roebroeks (2001) - hominid behaviour and the earliest occupation of europe: an exploration

8/7/2019 ROEBROEKS (2001) - Hominid behaviour and the earliest occupation of Europe: an exploration

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Wil RoebroeksFaculty of Archaeology,Leiden University,P.O. Box 9515,2300 RA Leiden,The Netherlands.E-mail: W.Roebroeks@

arch.LeidenUniv.nl Received 12 October 2000Revision received 23 May2001 and accepted 23 May2001

Keywords : Europe,Pleistocene, colonization,hominid behaviour, hunting,palaeolithic.

Hominid behaviour and the earliestoccupation of Europe: an exploration

The last decade has witnessed a heated debate over the age and thecharacter of the earliest occupation of Europe. This paper addressestwo aspects of the debate, one dealing with the chronology of occupation, which is put to use in the second issue, an exploration of

the behaviour of the earliest occupants of Europe. The review of thedebate on chronology concludes that a short chronology applies toEurope north of the large mountain chains of the Alps and thePyrenees, where the earliest traces of a human presence date back toabout half a million years ago. In this phased-colonisation model, theMediterranean, and especially Spain, saw an earlier occupation,starting around the end of the Lower Pleistocene. The archaeologicalrecord of these rst Europeans suggests that from the rst presence innorthern Europe onwards, regular hunting of large game was com-mon practice among Middle Pleistocene hominids. By situating thisarchaeological evidence in the context of ndings from a range of other disciplines I develop a behavioural scenario which suggests that,

at its latest by the Middle Pleistocene, increased forms of socialcooperation, exchange of information within larger groups and ingeneral forms of behaviour based on a release from proximity hadbecome a standard ingredient of the hominid behavioural repertoire.

2001 Academic Press

Journal of Human Evolution (2001) 41, 437461doi:10.1006/jhev.2001.0499Available online at http://www.idealibrary.com on

Introduction

The present day distribution of the humanspecies is the result of a complicated processof dispersal from the inferred African homi-nid cradle that started probably around2 mya. Given the age and the number of hominid remains from East Africa, fewwould question that beyond all reasonabledoubt hominids evolved there (but see

Dennell, 2001 for a critical review). Thetimetable of the subsequent dispersal ishotly debated though, with recent evidenceeven pointing towards the possibility of aLate Pliocene to Early Pleistocene rst Outof Africa radiation ( Swisher et al ., 1994 ;Gabunia et al ., 2000 b; Balter & Gibbons,2000 ; Lordkipanidze et al ., 2000 ; Hu ff man,2001 ; but see also Langbroek & Roebroeks,2000 ). Within the context of this larger

discussion, the last decade has seen anintensive debate over the age and thecharacter of the earliest occupation of Europe (see for instance: Bonifay &Vandermeersch, 1991 ; Roebroeks & VanKolfschoten, 1994 , 1995 , 1998 ; Carbonellet al ., 1995 , 1996 , 1999 a , b; Dennell &Roebroeks, 1996 ; Turq et al ., 1996 ; Villa,1996 , 2001 ; A. Turner, 1999 ; Oms et al .,2000 ; Balter, 2001 ). The debate is far from

over, and new discoveries both within andoutside Europe have fuelled its intensity andhave led to the formulation of various com-peting models favouring a wide range of chronological scenarios. In the middle of allthis uncertainty and controversy, however,the debate has resulted in a kind of consen-sus view that the European archaeologicalrecord changes signicantly around 500 kaBP, with an increasing number of sites

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probably indicating a more substantialoccupation than in the period before (e.g.,Roebroeks & Van Kolfschoten, 1994 ;Dennell & Roebroeks, 1996 ; Dennell, 1998 ;Bermu dez de Castro et al ., 1999 ; Villa,2001 ). Most authors also agree now thatnorth of the large mountain chains of thePyrenees and the Alps the rst unambiguoustraces of human occupation date from about500 ka BP; the Mediterranean witnessedearlier human presence than the morenorthern areas, though the exact agediff erence between the southern and morenorthern parts of Europe remains to beestablished.

This paper consists of two rather di ff erentparts, each addressing an issue related to thetopic of the earlier occupation of Europe.First, I will give a short review of the pastdecade s debate on the earliest occupation of Europe, in which I will focus on the shortchronology ( Roebroeks & Van Kolfschoten,1994 ) and the implications and the signi -cance of the Atapuerca TD6 and otherSpanish nds for the debate. The data on

the earliest occupation history of Europe willbe put to use in the second, more explorativepart. This discusses the behaviour of thehominids responsible for the earliest occu-pation of Europe, with an emphasis on therst substantial settlement of Europefrom about 500 600 ka onwards. Thepast decade has seen a wealth of cross-disciplinary studies concerning variousaspects of biology and behaviour of Lower

and Middle Palaeolithic hominids. By usingvarious independent, but convergent, linesof evidence from a range of disciplines tocontextualise the archaeology of these rstEuropeans, I will follow Aiello s (1998) leadand try to set limits to archaeological specu-lations on their behaviour and to sketch abasic outline of the social organization of these hominids and the way this might haverelated to the colonisation of various envi-ronments. While the review of the debate onthe rst occupation is heavily data oriented,

the part of the paper dealing with the secondissue is considerably more explorative, pull-ing together data and ideas from a widenumber of sources in an attempt to advanceour knowledge of the issue. The resultingscenario makes some predictions on thecharacter of the archaeological record andhence can be tested in some aspects, but italso contains elements that are more di ffi cultto verify, such as statements on language ordemography. The fact that these conceptsare di ffi cult to operationalise in studyingthe fossil record should not prevent usfrom occasionally attacking these ques-tions, especially by means of independentyet convergent lines of cross-disciplinaryevidence.

Earliest Europethe debate

Reviewing the evidence pertinent to therst human settlement of Europe, I haveargued earlier that the European archaeo-logical record changes substantially around500,000 years ago. Together with T. van

Kolfschoten I have taken this as indicatingthat Europe was not colonised until aroundhalf a million years ago ( Roebroeks, 1994 ;Roebroeks & Van Kolfschoten, 1994 , 1995 ,1998 ; Dennell & Roebroeks, 1996 ; cf.Dennell, 1983 ). This so-called short chro-nology was based on a reassessment of artefactual and dating evidence from a largenumber of European sites, carried out in theearly 1990s. In our reading of the evidence,

there was a diff

erence between the Europeanrecord from before 500 600 ka BP and thelater period (500 600 ka BP is the estimateddate of the appearance of a biostratigraphi-cally important vole, Arvicola terrestris can-tiana , in northwestern Europe situated inthe second part of the Cromerian complex,about 500 ka BP). A recent study in Spainhas shown that there, as elsewhere inEurope, this transition and hence theBiharian to Toringian boundary occurswithin the Brunhes epoch (cf. Agusti et al .,

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1999). Before 500 600 ka virtually all ndscome from a coarse matrix, afterwards wehave primary context sites in ne-graineddeposits. The small assemblages dating frombefore 500 ka are virtually all the result of selection by archaeologists of primitiveworked pieces from natural deposits; bycontrast, younger assemblages are oftenexcavated from knapping oors. We pro-posed two basic ways to interpret thesedi ff erences. The pre-500 ka assemblagescould re ect the sparse traces of intermittentoccupation of Europe by people withprimitive , Oldowan -type toolkits di ff er-ent from later ones, while substantial colon-isation of Europe took place from about500 ka onwards (cf. Turner, 1992 ). How-ever, in our reading of the evidence, manypurportedly early archaeological sites didnot contain rm artefactual evidence for anearly human presence. For instance, this isthe case for Le Vallonet in France ( DeLumley et al ., 1988 ; see for critical com-ments on the lithic assemblage also White,1995 ; Santonja, 1996 ; Villa, 1996 ) and

the lower levels of Ka rlich (A and B) inGermany (cf. Baales et al ., 2000 ). Othersites were at least problematic from achronological point of view (such as MontePoggiolo in Italy, cf. Mussi, 1995; Villa,2001 ) and/or shown to be younger thanoriginally thought, such as Soleihac inFrance (Raynal et al ., 1995) and Isernia inItaly (see also Cuenca-Besco s et al ., 1999 ;Villa, 2001 ). Hence, in view of the attributes

of the artefacts and the contexts of thepre-500 ka sites (see Roebroeks & VanKolfschoten, 1994 , 1995 ), we interpretedthese di ff erences as strongly suggestingthat there was no indisputable proof forhuman occupation of Europe prior to about500 ka BP.

Our scenario had several advantages.First, it is supported by a body of dataindependent of arguments concerning stonetools: the chronological distribution of human remains. From about OIS 13

onward we had Middle Pleistocene humanremains all over Europe, whereas from thelong period before we did not have a singleuncontested tooth, despite the huge amountof other mammalian fossils known from thistime range. That is, until the AtapuercaTD6 nds were discovered ( Carbonell et al .,1995 ).

As a second advantage, the pattern wasbacked by a long history of intensiveresearch, as can be assessed by a look at thehistory of European palaeolithic archaeologyand mammalian palaeontology. Thenineteenth- and early twentieth-centurysearch for early humans was a pan-Europeanphenomenon, and every country had its ownsearchers and sets of early sites. As inEngland, which I have treated in detail(Roebroeks, 1996 ), all over Europe count-less exposures were subjected to closescrutiny and many presumed primitive artefacts were discovered in Early Pleis-tocene and Tertiary deposits (cf. Boule,1921 ; Obermaier, 1912 ; Sollas, 1924 ).Many of these have, over many years,

yielded rich mammal faunas, but never anyconvincing traces of human presence. Thisapplies to such di ff erent cases as the Sommevalley in northern France ( Tu ff reau &Antoine, 1995 ), the Lower PleistoceneTegelen clay pits in the Netherlands (e.g.,Dubois, 1904 ; see also Luttschwager & vanBemmel, 1962 ; Peeters et al ., 1988 ), theUpper Val d Arno basin in Italy, whosemammal fauna was already known to the

seventeenth-century naturalist NicolausSteno (cf. De Mortillet, 1883 ) and the pro-lic early Cromerian site of Su ssenborn nearWeimar (Germany), a site where Goethecollected fossils ( Nolte et al ., 1969 ). Thefailure of the many investigations looking forLower Pleistocene lithic assemblages andhuman fossils strongly suggests the absenceof hominids from, especially, northernEurope prior to the Middle Pleistocene. Thesouthern and eastern parts of Europehave been investigated for as long but not

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nearly as intensively as northern Europe,and hence there is a greater probability of surprise there.

The third and most important advantageof the short chronology was that its basicelements were very easy to falsify. As wewrote earlier: The nd of only one Early

Pleistocene site of primary context woulddisprove it, and one would have to concludethat before about 500,000 occupationexisted (but was largely intermittent). Newstudies of some of the sites mentionedin our survey could lead to such a result (Roebroeks & Van Kolfschoten, 1994 : 500).The nds from Atapuerca TD6 have doneexactly this, while the nds from the GuadixBaza basin in southern Spain seem to beeven older than Atapuerca, judging from thefauna associated with the artefacts ( Turq

et al ., 1996 ; Oms et al ., 2000 ). The TD6assemblage was recovered from depositsthat on palaeomagnetic evidence ( Pare s &Pe rez-Gonza lez, 1999 ) and on the basis of ESR and U-series dating of teeth ( Falguereset al ., 1999 ) have been situated at the veryend of the Lower Pleistocene (cf. Carbonell

et al ., 1995 ). Furthermore, the TD6 faunacontains the vole Mimomys savini and isthus earlier than all archaeological sitesmentioned above. Hence the pan-Europeanvalue of the short chronology has beenfalsi ed, and the Mediterranean seems tohave seen some earlier hominid presence.Exactly how much earlier is di ffi cult to say,though. As stressed elsewhere ( Dennell &Roebroeks, 1996 ; Van Kolfschoten, 1998 ),full acceptance of the Lower Pleistocene ageof the TD6 assemblage is not unproblematic

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Figure 1. Locations of the European key sites discussed in the text. 1, Ambrona; 2, Atapuerca; 3, BarrancoLeo n; 4, Biache-Saint-Vaast; 5, La Borde; 6, Boxgrove; 7, Ceprano; 8, Coudoulous; 9, Fuente Nueva; 10,Hoxne; 11, Isernia; 12, Ka rlich; 13, Mauer; 14, Mauran; 15, Miesenheim; 16, Monte Poggiolo; 17,Neandertal; 18, Notarchirico; 19, Salzgitter-Lebenstedt; 20, Scho ningen; 21, Soleihac; 22, Su ssenhorn;23, Swanscombe; 24, Taubach; 25, Tautavel; 26, Tegelen; 27, Torralba; 28, Le Vallonet; 29, Val d Arno;30, Wallertheim; 31, Weimar.

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for some palaeontologists: despite the rangeof dating techniques applied at Atapuerca,the TD6 dating evidence is not entirelyconsistent with the evidence from the widersetting of the site, as its fauna containselements which elsewhere in Europecommonly appear above the Brunhes Matuyama boundary ( Van Kolfschoten,1998 , 1999 ). Relevant for this discussion isthe stratigraphical age of the species withinthe Allophaiomys pliocaenicusMicrotus (Sten-ocranius ) gregalis lineage. M. (st.) gregaloidesand M. gregalis forms date from the earlierpart of the Middle Pleistocene, and thepresence of M. gregaloides indicates that thereversed part of the Trinchera Dolina llmay represent one of the intra-Brunhesreversals, according to Van Kolfschoten(1999) , though this argument has beencountered by Pare s & Pe rez-Gonza lez(1999) . There is clearly a biostratigraphicalproblem here, which needs to be resolved.No matter how large the time di ff erencebetween the rst occupation of southern andnorthern parts of Europe will eventually turn

out to be, what is important here is that theTD6 nds are earlier than any other goodarchaeological site known in Europe beforeits discovery. The palaeomagnetic andfaunal evidence from Fuente Nueva-3 andBarranco Leo n in the Guadix-Baza basin insouthern Spain points to an even older pres-ence of hominids in southern Spain, nowestimated to be pre-Jaramillo, minimallyaround 1 million years BP ( Oms et al .,

2000 ).The Atapuerca TD6 case seems to haveclosed the debate around the age of theItalian archaeological site of Isernia, for along time presented as a late LowerPleistocene, late Matuyama site on the basisof both K/Ar and palaeomagnetic dating(McPherron & Schmidt, 1983 ; Peretto,1991 ; Peretto et al ., 1983 ). Apart from thesomewhat controversial nature of thesedates (cf. Villa, 1996 ), the presence of A. terrestris cantiana in the Isernia fauna

was di ffi cult to reconcile with a LowerPleistocene age of the site (cf. Roebroeks& Van Kolfschoten, 1994 , 1998 ; VanKolfschoten, 1996, 1998; Von Koenigswald& Van Kolfschoten, 1996; Agusti et al .,1999; Cuenca-Besco s et al ., 1999; Sala,1999). Others workers stressed the mis-match between the macrofaunal evidenceand the inferred Lower Pleistocene age(Petronio & Sardella, 1999 ; cf. Villa, 2001 ).In recent publications the Isernia team hasplaced the site in the (early part of the)Brunhes epoch ( Gagnepain et al ., 1996 ;Peretto et al ., 2000 ), without any discussionof the debate reviewed here. Coltorti et al .(2000) recently reported an Ar/Ar date onsanidine from the archaeological level of 605 10 ka, which agrees considerablybetter with the fauna from the site than theprevious age estimates. Discrepancies be-tween various sets of data, as in the case of Isernia, should be addressed explicitly, asthey show where our knowledge of the issuesat stake is inadequate and hence new insightcan be gained. At the same time the Isernia

case shows how powerful a tool thebiostratigraphy of small mammals is, andbiostratigraphical subdivisions of thePleistocene are becoming increasingly morene-grained. Recent studies of small mam-mal assemblages from northern and centralEurope by S. Par tt (London) and T. vanKolfschoten (Leiden) have shown that themiddle part of the Middle Pleistocene ismore complex than hitherto envisaged

(personal communication, 2000).It cannot be overemphasised thatAtapuerca TD6 and the Guadix-Baza basinconstitute, for the time being, a chronologi-cal loner in the whole of Europe. It is tobe expected that more traces of a hominidpresence of about the age of Atapuerca TD6and the Guadix-Baza basin sites will eventu-ally be uncovered elsewhere in southernEurope (e.g., in Turkey, Greece, formerYugoslavia and Italy), but these simply havenot been found, or rather identi ed, yet. All

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other European sites thought to date fromthe late Matuyama or the early part of theBrunhes are problematic, and the new datesfor Atapuerca TD6 cannot refute the earlierdoubts on these sites. Each piece of evidencehas to be considered in its own right, not byconsistency with a fashionable interpret-ation. The Atapuerca TD6 nds in no waylend credibility to the disputed stone assem-blage from Le Vallonet or to the age esti-mate for Ceprano, Italy, of about 800,000years BP ( Ascenzi et al ., 1996 , 2000 ). TheCeprano skull is a stray nd and its ageassessment is based on stratigraphical corre-lations only (cf. Carbonell et al ., 1999 a : 668;Villa, 2001 ). Villa s (2001) study of theItalian evidence agrees in general terms withthe results of Mussi s (1995) earlier survey.Of the Italian sites considered as importantoccurrences that might date to the upperMatuyama or the early part of the Brunhes,none has yielded solid evidence for humanoccupation prior to the Isernia Notarchiricotime range (but see Milliken, 1997 1998 ).This means that there is no evidence for an

occupation prior to the Middle Pleistoceneand even prior to the appearance of Arvicolafaunas (cf. Piperno, 1999 ; Sala, 1999 ) .

If the Iberian evidence really is an excep-tion, a chronological loner (yet), how arewe to weigh this evidence? Are the ndstraces of a late Lower Pleistocene circum-Mediterranean settlement that gaineddemographic momentum (and archaeologi-cal visibility) around 500,000 years ago,

with the increase in population density lead-ing to an expansion into new areas? Or arethe TD6 hominid remains the markers of aLower Pleistocene hominid incursion intosouthern Europe that just failed to survive?The current archaeological and palaeonto-logical evidence is too meagre to choose oneof these options. Some scholars, however,suggest that at least two successive disper-sals of hominids into Europe can be distilledfrom the archaeological and palaeontologi-cal record (cf. Carbonell et al ., 1999 b;

Milliken, 2000 ; Otte, 2000 ). They see themore substantial settlement, from aboutthe Isernia Notarchirico Miesenheim timerange, as the result of an in ux of new-comers who carried a toolkit di ff erent fromthat of the rst settlers, referred to as Homoantecessor (Bermu dez de Castro et al ., 1997 ;Carbonell et al ., 1999 b). In this view, therst settlers toolkit was characterised by thedominance of a core and ake technologyand the absence of handaxes (e.g. Carbonellet al ., 1999 b). Indeed, handaxes appear onthe European scene only at about 500 600 ka, i.e., about one million years aftertheir rst appearance in Africa ( Asfaw et al .,1992 ) and also considerably later than inIsrael, at Ubeidiya and Gesher BenotYa aqov (cf. Goren-Inbar et al ., 2000 ).Some see this as evidence for a successivedispersal of two di ff erent cultural traditionsin Europe, with Lower Pleistocene Mode 1people preceding Acheuleans and a laterAcheulean wave displacing Mode 1 groups(Carbonell et al ., 1999 b). There are someproblems with such a model though, for

instance arguments over the age of many of these sites (see above, and Villa, 2001 ), thedisputed arti cial character of some of thekey assemblages (see above), the variabilitybetween so-called Mode 1 assemblages(compare Atapuerca TD6 with FuenteNueva-3), and the large and well knownvariability between artefact assemblagesfrom 500 ka onwards ( Villa, 2001 ). Variousworkers (e.g., Dennell & Roebroeks, 1996 ;

Villa, 2001 ) have also stressed the small sizeof some Mode 1 assemblages, whichmakes an attribution to technical traditionvery unreliable: TD6 has yielded 268 arte-facts (including unmodi ed pebbles andchunks, cf. Carbonell et al ., 1999 a ), whileFuente Nueva-3 and Barranco Leo n yieldedeach about 100 artefacts, mostly akes andsmall aking debris ( Oms et al ., 2000 ).Paola Villa has taken up this very pointrecently in a survey of the Italian and West-ern European evidence, concluding that the

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evidence for a greater antiquity of core-and-ake industries is ambiguous, and that alsothe interpretation of biface and non-bifaceindustries as designating di ff erent groups of hominids lacks a sound archaeological basis(Villa, 2001 ).

In summary, a short chronology stillapplies to Europe north of the large moun-tain chains of the Pyrenees and the Alps.When these parts of Europe were rstcolonised, occupation took place rapidly,geologically speaking. The Iberian evidencesuggests that there was at least some inter-mittent presence around the very end of theLower Pleistocene in southern Europe, witha (possible) hiatus of 200,000 to 300,000years separating the TD6 activities froma more substantial human presencestarting around the Isernia Notarchirico Miesenheim time range, i.e., about half amillion years ago [this situation shows aremarkable similarity to the record fromAtlantic Morocco, as described by Raynalet al . (1995): while some traces of occu-pation may date from just before the

Brunhes Matuyama boundary (Thomas1-quarry, Level L), the main part of the richAcheulean sequence at Casablanca, forinstance, dates from the second part of theMiddle Pleistocene]. Virtually all workersagree that the European archaeologicalrecord changes signi cantly after 500 ka BP,a communis opinio which is certainly one of the positive results of the past decade sdebate (e.g., Roebroeks & Van Kolfschoten,

1998 ; Carbonell et al ., 1999 b; Turner, 1999 ;Villa, 2001 ). Sites increase in number in theMediterranean, while north of the largemountain chains of the Pyrenees and theAlps the rst unambiguous traces of humanoccupation appear. It is clear, nally, that inorder to set limits on speculations about thewhen and how of colonisation processes weneed better chronological frameworks thanthose at our disposal now. The discrepanciesbetween various sets of dating evidenceshould form the point of departure for re n-

ing the timetable of Europe s earliest settle-ment, as it is in these problematic cases thatthe largest progress can be made. Whilemany workers suppose that Europe mayhave seen various waves of Lower andMiddle Pleistocene immigrants (cf. Goren-Inbar et al ., 2000 ), it is only through acareful, multidisciplinary approach towardsthe chronology of sites that eventually suchseparate phases of colonisation will becomearchaeologically visible.

Continuity of occupation?

For the time being, there is little evidence tosuggest (or contradict) that Europe saw acontinuous presence of hominids from theend of the Lower Pleistocene onwards. Abetter case can be made for a continuouspresence from about 600 500 ka. Studies of the environmental tolerance of the earliestEuropeans north of the large mountainchains have shown that these hominids werepresent in a wide range of environmentalconditions, from full interglacial up to and

including cold climatic conditions and moreopen environments, as testi ed by the evi-dence from the loess sequence at Ka rlich,Germany ( Roebroeks et al ., 1992 ). Thesevarious environmental settings do notinclude the coldest phases of glacial interglacial cycles and likewise do not implythat the whole of Europe saw a continuouspresence of hominids in the Middle Pleis-tocene. Hominid numbers may have uctu-

ated signi cantly in a given area from onecentury to the next, especially for hominidsnear the margins of ecological tolerance.While the northern areas may have seenan ebb and ow of human presence (cf.Gamble, 1986 ), southern parts of Europecould have functioned as refuges during themost severe parts of the glacials. As feasibleas this possibility may sound, is not observ-able in the archaeological record, wherecontinuity of human presence is di ffi cult toargue anyway, especially for the time range

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at stake here, with all its problems of chronological resolution.

Recent developments in the eld of Nean-dertal studies can possibly be of help here.According to workers like Arsuaga et al .(1997 b) and Hublin (1998) , the typicalNeandertals of the last glacial were the resultof an accretion phenomenon, beginning inthe middle part of the Middle Pleistocene,around 500 ka, if not earlier. The major coldstages of the last half million years increasedthe climatic stress on the initial colonisersand resulted in the isolation of Europeanhominids and the evolution of cold adap-tations, already thought to be visible in theapproximately 500,000 years old tibia fromBoxgrove in southern England ( Trinkauset al ., 1999 ). Added to the founder e ff ectfollowing an initial colonisation of Europeby small populations, climatic uctuationsmay have produced genetic drift episodesresulting in xation of derived features andhence a decrease of variability through time(Maureille, 1994 ; Hublin, 1998 ; see also:Hewitt, 2000 ). Middle Pleistocene homi-

nids and the Neandertals are increasinglyseen as an ancestral-descendance sequenceof populations without rupture of reproduc-tive continuity ( Arsuaga et al ., 2000 ; but seealso Hawks & Wolpo ff , 2001 ). Hence, mor-phological studies suggest a continuous andisolated presence of hominids on the basis of a kind of allopatric speciation process.

Recent genetic studies of the Neandertaltype specimen from the Feldhofer Grotte in

Germany come to a somewhat comparableconclusion ( Krings et al ., 1997 , 1999 ).These studies indicate that Neandertals didnot contribute mtDNA to modern humans,at the same time not ruling out the possi-bility that Neandertals contributed othergenes to modern humans. Using themtDNA sequence from the Neandertalspecimen, Krings et al . (1997) estimated theage for the most recent ancestral sequencecommon to the Neandertal and modernhuman mtDNA sequences at 550,000

690,000 years BP. A second study of themtDNA sequence of the Neandertal typespecimen yielded an estimate of 465,000years, with con dence limits of 317,000 and741,000 years ( Krings et al ., 1999 ). Suchestimates are to be treated with caution, asthey rely heavily on the calibration pointof the chimpanzee human divergence andhave errors of unknown magnitude. How-ever, they strongly suggest that the age of thecommon ancestor of Neandertal and mod-ern human mtDNAs is four times greaterthan that of the common ancestor of mod-ern human mtDNAs. More important here,they unambiguously situate the most recentcommon ancestor of Neandertal and mod-ern human mtDNAs in the rst half of the Middle Pleistocene, a date that agreesvery well with the occupation history datareviewed above.

The archaeological record shows thatMiddle Pleistocene hominids were presentin a large range of environments, over large(but not all see below) parts of Europefrom about half a million years ago, and we

have many sites from the middle parts of theMiddle Pleistocene onward ( Roebroekset al ., 1992 ; Gamble, 1995 ). Combined withthe genetic data and the accretion modelwith its focus on the development of European endemicity, I interpret this assuggesting that from about half a millionyears ago, Europe saw a continuous occu-pation by occasionally very small and rather isolated 1 groups of hominids . From about

half a million years ago Europeans, as rep-resented by the Mauer and Boxgrove fossils,gradually developed into early Neandertals,such as those from the Sima de los Huesos

1 No matter how important a chronologically signi -cant geographical isolation may have been in this Nean-dertalisation process, contacts between Europe andother parts of the Old World must have existed. This isindicated by the in Pleistocene terms almost con-temporaneous appearance of the Levallois technique inEurope, Western Asia and Africa at around 300 ka BP.

If not genes, at least techniques were freely exchangedaround that time.

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of Taubach, near Weimar (Germany),strongly suggests that the major part of thewoodland rhinoceros assemblage there(mandibular MNI of 62) was the result of deliberate hunting of 1 1 5 year old rhinoindividuals. Bratlund discusses at lengthother workers scepticism regarding rhinohunting in the Palaeolithic; using archaeo-logical, ethnohistorical and ethnologicalinformation she concludes that this scepti-cism is not based on empirical evidence butrooted in the attitude of earlier Europeancolonial hunters towards these animals(Bratlund, 1999 a : 135 146). The best par-allel to Taubauch in respect to species rep-resentation and preservation is the OIS 7site Biache-Saint-Vaast, mentioned above,though the focus on age classes di ff ers.

The environmental settings of these sitesare highly variable, from cold to warm-temperate, from open subarctic type of envi-ronments to full interglacial wooded habitats(Roebroeks et al ., 1992 ). The sites are situ-ated in various landscape types, from dis-sected limestone valley systems in southern

France via the montane regions of theGerman Mittelgebirge up to the loess plainsof northern Europe. The large size of manyof the prey animals from these sites, such asthe bovids at the OIS7 site Biache-Saint-Vaast, is strongly suggestive of full- edgedcooperative forms of hunting (cf. Geist,1978 : 274 278).

Independently of the archaeozoologicalstudies mentioned here, studies of the stable

isotopes from Neandertal skeletal remainsstrongly suggest that they were top-levelcarnivores, animal protein constituting animportant part of their diet (cf. Bocherenset al ., 1999 , 2001 ; Richards et al ., 2000 ).

The evidence for hunting in the LowerPalaeolithic is not as clear cut as for theMiddle Palaeolithic, though it remains to beestablished whether these di ff erences reallyre ect behavioural di ff erences or whetherwe are simply dealing with sample biasesand processes of preservation as suggested

by the evidence from a few sites, andespecially from Scho ningen in Germany(Thieme, 1997 , 1999 ). The Scho ningen sitehas become well known through the discov-ery of a series of wooden throwing spearswhich date to about 300,000 400,000 yearsago. These were found amidst the concen-trated remains of approximately 20 horses,with almost no bones from other speciespresent, and together with a few dozenint tools as well as a large number of intchips, probably the result of resharpeningof tools ( Thieme, 1997 , 1999 ; personalcommunication, 2000). The faunal remainsare even more exciting than the spears, asthey yield precious evidence on subsistencestrategies of the earliest Europeans unknownas yet from any other Middle Pleistocenesite. An ongoing taphonomical study of theassemblage by B. Voormolen (Leiden)shows that a large number of the horsebones display cutmarks, and that manybones have been processed for marrowextraction. Voormolen s preliminary resultson body part representation and the virtual

absence of carnivore gnawing marks stronglysuggest that the horse remains are the resultsof hominid hunting activities (Voormolen,personal communication, 2000). TheScho ningen nds were recovered over anarea of approximately 1000 m 2 , from theborders of a former lake, which hominidsmay have used to disadvantage the horsesbefore killing them at close range with thespears (L. R. Binford, personal communi-

cation, 2000). The high sedimentary rateenvironment, where sedimentation was bothrapid and calm, enabled a perfect preser-vation of hominid activities. Scho ningen isvery clearly an exception, but in this sub-sistence discussion such an exception carriesmore weight than the normal type of Lower Palaeolithic sites, such as the onesavailable when Binford wrote his Look atthe northern temperate zone (1985 : 315):sites from where we do have indicationsfor some human interference with faunal

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remains, though mostly in small numbersonly, and usually accompanied by a nor-mal background fauna with abundant evi-dence for carnivore involvement. Sites asSwanscombe ( Binford, 1985 ) and Hoxne(Stopp, 1993 ) in England, Ka rlich-Seefer(Gaudzinski, 1998 ) and Miesenheim ( E.Turner, 1999 ) in Germany, and Torralbaand Ambrona ( Santonja & Villa, 1990 ;Villa, 1990 ) in Spain have complex tapho-nomic histories, with material introduced byvarious agents and accumulated over some-times longer periods of time than seems tohave been the case for the Scho ningensite under consideration here. Even at highrate sedimentation sites such as Boxgrove(Roberts & Par tt, 1999 a ), interpreted ascontaining evidence for hunting of largemammals such as horse and rhinoceros(Roberts & Par tt, 1999 b), the archaeologi-cal evidence is often not unambiguous(McNabb, 2000 ).

Apart from a few open air sites, evidencefrom cave sites also suggests that huntingmight have been common practice in the

Lower Palaeolithic, as can be inferred fromMoigne & Barsky s (1999) study of thefauna assemblage from Level L of the Caunede l Arago at Tautavel, southern France,which is dominated by reindeer remains,with an MNI of 40. They interpret thereindeer as hunted and butchered, with thebones of adult individuals systematicallyprocessed for marrow extraction.

If, as I believe, one can generalise the

Scho ningen evidence for the whole of Europe around 400,000 years ago, thisimplies that by the mid part of the MiddlePleistocene occupants of Europe were ca-pable and active hunters of large mammals.At least by the time we see the rst occu-pants of Europe north of the Pyrenees andthe Alps, these hominids must have solvedthe overwintering problem of the temperatezone by hunting and eating animals that hadsolved the overwintering problem longbefore the arrival of the rst Europeans.

The archaeological data for this statementneed to be reinforced, though, by detailedpublication of the faunal assemblages fromrelevant sites, Scho ningen and especiallyBoxgrove, where important taphonomicissues still need to be clari ed. However, theconclusion ts very well with the ecologicalstudies of Geist, who paid considerableattention to the possible adaptations of therst colonisers of the northern temperateenvironments, and in fact predicted thespeci c top-level carnivore (Richardset al ., 2000 : 7663) adaptation, which a widerange of recent studies have come toacknowledge:

Man could not sustain himself by gatheringthroughout the year. . . . There was no roomfor a slow, herbivorous ape. There was alsoan abundance of carnivores roaming about,even in winter, such as cave lions, wolves . . .cave hyenas and wolverines. Only the super-carnivore niche is open, since the aforemen-tioned predators concentrate preferentiallyon medium-sized and small-bodied ungu-lates . . . because ungulates of large size,such as moose but also horses, and pre-sumably bison, mammoth and woollyrhinos are dangerous, capable opponents.. . . The only niche open is that of thesupercarnivore, who can despatch very large,slow, dangerous herbivores, and also thelarge carnivores themselves (Geist, 1978 :281 282).

It is, nally, worthwhile mentioning herethat intrinsic nutritional factors may havelimited at least seasonally the extent to

which hominids could rely on these animalproducts for calories (e.g., Speth, 1987 ,1991 ). Total fat levels in temperate andnorthern latitude large ungulates may dropby late spring to 2 3% ( Speth & Spielmann,1983 ), and during cold seasons recenthunter-gatherers may focus more speci -cally on the acquisition of fat or carbohy-drates, while high protein food resources,especially lean meat, can take on a verymarginal role or even be avoided entirely(Speth, 1987 ).

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Contextualising the archaeologicalevidence

How can we go further from the inferencethat from about half a million years agohominids were hunting large mammals inEurope? What kind of information can wepossibly distil from these observations on thelives of these earliest colonisers and thebroader social setting within which thesehunting activities were situated? I suggestfollowing Aiello s (1998) important lead andtaking an exploratory look at other disci-plines dealing with the history of our species,in an attempt to contextualise the archaeo-logical data with the independent results of other approaches and to develop newinsights on or at least set limits to specu-lations about the social life of the earliestEuropeans.

For instance, as we know from animalstudies, the rst settlers greater dependenceon hunting must have had consequences formany aspects of their lives, including the sizeof their home ranges: among primates

higher quality diets tend to be associatedwith larger home ranges, and carnivoreshave much larger ones than herbivores andomnivores. Also carnivore ranges are largerin high altitudes than in lower ones ( Geist,1978 , passim; cf. Gamble & Steele, 1999 ).This is related to the general trophic pyra-mid of any environment, and together withhunter-gatherers ethnographic data show ageneral increase in the area of land exploited

as dependence on hunting increases: movingnorth, resources tend to become more spa-tially segregated along a gradient of decreas-ing temperature, and the average distancemoved per residential move of hunter-gatherers tends to increase with decreasingtemperature ( Kelly, 1995 ). As huntingbecomes more important towards the polesand most mammals need larger territories tosupport themselves in colder latitudes, totalareas exploited in general increase consider-ably. This suggests that the rst Europeans

north of the large mountain chains werehunters who operated over large areas.There is, however, no archaeological evi-dence on the speci c size of these hypo-thetical large areas. The small amountof data we have for raw material transfers inthe European Lower Palaeolithic ( Fe blot-Augustins, 1997 ) indicate that raw materialmovements were generally short, and canonly provide minimum estimates of theareas over which early hominids operated.The database is somewhat better for theearly phases of the Middle Palaeolithic,where we see transported lithics connectsuch di ff erent regions as the northern Euro-pean plains with the German Mittelgebirgefrom at least OIS6 onwards, with stoneartefacts being moved over distances of more than 120 km as the crow ies(Roebroeks et al ., 1988 ; Fe blot-Augustins,1999 ; Gamble & Steele, 1999 ). However,these Middle Palaeolithic data also provideonly minimum estimates.

A very important contextualisation of thehunting data is a ff orded by the so-called

Expensive Tissue Hypothesis. Developed byAiello & Wheeler (1995 ; see also Aiello,1998 ), this hypothesis focuses on the largebrains of modern humans and states thatstudy of the cost of these large brains can bevery informative about the evolution of thehuman niche. Human brains constitute onaverage only 2% of total body weight, butthey use about 20% of all energy. Withyoung children the situation is even more

extreme: in the nal phases of pregnancy thegrowth of the foetal brain consumes about70% of all energy a ff orded by the mother,and in the rst year after birth the brain stillneeds about 60% of the child s total energyexpenditure. Aiello & Wheeler posit that theway in which humans can maintain thesecostly large brains without also having highbasal metabolic rates is simply by reducingthe size of one of the other expensive tissues,the gastrointestinal tract, the gut. This isonly possible by adopting a higher quality

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diet, which would release the constraints onencephalisation ( Aiello & Wheeler, 1995 ).As detailed by Aiello (1998) , the relation-ship between the energy requirements forthe growth of the brain and a high qualitydiet . . . is likely to have had additionalprofound implications for hominin life-history and social behaviour and, therefore,for the human niche , encompassingchanges in gender relationships.

Women, especially, paid a high pricefor the increase in brain size, becomingincreasingly dependent upon the help of others grandmothers, males in the pro-curement of high quality food for themselvesand their o ff spring (cf. O Connell et al .,1999 ). The increase in brain size occurringafter 500 ka BP would have put bothfemales and juveniles under increasedenergetic stress, and it is around this timethat increased cooperation between caringfemales and high quality food in thiscontext: fat-rich game (cf. Speth &Spielmann, 1983 ; Speth, 1990 ; Kelly,1995 ) provisioning males is predicted by

the hypothesis.2

It is also around this timethat the archaeological record, as I havediscussed above, contains the rst evidencefor hunting of large mammals, as illustratedby the Scho ningen evidence. Though themorphology of the Scho ningen spearsresembles to a high degree that of thecurrent Olympian sportswoman s javelin(Rieder, 2000 ; Steguweit, 1999 ), Aiello s(1998) line of reasoning indicates that males

might have been the more common users of these weapons.The Expensive Tissue Hypothesis predic-

tions about female male cooperation are not

easy to test, as there are relatively little dataon this topic. However, a recent statisticalanalysis of the rich Middle Pleistocenehominid material from the Sima de losHuesos at Atapuerca ( Arsuaga et al ., 1997 a )corroborates to some degree the hypothesis predictions, by showing that the degree of sexual dimorphism of this group of about 30individuals mostly young adults is com-parable to that of modern human popu-lations ( Arsuaga, 2000 ). This is importantevidence for a modern type of male female relationships in the middle part of theMiddle Pleistocene. While the Sima de losHuesos material dates from about 300 kaBP, more speci c information relating to thesexual division of labour comes from studiesof Late Pleistocene Neandertals only, andeven there the evidence is very limited.Analysis of Neandertal humeri suggests thatmales and females used their arms di ff er-ently, with males routinely engaged in activi-ties that needed considerable strength fromtheir right arms ( Ben-Itzhak et al ., 1988 ).Di ff erences in traumas of upper body and

head between male and female Neandertalsform another possible signal of a sexualdivision of labour ( Berger & Trinkaus,1995 ).

While the (not uncontested: Hladik et al .,1999 ) Expensive Tissue hypothesis o ff ers avaluable tool for contextualising the huntingevidence and interpreting it in terms of thesocial behaviour of the early Europeans,Tooby & DeVore (1987) have likewise

developed an interesting behaviouralecology-perspective on the social setting of hunting. Hunting of large game is a boom-or-bust activity. The variability in huntingsuccess and the fact that, unlike vegetableresources, meat comes mostly in chunks inexcess of what an individual capturer canreadily consume, indicates that there musthave been a strong evolutionary selection forfood sharing, food exchange and risk shar-ing through deferred reciprocation amongthe larger social group (Tooby & DeVore,

2 Such a bringing home the bacon scenario needsto be put in the context of nutritional studies, though.For instance, as detailed by Speth (e.g., 1991) , preg-nant women may have to obtain up to 70% of theirdaily calories from nonprotein sources; high proteinintakes would especially have posed a problem duringwinter and early spring in the northern latitudes, with

pregnant women being a vulnerable segment of thepopulation.

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1987 : 224). In Winterhalder & Smith s viewit was only with the evolution of reciprocityor exchange-based food transfers that itbecame economical for individual hunters totarget large game ( Winterhalder & Smith,2000 ). And for sharing the positive andnegative outcome of hunting activitiesbetween independently foraging subunits totake place, people had to have meetingplaces: Such a meeting area to exchangefood makes sense if the supply is irregular:either more (in the case of success) or less(in the case of failure) than the hunterswould need for themselves (Tooby &DeVore, 1987 : 224). Hunting made foodsharing necessary, while food sharing madehunting feasible.

Active cooperation between individualsseems also indicated by the hunting activi-ties themselves, as already pointed at above.Compared to most other hunting animals,humans are equipped with limited olfactoryand auditory capacities, humans cannot out-run most game, and have to outsmart prey.As discussed at great length, for instance by

Geist (1978 : 308 310 and passim), modernhunters are successful by virtue of knowingthe animals and the various traces they leavebehind in the landscape, and by using thisknowledge of animal behaviour as a tool (cf. Liebenberg, 1990 ). It is obvious thatknowledge of animal behaviour must havelikewise played a key role in successfullyhunting large mammals in the Palaeolithic,especially with the limited technologies at

stake here. Even if experiments suggest thatthe Scho ningen throwing spears were superbhunting weapons usable up to a distance of about 25 m ( Steguweit, 1999 ; Rieder,2000 ), the most important tool must havebeen an extensive knowledge of a wide rangeof animal behaviour. Combined with a keeninsight in a variety of other factors such asdiurnal changes in wind directions humanscent triggers the immediate departure of most game animals that knowledge ena-bled them to get close enough to a large

mammal, considerably less than 25 m, forsome certainty of a hit. As stressed byGuthrie (1997) such stalking activities aremore di ffi cult by an order of magnitude outin the open, away from tree-stands in theforest or forest edge. Indeed, many workershave emphasised the di ffi culties and com-plexities of hunting, for instance Geist (1978and passim) and Frison:

Animal procurement requires a body of knowledge that comes only with long-terminvolvement in predator prey relationshipsand familiarity with animal behavior underall conditions and at all seasons of the year.

The factors that control the success or failureof any animal procurement event are com-plex and ever-changing. Every animalspecies and individuals within the speciesdemonstrate unique behavioral characteris-tics depending on internal conditions of sex,age and animal condition and on externalconditions of time of year, weather, topo-graphic features, and vegetative cover. Mostof these conditions can and often do changerapidly: for example, a decrease in rainfalland/or deep winter snow a ff ects the avail-ability of grasses, which subsequently a ff ectsthe location of the animals and ultimatelydetermines the optimum procurementstrategy ( Frison, 1998 : 14,578).

Hunting the variety of species we nowknow from Lower and Middle Palaeolithiccontexts, in a variety of situations . . .required considerable experience, qualityeducation, and years of intensive practice.

From what we know about hunting today,we can see that such hunting was verydiffi cult (Guthrie, 1997 : 107; see alsoGeist, 1978 ) . Hence, again, in all likelihoodthere was a strong evolutionary stimulus oncooperation, and on sharing informationbetween the older members of a group andthe young and judging from the evidenceon their life history pattern these hominidsmay have had a prolonged pattern of matu-ration similar to that of modern humans(Bermu dez de Castro et al ., 1999 ), while a

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recent study of the Boxgrove tibia suggeststhat archaic humans may have reachedquite high ages ( Streeter et al ., 2001 ). Thequality education needed to become anexpert Pleistocene hunter could not dowithout a complex form of informationtransmittal interaction in which the tran-scendence of the here and now , a release fromproximity ( Rodseth et al ., 1991 ) played akey role.

This brings me to the last exercise of thisexploration in contextualising the huntingdata, i.e., to a scenario for the social originof language, based on comparative mor-phology and the ethology of primates, whichts well with the implications of the archaeo-logical evidence for hunting in the MiddlePleistocene as discussed above. Dunbar s(e.g., 1992 , 1993 , 1996 ) research has shownthat there is a clear relationship betweenthe relative size of the neocortex of the brainof primates, their cognitive group size(Dunbar, 1998 ) and the time they invest inphysical contact, in grooming. The relation-ship between neocortex size and group size

holds up in at least four other mammaliantaxonomic groups, an important support forwhat has become known as the Social BrainHypothesis ( Dunbar, 1998 ), according towhich primates relatively large brains arisefrom the information-processing demandsof their complex social systems. Startingfrom the relationship between the relativesize of the neocortex, group size and timeinvested in individual contact, Aiello &

Dunbar (1993 ; cf. Dunbar, 1996 ) suggestthat by the time of the appearance of thegenus Homo , groups had become so largethat a nonphysical form of social contact language became necessary for mainten-ance of the group s social cohesion. While arather rudimentary language may havebeen at stake with early Homo , the MiddlePleistocene encephalisation mentionedabove would indicate a group size not muchsmaller than is common with modernhumans.

Using the human neocortex size ratio andextrapolating a value for human group sizefrom the primate equation produces a valuein the order of 150 for modern humans.This number agrees well with actual obser-vations of the number of individuals humanstend to associate with. As stressed byDunbar (1998) , with modern humans theselarger groups are not (or only very rarely)conspicuous as physical entities, but they dohave important social connotations for theindividuals concerned: 150 is the number of individuals most people know as an individ-ual on a personal basis, in fact the de nitionused with other primates ( Dunbar, 1998 ;see also Steele, 1996 ). The observednumber of 150 also ts very well withthe results of Wobst s (1974) computersimulations to determine the minimum sizeof a viable human breeding population,which suggest that biological survival isdependent on a group size of minimally175. 3 The actual size of such cognitive groups and prehistoric breeding populationsmust have varied widely, in various complex

ways, determined by environmental factorsamongst others (cf. Kelly, 1995 ). The keypoint here, though, is that there are goodarguments to infer a minimum cognitive

3 While these larger groups are only conspicuous asphysical entities to the ethnographer during periodicalaggregations of the well-documented minimum bandsof about 25 individuals ( Kelly, 1995 ), the archaeo-logical visibility of any group size is an entirely di ff erentproblem. Recent studies of some classical UpperPalaeolithic sites thought to re ect the remains of aggregations of smaller groups into larger ones (forritual or other purposes), e.g. Dolni Vestonice andPavlov in Czechia, Mal ta in Siberia and Oelknitz inGermany, show that such earlier interpretations are notunproblematic, and that the rich and large sites in facthave to be reduced in the rst instance to accumula-tions of isolated depositional events, without any clearrelationship between the individual events. In otherwords, small groups could have reoccupied much thesame locations in successive years, leading to the accu-mulation of large amounts of material (respectively,Verpoorte, 2000 ; Vasil ev, 2000 , S. Gaudzinski, per-sonal communication, 2000). In this sense large groupsizes remain invisible on the ground until the estab-

lishment of the rst sedentary communities with anarchitectural record.

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group size of around 130 individuals by themiddle part of the Middle Pleistocene.

Following Aiello & Dunbar (1993) , withsuch a large group size it is impossible toin uence other members behaviourthrough physical contact. Group integrationcannot be performed on a personal contactbasis along, and a modern way of grooming,vocal grooming, language, must by then,i.e., about half a million years ago, haveemerged for developing and maintainingstable social relations. According to theSocial Brain Hypothesis language cameinto existence as a kind of social glue thatenabled displacement in both place and time(see Kay et al ., 1998 for a comparablechronology , based on a very di ff erentdataset).

In summary, we can situate the archaeo-logical indications for hunting of large mam-mals within the context of the ndings fromother disciplines, i.e., the physiological needof a regular supply of high quality food and astrong selection pressure on the sharing of information and the products of hunting and

gathering activities. The three independentsets of evidence reviewed here, (1) occu-pation history, (2) archaeological evidencefor hunting and (3) comparative anatomy,converge in the suggestion that the smalland mobile groups of individuals, which weknow from the archaeological record fromthis period, were integrated into larger unitssuch as Dunbar s cognitive groups . Theindividual members of such loose larger

groups collected information on the where-abouts of other individuals and smallergroups and the availability of food resourcesas they roamed the landscape in a kind of embedded learning, and shared this infor-mation with other members of the largercognitive group. Language made suchexchange of information about resourcesand coordination of foraging activities poss-ible. At its latest in the middle part of theMiddle Pleistocene, intensive, languagebased cooperation between individuals and

between local small groups had become astandard ingredient of the social behaviourof early hominids. While Dunbar s largergroups are not visible on the ground , inthe actual patterning of nds on thearchaeological site level, I suggest that theirexistence is at its latest indicated by thepresence of hominids in temperate Europenorth of the large mountain chains,half a million years ago. And so we havegradually moved from a simple questionabout the diet of these early settlers to thequestion of what they did in between meals,simply because both topics are inextricablyintertwined.

Hominid behaviour and thechronology of colonisation

Where does this explorative exercise take usin terms of the larger picture of the colonis-ation of the Old World, or the long-termtrajectory of hominid adaptations? Fromsites earlier than the rst occupation of thenorthern temperate zones we have no unam-

biguous evidence for hunting a la Scho ningenyet, and the fascinating evidence for canni-balism at Atapuerca TD6 ( Fernandez-Jalvoet al ., 1999 ) is di ffi cult to interpret in thiscontext. However, from the very rstappearance of stone artefacts in East Africaonwards we see hominid interference withbones in the form of cutmarks and conchoi-dal fractures ( De Heinzelin et al ., 1999 ),and somewhere in the more than 2 Ma

separating these earliest archaeological sites(Semaw, 2000 ) from Scho ningen, hominidhunting a la Scho ningen must have devel-oped. If not for the freak conservation con-ditions at Scho ningen, the European earlierMiddle Pleistocene record would resemblethe earliest Palaeolithic record in terms of human involvement with animals to ahigh degree. While one can make a goodpoint that the Scho ningen taphonomicalexception signals the behavioural rulearound 400,000 years ago, it remains to be

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established when and where such behaviourstarted and, again, possibilities shouldnot be confused with observables in thiscontext.

We run the risk of becoming trapped ina chicken-or-egg problem when askingwhether hunting capacities enabled themove to the north or whether northern envi-ronments stimulated the development of new adaptations. The evidence from Iserniadoes show some hominid involvement withthe rich faunal remains there ( Anconetani,1999 ), but its exact character is too ambigu-ous to be of help in choosing between thesetwo options, and the same applies to othersites in Italy ( Mussi, 1995 ). On the otherhand it is clear that in due time the archaeo-logical record might yield information todistinguish between cause and e ff ect forinstance, further excavations at AtapuercaTD6, or the current studies of the well-preserved faunal remains from GesherBenot Yaqov in Israel ( Goran-Inbar et al .,2000 ). However, for the time being it isimpossible to choose, and there is the

additional problem that we have little to noidea how to evaluate the role of factors suchas demographic developments, adaptivelimitations, environmental changes or basicphysical obstacles in human range expan-sion, certainly as long as the chronology of that expansion is still unclear in so manyimportant aspects.

To illustrate the importance of a solidchronology I can sketch a scenario starting

from the communis opinio that the Mediter-ranean was colonised earlier than Europenorth of the large mountain chains, and thatthe chronological di ff erences between therst appearance of hominids in both areasmay have been in the order of about200,000 300,000 years. A comparable timediff erence seems to exist between the rsttraces of occupation in northwest and cen-tral Europe and its easternmost parts, wherethe rst sites date from the later part of theMiddle Pleistocene ( Gamble, 1995 ; Praslov,

1995 research bias may play a role herethough). If this is a correct chronology,Europe was not settled more or less at thesame time, as implied by an earlier versionof the short chronology. Instead, the chro-nology of occupation of the various regionsconveys an ecological message, as it enablesus to relate the history of colonisation tothe variations in key resources observedwithin Europe (cf. Gamble, 1986 , 1995 ). Ingeneral, resources tend to become morespatially segregated along a gradient of decreasing temperature ( Kelly, 1995 : Fig-ures 4 7). On a European scale, going fromthe Mediterranean to the north of Europedistances between resource patches becomegreater, and are more easily stretched bychanges in the environment. Distances be-tween feeding patches are critical for mobilehominids, increases in these distances in-creasing the size of their regular moves,and the size of the areas over which groupsforaged. In the sketched chronology we see alate Lower Pleistocene hominid presencestarting in the diverse environments of

southern Europe, where for Italy Mussi(1995) has stressed the high variety of envi-ronments over short distances, a factor alsocalled upon to explain the rich Lower andMiddle Pleistocene record of the Sierra of Atapuerca. A few hundred thousand yearslater hominids also show up in northernEurope, even in the northern plains, in awide range of environments, from temperatewoodland conditions to colder steppic

settings as documented at Ka rlich H(Germany) ( Roebroeks et al ., 1992 ). I haverelated their presence and aspects of theiradaptive strategies to large group sizes andhence to the large spatial scales of their net-works. For later phases of the Palaeolithic,raw material studies have made abundantlyclear that in the Middle, as well as in theUpper, Palaeolithic raw material transferstend to be consistently larger in the eastern,continental parts of Europe than in the west(Roebroeks et al ., 1988; Fe blot-Augustins,

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1993 , 1997 , 1999 ). The more continental,harsher conditions increased the size of theareas over which groups had to hunt and toforage and hence the spatial scale of thenetworks these hominids regularly main-tained. The fact that a few hundred thou-sand years seem to have passed between arst presence in the western central parts of Europe and the rst hominid presence inthe Russian plains, around OIS 9 ( Praslov,1995 ), might indicate that the spatial scale of the networks initially did not match the en-vironmental conditions of the continentalparts of Europe, with their extreme seasonaltemperature ranges. The di ff erences in dis-tribution and movement of animal resourcesin these continental environments wouldhave required matching mobility strategiesand information gathering over larger areasthan necessary in the more western parts,where the oceanic in uence created moremosaic environments ( Gamble, 1995 ). Wecan follow this chronological pattern intoSiberia, where even more extreme seasonal-ity must have confronted the rst settlers

with new challenges. As recently shown byGoebel (2000) , there is little evidencethat hominids entered Siberia during theMiddle Pleistocene. Early in the UpperPleistocene, middle palaeolithic populationscolonised Siberia s southernmost latitudes,but these were generally limited to theheterogeneous environments of the south,below 55 degrees north, as were the earlyUpper Palaeolithic inhabitants of Siberia

(Goebel, 2000 ).The palaeoenvironmental data from theLower Pleistocene site Dmanisi (Georgia),at the gates of Europe, ts very well into thechronology of the scenario outlined above.The site was in the middle of a highlyproductive ecotone rich in animal and plantresources. The upland environment com-bined diverse landscapes within short dis-tances from each other, a vertical zonationre ected in both faunal and oral remainswhich indicate the prevalence of a warm and

moderately dry Mediterranean type cli-mate ( Gabunia et al ., 2000 a ). Whereas inearlier work the hominid remains were situ-ated at the base of the sediments overlyingthe basalt and were associated with moreforested conditions, hominid remains andartefacts are now seen as intrusive into thesedeposits ( Gabunia et al ., 2000 b), which inmy reading implies that they date to periodswhen the forested areas were considerablyreduced in size (see Table 2 in Gabuniaet al ., 2000 a ). One site does not constitute apattern, but the Dmanisi evidence mightindeed be read as indicating that the rangeexpansion of hominids from east Africa wasassociated with adaptations to middle lati-tudes where the upland environmentsfeatured very diverse resources (Gabuniaet al ., 2000 a :799).

This is certainly a too simple scenario,which furthermore assumes that there wascontinuity of occupation in Europe from theLower Pleistocene onwards. I have alreadydiscussed above that the early Iberian tracesof occupation may have been the result of

one or more incursions that failed tosurvive within Europe, and that the moresubstantial settlement after about 500,000could have resulted from another migratorypulse into Europe. As stressed by Dennell &Roebroeks (1996 :540 541), hominids seemto have occasionally moved into southernEurope well before half a million years ago,as and when conditions permitted, but with-out living there continuously . In this

scenario, the earliest hominids remainedcon ned to areas south of roughly 45 north,whether in Iberia, Georgia or China. Theabsence of Lower and early Middle Pleis-tocene sites north of the Pyrenees and Alpssuggests that even if hominids were aroundthe Mediterranean perimeter from the lateLower Pleistocene onwards, it required sig-ni cant changes in their behaviour to takethem north, across the many inland barriersof Europe. Alternatively, the evidence fromareas outside of Europe seems to suggest

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that there may have been other factorsinvolved in the colonisation of Europe.Work in Atlantic Morocco indicatesthat there too, a marginal hominid(Acheulean ) presence just prior to theBrunhes Matuyama boundary was separ-ated from a substantial occupation in thesecond half of the Middle Pleistocene(Raynal et al ., 1995). The similarity to theEuropean patterns suggests that there mayhave been more at stake than adaptations tonorthern latitudes per se. Potts VariabilitySelection hypothesis o ff ers an interestingperspective here, as it relates both Africanand European developments to the increas-ing amplitude in environmental uctuationsapparent in deep sea records and in terres-trial regional records, especially from theMiddle Pleistocene onwards ( Potts, 1996 ,1998 ). Potts has pointed to the temporalcorrelation between these environmentalchanges and the spurt in hominid brainexpansion described above. In his view, thiscorrelation is one expression of the evolutionof complex anatomical, cognitive and social

factors capable of processing and respond-ing to intricate and variable environmentalconditions: in short, the evolution of thekind of adaptive exibility indicated bythe various types of environments docu-mented in the European Lower Palaeolithicrecord.

Conclusion

This review of the past decade s debate onthe earliest occupation of Europe concludesthat a short chronology still applies toEurope north of the large mountain chains.The southern parts of Europe must haveseen some earlier incursions, around the endof the Lower Pleistocene, as shown by theevidence from Spain. The absence of othersites from that time range strongly suggeststhat this was a marginal, intermittent occu-pation. A substantial settlement of Europetook place from about half a million years

ago. From that date onward, a good casecan be built for a continuous presence of occasionally small and isolated groups of hominids, gradually developing into theclassic Neanderthals of the last glacial. Thearchaeological record suggests that theseNeanderthals were capable hunters of largegame, most probably from the very rstoccupation of northern Europe onwards.This higher dependence on hunting hadimportant consequences for the life of thesehominids, as I have shown in a contextuali-sation of the archaeological data. At about500 ka BP, increased forms of socialcooperation over large areas, exchange of information between individuals withinlarger groups, and in general forms of behav-iour based on a release from proximity had become a standard ingredient of the behavioural repertoire of these rstEuropeans. At the same time the chronologyand ecology of the colonisation of thevarious European regions suggest that thesize and the scale of these networks weresubject to long-term changes, which in the

long run may have enabled the occupationof the eastern, more continental parts of Europe together with changes in other,e.g., technological, domains, not discussedhere.

The settlement history of Europe mayhave seen several pulses of immigration,including the arrival of modern humansabout 40,000 years ago. The geographicalimpact, i.e., the size of the areas settled,

must have been dependent on variousfactors such as the adaptive capacities of theimmigrants and their demographic make up.This last factor especially is extremely di ffi -cult to grasp in an archaeological context. Itseems clear beyond any doubt, however,that around half a million years ago bothdemographical momentum and behaviouralcapacities enabled a substantial colonisationof large parts of Europe, and a virtuallycontinuous hominid presence of more than400,000 years.

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Acknowledgements

Thijs van Kolfschoten (Leiden) kindlyprovided me with results of his biostrati-graphical research, as did Boudewijn

Voormolen (Leiden) with his work on theScho ningen fauna. An anonymous Journal of Human Evolution Associate Editor and threelikewise anonymous reviewers made exten-sive helpful comments on an earlier draft of this paper, as did Sabine Gaudzinski(Monrepos). I want to thank Juan LuisArsuaga (Madrid), Lewis Binford(Dallas), Raymond Corbey (Leiden), PaulHennekens (Maastricht) and Alain Tu ff reau

(Lille) for inspiring discussions on some of the topics addressed in this paper. RobertFranciscus (Iowa) provided me with somevery useful literature references. I amgrateful to Marco Langbroek, BoudewijnVoormolen (Leiden) and a dozen Leidenstudents for their active participation in a2000 seminar series Food for Thought ,where some of the ideas presented here weretried out.

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