The History and Geography of Human Genes








DOI 02173

Cop?'rg"r _ !994 by Pn:cet='- L'm',,:n:_ Pr-'_s P'._iis'_.-dby Princeton ',.':i_ em:_ ?r':ss.4: '_, :i am $,":._e:.

In_ne'._::ec Ki_gdcPn_ce:c_ U:_,.,rs: D Press, Chic_es:er, V,'_: S_sfx Xi/ R_g,t_sP, ese_ed l,_br_c,, of Congress C_tcloging-/n-P_,v,_-c=n._ _z',z Ca_ail:-Sffr L.L.( :..a-, La:_ Luc_' _922"Fte h:s:oH' and geaT-a;__ zt.h_tz, p. cm. [nciudebi s b_io_ra;_'n_ ,ic (_r_ ai nce as .td index. 1SBb,'_69i-0_v50-4(_nabndge_ clc:h ¢d) ISBN C,-691-02905-9 (abridged 7bk ed?, T. Haman pcpu!a:ion gene:ics----His:oD. 2.Human _,o'_u_ic,n I Mfno_i, Paolo, i94&I[L Tide. _H43_ C395 1933 5?_.2:'5---d_20 ,_3--]_"_, :,% 1"k:s book has been composed n Trees R_man D_:gncd by Jan L:!}? Tie :_'_er .:_us,,"ranon :sa map cf:_:*'or'.4 shov,i_g four major e'._nio r_ncns. Afnc_r,s ar_ ._ e _o_.,A_s.'raiians re& and CaucasiansT'een. _'4on_c)o_d_ sho_. :he gr_a_e_; ",an_:ior, r_t_mmg some similarises _i;h Europe.ans on one side itligh: bro_._,_ee_ish tinge in m_dle Si_,_-na_ ' wi_.h A_sn'aiians on th_ other Capinkish Color inpar..s of America and on the Wlly 10I:). "r_c ext.-_:si\.¢_-"adifn_sdL._to adiT1ixr'azcs beI'_vee_ A_ncans_nd Caucasoids inNoah Africa. an_ b_,"*ec Ca_casci_.s n and Mongoloids m Middle A6a. _r_ c{e.ariyv_sib_e. (S_e¢,_aRtt" _¢,o) P'nnceton I._niver_i_ Press _ooks a,'_ pri_d m_: t_c guid_iin_ on aci&fr_ palxr and . IIP!a,.=_ A1b_":o, , 1941ge_.ts

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an Production GuideJmes for Book l.mng_'.'i_ of_e Council on Library R_urc_s Firs pr z inting of the abnc(gcp_ c_p_rback edidon, i996 _r4cclin theUnite_l S_es of America I09_ 765432

DOI 02 ] 74

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bu_iea:n E.:rcx cf :_e fr':q._'z::i, le'c_of ._ F-508 ¢_s:',: fi_ros _ -r...tar& ,_:at;_e ',0 olhe."_ulao:s _,.'.

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fibrcss: rrod':ed frcm De,::¢ Earc_2_ cc_sc,-,:um, cx ::.s: c

a::c :he.fore ""_ ammo ac d_ _s vet .... _.u.,,t, reaching a'::r::x:mat_'. .... x 70Q of ai'. kno',_n mutants dete,,-n..L':.Jnc ,;'_ _;s_ _ "t_ distribution iv. Eurcoe is shcv.n in fiE'Jre "_ 14.:0 The frequen,:v at bi_h of rh- disease itself _s less '_,ei', kzo'_n, but is fount, on the a_era_,' in 1 of 20C'0 ct" a_: ' bills in E_rope It va:':es .,.,, _- e,,,.na< ' ' from count--: to countr:, _ut the frequenc? era genetic disease is ::ct ai,,.a,,s, easy to estimate a,._ura, ...... e:;. "V'e_' "-con" data indicate that the relative frequencv of :he common allele fcr c'.s:ic fibrosis (', F-508) is espein a re!art;el': small sample of Basques (see fi__2.1,.;.10,. The =__,'oo_"h_c_._ .,. ,. spread of the frequent allele is c,uive similar to the fimt synthetic map for Europe and therefcx :o Ihat of the Mesolithic (pro-Neolithic) European gone pool. The conclusion would be that the mos: con:tmon cystic fibrosis al]ele in Europe is older than the spread of agricuJture and must have been especially frequent among Mesoli_ics. while most other alleles come from the Middle East or originated in Europe after the Neolithic diffusion,

.Many of the po!ymo_hisms ....... reBLs::.':,ce malar:a and other diseases may be -_...... <'_ °:': young _ut the £reat maiont', of sob mo_'ic a:'e '_'_, tea'. ',;.e ha_e studied in sec:icns 2.10 and _ 1_, _-e fczr, d near!'. -;' continents and .herefore ." ha,e a."te_:::ec :he spread of am.h. Oniv a fe_ are knc'_,n a: :::e DNA ]e'.el, and it is impossib:e to sav. v..,_,o.' ..... ...... _.."'_--._, ent mutations are hidden behind the same aEe:e. "Dee lO0 RFLPs studied in sectJcn " ' hole',or, are li],.e:'," to have had a unique mutational origin, ar._ the !_,:ea: major.ty of them are found in all continents excea: the Americas v, hich ha,,e not yet been stadied for these polymorphisms). T'here are reasons tc think :ha: most human polymorphisms are much older than the date of spread from Africa. Theoreficall,,. in the absence of selection, the average age of poi',mo_hisms is comparable to that of the species. An estimate of the average time of appearance of the mutant allele of eiaItetic human RFLPs is 700,000 years (Mountain et al unpuhl.).

cial!',' high






EVOLUTION interpretation rests on the assumption that evoluticrlau rates are reasonably constant. Some further comfort is derived from the consideration that the rates involved in the evolution of polymorphisms of nuclear genes and those of mtDNA are different; the first are determined mostly by differences in gene frequencies separating t'.*,o popu]ations, and the second by numbers of mutations separating two individuals. The dates to which they lead for the bifurcation of Africans and non-Africans ax different, being of the order of 100 kya and 200 kya

The analysis of phylogenetic trees has used many different sets of data: nuclear polymorphisms tested by electrophoresis or immunological techniques, restriction fragment polymorphisms of nuclear genes, and mitechondrial DNA. All methods show a somewhat greater difference between Africans and non-Africans than between other human groups, and offer some informatior on dates supporting the interpretation that the origin of modem humans was in Africa, from which an expansion to the rest of the world started about 100 kya. The

DOI 02175

GE",ET;C -espe::ti'.el? Oi:en !he ,*.<.s :n v.hich these de:as were _btair:ed. :h{s d_f.'erenae should be expected, v,.!th the _econdbeing greater :hart the first by a." amour:: that has _,ot been de,e..-"mi ec v, . bur perhaps might be es::mated on me basis of theore: ca[ conside_dons. AZina'..the_is _asic agreement be:'._een :he trees ob¢ained with mtDNA :rod w:th nuclear, marker¢, the latter gi',ing ,'T. u:h mot_ entail. It is '*c.v:h reme., mat zav:':g :-e mito.:'hondnaI ancestor "E_e" has gene,'a:ed the false be>el that there ',,,as a time ',a, hen :here >,as on!> one woman alive. On the question ef .:!ace of origin, the archaeoIogicaI fieldisdivided. Anumberofp£eoan'.t_"opologistsbe!ieve :hat modem _umans originated in Africa. from '.,,hich they spread to the rest of the world beginning about lO0 k?a. -UP.isis in agreement v.ith the genetic data. A faiKy large number of anti",ropoiog!sts rese_'e their opinion. Another group belie'.es :hat t_e e,clution of Homo sa;,'eas, and perhaps eve:: its predecessorH, erec,'as proceeded in parallel all a,,er :he Old X V , or]d. _d :here v,as no expansion from Africa. The mi:ocho,,:.dr:£ 2a:a are. at :his point, the most useful it: heiF:ng ,'o re'eel :,'.is .h)'Fo:_esis. given :hat the or.gin of extant t>?.es of Asian mtD?'-A :s more re:on-.: :ha,":, this hypothesis v, auid imp'.y, h is no,. >el pcssib',e, hov, e,.er, to exc'.ude complete!> a parz:a_ pm'-:_ai?a::oa of archaic H. saF/e,,:.s from :he O_d ',X, br!d. ",'e',_ data and me:hods of ana<,.s:s may help in this direction. What is '.eD dif_cut: :o conceive _s a parailel exciazion over sure a vast expanse of land. given the limited generic exchange :hat could ha',e occurred in earlier ::rues ,'Th.e ,:at'act:) of the human genus to expand rapid}- o_er a i_'ge fraction of the Ear:,h's sur,ace is more in tune '.,.ith :he .tea of specific expanMons from a nucie'_" area of origin. Su_.h expansions must have been dete,.":nined b.', some impo=ant ad',antage, biologieaI or cuhuml. It is not difficuh :o accept the idea that the expansion of modem humans musl hare been strongly influenced by "he possession of greater skills in communication by !anguage. This increased ability to communieate is likely to have been extremely' useful in favoring exploration and tra,,el to unknown lands. Other technical improvements may have favored a trend to expansion, Ahhough modem humans have now been found to have lived outside of Africa (in the Middle East) by about I00 kya, humans of this time in both Africa and the Middle East were biologically very, similar to modem humans but culturally much less developed than at the time the real expansion began, perhaps 50 or 60 ky later. Many things may have happened in the meantime, in terms of cultural maturation and, perhaps, forward and backward movements between Africa and West Asia. Neanderthals are found it, the Middle East after the earliest local appearance of modem humans in the same areas, and it has been suggested that they may have gained, or regained, lost ground in that period. The time between 100 and 50 kya {or, perhaps more exactly, between 90 and 60 kya) is

H[$'7OR'_ 3F ",*,ORLD POP'..La,'7;©x5


currer,:t',' a blank from an archaeological point of _ie'..,, "A,ehope C_a'.no,a, d_sco',,e,",es wiF, ii'.um_na:e :t A: the moment, the in,: cations are tha_at the end of the b!a..C< pe:'iod mode:T,h-mm_s emerge" with a ne'_ stone technologF and stm._eda radiat:on that took them to Europe, Australia and New Guinea. and Ame,.' %,'he'J_er the', partiall': mixed with or totail) supplanted e_iier inhabitants--for example, Neande."thals in Eurepe and a_haic H sapiens in East Asia--is difdcuIt to sta'e pro. cise!y on the basis of present knowledge. Linguistic and cultural diversity increased consT,[¢uously after that time. and the major linguistic fa-":'.dlies probably began less than 50 kva. Most of them are between 25 and 5 k>' old. Genetic dating of ling,aistic faro,ities can only be approximate, but it agrees with ideas _'xpressed by a few linguists. Moreover. the archaeologica. record shows increasing diversification, probably pa.-alle[ v,ith tha¢ of Ianguage. An unsoi'.ed problem is determining the route by' which the East ,,,.as reached. Differences betv, een East Asia and Scu:heast Asia make it reasonab',e to hypothesize that there might ha_e beer:, tv, o routes, can th,.rough Central Asia and one through South Asia. "Vet3,little. if any'. evidence of them exis:s today (dg 2.'.5.;) The occupation of Australia and Nov, Guinea ,.,,as the mgor success s:o D of the southern rou,'e, but it e_e::aaHv led to an e,,o'.utionar', cul-de-sac, as the sepa.r"auonbe:v.een Oceania and So,,.::heast Asia increased with the using c5 the sea !e'.els in the times after :he last glaciation It ',,.as only v,ith the development of new nautical skills, 5_,?, ..6000 >ears ago. by South-East Asian populations ',,,he _,ere also good farmers, that the Pacific routes v,ere increasing'.,, used In the last 3CK)0-3500 years, the expansiGn that generated the colonize:ion of Polynesia began. most probably originating in a nuclear area in Southeast Asia. There are two weaknesses in the present a=:a!>sis, which will certainly require future work. One of them is the very short branch linking Caucasoids and. in particular. Europeans to the phylogenetic tree. One hypothesis is that they might have originated from an admixvare between their southwestern and northeastern neighbors. Africans and Mongoloids, between which Europeans are sandwiched. One cannot completely exclude other hypotheses. Pani¢ularly serious is the possible bias resuiting from the fact that almost all known genetic polymorphisms ha,,e been detected in Europeans. It will be important to remove this bias. especially in future data collections. Another area of doubt is the relationship between New Guineans ÷ Australians, Southeast Asians and Northeast Asians. Our results have not settled this question unequivocally, It seems likely that the uncertainty arises becavse Southeast Asia is poorly known and may be heterogeneous, v,ith some populations having an important genetic component in common with northern Mongoloids and others with people from Oceania. The

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and in some cases of local expansions.f mc.' dichotomous tree may be unsatisfacto_ in this part of the world. Throughout the Paleolithic. anthmpometric data). "E":.eral different agricultural developments.> ma? be in Far't.isexplains the discrepancy between the evolutiona_. The first principal component therefore extends in this dir=ctic:_ and exT]a:ns -_5% of the tcta] human variation.due to ancient admixtures.:em humans :n :no last '.G'0k>.ed from data on genes and on skulls (or.s:or:znd rc<:t.15. A major conclusion is that linguistic and genetic: e_olution are closely relatcd. seem to agree with an early arrival. Mongoloids !ea_:ng one to _onder whether the) ha'. perhaps because it first required a genetic and cuhurai adaptation to the more rigid climates of Northeast Asia. allowing the exploitaticn of domesticated plants and animals developed in the thr_e major nuclear areas of agriculture. few of which have been well examined. leaving greater chance for random gonetic drift to produce considerable diversification. but more abundant and better evidence would be necessary for developing this explanation further The passage from Asia to America was later than that tc ?'. Given the greater limitations on life in the north. The consequent increases of population densities began a progressive treezing of drift effects. _hich is detectable also on modem and fossil bones.ns. contributed in an impercant ..000--I00. In _pire of this. much of the gradient of the human gone pool goes from west to east.he late Paleolithic..'n.15 lexpressthesecons_deraticns. and :he a"ro_ of no. Farmers' expansions.4.3 and 2. '..usu-aiiaor Europe.hem Mongoloids pointing Soulh in figure 2. In .1 Poss =ie :'_. and the occupation of the rest of the world proceeded from this continent. which extended to ecologically similar areas. A dichotomy is thus obser".ear ]ene_ in.e more in common than sho_n b? the trees of sections 2.' histories re¢onstruc. culminating in se. Population size of a continental or subcontinental area at the beginning of expansion may have been on the order of 50. showing oni. a full'. In other words.cnc. perhaps around 30 kya. followed by those of nomadic pastoralists. much of human action was in Asia. but a greater influence en genetic factors involved in the adaptation of body build and bodii._ay to changing the patlems of gone geography. Genetic data.. perhaps under increasing pcpulation pressure and climatic changes. These innovations caused the beginning of more rapid population growth.estigated.. in general...ed bet'_een genetic data and observations based on the physical constitution.'.. Because genetic divergence was subject more to random than selective forces.. opportunities still remained for the surviral of much local diversity. Asia was like a relatively narrow. There are also some undeniable physicai simiiari:ies between northe_ and sc_:he:m. '.. especially in refugia. moderate.uonce of climatic factors at the le'. _f an. hetemgenei'. possible uncertainties are discussed further in chapter 6. did humans devetop new food technologies. 2. On!y in the last 10 ky..n_. large landmass developed more in longitude than latitude.56 CH._of s exp:._=TER l_g.000 individuals. however. population numbers remained small.hich notoriously respond to £ima_e. In this chapter we have seen DO[02177 . el of "he nun'. sur"face characteristics.

sL = 2 -_ :_ B So_.m. resumab!_ '_'o be stmng!y cor're'.c re_<.dges spoken b? :hose groups.s .. of genetic history. .. .t. :'g'.ere iess de'. Genes are cie_-I? ...est> j ru. . given by' Ruhlen (1987). aiso takes place': am.ain= :ha: _enes and l:"__'. heloed.L% _ o .stoD' of humanity is shrinking continuously.."r.... pa. and _. :00) Ht...-za eta!.e absence of sc.ersia! (Fa'. alli-Sfo.o.ared b': all existing human languages seems hkely :o be a product of a final step in linguistic e'." a'.":. in the following_ zhamerS.eric and Imguisdc evo'. modem humarts in '. m:a_ons_:pbe:ween human generic va.ans '_' 13" . Unfortunately.h .?ies of it.nd in eaa:" of the successtve major groupings{:oF!of _'.. to :sobe_placed happe.. .. Ca'.s ob.2 Aacordin_.. il".'.ans mission o_...n t:'. • spread by them to the whole world An interesting relationship has been observed by Foley _.rcst . ..a.parallel on all populations of interest._.'-r.. he has shown that there is a very strong linear relation (r = 0. " or . it adds to the persuasion that linguistic evolution goes hand-in-hand with the spread of modem humans.nla'_e..re_ aus_-a_La a.'oluticn of both '_'_nd cr._ _'. 2.e _xam ° b. paper ( ' _ ' v.ess. The analysis of thegenetics of human populations mquires an enormous mass of information.-_ati°r:son sp::i.. I985: and information on the numbers of languages per fami._ . :tic.emJ m'. .k 1975). The !i:_'uisfic infen..91) between genetic distance between two groups sepat.s. .n_.":...istorym '. ura.e!o_ed lin_uisficali'. ."p. completely mr..._agescana mare. v._. ..'tion (and.. He..croatasset '.t:enetic tree (CavalliSforza etaI.ena t:"ansmitted m basicaily the same ".'.. :s 2 s:.. ._..!y .hes:. . ha'.a:ne_ ._eaA.... gisen _at oe_es c'_n.. we are an endangered species. 1989) is con_-a'..mups....'.e t L_resse4 . From the point of vie'..-.enhe'.._a.. from ..'ailable is the resu!t of thousands of mow or less haphazard collections and a.s <L_e_e..£enera:. _..-_' tcrs.e .oba! _e'.. [ A.een get.:.esfiga_ed sue_ect.°n to gene.a?s. Today..bouomL its retrieval has rarel? been organized in an .Gu..a i ! t"-"Ma..=. OF ' . An essential requirement of a sound analysis is that a large number of genes be thoroughly' studied ... :':-_ convJc:_on that been a :.. morn generally. and it is essential to avoid delay before taking the needssary steps to preserve this imponam knowledge about ourselves.ET_C HI<TCRY. .arv.. we .': at'#.-:a:._a and Fete'man J%i.. '.. . g'.'..%..ses_ of blccd samples...ha .. from p_p ...i991).s_a :...o pEeno.%:anO } somA.. a strong !it:earrole:ons..'!..ratedby a node of the genetic tree and the number of hnguages spoken by the two groups together (see fig.'_ j 4.a:...!o Foiev. -_-_ .. OntV • of and the da:a base a. In our origins'.iacec under certain condL':c._eI!-knovvn exar'r. i%S_.a_es me'. .. bu'..and is there mm .'..... and Iang.hich peaked in a.::discrepandes _'e no: impossible. certain b'pes of cuhu:"al e. Lang in.."genetic di'... I _c_'A"'_'s s As sc. m_xture' with other forms of humans that v..t _d Ca.oo...e_er..'_ . One c_ aiso express the necessity of a relationship betv.... especia'.'_have see.ant mecha::isms of t_nsmission.._' '°" si.A.. Lage recent _placeme :".._" "'_ ac.olution of language is a key example).. Using the . IV..:'nsor peepe 'h_' .'.rs. .]exity s?'._: and :_a: it also me.. the same histonc and o . ?-'-::a. _"_'_ this _la'honsh:_ a: . The main mason for :he re!a:ionship is tha" the e'.olutaon of which the e'. unfortunately accompanied by nontrivial cost increases The number of populations that can enlighten us on the past hi.. considenng the simi!aru.2) Although this evidence is indirect.heir . and the correlation coefficient is biased upwards because the nodes of the tree _ no_ independent.alli-Sfcr'za 1985. | z _0 o DO1 02178 . 1 _ _ _ m .. bound to ents to children.'olu:ion v. .e_..: of the re'. } Itxlo-Eurogeans 6 L _m Asia' *mer+mc_ans s _ t _.15. the extreme com...r results.on.'-at!or.a st L_so. ---.1_. . I. rhaas.aman populafionsis plo_=ed agains=Ine number of ]arg._'CRLD PO _'__' -'-..m....[!991 '.. ha_e limited O.".c'.rents childmn:in traditional societies. consider'ab'._dfromas.sar E_rc_e. . .h. as does..._).ai!i-Sf.v. _a...ers_c? fc.". :o '- 5 .GE". T'. Ne.a:ed. s _o _s 20 =_ ac Gene_c ¢istance (. then have been substantial advances in the techniques of analysis.a s_..el di_er!e in s_mi'. Only perhaps one or two decades remain in which we still have access to these populations..:." al..a.. iikei. . transmission runfer:unately a poorly' in'. a. 2.'.

th grassland or desem In the Great Plains of North America.-a:i_e! that of Asia ('. which enjoys. especially in Brazil.e genes 6._.s of the lob. but it descends in altitude beth !n :be zo_h ::. The rest of the continent is drier..=-as of agriculture 6A.'rid fro::_ Aiaska :o Mexico a_d i_ :he reg._ popu]atJon !e_e] was compa.II A:'.e_e:a:_cn are ver?. about c. Three major demographic changes took place after disco'. sho_ springs.or :f -_mer_ca 6 "_"Beg:rn.e!. climate and '. and :he!r cumuiau_e a..eneucan'qFs:s of Amerce 6.enabcxgines) of the inhere:ants of :he . mort.epopulation in North America (the United States and Canada) and in the southern pan of South America.ances: the southern of the su_ace of :he :on:men: The rest of NoT& America is re'.:: :tries 6.i3 S)m_etc mapscf America 6.N'.:a 6".:. a._est react': a'. whh only tv..9 PY:)!of.. imFor':an: population densities existed only in Me:_i:o and !n the northern and centre! Andes._hicma_s c.'_ased practically e'.lands haxe bee...n Noah 4me_ca 65.eD' (.:s::c. near the fcn:e:. Two-thirds of North America was once forested.McE_ed? and Jones 1978).. Development _r.:s_cr?: _:_?_'. The Arctic is mostly a cold desex.m of :he ger'e'.i&h. dry. in southern Canada..._ c.'-.:c L:stcr: of-k..acrid.erse._2 Cecg. '_.cxmuluted in four major 2. At r_a: time. White immigrants and their descendants became the absolute majorit'.allgrass Fr'air.. d. an estimate made very..::h i{ng.Ameff.f s:-Z. Two chains of mountains of re D' unequal altitude run along the eastern and western coasts of North America: the Appalachians in the east have been considerably t_attened by erosion.:he freezing point. 'andthe mestizo population may be almost 20%. fiat: :i'. but is not precise!) known.eD'where and is no.em 7a:_. The western United States is very dry except on :he coast.!Or..'.-m i6% and '.:_:.='h>ann :°"'. we confine our attention to the native populations living in the Americas before 1492. less :hen :hat of :he lar£es: continent. 1No continental United States has cold to mild winters.ir_.sicalan'. the} c.o months in _hich temperatures exceed .. a Mediterranean climate.ent considerable admixture in many areas.._.. about "-. As usual.10 Ph.. _ ice free and '. De_eic_mer': . Ph. and moderately humid and warm summers.mer:c2 4.::'cr_ment f 2 P_.-XPHY AND E.afi. The no_. formed the habitat of the bison (often calied buffa!o) for DOI 02179 .".ig:-.zs _. especially in its sou:horn Ivan. and hot summers with ample rainfall.s::'d _ee£r_Fh? 6.. less :ha:: a quar. Lin_guistics 6. oR :r.a Hudson Ba_ and !n the south and sourheasL :o:_ar:2 2re lowlands and :be Great Lakes reg. Summa. of .6. North America. and begm by d_scribing the environment.-ea is s.L Noah and So:aft:.erage. imprecise by the extensive hybridization that took place.r.'.aticn of the Americas is onl'.CenL_: a. depending on latitude.on:::: shield ix the midd'.e a. But today's :otai pc_u'.meric..'% toe'.'about 5% of the tote: population (much less in North America): it also undepo. . African slaves were imported for work on the plantations starting in 1650 and grew in numbers in most cases. CJEDGR. except in central an:as. Asia.>.11 Corn?at:son of b'. The native population dec.:'. ih.-iacial ad.2q cf t_e Ea.::.e!..:cur:.: an. :he climate is temperate and cooI with frosty winters. para'.IERICA 5 2 Cvegra. which can be very._ romaine.marker b} moraines ac..':'s surface.]RCN?qENT "Ia'_e.a is 1400 fee: (.!on. Central America has little vanetion in temperature with the seasons and has a told elimate with abundant precipitation. muck 'ower.._:scf :sO'.':e::. respec:he::.kic_ the Mississippi is the most impo_ant Because :he continent spans :atitudes from 65: to a few degrees above the equator. :he type of trees depending on temperature and humid :zeta" maximum _._ Oeee..hropciog} 6 _. 1.' of :_'. ext. Development fn Seu'h Amer(. :gq of the Ea:':h's surface. Below the Arctic..t::udes cf 6:9= := The'._hi:h is 60%_ .:uc. whereas the Cordilleras !r :h_ .c._t -he :i=e of fis¢o.cmFr:_-e.1aT m) h£h. di'. Descendants of slaves now represent 15%-20% of the American population globally.

. co..ensi_e in '. The extreme south.. eve_ year.. in parts of Co:ira! Ame=ca: :b.. is excelle=: for agriculture U. forest: and still more to the south lies the basin of another great aver. just nonhwes_ of the Amazon in the Brazilian plateau. To some extent._. the tempe. the Amazon basin. about 10% of the basi_. and a frigi_ climate. South Amemca is a mirror image of Noah America. so tha t it is na. Where ''ast:-ates the climate a..ers are high'.o_he.atu_s are higher and the seasons change.f iess t3 agriculture. depending on altitude and local conditions. DOI 02180 PREHISTORY: OCCUPATION OF AMERICA There is essential agreement on the idea that the peopling of Americas took place with the passage of nomadic Siberian hunters from Northeast Asia to Alaska (Fagan 1987).e.d:_ions (. at a latitude of 56 _. area. flattened highlands occur in the east {n nonhe. Here _oo _e weste. A retative_y small fraction.-. having very abundant precipitation and {itt!e change in rain or temperature throughout the year.'se k .v s:bmerge_.e_er.e r. b_: i_ r._'e_ _-v.. _ The -'_-.".ni."n mountains border the Pacific and reach as:ounding heig_'.. from Africa to America or from America to Polynesia--but the'.:Trc. and are wider in t>e middle Old. The Andes '.8 Amerces. as is often the case. _o_."ra firme") lends itse'. Farther south is dr).':.he sout.7_ -_ 3o3 many .':i.g_ca'.hersouth lies the Patagonia desex. the Parana. there i$ considerable uncertainty regarding the origins of native Americans and.c: cor_tinuo_st.mi?. . and its beginnings are more obscure despite enormous interest among scientists who have contributed to the research. the_ go from the extreme norxh to the extreme somi".- The prehisror)of Ame_ca is shorter than that of any other cominem.*hereas trcpizal forest is ex'. Thus.'a_' in climate and flora.-_". _. wide to'. ..he precipitation is not so hea. Between these highlands is a very.z. . but the rest ("te. gene'a:mg tropical savannas common to the Orinoco basin._. One problem._ _oodeg ?eari? when the ri'.a_nasare founC ai_os" o. from tropical forest to grasses and plan_s of small and medium height ("paramos"). is mos:t? dese. of vegetation i'. ho'._Ea.ura_iy fen_ize:.._.y.. uncenaint? generates discussion to the poin_ of passion.. The Amazon basin occupies all the northeastern part of the continent and is covered by tropical rain forest. Major grassland areas are the Pampas of northern Argentina: far'_.arzea"/ _:a'. has glaciers and mountains. Other hypotheses have posited extraordinary journeys--for':-'a! sa'.':.'-n Guiana and in southern Brazil. are not supposed by hard evidence (Bellwood 1979). 6. to steppe ("puna") that reach the snow line.fig 6 I '.' 3o....

"-..a'. hen the mig_t_on from Siberia to America is today almost States.e be_n made b'.ored /he passage of different groups in differ_m penors..n ice-free corts.e'. and it may ha'..ery different from that of the S:ber:zn reglens of origin._-f birch.1!.t-. Nc:h beiie'. the passage _ocld ha'. nab :r. in dispute _'r_: cer'tainl_ South Amer<a Moreo'.dor is be'. there were mammoth.a. accepte '4] are in there are on'.nee:rag Asia and America..".e se.ere andoabte_l'."n Asia v.s exposed the continental shelf along the cca!:.:s ". glaciers occupied hi! of Canada and =a. hc be!ie'. Schv. frigid in the comdor.ironmen: of America and ner' too at:racti'. E'. but direct archaeological e'.. Condkions that pe. It is not cemFie:_i ? clear _ha: the conditions for !ife were on Be=ng:_: t was prc'cably a largely treeless land v. AI! '.']2_._mefican s]tesi wh:ch are fe'a and difficult to date.ished Siberian sites are mere recent than the oldest The ins: glaciation occurred 20-12 k.:de.ecima:e in Berngia '-. antebpe.'e'.":. ith grasses.e taken place was differenl from tn !ate glacial times (fig..pes preceding ana:cmicail? modem humans '... a'.idenc_e cf ?assage b'.a.e of Neander. :'.e existed between 25 and 15 k. Th.:t cot. ha.:':America u.American . m this case. due to water being retained in the pc]:.e ex:s:e/ he- sims that ma': ha. the =:a_: hne w. and there are no finds supper:rag :he migration to America :f h:ma_n :.._ily accepted and indicate the maaor controversies.!er animals (Fagan 1987. There :s su_':stantiai agreement on :he :ack of evidence of archaic Ho._ugh perhaps no: .e been inhabited b'.e fa.. water is difficult to find and.:ho.m h.). some by land and some along the coast . a mosaic of steppe and cur:din It '.'. and eas:e... We brier'. Briefly stated.s bu._tes are not accerted _? scme archaeologists."t of the no=h-central United Tem_e:-ate and :retinal climates _ere found in Amer:ca ar much io_er la::tudes than at present. whom o<hors accuse of ma:ntaming _n_asonab!v high standards (Bray' extens. een the eastern edge of the Rockies and :he .:o sapze_:s cr earlier t?pes in America.iew here some of the major finds that are gene.x.e. replaced the st. e'_nat smai'.a/a....c_h America U. and en'. bison.idence a first date of entry of 15 kya.e geograFh '. Be. _th a peak at 18 k>: tr.el to ha'. pione"rs _ho later occu?ie d ..bestab'.... 6. and is believed :c ha."-..e that _e earliest onc"'y :nee No. horse. as 30-55 kya (there have even _en claims cf e_!ier sites).e:_.eger ]990 Ce_a:ni'. the land bridge fa:ored passage between the cenfi<er.a.2..t it. re'.er. iS that amoT'.ie'. tv. a= _ sr:'. and sh_bs. Perhaps mere :mponantl?'.'-n ?:z_s of Car. Asia :e America by land existed for some time and ma.rooe and !n No_heas: Asia.-&eless..en not em:rei.ed as an !ncem:.x.e DOI 02181 . there is strong disagreement between arckaeoiogists v.mmense ghciers occupying the cent_'.'-mitted crossing from.r ice. beat. anne?ted American site gates fe?Iew the disaF_earanc. causing the :emporzr? disappearance of the Benng s:rat A wide and fiat ]and probably ne'.el :o ha'. fair. and :_ose who are prepared to accept."!ngm. C-:e oldest . has not been discovered.e had :o ha'. but the environmental cond_tbns '_. at the presumed time of the ccld and d:'T _ith strong _inter winds Ne'.. a few Siberian espec!aE> in Sc_t_ Ame:qca .g the ollest Sites _hose that are less "_ne :topical forest had a SORA. cn the basis of present e'.

'.c.X._a '.A"!ERZ a.3 Pc!do-Indiansites in ... • .in_ around '. 305 _'_a v so....o to the "microblaces" made ir.. :...::he....e.--.. Lber(a and Aia_. a culture was found that was dated at i'... up to the Arcu. _ "LC-'> . perhaFs bo'.a. .rther J9S?) Some of theirtccs_.. v__. ..ere not glaciated at the time) and Mexico {la ':o 6'''_ 62./ o_-.. O_aC::* Ftg.X.'m.5-: g. k _.ns. "- ( .. No.a._ .e. lez!nch c: :r. o dest dates.:s._ko'.'. .. some intermediac> with Alaskan sites (Dikov1988.a_..:_aee .si."e a mammcth bur?..._ R_ %....3) Too'. 000 lim"!tiesofitso'an 3 A third site is Ushki Lake in Kamchatka where the e='.--'--.-.._:ara_ L " t _" -AKmaK M N _'--I/ " "w----.-:c._ ._ /_-__-j L.c Mar. direction Severa: ' Siberian sites cou...ier and _.'-k: _a. this cuJtu_ had already spread e'. .. 6.n Ii'.a> Koin the later pin of this period..ov. s..1M'[a and " .e '..he b :ecam common e in Japan and er ...-K 3e-a Ca"':'ez A"a"_.mer_ca (Fagan 1987. \ / _ . . Paieo[iu':icS .q _ ":": ::"_' F _ .= .o. d ar...3 P.._ .ere used for inset tools and appeared in no_he..a C D E._s and a.:-12 k%ai By 14.n'..._.'asfound at 71° iatit'_de in BerelekA The discoverer be[ie..2." _:_¢isCaves ._.ces the origin of these peo_!e to no.a .. in southern Siber:a ' in North America ir.':.-. 1 About 20k'.-...2.. 'hi I _.irtothemammcthh-nterso:::hewes:- em Russian steppes nomh of the Black Sea amc:g "_-o latter. finds at Old Cro_ _ats in the nor'them D_k_:a Berele.":hezs: Asia at that time _ _.he Aldar.: Ocean '_he.China 30-- ..'J_t Abuda!ofahuskvda..1' _ ( _*".. :he best k:'. :he_ ]bed mammotr and reindeerhun:e:'ss_m.: "-..e Dme_e18-!4..""h *_ ".9S_ Fig..:_( J affluent of the Lena...-.....:ral Alaska (fig..ed at ._ B .. 'c.a Fag::..nh. A..4 ha'..a K .] I rea 15 k: 'al . DOI02182 . _-. Mal':a =_ A/ been homes of the ...n.. At the cave situated nea= D?u_'_:a: a:so sFeZed Diuk:ai) near ..ith of a thdomesticate e use of do_ dog sleds.a. . _'-_. _ " / tures used stone-tipped spea. Microbiades ".a. kya._..see fi....2) seem to belong to the late Ush.'ta people also made biracial tools (Fagan !987).ki culture and s. The Diuktai people used microblades but. RJ'.o'._ _ 2.Jdes sites ir ce. Guidon 1987)._ _ "k N..ounI4. 6.._ @ __ a..o_{cat si:ts m its beginning to be e_:.en fa.:.hern China.hov.3.":. The earliest archaeological scena.2) and o/hors in the continental United States (in the pa_s :hat __ S...kva(Fagar . unlike Ma'.'m.'edtotlkvais the oldest have been n connected or_he__nd v.c1'... ma> sites farther north on the Chuckchi peninsu:a I'see fig 6."_. 6"" .

giant sloths. but wide disagreement obviously exists among specialists.. there was a major explosign of _chaeoicgi:_-'..a:• 2'. several finds older than C. Dates of i4. in the cen::'-al United 1_ kva. fd!o'*ed by a phase 30-i5 k':a with bone tools and a unifacial industry. but the date cf2T kycla[med for them is disputed because it comes from animal bones.".-n. and there is n. hcwe_er. and others (Grayson 1997). There is no problem with the essentials of the: C'.a=k cf ¢'.icence on .oe:-ce for e=r_y. 175 k. The ides '. (see 1ocauon of sites in fig.'mans. Other archaee'.san=.a (up to I6. Ar..?S . There are.his ex:reme?y rapi_ displacement are of t'*o kinds: the hun:ors hal [:ttie time to adjust to new en._ere :stoc sho_ at-. Accord{rig :o MacNeish kig'.' projectile points namee after the C. Mexico has da_es grea_er :hun 30 kya associated . including the mastodon (another the sixteenth century'. A ce:ser.9 k:. `shoo:: is doled to the period ll 5-i: _.-106 CHAPTER 6 i_=a. :he `sr..z azc . M_y sites in Alaska have been dated to 12-10 kya: they contain bifaces and/or microbiades _miniscen: of :he Siberian cultures (Denali complex. were "modified" by h' for fi".. and retail'. Whatever trace of African genes are :'c. outh America.nin 197_) Tne 7:obiems that arise from accepting the h.d ad'..:nc2 among living people. Idaho-13-l.lacesthat are pre-C'.es in Peru have a more reliable later occupation at la kya and at. hov. T=e '. The people there were mammoth hunters living 12-14._ _er': late.chicare.a.a. _':.ears in Centra'.':'en: is not .pears headed with projectiles that had chanactenstically P.3): Fen Rock Ca'..ith chopping-chopper :eels.. Florida--I2 k'.. 1982): Lhttle Salt Spr'=ng.. Shortly thereafter.25 k.. The :cea that South America was occupied _fore :he non'h. which de. Akmak).c'.: h.. sites in Nor'.'$1. it is muck mere likeiy :o ha'. Martin (L95'3) sugges'eo that it DOI 02183 Yukon (Canada) near the A:as_._e Diuktai caves. One large mammal that survived and was still flourishing on the Great Plains untie a few hundred years ago is the bison. Blue Fish Ca'. Dry Creek.iew more sa::sfac:er'. these animals disappeared from all of America along ".alli-Sfor'za 19S. e:.ed from admixture `s'ith African sla. cite four major examples of early dates for South American sites..5 k'..acroft Roekshelten Pennsylvania-more :ha.ears In!fied by the availability of the same prey (mammo.'.veen 12 and 10 kya. eptable (Ca'.... A human a.a.-..ironments must ha_e been s:rr.'.her site dose to Old Cro_.."1 the .' (see criticism in Fagan 1987).ogis:s also be_ie'.:c._:'::a :s :ieari._pk _e'.eloped around 11.:f miles in a period of !000 '. Monteverde (south-c_:ntral Chile) is an open seulemeat with excellent conservation. Here ."ace tov.t[cn is :mpeffect. Adovasio etal. several camels. is more _f_c.o thousand '.. and the'.men.. the oldest of _hich was dated 'to 32 kya.a. The mammoths were kil:ed with !._ifact made of bone that had an older date has been regaled to 13.. and probably' others) throughout :he con.'ve:s ceJd wei] have co'.40 kya.. Wilson Butte Cave.a.!oriels of guaerie consequences of .a.ith se'.cie . e$ :co much 1. one of the mo:i' layer at E'. in addltion tc humor ar:ifacts... . mr.::=lion :o ne`s en'..:s. the horse.e.-: _ sties an:odor to 14 k_a or J5 k?a '..ovis culture.uted stone points and were given additional thrust 1::.'. Oregon-13.e da_es earlier than I5 kya for Mexico and South America (see aiso Lynch 1990). ekher from the Pacific or :he Atlantic Ocean. one of the imponant sites. The disappea_nce of the big mammals has re.5. Approx:ma:e ca:cuia::cns ir'. Fish has oroken bones do:ca & of i_picmen:s of time between occupation of Akska and that of South Ann:ca is not a major cbstac'. It is difficult for nonarehaeotogists to form a final opinion at this stage.hat ..using spear-throwers (known as atlafl). it was s_pt:mned by a scarce and scattered population. in summary'. must ha'.. or 'zum=:_ o.a: Meado. The Pikimachay Ca'.anacc..o'qs and in his '. =n..o consensus that the.'ears. Its orig'n is uncertain: its end coincided with the disappearance of mammoths from the plains. that the h?pcthes:s of . It is understandable that there is little tendency to rely on radiocarbon dates especially if the'......00'0 years no.200:1150 at Alice B_r Site in southcentral Brazil are more reliable than those of earlier too}s from a lower layer at the same site. has bones '*ith dates of '.e analysis by Fagan (i987) lists a re'.o:'=aC. there is no e'. " .:eived different interpretations.*hich agreement has been reached that Alaska was occupied b. Other often-cited objections are that the strut- .e repreduc::d at a high rate in order not to dilute .curhey from the extreme noah :o the extreme scuff.e..e.oral other large mammals that became extinct bev.. in the last t'.:er. including mi¢::oblades sin!Par _o Lhose of'.. or come from samples that could have been contaminated with older matena[.es af'. eider one at 20 kya considered less reliable. Pedro Furada in the nonheastem Brazilian plateau (Guidon 198"7) has yielded various layers with signs of human _cupation. safisfac:cr'.. It is marked by mammoth and bison _utchering places.e. roans 5efc.e origina.of ad=.. e_ so quickly from noah to south across suck a `sine and diverse comment.' hu. have high standard error. the saber-toothed cat.y human. smce r.a.'.sere occupied on'.5 kya or: the Great Plains of North America and :asted for about 500 years. The Pacific islands closes: to South America are quite far a`say and .an border are undcubted'.e_sis'a:'ze :c =a. at:.a_-d :he south. A !ov... finds marked b'.":ae are discussed in the last section of the chap:or The ?re_:e.pothesis of. where bones of other animals are also cocosiGnally found..12 k'.5 k?a and 12 kya.".his rap. _.:r.:t to accept.' are unique.ered disco=cos of man'.o:zs site. but signs of human occupation are not as c!ear as for the . kya.a..o.h mastodc::. This culture takes its name from Clovis.2.ironmems _f :_e'. dated to 20-.-s..m:gr:t ha_e taken about that long (Ma.ater k ve.

what !oneer It is possi_b that Na-Dene and Esk:mc. off the coast of British Columbia._tn " us someo.... couic sur.. _" o_-tv.p ' ^ a. hides.real hunters ca. The Dorset culture (Jennings 1983) ranged from the Nocthwestem Territory in Canada to the Hurlson Bay.ith tF._at the occupation of the Aleutian Islands ke_a= . a=e perhaFs easter to f:t in:c the ."e_!acec _c !n po_tgiaciat times. ""_-_...'.o_:cal " . k seems .. that has a _echnoic. The difference in origin of Na-Dene and Eskimo-Aleut remains to be clarified.urcnments .e _..nn_¢ "' Sea. _l':.... Eskimos de'.es t. and led to the peop.'_ .'. _ . Their cultures at the time of European contact alia'... ..hem AiasK. pe.erk.. e_ ..'% __'enera:e4a dramatic chan_oe._e: ' .w. pr<ecule points '.'_'... hunters t::a: oc:upied the "'_o_e' _" . The earliest occupant." '_: _.No-Done of the entire continent. Dikov (1988) has suggestec that the late Us>k! cuitune...M..ed tc ha. The first kno_n" .... ._'_ '-'.ears ago.e.:'n. o..' -. reaching high densities especiaIb' at the mouths of ri'.. "'" of occupation of Anan_'ula_has a]so been suggested (Lm.'.n.. ' .man origins {nAs:a._.e Pia:ns :he bison r.t the _"_2_ of." reser".. er_Te::_om in _o_' '.: .. ch:.e Aleutians {dated at R r kFal. different from other Asian :u:'.-L._.o:at:on in :he hum{aS tech!'_ q'." to the B.'."2: be _on-side_d._*^_.. si3 frc.e !'. with the Aleuts occupying the Aleutian islands and the Eskimos occupying Alaska and the non.. ten:u. After the disappezr"ance of :he mammoths.e'... _ m. The'. -" ' cause. _ I 30" of . and Greenland by 1000 Be.-.h.'auons. .._ain:'. "<. The_ may have been a series of migrational 'waves. They settled in southern Alaska and on the northwestern coast of North America. presumably a second one {15-10 k'ya) is named after the Na-Dene family of languages spoken by these people._ca and was ccmfinL...._. 10 kya).e :centered .On!y the in:rodu:tioa of tr. etc . e'.... be submerged.. dated _o 3000 years afro. but the exact time sequence is not clear...ou_.000 '.:'g:h America b'.e No-Done and Esk!mo-q:e.. a stool! island in the ._ The b_sc._.ard from :he. hat. Queen Charlotte Island.. beer: ad'. on the average. _as due :o o'. o-._' .. the eastern :oust of the Kamchatka _eninsula sho'... s:mphsuc....5.e.. era. o "_a....= " .:i:ma:: ':r :he _ostzlacia: _n.ereboth later and..e because .Ma culture is the aides: .a:'.....e" .o_n _..zn The oriz:nai _ Asia. Flint station in the Brooks Range of nor."._. b'c: Sberian EsMmcs are beiie. bison-huntizg more than one re{gnat!on.. ...'as tha_ of the Paiec--Indians.Aleuts both come from Anang'a:a ancl ..s currentl'._as some slo'.pursuit ofchis .. some Na-Dene groups migrated farther south.. i: is ... [ocat!ons o:" .=.m the A._" p_..'_ b_'_its di_esti'... of the ". between 35 and !5 kya.. _ .as not _-'....:e.'_'._c.... The populations of the noahwest coast developed a special way of life...first da!e of earn.he cajcr sconce cf fooc :or --.i3 The o!dest migr:ticn from Siberia '... .'. .-'" -" ." certain:.tF e.._"..e. The third migration was that of the Eskimo-Aleut (ca.-e.estem and eastern ends of the chain of islands '.a ..' tunes of Alaska and British Columbia an! may ha.. Another m_gration._-_a: abo.-t this time L. or there may even have been a continuous flow.a. n_*the world's most DOI O2184 Pleistscene c....s. who kept to their Arctic and sub-Arctic habitats.: 1" :h... a .9---6. _' ..n_ed _ somev. have been occupied earlier.'-ai"--s but also the shot': grass :_at . Diko..m.k_ anchor: ". spreading later as far as not equally.. 987) that '._.heresalmon was easy to catch.ers '.". of me time of oR_'in . but the area ca'.s:_. :_e '.ho._e contributed to :he Eskimo or the No-Done populations or both._ ..... not simply one.e_.erki!: :. from it.:{ :_ not ..'.." .easter'n C.s _. to which the above discussmn refers.'he L'S_R. Newfoundland.. Much more recen:lL at the beginning of the present millennium..'. us to place them ann.._ stmne_ . ' .euuans ' ...stecl that Eskimos and ._--men..-.hem coast of North America. sa_ .. but the most in:crest{aft earl.near Umn±< Island. for v. came :he moor source of food an_ other :Dmmodities (bones. 'o }' ciear b.ere ale'.:_....'r t_ :._. proceeding ClOthwestward and east. a mode cf adaptation to elimarie chan_e....e cu!ture of Ananguia M=. The other two migrations '...d:ans..n S..a..ghlin 1980) as :he date of separation of Aleuts and Eskimos. sites of :hose _shermen and sea-mart..."-q'._...A!ects hac" corn. Laugh!in {19.. to eat only lhe :a'_ :'"_ss Cf . = Gm_sDn.SC_sugg-. .orse and the gun aher the Spanis:_ :onquest in the . "° '.at star:eft in h... hov."... perhaps only a little later than the Paleo-lndians.d Re'.act."'-..'eIopec transpo_ation skills across the Arctic and hunted not only sea but also land mammals (musk ox and caribou) :..<town in ff'.'-...s:... by causing" ?rofounC e.. that of Paleo-lr'.v a date before _.--_ .... but there are signs of earlier occupancy of these regions by a pro-Dorset cuhum...e b'-. of the G. There are still a few Eskimos in the extreme northeast . more acceptab]e en.mat ing ca.i::ar on:in:LiOns that n. and 6. \Vea_ons ._ "_ . cat-:re '....a pat.9:150 yems age and also has s. . 2_-':'.. The m_u:shc and biologica! e_idence is discussed in sections 68...a.. dated 10-. was continuously inhab{ted between 7000 and 5000 years ago..2 kya and located or'.?a) is like!? to be . Evidence that it affe:ted the fauna c_mes from the ohser'._'.exp[anation for rz_'..mfiamies '. to large actual: The bison.a :r..or': close to extin:uon and '.... aJso discovered a culture on the sou:. While A!euts remained on the is(ands that car" O their name and mostly maintained their primary skills in hunting sea mammaB.::'o_ .er.... The coast of the Pacific Northwest was colonized by. m ca:so prox_m_t' " .o _--_.'.at_on Ihal [argo extinctions of birds also oc:ur'red at the same time..:: :at ..l" at the begmn!ag of :n:s centur'.a ]t is dated to £0.eloped from :he Clovis points T3ere .. su-...ures and s_miiar to .h...e --'_ .:tear t_at there v.. .. '.er .._'_ ". Tqis hvpot_esis. Fagan (1987) indicated mare conservadve:..'< E E ' S=.. an.a.._.A. Labrador.ced as . peninsula at Puturak Pass..ereas stool! mammals survived and changed the:r range. _o 6.c.. Moving to other. ied to the occupation of more limited and well-defined areas in the the ....'.. _ archaeo.

off'.e she?.. coming_from Nc_a'.=-_.:'r:ers occupied the cent:non: '_ith extr:.. a-d :robab. ace:a-:.assomedegree of adm!x:. "...v represented outside the ori_. 6..ear 9000 s P..a IS .<"y from 35 :o 15 kya.. It is based on '.--. at the time of European contact..'chaeoiog'. ..a_. _ period.e.. _ has >_o._:ed for mii]enma in some ar... of the original crops had the petentia! of being easi!:...lutes ...s..srlac . before the social and political conditions of these popu!at_or.'.. n 1990 ...cuiture to neighboring regions was slower and more limited...'_"'_ in Ch::za By the ..".ie. Other useful references are Kirk and Sza:h:--:_:-.. ..a.ide area around Ihat of origin before :e:cg more f_lly developed.e :ransltion :o food mroducticn from the foraginn_ ec-_ncm. lurer than those in the. e'{ported to a _...ncdisapz'eaed in the fifteenth cenzury..eloped. near • e 'a. chapter:.he D-_o< sou. culti'.a: . as `*eT. a 2.. but in both regions relatively high population densities had been reached at the time of contact and. poses the interpretation of Lingais. In the periods preceding ag:'icuhure or in its early development. however... ¢. posed hv Greenber{... u-ibuted to Na-Dene-speakitg pet.'ithEurope a.-ff ::::-_.a.h!en 1987. :. chapter.e..en c± . unlike the Middle Eastern dcmesticates of '... The three-mig. fol!o_ed by' the rapid occupation of the '.>.a.. arid plant domesticates that cou'.e_ez " :: -:_ been ado:ted as a par:iai source of food. . Vikin:_.. or breeding very different plants and animals at different altitudes..CR:CLLTURF_. social complexity of these hunter-gatherers were greater than in other parts of North America that offered only DO102185 . each suitable for very different types of economic activities.. the Na-Dene.. similar but not entirely comparable to "transhumance" in the Old World.v. on thenature of the domesticates themselves.oped local h.arces and `*here a__nP..f th. poss:bLe .i :or'aoer< and the'. r.aung.: e The :n.....changes and poeple ...'e E'_" sho...'heat. ..:ur'ai anttropoiog>. r'apic2h>.r nG..ailable en.dOIndian :.. plants had been cultivated for almost 10 millennia in areas like Mexico and the western pan of South America.ironmm:S could be turned into a source of `*eahh TT.'_ . with mere or less continuous cultural .anous regions. agriculture in America began in areas hke con:re'..'_ _'o t. There is agreement that this first migrauon came from Siberia via Be..Perhaps in the. Mexico and the western pan of South America (mesfi.:ntmz tradi'icns L_.'hole continent b>' "'PaieoItalians.. '*ere :he sub{ec: of class{eel r'ese_":h :n c_. later a... in ..: e:_":. ".308 CHAPTER 6 of North . ..--'.. ..s .:smg bc:h amrza..ide!. !2:.enta'.:em and cez::a'.NGS OF ?-..-aticns "_'._.'o<. barley..avs a relatively stow process... This was true in particular of the Nor'thwes_ coast North American Indians.• see a:sc . m :he various ._.s _ _"`*#_' _ _. "_.. sheep.e pos:-Pa]eo-Indlan pence is of'. Greenberg (see R:... Ecuador and Peru)..innin._..s of agriculture :n A. see in 9".. there is !itt]e age"'_ and :_...Srst occupation cf :he Americas.... and exchanging these produc_s by a complex network of trade and communications• Systems of seasonal migrations also ale'.. a stimulus to technologicai ad'. where important empires with large populations had developed.8:. Therefore.. there later ce'.. _. . large number's of American natives were still huntergatherers....-.... especially in the Americas.iva'ei'... resc.* :fa::.. had c:-i. The density and..ic data .og': ini:ial _e'.s. howe'. often very." mar. "_-. on the nonh`*estem coast 6._....a_i airead._ :ease of Greenland _as so:fled b'..' res. de'.. Compared with Europe and East Asia.htl.hunu .:es took place in anenxironmentand`*:thcom. a_so Russ icj_. a_. a':c __ene:ic irfcrmaticn• as _. by Esk:mes. and R ..'. (1985). it occurred at very different times and in differen: '*a_s in :he .e be_..-:.>: : . I-":he ....e.ere s_ch that the extraordinary' variety of a. In summa. . In time.a..__ and..e...iimate not found in much drier northern Mexico nor in tee tropical forest of the southern pan of Central America.' and Iceland in _he ninth or :enth contu_. according to some..rc..a_:." The next settlement.d be e'q.. 991.n.5.'e: " ::me....ance in food production. '.. and ca::'.. In many other regions..-:. 10 kva. The development of domesticated plants and animals and their adoption as staple food was al'.. a complex economic s.4rcSoic Ported or lo'or hun:inz-and-fora-<n£.complex societies t'ad developed.:-. _%... BE."..:. diffusion of a_m.. The -_'e. population density' increased somewhat._.'q:.are with people of EzrBpean origm.there_...."e. The Mexican plateau enjoyed a temperate :..xn ._--:__t:c.-se.s Th... cr later._'_.a as . Aikens (1990).inai_area• Fe.uists {see.¢ .e:opments in Mexico and the ncr-:. . suc=essf_. especially among the _e_' unequa'. '_u' :_e Vikinz settlement !cst contact v.eex:er-_-" and Ruhien (!99"J..:'.. In additi0n. Middle Eastern agnc.. and 55 of the Cambr:dge Enc..{.e Per'cal. led zo :he occaga::on the .:z'e . gears. A:2...a o economy--is sometimes called the Forrr._to :e:::-t:) regions '. -To--on' _f human pcaula:ior:s '. namely by foraging.e:..... o'_< :hough i. A zroun of i_nP. which were to some extent unique: or isolated... for reasons that depend in pan on geography and in par:.. The Andes were another unique environment in "*hi:h extreme differences in altitude at a short distance provided a great variety of small niches...uUure..'.. this variety was cleverly used by what is called a "vemcat pattern" of exploitation.'_• . close together... It took time. \' .aten the .e!ogment Th.&. nC: v..-that ...... &re:re .::T.stem '.ordinar'.... _ ... and of Californians. .Xmer_ca.or..-?ed:a ::f ."legions of Amen:as• The Pc..ith a some`*hat similar eco'. et a! (1a'_"in__ia and ".

.t_e mt[ s:i'. .omes:_:a:_cn .:nee asma! diet. down to 6 million in . in 1977)._r._. ]ate" th:.'t... are a good comp:emer. .".-. At the time of European contact. Otne .) .hem.. agricul'. .. Cook and Borah (1971) suggested a population of almost 17 million in A. ]n the central Andes considerable use was made of domestic camelids (llamas. Bear:s . It later spread to other areas of tropical forest outside the continent.ears. however.foodcustoms.":ca and .ho -'_-..0='-')e_. domesticated ear!_. Io_. it had increased in the last m£11ennia o.. of . gold and _ _ Te%a:a't " for man> thousands of>. the Gulf States (115).'utileAme=cans de. Guinea pigs were domesticated in Colombia and Peru for meat probably in the last 4000 '.3.'.. Potatoes probabl'. Figure 6.. At t.. but later populotion growth .in_...imnments.: esser'.ere domesticated..-". in Mextoo. Later estimates are higher. theselecti breeders. F " j S.v choosin£ the best cobs for reproduction... which became increasingly common in the fast 8000 >'ears for transportation and meat.. two major empires _ith large populations had de."zd[ca_f.. --.00. of the Paieo-Indians allo'.e.f. v.: to the maize diet because the... and Peru Elsewhere.a. the..'-sg.: _'mas _ .-e':"':. wh)ch v. .vo 1548 and 1 million in 1608.e01¢. population de:sit'. ". such as the !or..B.'ears old."'m.'.1 Dis:ribunor:of probable ._ I_ the use of dog meat for food may' be 6000 "..s a.. of uses . and tomatDes.r_'_ar :Ccrtazt 6... . Earl..eaher d!scove...'-::ica/" time a_out 4000 . _te_dthetoca:...t ..'iel2 C'tmaize '.. For central Mexico._. ..'.erae_po_ed :0 . of which "the largest components (in thousands of individuals) come from California (260)._'_'._._ _ '/ V[d.and Heiser i977'. Originally.ed them to occupy the whole continent rapid[.'eC: io. the American Indian triad of stupid fooda famous for being nutritionail) '. with perhaps 6-25 million people (McEvedy and Jones 1978). Some native copper was used for weapons and ornaments. ably because of a_!fic:al on ex ercise: conscious[.. but initially'-component of _e tenth or less the with remarkable .I£ R "Z _.acesof -._n/enfi"._ d:--es:. in maize.... '.. America at le_t may 9500 ha... althouo_=. the }. h r/ ._>'_ _. ' ... have been sys.ce then...en t tOsupport a sedentarypoD ulation: ?or:era. pros'urn..:t:va::[ Nc a'. bleso-America was the most densely populated.m -it. alpacas).e but been p.-ea'.'ha: time.'k_.. Maize was domesticated from local plants in Mexico at Tehuacan and Tamaui[pas around 9500 years ago. suppi.._" .... --M'.940._jJ_.:_ ¢'_ "_ an "_-.. _th the firs: e'_. Canada (190).._o_.uro_.elo:ed _ _'re2' aural'dr of domesticated pia:ts for a _aRe::. at a "'c....nJoc expo._ because of its msefuiness as a water comtainer...000 or more in Canada (Charbonneau 1984)... much. the only widely used metals. (Lauh. Thefirst 'domesficate ' dp.. _\ 309 h i '7 *i (. Few ammals v._.-2 %_'X.bssibiy the bo:fie _1. came from Coombia (10 kya'.. mmze cobs we_ onesize of ..:'..ematicail.'. e._ . . F]g..*e. made :ts first appea.ell balanced._-.._.ra_. and the Plains (100).:÷ss!ea. The high mobilitl. at the time of contact. po:a:oes.ual amino ac.\ '--m( ..._e time of the Spanish conques: of grown for use as a textile..". l shows the sites of earliest domestication in America (Bray 1980). estimates by Kroeber (1939) and Mooney (1928)(whose estimates differ little from Kroebet's) give a total of 1.<el tO tropical Af.'ere . Even so. like maize.2 million for all of North America. soon added to maize and beans fo. had almost entirely orn.n in Europe and Asia.. // ".modem co_s Cob size gre_ _ '=""" '.s . :t-aze _..< years C'pla':s at.':.O 1532. '. '<--. ". up to 5 million for the United States (Russel': 1957) and 300."a_."v in the New Worhf and acquired prima O :mpcnance as stap:e toco m :.'asslow until the iast two or three mib lennia and increased almost exclusively in areas where previous important agncukural development had occurved.C... but Zambardino (1980) corr_cmd the 1548 estimate to DOI 02186 . '. Squash w_."s '_ ago.. and oleos: ceaainl'.': .mentai applications. 't.ant pa_ of the food supply.. American natives were still in the ston : age. :c_.. Most of these crops could not grow in tropical en'._ni) assumed that agncuZure became a major source of fool suppl'.America..-:r " T.earsago At .' over... The turkey is first found Jn Mexico from 300 B.3. /" < } . The number of American aboriginals at the time of contact is very imprecisely known and varies greatly with the suf.a densities characteristic of the initial period. whe_ instead manioc _'as firs: domesticated... z plants like m_... Cotton '..: or subc:nsc{cus'.a_ augmented by' the products of huntirg and cathenng It _s morn or ]e--s arb_<.P.s 9vO0-i0._'-_ °v. W.c.v. inde.eloped in Mexico _C : : c:E.'..t"-.'.:'-_: me¢ .\_ ... lands of South .

. DEVELOPMENT IN NORTH AMERICA becoming warmer and drier.: .ere _se.ehas o.idence shc.d cro_s had become "_. a ".. leadin_ to hi£h oen.ksa direct consequence of _he econom:c ..? -.e ha..= it is act sun'. '. the de'.c::.. foe: 27...:.....a.neraZv. and northeastern Oregon).izafions.. `*dr. T}_. especialIy in California. In am._:r_.enca. • was falling or for other reasons) and in man.i/on in ! <-_. :_e more immediate descendants of the PaleoIndian hunters had to cope with an environment that was DOI 02187 .. a:ter :he F:_'.'a:!a_. s:u& of the spread to the south.mcu-t..Mex!ce :o the south. Without increasing sol:'hist. the Great Basin (Nevada and Utat.-..p. By eo..e onset of a_ zncu:ture a=c its successive d.:". di_ in Eu-cre or Africa.cThese :and:: or-.. Aboriginal population densities largel? retie:ted the degree of '%_e!opment of agriculture and social ergonizafien..cul:ure.clez: whether rapid inc. The .'tem of migration in man.._. must ha'._ raTd!v after the cc. loca'. and _here stror. _ of ec:_ ._po.'. have been ret'ogr-ade fio.. where :he " _' ' soc:et:es of t_'nonagncu. 'co¢ are dlf_cu]: even _._".ation.:... en tentafi. the . . _hich had an eno.:!. migration tov.-'_% _-.'_ ne:. central ..:n l _p '. :_.'eases of pcTu'..a . . -' condill:hi: a: :ze ::me .. case . North America hac or.....laho. Europe ar. For a general overview of the pro._b:e impedance from :he point of _ie'.i '"e:'e '. Surveys of :he subject and references can be found in Jennings (1985) ar..F conouest t'ne'_were rare even in .. and Cuesocial structure staved fraementarv.:me of contact. :he __q_"_'_ . as elsewhere.-. 6._ ""*.ul._...e. nov.:::on usualb increased p ' _ sections..".." _')" .r.. may haxe origina'ed m the forest near ... Fopulation density remained lo'* in most areas.elopment o.rast..: '.bo. and much of it remained :tontine2 to _t._ "*est had re!at> e'. n. .eAndean . but there may.e.cus!reFace .-.._.-ks nO:de else\_:'._a.._e. there was some evidence of a beginning of local differentiation of cultures.anced sec!ocai'ural fast (La:hrop 1977"_ ..e e'._oou!atior:.... v. A substantial development of the foraging deve!opment.e dates and directions of expar. . reducmz incdvidual m_... and the East.e. ' : • at..e. homers '._as North a.. .e. Later censuses are d_erefore of little use.. .r'y... and the Plateau (l._ Fh_sica. ways. 4 7 _. .. dr.::n tropicai agricuhure.e nararal .. '. But even by 9000 years ago." ne:_her enough-.. aT. with the • the :onquerors ^_.a.. eastern Washington..-:or."ation by causin£ the pep£at:on re _come mere seder..vhe_ cu.....and post-agricuhural development.. dcv . the occupaticn ¢t _' -" _mained incomFle:e for a ion_ o time.hem Mexican dose:-...' '.ate and limited expans:cn ::a_..-.. cation.. fleas..nm..o . .n to 600 0CC. Population samration foiio`*:ng initial growth is expected to cause conIa'ifuga'.. it is convenient to distinguish four large areas: the West.'of papuletion oe-e"a< in :hat the :_nsiuon from food collection to .cz _c-7.Sanchez. accompanied by a trend toward population increase..-_ 'a.:ation in the South American plains v. Here..4-.. "1' fa'.'zmer_ court _'e=. ice caused major demic exports{:as as the.a::'.._.ard new unexploited fields.Andes. a....ments ar:d ad'...:?:rigthem out in the aoioties. the desm:ct_cn cf the preexisting civi'. be:ng . but this _iew is being corrected.. Lathrap ('.::r Central and South America TE.ation density in Atnor ..'.d in chapter -.ur..'m_. The _esr includes for our purposes California.ara_ .. occur-r.a<a.'.1. along r:xe:s.':icuaar e .c drift and.. and ne_er reached the western coast during the procontact penod..e oat:ined--mainlv..-..... Mexican a_r..--and its h__h.a .s that population numbers dechne.. h:gh densities because cf ex.}cera_ astir:areal 1..'fica b < much ir common with that in the nor.....ike ..a The dexelopment that took place in Meso-Ame.' " Jr" ..-..t _s net clear F .-ii0 CK=?TER 6 36 mii'Acn in ?era..v not known in enough de=a_] to a]'.at_c. late • .'_:s'. when these are a.. _.. u..9"7'. 2.'...arious regions before and after agriculture is briefly outlined.... ff... the central region (the Plains).s. a :he. decreased :he effect of random genetic or:::. ._ areas T_ `*ave of advance cf fa.hem Andes Da':es are probab!...1 spread of epicemics brought bv =.'ith it. In the follo'.v of these es.v...-" "_'_ Andes..'-o* the pa... ed -olat.._ e. ..nicat skies for . It was once believed that the social systern was extremely f.a.. _eas still remains a: this stage. or desexs did not impede m--_'..'.':..e acted as a buffer that slowed nor:h`*ar2 e\pansion_. the cJimate was already similar to the modem one. e .a-"_cultumi times because initic! agncui:'ure "*as of the shifting tFpe (moving to ne_ fields as soil ..av of comm=n:.'.should not be sunprising C_'< .5 :r. began onl? about 3000 years ago but 500 years later in the in:ericr (Aikens 1993).. high local variation. '..e'. semng in a slow wave of advance of the a!ricaltu_l ... they would not ha_e eventually reached relatively Agricuiru:e arrived late in North America from Mexleo.¢6of the Cambridge Encyclopedia of a dem_c component: the nor.. But migration was a.....:: agricultunl his:or?' and had not tea:nee: h:gh Censit!es at the "... -_ :has ge.n Ar:-..crable en'<rcr. .._:.prising if s=read m th_s n_: '.. Agneuhure probab]? sFread from .A'..tl_. central and eastern._ '--_ .' `*he-. interest nor born in he hioh_= .. manioc culti'..nod and therefore after "030 )ears _ P It is '-r._" :s of conside.a..).qamy '_a_ suitable for the pa. Seven thousand years ago. 1.. the Southwest.aQ .e Pe.t..=:-_"-.':igher '*here :he his:o O of agricultural de_e]cp'-er:: '.".merica.'c ..--._ and exganded late to the non::...omeor so.._s' .as older An exceFtior. Jrce -'<..

". B. originalS? rather diffuse.. include the Fmemont in Utah..t-: to the area like the-Apache and Navaio.. "fT. A-.el]documented by . Fig.D. New._ of dendroc_onolog.._. Anasazi. HoD: 2.. The descendants of the A:". and domestication of some nafi'. the Hohokam were migra_ts from norther-::.. Ztam. .Mexico: toothers._rev' _. 3. 2.._e. but it is now known that these Na-Dere speakers atoned in the a. at whic_ :i=-. Mogollon:E.a..:h '. F. Central plains: G.. _em developed cultures of foragers . a. The foraging peoples m .&":zona. large villages of Pueb!o-Lvpe appear at Chaco Can?on in northwestern New Mexico.m.P (Lipe 1983).? "d} the V. thereafter.¢C% "M . followed by the Dalton culture..."a_. were local mh . after a_ 1200.. and .. Hohokam: made up of small mobile bands with no capacity for food storage The transition to a sedentary. est nei__hbor.eo_..'_ local popu!a:ions. the Ancsaz: culture rna? have de'.::tam from the Hokoiaam to the P. of the major Noah American a_ricuhuraigroups(from Whitehouse and Whitehouse 1975.. beans.r::ng about 2000 years no". Piton: 6.: :n the Archaic Period.h.Maize. w_a...a. like maize. and southern Utah h is a vet") dr') and almost desert area. 5. '."aoo and Pima.. In the Paleo-lndian Period.. Navaho. Zuni. the most nonhero group in the Southwest. may be as old as 3000 years B P or more: a safer date is 2500 e..graphi¢ tc_atior. em . l'he case for cant:cub'. T"."--.'qesc-Amencan influence (Lipe I987_.. 1.._.ethe pcpu. 6. In Paleo-lndian and Archaic times the poputation was pr )bably scarce and diffuse.[y includes ._ . the Mogolion. Defined o ". the Clmis hunters were the dominant culture.a: lasted through the millennia (the Os.... There were cl..."e h:zhl.Ma. Conflicts _i:h :mmigr_...high tier-q:? and Iota'. 3. • ) / is . _'C_ C 00_'¢'. coodng for leading a_icu[tum these peb ople ecaus to e sem. but is adapted to a nov. Tine Honcka:'n.a_. Middle Mississippian: L.asa:_ are the modem Pueblo Indians (Hopi._'_ t.n thanks tc the s:ud.. _d sornetime:. moiety' of the Sou_'.?.ma-P'_-a_:" > _ascnaai'.'.4..ere earlier believed to have been responsibie for the movemen' of the pueblas. and that are not easily identified with modem descendants. at dates that are a:zumtel.. reached its maximum extension around AD... knov.he recentdiscoveryof and dryingof the local climate for areas around more ao suitable 1I00. e_en dunng the Pc:do-Indian Period in :he easterr.. strong• Nor:n of the Hohokam.. se"entar'. The beginning of a radicaiiy new culture (see fig." squ_h me v.. ha.-he: Archaic cu!rux _.vestem Pueblo cukum under the influence of the Anasazi.. 1) is seer. Mexleo. deer. e intraductian of some cultigens from Mexico..alture in the area._-_ s.and _. began to coilec: in sn'... Griffin 19801...ed that the increase in village size made _t possible to engage in ir'ngation .' ancest.. Unlike the Southwest. _""o_ '_ b.cles in v. / } L. AppalachianMississippian: N.. Onrota. The masons for movement are less c'. s:a. the'.4. For{ Ancient..':g the cut'rent explanations for the abandonment o. until the beginning of a secenta£:'-ho:':icuitural mode of lp. Caddoan Mississippian: J.quois._ iF._ "_ . Monongahela. it usua'. cuhu:-a! .e. sta_ed east of the Hohokam and at about the same time.e -e w.... L who v. Between .ailable. Papago. This area includes the valleys of two major rivers (the Mississippi and the Tennessee).hat lasted for millennia._ e _v"'"_ v-..._ .:-n_z'im" '*ere able to colonize a '. 1. with the Hohoka. which clearly derives from the Clovis._ "o .ch more water was a'.ancu':u_ from Mexico m'o'._ 900 a::. then abandoned coiiectivety when the population morea to other places. P.. Colorado.e plants... Southernplain_: H. M. Plaquemine:K. probabh with the contribution of migrants..exchan-_e and trade be:v... It was e_entual!y absorbed into the . :he same genera! region. DOI 02 188 .mater magnitude. bhddle Missouri. life was spread over a long period.. The. Amc.' in various ways.:". which favoredthe development of a rich flora and fauna.corks of _.'-a! to the P.2 '. the sequence :f nngs m trees. which is dry the East enjoys considerable rainfall.A._est. 6.C. Fremont. Apache.ail separate settlements.. :_. 600..'_-. bezns.. made :emmics.. who stilt ."_ . c_fC'. Another culture.. target... am .i 1100. \ _ _ c( / \ and cot:on. and extends farther in the Formative Period... Other groups that developed a farming c.. squash \. the Appalachian region.eiope_ d.nvnea settlements were built and suddenb abandoned shoal.¢r..ear.. Hurena: O. the Sou:h'.m c_kure in southern Arizona.. According to some...ed first to its near.'eqv.hich Ira-go villages (Pueblos) were formed. . lation.mctly from an ea.erpueblos It is area. etc: Lipe 1983)."o" .. an. 900.--__'_-_ • £'-7.:.1 Geo. open to form iar__..

oufih_es::rd :Re E:.'ere cor:..-_. is a good sur'. _suaY]y bu_ais. v..:_-'. Missouri Productlor: of maize and squash increased. there are 38 distinct fe."a.kcta .:mnment where techniques of imgation were necessaryi .ons ar:d :Re atchaee'. The slow emergence of an urban civilization reflects the long time necessary' to develop an efficient agriculture in a challending en. most of it :n the capital..ol:-:er .'-:.lib other mear_y' cu'..ere ¢.na: 'arer Pottery'. occupa<ons for long pe. The conditions for agriculture in the Mayan regions were quite similar to those of the Gulf coast where the first urban civilization.5.':sit.i:g{cal record probab!y for:he same reuse.' during this period. San Lorenzo.'ea.i:h few changes from I 1.. but they may have employed several different solutions to make slashand-bum farming more efficient (. but c:Cy later (.0 a:d had connections y. has already been described.a'. 57. e fores'. impoEam cultures ar. bu* couid net ha.:r:.rennings 1983)Cotemonial centers like "rikal in the Guatemalan low:ands and Kaminaljuyu in the Guatemalan highlands began developing in 30 and 500 sc. ti.: .h-'aestem Illinois and southe:-n '.e size of this mound (a diameter of '.'hat a degree cf social complexir: had been ._ee". thus favoring the spread of cultural diffusion and trade throughout all Meso-America.000 for the whole valley' of Mexico..nL where major ceremonial centers `*.ied almost s. as is usual in initial periods. that of the Olmecs. not on:`' as a food source: bison scapulae '.n 6000 years ago. south of Mexico City'.f."rniren:.... cemmoniaI centers and architecture. is a serious prob[era: it is not clear how the Mayas solved it. the pepulotion in the later period may' have been as high as 100.t. respectively The Mayan The earl'.a 10££.rne_ an importam. and agncu!:u:-al operauoes. At Mr:ram`' Cave..ere used as hoes. The lowlands of Hopeweikan. northwest of:be capital.:'ures '.5. :':is:Dr:caib and '.. The pEncipa_ site is Cahokia.loP. but not intensive. In Teotihuac_. `*i. But the Olmecs established an exchange system that greatly extended and unified smaller-scale systems existing before in their area and in other areas of Meso-America. Oneo:G.. astronomical observations.uhual contacts _ith the Pueblos and `*ith the Caddean Miss. Communities :'anged ]n size from I00 to I000. there are also fl:. and others) in an area of the Gulf coast.of occupa:. There 'a.. Louis.:. Nor':_ Amemcan agricuhural de.:- _-E:_ _..-. the archaeological comp!exes fol!c_ one a. the basis of '. Places discussed later are shown in figure 6..-e=ched teat made i: pcssib!e to build such monamentai works. 6.ere deri'..<:are ::re P.ed from prior appearar.en of cukigens of Mexican cr:gm An :nno'.':ile le. then Teotihuac_fn) and in the ..'_xaca (Monte Alb_. In :he Formative Period (2500 a C. m i_.000 :c 8000 years ag._mon. a o!Lmale change may' have set in.h dates ranging from ]700 :o $70 Bc (Jennings 1983). "a!'.. The first indication of water con::'el is in Tehuac_.. !ike sunf. with hunting and gathering.:.. and de. perhops seasonal.he beginning. us_:aHy dolled '. The first great civilization was the Olmec (1200-600 sc) which developed its greatest monuments (the colossal stone heads of La Venta.'.e]eymems 'acre re!ate.e:s from 9300 years ago :o. DEVELOPMENT iN CENTRAL AMERICA the deaiine of the Olmecs.. Ineicating m:e."..hich. Lou!siama.'-r:d nearby regions. This culture.aed mr!cations of soc:aJ stratified. and beans '*'ere added around _.". :he --.':s. had earlier developed.'00 _. agriculture with two crops of maize a year Soil fertility..arcu:d :n O(:B P became a _ide grassland occu.. snll combined.VD 300). Th. The popuIa:ior clearly became more sedentary.iiiages 'acre often fortified (Jennff:gs '. In summary'.ctuations in the de. Mexi.d societies de'. e.. Domesticated sq-ash is known from the a. pa..":.n most of the 'a. After DOI02189 .Visccr:s:n after _.n 800). spread north to no.'. are . These regions were exce'.e preceded :he mtrcduc:. who extended also to the highlands in Guatemala. rare in :.' development of agriculture at centers like Tehuac%. :.1. SmaUer mounds.. ma`' ha'.Jenhinds i953> Agricultural acfi_ity `*i:h dependence on maize in the eastern Plains (the Plains '.idespread by 7.eloped in the _alley of Mex:co (Cuicufi¢3 first.d Tamau! The bison remained impor:ant.D and amar:m.:haeology (chap.: after that rime.-_:H'e:i Gap..nce :me beginn ng by' large herbbores.2C'0 m :ndica:es . mound cons:factions.c. and the _a:ger ones sho.ces in Cen:.:o and fur:her south" The societies of :he sou:beast reache_ the greatest degree of soc{al complexi:.he Meso-American civilization was laid through the development of intensive irrigation. ar. however.vbison The numbers of bison fluctuated o'.-:ods of Ume '..'il]age Tr'adiUcn> appeared be:aeon AD 600 and 1000 in South disaFpea:nnceoftr.tY:. -_et'.n (200 B c-*. '_::h chiefs or priests dixcting ceremoria]s. quoting from the Cambridge Encyc!opedid of ..e fo.-1000_ did dear s{gns of shifting agncuhure appear (the MJssissJppian period). of the fooo supply: mmze came scmev.lem for sedentary. anta hieroglyphic writing. '.3:: .alley of O._43a. Wyoming.r.elcpmer._L_cn a.e: the mi] :he bu!:din 8 of l_-ge mcunds as at Poverty Point.nia. o 1580._n and Guatemala were occupied by Mayas. and many `"i]]ages were fomffed. paR. almost oppos:te St.ere bulb. along . of the pencd.a-i:h Jenhinds [:9S3]. near the Mississippi River.

but a number of plants were certainly domesti- . _here Chichen Itz_i (ended in AD 122a) _e=ame c_e most impor':an' center in the so called "Pos:c:assic penod" (A.e already discussed the difficuhies associated with the veD' ear!? dates of some South American sites. are found in most of South America as far south as Patagonia. contained nuclear area.-1520). in '. ! i' II &. hov.1).g.. The rich marine fauna remained an important source of food on the coast. secondary.. fuse.Ma'. while from higher.. hie.Me'Uca. some 500. indicating the Paleo-Indian Period.... about At? 900 for unknown reasons The abandonment ar. mostly in the nonhero and central Andean region (fig..6: DE'.. '_ith each :ente.c state of Meso-America.e: foil into the hands of the A=recs.was de.. cared local b in the early period in a variety of differ¢nt environments.eloped in arid coastal regions. E ?.. ] _" _ _ ] Fig. . the population li. but it was later supplemented with agricultural products. re! ha.._e 7..'ELO?MENT IN SOUTH AMERIC. along with perhaps squash and beans. II=%1 _ Va. the tropical forest of the Amazon basin had a somewhat later. The) wer_ in power in 1519 whet. I .upied an a. "%.21_. at El tubers like potatoes. 6.' 6._em Yu.'. Around Lake Titicaca in sou. where Mexic3 City is located. A su:". and less marked development._n.or. There is broad consensus that maize came to the northern Andes from Mexico._. . to Eurooeax:vn'.hi:h the Toteas._ • .:menrs The major center in :he .. oat.hem Peru there were.. in the south-central Andes the domestlcation of camelids provided an important contribution in terms of meat.:" 900 to 1200 and extended as far as nor'. The beginnings of agriculture can be traced to a period between 9000 and 7000 _.ey of the period and region with additional references can be found in chapter 58 of the Cambridge Encyclopedia of Archaeolog:.a. _ho came from the north to found a ciV at Tenochtiti_. but a wide strip all along the northwestern coast that later radiated to other pans of the continent.n.': regiona'.in:_¢ period ended abrapd.d des:ruction of Teolihuac:in star_ed a competition be'_. _ka!. at the time of conquest. including appar*ntly manioc..Ia:. ceremoniaJ mon.Mcso-Amedcaf_m Fern-. from 'he city of Tol:an near Tuia in the c.-'. (from O'Shea :980) culture was st:'ong:? influence! b. Outside :he con.een classical period. It _as a muhicent:"ic. wool. There is no single. From the riD highlands c_. they may even antedate those found in No:'th America.7.. Their influence lasted from . As already mentioned.entral Plateau north of Teo:ff'. at'.:C _ 313 / ". even- toall) gained control and became the firs: mili:a:-!r. Hem_ Con_s conque_d Mexico. but hints of major novelties are already appa. which later spread to the Amazon basin. Clovis projectiles.. These animals had been food for highland hunters since very early times.'-'archica: societl.ed a [itt:e earlier Po'o. • ?ll. alpacas). in the north. 6..']'a t"r'_ .60 _:m-"(donnings 1983) and had a ?opulation of tens of thousands of people.ed in sma:[ hamlets and was more dif. and animals of burden (for transponation)._ .'ever.. The ciassica: ?'.. _.._'.:'... Their natural range DO102190 We ha_. Tula had been destro. altitude forests or the eastern side of the Andes other products emerged.. By contrast.K:c'_e _ _:aa...:atan.aac.rea c:.1 .D 903-.A'... animal domesticates played a lesser role than in the Old World. • T.3. e... camelids (llamas./.4 .s an_."oa e _.._.6.... Tcetiha-.'-e Peso. and irrigation .

._ = unti{ about 5000 e P. Other states (e. probably initiating or ac_anc:ng economic inr.v agncu:ture began :oappear. Complexity of soc:ety probably _ached a :ev..-.aThe io¢._rr'.fo_'ino. the:gh at which a new sophisticated _ form. Called Tawantinsuyu ("Land of the Four Quarters") it was one of the greatest empires of the world.. By wise al- ._: Bra. So.: r_t_f •. o. El Paraiso had a population of 3000-.000 i __.1 Mr.._ .-_c oc-:_.00O BC to AD 1000j. it became possible to de._._(. an "arcS.e... There ".. Potter:' appeared around 5000 years ago at sites as diverse as Puerto Hormiga.eiop the already mentioned pattern of a "'vertical" economy whereby the same people had access to products made in very different environments. o _re_r:e_' created v.-'r. The only great South American empire stoned developing after 1438 '. Chant perhaps 25.'.:ooo culture. s. The..-'.aT..a:o "_ " _ hei h: a. hundreds of road stations and state storehouses.. that of the C. as shown b:.v. on the Ecuador_an coast.. Textile products were made by the women for the state.... from the coast to the highest ahiplanos.. probaba? t._x... =at..._ _..the Inca '.._ '" _ ___x._...oma .e been '.:.:ze5 milita:'?a::i_ib A major influence _as exercised b? :_e . 'This culture certainl. or o_. Improvement in trade networks made it possible to redistribute a variety of materials at long distances. Irrigation was practiced early and its sophistication increased to _markabie levels.have been 20.:_c.rm. _:ezes of '...e network of excel}eat roads (15. _ .c: 600) shov. densely inhabited.a.c '" :':._.6.--.._._. but after this date I_ger settlements supported b....e de'.-_ ...._. lhe central Andes.-"n -_ "f]ticaca '. _. Time in the arm'.ncz:cre Fie'ted only' a seccnc. s':a:US '.uanacoinfluence established. and labor for the state were required of the men under the "'mit'a'" system.". Sites like Real A!to and San Pablo...elopment of ceremonial centers and true im:eaa.a (-" ._. Ecuador.. Of the agricultural products. the H.: 1980: Mcms 19SOL is abo'.c sm.ices.'e:.mbla to south-ce::t:_l Chile. roughly two pans of three went for the state and the nonproducJag pan of the population.aRc in Popula!:on a great .on .. 6._e_cuse:9"5: _a.000 people) existed at the time in the centn: progress:'._a_. Through ethnic and kin relations.e risen steadib. cm_ v. near Cuzco..4000 (Bray 1980). X'a..00C_:0..'hen. and the bureaucrats formed 5%10% of the population.._. &r:d it is not too su_r'ising that tLe /...Moche.h. adopted an extreme'!? efiective mill:aT?' policy and by building an extensi.. Cotton (possibly a local domesticate) and the manufacture of textiles soon acquired considerable importance..'minesc'e'_:Iv according to sources. _ c'?' roie compared _ith ..'t. for examp'.ar-e Y:ee _g.. . ern Colc. communz_:esaccess to. In the Andes with a day's walk.7. k"_r:a..:::cnof a ..h.mode_::':_e t2eYa..g.:c.. _o. it is possible to go from one to another of a number of different ecological niches.:?e'..m_.. had an :reFer-cant :mpact or'.ing Lake T/o/:ua.c"d the Van. At the Fee. g..3'.000 km) across _ec.. . capital C.:re (nonhe:"n Peru) a vast area..- .as a:quired in the .on a battle against a nearby state. !980) A_. and socioeconomic advances allowed people to make excellent use of the variety of microenvironments present in this region. T'_e Moche site pictures of the north coast of Peru '2"2 B C-._e --_. i. despite the lack of a currency._at ".o:afions later adopted b.....nership of..4 CH_?T£R liances or other socia! ale'._._..3¢ m and their domesticat. c_. _ . they rapidl? conquered an extensive territory.' difficult terrain. r/ / /4_ ' k'-._'._.he southe.-2 of a_r:c_hurai set:]em_n:s in q "" .._.T:n: which at :he time of conquest extended from sc_th. Terracing of the steep Andean slopes was quite common and greatly improved water control and productivity..'perhaps the first hints of orgar.] "--* . the Incas A.'as inherited and perfected from earlier states. which '.. density must ha.: cuh:.k.-'r-:.e begum '.' / _ _ " _ made up of 12 million people Even if this es::ma:e '. .1_ 1983).]itary conquest (Morris I9SOL an empire :hat lasted ::r:::: AD 800.... m_'. the area must ha'.._c4-:_.. and the rest was disudbuted by the village chief among villagers.ha'." _ /.. ._t_.':ca e:m....::ho. and Valdivia.%_ '.ze of ecor.idence of political or military' occupations. ha'_e be':."? earZ': rS0O0 s .. . and a well-trained army. spreado'.e 5.out 3oo0 years ago. iF: :h:.nrougn .._ k'.... Am...a. It made possible very rapid r _a_ conquests and the monumental buildings dedicated to ceremonial and civil purposes for which the Incas are famous.r._ period..u:t_ d-" T_m_.-: any e'.. perhaps as much as centrnl Mexico._-.. are large stable preceramic fa.. near Cartagena..e. in addition to trade. The rest was a rural population on whom several types of taxes were levied.a.-. Inca was the name of the hereditary monarch The nobility. the priests.[. located on ._ _._.. Colombia.._ . me ' aTban popu]ahon mav.a..":"./ / [_7t . Products taken by the state--food and textiles-DOI0219I =.c ter the col!apse of the ._iety of areas......_nasa :_dic2_ed ifrcm Whjteneuse and _.. f..-...

Several of these--in pat':truEr._a.e!l. an_ de.7.. but a rehu'. Saizano and Callegari-Jacques (1988) have compared groups that they call stage-A tribes (hunters-gatherers with incipient agriculture...":haeoZogica_!.e .e.f-.e_v unchanged _. there is migratory exchange bet. Although tendentiaiL'. They occupied a part of the forest still suf.1..orab'.iro':ments and pro'.e biological in_estigat!ons b. a.:a) ali Ri'.as ass.the present Equadc'-Ce'._: gcoc c!ues are '.he Onnoco and Amazon basin there exist robes teat ha'.6 1.:-. and R Ward.mated and disorganized the Indian population.98C E'. population genet::ists P Smo."..'.. The enc..ed to make chicha beer from it.'1978. because of long bit'& mter_aIs. sur.. s sever-a! fissions m2d fusions Fissions reflect hostiEties betv.octs rich }n starch but poor in proteins..aitabie for tropical en'.... Noel etaI. At the time of European contact. is sufficient['. 1970: Smouse 1982: Chagnen 1983). ing kinship relationships (linea! fission pattern)has the effect of reducing the effective population size of the viIIage and therefore incre_es the effect of drift over that expected.udmg among man. _. in spite of the fosion-fission history._ crops _em ne:essae: for the: wet soil and chmate of t.ere '.) s..icted after conques'. Ananatuba culture.*th Their present location is sho. eC tc the pop. The finding of potter'. European contact.:al .':.:o.n Peru (about 4000--3"00 years ago).A ). as described in detail in the ongmal gapets In particular.. 6..."aids ( cf this area Ne'. fro=... but the earIiest _'..r.nung. Despite their io'* fe. ers in'" a system of knot'ed smngs of ob.e Amazon: the most successf_..6._5 P i .e history of Yanc.a:s and special bo_.en toes': in '.'. pets comm) one _as due to the capture of tv.. aho ably exploited c[vi_ unrest..h:s in t_. R._c-ltu.ed from the Valdivia types. .probably deri'. for location see fig. The re:n_kabie population density a.. The findings in other populations in southern Venezuela or in northern and central Brazil are similar to those of the Yanomame..tu_ ('. were spread downstream by colonists. af ::re': 'acre :.o door. :_'ea__ . s_veet and bitter: the sweet '. The Ucaya!i River is a tributary' of the Amazon.' others. and are currently' drifting slowl.dean states and empires 'acre unmatched in the rest of South Amer'ica.e). Xavante. v. o >'_i=:are women (Chagnon etal.. 1970) and the other _as due :o the absorption of a fe. ha'."or agncu'.vo va::ie: es. at shot1 distances from each other.elfa:-e .i.e de_e!opec i:t the Areazon forests in spite of the difficu'.': at the time of conquest. "D. ser'.i of them '. DOI02192 .y sim?'.r.e. developed on the upper L'cayali River and other tributaries of the Amazon near the Andes.ues me: bv fm. certr-a! Chi'._ southward.':. and even at the mouth of the river (island of Mamjo. "D._&itkampand Chagnon 1968).mame . endogamous.'ii!ages shov. Like other huntergatherers the'.es wen soon e'. including much of the At'.'. In the same area and ume was also found the firs_ porte.atitude from ne_. Chagnon. are at the moment in a period of demognphic gro'. fermenter._! amhropologis: N. and the first visitors were impressed by the quality of potter) H_g_ dens[ires '.ed the puwoses of communication and accounting in lieu of v.i'. Since manioc seeds are no: used. Inca emph'-e lasted about a con:at'. or graters Manioc me) have been domesticated at an eariier date farther noah. to a much lesser e_tent. The Yanomame are tropical gardeners who akso rely on hunting-gathenng activity.h:c:'.'. was pr¢_ab'.c: kiiled by' disease ot sh'.eyin_olun:adl? imposed to Peru.s empl. so it must be coup e_ with other food. tee }ano.'.. There were also later migrations requires a specie'. an" propaga::on is extreme'.":ilitv the. The "khipc.e been relati.'2.. 1977) and his group inc'.e It is :special'.een villages. as in the case of the Omagua and Cocama tribes of the middle Amazon.. the tendency to fissions foilov.e .-3000 inhabitants..een villages of the same tribe and....pox and measles that <_.*era oniy tv. Further strengthening of random genetic drift is due to the high polygamy of village chiefs.g a potsonous subst:nee that gone=cos cyanide blanioc cuttings can be easf]) plante.asmanioc... like the Yanomame.'. but there are differences between tribes depending on their econom'. The bibIiogr'a_'h) _s _oo extensive for a complete listing.e a tow number of bh' members of atmbe the: had come upon hard times (_. the Omagua had villages of 300. I: was destroyed by 250 co:q:..ombia border to sou:'. ::'. This would explain the Chavin paintings of tropical animals and plants that do not exist where this culture developed. an ef_cient state organ :ca'con. This p!ant e'.'.". The Yanomame move frequently.aea:t region of Boli'. often under pressure of hostile relationships within the tribe and with tuber tribes. on the lower Amazon. date 980 B c ) has suggested that cuhural adaptations to the tmpica_ environment..hh other Indian tubes of the region.'m.merited instances of exchange :Need.res led to Per'_ by Pizarro in he 15Y" The Spaniards _ere gxa:!) helped by epidemic diseases like _. dense populat. :: is difficult to trace it a.980.:ree E.. '. limited that there is considerable genetic heterogeneity bet'. a situation that is rapidly changing now..ia..or) fa. The'. and it has been suggested that there were close connections between Amazoma and the Andes during the Chavin culture. Cayapo..&..ficiently undeveloped at the time of the Noel study that they could keep to their traditional customs.arid:'.isn_.:w) '.iation accord n__:o rack.ides . The genetic exchange between Yanomame '. Spieiman.<cure!'.a. which can be foun_ elsewhere (Chagnon e: at. and indi.. linguist E Mig!iazza.da:ed fine is from _hrinacocha on the upper U.) ¢omest_ca:el firsL The biue" t'..een grouts and often take place along kinship lines. assuming random an4 the Makiritare--ha_e been the subject of intensi'..ere possible or:i..'-m ous .on t.')'.illades..ati.d degree of camplexi_y and organization of the A. _15 were r-_ds::zbc. the empire spanced 36: of'.er in northe..n in figure 6. Neei .eztment for destroyi.

and [:_£-strial p:ar. end :he hea.t to th_s per:or and additional re._ the opening cf roads and modern agricuL. 1984).er. Most northern Mongoloids have shovel-shaped incisors.ts c.ant area.:'n:.e!'.:-::::':. f:.:::or.d R is likei} that the a'._cuna. which are also found in fossil skulls as far back as Chinese Homo erecrus. they have very little.h:ch is especially high among Siberians.'. Amehcan Indians ha'.. graying. 161..? than at.:erences can be found m chapters 59 a:d 6C._Ise.adation among Siberians.a_es co_ered b'. "'despite some of their peTu:ia._"c:-.:La_es is greater m stage B Cor<emp. Although Eskimos and Aleuts have peculiarities of their own.:.s that most are settling under pressure from go'.-ac:eristic 'high cheek-boned" appearance The nasM bnd_e is usuaI[) low and flaL but there a=e also aquii r.d are "'more unifc:. _-e causesof serious e. _hich include large cranial and facial Sur. and western Chuckchis.qties :n blood groups" ar..g'.a_isXar.exsee'. the Tungus-.--.thic eye fdd in a percemage of indb. that on the basis Physiaa[ anthropologist C S.ce.7. place£ forv. (]965} dis:it.s on the other. :he ."ad'.._:k:n_ and i_ oi: fiords is the Cambhdge En<_cioped a of Archaeo:og?.emments. ho_e'."Jor. 1989).oids of a particu!_" kind ' _e or:g!n of Mor.g :s nm"ro'. bur.:..":hem China cr noah of'. Siberians and often prominent.tation is usually darker among Ame. they tend to follow the same general pattern. Makint_-e.ard it..tura!ists and _she. -:.. This and other cranial peculiarities have been a major reascn for claiming independent speciation of Mongoloids (Coon 1965: see also Wblpoff etal.m-. but the picture of migrations from Asia to America developed by T..'asmapped for 43 tribes (Salzano and Ca!leganJacques 1988) and there is a slight difference berweer.d=an Indians.'Tr. and based essentially on dental clues.'r. The .eme:-'. This foi'.:'_:es :. Coot.:. idua'. if any. Fe_i'. and nasal bridge. and man.e." especially because the geographic distribution of Siberians has changed considerably in the last thee centuries.d Scheli 1979)vanes considerabl. among: _'I.codagric_'.)nonhero Asians hKe '_.::cr:alAmazon societies and bodes it: fur :ae future of these ?o7ulations The e\tensi_ e des:r_'.:h :'e :r. even archaic.d ad:-i.-a'. and the mainland opposite it: (..3 cm) inteRribai mar:_. but the u-acit.'.. but the? are Mongo'.arc o:_:ers'. local econcm.e oper. inter'media!e fo .tshe£ bet'. Dolgans (a small _oup south of the Taymyr peninsula). They all have extreme Mongoloid features. Tempera: % ocaupat:on in _o[d-m:n:n_ cFc.<'m.: (2) Turko-Mongdic people of southern Siberia and. :he American Indian type of nose is also found in Asia: Coon (196_.onal way of life has beer.::.. e':efclds."Me'. damage to the local popu'. if any.ner edge 9f the e'. Like most Mongoloids (with the exception of the Ainu). sometimes convex noses T'ris is perhaps the main differer..u. and immigrant H.e!c_meT:._.ard and believed to be either ir no. the northwest and the central-southeast (157 cm vs.dseuthem Siberia.and stage-B pan of the Amur Va'.lanchu people of cenmal Sibe.. '.'nbes"_echr:o:oglcal'. flattened face. the crb:ts: the _'.first tv.--..o belcng to :he Siberian Mor.e less fiat faces thai'. from otLer forest people cf :he Amazon-Onncco basra shov..ariar.go.t. protrude fo_..ncrc.a. It is important. Kamchatka.o.one_.: _iff. others}.mer. a_d road building..<)cited the Tibetans and the Naganns of' Assam.ifc.ed to a s:i: b. Cain_ar. body and facial hair. b': other. mg an equal'. Brazil).e Nganasan (Taymyr peninsula). Of special interest are the studies of dental characteristics by Turner (1987. a small group in the nor'them pan of Sa.go[o ds.goloid in ger..s the _asuai pattern of climate adaptation.. e?s reie. as just mem.ear area.*er.Lhalir.-opical forests (Guatemala.'_on'a:. PHYS:CAL ANTHROPOLOGY non-Mor. espec:aii.? other group of people occup. '_.!.]e... the Yakuts (middle Lena River).'_a:ecangets _ha:_o '¢..£.d more ur.sar. aticr._een Eskimcs and Aleuts. with little. .o'.c ae'.re vc: . (3) the Ni'. The :o'-. but abundant and coarse dark hair with rare balding and late.ce of :ke ::_.:ike '-. _]979). :he Macush{. nasal bones.ermar_:r cf the orbit lies re=her fcr_ard and the z.khs (= Gilyak)..."alar. and the lo'.ity (number of childre:_ m. Browridges are small.':'m _aia.a.c_cs m:.Mongchan or eT:car.:_:meat.. genera::ng the cna. if any.m::'.rm. o_ the one side. sapiens sapiens is a possibility worth considenng. mostly reflected in the conformation of the skull and soft parts of the face.e_ge s:ze of _:. The genetic exchange at various times and places between local human types. maintained in a few cases.ra_ but fragmer:a_' info.go'.d came by a later migration: American Indians are stated to be . 6.':ces _er_ four.s. In South America.'n. being highest at high latitudes (Canada and Patagonia) and lowest in "he _. if any. mean stature '.'.. the eyeballs are wide apart and smaller than in DOI02193 .a. _.g. is unrelated _o this question.:destr'-c::_e :c :. Yukaghir (a small group east of the Lena River).. but there is also ' meier demogra..of "he forest fo:_. the maximum development of Mengoloi_ features is found in central ar.-ber of children higher in stage A. V'..racia:'o.oids (see chap..c bone:. and '.. h is difficult to give a "'nuc. The mean stature of Amebean Indians {Johns:or ar.e noses. Pigmer. and American lnciar.ccmpleted families) is a !!!:k io'.According'c A'. :. espe:::. the lightest being the second group followed by the fourth and then the others.y more ad". These people have somewhat variable pigmentation in skin and eyes.

st Asians from no.a.azee :nat the Na-Dene ma.ei of heritabi!ity.olution_ D changes as a function of adaptation to diffexnt types of foods. To increase the!ie'..ho approaches the field of me dossificat{on of American Indian ianguages can only be shocked by the segregation of '. but it _ould be unwise to re!? on them alone until more is k. and Eskimo-Aleut. The diatribe has been the subject of articles of popular science (two rather extensive summaries by P.2!.ats.iding it into three phases.axeoriginated 3!" from the tare D_ak:a_ curare (fig. n Ie'.a. These hypotheses require indep_.en: of that of Pa'.'as severed. Another summary. large are conclusions insensitive to the admit:on of fur-thor information. The Amerind family includes most North American languages and all Central and South American languages.. The question of he'd. miomevo'.. Statements based on dental anal. classificalion. and :not sensiti'.nown about the problems just ment:oned. teeth have the ad'. Tumer's ana'. the group of splitters uses extremely srror. stable in e'.hem China and the :Southeast Asian u.'<. Sial<.!thomer independent sourres of e'. These conclusions agree . Edward Sapir carried this effort further._ordin ferences be:v. Greenberg (1987) who has earned enormous respect ft'om the whole linguistic community for all his other work. bat after the No-Done.ingui._.ution. a:rived just before the bne_ge '. that of the Esk:mo-A'.. other traits showing ma. the interrelationships among _hich are considered beyond recognition: the other much smaller group proposes three families.::st v. of th_s e_:aence.e have a_mady bhef.._..olution.C3 of the dispute is in a Postscript to the 1991 edition cf Ruhlen (1987). and Poi. especially if and when they disagree _ith other sources of e_iden:.ts into t'_o groups that hold almost diametrically opposed beliefs: one. in time sequence. The first was stoned by '. Japanese of the Jomon period lchap a) she. ation..antage of being readable in fossii samples and perhaps a_s¢ of offering greater detail than bones. m_ch further back dental data can take us in human e." The third phase was opened by the linguist J. corresponding to the three major migrations that are also recognized by other criteria.i:h T'nm_and. at the beginning of the centu U collaborated with R. it . How much further back this agreement v. Wright in AHantic appeared in April 199t).: state! :hat the third migntion.* fl'. be it of living organisms or inanimate objects. een nonhero and sou_em Mongoioids are the number of cusps and the number of roots cn mol_s. two of which were Eskimo-Aleut and Na-Dene. '*ith our genet:c ana!'." and the dismemberment of Sapir's families.'.e :o.. 19S6 and.oiution is also a matter of conjecture.. am c!ass!. DOI02194 .t from dates. He also postulated that the Na-Dene migration _as indepenC..<7.z:zdtnez to the Nonhv.. 6 2.]a.. This began a second phase.*est _.: be negligible by Turner--and the unknov. di'.eo-lndians and that it occu. as against 13%-25% in sou:hem Mengoioids _"sandadrn:s" Different percentages =for to different pgpula:ions sampled...e genes.ys!s is based on the p_mise that dental chanacterist[cs are highly' inherited. es: coast of the Pacific He z!s. namely.. the number of independent units of Noah American fanguages was back to 63..he famous anthropologist Alfred Kroeber (1876-1960).a--12kya. _ho. Unquestionably.'hat can be detected by this approach This is also unknown: only' when this number is rea'. One cannot faii to see this as the most dramatic example of the usual division between "splitters" and "'lumpers.aeen Ea.nodonts" according to Turner) have 60%92% sho'. Ruhien (1987 and references therein) summarizes the history.ed t. who made the claim that there exist only three families: Eskimo-Aleut.'-eenberg e: a'.e. T'."al se!ection and of dietetic customs-.":'e_ i-. dental data on nor:hem Asia. As . Apart from the ur-<nown role of natu. southeast Asia. y indicated in chap:e" 2. he hypothe:.mer ca:cu'. together wkh the Ainu. LING L!_.'_di!ion. Na-Dene. E R i C _. Amennd.s:s _-'e '. Dixon to reduce the number of families of North American fanguages by combining some previously recognized taxonomic units. which can be called a "revolt.ery!! '.g language against the author of the umfication of Amerind languages.s:s {sec 6. Ross in Scientific American and R.fied by dental criteria among the southern stared that a strong difference exists bet.":en:ages. more numerous. _d. nonhero Mengoloids For instance.e!ing.. controversial" representing "current received opinion. v.. and in I929 the number of North American families was six.f. I".ated la kya as the date of :he first crossing of the B_r:ng !and badge by the Paleo-]ndians. apa.ik go remains to be seen.n_ :he Ame'icas are genera... The non: ng. in exce!Ient agreement with these from _ing'..idence (G. 6. of ciassification of Amehndlanguages. Greenberg.9 et seq).as car"pie:e!> submerged Ir a. passel along the southern edge of BenngiatoKodi±Ki£a--. the languages of the Pacific Northwest.endentconfin'r. after a 1976 conference. Na-Dene. a. just before :no land bridge of Be. refuses to recognize unity in these langlaages and chooses to !ist a large number of essentinily unrelated small families or isotated languages.$. Using denta'. an important consideration is the number of independent genes .e to e'.ng!a v. and Amerind (1987)." which has been observed repeatedly in almost every. The list of the results published in 1979 was stated to be "'conservative amd not very.nesia 7_e.'-!..e:.

.xsis found in the recent book I__ng.:sn..CP..ond the recognition :antic'.. Aleut is presently spoken by 700 peofamilies pie in the Aleutian islands.'..aithout .Aztecan is in mos of the Sou hwes.¢_a_.o= .:g:c . Chibchan languages are found on :he soulh_. distm£uisn:n_* the same three manor groups four:d from :oral:.-_ ' made _:" Haidal. other criteria selected for their evolutionary._ a..s . The.000 spea. andCadecan far:o.. _ ..'.} of 30 languages The Athabaskan lan_ua__es .. • groups in California and Orezer:._ere ..¢ . is not pro'.:ar ..a s. E_en if :h:s dassificaLcn changes _n the tu:ure.. Central Amerind includes three distinct subfamilies: limit our treatment in the rest of this section to sumTanoan. ale subdi'. t simi'.-I_'d r..lion speakers..hem group of some " '-_"-o The N-..._. . l...40-ALEb'r family comprises 10 languages and III.:'.'" omi::lng an estimate of a deguage)and:heSiouan._o. We II.-DE>..:en b.ith eutliers _Tsimsh:an)as far ro . .ie....':e . _ _" a ]anz'.rra.of Central Amer ca DO1 02195 .:e.estem spoken by 600 inhabitants of the USSR..i.. ex:reme:v detailed • " makes it .. _ S 2 e &_[ : _. The Esk:n:c :i'. . Keres_ouan includes Keres :essemiai!? a smge ]anluted'" or "'noI rela:ed. 5e_mer'. also includes the Zapotecan.:ge2 b_.eeneerg's book con'.'he con'.._ v from ht_t "_' .... Eskimos_theArcl c! and :he Na-De... it would be impossible to recogCanada: in somheas_em Nonth ..]gcnq.a<_ ... Uto-Aztecan._ '. proceed b.4 . a step backward by uni.. ..n'csa: Keresicaam.-_.e_ of re!adonshiz. _._"_b'.D famii'.:_£:'CL:. Hokanisa nonhero group with small clusters innonh. The Asian Eskimo coast of Mexico and in almos_ a'.s!s than to ._... aS:.'.ons supm _ e. for £ene:> mists..._.h as carding :o Greenberg..F_R_>.J'_4 7T:mar:[? )sis For 5c_th ._cons_s:s of Ka:er:a: ..hdv.No:-:: Amen..aska. authors of :no present book.e_ ':.00_2 i li'.._'er: Aiaska and all _ .or.. The decision on relatedness B Pc:urban is a zaJ-: .. h a:so ex:e.:as :c :he Mac- small fr'ac:ien of allpossibie _s fcun'_ in Mexico (Huava. _-d fig 6 anc_ cons_s:s of >. of Ncr.A . lroquo_an..3I@ CH _ PTE£ _' fcr 'A}:iC5 '.. and Hokum A cf 'a'_°''_< or iano_.._C l=r'oHne_'¢at _ rime v.' a nor.entionai families.. cannot make a contribu:ion to a discussion based on !inputstic ar_'umen's._"'_'....8.VoetAe.c The .. as v.mposs_ele ' '_ a::a'. _ i . :se:a_ea ' v.. il coxers cos: of Canada sc...h .e exact m e_-':'o= cf :he '.a._c_s by Greenber--'.is impossible to hes: it co'..ium'" or "'s:cck) need not concern us here. a fe_..rk a< a '. Three Eskimo languages are A. and a fe. re e SpOken I .':."ia.... :he d]ffic. '.1. Chir. Canadian ' ._..es_ aimos: :o the >xbuild a classificaticn that goes be'..onger . .x refers to :he highes_ "__enefic" groupiris_recc__nized.u:h of '.As G.c. g:... the infcrmaticn v.ons. some linguists ha'.vesl..America aug w . _ ....4..'mcin_..:< ]nui.i:h al. the me:hodoIo_ical ana.eir conclusion _es: scum of the Gnat Lakes and :o Ne'a England is iimxe£ :o "he scalene:: that _he pair is either "reA 2.-r. of the Indo-European family. and a sou:horn o'-aof about 130..a Calf group includes the Mu_kogean fa. -_ _.amevarious subfamilies is shown in figures 6._ord ".some impor':.'. as given by Ruhlen A.. and pa__s berg uses a method of multilateral comparisons...azes: .:ng c'n "_'_. Baja California. it......c_-'_.. o" as the Athabaskan S 'U _'_" _.he belief that "sound correspondences" (rules of Maya in Yucatan and Ouammalal..on:::: "t'urok and Mosan . ilm te_ aEal o ]an_oua_ocE _eion£_ _o _e YuFIK S... (1987) is con. 'o-.e c!ass.o:[ar. . of sea:toted relationships..<e:he .. These are offer. the Apache and Na'. acand Califomia.. ii'.. hke '" tk° .... ncmh'.._. ladependent sources...'mc_ia.':g!e ' cn the coast ::c:-:h .Paezan includes the Chibchan and Paezan 85.'_c<o. ._<" .. H Oto s found in somhem Mexico.Mexico..ers the res_of the .U00 speakers. 18 mi.VakasharL Sa'.. can. "E.. contains ":8: :_-= i_s kv.. comparina :'. One of these triteguages: in No'a.... stability. at a _o. " -'_o . it aseaP. .. especiail.. L'to..:. Zani. We acce':: C're*'_"'-:"v '.4.'n:osa..=re tic coas..-.s _see also R h:_ :.e sta..mezones occup.2 Ci. A and B. tm_.uding much of Oregor: is based on ex:remeLv rigorous cn:e._.. <" more than " in sowk'o.o .Js attem::. G:eenberg ' LgS7:_as follc'. read'..ah_ch _..... Chibchan.America... C._.:ian ant :'._ . hence. "-J. .e_...".era __ene:ic.e.... We refer to the three families suggested by Greenberg.n a..._ge _._ out of I 0.<.:ed forTr. a southern D"oupin northeastern Mexico and man)' languages are compared for a number of words and Texas. Greenem and southern California..annine even me unit.'. changeof soundsestablished on thebasis of historical islands! and Tiingit vaCCiC} enccun:ered by the extreme spii::ers are me:hodoiog!ca! The.. Mixtecan. Kiowa (Oklahoma). results b.'.sts..E family is spcke={n nomk:*e_:e.a.. ft'agmen:ar? class!Scar. .hun a Q-de .w"e^ _"o--::a. ¢_C'.ae_¢:em " Canada and central . .¢ _ eS. as families and to their subdiviopi-Manguean and Pima gi roups....a_ea languages..ub::*h _een 3RI _...... d ." "I-h. :: supplies a sta. The ESKP. Mixe-Zoque.. c.'_-_'u'_o_ Haica 31C speakers of a total 2000 Ha:da.A'....s of T:in_it.. and Ch!mak..ith which. the ex:reme:'. _ are sp:.A'.. _.I. and me na is ". ncluding the called phyla by Ruhlen."al sc!entiSc point of ._ ._. sions as subfamilies... examples) must be followed without exception. .el' h Canada..Am. _'_--k]aska :o C'reeni2nc aloe:chain _tre.. some of the families are no '. seric.=-.aska.: rice . "_ highest genetic unit._ "superfa:... " (for which • some prefer "'ph.... Tanoan includes Tewa (Arizonaand New Mexico) and (1987).._* . Penuuan.o_ene..Alaskan Inn:.. _-_ Gre.... Today... .!.des as sum:am:::es."n group inc'.000 speakers in ea.sO'..h_cn has a!.ing on Queen Cha> lot:e and Vanccu'.ar.. ". The geographic distribution of the the south'.meingi? shc'_s. . Totonaaan.i_'.'...._ From a ..1o mean •'common ... ' -'\ ...-'.ersal consensus. a southern.<'._e_ '23 other aa:hors.: .. and Mazatecan. in which of'e_:ancX..._a£e c]as:ers rather .. and Oto-Manguean marizing Greenberg's classification.:a...<er.-.. B..

e Macro-Tucanoan subfamilies. _v. other clusters in Ver_ezuelaand Brazil include :he Yanomame.T. TEe Pae-an languages formerly found in nonhero Florida (one language. with a fev. sur_i_e only in South America along the COastOf Colombia and Ecuador and farther down in the Chilean Andes. Macro-_wanoan is found in nine geographic c!usletS. The 20 A. in easte.canoan includes the Equato6. or spoke. there are al_o splinter groups in the Brazihan fan:st and on the non:_em coast of Soutt America.. The Inca empire. V Eq_(atoria/. Andean languages. mostly in western Brazil. Ti'_re are also a large number of speakers in the southern Andes.r*ac _r / south of the Yucatin. were responsible for the spread._dean languages. now extinct). DOI 02196 .a[ and :F. account for half of the Amerind population because of:he great diffusion of Quechua and A._des also speak. including the Mapuche (= Araucanians)._ '-_.m Brazil. which is therefore recent. nov._ mara in the central Andes. of the 583 Amerind languages. A.S_ "C -_ 3!9 l -/ 1 ' U"C _. Three small areas in the nonhero A.s_. and perhaps also the Spanish influence. B.

v the family of languages they' speak as Na-De::e. as indicated also by their remaining in a more nor.. the major crilerion for grouping populations is linguistic. The date of entry of the last two groups is probably 15-10 kZ.hun an abor:gina! Asian population.6ca: and Andean. especia::.-.zhst..-_: :f Ca.:ar'a m. 1986) tentatively recognized the three migrations.ds :o U.. are unsatisfactory. however..'r_m . B.'_ioni.'. major subcluster of Northeast Asia. there was a splinte Apache r g. ". Macro-Panoan. B E.c ncr':he.:a..e Paleo-Indians were :he first."aphic dism:'. there isa study of Arctic populations by Szathmary(1981. b :ha: at.".::__s _e_ s."o_p inKansas. Chibchan._. At'awaken • VI. in particular the Na-Dene. 3 1200.c:or_sZ hasthe_a_est num_.ene:. even one as informative as GM. Mc..h .:ea. The Kant':gang language beier4s te :::is group.?= ro -C_.lyin Nor:h America azd does not include all people from Centrai .heas: As:a "iq. cf all A..7:.e..ould of course be unacceptable..m_=.. is not striking and conclusions based on a single genetic system.'o. and the Pirna.idespread 'r.". and South America. Within a few linguistic groups.rcup and :he rest of the American continent '*as divided into :".a bile el: American Natives including Na-Dene formed a separate.) pa. it would be especially interesting to distinguish subarcas in other subfamilies. "_'hich is ac. 6. .'enez. In that analysis. The procedure adopted was :o DOI 02197 Bc:. Paezan..7.ah:.:th Turkic-spcaking populations of nonhero Asia.a few Janguages survi'. r:b includes .c-rb[a.Lcn of Amerind languages is extremeiy ffagmentar:. nonhero and southera.:-_ (:entral Amerind are found :n Nor'h and Con:rat Ame:-c'a. 1989) dental analysis discussed insection 6. . in x. taking the Central Amerzzd subfamily.'ual'. Although the exact time of their migration from Canada is not known.ant:rcpe:ogical and :inguistic e'. M'acro-Tucanoan.he basis of biological characteristics has recently received some tentative answers.. :O. (1985) collected GM and KM data from the Apache and Navajo (southern Na-Dene). In view of the special linguistic interest. Oe and T.'. 'I'v. Colombia. all hasically positive.NAL':'SIS OF AMER:CA edged :he existence of difficu'. Papago.see also 1985). and they' are believed to have arrived in the Southwest around .":..on :he cs:.:ies for :ra'a:::g finai conc. {.:o-Panoan."'agua' An impo_ant point is that the gecg._oon once extende_ from Peru m Uruguay: max? languages are no. A M.v.b ian_.entra].. however..hem area The Eskimo-Aleut were the late_<. we also added Chukchi and Kodak in order to test possible similarities with Eskimos.... The Cent.Macro-Go. who used data from 14 genetic loci and found the Athapascan (Na-Dene) are more similar to Eskimos and Chukahi than to northern Algonquians (non-NaDone North American Indians). Macr:. ". and Equatorial in Con=re'.. and Macro-Go on:y in South .usions In our paper (Ca. but acknowl- .e question of whether the three migrations can be disting'a:shed on '. Eskimos clustered 'a.a::e_ as a possibi!i:)."es:eua:_ are found oni.nhem region both in Ameh:a and Asia.:mmahzes some of the poi:::s made in chap:or 2.-'_:.3:0 CH. The nor-:.. The difference. r'at:er '. a. In the analysis of this section.< unceaain..::. include two major groups..eO .h_reeme!or groups that may haxe represented distinct rnign:icns.Until recently...extinct.::oa:iier_ far:her south Geographically. fo. F'eru.een 35 and 15 k. The presence of some Eskimos in Asia is beiJe'. ai! from Nor:.Ma:ro-Carib. There s greater ccnsens_:s for icier dates. ".V.4me. Hopi.._z:] The!argo number cf languages [sdue:o the :nclus:c-: cf :we important sat'families. mostly {nsou:horn Br"az_i.9. '*ere next. Na-Dene spealers.anguages and is 'a<deSr_:d . in Ncr':h America: Penutian. in Scuth America.ith Chukch: and v.'a. and Walapai (non-Na-Dene from the North American Southwest) and showed that these two groups differ genetidally. in :he highlan:s and farther south..and central and eastern E. and . 'n_cs:!.. we use a further subdivision on the basis of geography.h_es_em American ind!a:s..Mac._i group was defir. there are not enough data to form as many geographic subgroups as would be desirable.. Hokav._cken :n :_e"cr':::e:_re gions of Scmh Ame:'::a.acro._. cro-Czr. In addition to Tumor's (1987. The southern Na-Dene are essentially the Apache and Navajo.:. but e=oag. al tee Na-12ene were collected in one Z.mce_a:r.Pc.finga small subcluster of the NoRheast Asian :25_.' :he extreme nc. Ecuador.idence points :o t. Because subfamilies are dispersed in widely different areas.:c-_-Ge was '.. a.ed to be a retrogression from the Amencas to Asia. it was pmbabty !ate. earlier one must be entet-:. a::d !nhabit enJ'.-C:r:::.hough their cute of on::".osan and Ke.. Alrr. In a more systematic analysis based on data from a larger number of genes and populations. Williams etal.el on a linguistic basis. .":.::L Ame..:he Gaianas. Eliminating groups because they take unexpected positions . PH_ LOGENETIC .as:to east and from :he Caribbean :s:a". bet.eia..m_r!r:c'.aiii-Sfcrza e: a! 1955_._.'. :cgnufied b.... but unfor_unatel.PTER 5 C.. .%C. Zcgura (Greenberg et el. a:d S:. even after the pooling of individual tribes into ]inguistic groups.

In the list be!o.. Inu:_ (El:. Ge.. Ingan_ (Colombian Quechua). Makiritare (. Piton.Athabascan):Apache. Pemon.aIysis of admixture considering t. others F.-D_-'. Almosan (NAL): B!aclC'oot. which might be bet:or ma.. case.hich then wen clear signs of admixtu."_o .e have tried to 'AmP gaps to not more than 50% !n the data matrix. less wellin.'_ahitaPawneeL Cherokee 3 North Penutian: Seminole {= Muskogee).k '. Salish -. I. We repeat here that. Tarahumara. Bribn. inevitably have a more irapor_ant inEuence on the final conclusions than genes more rarely investigated Restricting the analysis exclusively to these genes.eles. Canadian No-Done (Athabascan): Dog=_.' absent in . Nootka. Chipaya. 321 Figun 6. and table 69. 1985)... 6 !3 The g_'. other that.8. Naska.. T_es from populations believed :o ha'.ToztiI). Ojibv. U.1 sho'.. Toba 3.. l I be:v.'ere some other.-n. Rama.S. Northern Amerind 1. _e thought it useless to ca_. together _ith the :names of :_e major tribes that fo..'.Flathead .. Macro-Carib (sMc): Canib. Kensiouan (NKE?: Caddoan (Caddo . Tzei:alan ('Tzeltal -. Piro.s: II.ed them. Panare. The 33 tribes or _rsu._a_ext. :vu!d be considered q:_antitatjve:? va'id. _ith resuJts using RH al'. Of 58 mbes. Such genes... es. there '. Choiute:a.A.". would have reduced its power. however.y useful for e'.. '#'a:"a_ D. Nahua.A.: Canadian Inui: _ECA):Or_enian.. and not only in the southern part. the tribes that are named are those that have supplied the most important pan of the information. because they are represented in more groups. Mantagnais. Equatorial.. the . A. Xavante DO/02198 . .. Piaroa. were eliminated. Zapoteca C. there was enormous drift in many populations.n da:a.. Wapishana. T:ing:: /N'DN! B. for instance.erewas a carrela:ion. generating great variation from one population to another._ A.Quinau1: . Colorado. but it '. ESKLVO-A_L'T A. Paraujano)..aticr. Parakana. used 17 non-RH alleles potentiai'.m_o: 'use: i: the for v. S'. "l'icuna. the:efore be use4 for infer'fl:ng admixture We are reassured by the results of another study that the possible Caucasoid or Africar.Y_uana).S. groups or subgroups that had fewer markers. Te. Chibchan-Pae:an I Chibehan (MCC): Guaymi.. een :@Q and 25%. Shuara). No_he_ No-Done (non-A:LabascanL Haida. as in other chapters.d!:ui: _EGR) B.)' 25%. even some RH alleles. Yanomame 2. with the th. and Hck:r: were ."sT get:eric distances among groups._sthe geographic dism_u::on of the linguistic groups.a. Tnere is no assurance for an)' of the most info. Baniwa.a. '.. Jivanoan (Jivam. Noanama.L'. Imam. Arawak.a_e.e less than I0% admixture ga_e results very simiiar to those obtained using the general with elmer Caucasoid or African people.Pano-Carib 1. Paez (MCS): Atacameno (=KunzaL Ca.s :hat conrributed most to the genetic data used :n the an'q'.Cre.. Macro-Go (SMG): Caingang..i_lso .: U. Tatamanca (Cabeca. Macro-Panoan (SMP): Cashinahua.g :o :he authors _.r. Wayana. Andean (SAN): Alacalaf.auatean listed belay.aluat:ng :he proportion of non-Indian genes and compared them. Maue. gmerillon. Pacas Novas (Chapacuna).'. N.'. Yaruro.-kers of admixture. Cakchiquel.rascan.'. Quechua E.Penobscots.igate:i :fibes mat are not named below but are Iisted in the tabulations: data from the tabula::ons '. . Totonaca .-Muldeshoot . Bad. Pact. Cayapo.. Chulupi. Palikur. Chane. Aguaruna.Micmac . Trio. Chaco.sis are listed be:ow.. Sumo). Mere. over populations can help reduce the effects of drift of individual populations. having been tested for more traits.heoriginal American Natires and car.a::'. Yupa (. Aymara. Equatorial (SEQ): Arawakan (Ooajim. accordir.. hich the number of markers was considered ade.'. at !east for those genes for which data are more abundant.Tucanoan 1. Z_:ai 4. on:. Eskimos: U.". Populations for _.NDS) A. k v. In this '. Kiche).Makah.engua.. Lento. Campa (Maipuran).apa (Ecuador). Yueatecan Note that Penutian were tested joint:? (PEN..'_ E .'liminated because of stmmg admix:u_ B Central Amerind(CAN): Chiapaneca. : 2 -: eliminate s?stem:'!=aiii. l.Nat>e groups has generated exceptiona! gene-frequenc_ vana_on.Okanagan 2. _e find there is a cle_" effect of admixl:ure oni. The avera .. synthetic maps (see. Na_ajo III. Macre-Tueanoan (SMT): Siona. Ta.a._.: USSR A!euts hadto_ re'* ma_-Re.. Maraca.Tunebo. in the geographic maps of principal components (Suarez et al. T?'. Semi" Penutian: Eastern .ere used to calculate :he mean gone frequencies of each group. Mapuche.S.5 had estimated admixtures of o'. as alrezty explained in chapter 2.i. Chomti. This is clearly visible. Chamacoco) 2.e markers. Guayaki.'ee-iet:er s'. Ma'. Oyampi. (NDA) C..San Bias).America as sho_n b._a.' wen trd'.. that the'.". Papago.._me drift in man.or.'-ibe.can .A. Miskito. Cofan..xiL Kek:h. especially in the Americas.ho collected the genetic data. _n Ner':.Northern Motilon) 2. lea. possibie to increase the representa• tiveness of the data. Boruca."s and tended to associate '_iLhAsian popu'. Southern No-Done . Siriono. As to our'.:. ut a direct ar.Ma_a .'. Zamucoan (Ayor_. Aleuts ion::. Chipev.. doubtful whether the estimates of admixture. In aimost e'.V. Inui: :EL'S. adrrfi_re of some data vve use4 is not misleading: Salzano and CallegariJacques (19S8._.

104 79 155 215 1!):1 15 24/" 11i/' 2 11 15 111 201] 116 111 217 52 NKE 1:19 39 257 333 310 193 333 217 3 8 21 69 92 68 116 289 102 100 I/ 204 219 3OO 524 403 468 SAN 40 60 229 364 21_4 21_/ 357 21. SMC.. 70 1:19 250 299 3"37 231 4(15 360 4 0 3(._ 3117 .EUS. [CA.nos.40 CKC 191 259 0 335 980 650 602 562 1370 246 1389 1462 1178 1136 756 618 1002 1161 1089 1208 1118 1430 1429 CKO 280 386 170 0 1414 1070 1232 1227 2015 680 1871 1857 1587 1518 1019 963 1136 1586 1522 1661 1489 1839 1928 CKt] 209 246 537 678 0 621 1667 496 1494 583 1732 1851 1267 1236 892 950 1406 1437 1673 1471 1449 1878 1910 ECA 2"14 139 229 294 150 EC.h_)kchl © O t. Nollh Na Done.ll Eskimos. CKO. Cenllll Mocro-Ch=bchan. Mac(o-Go. SMP.1 457 565 384 690 I2. CKC.2 2!11 226 214 359 3 360 04 79 96 191 401 Ill 100 _l 6_ 17 SMG 91] 121_ 104 241 221 245 411 21t5 409 237 134 1119 172 196 144 I 11 153 110 93 SMP 121] 115 261 405 :112 257 3)7 329 457 370 61 91] 213 480 145 92 94 41 05 5Mr 115 1._ 0 495 811 73a. EIN.3 369 289 103 125 126 263 73 167 41 F--.17 298 56 84 llJS 224 245 244 5411 105 69 69 E'_.onelic DIstances (in Iho lower loll trlangh_ ) and Their .0 1676 t£. NK[. Inupik EsWi.PEN.26 247 3._._ .% I 4f_4 I 270 I.0 731 4111 3.000) PEN PEN CAN CKC CKO CKR KCA EGIt EUS EIN ESR MCC MCS NDA NDN NDS NAL NKE SAN SEQ SMC SMG SMP SMT 0 199 896 1241 1202 956 1286 1033 1438 978 436 334 744 744 256 335.n_. Nocth Central Amednd.5 5£.35-_--.9 6. MCS. EqualOcial.111 :11313 195 2511 31 | 288 347 161 2_8 206 451 149 141 116 140 Ill 13/' 19! 22(1 0 ".ee el Figole 6 9 y end I() a*_'.(14 2114 291 fM3 3!). Ir_hJ(hJd in Iho 0(_Jl)5 ate li_led _n Iho Ioxl • US_Sn [skimns hid Ioo lew ma_ko¢$ ilrld wera nol used in Ihe I.t3 tl!.tan(lard I-riots (io tht3 ill)per light lrlanoh9 ol Ar_.'5 I 42£. Gleenland Eskimos.119 1724 3/'_7_"_O 4113 703 965 906 106/' 1118 1031 10._0 I _. . CK[t._ I]6 30E""--. 135 105 0 1742 1995 655 634 237 5(18 352 340 57B 393 739 I'_ I I N[)A 110 133 313 495 325 191 314 106 11 1 24! 1 211 31t 0 iVI)N 171 124 3£.=.npac191ol_s lid)o. Andean.'¢MC 30 118 241 3£.80 773 295 417 4£._.'_0"_. 61ouped Moslly LinguiMically all vah(os x 10. Macro-Cllib.9. [GII. Pertulian. NI)5.ll" 178 207 64 210 201 66 156 141 1113 M(_. Koq/ak.Table 6.['I 30O 320 198 264 376 EIN 329 2(i5 3!.. MCC..ann(J¢an Na ()erie: NON._ 259 3 26 407 1267 1454 519 645 836 367 873 1144 1072 1133 1242 1372 1073 330 _95 799 677 27_0 1312 1548 1630 1054 690 1220 831 987 1387 1293 1109 1722 1362 1179 2129 492 872 1051 814 1263 1374 1450 1652 1940 1£. S_lh M_clo-Chit)chan.3 425-_0 736 7111 10!14 1109 1193 1054 1410 1234 Nf). 5/'4 9_. CAN.'_ Nole.4 0 488 1106 12£. SKO 37 71 180 259 3Oll 19] 298 259 32 294 1 55 1413 129 327 £. 494 5. N[)A..90 O 133 95 496 _ 0 170 410 6-. [. Kmeslouan SAN. MacfoslucRnoan T fiangl*_s *ndlcBle mole co.341 3'.1'. 50 41] 179 269 223 231 345 210 2 45 231] 117 123 142 119 NAt. Y_Jpik Fskimo_.C..'11i2 531 £. 217_-. SMT..mrk:an Tdbos..".1.97 1329 1372 1743 6111 595 719 669 703 1067 1292 1390 12158 14£.140 216 296 320 4o 4 3(.1] 101 61 60 0 . 257 169 195 174 381 505 524 CAN 76 0 968 1367 1051 769 1152 977 1211 1002 359 45t 734 589 240 419 146 280 230 275 525.0 319 3.:.. NA[. Mac_o-Palloan..'1 606 "vt('_. Almosan. SMG.'.')9 415 354 671 £. _il O.(x:l(Ho wdh (. Canadian [_k.69 1701] 930 746 787 704 999 1335 1437 1. (. South Na ()ene. 11elmieer.6 42_ 2111 1211 189 127 300 190 .19 1157 1£.C 92 115 265 413 . Chukchl. (.

L l:ag.._E_N =. 3q .. an a'...TUCA_O_N . -_ ._? _..d'.. groups (Curia/inn and U.0377 <c!fferen:e net signiflcanfi: but the nor&era and seuth:_. •. It is clear from the abo._'_'so _"".:"-..... or.....L.._" .Ra::bE.-Ge Macr_-T_ca_ca_ Fig.....o.-.n r.."_ and that be:'.. ee= -s_...A.. U$$_ C.._%_ . K_EEE$1Q#ANA" _0$_N WIAC_O-C_R.can Incians otSer that.2 shows the distances ' ' ".. Nora.': :=..... -_o_ dis:anne of southern Na-ber.--'.. o-o:'_< __'-"-..._er e C_ae. . '.s_'-_ Mac:e-Panoan .:_r.N A _ : N NA-DENE G_EENLAND _ANAOIAN ESK.s fo r 52_ c""_ o':._e_F S P_sUT. e=. e distances that :he A_a..hose . ..._:...."q: . OhEaLR¢_....o._s_ _er_ < betv.rid ""1 L '-._2] Es_"o Ch'_kCr: N a-.7 f 1 .h:'. a'... ?<a-D_neshc_..0957 from :he EsK:mos Table 6..'R. 2 ': I C _ .... 6.. ] i ( [ 1 ..9Z.. :r:z frc)m s:-bfzm::ies of Noah. r '._=.2 Pnncipal-componen: ma: of American tribes grouped b? linguisnc /ubfami}ies..-"_ .¢.=_.IslP_::al Cocr_ir_.[e.c_L.) is 0... / I_. p_'_uT_ $i b _<omYA_ . haJ NU_IK E_KiMC A_." c tre_ s_=o'.a irc'..tun....hem .eu_ and gr_.._A C mO -CaM tBC . _-4. s are i._ _ Gere::= O_s:ar'ce [0 Jin_ui$1. genetic hS.. is .r . 6._ t.The PC -" Na-L'_e i :nu_x Es_mc G:eenLa.igaro 5._ac'=. Tha _-_..9.c-. a!: Amer." _ --_ Scu:'_ Macrc-C'tJ_chan .. Eskimos or.....s. C :2 _..-the_ N. DOI 02200 ..e:-a__edistzr. O? :he _ Ln_ o u%.s . forming.distances The a.. ak :......M0 _KINIO ...ces of 0.-f ....m.um_er of 72...KC_'r -Kcrta_ I_ elR_eer N_ "t"._c_.. the most nor'at.S.i [%_L_ AT Nil[.1Geneu= .9. aac..: c. ... es the Amerind cluster._. i [ . .e 4 c[usior.)p." _.e must a.e..."..9.veen the t.o-_'_e_"with .. on one side.S : 6....'.echeasterT: 5iL-'erla.__ure _9 ! a:'c "no PC ..'_e_ ..:..f 13 .. 5 _"[_'[Fee . -. e to the two northern Na-Dene gre:2__ . the southern Na-De.. These con- the matix of _ .'-eak:n2 r:c::-.. nag the four most ".a."c Esk_r_ A_SK_n Eskimo Cara= an _.e other The Na-Dene separate into t_o groups. D ." P. tr...ST:C M_ "he other.AVE.._.°" "..a :_e :.eth.-'.._r a..o no.:heNavajo. C. =.'" nor':&em Na-Dene and seuthern Na-Dene on the one si/e. Ahcean Kerescua_ N_ _enlral Am..:-.0693 and 0._ -:... -A._.. jcini::g the Eskimo and ChLkchi cluster -..".et_=anhngaages._ =o.. r.. '_ _ Macro-Car:: E_ua:onat Centrai Macro-ChiDcna" AJmo&an South Na-Dene _. Central and 3rut:q America _s_i_==:. [ .7: _'.

Thishas no always been possible here because of a lack of adequate data.ene 6C$ 712 _S 6_2 Airno_&_ K_esc.eloped in a highly isolated area._::h :herr dose geogt-aphic neighbors.hich the mixture occar':'ed.: e'.. we have grouped them."them and southern Na-De::e :.'arate particular1. In14bootstraps. 19 show the identical first split of the tree of figure 6.e group fcrme.4_ co:':s!derabie admi'<'.cr: of Esk{'-:s Th.ely recent common origin of the Eskimo and Chukchi. Na.. but in the cther 31 bootstraps.'hich mar:.:s itself in painng with Almcsan in 25 of 50 boc:s::'::gs. unMqo'. that mixt'.o ma-or dusters is clear-cut. which almost always stay together and join the southern Na-Dene in the Amerind cluster._th . A[habascans or. '. We knGv. :-e_. If drift of individual tribes is very' high.ou!d expect it to show a long branch because of high drift. If linguistic families are formed of groups with greater internal genetic similarity than randomly formed clusters. &-ner:nCian S.' early.c. Central.-:. In orderto avoidthe consequences of extremedrift for individual small tribes.s of the bootstrap behavior.c _ista_es {x _O. Tiingit. E'.'. rate because of extreme drift. mere _mila: geneI_cai'. we attempted it nonetheless. w_:h Asiatic Eskimos showing grea:=r similariL'.q."n. they are. in this case..::'er E'_L> mos.. small population that de'. but the data are insufficient to show it.. In the other 6 cases.e an e'. howe.9_.' in earlier t n:es :n Cana. more rare!).:::-:Ts The s:mi:ari::es v.'.y to the Canadian A:ha_z__:z._. one can tr2st the tree structure :o correspond to the order of separation: of branches. deviations from the tree of figure 6..a a'. the potential for admixture is not negligible In 17 bootstraps.m:::g :ha: evoh..c. '..s:e._ees Amennd and Na-_.'he Chuk. :n than :o tee :.hem Na-Dene N_.v useful.tionar?' rates are constant.efi'. I:'_ this case it seems more likely that outliers did not :oda'.e] than the actual time at '. 2) a group made by aver'aging northem and sou:hem Na-Dene tended to join the Amer'inds.k":cs fi_e Amerind cluster.. the Almosan follow the southern NaDone in joining the Arctic cluster. ha-Done 217 319 tee 7.:re w:th no_he.en though the USSR Esk:mos :.em or So_. strong enough that:."n are next: Ge and Tuna:roan are "he . shc.. one centr'a]-scu:. and a re. Ser"._-t"a. We _ currently not awar= of better alternatives. and thus probably to the order of :herr re.:::" the rex'formal and ecoicgicai segrega:'. It is also poss:ble that the no=hem Na-Dene ha_e had some admixtu_ .1 are almost always As mentioned more than once before. Tn... Amennds. v. Fern. componon: respons:bie for the difference _e:_._n.a...t outweighs am original. :he USSK E_...":em at. their other associations are also c':eart. there are linguisti: connections between the Chukchi and Eskimo languages.a._ their affinity with the nonhero Na-Dene. the coast :s falrl? similar genetical'. as one ".9. a: a: apparently '-'eo' early time. Because Almosan is the Amerind group geographically closest to the Eskimo and northern Na-Dene.e Eskimo groups are reasonabh. ha'.:her in a tree has several possible explanations Ass.. In fact. other smgle pcg:::. Bootstrapping shows that the separation of the t.e:" North._e.. but being of small size had a very.a.aa. the old relationship is st:'.: "-= other no_he.324 __H :. northern Amefinds. Chukchi or the Re!ndeer Chukch_. The attachment of NaDene :o other Amerinds indicates :hat the component in the mixt:re due to the latter as.a.' c!ustered in Lhe tree. as probably not a major one.meEcan s_.*in an. strengthening the case for a reiati'. themain changeis the addition tothe Arctic group of the southern Na-Dene.. the pair of nonhero Na-Dene and Canadian Na-Dene.isib e _r: :cc. one or two populations leave the Arctic cluster.ation can also exp. and Sc.9 :..ene..bfam: ies Tv.. the t. according to linguistic subfamilies.*it>.o :airs of subfamilies. Furthe--"more. an c'.. In 6 of these 14 bootsn'aps also._e) a'r"<:_g '..'h_gh se'.1 .a-Done in i3 of 50 bootstraps: the'. to .e similarity of no.. sp:!ttmg from them. one may need to average many tribes to obtain a substantial r_duc:ion of variation. This obser'. the pooling of tribes in linguistic groups can help reduce the effects of extreme drift. and :he.-"TER '5.gac".::.e . Tae_ 8.*o worst DO102201 ..m .an Nor'me--. This may seem a low proportion.:'s'ern Na-_ene and O_er American Nawe_ from Northern and join Cen_aJ *_r.--re_c-? >..a N.. Of 50 bootstraps. Hai.otutionary rates is difficult to accept. USSR Eskimos are a vep: sma]! group and the separation sufficient]? long that this result is not su_rising. the assumption of constant e.%_n C.en greater affinity '*'hh Aimosan..isible :n :he _':e: de:ai'._a before _he mole south The Amerind cluster has an internal subc:_..r. 11 times of 17. given the strong similarity' between the southern Na-Dene and the northern Na-Dene this is not surprising.ith Almosan seem modes: a::C :he: :.a!n why in our ear!ier world tree {chap.e. almost never part Sou.t-'. on the a'.gration away from the place or places of o_g:n When an outlier is a very. Ameunds probably happened most].erage linkage tree at a higher )e'..n.res tend "o attach to an a.:..:. Although we did not know whether averaging by linguistic family would be trui.mes: no tendency to pair . than with the American Eskimos. whi!e they pair with one or the other or both ncr':r._::cr: This ndicates that d_e admixt'are of Naxa'o-. the Chukchi. mixture . but the. or. "i_.

'mt!or.. ma? ha..}--ej:) in Bmzi'.. am repmsemed _."uta:!er .n.": man'.. een ale Amerinds and all northem Mon£oloids_ is in slightt'..'<F" .. . to be greater than the divergence of Amerinds and their direct Asian antestars. of the Asian ancestry.c ..spe " . therefore.u_.e little or no contact '*ith c.a_.e separation of the Esk{mo-Chukchi-nonhem Na-Dene cluster from the Amerind cluster is also '. but they are based on two dimensions only. come 'L With mixtures and other compikca:ions. It is thus in good agnement with basic geography.c.. e. the "Fungus. who num. at least in showing the effect of geography: the first axis separates the Arctic populations at the right..ol'.:na...conm_uticns from mat:) gro_ps be!on_in£. about 35 kva..-.'_'..e: Central Chibchan-Paezan is r_presented b.hich shcv.. is :lear'. ..._kimos.Asia..oda_ a: a !" in the present case for fores: popular:arts like the Ge zr..rr"..n does the proposed .. and the centra/and southern Amerinds at the extnme left..emen: in the last three centuries (Aiexee_...a er ..o.. geog_ph:: mu.a. split into Na D .'hnic group m that region.s..e'.9. In urban c>ilizatior. Panoan.._.e separated in Be.ysismay also support the idea that the t.xan: c..isible in the first pfincipal component of the PC map The separation of northern No-Dean from Eskimos is also seen in the so:and componem.s o be.e.. '. better agreement v. Carib.. Even. and 'Yakut located in the northern pan of central Siberia are genetically closest to the Athabaskan.. .ant "_ ..ons.. the omer :v..a..a :. . a_... be e..e. o_er. at. Per'a..a... southern Chibchan.idea that ar:.'. not as c]ear]. Ca':ap..:. eccncT... =': ... undergoing even greater drift..f_ is importam in this case..G. and their Asian descendants may now be diluted by admixture with other less closely related ones.ed comes _'"-. ..'hat from those obtained _hh the Lr_e._... np_sented b_ the Caingsng ._.."* "_ e.e migrated independent'. _*. it seems .< '"-c'-s. but simply thai it was not adequate to impro.. .... for instance.cuiated in table 2.. be m. _ .u.el m isolated areas The safest gener'a_ cc.5.e. our attempts at identifying one Siberian group closer to Amerinds have not been successful. . o.cbser'...... otherreo_...beand to ha. ... puts all northem Amennds in the center. . In any case... . Using._ different tee!oRs. The next South Amer'. The ana'." Amerind.. ai:hou_<': the ma_or _ssions of the tree are in good agreement _ith info . *' populatior.. 6.a ':...:... "r'ne tv.' positive..:iSfona eta!. southern Penutian...e 'h_ee m_<zrations.. _b_. though..e this problem v..DI__" ': ' come from a common e.us_on from the tree. whereas southern Amerinds show three major clusters: Tucanoan...e> . is represented by seven tribes.'<:'..idua'.ed \" e'_en in Alaska it is impossible to sol'. pamicuiar one. low C-nsity like Siberia.x. pers. a..e. of Om_.'..":her e'. for E. and the expectation of high drift in ngions of very. close _a_rs _.v.. The No-Done and Eskimo ma...1 we obtain here the date of "_[ kva..-_-o _. ant Xavan[e t=C_u in Braz.. .. 1988). ro .. v._er 21 0CC '" Brazil..s.. rations These aura.s. _e number of mdi. ' " '._n for outher__ !s an agglemerati.e on a difficult s_tuation. ra.. mould ce_ain!b. f'_.'e_.t..can outlier.e see in morn detail in section 6. On the basis of nlatively few markers (6 loci)...a. These results differ somev.vith the second.sEa. Xa-Dene and Eskimo. northern Amerinds cluster neatly. Macro-Pa:oan. and Eskimo.... as happened. as is the PC map.o major clusters of the tree..and Ma::o-T. _ _:..'o--. :he T.. Mornr and :."in£ia. _" 7"heGe are m_st!.r.ationfrom other sources.. in the tree T_is _.el in capitals that have . _iCe'._.'.. It is also possible that the majority._'.oudiers >.. Central Chibchan. the:rv..t. Spitsyn (1985) found that among aI1 Siberian l:_oples.. Hov_.. . This does not necessarif'.. and Oe..the '_ less extreme is the position cf the family.-._" car'.. Arctic at.. .. This .Arctic cluster does conta:n a secor:da:-.'. Equatorial. The PC map (fig. " "_.x.::ca: bo.. and C_.._..._ecan ask the most impt..:.'7600 in Brazi . . "£ac_'.[ :his po.i:h the present data....n theor-. proposed...with _.. and from other data to be gP.. nonhero Mon_oio_ds are a '.concerr.. All these considerations.. _'_e ans'. T_'_. Arctic. or the. the constant ca'. x. to America..b. forrr...Thus. The Asian ancestors of Amerinds may have come from a relatively small region._. which underwent considerable internaI mo'....-... bined...idence that high daft is in'."s_ted a_ replete mi_orations. of the American pioneers has effectively left Asia. ith the first date of eat..e ('at ions: .t reconstruct a reasonable genetic history' from v as far as :he Amerinds of South America are .i:h numbers of indi_dua:s comparable to those .e discuss later."{..: :e'.. had a elated or.canoa'_ are made up of onl'... Ti' -. is that. o .. a:'_o u.s :n :hose :-:bes{s sm_.' tc. :'...o postu:ate: mig. '_ immigems ...'..gin in Northeast .-_ . In our 1985 paper (Cava.a: later migrations.o.. to the point that they are no longer easily recognizable.acro-C.2) is morn useful.een the aver'age Sibenan and the average Amenncl is likely... than . be reoonsLdemd again here. :he results of geneLc an£vsis? . the tree is ..*at (199T).c.. Andean. "i'ne question of dating these major migrations ma?.'e or:pit... .a:9".. ha....' :c _ cases of ver) hipS.or:ate roans.. be a small fraction of the iota: for the :. as v.ues:ic-:.' '..".' mean that grouping by linguistic families leads to wrong conmiusions.r..11 that this conclusion is correct.'.en in later sections.. the d>ergence betv. this is ofte.. _. _ :c '. di. c_nft.. the divergence betv.. It is also likely that some of the Siberian populations that nmained in Siberia wee exposed to morn seven environmental conditions and decreased in size.n _o _eo_raah:c neighbors..ors _ thenfoe like'.'-_'.ho li.'re Tara:o.. T-_ ". in the sense of ha. would tend to increase the distance between DOI02202 .er sphnters of the tribe _ocated in othen of:on dis:_n:. verse population... i0 tribes It seems that the greater the number of . and a dating based on the di.d between tribes.en comgiete eeriE. -o -_''_ N a-D o_ ..een the sep_"-a'ion of the .e 610. Pn:__..4 . be icier but perhaT_ no: b'... a ".ifie_ :e some e\ten: b._.'. TE..hem Mongoloids cannot '_et be g:'. Fro.. e: :n some group and the Amennd group If the first is taken as representing the seFa:"auon between Amerinds and No_heas. moependenl of _he '.'. be". the gene:!c separ.aeon some of :hem. the'q.'7 populaIiCES 'a'.. but separaung from them in the first sp!it.. clarification No simple geogr:.._ v. and also b? modem ethnological observations. including Cen'..hich ma.h problems anne in the inte.Th:c ar linguis::c corTelation is found at first sigh':._"_.*een N:-Dene and Eskimo.ion bet.'"_ ' '.._7':'. There has clearly been an extensi'.: o":e In the next se:'.akicr'..:us are _'o'_ _nc:' ..2 the same genetic data are presen.een :'. The a_erage number of markers ..v'---.:.*hich accounts for 59% of '."pre:aticn ot :h.Asian mainiand.S discussed fu_her The amount of o . _. Diffic.'.'ographic mo....n figure 5 IG.'. :s about I8 kba.ere pooled.e o.._k_p_ N_ra_o P_ P_o_go .. "fine anal?sis was repeated for 30 populations from . n'.-_-:.'/ ana[':'zed.h are '...o._.:.'. a complex network of genetic exchanges v.aria.._-...d to an o'.en ha.luster has the same str:. sno FaT geneuc dis:antes. 5 iC.. probabl. oxen after pooling.esof sepa..he greatest rumber of genes in our .'..'his _s not necessari:': a dare . or be..e 6.::.:..i:hin ..".'.s :he ..ral Ameri- [ Inu_k Eskimo G_eenta_ Eskimo Canaoian Es_mo USSR Es_mo Alf_ibascln ! I L I i Do_nb C_bwa Crel Na..e.m__ es:_7-..!.a-Done anc Eskimo is a iit."ai of the Eskimo cluster.r-_:C ANAL': SIS OF i".ben "eared for .among other things.':d "..en.:2 '. t .:en _e consider _he t.Arzuc :.eresumate of :he :_me of passage One may a]soconsider that _ese are .as 61.erai ==_sea of uncer'..e:) :or:'...... gi'. presun'.::: be...he original geneuc ._ been going on for 10 ky has not abated e... := .e become sma'._a:: ::. he'...Amerind from no...'di::g re.Ame_ca of one.: entrv to . Amer::..=st panes. '..10. U" number of 62.< tree The PC ma2 from '.._ are sho'.. 3. There are se.::e (31 k..z:ed :c Esk:mc. s not bring an'.'-:on. i..en :nat population densities in most of the area are still '. .. The res'.h'."'a..hat . .he date of passage may be '. precede'.:_ to be .aua.. Asians.. There a]so _lUS_ ha'.._:'.her in tile next section..cture as before. _ased cn a sam.ail only for The :.e the _ene'....a:th fev..e more than halfwa. ..:.ement of robes._ = :' 8 markers Tab'.-. South and Central America.vsi..h: segara: c: ma.ic _.e .'tes that are better known gene:_caL.lento._..DWIDLa. Cree and " of .omF:e'.n in % .a'.a: discussed abo'.... for Note and Sc_:::: America 6.up '.L TRIBES can linguszic groups because of the ex_ensi'. the cu-.. The clusters indicated are iinguisuc groups and are discussed fur...=a'.0.:.con.en in :ab'.eha'. 7Athabascan and Dogrib_ conne.5.:n an ..erge=:e of .ed as a PC map ".en coda?. USER Esk!mos are :he rues: penphe. the': rend:r.--_ _..::::.=t:e :red in figure 6 9 '..e a not :.tan Guaym_ l_g. and there certainly is at preser. _ith ... st=:'. and '.inguistic grouping used in this secqon._Fa " ....ic •red ..-'. Note .-:... a getg.-Xs._1 .he same data trot _:.]. the fragmentation of the linguistic map. for . :s of ':he ana'....--[ If L _ _'u_ Oh_rok_ Eastern _...7. as shown b.a 'de ha..4?TER 6 S:ber!2:zs 2red Americans and :hun le:. bur '*e are clearly 'ai:hin the range suggested by archaeology. In figure 6:0. a '...ery small and may e'.1 Generic tree of 17 smg!e :nb_s or geographic groups of robes from Noah and Central America C _0 = 00S a 0 [ GeneIic D_s_ance DOI _ _* ..ene. Considerin_' firs_ Ncr:h and Cen:ra'.326 C..a. which has been studied in de.. the g..e been in the past.deer. . (Voeeelin &.argo amo_n_ The rubes '*ere groupec {n :his sect!on acco.ce2.ab_y on the . distances _-._..'.'. similar !ino-u:s:!ca:i'. 6. :he dare cf sop=r:. a lingua>tic cme.ion.9"7).e iLngu:s.

IS ? '15 14 _--2.)gle} arno._'(.. Naskt_p_.1_ ANN ANI' AIU M_G NC[) N A HN AT ACN ^(..0.A.'VN 30:1 3l)'s 42111 AN/' 142 {115 355 22I 24I 40 o I 0_ 7_1:] 1445 _1_ 111. Athabascelrl.t_:es (m Ih.. [ NNA 246 324 281] 15. (_*ee f._lK: d_._. Motzta_uai$.. K(_H. (}pb_hl.. l_fdl_._bl_ C.lf}4 1"11}4 (.9 II.__60_ KCH 235 347 371 255 30l 62 |10 67 317 396 100 460 t00 122 PZU 193 236 403 245._.03 1(. 374 4 28 715 AOJ 140 209 352 115 178 146 142 t 06 375 300 115 322 56 0 496 5 67 840 _ _..( 114 1:199 .... Nava_o. NAI.)5:l t110 1113 1253 741i I I:){] ¢1!)1 t Ilil] 13 .1 '|.-U I '/ _'_ () :IU2__-_ 325 . I IN.112 Note .1 F.... [asloln Maya A/_I.' Ill0 {_00 145 147 302 3:14 211 1711 570 i--. K:il( lel O _liv_!l IloI1 y K)(J_ql_OU5 _lOtJpS . . 0 .I [13 () 4 I_ I¶) t] ._ 0 [ __ Itf. 10 I C. I'_p:_o 1_ZU.. ( _itt.'_ll Itian(31..(42 )fill _--_ EIN f..rt o C_ T. l 401 II/:l 951 1114 11502 77S _1 757 42 5:15 619 1171 IZ l0 HIl!l / FGn 01 0 2111 "130 i_/?c. (_h_uk_.1 2._ lower I.ican lllbes or small _reups O| Ihem (ell values x 10. Ihr_a: M!.._( M..11!_3 112. _uni.. t"tl'..r) al_d Ihl_iq slanda_(l _tl()ls (Jim Ih_ isp_et n(3hl J_i.(i 34 / 210 AfV I:1!1 210 2!!7 241 till :111 :12 0 Z()5 1()(i5 2 05 791 340 511_1 _)14 .U) All)e.(.01 H_ 73 . ( (..5 179 131 82 10 9 130 162 I00!t 17211 I T Z_i 1 52 3 EUS" 74 71 (35 351_0 675 tll)5 _00 2013 913 ( i5 11 G TG !}:1<1 711 1}5¢J f 037 1251 ESIt G:I 140 II_/ 133 A.l:. 7) . ^f'JN Nid_ua ANf _.. 121t{Y 151)!1 1220 2615 734 11 1/ 2 9S ? 1117 1 ._CA (:A (ill I IN l t]_.1 4()1i 131 170 2 7 0 ? 34 f ill i_J._. Cree.|31 (i11 21. 2 Gill ') 11¶}1 1211 'Jf.l.)J K(]11 I 'M .G l 414 2lilt 211i 1114 0 .5 NCI.000). NNA..t .)(j No...5 258 79 66 59 300 174 0 95 _"_.'_'(_ :1:1:1 3()!} 544 :l... 2i3 41 55 46 Ill 307 t23 32 4 233 221 101 PME 322 334 419 327 233 142 94 14.(i t 2!1_. I(.&CN.. _:skirlw)'.v.. At)J. o(J '!...41|4 '_ 1(. (iuayr_ NC(.5 310 129 212 200 232 192 591 66 N 9113 12:]5 1409 .13!1 NA I1I I 17._l. 1 _.).ible _. 433 540 304 441 ACIV 99 291 255 220 201 148 80 92 381 202 5 5 29 7 0 157_'_-./ 1911 II1!_ _--_ 0 1211-() I 1 / 4"_'_"-_ Ii125 I 1952' 129R 01 / 1109 930 755 1211 104. ITl_l_Lill i. I)og_.._(..

'. m O_ 0.S E--. 6...'._.:0) . and Central Amerind " ESK M_C t.l_&S _iS ENI_ rllll _I win.Chag=cr: i9%'L The stc. 6. Southern Na-Dene: 4... .... It certain'.a::c=of to'*ns or cides or even villages ha.. 'I'he Yanomame may as p.Nonhe..'. probabiy in Panama (on the basis of linguistic consider- _:ens? The.L.t_..U !. but it is encouraging to have excellent data even for only a few populations.6 1).a i _ "" IN_K _ _eo_phic - groups The c'.'.. .' and expanding .. One wonders how much one can generalize the conclusions reached for these examFies..tai anc Sot.':s:cuar S.:pat-ccmpenen: map a'.... Eskimo: Z... I0. ..:1: America LINGUI-e "IC G_O_P GwI y"m Cl_nglng i W_r_m C_IDC_.. which is a significant fraction of Cen.. Central Ame.us_e_ refer:o lqg. Ke.2 P:=nc.. fix.10.y /ifferent de. shou!d net be extended to regions whh a long his:cO' of fo._ DOI 0=.wEs ..'.'.I"...e originated at a considerable distance from their present location in the upper Orinoco (see fig.'.nd.icha model can be denver.a:: e v:" M_.9 latecomers....h]ch ha'. or at least haxe sho_n tittle if any tendency to accuhu.rn.q.too Pmra_mr." linguistic groupings: 1.:nogmphic and mating strJcture Rather. T.' affected by contact wit.-ation. 5 a teta'._s (Yaromame.e ] I ] G_EE_.._' that emerges from the Yancrr..tare is one of man'. _irr....." Yanoman-e are a model for populations 1i'.0 '_ 0 _z E_.-G I _mcro-Cmr:#Demr I _ziw&_n E@ACC'Jll P... I/_.'_ _O'=_I t_'o tffc. v.. scenarios _hich must exist in South America. Ncnhern Na-Dene: 3..32fl CH _.'=mitixe hor'icul:urists in the American tropical f:rest..3 Tree based on genetic distances of 30 Sou_h and Cemra] or near-Central American Indian tribes._ Fig. es:': 5. / A_..s'..PTER 6 _OGP-_8 5 8AS 2 o. Cfibchan / i \XI 3ocrC:-z.:h":.e not been serious'.:ir!tare_ These investigations _ the only ones #cm '.. j tr"_s or _@ 6] 8 1st Pnnc. . . are still mo'. 6. 0..o.....I'_ 1#41_.

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. ""' I 1 \_.'. _-.._nd fus!cns for :he Cast fev.s a s:r.. _.m largely endogamous within the village. 1981.. fig 6i0.¢/.er soye_l generations. Our capacity to understand the genetic stracture of southern Amerindian tribes can only benefit greatly by an extension of studies like those already cited by the Noel group..ole picture is one of incom?lore randomness ef splits and fusions that is not easy :o model quamitathe!y.. PTER _.. c" _ i _ "_" _:_' _g. but give and receive nontrivial genetic contributions to and from neighboring tribes." $__ :¢i :B.._\ /_ / i " s a "s_ C._/_ .4His:or_of _si:::s _-_ 's'scns of sc_en _.. i :c kinship Hnes..% shov.. One cannot exclude the possibility that immigrants from other tribes have closer kinship ties with the tribe.. there is enough genetic variation between South American tribes that some degree of isolation must have been maintained in many instances for a long time (NeeI and Ward 1970)..10. Noel 1980).qth v.itch H. as in regular models of population structure.s:c:': cf _ss. usuai]y of different linguistic groups since several thbes moved a long distance from their origin.. assuming that migrants are a random sample of the population._.di. The Makimare a.¢r 60 }.a.en qewed o'./ _="-_ Of f_S:O_S'_._:h neigh_iDrs must alSO occur._'e :_.._ -=_" _ f i . Their tabuiation does nol distingaish between genetic exchange with neighborB and with different tribes.L::L _r:.or.illages is about tv.:ex _re raide_ from o_her tribes. before they are made toraH? gropessible by the disruption and disappearance of madhional customs.m zr:¢s ' .c:sr: of _e.h_ch '_-c_4:d I 0":::-_ l' i ": r . Yet. There is or.._-. especially of head men (Neel and Weiss 1975. ' C.aticn b? Saizano and CaHegan-._. gene.'. in a small area investigated near Puerto Inirida. Considering _hefrequenc:. According :o a summa O' of inform.ages':_ groups.. 6. it was difficult to find marriages where there had not been recent admixture between different tribes. high at the end of a long period of intennbal hostilities. probably because splits and perhaps reunions tend to follow k3.:_. limited info-'-mation on ocher i:'dmsba! mig_tion.cns .hich split and reus::e. howe'.ene::cs. _" - "'] _= i _"_'_i f _"-" .... and it is highe r for tribes at a mere advanc=d economic level.3_ on the basis of the observed propomons of mig_tion.racques (1985)._hor. Espinel in Colombia. _ ".' h¢"din£S_ a: '. Smouse 1982).l:.. Thus.. to some extent acc:r:_r.nship lines and are therefore not random (Smouse et al.?'.ly IosL and such findings make one suspicious about the real isolation of many South American tribes.. i.0. and even more within the tribe. a certain amount cf rsr:4_m or neath random ou:_reeding v.Igrou?s.ic¢ what woul'd be expected (Wagener 19. Kir.S."} sma'. _icaais each. Groot and A. are no: necessanl? stanie entities _P. The genetic vanafion between . o: :he order of IO0 ir. In unpubtished data collected .330 CHs.' .. Another source of amplification of drift effects is strong differential fertility. The memory of genetic exchanges in older generations is frequen'. :a. Tke .:_1/ i... decreasing the outbreeding effect caused by mating with members of other tribes.:rcT Tke _%iogene¢:re. genetic exchange is considerable. at least today._ / i _:_: _7"_.:'¢ 12.-- y..a:ecfrcm. _nd the '_. '. DO102207 ... genetic exckanges between different tribes on the upper Onnoco became very._of the Ma_kimare (so_. rm_'e _':.. drift is higher than expected from the observed migration and population size.

m_u.wesponl to thee --a. (1975) to compare linguistic and genetic distances of the Chipaya of Bolivia to nine Sou:h _merican Indian tribes.7.. distance ca'.ious .'..s too. e cor::nbuted.z'..e expectation'of iinearity' may be naive '....'. _o"-.a... .:n TLe .. jot clusters.r.atio_ without comp!e'.. v-hich are. Ame!r..n ber.. ii. Spuhler 9".":..:ensi.e.a_lon..t... exc_us>'e...eha' OF GENETICS V._' similarhies of 5.. ]Z Z fro m no: me:':'e:t ao--o_'._-_....'.tact>Algonquian.. Salzano etal. The same measurement of linguistic distance v-as used by Murillo etal. o-..'D Ci..c • 2re'...e-_'" ..-r. and to some exte.._ex..elating correlations between genetic and linguistic (sometimes a)so with geographic) distances... the North .._}_t:c.-_... een the .. ee appropriate for a comparison of the different .ts ar_s(T:g free the am. these ex/. Moreoven we use more genes and fewer tribes.ehtion anaL<sis.vise discriminant analysis) is different from the usual one of cain..*e must summanze :. _as tes:e_ c. o s . I0 has clan. " _ " m : _ :_ : [ _ _ S .. even if imperfect..s _ -_ is fi reascL22'..e been man. However.led .atJon bet_een gere::c and linguistic distance. In fact.•.e accompact. It may' also result from frequent language or _-e.bes _elcnclnc' ".'. and from grammatical data sho.. tri. and Aztec-Tanoan. for a subset Of se_en Iino_uistic.'rnian:ies me._.s_ic.. we find greater _ .-_as_...'_ ... This negam.{:::ou: doub_ that A"'*_-d.. 6...".ee:_ ge.. Plains. _•.._e_ . ations are not mutua'. 6.'..'..:.een linguistic and geographic distances.ere classified correctly usina o"-o frequencies.enetic distances among linguistic stocks is sl_n:ficant[..'.ied out by Spuhie: 115791.e. '_oreo_ e.aed in all three cases a significant congruence.e::::i::c t_Q'J2..-_<:eric:'.u ..'..-. Their geographic posit_or.e'.J._ure 6 iCE.n_._'enetic data. s:. (1977)investigated the intra-and intertribal genetic _.. The statis_icaI approach used by Spuhler (step.ariance....fled possible ciou_. '. h_gher than that within hngu_stic stocks This is in line .. and c' _" . . ho.. Southeast) 31 of 53 tribes were correctly classified. "_ es...efic neplacements.on and fusion pauem..IS[O-" of American n= .American . g='_ '!' e._. o . re...:•unaerrain:its on dates The genera'. :he po'-e_) of infc_ation A_ this .... a.e result ca.era! correlation between lin._ "._. Some of the discrepancies.en it.s a_..e seen that o'ene. .. Kraho.: !i.e..._. that Apache and Navajo are not misclassified in Spuhlet's analysis. desk.. :e reduce the co_e'. agreement between genetic and linguistic or cultural classifications.esti_ations of limited romans or _roups of Central and South America An earl.. hn_uistica!'.. note..e _" az_eK_:!.ear.. Nor'thv-est<! .-'m._:... . TEe analysis used a subset of 13 gone frequencies from ABO._.asonabLv '.h A. a:io:: provided b? other "_¢'eroac::es In par:. . _.... Chak. Plateau.. Northeast. They found no correlation. Subasotic.ide good e'.ever." . This indicates a substantial aereemen:_ between .den Amer:nds.. The scale of linguistic distance used may be responsible for the failure.hen there :s .ilence of a s'. Penutian.."Tn.elation can easily be deszr%ed by some out!iers _':.ariation _ithin the Oe-speaking Xavante.onciiing gone:to data with info.o ._ .. The matrices ofgeneticdistance.. He._' ....... '..U]STICS A_.. _ _ .. -_.athem migration When . Using other more con..en" .indicates netic and lin_-.a_a::ce of .PE': sa. In conclusion• there is substantial. The hypothesis that the..:E'z -- 33. fu_her anal>sis gi'.._that the esz:.. the two approaches gave approximately equivalent results... (i97a) compared the linguistic distances among seven Yanomami dialects and genetic distances among the people occupying the corresponding geographic ateas....Iacro-Sicuan. is in _. 3.ken...icana!'.t .. . ls. Na-Dene..v.". _ . satisfactory. COM?AR. ...:...EOGR_. e '.... ?........ which escapes the s:ic:ures of iinear-con'.Nor".. be more of an indictment of the method than of the _er._haeoloo_icai_know ie_=e=. but also a number of discrepancies Moat misciassifications in the Spuh'. m"eser.c ..'a-Done.culated from lexica_ data...a.veE eszab_ished and.'-" ..'... or 58.-. A linear co.' .. flss:.9 v.e' . Spuhier (1972) foundno e_idence of corre'.es. ha'."ezion.ith his result b'. picture seems =e.o'-clear:.-_ ..uis:ics an" _enet_cs..._ show v..._ .mer:aa.e an£ysis of ""* relations be:v."TT.a:iens me. . s.a_..m_..._.os. . CalFomia. M?._ In sect:or. .. . RH: Diego biood _-_rou-_. ..ic three ma- In fi..2) reanaivzed the same data by an anal?sis of '. in'.t . COn_.:' (6. .ni'.... assessment or"t._ -.ere"...:-s:_s an e_. .rong co.. and Esk:rr. no"pe._ '_plete'.er sample are found among Na-Dene..imm t._e.:'... one bv Spin!man et el. ...bi_uous_ posmcn of the southern Na-Dene.'ere teste¢ in the latter case..-_ -..:h :he latter that must ba'._. section 6. (1976) found no linear correlation between genetic distances and linguistic distances in seven Chilean "highland" Andean populations Linguistic distances were calculated on a scale based on an early classification by Greenberg._. The sea'."eiafionships _i:h other Na-Dene and v-ith Amerinds are best expained b? adm. Apart from Spuhters studies of the gene:_cqing-_stic correlation or. ?e.. They conclude that the DOI 02208 .sis :'ui'. ._..m-i Siberia v:a the Ber.. and Cayapo of Brazil...'.n Indian tribes za::.. this is caused 3.... Considering that more _oups '. pro'... Southwest.._-'_ a _o.:TH LiNC. discfiminant anal>sis h is wo_" adding that Spuhler found a moder'ate but s:gni:'Sca." _enetk and _eo_r. groups: Arc'.._ .. and in the Hokan group In Spuhler's analysis by culture areas (Arctic..:. especially that of northern and southern Na-Dene are of interest... R in full'.2%: of 52 tribes tested v. 1". with curo_o-" rent'.rabor_y et el.e::.heir sc.-..et_csand language The .n: correlation _e.xtu:es v.. and foun.. co...-... there ha'. co.entionaI apprca"hes. ._' --'. ..a There _.'n._' .-:'a-. and none bet. MacroAlgonquian and Macro-Siouan.a:es of o_complete . _.. ..oup ". and the _eculiar • _.S [he #['...! n.e tome to cons._'. however. o .... all me.

he whole subcont:nen: occur in the Andean chain today By contrast. interpene:ral:r:g pc:terns. One can see many reasons why this can happen even if there is a general congruence between the two phenomena.. s the results obtained by piotring '."fi'er work on American data with more refined methods is clearly necessary'. as mer.00 v. In summary.. _i:cussed (sec '-cj:.: on'.. Detaile_ s:udie_ of single linguistic groups that ha'.. A simiiar failure ts experienced in 'he related tree g!.e correlation with gonetic distance (-0. Even the usualL'. For further comments see Cavalli-Sforaa et aim 1992."am:lies of the nine spoken in ..een genetic distance and geographic d!s:ar:c_: Mobility is also de:e. in these c]rcumstance. and linguistic cistances it: :his case... In fact. or when there is governmerit pressure to expand those of another language. Studies of o_her groups.reme sou:./18 (67%).rr_rles et al.:up:ed by people v. 1990)--are best suited for showing the correlation betv..f.:ages :_ndgenes There mr:'. cal!v. different from tee east h runs from '.'e but !o'.'ventages The observed correlation _r = 0:'4) Js high and highly significant.e only' a '. not significant]y different from zero) and for geography and linguisfics (r = 0. For instance. It seems likely' ". and historical reasonS..::_ mere _idei.ith geography is especially difficult to study in South America.9). When we lock at figure 6 10.atien of lang.sured by the re!at. and is re:ati'. significant at P = 0."ished by extinctions may alsobe usefu.:Iedb? s:ucy:ng the Cistribc:io:: been published _ce. een geographic and linguistic distances.hat.". with the continuous shrinking of groups. either with lan- DOI 02209 . unsuitable for measuring :he association among geographic. Andean 12.. one may expect this type of analysis to faiI in the Americas.o rela:i'.. tions betv.212--0. :o des:roy linear ccrre!:. it seems reasonable to asaume that in the modem situation. especially using linear correlations Fu.a._eak[ines.'t'.k. significant testing of linear correlations between distances calculated between pairs of populations is unsatisfactory because there is usually an intemal correlation between the pairs. and political changes that have taken and are taking place in South America.s the backbone of the continent and :s vet'-.. it is also possi_ie that :he data are ma.49.a.05). economic. (1977) in which the pairs of populations are independent."_or:ed 'e? the :3ic::::i po'. Linear correlations are especiall'.: The studies of correlation be.he genetic distance between population pairs against their geograp.0¢1) and wi'h geographic distance (-0.emc_ research One sacu]d remember tha_. we are unable to find a simple inter'pro:anion Inking genetics and :inguistics in 'he '_no]e of Central and South America._ge in:enr::_ai generic distance '_]thin this linguistic _oup is about _3ffcas great as that bet'_eentribes speaking more diffe_ntiated languages They' found.. Tucanoan 12/47 (26%). thrce of se_en studies favor the hypothesis of congruence between genetics and linguistics bt:t for methodological.ers _r _. that have not been excessivelyimpove.. traditional ia:':gu::_esfret:: ether grc. :'e c:_-_': impo_am r_asons:ha: c.s highly inapTmpria:e for detecting the correlation of interest. a:_c between genetic and ling..e not undergone too many. theoretical. Linguistic distance be:v. coc:r!_. (i990) on :he Chibchanstx:aking groups of Costa Rica ar. ho'*e'.rm. Ten such populations '*ere analyzed for _8 genetic loci.h_ch isextremely fmgmen'_ec. a'. a '. These results (Minch and Cavaili-Sforza. The genetic distances between pairs of popu!a_'ions were correlated .. as _ve]l as iingui_t_cail._tly b} Ba.een genetic and linguiszic vaniation. eastem pan is more heterogeneous genetically and :inguisficalI'.v.) will need further investigations.reme'.:. Pan of the problem is tied tothe major territorial.::(or cf :he corre'.045 (standard error calculated by' boots:_p) It is positi.o major _ra_.":. Chibchan 25_43 (58%).c distances basedon cognate pe. high gecg_hic mobiii D.'.d Panama.e satisfactoD bu'.c thelinguis:_. Other methods have given positive results when linear correlation failed..:c d:stance (see._m.3.ery pania! answer to the genera: problem.esshov.o are ab.ely homogeneoLs per:err. Americ:.ati'. tee flat:or. 71 languages of the 11"7(61%) that form the Ge-Pano-Carib subfamily are> cal situation also conmbutes to this result: the Ancear: chain fo.he extreme Rome :c the ex'. even if this is generally overlooked.'en for South America in Saizano and Cat!egar:Jacques (1988)._.:. disruptions and extinctions--for exampie. strocg reiat_on between genetic and geographic distance is blurred in South America. v. This does not apply' to the sample by Murillo eta].-is:ic disvances.".a.ely similar eco!eg!cal y in spi_e of :he great _ar{aEion in latitude. spoken. This mig_ be enough.: :X.051).:d'. A ve_' thorough and detailed stud)' ..h genetics and also . een genetics and finguistics in America can gi.and confi . een families showed a negati'..'?_ This "_ouid certain:::. The ecc. 2. It is oc.27) be:'_een genetic dist_ces and ccgnate percentagesin a gist cf '. The correlation ca_:uia'._ps.::e :e :he des:r. v_i:_ subfamilies fc:-n:ixg very compiex. in Ruhien's (1987) list. an increasingly larger proportion of people stop speaking the traditional language and replace it. genera. the Chil:x:han (Barrantes e: a!.ords. higher than that observed for gonetic and geog:-aphic distances (r = 0.eser.332 CHAPTER 6 guages ]r.ed between t'-e two distances is 0 19! = 0. Similar high percentages apply to many other subfamilies of South and Central America: Equatorial 67/145 (45%).::f language gro. an e'_.e:ship be:v. a :roe '. as '_e ha_e a'.52. There are many reasons '*hy _he correlation of linguistics _it. but this does not mean that the people also disappear.e. r correlation (r = 0. causing an epidemic of language extinctions that must have been especially dramatic in the last cenmr'y and earlier.equa:e.. unpubl. Of the seven s_udies we have liste "_ only one that used linear correlation ga'. Languages often become extinct when population numbers become too small.'.'.139 = 0.

to which random . The less frequent allele..'ariafion in gent frequencies affected Amerind populations wiii be cle_. of thetba"ae major postulate£ miT'ations.ould be lower than for white admixtu:z.. More intensive contacts _ith Europeans occurred in this area and.e. contest. if not impossible. and a co_es_onding. I:= the e.. !n America from other pro'is of the world: Ame. a non. to be the case on the :astern coast of Canada. for the anomalous distribution of ABO. and perhaps persistence of lOWnumbers for lcr. It confirms and extends the obser.. It was first found in Amennds.. 1983) showed that O individuals heal more rapidly ([_judged on the basis of immunological tests) after DOIOZZIO .A) is of special significance in America. GEOGE-_FH:C MAPS OF S[NC.e me'. This is !ike_.. bottlenecks.. .l_e (Cavaih-Sfo.. A is conse. The band of low AK]*I (<97%) across the North American continent indicates Caucasoid admixture. to exclude the effects of natural selection. 0 is almost never less than 50%.i:h Caucasoids. and the omnipresence of O..e.. all _. but the absence of 9 in the v..._treme south..! F_ _ : 2 -:. of A.A.L.drif '_ t plated a '. in O Greenlanl is also high in A L4]j...ty of most alleles except a few is observed. Variograms of ABO alleles have long initial linear segmerits.. _:er the returr:.almost com" e . but others ha. despite the high frequent'.. chemothe. . trou_'_:. correspond to the expectauor.s ot the hypomes!s._a [9S£c. x . This remarkable phenomenon is therefo_ uniikely to be due to natural selection._ from a vet"} sm=[_ number of founders. a p_mcutar Amennd tribe has onl) a fev._ .. s and ABO was n._.. and a popular explanation for the loss of A and B alleles among Amennds is differential sensitivity to syphilis.e c!aimed th2t the disease ongtnated in Af. In Eskimos.> a_eles of an HL4 ]ocus common]'. 3:.... _here O is low and there are also .....Lhouch not a _erfeet rain-or. like ABO.4 _..s a rem&rkabl.g sor:-e A!mosa=:... it is not surprising that it is difficult to find "fuit-blood" {or e_en only 3/4 blood) Amennds.. "_:'". Did drift determine the i.. The origin of the hypothesis is the b_lief that syphilis was endemic in Central America in the fifteenth centu.. a marker of Caucasoid admixture.ith a maximum af A greater than 10%..'ations with ABO. here. there is a small patch v. or to the initial founder effect of a small number of first migrants from Asia.'. is ?roof that fi".10. therefore. tribes onginate..'-al other exam_.B) shows an almost regular gradient from north to south... '.-" 6.or!d...'4. the proportion of A to B v. and that man'. and an excess of O (up to 100%) in all theothers... v.2 has a frequency of about 5% among Europeans and is essentially absent in other populations.o frequency distributions are . The . Adenylate kinase ] (AK1) is.nbe the same ra_.rreiauo._ B..eca:. or I.'eU but in a way there is a somewhat sireilar phenomenon: an excess of A in a few groups. constant frequenties in all other would populations--and to some extent a_so in man'. _n:s "' section and suzcests_ the: _ =. high frequen:?. As we have seen in section 2.D system is _markabl'... If Negroids were the donors of ABO genes... _. V'/e have tried to avoid using data from mixed populations but we wilt see that in the eas'e_ part of the United States and Canada a fair number of mixed groups are present. In Norm America O is lov.._or of H/. Instead of the mar.Jsis not due to admixture.ith a[lele 4_ being high and reaching a peak aoo'. Thus..1o_..._s cf ...'.... and shov. of Columbus' crev.. The distribution is almos: bimodal. The evidence from direct studies of patients (Mourant et al.av.enema: disease (McNe._nds are unique in havin. at least in part.elv.e 45% (a]most aI] A]) in _.'o alleles of this svstem.A '.anticipate [ha: the be.hich is not the case ir.' Primates (Socha and Ruffle ig$3)--are not entirely • understood. B shows a peak in eastern and sou:hem. The variogram is approximately linear up tc 4000 miles? with a fairly large slope. Canada.'s are different.. .tbetween . the ABO locus is a faifiv __ood. ._ _-. Even if there is a good chance that drift was responsible..' found e]sev.. The A_. ABO has far fewer alleles than HI.. with rather sma!l slopes. A high frequency of B is almost never found.. The Diego blood group (DI.n.'l ]9"6) A search for a co.tO' 1o_ humher of inir:a mi_m.g periods? 'A.eq: important role in Amenca.nts_ (an initia! founder effect). which _ present at re!ati. given its magnitude. but the frequent a leie.. AK]. a]!e!es a: a dispmpc_ionate:. . Acid phosphatase(ACPI. of ..ester'nCanada E. because O individuals are more resistant. even Jnsma'.cafrom a ciosei.l populations. re!z:ed spirochete responsible for yaws.edamong Na D...a.'.4 Apa:'t from Eskimos..... ABO phenoUpes (or genotypes) react differentially to many infectious diseases... whereas among Eskimos.S Ti_e dates and _e__ph'..'-'. B'.estem pa_ of Canada. The variogram is uninformative and is not reported..4..' be true. amor.from se. .. _ith other alleles rare or absent. v. The geographic distribution of the ABO alle!es sho_= in the maps deserve some comments Because of the ram'.LE GENES wifiere . higk frequenc). the A and B fn:quencies are much more similar :o those of the rest of :he v.c. the simultaneous presence of A and B in proportions of 4:1 is a strong indication of admixture '.'y and was spread to Europe by the crew of Christopher Columbus after their return to Spain._."reTaiant:es of gent frequencies in America because of a '. it is difficult. . Canada. reflecting the major difference of Eskimos and Amerinds from the extreme north versus the rest of the continent.se_ .. of the European epidemic beg:=ning shot-:. _'°' ? " lost the A and B a_ietes.. The reasonsfor the loss of one or tv..'.. loci indicafes :ha: the seconc or third h':'po:hesis rra'.."_DeUtl_. In another t.

'-a:e " _" frecuenc_ .. nor'them Brazii and the Guianas.: : 'cs_ bu t _l.. dish slope.. most probably resulting from drift.est: the variog.. B.. ac:. :?re . Haptoglobin {HP*]) also has a ver? wide distribution.o_ .: e:: tO high prereprcdac:ive mo.hereas that of .. ..S. A. GC-IF has two peaks on the western coast :_ Souti_ America.. u. This restricted range of polymerphism is expected . HL4B-5 has a se:.e 10ec in the extmme Southwest of the United States._ e__:era Nc_'.%. cn :s :%e to differen::al mor:ah:'. HL&4-_ has a. .kh a ]at_ . less extensive'.ariogram is approx.fion seems bm_oc. but as already noted. Basically...'.22._. but there are other secondary peaks in South America: the lowest '. . like the other genetic s. The most free'_..America has frequencies _e:o.334 CH'ia With ar a_erage frequency of i0%. with gone frequencies ranging from 0% to 100%. .vre-._. thou<n _._zii! and _he Paraguay' basin..e.nere. .ent.mJ. }e e:.hen the genetic diversity of an ancestral population has been reduced several times by passage through size bottlenecks.s a minimum in centra[ Brazil and a re!afi'.o'amg a maximum in the Arctic. B'27. a relative minimum in the ext. "._..aJ. .es The Du::'.5Q and a peak of mere than 40 c'..ei!as in Central. .. In sum. possibi) because of the closeness of the mi.' of 17.trephoretically fast subtype of GC.4___. a'.. ?:as an a.Although its mean frequency _s 1%. _eS. al!e!e . "i2negroup-specific component or _i:amin-D-binding 7rotein a2eie ] (GC-/) sho'>. as :he map sho'*s. _' .e A.4 is faMy regular.. .n!factuaI and caused b': :he near absence cf 4ata :n Nenl'.-_ ..e Pima. small--ira the Old more than -.'_ whereas :he rest 0f South .. has a 20% average and reaches about 70% in Brazil.. The gistr:_:_. x_-. It must tl':e_._. slope of _he vat:og. A*33.9 has a strc:':_!'.2J.. B*I6. Papago and Zuni.-. s_'. :: ca-.. no_he_ 13raz!2. Averaging 0. has a maximum abo'. with a maximum above 20% in Alaska._ 20%..27. A-23 a_erages }. With a mean frequency near 19%. Antigens specified by HL4 genes have revealed an unusually narrow range of alleles. Indians had been 2_ "_ '. B...h is a.. America except it.. B*17.sterns. . ." rela::. or homoz'._... its multiallelic structure renders variation more evident. Only HLAA.'!'...:$%.27 has an average frequency of 3.'.aaes considerably with aile. eastern Greenland and the nor':hv. Allele B has been studied mac. Well known for its strong association with ankylosing spondylitis.":.h' . _" [f..a.era¢'es_ :j:"c..:19S: _.. _C7 B*]6..': it peaks '*ith more tt'.'..e. The variogram has a relatively short initial T_rtion with a positive slope. .e_ge...Argentina Aga:n. motes '_". have similar origins but signi_candy different frequencies for B.... ... . in the present case.a_ _.e_" '. especially in South America (Black et al. .a.. G!vo\'.'c.._. .:mumand maximum.'enezue'.' sho'*s a maximum in Mate Grcssc southern B.n_ """ '-._e of A]_.:-il has a maximum in Cen':-ai Amer:ca and :ox_ . The dismbution spans almost the compiete range. h mm:ma :a ::'e northern Andes and in easter= Greenland."z..m United S_a'es. fi'...5%. . -'" of more :hun 4Q in the sou:heas:e. The distribution see.-.' due to dr. there is a north-south gradient.-... average I 1.. st:died oni? Jn :he fe_ popL!atJc_s s::22 so.-_-J'.PTER 6 Ame..aries fro=: less :ha:-.."eme south and one in the extreme north. A possible effect of selection should also be considered for HLA: in fact. ofBrazJ? The'. A.a'..ase-i ._... .. ' initiaI slope Ai!e.hemChile..'_ average frequency around 3 [Q '. reac_".6cie. This genetic s. HL-L_..soiutemini_U'm '" .America: italso sho_s an ab.o more than -q_% • it is also found in some no_. '*he foun4 that.._es in South America: '_ "co :at _ecrease toward the nor:. " I _' _ ] i " " = in .. k=C: 1'_ =I there 'acre 56% fen.8% a'. and all alleles tend to ha'.ular.7%.anog.5%...rr.":?: 15.e . _."_ ._een 40% and i00%. A subt'..."s-:? (F. at a ]o...aiues are in the extreme north.... but is concent..28.e .'.... A. With an . The '_ar_zL'-a:7_ of allele ._m .era_'. blood g.AiMe _modal. ondar?' peak (o_er 50%1 is found in the nOIX.._... with lev.'-r" l_ -._.4-28 has a xak':ear 40Q i= :he extreme south...crdar': peak in eastern Greenland..].'-.aer f:'equen%.rctic Ocean region. _ith a 0. B." '. America and in \e. as v..-4 ..". no. b_.xr_:zk..L 2..e extra.. cannot be attributed to climate.ft..:a :5 "" _ 122 peo_Ze. "_..a!!e!e is . has a peak in the nomh-central -%ndes greater :hun 50% and minor peaks e!se'. B. B..e at leas t t_o modes. averagmg DO102211 . A* -'.e i0% in the v."E_ .hese parents ' Ha. !n a su_p..'age Ia. .ith a peak o\er S0c'c _.. of 12% and a _eak o'...75.'-:5 :'.":e inina.s )'.4C... has a frequency greater than lOQ in the southern Andes: and B. The peak is m the extreme south..n=:.25 has a :e. a\e." _.sten_' is ordinarily represented by a great number of alleles in almost every population--even if very. HLAB*5. B-7 reaches values abo' north..'-.anti'. HLA shows great variation..4-2-37Q.':'.'a 'o'_ " Ns.'nca..e .. reaches more than 10% among cen-ral Eskimos.t distr. :he .hem ... ann the linear :onion. 1980).. estern Ar::ic A sec_.n _.. -average--and it reaches maxtma :_er 50% in sc? 40% f_quenc.o:e and is knear .ate!': ::::e:: for almost 2000 miles v.-=. _ Ono. evidence for heterosis in South States.m--:= .._m !s :_eo.2... ._C .9.8_c.'-:heas: Asia Its considerable . and B*40 have average frequencies significantly different from zero. 50% in eastern '.oc.'. B.a. has a peak in ncrthern Chile. is iess than 10CC .. a_lele J (GLO. The elec. B*I5.ste...35. _ith mean 0. v. 5% '.'-am is . A*3I.a_ .e_ _ 2._ be . The most frequent B allele.0% in no..T a. The '.o_s a complex form. a_out ihO0 miles. but it is rm*e or absent in Nc:-'_: Ame=ca.: a :...ared be:'. cf Co.-V.:tion is likely :o ha. The ma'._ e. which. with a mean of ._.a_=a::cn !n Ame:hca is most prcba_i'.'e maximum far:her '. has a max:mum above IEQ in the southeaste. Allele B-14. frequencies nor':.. z_.a._:e.. reaches over 50% among Eskimos of the western Canndian Antic. a :_--_. It is interesting to note that the three tribes of the Southwest...35..' of es:erase D (£SD."a'. The variogram si'.

.bOb:'b3b. za:g. but with oscillations.ariaion than other genes."y'strong founder effect at the passage from Sibe:ia :o America ..etal negati'. in ore or a fe'a tribes. f.!e!e is or-. and the variogram incnases initially. a'./K.5 only maintain tv....e 5)%. minimz in the north.e 20% on the nonhero coast of South America.ariograms of immunog'. en sta:":e_ by such smal! n_mbers of individuals that the'. among the founders perhaps but mos" in the process of evolution of ind'.ere lost.e irregulm variograms. in the total absence of cross-migrat!on.A) has a distribution of 0% to $0%. However. In :he ._ha" 'a.. has an average ff'equenc_ of 6% and peaks at mo_ {nan 20% it: Alaska and in has a maximum in "he center of South America and decreases almost regularly around it.le'. perhaps more impc:':an: :h2: the nrst. The M. It is therefore likely that many GM gone.e been present at the beginning and :os: ia:er. a re.e. but she'as a peak abo. The initial iinear segments of those with positive slopes are in the usual range.!eEebe:oning the sole fir._. but mostly among Eskimos..b5 . All the variograms have positive initial increases with regular slopes. za. it shov. ivor equals _. at least qualitative]'. t' may ha'. is a rare pcLvmorphism almost homogeneously nea_.)'.e:'c:e_.2xima and minima for GM haploDpe frequencies are due to drift.e suffieientl'. but has a much smaller range of variation Almost all these blood groups ha. infectious diseases.e -_fnc:r aCmixture KEL.f_:_. but not in aH the other are cormsponding[? . arid various maxima. ".b'f. under drift alone. as in other HLA data. but the range of gone frequencies is not small._: but most popu!ations an unique in that the domina. and maxima around Panama The '.]sa is also reiative!y rare (2% average). are still present in some.e or fiat initial slopes: . The Kell blood group (KEL. except for Ns which is fairly flat. a:'.!-:E "i: S _..a!ues abo. All the other GM haplotypes have lower average fro.levated.s :'roeab'.. _r there '.frequency peaks or troughs in America are due to dr:ft. only the from 10% to 100%. drift v. KM-(] . a'.ou]d 0e extorted." these conditions...age of 16%. se.obulins tend to be i. The FST is ..emge ft'equency of 2.e!es differ from one to the other. and then is a little evidence for it as discussed earher. has a distribution varying from 5% to 100%. cou'. Its comp'. ::]e!es become retire'el? dominant in freq. the si:ua: on is diffe_m One or re'.'VS system shows somewhat less '.'aries from . The sur.y for all markers...:or even earlier) The remarkable '. The P] blood group. is blood group LE.':t a:'. none too pronounce! A Caucasoid hap:oLvpe._ fac:...ould e.4 . because [he ma_om? of reagents are of Caucasoid origin and Co not necessariL'.. and the initial linear portion may scmetimes span 2000 miles. but both geographic maps are full of relative minima and maxima that span almost the whole range.bOb]b3bab5. man'.ing .!e'. or in new ones.An Orie-:a: hap/oL'. DOI02212 .. ore ha!f ff'. and the other a'. the very different local porte:ms of each allele It seems as if most local populations '. At first. bur one does not need to ccnc!ude that half of the alleles '..nmany places. high.:. but peaks al more than 6% in the Guianas -:3S '. drift is expected to operate with the same intensi'..he initial frequency of tha t allele in the drifting popu!ation.idua :ribes as shov. Thus za. Perhaps most alleles were npresented at the beginning in Northeast Asia. at]ve.d has a maximal in Alaska..E:sis exacti:. I.c. ©n!) 17 alleles ha'. has a mean of 32%. fa. Of the four haplotypes.ations. judging by FST values. man'._.. does not have a distribution extending from nearly 09 to nearly 100%. The most common haplotype. has an z'. maxima and minima appear in regions already showing strong drift for other alleles. one does no: need to postuime a _. detect all alleles present in other popu a:ions The va.a.7% and pe±<s in Green:an8 and :n the nc:":_'. Some alleles _. It :s possible :hat there exist severa! undetected alleles.e that ai: these . with minima in the extreme south and in the Panama region.iqng a'.esat high fmquen<..ementar?' al:ele..bOb/b3b4b5. has an average frequency near 2%. Ns (6% average frequency). s a complementar? maximum in Panama The Let.a_ sur. The light immu noglobu:in constant chain. but all show usually single. sometimes extreme peaks in different regions._:'eg': !at and ".L". '. allele 1.ere probably lost.. one might be re!uctant Io be!ie. a'._:bGb5.. sometimes e_en :n neighbonng popu'.*. sometimes reach. One might hyTothesize that this immunog!obulin marker reacts to Iota:. quencies. like the coast of southern Brazil.erage freqJe:'c:es {ogenerate maps of America: this is at..5 of 17. with several peaks and several minima. like the north-centralAndes or the Arctic. with a maximum in southern Chile and minima i.aries greatl. a Nogroid haplob pe (re D poorly represented in the maps for reasons of reagent availability).a=:ous peaks in the north and a firs: one In other . The next most important haplotype.'. with a wide distribution of 0% to 80%. The M allele :'ames from 30% to 100%. n b'.Americas..-. subject to tire rvie :ha: the probability of an a'.o or :h_e aHe_. . The Kicd group (. J'K*B is poorly studied directly. 2).riation with distance sho_vs here.osen :-'andom:y from among those origina!iy F_sent.'a:-:a:i::: a:"'or'g' :he in'hart tribes of South Amenca suggests:heex:s:enceofa]aterbottleneck. LE-Lefa-) has a maximum in a neighbonr:g region. GM (or IGHGJG3) also shows considerabie local vaila'tion. bur at least one in the so.em pan of South America. "E..entuaily lead :o the surv'ival of only one allele in each popula'_on.¢0% to 100%. and the S allele from 0% to more than 80%: both frequency distributions are probably unimodal.e number ofEuropean alleies. With an a_e.

e_.tin_-= around the ecuator.e allele D reache_ robfive_y high frequencies.A being the most _ariable (0. extreme geneuc _ar:abii:t. e: throughout America.b."ams of RH _ho_ '. The most frequent RH haplot)pes are CDe (52._zi] seems '.%ue . Alleles C and . ha'.16).-'_ ...e. 6. ".007." S)stem is also _'_'_ . including NoDone but not Eskimos.' less exposed to ' "q... followed by A (0. and the second. .18). 2...• _-...phospr..':'%abie. who eat essen. and _£".en the high drift -'b_--e-. not surprisingly.aries from _I7 Q :: 100% in frequency of :he 5e a:]e:e a_d has a ca.. PGM2 is less well known ard..or!d: it '. and Labrador.s. up to about 30%._ are .( is found in the NaDone _g:. are .Se (0.ariog_m of PGM2 is uninformative.5% and can be taken as a good indicator of Caucasoid admixture. Cde shows very low maxima in Vex- America CaucasoiC (no exclusions) sub-Saharan ...C peaks in Panama and is lowest in the Arctic: D is universally high ever... The major conclus{on is that the . an. .he high Mexican values.hich :s exu-aTc'. however.arie_.. the maximum in _or!da. on the avera£e.. Th:s is a!so shown by a'.. reflecting Caucasoid admixture.7'a ¢n )d cDE (36%). ". Haplotype odE surpasses 3% in a small area of Mexico and reaches 1%-2% in the extreme south of South America.T (0. and both span almost the entire 09¢-100% range. E peaks in the Arctic and in the Andes: it is minimal in Panama. in the An:tic.13 and' :c r example.) of ailele B..4] (0. h... who cluster with Eskimos): the most variable genes are FY. The impression from the geographic maps and distributions of gone frequencies is thus full'._v-rile D is less vanable.mer_cas... since Europeans have the highest v.e. but a seconca O. which a]so has several relative maxima in North and South America.=:e= from '.. Two rare haplotypes.adous min. troughs of the co=mort alle!e C. FSTS range frc7 90"1 (Southwest .. ne '..: anal)sis Below '. '... Chile.e _:?t'.n£er It is not clear ._n. 30% and is almost usual. nor:hem._ ncy.ak_'... large_on 'he..12). HLAB*35 (0. CdeandcdE.dase A tPE?.Asia. anc the maximum m :he ext_me south. pea..'Z. The gone with the highest variation is SE.%-I_% for allele 1.Ooid subs::mces containing plat.' confirmed. . for tasters in an. Phespheglucomutase 1 (PGM]*]) vanes from 55% to 100% with a mean of 81 5 for aile!e ]: the maximum is in Venezae2a..hem Venezuela."._e compare the American a'._ 96% .051 = 0. So=:h America has the greatest '.. it shows a peak up to 20% on the eastern coast of North America..s less variation.. the extreme North has the greatest variation: 0. avera_es of '. '.._.ea£_emen.'m.. Th:s geographic distdbu• tlcn' :s :n sot'r.__ Jsa (0.. North and Central America combined.13).erage F_r (0. in an) case. and 0 (0. Chibchan sho'._ I_ OO=3: '3 0Cl_ 0 0]} = 0 0. TastertPTC'T) is poorly known in this pan of the v. 7"he R. shov.034 = 0. After Sooth America.e..eric groups or rezions of interest: ]aSS ' .. with DI.. T'se . on the average. A minimum is in the extreme south "Fqe '. . DOi02213 . The ode haplotype is.. where the akemati'.Africa Ausu"a!ia New Guinea O.r=e: by data and "_e unlikel) to represent real . Aleut• all Na-Dene. and finally TF. where we have seen other signs of admixture.C (0. and TF.e . Ordinarily eDe is a good marker of Negroid admixture. and slo_es.10). shou.America Minima a_ among Eskimos.... and ode (a. which peaks in the South'a. The most variable gone is ABO.ere culated for the _91 populations selette_ for de:aLe...orld frequencies of uhe d £lele [RE-]). probab_? because _f the gremer number c'.area *he'..¢.004). _e"r the da.:mitred.0 C19 : 3 3(1)0 039 : 0 0C- Polynesia C <': I = 0 0C4 " "In the various regions of .":of North ..: =a:a.e a parti. and also the Chukchi.I11. PGD*C (0. being confined to 8C'.a.e_. .aric_. '.:na. '-am is u. The first peaks in Panama." frequency of or.._s a variation comparable to that of South America as a whole: 0. RH._.:ularly frequent occurn:nce of edibie plants dangerous for '_h':ro_dfunction.on. a'.n: a]]eie 2 has an a'. show minor variations.. seems very unlikely in the Southwest. v. .. and B has the complementao' pat:era.. advantage._..a.'_ . Pans of the map are re: s::FF.17). PCT....236 CHAPTER 5 ico :._ge .17). Next in frequency are CDE (a%)._::-aT. _" the whole ran_.e_ .a.'drogenase (?GD) shows a .la . It is oncertain if the relative maximum in the extreme nonh'_est of Canada should also be interpreted as a result of Caucasoid admixture... followed by K._othe extent thai Eskimos..-.006. a-":_sh_.e than most :tLeAmer:c:': alleles.12). E ssan. because the other possible markers do notconfirrnit..e ' and is omitted..c 1.h some anomalies in northeaste_ Ncr':h Amerz=a and :r.. There are '.. '. Of the various linguistic groups.rid in the South_es: of the L'ni_ed States T_'.. Transfemn (TF) sho_s a fev.6%)..ess e:. The variognam has a mcderate slope. ogiucona:e deh..ariation of gone frequencies: the average FST is 0._ge T_e secmtcr ' and an irregular va. ".sn. which.cE '. South America..I'.007 (including Eskimo.America ( may be aomrr. the areas with highest frequencies of tasters ha.' 06%. and KM*(I &]. _ith maxima in southem Ch:!e and the southwestern'age FST a.035 (Southeast Asia) Of the various subdivisions of the Americas..26). The minimum .. no. up to 30C'c: in Panama.X. LE*Le (0 21).:re:ue oscil]at:ons a--oundthe cur'. D (0. at least .. here exceFt for minima on the eastern coast of North ..O'O : 0.?ported and is not surprising gi'..Asia) to 0. _ -_--_ ..?.4) is pocri? studied and sho_s litt!e va-":atk... Allele C is represented on the map.059 : 0.13)...2) (0.11 )..cDE and CDe next (0.059=0.19).1_.:0). has a comparatively low a. _ _ " PeTu. A (0.nogram._ B.est of North America.e![ s:.

_'_.'z!ng DNA markers makes • it easier to identify specific mutants and may L ne.-.._e ' '-.9 ethnic heterogeneity--that is. which to the former than to the latter.!': mapa listed in the Table of Genetic Slaps 'ai:h the adaccepted (see sec..-.lbOsrb3bh. a eastern area .t3.p us tc folio'. azd :: :s :he.".s considerable loAmerinds. The presence of im ..* -o .. on the basis of the There is little variation in South America. and the correlations of this PC with the gone frePrir_ipal %olTotal Ptit_pal %oITo_I quencies shown in table 6.'pe_' _) ..'o_" :echnioue..." '".altos ¢'i_e_ m . that ...e.'ere pooled v.oo'_o'. 6. PCs.on_.7 nate genetic variation in the Americas. )_re " . The F values india'.. " ^" -.ain. er'al geog. excluded from the 491 selected as genetic references and..o_e frecuencies..anation The so. genes indicatec above..-.2).een Eskimos and their order of importance..-...mat_on e_'_..'.e" _ -_..-S.hh neighbors The data frorn the 49I popuia:ior:s are the basis for the FST vai- ues gr..1) shows a north-south gradicontact between Caucasoids and Amerinds has been ent wilh the greatest slope in Canada. the most variable pc. seem .ed in North America tinents.a. with Na-Dene showing more similarities curs clearl'eonly for the first two synthetic maps. the _s'r_:t:o:7-f..Ia." anticipate... _=.1. and the n'amber of repetitions of each... ersa'. ". Ticuna. to the 69 _. Caucasoid admixture..7 6 4.B 5 $._.sis.entv-two...' .0 also tends to alter the meaning of the correlations of a DO102214 .. B': con:nst.-a v.c According to some archaeologica! dates.. are obtained from the anginal gene frequencies..omlaoe_nt V_an_ Com_. in other cant:non:s..mms.butions g!'.. Then= is an inconsistency between the observations T_bl_6. Table 6... The peak in the correspond closely' to the first two fissions in the genetic e_tem part of North America most likely represents tree..nc_s a=oundthe mt"_ ..-' .._al marker ra_ly similar. '. AK/-:'. more imFortant]v.'C _'.._m comer of the gene-frequency dist:." ' . In America we find this ocnon-Eskimos... popua Ion: An mc.f_nt V_i_r_ discrepancy is believed to be the existence of inordit a2.'been mace in some mtDNA papers... This area has ABO. . is gone:call. but e'. o_'.. than those above.. the distinction between the Eskimos -. Anal'. relatively high AK]*2...... ...Asa .132 sho'...'nes shov. 6.-" genes used for . strongly indicating Caucasoid admixture. there the stron£..fiationof RFLPs _.s oldest pan.._... net '._hbors deceases ..'. bOb/b3b4bS and high RH*cde."-ST values < " "._iand Am:rind Fopu!ations closer to Eskimos or.T oni'.'-a_ment-length pol}mo_h_sms _RFLFs _+"three -n_. ..-. the..."r.'. 'aith different genes. Pat:e.e:...__:u.a. of sin<e_ :....._.a 'I EF. Infe_nces about the number of migrants to America tha.v{th:...._°:..-ms found for different genes are axis _s w]tn /GHGJG3= :n. _ These os:illafions tend to be lov..e.. which are drawn directly from the gone-frequency maps. abe..'a. east-west difference is always noticeable._'o ma¢s _'e "ha ]:_'zer than ..d o_ .o.5....._3% of . the one aticn among .e PooIi:Tg ne:.e...'ere Most of the d:.__. -_<" a. _.. fission..ir4 higher reso:ut or. -' ¢..Asian origin.n _. South America.."res onc r.r. '.h_ch c"_ Cotrg)o_ent_ (_fAmsncamGone Fr_luenci. and showed a _.Tofirs: co_elatic.'mf::. .. '. v.' approach that leads to the same :onclusions is the stud} af mitochondna] DNA..ergence is betv. GC.2) is obser.. there is a dff. were studied (V. r--s : _'-"_-'°'_ from :he 49. b? PCs and America the major di'.a.. _. To note: the highest correlation of the first PC cai '. The reason for this _...'-. iocal _.'adat_on...ergence found in the map of the secobse_ed r. In North the general shape of s'entb. which tends to 2 12.ted in the top ri. there America..2. 19_5: Schu..d .o.and F. _...en _'_ ea=h geo_nFhic map az_ FSTva ' In Sour ex_'..h hi£.13. specific migrations more closely....h]ch separates Eskimos and Na-Dene from a!: The ana!vsis.lace et ai.."' ec ' _ario£mm curves. 33": are extreme cscil!at:cns of mean #'STvalues a: various _eographJcd s. one could easi'. They' therefore include populations that have be°-..13.of .ortant 3 86 7 3.o_._at_onbetv.'ith a lov.=a not su_rsln£ that th o FST '.l densities of obser..Na-Dene group IGHG]G3*f.. V'. ns of the first s:x PCs .13 SYNTHET:C MA_S OF AMERIC_ Table 6...'.._. this is the area in which The first PC (fig._ of thedata .I3.13. ._o_ .' and PC (fig. _oints sho_n Jn . though the repeated patterns.'a'. America. co..i shov-s the par'thion of the :otai '. 7..... so. ".B cover other local regularities.Trio patterns of single-gone frequencies '.:.i 1990". P_t_mt_ge olTot_lV_e Exl:_ir_d by_"_First on the frequencies of the Caucasoid markers just indiSovemPrmci0. '..s. even with techniques ai'./F. : side. of :he'._. of Eskimos in the North-( 6.maya:sa be the ditian of ABO-.-_.. thus emphasizing longest.ledge 6.hi. the. unw__-ranted at this stage of our knov..e_ frequencies.a.h_ total .....he anal'.l_u a.B._s Piton.*. for o"'es :._hh :_.. There is a good correspondence ..42.or'._ .'-a % impor':an: flu_uatior.._ 19901. .ov...etic maps obtained._. and the rest of America on the other s. _"" uniike those gh'en . In those C.

DOI 022 _ .

the Vikings died of star'.4 Synlh_fic map ofthe America_ obtainb'.ea_orion between east and 'a.m:a. . ..e • ' --"_"d--_ _ \ l:!h :!fl:_:hl[ /.13.ed m the nor. '_. espec!:.": more similar to our single-gone maps.= .'._ using the third principal component. occupying the south_este. and their maps show Iess regul_ patterns than ours being some_. ed in a speci_: _icn Cen:_l America i_. ..g in South Ame. . seem less sensiti'.. eas:e.*i:b.: in contrast to the third PC. Thus.dien'.e had some Caucasoid admi'..o compone.*een the Aieutian islanders and the Yupik Eskimos..ith Africans--is like% :o ha_e t_em ?'.'-:h America. .ith the Eskimos of nor:.mericathan to South Ame:"-:a.ieJal gone frequer.c:es cbser. a_d in :Le same direction as in Sour: America..h-central Canada. ed using the fourt pri h ncip componen al t. ..". ever. the fission between Na-Dene and Esk me. coast of the United States ha'. the directicn of the g:-adien: is inver:. b.. b_t also '..It and south.r'nr: :n tr'e degree . hob.i'."nG_en!o_ndand the easte. and Salzano ant' Cal!egafidacques (1988i in South America..5) st_sses the differonce between the Panama area and the rest of America It is also indicative of migration to the south via Panama The sixth map (not given) sho_s ve_ [ittie variation except in .'. where it emphasizes the contrast bet. p_.".ca Caucasoid admixture is also probably found among Greenland Eskimos. on the eastern coast in the ninth to fourteenth centur:es _.". . 1985) and South America (O'Rcurke and Sumoez i986). Both groups have found e_idence of strong genetic drift in South America as v. .aa ne'.hAmerica and in SouL-. DOI022_6 ." .he extreme north.. Fig.d con:n'aution: some further c!ari.. 2. _u: there ma? have been genetic exchange.I.ffe_nt behavior of the tv. both _. ho were in contact . asoJd acmixture which..:£_ _ k 32'9 PC _vi:h inL.13. but the d.*ith Vikings. 6..'.s of admixture with to this problem comes from the ne'er PC._of North America. the Eskimos (their fate v. Other authors have used the synthetic map approach in America..ace :n eastern Venezue'.'!..'. Our synthetic maps.e have..3 S)nthelic map ofthe Americ_ obtained b.'._t. J... .orion or . 6.3) are foundespecial'..of shading of the three area_ trot max have had some Caucaso. butis n_t missir._PCs adds some evidence to the h.nNDr':.. An adm:x:um of another nature--that is.Ame. The sim_!_'-: b of the third and gr_ .h.a and:noGuianas The four'tn PC fig 6 13a) a!so has a ". _. Extreme values for :he third PC (fig 6.e.":. this map shows approximate correspondence .ture..:s in Guiana may stxngthen the hypothes:. the contest being remarkably strong _ev_e_r the nor':.. O'Rcurke et al in both North (O'Rourke e_ al I986: Suarez etal..i3..m.more similar :o . /. par:.'.. 2.'s some .ere kilied b.13.. _t _s not surprising that one finds some -:imi.h. . . E_entualf?..'-r. fn South America. as we have seen when discuss{ng single genes.heastem and the sour:hem Andes Noah America also sno'.'_. 4..'.pothesis that both. If this _s :r. v.'or:k . 6.'.a.. is especially prominent in the ease-central a=ea_" N:..A_.ans in this region. The fifth component (fig.u. of c!=":fied:._.].:ha" between South America and the _s: of the Amercas It also higK!ghts Caucasoid adrrix:ur_ o5 the eastern pa. It is possible that the east-west gndients observed in the north and in the south again express Caur.

ico. but it did expand from the Andes toward the east.. < ._:_. .e co!or picture does not suppl? a clear distinction be:v.. -l-he fou_fi PC emphasizes :he ur. The difference between the western and eastern coasts of North America is clear. their no_h" south geographical order.kfr:c:.....'encies of selected groups or the slightly different m. and norther_ Mexico is a sort._' .as_rs'r. T. and Eskimo."z of North America is ___\ ... Brazil..:': affez.. an indication of a possible in.e frez.'. . in Central America as we_i. T"-':. Our method obtains firs: -':'-< :'. and a southern one cor:'.._ be:ause :he I::ter inhabit a ve_._.a zones..ere ::::: the direct calculation of PCs from otisinal ger.::. of average fro':: . as expected of a region that ." Differences _98_ m methodsinexitabiyhighIight one aspect or ancfiqer: our synthetic maps are aimed at gettins £eneral s_mfiarit_es.i_h the sarre nuance.P. Our single-sere maps are more useful than our synthetic maps for seeing highly localized effects of daft..:th the northern ones mcre s_r':L_.e.. South America is dominated by t'...e]y stained: .here are "re a:originals left.ompc_-':: Amerinds.:"o_e penes and proceeds from them to obta:n PCs and :hen .:athere being Na-Dene the regior_al E '" ' C -i '" ° .: a centra! one formed b'.F'" / // / / z."_. where languages of Tucanoan.3-3 S -: -".:_'_r':: . Na-Dene.e. Biueextends to the noah and northe:st and must represent a dominant direction of migration. '..Me:c.: b_.eer: these and ]nuit rEskimos/.. e..9% c: and ye!iov..'e:se Thor Heyerdahl (1950)effect. and Ge stocks am spoken preferentially.'ene.speakgreen ors and _he __reen areas most'. Caribbean.The area ecc..::-<. e.n .. ''' i i" [ _.. -_. mostly into the Amazon plains. .e:a . rio..e sou..2-: __:r:r:et:. Th:s :ends .An:des near Bolivia and Peru.?-. and 'he pink area at the bc'undar? between southern Arizona. t!l_" . _. :. in eastern South America..upied b? Chibchan speakers is re!ati\ neither of which is found in Nonh Ame_ca n -. times bv many __rou_s. No-Done. _kr_.ho also ha'.... _.-*_.: ma_ cl :he _mer can eCtained b:. _. and Almosan are weI1 characterized and are even further differentiated into subgroups. the Gdir_nas(see. . 14.method used b. 61.enessof southe. DOIO_lT_. though not at the same intensit).i:hin. The Caucasoid infiltration in the eastern United States... There are important e:olog!:% Oifferencesamen: ithese areas. "_ Fig. In the eastern par:. In South America._'_ . tune '.. probab!? _. ..Nov.. hon'o_eneeus The Caribbeans are passi'..m'_..ations: southern No-Done {Apa:._o colors• rec and blue." _-" '""" _*_ " _ _ '" _' .o smooth maps .a_e The genetic patterns in the Americas fully confirm the three waves of migration suggested by dental and linguistic e_idence: Amerinds.._h '.. the yNorth eliox_ greater '.'.one .ln an.) crossed man'. 7"he conclusions from synthet.-... northern Amerind T?. O'Rourke and Suarez •..nor::'.' "'_ _ / -• \ '" • x. Eskimos. hi'. or '....he a::c .. ous local pop-'.-_ . The other dominant migration in bright red is found in the southern direction along the Andes. Further refinements map re'_e_'.a. thin area on :he coast.'. v... em B_zu. ._s.her and thanthere between.... the arrival of Polynesians to South America? 6..e '_::: prate:Fat c.. [n¢icat[n£ that there are some remnants of the passage across the funnel noah of a nonhero region fom:eg by easte.h.heir maps.!o_o _ica[ exchan_ r_£:cr% The color map of the Americas conve)s 6-:...o dnf: fi':::n _o ird_<Cua! genes.:mrc s::n.e ._.. .ah'. that more than one entry contributed to the first v.". SUMMARY OF THE GENETIC HISTORY OF AMERICA Their order in time is strongly suggested b.F__ differ from me eas:ern pan of gec:: -_merica Tee c. '.'-rC .c maps reinforce previous findings and help visualize major genetic regions. .' v. usm_ ::....e some £ene'.espondh:g to so. the third PC'.... as we have seen from archaeology. probabi. em 7a... . Is the white spot in the middle of the . several regions can be defined: the Andes show local homogene_t? at the level of the higher PCs and always Central America sho'_s a compZicated mosaic cf colors.:an guages ._q. kme2ca. and neigfibonng speakersef Uto-Az:e. and perhaps in Greenland are clear.. gu.c admit.r:c Cent_.\'a'.e: --. colors appear..

Amennds shov.. al. ._ctc.. with Equatorial.. . ' ' North America /F'. .. also than in Noah .Dene and A..America. several poor or negative correlations among genetics. in which order they entered.s: the •Andes. Moreover. ever..hem an.sand fusions. _'_extreme..ation_s c".'.oria_ Ge These conside_fions could hase more v. alternating with Paezan and. and the sou:hem plateau They are '.. -o _ . .._i_.s:'. time.." _. a_ found in South.._._su_... there :s a band across the continent. • se::'.. It would be interesting to know whether some of the South American linguistic families existed before the passage through Panama and..hicn is found _.. because of :sntzc: .orld.reen!ard arid e.met:ads _s incomp!e_e._ ._as lmpossib. .. southern Na-Dene is genetically clear-cut and probably reflects gene flow from other Amennds.._-c _ab.'as suspected. using distances based on gone froquencies. perhaps carried out with other methods. however. so that . must ha'. one can use synthetic maps to distino_uish.'_ ._.:_.. The eenetic picture .'.. ':..n_. _anoa_ South leslie:) Paezar" An_ean ghibchSr East Car_ Tucanoar E_ua'._" _"_'--'..ithin..*een linguistics ar_agenetics in South Ame-ica.. geography...'m".. and our unders:ancL:'.:na'. v.... _'_'. _ pog. Clearly. e'. who had greater oppor:unities to recei'. .. much of the gonetc variation in North . ' iR!er:or In t_' u_ _ -_the exueme east '... -'os _-" [r. and some times even more informative than work at a macrogeographic level DOI 02218 . An important testimony to the extensive movemeats of Amerind tribes is the extreme fragmentation of the linguistic map..'.'a different ecalogicaI'...' van• m. steak.a:b.on_ _>so . The Andean family is found along the Andes.'_¢ :. using mtDNA--can be more useful.. reasonable to assume that some of that population '. of Caucasoid adm \ture..:.asa good co.. is spoken from Venezuela to Un. This admixture is also found e!se'. maintained o'._._ Vikings the millennia by ecological. of the world.' aiso ' :n South.Green and."'__.'.. u: introducing an'-)_ ....ex kI_.:ch mo_ comp'...._ rela'._.' ma_ .'.e of e' been found. Na. ha'... of . in some places.. the considerab!e _...e been numerous. justifying the enormous intertnbal and interregional drift: they must also have moved around. 1990) reassure us that we are on the right track in assuming a parallelism of genetic and linguistic differentiation in America._ are _qui:'ed to eat .:mfc. _ is most ::esei'._ '"..J'._..'.'£-'2but the archaecio:...fairl. and social separation among the m'oups.c. 'ff'.'. Some subfamilies. The research by Spuhler in North America and that on the Panama Chibchan (Barrantes et al.. _.e_ :o Chibchan. there is not.:..ajo). ... '.the Amazon bas'.X O_.'e'. '.... behavioral. ..i/bin the reg. v.. It is not unreasonable to think that the 3..... ...a.'.e. e..ana_!e that a.. • settle this _..hc '-'_'.".-iable size. at a micmgeographic level. K_ inferencesabout '"_ cr'%r of er. especially in areas like the Amazon and Orinoco basins. Equatorial• Go.orica] ar... it seems natura! to sucres: t_a: the. r.e 'mixed v.bud. into Sou:.. ci...': enormous amount of genetic drift clearl.olutionar'.ot.._:..e:_h: .._e ha_e tried to avoid using populations in v..-e U small populations.America..e_ bias.. that this research model is productive.. Iin_ and -_-.. r. -n ' in C . _s _ess _crer..erm_..n. Unfortunate:.._c pa::er:" of"-.". and probabl. .gua': w_t North (:ate. regior. makes an historica! inte:"Fretat_on of ti". besides.'.hich admixture of somemagnitude v. which Js "_1_"-' m :he easl.in_.America The but _.. Jn South south The Paez3': faro.e occunred. and probabl. v._"_. those located farther south being first.... rate from drift is so ". hat '..m_xed _.America._ to .. but it is somev. and linguistics show the need for more detailed research.ic. because ihe. fissions of tribes.e _enetic tree Iess credible in South America thar.u _ .... three maor =_.-'_" -_':. Some regularities emerge from the genetic analysis of major geographic regions in South America but. _ ]n Noah America.... Car:b. . most :..At'tlC cocsta'.-d.1 is difficult to say if other approaches--for example.ho serried ."oblem Eskimos...veen no._ .'-relationbet.. the simplest hypothesis is that fissions and movements of tribes.n G.! Andeans entered before "_* Paez2c. and genetic exchange may have been less f_qaent "_m'"o them than _.'e to a. in other pans of the v.. At the moment. their complex gone flo'.'am. extend fan_". *" -'.ies basis of the geo=t"arhicd.'. The linguistic and geographic splh bep. or it has not . as they still do.'e=t. No such obvious original ethnic differences exist in the rest of the continent. .e it. especia!ly in the southern Na-Dene (Apache and Na.stenous circumstances It seems ished under fa_rl. on the basis of genetic da_a On the _' _ " ..ith Caucasoids.e.'..rra.. if so._ in the order in v-'hJchthe'. and the contrast that can be expected between the genetic and linguistic effects of fusions between tribes all contribute to dissociate genetic and linguistic evolution and to some extent even their relation with geography in this part..._..._.itho that the length of the branches of the tree is hardly indicati.America is u direct consequence of major differences among ethnic groups like Eskimos. ha. ulations ent:rei. V¢ith '.'as _a_sorbed b.. especially in South .._ ._variab!e from prate to place.. Man)' tribes have probably originated from a small number of founders. . -_ .. but it '. and Panoan. th .ene:ic noise caused b'_ daft..olutJonar).. m_id:.._ high_.ere in more direct contact..a-' _ . In general. have a very wide r'ange: the Eqaatoria! fami!L for instance.. Green:an.:" ltisvery "_'-.av an!e_ate their entr'._ hov.z of the £ene'... This varianon is also found in North and Central Ame_ca. a m.'. the last '.. the'• .

: they' therefox reached ".ents would become more common. The idea that a single band wandered across from ._<.2 CH _.atively large size.oIves a specific beha_ ior that :s not t." _..'O rose =n the Andes to levels much higher than in the res_ of the subcontinent.. domes :n regions behind _he advancing frontier v..en: of theAme::cas couZd ha_eproduced thehigh__enetic _ariatioir.0.. :x.he initial dome.. k is difficult to e'.'. closer to saturation of poF_Za:i.orable.. In the past. v.'een demos would e'. The most successful ci'. ant part because the =end..`'. reduced. But in the northern subcontinent.:ere of census siT"e500: ("'. In a m. fusion e'._ur. Perhaps increasing m. een domes...'. and comfort.. the process would be closer to unidimensional • and unidirectional._".th is rapid at the fron'Zer and ceases or s]o_Is dev.v u.. cenainiy un_u_ed except by the search for game. The average rate of (random) move- This DO[02219 . in times befcre European contact.. because :_s ::. n considerably back of the frontier {..he...'aTER 6 merit of 5 miles per _ear _s fas. Along coas_s and fi'..' '. m.ears(a rate of grov. ere urban communities arose• they were e' more fa' in environments either not contaminated or less contaminated by previous inhabitants.d probably" be random in direction.tzh America. fi_e assumptions were made: ( 'J demos (tfibes) '. to ne.pical of _...-ro< -_ ... obse-%e -a (Cavaili-Sfor-za ]9_6)...." cer':ainLv mc._e tcr so•oral _."" li.__e. import.size of a fuii dome in 50 . Pygmies ha'.here the c'..[-kn:.mme.. reduce genetic variation.. an al:: _rate simulation could give more realistic values Per:-a7s only at a later stage.. the final genetic ..en be too high with 3.. but nct .ouic. "--" _- Lob*ever.' = -_¢._eciera:_-.. the population numbers ere'.ears in a ro'_II pattern of repeated mc'...xm ) a nudging cycle (two generations).. of course.e'::er: in'......hess means that often. ploitation of the ".``. c variation . in en'.. It is a necessar': assurr. ._:'c'> merits as di\ers¢ and poorly knov.. and. on the contraQ'.).-.'. in the absence of admixture betv."man and Ca'.. _ . h is.:n density. Plains tribes were proba_1? of re'. for Atnoun P.not e_eo roe:on v_ii] be equal]? favorable for m_cro.__.'th supported by man.entuall> drascica_l'.. the occupation of the Americ' abodes.he move for long distances . pans of Central America v. ae excessive if the models above are fight.'ear).alua_e the sa*uration densiL'.. Man`` urban developments.v that buds advancing in new rotatory. .e'_ a. .. (5) It is iikel.'. desio_ned_ to tes_ _hether the setflem._. The mode! is _I e _.een demos would.n as those in Sc:.e Andean region in the last few m_ilennia. '.n">ing capac:t.s :e.i]enma.the....'_'aS b_ leaps andhounds... More sedentary.'.ou!d soon slo.. it _ C _ ]4 .Gone Row'.'._at:on _t'.a..3. a dome moved an a.e gradual!'.g4 .:he .ed far long distances.Andes and in man...-ksJato America seems unrealistic.. :sat mo_e_ en..ears:. A detain budding and expansion _. expansicn that population gro_. especial'. o dimensions _ou'._Ce!._ the .as caused by rapid saturation of local population dens::':.. e"day hunter-gatherers (e.::Ta:e '.gm es: Ca'. _..'id be completed in fev.g..L: to find comparable modem situations. observations on popular:arts in free growth...ah ._ year but on established paths and repetRive.. in search of no'.. No such developments e_er rock place outside the Andes or other pans of Con:re'. v.. of :gurse. a I:_I Sfer-za 1986).g':ion of an. population density gradua'.. :roCuced "buds" -_'_<% of t>e size of t.Iitizations arose i'._ down pepuia_on increase.'-z::. *73)buds doubled in size every generation of 25 .. see see 2 %.. groups lixed in ::ommunities that reached numbers in the thousands (sec 614)i Thus. Under these conditions. o_ "Y.rocess in tv... but it is diffz. '. Clearly.._. Tubal fusions are bound to i-. _"-_ per mo_e if people stayed in the same D.:.A::erica.he elfonts of drift were buffered and.ic anahsis.erage of 250 miles (5 miles _r ...v teeter. had low mona.'ariet`` of ecological niches and a:::_:e social management in organized states must ha.a_e . but greatly increased the ca.. safety.'. _ ] i_ S . circuits._roum. pia. and onh.aIli-Sforza 1984)... approximate.r.ers.