Eurasian gates: the earliest human dispersals

EURASIAN GATESThe Earliest Human Dispersals

Eudald Carbonell, Marina Mosquera, Xose Pedro Rodriguez

Jose Maria Bermudez de Castrot

Francese Burjachst

Jordi Rosell, Robert Sala, and Josep Vallverdu

All authors except J. M. Bermudez de Castro are affiliated with IPHES.Area de Prehistoria, Universitat Rovira i Virgili, Plaea Imperial TiLrraco I,

Tarragona 43005, Spain. Email: [email protected]

t Centro Nacional de Investigacion sabre Evolucion Humana (CENlEH),Avenida de la Paz, 28. Burgos 09004. Spain

t Instituci6 Catalana de Recerca i Estudis AvaDlyats (ICREA)

KEY WORDS: Eurasia; Hominin dispersals; Socialization of technology; Generalist diet;Climate; Subsistence

This paper revises the current slate of the debate about the earliest hominindispersals out of Africa. First we review the archaeological evidence for theearliest occupation of Asia and Europe. Next we summarize the environmentalparameters relatedto the earliestphases afhuman evolution-specifically, climaticimplications for human adaptations andfaunal dispersals. We discuss which werethe first hominins to leave Africa, and we propose the invention oftechnology asa fundamental step for the development of our genus, likely related to changesin subsistence and diet during the late Pliocene and early Pleistocene. From ourpoint of view, hdminins were able to generalize the use of technology, as wellas to generate, integrate. and diffuse new information into their collective socialbehavior. We refer to this concept as "socialization. " Hence, technology and thesocialization thereofbecame integral aspects ofthe ecological niche ofhominins.

SINCE THE LAST DECADE OF THE TWENTIETH CENTIlRY there have been a plethora ofscientific publications discussing when and where the first hominin dispersalsout of Africa aod into Eurasia took place (e.g., Aguirre and Carbonell 2001;Anton et a!. 2002; Arribas and Palmqvist 1999; Bar-Yosef and Belfer-Cohen2001; Carbonell et a!. 1996, 1999b; Dennell 2003; Gabunia et a!. 2001; Mithenand Reed 2002; Roebroeks 2001, 2006; Roebroeks and van Kolfschoten 1994;Rolland 200 I; Turner 1992). This period also witnessed important discoveriesthat led to the modification ofprevious proposals. In some cases, new hypotheseshave been tested empirically. Such is the case of current scientific opinionsregarding the Mature Europe and Old Eurasia scenarios (Carbonell et a!. I999b).New alternative hypotheses suggest that Asia, rather than Africa, may have beena center for earlier dispersals (Dennell and Roebroeks 2005). In this paper we

Journal ofAnthropological Research. vol. 64, 2008Copyright © by The University of New Mexico

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THE EARLIEST EURASIAN SITES

revisit the issue and propose new explanations concerning human evolution andthe occupation ofour planet. Here, we consider and analyze several parameters inorder to offer explanations for the so-called Out-of-Africa or Hominin Dispersalphenomenon. We propose that the "socialization" of technological improvementsis a key issue in the explanation of hominid dispersals. The "socialization" ofa technological system consists of adopting technological development in ageneralized manner within the hominin group. Once the use of a technologicalsystem has been generalized in a community, new opportunities emerge andimprovements are easier to adopt. That is, homlnins are able to generate, integrate,and diffuse new information into their collective social behavior.

The Earliest Hominin Occupations in AsiaThere are several tenninal Pliocene or Early Pleistocene sites in Asia (Figure

I; Table I). Paleomagnetic analyses carried out in Riwat (north Pakistan) haveyielded dates assigned to the early Matuyama (between 2.4 and 1.9 mya; Dennellet aI. (988). A few quartzite objects were discovered at this site. The anthropogenicnature of these lithic objects is controversial (Hemingway and Stapert 1989),although one of the authors (E.C.) analyzed a sample and considered them to beartifacts. However, the association of these tools with the conglomerate dated toa minimum of 1.9 mya needs further confirmation. We face the same situation inthe nearby site of Pabbi Hills, which could date to either 1.4--1.2 or 1.8-1.6 mya(Dennell 2003).

An age between 2.5 and 2 mya has been proposed for the fauna and lithicartifacts found in Renzidong (Anhui, China; Jin et aI. 2000; Weiwen and Pu 2007).However, the evidence from this site has also not beenvalidated owing to the scarceinfonnation about the lithic tools (Dong 2006). Faunal remains, lithic artifacts, andsome possible hominin specimens have been recovered from Longgupo (Wushan,China; Wanpo et aI. 1995). This assemblage has beeo dated to the Olduvai subchron(between 1.95 and 1.77 mya) on the basis nf paleomagnetism, biostratigraphiccorrelations, and ESR dating methods. These results remain questionable (Zhu etaI. 2003). Nevertheless, the most debated issue regarding this site is whether ornot the hominoid remains belong to Homo (Eller et aI. 2001). In our view, theanthropogenic nature of the Longgupo lithic objects is indisputable (E.c., personalobservation), despite Dennell's (2003) questioning.

The stratigraphic position, age, and anthropogenic nature of some of the lithicobjects (in particular, some of the core tools) from Erq-EI-Ahmar (Israel) are stillunder discussion. Paleomagnetic analyses ascribe these objects to the Olduvaisubchron (1.95-1. 77 mya; Ron and Levi 200 I).

The earliest convincing evidence of hominin occupation in Eurasia is that ofDmanisi (Georgia), dated to around 1.8 mya (Gabunia and Vekua 1995; Gabuniaet aI. 2000b, 2000c; Lordkipanidze et aI. 2007). The archaeological record ofDmanisi includes lithic artifacts, faunal remains, and hominin specimens. Theartifacts, made on silici.fied volcanic tuff, basalt, and, to a lesser extent, quartz,can be classified as Mode I (Le., choppers and flakes). There is no consensus

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TABLE IThe Oldest Sites in Asia

Lithic Faunal HomininSite Location Dale (mya) Dating Method lndustty Remains Remains

Riwatt Pakistan 2.4-1.9 PM Mode 1 No No

Renzidongt China 2.5-2.0 BS Mode 1 Yes no

Longgupot China 1.95-1.77 PM, BS, ESR Mode I Yes Yes?

Erq-EI-Ahmart Israel 1.95-1.77 PM Mode I No No

Pabbi Hillst Pakistan 1.95-1.77 PM Mode I No No

Mojokerto Indonesia 1.81 ± 0.04 4°Arj39Ar No Yes

Dmanisi Georgia 1.8-1.6 PM, "'Ar/"Ar Mode 1 Yes Yes

Sangiran Indonesia 1.66 ± 0.04 "'Ar/"Ar Yes Yes1.07-0.78

Majuangou-Goudi China 1.66 PM,S Mode i Yes No

Vbeidiya Israel ca. 1.4 BS Mode 2 Yes Yes

Xiaochangiiang China 1.36 PM,S Mode 1 Yes No

Xiantai China 1.36 PM,S Mode 1 Yes,

No

Xihoudou China 1.27 PM, S Mode I Yes No

lsampur India 1.2 ESR Mode 2 Yes No

Gongwangling-Lantian China 1.2-1.15 PM, S Mode 1 Yes Yes

Donggutuo China 1.07-0.99 PM Mode I Yes No

Bitzat Ruhama Israel ca. 1 PM, RTL Mode 2? Yes No

Evron Quarry Israel 0.99-0.78 PM,BS Mode 2 Yes No

Akhalkalaki Georgia 0.98-0.78 PM, BS Mode 1 Yes No

Kuldara Tajikistan ca. 0.85 PM, TL Mode 1 Yes No

Boa Lesa [ndonesia 0.84 ± 0.07 FT Mode 1 Yes No

Bose China 0.803 ± 4fJArP9Ar Mode 2 No No0.003

MataMenge Indonesia 0.800± IT Mode I Yes No0.007

Gesher Benot Ya'aqov Israel ca. 0.78 PM,BS Mode 2 Yes Yes

t These sites are controversial.

Dating methods: PM = Paleomagnetism, S = Stratigraphy, rates ofsedimentation,5S = Biostratigraphy, ESR = Electro-spin resonance, RTL = Radio-Thennoluminescence.TL = Thermoluminescence, 44ArP9Ar = Argon-Argon, FT = Fission-track


regarding the taxonomic interpretation of the hominin specimens, which arevariously assigned to H. erectus (Brauer and Schultz 1996), H. ergaster (Gabuniaet al. 2000b), or a new species, Homo georgicus (Gahunia et al. 2002). In anycase, the Dmanisi hominin remains reptesent the oldest reliable record of Homooutside Africa, with a very primitive morphology and associated with a Mode Itechnology (Lumley et aI. 2005).

Some fossils discovered in Java (Indonesia) could be as old as Dmanisi. Adate of 1.81 ± 0.04 mya has been proposed for ~ojokerto, and 1.66 ± 0.04 myafor Sangiran, using 40Arf"Ar (Swisher et al. 1994). Nevertheless, the controversysurrounding the dating of these sites has not yet been resolved (Huffman 2001;Langbroek and Roebroeks 2000; Semah et al. 2000). Overall, there are twoopposing views, with some researchers accepting the antiquity of these fossils,which could date to between 1.8 and 1.5 mya (Dennell 2003; Huffman 2001;Larick et al. 2001; Swisher et al. 1994), and others rejecting these dates andsuggesting a maximum of around 1.2 to 1.0 mya (Bergh et al. 2001; de Vos andSondaar 1994; Langbroek and Roebroeks 2000). The key issue is the interpretationof the stratigraphic provenience of the fossils, together with the dating of thesestrata (Huffinan et al. 2006). For the moment, the earliest presence ofhominins inJava is still a topic ofdebate.

A bit later and again in the Near East, the site ofUbeidiya (Israel) has yieldeda rich collection of faunal remains and lithic artifacts, as well as three isolatedhominin remains, from different layers (Belmaker et al. 2002). There is generalconsensus regarding the antiquity of the site,'which dates to around 1.4 myabased on biostratigraphic correlations (Tchemov (988). The oldest levels containabundant core-choppers, polyhedrons, and spheroids, but lack handaxes. Theupper part of the stratigraphic sequence contains handaxes in varying quantitiesand may be classified as early Acheulean (Bar-Yosef and Goren-Inbar 1993).Using Mary Leakey's (1971) terminology, the levels with few handaxes could beclassified a~Developed Oldowan. However, given the strong similarities betweenthe Acheulean and the non-Acheulean basic knapping techniques, the presence orabsence of handaxes may be due to different occupational functions at the site.According to Bar-Yosefand Belfer-Cohen (2001), the previous supposition thatthe lower levels had an exclusively core and flake industry is no longer validsince a fragment of a trihedral pick was found in one of these levels. This factreinforces the opinion of Bar Yosef and Goren-Inbar regarding the classificationof the entire Ubeidiya assemblage as early Acheulean (or Developed Oldowan,following Leakey's classical terminology).

In India, a preliminary ESR date of 1.2 mya has been assigned to the site ofIsampur, which contains a rich lithic assemblage associated with faunal remains(bovid, cervid, equid bones, and fragments oftortoise carapace; Paddayya etal. 2002).This site has been interpreted as a handaxe and cleaver workshop. Consequently,it corresponds to a Mode 2 technology (petraglia et al. 1999, 2005). In Pakistan,handaxes have been found at Dina and Jalalpur in contexts dated paleomagneticallyto ca. 700-400 kya (Denne1l2003; Rendell and DennellI985).

In the Nihewan Basin (China) at least four sites have been dated to theEarly Pleistocene (see Table I for dating methods): Majuangou (now renamed

"


as Goudi; Gao et aI. 2005), Xiaochangliang, Xiantai, and Donggutuo. All of themhave yielded faunal and lithic remains. The oldest one is possibly MajuangouGoudi, which has fou~ layers with stone tools and processed faunal remains. Thetime span represented in the stratigraphic sequence is around 340,000 years, andbased on magnetostratigraphy, sedimentary layers formed between the Olduvaisubchron and the Cobb Mountain event. Therefore, it has been assigned an ageof ca. 1.66 mya for the oldest level and ca. 1.32 mya for the youngest one (Zhu etaI. 2004). The next-youngest sites are Xiaochangliang and Xiantai, both dated to1.36 mya (Deng et aI. 2006; Zhu et aI. 200 I), and finally Donggutuo, with a faunaland lithic assemblage ascribed to the Jaramillo subchron (0.99- I.07 mya; Zhu etaI. 2002, 2003).

At Xihoudou (Shanxi, China), associated lithic and faunal remains have beenfound. Paleomagnetic analyses for this site have provided a date of 1.27 mya(Zhu et aI. 2003). The human skull from Gongwangling (Lantian, China) coulddate to between 1.2 and 1.15 mya (An and Ho 1989; Hyodo et al. 2002). This sitealso yielded lithic artifacts, some of which came from the hominin-bearing layer.Others were found in younger layers, slightly older than 0.78 mya.

Near the end of the Early Pleistocene, between around I and 0.78 mya, theamount of data increases. Evron Quarry (Israel) has yielded lithic artifacts andfaunal remains, but all of the handaxes were collected from the surfa~e. They aresimilar to some of the handaxes from Ubeidiya and Latamne (Bar-Yosef 1994).The Evron Quarry assemblage has been technologically classified as MiddleAcheulean, and the site could be Early Pleistocene in age, between 0.99 and 0.78mya (Ron et aI. 2003). However, TL and ESR datings have yielded more recent,Middle Pleistocene dates (Porat and Ronen 2002). At the site of Bitzat Ruhama(Israel), lithic and faunal remains with a date of about I mya were documented(Table I). The lithic industry shows very particular traits, comprising diminutiveartifact types (Zaidner et al. 2003).

One of the most interesting sites in the Near East is Gesher Benot Ya'aqov(Israel), which dates to around 0.78 mya. Here, lithic, faunal, human, plantremains, and even anthropogenic fire have been documented (Goren-Inbar etal. 2004). The lithic industry corresponds to Mode 2, with traits resembling theAfrican Acheulean tradition (Goren-Inbar et aI. 2000, 2004).

Akhalkalaki (Georgia) has yielded a rich faunal assemblage dated to between0.98 and 0.78 mya. The association of a very small lithic assemblage with thefaunal remains is not clear, given the absence ofclear hominin modification ofthebones. The faunal remains were likely collected and modified by carnivores. Theartifacts may have been deposited into the faunal layer from the surface via rodentburrows, and thus were not really associated·with the faunal remains (Tappen etaI. 2002: 1388).

The oldest site in Central Asia is Kuldara (Tajikistan). It comprises a loesspaleosoil deposit containing severarfaunal specimens and lithic artifacts dated to0.85 mya by paleomagnetism and TL. The bulk of the artifact assemblage consistsof small flakes (Ranov et al. 1995).

In Bose (China), several artifacts classified as similar to Acheulean handaxes(Mode 2) have been documented (Hou et aI. 2000). Tektites found in the same

.,


level in which the archaeological material was found have heen dated to 0.803 ±0.003 mya. However, these rocks could have heen redeposited into their currentposition, so the dates may not be related to the archaeological site (Kocberl and Glass2000). Furthermore, some authors assert that East Asian handaxes are not similar inmorphology to the typical western Old World Acheulean implements (Norton et al.2006). In our view, they rellect the wide variability of Mode 2 technology.

The island of Flores (Indonesia) has yielded two archaeological sites with fewlithic and faunal remains, which possibly belong to the upper Early Pleistocene,Boa Lesa and Mata Menge. Boa Lesa dates to 0.84 ± 0.07 mya and Mata Menge,to 0.800 ± 0.007 mya (Moore and Brumm 2007; Morwood et al. 1998, 1999).

The Earliest Occupations in EuropeThe earliest hominin occupation of Europe is represented by a significant

number of sites (Figure I; Table 2) throughout the entire continent, especiallythe South (Carbonell and Rodriguez 2006). Two areas in the Iberian Peninsulaare noteworthy: Guadix-Baza (Granada), with the Fuente Nueva 3 and BarrancoLeon sites, and the Sierra de Atapuerca (Burgos), with the Sima del Elefante andGran Dolina sites.

The fossiliferous deposits from the Guadix-Baza basin are composed ofclayey paleo-lakeshore sediments. Fuente Nueva 3 and Barranco LeOn havebeen dated both biostratigrapbically, based on the presence of Early Pleistocenemicromarnmal assemblages, and by magnetostratigraphy. Fuente Nueva is slightlyolder than 1.07 mya, around 1.2 mya, while Barranco Leon is dated to between1.2 and 1.3 mya (Oms et al. 2000; Toro et al. 2003; Turq et al. 1996).

TABLE 2The Oldest European Sites

Site Location Dale (mya) Dating Method Lithic Faunal HomininIndustry Remains Remains

Pirro Nord Italy 1.7-1.3 BS Mode I Ves No

Sima del Elefante TE9 Spain 1.2 PM, BS, eN Mode I Ves Ves

Barranco LeOn Spain 1.3-1.2 PM,BS Mode I Ves No

Fuente Nueva 3 Spain 1.2 PM,BS Mode I Ves No

Pont-de-Lavaud France 1.2-1.1 RPE Mode 1 No No

Vallonoet France 1.07-0.99 PM, BS, ESR Mode 1 Ves No

Soleihac France 1.07-0.991 PM BS Mode I Ves No0.61 '

Ceprano Italy 1.06-0.7 KiAr, PM, Mode I Ves Ves

Monte Poggiolo Italy 0.9 PM, ESR Mode 1 No No

Gran Dolina TO 4 Spain >0.85 PM, BS, ESR Mode I Ves No

Gran Dolina TD 6 Spain 0.85-0.78 PM, BS, ESR Mode I Ves Ves

Dating methods: PM ". Paleomagnetism, 5S = Biostratigraphy, CN = Cosmogenic nuclides, RPE =Electron paramagnetic resonance, ESR = Electron spin resonance, KJAr = Potassium-Argon, Ff =

Fission·track

The Atapuerca site complex has yielded two Early Pleistocene fossiliferousdeposits containing evidence for early hominin presence in Europe: Sima delElefante and Gran Dolina. The former has an Early Pleistocene sequence made upof alternating sandy and clayey layers. Three of these layers contain lithic artifactsand bone fragments with cut-marks, indicating hominin activity. One layer, TE9, hasrecently yielded a hominin mandible assigned to Homo antecessor, with a date of1.2 mya based on paleomagnetism and cosmogenic nuclides (Carbonell et a!. 2008).The sediments from the lower levels of Sima del Elefante reveal a geomagneticreversal, interpreied as the Matuyama-Brunhes boundary (Pares et al. "2006), andmicrofaunal taxa identified indicate a minimum age of I mya (Rosas et a!. 200 I).

The lower halfofthe Gran Dolina sequence is also formed by Early Pleistocenedeposits, with four levels containing evidence of anthropogenic activity. TD6 isthe most complex and richest of these fossiliferous records from the lower levelsof Gran Dolina (Figure 2; Carbonell et a!. 1995, 1999a). All Gran Dolina lowerlevels are situated under the Matuyama/Brunhes reversal, thus belonging to theEarly Pleistocene (pares and Perez-Gonzalez 1995, 1999). Furthermore, TD6 hasbeen dated to between 0.857 and 0.780 mya through ESR (Falgueres et a!. 1999;Falgueres 2003), while micrommamal biostratigraphy indicates that the levelbelongs to the Upper Biharian zone, possibly within oxygen isotope stage (OIS) 19,around 0.79 mya. Cuenca-Bescos and colleagues (1999) have proposed that TD5correlates with OIS 20, around 0.81 mya, and TD4 with OIS 21, around 0.85 mya.

In Italy, Pirro Nord has provided Early Pleistocene lithic tools and fauna(1.7-1.3 mya; Arzarello et al. 2007). The lithic assemblage of Monte Poggiolo(northern Italy; Peretto et a!. 1998) is found in a sandy/gravelly regressive marinebeach deposit. It contains lithic artifacts and gastropods, and it was dated bypaleomagnetism (Gagnepain et a!. 1998) and ESR (Falgueres 2003) to around 0.9mya-that is, younger than the Jaramillo subchron.

The fossiliferous locality of Ceprano is part of the long sequence of theCeprano Basin, in the Lacio region (central Italy). This stratigraphic sequencecomprises a lower lacustrine complex and an upper colluvial-alluvial complex: thelatter yielded a hominin skull fragment. This find was correlated with definitivearchaeological assemblages from the same region ascribed to Mode I and Mode2. Thus far no absolute dates have been assigned to the layer containing the skullfragment. However, through chemical analyses, regional chronologies for thevolcanogenic sediments have been proposed, situating the deposit hetween 0.7and 1.06 mya. The assemblage correlates with Mode I lithic assemblages fromthe same region (Ascenzi et a!. 1996; Manzi et a!' 2001).

Recently, dates have been published for the open-air terrace site of Pont-deLavaud (France), where, although no bones were preserved, a lithic assemblagewas found composed of quartz choppers, chopping tools, denticulates, andscrapers. The ESR dating points to 1.2 mya for the hominin occupation of thisarea (Despriee et a!. 2006).

The French site of Vallonnet has yielded lithic and faunal assemblagesstudied by Lumley and colleagues (Yokoyama et a!' 1988). Paleomagnetic,biostratigraphic, and ESR analyses at this cave have yielded a date between 0.99and 1.07 mya, within the Jaramillo subchron (Lumley et a!' 1988). The French


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site of Soleihac is disputed. This site yielded abundant paleontological andarchaeological remains. It is located at the lakeshore and consists of lacustrinesediments in a volcanic crater. Soleibac was assigned to the Jaramillo subchron(Bonifay et al. 1976, 1989), but biostratigraphy points to a Middle Pleistocene ageclose to 600 ka (Lacombat et al. 2003; Roebroeks and van Kolfschoten 1994).

Climatic Implications for Human EvolutionThe present-day distribution of African plants and animals is a result

of particular environmental changes that took place during the Pliocene andPleistocene (Figure 3). Contractions and expansions in response to alternating dryand wet seasons fragmented primitive forest habitats. As a result, we encountertoday a rich diversity oflandscapes and habitats, dry-adapted Acacia commiphora.deciduous bushland, savanna, semidesert scrub, grassland, and mesic Afrom~ntane

forest (Bonnefille 1985).Pliocene and Pleistocene environmental changes in Africa had several

implications. Habitat diversification, niche partitioning, guild or niche expansion,biogeographical rearrangement (immigration/migration), speciation, extinction,coevolution, and disease are processes dependent on climate change, as well ason sea level oscillations, soil formation, and to a lesser extent meteoric impacts,meteorological phenomena (e.g., hurricanes), natural fires, volcanism, and otherprocesses.

The transition to Quaternary glaciations was gradual. As a matter of fact, thedates ofthe earliest glacials have not yet been well established. A cold episode at 3.5mya has been recorded (Haug and Tiedemann 1998), along with an intensificationof cold conditions in the northern hemisphere toward 2.75 mya (Shackleton etal. 1984). However, the earliest documented glaciation in Europe dates to 2.3mya (praetiglian) and roughly coincides with biotic changes, including the firstextinctions ofaustralopithecines in Africa and the emergence of the genus Homo.

Along with global cooling, we must consider the aridization trend that tookplace concurrently. Increasingly drier conditions began at the end of the warmperiod (ca. 3.5 mya), when mangroves, jungles, and tropical forests occupied alarge part of Africa. This trend reached definitive cyclicity at 2.5 mya. Duringthis period (after 3.2 mya), the present-day dry summer pattern typical of theMediterranean zone was established (Sue 1984; Sue and Zagwijn 1983).

The evolution of the African landscapes at the borders of the standardclimatic zones, up until 2.5 mya, entailed a gradual transition toward more opentropical forests with perennifolium bushes, savannas, grasslands, and steppes. Itwas toward 2.5 mya and within this context of mosaic landscapes and increasedecological diversity that, most authors agree, major biotic changes took place,including a major faunal dispersal event and the emergence of the genus Homo.

We should also consider the critical date nf 2.6 mya, which marks thecreation of the Saharan desert (Leroy and Dupont 1994; Potts and Behrensmeyer1992). This implies the isolation of North Africa from tropical Africa, though notcompletely, as the Nile corridor remained open.


• major biota changes

.. faunal movements and changes

...- Westemoven interglacial<Of--- g\&cial A (Cromerian)

...- Dorst glaciation (MIS 18)

+- Unge glaciation.-- savanna (Afaf)

.-Weichselian glaciation (MIS 4-2)

.-Saalian glaciation (MIS 61

...- E1stenan glaciation (MIS 10)

.- Menapian glaciation

.- cold biotic changes in Mediterranean region(MIS 50-48) -+-- dry grassland (Konsa)

+-- coki biotic changes in Mediterranean region(MIS 62-581

4- EburOnian glaciation

~ +- cold biotic changes in Mediterranean region:ot (MIS 82-18)

:::::. _ PrBeliglian glaciation

.~tHomo'" +- __ (Omol

1+- +- glBSSiand (E_)

3 severe dry periods, 1st North Amcan desert

z:- it .- intensification of +- limit mangrove & tropicalis j Northern Hemisphere forest c;21ON. W Africa)'C i!: laciation<: • 9-

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(MIS: Marine Isotope Stage; W: warm; C: cold).

It seems obvious that the emergence and dispersal within Africa of our genus was associatedwith global cooling and aridization. In fact, the climatic crisis of the late Pliocene, which becameaccentuated around 2.7 mya, is key to understanding the diversity of hominin genera that emergedin Africa during the Pliocene and the early Pleistocene. The humid, temperate conditions and relatedpresence of abundant lacustrine zones with rich mosaic vegetation prior to 2 mya and between 2 and1.9 mya enhanced biodiversity and, consequently, led to an increase in the available environmentalresources (Bchrensmeyer and Bobe t999). Somewhat later, climatic conditions deteriorated, becomingcolder and drier, with a lowering of the sea level. According to some researchers, this was the firstopportunity for homjnins to leave Africa (Aguirre and Carbonell 2001 ; Anton et al. 2002).

Faunol DispersalsSeveral faunal movements occurred around 2.6-2.5 mya (Azzaroli et al.

1988). Elements of African fauna have been identified in the Caucasus (giraffids,procavids, and ostriches), and in the Balkans and the Iberian Peninsula (giraffids),coinciding with the so-called Eurasian Elephant-Equus event (Martinez-Navarro2004). Also, several African antelopes dispersed into Central and SouthernAsia at around these dates (Vrba 1995). In contrast, this sarne period witnessedmovements of Eurasian faunas toward Africa, including the presence of Equus;Ursus cf. etruscus; the raccoon dog, Nyctereutes abdeslami; and Capra primaevain North Africa (Arambourg 1979; Geraads 1997).

Another period of faunal change and dispersals occurred between 2 and 1.8mya. It included an almost cpmplete replacement of carnivores and most of theungulates in Europe, with the arrival of species such as the large canids, Canisarnensis. Canis elruscus, or Lycaon falconer;; the giant hyena, Pachycrocu/Qbrevirostris; the European fossil jaguar, Pontheragombaszoegensis; the sabertooth,Megantereon white;; and ungulates, such as the ovibovine Praeivibos sp.; Equusaltidens; the rhinocerotid Stephanorhinus hundsheimensis; Hippopotamusantiquus; and the cervid Pseudodama sp. (Martinez-Navarro 2004).

The time around 1.5 and 1.4 mya witnessed other faunal dispersals, as shownby the paleontological record of the Levantine Corridor at the site of Ubeidiya.

From 2.5 mya onward, coinciding with the first recorded European glaciations(Praetiglian), the landscapes inhabited by the earliest Homo species consisted ofdry, open savannas or dry grasslands. This was also the time of the last majorhumid period in East Africa, during which some authors place the origin ofHomo(deMenocal 1995).

As a consequence of the beginning of glaciations, we can observe at midlatitude biotic and isotopic changes from MIS (Marine Isotope Stage) 82 to 78 (2mya), 62 to 58 (1.6 mya, Eburonian glacial), and 50 to 48 (1.35 mya), with coldepisodes similar to MIS 2 (with the expansion of Artemisio; Combourieu-Neboutand Vergnaud-Grnzzini 1991). During this glacial stage, at 1.6 mya, we observe theentry of Atlantic foraminifers into the Mediterranean via the Strait ofGibraltar.

On the other hand, at Dmanisi (Georgia) between 1.8 and 1.6 mya, a mosaicof vegetation integrating different habitats prevailed, from more or less opensteppes to mountain woodlands, as well as gallery forests along the river coursesand small lakeshores. Many of the plant species that have been recorded forDmanisi at the end of the Pliocene and beginning of the Pleistocene still occur inthe Caucasus region today (Gabunia et al. 2000a). In other words, this part of theplanet has been functioning as a refugium for biota since the Pliocene, and hencean important locus for species maintenance, diversification, and distribution.

This period witnessed a progressive increase in the C4-type de~rt plants inAfrica, which stabilized around 0.7--{J.6 mya. At this time, major periods ofaridityin the Mediterranean have also been documented (Bertini 2000). Therefore, itwas toward the end of this period of aridity (0.9--{J.8 mya) that the marked bioticchanges occurred, coinciding with the beginning of the dominant Milankovitchcycle of 100 kya.



It involves the appearance of such species of African origin as Pelorovisoldowayensis/bubaloides, Oryx cf. gazella, Equus cf. tabeii, Kolpoehoerusolduvaiensis, Sus sp., and Theropitheeus oswaldi. In western Europe, thisfaunal change is detected by the arrival of some ungulates, such as the caprineAmmotragus europaeus, found at Vallonnet (southeastern France) and FuenteNueva-3 and Cueva Victoria (southeastern Spain; Moulle et al. 2004), or thelarge Equus cf. bressanus (Martinez-Navarro and Rook 2003).

Finally, the time around 900-800 kya may be considered the last great periodof faunal changes, especially concerning European extinctions of large carnivores.These are the cases ofLycaon. Megantereon, Pachycrocuta, Acinonyx pardinensis,Puma pardoides (Vallonoet, Gesher Benot Ya'aqov; Martinez-Navarro 2004)and the last European record of the jaguar, Panthera gombaszoegensis, at thelowermost levels of Gran Dolina (Atapuerca, Spain; Garcia 2003).

The arrival ofBos primigenius from Africa around 0.6-0.5 mya in associationwith the appearance of Acheulean culture in Europe is especially siguificant(Martinez-Navarro, et al. 2007).

HOMININS

Was Homo the Only Hominin Involved in the First Out-ol-Africa Dispersal?Most authors agree that the genus Homo, and in particular H.-ereetus, was

the first bominin to leave Africa. However, as we will discuss, there are objectivereasons for posing alternative hypotheses.

First, we know that the first out-of-Africa hominin dispersal occurred atthe end of the Pliocene, perhaps about 2 mya (Aguirre and Carbonell 2001).Geochronological data from Dmanisi support this hypothesis (Gabunia et al.2000c, 200'1; Lumley et a1. 2002). At that time, hominin diversity in East Africaincluded the genera Homo, Paranthropus, and possibly Australopitheeus-ifthe species A. garhi (Asfaw et al. 1999) persisted in the region until the latePliocene. The Ethiopian sites of Konso (P. boisei; Suwa el al. 1997), BouriFormation (A. garhi),. and Hadar Formation, A.L. 666 (Homo sp.; Kimbel et a1.1996) bear witness to the northernmost presence of the three groups in Africa.Hence, if ecological conditions in Northeast Africa and the Near East werefavorable during the late Pliocene, the three groups of hominins could haveexpanded their range northward, and they might have crossed the boundariesof the African continent. In other words, the entire ecosystem could havemoved northward.

The evidence from Dmanisi compels us 10 reopen the debate posed with regardto the finding of the Sangiran 6 mandible in 1941, as well as other specimensespecially Sangiran 5, 8, and 9-from this site in Java. The great variabilityof the Sangiran hominins has been recognized by numerous scholars (Franzen1985; Koenigswald 1960; Kramer 1989; Kramer and Konigsberg 1994; Pope andCronin 1984; Tobias and Koenigswald 1964; Weidenreich 1944; Wolpoff 1980).Von Koeni'gswald proposed Meganthropus palaeojavanieus to accommodate thesize and shape of the Sangiran 6 mandible, this taxon having been published forthe first time by Weidenreich (1944).

With the finding of a diversity of Pliocene hominins in South and EastAfrica during the first half of the twentieth century, some authors referred theSangiran 6 specimen to the robust australopithecines (first Robinson 1953, andnnt later than Franzen 1985). However, on the basis of dental morphology LeGros Clark (1955) placed Sangiran 6 in the Pithecanthrapus erectus group. Morerecently, other authors have contended that the variation in the Sangiran mandiblesamples is within the range of a single species (H. erectus) with a high degree ofsexual dimorphism (e.g., Kramer 1989; Kramer and Konigsberg 1994; Pope andCronin 1984; Wolpoff 1980). Wolpoff (1980: 190) writes, "Sangiran 6 (formerlyattributed to Meganthropus) has a mandibular body as thick and tall as the Natronaustralopithecine female (MLO 40), although the posterior teeth are much smallerand the P4 is not molariform. Australopithecine-like features are also seen inSangiran 9; the shape of the tooth row forms a 'y' similar to that found in muchearlier hominin specimens from the Afar and Lake Turkana (ER-1482)." Thus, inspite of most authors preferring to include all specimens in H. erectus, they cannotavoid noting the special features of the Sangiran hominins. Yet, some researchershave attributed Sangiran 6 and other Sangiran specimens to H. habilis (Howell1978; Tobias and Koenigswald 1964; Tyler 1991, 1994).

The published hominin mandibular sample from Omanisi includes the gracilespecimen 0 12 and 02735, and one robust specimen, 02600 (Gabunia and Yekua1995; Gabunia et al. 2002). Gabunia and colleagues have attributed the homininfossil hypodigm from Omanisi to a new species, Homo georgicus. In particular,these authors describe the large 02600 mandible as the key specimen (holotype)for naming the new species, which would have had marked sexual dimorphism.In fact the dimensions of 02600 are remarkably larger than those nf 0211, and02735, but Gabunia and colleagues (2002) think that the shape is similar in allspecimens. The height of the mandibular body of 02600 at any level exceeds therange obtained in all Homo specimens (Rosas and Bermudez de Castro 1999) andis comparable to that of Paranthropus. However, the breadth of the mandibularbody, as well as the relative size of the anterior teeth (especially that of the largecanines) witl] respect to the posterior teetll, clearly separates 02600 from thisgenus (Gabunia et al. 2002). Gabunia and collaborators (2002) also point to thesimilarities between 02600 and the specimen attributed to H. mdolfensis, KNMER 1802, and UR 501 (from Uraha, Malawi), as well as Sangiran 6, concerningthe size and shape of the first premolar. The P, of 02600 is asymmetrical and hastwo (mesial and distal) roots, as do these specimens and Paranthropus (Bromageet al. 1995; Wood et al. 1988). According to Gabunia and colleagues, H. georgicusis related to both H. habilis and H. mdolfensis, and represents the origin ofa longEurasian lineage.

From all of these observations, it seems reasonable to conclude that Homowas probably the only genus that moved out of Africa. However, there is evidenceto suggest that the protagonists of the first dispersal were more than one Homospecies, perhaps some time before 2 mya, when this genus was emerging in EastAfrica (Aguirre and Carbonell 200 I).

The variability in the oldest Eurasian hominin fossil record may be related tothe variability observed in the early Homo record from Africa (see Wood 1992).



We believe that the new ecological niches associated with the diversity ofPliocenePleistocene Homo types entailed wider and more diverse habitats. [n other words,the adaptive radiation of the genus Homo at the end of the Pliocene was probablyassociated with a demographic phenomenon. It is possible that the Paranthropusniche was more restrictive and that this genus remained tied to its Aliican habitat.

The Dawn o/TechnologyThe earliest lithic industry recorded thus far comes from the Ethiopian site

of Kada-Gona (Semaw 2000; Semaw et a!. 2003), with an age of 2.6 mya. TheEthiopian sites of Kada-Hadar (Kimbel et a!. 1994, 1996) and Omo-Shungura(Howell et a!. 1987), together with the sites of Lokalelei (Kenya; Roche et a!.1999) and Senga 5A (Zaire; Harris et a!. 1987), follow it, with dates of around2.4-2.3 mya. The industries of these sites have been included in the Mode I orOldowan, but they show a significant feature--a scarcity of bifacial reductionin flaking. Mostly unifacial and orthogonal reduction are represented in theseassemblages. These lithic records are abundant and diversified, which suggeststhat technology was not newly formed, but had already been generalized by thistime. The first hominin dispersals out ofAfrica might have occurred before 2 mya,as suggested by the cases of Riwat (Dennell et a!. 1988) and possibly Renzidong(Dong 2006; Weiwen and Pu 2007). Lithic technology evolved in Aliica towarda bifacial conception of both flaking and tools, which was fully developed andgeneralized (and thus socialized) from 2 mya onward, as shown by the sites of01duvai Gorge Bed I (Leakey 1971), Am Hanech (Sahnouni and Heinzelin 1998;Sahnouni et a!. 2002), Gombore I (Melka-Kunture, Ethiopia; Galloti and Piperno2003), Peninj (Tanzania; Dominguez-Rodrigo et a!. 2001), and Swartkrans andSterkfontein (Brain 1981), among other technological assemblages.

The first appearance of Mode 2 occurred at 1.7-1.6 mya in Kokiselei 4 (WestTurkana, Kenya; Roche et a!. 2003) and Konso-Gardula (Ethiopia; Asfaw et a!.1992), but again this industrial record seems to be fully developed, instead ofemergent. Therefore, it seems that there had been an earlier emergence of thistechnology. Mode 2 produces large flakes to configure big tools, diversifiesthe morphotypes of small retouched tools, and standardizes knapping methods,among which the bifacial centripetal technique stands out.

Mode 2 seems to have rapidly expanded along the Rift system once thistechnology was routinized or "socialized" by hominins, as indicated by the siteofUbeidiya (Israel), which dates to 1.4 mya (Bar-Yosefand Goren-Inbar 1993;Tchernov 1988). Around 1.5 to I mya, Mode 2 is found at the Ethiopian sitesof Garba IV (1.5-1.4 mya, classified as Mode I, although it contains handaxes;Galloti and Piperno 2003) and Kesem-Kebena (Wolde-Gabriel et a!. 1992), in theTanzanian site ofOlduvai-Upper Bed II (Leakey 1971), and in the Moroccan siteofThomas Quarry I Unit L (Raynal et aJ. 2004). Thus, Mode 2 has been found inexactly the same area as Mode 1 assemblages.

In Aliica, Mode I and Mode 2 developed alongside one another after 1.6 mya.Both technological systems therefore coexisted for tens of thousands of years. Itappears that certain communities ofHomo adopted this new bifacial industry, butothers continued to use Mode I techniques (Bar-Yosef 1998). Alternatively the


coeKiltence ofMode I and Mode 2 can be explained as tbe result ofdifferent tasksor needl in combination with variations in tbe local availability of suitable lithicraw materials (Clark et al. 1994).

Asia shows Mode I assemblages by around 2 mya, and isolated evidence ofMode 2 appears at lsampur (India) (1.2 mya; Paddayya et al. 2002) and at Bose(China) (800 kya; Hou et al. 2000), but it is not until the late Middle Pleistocenethat Mode 2 seems to have spread across this continent (petraglia 2007). Therefore,we must point out the persistence of Mode I in Asia.

The earliest European occupations with Mode 1 technologies are recordedby 1.2 mya, but is not until ca. 0.6 mya that Mode 2 enters the subcontinent, atwhich point it spreads surprisingly fast. As a result, we find evidence of Mode2 assemblages from England (Boxgrove; Roberts and Parfitt 1999) to Italy(Notarchirico; Piperno 1999) and from Germany (Karlich Secufer; Gaudzinski etal. 1996) to the Iberian Peninsula (Sima de los Huesos and Galeria, at Atapuerca;Carbonell et al. 2001, 2003). Strong similarities between certain lithic types(particularly cleavers) from North Africa and the western Mediterranean (Alimen1975; Giles and Santiago 1988) suggest that Mode 2 came to Europe directly fromAmca, the Strait of Gibraltar being one of the possible dispersal routes for thisMiddle Pleistocene event.

Subsistence and Diet during the Pliocene and Plio-Pleistocene: Hominins andLarge Predators

One of the most determinant elements that led to the increase in the size ofthe hominin brain was the inclusion of meat in the usual diet (AieUo and Wheeler1995), something which also allowed development of the complexity of humanculture. The emergence of subsistence strategies based on dietary generalismoccurred prior to the emergence of the genus Homo. Some studies suggest thatAustralopithecus alricanus had already acquired a diverse and high-quality diet,which included the habitual consumption of animal biomass (Sponheimer andLee-Thorp 1999). This omnivorous diet is also documented among the robusthominins (d'Errico and Backwe1l2003; Lee-Thorp 1994). Therefore, the possibleemergence of this subsistence strategy during the Pliocene must not be rejected,especially considering the fact that wild chimpanzees are able to organize complexhunting activities (Boesch 1994a, 1994b; Stanford 1996; Teleki 1981).

Most models about early dispersals of Homo out of Afiica and into Eurasia(or, specifically, into Europe) try to correlate these dispersals with the dispersal oflarge mammals. In such models, either the dispersals of large mammals modifiedthe environment and thus made it possible for Homo to disperse, or they reflect aclimatic change that modified the environment and thus enabled Homo to disperse.Some of the latest models linking faunal and Homo dispersals are the following:

Turner (1992) proposed a model in which Homo dispersed along withPanthera leo, P. pardus, and Crocuta crocuta from Africa through Asia and intoEurope as part ofa larger event around 1.2-0.9 mya, which marked the transitionfrom the Villafraochian to the Galerian and the onset of an alternation of glacialand interglacial faunas in Europe. The appearance ofthese "new" large carnivoresand the survival of archaic Acynonyx, Pan/hera gombaszoegensis, Megantereon

EURASIAN GATES: THE EARLIEST HUMAN DISPERSALS ZII

cultridens, and Homotherium latidens were supposed to provide many "hllh.quality carcasses" for scavengers, which encouraged Homo to colonize Europe.although such a siiuation could not be maintained owing to competition, wl~,Pachycrocuta. The extinction of Megantereon around I mya was supposed tdhave decreased scavenging opportunities for Homo, whereas the replacement ofthe large Pachycrocuta by the smaller P. perrieri (hyena) around 0.5 mya led tothe permanent presence of Homo in Europe.

Martinez-Navarro and Palmqvist (1995, 1996) presented a model in whichthe hominiD Out-of-Africa dispersal was a consequence of their dependence onlarge predator behavior. According to them, the main agent for the expansion wasMegantereon whitei. The masticatory apparatus of saber-toothed felines allowedthem to specialize in the acquisition of large prey, but they were not capableof extracting all of the nutrients from them, or of fracturing the bones (Marean1989). Consequently, scavengers that had access to the carcasses abandoned bythese felines would have benefited from a great amount of meat and marrow.Following the three-stage model (carnivore-hominid-carnivore) developedby Capaldo (1997) and by Selvaggio (1998) using the FLK Zinjanthropus site(Tanzania), Arribas and Palmqvist (1999) make a direct link between these threeanimal groups, in which the large felines acted as hunters and hominins andhyenas were the scavengers of their prey. .

From this perspective, Arribas and Palmqvist (1999) looked at the sabertoothdispersals during the Early Pleistocene as the main cause for the first hominin'dispersal into Europe. The arrival of hominins and Pachycrocuta into Europewould have been conditioned by their ecological association with Megantereonand their prey. Likewise, this association would have been maintained during theentire Early Pleistocene and part of the Middle·Pleistocene.,

The model adopted by Martinez-Navarro and Palmqvist (1996) and Arribasand Palmqvist (1999) has several weak points regarding hominin subsistence.First, Mesantereon cultridens existed in Eurasia long before the dispersal ofHomo. It too was a hyper-carnivorous species, so there is no reason to believethat they behaved differently than M whitei with regard to the abandonment ofcarcasses. In this case, there is no reason to believe that the dispersals of Homowere a product of the Megantereon whitei dispersal.

Second, the evidence from the Early Pleistocene Aurora Stratum of levelTD6 at Gran Dolina (Sierra de Atapuerca, Spain), which dates to around 800 kya,points to hunting as the main strategy for carcass acquisition (Diez et al. 1999). Atfirst, Diez and colleagues considered the possibility of a scavenging componentgiven the marked presence of young individuals among the larger animals.Nevertheless, the skeletal representation pattern and evidence of butchering (cutmarks and intentional bone fracturing) suggest primary and immediate accessto the carcasses. Therefore, the evidence supports the use of hunting for meatacquisition as acommon practice among the hominins (H. antecessor) representedin the Aurora Stratum, at least regarding the small and medium-size animals suchas red deer, wild boar, roe deer, and the hominins themselves (Diez et aI. 1999).

To the contrary, we first thought that the skeletal selection of large animalswas the result of secondary access by hominins to the remains abandoned by large

L

felids. Modem lions generally leave only a small number of tootb-marks on thebones of their prey, and they discard whole carcasses without having fracturedthe bones to any appreciable extent (Bailey 1993; Schaller 1972; Treves andNaughton-Treves 1999). Scavenging of carcasses abandoned by these predatorswould have offered hominins the opportunity to harvest a large amount ofundamaged marrow. This type of faunal accumulation would contain many longbones and skulls of large animals with few tooth-marks, as is actually found inthe Aurora Stratum of TD6. However, from an ethnographic point of view, theskeletal profile of large animals may also correspond to primary and immediateaccess by hominins (O'Connell et al. 1988a, 1988b, 1990, 1992).

The hypothesis proposing that the TD6 homillins had a substantial huntingcapability is supported by two central arguments (Rosell 2001). First, the bitemarks of carnivores identified at this level seem to match those ofa single, rathersmall predator (fox). Although large felids do not nomnally leave many markson the bones of their prey, contact between their cusps and any bone elementis unavoidable, and therefore some marks are fomned. None of the tooth-marksidentified in the TD6 assemblage falls outside the average established by extensivemeasurements. Second, the signs of hominin processing on bones oflarge animalsare similar to those found on other animals. Humans would not have processed thepartially eaten large carcasses that were the felids' prey in the same way that theyprocessed the results of their own successful hunts of animals of any size.

Therefore, there are currently more reasons to support the idea of primaryaccess to large animals than to defend the hypothesis of systematic scavengingof carcasses abandoned by large felids. In any event, secondary access may haveoccasionally occurred, as indicated by the remains of a single proboscidean andtwo rhinos. However, primary and immediate access by most of the individualsthat fomn the TD6 assemblage (including hominins) must be understood within anetwork of a well-organized hunting activity.

Hunting also seems to have been the preferred strategy for acquiring meat atthe French site of Soleihac, which it is claimed was similar in age to TD6 (Fosseand Bonifay 1989; but see the remarks above on the fomner site's age). However,in the French case there is clearly a scavenging component based on the size ofthe animals represented. According to Fosse and Bonifay, hominins scavengednaturally deceased elephants (Palaeoloxodon anliquus), while the remainingtaxa-,predominantly cervids-are habitual hominin prey.

In the third place, recent work demonstrates that there was greater variabilitythan ouce supposed in animal biomass acquisition strategies among African PlioPleistocene hominin communities (Dominguez-Rodrigo and Pickering 2003).During the 1980s and early I990s, Blumenschine (1988, 1995) and Blumenschineand Marean (1993) developed a new hypothesis according to which the earliestAfrican hominins were passive scavengers, specialized in the extraction ofmarrowfrom the carcasses left behind by large predators in savanna environments. Butfrom a more actualistic, methodological perspective, Dominguez-Rodrigo andPickering (2003) now argue that the data are insufficient to support this hypothesis.According to these authors, Plio-Pleistocene African hominins were capable ofobtaining meat by primary, immediate, and possibly active means.


DISCUSSION AND PERSPECTIVES

• Technology already socialized.t Date of the Acheulean site ofThomas Quarry I Unit L (Morocco) (Raynal et at. 2004).

The earliest lithic industry known is 2.6 mya old, and it does not appear to bean emergent technology, but rather a developed one (Table 3). This means that

Mode 2

Proposed Earliestemergence evidence·

ca. 2 mya 1.7 mya •

I.Smya· 1 mya·t1.5--1.4 mya •

ca. I mya? • 0.8 mya •

ca. 0.6 mya •

Earliestevidence*'

2.6 my.

2 mya

1.8 my.

1.66 mya

1.3 mya

ca. 3 mya

>2 mya

2.5--2 mya

Mode I

Proposedemergence

East Africa

Rest of AfricaWest Asia

East Asia

Europe ca. 1.5 mya

TABLE 3Earliest evidence and suggested emergence of Mode 1 and Mode 2 technologies

during the Pliocene aod Pleistocene


The ability to make a variety ofdecisions in response to an external stimulusis an old and particularly important human feature. We can extrapolate this fromhominin subsistence strategies, as well as from lithic technology and settlementpatterns. Shipman and Walker (1989) and Shipman (2000) have argued that braingrowth and the subsequeot need for an increasingly nutritious meat intalce led tothe systematization of hunting techniques among African hominins, particularlyHomo ergaster. Tbe newly acquired bunting habits allowed for expansion intolarger territories, leading eventually to bominin dispersal outside Africa. Fromthis perspective, we support the bypothesis suggested by these authors, but onethat is focused on primitive Homo forms instead ofon H. ergaster.

This versatility, whicb is typically human, bas also been noted by Bar-Yosefand Belfer-Cohen, wbo suggest that hominin dispersal out of Africa may haveoccurred owing to "the absence ofthe zoontic diseases tbat plagued and constrainedhominins in their African •cradle ofevolution'" (200 I :26), even as they recognizethe indisputable "flexibility in food procuring techniques" of hominins. In ourview, tbe absence ofdisease may be a very important point to consider regardingthe success ofa species in a new environment, but it presupposed that individualsof that species are able to survive there.

Nevertheless, in times of ecological shifts, omnivorous hominins, as welladapted generalists, were capable of maximizing resources by acquiring acomplementary plant-meat diet. Moreover, tool-making is an additional strategyfacilitating the acquisition of many kinds of available resources, with the goalof stabilizing the population structure, maintaining demographic growth, andavoiding extinction. In sum, hominin adaptive strategies during dispersal eventsdid not follow the steps of large carnivores, and in tbis sense, an omnivorous dietmay have bloen part of the generalist subsistence strategy developed by bomininsto better adapt to any kind ofenvironment.

technology may have origioated in Africa some time before this date, maybeas early as around 3 mya (Carbonell et al. 2007). According to current data, wesbould not reject tbe possibility that the real inventors of technology may havebeen australopitbecines. Actually, the technical behavior now acknowledged forcbimpanzees (Mercader et al. 2002)--although clearly far from "real" technologymust be taken into account when we consider hominin species with even largerbrains. Otherwise, technology would bave been invented by primitive Homo, whicbpoints to tbe possibility offmding remains of this genus at around 3 mya.

This period also marks tbe beginning of a climatic crisis, whicb pushedseveral faunal species to move into and out of Africa around 2.6-2.5 mya. Thissame climatic event may have pushed hominins to search for new strategies-tbeinvention of technology and the adoption ofa generalist diet, both allowing fulleruse of more and mbre dispersed, diversified resources, and therefore facilitatingadaptation to the new environments.

The socialization of technology was achieved around 2.6 mya, as thedeveloped record from Gona suggests. Between 2.6 and 2.3 mya there wereindustries in several sites: Omo-Shungura, Lokalelei, and even in more distantareas, such as Senga 5 in the southeastern Democratic Republic of Congo, andtbe somewhat later site ofEI-Kherba in Algeria (Sahnouni et a1. 2002). Therefore,technology had actually dispersed across much ofAfrica by this time. [n tHis sense,we propose that new strategies and "inventions" can only expand to other areasonce they have been generalized-that is, "socialized"-among the communitieswhere they emerged. Accordingly, we propose tbat industries dating to around2.4-2.3 mya may be found all across Africa.

This expansion went beyond the African continent and arrived at theeastern end of Asia around 2 mya or maybe even earlier. The sites of Erq-EIAhmar ([srael), Riwat (Pakistan), Longgupo (China), Dmanisi (Georgia), andSangiran (Java; if we rely on its older chronology) represent this dispersal event.Unfortunately, only Dmanisi and Sangiran bave yielded definitive homininremains, botb showing primitive features belonging to habiline forms. That meansthat both occupations may have occurred a short time after African Homo habilisacquired its own features.

Altogether, these indications suggest that the earliest hominin dispersals outof Africa occurred between 2.5 and 2 mya ago, taking tbe former date from theAfrican technological "socialization" showed by Kada-Gona.

Later Asian occupations are chronologically situated around 1.6 myaonward, which appears to represent a gap between the oldest and the later ones.It is not currently possible to determine whetber tbe earliest Asian occupationswere successful or unsuccessful in terms of demographic continuity, since thisgap may just be the product of differential preservation andlor different researchmethodologies. However, the new evidence coming from Flores may help: if theFlores settlers were the product of as primitive an origin as has been claimed(Brown et al. 2004; Morwood et al. 2004), then we have to admit that ihere wassome kind of continuity in tbe peopling of Asia.

Mode 2 (or the Acheulean) appeared for the first time at 1.7-1.6 mya inKokiselei 4 and Konso-Gardula. Again, this technology does not suggest an



emergent form, but ratber a developed one. Once more, this means thaI Mode2 may have originated some time around 2 mya in Africa, being both socializedand flISt diffused along the eastern part of this continent around 1.5 mya. In thissense, we propose that Mode 2 industries will be found at around 2 mya in thenear future. The "inventor" ofthis technology may have been either Homo hobilisor Homo ergaster. 10 our view, the hiological complexity of Homo ergaster (interms of increased body and brain size) would match better with the complexityof the Mode 2 technology. In this case, the emergence of both Homo ergaster andMode 2 technology is older than previously thought.

On a geological scale, the emergence and sncialization of Mode 2 maycorrelate with another episode of climatic crisis around 2-1.8 mya. The greatestfaunal change concerns the replacement of carnivore species in most of Eurasia.Again, the hominin response to climatic crisis came from biology, with theemergence of a new species, and from cultural adaptations. with the emergenceand socialization of a new technology.

Mode 2's earlier appearances outside Africa are at Ubeidiya (1.4 mya),Isampur (1.2 mya), and Bose (800 kya). In our opinion, Mode 2 industries of 1.5mya onward may be found in Asia. For its part, Europe was peopled with ModeI communities at around 1.2 mya, for this is the date of the oldest Europeanoccupations (Pont-de-Lavaud, Fuente Nueva, and Barranco Leon). This peoplingphenomenon may have come from both Africa and Asia. However, currentresearch suggests that the European Homo antecessor-the oldest hominin foundto date in Europe, at around 1 mya-<!id not come from Africa, but from Asia(Carbonell et al. 2005). That is, it is closer to the Asiatic-ereetus forms than to theAfrican-ergaster ones. This means that both Homo antecessor and the EuropeanMode I shOJ.lld have come from Asia at some point around 1.5 or 1.4 mya. Thecurrent state ofknowledge about the European lithic industries can neither supportnor reject the hypotheses regarding African or Asian origins.

The period ofthe Mode 2 entrance into Asia and the earliest European peopling(I .5-1.2 mya) coincides with the minimum obliquity cycle ofMilankovitch, whichcaused the Menapian glaciation. This period also correlates with a stage of faunaldispersals involving the appearance ofsome species of African origin in the NearEast, and the appearance in Western Europe of some ungulates of possible Asianorigin, all ofwhich says nothing about that hominin-technical dispersal.

Another stage of great faunal changes is documented around 900-800 kya,especially concerning European large-carnivore extinctions. Again, this periodcoincided with the change in the astronomic cycle to 100 kya, but it has no specialreflection in hominin behavior. European peopling has occurred since 1.2 myawith apparent continuity, as shown by the chronological chain represented bythe sites of ·Fuente Nueva, Barranco Leon, Pont-de-Lavaud, Sima del Elefante,Vallonnet, Monte Poggiolo, and Gran Dolina.

Mode 2 in Europe appeared no earlier than 600 kya, at Arago lower levels(Lumley and Barsky 2004) and Notarchirico. Once there, it expanded acrossthe subcontinent extremely fast, as already explained. South European Mode 2reflects some features resembling the North African ones, such as the presence ofcleavers, which are less frequent in the rest of Europe. That means that southern

CO eLUSIONS

European Mode 2 came directly from Africa, instead of via Asia. This technologyis associated in Europe with the Homo heide/bergensis chronospecies, which mayhave descended from Homo ontecessor (Bennudez de Castro et al. 1997). In thiscase, the rapid adoption and socialization of the Mode 2 in Europe by Homoheide/bergensis populations may have been the result oftheir high levels ofsocialorganization and well-established communication networks. Alternatively, thehypothesis supporting the African origin of Homo heide/bergensis would answerthe question of how both this species and the Mode 2 technology entered Europe(Foley and Lahr2003). The entrance ofMode 2 into Europe, with or without a newhominin species, coincides with the Westerhoven interglacial (MIS 15) into theCromenan period. It has no chronological relationship with faunal dispersals.

Climate cbanges push all species to adapt by means of moving or changingstrategies, both of which occurred among hominins. In this sense, it is absolutelyconsequent that some faunal dispersals temporally co-occurred ·with somehominin ones. However, hominins did not accompany faunal dispersals owingto any kind of subsistence subordination. Simply put, both fauna and homininscoincided, but not always moving in the same ways. It was the non-standardizedbebavior of hominins, as well as their capability to adapt through both somatic(generalist diet) and extrasomatic (technology) means, that allowed them tomaximize the supply of resources in new and changing ecosystems. In fact., theevidence suggests that technology made all hominin dispersals possible, sincenot a single hominin fossil has been found outside Africa in contexts withoutassociated lithic assemblages.

But moving is not successful in tenns of continuity, nor is it adaptive toadopt new strategies until they are integrated into the cultural background of eachcommunity-that is, until the new ones have been socialized. It was the socializationof both technology and more versatile strategies of subsistence and behavior thatallowed Plio-Pleistocene hominin groups to expand across the planet.

After their emergence, new ideas or inventions may be generalized withina community. In this sense, the individuals of a group will observe and evaluatethe potential and advantages of the inventions. If tbey are welcomed and becomegeneralized in use, then the new concepts have been socialized, because theyhave entered into the behavioral core of the community. Obviously, socializationrequires a capacity to generate and integrate new information into the knowledgesystem of a group, as well as the capacity to diffuse this knowledge to othergroups, through effective contact and communication. Finally, the developmentof all these components enhances the generation of new social formulas forsubsistence, culture, territoriality, and lifestyle-that is, the social reorganizationof the community. This is the way through which all cultural novelties becomehistorically adaptive.

. Therefore, the chain ofprocesses leading to the earliest Homo dispersals out ofAfrica can be summarized as follows: First, the emergence ofa new technologicalmode among the hominin groups; then, the generalization and socialization of


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EURASIAN GATES: TIiE EARLIEST HUMAN DISPERSALS 217

tbis tecbnology, wbicb implies the adoption of new bebaviors and reorganizationof adaptive strategies; later, demograpbic growtb and evolutionary success oftbebominin groups as a consequence of baving improved tbeir adaptive capabilities;and finally, the occupation of new territories.

NOTE

The authors are grateful to Carolina Mallol for her help in translating part ofthis paper. Wealso thank Bienvenido Martinez-Navarro, four JAR reviewers, and the Editor (who alsocorrecJed the English) for their comments. The authors belong to the Atapuerca ResearchProject MEC (001) CGL2006-13532-C03-01.

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