Ecological Modelling 110 (1998) 5 17

Analysis of hunting options by the use of general food taboos

Johan Colding *
Department of Systems Ecology, Stockholm Uni6ersity, S-106 91 Stockholm, Sweden

Abstract
A hypothetical model was built, using the STELLA II software program, to test several hunting options for a
human hunting group. Different outcomes of possible hunting modes are analysed, such as a change in hunting rate,
prey hunted, or species avoided or not avoided by taboos. The model consists of five sectors that reflect a short food
chain in an upper Amazonian ecosystem. There is a vegetation sector, a predator sector, and two sectors consisting
of browsers and grazers. The last sector represents a human group, known as the Ecuador Achuar. The critical factor
analysed is how differences in hunting rate affect a target resource, and how this resource may be affected by general
food taboos. The major results of the model are that general food taboos may not be an adaptive short term strategy
for hunters, but that a moderate hunting mode may be the most effective option for the human group. Since the
model is a simplification of the real world, no general conclusions for management should be drawn from the results.
1998 Elsevier Science B.V. All rights reserved.
Keywords: General food taboos; Hunting options; Amazone ecosystems

1. Introduction
Many traditional societies employ food taboos
on animal species for a number of reasons. Some
researchers suggest that there are nature management motives behind them (Rappaport, 1967,
1968; Reichel-Dolmatoff, 1976; McDonald, 1977;
Johaness, 1978; Ross, 1978; Harris, 1979), while
others resent any such ecological motives (Rea,

* Present address: The Beijer International Institute of Ecological Economics, The Royal Swedish Academy of Sciences,
PO Box 50005, S-104 05 Stockholm, Sweden. Tel.: +46 8
6739500; fax: + 46 8 152464; e-mail: johanc@beijer.kva. se

1981; Edgerton, 1992). Hunting among some traditional groups may be conducted in a highly
efficient way, consistent with predicators of foraging theory (Alvard, 1993, 1994). While species are
thus pursued with short-term harvest rate maximization, one may very well ask why some native
hunting groups employ food taboos. McDonald
(1977), in a study of 11 South American tropical
groups, suggests that taboos may function as conservation mechanisms for reducing the hunting
pressure on larger mammals in environments
where such species are low in abundance. General
food taboosapplying to all members within a
communitymay play a role in biodiversity con-

0304-3800/98/$19.00 1998 Elsevier Science B.V. All rights reserved.

PII S 0 3 0 4 - 3 8 0 0 ( 9 8 ) 0 0 0 3 8 - 6

J. Colding / Ecological Modelling 110 (1998) 517

Fig. 1. Approximate location of the Ecuador Achuar territory (marked in grey).

servation. For example, Colding and Folke

(1996), found that a number of threatened populations of species, including endemic and keystone
species, benefit from such taboos.
The purpose of this modelling exercise with
STELLA II, has been to analyse possible relations
between general food taboos and different hunting modes. The STELLA II program may be a
suitable tool for an investigation of this kind.
While real world data on wild populations are
available in the literature (see Emmons, 1990;
Nowak, 1991; Redford and Eisenberg, 1992), it is
nevertheless problematic to generalise from such
data. For example, data on numbers of density
are measured in habitats where species thrive. To
generalise from such data, from one habitat to
another, may be highly awkward. Among other
things, this is due to the differences in the heterogenity of landscapes within and between ecosystems. Perhaps more realistic data for the
presented model could be obtained using site-spe-

cific data from field studies. Unfortunately, this

has not been possible. Thus, the estimates used
here are highly artificial.
The Ecuador Achuar represents the human
group of this model. This group, inhabiting a vast
region at the Ecuador/Peru border (see Fig. 1),
has been well studied by DeScola (1986). The
Ecuador Achuar impose general food taboos on
several species of mammals. They abstain from
the consumption of all carnivorous mammals and
regard several other such species as inedible, referring to them as sickening meat (DeScola, 1986).
Among such species is the rodent Capybara, Hydrochoerus hydrochoerus, which is widely hunted
by other neotropical forest groups (see Redford,
1993; Bodmer 1994). However, other species of
rodents are hunted, e.g. agoutis (Agouti paca) and
acouchis (Dasyprocta and Myoprocta). Ungulates,
such as Brocket deer (Mazama Americana) and
Tapirs (Tapirus terrestris and T. pinchaque) are
avoided by general food taboos, due to reincarnation beliefs (DeScola, 1986).

J. Colding / Ecological Modelling 110 (1998) 517

narios are presented in Section 3. Individual

model runs will also be discussed in Section 3,
while an overall discussion will follow in Section
4.

2. Basic structure and data

Fig. 2. The basic structure and key actors of the model.

The results obtained in this study are highly

artificial and should not be taken as indicators
of the Ecuador Achuar hunting mode. The
model is far too incomplete for any such inferences to be made. This complies for both the
structure of the ecosystem, as well as for the
various populations of species described in the
model. Some possible hunting modes and sce-

The main sectors and key actors of the model

are presented in Fig. 2. This figure illustrates the
basic food chain investigated. The population of
jaguars is regarded as the top-predator in this
extremely simplified ecosystem. Jaguars prey on
both the rodent and tapir populations respectively, that, in turn, feed from the vegetation
sector. Although all carnivore species are
avoided by the Achuar hunter, the jaguars in
this model are thought of representing a fraction
of all the carnivores in this ecosystem. The same
holds true for both the rodent sector and the

Fig. 3. A model for examining different hunting options for a human group.

J. Colding / Ecological Modelling 110 (1998) 517

Table 1
Sensitivity runs on population stocks of the model, using STELLA II
Months

1: Vegetation

2: Vegetation

3: Vegetation

1: Rodent

2: Rodent

3: Rodent

0
50
100
150
200

3 000 000
6 617 864
6 605 182
6 557 093
6 620 185

6 000 000
6 618 420
6 605 278
6 557 171
6 620 226

9 000 000
6 618 413
6 605 277
6 557 170
6 620 225

9000
11 955
11 772
14 512
12 272

18 000
12 009
11 785
14 521
12 279

27 000
12 035
11 791
14 525
12 281

0
50
100
150
200

1: Tapir
300
300
294
335
338

2: Tapir
600
541
511
530
503

3: Tapir
900
651
566
561
521

1: Jaguar
10
28
28
31
27

2: Jaguar
25
28
28
31
27

3: Jaguar
40
28
28
31
27

0
50
100
150
200

1: Tapir*
300
218
67
0
0

2 Tapir*
600
485
410
400
333

3: Tapir*
900
603
487
463
392

An increase in the rate of tapirs per jag from RANDOM (0, 0.4, 0.4) to RANDOM (0, 0.5, 0.5) is marked as * in the table.

tapir sector. The former represents mainly a

lumping of larger rodents that the Ecuador
Achuar pursue, and the latter a fraction of the
ungulates avoided by general food taboos.

2.1. Climatic conditions of the ecosystem

The ecosystem is a simplified version of the
Ecuador Achuar territory, situated at the
Ecuador/Peru border. It lies north of the Pastaza
river, and covers an area of about 9000 km2 (see
Fig. 1). The region has a typical equatorial climate, constantly humid, no dry season, and a
monthly rainfall always over 60 mm. It is situated 2 south of the equator, where days and
nights are nearly the same length, and temperatures remain constant throughout the year. Despite its proximity to the Andean barrier, this
ecosystem is not directly affected by the special
meteorological conditions of the foothills. Mean
annual daytime temperatures vary between 24
and 25C. The annual average low temperature
fluctuates between 19 and 20C, depending on
altitude, and the average annual high tempera-

ture is between 29 and 31C. It is marginally

warmer from October to February.
The average annual rainfall is no more than
3000 mm for the highest latitudes, and not less
than 2000 mm for the lowest. However, rainfall
varies from year to year. Periods of too little or
too much precipitation have no noteworthy effect on the vegetative activity of wild and cultivated plants, since the duration is too short to
have any long-term influence. However, animal
populations may be affected by this variation,
since droughts rapidly dries up secondary
branches of rivers, normally filled with water,
affecting the fish that live there, and other species of animals that normally receive water at
these places. In the opposite case, heavy continuous rainfall tends to accelerate the decomposition
of organic bedding, rapidly destroying the fruit
and seeds eaten by large terrestrial herbivores
such as tapirs and peccaries.
The ecosystem is in a state of dynamic equilibrium, as its system of energetic exchange operates theoretically in a closed circuit (Odum,
1971). The organic matter and minerals are con-

J. Colding / Ecological Modelling 110 (1998) 517

Fig. 4. A case with a zero hunt on all the three populations of species. The ecosystem is in balance.

A detailed structure of the model, describing

the various in- and outflows between and within
stocks and sectors is presented in Fig. 3. The
model has been run at a monthly time step for
200 months. This length was chosen on the basis
that it would allow a human population to adjust
to changes in hunting modes, since possible effects
of a taboo would thus be known in a fairly short
notice of time.

Section 2.1 the vegetation sector is considered to

be largely unaffected by climatic variability. Nutrients are thus tightly recycled and closely associated to the standing living biomass. This has been
accounted for in the model by the design of a
nutrient cycle circuit (see Fig. 3). The vegetation
sector was constructed to be robust, and largely
unaffected by different levels of population
changes within other sectors. Sensitivity runs indicate that population levels of the herbivores (rodents and tapirs) are constrained by their
respective carrying capacities, and do not pose a
threat to the vegetation, even if initial numbers
greatly overshoot carrying capacity levels (see
Table 1). This sector is thought to consist largely
of evergreen leaves, flowering plants, fruits and
seeds, and constitutes the first trophic level of this
ecosystem (for information on input data used
throughout Section 2.2, see equations in the Appendix).

2.2.1. The 6egetati6e sector

Based on the climatic conditions described in

2.2.2. The rodent sector

The rodent sector of the model represents ro-

tinually recycled by a complex network of microorganisms and specialised bacteria. Given that
almost 90% of the nutrients of tropical forest
ecosystems are stored in the standing plant
biomass (Begon et al., 1990), this closed circuit
system is sensitive to larger clearings and deforestation activities.

2.2. Description of model sectors

10

J. Colding / Ecological Modelling 110 (1998) 517

Fig. 5. A case where rodents are hunted at the hunting rate of 0.17, equal to an average of pursued rodents by other neotropical
forest groups (Redford 1993). Jaguars and tapirs are not hunted at all.

dents that may be hunted by the Ecuador

Achuar (mainly those of a weight between 2
and 10 kg). Species under consideration include
Agouti paca, and Dasyprocta spp. and Myoprocta spp. They represent part of the second
trophic level. Rodents are widely hunted among
tropical Amazonian groups (Redford, 1993) and
are important sources of protein in the Ecuador
Achuar diet.
Initial population of rodents were set at
18 000. This estimate is based on adding the
density numbers for the above rodent species
(see estimates of Nowak (1991), Redford (1993),
and Bodmer (1994)) (Density of Agouti is 5.1
per km2, for Dasyprocta and Myoprocta 6.1 per
km2.) For an ecosystem 9000 km2 in size, the
estimate will end up being about 110 000 rodents, which is about 12.2 individuals per km2.
This is considered as an extremely high estimate.
Densities for rodents such as hares in Sweden,

are considered very high at 4 per km2 (ibid),

which is about 1/6 of the number above. For
this reason the estimated number on density has
been set at about 2 rodents per km2, which
gives a total value of about 18 000 rodents.
Mean litter size of rodents was set at four
(Eisenberg, 1981), with 34 litters per year
(ibid). Based on these numbers the specific birth
rate was set at about 6.0, which gives a monthly
birth rate of 0.5. The death rate was estimated
by using the survivorship curve based on that of
Odum (1983), and set at a monthly rate of 0.46.

2.2.3. The tapir sector

Tapirs are both grazers and browsers. They
represent part of the second trophic level of this
model. The Ecuador Achuar employ a general food
taboo on the killing of tapirs. Densities for tapirs
have been estimated at 0.4 per km2 (Bodmer, 1994).
As was the case for rodents, 1/6 of this estimate has

J. Colding / Ecological Modelling 110 (1998) 517

11

Fig. 6. A case displaying the maximum hunting rode on rodents. At a hunting rate of 0.92, both rodents and jaguars are severely
affected. Above this rate these populations go extinct. Jaguars and tapirs are not hunted at all.

been used as an input value for this model. Therefore, the initial population of tapirs is set at about
600. For the calculation of birth and death rates,
the survivorship curve of the species of Black-tail
deer was used (see Odum, 1983). This species is a
herbivore of approximately the same size, and was
assumed to have a survivorship curve of the same
kind as tapirs. Litter size of tapirs was set at one,
with an average of about one litter per year
(Eisenberg, 1981). Monthly birth rate was
calculated at 0.04, and monthly death rate at
0.017.

2.2.5. The Achuar sector

The Achuar of this particular region of
Ecuador has a steady population of about 1000
(DeScola, 1986). The group has many options for
hunting which will be analysed below.

2.2.4. The jaguar sector

Jaguars represent the top-predator of this
ecosystem, preying on both rodents and tapirs.
No information on rates of predation, birth and
death were found in the literature. Ecological
density of jaguars is estimated at 0.013 per km
(Redford and Eisenberg, 1992). One sixth of this
value sets the initial jaguar population at about 20
individuals.

3.1. Scenario one

3. Scenarios and individual outcomes of runs

The results of the different runs are presented in
Figs. 49. Following are three different scenarios,
each representing possible hunting options for the
Ecuador Achuar population.

As has been stated, the Ecuador Achuar obey

general food taboos on the hunting of tapirs and
jaguars. In this first scenario, this has been accounted for by setting hunting rates on these
species at zero (see Figs. 46). In this scenario,
consisting of three runs, the Achuar only hunt
rodents. Different outcomes will be obtained, de-

12

J. Colding / Ecological Modelling 110 (1998) 517

Fig. 7. A case where jaguars are pursued at a hunting rate of 0.0036, which turned out to be the maximum rate allowed for not
creating a collapse of the jaguar population. Rodents are pursued at a hunting rate of 0.17. Tapirs are not pursued at all.

pending on the different hunting rates of rodents.

Fig. 4 represents a case where the Achuar employ a zero value in the hunting rate of rodents.
This run indicates that the model ecosystem is in
balance.
In the second run (see Fig. 5), the hunting rate
on rodents employed by the Achuar was set at
170 per month, which is about the normal hunting rate on rodents by other neotropical forest
groups (Redford and Robinson, 1987).
In the third run (see Fig. 6), the numbers of
rodents hunted was set at 920 per month, which
was the highest number of rodents that could be
hunted without collapsing this population. If the
rodent population collapses it will make the
jaguar population collapse as well, since jaguars
predominantly prey on rodents in this model.
These results indicate that the Ecuador Achuar
sector of the model may hunt rodents sustainably
at a rate of \0.17 to 0.92.

3.2. Scenario two

If rodents are hunted at a rate of 170 per
month, and the taboo on tapirs is still employed
but the taboo on jaguars is not obeyed, the run
presented in Fig. 7 is obtained. In this run the
hunting rate on jaguars is set at 3.6 per month,
which is the maximum hunting rate allowed
without collapsing the population.

3.3. Scenario three

In this last scenario two different runs were
made. In Fig. 8 the taboo on the hunting of
jaguars is observed by the hunters, but not the
taboo on tapirs. The result from this run indicates that if tapirs are hunted at a rate of two
per month, or greater, the number of the tapir
population will gradually decline and eventually
go extinct. Additional runs were made in which

J. Colding / Ecological Modelling 110 (1998) 517

13

Fig. 8. A case displaying a zero hunting rate of jaguars (by taboo). With a hunting rate on tapirs of 0.002, the population declines
steadily over time. Rodents are hunted at the rate of 0.17.

it was possible for hunters to hunt all three populations of species simultaneously at rather high
hunting rates in a sustainable manner. For example rodents, jaguars and tapirs, may all be pursued at corresponding hunting rates of 170, 3 and
2 per month, without threatening the long time
survival of these populations.
Fig. 9 shows that over-hunting of jaguars
may be a rational hunting option for groups
that hunt species, such as rodents and ungulates. The interspecific competition between the
human hunters and jaguars is thus reduced. This
situation does not pertain to the Ecuador Achuar.

4. Discussion
There are several limitations to this exercise.
For example, input data on consumption rates
are simply estimated without any real references. The system was sensitivity tested and
analysed in this regard, which demonstrated

that if the numbers of tapirs taken per jaguar

are slightly increased from a random value of
0.4 to 0.5, it could have a dramatic impact on
a tapir population of about 300 individuals (see
Table 1). At this level the whole tapir population collapsed and became extinct.
Estimates of vegetation, below and above
values in the model, revealed that carrying capacity is set at about 6.6 million vegetative
units. Sensitivity runs on the rodent population
at numbers below or above the estimated population size, showed that the rodent population
established itself close to a carrying capacity at
about 12 00014 500 in most runs (see Table 1).
Despite the limitations of this exercise, what
can be said about the overall results of this
modelling exercise? Firstly, the results indicate
that general taboos imposed on species of
jaguars and tapirs protect them from becoming
extinct. However, the taboos of this model
seem not to have any hidden ecological functions by, e.g. enhancing other populations of

14

J. Colding / Ecological Modelling 110 (1998) 517

Fig. 9. A case showing that it may be advantageous for hunters to pursue a predator species such as the jaguar. This may reduce
the interspecific competition between two populations feeding on the same resource. Hunting rate on jaguars is set at 0.0036, on
rodents at 0.17, and on tapirs at 0.002.

species pursued by the Ecuador Achuar. It seems

to be more efficient to pursue jaguars whenever
encountered, than not to. As Fig. 8 also indicates
(and several other runs have demonstrated), it may
be most efficient to pursue all three species of
mammals at moderate hunting rates.

based on mythologies and perceptions of reality,

may be the reason behind these taboos in the real
world (see also DeScola, 1986). On the other hand,
for the preservation of species that may go extinct
from over hunting, the use of a moderate hunting
mode, or the use of general food taboos, seem to
be viable options. Any final conclusion on these
relationships cannot be drawn from this model.

5. Conclusion
General food taboos may be ecologically adaptive for hunters in ways which are not disclosed by
the nature of this simplified model. The STELLA
model used here indicates that a moderate hunting
mode may be the most effective option for the
neotropical group of this model. However, no
inferences should be drawn based on this result for
tropical forest groups such as the Ecuador Achuar.
In the model of this paper, general food taboos do
not increase the yield for hunters of species which
are not surrounded by taboos. Cultural reasons,

Acknowledgements
I would like to thank Professor Robert Costanza.
at the Institute for Ecological Economics, University of Maryland, for giving me valuable advice and
feedback in the construction of this model. Without
his insistence this paper would never have been
completed. Warm thanks also to Professor Carl
Folke, at the Beijer Institute of Ecological Economics, Stockholm, for valuable suggestions and
thoughtful considerations.

J. Colding / Ecological Modelling 110 (1998) 517

Appendix A. Equations for the model

Human
Human group(t)= Human group(t
dt)
INIT Human group = 1000
 Jaguar kill per human = 0
 Rodent kill per human = 0
 Tapir kill per human = 0
Jaguars

Jaguar pop(t) = Jaguar pop(t

dt)
+ (births jaguar deaths jaguar)*dt
INIT Jaguar pop = 20
INFLOWS:
births jag = Jaguar
pop*(birth frac + jag of
tap+birth fract jag of
rod)
*(1(Jaguar pop/car cap jag))
OUTFLOWS:
deaths jaguar = Jaguar
pop*(death frac jag of
rod+death fract jag of

15

(40.0, 0.245),
(50.0, 0.305),
(60.0, 0.35),
(70.0, 0.37),
(80.0, 0.383),
(90.0, 385),
(100, 0.385)

*
death fract jag of tapirs=
GRAPH(Tapir pop)
(0.00, 0.11),
(60.0, 0.105),
(120, 0.09),
(180, 0.075),
(240, 0.06),
(300, 0.05),
(360, 0.04),
(420, 0.032),
(480, 0.0235),
(540, 0.016),
(600, 0.00)

*
death frac jag of rod=
GRAPH(Rodent pop)
(0.00, 0.91),
(1800, 0.855),
(3600, 0.76),
(5400, 0.625),
(7200, 0.45),
(9000, 0.295),
(10 800, 0.18),
(12 600, 0.105),
(14 400, 0.06),
(16 200, 0.025),
(18 000, 0.0005)
Rodents

Rodent pop(t)=Rodent pop(t

dt)+(birthsdeaths)*dt
INIT Rodent pop=18 000
INFLOWS:
births=Rodent pop*birth
fraction*(1 Rodent pop/car cap
rod))
OUTFLOWS:

tapirs)+Human group*Jaguar kill

per human
 car cap jag = 40

*
birth fract jag of rod =
GRAPH(Rodent pop)
(0.00, 0.0265),
(100, 0.0285),
(200, 0.033),
(300, 0.0375),
(400, 0.0425),
(500, 0.048),
(600, 0.0515),
(700, 0.0575),
(800, 0.059),
(900, 0.06),
(1000, 0.06)

* birth frac jag of tap = GRAPH(Tapir pop)
(0.00, 0.13),
(10.0, 0.145),
(20.0, 0.174),
(30.0, 0.205),

deaths=Jaguar pop*rod per

jaguar+Rodent pop*death
fraction+Human group*Rodent kill
per human
 car cap rod=36 000

rod per jaguar=RANDOM(0.100, 50)

*
birth fraction=GRAPH(Vegetation)
(0.00, 0.12),
(700 000, 0.165),
(1.4e+006, 0.23),
(2.1e+
006, 0.285),
(2.8e+006, 0.33),
(3.5e+006, 0.395),
(4.2e+

J. Colding / Ecological Modelling 110 (1998) 517

16

006, 0.43), (4.9e + 006, 0.455), (5.6e +

006, 0.485), (6.3e + 006, 0.5), (7e +
006, 0.5)

* death fraction = GRAPH(Vegetation)
(0.00, 0.815),
(80 000, 0.63),
(160 000, 0.49),
(240 000, 0.395),
(320 000, 0.32),
(400 000, 0.27),
(480 000, 0.24),
(560 000, 0.215),
(640 000, 0.213),
(720 000, 0.215),
(800 000, 0.213)
Tapirs
Tapir pop(t) = Tapir pop(t dt) +
(births tapir deaths tapir)*dt
INIT Tapir pop = 600
INFLOWS:
births tapir = Tapir pop*birth
fract tapir*(1
(Tapir pop/car cap
tapirs))
OUTFLOWS:
deaths tapir = Jaguar
pop*tapirs per jag + Tapir
pop*death fract tapir + Human
group*Tapir kill per human
 car cap tapirs = 1200

tapirs per jag = RANDOM(0, 0.4, 0.4)

* birth fract tapir = GRAPH(Vegetation)
(0.00, 0.0005),
(80 000, 0.0185),
(160 000, 0.0275), (240 000, 0.033),
(320 000, 0.0358), (400 000, 0.0388),
(480 000, 0.0403), (560 000, 0.0425),
(640 000, 0.0433), (720 000, 0.0438),
(800 000, 0.044)

* death fract tapir = GRAPH(Vegetation)
(0.00, 0.9), (600 000, 0.8), (1.2e +
006, 0.7), (1.8e + 006, 0.4), (2.4e +
006, 0.0498),
(3e + 006, 0.0305),
(3.6e+006, 0.019),
(4.2e +

006, 0.016),
(4.8e+006, 0.0155),
(5.4e+006, 0.015), (6e+006, 0.0145)
Vegetation
Vegetation(t) =Vegetation(t dt)+
(regeneration consumption)*dt
INIT Vegetation=6 000 000
INFLOWS:
regeneration
=seeding+(Vegetation*regen per
plant)
*(1(Vegetation/car cap veg))
OUTFLOWS:
consumption =Rodent pop*veget
per rodent+Tapir pop*veget per
tapir
 car cap veg=7 000 000
 seeding=1
 veget per rodent=20
 veget per tapir=20

* nutrients=GRAPH(Vegetation)
(0.00, 0.015),
(600 000, 0.07),
(1.2e+006, 0.135),
(1.8e+
006, 0.23), (2.4e+006, 0.365), (3e+
006, 0.565),
(3.6e+006, 0.745),
(4.2e+006, 0.815),
(4.8e+
006, 0.855), (5.4e+006, 0.88), (6e+
006, 0.9)

*
regen per piant=GRAPH(nutrients)
(0.00, 0.085),
(0.1, 0.125),
(0.3, 0.145),
(0.3, 0.185),
(0.4, 0.215),
(0.5, 0.275),
(0.6, 0.385),
(0.7, 0.505),
(0.8, 0.645),
(0.9, 0.705),
(1, 0.735)

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