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Journal of Human Evolution 68 (2014) 1e13

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Journal of Human Evolution

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

On the chronology of the Uluzzian

Katerina Douka a,*, Thomas F.G. Higham a, Rachel Wood a,b, Paolo Boscato c,Paolo Gambassini c, Panagiotis Karkanas d, Marco Peresani e, Anna Maria Ronchitelli c

aOxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South ParksRoad, OX1 3QY Oxford, United KingdombResearch School of Earth Sciences, Australian National University, AustraliacDipartimento di Scienze Fisiche, della Terra e dell’Ambiente, U.R. Preistoria e Antropologia, Università degli Studi di Siena, Via Laterina 8, 53100 Siena, Italyd Ephoreia of Palaeoanthropology-Speleology of Southern Greece, Ardittou 34b, 11636 Athens, GreeceeDipartimento di Studi Umanistici, Sezione di Scienze Preistoriche e Protostoriche, Università di Ferrara, Corso Ercole I d’Este 32, I-44100 Ferrara, Italy

a r t i c l e i n f o

Article history:Received 10 April 2013Accepted 10 December 2013Available online 7 February 2014

Keywords:Modern humansNeanderthalsRadiocarbon datingItalyGreece

* Corresponding author.E-mail addresses: katerina.douka@rlaha.ox.ac.uk (

edu.au (R. Wood), paolo.boscato@unisi.it (P. B(P. Karkanas), psm@unife.it (M. Peresani), annamM. Ronchitelli).

0047-2484/$ e see front matter Crown Copyright � 2http://dx.doi.org/10.1016/j.jhevol.2013.12.007

a b s t r a c t

The Uluzzian, one of Europe’s ‘transitional’ technocomplexes, has gained particular significance over thepast three years when the only human remains associated with it were attributed to modern humans,instead of Neanderthals as previously thought. The position of the Uluzzian at stratified sequences, al-ways overlying late Mousterian layers and underlying early Upper Palaeolithic ones, highlights its sig-nificance in understanding the passage from the Middle to Upper Palaeolithic, as well as the replacementof Neanderthals by modern humans in southeastern Mediterranean Europe. Despite several studiesinvestigating aspects of its lithic techno-typology, taxonomy and material culture, the Uluzzian chro-nology has remained extremely poorly-known, based on a handful of dubious chronometric de-terminations. Here we aim to elucidate the chronological aspect of the technocomplex by presenting anintegrated synthesis of new radiocarbon results and a Bayesian statistical approach from four stratifiedUluzzian cave sequences in Italy and Greece (Cavallo, Fumane, Castelcivita and Klissoura 1). In addition tobuilding a reliable chronological framework for the Uluzzian, we examine its appearance, tempo-spatialspread and correlation to previous and later Palaeolithic assemblages (Mousterian, Protoaurignacian) atthe relevant regions. We conclude that the Uluzzian arrived in Italy and Greece shortly before 45,000years ago and its final stages are placed at w39,500 years ago, its end synchronous (if not slightly earlier)with the Campanian Ignimbrite eruption.

Crown Copyright � 2014 Published by Elsevier Ltd. All rights reserved.

Introduction

In 1964, Arturo Palma di Cesnola coined the term ‘Uluzzian’ todescribe the technocomplex he identified a year earlier at CavalloCave (Grotta del Cavallo) in southern Italy (Palma di Cesnola andBorzatti von Löwenstern, 1964). The distinct, dark Uluzzian layersoverlay a long series of Mousterian deposits and were superposedby a late Upper Palaeolithic (Late Epigravettian) phase from whichthey were separated by a thin stalagmitic crust and a tephra layer.

The Uluzzian layers were rich in lithic elements of UpperPalaeolithic character, terrestrial and marine faunal remains andseveral hearths. They contained bone points, perforated shell

K. Douka), rachel.wood@anu.oscato), pkarkanas@hua.graria.ronchitelli@unisi.it (A.

014 Published by Elsevier Ltd. All

beads, mineral pigments, as well as two human deciduous teeth.The technocomplex was assigned to the Upper Palaeolithic (‘Lep-tolithic’) tradition, yet archaic at its lower layers, due to the pres-ence of particular tool-types (lunates) that do not exist in thepreceding Mousterian levels. Palma di Cesnola immediatelyrecognized parallels with the Franco-Cantabrian Châtelperronian,and efforts to elucidate the geographical, techno-typological andchronological span of the Uluzzian, as well as its role in the Middle-to-Upper Palaeolithic transition, began.

In 2011, almost 50 years after the initial discovery of the tech-nocomplex, the Uluzzian was brought to the forefront again. Thetwo deciduous teeth from Cavallo Cave, the only human remainsassociated with the Uluzzian so far, were identified as belonging toanatomically modern humans and not Neanderthals, as was theconsensus until then (Benazzi et al., 2011). The parallel radiocarbondating of associated shell beads found across the Cavallo Uluzzianstratigraphy rendered the teeth the oldest currently-known re-mains of modern humans in Europe (Benazzi et al., 2011).

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K. Douka et al. / Journal of Human Evolution 68 (2014) 1e132

The geographic span of the Uluzzian has remained almost un-changed over the last decades and several authors have describedthe techno-typological features of the assemblages assigned to it.The chronology of the Uluzzian, however, remained poorly under-stood. Given the importance of the technocomplex in our under-standing of the appearance and spread of modern humans inEurope, a chronological synthesis is long due and is the main focusof the present article.

Background

Geography

The presence of the Uluzzian has been reported in about 20caves and open-air localities of peninsular Italy (Palma di Cesnola,1989, 1993; Riel-Salvatore, 2007, 2009; Ronchitelli et al., 2009).These include sites mainly in Apulia (Cavallo, Uluzzo, Parabita,Mario Bernardini, Serra Cicora, Torre Testa, Falce del Viaggio, For-esta Umbra), Basilicata (Atella Basin), Campania (Cala, Castelcivita,Tornola), Calabria (San Pietro a Maida, Punta Safò), Tuscany (LaFabbrica, Poggio Calvello, Val Berretta, San Romano, Indicatore, SanLeonardo, Salviano and Maroccone) (Palma di Cesnola, 1989, 1993and references therein). More recently, the presence of the Uluz-zian at Fumane, Verona, was reported and this is currently thenorthernmost occurrence of the technocomplex (Peresani et al.,2008; Peresani, 2012).

A single occurrence outside Italy, at the Greek cave of Klissoura1, Argolis, was described by Koumouzelis et al. (2001). This waslater questioned by Papagianni (2009), but has since been re-confirmed by the excavators (Kaczanowska et al., 2010). AnotherGreek Uluzzian context from Kephalari Cave, also in Argolis, wasreported on anecdotal evidence (Hahn, 1984). This is currentlybeing re-examined and more data will become available soon (G.Marshall, Personal communication).

Lithic technology-typology

The Uluzzian is a flake-dominated industry (Palma di Cesnola,1993) that brings together a set of technological innovations (e.g.,De Stefani et al., 2012). The production scheme is characterized byvariable methods and techniques and a significant reduction inpredetermined forms, when compared with the final Mousterian.The increased importance in the launch of various reduction se-quences cannot be attributed to the qualities of the raw materialonly, but must assume behavioural significance too.

Blanks are detached using a unipolar method, with a singlestriking platform by direct percussion, and from a bipolar knappingon an anvil, which produces splintered pieces usedmainly aswedgeson medium-hard material. The principal objective was the produc-tion of average-thickness flakes and laminar flakes, sometimesnaturally backed. Small blades and a small number of bladelets werealso obtained. Few orthogonal, multi-directional and surface cen-tripetal cores are also present (Palma di Cesnola, 1965a, 1966a, 1989;Peresani, 2008; Moroni et al., 2013; De Stefani et al., 2012).

Several tool-types are identified as part of the technocomplex:various end-scraper types, side-scrapers, few burins, denticulates,retouched blades and bladelets. The geometric crescent-shapedbacked piece, normally referred to as ‘lunate’ (or ‘semi-luna’ inItalian) is the fossile directeur of the industry; an innovationsometimes of microlithic dimensions (Palma di Cesnola, 1993). Thistool is particularly abundant in Cavallo Cave. Lunates do not seem tooriginate from specific operational chains.

The high incidence of splintered pieces (‘pièces esquillées’) isanother characteristic feature of the Uluzzian. Small blades andbladelets are irregular and sometimes with traces of cortex, while

there is low incidence of lamellar débitage. Local raw materialprocurement, including thin slabs of siliceous limestone for theproduction of retouched pieces, is a further characteristic of theearliest Uluzzian phases (Gambassini, 1997; Riel-Salvatore andNegrino, 2009).

Based on the most complete Uluzzian complex known to date,the stratified sequence of Cavallo Cave, and the typological vari-ability observedwithin it, Palma di Cesnola (1965a, b, 1989) dividedthe Uluzzian into three distinct industrial phases: Archaic (Cavallolayer E III), Evolved (Cavallo layers E II-I) and Final (Cavallo layers DII-D Ib). Current techno-typological and functional analyses areunderway to clarify issues of origin, internal evolution, externalinfluences and adaptations of the Uluzzian, as well as its relation-ship to the succeeding Aurignacian (De Stefani et al., 2012; Ranaldo,In press), broadly contemporaneous Châtelperronian and precedingMousterian. What has become clear is that the one-to-one analogywith the Châtelperronian cannot be supported anymore as thetechnological details for tool production, as well as the final prod-ucts and their typology, are indeed very different.

Bone industry, shell ornaments and pigments

Along with the distinct lithics and microlithics, several Uluzziansequences (Cavallo, Castelcivita, La Fabbrica) have yielded workedbone industry. About 15 bone implements in cylindro-conicalforms, mainly awls, have been discovered and described so far(Ronchitelli et al., 2009; d’ Errico et al., 2012). When identifiable,these were manufactured on horse and red deer metapodials andon horse fibulae. While they incorrectly associated them with Ne-anderthals, d’ Errico et al. (2012) suggested that the Uluzzian bonetools reveal a techno-typological complexity from their earliestappearance. In addition to these tools, a recent publication reportson the presence of bone retouchers in the Uluzzian layers ofFumane Cave (Jéquier et al., 2012).

The use and manufacture of beads is another key behaviouralcharacteristic of the Uluzzian. Perforated marine shells have beendiscovered at the Uluzzian layers of Cavallo, Klissoura (and prob-ably Cala, which is not part of the current study). They make theirfirst appearance at the Archaic Uluzzian phase of Cavallo and in-crease in number, up to a few dozens, at its final phases (Palma diCesnola, 1964, 1989). Several shell beads appear in Klissoura(n¼ 32; Stiner, 2010) but no ornaments have been discovered in theUluzzian of Fumane and Castelcivita.

Beads are almost exclusively manufactured from the shells ofmarine molluscs, mainly scaphopods (Dentalium sp.) and gastro-pods (e.g., Columbella rustica, Cyclope neritea). A small number ofbivalves have also been identified in the assemblages and, with theexception of a series of perforated Nuculana cf. sp. from Cavallo, therest are not humanly modified (e.g., Pecten sp. from Castelcivita andother unidentified fragments). Just as in the Aurignacian, theUluzzian shells are transformed into beads through perforationwith the aid of lithic tools using direct or indirect percussion. In thecase of the tubular Dentalium shell, the apical end is snapped offand the shell is sawn to create smaller sections.

Finally, evidence for the use of colourants (ochre and limonitepieces) has been revealed in Cavallo and Mario Bernardini caves(Borzatti von Löwenstern, 1970; Palma di Cesnola, 1989). At themoment it is not possible to tell whether these minerals were usedsolely for symbolic practices or other activities (e.g., hide prepara-tion, as grinding medium or hafting agent).

Taxonomy

In terms of biological attribution, the makers of the Uluzzianhave remained elusive, until very recently. The only known fossils

Table 1Previously available chronometric determinations for the Uluzzian layers at the four studied sites: Fumane, Castelcivita, Cavallo and Klissoura.

Site Lab code Age � Context Material Reference

Fumane Cave LTL-1830A 29,602 240 A3 e str. I Charcoal Higham et al., 2009Fumane Cave LTL-1831A 29,233 350 A3 e str. II Charcoal Higham et al., 2009Fumane Cave LTL-1796A 29,361 320 A3 e str. IV Charcoal Higham et al., 2009Fumane Cave LTL-1795A 37,828 430 A3 e str. IV Charcoal Higham et al., 2009Fumane Cave OxA-8021 33,300 400 A4 II Charcoal Peresani et al., 2008Fumane Cave OxA-6462 33,150 600 A4 II Charcoal Peresani et al., 2008Fumane Cave LTL-566A 33,700 350 A4 II Charcoal Peresani et al., 2008Fumane Cave ESR 44 � 7 ka A4 Charcoal Peresani et al., 2008

Castelcivita Cave F-71 32,470 650 rsa Burnt bones Gambassini, 1997Castelcivita Cave F-106 >34,000 ? Burnt bones Gambassini, 1997Castelcivita Cave F-107 33,220 780 pie Burnt bones Gambassini, 1997Castelcivita Cave GrN-13985 33,300 430 rpi Charcoal Gambassini, 1997

Cavallo Cave RM-352 >31,000 E II-I Charcoal Palma di Cesnola, 1969Cavallo Cave AA-66812 34,900 1900 E III Burnt bone Riel-Salvatore, 2007; Ronchitelli et al., 2009Cavallo Cave AA-66819 36,600 2300 EIII Burnt bone Riel-Salvatore, 2007; Ronchitelli et al., 2009Cavallo Cave AA-66814 36,510 2300 EIII Burnt bone Riel-Salvatore 2007; Kuhn et al., 2010Cavallo Cave AA-66813 32,300 2700 EIII Burnt bone Riel-Salvatore, 2007. Kuhn et al., 2010Cavallo Cave 6 AMS 14C dates w21e31 ka BP EIII Burnt bone Riel-Salvatore, 2007

Klissoura Cave Gd-7878 17,430 100 V Carbonates Koumouzelis et al., 2001Klissoura Cave Gd-12037 26,250 310 V/hearth 42 Carbonates Koumouzelis et al., 2001Klissoura Cave Gd-12027 27,100 600 V/hearth 53 Carbonates Koumouzelis et al., 2001Klissoura Cave Gd-10714 >31,100 V/hearth 42 Soil organics Koumouzelis et al., 2001Klissoura Cave Gd-10715 >30,800 V/hearth 53 Soil organics Koumouzelis et al., 2001Klissoura Cave RTT-4790 29,660 360 V upper Charcoal Kuhn et al., 2010Klissoura Cave RTT-4791 30,774 410 V upper Charcoal Kuhn et al., 2010Klissoura Cave Gif-99168 40,100 740 V/Sq. B3/H42/175-180 Bone Kuhn et al., 2010

With the exception of an ESR determination from Fumane, all other results are 14C.

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e13 3

associated with the technocomplex are two deciduous molars fromCavallo Cave. Cavallo-B was found at a hearth at the base of theArchaic Uluzzian phase (E III), and Cavallo-C, 20 cm above, in theEvolved Uluzzian phase (E II-I). They were initially described andpublished by Palma di Cesnola and Messeri (1967), who classifiedthe earliest, Cavallo-B, as modern human and Cavallo-C as Nean-derthal. Churchill and Smith (2000) were unable to link the molarssecurely to either Neanderthals or modern humans but, unlikePalma di Cesnola and Messeri (1967), they favoured a Neanderthalattribution for Cavallo-B. This is most likely due to a mistake(reversed values) in the published dimensions these authors usedin their analyses (see Benazzi et al., 2011).

The most recent and definite work over the taxonomy of the twodeciduous teeth is that of Benazzi et al. (2011). Two independentmorphometric methods were used to assess the taxonomy ofCavallo-BandCavallo-C, namelygeometricmorphometric analysis ofdental crown outlines and 2-D enamel thickness based on micro-CTscanning. Both lines of investigationplaced the two specimens in themodern human spectrumwith certainty and therefore transformedthe traditional interpretation for the makers of the Uluzzian, whichare now considered to be modern humans and not Neanderthals.

Finally, a deciduous central incisor found in the faunal assem-blage of the lower Uluzzian layer EIII, and tentatively reported byRiel-Salvatore (2007) and Gambassini et al. (2006) as belonging to aNeanderthal, is not human (Benazzi, personal communication;contra Zilhão, 2013). Recent criticisms on the basis of thismisidentification that “the issue of whomade [the Uluzzian] cannotbe answered simply because both groups were implicated” (e.g.Zilhão, 2013) is therefore unsupported.

Previous chronological framework and problems

Prior to our work, only a small number of Uluzzian contextswere dated (Table 1, Fig. 3).

An infinite age (RM-352: >31000 years BP [before present]) wasuntil 2009 the only published chronometric information for Cavallo

Cave. Palma di Cesnola obtained this determination from charcoalhe found at the Evolved Uluzzian layer (E II-I) (Palma di Cesnola,1969; Alessio et al., 1970). A series of ten radiocarbon dates fromthe Archaic Uluzzian layer E III were obtained by J. Riel-Salvatore(2007). The four oldest determinations, and thus favoured, rangedbetween 32 and 36 ka BP (thousands of years ago before present)and were published by Ronchitelli et al. (2009) and Kuhn et al.(2010) (Table 1). However, the remaining six dates from thegroup, all from layer E III, fall between w21 and 31 ka BP (Riel-Salvatore, 2007) and are completely inconsistent with the strati-graphic position of the samples (see discussion below).

Four previous determinations from the Uluzzian layers at Cas-telcivita (‘pie’, ‘rpi’, bottom of ‘rsa’), again on burnt bone, clusteredat 33.3e32.5 ka BP (Gambassini, 1997) (Table 1). These de-terminations are again inconsistent with the position of the sam-ples well below the Campanian Ignimbrite that seals thearchaeological series.

At Fumane, the presence of Uluzzian was only recently formal-ized in layers A3 and A4 of the site (Peresani, 2008, 2012). Theselayers were associated with several radiocarbon determinationsranging from 29 to 37 ka BP (Table 1) and one on ESR of a tooth fromA4 at 44 � 7 ka (Peresani et al., 2008; Higham et al., 2009).

In Greece, Uluzzian layer V at Klissoura Cave 1 was associatedwith five determinations on carbonates and organic fractionsranging between 29 and 31 ka BP (Koumouzelis et al., 2001) andone bone determination at w40 ka BP (Gif-99168) (Table 1). Thelatter has no secure context as it was collected from an OSLdosimeter tube. Given the small thickness of layer V, it may wellhave derived from underlying Middle Palaeolithic layers (Kuhnet al., 2010).

A good number of these previously available determinations arestatistically identical, falling around 32.0e33.5 ka BP or, when cali-brated, between 34 and 38 ka cal BP (Fig. 3). Due to this seeminglyinternal consistency, prehistorians have accepted them as reliable.

Many researchers fail to acknowledge the limitations posed bysample selection and pretreatment chemistry in the successful

Figure 1. Map of southeastern Mediterranean Europe showing the location of the four studied Uluzzian sites. The location of some of the Aurignacian and Mousterian sitesmentioned in the text is also shown with open squares: 1e2, Mochi-Bombrini; 3, Mezzena; 4, Paglicci; 5, Serino; 6, Theopetra; 7, Asprochaliko; 8, Franchthi; 9, Lakonis I; 10,Kalamakia. The dashed line indicates the exposed continental shelf at sea level regression of about 100 m; this corresponds approximately to the coastline at w40,000 years ago.

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e134

application of radiocarbon dating, especially towards the age limitof the technique. Some of the aforementioned Uluzzian de-terminations were produced on material known to be very difficultto date. For example, most measurements from Cavallo and Cas-telcivitawere performed on burnt bones, a highly unreliable type ofsample (van Strydonck et al., 2005, 2009), especially forPalaeolithic-aged material. This is because in burnt bone themeasured carbon (C) comes from the mineral matrix, a highlyporous and non-homogenous phase prone to external contamina-tion. In contrast, when fresh (i.e., non-burnt) bone is dated, themeasured C derives from the more resilient organic (collagen)fraction.

Figure 2. Composite stratigraphic columns of Fumane, Castelcivita, Cavallo and Klissoura 1 (et al., 2008; Giaccio et al., 2008; Karkanas, 2010). The location of the dated samples is indi

In addition to problems relating to dating burnt bone, manyprevious radiocarbon determinations for the Uluzzian were pro-duced on charcoal/organic matter cleaned using the Acid-Base-Acid(ABA) method. This chemical protocol has been shown to under-estimate systematically the age of samples older than 30 ka BPwhen compared with more rigorous new methods, such as theABOx-SC protocol (Bird et al., 1999; Santos et al., 2003; Highamet al., 2009; Douka et al., 2010b; Higham, 2011; Wood et al.,2012). In turn, this means that the ABA determinations must un-derestimate the true age of the Uluzzian.

These methodological, 14C-related, limitations are corroboratedby another fact. In at least three stratified Uluzzian sequences

modified from Palma di Cesnola, 1964; Gambassini, 1997; Martini et al., 2001; Peresanicated. Columns not to relative scale.

Figure 3. Previous available determinations for the Uluzzian in Italy and Greece. Theconventional measurements are calibrated using the IntCal/Marine 09 curve (Reimeret al., 2009) on the OxCal platform (Bronk Ramsey, 2009). Infinite determinations, aswell as the group of burnt bone dates from Cavallo, were added separately and arepresented here as rectangular bars. Gd-7878 is out of scale, calibrated at w20 ka cal BPand is not shown. On the top of the graph, the NGRIP d18O record is shown (Andersenet al., 2006; Svensson et al., 2006) and the Greenland Interstadials, corresponding toameliorated climatic conditions, are numbered. The age for the CI ash is indicated onthe top of the sequence as a continuous vertical grey line at the sites it occurs, and as adashed line in Fumane where its presence is not confirmed.

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e13 5

(Cavallo, Castelcivita, Klissoura), a tephra layer identified as theCampanian Ignimbrite (CI) on stratigraphic and geochemicalgrounds overlies the Uluzzian or, in the case of Castelcivita, theProtoaurignacian deposits, occurring directly above the Uluzzian(Gambassini, 1997; Giaccio et al., 2006, 2008; Fedele et al., 2008;Lowe et al., 2012) (Fig. 2). The age of the Uluzzian at these sitesshould be older than w39.3 � 0.1 ka, the calendar age for the CIeruption (De Vivo et al., 2001). Clearly, the dates from these sites aremuch younger, often by several millennia when calibrated (Fig. 3).In fact, of the 18 finite determinations previously available for theUluzzian (Table 1), 12 postdate the CI chronological marker at 39e40 ka (Fig. 3). Only four of the burnt bone determinations fromCavallo (Ronchitelli et al., 2009; Kuhn et al., 2010) are statisticallysimilar to the age of the CI and predate it. However, a more carefulexamination reveals that these dates come from the lowermostUluzzian layer E III, about 80 cm below the tephra layer, for whichan older age should be expected.

Studied sequences

Given the distinct techno-typological features and importantstratigraphic position of the Uluzzian and, as of recent, itsconnection with the earliest modern humans in Europe, a reliablechronology is of uttermost importance in understanding aspects ofthe initial appearance, expansion and eventual demise of thetechnocomplex, which have remained unknown thus far. The reli-able dating of the Uluzzian has been one of the major goals of aproject initiated by the authors and performed at the OxfordRadiocarbon Accelerator Unit between 2008 and 2011. We decidedto date four of the best-known Uluzzian sites (Cavallo Cave, Cas-telcivita Cave, Fumane Cave and Klissoura Cave 1; Fig. 1). Despitethe presence of several open-air localities across Italy identified asUluzzian on the basis of surface collections, here, priority was givento stratified sequences.

Cavallo Cave is situated on the rocky coast of the bay of Uluzzo,in Puglia, SE Italy. It was discovered in 1960 and excavations tookplace during 1963e1966 (Palma di Cesnola, 1963, 1964, 1965b,1966b) and from 1986 to 2007 (Sarti et al., 1998e2000, 2002).Salvage excavations were conducted in the late 1970s and early1980s in the process of installing a gate at the entrance of the caveto protect it from looters, who had ravaged most of the standingarchaeological deposits.

The site contained a long stratigraphic succession comprisingabout 7 m of archaeological deposits, directly based on a marineinterglacial beach conglomerate (layer O) (Romagnoli, 2012). Thearchaeological sequence of Cavallo (Fig. 2) is dominated by Mous-terian layers (N-F I), followed by a thin layer of green volcanic ash(Fa). The ash is overlain by the Uluzzian layers (E III to D Ib). Theseare capped by a stalagmitic floor (D Ia) and a further layer of vol-canic ash (C II) altered at the top (C I). This tephra has been iden-tified as deriving from the Campanian Ignimbrite eruption (R.Sulpizio, Personal communication). Above this are layers B IIeB I,with Late Epigravettian (Romanellian) remains, and the sequence issealed by Neolithic layer A.

Castelcivita is situated in the Calore river valley, at the foot of theAlburni Mountains (Salerno), w100 m above sea level. Excavationswere conducted from 1975 to 1988 when a 3 m-long sequence wasdiscovered and investigated in detail (Gambassini, 1997). Thearchaeological succession contained several Mousterian, UluzzianandProtoaurignacian levelsandwassealedbythickvolcanicdepositswith the distinctive two-phase structure of the CI (Giaccio et al.,2008). Within the stratigraphic sequence, eight human-derivedphases were recognized. The three lower phases are attributed tothe Mousterian and correspond to levels XIIIeVII (¼ cgr-gar-rsi00).They are characterized by blocks of limestone of different size and bycemented layers. Anerosional discontinuity separates the lowermostMousterian from the two middle parts of the sequence (levels VIeIIIb ¼ rsi0-pie-rpi-rsa00), which contain Uluzzian remains. The Uluz-zian levels are characterized by fine red sediment mixed with lime-stoneblocksderived fromthecollapseof theentranceof thecave. Thesucceeding three phases (levels IIIa-ars ¼ rsa0-gic-ars), containedProtoaurignacian remains within sandy sediment (see also SOM).

Fumane is a dolomitic limestone cave at the southern slope ofthe Venetian Pre-Alps, between the low alluvial plains and the highplateau of Mount Lessini. Systematic excavations started in 1988and continue to this day (Bartolomei et al., 1992; Cremaschi et al.,2005; Peresani et al., 2008). The excavations have revealed adeep, 12 m thick sedimentary sequence. On the basis of distinctlithological composition, pedological features and the density andnature of cultural evidence, the stratigraphy has been divided intofour main macro-units labelled S, BR, A and D (Martini et al., 2001)(Fig. 2). Layers A4eA3 from macro-unit A were recently charac-terized as Uluzzian (Peresani, 2008, 2012; Peresani et al., 2008; see

Table 2New radiocarbon determinations for the Uluzzian layers of Cavallo, Fumane, Castelcivita and Klissoura.

Site/sample ID OxA-code 14C age � Context Material Genus/species Cal yr BP (95.4%) Analytical parameters

From To

Collagen qualityFumane Cave/RF44 21736 39,100 1000 A3/69a/53 Bone Large mammal 44990 42030 C/N: 3.4, % Coll.: 2.9,

d13C: �19.6, d13N: 6.3Fumane Cave/RF49 X-2295-52 41,300 1300 A3/78-79/R818 Bone Large mammal 48140 43030 C/N: 3.3, % Coll.: 6.2,

d13C: �18.8, d13N: 4.7Fumane Cave/RF28 21733 41,000 1300 A4II/86g/R367 Bone Large mammal 47740 42800 C/N: 3.3, % Coll.: 2.9,

d13C: �18.8, d13N: 7.6Fumane Cave/RF29 21734 42,000 1400 A4II/100dþg/R1120 Bone Large mammal 49100 43610 C/N: 3.4, % Coll.: 3.3,

d13C: �20.5, d13N: 3.8Fumane Cave/RF30 21735 42,000 1700 A4/struct. II/744 Bone Mammal 49490 43430 C/N: 3.2, % Coll.: 0.8 (low yield),

d13C: �19.7, d13N: 5.8

Charcoal qualityCastelcivita Cave/No. 7 22622 36,120 360 rsa”, spit 11 Charcoal c.f. Ilex aquifolium 41910 40560 % C: 76.7, d13C: �24.0

% Aragonite-calcite / % C yieldCavallo Cave/Cvl 10 21072 19,685 75 D I Marine shell Cyclope neritea 23,380 22,470 100-0 / 9.3Cavallo Cave/Cvl 10 Dupl. 19,235 75 D I Marine shell Cyclope neritea 23,220 22,060 100-0 / 9.9Cavallo Cave/Cvl 2 19254 35,080 230 D 1 ¼ DIb Marine shell Nuculana sp. 40,450 38,860 100-0 / 9.8Cavallo Cave/Cvl 4 19255 36,260 250 D 2 ¼ DIb Marine shell Dentalium sp. 41,570 40,390 100-0 / 9.5Cavallo Cave/Cvl 11 20631 36,780 310 D II Marine shell Dentalium sp. 42,010 40,880 99.8-0.2 / 9.9Cavallo Cave/Cvl 6 19257 42,360 400 D 3 ¼ D II Marine shell Bivalve fragment 45,990 44,640 100-0 / 10.2Cavallo Cave/Cvl 8 19258 36,000 400 D 8 ¼ DII? Marine shell Dentalium sp. 41,570 39,560 100-0 / 9.7Cavallo Cave/Cvl 5 19256 39,060 310 E 1 ¼ E-D Marine shell Dentalium sp. 43,510 42,350 99.7-0.3 / 10.1Cavallo Cave/Cvl 5 X-2280-16 38,300 400 E 1 ¼ E-D Marine shell Dentalium sp. 43,380 42,080 100-0 / 6 (low yield)Cavallo Cave/Cvl 3 19242 39,990 340 E 4 ¼ EII-I Marine shell Dentalium sp. 44,300 43,000 50-50 / 8.5

Klissoura Cave 1/Klis 19 19936 27,950 160 V/AA4b/185-190 Marine shell Cyclope sp. 32,100 31,260 100-0 / 6.8 (low yield)Klissoura Cave 1/Klis 21 21068 34,580 220 V/AA4b/180-185 Marine shell Dentalium sp. 39,930 38,540 15-85 / 9

Conventional ages are expressed in 14C years BP and are calibrated in OxCal (Bronk Ramsey, 2009) using the IntCal09/Marine09 curve (Reimer et al., 2009). The right handcolumn lists analytical parameters for collagen and charcoal quality as well as percentages of aragonite to calcite (based on high-precision XRD analysis) for shell samples. Thedates for sample Cvl-10 are too young to be considered Uluzzian and are therefore excluded from the Bayesian models.

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e136

also SOM). Layers A2-A1 are classified as Protoaurignacian whilelayers D3d, D3b and D3a are later Aurignacian units (Broglio et al.,2003). D1d has yielded a small number of Gravettian artifacts and isattributed to the Gravettian period (Broglio et al., 1996e1997) (seealso SOM).

Klissoura Cave 1 is a small limestone cave located at the north-eastern edge of the Argive plain in the Peloponnese, southernGreece. It was investigated between 1994 and 2006 by Greek andPolish teams (Koumouzelis et al., 2001; Karkanas, 2010) thatrevealed an 8m long sedimentary succession. The cultural sequencewas divided into two broad phases: the Middle Palaeolithic phase,about 6.5 m thick, and the Upper Palaeolithic-Mesolithic phase,about 2.0 m thick, separated from each other by clear erosionalcontacts. A series of Middle Palaeolithic layers (XXa-geVII) is fol-lowedby layerVI, amixofUpperPalaeolithic andMiddle Palaeolithicelements since this layer isdirectlyoverlainby theUpperPalaeolithicsequence in several areas. Layer V is a thin, discontinuous lensoidhorizon, its contents ascribed to the Uluzzian and roughly comparedwith the ‘evolved’ phase of the technocomplex. It is a dark grey,clayey silty layer comprising mainly reworked and locally in situburnt remains. A series of Aurignacian layers (early: IV, middle: IIId-g, late: IIIa-c) overlaid layer V, and were followed by Gravettian àpointes à dos indifférencié or Mediterranean backed bladelets (III0),Epigravettian (IIa-d) and Mesolithic layers (3e5a) (Fig. 2)(Koumouzelis et al., 2001;Kaczanowskaet al., 2010;Karkanas, 2010).

Results

New chronological framework

In the current project, 18 radiocarbon measurements were ob-tained for the Uluzzian layers of Cavallo, Castelcivita, Fumane andKlissoura 1 (Table 2). Shell, charcoal and bone collagen were the

three types of dated material. Bones were dated only in the case ofFumane. In all other sites tested bones preserved no collagentherefore alternative material was used. All determinations wereproduced using the latest preparative methods (ultrafiltration forbones, ABOx-SC for charcoal, CarDS for shell), which have beendescribed in detail by Brock et al. (2010), Douka et al. (2010a) andWood et al. (2012). For the vast majority of samples, carbon yieldsand sample size were within the expected standard ranges. In thecase where an unusual value was observed (variable d13C, low %C orhigh C/N), these determinations got an OxA-X rather than theroutine OxA- prefix (Table 2, see with notes on last two columns).

We obtained ten radiocarbon dates for Cavallo Cave, the refer-ence Uluzzian sequence (Table 2). With one exception (Cvl 6), alldeterminations were produced on marine shell beads (Dentaliumsp., Nuculana sp. and C. neritea), most likely used as personal or-naments. They come from the two upper phases, Evolved Uluzzian(E II-I) and Final Uluzzian (D II-Ib) layers. These determinationswere initially reported by Benazzi et al. (2011).

Most dates (OxA-19242, OxA-19256, OxA-19258, OxA-20631,OxA-19255, OxA-19254) are consistent with the stratigraphic po-sition of the samples and range from 35 to 40 ka BP, from top tobottom. Only OxA-19257 (D II), a small, indeterminate fragment of abivalve shell appears too old for its context (42 ka BP). It is notunusual for bivalve shells to be older than their context (see forexample discussion in Sivan et al., 2005; Douka, 2011) and wesuggest that this is indeed an old valve, either collected on purposeor accidentally brought to the site long after the death of themollusc. On the other hand, OxA-21072 (and its duplicate) obtainedfrom a C. neritea shell from D I, was much younger than expectedconsidering its position in the sequence. According to the excavator,D Ib was the latest Uluzzian layer at the site. This layer was sealedby D Ia, a stalagmite crust covered by volcanic ash (C), both forminga rather continuous layer across most of the excavated area. What

Figure 4. Bayesian modelling based on the new radiocarbon determinations from fourUluzzian contexts in Italy and Greece. There appears to be a clear chrono-stratigraphicprogression of the technocomplex, and close correspondence between the Italian sitesand the Greek case. The raw determinations are calibrated using IntCal/Marine 09curve (Reimer et al., 2009) and the OxCal platform (Bronk Ramsey, 2009). On the top ofthe graph, the NGRIP d18O record is shown (Andersen et al., 2006; Svensson et al.,2006) and the Greenland Interstadials, corresponding to ameliorated climatic condi-tions, are numbered. The age for the CI ash is indicated on the top of the sequence as agrey line at the sites it occurs, and as dashed line in Fumane where its presence in notconfirmed.

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e13 7

makes this determination more curious is that no other archaeo-logical material ever reported from the cave could be assigned tothis time period (w20 ka BP). We have no further details tomake aninformative suggestion or better explain this young determination,but we consider the shell as unrelated to the Uluzzian of Cavallo.The latter clearly ends around the time of the CI eruption whenairborne tephra was deposited and covered the top of layer D.

We obtained no dates for the lowermost Archaic Uluzzian phase(E III) due to the scarcity of suitable shell for dating, the absence ofbone collagen in all bones we tested, and the lack of charcoal in thestored collection. The antiquity of Archaic Uluzzian phase is,therefore, unknown but based on the corpus of the current de-terminations, independent stratigraphic evidence and estimationsof sediment deposition rates, E III must predate E II by at least 1e3millennia. The age of the first Uluzzian occupation at Cavallo can bedetermined securely through direct dating of E III or, failing that,dating of the latest Mousterian layer, which will provide a usefulterminus post quem for the start of the Uluzzian at the site. Ourgroup is currently working towards these possibilities.

At Fumane, we dated bone remains from layers A3 and A4, bothassigned to the Uluzzian (possibly Archaic and Evolved, respec-tively). The position of the samples and their relation to structuresand other cultural remains are discussed in the SOM. We haveobtained five new determinations on modified (smashed and cut-marked) bone, all treated with the collagen ultrafiltration method(Higham, 2011). The ages range from 39 to 42 ka BP (Table 2). Itappears that the phase in Fumane was relatively short and it ischronologically indistinguishable from the late Mousterian at thesite for which several radiocarbon determinations also exist(Higham et al., 2009; Higham, 2011).

The single determination we obtained from Castelcivita comesfrom the uppermost Uluzzian deposits at the site (rsa00, top of spit 11,square H14 I), which is normally associated with the final Uluzzianphase at Cavallo. This layer is found 50e55 cm below the well-characterized CI deposits (Giaccio et al., 2008) and therefore its ageshouldwell predate the CI eruption. The layer contained evidence ofhearths and the particular charcoal sample (cf. Ilex aquifolium) wasdated twice, as part of experimentation with two radiocarbon pre-treatmentmethods (ABA and ABOx-SC;Wood et al., 2012). The less-refined ABAmethod gave an age much younger than expected at 33kaBP (orw37.0e38.8ka calBP),whichpostdates thecalendarage forthe eruption. Pretreatment of the same charcoal samplewith ABOx-SC gave a reliable age of 36.1 ka BP (w40.5e42.0 ka cal BP) (Table 2).This newmeasurement is around threemillenniaolder thanourABAresult, as well as all other determinations previously obtained forUluzzianmaterial from Castelcivita (Table 1). This is the first time anUluzziandeterminationat the site predates the estimate age of theCIlayer that seals the sequence, and agrees well with the stratigraphicposition of the sample.

Just as with Cavallo, the start of the Uluzzian in Castelcivitacannot be established because there are no reliable chronometricresults below rsa00. Three previous determinations from theMousterian were obtained in the 1990s (GrN-13982: 39100 � 1300and GrN-13984: 42,700 � 900 from level 11/cgr, spit: 29e30; GrN-13983: 33800 � 1300 from level 11/cgr, spits 27e28) (Gambassini,1997). These, however, were made on burnt bone and, just as withthe previous young Uluzzian determinations, they should also beconsidered minimum ages for the Middle Palaeolithic at the site.

In Klissoura Cave, Uluzzian layer V appears locally contaminatedwith younger and older material at its contact with other layers(Aurignacian IV and Mousterian VIeVII). It compares best with thelater stages, Evolved or Final Uluzzian, of the Cavallo phasing.Recently, microtephra occurring as a sharp peak at the interface oflayers V and IV but also spreading further up the stratigraphic col-umn was identified as the CI (Lowe et al., 2012). This microtephra

peak acts as a terminus ante quem for the deposition of the Uluzzianlayer V, for which we obtained two dates on shell beads (Table 2).Both beads were reported as coming from square AA4, outside thedripline of the cave. OxA-21068 at 34.6 � 0.2 ka BP (39.9e38.5 ka cal BP) agrees with the position of the sample beneath the CI.The fact that this datewas produced on aDentalium sp. shell is all themore important given the presence ofDentalium beads also in ItalianUluzzian contexts, such as in Cavallo Cave. The second determina-tion, OxA-19936, obtained from a shell found 5 cm below the pre-vious one, is much younger (w28 ka BP). When we questioned thecertainty of this level assignation and looked into the excavationplans it became clear that in squares AA1eAA3 (and their extensionin AA4), and between 175 and 185 cm below datum, Aurignacianlayer IV truncates Uluzzian layer V at places and lies in direct contactwith Middle Palaeolithic layers VI and VII. Given this very young ageand the rest of the determinations from overlying Aurignacian layersthat centre between 33 and 30 ka BP, OxA-19936 originally assignedto layer V, is most likely to belong to layer IV or a later occupation(sub-layers IIIeeg).

Bayesian framework for inter-site comparison

In an attempt to place the Uluzzian in its most likely calendarage and palaeoclimatic context, as well as assess its temporal

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e138

evolution across the dated sites, all new radiocarbon de-terminations were calibrated using the INTCAL09 (atmospheric andmarine) curve (Reimer et al., 2009) and a Bayesian model was builtusing OxCal 4.1.7 (Bronk Ramsey, 2009).

The presence of the CI as a chrono-stratigraphic marker is alsoincorporated at the sequences where it occurs and is shown inFigs. 4 and 5. The age of the CI is taken as 39,280 � 110 years (DeVivo et al., 2001), an age estimate that has been produced froman average of 36 40Ar/39Ar measurements from 12 proximal de-posits. No macro- or microtephra has been identified in Fumane.

Sixteen determinations from Cavallo, Castelcivita, Fumane andKlissoura were included in the generated model (the two deter-mination on Cvl-10 were excluded since we hold that they are notreally related to the Uluzzian occupation of Cavallo). We set theresolution of the model at 20 years, and also ran an outlier detec-tion analysis. The modelled output is shown in Fig. 4. In reality, thestructure of the model consists of a few more determinations fromolder and younger phases, i.e., Mousterian and Aurignacian inFumane and Klissoura, published by Higham et al. (2009), Kuhnet al. (2010) and Higham (2011). In Fig. 4 we show the relevantUluzzian phases and determinations therein. We compare themodelling output against the NGRIP d18O record (Andersen et al.,2006; Svensson et al., 2006) tuned with respect to the Hulu CaveU-series chronology (following Weninger and Jöris, 2008) (Fig. 4).

Overall, the modelling of the results suggests that the Uluzzianat all sites started much earlier than previously thought, at orshortly before 45 ka cal BP, and persisted for as much as 5e6millennia, until about 39 ka cal BP. The modelled results are more

Figure 5. Probability distribution functions (PDFs) for the start and end boundaries ofthe Uluzzian at the studied sites, as well for the start of the Proto-Aurignacian at theItalian sites mentioned in the text. These were generated after modelling the newradiocarbon determinations, shown in Fig. 4, using the OxCal platform (Bronk Ramsey,2009). The start of the Uluzzian in Cavallo is given here as the PDF for the startboundary of the Archaic Uluzzian (E III). On the top of the graph, the NGRIP d18O recordis shown in blue (Andersen et al., 2006; Svensson et al., 2006) and the GreenlandInterstadials, corresponding to ameliorated climatic conditions, are numbered. The agefor the CI ash is shown on the top of the sequence as a grey line at the sites it occurs,and as dashed line in Fumane where its presence in not confirmed.

precise and the newly produced chronology for the technocomplexis older and more consistent than previous estimates (compareFigs. 3 and 4).

Discussion

The new chronology has wider archaeological implications forour understanding of the Middle to Upper Palaeolithic transition insouthern peninsular Europe (limited here to Italy and Greece).Below we discuss key points arising from this new chronologicalframework.

Origins and geographic spread of the Uluzzian

Two of the big questions surrounding the appearance of theUluzzian are its origins and the local pathway(s) of expansion.

Given that its makers were traditionally suspected to be Nean-derthals, the origins of the technocomplex were originally thoughtto derive from local denticulate Mousterian (e.g., Palma di Cesnola,1993). The recent disambiguation of the biological affinities of theCavallo teeth and their attribution to modern humans (Benazziet al., 2011) means that an external source of origin should besought, a wave of modern human dispersal responsible for theappearance of the Uluzzian.

The traditional east to west scenario for the spread of the UpperPalaeolithic in Europe, especially that of the Aurignacian culture(e.g., Mellars, 2004, 2006), cannot be sustained in the case of theUluzzian because no similar technocomplexes have been discov-ered in the Levant or further to the east of Europe. Recently, Moroniet al. (2013) suggested that the origins of the Uluzzian should besought in sub-Saharan Africa on the basis of distinct techno-typological similarities of the Uluzzian with assemblagesbelonging to the Middle Stone Age to Late Stone Age transition.These authors also suggested that the Uluzzian spread in southernEurope via an Adriatic coastal route (Moroni et al., 2013). The factthat both the Adriatic and the Peloponnese are part of thisgeographic spread may relate to the emergence of coastal areas ofthe Ionian-Adriatic during glacial periods, which could have madesuch a dispersal route available. Ferentinos et al. (2012) discuss thepossibility of seafaring around western insular Greece during theLate Pleistocene. The authors suggest that hominids (both Nean-derthals and early modern humans) were capable of seafaring ac-tivities, possibly encouraged by favourable regional coastalconfiguration during the Middle and Upper Palaeolithic. Unfortu-nately, the absence of reported Uluzzian sites outside the restrictedcircumference of Italy and southern Greece and in the intermediateregions between Africa and southern Europe, as well as the inun-dation of large tracks of land in the Adriatic and other coastlinesduring the Holocene, render the East Africa-to-southern Europe ahypothesis only based on techno-typological affinities of the lithictoolkit and other elements of the archaeological record (productionof beads, use of colourants, modern human affinities). Only theexcavation of additional sites and the identification and study offurther Uluzzian assemblages within and outside the tech-nocomplexes current geographical span will elucidate exactdispersal pathways and areas of Uluzzian influence.

Even within the limited geographic area of Italy, where the ma-jority of the archaeological evidence stems from, the debate onwhether the Uluzzian first appeared in the south, centre (Palma diCesnola, 2004; Ronchitelli et al., 2009) or the north (Peresani,2008) of the peninsula remains open. Scenarios of delayed coloni-sation of the south by northern Uluzzians (e.g., Kaczanowska et al.,2010) have been recently put forward. Here, we attempted to useour new data to answer this question. Ordering of the probabilitydistribution functions (PDFs) that were generated by the Bayesian

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e13 9

model for the start of the Uluzzian phases at each site (Fig. 5) allowsus to estimate the chances for an Uluzzian layer at a site being olderthan that at another site (Table 3). In the absence of independentstratigraphic markers to confidently identify one context as beingcontemporaneous, younger or older than another, we require a startboundary to have significantly more chances of being earlier thananother start boundary. The results of this ordering exercise suggestthat the generated age estimate for the start of the Uluzzian in theItalian south, represented by Cavallo, appears to be older (70%chances) than that of Fumane in the north of Italy. However, thisshould be seen only as a tentative hypothesis given that the lower-most Uluzzian levels and final Mousterian levels at Cavallo (andCastelcivita) remain undated and unconstrained in absolute terms,while those of Fumane are. Improved chronology might change thepicture. With regards to the other sites, the Uluzzian in Klissouraremains the youngest of all.

Based on the current data, the only valid conclusion one canreach is that the Uluzzian is already present both in southern andnorthern Italy by 39e40 ka BP (43e46 ka cal BP). Howmuch earlierbefore that, is not known. The evidence fromGreece is limited so farand appears rather late. It is expected that comprehensive dating ofmore existing sites will help to answer questions revolving aroundthe origins of the Uluzzian technocomplex.

From a climatic point of view, the appearance of the Uluzzian aswe currently define it, falls at the end of Greenland Interstadial 12(GIS 12) and the cold stadial between GIS 12 and 11.While Cavallo EIII lacks direct determinations, the Bayesian model tentativelyplaces its age well within GIS 12. GIS 12 follows immediately afterHeinrich Event 5 (HE5), a climatic low at w48 ka. Müller et al.(2011) have suggested, on the basis of environmental proxies,that the initial arrival of anatomically modern humans is very likelyto have occurred post-HE5 and during GIS 12. The independentnew evidence from the Uluzzian sites in our study seems to agreewith this suggestion. The decrease in temperature inferred from thecomposition of the faunal assemblage in Cavallo layer E III (Boscatoand Crezzini, 2011) and at levels A4 and A3 in Fumane (Tagliacozzoet al., 2013) may therefore correspond either to the onset of HE5 orthe stadial conditions between GIS 12 and 11. If the latter is correctthe start of the Uluzzian would slightly postdate GIS 12.

The Uluzzian and its relationship to the Aurignacian in Italy

In southern Italy, Castelcivita (and Cala, not examined in thisstudy) is a crucial archaeological sequence because the Uluzzian isdirectly followed by a series of Aurignacian layers. Although these

Table 3Likelihoods generated after ordering the posterior probability distributions (pdfs) of thboundaries for three Italian Aurignacian sites mentioned in the text.

Probabilityt1 < t2

ULUZZStart and end

t1

Fumanestart

Fumaneend

Cavallostart

Cavalloend

Castest

U Fumane start 0.0000 0.9994 0.2748 1.0000 0.8L Fumane end 0.0006 0.0000 0.0019 0.9960 0.2U Cavallo start 0.7252 0.9981 0.0000 1.0000 0.8Z Cavallo end 0.0000 0.0040 0.0000 0.0000 0.0Z Castelcivita start 0.1875 0.7468 0.1235 0.9989 0.0I Castelcivita end 0.0093 0.2670 0.0058 0.9519 0.1A Klissoura start 0.0063 0.2184 0.0056 0.7462 0.1N Klissoura end 0.0000 0.0004 0.0000 0.1620 0.0A Mochi start 0.0268 0.8466 0.0194 1.0000 0.5U Fumane start 0.0005 0.5267 0.0017 0.9925 0.2R Serino start 0.0001 0.0143 0.0001 0.5854 0.0

The pdfs corresponding to this ordering are shown in Fig. 5.

Aurignacian layers are currently undated, their age is constrainedby two ‘known-age’ horizons: the uppermost Uluzzian layer rsa00

dated here between 40.6 and 41.9 ka cal BP (Table 2) and the CIvolcanic debris of estimated calendar age around w39.3 ka. Thissuggests that the Aurignacian in Castelcivita arrived soon after 41e42 ka and advanced in two successive forms: initially as Proto-aurignacian with Dufour bladelets, and later as a local variantincorporating tiny ‘Castelcivita micropoints’ (Gambassini, 1997).The Aurignacian lasted at the site for the next w2e3 millenniawhen the sequence was completely sealed by the pumices and greyash of the CI eruption (Giaccio et al., 2006). At a nearby open-airsite, Serino (Fig. 1), the CI covers Aurignacian deposits with raremicropoints similar to those of Castelcivita’s Protoaurignacian(Accorsi et al., 1979). Recently, four charcoal samples from a distincthearth in Serino were dated using ABOx-SC at w34.5 ka BP (Woodet al., 2012). When calibrated, these determinations correspondexactly to the accepted calendar age of the CI eruption at 39.3 ka(De Vivo et al., 2001). Based on these two archaeological sequences,we can suggest that Protoaurignacian groups had spread intowestern and central Campania by 39 ka and even earlier, but notbefore 41e42 ka.

Moving to northern Italy, the Aurignacian has been securelydated in two sites: Fumane in Verona andMochi inwestern Liguria.The Uluzzian in Fumane seems to have ended shortly after 42 ka calBP, and there seems that occupation of the site came to a halt forabout a millennium, and possibly more, until the arrival of the firstAurignacians. The Aurignacian chronology for Fumane wasdescribed by Higham et al. (2009) and Higham (2011) who, on thebasis of new determinations for ABOx-treated charcoal and ultra-filtered bone collagen, placed the earliest Protoaurignacian layer A2at 40.5e41.2 ka cal BP. This mirrors very well the estimated age forthe spread of the Protoaurignacian in Castelcivita. Unlike Castelci-vita, where occupation terminates with the CI eruption, Fumane isvisited by Aurignacians until about 36e38 ka cal BP as evidenced bythe cluster of dates at around that time (Higham et al., 2009;Higham, 2011).

In Riparo Mochi (Fig.1), evidence for older Aurignacian presencewas reported by Douka et al. (2012). A series of 15 radiocarbondates, obtained mainly on shell beads and one charcoal sample,were used to date the largest part of the stratigraphic sequence. NoUluzzian is found at the site, and a semi-sterile layer, currentlyunder study, follows the final Mousterian layer. The new radio-carbon determinations place the start of the Protoaurignacianoccupation at the site (layer G) at about 37 ka BP or betweenw42.7and 41.6 ka cal BP. Layer G of Riparo Mochi is therefore the oldest,

e start and end Uluzzian boundaries for the four studied sites as well as the start

IANboundaries

AURIGNACIANStart boundaries

t2

lcivitaart

Castelcivitaend

Klissourastart

Klissouraend

Mochistart

Fumanestart

Serinostart

125 0.9907 0.9937 1.0000 0.9732 0.9995 0.9999532 0.7330 0.7816 0.9996 0.1534 0.4733 0.9857765 0.9942 0.9944 1.0000 0.9806 0.9983 0.9999011 0.0481 0.2538 0.8380 0.0000 0.0075 0.4146000 0.8987 0.8676 0.9998 0.4328 0.7273 0.9947013 0.0000 0.6775 0.9950 0.0317 0.2458 0.9257324 0.3225 0.0000 0.9385 0.1138 0.2138 0.6885002 0.0050 0.0615 0.0000 0.0000 0.0044 0.1033672 0.9683 0.8863 1.0000 0.0000 0.8299 0.9980728 0.7542 0.7862 0.9956 0.1702 0.0000 0.9830053 0.0743 0.3115 0.8967 0.0020 0.0170 0.0000

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e1310

directly-dated Aurignacian assemblage in Italy. A second group ofyounger dates relates to an Early Aurignacian facies at around 32 kaBP or between 37.3 and 36.4 ka cal BP (Douka et al., 2012). Thesynchronous shell and charcoal radiocarbon determinations fromRiparo Mochi suggest strongly the reliability of marine gastropodsas an appropriate substrate for 14C dating, and dismiss large vari-ations of the local marine reservoir during the period at issue(contra Banks et al., 2013).

All other Protoaurignacian or Early Aurignacian industries inItaly either lack secure absolute determinations or have beenshown to be younger by several hundreds to a few thousand years(e.g., Bombrini, Paglicci, Fumane).

There is a clear chronological and stratigraphic separation be-tween Aurignacian and Uluzzian assemblages at the sites where thetwo co-occur. At no site is there evidence of interstratification be-tween the two technocomplexes and only the presence of some‘aurignacoid’ tools in the final phase of the Uluzzian offers limitedsupport that the two populations (Uluzzian and Proto-/Aurignaciangroups) could have met. however, on the basis of the observedspatio-temporal distinction we observe on our results we maysuggest that the transition from the Uluzzian to the Aurignaciancould have its roots in a secondary modern human populationmovement, which ultimately spread across Europe. The processesinvolved in this succession, whether those of population replace-ment, elimination or incorporation/absorption, remain unknown.

With regards to the advance of the Aurignacian, and thereforeits relationship to the final Uluzzian in the studied regions, theolder determinations from Mochi and the well-established con-nections of its Protoaurignacian inhabitants with both Italy andFrance, may hint at a northwest, possibly to northeast, and finallysouthern Aurignacian spread (i.e., from southern France to north-ern Italy and finally to southern Italy), from w37 ka BP(w42.5 ka cal BP) onwards. Although there is little evidence fromintermediate regions, this hypothesis corroborates the later agesthat exist for the Aurignacian in southern Italy (Paglicci, Serino,Castelcivita, Cala), where the industry does not appear beforew35 ka BP (39e40 ka cal BP).

The very small number of excavated and dated Aurignacian sitesin Greece does not permit a broad discussion on the chronologicalposition of the technocomplex, neither on its origins. Similarly tothe Italian south, a relatively late arrival for the Aurignacian may besuggested, since nowhere does the technocomplex predate 35 kaBP (e.g., Franchthi, Klissoura) (Kuhn et al., 2010; Douka et al., 2011).In terms of origins, a northern (Balkan) route is the most prevalentmodel, however, western/north-western influences from across theAdriatic cannot be excluded either.

Scenarios for co-existence between Uluzzians (modern humans) andlate Mousterians (Neanderthals)

We wanted to investigate how the older age estimates for theappearance of the Uluzzian in Italy and Greece may influencecontact scenarios between modern humans and Neanderthals.Reviews on the topic of Neanderthal and modern human contacthave focused almost exclusively on spatio-temporal overlap toverify or negate population contacts. Long periods of regional co-existence are bound to facilitate population contacts. This, inturn, would give rise to uni-/bidirectional flow of knowledge(acculturation) and/or of genetic material (interbreeding) leadingto the development of industries possibly with both MiddlePalaeolithic and Upper Palaeolithic characteristics, as well as hy-bridized human forms. One of the main points of refuting accul-turation scenarios between Neanderthals and early modernhumans has been the claim that there was no significant period ofcoexistence between the two populations (e.g., Zilhão et al., 2008).

It is now thought that Neanderthals may have survived in widerEurope until about 40 ka (e.g., Pinhasi et al., 2011), several millenniaafter the appearance of the Uluzzians in Italy and Greece, placed atw 43e45 ka. The claim of insufficient time for co-existence/acculturation therefore can no longer be sustained withoutfurther examination of the record for Neanderthal presence syn-chronous to the Uluzzian occupation of southern Europe.

Sadly, both in Italy and Greece final/terminal Mousterian con-texts, thought to represent the latest Neanderthal populations, arevery badly dated. In Italy, only Mochi and Bombrini in Liguria(w36e38 ka BP/41e43 ka cal BP) (Douka et al., 2012), Fumane Cave(41e42 ka BP/44e46 ka cal BP) (Higham et al., 2009) and, lesssecurely, Mezzena rockshelter (35 ka BP/w40 ka cal BP) (Longoet al., 2012) have been dated using the latest preparativemethods. In the case of Mezzena rockshelter, claims for lateNeanderthal survival were made on the basis of a single 14C mea-surement on mammal bone. However, a single measurementcannot provide the precision required for drawing strong conclu-sions and more extensive work is needed. In all other instances(Broion, S. Bernardino, Rio Secco, all’Onda, Buca della Iena, Breuil,Sant’Agostino, Reali, Castelcivita, Poggio shelter, Oscurusciuto)(Hedges et al., 1994; Gambassini, 1997; Caramia and Gambassini,2006; Milliken, 2007; Peresani and Gurioli, 2007; Riel-Salvatore,2007; Boscato et al., 2009; Ronchitelli et al., 2009; Peresani, 2011;Peretto, 2012), Mousterian determinations should be considered atbest minimum ages.

In Greece, the situation is worse since the record is patchy anddiscontinuous, and only a handful of poorly-dated sites have beenassigned to the Middle Palaeolithic period (Fig. 1). Mousterian siteswith absolute ages include Theopetra, dated to the last Interglacialor before (end of OIS 6/beginning OIS 5; Valladas et al., 2007),Asprochaliko imprecisely dated at w40 ka BP (Bailey et al., 1992),and Kalamakia at >40 ka (Darlas, 2007). Some later ages for find-spots close to River Peneios, between 28 and 44 ka BP (32e48 ka cal BP), have been attributed to a late persistence of Nean-derthals in the area (Harvati et al., 2009). Caution is advised,however, since determinations of such antiquity, produced usingconventional radiocarbon methods and less refined chemical pre-treatment protocols, must represent minimum ages at best. Klis-soura 1 and Lakonis I both contain final Mousterian assemblagesand they are currently the best-dated sites for this period in Greece.In Lakonis, late Middle Palaeolithic Unit Ib was dated between 39and 43 ka BP (43e48 ka cal BP) (Panagopoulou et al., 2004; Elefantiet al., 2008), although none of the determinations were treatedwith modern preparation methods for charcoal (ABOx) and there-fore may also correspond to minimum ages. At the same site, a‘transitional’/Initial Upper Palaeolithic (IUP) complexwith evidenceof Upper Palaeolithic tool types (Unit Ia) is associated with statis-tically identical determinations (38e42 ka BP/42e48 ka cal BP). Theexcavators have suggested that the industry of Unit Ia was theproduct of Neanderthals on the basis of a Neanderthal tooth foundthere (Harvati et al., 2003). The determinations fromUnit Ia predatethe Uluzzian evidence of Klissoura and there appears no obviousconnection between the IUP of Lakonis with the Uluzzian of Klis-soura in terms of material culture or lithic techno-typology.

Finally in Klissoura, some recently obtained radiocarbon de-terminations place the end of the Mousterian at about 41e40 ka BP(AA-73819: 40920 � 580, AA-73818: 41480 � 810; Kuhn et al.,2010) or at w44e45 ka cal BP. Further down that stratigraphy,layer VII is dated with one measurement (AA-73820:48990 � 1770 BP) at w47e51 ka cal BP. A chronological gap ofseveral millennia is identified in the determinations available forthe terminal Mousterian and the Uluzzian in Klissoura, whichcorresponds well with the stratigraphic discontinuities observed atthe interface of Middle Palaeolithic layers VIIeVI and the Uluzzian

K. Douka et al. / Journal of Human Evolution 68 (2014) 1e13 11

layer V. Again, Klissoura refutes scenarios of population overlap inthe Peloponnese. We conclude that no evidence to support modernhuman and Neanderthal co-existence/interaction can be found inGreece.

Overall, at all sites where a succession of Mousterian-Uluzzian-Aurignacian occupations exists, this always follows the indicatedsequence with no interstratifications. In addition, at most sites(including the sequences studied here (Cavallo, Castelcivita, Klis-soura)) the Mousterian is separated from the Uluzzian by a sedi-mentary break, a layer of volcanic ash or other stratigraphic hiatus,which may suggest that a period of time has elapsed between thetwo phases.

Taken together, while the antiquity of modern human presencein southern peninsular Europe (Italy and Greece) has beenextended significantly with the attribution of the Uluzzian tomodern populations, neither the available chronology nor thestratigraphic evidence of late Mousterian contexts may validateparallel existence and overlap between late Neanderthals and earlymodern humans in the region concerned (contra Longo et al., 2012;Zilhão, 2013). In turn, this may lead us to suggest that the Uluzziansarrived at ‘virgin lands’where Neanderthal numbers were very low,if at all present. This is in agreement with the hypothesis expressedby Moroni et al. (2013).

A further implication of this scenario, which (i) identifies theUluzzian as a product of modern humans, and (ii) allows itsdevelopment far from and independently of Neanderthal in-fluences, is the potential support it offers to the suggestion thatother transitional industries reported from Europe and the Levantmay indeed be the product of modern humans and not Neander-thals (Bar-Yosef and Bordes, 2010; Bordes and Teyssandier, 2011;Douka et al., 2013). We need, however, to remain cautious andsuggest that further detailed work is required if we were to un-derstand aspects of other transitional technocomplexes exhibitingsimilar characteristics to the Uluzzian, such as the Balkan Bach-okirian (e.g., level 11 of the Bacho Kiro) or the Bohunician (orEmiro-Bohunician) of Central Europe. A good start would be a freshand reliable absolute dating of these contexts.

Conclusions

We present here the first integrated chronological synthesis forthe Uluzzian, currently assumed to be the oldest modern humantechnocomplex in Europe. By using a series of new radiocarbondeterminations on shell, bone and charcoal substrate, all producedwith the latest preparative methods at the Oxford Radiocarbonlaboratory, and a purpose-built Bayesian statistical framework, weconstrained the age of the Uluzzian and compared it with that forprevious and succeeding technocomplexes (Mousterian, Aurigna-cian). We conclude that the Uluzzian arrived in Italy and Greece,where its only occurrences are identified so far, shortly before 45 kaand continued evolving until its final stagesw39 ka. Issues of originare not easy to disentangle at the moment, but similarities of theUluzzian tool techno-typology with assemblages in East Africa havebeen drawn. Yet, given the lack of intermediate assemblages be-tween Africa and southern Europe, as well as the absence of theUluzzian outside the limited geographical circumference of Italyand Greece, any quest for roots will remain challenging.

Regardless of where the Uluzzians arrived from, in addition tomodern human anatomy they also convey fully-modern behaviourexpressed through techno-typologically complex lithic toolkits,highly-adaptive subsistence strategies, bone tool industry, use ofpigments and productions of ornaments. These are part of the‘Uluzzian package’ since its earliest appearance in southeasternEurope and many of these features, such as bone tools and shellbeads, do not belong to the behavioural package of previous

populations (Neanderthals) living in the same region. The securechronology presented here for the Uluzzian elucidates the least-known aspect of the technocomplex and attempts to place thearrival of early modern humans in Europe at the right period. Thiswe hopewill enable convincing future comparisons of the Uluzziantechnocomplex with other transitional and early Upper Palaeolithicindustries in wider Eurasia.

Acknowledgements

We would like to thank R.E.M. Hedges for his support andguidance throughout the period that this work was undertaken.Several colleagues were instrumental to our study, and are speciallythanked: A. Palma di Cesnola, H. Klempererova, M. Koumouzelis, A.Moroni, F. Ranaldo and J. Riel-Salvatore. We would also like toacknowledge the continuing support of the Puglia and CampaniaArchaeological Superintendence branches, for facilitating excava-tion and study of the material. The dating work has been funded bya NERC-AHRC NRCF grant, as well as a major NERC grant (NE/D014077/1) to TH. KD was funded by IKY-Greece, the LeventisFoundation, the Leverhulme Trust and the European ResearchCouncil Grant “PALAECHRON” (ERC-2012-AdG–324139) during thecourse of this work. Research at Fumane is coordinated by theFerrara University in the framework of a project supported by theItalian Ministry of Culture e Veneto Archaeological Superintend-ence, public institutions (Lessinia Mountain Communitye RegionalNatural Park, Fumane Municipality, Veneto Region e Departmentfor Cultural Heritage), and private associations and companies(Cariverona Foundation, Banca di Credito Cooperativo dellaValpolicella).

Appendix A. Supplementary data

Supplementary data related to this article can be found at http://dx.doi.org/10.1016/j.jhevol.2013.12.007.

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