• Save
Application of the biological model of diversification to cultural distributions in tropical lowland south america
Upcoming SlideShare
Loading in...5
×
 

Application of the biological model of diversification to cultural distributions in tropical lowland south america

on

  • 171 views

 

Statistics

Views

Total Views
171
Slideshare-icon Views on SlideShare
171
Embed Views
0

Actions

Likes
0
Downloads
0
Comments
0

0 Embeds 0

No embeds

Accessibility

Categories

Upload Details

Uploaded via as Adobe PDF

Usage Rights

© All Rights Reserved

Report content

Flagged as inappropriate Flag as inappropriate
Flag as inappropriate

Select your reason for flagging this presentation as inappropriate.

Cancel
  • Full Name Full Name Comment goes here.
    Are you sure you want to
    Your message goes here
    Processing…
Post Comment
Edit your comment

    Application of the biological model of diversification to cultural distributions in tropical lowland south america Application of the biological model of diversification to cultural distributions in tropical lowland south america Document Transcript

    • Application of the Biological Model of Diversification to Cultural Distributions in TropicalLowland South AmericaAuthor(s): Betty J. MeggersSource: Biotropica, Vol. 7, No. 3 (Sep., 1975), pp. 141-161Published by: The Association for Tropical Biology and ConservationStable URL: http://www.jstor.org/stable/2989620 .Accessed: 21/06/2011 20:45Your use of the JSTOR archive indicates your acceptance of JSTORs Terms and Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp. JSTORs Terms and Conditions of Use provides, in part, that unlessyou have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and youmay use content in the JSTOR archive only for your personal, non-commercial use.Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at .http://www.jstor.org/action/showPublisher?publisherCode=tropbio. .Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printedpage of such transmission.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact support@jstor.org.The Association for Tropical Biology and Conservation is collaborating with JSTOR to digitize, preserve andextend access to Biotropica.http://www.jstor.org
    • Application of the Biological Model of Diversification to CulturalDistributionsin Tropical Lowland South AmericaBetty J. MeggersSmithsonian Institution, Washington, D.C. 20560, U.S;A.ABSTRACTEvidence from several fields suggests that Amazonia has not remained free from the drastic type of climatic fluctuationsexperienced in temperate and highland portions of the western hemisphere during Pleistocene and Recent times. Zo-ologists and botanists have employed a model of climatic and vegetational cycles, in which the lowland tropical forestwas periodically reduced to enclaves isolated by savanna or parkland, to explain the abundance of species and theirdistributions. Since the two most recent episodes occurred subsequent to mans arrival, their impact should be observablein the anthropological evidence. Examination of linguistic, ethnographic, and archeological data reveals patterns inharmony with the biogeographical model, opening a new avenue for interpretation of the history of human adaptationto the tropical lowlands prior to European contact.THE LOWLAND TROPICALFOREST of South Americacoverssome six million squarekilometersof remark-ablylevel terrain(fig. 1). It is bisectedby the Ama-zon,which flows generallynortheastward,leavingtheAndes at about 5 degreessouth latitudeand crossingthe equatoras it emptiesinto the seanorthof MarajoIsland. As a consequenceof its miminal elevationand equatorialposition, Amazoniaexperienceslittleannual temperaturevariation,the difference in theaveragesfor the warmestand coldest months beingabout 3 degreesC. Exceptwithin a band extendingdiagonallysoutheastwardacrossthe central Guianasand into the Brazilianstate of Para, yearlyrainfallnormallyexceeds 2000 mm. In spite of an annualdischargefive times greaterthan the Congo,alterna-tion of the rainy seasons to the north and southdistributesthe influx over a sufficientperiodof timeto reducethe normalcrest at the mouth of the RioNegro to about 10 meters,or half of that reachedby the Ohio River in an areawhere rainfallis onlya third as much.The climatic"monotony,"the presenceof streamsand ponds with black,white or clear water in mostparts of the lowlands,the morphologicaluniformityof the vegetation,and the unobtrusivenessof animalsother than birds and monkeys give Amazonia anappearanceof homogeneitythat is the more strikingbecauseof its vast extent. The two most distinctiveecologicalsubregionsare the varzeaor floodplainofthe Amazon and its white-watertributaries,and theterrafirme or land not subjectto annualinundation.Both extend acrossthe lowlands,uniting ratherthandividing them. The antiquity attributed to thisbiome and the absence of climatic, topographic,orother natural barriersconducive to isolation, selec-tion, and speciationhave made it difficult to explainthe tremendousnumbersof taxa. A hypothesisre-centlyproposedby biogeographersseemsto offer thefirst satisfactorysolution to this enigma. It postu-lates several cycles of forest fragmentationof suf-ficient length and intensity to allow differentiationbetween formerly homogeneousgroups. The datesand durationsof the periodsof aridity,the locationsandextentsof the forestrefuges,andotherimportantdetails are still poorly defined, but the two most re-cent episodes appear to postdate mans arrival sothat it is worth examining whether the model canshedlight on the equallyheterogeneousand puzzlingculturaldistributionsin the Amazonianlowlands.THE BIOLOGICAL MODELZoological evidence for environmental change inAmazonia has been provided by Haffers (1969)analysisof avianspeciation,Vanzolinis(1970) workon lizards, and Mullers (1973) reconstructionofdispersalcenters for terrestrialvertebrates. Similarpatternsare supportedby racedistributionin butter-flies (Brown, Sheppard,and Turner 1974). WhenHaffer reviewed the distributionsof severalgenera,superspecies,and species of forest birds, he foundthat the zones of secondarycontacttendedto be thesame for unrelatedgroups and that their locationsdid not coincide with existing naturalbarriers. Heinferred from this the existence of barriersin thepast that isolated ancestralpopulations for a suf-ficient time to permit their differentiation.Since thespecies were forest adapted,one or more episodesduringwhich the forest was disruptedby more openformsof vegetationwould have impededtheir inter-1 This paper was prepared for the "Smithsonian Conferenceon Biological and Biogeographical Concepts in Archeologyand Anthropology," held at the Smithsonian Institutionfrom 30 April through 2 May 1974.BIOTROPICA7(3): 141-161 1975 141
    • FIGURE 1. Boundary of the lowland tropical forest (solid line). Average annual precipitation exceeds 2000 mm ex-cept in a diagonal strip across the eastern portion of the region, where it drops below 1500 mm. Large enclavesof savanna are characteristic in this corridor, especially north of the Amazon. The Jivaro, Waiwai, and Kayap6 areamong the numerous tribes that occupied the forest in pre-European times.action. The existence of relict populations in now-isolated patches of savanna in eastern Amazonia(Haffer 1969: 134) also supports the view thatgrassland once extended from central Brazil to theOrinoco Basin and the Caribbean coast.Haffer offered a generalized reconstruction of thenumber and location of past forest refuges based ontwo principal types of data: (1) the ranges of sev-eral avian superspecies (fig. 2) and (2) the rainfalldistribution, which he considered to have been ap-proximately the same as at present but lower inintensity. Six primary regions were postulated with-in Amazonia, one composed of several disjunct seg-ments (fig. 3). He also suggested that small patchesof forestprobablysurvivedon mountainslopes,riverbanks, and portions of the southwestern lowland.Data are insufficient to differentiaterefuges duringsuccessiveperiodsof fragmentation,but Haffer com-ments that "it seems possible that the rupturingofthe Amazonianforest was most markedduring thearid periods of the Pleistocene. During the post-Pleistocene merely a separationof an upper Ama-zonian forest from lower Amazonian forests mayhave resulted from the disappearanceof forestgrowth in the dry, transverse zone through theObidos-Santaremregion"(op. cit.: 134).A similar conclusion was reached by Vanzolini(1970) from the occurrencesof two generaof forest142 Meggers
    • FIGURE 2. Distribution of species within the superspecies Seleniderramaculirostris, a toucanet also reported from iso-lated parts of eastern Brazil. The existence of similar geographical patternings in a number of groups forms the basisfor reconstruction of the location of refuges in the lowlands during Pleistocene and Recent periods of forest fragmenta-tion (after Haffer 1969: fig. 4).lizards: Coleodactylus,which is restricted to dryleaves covering the forest floor, and Anolis, whichlives a few metersabovegroundon tree trunksandassociatedplants. Disruptionof formerlycontinuousranges is implied by disjunct distributionsin onespecies of each genus and by differentiationin twoAnolis species sufficient to permit sympatry. Thecomplexitiesof these cases lead Vanzolini to postu-late two cycles of forest fragmentationand recoa-lescence. He regardsrelief an importantfactor indeterminingwhere forest survivedand suggestsfourprincipal refuge areas during the most recent pe-Cultural Distributions in South America 143
    • FIGURE 3. Principal forest refuges during warm and dry periods of the Pleistocene, inferred from the ranges of sev-eral species of Amazonian birds. River banks and upland slopes probably also remained forested. The arrows indicateintrusion of fauna from open habitats to the south. The refuges are as follows: (1) Choco, (2) Nechi, (3) Catatum-bo, (4) Imern, (5) Napo, (6) East Peruvian, (7) Madeira-Tapaj6s, (8) Belem, and (9) Guiana (after Haffer1969: fig. 5).riod: (1) the centralGuianas, (2) the north coastof Venezuela, (3) the easternAndes in Colombia,Ecuador,and Peru, and (4) centralBrazil,near theheadwatersof the Tocantins. Consideringthat thisanalysiswas independentof Haffers and relies ondifferent zoologicalevidence and environmentalcri-teria, the result is remarkablysimilar (Vuilleumier1971: fig. 4).Mullers more comprehensivestudy covers theentire Neotropical realm. He recognizes40 disper-sal centers (also consideredcenters of speciation),which representthree general biomes: (1) unfor-ested lowland, (2) arboreal,and (3) oreal. Hisarborealor rain-forestcenters are fewer and largerthan those proposedby Haffer and Vanzolini (Miil-ler 1973: fig. 101) but he supportsthe view that144 Meggers
    • "the Neotropical forest fauna evolved in forestrefugesduringarid phases"(p. 206).Botanicaldataappearconsistentwith the generalmodel, but suggest the refugeswere not as small aspostulatedby Haffer and Vanzolini. A review ofthe distributionsof the generaandspeciescomposingfour families of woody plants widely distributedinthe Amazonianlowlands led Prance (1973) to ac-cept past dimatic changeas the primaryfactor un-derlying modern floral diversity. He believes thatthe refuges suggestedby the zoologists are too re-stricted to have permittedsurvivaland reexpansionof primary vegetation, however, and proposes 16(fig. 4), some of which coincide with Haffersbutare more extensive and some of which were notapparentfrom the zoologicalevidence.FIGURE 4. Principal late Pleistocene and post-Pleistocene forest refuges indicated by the distribution of lowland spe-cies of four families of woody plants. Although the areas are larger and more numerous than those postulated by zo-ologists, there is general agreement. The refuges are: (1) Choco, (2) Nechi, (3) Santa Marta, (4) Catumbo, (5)Rancho Grande, (6) Paria, (7) Imataca, (8) Guiana, (9) Imern, (10) Napo, (11) Olivensa, (12) Tefe, (13)Manaus, (14) East Peru, (15) Rondonia-Aripuana, and (16) Belem-Xingu (after Prance 1973: fig. 24).Cultural Distributions in South America 145
    • Analysis of the ranges of species and varietiesof Hymenaea,a resin-producingplant, led Langen-heim, Lee,and Martin (1973: 33) to the conclusionthat "evolutionwithin the genus has respondedtodry environmentalconditions." They also questionwhetherthe forestreductionwas as extremeas postu-lated by Vanzoliniand Haffer becausethe reproduc-tive behaviorof trees adaptedto tropicalrain-forestconditionslimits their ability to reinvadelarge openareas. The sameargumenthas been presentedmoreforcefully by Gomez-Pompa, Vazquez-Yanes, andGuevara(1972), who stressthat mass extinctionoftropical rain-forestspecies in several parts of theworld as a result of human activity supportsotherbotanical evidence that many primarytrees cannotmaintainthemselvesin habitatsthat have been sig-nificantly reduced in size or recolonize extensivedisturbedareas. The maximumclearingcompatiblewith survivalhas not been established,but it is evi-dent that temperate-zonecomparisonsare inappli-cablebecauseof the multitudeof variablesin growthpatterns,seed dispersaland survivorship,plant asso-ciations, predator susceptibility, and edaphic re-sourcesthat differentiateprimarytropicalfrom tem-peratevegetation.Periodsof forest fragmentationare also impliedby the occurrenceof geological features associatedwith arid conditionsin regions now heavilyforested(Vanzolini 1970: 41-42). Layersof laterite,cobblesand otherwater-depositedformations,and stone lineshave been reported in the soil profiles of Belem,Marajo,the lower Tocantins,Roraima,Cuiaba,andmany parts of centralBrazil (northern Mato Gros-so). Similarformationshave been observedon theRio Caroniin VenezuelanGuiana,in the valleys ofthe CordilleraOriental in Peru, and on the llanosof eastern Colombia. The stratigraphyin the vi-cinity of Santaremand in Amapa,Brazil,shows twoperiods of aridityseparatedby a wet interval. Pol-len profiles from the Andes, the Colombian low-lands,and northernGuyanaindicatethat wet periodswere interruptedby droughtsof sufficient durationto causedevelopmentof open vegetationover exten-sive areas. The same interpretationhas emergedfrom analysis of ocean sediments originating fromthe Brazilianand Guayanashields. The absenceofsoil formationsindicativeof aridityin elevatedpor-tions of eastern Brazil, Mato Grosso, Goias, andMaranhao implies these regions remained perma-nently forested.Few carbon-14dates are availablefor estimatingthe antiquityand durationof these episodes. Threefrom southern Brazil place the beginning of themostrecentone between3513 ? 56 and 3284 ? 48years ago and its terminationabout 2680 ? 150yearsago (Vanzolini 1970: 42). In easternColom-bia, open vegetation prevailedbetween about 3095and 1990 yearsago (Van der Hammenin Vanzolini1970: 42). Considerationof the dates for recentvegetational fluctuations in Africa and post-glacialeustatic changes in sea level on the Braziliancoastleads Muller to bracketthe forest recessionbetweenabout 5000 and 2400 yearsago (1973: 189). An-other arid interval has been estimatedfrom glacialevidence to have occurredabout 11,000 years ago(Damuth and Fairbridgein Vanzoliniloc. cit.).In summary, zoological, botanical, and geologicalevidence suggests that the lowland tropical forest ofSouth America suffered several periods of fragmen-tation followed by recoalescence and indicates that thetwo most recent episodes probably took place subse-quent to mans invasion of the area. It is conse-quently of interest to see whether available culturalevidence exhibits patterns of distribution or othercharacteristics compatible with the evolutionarymodel proposed by the biologists.TYPES OF CULTURAL EVIDENCEReconstruction of the prehistory of tropical lowlandSouth America is handicapped by the fragmentary,uneven, and often unreliable nature of the culturaldata. Vast areas are unknown archeologically; hun-dreds of languages remain unclassified because of in-adequate information or have been assigned to cate-gories on the basis of a few words; detailed ethno-graphic studies are few and often limited to selectedaspects of the culture. Even so, the cultural data areprobably no worse than those from the natural sci-ences. The fact that they were collected withoutknowledge of past climatic fluctuations assures thatany agreement with the model derived from bio-geography cannot be attributed to bias.Before reviewing the distributional evidence, aword should be said about the reliability of culturaldata for historical reconstruction. The three generaltypes, linguistic, ethnographic, and archeological,have different advantages and disadvantages. Lin-guistic data are most amenable to systematic treat-ment and least subject to adaptive pressures. Lan-guages follow rules of change that can be used todetect past relationships and their relative closeness,much as the biologist employs evolutionary theory toreconstruct phylogeny. The linguists have gone far-ther, however, and devised a method of estimatingthe length of time since separation of two languages,families, or stocks. Although lexicostatistical datingremains controversial, the results are useful for com-parison with dates obtained by other methods. Un-146 Meggers
    • fortunately,unwritten languages leave no physicaltraceso that it is impossibleto demonstratewithoutotherevidencewherethe speakerswere living beforethey becameisolatedfrom one another.Ethnographicaldata have other shortcomings.Becausecultureis a meansof behavioraladaptation,it is potentiallysensitiveto environmentalinfluencesand capableof rapid alteration. These characteris-tics promotethe formationof cultureareasthat cor-respond in general to natural regions. The sameprocessesunderly the appearanceof floristic zones(fig. 5) andzoogeographicprovinces(Fittkau 1969:fig. 1). The distributionsof traits not subject toadaptivepressures,such as art motifs, myths, songs,and even some technologicalelements,may provideclues to past relationships,but the existenceof num-erousvariables(among them easeof diffusion,avail-ability of raw materials,and differentialratesof re-tention) and gaps in informationreduce their re-liability for historicalreconstruction(e.g., Norden-skiold 1919; Ryden 1950).Archeologicalremainshave two advantagesoverlinguisticand ethnographicalevidence: (1) they arefixed in spaceand (2) theyoften canbe dated.Evenwhen informationis minimal, it may be sufficientto permitthe recognitionof past culturaldifferencesand their chronological position and geographicalrange,as well as to identify the generaltype of cul-ture and its level of complexity. Unfortunately,inwet regions such as Amazonia, where tools andweapons were typ,icallyof perishablematerials,thearcheologicalrecorddoes not begin until the intro-duction of pottery. A further complication is theuncertaintyas to what kind of ethnographicunit isrepresentedby an archeologicalcomplex or "phase";we do not know whether it is the equivalentof atribe, a sub-tribe,a group of interrelatedfamilies,or someother kind of socialconfiguration.Addedto the difficultiesin interpretingthe vari-ous kinds of anthropologicaldata is the generalproblem that race, language, and culture are in-dependentvariables. A group can changeculturallywhile retaining its language, or adopt a new lan-guage without modifying its patternsof daily life.People of any race can learn any languageand par-ticipate in any culture. As a consequence,mappingof linguistic, ethnographic,and archeologicaldistri-butions frequentlyproducesdisparateresults. Thisindependencehas the advantage,however,that whencorrelationsexist they are likely to have historicalsignificance. A review of some of the culturalevi-dence from the tropical forest region of lowlandSouthAmericarevealssomeinterestingparallelswiththe situationobservedby the biologists.LINGUISTIC DISTRIBUTIONS: Several efforts have beenmade to classify and map the aboriginal languagesof South America. We will consider only two ofthe best known, one by Loukotka (1967) and theother by Mason (1950), both of whom employedsecondary sources. Although the results differ indetails, they show a much greater diversity in Ama-zonia than in the southern and eastern portions ofthe continent. Some of the heterogeneity reflectsinadequate information, but even when this is takeninto consideration the situation appears remarkablycomplex. Its reliability is enhanced by the work ofNimuendajui, who spent many years in Amazoniaand had first-hand acquaintance with numeroustribes. He recognized 42 stocks and 34 isolatedlanguages in the tropical lowlands, in addition tohundreds of languages that he could not classify be-cause of insufficient information (Mason 1950: 166-167).When mapped, the linguistic distributions ex-hibit several interesting features (figs. 6-7). First,diversity is greatest in western Amazonia, especiallyin a band adjacent to the Andean foothills. With afew notable exceptions, these languages are spokenby small and apparently relict populations. Second,several stocks, families, and subfamilies have disjunctdistributions implying that speakers became isolatedfrom one another by migration or that their once-continuous territory was fragmented by intruders ofdifferent linguistic affiliation. Third, the three majorstocks have one or more widely dispersed families,suggesting long-range population movement. Ara-wakan and Tupi-Guaranian have been systematicallyanalyzed, the former by Noble (1965) and the latterby Rodrigues (1955, 1958), and both classificationsare based on quantitative differences that permit in-ferences about degree of relationship and length oftime since separation; Cariban has not received simi-lar attention to my knowledge.Following the criterion suggested by Swadesh,Rodrigues (1958: 234) assigned all languages thatshared 12 percent or more standard vocabulary tothe Tupi-Guaranian stock. He divided the stock intoseven families, each composed of groups of lan-guages that shared between 36 and 60 percent basicvocabulary. Languages with 60 percent or morecognates were placed in subfamilies, and those withmore than 81 percent cognates were considered di-alects. Six of the families are represented in south-western Amazonia, north of the Rio Guapore andeast of the Rio Madeira, and four are restricted tothis region (fig. 8). Yuruna has a disjunct dis-tribution on the upper and lower Rio Xingu. Tupi-Guarani, by contrast, is very widespread both withinCultural Distributions in South America 147
    • FIGURE 5. Floristic zones distinguished by Duke and Black within the lowland tropical forest. The longitudinal di-vision is related to rainfall, but edaphic and historic factors prevent a close correlation with climate. The eastern,central, and western regions are subdivided latitudinally, with the Amazon River forming the boundary between thenorthern and southern portions of the eastern and central zones. Although variation in relief, climate, and soil createsconsiderable heterogeneity within each zone, three general patterns of plant distribution have been noted: (1) species ex-tending over all three regions, mainly forms adapted to the floodplain; (2) species restricted to one of the zones orsubzones, and (3) species present in the eastern and western but absent from the central zone (after Langenheim, Lee,and Martin 1973: fig. 3 and pp. 10-11).Amazoniaand along the Braziliancoast.Territorialmagnitude correlateswith degree ofinternal differentiation. The Tupi-Guaranifamilycontainssix subfamilies,20 languages,and numerousdialects,whereasfive of the other families have nosubfamilies and only one to five component lan-guages. Yuruna is divided into two subfamilies.one with two and the other with a single language.148 Meggers
    • 4 m ~~~~~~~~~~~~ARAWAKATUPI-GUJARANIAN.0 9m CARIBANm X -<gPANOAN)!m CAINGANGamxm TUCANOANPUINAVEAN-MACUFIGURE 6. Distribution of the aboriginal languages in lowland South America according to Loukotka. Althoughdetails differ, the pattern agrees with that obtained by Mason (fig. 7) in showing a concentration of isolated languagesin the western portion of the lowland, disjunct distributions in languages belonging to several minor families, and awidespread dispersal of the three major stocks: Arawakan, Tupi-Guaranian, and Cariban. These patterns are consistentwith successive periods of population displacement, which could have been provoked by cycles of fragmentation andrecoalescence of the tropical forest (after Loukotka 1967).Tupi-Guaranisubfamiliesoccur on the upper Ama-zon (Omagua-Cocama),the lower Amazon (Maue,Mundurukui),in southeasternBrazil (Guayaki) andlowland Bolivia (Siriono), as well as numerousin-tervening regions. The most widespreadlanguage,also called Tupif-Guarani,was spoken throughoutal-most the entire rangeof the stock. Becauseof thisextensive distribution, it became the lingua francain most of Brazilduringthe earlycolonialperiod.Linguistic diversification,like biological diversi-fication,requiresisolationand time. In makinghis-torical reconstructions,linguists considerthe regionwith greatestvariabilityto be the homelandof thestock and regions with little diversityto be recentlyinvaded (Dyen 1956). By this logic, the Tupi-Guaranianstockoriginatedin the southwesternAma-zonianlowlands (fig. 8). The disjunctdistributionsexhibited by most of the families, severalsubfami-lies, and even a few languagessuggesta complicatedhistory involving several periods of displacement.Cultural Distributions in South America 149
    • :~~~~~~~~~~~~ -lX 0~~~~~~~~~~~~~~ARAWAKAN; < g = + m V~~~~~~~~~~~~~~CRIBAN0 2 | / m ~~~~~~~~~~~~~PANOANtX ) /~~~~~~~~~~~~~~~CAINGAN-_ PUINAVEAN- MACFIGURE 7. Distribution of the languages of lowland South America according to the classification of Mason. Note-worthy are the concentration of isolated languages or relict families around the periphery of Amazonia, the disjunctdistributions of languages belonging to four minor families (Caingang, Panoan, Tucanoan, and Puinavean-Macu),and the widespread dispersal of the three major families (Arawakan, Tupi-Guaranian, and Cariban). Lexicostatisticalestimates for the separation (and presumably dispersal) of Arawakan and Tupi-Guaranian coincide with carbon-14dates for the most recent period of forest retreat (after Mason 1950: map).Population dislocation is also implied by the evenmore widespread dispersal of Arawakan familiesand subfamilies from a postulated "homeland"insoutheasternPeru (Noble 1965: 107 and map), aswell as by the large numberof smallerfamilies andisolatedlanguages.The time of separationof stocks, families, lan-guages, and dialects can be estimated by lexicosta-tistical dating (also known as glottochronology).The method is basedon the observationthat wordsreferringto culturaluniversalstend to change at arelativelyuniformrate (Swadesh1955: 1007-1011).150 Meggers
    • FIGURE 8. Distribution of the families, subfamilies, languages, and dialects of Tupi-Guaranian recognized by Rod-rigues (1958). The locations are those shown by Mason (fig. 7). The presence of six of the seven families southof the upper Madeira suggests a homeland in this part of the lowlands. The seventh family, Yuruna, has a disjunct dis-tribution on the upper and lower Xingui. The only family tO achieve wide dispersal is Tupi-Guarani. It has differen-tiated into six subfamilies, which are located on the upper Amazon (B), the lower Amazon (D), the right bank of thelower Madeira (E), the southern lcowlands (F), southern Mato Grosso (C), and along the Brazilian coast from the Uru-guayan border to the mouth of the Amazon, with enclaves in the eastern Guianas (A). Seve,al languages of the lattersubfamily ate also spoken in south-central Amazonia. Lexicostatistical dating indicates that the Tupi-Guaranian familieshad separated about 2500 years ago and the subfamilies some 1200 years ago. Archeological evidence places the ap-pearance of the Tupiguarani ceramic tradition on the south coast of Brazil about the beginning of the Christian era.There iS insufficient archeological evidence from the region between the Madeira and the lower Amazon to demonstratewhether all the speakers reported from that area are post-contact intrusions or some represent a displacement northwardsimultaneous with that toward the coast.Cultural Distributions in South America 151
    • Although the rate of replacement, the size and con-tent of the standard vocabulary used for comparison,and even the validity of the general hypothesis havebeen criticized (e.g., Chretien 1962), good agree-ment with archeological evidence has been obtainedin several cases (Swadesh 1954). Application ofthe method to Tupi-Guaranian and Arawakan ishampered more by differences in the degree of simi-larity used to define subcategories than by the lexi-costatistical formulas, since the rates proposed bySwadesh (1955: 1009-1010) and Lees (1953) pro-duce almost identical estimates within the past 4000years, although they incorporate different types ofcorrections. Noble (1965: 107) estimated thatProto-Arawakan began to differentiate between 5000and 3500 years ago; Rodrigues includes within theTupi-Guaranian stock all languages sharing at least12 percent cognates, which converts into ? 5000years for its inception. Noble (op. cit.: 111) hasused a minimum of 25 percent cognates as the cri-terion for distinguishing Arawakan "languages" andestimates their separation to have been underwayabo-ut 3300 years ago. Rodrigues defines Tupi-Guaranian families by the possession of at least 36percent cognates, which implies a separation about2500 years ago. This is comparable to the 35 per-cent shared cognates detected by Noble (op. cit.:110-111) for the Maipuran division of Arawakan.Tupi-Guaranian subfamilies, sharing more than 60percent cognates, would have begun to differentiateabout 1200 years ago, and their component languages500 or more years ago. These data can be inter-preted as indicating that Arawakan families beganto diverge somewhat earlier than Tupi-Guaranianones, an inference that would be consistent with thewider dispersal and greater habitat diversity of Ara-wakan speakers.In summary, the linguListicpatterning has severalgeneral characteristics: ( 1) there is a high degreeof diversity, particularly along the base of the Andes,(2) disjunct distributions are common and occur atthe family, subfamily, and language levels of differ-entiation, (3) languages belonging to the samefamily are spoken in widely separated parts of Ama-zonia, and (4) the three major stocks are dispersedthroughout the lowlands, with Arawakan havingspread into the Antilles and the Andean highlands.Lexicostatistical dating suggests that major disloca-tions took place between about 5000 and 3500 yearsago, around 2500 years ago, and around 1200 yearsago.ETHNOGRAPHIC DISTRIBUTIONS: A number of at-tempts have been made to recognize culture areasin the tropical forest region. Those by Steward(1948) and Murdock (1951) are best known andwill serve to illustrate the type of patterning ex-hibited by ethnographic evidence.Tropical Forest culture in general is characterizedby subsistence reliance on hunting and/or fishing,slash-and-burn cultivation, and gathering of wildfoods. Villages are composed of one or more com-munal houses, each occupied by an extended family,and are moved approximately every five years. Socialinteraction is regulated by kinship, sex, and age. Theoldest male is usually the head of the household orvillage, but his influence depends upon his personalqualities and he receives few if any privileges be-cause of his status. Typically, the only specializedoccupation is shamanism, and shamans are seldomrelieved from the daily tasks traditionally assignedto members of their sex. Warfare and sorcery werewidespread and generally motivated by revenge. Re-ligion emphasized the unity between man and hisenvironment by endowing not only human beingsbut other animals, plants, topographic features, andnatural phenomena with spirits capable of doinggood or evil. Dietary taboos, magic, and ceremonieswere the principal methods of dealing with the spiritworld. Birth, puberty, marriage, death, victory inwarfare, and maturation of certain wild or cultivatedplants were occasions for feasting and dancing, aswell as events requiring observance of special taboosand rituals. Since several villages usually partici-pated, these festivals provided opportunities fortrade, arrangement of marriages, and other kindsof social interaction. Characteristic material cultureitems included pottery, hammocks, twilled basketry,cotton cloth, bows and arrows, ornaments, trumpetsand drums, all of which were usually made by theindividual who used them.Steward (1948: map 8) distinguished six re-gional varieties of Tropical Forest culture and fivedisjunct enclaves occupied by groups that made littleor no use of domesticated plants, and whose generalway of life was more similar to the non-agriculturalMarginal tribes than to their Tropical Forest neigh-bors (fig. 9). He commented, however, that "Froma technological and ecological point of view, thebasic Tropical Forest culture is strikingly uniform sofar as present data reveal. . . . The more conspicu-ous and the most often mentioned differences be-tween the Tropical Forest peoples are such readilyobservable items as dress, ornaments, body painting,tattoo, and featherwork. These external features,however, distinguish tribes and individuals evenmore than major areas; the cultural elements in-volved have highly diversified distributions. The152 Meggers
    • FIGURE 9. Subdivisions within the general Tropical Forest culture area recognized by Steward. Those occupied byagriculturalists are: (1) Guianas, (2) Northwest Amazon, (3) Montania, (4) Jurua-Purus, (5) Mojos-Chiquitos, and(6) Tupian with three regional variants. Hachured areas represent hunter-gatherers and stippled ones, incipient agricul-turalists (after Steward 1948: map 8).same is probablytrue of ornamentation,form, andother secondaryfeaturesof bows, basketry,ceramics,andthe like. . . . In drawinglines betweenthe mainculturalsubdivisions. . ., therefore,we arebroughtto sociological and religious patterns"(1948: 885-886).Murdock(1951) dividedSouthAmericainto 24culture areas, 11 of which fall within the TropicalForest area defined by Steward (fig. 10). Nineclassificatorycriteria were employed, among themlinguistic affiliation,subsistencetechniques,incidenceof selected crafts, house types, kinship terminology,rulesof marriage,and the relativedegreeof develop-ment of sociopolitical institutions (op. cit.: 416).When the resultingmap is comparedwith Stewards,the disagreementsare most markedin the westernCultural Distributions in South America 153
    • FIGURE 10. Culture areas recognized by Murdock based on the clustering of nine types of cultural information. Thosefalling within the general region occupied by Stewards Tropical Forest area are: (1) Guiana, (2) Savanna, (3) Caqueta,(4) Loreto, (5) Amazon, (6) Jurua-Purus, (7) Montafia, (8) Bolivian, (9) Xingu, (10) Para, (11) Eastern low-land (after Murdock 1951: fig. 1).portion. In spite of the different classificatorycri-teria, the results exhibit considerablesimilaritybe-causetheyreflectthe pressuresfor adaptationto localconditionsand the homogenizingeffects of interac-tion with neighboringgroups.Stewardsmap emphasizesone aspectof the dis-tribution of Tropical Forest culture that Murdocksdoes not; namely,the presenceof enclavesoccupiedby groupsplacing little or no relianceon cultivatedplants. Steward (1948: 883) noted that their dis-tribution tended to be aroundthe peripheryof theAmazonBasin,in regions"difficultto accessin pre-Columbiantimes no less than in modern times toessentiallyriparianpeoples." This led him to inferthat "whatis thoughtof as a typicalTropicalForestor silvan culture . . . flowed along the coastand up154 Meggers
    • the main waterways, stopping where streams wereless navigable and leaving the hinterland tribes ona more primitive level. Some of these tribes . . . re-mained preagricultural nomads. Others . . . adoptedsome agriculture but otherwise acquired few of thebasic Tropical Forest traits" (ibid.). In other words,he attributed the persistence of a hunting and gath-ering way of life to isolation, except in those areaswhere the environment precluded adoption of agri-culture. This interpretation deserves reexaminationin the light of the new model of ecosystem in-stability.Another way of approaching the ethnographicdata is to plot the distribution of individual ele-ments. Nordenskiold (1919) used this method todemonstrate that the Ashluslay and Choroti, twotribes of the Gran Chaco, had been influenced bycultures of the Andean highlands. A similar ap-proach was employed by Ryden (1950) to recon-struct the origin and dissemination of hunting traps.Although many of his maps indicate a nearly univer-sal occurrence, two exhibit the type of disjunct dis-tribution manifested by certain biological taxa (op.cit.: figs. 10 and 25), suggesting that application ofthis type of analysis to a wider variety of culturalelements might reveal coincidences of patterning use-ful for historical reconstruction (fig. 11).The principal characteristics of the ethnographicevidence can be summarized as follows: (1) cultureareas tend to follow the patterning of floral andfaunal areas, but are smaller and more irregular sug-gesting that historical and local environmental factorsplay an important role (alternatively, more refinedbiological definition might produce closer correla-tion); (2) the distribution of individual elementsoften appears erratic, but a few disjunct distributionsof possible historical significance have been reported;(3) the largest regions occupied by hunting andgathering or incipient agricultural groups are alongthe western and southern margins of the TropicalForest area. There is no method of estimating theantiquity of culture areas from ethnographic evi-dence; among component traits, those with thewidest distribution are generally assumed to be theoldest.ARCHEOLOGICALDISTRIBUTIONS: Intensive surveyand stratigraphic excavation has permitted recon-struction of prehistoric sequences along the RioUcayali in eastern Peru (Lathrap 1965), the RioNapo in eastern Ecuador (Evans and Meggers1968), the middle and lower Orinoco (Sanoja andVargas, ms.), northern and southern Guyana (Evansand Meggers 1960), and the mouth of the Amazon(Meggers and Evans 1957, Simoes 1966). Infor-mation on extinct cultures along the varzea comesmainly from collectionsof surface samples of pot-sherdsobtainedby Nimuendajuiin the 1920s fromsome 85 sites on the middle Amazon,investigationsby Hilbert during the 1950s on the Solimoes andJapura(Hilbert 1968), andmorerecentlyby Simoes(1974). Complete pottery vessels, stone carvings,and potsherdsexist in museumsand private collec-tions, but most are isolatedfinds of unknownor in-exact provenience. Major rivers, among them theMadeira, Jurua, Purus, Negro, and Tapaj6z, havenever been surveyedand the interveningterrafirmeis equally unexplored. No preceramicsites havebeen identified, although a few stone projectilepoints have been encountered. The paucityand in-consistency of the carbon-14 dates has forced ar-cheologiststo rely primarilyon typologicalsimilari-ties for interpretationof the prehistoric situation.In spite of these inadequacies,certaingeneralizationsseem apparent.The relativesequencesestablishedfor MarajoIs-land,easternEcuador,and easternPeruarecharacter-ized by discontinuity. Marajoappearsto have beeninvaded by five successivegroups, the Napo basinby four, and the Ucayaliregion by at least 12 (Lath-rap 1965: 12). The small amountof stratigraphicinformationavailablefrom the middle Amazon alsosuggestsdiscontinuity. By contrast,intensive surveyon the upperOrinoco (Evans,Meggers,and Cruxent1960), in southern Guyana (Evans and Meggers1960), and on the upper Xingu (Simoes 1967) re-vealed only one pottery-producingculture in eacharea (fig. 12).A secondcharacteristicof Amazonianarcheologyis the wide distribution of several ceramic tradi-tions. The best known,defined by red and/or blackpaintingon a white-slippedsurface,occursalong theentire Amazon from its upper tributariesin easternPeru and Ecuadorto MarajoIsland. Another dis-tinctive tradition, which combines very straight,closelyspaced,parallelincisionswith ringsor puncta-tions, extends along the middle Orinoco,the middleand lower Amazon,and the Guianacoast. A third,characterizedby zones of fine hachureoutlined bybroad incisions, has been reported only from theperiphery,in easternPeru, eastern Ecuador,coastalVenezuela,and the mouthof the Amazon. At mostsites,these "horizonstyles"coexistwith otherdecora-tive techniques,amongthem incision,excision,punc-tation, modeling, applique, and red slipping. Thediversity is illustratedby surface samples collectedby Nimuendajuifrom the middle Amazon and de-posited in the GdteborgMuseum. About one-thirdCultural Distributions in South America 155
    • FIGURE 11. Ethnographic occurrence of two types of hunting traps, the pole snare (circles) and the simple noose(squares). The disjunct distributions resemble those employed by biologists to infer successive periods of isolation anddispersal (after Ryden 1950: fig. 25).of the sites are in the vicinity of the mouth of theTapajoz and representthe Santaremculture, whichsurviveduntil Europeancontact. The remaining55collectionsexhibit considerablevariationin the num-ber and kinds of techniquesand motifs. Some con-sistent combinations(for example,painting and in-cision, or incision,punctation,and modeling) prob-ably denote an ancestralcomplex that diversifiedwhen the constituent populations became isolatedand were exposedto dissimilarinfluences. Age dif-ferences, site reoccupation,cultural amalgamation,and regional specializationare among other factorslikely to be involved but which cannot be assesseduntil more researchhas been done.Carbon-14dates from lowland sites are few andoften difficult to evaluate. The oldest from easternAmazoniais 980 ? 200 B.C. It is associatedwiththe initial ceramiccomplex on MarajoIsland (Sim-156 Meggers
    • /~~~~~~~~~~~~5FIGURE 12. Distribution of archeological sequences composed of single (outline) and multiple (hachure) phasesor cultures. Superposition of distinct ceramic complexes is characteristic of: (1) Marajo Island, (2) the varzea of theAmazon, and (3) the lowlands of eastern Peru and (4) eastern Ecuador. Areas where intensive survey has shown pot-tery to be more recent and less diversified are: (5) the lower Japura, (6) the middle and upper Orinoco, (7) theRupununi savanna, (8) the upper Essequibo, and (9) the upper Xingu. The complexity of the situation at the centerof the Amazon basin implies repeated intrusion of disinct traditions, followed by amalgamation, replacement, isolation,and/or emigration (after Evans and Meggers 1969: fig. 80).oes 1968), characterizedby zoned-hachuredecora-tion. Its similarityto earlypotteryfrom the AndeanArea suggests it was introducedinto the lowlandsfrom northwestern South America (Meggers andEvans 1961). The oldest potteryfrom the Ucayali,associatedwith the TutishcainyoPhase,employsthesame decorative technique. Although it has notbeen dated,relatedmaterialfrom the adjacenthigh-lands falls within the second millenniumB.C. Car-bon-14 dates for other Amazonian sites and com-Cultural Distributions in South America 157
    • plexes are more recent, the majority being withinthe Christian era. The geographical patterning ofthe few available dates is sufficiently ambiguous thatit has been used to support conflicting reconstruc-tions of the sources of innovations and their direc-tions of spread (cf. Meggers and Evans 1968, Lath-rap 1970).The archeological evidence can be summarized asfollows: (1) discontinuity and heterogeneity arecharacteristic of sequences on the eastern and west-ern margins of the lowlands and of sites along themiddle and lower Amazon; (2) the existence of sev-eral widespread ceramic traditions suggests intrusionof people and/or cultures from different directionsat different times; (3) pottery was being made atthe mouth of the Amazon by about 3000 years ago;(4) ceramic complexes and chronological sequencesare much more diversified and earlier along theAmazon River and its Andean headwaters than inthe other parts of the lowlands that have been ex-plored.APPLICABILITY OF THEBIOGEOGRAPHICAL MODEL TOCULTURAL DATAHeterogeneity is one of the most frequently men-tioned characteristics of both biological and culturalphenomena in the tropical lowlands of South Ameri-ca. Fittkau (1969: 652) has said of the rain-forestfauna that "among almost all animal groups thediversity of species and the abundance of differentlife forms is strikingly high in comparison withfaunas of other equatorial areas of the world."With regard to languages, Mason (1950: 163)observed that South America is "probably the re-gion of greatest linguistic diversity in the world."This situation was difficult to explain as long asAmazonia was viewed as a stable environment dur-ing the Pleistocene, because both biological and cul-tural alterations are usually responses to changingconditions. Evidence is accumulating to challengethe assumption of stability and to indicate not onlythat there were several periods when large sectorswere converted into savanna or parkland, but thatthe two most recent episodes occurred within thepast 12,000 years. The impact on the fauna musthave been drastic, since few tropical South Americanmammals are adapted to grasslands. African savannashave 68 species of ungulates, for example, comparedwith 6 in Latin America. The discrepancy betweenforest ungulates is considerably less, with 27 speciesin Africa and 9 in Latin America (Bourliere 1973:Table 5; cf. Fittkau 1969: 652). By contrast, thetotals for African and South American forest mam-mals are almost identical: 92 species of rodents inthe Congo (Zaire) versus 95 species in Brazil; 44species of primates in Africa versus 42 in LatinAmerica (Bourliere 1973: Tables 3-4). Savannaplants suitable for human consumption are alsouncommon in the New World tropical lowlands. Asa consequence, periods of forest reduction wouldhave exposed human groups to the threat of mal-nutrition or even starvation. Those unable to remainin the forest refuges would have had few alternatives.They could have increased their mobility and re-duced group size, expanded the number of wildfoods consumed, or emigrated. Decline in popula-tion size and density, increase in spacing and con-sequent isolation between bands, and widespread dis-persal are predictable consequences.When the cultural data are examined in terms ofthis model they appear to fit. The linguistic distri-butions imply widespread migrations by some speak-ers, reduction in group size and isolation among oth-ers. Successive episodes of forest reduction sep-arated by several millennia could explain much ofthis heterogeneity by the same logic employed bybiologists. Speakers of the same language that wereseparated from one another would diverge in speechand other aspects of their cultures. Differentialrates of retention would account for the widespreaddistribution of some traits and the restricted occur-rence of others. The archeological evidence for suc-cessive intrusions and high intersite heterogeneityalso implies population instability.Parallels are evident too in the patterning of bio-logical and cultural phenomena. The complicatedarcheological situation along the middle Amazon iscomparable to and probably a reflection of the cir-cumstances adduced by Vanzolini (1970: 44) to ac-count for the biological complexity: "The most im-portant fact about Amazonia is its soup-plate form:the refuges are elevated and peripheral. This ex-plains perfectly why the patterns of differentiationin the center of the basin are generally complicatedand confused. The latter area was one of fusion ofmany stocks differentiated on the periphery andbrought into contact during a period of ecologicalcomplexity, such as the reforestation of the region.This situation also has a practical importance, be-cause it makes clear the impossibility off studyingany group in only part of the area-the phenomenaof differentiation can only be understood as awhole."The geological dating of 3500 to 2000 years agofor the most recent episode of forest fragmentationcoincides with lexicostatistical estimates for major158 Meggers
    • separationsin the Arawakan and Tupi-Guaranianstocks. The carbon-14date of about3000 yearsagofor the introductionof pottery makingto the lowerAmazonalso falls within this time span. The spreadof savanna-adaptedflora and fauna into Amazoniaduring arid intervals (fig. 3) implies that humanbeings could have behaved in the same way. Theaffiliation of the oldest potteryto a non-Amazoniantradition suggests that it was introducedby immi-grantsfrom a region of open vegetation,who wereattractedinto the lowlands during the most recentperiodof deforestation. Althoughthereareno datesfor the spread of the Caribanlanguages,the coin-cidencebetween their distributionand the areawiththe lowest rainfalland largestenclavesof savannainmodern Amazonia suggests an intrusion from thenorth during this period. Concurrently,the forest-adaptedArawakanand Tupi-Guaranianspeakersap-parentlymigrated in searchof regions where theirfamiliar habitat survived. Carbon-14dates aroundthe beginning of the Christianera for the earliestTupiguarani-traditionpottery from the coastal Bra-zilian state of Paranaagree with the lexicostatisticalestimateof ? 1500 yearsfor the separationof thelanguages in the Tupi-Guaranisubfamily. Coastalsites are associatedwith forest vegetation and be-come progressivelymore recentfrom south to north.A forestorigin and migrationin searchof an earthlyparadise were a prominent part of Tupi-Guaranimythologyat the time of Europeancontact.Variations in the antiquity and permanenceoftropical forest vegetation in different parts of theSouth American lowlands might account for someof the differences in the degree of dependenceonwild foods (other than game and fish) by groupspracticingsubsistenceagriculture.Where nuts, seeds,roots, shoots, fungi, fruits, insects, and other suchresourcesare importantin the diet, it might be in-ferredthat relianceon domesticatedfoods was com-paratively recent. Such a contrast exists betweenthe Kayapoof southeasternAmazonia,who adoptedagriculturewhen they moved into the forest in post-Europeantimes and who subsistentirelyby hunting,fishing, and gatheringduring severalmonths of theyear (Meggers 1971: 70), andthe Waiwai of south-ern Guyana,a Cariban-speakinggroup that makesminimal use of wild plants for food (Yde 1965:67). The Yukpa-Yuko,who inhabita forestedareaon the Colombia-Venezuelaborder,consume insectsbelongingto 22 generaand 7 orders (Ruddle 1973:94).Another indicationof the wild-food potential isthe availabilityin marketsin the state of Para ofsome 70 species of native fruits (Cavalcante1972).This raises the question whether the persistence ofhunter-gatherers in the tropical forest can be at-tributed to isolation or whether it is a reflection ofthe evolution of a schedule of wild-food exploitationthat is more reliable than slash-and-burn agriculture.The tendency of these groups to be located in the vi-cinity of the postulated forest refuges suggests theymay have enjoyed the long-term habitat stability con-ducive to efficient adaptation. The fact that, asSteward (1949: 691) noted, "More advanced tech-nologies were absent to a surprising degree, evenamong the tribes who adjoined or formed enclaveswithin the Tropical Forest peoples and would seemto have had considerable opportunity for borrowing"is compatible with this hypothesis.Another intriguing line of inquiry is the possi-bility that differences in the internal complexity ofcultural configurations comprising Tropical Forestculture reflect differences in the length of timeavailable for perfecting adaptation to the modernhabitat. The Jivaro and the Waiwai, for example,subsist almost exclusively by hunting, fishing, andagriculture, and their sexual division of labor isapproximately the same (Meggers 1971: 115). Acommunal house occupied by an extended familyconstitutes the village, which is economically self-sufficient and geographically isolated. Several vil-lages form a social unit within which marriages, fes-tivals, raids, and other activities requiring interactionbeyond the family level are arranged. This commonfoundation supports two very different configura-tions.Jivaro culture is a complicated web, in whicheconomic, social, religious, and technological aspectsare tightly interwoven. The staple food is a slightlyfermented beverage made by the women from sweetmanioc. Since it must be prepared daily by a time-consuming process, it is practical only in the contextof polygyny. Polygyny requires an unequal sexratio, which in turn depends upon elimination of asignificant portion of the adult males. This is ac-complished by warfare and blood revenge, in whichall males must participate or suffer severe penalties.Integration is so complete that interference with oneaspect can destroy the whole configuration. This, infact, has happened where head hunting has been sup-pressed by national authorities. Waiwai culture ap-pears much less complex. The staple is bittermanioc, which is time-consuming to prepare but canbe stored after processing. Monogamy predominates,although polygyny is allowed. Warfare is no longerpracticed and there has been an imbalanced sex ratioin recent years because of a higher birth rate ofmales, but these changes were accommodated withoutCultural Distributions in South America 159
    • significantdisruptionof the aboriginalway of life.It so happens that the Jivaro live in a region thatprobablyremainedforested during the most recentaridperiod,in which case they would have had am-ple time for culturalspecialization. The fact thatthey speak a language with no known relatives isanotherindicationof antiquity. The Cariban-speak-ing Waiwai by contrastare recent intrudersin theirpresentterritory,and their linguistic affiliation sug-gests that they may be relative newcomersto Ama-zonia.CONCLUSIONAlthoughthe evidencefrom everydisciplineis mini-mal, the fact that existing botanical,zoological,andculturaldatapresentsimilarcharacteristicsand typesof patterningsuggeststhat they have a commonex-planation. The alternationbetweenperiodsof forestfragmentationand coalescencepostulatedby the bi-ologists to accountfor species diversitywould haveposed subsistence problems for human beingsadaptedto the forest becauseof the relativepaucityof savannafood resources.The inferencethat groupsunable to remain in the forest refuges would havebeen obliged to adapt to the savannaor to searchfor otherforestedhabitatsis compatiblewith linguis-tic and archeologicalevidence for widespreadpre-historicpopulationmovements. Lexicostatistical,ar-cheological,and geological dates all indicate that amajor disruptionoccurredbetween about 3500 and2000 yearsago. Although there is a great deal yetto be learnedbefore the size, location,and durationof the refuges can be accuratelydefined, this modelof environmentalfluctuationprovidesa new and ex-citing basis for interpretationof the diversity anddiscontinuitythat characterizethe culturalevidence.LITERATURECITEDBOURLIiERE, F. 1973. The comparative ecology of rain forest mammals in Africa and tropical America: some in-troductory remarks. In B. J. Meggers, E. S. Ayensu, and W. D. Duckworth (Eds.). Tropical forest ecosystemsin Africa and South America: a comparative review, pp. 279-292. Smithsonian Institution Press, Washington,D.C.BRowN, K. S., JR., P. M. SHEPPARD, AND J. R; G. TURNER. 1974. Quaternary refugia in tropical America: evidencefrom race formation in Heliconius butterflies. Proc. R. Soc. 187: 369-378.CAVALCANTE, P. B. 1972. Frutas comestiveis da amazonia, I. Publcoes avuls Mus. Paraense Emilio Goeldi 17. Belem.CHRE1TIEN, C. D. 1962. The mathematical models of glottochronology. Language 38: 11-37.DYEN, I. 1956. Language distribution and migration theory. Language 32: 611-626.EVANS, C., AND B. J. MEGGERS. 1960. Archeological investigations in British Guiana. Bull. Bur. Am. Ethnol. 177.Smithsonian Institution, Washington, D.C.- AND . 1968. Archeological investigations on the Rio Napo, eastern Ecuador. Smithson. Contr. Anthrop.6. Smithsonian Institution, Washington, D.C., AND J. M. CRUXENT. 1960. Preliminary results of archeological investigations along the Orinoco andVentuari Rivers, Venezuela. 33mo Congreso Internacional de Americanistas, Actas 2: 359-369. San Jose.FITTKAU, E. J. 1969. The fauna of South America. In E. J. Fittkau, J. Illies, H. Klinge, G. H. Schwabe and H.Sioli (Eds.). Biogeography and ecology in South America 2: 624-658. Monographiae Biologicae 19. W. JunkN.V., The Hague.GOMEZ-POMPA, A., C. VAZQuEEZ-YANES, AND S. GUEVARA. 1972. The tropical rain forest: a nonrenewable resource.Science, N.Y. 177: 762-765.HAFFER, J. 1969. Speciation in Amazonian forest birds. Science, N.Y. 165: 131-137.HILBERT, P. P. 1968. Archaologische Untersuchungen am mittleren Amazonas. Marburger Studien zur Volkerkunde1. Reimer, Berlin.LANGENHEIM, J. H., Y.-T. LEE, AND S. S. MARTIN. 1973. An evolutionary and ecological perspective of AmazonianHylaea species of Hymeneae (Leguminosae: Caesalpinioideae). Acta Amazonica 3 (1): 5-38.LATHRAP, D. W. 1970. The upper Amazon. Praeger, New York.LEEs, R. B. 1953. The basis of glottochronology. Language 29: 113-127.LOUKOTKA, C. 1967. Ethno-linguistic distribution of South American Indians. Ann. Ass. Am. Geogr. 57 (2): mapsupplement 8.MANN, G. 1969. Die Okosysteme Siidamerikas. In E. J. Fittkau, J. Illies, H. Klinge, G. H. Schwabe, and H. Sioli(Eds.). Biogeography and ecology in South America 1: 171-229. W. Junk N.V., The Hague.MASON, J. A. 1950. The languages of South American Indians. Bull. Bur. Am. Ethnol. 143 (6): 157-317 andmap. Smithsonian Institution, Washington, D.C.MEGGERS, B. J. 1971. Amazonia; man and culture in a counterfeit paradise. Aldine, Chicago.160 Meggers
    • - AND C. EVANS. 1957. Archeological investigations at the mouth of the Amazon. Bull. Bur. Am. Ethnol.167. Smithsonian Institution, Washington, D.C., AND - . 1961. An experimental formulation of horizon styles in the tropical forest area of South Ameri-ca. In S. K. Lothrop and Others. Essays in pre-columbian art and archaeology, pp. 372-388. Harvard Uni-versity Press, Cambridge.MULLER, P. 1973. The dispersal centres of terrestrial vertebrates in the Neotropical realm. W. Junk B.V., The Hague.MURDOCK, G. P. 1951. South American culture areas. SWest. J. Anthrop. 7: 415-436.NOBLE, G. K. 1965. Proto-Arawakan and its descendents. Int. J. Am. Linguistics 31 (3): part II. Bloomington.NORDENSKIOLD, E. 1919. An ethno-geographical analysis of the material culture of two Indian tribes in the GranChaco. Comparative Ethnographical Studies I, Goteborg.PLATr, R. S. 1942. Latin America; countrysides and united regions. McGraw-Hill, New York.PRANCE, G. T. 1972. An ethnobotanical comparison of four tribes of Amazonian Indians. Acta Amazonica 2(2): 7-27.. 1973. Phytogeographic support for the theory of Pleistocene forest refuges in the Amazon Basin, basedon evidence from distribution patterns in Caryocaraceae, Chrysobalanaceae, Dichapetalaceae and Lecythidaceae.Acta Amazonica 3 (3): 5-28.RODRIGUES, A. D. 1955. As linguas "impuras" da familia Tupi-Guarani. 31mo Congresso Internacional de Ameri-canistas, Anais 2: 1055-1071. Sao Paulo.. 1958. Classification of Tupi-Guarani. Int. J. Am. Linguistics 24: 231-234.RUDDLE, K. 1973. The human use of insects: examples from the Yukpa. Biotropica 5: 94-101.RYDEN, S. 1950. A study of South American Indian hunting traps. Revta Mus. paul. 4: 247-352.SIM6ES, M. F. 1966. Resultados preliminares de uma prospecsao arqueologica na regiao dos Rios Goiapi e Camara(liha de Marajo). Simposio sobre a Biota Amazonica, Atas 2: 207-224. Rio de Janerio.1967. Consideraq6es preliminares sobre a arqueologia do alto Xingu, Mato Grosso. Programa Nacionalde Pesquisas Arqueologicas; Resultados preliminares do primeiro ano, 1965-66. Publcoes avuls Mus. ParaenseEmilio Goeldi 6: 129-151. Belem.. 1968. The Castanheira site: new evidence on the antiquity and history of the Ananatuba Phase (MarajoIsland, Brazil). Am. Antiq. 34: 402-410.. 1974. Contribuiqao a arqueologia dos arredores do baixo Rio Negro, Amazonas. Programa Nacional dePesquisas Arqueologicas; Resultados preliminares do quinto ano, 1969-70. Publsoes avuls. Mus. Paraense EmilioGoeldi 26: 165-188. Belem.STEWARD, J. H. 1948. Culture areas of the tropical forests. Bull. Bur. Am. Ethnol. 143 (3): 883-899. Smith-sonian Institution, Washington, D.C.. 1949. South American cultures; an interpretative summary. Bull. Bur. Am. Ethnol. 143 (5): 669-772.Smithsonian Institution, Washington, D.C.SWADESH, M. 1954. Time depths of American linguistic groupings. Am. Anthrop. 56: 361-364.1955. Towards a satisfactory genetic classification of Amerindian languages. 31mOCongresso Internacionalde Americanistas, Anais 2: 1001-1012. Sao Paulo.VANZOLINI, P. E. 1970. Zoologia sistematica, geografia e a origem das especies. Instituto de Geografia, Universi-dade de Sao Paulo, Serie Teses e Monografias 3. Sao Paulo.VUILLEUMIER, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Science, N.Y. 173: 771-780.YDE, J. 1965. Material culture of the Waiwai. National museets Skrifter, Etnografisk Roekke 10. Copenhagen.Cultural Distributions in South America 161