Protein Capture and Cultural Development

in the Amazon Basin

DANIEL R. GROSS
Hunter College, CUNY
This paper examines the proposition that availability o f animal protein limits the
size, density, and permanence o f settlements o f the aboriginal societies o f the
Amazon Basin. Previous discussions have focused mainly on agricultural potential.
Evidence is presented from ethnography and ecology suggesting that fish and game
are scarce, particularly away from major rivers. Nine relatively unacculturated
societies have relatively low but probably adequate per capita intake o f animal
protein. The effects which this limitation may have on Amazonian culture are
discussed.

THERE IS WIDE AGREEMENT among anthropologists that a large, nucleated, sedentary

population is a necessary condition for the development of complex society. A number of
investigators have concerned themselves with the capacity of a given techno-environment to
support a population of sufficient size t o permit higher socio-cultural forms t o evolve. South
America has been an important testing ground for these ideas. Betty J. Meggers (1954)
suggested a relationship between attained levels of cultural complexity and the agricultural
potential of different regions of South America. In particular she argued that the thin, easily
leached soils of the Amazon basin limited cultural development to a tribal stage (1954). In
support of this assertion, Meggers cited the Marajoara culture which appeared to be intrusive
in its lower Amazon island habitat (1957). She interpreted the sudden appearance and
relatively rapid decline and disappearance of the Marajoara culture as indications that a
relatively complex society with temples, class differences, and craft specialization could not
sustain itself in the tropical forest.
The scant demographic data available from Amazonia seem to support Meggers
assertions. Taken as a region Amazonia upon contact with Europeans is estimated to have
had a population density no higher than l/km2 (Steward and Faron 1959:53), and even
today the density is not much greater. The population of individual settlements (nucleated
or not) is also small, judging from relatively recent data. Twenty-four of thirty Amazonian
societies listed in the Ethnographic Atlas (Murdock 1967:227-231) have a mean settlement
size of less than 100. Large settlements with nucleated populations of up to 1500 existed in
the past, particularly along the main stream of the Amazon (Metraux 1945:698; Denevan
1966). Nevertheless, Meggers is apparently correct that most Amazonian settlements were
quite small even before devastating European contact.
Robert Carneiro (1961) challenged Meggers 1954 views-which have since been
considerably amplified and modified (Meggers 1971). Using data from the Kuikuru of
Central Brazil and employing a formula he had derived to estimate the carrying capacity of
swidden agriculture (Cameiro 1960), he suggested that the Kuikuru had the agricultural
potential to maintain a sedentary village of up to 2000 with low labor inputs and without
degrading the environment (but cf. Street 1969). He presented other evidence to show that
tropical soils were, in some instances, capable of supporting densely settled populations.
Submitted for publication August 26. 1 9 7 4
Accepted for publication January 23, 1 9 7 5

526

Gross]

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

527

Cameiro concluded that low agricultural potential was not an obstacle to the development
of complex societies in the Amazon basin.
This study considers additional ecological factors which may have limited the size and
permanence of precontact Amazonian settlements. Following the suggestion of a number of
other investigators (Lathrap 1968; Denevan 1966; Meggers 1971; Cameiro 1970c; Spath
1971), I examine the evidence that dietary protein was limiting on aboriginal settlements
throughout most of Ammonia. I further suggest that small and frequently shifting villages,
warfare, and population control were adaptations to this limitation.
The principal source of calories for most Amazonian groups was and is cultivated crops.
In general, cultivators rely on vegetatively reproduced crops like manioc, sweet potatoes, and
the plantain, rather than on the more nutritious seed crops such as maize and beans which
are also known in the region (D. R. Hams 1971). Root crops and the plantain are
characteristically high in calories but low in protein and other essential nutrients (see Table
1).
Protein is provided primarily by meat and fish. Prior to European contact, no animals
were bred domestically for food. Thus the only sources of animal protein were the many
wild species inhabiting the forest and streams, especially fish, caymans, tortoises, snakes,
deer, sloth, monkey, peccary, tapir, capybara, armadillo, parrots, manatees, various water
fowl, and many others. The larvae and adults of many insects and turtle eggs are also
gathered, but there is little information as to how significant they are in the diet.2
The Kuikuru are perhaps typical in their reliance upon manioc for 80-85%of their diet,
other cultivated plants (including maize) less than 5%, fish 10-15% and meat, negligible
(Cameiro 1961).
Since they provide an important test-case, the Kuikuru may serve as a paradigm for
certain hypothetical exercises. Manioc, although it provides the bulk of the diet, cannot
begin to meet the average individuals requirement for dietary protein because of its low
TABLE I. NUTRIENT VALUES OF SOME TYPICAL SOUTH AMERICAN FOODS
PER 1 0 0 g EDIBLE PORTION

Food

Food Energy*
(kcal)

Protein*

3 20
97
122
116
79
100
129
194
101
146
144

1.7
1.2
1.0
1.3
2.8
2.0
4.1
17.5
17.9
29.5
11.3

Manioc Flour
Banana
Plantain
Sweet Potato
Potato
Yam
Maize (Whole Grain)
Pork
Fish
Venison
Fresh Hens Egg
*Source: INCAP-ICNND 1 9 6 1
tSource: F A 0 1 9 7 0
$Value for beef

(g)

Essential
Amino Acidst
(mg)
404
352
.-

414
667
821
3820
5203
8465
7875$
6338

Biologicalt
Value
-

66.7
59.4
74.0
76.0
74.3$
93.7

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AMERICAN ANTHROPOLOGIST

[77,1975

protein content (see Table I). The small trace of protein in manioc also has a low Biological
Value (a measure of efficiency of utilization of dietary protein). Thus the Kuikuru could ill
afford to consume less fish than they do (see below, p. 534). To increase their population,
the Kuikuru villagers would have to increase their production of protein. Were they to
achieve the asymptotic !eve1 of 2000 suggested by Cameiro, they would collectively require
about 100 kg of protein per day to meet a requirement for high quality protein of 50
g / p e ~ o n / d a y .This
~ might be derived from a daily average catch of 500 kg/day, or an annual
catch of 182.5 metric tons. Clearly, this level of production could not be sustained by
fishing lakes and streams close to the village, so fishermen would have to travel some
distance from their homes. The unit cost in mandays of work would increase rapidly with
distance away from the village. The area of rivers and lakes accessible to the Kuikuru is not
known, but let us make the following assumptions. Based on values calculated from Cable
(1971) and Hickling (1971), one may speculate that Upper Xingu rivers are capable of
reliably yielding 5000 kg/km2 /yr of fish. The annual fish requirement of 182.5 metric tons
would require that the 2000 Kuikuru have uninhibited access to 36.5 km2 of riverine
waterways, or about 365 linear kilometers of river channels averaging 100 meters in width.
Thus the distances involved would probably make it impossible, given the Kuikurus
technology, to exploit and defend this vast extent of river.
A group like the Kuikuru, then, with the technology and diet described by Carneiro,
could probably not sustain a healthy sedentary population of 2000, primarily because such a
village could not meet its daily protein requirements. The story is probably different for
groups living on the banks of large, highly productive rivers such as the Amazon, the
Araguaia, etc. If protein were a limiting factor on the size and density of a riverine
population with neolithic technology, it should be even more so for populations living away
from major streams. There are few studies on the abundance of animal species in Amazonia,
but the following data, culled from the literature, may be suggestive.
In general ecological terms, we may note that although the evergreen tropical rainforest
has the highest level of primary productivity of all terrestrial ecosystems (Odum 1971:51),
this abundance is not matched by the non-detritivorous fauna. Petrusewicz and Macfayden
point out that
despite a tenfold increase in solar energy reaching the earths surface in tropical forests,
the amount of energy available to secondary producers has hardly increased at all. This
appears to be largely due to increased respiration by tropical plants [1970:154-1551.
In one of the few holistic studies of living communities in the Amazon basin, Fittkau and
Klinge made the following observations on a section of rainforest about 100 km south of
Manaus:
The amount of animal biomass in the central Amazonian rainforest is extremely small
when compared with the plant biomass. . . the rarity of rodents can be established from
the fact that collections. . . had to be suspended because of the difficulty of securing the
animals. Hunting for human food is never effective (even by experienced
people). . . / . . . The small percentage of animal biomass compared with the total
biomass of the central Amazonian rainforest becomes much more evident if we compare
similar figures for African steppes and savannas, or for other tropical forests. In a
montane tropical rain forest in Puerto Rico, the animal biomass comprises 0.1% of the
total biomass, while in central Amazonia it comprises only 0.02% of the total
biomass. . . . in Ghana forests there are 0.72kg/ha of ungulates and primates [1973:8-91.
These authors also found that a majority of Amazonian fauna are detritivores feeding on
dead plant matter and fungi, while the amount of animals grazing on live plants is rather
small (1973:9). They conclude that The main energy flow is undoubtedly through the
detritus food chain (1973:9). They attribute this state of affairs t o the scarcity of nutrients
available.

Gross]

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

5 29

Odum and Pigeon (1970)studied a montane forest in Puerto Rico and reported an animal
standing crop of 39,300 kg/km2. But of this total, only 17% consists of species ever eaten by
humans and less than 1% are birds and mammals. In perhaps the most thorough census of
animals in a neotropical forest to date, Eisenberg and Thorington (1973)provide a value of
4400-5300 kg/km2 for nonvolant terrestrial mammal biomass on the protected Barro
Colorado Island, Panama. If these animals could be harvested by predators at the relatively
high rate of 20% of biomass per year without depletion, such a biomass would yield about
1.2-1.45kg/km2 /day of meat.
The proportion of this meat going to any one predator would depend on its numbers and
relative efficiency. Hesse (1937:430) observed that, animal populations in the tropical
forests are usually larger than is assumed because so many animals are either hidden in trees,
burrowed or buried in the ground or are nocturnal. Indeed, in the Eisenberg and
Thorington study (1973),27.4% of the mammalian biomass is exclusively nocturnal, and
72.3% is entirely arboreal. Human predators are relatively disadvantaged because their size,
anatomy, and sense organs equip them poorly for locating and capturing prey. Consider, for
example, the highly abundant three-toed sloth (Bradypus infuscatus, 48% of mammalian
biomass on BCI): In spite of its high density, the animal is so cryptic that it is rarely seen
and impossible to census visually (Eisenberg and Thorington 1973:156).Thus humans can
expect but a small share of whatever meat may be available to predators in the tropical
forest.
Land hunters armed with bow and arrow or blowgun, and perhaps poisoned projectiles
must spend a high amount of energy per unit food energy captured as meat, compared to
fishermen or animal domesticators. Many studies attest to diminishing returns on hunting
effort in the area surrounding a given settlement (see Johnson 1974 and Cameiro 1970c for
constructive suggestions for dealing with this problem).
But hunters who move frequently may encounter another problem as they travel from
one area of tropical forest to another: different species may require changes in hunting
techniques. The density of any particular species is usually quite low and its distribution
may be patchy and unpredictable (Diamond 1973). J. Haffer (1974)suggests that the
uniformity of the dense forests presently blanketing much of Amazonia may be deceptive.
He explains the discontinuous distribution and high species diversity of avian fauna over
much of Amazonia as the result of climatic fluctuations which periodically created isolated
forests (refugia) within which speciation took place. The last severe arid phase in Brazil
occurred between 4000 and 2500 B.P. (Haffer 1974:142). Diamond (1973)points out that
tropical fauna are often non-migratory, so that areas left vacant may fail to be colonized for
long periods. These factors would increase the risk of hunting out a particular area
subjected to prolonged h ~ n t i n g . ~
Certain widespread areas of Amazonia may also be particularly poor in both fish and
game because of a phenomenon described by Daniel Janzen (1974).In these areas of white
sandy soils (of sedimentary origin), primary productivity may be exceptionally low, limited
by low availability of nutrients in the soil. In such mas, plants are highly susceptible to
damage by grazers, a condition which may have led to their being exceptionally high in
chemical defenses, particularly phenolic compounds in leaves. These leaves, when deposited
on the forest floor, decompose very slowly, releasing their poisons t o runoff and ground
water as they do. Food chains in such areas tend to be truncated, limited to species which
are able t o detoxify the phenolic compounds from their food. Rivers are particularly
affected. Such rivers are characteristically filled with dark colored water which is low in
dissolved oxygen and suspended matter, making them extremely inhospitable t o fish. Thus,
according to Janzen, these black water rivers may flow through areas virtually devoid of
animal life (cf. Sioli 1964;Fittkau 1970).
In the literature on Amazonia, there are many statements by zoologists, explorers,

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AMERICAN ANTHROPOLOGIST

[77,1975

ethnologists, and others to the effect that game is scarce, at least in certain areas. J. Wilbert,
an ethnologist, states that the territory of the Yanomamo of southern Venezuela and
northem Brazil is now virtually devoid of animal life . . . (1972:15). N. Chagnon, writing
about the same area, reported that
game animals are not abundant, and an area is rapidly hunted out, so that a group must
keep constantly on the m o v e . . . . I have gone on five-day hunting trips with the
Yanomamo in areas that had not been hunted for decades, and had we not brought
cultivated foods along, we could have been extremely hungry at the end of this time
[1968b:33].
It is noteworthy that until very recently, the Yanomamo territory had been hunted with
indigenous weapons only, not with firearms.
The upper Xingii region on the Central Brazilian Plateau was visited by a Brazilian
scientific expedition in 1948, also before substantial numbers of firearms had been
introduced. A zoologist accompanying the expedition observed,
Our attention was drawn to the fact that the Indians burned off the surrounding
savannas.. . . This practice impoverishes the savannas and its outcome is very
evident., . . the fauna of the savanna already naturally poor, is even further reduced,
reaching a point such that in some places it is possible to travel over an appreciable area
without meeting with so much as a spider. . . [Carvalho 1949:s; emphasis added, my
translation].
Thousands of miles to the Northwest in Ecuador, in Jivaro territory, M. Harner observed
that
a great diversity of botanical and zoological species exist in this region, but this variety is
accompanied by low density of population for any particular species. This low density
sometimes poses a problem for Indians engaged in collecting a specific kind of wild fruit
or hunting a particular species of game. This situation is aggravated for the hunter by the
fact that virtually all of the Jivaro territory has been hunted efficiently for a long period
of time, with the result that game is not as abundant as in regions unoccupied by the
Indians [ 1972: 55-561.
Allan R. Holmberg, writing of the Sirion6 of the tropical forests of Eastern Bolivia, states
that, Game is not plentiful (1969:70), and that, it is not uncommon for the Siriono to
go for several days at a time without eating meat (1969:73). At least 25 percent of the
time [the hunter] returns to camp empty-handed, or with insufficient food to completely
nourish his family . . . (1969:249). Writing on the Vaupes region of Colombia and Brazil,
Silva states that The fauna of the Northwest Amazon is poor, qualitatively and
quantitatively, in animals for food [my translation] (1962:242).
To be sure, other sources can be found attesting to plentiful game resources in some
places (e.g., Cameiro 1970a) but most of the accounts I have seen by persons who have
spent long periods with indigenous groups, hunting by traditional means or with firearms,
seem to agree with the statements recorded above.
Another means of trying to get at the availability of protein in Amazonia is to estimate
the contribution of animal protein to the diet from the scant data available on fishing and
hunting by native peoples. Table I1 shows such estimates for nine relatively unacculturated
tropical forest societies in the New World. Note that one case (Miskito) is Central American,
and another (Kaingang) is located in semi-deciduous forests, but all are roughly comparable
in other respects, e.g., faunal species present, type of forest cover, etc. (See Appendix for a
discussion of how the data were estimated.) In each one, a number of assumptions was made
(see Appendix), but every effort was made to squeeze the maximum precision from
imprecise data and to avoid underestimating protein yield. If anything, the values stated for
animal protein capture are too high because of what I suspect to be systematic sampling
error (see Appendix).

E. Peru

Panama

Eastern
Nicaragua
Southern
Brazil
Eastern
Bolivia
GuyanaBrazil
E. Peru

E. Peru

GuyanaBrazil

Campa

Bayano Cuna

Miskito

Sharanahua

Shipibo

Waiwai

interfluve
forest
riverine
forest
tropical
forest (coastal)
su b-tropical
forest
interfluve
forest
riverine
forest
riverine
forest
riverine
forest
riverine
forest

*See Appendix f o r sources and details of calculations.

Wayana

Siriono

Kaingang

Location

Society

Habitat
T Y Pe

54

35 g
mean =

48

63

31

44

19

20

18

15-20

Animal
Protein
per capita
Per day (9)

manioc

plantains

+
-

manioc

maize and
manioc
manioc

pine nuts

manioc
(72%)
bananas
(89%)
manioc

Firearms
Present?

Principal
Food
Staple

varies

365

28

365

92

38

365

14

Length
of Study
(days)

77

107

90

16

60-154

106

997

170

5-15

Average
Size of
Pop. Unit

TABLE 11. ESTIMATED RATES O F ANIMAL PROTEIN CAPTURE FOR NINE TROPICAL SOUTH A N D CENTRAL AMERICAN SOCIETIES*

532

AMERICAN ANTHROPOLOGIST

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With all the roughness and inconsistency of measurement, it is remarkable how little
variation there is among the cases. The highest value (63 g) is less than twice the mean and
the lowest (15 g) is just under half. Most nutritionists agree that a daily intake of 63 g of
animal protein would be adequate, but that the lowest value would not be unless
supplemented by high-quality proteins of vegetable origin. The data suggest that the protein
intakes of native Amazonians approach or fall short of a minimal acceptable level. Fairly
small changes in the availability of dietary protein could be quite crucial to the health status
of these groups. Of course, these values are rates, not necessarily potential maxima, but there
are indications that most of these groups would eat more fish and game if they could. In
fact, one could state that, in their food preferences, these peoples show a healthy regard for
foods with high quality protein. Later, I shall marshal1 evidence t o show that actual cultural
practices may be seen as responses to potential protein scarcity.
A good deal of anecdotal evidence exists suggesting that native peoples feel meat is scarce
or lacking in their diets. They express this in preferences for meat over other foods,a special
sense of hunger which refers especially to meat, and a tendency for women to grant or
withhold sexual favors or approval in accordance with a mans ability as a hunter. For
example, Jules Henry reports that, to the Kaingang, meat is the principal article of diet,
everything else is garnish (1964:159). Robert Carneiro points out that meat, abundant in
the Amahuaca territory, is an important part of the Amahuaca diet, and no meal is
considered really complete without it (1970a: 332). Allan Holmberg observed of the Siriono
of Bolivia, Although meat is the most desired item in the diet of the Indians, it is by no
means the most abundant (1969:76). For the Yanomamo of Southern Venezuela, Meat is
always the most desirable food and is always considered t o be in short supply (Chagnon
1968:91). William Crocker reports that among the Canela (Timbira) of Maranhio that The
ordinary phrase to express hunger is ii.md pliim, or literally, in me need, but hunger for
meat has a special term, iiyate, I am hungry for meat (1972:258). Among another
G-speaking group inhabiting the cerrado in Central Brazil, the Shavante, David MayburyLewis observed that Meat far and away transcends other forms of food in the Shavante
esteem and in their conversation (1967:36). In her research in Eastern Peru, Janet Siskind
found that
The Sharanahua are continually preoccupied with the topic of meat, and men, women
and children spend an inordinate amount of time talking about meat, planning visits to
households that have meat, and lying about the meat they have in their households
[ 1973a:84].
Alongside this preoccupation with meat, we find that prestige and other goods accrue to
the good hunter. Siskind observes that Sharanahua women not only goad their men into
hunting for meat but also bestow or withhold sexual favors in accordance with a mans
hunting prowess (1973b). She generalizes this to other societies:
Pride in hunting is a benefit to the society, and it is rewarded . . . by prestige . . . . Prestige
is not a vague goal. . . it brings a definite reward, the possibility of gaining women as
lovers and/or wives. It is a common feature that the Sharanahua share with all tropical
forest hunters: The successful hunter is usually the winner in the competition for women
[1973a:95-96].
Among the Siriono, Allan Holmberg reports that,
Not infrequently the unlucky hunter, while resting from an unsuccessful chase, is
reproached by his wife for not having brought home more game, and, invariably as one
leaves for the hunt, the women and children call after him such commands as, Bring me
back the leg of a peccary, or Bring me back some tapir meat [1969:71].
Shavante women appear to be no more understanding than the Siriono for they receive an
unsuccessful hunter with a marked coldness, even when there is plenty of other food . . .
(Maybury-Lewis 1967:36).

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PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

533

Not only marital but also extramarital delights await the successful hunter:
Extramarital sex partners [in SirionB] are seduced primarily through rewards of
food. . . . This is clearly observable among the women who prefer good hunters to all
other sex or marital partners [Holmberg 1969:255].
There is even a cultural analog to the telltale lipstick stain on a western husbands collar:
In general, the [SirionB] wife supervises the distribution of meat, so that if any part of
her husbands catch is missing, she suspects him of carrying on an affair on the
outside. . . . [Holmberg 1969:167].
Elsewhere, meat is exchanged for extramarital sex among the Mehinacu of the upper Xingu
(Gregor 1973:245) and the Canela (Crocker 1972:258).
To summarize, there is some evidence to support the conclusion that hunting and fishing
productivity are limiting on the size and permanence of Amazonian settlements. There are
studies by ecologists and other naturalists showing that game is scarce in the tropical forests
of South America. There are estimates of animal protein availability in some Amazonian
societies which tend to be suboptimal when measured against minimum recommended
allowances. Finally, there are various cultural manifestations of a cognized sense of meat
scarcity in a number of Amazonian groups.
In view of the low availability of animal protein in Amazonia, we may inquire why other
resources were not exploited or other techniques not developed, e.g., animal domestication
for food, or increased use of protein-rich cultigens. Protein intake from meat and fish, at the
rates indicated in Table 11, would probably not be cause for alarm in most European or
North American populations. This is because additional foods are present in the diets of
these people which bring protein up to high levels. In particular, most Westerners daily
consume foods containing substantial amounts of cereal grains, milk, and eggs. All of these,
particularly milk and eggs, are high in essential amino acids (see Table I). In Amazonia, the
principal staples of horticultural groups tend to be starchy root crops or (since contact)
plantains, foods with low protein content of low Biological Value (i.e., low in essential
amino acids). Milk, except for mothers milk, was not available in precontact Amazonia.
While turtle eggs are avidly consumed by Amazonian peoples, these are available only during
relatively short seasonal periods. Meat and fish are thus all the more crucial, especially for
small children whose growth requires protein but whose size does not permit them to ingest
the large volume of vegetable food necessary to achieve adequate protein nutrition (BChar
1968; see note 2).
One can only speculate as to why, over the long period of Amazonian prehistory,
productive techniques were not adopted which might have yielded larger and more reliable
protein sources outside the major riverine locations. One alternative might have been to
domesticate wild animal species for food. Lowland South Americans often kept birds,
monkeys, and dogs as pets but not for food. Land tortoises and peccaries were sometimes
captured alive and held for later consumption, but they were never bred in captivity
(Bennett 1962:40n; Steward 1945:18). If peccaries could have been bred in captivity
(Gilmore 1948:382), they might have provided small but significant increments in protein,
as seems to be the case with New Guinea pigs (Rappaport 1968:74-87). Peccary breeding
would have led t o a decline in mobility and an intensification of agricultural effort in order
to feed them. But these are the very features which are dependent variables in the present
analysis. In any case, the required kick in the productive system of lowland South
Americans which might have impelled them in this direction did not occur. Perhaps rising
population densities in relatively circumscribed locations (Cameiro 1961, 1970b) would
have been a sufficient condition for animal domestication, but I am inclined to doubt it
since nowhere is this reported to have occurred.
Another alternative which would have provided more high-quality protein would have

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AMERICAN ANTHROPOLOGIST

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been greater reliance on seed crops as a dietary staple. Compared to meat, maize has
relatively low Biological Value; its amino acid composition is nevertheless superior to manioc
and plantains (Table I). A high maize diet, with alkalai processing, and relatively small
supplements from animal protein and beans, provides a fairly high quality diet for older
children and adults (Katz et al. 1974; Bkhar 1968). There is some evidence that lowland
Maya civilization was based on such staples (Turner 1974; BLhar 1968; but cf. Puleston
1971; Bronson 1966). High dependence on maize has been observed only rarely in
Amazonia, although maize is widely cultivated as a supplemental crop.6
A few tentative reasons may be advanced as to why vegetatively reproduced plants were
preferred over more nutritious maize: (1)Maize is less productive of calories per unit area
and per unit labor than manioc (Carneiro 1961). (2)Manioc seems to be a more reliable crop
than maize. The latter may be more susceptible to poor drainage, rainfall fluctuations, low
soil fertility, etc., than root crops (D. R. Harris 1971). Harris also observed that South
American tropical swiddens based on vegetatively reproduced species tend to have greater
species diversity and a more completely closed canopy of vegetation, exposing the soil to less
cf.
of the destructive influences of sunlight and rainfall than do seed crops (1972:253-254;
Geertz 1963:24-25).(3)Maize may be more difficult to cultivate in the humid tropics
because it is
more dependent than manioc on the efficiency of clearing and burning. . . [ A ] s the dry
season progressively diminishes in duration, intensity, and regularity toward the equator,
the task of burning the cleared vegetation thoroughly to ensure an adequate yield of
maize and other seed crops becomes progressively more difficult [D. R. Harris
1971:495].

Root crops, like manioc, which mature more slowly can draw nutrients from the slowly
decomposing litter left on the surface by a partial burn, while maize requires a more
concentrated dose of nutrients. (4)Finally, manioc has superior keeping qualities, both in
the ground where it may be kept indefinitely (barring inundation), and in the form of
processed flour. Maize, by contrast, must be harvested soon after maturing and is more
susceptible to mildew and pests (cf. Spath 1971;Puleston 1971).
High dependence on so-called bitter manioc may aggravate the potential scarcity of
animal protein in the Amazon area. Bitter manioc is preferred over sweet varieties because of
its superior keeping qualities in spite of its high content of toxic cyanogenic glucosides. No
matter how efficient the processing of the tubers, these compounds may remain in
concentrations which are dangerous to health, particularly where large amounts are ingested.
Dietary cyanides may be detoxified by the body, but this requires the presence of amino
acids containing sulfur, cystine, and methionine. These are particularly lacking in starchy
root crops. One kilogram of manioc flour contains about 450 mg of sulfur-containing amino
acids (FA0 1970)while the average minimal daily requirement for adult males is set at 1100
mg/day (Hegsted 1964:148).This requirement could be met by about 150 grams of beef,
fish, or eggs (FA0 1970). Thus upland tropical human communities forced to rely heavily
on bitter manioc by soil and climate conditions are particularly dependent on reliable
sources of animal protein (Spath 1971; Lowenstein 1973:306).The symptoms of manioc
poisoning (visual difficulty, burning, prickling of the skin, impaired hearing, and even
crippling) have been observed in Nigerian populations depending heavily on manioc
(Osuntokun et al. 1969), but not, to my knowledge, in Amazonia. What is ultimately
limiting, if the above is correct, is the amino acid methionine (Spath 1971).
The most remarkable thing, perhaps, is that the peoples of the Amazon basin have
adapted so well to an environment with relatively poor soils, few sources of animal protein,
cultivated foods low in essential nutrients, and even high toxin levels in the principal staple.
Perhaps the best evidence for my claim of a successful adaptation to protein scarcity is the
fact that symptoms of protein deficiency disease have never, to my knowledge, been

Gross]

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

535

reported for relatively unacculturated tribal peoples in the Amazon region. Health and
nutritional surveys have reported varying levels of illness, including especially introduced
respiratory diseases such as tuberculosis, measles, and influenza, among others. Some
deficiency diseases, particularly goiter, have also been reported, but I have seen no reference
to marasmus or kwashiorkor (see, e.g., Lowenstein 1973; Nee1 et al. 1964). How may we
account for the apparent absence of protein deficiencies among the native peoples of the
Amazon basin?
A number of cultural mechanisms may have been favored by selection to enhance a
societys adaptation to low protein availability, especially scarcity of game (cf. Meggers
1971:97-113). Among these are the following: (1) Maintenance of small settlements which
minimizes the impact of human predation on fish and game within the area accessible to
settlements. Some cultural traits favoring small settlement size are lack of strong political
leadership, disputes over women, witchcraft accusations, and seasonal dispersals, all of which
favor the fissioning of settlements. (2) Dispersion of settlements, as opposed to clumping, to
avoid overlapping areas of exploitation. This is favored by warfare where a constant danger
of attack exists. (3) Maintenance of no mans lands between settled areas forming
preserves where prey species may reproduce unhindered by human predation. This is also
favored by raiding. (4) Frequent movement of settlements to avoid overexploitation. This is
favored by warfare, fission, and the practice of shifting agriculture. (5) Low rate of
population growth which eases the rate of growth of pressure on resources. Some traits
favoring low natality are infanticide, particularly female infanticide which lowers the
number of reproducers, abortion, contraception, and taboos on sexual intercourse (cf.
Whiting 1964).
Many of these practices are found in Amazonian societies and often they are found
together. Some reinforce each other or, at least, are compatible. For example, in his
ethnography of the Yanomamo, Napoleon Chagnon shows that polygyny and infanticide
create a scarcity of women, promoting disputes which may lead to village fissioning, and
ultimately to warfare between villages, which, in turn, precipitates relocations to distant
areas to escape attack (Chagnon 1968a, 1968b, 1973). For a population of a given size,
female infanticide may change the sex ratio in favor of males, satisfying the demand for male
warriors (Divale and Harris 1974). Long post partum sex taboos are linked culturally to long
periods of lactation which, in turn, may be a direct response to low protein availability
(Whiting 1964; G h a r 1968). While Whitings suggestion that polygyny is causally linked to
the sex taboo may be implausible (Ember and Ember 1973:337), it does seem plausible that
such a taboo could encourage philandering which creates disputes over women.
Before concluding, we must return briefly to the subject of tropical horticulture since it is
intimately related to the total subsistence system of any Amazonian group. Recent studies,
beginning with Carneiro (1961), have stressed the variety and flexibility in tropical
agriculture. Some authors still emphasize limitations inherent in tropical soils (Meggers 1971;
D. R. Harris 1972). This may be particularly appropriate for the Amazon basin where
enormous areas are covered by Orthos soils of the Oxisol order, derived from very old parent
material, poor in nutrients and low in organic matter (Aubert and Tavernier 1972:18, 28).
Therefore, comparisons to other tropical areas like Nigeria or New Guinea may not be
appropriate. Nevertheless, recent studies in the American tropics, New Guinea, and Africa
have suggested that techniques such as ridging, mounding, and terracing permit greater
capacity for intensification in tropical agriculture than heretofore accepted among students
of the subject (Denevan 1966a, 1970; Turner 1974; Waddell 1972; Netting 1968).
Granting that agricultural potential does not limit size and density of Amazonian
populations, we must look to other limiting factors. This paper focuses on protein as a likely
one, although others may also be responsible. But even protein does not limit population
size and density under all conditions. Incorporation of Amazonian societies into larger

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AMERICAN ANTHROPOLOGIST

[77,1975

polities could effectively block the adaptive responses to local protein scarcity discussed
above. Such polities are unlikely to have arisen autochthonously in Amazonia for reasons
discussed above. Since 1500, modern states have suppressed warfare, established missions,
and confined native peoples to reservations. Commercialization has transformed tribal
Amazonians into rubber gatherers, Brazil-nut gatherers, pelt hunters. All these changes tend
to sedentarize and concentrate populations in a manner prejudicial to the optimal capture of
wild protein resources. Thus contemporary Amazonia illustrates the process whereby small,
relatively autonomous systems are deprived of their capacity to respond to local fluctuations
by incorporation into larger, more coherent politico-economic systems (cf. Lees 1974).
For those populations remaining substantially free of state controls, shifting cultivation is
advantageous, not only as a source of food requiring low effort, but also with regard to
protein scarcity. This is because abandoned swiddens may help to attract and maintain
certain game populations, at least temporarily, in areas close to human settlements. Whether
because of weed growth or soil exhaustion, swiddens in Amazonia are rarely cultivated for
more than two years. A small settlement may create hundreds of hectares of secondary
growth in ten years. Secondary growth forests are preferred habitats for a number of animal
species (including invertebrates) some of which are rarely found in mature forests (Beebe
1971, Ch. 5; Richards 1952). The earlier stages of succession in tropical seres abound in
grasses and shrubs, in contrast to the tough woody stems of the mature forest. With
reference to domesticated grazers, G. W. Arnold states
physiologically younger material may be preferred because it is generally shorter, or
because it differs in chemical composition from older material. The selected material
is . . . usually higher in nitrogen. . . phosphate.. . sugars. . . and gross energy
[1964:135].

Ground grazing species like the peccary or deer will find more promising leafy material and
even tubers in an abandoned swidden than in a mature forest where so much of the plant
biomass consists of upper story leaves and woody stems. A few examples are pertinent.
Bernard Nietschmann (1972:57) observed that for Miskito hunters in Eastern Nicaragua,
palm swamps, old and new swiddens, and secondary forests had the highest return of
meat. Whitetail deer is hunted particularly around abandoned swiddens (1972:51). Not
only abandoned swiddens but also burned-over savannas or cerrado may serve to attract
game animals. H. Baldus reports that the TapirapC burned off grassland areas as they made
their way to the river, explaining the practice as a means of clearing their path for the trip
back. Brazilian backwoodsmen and Karaji Indians, however, stated that this practice would
attract deer who prefer the fresh young grasses (Baldus 1970:174). Other groups living in
non-forest areas practice communal surround hunts with the use of fire, a technique that
could have a similar effect. The burning and clearing of land for horticulture or other
purposes may also be seen as responsive to protein scarcity.
DISCUSSION

I have argued that amino acids, more than calories or social dynamics, are limiting on the
size and permanence of native settlements in Amazonia. Below, I discuss some possible
implications of this view.
(1) The Amazon basin is not a homogeneous habitat from the point of view of human
utilization. Soils, fauna, flora, rainfall, and seasons vary considerably. At least three broad
types of habitat may be distinguished: (a) Riverine habitats, characterized by relatively
fertile soils, replenished by alluvial deposits during river flooding. They are also relatively
rich in animals, particularly aquatic ones. Areas drained by black-water rivers will conform
to this generalization to a lesser extent, depending on the concentration of the life-inhibiting
compounds in the soils and waters. (b) Interfluvial or upland habitats in forested areas away

Gross]

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

531

from major streams, characterized by relatively impoverished soils, easily leached, and
relatively limited in animals sought by humans, especially aquatic ones. (c) Non-forested or
savanna habitats,8 characterized by low dry-seaon rainfall, poor soils, scrub or grassy
vegetation, interrupted by sinuous bands of gallery forests along watercourses.
(a) Donald Lathrap (1968:26) has suggested that the earliest inhabitants of Amazonia
were riverine horticultural peoples and that the upland hunters- of recent periods are
formerly horticultural peoples pushed out of the prime river-bank areas by successive waves
of militarily superior immigrants. In spite of the low average availability of protein which my
data suggest, I do not believe that Amazonia was uninhabited before the arrival of
horticulture. It seems possible that Amazonia was once settled thinly but extensively by
populations subsisting on wild plants, fish, and game. Fish and game were probably more
available and plentiful since nomadic food collectors would not have had to compete with
more densely settled horticulturalists for these resources. In other words, the carrying
capacity for hunters in Amazonia is probably low, but it is not zero. Most groups are
probably stabilized at levels below the capacity of their local habitats to provide protein,
even sedentary horticulturalists.
I am more amenable t o Lathraps suggestion of competition among riverine groups for
control over the highly productive riverine habitats with their ample protein resources and
annually replenished soils. Archaeological research should eventually be able to confirm or
disconfirm military competition and succession. The relatively sharp ecological boundaries
of these habitats might have provided the circumscription which Cameiro (1970b) saw as a
condition of the rise of centralized polities. Although it is now recognized that some groups
like the Tapajos and the Omagua had large settlements and complex social systems with
apparent social stratification, craft specialization, priest-temple-idol cults, etc., the annual
flooding, the seasonality of resources and perhaps the scarcity of accessible and defendable
habitation sites may have set a limit on the size of groups occupying the banks of the
Amazon (Meggers 1971). Nevertheless, it is evident that such groups occupied considerable
portions of the lower and middle Amazon and sociologically were qualitatively different
from most other Amazonian societies.
(b) In the vast uplands, covering an area far greater than the riverine lowlands, I suggest
that the availability of protein from all sources placed a low upper limit on population size
and density and permanence of settlement. Higher levels of political centralization would be
unlikely to have emerged here. Some settlements may even have depended on trade with
riverine groups as a source of protein. Archaeological studies may reveal the existence of
satellite groups or trading partners in upland areas with links t o specific riverine groups. One
may speculate that stone, stone tools, pottery, baskets, bows and arrows, and items of
magical and aesthetic use would have been exchanged for fish. I would expect relations of
dominance and subordination between riverine and upland groups. There is some suggestion
of this in Goldmans account of the riverine Cubeo and the upland Macu (1963:105-107,
292). Trade could have taken place on an individual basis, but t o the extent that a riverine
group was ranked socially, trade was probably centrally controlled. The social organization
of the upland groups was much more egalitarian than among the riverine groups.
(c) In the unforested areas, particularly in Central Brazil, a different pattern existed,
partly as a result of the peculiar pattern of protein availability here. Many of these groups
subsisted primarily by hunting and gathering, and there are some indications that game was
more plentiful (at least easier to find) on the open savannas than in the forests (Wagley and
Galvi5o 1945:169; cf. Hopkins 1967). Horticulture was practiced by most of these groups,
but it was restricted to the limited gallery forest lands, and thus was not productive enough
to support large settled populations the year round. Many of these groups were sedentary
only during the period following the harvest. For the rest of the year, the large villages (up
to 1200 in population) dispersed into smaller, functional hunting units (much like the bison

538

AMERICAN ANTHROPOLOGIST

[ 77,1975

hunters of the Great Plains in North America) in order to exploit the game of the savanna.
The social complexity of the Central Brazilian groups may be due to alternation between
dispersed, mobile groups and large village aggregates. In particular, the circular village plans,
crosscutting moieties, elaborate systems of name transmission, age grades, ceremonials, and
sporting events for which these societies are known, may have served as cultural means for
integrating semi-autonomous foraging units into unitary, annually reconstituted villages,
providing for the control of conflict between groups, the distribution of garden produce
throughout the village, and the mobilization of warriors for defense and raiding (Gross
1974).
(2) The three habitats distinguished may help to elucidate some anomalies of
contemporary ethnography. As Lathrap (1968) suggested, a number of societies known
ethnographically may be former riverine groups driven into upland areas where they
dispersed into smaller villages undergoing depopulation and deculturation. The Amahuaca
(Cameiro 1964), the Isconahua (Whiton et al. 1964) and the Sirion6 (Holmberg 1969) may
be such groups. The distinguishing characteristic, in the absence of some direct evidence,
may be traces of former social complexity, e.g., inherited chieftainship, not ordinarily
associated with such small, simple societies.
Other upland groups may have moved recently to the edges of rivers, following the
destruction of riverine peoples by warfare, introduced disease, and exploitation brought by
national populations in recent times. This process was certainly hastened by the great rubber
boom of 1882-1910 which sent hordes of rubber tappers into the forests near rivers to
extract latex from Hevea brasiliensis (Murphy 1960; Wagley 1964). Some examples of such
groups may be the Yanomamo (Lizot 1971; Chagnon 1968b), the Sharanahua (Siskind
1973a:39-41), and the Tukuna (Nimuendaju 1945:713). These societies, in contrast with
those above, may exhibit characteristics less appropriate to complex sedentary life than to
more migratory upland life. They may lack the ability to construct water craft and fishing
techniques, indicating their recent arrival in riverine habitats.
(3) The foregoing may also have some relevance for the study of warfare. Prior to
European penetration, I suggest that two fairly different patterns of warfare existed in
Amazonia. The first was a relatively organized form of military campaign, perhaps involving
multilocal mobilization of warriors, and having as an outcome the occupation and control of
territory. This pattern could have been beneficial to groups in the context of a circumscribed
riverine habitat depending on fishing and farming of alluvia (cf. Vayda 1961). There is some
indication that such warfare patterns prevailed among the Omagua of the Central and upper
Amazon (Metraux 1945:689) and the Tupinambi of Brazil's East Coast (Fernandes 1952).
The second pattern of warfare, characteristic of upland areas, was the organized raids
and counter raids motivated by vengeance, sorcery, headhunting or the capture of women
which was virtually ubiquitous in the Amazon basin. This pattern probably did not have the
consequence of the taking and holding of territory for agricultural purposes (Chagnon
1973). Given the shortage of game in upland areas, a more important outcome of such
raiding might have been the dispersion of settlements over wider areas and a consequent
relief of pressure on game resources.'
CONCLUSION'

I have suggested that the size, form, and permanence of settlements, social complexity,
and warfare patterns may vary in the Amazon basin with differences in the availability of
animal proteins in the diet. The forms observed were not the only forms which are possible.
A number of processes could have yielded different outcomes. For example, greater reliance
on a seed crop or domestication of a native animal species for food could have had striking

Gross J

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

539

effects. The encroachment of powerful territorial states from outside can and did have
striking effects on local populations in terms of the variables discussed here. The imposition
of wages, contracts, rents or tributes can lead a population to exceed crucial tolerances of
parameters like fallow periods, intensity of hunting, or land clearance, leading to such
responses as malnutrition, migration, or depopulation. Cultural practices which had, as
consequences, the prevention of such excesses may therefore be characterized as the
unconscious wisdom of a people, even though the particular practice might appear
irrational t o the ecologically unsophisticated observer.
It has become fashionable in recent human ecological thought to downgrade the
importance of the natural environment in favor of other factors taken as independent
variables such as demographic change, social dynamics, or even the ideological environment. These are taken as though they were somehow exogenous to the population under
consideration. This is unfortunate, first of all because it tends t o blur the concept of
environment, making it less useful for ecological analysis. Secondly, this approach may
prematurely deprive us of one of the few firm bases on which to begin analysis of a
sociecultural system. It is undeniable that the natural environment has often been dealt
with simplistically and that specific kinds of population interactions with the environment
are more important than the environment taken alone. But the environment, nevertheless,
does provide a set of limits and possibilities that is external to the population itself. This is
true, even when we deal with the application of modern industrial technology to the tropics
(cf. Janzen 1973).
In the case under discussion, I have suggested that the Amazonian environment imposes
limitations on cultural development of a society dependent on a neolithic technology. That
the indigenous populations of South America may have developed mechanisms which
prevented their degrading their environments is a tribute not so much to their rationality as
to their responsiveness to signals and cues in the environment comparable to that possessed
by other animal populations. Recently, many of these populations have been compelled to
abandon or modify these practices. The results have been, in many cases, malnutrition,
disease, runaway population growth, etc. As the large, politically centralized societies
extended their control over smaller, autonomous groups the latter have been forced to
ignore the accumulated wisdom of their ancestors.
APPENDIX
This section explains how the values for daily per capita protein intake in Table I1 were
obtained. Although it is less desirable, a per capita value was calculated rather than one
based on body weight or the actual distribution of age and sex in a population. This was
done for the sake of uniformity since the latter data were not available for the cases used.
Including infants below one year of age has the effect of lowering the overall average value
since most of them would not ingest protein of animal origin apart from mothers milk.
Nevertheless, I believe that there may be a systematic bias which inflates the estimates of
animal protein in the diets of these groups. This is so because in all but three cases (Wayang,
Miskito, Shipibo) the measurements were made during periods of normal hunting activity
and do not reflect the decline in hunting and fishing which usually occurs during the rainy
season and during the clearing and burning of new swiddens. No attempt was made to
correct for this probable bias in the measurements. Thus the data do not reflect seasonal or
annual variation in protein capture rates.
As will be seen, the calculations depend upon a number of assumptions, e.g., the
comparability of average size for species from one area to another. Obviously, it would be
far more desirable t o work with more and better data in arriving at these estimates. One
hopes that such data will be collected by field workers in the near future. Nevertheless, in
working with these scattered and imprecise data, the values for protein capture were
remarkably consistent with each other and with the expectations discussed in the text.
As an aid to estimating rates of protein capture I offer the following formula:

AMERICAN ANTHROPOLOGIST

540

Q =

[ 77,1975

D~ x . 2 n ~ ~
TP

where

Q = protein available from meat and fish per capita per day in grams;
D = number of man days hunting or fishing for period T ;
I$=
the dressed weight in grams of the average days catch of one hunter or fisher
for period T ;

T = length of observations in days;

P = the average population subsisting on the meat and fish obtained over period
T.
The constant (0.2) is a rough value indicating the protein yield from any given quantity of
fresh, dressed meat or fish. This was obtained by inspecting the values for protein in 100 g
edible portions of several kinds of meat and fish in a food composition table prepared for
use in Latin America (INCAP-ICNND 1961; see Table I). The formula may appear
unnecessarily complicated by separating the values Dt and M . They are presented as such to
t
provide for those cases where the investigator cannot monitor all the yields of all hunting or
fishing activities. If total monitoring is feasible, the numerator may be expressed simply as

.2Ft where Ft

to population P over period T.

In the sections which follow, I shall explain the derivation of a value for Q for each case in
Table 11. The reader will note that it has not been necessary or feasible to utilize the above
formula for every case.
Campa: William Denevan (1971) gives these values for Q as the result of his own survey
and measurements of Campa diets. Although during a three-day sample period during the
dry season, Denevan observed that an adult male consumed about 9 1 g of animal protein per
day, he states,
Observations of many Campa meals indicate. . . on the typical day a family will have a
few birds and small animals that provide each adult with about 90 grams of meat
containing between 15 and 20 grams of protein daily [1971:514].
= the edible weight in grams of all fish and game available

Bayano Cuna: The value is given by Bennett (1962) who measured food intake in 15
households over a 14-day period. Fish and meat provide approximately equal amounts of
animal protein in the average diet.
Miskito: Nietschmann (1972) kept daily records of game, meat, and fish brought into a
Miskito village including marine and riverine fish and turtles. Actual diets were sampled in
three households in the same period, and the results extrapolated to the entire village. The
value given in Table I1 was calculated from Nietschmanns (1972) Table IV by dividing the
total estimated consumption of animal protein for the village by the population. Purchased
food is included.
Kaingang: Henry (1964) describes a 5-week hunt by a party of 18 which left the base
camp. Henry states that meat from a hunt would last up to 3 days, so I added 3 days to the
hunting period for a total of 38 (T). During this period, 8 tapirs estimated to weigh 200 lb
(90.90 kg) each were killed. Ten monkeys and about 20 birds were also bagged but weights
are not given. From Nietschmann (1972) and Eisenberg and Thorington (1973) I took
average live weights of monkeys a t about 5 kg each and birds at 1 kg each, summed these
values and deducted appropriate amounts for butchering. This yielded 389 kg of meat for
the period of the hunt, which may be substituted in the simplified formula as F. Henry
states that each adult consumes 2 Ib (909 g) of meat per adult each day (1964:159), but he
clearly has not deducted enough for butchering and also ignores the fact that a considerable
portion of [the meat] was taken away by Indians returning to the post (1964:159). My
solution was to assume that the entire Kaingfing population of 106, both those at the post
and those on the hunt, shared the game and that no other game was coming in a t the post.
Thus

Q =

o2
389i000 = 1 9 g proteinlcapitalday.
38 x 106

Henry also states that the hunt took place during the optimal hunting season and that they
were assisted by his own well trained, well fed tapir-hunting dogs (1964:158).

Gross]

PROTEIN CAPTURE AND CULTURAL DEVELOPMENT

541

Siriono: The ethnographer, A. Holmberg, gives estimates of meat consumption per person
over a three month period during 1941. The amounts were estimated by Holmberg himself
and a Bolivian employee while they were situated at Tibaera, a site selected by Holmberg
and his assistants (1969:xx). In the third month of observations, Holmberg brought an
additional group of Sirion6 to the site enlarging the settlement from 60 to 154. Holmberg
warns the reader that these are rough estimates, not precise measures (1969:75). The
values reported (converted to g ) are as follows (1969:74):
August
254
g/individual/day
September
240
g/individual/day
October
163
glindividuallday
These values yield a mean of 219 g meatlcapitalday or 44 g protein. Holmberg attributes the
October drop in meat availability to garden clearing activities, but the enlargement of the
settlement may also have been pertinent. Holmberg states that meat consumption declines
from January to April, because of the difficulty of travel during the rainy season
(1969:75).
Wayana: La Pointe (1970) made exceptionally complete observations on hunting and
fishing yields in a single small settlement of fluctuating size. The data are reported as pounds
of meat or fish taken per month over the period of one year. Unfortunately, he does not
state the size of the settlement depending on this catch, but we may estimate it as
fluctuating around a mean of 16(P) by calculating from La Pointes tables that an average of
4 hunters per month were present (1970:52-53), and assuming a ratio of 1:4 for
hunters:nonhunters. This is close to the average Wayanfi settlement size of 1 2 reported by
the ethnographer. During the year 802.727 kg of fish and 1038.636 kg of game were
produced. Summing and converting to butchered weight yields 1104.82 kg (Ft). Substituting
in the formula
o.2
=

lo4820 = 31 g proteinlcapitalday.
365x 16

Sharanahuo: J. Siskind (1973a) recorded all the game brought in to three households (in a
village of eight households) in which ten hunters (men over 16) lived for a period of 28 days
during April 1966. According to the ethnographer, this month was, toward the end of the
rainy season, a fair period for hunting (1973a:205). The ten hunters represent just half the
men in this age category for the entire village with a population of 90. Since during this
period, game was exchanged in both directions between these three and the other five
households I shall assume that it represents about half the game brought in to the entire
village during the period. Accordingly, I shall calculate protein intake for the entire village of
90. Siskind lists only the species or type of game brought in so it was necessary to estimate
their weights using Nietschmanns (1972) tables and a few other sources. The dressed weight
of the game reported by Siskind was thereby estimated at 339,525 g, giving an estimated
total of 679,050 g for the village. Siskind did not estimate the fish catch but she states that
the diet consists of 30% game and 5% fish (1973a:84). Assuming that this ratio applies to
weights, I added one-sixth to the value for game, giving a total of 792,225 g Ft. Thus
o2 792 225 = 63 g proteinlcapitalday.
28
90
Shipibo: Roland Bergman (1974) lived over one year in a Shipibo village of 107
population, intermittently sampling hunting and fishing yields. He reports estimated values
directly in grams protein for a one year period as follows (1974:133): Fish--1,511,619 g;
Game-366,411 g. He also reports a value of protein from cultivated food as 679,665 g,
which I record here just for its intrinsic interest but omit from the following calculation:

878 030 = 48 g protein/capita/day.

107
Waiwai: Ms. Barbara Meltzer combed through four ethnographic reports on this small
group (Evans and Meggers 1955; Fock 1963; Guppy 1958; and Yde 1965), finding partial
observations on six different hunting or fishing expeditions. Multiplying the number of
hunters on each hunt by the length of each hunt in days gives a value of 39 man days for a
yield of 147.7 kg of fish and game. Thus Mt may be calculated at 2272 g. From the
= 3k5

AMERICAN ANTHROPOLOGIST

542

[77,1975

ethnographies, Meltzer estimates that Waiwai men hunt every third day on the average. Thus
each Waiwai hunter goes hunting or fishing about 120 days per year. In a Waiwai village of
77, with 28 hunters, Dt takes a value of 3360. Substituting in the formula,

3360
o.2 22721 = 54 g protein/capita/day.
3 6 5 x :7

NOTES
This study had its beginnings in a brief, preliminary field study of the Western Gavioes
of Par6 State, Brazil which was supported by a grant from the Research Foundation of the
City University of New York. An earlier version of this paper was read a t the 38th Annual
Meeting of the Society for American Archaeology in May 1973 in San Francisco. It was part
of a symposium entitled Ecological Anthropology which received support from the
Wenner-Gren Foundation. Early versions were also read at Wayne State University and the
State University of New York, Stony Brook. I thank the students in my course in cultural
ecology in the Ph.D. Program in Anthropology of the City University of New York,
especially Honora Ban, Barbara Meltzer, Fred Roberts, and Ann Sheedy.
The following persons read and commented on a draft of this paper: Joan Abelove,
Daniel Bates, Lucile Brockway, Robert Carneiro, Francis Conant, William Denevan, Carol
Ember, Melvin Ember, Nancy Flowers, Sildde Gross, Marvin Harris, Daniel Janzen, Allen
Johnson, Susan Lees, Betty Meggers, Bernard Nietschmann, John Pfeiffer, Eric Ross, Jane
Ross, and John Speth. I am most grateful for their invaluable criticisms and encouragement.
I am solely responsible for any errors of fact or interpretation.
In this paper, the terms Amazon Basin or Amazonia are used interchangeably and are
intended to include the drainages of the Amazon and Orinoco Rivers and thereby most of
Central and North Brazil, Eastern Bolivia and Peru, Southeastern Colombia, and interior
Venezuela and the Guianas, but excluding coastal areas and the Andes range.
River turtle eggs (Podocnemis expansa) are available in vast quantities along certain
stretches of river bank and they are avidly consumed by some groups during the brief season
of availability (Carneiro, personal communication). Another seasonally available source of
concentrated protein is in the seeds and nuts of wild plants, such as the Brazil nut
(Bertholletia excelsa).
K. Ruddle (1973) conducted a study of subsistence among the Cariban Yukpa of
Colombia and Venezuela and concluded,
Although invertebrates supply a small percentage of the products of animal origin
consumed by the Yukpa, they nevertheless comprise an important element in the food
supply of the tribe. During limited seasons of particular abundance, large quantities of
insects can readily be collected, a t which time they serve to supplement the products of
hunting, which are obtained only after the expenditure of much greater effort
[ 1973: 991.
G. Morren (1973) recently presented data from tropical New Guinea showing that food
from rodents, frogs, and lizards, hunted by women and children and perhaps even dragged
into the village by pet dogs yielded small but significant increments to the animal protein
capture of a group of swiddeners of the Sepik headwaters. He points to a serious matter of
relevance to this study:
The problem presented is one of measurement: both the anthropological and the folk
version of the stereotype result in low counts or no counts when it comes to assessing
the objective roles of women [Morren 1973:14].
3
A variety of values have been proposed for minimum protein need (Hegsted
1964:154-155), and recommended daily allowances usually have built-in safety factors of up
to 50% to account for possible variation within a population. Needs vary within a population
depending on age, sex, body size, health status. Perhaps the lowest values proposed by
responsible scientists have been F A 0 recommended minimum needs of 0.30-0.35 g
protein/kg body weight/day (Taylor and Pye 1966) for adults when the protein ingested is
of high Biological Value. The F A 0 recommends substantially higher values of 0.65-0.80 g/kg
BW/day when proteins are of low Biological Value (Taylor and Pye 1966:130). The latter
condition certainly prevails when more than 70% of calories are provided by starchy foods
like manioc, potatoes, o r plantains (see Table 11), which are extremely low in certain

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PROTEIN CAPTURE A N D CULTURAL DEVELOPMENT

543

essential amino acids (Table I). Thus, I suggest that the average minimum per capita protein
need for Amazonians subsisting on starchy cultigen wild game and fish should be set a t
40-50 g/day, thereby allowing for the particularly high needs of active adults and rapidly
growing adolescents. This requirement could be met by adults with a daily diet such as the
following.
Amount
Calories
Protein
( kcal)
(grams)
Food
(grams)
150
225
30
Meat or Fish
Manioc Flour
700
2240
12
Sweet Potato
400
464
5
4
Maize
100
129
Totals
1350
3058
51
Note the critical importance of meat which supplies 59% of the protein but only 7% of the
calories and 11% of the bulk.
Given the patchiness phenomenon and the rather good possibility of local extinction
from over efficient predation, one might undertake a fresh look a t the many taboos on the
killing and eating of certain animal species among Amazonian tribes. These taboos, which
sometimes are general, sometimes specific to particular statuses o r life stages, may have the
effect of averting the hunting out of certain animals.
Two species of peccary (Tayassu spp.) are found throughout the Amazon Basin, the
collared peccary ( T . tajacu) and the larger white-lipped peccary ( T . pecari). Gilmore asserts
that
either species would seem to be potentially domesticable because the young tame easily,
but unless castrated, the males soon become vicious and unmanageable [ 1948:3821.
According to Crandalls authoritative work (1964),the collared peccary breeds frequently in
captivity (i.e., in zoos), the white-lipped species less often. But, he cautions,
while there is still some question concerning the reputed ferocity of peccaries in the wild,
there can be no doubt that captive animals, especially males, may sometimes be
aggressive . . . [ p. 5291.
Thus it appears that the full domestication of these species would present serious, perhaps
insurmountable problems.
6 1 have seen no evidence for the assumption made by some writers, that maize reached
the Amazon later than manioc and that it had not yet been adopted as a major staple because
of its tardy arrival.
A passage from M. Harners description of Jivaro subsistence (1972)illustrates this
mechanism:
virtually all of the Jivaro territory has been hunted efficiently for a long period of time
with the result that game is not as abundant as in regions unoccupied by the Indians. This
fact was particularly driven home to my Jivaro companions and me in 1969,when, in
travelling through an unhunted no-mans-land between the Jivaro and the Achuara
along the lower Rio Cangaimi, which had not been exploited because of the enmity
between the two groups, we encountered unprecedented quantities of monkeys and birds
[1972:56].
See also H. Hickersons ethnohistorical study of The Chippewa and Their Neighbors where
he documents the existence of a similar no-mans land between the Dakota and Chippewa
(1970:91-119),
attributing to it the same function as I suggest here. .
Roberto Goodland (1971)and others have suggested that the term savanna, in the
sense used in Africa, is not generally applicable in South America. For central Brazil he
prefers the Portuguese term cerrado, usually translated as scrub forest. South American
cerrados are certainly not comparable to the African savannas. The enormous diversity of
large, specialized savanna grazing ungulates with their extraordinarily high densities did not
occur in South America (cf. Bourliere and Hadley 1970).
The Yanomamo of Southern Venezuela exemplify this pattern. This is a large group,
expanding rapidly (Chagnon 1974) from a relatively recent origin in an unforested highland
habitat (Lizot 1971 :141), perhaps supplanting riverine groups decimated earlier by

544

AMERICAN ANTHROPOLOGIST

[ 77,1975

introduced diseases. The introduction of plantains and steel tools may have set the
expansion off, creating an imbalance between the ability to produce calories and protein
availability. This expansion may have intensified a pre-existing raiding pattern. Differential
infanticide favoring males provided a mechanism which motivated and sustained raiding as a
prominent feature of Yanomamo life (Chagnon 1968a, 1968b). Continuous and ruthless
conflict within the boundaries of the society preadapted them to fierceness against other
groups on their border, facilitating expansion. Chagnon (1973) suggested that warfare-not
soil exhaustion-was the proximate cause of distant relocations (or macro moves) of
villages, but these moves could still have the function of preventing the depletion of game
stemming from the overconcentration of this expanding society.
Chagnon has rejected this interpretation, and referred back to his circular chain of
processes in which male fierceness generates a preference for boy babies, female infanticide,
thus a shortage of women, raiding for women, more warfare, and consequently a status
structure offering high rewards to the fiercest. Arguments over food, states Chagnon,
sometimes precipitate mortal confrontations:
I could easily fight with Moawa over a banana, and one of us might get gravely injured
or killed. But the banana would have been only a very insignificant aspect of the fight.
The real reasons for the fight,. . . would have to d o with the status system. It would
be . . . a gross misrepresentation to say that the cause of fight X was bananas . . . .
Recent trends in ethnological theory are tending more and more to crystallize around the
notion that warfare among swidden cultivators must always be explainable in terms of
population density, scarcity of strategic resources such as territory or proteins. . . . The
Yanomano are an important society for their warfare cannot be explained in this way
[ 1974: 194-1951.
Chagnon seems to impute to some the notion that the Yanomamo are fighting over
territory or proteins. No one, to my knowledge, has suggested this. The proposal made by
myself and also by M. Harris (1974:61-80) does not invalidate Chagnons political and
motivational analysis, but it does provide a plausible causal explanation. Improved
opportunities to secure resources may be an unintended or latent consequence of
Yanomamo warfare, and it does not depend on the actors perception of this as a goal. To
discredit this suggestion would require a demonstration that the settlement and migration
patterns encouraged by Yanomamo have a negligible or negative effect on protein capture,
and this Chagnon has not yet provided.
The conclusions are presented as if it had been adequately demonstrated that proteins
of animal origin are a limiting factor on native population densities and sizes in South
America. In fact, it has not. One of the purposes of writing and publishing this paper has
been to stimulate fieldworkers to attempt to measure such variables as dietary intake in the
societies in which they conduct research. A number of people who have commented on this
papers findings have suggested that members of one or another society obviously d o not
suffer from protein scarcity since they have been observed gorging themselves on meat. One
may easily gain this impression from Holmbergs eyewitness account of the Sirion6
(1969:74-75). But without careful measurement, even first-hand observation can be
misleading as may be seen from the per capita animal protein intake value calculated from
Holmbergs own data. The dialectic between theory and data has now brought us to the
point where only more and better measurements will yield satisfactory results.
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