Amerindians - Testing the Hypothesis - Sardi Et Al

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Human Palaeontology and Prehistory

Amerindians: testing the hypothesis about their homogeneity

Marina L. Sardi a,b,*, Fernando Ramrez Rozzi a,Silvia L. Dahinten c, Hctor M. Pucciarelli b

a UPR 2147, Dynamique de lvolution humaine , CNRS, 44, rue de lAmiral-Mouchez, 75014 Paris, Franceb Departamento Cientfico de Antropologa, Facultad de Ciencias Naturales y Museo, Universidad Nacional de la Plata,

Paseo del Bosque S/N, 1900, La Plata, Argentinac Centro Nacional Patagnico. CONICET. Boulevard Brown S/N, 9120, Puerto Madryn, Argentina

Received 20 October 2003; accepted after revision 19 April 2004

Available online 29 July 2004

Presented byYves Coppens

Abstract

One of the postulates of the most accepted models about the peopling of the Americas proposes that Amerindians constitutea homogeneous population. The aim of the present study is to assess the homogeneity of the craniofacial morphology of SouthAmerican Amerindians in a worldwide context. Variance of each geographical region and FST values among local populationswere obtained. Results indicate that the southern Amerindians present a high level of morphologic variation, even whengeographically closer populations are compared. The origin of Amerindians as well as micro evolutionary processes, whichwould explain the high variability, is discussed. To cite this article: M.L. Sardi et al., C. R. Palevol 3 (2004). 2004 Acadmie des sciences. Published by Elsevier SAS. All rights reserved.

Rsum

Amrindiens : valuation de lhypothse sur leur homognit. Les modles les plus accepts pour expliquer lepeuplement desAmriques supposent que lesAmrindiens constituent une population homogne. Lobjectif de cette tude est detester lhomognit de la morphologie crniofaciale des groupes Amrindiens de lAmrique du Sud par rapport unchantillon mondial. Nous avons calcul la variance pour chaque rgion gographique, ainsi que les FST entre populations

locales dune mme rgion. Les rsultats suggrent que les Amrindiens sud-amricains prsentent une variation leve.Lorigine desAmrindiens et les processus micro-volutifs qui pourraient se trouver la base de la variabilit amrindienne sontdiscuts. Pour citer cet article : M.L. Sardi et al., C. R. Palevol 3 (2004). 2004 Acadmie des sciences. Published by Elsevier SAS. All rights reserved.

Keywords: Amerindians; Craniofacial morphology; Variation

Mots cls : Amrindiens ; Morphologie crniofaciale ; Variation

* Corresponding author.E-mail address: [email protected] (M.L. Sardi).

C. R. Palevol 3 (2004) 403409

2004 Acadmie des sciences. Published by Elsevier SAS. All rights reserved.

doi:10.1016/j.crpv.2004.04.001


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Version franaise abrge

Les modles de Trois (TMM) [14] et Quatre(QMM) migrations [25,26] qui expliquent le peuple-ment des Amriques suggrent que les Amrindiensrsultent dune seule vague migratoire provenant delAsie du Nord-Est et quils sont donc morphologique-ment homognes. Lobjectif de cette tude est de testerlhomognit de la morphologie crniofaciale desAmrindiens dans le cadre de la variation mondiale.

La variation morphologique a t value par un modle dvolution neutre [31,32]. Daprs ce

modle [30], la variance rgionale peut tre value enprenant, soit les rgions gographiques (ARG), soit lespopulations locales de chaque rgion (APL) commeunits danalyse. Dans ARG, la variation dune rgionest dduite par sa variance observe. Dans APL, le FSTindique la quantit de variation dune rgion qui estdue aux diffrences entre populations locales. Septrgions gographiques ont t incluses dans ARG (Eu-rope, Asie, Australasie, Polynsie, Afrique sud-saharienne, Amriques et Patagonie) (Tableau 1) ettrois populations locales de chaque rgion ont t uti-lises pour raliser un APL. Trois autres populations

amrindiennes ont t ajoutes aux prcdentes poureffectuer un APL seulement entre des Amrindiens.

Quarante-trois mesures de Howells [18] (Ta-bleau 2), aprs standardisation Q [5], ont t utilises.Les matrices de variance-covariance ont t calculesavec une hritabilit de 0,55 [7].

Dans ARG, le FST non-biais est de 0,1492 (erreurstandard = 0,0015). Les rgions de Patagonie et dAfri-que sud-saharienne montrent les variances les plusleves (Tableau 3), ce qui indique que la variationdans ces rgions est plus leve que celles des autresrgions. Dans APL, le FST pour les populations de

Patagonie occupe la troisime position indiquant unevariation plus leve que celle dautres populationslargement disperses (Tableau 3). Les FST pour lesautres rgions sont trs semblables ceux obtenus parRelethford [30], dont la diffrenciation est associe ladistribution gographique. LAPL fait avec les chan-tillons dAmrique du Sud montre une distribution deFST avec une moyenne de 0,1573, un FST minimum de0,0890 (pour les populations de Patagonie) et un FSTmaximum de 0,2180. Bien que les populations de Pata-gonie aient une variation leve par rapport aux popu-

lations dautres continents (Tableau 3), la variationentre elles correspond la plus faible parmi les Am-rindiens.

Les rsultats indiquent que les Amrindiens ne pr-sentent pas une morphologie crniofaciale homogne,comme le veulent les modles plus accepts. La varia-tion leve des Amrindiens peut tre due : (a) unpeuplement plus ancien, (b) plusieurs vagues migratoi-res, (c) des populations grand nombre dindividus,(d) un taux lev de croissance dmographique, (e) undegr lev disolation et (f) des facteurs environne-mentaux. Le point (a) tant propos par quelquesauteurs [11,15,16,27,33] semble probable, mais lvi-dence archologique doccupation est faible [8]. Lop-tion (b) est carte, si on considre la variation enAsie,car la diversit observe en Sibrie [20] ne peut pasexpliquer la variation leve des Amrindiens. Destudes en gntique ont trouv des rapports entre lesAmrindiens et les Asiatiques [6,23], mais la localisa-tion gographique du groupe anctre reste douteuse[11,22,23,38]. Par ailleurs, la variation leve peutrsulter du fait que certains groupes amrindiens soientdes descendants des Paloamricains (premiers grou-pes peupler lAmrique la fin du Plistocne) [13],ce qui nest pas considr par le modle de Quatre

Migrations [25,26]. Les points (c) et (d) devraient trecarts, car la taille des populations des chasseurs-cueilleurs augmente rarement et encore moins dans unenvironnement instable [33,37]. Les points (e) et (f)sappuient sur lvidence archologique qui suggreque les Amriques taient largement peuples vers13000 AP [8]. Lisolement des groupes d un envi-ronnement aride qui aurait rduit la mobilit auraitfavoris la drive gntique en augmentant donc lavariation (point (e)). Mais ladaptation mise en vi-dence par le registre archologique nest pas unepreuve de variation morphologique (point (f)). Mais,

bien que linfluence de lenvironnement ne puisse pastre nglige, il reste difficile de comprendre pourquoila pression de lenvironnement aurait t plus impor-tante en Amrique que dans le reste des continents.

En conclusion, la variation des Amrindiens du Sudest trs leve. Elle peut tre explique par : (1) unegrande dispersion des populations, avec un faible nom-bre dindividus, dans un environnement instable etfragment ; (2) la prsence, parmi ceux-ci, des descen-dants directs des premiers habitants, les Paloamri-cains ; (3) lexistence de plusieurs vagues migratoires

404 M.L. Sardi et al. / C. R. Palevol 3 (2004) 403409


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vers lAmrique en provenant de lAsie, mais dautresrgions gographiques ; et (4) un peuplement plusancien que 13000 ans. La contribution faite dans cettetude, du point de vue de la morphologie crniofacialechez les Amrindiens du Sud, ne permet pas de rsou-dre le dbat sur leur volution, mais il oblige recon-sidrer les modles plus accepts.

1. Introduction

Among the theories concerning the peopling of theAmericas, the Three [14] and the Four [25,26] mi-

gration models (TMM and FMM, respectively) are themostly accepted. In both models, the term Amerindi-ans refers to all American Indians, with the exceptionof Na-Dene, Eskimos and, according to the FMM,Paleoamericans the most ancient inhabitants ofAmericas. They propose that Amerindians are descen-dants of a unique migratory wave coming from theNorth-East of Asia, so they would constitute a homo-geneous population. Nevertheless, some genetic [3,9,28,35], linguistic [27] and morphologic [2,17,21,34]studies suggest that Amerindians do not behave aspredicted by these models. The aim of this study is to

assess the homogeneity of Amerindian craniofacialmorphology in a worldwide context. According to theTMM and the FMM, it is expected to find low levels ofmorphologic variation among southern Amerindians.

2. Materials and methods

Assessment of morphologic variation was based ona neutral model bound approach [31,32]. Accordingwith Relethford [30], there are two ways to assess theregional variance: the geographic regions or the localpopulations of each region can be taken as the unit of

analysis (among-regions and among-local populationsanalyses, respectively). In the first case, a unique FSTfor the entire comparative context is obtained whichmeasures the proportion of the total variation that isdue to differences among the units analysed (regions).The variation is deduced from the observed variances.In the second case, a FST for each region is obtained. Ithas the advantage over the first case that it allows todeduce which part of the entire variation in a geo-graphic region is due to the among-local populationsdifferentiation.

For the among-regions analysis, seven world re-gions (South-Saharan Africa, Europe, Australasia,Asia, Polynesia, Americas, and Patagonia) each onerepresented by three local populations were compared.For the first six regions, data come from the Howellsdatabase and the selected populations (Table 1) arethose used by Relethford [30]. For the seventh one,Patagonia, measurements were obtained by one of us(MLS). Patagonia seems to have been occupied by aunique migratory wave. Even when the contact withnorthern groups has been demonstrated [12], an in-tense gene flow was improbable and moreover, thisregion never underwent an important demographicgrowth [1]. Thus, a low variation is expected amongPatagonians. They are represented by two northernsamples: Chubut River Valley (CH, n = 99) and RioNegro River Valley (RN, n = 72), and a southern one:Tierra del Fuego (TF, n =45)(Fig. 1).The among-local

populations analysis was made with the samples in-cluded in the among-regions analysis. A secondamong-local populations analysis was carried out withPeru, CH, RN and TF and with three additionalsamples (Fig. 1): the Bolivian sample (BO, n = 19) ofaymar-quechua origin, localised in the Andeanhighlands; the hunter-gatherers of the Paran River

Delta (PD, n = 38), with an antiquity estimated byarchaeological associations around 2000 years BP[39]; and the western Pampas sample (WP, n = 37)which represents a Mapuche population that settled inthe Pampas after migrate from the southern Andeanregion [4]. Data for these three samples were collectedby one of us (MLS).

Forty-three craniometrical linear Howellsvariables[18] were used in the study (Table 2). They weretransformed in dimensionless vectors through theQ-standardisation [5] to remove size, whose variationamong Amerindians is attributed to environmental fac-

tors [17,21,34]. Following previous studies [31,32]Table 1List of compared samples derived from Howellsdatabase.chantillons compars, drivs de la base de donnes de Howells

Region n Populations

South-Saharan Africa 283 Teita, Dogon, ZuluEurope 317 Norse, Zalavar, BergAustralasia 298 Australia, Tasmania, TolaiAsia 261 Hainan, South Japan, North JapanPolynesia 294 Moriori, Mokapu, Easter IslandAmericas 281 Peru,Arikara, Santa Cruz

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additive genetic variance-covariance matrixes, fromthe phenotypic ones, were calculated with a heritabilityof 0.55. This value was estimated by Devor [7] andrepresents an average proportion for metric characters.

In the among-regions analysis, the regional variationwas calculated through observed variances; in theamong-local populations analysis, the regional varia-tion is represented by the FST values.

3. Results

In the among-regions analysis, an unbiased FST of0.1492 (standard error = 0.0015) and a mean variance

of 0.819 between all the regions were obtained. Pat-agonia and South Saharan Africa show the greatestobserved variances (Table 3). In the among-local

populations analysis, the FST values are very similar tothose obtained by Relethford [30] who found a signifi-cant association of the among-populations differentia-tion and the geographic distances. In this context, Pat-agonia occupies the third position after Polynesia andthe Americas showing a higher variation than largerregions (Table 3).

FST values were obtained with BO, WP, PD, CH,RN, TF and the Howells Peru samples. Each combi-

nation of three South American samples producedthirty-five normally distributed FST values. The mini-mum value is that of Patagonia (0.0890), the maximum0.2180 represents the comparison of WP, PD and Peru.The mean of the distribution is 0.1573 with a standarddeviation of 0.0323. FST values for the geographicallyclosest groups are: 0.135 (CH, RN, PD), 0.162 (CH,RN, WP), 0.179 (RN, PD, WP) and 0.203 (CH, PD,WP).

4. Discussion

Results indicate that the South Amerindians are farfrom being a homogeneous group in cranial morpho-logic terms (Table 3). Gene flow with Europeansshould be discarded due to the absence of morphologicaffinities between Europeans and these Amerindiangroups [34]. The low variation among Patagoniansregarding otherAmerindians supports their unique ori-gin inside America. However, it is worth to know thatthese same groups from Patagonia show the first andthe third highest variations in a worldwide context(Table 3). The high variation in Amerindians can beassociated to: (a) a greater antiquity of the peopling,(b) several waves of migration, (c) greater effectivepopulation size, (d) greater rate of population growth,(e) greater degree of isolation, and (f) adaptative fac-tors.

The option a), a peopling before 15000 years ago,has been suggested by many authors [15,16,27,33].This option seems probable even when archaeologicalsites older than 13000 BP [16] are not accepted withoutdoubts [8].

Option b) is difficult to accept in the context of theTMM and FMM models that suggest that Amerindian

Fig. 1. Geographical localization of South Amerindian populations.Fig. 1. Localisation gographique des populations sud-amrin-diennes.



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characteristics must reflect the northeastern Asiaticones. However, in Siberia or Beringia a high variationis not reflected before Holocene times. Kozintsev et al.[20] mentioned that Baikalians, about 8000 years BP,had extremely flat faces and the later inhabitants wereless mongoloids due to gene flow with European

populations. But for both the TMM and FMM thearrival of Amerindians was at the Terminal Pleis-tocene, so the Siberian holocenic diversity does notexplain, therefore, the Amerindian one.

If another than an eastern Asiatic origin is consid-ered, the morphologic diversity is easier to be ex-plained. Genetic studies support the Asiatic origin ofAmerican Indians [6,23,24] without accord about thegeographic localization of the ancestor [11,22,23,38].However, some Amerindian tribes show genetic char-acters that do not accommodate with the suppositions

of homogeneity. Brown et al. [3] found a haplogroupX extensively distributed in America and present inAsia only in theAltaians of South-Central Siberia [10].Even when there is not conclusive evidence that Amer-indians have more than one ancestor non-Asians the Amerindian diversity does not seem to be com-pletely derived from the Asiatic one.

By other hand, if the double-migratory event (Pale-oamerican and Amerindian) for the main part of theAmericas is accepted, the eventual genetic relation-ships between both waves must be discussed. Morpho-logic divergence between Paleoamericans andAmerin-dians would suggest that the former have notoriginated or at least did not genetically contributed tothe later [19,25,26,36]. However, Powell and Neves[29] proposed that micro evolutionary processes couldbe responsible for that morphologic divergence. It isalso probable that some South American groups aredirect descendants of Paleoamericans as it was sug-gested for the Perices of Baja California in Mexico

[13].Options (c), (d) and (e) are proposed by the genetic

neutral models. Options (c) and (d) do not seem prob-able: according with Rogers et al. [33], the hunter-gatherers increase very rarely their population size in ashort period of time moreover during climatic instabil-ity. Steele et al. [37] developed a model for Pale-oamerican dispersion in North America during thePleistocene-Holocene transition and they deduced thatit was not followed by an increase of the populationsize.

Table 2List of Howells craniometric measurements.Mensurations crniomtriques de Howells

Glabello-occipital length Orbit breadth Cheek heightNasio-occipital length Bijugal breadth Supraorbital projectionBasion-Nasion length Nasal breadth Glabella projectionBasion-bregma height Palate breadth Foramen magnum lengthMaximum cranial breadth Bimaxillary breadth NasionBregma chordMaximum frontal breadth Zygomaxillary subtense NasionBregma subtenseBistephanic breadth Bifrontal breadth NasionBregma fractionBizigomatic breadth Nasio-frontal subtense BregmaLambda chordBiauricular breadth Biorbital breadth BregmaLambda subtenseMinimum cranial breadth Dacryon subtense Bregma-Lambda fractionBiasterionic breadth Interorbital breadth LambdaOpisthion chord

BasionProsthion length Simotic chord LambdaOpisthion subtenseNasionProsthion height Malar length inferior LambdaOpisthion fractionNasal height Malar length maximumOrbit height Malar subtense

Table 3Estimation of variance on the seven geographic regions.Estimation de la variance dans sept rgions gographiques

Unit of analysis Regions Local populations

Region Observed

variance

FST Mean

variance

South-SaharanAfrica

0.87 0.0825 0.938

Europe 0.824 0.0549 0.925Asia 0.816 0.0371 0.961Australasia 0.742 0.0686 0.924Polynesia 0.793 0.1205 0.836Americas 0.789 0.1005 0.893Patagonia 0.901 0.0889 0.926



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Option (e) is partially related with population size.The archaeological record shows that around13000 BP South America was widely occupied. Due tothe arid conditions after 13000 BP, populations wouldremain near waterways sites limiting their mobility [8].Thus, the fragmentation of the geographic range wouldcontribute to variation through genetic drift.

The option (f) was exceptionally mentioned [17,21]and it is discarded. The effect of environmental factorson morphologic traits is not denied but its contributionto inflate or reduce the distances among populationshas not been proved. Moreover, it is difficult to under-

stand why the environmental pressure would be greaterin the Americas than in any other region.

5. Conclusions

These results show a high level of craniometricdiversity among the southern Amerindians. This factwould result by different but not exclusive processes:(1) a low population size of first migrants followed by agreat dispersion in different and unstable environ-ments; (2) the genetic contribution of the ancient Pale-oamericans to the most modernAmerindians; (3) many

migratory waves coming from different geographicregions; and (4) an old settlement (older than13 000 yr) of Americas. The high craniometrical varia-tion found in this study is insufficient to solve theproblem of their evolution but it constitutes an elementthat questions the established idea about Amerindians.

Acknowledgements

This work has been possible by Scholarships of theUniversidad Nacional de La Plata (Argentina) and aFondation Fyssen Grant held by MLS and by the

French-Argentine cooperation CNRSCONICET: Laplace des Amrindiens face au contexte mor-phologique et gographique de lEurasie.

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