Application of the Biological Model of Diversification to Cultural Distributions in Tropical
Lowland South America
Application of the Biological Model of Diversification to CulturalDistributions
in Tropical Lowland South America'
Betty J...
FIGURE 1. Boundary of the lowland tropical forest (solid line). Average annual precipitation exceeds 2000 mm ex-
cept in a...
FIGURE 2. Distribution of species within the superspecies Seleniderramaculirostris, a toucanet also reported from iso-
FIGURE 3. Principal forest refuges during warm and dry periods of the Pleistocene, inferred from the ranges of sev-
eral s...
"the Neotropical forest fauna evolved in forest
refugesduringarid phases"(p. 206).
Botanicaldataappearconsistentwith the g...
Analysis of the ranges of species and varieties
of Hymenaea,a resin-producingplant, led Langen-
heim, Lee,and Martin (1973...
fortunately,unwritten languages leave no physical
traceso that it is impossibleto demonstratewithout
FIGURE 5. Floristic zones distinguished by Duke and Black within the lowland tropical forest. The longitudinal di-
vision ...
4 m ~~~~~~~~~~~~ARAWAKA
0 9
FIGURE 6....
:~~~~~~~~~~~~ -
X 0
; < g = + m V~~~~~~~~~~~~~~CRIBAN
0 2 |  / m ~~~~~~~~~~~~~PANOAN
X ) /
FIGURE 8. Distribution of the families, subfamilies, languages, and dialects of Tupi-Guaranian recognized by Rod-
rigues (...
Although the rate of replacement, the size and con-
tent of the standard vocabulary used for comparison,
and even the vali...
FIGURE 9. Subdivisions within the general Tropical Forest culture area recognized by Steward. Those occupied by
FIGURE 10. Culture areas recognized by Murdock based on the clustering of nine types of cultural information. Those
the main waterways, stopping where streams were
less navigable and leaving the hinterland tribes on
a more primitive level...
FIGURE 11. Ethnographic occurrence of two types of hunting traps, the pole snare (circles) and the simple noose
FIGURE 12. Distribution of archeological sequences composed of single (outline) and multiple (hachure) pha...
plexes are more recent, the majority being within
the Christian era. The geographical patterning of
the few available date...
separationsin the Arawakan and Tupi-Guaranian
stocks. The carbon-14date of about3000 yearsago
for the introductionof potte...
significantdisruptionof the aboriginalway of life.
It so happens that the Jivaro live in a region that
- AND C. EVANS. 1957. Archeological investigations at the mouth of the Amazon. Bull. Bur. Am. Ethnol.
167. Smithsonian Ins...
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Application of the biological model of diversification to cultural distributions in tropical lowland south america

  1. 1. Application of the Biological Model of Diversification to Cultural Distributions in Tropical Lowland South America Author(s): Betty J. Meggers Source: Biotropica, Vol. 7, No. 3 (Sep., 1975), pp. 141-161 Published by: The Association for Tropical Biology and Conservation Stable URL: . Accessed: 21/06/2011 20:45 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may 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 . . Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact The Association for Tropical Biology and Conservation is collaborating with JSTOR to digitize, preserve and extend access to Biotropica.
  2. 2. Application of the Biological Model of Diversification to CulturalDistributions in Tropical Lowland South America' Betty J. Meggers Smithsonian Institution, Washington, D.C. 20560, U.S;A. ABSTRACT Evidence from several fields suggests that Amazonia has not remained free from the drastic type of climatic fluctuations experienced 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 forest was periodically reduced to enclaves isolated by savanna or parkland, to explain the abundance of species and their distributions. Since the two most recent episodes occurred subsequent to man's arrival, their impact should be observable in the anthropological evidence. Examination of linguistic, ethnographic, and archeological data reveals patterns in harmony with the biogeographical model, opening a new avenue for interpretation of the history of human adaptation to the tropical lowlands prior to European contact. THE LOWLAND TROPICALFOREST of South America coverssome six million squarekilometersof remark- ablylevel terrain(fig. 1). It is bisectedby the Ama- zon,which flows generallynortheastward,leavingthe Andes at about 5 degreessouth latitudeand crossing the equatoras it emptiesinto the seanorthof Marajo Island. As a consequenceof its miminal elevation and equatorialposition, Amazoniaexperienceslittle annual temperaturevariation,the difference in the averagesfor the warmestand coldest months being about 3 degreesC. Exceptwithin a band extending diagonallysoutheastwardacrossthe central Guianas and into the Brazilianstate of Para, yearlyrainfall normallyexceeds 2000 mm. In spite of an annual dischargefive times greaterthan the Congo,alterna- tion of the rainy seasons to the north and south distributesthe influx over a sufficientperiodof time to reducethe normalcrest at the mouth of the Rio Negro to about 10 meters,or half of that reached by the Ohio River in an areawhere rainfallis only a third as much. The climatic"monotony,"the presenceof streams and ponds with black,white or clear water in most parts of the lowlands,the morphologicaluniformity of the vegetation,and the unobtrusivenessof animals other than birds and monkeys give Amazonia an appearanceof homogeneitythat is the more striking becauseof its vast extent. The two most distinctive ecologicalsubregionsare the varzeaor floodplainof the Amazon and its white-watertributaries,and the terrafirme or land not subjectto annualinundation. Both extend acrossthe lowlands,uniting ratherthan dividing them. The antiquity attributed to this biome and the absence of climatic, topographic,or other natural barriersconducive to isolation, selec- tion, and speciationhave made it difficult to explain the tremendousnumbersof taxa. A hypothesisre- centlyproposedby biogeographersseemsto offer the first satisfactorysolution to this enigma. It postu- lates several cycles of forest fragmentationof suf- ficient length and intensity to allow differentiation between formerly homogeneousgroups. The dates and durationsof the periodsof aridity,the locations andextentsof the forestrefuges,andotherimportant details are still poorly defined, but the two most re- cent episodes appear to postdate man's arrival so that it is worth examining whether the model can shedlight on the equallyheterogeneousand puzzling culturaldistributionsin the Amazonianlowlands. THE BIOLOGICAL MODEL Zoological evidence for environmental change in Amazonia has been provided by Haffer's (1969) analysisof avianspeciation,Vanzolini's(1970) work on lizards, and Muller's (1973) reconstructionof dispersalcenters for terrestrialvertebrates. Similar patternsare supportedby racedistributionin butter- flies (Brown, Sheppard,and Turner 1974). When Haffer reviewed the distributionsof severalgenera, superspecies,and species of forest birds, he found that the zones of secondarycontacttendedto be the same for unrelatedgroups and that their locations did not coincide with existing naturalbarriers. He inferred from this the existence of barriersin the past that isolated ancestralpopulations for a suf- ficient time to permit their differentiation.Since the species were forest adapted,one or more episodes duringwhich the forest was disruptedby more open formsof vegetationwould have impededtheir inter- 1 This paper was prepared for the "Smithsonian Conference on Biological and Biogeographical Concepts in Archeology and Anthropology," held at the Smithsonian Institution from 30 April through 2 May 1974. BIOTROPICA7(3): 141-161 1975 141
  3. 3. 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 enclaves of savanna are characteristic in this corridor, especially north of the Amazon. The Jivaro, Waiwai, and Kayap6 are among 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 that grassland once extended from central Brazil to the Orinoco Basin and the Caribbean coast. Haffer offered a generalized reconstruction of the number and location of past forest refuges based on two principal types of data: (1) the ranges of sev- eral avian superspecies (fig. 2) and (2) the rainfall distribution, which he considered to have been ap- proximately the same as at present but lower in intensity. Six primary regions were postulated with- in Amazonia, one composed of several disjunct seg- ments (fig. 3). He also suggested that small patches of forestprobablysurvivedon mountainslopes,river banks, and portions of the southwestern lowland. Data are insufficient to differentiaterefuges during successiveperiodsof fragmentation,but Haffer com- ments that "it seems possible that the rupturingof the Amazonianforest was most markedduring the arid periods of the Pleistocene. During the post- Pleistocene merely a separationof an upper Ama- zonian forest from lower Amazonian forests may have resulted from the disappearanceof forest growth in the dry, transverse zone through the Obidos-Santaremregion"(op. cit.: 134). A similar conclusion was reached by Vanzolini (1970) from the occurrencesof two generaof forest 142 Meggers
  4. 4. 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 basis for 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 dry leaves covering the forest floor, and Anolis, which lives a few metersabovegroundon tree trunksand associatedplants. Disruptionof formerlycontinuous ranges is implied by disjunct distributionsin one species of each genus and by differentiationin two Anolis species sufficient to permit sympatry. The complexitiesof these cases lead Vanzolini to postu- late two cycles of forest fragmentationand recoa- lescence. He regardsrelief an importantfactor in determiningwhere forest survivedand suggestsfour principal refuge areas during the most recent pe- Cultural Distributions in South America 143
  5. 5. 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 indicate intrusion 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 Haffer 1969: fig. 5). riod: (1) the centralGuianas, (2) the north coast of Venezuela, (3) the easternAndes in Colombia, Ecuador,and Peru, and (4) centralBrazil,near the headwatersof the Tocantins. Consideringthat this analysiswas independentof Haffer's and relies on different zoologicalevidence and environmentalcri- teria, the result is remarkablysimilar (Vuilleumier 1971: fig. 4). Muller's more comprehensivestudy covers the entire Neotropical realm. He recognizes40 disper- sal centers (also consideredcenters of speciation), which representthree general biomes: (1) unfor- ested lowland, (2) arboreal,and (3) oreal. His arborealor rain-forestcenters are fewer and larger than those proposedby Haffer and Vanzolini (Miil- ler 1973: fig. 101) but he supportsthe view that 144 Meggers
  6. 6. "the Neotropical forest fauna evolved in forest refugesduringarid phases"(p. 206). Botanicaldataappearconsistentwith the general model, but suggest the refugeswere not as small as postulatedby Haffer and Vanzolini. A review of the distributionsof the generaandspeciescomposing four families of woody plants widely distributedin the Amazonianlowlands led Prance (1973) to ac- cept past dimatic changeas the primaryfactor un- derlying modern floral diversity. He believes that the refuges suggestedby the zoologists are too re- stricted to have permittedsurvivaland reexpansion of primary vegetation, however, and proposes 16 (fig. 4), some of which coincide with Haffer'sbut are more extensive and some of which were not apparentfrom 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
  7. 7. Analysis of the ranges of species and varieties of Hymenaea,a resin-producingplant, led Langen- heim, Lee,and Martin (1973: 33) to the conclusion that "evolutionwithin the genus has respondedto dry environmentalconditions." They also question whetherthe forestreductionwas as extremeas postu- lated by Vanzoliniand Haffer becausethe reproduc- tive behaviorof trees adaptedto tropicalrain-forest conditionslimits their ability to reinvadelarge open areas. The sameargumenthas been presentedmore forcefully by Gomez-Pompa, Vazquez-Yanes, and Guevara(1972), who stressthat mass extinctionof tropical rain-forestspecies in several parts of the world as a result of human activity supportsother botanical evidence that many primarytrees cannot maintainthemselvesin habitatsthat have been sig- nificantly reduced in size or recolonize extensive disturbedareas. The maximumclearingcompatible with survivalhas not been established,but it is evi- dent that temperate-zonecomparisonsare inappli- cablebecauseof the multitudeof variablesin growth patterns,seed dispersaland survivorship,plant asso- ciations, predator susceptibility, and edaphic re- sourcesthat differentiateprimarytropicalfrom tem- peratevegetation. Periodsof forest fragmentationare also implied by the occurrenceof geological features associated with arid conditionsin regions now heavilyforested (Vanzolini 1970: 41-42). Layersof laterite,cobbles and otherwater-depositedformations,and stone lines have been reported in the soil profiles of Belem, Marajo,the lower Tocantins,Roraima,Cuiaba,and many parts of centralBrazil (northern Mato Gros- so). Similarformationshave been observedon the Rio Caroniin VenezuelanGuiana,in the valleys of the CordilleraOriental in Peru, and on the llanos of eastern Colombia. The stratigraphyin the vi- cinity of Santaremand in Amapa,Brazil,shows two periods of aridityseparatedby a wet interval. Pol- len profiles from the Andes, the Colombian low- lands,and northernGuyanaindicatethat wet periods were interruptedby droughtsof sufficient duration to causedevelopmentof open vegetationover exten- sive areas. The same interpretationhas emerged from analysis of ocean sediments originating from the Brazilianand Guayanashields. The absenceof soil formationsindicativeof aridityin elevatedpor- tions of eastern Brazil, Mato Grosso, Goias, and Maranhao implies these regions remained perma- nently forested. Few carbon-14dates are availablefor estimating the antiquityand durationof these episodes. Three from southern Brazil place the beginning of the mostrecentone between3513 ? 56 and 3284 ? 48 years ago and its terminationabout 2680 ? 150 yearsago (Vanzolini 1970: 42). In easternColom- bia, open vegetation prevailedbetween about 3095 and 1990 yearsago (Van der Hammenin Vanzolini 1970: 42). Considerationof the dates for recent vegetational fluctuations in Africa and post-glacial eustatic changes in sea level on the Braziliancoast leads Muller to bracketthe forest recessionbetween about 5000 and 2400 yearsago (1973: 189). An- other arid interval has been estimatedfrom glacial evidence to have occurredabout 11,000 years ago (Damuth and Fairbridgein Vanzoliniloc. cit.). In summary, zoological, botanical, and geological evidence suggests that the lowland tropical forest of South America suffered several periods of fragmen- tation followed by recoalescence and indicates that the two most recent episodes probably took place subse- quent to man's invasion of the area. It is conse- quently of interest to see whether available cultural evidence exhibits patterns of distribution or other characteristics compatible with the evolutionary model proposed by the biologists. TYPES OF CULTURAL EVIDENCE Reconstruction of the prehistory of tropical lowland South America is handicapped by the fragmentary, uneven, and often unreliable nature of the cultural data. 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 selected aspects of the culture. Even so, the cultural data are probably no worse than those from the natural sci- ences. The fact that they were collected without knowledge of past climatic fluctuations assures that any agreement with the model derived from bio- geography cannot be attributed to bias. Before reviewing the distributional evidence, a word should be said about the reliability of cultural data for historical reconstruction. The three general types, 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 to detect past relationships and their relative closeness, much as the biologist employs evolutionary theory to reconstruct phylogeny. The linguists have gone far- ther, however, and devised a method of estimating the length of time since separation of two languages, families, or stocks. Although lexicostatistical dating remains controversial, the results are useful for com- parison with dates obtained by other methods. Un- 146 Meggers
  8. 8. fortunately,unwritten languages leave no physical traceso that it is impossibleto demonstratewithout otherevidencewherethe speakerswere living before they becameisolatedfrom one another. Ethnographicaldata have other shortcomings. Becausecultureis a meansof behavioraladaptation, it is potentiallysensitiveto environmentalinfluences and capableof rapid alteration. These characteris- tics promotethe formationof cultureareasthat cor- respond in general to natural regions. The same processesunderly the appearanceof floristic zones (fig. 5) andzoogeographicprovinces(Fittkau 1969: fig. 1). The distributionsof traits not subject to adaptivepressures,such as art motifs, myths, songs, and even some technologicalelements,may provide clues 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 advantagesover linguisticand ethnographicalevidence: (1) they are fixed in spaceand (2) theyoften canbe dated.Even when informationis minimal, it may be sufficient to permitthe recognitionof past culturaldifferences and their chronological position and geographical range,as well as to identify the generaltype of cul- ture and its level of complexity. Unfortunately,in wet regions such as Amazonia, where tools and weapons were typ,icallyof perishablematerials,the archeologicalrecorddoes not begin until the intro- duction of pottery. A further complication is the uncertaintyas to what kind of ethnographicunit is representedby an archeologicalcomplex or "phase"; we do not know whether it is the equivalentof a tribe, a sub-tribe,a group of interrelatedfamilies, or someother kind of socialconfiguration. Addedto the difficultiesin interpretingthe vari- ous kinds of anthropologicaldata is the general problem that race, language, and culture are in- dependentvariables. A group can changeculturally while 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,mapping of linguistic, ethnographic,and archeologicaldistri- butions frequentlyproducesdisparateresults. This independencehas the advantage,however,that when correlationsexist they are likely to have historical significance. A review of some of the culturalevi- dence from the tropical forest region of lowland SouthAmericarevealssomeinterestingparallelswith the situationobservedby the biologists. LINGUISTIC DISTRIBUTIONS: Several efforts have been made to classify and map the aboriginal languages of South America. We will consider only two of the best known, one by Loukotka (1967) and the other by Mason (1950), both of whom employed secondary sources. Although the results differ in details, they show a much greater diversity in Ama- zonia than in the southern and eastern portions of the continent. Some of the heterogeneity reflects inadequate information, but even when this is taken into consideration the situation appears remarkably complex. Its reliability is enhanced by the work of Nimuendajui, who spent many years in Amazonia and had first-hand acquaintance with numerous tribes. He recognized 42 stocks and 34 isolated languages in the tropical lowlands, in addition to hundreds 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, especially in a band adjacent to the Andean foothills. With a few notable exceptions, these languages are spoken by small and apparently relict populations. Second, several stocks, families, and subfamilies have disjunct distributions implying that speakers became isolated from one another by migration or that their once- continuous territory was fragmented by intruders of different linguistic affiliation. Third, the three major stocks have one or more widely dispersed families, suggesting long-range population movement. Ara- wakan and Tupi-Guaranian have been systematically analyzed, the former by Noble (1965) and the latter by Rodrigues (1955, 1958), and both classifications are based on quantitative differences that permit in- ferences about degree of relationship and length of time since separation; Cariban has not received simi- lar attention to my knowledge. Following the criterion suggested by Swadesh, Rodrigues (1958: 234) assigned all languages that shared 12 percent or more standard vocabulary to the Tupi-Guaranian stock. He divided the stock into seven families, each composed of groups of lan- guages that shared between 36 and 60 percent basic vocabulary. Languages with 60 percent or more cognates were placed in subfamilies, and those with more than 81 percent cognates were considered di- alects. Six of the families are represented in south- western Amazonia, north of the Rio Guapore and east of the Rio Madeira, and four are restricted to this region (fig. 8). Yuruna has a disjunct dis- tribution on the upper and lower Rio Xingu. Tupi- Guarani, by contrast, is very widespread both within Cultural Distributions in South America 147
  9. 9. 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 the northern and southern portions of the eastern and central zones. Although variation in relief, climate, and soil creates considerable 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 or subzones, 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 of internal differentiation. The Tupi-Guaranifamily containssix subfamilies,20 languages,and numerous dialects,whereasfive of the other families have no subfamilies 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
  10. 10. 4 m ~~~~~~~~~~~~ARAWAKA TUPI-GUJARANIAN '. 0 9 m CARIBAN m X -<gPANOAN )!m CAINGANG amxm TUCANOAN PUINAVEAN-MACU FIGURE 6. Distribution of the aboriginal languages in lowland South America according to Loukotka. Although details differ, the pattern agrees with that obtained by Mason (fig. 7) in showing a concentration of isolated languages in the western portion of the lowland, disjunct distributions in languages belonging to several minor families, and a widespread dispersal of the three major stocks: Arawakan, Tupi-Guaranian, and Cariban. These patterns are consistent with successive periods of population displacement, which could have been provoked by cycles of fragmentation and recoalescence of the tropical forest (after Loukotka 1967). Tupi-Guaranisubfamiliesoccur on the upper Ama- zon (Omagua-Cocama),the lower Amazon (Maue, Mundurukui),in southeasternBrazil (Guayaki) and lowland 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 this extensive distribution, it became the lingua franca in most of Brazilduringthe earlycolonialperiod. Linguistic diversification,like biological diversi- fication,requiresisolationand time. In makinghis- torical reconstructions,linguists considerthe region with greatestvariabilityto be the homelandof the stock and regions with little diversityto be recently invaded (Dyen 1956). By this logic, the Tupi- Guaranianstockoriginatedin the southwesternAma- zonianlowlands (fig. 8). The disjunctdistributions exhibited by most of the families, severalsubfami- lies, and even a few languagessuggesta complicated history involving several periods of displacement. Cultural Distributions in South America 149
  11. 11. :~~~~~~~~~~~~ - l X 0 ~~~~~~~~~~~~~~ARAWAKAN ; < g = + m V~~~~~~~~~~~~~~CRIBAN 0 2 | / m ~~~~~~~~~~~~~PANOAN t X ) / ~~~~~~~~~~~~~~~CAINGAN -_ PUINAVEAN- MAC FIGURE 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 disjunct distributions 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). Lexicostatistical estimates for the separation (and presumably dispersal) of Arawakan and Tupi-Guaranian coincide with carbon-14 dates for the most recent period of forest retreat (after Mason 1950: map). Population dislocation is also implied by the even more widespread dispersal of Arawakan families and subfamilies from a postulated "homeland"in southeasternPeru (Noble 1965: 107 and map), as well as by the large numberof smallerfamilies and isolatedlanguages. 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 words referringto culturaluniversalstend to change at a relativelyuniformrate (Swadesh1955: 1007-1011). 150 Meggers
  12. 12. 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 south of 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 the lower 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 latter subfamily ate also spoken in south-central Amazonia. Lexicostatistical dating indicates that the Tupi-Guaranian families had 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 demonstrate whether all the speakers reported from that area are post-contact intrusions or some represent a displacement northward simultaneous with that toward the coast. Cultural Distributions in South America 151
  13. 13. 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 have been criticized (e.g., Chretien 1962), good agree- ment with archeological evidence has been obtained in several cases (Swadesh 1954). Application of the method to Tupi-Guaranian and Arawakan is hampered more by differences in the degree of simi- larity used to define subcategories than by the lexi- costatistical formulas, since the rates proposed by Swadesh (1955: 1009-1010) and Lees (1953) pro- duce almost identical estimates within the past 4000 years, although they incorporate different types of corrections. Noble (1965: 107) estimated that Proto-Arawakan began to differentiate between 5000 and 3500 years ago; Rodrigues includes within the Tupi-Guaranian stock all languages sharing at least 12 percent cognates, which converts into ? 5000 years for its inception. Noble (op. cit.: 111) has used a minimum of 25 percent cognates as the cri- terion for distinguishing Arawakan "languages" and estimates their separation to have been underway abo-ut 3300 years ago. Rodrigues defines Tupi- Guaranian families by the possession of at least 36 percent cognates, which implies a separation about 2500 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 60 percent cognates, would have begun to differentiate about 1200 years ago, and their component languages 500 or more years ago. These data can be inter- preted as indicating that Arawakan families began to diverge somewhat earlier than Tupi-Guaranian ones, an inference that would be consistent with the wider dispersal and greater habitat diversity of Ara- wakan speakers. In summary, the linguListicpatterning has several general characteristics: ( 1) there is a high degree of diversity, particularly along the base of the Andes, (2) disjunct distributions are common and occur at the family, subfamily, and language levels of differ- entiation, (3) languages belonging to the same family are spoken in widely separated parts of Ama- zonia, and (4) the three major stocks are dispersed throughout the lowlands, with Arawakan having spread into the Antilles and the Andean highlands. Lexicostatistical dating suggests that major disloca- tions took place between about 5000 and 3500 years ago, around 2500 years ago, and around 1200 years ago. ETHNOGRAPHIC DISTRIBUTIONS: A number of at- tempts have been made to recognize culture areas in the tropical forest region. Those by Steward (1948) and Murdock (1951) are best known and will serve to illustrate the type of patterning ex- hibited by ethnographic evidence. Tropical Forest culture in general is characterized by subsistence reliance on hunting and/or fishing, slash-and-burn cultivation, and gathering of wild foods. Villages are composed of one or more com- munal houses, each occupied by an extended family, and are moved approximately every five years. Social interaction is regulated by kinship, sex, and age. The oldest male is usually the head of the household or village, but his influence depends upon his personal qualities and he receives few if any privileges be- cause of his status. Typically, the only specialized occupation is shamanism, and shamans are seldom relieved from the daily tasks traditionally assigned to members of their sex. Warfare and sorcery were widespread and generally motivated by revenge. Re- ligion emphasized the unity between man and his environment by endowing not only human beings but other animals, plants, topographic features, and natural phenomena with spirits capable of doing good or evil. Dietary taboos, magic, and ceremonies were the principal methods of dealing with the spirit world. Birth, puberty, marriage, death, victory in warfare, and maturation of certain wild or cultivated plants were occasions for feasting and dancing, as well as events requiring observance of special taboos and rituals. Since several villages usually partici- pated, these festivals provided opportunities for trade, arrangement of marriages, and other kinds of social interaction. Characteristic material culture items included pottery, hammocks, twilled basketry, cotton cloth, bows and arrows, ornaments, trumpets and drums, all of which were usually made by the individual who used them. Steward (1948: map 8) distinguished six re- gional varieties of Tropical Forest culture and five disjunct enclaves occupied by groups that made little or no use of domesticated plants, and whose general way of life was more similar to the non-agricultural Marginal tribes than to their Tropical Forest neigh- bors (fig. 9). He commented, however, that "From a technological and ecological point of view, the basic Tropical Forest culture is strikingly uniform so far as present data reveal. . . . The more conspicu- ous and the most often mentioned differences be- tween the Tropical Forest peoples are such readily observable items as dress, ornaments, body painting, tattoo, and featherwork. These external features, however, distinguish tribes and individuals even more than major areas; the cultural elements in- volved have highly diversified distributions. The 152 Meggers
  14. 14. FIGURE 9. Subdivisions within the general Tropical Forest culture area recognized by Steward. Those occupied by agriculturalists are: (1) Guianas, (2) Northwest Amazon, (3) Montania, (4) Juru'a-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, and other secondaryfeaturesof bows, basketry,ceramics, andthe like. . . . In drawinglines betweenthe main culturalsubdivisions. . ., therefore,we arebrought to sociological and religious patterns"(1948: 885- 886). Murdock(1951) dividedSouthAmericainto 24 culture areas, 11 of which fall within the Tropical Forest area defined by Steward (fig. 10). Nine classificatorycriteria were employed, among them linguistic affiliation,subsistencetechniques,incidence of selected crafts, house types, kinship terminology, rulesof marriage,and the relativedegreeof develop- ment of sociopolitical institutions (op. cit.: 416). When the resultingmap is comparedwith Steward's, the disagreementsare most markedin the western Cultural Distributions in South America 153
  15. 15. FIGURE 10. Culture areas recognized by Murdock based on the clustering of nine types of cultural information. Those falling within the general region occupied by Steward's 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 local conditionsand the homogenizingeffects of interac- tion with neighboringgroups. Steward'smap emphasizesone aspectof the dis- tribution of Tropical Forest culture that Murdock's does not; namely,the presenceof enclavesoccupied by groupsplacing little or no relianceon cultivated plants. Steward (1948: 883) noted that their dis- tribution tended to be aroundthe peripheryof the AmazonBasin,in regions"difficultto accessin pre- Columbiantimes no less than in modern times to essentiallyriparianpeoples." This led him to infer that "whatis thoughtof as a typicalTropicalForest or silvan culture . . . flowed along the coastand up 154 Meggers
  16. 16. the main waterways, stopping where streams were less navigable and leaving the hinterland tribes on a more primitive level. Some of these tribes . . . re- mained preagricultural nomads. Others . . . adopted some agriculture but otherwise acquired few of the basic 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 areas where the environment precluded adoption of agri- culture. This interpretation deserves reexamination in the light of the new model of ecosystem in- stability. Another way of approaching the ethnographic data is to plot the distribution of individual ele- ments. Nordenskiold (1919) used this method to demonstrate that the Ashluslay and Choroti, two tribes of the Gran Chaco, had been influenced by cultures 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 of this type of analysis to a wider variety of cultural elements might reveal coincidences of patterning use- ful for historical reconstruction (fig. 11). The principal characteristics of the ethnographic evidence can be summarized as follows: (1) culture areas tend to follow the patterning of floral and faunal areas, but are smaller and more irregular sug- gesting that historical and local environmental factors play an important role (alternatively, more refined biological definition might produce closer correla- tion); (2) the distribution of individual elements often appears erratic, but a few disjunct distributions of possible historical significance have been reported; (3) the largest regions occupied by hunting and gathering or incipient agricultural groups are along the western and southern margins of the Tropical Forest area. There is no method of estimating the antiquity of culture areas from ethnographic evi- dence; among component traits, those with the widest distribution are generally assumed to be the oldest. ARCHEOLOGICALDISTRIBUTIONS: Intensive survey and stratigraphic excavation has permitted recon- struction of prehistoric sequences along the Rio Ucayali in eastern Peru (Lathrap 1965), the Rio Napo in eastern Ecuador (Evans and Meggers 1968), the middle and lower Orinoco (Sanoja and Vargas, ms.), northern and southern Guyana (Evans and Meggers 1960), and the mouth of the Amazon (Meggers and Evans 1957, Simoes 1966). Infor- mation on extinct cultures along the varzea comes mainly from collectionsof surface samples of pot- sherdsobtainedby Nimuendajuiin the 1920's from some 85 sites on the middle Amazon,investigations by Hilbert during the 1950's on the Solimoes and Japura(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 the Madeira, Jurua, Purus, Negro, and Tapaj6z, have never been surveyedand the interveningterrafirme is equally unexplored. No preceramicsites have been identified, although a few stone projectile points 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,certaingeneralizations seem apparent. The relativesequencesestablishedfor MarajoIs- land,easternEcuador,and easternPeruarecharacter- ized by discontinuity. Marajoappearsto have been invaded by five successivegroups, the Napo basin by four, and the Ucayaliregion by at least 12 (Lath- rap 1965: 12). The small amountof stratigraphic informationavailablefrom the middle Amazon also suggestsdiscontinuity. By contrast,intensive survey on the upperOrinoco (Evans,Meggers,and Cruxent 1960), in southern Guyana (Evans and Meggers 1960), and on the upper Xingu' (Simoes 1967) re- vealed only one pottery-producingculture in each area (fig. 12). A secondcharacteristicof Amazonianarcheology is the wide distribution of several ceramic tradi- tions. The best known,defined by red and/or black paintingon a white-slippedsurface,occursalong the entire Amazon from its upper tributariesin eastern Peru and Ecuadorto MarajoIsland. Another dis- tinctive tradition, which combines very straight, closelyspaced,parallelincisionswith ringsor puncta- tions, extends along the middle Orinoco,the middle and lower Amazon,and the Guianacoast. A third, characterizedby zones of fine hachureoutlined by broad incisions, has been reported only from the periphery,in easternPeru, eastern Ecuador,coastal Venezuela,and the mouthof the Amazon. At most sites,these "horizonstyles"coexistwith otherdecora- tive techniques,amongthem incision,excision,punc- tation, modeling, applique, and red slipping. The diversity is illustratedby surface samples collected by Nimuendajuifrom the middle Amazon and de- posited in the GdteborgMuseum. About one-third Cultural Distributions in South America 155
  17. 17. 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 and dispersal (after Ryden 1950: fig. 25). of the sites are in the vicinity of the mouth of the Tapajoz and representthe Santaremculture, which surviveduntil Europeancontact. The remaining55 collectionsexhibit 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 diversified when the constituent populations became isolated and were exposedto dissimilarinfluences. Age dif- ferences, site reoccupation,cultural amalgamation, and regional specializationare among other factors likely to be involved but which cannot be assessed until more researchhas been done. Carbon-14dates from lowland sites are few and often difficult to evaluate. The oldest from eastern Amazoniais 980 ? 200 B.C. It is associatedwith the initial ceramiccomplex on MarajoIsland (Sim- 156 Meggers
  18. 18. /~~~~~~~~~~~~ 5 FIGURE 12. Distribution of archeological sequences composed of single (outline) and multiple (hachure) phases or cultures. Superposition of distinct ceramic complexes is characteristic of: (1) Marajo Island, (2) the varzea of the Amazon, 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) the Rupununi savanna, (8) the upper Essequibo, and (9) the upper Xingu'. The complexity of the situation at the center of 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 Andean Area suggests it was introducedinto the lowlands from northwestern South America (Meggers and Evans 1961). The oldest potteryfrom the Ucayali, associatedwith the TutishcainyoPhase,employsthe same decorative technique. Although it has not been 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
  19. 19. plexes are more recent, the majority being within the Christian era. The geographical patterning of the few available dates is sufficiently ambiguous that it 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 as follows: (1) discontinuity and heterogeneity are characteristic of sequences on the eastern and west- ern margins of the lowlands and of sites along the middle and lower Amazon; (2) the existence of sev- eral widespread ceramic traditions suggests intrusion of people and/or cultures from different directions at different times; (3) pottery was being made at the mouth of the Amazon by about 3000 years ago; (4) ceramic complexes and chronological sequences are much more diversified and earlier along the Amazon River and its Andean headwaters than in the other parts of the lowlands that have been ex- plored. APPLICABILITY OF THE BIOGEOGRAPHICAL MODEL TO CULTURAL DATA Heterogeneity is one of the most frequently men- tioned characteristics of both biological and cultural phenomena in the tropical lowlands of South Ameri- ca. Fittkau (1969: 652) has said of the rain-forest fauna that "among almost all animal groups the diversity of species and the abundance of different life forms is strikingly high in comparison with faunas 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 as Amazonia was viewed as a stable environment dur- ing the Pleistocene, because both biological and cul- tural alterations are usually responses to changing conditions. Evidence is accumulating to challenge the assumption of stability and to indicate not only that there were several periods when large sectors were converted into savanna or parkland, but that the two most recent episodes occurred within the past 12,000 years. The impact on the fauna must have been drastic, since few tropical South American mammals are adapted to grasslands. African savannas have 68 species of ungulates, for example, compared with 6 in Latin America. The discrepancy between forest ungulates is considerably less, with 27 species in Africa and 9 in Latin America (Bourliere 1973: Table 5; cf. Fittkau 1969: 652). By contrast, the totals for African and South American forest mam- mals are almost identical: 92 species of rodents in the Congo (Zaire) versus 95 species in Brazil; 44 species of primates in Africa versus 42 in Latin America (Bourliere 1973: Tables 3-4). Savanna plants suitable for human consumption are also uncommon in the New World tropical lowlands. As a consequence, periods of forest reduction would have exposed human groups to the threat of mal- nutrition or even starvation. Those unable to remain in the forest refuges would have had few alternatives. They could have increased their mobility and re- duced group size, expanded the number of wild foods 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 of this 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 of this heterogeneity by the same logic employed by biologists. Speakers of the same language that were separated from one another would diverge in speech and other aspects of their cultures. Differential rates of retention would account for the widespread distribution of some traits and the restricted occur- rence of others. The archeological evidence for suc- cessive intrusions and high intersite heterogeneity also implies population instability. Parallels are evident too in the patterning of bio- logical and cultural phenomena. The complicated archeological situation along the middle Amazon is comparable 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 differentiation in the center of the basin are generally complicated and confused. The latter area was one of fusion of many stocks differentiated on the periphery and brought into contact during a period of ecological complexity, such as the reforestation of the region. This situation also has a practical importance, be- cause it makes clear the impossibility off studying any group in only part of the area-the phenomena of differentiation can only be understood as a whole." The geological dating of 3500 to 2000 years ago for the most recent episode of forest fragmentation coincides with lexicostatistical estimates for major 158 Meggers
  20. 20. separationsin the Arawakan and Tupi-Guaranian stocks. The carbon-14date of about3000 yearsago for the introductionof pottery makingto the lower Amazonalso falls within this time span. The spread of savanna-adaptedflora and fauna into Amazonia during arid intervals (fig. 3) implies that human beings could have behaved in the same way. The affiliation of the oldest potteryto a non-Amazonian tradition suggests that it was introducedby immi- grantsfrom a region of open vegetation,who were attractedinto the lowlands during the most recent periodof deforestation. Althoughthereareno dates for the spread of the Caribanlanguages,the coin- cidencebetween their distributionand the areawith the lowest rainfalland largestenclavesof savannain modern Amazonia suggests an intrusion from the north during this period. Concurrently,the forest- adaptedArawakanand Tupi-Guaranianspeakersap- parentlymigrated in searchof regions where their familiar habitat survived. Carbon-14dates around the beginning of the Christianera for the earliest Tupiguarani-traditionpottery from the coastal Bra- zilian state of Paranaagree with the lexicostatistical estimateof ? 1500 yearsfor the separationof the languages in the Tupi-Guaranisubfamily. Coastal sites are associatedwith forest vegetation and be- come progressivelymore recentfrom south to north. A forestorigin and migrationin searchof an earthly paradise were a prominent part of Tupi-Guarani mythologyat the time of Europeancontact. Variations in the antiquity and permanenceof tropical forest vegetation in different parts of the South American lowlands might account for some of the differences in the degree of dependenceon wild foods (other than game and fish) by groups practicingsubsistenceagriculture.Where nuts, seeds, roots, shoots, fungi, fruits, insects, and other such resourcesare importantin the diet, it might be in- ferredthat relianceon domesticatedfoods was com- paratively recent. Such a contrast exists between the Kayapoof southeasternAmazonia,who adopted agriculturewhen they moved into the forest in post- Europeantimes and who subsistentirelyby hunting, fishing, and gatheringduring severalmonths of the year (Meggers 1971: 70), andthe Waiwai of south- ern Guyana,a Cariban-speakinggroup that makes minimal use of wild plants for food (Yde 1965: 67). The Yukpa-Yuko,who inhabita forestedarea on the Colombia-Venezuelaborder,consume insects belongingto 22 generaand 7 orders (Ruddle 1973: 94). Another indicationof the wild-food potential is the availabilityin marketsin the state of Para of some 70 species of native fruits (Cavalcante1972). This raises the question whether the persistence of hunter-gatherers in the tropical forest can be at- tributed to isolation or whether it is a reflection of the evolution of a schedule of wild-food exploitation that 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 they may have enjoyed the long-term habitat stability con- ducive to efficient adaptation. The fact that, as Steward (1949: 691) noted, "More advanced tech- nologies were absent to a surprising degree, even among the tribes who adjoined or formed enclaves within the Tropical Forest peoples and would seem to 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 of cultural configurations comprising Tropical Forest culture reflect differences in the length of time available for perfecting adaptation to the modern habitat. The Jivaro and the Waiwai, for example, subsist almost exclusively by hunting, fishing, and agriculture, and their sexual division of labor is approximately the same (Meggers 1971: 115). A communal house occupied by an extended family constitutes 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 interaction beyond the family level are arranged. This common foundation supports two very different configura- tions. Jivaro culture is a complicated web, in which economic, social, religious, and technological aspects are tightly interwoven. The staple food is a slightly fermented beverage made by the women from sweet manioc. Since it must be prepared daily by a time- consuming process, it is practical only in the context of polygyny. Polygyny requires an unequal sex ratio, which in turn depends upon elimination of a significant portion of the adult males. This is ac- complished by warfare and blood revenge, in which all males must participate or suffer severe penalties. Integration is so complete that interference with one aspect can destroy the whole configuration. This, in fact, has happened where head hunting has been sup- pressed by national authorities. Waiwai culture ap- pears much less complex. The staple is bitter manioc, which is time-consuming to prepare but can be stored after processing. Monogamy predominates, although polygyny is allowed. Warfare is no longer practiced and there has been an imbalanced sex ratio in recent years because of a higher birth rate of males, but these changes were accommodated without Cultural Distributions in South America 159
  21. 21. significantdisruptionof the aboriginalway of life. It so happens that the Jivaro live in a region that probablyremainedforested during the most recent aridperiod,in which case they would have had am- ple time for culturalspecialization. The fact that they speak a language with no known relatives is anotherindicationof antiquity. The Cariban-speak- ing Waiwai by contrastare recent intrudersin their presentterritory,and their linguistic affiliation sug- gests that they may be relative newcomersto Ama- zonia. CONCLUSION Althoughthe evidencefrom everydisciplineis mini- mal, the fact that existing botanical,zoological,and culturaldatapresentsimilarcharacteristicsand types of patterningsuggeststhat they have a commonex- planation. The alternationbetweenperiodsof forest fragmentationand coalescencepostulatedby the bi- ologists to accountfor species diversitywould have posed subsistence problems for human beings adaptedto the forest becauseof the relativepaucity of savannafood resources.The inferencethat groups unable to remain in the forest refuges would have been obliged to adapt to the savannaor to search for otherforestedhabitatsis compatiblewith linguis- tic and archeologicalevidence for widespreadpre- historicpopulationmovements. Lexicostatistical,ar- cheological,and geological dates all indicate that a major disruptionoccurredbetween about 3500 and 2000 yearsago. Although there is a great deal yet to be learnedbefore the size, location,and duration of the refuges can be accuratelydefined, this model of environmentalfluctuationprovidesa new and ex- citing basis for interpretationof the diversity and discontinuitythat characterizethe culturalevidence. LITERATURECITED BOURLIiERE, 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 ecosystems in 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: evidence from 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 and Ventuari 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. Junk N.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 Volkerkunde 1. Reimer, Berlin. LANGENHEIM, J. H., Y.-T. LEE, AND S. S. MARTIN. 1973. An evolutionary and ecological perspective of Amazonian Hylaea 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): map supplement 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 and map. Smithsonian Institution, Washington, D.C. MEGGERS, B. J. 1971. Amazonia; man and culture in a counterfeit paradise. Aldine, Chicago. 160 Meggers
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