Dispersal and colonisation, long and short chronologies:how continuous is the Early Pleistocene record for hominids

outside East Africa?

Robin Dennell*Department of Archaeology and Prehistory, University of She!eld, She!eld S1 4ET, UK

Received 14 October 2002; accepted 1 September 2003

Abstract

This paper examines the evidence for hominids outside East Africa during the Early Pleistocene. Most attention hasfocused recently on the evidence for or against a late Pliocene dispersal, ca. 1.8 Ma., of hominids out of Africa into Asiaand possibly southern Europe. Here, the focus is widened to include North Africa as well as southern Asia and Europe,as well as the evidence in these regions for hominids after their first putative appearance ca. 1.8 Ma. It suggests thatoverall there is very little evidence for hominids in most of these regions before the Middle Pleistocene. Consequently,it concludes that the colonising capabilities of Homo erectus may have been seriously over-rated, and that even ifhominids did occupy parts of North Africa, southern Europe and southern Asia shortly after 2 Ma, there is littleevidence of colonisation. Whilst further fieldwork will doubtless slowly fill many gaps in a poorly documented LowerPleistocene hominid record, it appears premature to conclude that the appearance of hominids in North Africa, Europeand Asia was automatically followed by permanent settlement. Rather, current data are more consistent with the viewthat Lower Pleistocene hominid populations outside East Africa were often spatially and temporally discontinuous,that hominid expansion was strongly constrained by latitude, and that occupation of temperate latitudes north oflatitude 40( was largely confined to interglacial periods.! 2003 Elsevier Ltd. All rights reserved.

Keywords: Hominid dispersals; Colonisation; Lower Pleistocene; Asia; Europe; Eurasia; North Africa

Introduction

There has been considerable recent interest inwhen hominids first dispersed out of Africa andinto Asia and Europe. Most discussion of early

hominid dispersals has been about when theseoccurred, with some attention being paid to howoften. A third issue, of how successful these disper-sals might have been, has received much lessattention, and is the main focus of this paper.These three issues of timing, frequency and successare inter-related (see Fig. 1): for example, an earlydispersal allows more time for subsequent ones,

* Corresponding authorE-mail address: r.dennell@she!eld.ac.uk (R. Dennell).

Journal of Human Evolution 45 (2003) 421–440

0047-2484/03/$ - see front matter ! 2003 Elsevier Ltd. All rights reserved.doi:10.1016/j.jhevol.2003.09.006

but none might be successful, or a later dispersalevent might be isolated, but nevertheless result inpermanent colonisation. The success (or failure) ofa species to disperse into and colonise new areasprovides much information about its adaptivebehaviour, and thus the continuity of early homi-nid populations outside Africa deserves as muchattention as when they first dispersed, and howoften.

This paper examines the evidence for long-termcontinuity of occupation by hominids in areas intowhich they may have dispersed in the Pleistoceneprior to ca. 500 ka. It explores the possibility thatmost of the Lower Pleistocene record for hominidsoutside East Africa is one of occasional visitors,rather than long-term residents. Since the firstappearance of hominids in a region does notnecessarily mean that they became a permanentpart of the fauna, continuity of occupation needsto be demonstrated, rather than assumed. To date,most researchers have assumed that the earlyPleistocene fossil and archaeological record can be“read” in the same way as for the later Pleistocene,i.e. as a record of human populations that werelong-term residents in an area. This assumption isimplicit in numerous distribution maps, hominidphylogenies and summaries of lithic traditions.The underlying assumptions in these types ofsummaries are first, that the ability of earlyPleistocene hominids to colonise new areas wasessentially as “human” as that for the laterPleistocene; and secondly, that further fieldworkwill eventually fill the many gaps in what are oftenvery sparsely documented regional records. An

alternative viewpoint is that the distribution of, forexample, Acheulean and non-Acheulean industriesacross Eurasia is not comparable to a distributionmap of, for example, early Aurignacian and lateMousterian assemblages in Europe, as the data arederived from hominids with fundamentally di"er-ent technologies, modes of subsistence, socialbehaviours, and cognitive abilities. With Homoerectus, we are investigating an extinct species, thebehaviour of which is still very imperfectly under-stood; with humans of the last 40,000 years, we canobviously assume far more points of similaritywith ourselves. Just as studies of early hominidsubsistence in the 1970s were flawed by theassumption that modern hunter–gatherers were anappropriate analogue of Plio-Pleistocene homi-nids, so studies of early hominid dispersals mayhave been flawed by the assumption that thecolonising abilities of modern humans were anessential attribute of Homo erectus and otherarchaic hominid populations. (Contra Tattersall,1997: 49), we might question whether earlyPleistocene hominids had “a typically humanwanderlust” that prompted them to colonise Asiaand Europe). Instead of assuming that a fewhominid remains and archaeological finds that arewidely-spaced geographically and temporally indi-cates continuous regional occupation, it may bemore realistic to assume that they indicate apalimpsest of intermittent dispersal events, onlysome of which resulted in long-term colonisation.

Before we examine the evidence for the LowerPleistocene colonisation of Eurasia, we need someassumptions as to what we might expect fromHomo erectus.

Early hominid dispersals into Eurasia: someassumptions

Hominid dispersals across Eurasia can bemodelled in terms of a number of basic assump-tions derived from what is already known aboutearly Pleistocene hominids in Africa:

1. The earliest Eurasians preferentially occupiedgrasslands and open scrub- and wood-lands, as inEast Africa.

This assumption seems reasonable: Homoergaster/erectus in East Africa after 1.7 Ma is

Timing

Frequency Success

Fig. 1. Timing, frequency and success: the three key issues ininvestigations of hominid episodes of colonisation and disper-sal. Most attention has been given to evidence of when disper-sals occurred, rather than to evidence of subsequent settlement.

R. Dennell / Journal of Human Evolution 45 (2003) 421–440422

associated with hot and dry conditions, and opengrasslands; its post-cranial anatomy, with its longlimbs was geared to long-distance walking acrossopen ground, and to heat dispersal throughupright posture. H. erectus also utilised a widerrange of habitats than previous hominids, andmay have been a “weed” taxon that could takeadvantage of disturbed habitats (Cachel andHarris, 1998: 123). Although well-adapted tograsslands, it appears to have preferred mosaicenvironments that also contained permanent waterand some wood- or scrubland as well as grass-lands. Additionally, it appears to have avoideddiverse habitats, such as forests, in which potentialavailable biomass was higher, but more dispersedand less easy to extract.

2. Early Pleistocene hominids were primarilycarnivores, and thus a) annual ranges would haveincreased northwards and during periods of aridity,as food resources diminished; b) their ability tobecome long-term residents in a region would havebeen strongly influenced by their ranking in theprevailing carnivore guild as well as by the type andabundance of prey.

Although there is still active discussion ofwhether H. erectus was a hunter, a confrontationalor a passive scavenger, there seems general agree-ment that it was not primarily a vegetarian, andthat meat was a prized resource. Current researchappears to favour aggressive scavenging, possiblywith some predation, as the most likely mannerof food-procurement by early Pleistocene homi-nids (e.g. Dominguez-Rodrigo, 2002), ratherthan the type of passive scavenging proposed inrecent decades (e.g. Binford, 1981; Cavallo andBlumenschine, 1989). If we assume that H. erectuswas carnivorous, we can additionally assume thatthe size of its annual territories would haveincreased towards the northern limits of its range,because the quantity and quality of plant foodsupon which herbivorous prey depended would alsohave decreased from south to north. Likewise,annual ranges would have increased under con-ditions of increased aridity and/or in the driestparts of glacial-interglacial cycles.

Turner (1992) in particular, and also Hemmer(2000) have emphasised the importance ofcarnivore competition as a factor influencing the

ability of hominids to colonise a region. Morespecifically, Turner (1992) argued that the largefelids Megantereon and Homotherium, and thelarge hyaenid Pachycrocuta would have out-competed hominids in Europe between 1.8 and0.8 Ma. Only after their extinction, and thereplacement of Pachycrocuta by Crocuta wouldhominids have gained ready access to carcasses.

3. Hominid dispersals would have been addition-ally constrained as they dispersed northwards intoEurasia by a) the di!culties of maintaining social/mating networks whilst maintaining large annualranges under levels of low population density and b)winter day-light length, and its obvious restrictionson the time available for locating, obtaining andprocessing prey.

The ultimate limits of hominid dispersals north-wards would have been constrained by increasingdi!culties of maintaining viable mating networksas population densities decreased as well as theamount of food available in winter. These con-straints would have been less severe at lowerlatitudes, where the e"ects of seasonality and cli-mate change are more muted. Regarding winterdaylight lengths, African evidence indicates that by1.8 Ma, hominids were able to live at least 35(Nand S of the equator (i.e. from Ain Hanech,Algeria to Sterkfontein, S. Africa), and were thusable to tolerate mid-winter daylight lengths ofca. 7.5 hours. (Mid-winter daylight lengthdecreases by roughly 8 minutes per day with eachdegree of latitude). Latitude appears to haveseverely constrained early hominid expansion, as itwas not until 1.5 million years later that hominidscould survive at 55(N (at Pontnewydd, N. Wales)(Green et al., 1981).

4. Hominid abilities improved over time, and thusthe range of environments occupied would also haveincreased.

Hominids “were not simply dancing to therhythms of the Pleistocene” (Gamble, 1999: 125),and the range of environments and climates thathominids could tolerate increased over time astheir survival skills increased, and as their abilityto maintain social networks over large areasimproved. One would thus expect greater evidenceof continuity (whether in population, culturaltradition, use of a landscape) in the Middle than in

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 423

the Lower Pleistocene. This assumption appearsreasonable: evidence from Olorgesaillie (Pottset al., 1999) shows that hominids between 992 and790 ka were able to re-occupy areas after majorchanges in the environment, and able to cope witha wider range of environments than their latePliocene predecessors at Olduvai and Koobi Fora.Likewise, hominids in Europe south of 50(N wereable to cope with almost the full range of environ-mental conditions after 500 ka, except for theseverest glacial maxima, even if population densi-ties undoubtedly fluctuated in response to climaticchanges (Roebroeks, 2001). Hominids are evi-denced in a wider range of environments in theMiddle Pleistocene, and at times were expandingnorthwards beyond 50(N. Nevertheless, thesegains in adaptability would have been o"-set bythe climatic regime of the Middle Pleistocene, asindicated below.

5. Environmental disruption caused by glacial-interglacial shifts increased after 800 ka.

Hominids might not have been dancing to therhythms of the Pleistocene, but these rhythmschanged dramatically after 735 ka. Both theloess and deep sea records indicate that in thelower Pleistocene, there were numerous, lowamplitude climatic fluctuations, with a periodicityof ca. 41 ka, but in the Middle Pleistocene, after735 ka, there was a shift towards low-frequency,high amplitude alternations of glacial/interglacialclimate, occurring on a cycle of 100 ka (Ruddimanet al., 1986; Ruddiman and Raymo, 1988). Conse-quently, hominids might have improved theirsurvival skills, but also faced much greaterenvironmental disruption. Animal and plant distri-butions would have been subject to greater alti-tudinal and latitudinal shifts, and the ebb and flowof hominids would probably have been as marked,even if bu"ered by improved methods of obtainingfood, maintaining social networks, and survivingin harsh environments. However, in some situ-ations, environmental disruption might have ben-efited hominids. As Gamble (1999: 105) notes ofEurope “the long-term trend was towards longerperiods of cooler, more open continental con-ditions” which resulted in the breaking down offorest communities, and the creation of a mam-moth steppe, not particularly diverse, but rich in

large and medium-sized prey such as mammoth,horse, bison that were easily targeted by a profi-cient, carnivorous hominid. Anton (2002) has alsorecently pointed out that land-bridges broughtabout by the lowering of sea-levels duringglaciations would have lasted longer in theMiddle than in the Lower Pleistocene, and areassuch as Indonesia would thus have been lessisolated.

6. At any one time, there would have been coreand peripheral areas of hominid occupation; coreareas would have expanded as hominid survival skillsincreased and/or as climatic conditions permitted.

At a continental level, most pre-modernPleistocene hominids are thought to have lived inAfrica (e.g. Relethford and Harpending, 1994).Inter-regional hominid population densities wouldalways have been variable, and some areas (e.g. theRift Valley, the Levant, S.W. France) are oftenregarded as being core areas at various criticaljunctures in human evolution. At levels of very lowpopulation sizes and densities, hominid popu-lations in Eurasia are also likely to have beenspatially discontinuous, or only tenuously con-nected. Reasons might be because of mountain,desert or sea barriers; stochastic failures of repro-duction, disease (e.g. Bar-Yosef and Belfer-Cohen,2001), or climatic change. Discontinuities wouldalso have been temporal, and may have variedfrom a few millennia to substantial parts of glacial-interglacial cycles. Whatever the reason, the under-lying point is that continuity of residence needs tobe demonstrated, not assumed. As a salutaryreminder, Ashton and Lewis (2002) have recentlysuggested that Britain may have been uninhabitedin oxygen isotope stage 5 and possibly much ofstages 4 and 6, i.e. between 60–150 ka; and evenfully modern humans had to abandon much ofnorthern Europe at the height of the last glaci-ation. Discontinuities of occupation are even morelikely in the earlier Pleistocene.

The Pleistocene record for hominids in the OldWorld can thus be seen as a game between one sideof increasingly proficient hominids, and anotherside of an increasingly disruptive climate. If earlyHomo erectus was a skilled carnivore, it shouldthus have enjoyed considerable success as acoloniser of Eurasia in the Lower Pleistocene,

R. Dennell / Journal of Human Evolution 45 (2003) 421–440424

when the severity of climatic fluctuations was lesspronounced than after 0.735 Ma.

There is no general agreement over how wemight recognise continuity of regional hominidrecords over a million-year span of the LowerPleistocene. One indicator is an assessment ofspatial and temporal frequency of hominid occur-rences. Villa (2001: 126), for example, has recentlyestimated that in Italy (covering ca. 100,000 sq mi.)during the Middle Pleistocene, there is an averageof only one site per 100 ka, and existing data “fitan ad hoc interpretation of multiple, sporadic anddiscontinuous episodes of settlement”. She wouldthus suggest that there has to be more than one siteper 100 ka per 100,000 sq. mi. as a minimumrequirement of site density before continuity ofoccupation can be assumed. Another approachwould be to consider whether evidence of homi-nids occurs in both cold and warm periods ofglacial cycles; i.e. if hominids are present in both, itmight be reasonable to assume that they werecontinually resident. A variant of this approach isto consider the frequency of occupation in a localeunder di"erent conditions, as an indicator of theresilience and flexibility of a local hominid popu-lation: an example is the study by Potts et al.(1999) of the Olorgesaillie sequence. Anotherapproach might be to consider evidence of conti-nuity in artefact design, but this is unlikely to berewarding when dealing with the unstandardised,informal type of lithic assemblages found in mostof the Lower Pleistocene. Likewise, evidence ofcontinuity in regional fossil hominid records isonly possible when there are numerous well-datedobservations (as in later Middle PleistoceneEurope, and possibly Lower Pleistocene Java). Inthis paper, a combination of these approacheswill be used, according to whichever is themost appropriate for the regional record underconsideration.

In order to assess whether Lower Pleistocenehominid dispersals from sub-Saharan Africaresulted in long-term colonisation, we can begin byconsidering the evidence for hominids dispersalsacross Eurasia and North Africa ca. 1.8 Ma andthen their environmental contexts. This time-line isselected because it may indicate the earliest disper-sal of hominids into Eurasia, and is also the

base-line of the so-called “long” chronology (e.g.Bosinski, 1992; Gibert, 1992; Carbonell et al.,1995a, 1999; Tattersall, 1997; Gibert et al., 1998,2001; Aguirre and Carbonell, 2001). Eurasia isshown in Fig. 2, along with relevant archaeologicaland fossil hominid sites.

The Late Pliocene of Eurasia and North Africa

Given the poor quality of current data fromAsia, the best base-line for assessing environmentalconditions at a continental level during the earlyPleistocene is the PRISM2 reconstruction ofnorthern hemisphere climate and vegetation in thelate Pliocene, between 3.29 and 2.97 Ma (Dowsettet al., 1994; see Fig. 3). As with most reconstruc-tions of Pleistocene global climatic conditions,the primary data are from deep-sea cores, withterrestrial data incorporated when su!ciently well-dated and environmentally-sensitive. Asia is atpresent very poorly sampled, but doubtless detailwill improve as more data become available. Interms of studying human origins and dispersions,the most interesting feature of the PRISM2 recon-struction is that grasslands ca. 3 Ma were probablycontinuous from West Africa right across to north-ern China; in other words, the present-day desertbarriers of the Sahara and South West Asia didnot then exist. Additionally, the Red Sea barrierbetween Africa and Arabia would have beenmuch less e"ective than now, particularly at thesouthern end, at the Straits of Bab El Mandeb(Tchernov, 1992: 116–117), and this might havebeen an important crossing point between Africaand Asia. We can also note that for much ofthe Plio-Pleistocene, the main part of the Niledraining northern Ethiopia and eastern Sudanflowed eastwards to the Red Sea, and was thus lessof a barrier to faunal migrations between Africaand Asia (Butzer and Hansen, 1968). If hominidshad already dispersed across the African grass-lands in the Late Pliocene (Brunet et al., 1995),there is little reason why they might not also havedispersed into similar grasslands in Asia, as sug-gested recently by Dennell (1998) and Turner(1999).

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 425

The Late Pliocene/Early Pleistocene (2.5–1.8 Ma)in Eurasia and North Africa

The main climatic change in the late Pliocenewas the inception of northern hemisphereglaciation (Shackleton et al., 1984). This wasaccompanied by a major cooling ca. 2.6 Ma in theLake Baikal region, central Siberia (Williams et al.,1997; Grachev et al., 1998; Muller et al., 2001;Demske et al., 2002), and the onset of loessdeposition in north-central China at 2.6 Ma atLingtai (Ding et al., 2000) and Yanyu (Han et al.,2003), 2.47 Ma at Luochuan (Lu et al., 1999), andin Central Asia at 2.5 Ma (Dodonov andBaiguzina, 1995), as well as a dramatic increase inthe deposition of aeolian dust into the Sea ofJapan ca. 2.5 Ma (Xiao and Zhisheng, 1999).These Asian changes all indicate a strengthening ofthe winter monsoon in East Asia, and the strength-ening of high pressure systems over Siberia. InEurope, the first glaciation occurred ca. 2.3 Ma(Zagwijn, 1992), although the vegetational conse-quences of glacial-interglacial cycles appears tohave been very slight in northern Europe prior to

1 Ma (de Jong, 1988; Gamble, 1999: 102). AMediterranean-type climate was established in thesouth-central Mediterranean in the early Pliocene,but only at 3.2 Ma in the north-west part; bothregions show glacial-interglacial contrasts after2.4–2.3 Ma (Suc, 1984; Bertoldi et al., 1989).

The palaeoenvironmental record for the EarlyPleistocene of southern Asia south of latitude40(N is patchy, but generally indicates that thePliocene grasslands of 3 Ma were still largely inplace in North Africa, South West and South Asia,and that conditions were moister than at present.These regions should therefore have been suitablefor Homo erectus. Their evidence for LowerPleistocene hominids in these regions can now bereviewed.

North AfricaThe earliest evidence for hominids north of

the Sahara is Ain Hanech, Algeria (36(N). Thelocality was situated in a flood-plain; grassland isindicated by remains of Equus and gazelle, andopen water by Hippopotamus. The artefacts areOldowan-type, and are dated palaeomagnetically

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Fig. 2. The earliest hominid and archaeological localities in southern Eurasia. The circles indicate the earliest evidence for hominidsfrom each area considered in the text: 1) Ain Hanech; 2) ’Ubeidiya, 3) Evron; 4) Dmanisi; 5) Kashafrud; 6) Riwat; 7) Pabbi Hills;8) Xiaochangliang; 9) Gongwangling; 10) Sangiran; 11) Modjokerto. The squares represent the next youngest indication of hominidsin each region: A) Casablanca; B) Atapuerca; C) Orce; D) Monte Poggiolo; E) Ceprano; F) Gesher Benot Ya’qov; G) Treugol’nayaGora; H) Kul’dara; I) Dina, Jalalpur; J) Bori; K) Nihewan; L) Zhoukoudian; M) Chenjiawo; N) Bose.

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to ca. 1.8 Ma (Sahnouni and de Heinzelin, 1998;Sahnouni et al., 2002) although Raynal et al.(2001: 68) suggest (without giving reasons) that itmay be only ca. 1.2 Ma. There is no other evidencefor hominids in North Africa in the Lower Pleisto-cene. They are absent from the excellent Plio-Pleistocene sequence at Casablanca, Morocco(33(N), until ca. 0.7 Ma BP (Raynal et al., 2001),and this may be evidence for the absence ofhominids, rather than the usual absence of evi-dence. Ain Hanech may therefore indicate a short-term Early Pleistocene dispersal event north of theSahara under suitably moist conditions.

South-West AsiaThe earliest evidence for hominids in Israel

dates from the Early Pleistocene, as claims ofartefacts under a basalt 2.4 Ma at Yiron (Ronen,

1991) have so far failed to convince their critics.According to recent palaeomagnetic investigationsof the artefact-horizons at Erq el-Ahmar, theOldowan-type artefacts there may date to theOlduvai Subchron (1.77–1.95 Ma) (Ron and Levi,2001). The Early Pleistocene fauna from Bethelem(31(N) indicates a largely grassland environmentsimilar to an African lowland savanna at this time(Bar-Yosef, 1994). Slightly later are the betterknown archaeological sites from the 150m-thicksequence at ’Ubeidiya (31(N), dated to ca. 1.4 Ma(Tchernov, 1987). This site is remarkable for con-taining the oldest Acheulean assemblages outsideAfrica, and for showing repeated frequency ofuse, with no less than 15 major archaeologicalhorizons from swampy layers, beach deposits andstream conglomerates (Bar-Yosef, 1994: 231). Thelocal environment was open and semi–arid, with

Fig. 3. Northern hemisphere vegetational zones in the late Pliocene (after Dowsett et al. 1994).

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 427

East African species such as Oryx, Kolpochoerusolduvaiensis, Hippopotamus gorgops, Pelorovisoldowayensis and Crocuta crocuta (Tchernov,1992). Other late Lower Pleistocene Acheuleanassemblages are known from Evron Quarry(Israel) (0.78–1.0 Ma (Ron et al., 2003)), andLatamne and Sitt Markho (Syria) (Tchernov et al.,1994), and the extraordinarily-rich horizons atGesher Benot Ya’aqov (Israel (ca. 0.8 Ma; Goren-Inbar et al., 2000)). Taken together, this evidencemakes the Levant one of the few areas outsideAfrica where there is an average of one later LowerPleistocene archaeological locality per 100–150,000years, two of which (’Ubediya and GBY) showrepeated usage. Even so, it is significant thatSaragusti and Goren-Inbar (2001) note that theseearly Acheulean sites have more in common withAfrican assemblages than with each other, andinterpret this phenomenon as indicating “adynamic system in which populations and/orideas dispersed out of Africa during the Lowerand Middle Pleistocene in repetitive, distinct andseparate waves” (ibid. 85; emphasis mine).

Arabia, Turkey and IranThere is no unambiguous evidence for hominids

in Turkey, the Arabian peninsula and Iran (total-ling over 1.8 million square miles) during theLower Pleistocene, and this enormous regionremains one of our largest areas of ignoranceconcerning early hominid dispersals into Eurasia.Conditions were probably favourable for homi-nids, as the region was moister than today.As an example, fossil remains from three EarlyPleistocene localities in lacustrine deposits in theAn Nafud desert (27(N), N.W. Saudi Arabia(Thomas et al., 1998), include those of Crocutacrocuta, Pelorovis cf. oldowayensis, Oryx (allpresent at ’Ubeidiya), a large felid Panthera cf.gombaszoegensis, fox (Vulpes cf. vulpes), anelephant, (probably Elephas recki), Equus, ahippopotamid Hexaprotodon, as well as varioustypes of Alcelaphines, Bovids and Camelids. Fishand turtle are also present, indicating perennialopen water. Carbonate analyses of herbivore teethindicate a diet of C4 plants, implying open grass-land. Early Pleistocene Anatolia also appears tohave been moister than today, with evidence of

lakes from the Lake E!irdir area of Anatolia, andterra rossa soils indicative of warm temperate tosubtropical climatic conditions with both humidand dry seasons (Nemec and Kazanci, 1999).Faunal remains from the Lower Pleistocene local-ity of Dursunlu, Anatolia, indicates open steppeconditions with some woodland (Gulec et al.,1999). In the Kashafrud Basin (36(N) in NorthEast Iran, Oldowan-type stone artefacts werefound in the palaeo-shore deposits of a vast,shallow Late Pliocene/Early Pleistocene lake, andmay thus be late Pliocene or Early Pleistocene inage (Ariai and Thibault, 1975; Smith, 1986); thislocality would repay further examination.

CaucasusThe best known Early Pleistocene hominid and

archaeological occurrence is, of course, Dmanisi,which has an Oldowan-type lithic assemblage,several hominid remains, and is dated to or shortlyafter 1.8 Ma (Gabunia L. et al., 2000a,b). Atlatitude 42(N, it is the most northerly Early Pleis-tocene hominid locality in Eurasia. Palaeo-environmental indicate a semi-arid to warmMediterranean-type climate, with abundant localwater resources and a fairly open landscape withforests found on river banks and nearby mountainslopes (Gabunia L. et al., 2000a). When occupied,it was probably only 60 km from the Caspian,which may then have joined the Black Sea(Gabunia et al., 2000b) and prevented dispersalfurther north, although Mitchell and Westaway(1999) argue that this did not occur until 1.2 Ma.Pollen analysis indicates increasing aridity afterthe hominid occupation, leading to the expansionof steppe grasslands at the expense of foresthabitats (Gabunia L. et al., 2000a).

There appears to be a million-year gap betweenDmanisi and the next evidence for hominids in theCaucasus (Lioubine, 2000: 147). They are prob-ably absent from Akhalkalaki, a rich faunal LowerPleistocene locality in Georgia (Vekua, 1986), asthe few artefacts found are probably intrusive(Tappen et al., 2002) (Contra Oms et al. (2000):10666), there is no Acheulean assemblage atAkhalkalaki). Hominids may be present at the lateLower Pleistocene locality of Amiranis-Gora(Gabunia M., 2000), but are certainly present at

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the cave site of Treugol’naya, in which the earlieststone tool assemblage (layer 7) has an average ageof 583"25 ka (Doronichev, 2000).

North India and PakistanEvidence of hominids in this region during

the early Pleistocene is sparse and contentious.Convincing examples of intentionally flaked stonewere found in a sandstone/conglomerate in theSoan Valley at Riwat (33(N), Pakistan and datedto a minimum of 1.9 Ma (Rendell et al., 1987;Dennell et al., 1988). The folding of the SoanSyncline, which contains this artefact-bearinghorizon, was dated to the late Pliocene by anearlier, independent geological team (Burbank andReynolds, 1984). Stone artefacts were also foundon the erosional surfaces of fossil-bearing depositsca. 1.2–1.4 and 1.8–2.2 Ma in the nearby PabbiHills, and in the absence of any obvious immediatelater context, some at least are assumed to becontemporaneous with those deposits (Hurcombeand Dennell, 1992). Those circumstantially datedto 1.2–1.4 Ma may indicate that hominids weretherefore present in northern Pakistan in the laterLower Pleistocene. Nevertheless, no hominidremains or large concentrations of in situ artefactshave yet been found in the Indian sub-continent inearly Pleistocene contexts, and in India, there is nodefinite evidence of hominids until the MiddlePleistocene (see below), if we leave aside an uncon-firmed ESR date of 1.3 Ma for an artefact horizonat Isampur.

The virtual absence of evidence for hominids inthe northern part of the Indian sub-continentdurng the Lower Pleistocene is not caused by theabsence of the type of grasslands they preferred.Soil carbonate analyses of the 2.2–0.5 Ma sequencein the Pabbi Hills (Quade et al., 1993) show anoverwhelmingly C4 vegetation throughout thelower Pleistocene, indicative of open grassland.The fauna (as in the Upper Siwaliks of northernIndia) is dominated by grassland types, notablyEquus, Rhinoceros, a variety of medium-sizedbovids, Elephas, Stegodon and ostrich. The maincarnivores were Pachycrocuta, followed byMegantereon, Panthera, and a small jackal-sizedcanid (Turner, in prep.). The older horizons,dated to ca, 1.8–2.2 Ma, contain remains of

Anthracoceres, the gira!d Sivatherium giganteum,a very large bovid (type unknown), and a smallprimate, as well as a large canid, Canis cautleyi.None of these is present in younger horizons(Dennell, in prep.). The herbivores in this groupwere probably browsers, and their disappearanceafter 1.8 Ma may indicate a contraction of wood-land and more arid conditions similar to thatobserved at Koobi Fora (Behrensmeyer et al.,1997).

ChinaEarly Pleistocene environments of southern

China, Southeast Asia and Indonesia during theEarly Pleistocene are still poorly documented.According to Jablonski et al. (2000), sub-tropicalconditions prevailed over much of northern China,and tropical ones south of the Qinling mountains.When these were first occupied is unclear as vari-ous claims for early Lower Pleistocene hominidsand/or stone tools remain unverified. The cavefissure of Longgupo (ca. 30(N) (Wanpo et al.,1995) has been reported as containing evidence ofHomo, flaked stone and a hammer-stone as wellas some teeth of Gigantopithecus, all dated toc. 1.8 Ma. However, the hominid remains areunconvincing and probably hominoid (Schwartzand Tattersall, 1996) or an ape (Wu, 2000), and theflaked stone “artefacts” lack clear flake scars,ripple scars and bulbs of percussion or otherevidence of intentional flaking, and are equallyunconvincing (personal observation). Likewise,claims of hominids at Dongyaozitou andRenzingdong, at or beyond 1.5 Ma are also re-garded as not proven (Bar-Yosef and Belfer-Cohen, 2001: 22).

The earliest, unambiguous and well-dated evi-dence for hominids in China appears to be lateLower Pleistocene. At present, the earliest evidenceis from the Nihewan basin (latitude 40(N), whereZhu et al. (2001) have recently dated the hominidoccupation at Xiaochangliang to 1.36 Ma. Itsclimatic context is unclear, and it is not yet clearwhether the occupation occurred during an inter-glacial or glacial period. The faunal remainsinclude Cervus and Gazella, which do not implycold conditions. Given the evidence fromGongwangling as well as Central Asia (see below),

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 429

the occupation at Xiaochangliang 1.36 Ma-agowas probably during a warm episode. Thiswould be consistent with the suggestion byWang et al. (1997) that the first hominids atGongwangling north of the Qinling mountains(34(N) ca. 1.15 Ma were only there in warmseasons. This age-estimate is also similar to arecent re-dating of the Lantian calotte (34(N) to1.2 Ma. by Hyodo et al. (2002), which reinforcesthe evidence for a late Lower Pleistocene presenceof hominids in China.

IndonesiaThe dating of the Javan hominids is proving to

be a re-run of the famous “KBS controversy” ofthe 1970s over whether the large-brained specimenKNM-ER 1470 was 2.6 or 1.8 Ma. As many willbe aware, the contest was largely between defend-ers of the absolute dating of 2.6 Ma, and thosewho placed more reliance on biostratigraphic evi-dence, and preferred (correctly, as it turned out)the younger dating (Lewin, 1987). As in the KBScontroversy, the case for an early dating rests onabsolute dates, this time derived from Ar40/Ar39

dating of hornblende grains from pumice deposits.Swisher et al. (1994) dated the volcanic layer fromwhich the infilling of the Mojokerto cranium isthought to have been derived to 1.81"0.04 Ma,and the earliest Sangiran hominids (S27 and S31)to 1.66"0.04 Ma. Hu"man’s (Hu"man, 2001)detailed re-examination of Perning/Mojokertosupports the original observation that the hominid(found in 1936) was penecontemporaneous withthe volcanic unit subsequently dated by Swisherand his team to 1.81 Ma. This result is broadlycompatible with other recent Ar39/Ar40 datesobtained by Larick et al. (2001). They examinedsediments from the South East quadrant ofthe Sangiran Dome where the Bapang (Kabuh)Formation has yielded almost 80 H. erectus speci-mens, over 50 of which have known find-spots,and obtained dates ranging from 1.51"0.08 Mafor the Bapang/Sangiran Formation boundary to1.02"0.06 Ma at the top of the hominid fossilifer-ous sequence. The earliest date of 1.51"0.08 Malay above an unconformity, and also 10 m abovethe earliest hominid remain (Brn 1996.04) in thesequence, so hominids were clearly there before

1.5 Ma—assuming of course that the dates arecorrect.

These findings are at odds with several years ofdetailed palaeomagnetic, biostratigraphic andlithological studies by a variety of (non-American)teams, who consistently argue for a younger datingof ca. 1.3–1.0 Ma for the first appearance ofhominids in Java. Work by Semah et al. (2000)suggests that the ‘lower lahar’, a volcanic sedimentlayer at the base of the deposits in the Sangirandome that marks the emergence of dry land in thearea dates to ca. 1.7 Ma., and as such circum-scribes the first appearance of hominids to afterthat date. Palaeomagnetic studies (e.g. Itiharaet al., 1994; Hyodo et al., 2002) place the hominidsin the late Matuyama; and on faunal grounds,Leinders et al. (1985), Sondar (1984), van derBergh et al. (2001) all place the earliest hominidsin the Ci Saat fauna, which they date to ca.1.2–1.0 Ma. (The scale of current discrepancies indating is evident by Larick et al., (2001) assigninghominid Brn1996.04 (the one 10m below the dateof 1.51 Ma) to the Ci Saat fauna, which van derBergh et al. (2001) date to 1.0–1.2 Ma—at least300 ka younger than Larick et al.’s estimates).More recently, Hyodo et al. (2002) has re-assertedthe primacy of palaeomagnetic evidence, andclaimed that the dates obtained by Swisher et al.(1994) and Larick et al. (2001) are “too old”,and the result of reworking, as also argued byde Vos and Sondaar (1994) and Langbroek andRoebroeks (2000), who also argue for a muchyounger dating on the basis of the tektite evidence.(According to Anton (2002), however, this particu-lar criticism is based on a single date of a tektite ofuncertain provenance).

The present impasse will not be resolved with-out further research. Current evidence may beinterpreted therefore as showing either, a short-lived, early dispersal of hominids into Indonesiaca. 1.8 Ma (Swisher et al., 1994), but withoutresulting in colonisation; or a population that wasresident after 1.5 Ma (Larick et al., 2001) or amuch later, and probably more successful dispersalbetween 1.2–1.0 Ma. On circumstantial grounds,the “traditional” dating of the Javan hominidsto the later Lower Pleistocene is consistent withcurrent data from China (especially now that

R. Dennell / Journal of Human Evolution 45 (2003) 421–440430

Longgupo is no longer a credible hominidlocality), whereas the very early Javan datesprovoke the need to explain why hominids reachedIndonesia ca. 500,000 years before China.

Whatever the age of the oldest Javan hominids,the large number of hominid finds (over 80) fromthe 500 ka sequence at Sangiran, and their appar-ent homogeneity, are a better indication of aresident hominid population than in most otherregions of southern Asia. The faunal evidence(Sondar, 1984) suggests that hominids migratedinto a region of open grassland and woodland, inwhich Stegodon, Hexaprotodon and cervids werethe main herbivores, and in which rain forestlike the present-day did not exist until UpperPleistocene times. There appears to have been littlecompetition from carnivores (apart from a largefelid). Although an extremely large continentalshelf was exposed across Sundaland at times of lowsea level, it is curious that the only record forhominids in the Lower Pleistocene is from thewestern side, as hominids are not evidenced inSarawak until late in the Upper Pleistocene(Barker, 2002).

Southern EuropeCould hominids have reached southern Europe

in the Early Pleistocene? Advocates of a “long”chronology (e.g. Bosinski, 1992; Gibert, 1992)believe this is probable, and a number of localitiesare often cited as supporting evidence. Asone of the proponents of the opposing “short”chronology (Dennell, 1983: 21–39; Dennell andRoebroeks, 1996), I find very few, if any, of theclaims for Early Pleistocene Europeans convincingin terms of their dating, context and identification.Additionally, Turner (1992) has pointed out, com-petition from carnivores was more severe inEurope in the Lower than in the MiddlePleistocene, and it was only after the large sabre-tooth felids Homotherium and Megantereon, andthe large hyena Pachycrocuta became extinct thathominids would have been able to gain readyaccess to carcasses, an argument recently re-iterated by Hemmer (2000). Hominid dispersalsinto Europe thus seem unlikely between 1.5 and0.8 Ma. Nevertheless, hominids might have dis-persed into southern Europe ahead of these large

carnivores in the early Pleistocene, either via theDardenelles or the Straits of Gibraltar, eventhough there is no unambiguous evidence that thishappened.

The evidence (or lack of it) for hominids inEurope before 1 Ma has been extensivelyreviewed in recent years (e.g. Roebroeks and vanKolfschoten, 1994, 1995; Gamble, 1999: 115–125),and need not be revisited here. Atapuerca TD6 at780 ka is often viewed as the clearest evidence ofhominids in southern Europe before 500 ka,although other localities such as Barranco Leonand Fuente Nueva 3, Spain (Oms et al., 2000),Monte Poggiolo (Peretto, 1992) and Ceprano,Italy, (Ascenzi et al., 1996) may be slightly older.Grasslands and prey might have been availablenorth of the Mediterranean, but they were notutilised on a permanent basis until the MiddlePleistocene.

Implications

This review suggests that there is remarkablylittle evidence for the “conquest of con-tinental Eurasia” (Boaz, 1997: 189) that has beenattributed to Homo erectus after 1.8 Ma. Althoughthe dispersal of H. erectus out of Africa has beendescribed as a “damburst”, by which populationsmigrating at the rate of only 10 kilometres ageneration could have reached Java from EastAfrica in only 25,000 years (Lewin, 1994), theevidence suggests occasional trickles rather than aflood. Hominids probably dispersed into SouthWest Asia, the Caucasus and South Asia in theEarly Pleistocene, ca. 1.8 Ma but for the most part,their presence across South-West and South Asiaca. 1.8–1.5 Ma can be regarded as simply a latitu-dinal dispersion into the type of habitats alreadyutilised in East Africa. Depending upon one’schoice of dates, North Africa and Java might alsohave been first occupied at this time, or later in theLower Pleistocene, perhaps ca. 1.3–1.2 Ma. Homi-nids in eastern Asia appear to be late LowerPleistocene in age, with Xiaochangliang at 1.35 Macurrently the oldest. However, the only evidencefor resident hominid populations during the LowerPleistocene is from Israel, with at least six localities

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 431

for the period 1.8 Ma (Erq el-Ahmar) to 0.8 Ma(Gesher Benot Qa’aqov), and evidence of multipleusage of at least two of these (’Ubediya and GBY),and also Java, with an average of one hominid findper 62,500 years over 500 ka. There is no reason toassume that populations were geographically ortemporally continuous between these two regionsthroughout the Lower Pleistocene. Indeed, as theJavan H. erectus populations are regarded bymany as isolated from those of H. ergaster inAfrica, the scarcity of evidence for hominid popu-lations in South and South West Asia in the LowerPleistocene is perhaps not surprising.

The scarcity of evidence for hominids acrosssouthern Eurasia and the Mediterranean cannot beattributed solely to an absence of their preferredgrassland habitats. The type of grassland environ-ments used at Ain Hanech, Ubediya, Dmanisi werealso present at Casablanca, the Caucasus, northernIndia, the Arabian peninsula, and southernEurope, but hominids appear to have been absentor very thin on the ground in these regionsbetween 1.8 and 1.0 Ma or even later. Nor can it beattributed to climatic instability as glacial-interglacial contrasts were much less in the Lowerthan in the Middle Pleistocene. It may be signifi-cant that Dmanisi and ’Ubediya were both insmall-scale mosaic environments, with a varietyof resources nearby, and in regions with aMediterranean-type climate. This would imply thathominids’ colonising abilities were constrained to aparticular mix of habitats within a small area. Thefossil hominid evidence from Dmanisi lends someindirect support to this suggestion. The surpris-ingly small size of cranium D2700 suggests that“the first humans to disperse from the Africanhomeland were similar in grade to H. habilis(sensu stricto)” (Vekua et al., 2002:85). If theirbehavioural abilities were similar, its abilities as acoloniser might have been very limited.

The virtual absence of hominids across south-ern Asia in the Lower Pleistocene may well reflecta lack of fieldwork and/or a lack of suitable LowerPleistocene exposures and sections. It is undeniablethat there has been little exploration of theLower Pleistocene in much of SW Asia outsideIsrael, northern India/Pakistan, much of Chinaand mainland SE Asia. On the other hand, homi-

nids appear to have been absent in areas wherepeople have looked, and where good sections exist.Examples are the Ebro Valley in Spain, Seneze inFrance, the Val D’Arno sections in Italy (seeRoebroeks and van Kolfschoten, 1995: 308),the Matuyama loess sections of Central Asia(Dodonov, 2002), the loess plateau of North China(where the absence of hominids may result froman absence of suitable stone material for makingtools (Dodonov and Zhou, 2003)); and theUpper Siwaliks of northern India and Pakistan.Additionally, the record improves considerablywith the Middle Pleistocene, as seen below.

The onset of permanence: the evidence forhominids in Eurasia 1 Ma–500 ka

At what point do we see evidence for con-tinuous occupation in Eurasia? Although thearchaeological evidence for Eurasian hominids inthe later lower and early middle Pleistocene is stillextremely poor, two major developments can berecognised between 1.0 and 0.5 Ma.

Hominid dispersals into cooler environments

Hominids appear to have dispersed into temper-ate latitudes as far north as 40–45(N around1 Ma. To take a continental transect acrossEurasia, the earliest current dates in these latitudesare from the Orce Basin, Spain (37(N), at ca.1 Ma (Oms et al., 2000; Atapuerca TD6, also inSpain, at ca. 780 ka (42(N; Carbonell et al.,1995b); possibly Ceprano, Italy (Ascenzi et al.,1996) and Monte Poggiolo (44(N), 900 ka,Milliken, 1999); Kul’dara, Tadjikistan at 900 ka(38(N, Ranov, 1995); Donggutuo in North Chinaat 0.9 Ma (40(N; Lanpo and Qi, 1987). It isprobably significant that this northward expansionoccurred around the time of a major faunal turn-over marked by the appearance of the Galerianfauna (Turner, 1992), and the expansion of a“mammoth steppe” across northern Eurasia(Guthrie, 1990).

Most of these initial forays into more northerlyareas were probably confined to interglacials. Inthe Central Asian loess sequences, for example,

R. Dennell / Journal of Human Evolution 45 (2003) 421–440432

sites such as Kul’dara (and later ones such asLakatui and Karatau) are found only in palaeo-sols, i.e. interglacials (Dodonov and Baiguzina,1995: 712). These probably indicate “discontinu-ous sequences of occupation” (Davis, 1987:130) resulting from “a back-and-forth pattern”(Velichko, 1999:13) of hominids, and imply theexistence of a parent population further south(probably in northern Iran or Afghanistan) thatwas continuous in cooler periods when loess wasbeing deposited in Central Asia. Evidence fromChenjiawo (34(N) in North China suggests thathominids were present in both cold and warmperiods only after 650 ka (Wang et al., 1997). Theearliest occupation in layer TD6 at Atapuerca,Spain, is attributed to an interglacial period, cor-related with OIS 21 (Antonanzas and Bescos,2002). At Monte Poggiolo, Italy, however,environmental conditions are described as a coldand arid steppe (Milliken, 1999: 11).

As has been extensively discussed in recentyears, hominids dispersed into northern Europeup to 50oN ca. 500 ka (Roebroeks and vanKolfschoten, 1994, 1995; Gamble, 1999;Roebroeks, 2001), after which Europe was prob-ably occupied continuously, excluding Scandinaviaand the Russian Plains.

The Acheulean

The second development was the appearance ofAcheulean, bifacial assemblages in Europe, NorthAfrica, South West Asia and the Indian sub-continent in the early Middle Pleistocene. InEurope, the appearance of Acheulean, bifacialassemblages is a major part of the evidence for thecolonisation of Europe, and has been extensivelytreated elsewhere (e.g. Roebroeks and vanKolfschoten, 1995; Gamble, 1999). Sites such asBoxgrove (50(N), occupied ca. 478–524 ka in thelast interglacial before the Anglian (OxygenIsotope Stage 12) provide clear evidence of thislithic tradition in northern Europe by 500 ka. Ingeneral terms, the Acheulean in Asia appears to berestricted to the same type of open grasslandssouth of 35(N that were preferentially occupied inthe early lower Pleistocene. When viewed moreclosely, the record for the Asian Acheulean is

very patchy, with most dates falling in the earlyMiddle Pleistocene. The oldest are in Israel, withthe earliest Acheulean outside Africa being at’Ubeidiya (31(N), at ca. 1.4 Ma (Tchernov, 1987),and sites like Gesher Benot Ya’aqov (31(N) at0.8 Ma (Goren-Inbar et al., 2000). There are noother comparable absolute dates for the EarlyAcheulean over the 3500 kilometres between Israeland Pakistan, where hand-axes have been found atDina and Jalalpur in contexts dated palaeo-magnetically to ca. 700–400 ka (Rendell andDennell, 1985). The appearance of the Acheuleanmay be further constrained in Pakistan by itsabsence despite substantial searching-in the PabbiHills before 1.2 and probably 0.9 Ma (Hurcombeand Dennell, 1992). In India, the earliestAcheulean dates are from Bori (19(N) ca. 670 ka(Mishra et al., 1995).

The recent discovery of Acheulean-type hand-axes at Bose, South China, dated to 800 ka (Yameiet al., 2000), well to the east of the Movius Line,suggests that sites of a comparable age should befound further to the west. If the dating is correct,this discovery may represent a short, atypicaldispersal event from areas further west related toand possibly immediately following widespreadforest-burning after the great South East Asiantetkite shower that dates the assemblage (Yameiet al., 2000). The dating is problematic, however(Langbroek, 2003). Tektites are present over a verylarge area of S.E. Asia, but typically in muchyounger lag deposits resulting from denudation(Fiske et al., 1996, 1999; Koeberl and Glass, 2000),and have not been found in their original contexts.The age of the Bose artefacts therefore requiresindependent corroboration. Even if confirmed,however, they do not greatly alter the long-observed overall restriction of the AsianAcheulean to South West Asia and the Indiansub-continent.

The restriction of the Asian Acheulean to theLevant until the Early Middle Pleistocene hasmajor consequences on assessments of the“Movius Line”, the boundary between the simple(Mode 1) flake-core technologies in South Eastand Eastern Asia, and the Acheulean (Mode 2)assemblages to the West. On current evidence, the“Movius” Line was e"ectively immediately east

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 433

of Israel until the early Middle Pleistocene. Theabsence of bifaces in eastern Asia has beenexplained in various ways: because bamboo andother woods could have fulfilled the same function(Pope, 1989), because of greater reliance of plantfoods (Watanabe, 1985); or because the complexrules of manufacture made the Acheulean vulner-able to discontinuities in space and time Schick(1994). However, the question is less one of havingto explain why the Acheulean was not adopted ineastern Asia as why it did not manage to expandits distribution into South Asia until the MiddlePleistocene. For around half its total duration inAsia (1.4–0.2 Ma), the Acheulean appears to havebeen restricted to the Levant, several thousandkilometres west of the Movius Line. Recently,Gamble and Marshall (2001) have suggested thatthe Acheulean world can be divided into twoprovinces along a “Roe Line” that runs throughIsrael to West Africa. To the north, hand-axeswere made largely by faconnage, or shapingfrom a core, and to the south and east (includingIndia) a zone where hand axes were made largelyby debitage, or on large flakes. A point notdeveloped by Gamble and Marshall (ibid.) is thatthe “Roe Line” is also (on current evidence) theboundary of Movius Line until the early MiddlePleistocene.

Summary and conclusions

A number of conclusions are drawn from thispaper:

1. The skeletal and archaeological record forhominids in Eurasia before 1 Ma is probablyone of repeated, short-lived and modestdispersal events, rather than of continuousresidence (see Fig. 4). Although there aremany reasons why the Lower PleistoceneEurasian hominid is so poor (e.g. a lack offieldwork; a shortage of accessible fine-grained early Pleistocene exposures; problemsof dating poorly-provenanced finds, especiallyin areas lacking volcanic ashes; di!cultiesof dating sites palaeomagnetically that liebetween the Olduvai and Jaramillo normalevents; and the submergence of coastal areas,particularly in S.E. Asia and eastern China),it seems unwarranted to assume that therewas continuity of occupation during theLower Pleistocene, given that many regionsexhibit discontinuities as long as 500 ka oreven one million years.

2. The lack of evidence for continuity in theEurasian Early Pleistocene record implies thatthe colonising abilities of early H. erectus were

0.5 Ma0.60.70.80.91.01.11.21.31.41.51.61.71.81.92.0

Europe N.Africa Levant Caucasus S. Asia S. China N. China Indoneisa(long) (short)

Ain Hanech Erq el-Ahmar

CasablancaAtapuerca TD6

Orce

CepranoM. Poggiolo

'Ubediya

GBY

LatamneEvron

Dmanisi

Amiranis Gora?

Treugol'naya 7

Riwat

Pabbi Hills?

Pabbi Hills?

Bori

Xiaochangliang

Bose?

LantianGongwangling

Chenjiawo

Mojokerto

Sangiran

Fig. 4. Discontinuities in the fossil and archaeological records for hominids in Eurasia during the Lower and Early Middle Pleistocene,prior to 500 ka. Shaded areas indicate periods for which there is no definite evidence that hominids were present. Controversial datesor finds are indicated by a “?”.

R. Dennell / Journal of Human Evolution 45 (2003) 421–440434

very limited. Although it has been modeledas a highly successful coloniser (Anton et al.,2002; Mithen and Reed, 2002) that could notonly disperse rapidly but also remain perma-nently, the evidence at hand suggests other-wise. It may be significant that the ability ofhominids to colonise temperate latitudesnorth of 45(N did not occur until theircranial capacity moved into the modernhuman range. However, even in the MiddlePleistocene, and notwithstanding evidence forgreater encephalisation, controlled use of fire(James, 1989), improved hunting techniques,including the use of spears (e.g. Thieme,1997), evidence for continuity of populationappears weak in Italy (Milliken, 1999) andBritain (Ashton and Lewis, 2002). It shouldnot be assumed that there was any greatercontinuity in populations and lithic traditionsin earlier periods, or in other less well-documented Middle Pleistocene areas. Even ifdispersal may have been rapid longitudinallyacross southern Asia, there is little evidenceof sustained occupation until the MiddlePleistocene.

3. The discrepancy between Europe and Asia inthe length of their respective hominid recordsseems more apparent than real. Although somehave wondered why hominids waited so long at“the Gates of Europe” (Gamble, 1999: 123),the records of Asia and Europe are broadlysimilar, given that the main long-trend ofhominid dispersals was latitudinal ratherthan longitudinal. Hominids were not kickingtheir heels at the Gates of Europe for most ofthe Lower Pleistocene. It was probably notuntil the Middle Pleistocene that hominidsbegan habitually to utilise latitudes up to45–50(N, which happen to include most ofEurope.

4. Both “long” and “short” chronologies for earlyhominid occupance of Eurasia may be valid,but may also be telling us di"erent storiesabout the process by which most of Eurasiawas eventually colonised. “Long” chronologiesmay record intermittent, short-term dispersalevents, under favourable conditions, into areasgenerally similar to those already occupied

elsewhere. “Short” chronologies may be tellingus histories of more permanent occupation,and of how hominids were able to colonise newareas.

5. “Out of Africa 1” is a grossly simplistic con-struct for discussing the dispersal of hominidsinto Eurasia. There were probably severaldispersal events in the Lower Pleistocene: theone recorded at 1.8 Ma might not have beenthe first; Israeli evidence suggests at least twobetween 1.4 and 0.8 Ma, and the expansionof the Acheulean across southern Asia andinto Europe may imply a further series. Thecolonisation of Europe after 800 ka wasprobably complex, with dispersals fromseveral regions (Rolland, 2001). The lowerPleistocene may thus have experiencednumerous dispersals from Africa, and severalothers within Eurasia, as earlier suggested byRolland (1992). As Straus and Bar-Yosef(2001: 1) point out, dispersals may also havebeen two-way between Africa and Asia.

6. The model proposed here of intermittent dis-persal events that gradually encroachednorthwards is capable of further testing, falsi-fication and elaboration. In arguing that mostof the Eurasian archaeological and fossilhominid record is one of occasional visitors,I have tried to show there is little support forthe common assumption of continuity inpopulations after their first appearance. Justas the “short” chronology was useful in clear-ing away “several exaggerated claims andmuch weak evidence” (Gamble and Marshall,2001: 8), so it may be useful to question thevalidity of models of Early Pleistocene lithictraditions or hominid phylogenies that arebased on an assumption of continuity. Whatare needed are not just more well-datedLower and Middle Pleistocene sites for di"er-ent parts of Eurasia, but far more contextualdata on their climatic and environmental con-text, and greater consideration of how conti-nuity might be defined. Only then will we beable to see how hominids responded to theebb and flow of temperature shifts through-out the Old World during the EarlyPalaeolithic.

R. Dennell / Journal of Human Evolution 45 (2003) 421–440 435

Acknowledgements

I am grateful to Kathryn Holmes for referenceson Saudi Arabia, Marco Langbroek for drawingmy attention to the problems of dating tektites,and Andrew Chamberlain for critical attention tothe text.

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