application of the biological model of diversification to cultural distributions in tropical lowland...

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 20:45

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's 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 [email protected].

The Association for Tropical Biology and Conservation is collaborating with JSTOR to digitize, preserve andextend access to Biotropica.

http://www.jstor.org

http://www.jstor.org/action/showPublisher?publisherCode=tropbio

http://www.jstor.org/stable/2989620?origin=JSTOR-pdf

http://www.jstor.org/page/info/about/policies/terms.jsp

http://www.jstor.org/action/showPublisher?publisherCode=tropbio

Application of the Biological Model of Diversification to Cultural Distributions 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 TROPICAL FOREST of South America covers some six million square kilometers of remarkably level terrain (fig. 1). It is bisected by the Ama- zon, which flows generally northeastward, leaving the Andes at about 5 degrees south latitude and crossing the equator as it empties into the sea north of Marajo Island. As a consequence of its miminal elevation and equatorial position, Amazonia experiences little annual temperature variation, the difference in the averages for the warmest and coldest months being about 3 degrees C. Except within a band extending diagonally southeastward across the central Guianas and into the Brazilian state of Para, yearly rainfall normally exceeds 2000 mm. In spite of an annual discharge five times greater than the Congo, alternation of the rainy seasons to the north and south distributes the influx over a sufficient period of time to reduce the normal crest at the mouth of the Rio Negro to about 10 meters, or half of that reached by the Ohio River in an area where rainfall is only a third as much.

The climatic "monotony," the presence of streams and ponds with black, white or clear water in most parts of the lowlands, the morphological uniformity of the vegetation, and the unobtrusiveness of animals other than birds and monkeys give Amazonia an appearance of homogeneity that is the more striking because of its vast extent. The two most distinctive ecological subregions are the varzea or floodplain of the Amazon and its white-water tributaries, and the terra firme or land not subject to annual inundation. Both extend across the lowlands, uniting rather than dividing them. The antiquity attributed to this

biome and the absence of climatic, topographic, or other natural barriers conducive to isolation, selec- tion, and speciation have made it difficult to explain the tremendous numbers of taxa. A hypothesis recently proposed by biogeographers seems to offer the first satisfactory solution to this enigma. It postu- lates several cycles of forest fragmentation of sufficient length and intensity to allow differentiation between formerly homogeneous groups. The dates and durations of the periods of aridity, the locations and extents of the forest refuges, and other important details are still poorly defined, but the two most recent episodes appear to postdate man's arrival so that it is worth examining whether the model can shed light on the equally heterogeneous and puzzling cultural distributions in the Amazonian lowlands.

THE BIOLOGICAL MODEL Zoological evidence for environmental change in Amazonia has been provided by Haffer's (1969) analysis of avian speciation, Vanzolini's (1970) work on lizards, and Muller's (1973) reconstruction of dispersal centers for terrestrial vertebrates. Similar patterns are supported by race distribution in butterflies (Brown, Sheppard, and Turner 1974). When Haffer reviewed the distributions of several genera, superspecies, and species of forest birds, he found that the zones of secondary contact tended to be the same for unrelated groups and that their locations did not coincide with existing natural barriers. He inferred from this the existence of barriers in the past that isolated ancestral populations for a sufficient time to permit their differentiation. Since the species were forest adapted, one or more episodes during which the forest was disrupted by more open forms of vegetation would have impeded their 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.

BIOTROPICA 7(3): 141-161 1975 141

FIGURE 1. Boundary of the lowland tropical forest (solid line). Average annual precipitation exceeds 2000 mm except 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 several avian superspecies (fig. 2) and (2) the rainfall distribution, which he considered to have been approximately the same as at present but lower in intensity. Six primary regions were postulated within Amazonia, one composed of several disjunct seg- ments (fig. 3). He also suggested that small patches

of forest probably survived on mountain slopes, river banks, and portions of the southwestern lowland. Data are insufficient to differentiate refuges during successive periods of fragmentation, but Haffer com- ments that "it seems possible that the rupturing of the Amazonian forest was most marked during the arid periods of the Pleistocene. During the post- Pleistocene merely a separation of an upper Ama- zonian forest from lower Amazonian forests may have resulted from the disappearance of forest growth in the dry, transverse zone through the Obidos-Santarem region" (op. cit.: 134).

A similar conclusion was reached by Vanzolini (1970) from the occurrences of two genera of forest

142 Meggers

FIGURE 2. Distribution of species within the superspecies Seleniderra maculirostris, a toucanet also reported from isolated 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 fragmentation (after Haffer 1969: fig. 4).

lizards: Coleodactylus, which is restricted to dry leaves covering the forest floor, and Anolis, which lives a few meters above ground on tree trunks and associated plants. Disruption of formerly continuous ranges is implied by disjunct distributions in one species of each genus and by differentiation in two

Anolis species sufficient to permit sympatry. The complexities of these cases lead Vanzolini to postu- late two cycles of forest fragmentation and recoalescence. He regards relief an important factor in determining where forest survived and suggests four principal 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 several 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 central Guianas, (2) the north coast of Venezuela, (3) the eastern Andes in Colombia, Ecuador, and Peru, and (4) central Brazil, near the headwaters of the Tocantins. Considering that this analysis was independent of Haffer's and relies on different zoological evidence and environmental criteria, the result is remarkably similar (Vuilleumier 1971: fig. 4).

Muller's more comprehensive study covers the entire Neotropical realm. He recognizes 40 dispersal centers (also considered centers of speciation), which represent three general biomes: (1) unfor- ested lowland, (2) arboreal, and (3) oreal. His arboreal or rain-forest centers are fewer and larger than those proposed by Haffer and Vanzolini (Miil- ler 1973: fig. 101) but he supports the view that

144 Meggers

"the Neotropical forest fauna evolved in forest refuges during arid phases" (p. 206).

Botanical data appear consistent with the general model, but suggest the refuges were not as small as postulated by Haffer and Vanzolini. A review of the distributions of the genera and species composing four families of woody plants widely distributed in the Amazonian lowlands led Prance (1973) to ac-

cept past dimatic change as the primary factor un- derlying modern floral diversity. He believes that the refuges suggested by the zoologists are too restricted to have permitted survival and reexpansion of primary vegetation, however, and proposes 16 (fig. 4), some of which coincide with Haffer's but are more extensive and some of which were not apparent from the zoological evidence.

FIGURE 4. Principal late Pleistocene and post-Pleistocene forest refuges indicated by the distribution of lowland species of four families of woody plants. Although the areas are larger and more numerous than those postulated by zoologists, 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).


Analysis of the ranges of species and varieties of Hymenaea, a resin-producing plant, led Langen- heim, Lee, and Martin (1973: 33) to the conclusion that "evolution within the genus has responded to dry environmental conditions." They also question whether the forest reduction was as extreme as postulated by Vanzolini and Haffer because the reproduc- tive behavior of trees adapted to tropical rain-forest conditions limits their ability to reinvade large open areas. The same argument has been presented more forcefully by Gomez-Pompa, Vazquez-Yanes, and Guevara (1972), who stress that mass extinction of tropical rain-forest species in several parts of the world as a result of human activity supports other botanical evidence that many primary trees cannot maintain themselves in habitats that have been sig- nificantly reduced in size or recolonize extensive disturbed areas. The maximum clearing compatible with survival has not been established, but it is evident that temperate-zone comparisons are inappli- cable because of the multitude of variables in growth patterns, seed dispersal and survivorship, plant asso- ciations, predator susceptibility, and edaphic resources that differentiate primary tropical from temperate vegetation.

Periods of forest fragmentation are also implied by the occurrence of geological features associated with arid conditions in regions now heavily forested (Vanzolini 1970: 41-42). Layers of laterite, cobbles and other water-deposited formations, and stone lines have been reported in the soil profiles of Belem, Marajo, the lower Tocantins, Roraima, Cuiaba, and many parts of central Brazil (northern Mato Gros- so). Similar formations have been observed on the Rio Caroni in Venezuelan Guiana, in the valleys of the Cordillera Oriental in Peru, and on the llanos of eastern Colombia. The stratigraphy in the vicinity of Santarem and in Amapa, Brazil, shows two periods of aridity separated by a wet interval. Pol- len profiles from the Andes, the Colombian lowlands, and northern Guyana indicate that wet periods were interrupted by droughts of sufficient duration to cause development of open vegetation over extensive areas. The same interpretation has emerged from analysis of ocean sediments originating from the Brazilian and Guayana shields. The absence of soil formations indicative of aridity in elevated portions of eastern Brazil, Mato Grosso, Goias, and Maranhao implies these regions remained perma- nently forested.

Few carbon-14 dates are available for estimating the antiquity and duration of these episodes. Three from southern Brazil place the beginning of the most recent one between 3513 ? 56 and 3284 ? 48

years ago and its termination about 2680 ? 150 years ago (Vanzolini 1970: 42). In eastern Colom- bia, open vegetation prevailed between about 3095 and 1990 years ago (Van der Hammen in Vanzolini 1970: 42). Consideration of the dates for recent vegetational fluctuations in Africa and post-glacial eustatic changes in sea level on the Brazilian coast leads Muller to bracket the forest recession between about 5000 and 2400 years ago (1973: 189). An- other arid interval has been estimated from glacial evidence to have occurred about 11,000 years ago (Damuth and Fairbridge in Vanzolini loc. cit.).

In summary, zoological, botanical, and geological evidence suggests that the lowland tropical forest of South America suffered several periods of fragmentation followed by recoalescence and indicates that the two most recent episodes probably took place subsequent 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; hundreds of languages remain unclassified because of inadequate information or have been assigned to cate- gories on the basis of a few words; detailed ethnographic 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 biogeography 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 comparison with dates obtained by other methods. Un-

146 Meggers

fortunately, unwritten languages leave no physical trace so that it is impossible to demonstrate without other evidence where the speakers were living before they became isolated from one another.

Ethnographical data have other shortcomings. Because culture is a means of behavioral adaptation, it is potentially sensitive to environmental influences and capable of rapid alteration. These characteristics promote the formation of culture areas that cor- respond in general to natural regions. The same processes underly the appearance of floristic zones (fig. 5) and zoogeographic provinces (Fittkau 1969: fig. 1). The distributions of traits not subject to adaptive pressures, such as art motifs, myths, songs, and even some technological elements, may provide clues to past relationships, but the existence of numerous variables (among them ease of diffusion, availability of raw materials, and differential rates of retention) and gaps in information reduce their reliability for historical reconstruction (e.g., Norden- skiold 1919; Ryden 1950).

Archeological remains have two advantages over linguistic and ethnographical evidence: (1) they are fixed in space and (2) they often can be dated. Even when information is minimal, it may be sufficient to permit the recognition of past cultural differences and their chronological position and geographical range, as well as to identify the general type of culture and its level of complexity. Unfortunately, in wet regions such as Amazonia, where tools and weapons were typ,ically of perishable materials, the archeological record does not begin until the introduction of pottery. A further complication is the uncertainty as to what kind of ethnographic unit is represented by an archeological complex or "phase"; we do not know whether it is the equivalent of a tribe, a sub-tribe, a group of interrelated families, or some other kind of social configuration.

Added to the difficulties in interpreting the vari- ous kinds of anthropological data is the general problem that race, language, and culture are independent variables. A group can change culturally while retaining its language, or adopt a new language without modifying its patterns of daily life. People of any race can learn any language and participate in any culture. As a consequence, mapping of linguistic, ethnographic, and archeological distributions frequently produces disparate results. This independence has the advantage, however, that when correlations exist they are likely to have historical significance. A review of some of the cultural evidence from the tropical forest region of lowland South America reveals some interesting parallels with the situation observed by 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 because of insufficient information (Mason 1950: 166- 167).

When mapped, the linguistic distributions exhibit 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 similar 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 languages 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 dialects. Six of the families are represented in southwestern 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 distribution on the upper and lower Rio Xingu. Tupi- Guarani, by contrast, is very widespread both within


FIGURE 5. Floristic zones distinguished by Duke and Black within the lowland tropical forest. The longitudinal division 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 extending 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).

Amazonia and along the Brazilian coast. Territorial magnitude correlates with degree of

internal differentiation. The Tupi-Guarani family contains six subfamilies, 20 languages, and numerous

dialects, whereas five of the other families have no subfamilies and only one to five component languages. Yuruna is divided into two subfamilies. one with two and the other with a single language.

148 Meggers

4 m ~~~~~~~~~~~~ARAWAKA

TUPI -GUJARANIAN

'. 0 9

m CARIBAN

m X -< gPANOAN

)!m CAINGANG

amxm TUCANOAN

PUINAVEAN-M ACU

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-Guarani subfamilies occur on the upper Ama- zon (Omagua-Cocama), the lower Amazon (Maue, Mundurukui), in southeastern Brazil (Guayaki) and lowland Bolivia (Siriono), as well as numerous intervening regions. The most widespread language, also called Tupif-Guarani, was spoken throughout almost the entire range of the stock. Because of this extensive distribution, it became the lingua franca in most of Brazil during the early colonial period.

Linguistic diversification, like biological diversi-

fication, requires isolation and time. In making historical reconstructions, linguists consider the region with greatest variability to be the homeland of the stock and regions with little diversity to be recently invaded (Dyen 1956). By this logic, the Tupi- Guaranian stock originated in the southwestern Ama- zonian lowlands (fig. 8). The disjunct distributions exhibited by most of the families, several subfamilies, and even a few languages suggest a complicated history involving several periods of displacement.


:~~~~~~~~~~~~ - 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 southeastern Peru (Noble 1965: 107 and map), as well as by the large number of smaller families and isolated languages.

The time of separation of stocks, families, languages, and dialects can be estimated by lexicostatistical dating (also known as glottochronology). The method is based on the observation that words referring to cultural universals tend to change at a relatively uniform rate (Swadesh 1955: 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 south of the upper Madeira suggests a homeland in this part of the lowlands. The seventh family, Yuruna, has a disjunct distribution on the upper and lower Xingui. The only family tO achieve wide dispersal is Tupi-Guarani. It has differentiated 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 appearance 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.


Although the rate of replacement, the size and content of the standard vocabulary used for comparison, and even the validity of the general hypothesis have been criticized (e.g., Chretien 1962), good agreement 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 similarity used to define subcategories than by the lexicostatistical formulas, since the rates proposed by Swadesh (1955: 1009-1010) and Lees (1953) produce 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 criterion for distinguishing Arawakan "languages" and estimates their separation to have been underway abo-ut 3 300 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 percent 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 linguListic patterning 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 differentiation, (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 exhibited 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 communal 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 because 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 regional 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 between 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 involved have highly diversified distributions. The

152 Meggers

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 agriculturalists (after Steward 1948: map 8).

same is probably true of ornamentation, form, and other secondary features of bows, basketry, ceramics, and the like. . . . In drawing lines between the main cultural subdivisions . . ., therefore, we are brought to sociological and religious patterns" (1948: 885- 886).

Murdock (1951) divided South America into 24 culture areas, 11 of which fall within the Tropical

Forest area defined by Steward (fig. 10). Nine classificatory criteria were employed, among them linguistic affiliation, subsistence techniques, incidence of selected crafts, house types, kinship terminology, rules of marriage, and the relative degree of development of sociopolitical institutions (op. cit.: 416). When the resulting map is compared with Steward's, the disagreements are most marked in the western


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 lowland (after Murdock 1951: fig. 1).

portion. In spite of the different classificatory criteria, the results exhibit considerable similarity because they reflect the pressures for adaptation to local conditions and the homogenizing effects of interaction with neighboring groups.

Steward's map emphasizes one aspect of the distribution of Tropical Forest culture that Murdock's does not; namely, the presence of enclaves occupied

by groups placing little or no reliance on cultivated plants. Steward (1948: 883) noted that their distribution tended to be around the periphery of the Amazon Basin, in regions "difficult to access in pre- Columbian times no less than in modern times to essentially riparian peoples." This led him to infer that "what is thought of as a typical Tropical Forest or silvan culture . . . flowed along the coast and up

154 Meggers

the main waterways, stopping where streams were less navigable and leaving the hinterland tribes on a more primitive level. Some of these tribes . . . remained 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 gathering way of life to isolation, except in those areas where the environment precluded adoption of agriculture. This interpretation deserves reexamination in the light of the new model of ecosystem instability.

Another way of approaching the ethnographic data is to plot the distribution of individual elements. 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 reconstruct the origin and dissemination of hunting traps. Although many of his maps indicate a nearly univer- sal occurrence, two exhibit the type of disjunct distribution 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 useful 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 suggesting that historical and local environmental factors play an important role (alternatively, more refined biological definition might produce closer correlation); (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 evidence; among component traits, those with the widest distribution are generally assumed to be the oldest.

ARCHEOLOGICAL DISTRIBUTIONS: Intensive survey and stratigraphic excavation has permitted reconstruction 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 collections of surface samples of potsherds obtained by Nimuendajui in 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), and more recently by Simoes (1974). Complete pottery vessels, stone carvings, and potsherds exist in museums and private collections, but most are isolated finds of unknown or in- exact provenience. Major rivers, among them the Madeira, Jurua, Purus, Negro, and Tapaj6z, have never been surveyed and the intervening terra firme is equally unexplored. No preceramic sites have been identified, although a few stone projectile points have been encountered. The paucity and in- consistency of the carbon-14 dates has forced ar- cheologists to rely primarily on typological similari- ties for interpretation of the prehistoric situation. In spite of these inadequacies, certain generalizations seem apparent.

The relative sequences established for Marajo Is- land, eastern Ecuador, and eastern Peru are characterized by discontinuity. Marajo appears to have been invaded by five successive groups, the Napo basin by four, and the Ucayali region by at least 12 (Lath- rap 1965: 12). The small amount of stratigraphic information available from the middle Amazon also suggests discontinuity. By contrast, intensive survey on the upper Orinoco (Evans, Meggers, and Cruxent 1960), in southern Guyana (Evans and Meggers 1960), and on the upper Xingu' (Simoes 1967) re- vealed only one pottery-producing culture in each area (fig. 12).

A second characteristic of Amazonian archeology is the wide distribution of several ceramic traditions. The best known, defined by red and/or black painting on a white-slipped surface, occurs along the entire Amazon from its upper tributaries in eastern Peru and Ecuador to Marajo Island. Another distinctive tradition, which combines very straight, closely spaced, parallel incisions with rings or puncta- tions, extends along the middle Orinoco, the middle and lower Amazon, and the Guiana coast. A third, characterized by zones of fine hachure outlined by broad incisions, has been reported only from the periphery, in eastern Peru, eastern Ecuador, coastal Venezuela, and the mouth of the Amazon. At most sites, these "horizon styles" coexist with other decorative techniques, among them incision, excision, punctation, modeling, applique, and red slipping. The diversity is illustrated by surface samples collected by Nimuendajui from the middle Amazon and deposited in the Gdteborg Museum. About one-third


FIGURE 1 1. 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 represent the Santarem culture, which survived until European contact. The remaining 55 collections exhibit considerable variation in the number and kinds of techniques and motifs. Some consistent combinations (for example, painting and incision, or incision, punctation, and modeling) probably denote an ancestral complex that diversified when the constituent populations became isolated

and were exposed to dissimilar influences. Age differences, site reoccupation, cultural amalgamation, and regional specialization are among other factors likely to be involved but which cannot be assessed until more research has been done.

Carbon-14 dates from lowland sites are few and often difficult to evaluate. The oldest from eastern Amazonia is 980 ? 200 B.C. It is associated with the initial ceramic complex on Marajo Island (Sim-

156 Meggers

/~~~~~~~~~~~~

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 pottery 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), characterized by zoned-hachure decora- tion. Its similarity to early pottery from the Andean Area suggests it was introduced into the lowlands from northwestern South America (Meggers and Evans 1961). The oldest pottery from the Ucayali,

associated with the Tutishcainyo Phase, employs the same decorative technique. Although it has not been dated, related material from the adjacent highlands falls within the second millennium B.C. Car- bon-14 dates for other Amazonian sites and com-


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 reconstructions of the sources of innovations and their directions 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 western margins of the lowlands and of sites along the middle and lower Amazon; (2) the existence of several 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 mentioned 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 region of greatest linguistic diversity in the world." This situation was difficult to explain as long as Amazonia was viewed as a stable environment during the Pleistocene, because both biological and cultural 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 mammals 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 reduced group size, expanded the number of wild foods consumed, or emigrated. Decline in population size and density, increase in spacing and con- sequent isolation between bands, and widespread dispersal are predictable consequences.

When the cultural data are examined in terms of this model they appear to fit. The linguistic distributions imply widespread migrations by some speakers, reduction in group size and isolation among others. Successive episodes of forest reduction separated 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 occurrence of others. The archeological evidence for successive intrusions and high intersite heterogeneity also implies population instability.

Parallels are evident too in the patterning of biological 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 account for the biological complexity: "The most important 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, because 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

separations in the Arawakan and Tupi-Guaranian stocks. The carbon-14 date of about 3000 years ago for the introduction of pottery making to the lower Amazon also falls within this time span. The spread of savanna-adapted flora 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 pottery to a non-Amazonian tradition suggests that it was introduced by immi- grants from a region of open vegetation, who were attracted into the lowlands during the most recent period of deforestation. Although there are no dates for the spread of the Cariban languages, the coin- cidence between their distribution and the area with the lowest rainfall and largest enclaves of savanna in modern Amazonia suggests an intrusion from the north during this period. Concurrently, the forest- adapted Arawakan and Tupi-Guaranian speakers apparently migrated in search of regions where their familiar habitat survived. Carbon-14 dates around the beginning of the Christian era for the earliest Tupiguarani-tradition pottery from the coastal Bra- zilian state of Parana agree with the lexicostatistical estimate of ? 1500 years for the separation of the languages in the Tupi-Guarani subfamily. Coastal sites are associated with forest vegetation and be- come progressively more recent from south to north. A forest origin and migration in search of an earthly paradise were a prominent part of Tupi-Guarani mythology at the time of European contact.

Variations in the antiquity and permanence of tropical forest vegetation in different parts of the South American lowlands might account for some of the differences in the degree of dependence on wild foods (other than game and fish) by groups practicing subsistence agriculture. Where nuts, seeds, roots, shoots, fungi, fruits, insects, and other such resources are important in the diet, it might be inferred that reliance on domesticated foods was com- paratively recent. Such a contrast exists between the Kayapo of southeastern Amazonia, who adopted agriculture when they moved into the forest in post- European times and who subsist entirely by hunting, fishing, and gathering during several months of the year (Meggers 1971: 70), and the Waiwai of southern Guyana, a Cariban-speaking group that makes minimal use of wild plants for food (Yde 1965: 67). The Yukpa-Yuko, who inhabit a forested area on the Colombia-Venezuela border, consume insects belonging to 22 genera and 7 orders (Ruddle 1973: 94).

Another indication of the wild-food potential is the availability in markets in the state of Para of some 70 species of native fruits (Cavalcante 1972).

This raises the question whether the persistence of hunter-gatherers in the tropical forest can be attributed 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 vicinity of the postulated forest refuges suggests they may have enjoyed the long-term habitat stability conducive 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 villages form a social unit within which marriages, festivals, raids, and other activities requiring interaction beyond the family level are arranged. This common foundation supports two very different configurations.

J ivaro 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 appears 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


significant disruption of the aboriginal way of life. It so happens that the Jivaro live in a region that probably remained forested during the most recent arid period, in which case they would have had am- ple time for cultural specialization. The fact that they speak a language with no known relatives is another indication of antiquity. The Cariban-speaking Waiwai by contrast are recent intruders in their present territory, and their linguistic affiliation suggests that they may be relative newcomers to Ama- zonia.

CONCLUSION Although the evidence from every discipline is minimal, the fact that existing botanical, zoological, and cultural data present similar characteristics and types of patterning suggests that they have a common ex- planation. The alternation between periods of forest

fragmentation and coalescence postulated by the biologists to account for species diversity would have posed subsistence problems for human beings adapted to the forest because of the relative paucity of savanna food resources. The inference that groups unable to remain in the forest refuges would have been obliged to adapt to the savanna or to search for other forested habitats is compatible with linguistic and archeological evidence for widespread prehistoric population movements. Lexicostatistical, archeological, and geological dates all indicate that a major disruption occurred between about 3500 and 2000 years ago. Although there is a great deal yet to be learned before the size, location, and duration of the refuges can be accurately defined, this model of environmental fluctuation provides a new and ex- citing basis for interpretation of the diversity and discontinuity that characterize the cultural evidence.

LITERATURE CITED

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

- 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 Gran

Chaco. 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, based

on 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 Nacional

de Pesquisas Arqueologicas; Resultados preliminares do primeiro ano, 1965-66. Publcoes avuls Mus. Paraense Emilio Goeldi 6: 129-151. Belem. . 1968. The Castanheira site: new evidence on the antiquity and history of the Ananatuba Phase (Marajo Island, Brazil). Am. Antiq. 34: 402-410. . 1974. Contribuiqao a arqueologia dos arredores do baixo Rio Negro, Amazonas. Programa Nacional de Pesquisas Arqueologicas; Resultados preliminares do quinto ano, 1969-70. Publsoes avuls. Mus. Paraense Emilio Goeldi 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. 31mO Congresso Internacional

de 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.


application of the biological model of diversification to cultural distributions in tropical lowland...

Travel

conditions of use

tropical lowlands

model of climatic

jstor archive

noncommercial use

lowland tropical forest

tropical biology

jstor transmission