carbonation is intense and massive with some laminarstructure on top (at the contact with unit LS01); the porosity is low and formed of very fine channelswith carbonate hypocoatings; the archaeological material is common and shows random distribution andorientation patterns; the lower boundary is clear, poorly distinct, dipping 25 E; 60 cm thick.

LS03 Fine breccia with moderate sorting formed of subangular and angular (rare subrounded) fragments of thesame lithologies as above with good degree of isorientation of the tabular and platy elements, which areparallel to the lower boundary; the size ranges from 2 mm to 3 cm, with an average of about 1 cm; very

rare fine (5 mm) rounded pebbles are observed; the matrix is 7.5YR4/2.5 silty-sand; the other featuresare the same as in unit LS02 but for the archaeological material, frequent; the lower boundary is clear,poorly distinct, dipping 25 to E; 32 cm thick.

LS04 Sandy loam with abundant angular and subangular stones from 2 mm to 6 cm with random distributionand orientation pattern; the matrix is the same as in unit LS03 but some richer in sand formed of angularand subangular grains of local dolomite and limestone; the archaeological material is common; the lowerboundary is clear, poorly distinct, dipping 25 to E; 30 cm thick.

LS05 This unit shows the same sedimentary characteristics as the overlying one (unit LS04) except for: thematrix is composed of coarse to very coarse sand with some very fine gravel; some vague stratification,due to the presence of discontinuous fine-gravel partially-open-work intercalations is recognised;common carbonate concretions are found at the base (sub-unit LS05k); the lower boundary is clear,poorly distinct, dipping 25 to E; 25 cm thick.

LS06 Fine breccia with moderate sorting formed of subangular and angular (rare subrounded) fragments oflocal lithologies with poor (locally moderate) degree of isorientation of the tabular and platy elements,parallel to the lower boundary; the size ranges from 2 mm to 3 cm, with large (max. 20 cm) subroundedand subangular fragments of dolomite, limestone and limestone breccia particularly common at the baseof the unit (sub-unit LS06l); the matrix is 7.5YR4/2.5 silty-sand; the archaeological material is frequent;the lower boundary is clear, poorly distinct, dipping 25 to E; 55 cm thick.

LS07 Fine-medium poorly sorted breccia formed of angular and subangular stone fragments from 2 mm to 2cm, with random distribution and orientation pattern; the is matrix 9YR5/6 sandy-silt, with the sandpredominantly in the fine sand sub-class; the carbonation is very intense with strong cementation; noarchaeological material is present; the lower boundary is diffuse, poorly distinct; 25 cm thick.

LS08 This unit has the same characteristics as unit LS07 but contains larger stone fragments (max. 10 cm),

sometimes with an orientation parallel the lower boundary; no archaeological material is present; thelower boundary is sharp, discontinuous; 10 cm thick.

LS09 Discontinuous thin layer formed of breccia with same characteristics as unit LS07 with 10YR6/5 sandymatrix; no archaeological material is present; the lower boundary is sharp, discontinuous; 4 cm thick.

LS10 Fine clast-supported breccia with occasional stone fragments up to 3 cm, random distribution andorientation patterns; the other characteristics are the same as for unit LS07; the lower boundary is sharpirregular to bedrock or to the complex CC; 20 cm thick.

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SI Figure 1. Murcia and its Middle Paleolithic. Top. Location of the Region of Murcia in a topographic map of

Iberia. Bottom. Location of the Middle Paleolithic sites of Murcia with a stratigraphic or site formation context in

a topographic map of the region (elevations in meters).

20 km

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SI Figure 2. Cueva de los Aviones. Top. Clockwise: site setting, April 2008; the 1985 excavations; the remnant,

prior to excavation. Middle. Site plan and profile (3-4). Bottom. View of the cave, April 2009.

A

A

0 5

N

AA

m

Middle

Paleo

lithic

brecci

atedr

emnan

t

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LS02-03 (I)

LS04 (IItop)

LS05 (IIbase-III)

LS06 (IV-V)

CC

flowstone

US

LS07-10 (VI)

reworke

d

A1

A1

B1 B1B2

C1 C1C2

A1

A1

B1 B1B2

C1 C1C2

A1

A1

B1 B1B2

C1 C1C2

A1

A1

B1 B1B2

C1 C1C2

Level I

~0.4 m

Level II

~1.1 m

Level III

~0.9 m

Level IV

~1.7 m

SI Figure 3. Cueva de los Aviones excavation and stratigraphy. Top. View of the remnant in April 2009, withindication of stratigraphic limits (in parenthesis, the archeological level designations used in 1985); the small

yellow ruler at the center measures 20 cm. Bottom. The 1985 excavation grid with indication of the areas and

volumes affected, per recognized stratigraphic unit.

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SI Figure 4. Cueva de los Aviones stratigraphic column. A. distribution of archeological finds as observed in

the remnant (B=bone fragments, C=charcoal fragments, L=lithic artefacts, S=shells). B. stratigraphic column (the

width is proportional to the grain size of the units matrix; C=clay, Si=silt, Sa=sand, G=gravel). C. units describedaccording to this study (US, LS and CC stratigraphic complexes; LS01 to LS 10 units of the LS complex;

numbers correspond to the average depth in centimetres recorded in the remnant). D. units defined during

excavation (the numbers indicate depth of boundaries in the profile as illustrated in refs. 3-4).

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SI Figure 5. Cueva de los Aviones finds. Top and Middle. Flint tools (from the excavations photo archive).

Clockwise: point (level I), sidescraper (level I), point (level II), sidescraper (level II), sidescraper (level III),

sidescraper (level IV). Bottom. Pages of the excavation diary noting the discovery of the perforatedGlycymeris.

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SI Figure 6. Cueva de los Aviones level II perforated shells, prior to cleaning for this study. Top.

Acanthocardia tuberculata. Middle and Bottom. Glycymeris insubrica.

1 cm

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SI Figure 7. Cueva de los Aviones perforated shells from level II. The perforations prior to (left) and after

(right) mechanical cleaning (preceded by removal of the brecciated clutters in an ultrasound vat). Top.Acanthocardia tuberculata. Middle and Bottom. Glycymeris insubrica. Note, especially in the bottom specimen,

the heavy chemical micro-weathering around the perforations, which relates to the formation of the carbonate

coatings and prevented the presevation of any use wear evidence.

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SI Figure 8. Cueva de los Aviones non-perforated Glycymeris insubrica shells. Top. Fragments from level I.

Middle. Whole shell from level I (from excavations photo archive, missing in the Museum of Cartagena) (left)

and fragments from level II (right). Bottom. Whole shell and fragment from level III.

1 cm

1 cm

1 cm

1 cm

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SI Figure 9. Cueva de los Aviones complete non-food bivalve shells. Top. Upper valve ofSpondylus

gaederopus (level IV). Middle. Lower valve ofSpondylus gaederopus (level I). Bottom.Acanthocardia

tuberculata shells from level III (left) and level IV (right; excavation-broken).

1 cm

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SI Figure 10. Cueva de los Aviones non-food shells. 1. Callista chione (level I). 2. Charonia lampas (level IV).

3. Pecten maximus (level III). 4. Chama gryphoides (level I). 5. Thais haemastoma (level II). 6. Pecten maximus

(level III). 7.Laevicardium oblongum (level IV). 8. Trunculariopsis trunculus (level III).

1

2

4

5

6

7

8

3

1 cm

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Supporting Information II:The site of Cueva Antn

Cueva Antn is a rockshelter excavated into the base of an E-W, 25-30 m high, Eocenelimestone escarpment located towards the tail end of the La Cierva reservoir, on the Mula river(SI Fig. 11). Salvage work undertaken in 1991 exposed a >4-m-thick sequence, at the base of

which Middle Paleolithic levels were identified (9). After two seasons (2007-08), newexcavation work has also exposed Middle Paleolithic occupations in the uppermost units.

In the areas currently under investigation, the Antn deposits can be divided in four maingeoarcheological complexes: DD (Dam Deposits), made up of fine material with some coarserintercalations, and accumulated over the last decades at times when the level of the La Ciervareservoir raised above the base of the shelters overhang, fully submerging the site; TL(Transitional Layers), formed of disturbed layers of uncertain age found to the north of row 18of the grid in intermediate position between the DD deposits and the underlying Pleistocenesediments; AS (Archaeological Succession), formed of a number of superposed alluvialsequences featuring significant lateral variation and distinct sedimentary facies, and including

intercalations of slope material, particularly against the back wall and especially so in its upperpart (SI Fig. 12); FP (Fine Palustrine), a fine organic sediment forming the base of the exposedsuccession.

Geomorphologically, the AS complex can best be described as a fluvial terrace created froman alluvial sequence whose sheltering under an overhang limited the impact of post-depositional modifications and soil formation processes. Burrowing features created by rabbitsand small carnivores, a major cause of disturbance in all cave and rockshelter sites ofMediterranean and west Atlantic Iberia, are conspicuously absent from the 1991 profiles andhave not been observed in current excavation work. Where the uppermost levels of the AScomplex are concerned, these observations are consistent with the fact that the abundant rabbit

bone component of the deposits is almost exclusively formed of juvenile remains bearingtraces indicative of accumulation by eagle-owls (and individual, intact regurgitation pelletshave even been recovered in the finer, silty-loamy units). From a taphonomic perspective, thesequence is therefore pristine, with no evidence for vertical post-depositional dislocations andwith only limited, horizontal syndepositional displacement of finds having occurred in thecoarser levels (those made up of fine alluvial sands or of cm-size cryoclastic wall debris).

Documenting the shifting location of a diverse range of valley bottom environments overthe time of accumulation (river channels, gravel bars, sand beaches, bogs, etc.), 48 differentunits, grouped in five subcomplexes, are currently recognized within the 2-3 m spanned by theAS complex (SI Table 3). Attempts at radiocarbon dating its base have so far failed due to theinsufficient yields of the charcoal samples used, but a consistent chronology exists for the

upper part, above an erosive unconformity at the base of level II-m. This level corresponds to agravel/sand bar accumulated at a time when the Mula river ran through the shelter, and theoverlying sand beach (level II-l) yielded characteristic sidescrapers and cores for which wehave a terminus ante quem of ~43.5 ka cal BPthe age obtained for overlying level II-h/i, anarcheologically sterile massive silt deposit accumulated by decantation in the framework of asubsequent episode of low-energy inundation of the site. Carbonate-incrusted level II-dseparates the similarly sterile alluvial units that overlie II-h/i from the uppermost archeologicaloccupation, which is contained in level I-k. This unit is a clast-supported breccia formed ofsmall angular limestone debris with a clayey silt matrix and has been dated to ~32.9 14C BP(~37.4 ka cal BP) by a pine charcoal sample pretreated with the ABOx-SC technique(Supporting Information III).

In the basal units of the AS complex, the archeological levels correspond to well-preserved,single-event Mousterian occupations organized around hearth features; stone tools are

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abundant (SI Fig. 12), and the bones, often cut-marked, are of large herbivores. In theuppermost units (II-l and I-k), however, finds are scarce, suggesting sporadic and highlyephemeral visits only, but, technologically and typologically, the lithics are of Mousterianaffinities (SI Fig. 12).

Where level I-k is concerned, the handful of artifacts (including the Pecten shell) wasrecovered in close association with the dated charcoal samples (Supporting Information III),

and these dates place the human occupation represented therein towards the end of the periodof late persistence, in Iberian regions located to the south of the Ebro drainage, of a Neandertal-associated Middle Paleolithic (10), which, at Gorhams Cave (Gibraltar) and Gruta da Oliveira(Portugal), is well documented until ~32 ka 14C BP. Assignment of the level to the latestMiddle Paleolithic is further supported by the fact that its characteristics fall outside whatwould be expected in a context of the regional Upper Paleolithic (for which, moreover, theearliest reliable dates are, at present, 3 m of it having already been excavated.

SI Table 3. Stratigraphic layout of complex AS of Cueva Antn.

Subcomplex Short description Base

Stratigraphic

span Archeology Age

AS1

upper part, with slope/wall inputs and finealluvial

paraconformityI-g (andunnamed unitsabove) to II-c

I-k(artifacts)

I-k:~31.1 ka 14C BP

AS2

mainly alluvialsediment, forming atleast three minorcycles

erosiveunconformity II-d to II-m

II-l(artifacts)

II-i:~39.6 ka 14C BP

AS3

mainly alluvialsediment, forming atleast three minorcycles

erosiveunconformity

II- to II-t OIS-3

AS4

sand layers preservedin West profile ofsquares L21-L22

paraconformity II- OIS-3

AS5

mainly alluvialsediment with a poorly

developed buried soilon top, forming at leastfour minor cycles

paraconformity II-u to III-n

II-u/y(artifacts)

III-b/d, III-i/j(artifacts,features)

OIS-3

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SI Figure 11. Cueva Antn. Top. The La Cierva dam and reservoir, April 2009 (the circle indicates the location

of the rockshelter). Middle. View of the excavation at the end of the September 2008 field season. Bottom. Site

plan and grid, with indication of the excavated areas.

Nm

Extant profiles

2007 geological trench

2007-08 archeological trenches

1991 Trench 1

1991Trench2

Elevation of exposure ofthe 1991 unit IIIdeposits

JKL

22

21

20

5 m +4

+2

0

+3+5 +7

-2

-4-6

0

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26/79

SI Figure 12. Cueva Antn. Top. Flint artifacts from level I-k; left to right, centripetal core on flake blank,

denticulate and splintered piece. Middle. Flint sidescrapers from level III-f. Bottom. East profile of the J/17-19

squares, with indication of the stratigraphic units recognized in the AS complex and of the age of dated levels.

II-e

II-

II-p II-r

II-q

II-u

II-m

II-l

II-k

II-i

I-k

~43.5 ka cal BP

~37.4 ka cal BP

J>I17 J>I18 J>I19

218 cm

[from Martnez et al., 1991: Fig. 8]

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7/28/2019 Zilhao Et Al 2

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Zilhao Et Al 2010 PNAS Neandertal Shells

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