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THEODOREC. FOIN
WILLIAMG. DAVIS
University
of California,Davis
THE
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[89, 1987
zation of population fluctuations; the second is one form of qualitative stability; the third
is quantitative stability; and the last is another form of qualitative stability termed "persistence," or "resilience." The opposition of constancy and resilience is but one example
of mutually exclusive definitions of stability (May 1975).
A second problem is that field investigation of stability is very difficult, since it is often
impossible to know, a priori, what the relevant measures are, irrespective of the length of
investigation needed to develop precise estimates of stability properties (for a recent and
fuller discussion of both of these issues, see Connell and Sousa 1983).
The magnitude of the field measurement problem has meant that many of the pioneering studies in stability analysis in ecology have placed heavy dependence on models and
model ecosystems as the only tool available for stability analysis (May 1974; Pimm 1981).
This situation does not seem likely to change much in the near future because of the limitations of field ecology.
The study of stability in human ecosystems suffers from exactly the same kinds of defects; if anything, they seem even more difficult to overcome. Ecological anthropologists
have been interested in the measurement of stability for some years (Moore 1957; Sahlins
and Service 1960; Piddocke 1965; Vayda 1961, 1969; Leeds and Vayda 1965; Rappaport
1968, 1979, 1984; Thomas 1972), but have not given much attention to the details of
measurement and definition. Human ecosystems are characterized by a rich, complex
feedback loop structure, even in simple models (Forrester 1972), and by limited human
population growth rates compared to other populations. Together, these two factors ensure that it will be very difficult to determine what factors regulate stability in human
ecosystems, and it will take a great deal of effort to estimate stability with any degree of
precision.
The shifting agricultural systems of Highland Papua New Guinea probably have been
studied by more investigators than those in any other place in the world (Vayda 1971;
Rappaport 1968;Clarke 1971; Strathern 1971; Buchbinder 1973; Moylan 1973; Salisbury
1975; Manner 1977; Meggit 1977; Lowman 1980; Boyd 1985). The information available
for the Maring speakers is particularly rich and suitable for an analysis of stability properties. These data permit simulations of Maring population dynamics, which enable us
to gain further insight into the meaning of the vast body of empirical data that already
exist. In turn, this leads to an examination of the stability properties of the simulation
model, and hence the system itself. In this paper we present an analysis of population
stability based on the Maring data. We begin by summarizing the key features of each of
three population dynamics models that have been proposed for these agroecosystems,
then analyze the behavior of each in an attempt to determine which of the three models
best describes the Maring data.
The Maring Agroecosystem
The Maring population consists of approximately 7,000 persons who reside in the Jimi
and Simbai River valleys in the Bismarck Range of Highlands Papua New Guinea. The
population is organized into 20 more-or-less politically autonomous, local groups which
range in size from roughly 100 to 900 persons (Rappaport 1968). The main zone of habitation lies between 1,000 and 2,000 m and is characterizedas being more heavily forested
than is usual for similar altitudes elsewhere in New Guinea (Buchbinder 1977).
Shifting cultivation is the main source of subsistence, but pig husbandry and foraging
also are economically important. New fields are cut from the forest each year. Fields usually are cropped for 14 to 26 months, then returned to fallow for 8 to 20 years. Mature
secondaryforest is favored for agriculture over primary forest. The major crops cultivated
are taro, sweet potato, yams, manioc, and various kinds of leaves and grasses (Buchbinder 1977).
Maring local populations are (or were) characterized by a complex cycle of warfare
and truce. Each local population frequently is at war with some groups and allied with
FoinandDavis]
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group moves through a developmental cycle in which population density varies over time.
Groups begin in a pioneering, low-density phase in which environmental quality and nutritional status are high. They mature at a high-density phase in which the forest environment is degraded, productivity declines, and nutritional status is poor. As nutritional
status declines, the population becomes more vulnerable to malaria, which is the factor
responsible for compensatory mortality. Buchbinder's hypothesis is based on data and is
thus plausible, although some authors (Scrimshaw, Taylor, and Gordon 1968; Murray
et al. 1978a, 1978b; Lepowsky 1984; see also the review by Beisel 1982) have argued that
severe malnutrition will halt the growth of Plasmodium
and prevent a serious clinical manifestation of malaria. These findings cast some doubt upon the effectiveness of malaria as
a mortality agent when nutritional status is poor. As a result, we examined the influence
of malarial mortality on model behavior more closely.
The ritual regulation hypothesis of Rappaport (1968, 1984) is least conventional but
is the best known of the three discussed here. His model is based explicitly on ideas about
population regulation advanced by Wynne-Edwards (1962). Wynne-Edwards argued
that many social animals, especially in their optimal conditions, practice self-regulation
by assessing numbers and consequently limiting density to average values below those
which would damage essential resources. Self-regulatedpopulations are supposed to have
one or more means for sensing excess density ("epideictic signals"), and an effective
group response for limiting further increase in density.
Rappaport proposed that the key epideictic signal for the Tsembaga Maring is the intensity of female labor. In the Maring division of labor, females are principally responsible for pig husbandry. Women tend the gardens, prepare the food, and feed the pigs.
These are labor-consuming tasks; Rappaport estimated that immediately before the ceremonial pig slaughter that he witnessed, pigs were consuming 80% of the manioc harvested and 50% of the sweet potatoes produced by the Tsembaga. Gardens were 36%
larger before the pig sacrifice than afterwards. The intensity of female labor is directly
proportional to pig density and thus is an attractively simple index of environmental
quality. Rappaport argued that as labor devoted to pig husbandry increased, complaints
about the workload would also, thus triggering a kaikoas the only response that could
relieve the workload. An incidental, but crucial, consequence of the kaikois that warfare
usually resumes shortly thereafter.
Thus, the ritual cycle is a means for reducing both pig and human numbers, and hence
pressure on the environment and resources. The time required to rebuild the pig herd to
the level necessary to support a ritual festival, on the other hand, prevents excessive warfare. The net result of the ritual cycle, therefore, is establishment of population equilibria.
In higher-quality environments pig populations grow more swiftly, kaikosare held more
frequently, warfare occurs at shorter intervals, and war mortality is higher. In lowerquality environments, exactly the converse situation holds. It follows that the ritual cycle
is a homeostat that functions to regulate the size of both human and pig populations,
population dispersal, nutritional states, and environmental quality.
RegionalStability,LocalInstability
Moylan (1973) and Lowman (1980) are the principal proponents of this model. A regional stability, local instability model is based on the notion that local groups are unstable (neither point stable nor subject to a limit cycle), but that local populations persist
in time and space at some points in the region and recolonize. Moylan developed a general, multiple-causation hypothesis of regional-local interactions, rather than a more specific proposal for a group of populations. His major contribution was to point out that
local instability is a necessary feature of regional stability. Indeed, it is unnecessary to
invoke regional stability if local stability is commonplace.
Lowman developed a more specific model, but one along the same lines as outlined by
Moylan. She termed her hypothesis "the structure of impermanence," in contrast to
Clarke (1977). She postulated that individual populations exhibit a developmental cycle
ANDNONEQUILIBRIUM
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13
much like that portrayed by Buchbinder, but differs in that each population ultimately
decreases to local extinction. A complete life cycle for a population, estimated by Lowman
to occur in approximately 200 years, begins with establishment of a small group of migrants in an unused forest environment. With time, the group expands as new migrants
join it; when the group reaches a critical minimum size, it will have established effective
self-defenseand social rules. As it continues to grow, partly through natural increase and
partly through continuing immigration, forest regeneration is impaired and the resource
base suffers. Immigration becomes emigration, fewer resources are available to reward
allies, and a military defeat becomes inevitable. Each subsequent defeat worsens the situation, as the group no longer can attract new brides or warriors, until ultimately the
group is routed from the land. The group is forced to seek refuge at lower altitudes where
malaria is endemic, and which ultimately leads to the extinction of the social unit in the
lowland environment. Lowman's regional stability model is plausible and overcomes a
number of the objections that have been set against the local equilibrium model
(MacArthur 1974; Salisbury 1975). However, as Lowman points out, the data needed to
confirm her model do not presently exist, nor is it clear that they ever will, given the long
time frame of her hypothesis. Furthermore, a logical concern about the model may be
raised here: it is not clear why a population would not, while it still possesses its greatest
numerical strength and political power, simply use its position to annex higher-quality
territories occupied by weaker neighbors, rather than resigning itself to inevitable decline.
"Model"
TheDisequilibrium
Several students of Highland New Guinea ecology have expressed skepticism about
the validity of equilibrium models (Watson 1965; Salisbury 1975; Golson 1982). The two
alternative states are nonequilibrium (which refers to the absence of any equilibrium
point) and disequilibrium (which recognizes the existence of an equilibrium state, but
argues that the system is seldom if ever in this state). We know of no authors who argue
for nonequilibrium as defined here. Of those authors cited above, Salisbury comes closest
to presenting a comprehensive qualitative model for disequilibrium for Highlands populations. He begins by explaining how cultural rules and environmental reality on which
they are based can be seriously out of phase. He argues that culture consists essentially
of sets of rules, each of which may permit a variety of behavioral outcomes, with no alteration in the rules themselves. As many aspects of culture are sensitive to resource availability, any given set of rules may be expressed in manifold ways, depending upon population density. Thus, retention of the categories of traditional culture by a population
may easily conceal the fact that the actual behavior associated with those rules has undergone profound transformationas density or resource availability varies.
Having shown that cultural stability does not necessarily imply population stability,
Salisbury outlines a model to explain disequilibrium. The essence of the model is that
exogenous inputs, typically new technologies for food production or more efficient organization, can be expected at a frequency such that resource limitation is rarely a serious
factor. Even a temporary limitation serves mainly to increase the likelihood of a technological or organizational innovation. Furthermore, occasional episodes of disease, war
mortality, or deaths from other causes occur. Consequently, the population receives no
selective pressure to stabilize. Salisbury's model, then, is a nonequilibrium and a disequilibrium model: the population never reaches a true equilibrium because of continuous
change, nor is it possible to define what the equilibrium population size is-if indeed
there is one.
Golson (1982) has provided one example of the impact of an exogenous factor on the
Maring. He showed that the sweet potato had a dramatic and lasting impact on Maring
culture, since it was so highly productive, was good for feeding pigs, and could be grown
at higher elevations than traditional crops. The sweet potato opened new environments
and modes of existence for the Maring and is but one illustration of the effects of exoge-
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nous factors on social systems. Other examples include the intervention of Australian
authority and the introduction of market agriculture and commercial forestry to New
Guinea.
Using Simulation to Evaluate Stability
The empirical difficulties associated with the field assessment of stability properties in
human ecosystems suggest that mathematical models of the Maring are appropriate tools
for the problem. Mathematical modeling of Highland New Guinea agroecosystems addresses both the time and multivariate problems, and simulating the dynamics of shifting
agroecosystems permits experimental manipulation of the simulation. By identifying the
predictions of each stability model, it is possible to compare them to the predictions of
the simulation model in order to determine which stability model best describes reality.
The Local Equilibrium Simulation Model
The Maring simulation is based upon a synthesis of the literature, particularly Buchbinder's work and many of the provisions included by Shantzis and Behrens (1973) in
their simulation. The local system is defined as the population or social group that acts
as a unit in warfare, and is most commonly seen as a village or group of villages. The
causal-loop diagram for this system, displaying the main variables and their relationships, is shown in Figure 1. The main sectors are:
1. The population sector, which contains provisions for an average net growth rate,
death rates set by war and by disease mortality, and negative feedback on birth rates.
The major interactions of the population sector are with the forest succession and the food
production/diet sectors; the latter mediates the severity of disease mortality. In this
model the population was not disaggregated by age or sex, following the practice of
Shantzis and Behrens (1973). Although we recognize that significant effects on the local
population are traceable to age and sex (e.g., labor available for specific tasks), close
inspection of the results of the simulations indicated that our conclusions about stability
properties would not be affected by further disaggregation of the population sector. For
this reason, we chose not to do so.
2. The forest succession sector, which tracks the composition of forest, forest productivity, productivity of the swiddens, and changes in productivity and recovery rates depending on swidden practice and population size. The major function of this sector is to
account for changes in forest succession and its impacts upon restoration of productivity.
The succession sector is sensitive to various human actions, such as forest cutting rate
and delayed abandonment of gardens.
3. The food, diet, and disease sector, which translates productivity into caloric output,
calculates caloric availability per capita and sets the level of malarial impact on the population.
4. The pig population sector, the main functions of which are (1) to serve as a sink for
some part of the productivity of the system and (2) to serve as a trigger mechanism for
the ritual festivals characteristic of Maring society.
5. The festival sector, which is the trigger for a period of warfare with neighbors. In
this simulation warfare never leads to a rout from the territory; its role is restricted to
being a source of mortality.
The principal causal loops in this simulation were developed from a variety of sources,
but the simulation constructed by Shantzis and Behrens (1973) was the original source
of the structure of the program. This simulation was reprogrammedwithout substantial
change and its behavior investigated in an earlier paper (Foin and Davis 1984). Although
the general orientation of the original simulation was retained, the present one differs in
several important ways:
1. The population sector has specific loops for malarial effects. As the population grows
and experiences declining food supplies per capita, the death rate due to malaria in-
MODELS
ANDNONEQUILIBRIUM
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SWIDDEN
SWIDDEN
LANDS
PRODUCTIVITY
FOREST CUTTING
DECISIONMAKING
FOOD SUPPLY
PER CAPITA
MALARIAL
DISEASE
IMPACTS
FOREST SUCCESSION
AND PRODUCTIVITY
HUMAN POPULATION
DYNAMICS
WARFARE
FESTIVAL-WARFARE
TRIGGER
PIG
POPULATION
DYNAMICS
Figure 1
The principal causal loops of the local simulation model. The arrows indicate the major
causal flows in the model. By convention, the variable at the tail of the arrow has one or
more specific impacts upon the variable at the head of the arrow.
creases as a consequence of reduced host resistance and increased local endemicity of the
mosquito vectors, following the work of Buchbinder (1973) and Lowman (1980). In addition, malaria also reduces vigor such that fertility and early infant mortality also increase (Buchbinder 1973, 1977). Malaria acts as a classic regulatory mechanism, operating on population density through the effects of dietary adequacy. Samuels (1982) has
developed a simulation for the Maring, which also depends upon a significant role for
disease in population control.
2. The forest productivity sector was completely reconstructed. Shantzis and Behrens
built in strong sensitivity to overuse of the forest such that collapse was inevitable once
forest degradation reached a certain point. In this simulation we have developed a successional sequence in which the forest is partitioned into a number of categories. Overuse of
the forest affects recovery both quantitatively and qualitatively, but does not necessarily
lead to irreversible forest destruction. The evidence available on Asian montane forests
strongly supports this view rather than the more extreme scenario put forward by
Shantzis and Behrens (Paijmans 1976; Manner 1981; Dove 1981).
3. The forest succession and cutting sector features explicit decision-making behavior
absent from the Shantzis-Behrens simulation. The behavioral variables include a preference function for forest type and age; limits on per capita ability to clear forest; adjustment of cutting rate as a function of dietary quality; and control over swidden retention
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and fallow period intervals. According to the literature, flexible decision making is commonly associated with shifting cultivation (see, in particular, Conklin 1954, 1957; Dove
1981).
4. The ritual festival-warfare cycle differs between the two simulations. Shantzis and
Behrens followed Rappaport (1968) in utilizing pig/human ratios and incidents of pigs
raiding gardens as ritual festival triggers. Both of these triggersdepend upon pigs becoming nuisances. The present simulation views pigs as a valuable economic commodity; the
ritual festival is triggered only when there are sufficient numbers of pigs to support an
adequate festival (Salisbury 1975; Peoples 1982; Boyd 1985). The warfare phase is substantially the same in both simulations, differing principally in the rate of mortality per
episode (Foin and Davis 1984).
Detailed Structure of the Simulation Model
Regulation
Population
The Maring population grows or decays exponentially when rates are fixed. The equation is
(1)
H, ,=
H,+
r-H,-
(D, + D)
where H is the Maring population, rH is the net growth rate, D. is the number of deaths
in the interval t to t + 1 due to warfare, and Dd is the number of deaths due to disease.
The parameter rHis calculated from a range of variables (- 15% to 1.5%), using a TABLE function based upon dietary adequacy and forest stocks, equally weighted. Dietary
adequacy is normalized on 742,000 kcal/capita/annum, the number used by Shantzis and
Behrens. The forest stock function is % total mature forest as fraction of total territory.
Denters episodically with warfare at 3% of the total population. Ddis also calculated
with a TABLE function based on dietary adequacy; the mortality rate ranges nonlinearly
from 0.0% to 20%. There are no data to support these values, so they were subjected to
extensive sensitivity analysis.
ForestSuccession
The forest succession sequence consists of three categories: mature primary forest, mature secondary forest, and immature forest. The distinction between primary and secondary forest is principally one of historical use: primary forest has not been cut within
recent memory, while secondary forest is cut on a regular rotational cycle (Dove 1981).
Botanical differences tend to be quite minor (Whitmore 1975). Immature forest refers to
early stages of regrowthand recolonization of abandoned swidden plots. This is estimated
to be 8 to 15 years following abandonment; in the simulation the longer time was used.
There is also provision for an anthropogenic grassland succession when swiddens are kept
in production too long.
Succession is an input-output process for any one vegetation group. In mathematical
form succession is
(2)
F, ,= F, + (I - O)(t)
where Fis the number of acres of vegetation type F, I is the sum of the input rates (succession from less mature vegetation types), and O is the sum of the loss rates (succession into
the next higher category and losses to cutting for swiddens). All vegetation categories are
transitory in this simulation except for primary mature forest, which will persist indefinitely in the absence of cutting. Normally succession is from old swiddens to immature
forest, as the forest invades the site. However, with extended use of the swiddens (subject
to a maximum of four years), some proportion of the abandoned swiddens will convert
to anthropogenic grasslands typified by Imnperata
cylindrica.If such grasslands are scat-
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tered, they represent only a temporary delay in return to forest. Dove (1981) found that
grasslands in Kandep territoryin Kalimantan would be overtopped by forest and shaded
out in four years). Under very heavy cutting rates, when acreage under total mature forest is depressed, then grasslands are more persistent as a consequence of diminished ability of the forest to recover.
Literatureestimates on succession rates are not very precise. Succession to mature secondary forest was estimated to be 15 years, but it could be as short as 8 years (Davis 1973;
Sabhasri 1978); it is unlikely to be very much longer than 20 years. The least certain time
is for succession into primary forest, estimated in this simulation to require 100 years.
SwiddenCuttingandDecisionMaking
In the simulation, swiddens are cut according to certain decision rules. Each member
of the population is assumed to share a preference function for forest type. Thus, Iban
are primaryforest swiddeners (Freeman 1955) while Karen and Maring prefersecondary
forest (Kunstadter, Chapman, and Sabhasri 1978; Clarke 1977). The cutting preference
for the population is easily modeled by postulating various forms of relationships between
the preferencefor type and proportion of secondary in mature forest. The Maring prefer
secondaryforest and will switch to primary forest only when secondary forest is rare compared to the former type. This preferencewas established using a TABLE function in the
simulation.
The acreage cut in the simulation depends upon dietary quality in the previous time
step. If the harvest (measured in calories) at time t - 1 is adequate, the simulation only
replaces the swiddens returning to forest. If caloric intake is not adequate, then more
forest is cut, subject to a maximum of 0.085 ha per capita which, following Dove (1981),
we take as the limit set by labor. This is an action taken to restore caloric output from
the forest to adequate levels. Cutting rates ultimately affect the average fallow time for
the average plot of forest.
Finally, the simulated swiddeners also can control the time period in which they use a
given plot (swidden retention time). This generally occurs simultaneously with expansion
of cutting, since it is also a function of dietary quality, subject to a maximum retention
period of 4 years. As noted above, the longer the retention time, the greater the subsequent environmental degradation, expressed in delayed forest recovery and grass invasion of the swiddens.
Swiddens can be cut from any forest type and/or from grassland. Swiddens from each
type are accounted for separately, since each type has a different initial value for productivity. In the present simulation these values (in 106 kcals/acre/yr) are 5.2 for primary
forest, 4.4 for secondary forest, 2.2 for immature forest, and 0.5 for grassland-derived
swiddens. These values are assumed to hold for 1.5 years, the base value for retention
time, but to decrease as average fallow times decrease. This provision incorporates the
effect of declining soil fertility known to affect swidden productivity (Nye and Greenland
1960; Sanchez 1976), although not directly. The literature suggests that productivity is a
function of vegetative biomass (Pelzer 1978; Harcombe 1977; Manner 1981).
andRitualFestivals
Pig Populations
The pig population grows exponentially at rates ranging from 6% to 14%. It will grow
at lower rates in the absence of human husbandry, but achieves maximum rates only
when a portion of garden produce is fed to the pigs. The specific forms of mortality imposed upon the population are (1) a low annual rate of killing (1%) for use as sacrifices
during illnesses and for those pigs caught raiding gardens; and (2) catastrophic mortality
(75% to 90% in the literature) when a ritual festival is staged. A ritual festival is staged
only when the pig population meets or exceeds 100 animals (an imprecise figure but approximately the same argued by Rappaport 1968). If the herd is slaughtered, 75% of the
herd is killed, the remainder being reserved as a starter herd for the next generation.
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Simulation Strategy
The simulation was used to evaluate the inherent stability of the agroecosystem. Thus,
we constructed the simulation as described above with data taken from the cited literature, supplemented as needed by our own estimates. The model output obtained with the
best estimates for parameter values is referred to as the "baseline" version. Once the
simulation was debugged and verified, it was further modified to include a standard set
of response variables (behavioral indicators), in addition to state variables already present in the outputs. The additional response variables are the derivatives of selected variables.
The simulation strategy used was standard for simulations of this type. We compared
the effect of a specified change in simulation structure or parameter values by comparing
the response variables of the experimental run to the baseline, and where appropriate,
by normalizing using the baseline version, in a fashion similar to that used by Miller
(1974) and Miller, Butler, and Bramell (1976).
All runs of the swidden simulation were carried out using a period of 400 years, with
integration step size of 1 year and print/plot intervals of 10 years.
Simulating Regional Equilibrium
Evaluation of Lowman's (1980) regional stability model requires a simulation that is
sectored into several local units. The simplest model that accomplishes this would consist
of several local units that are dynamically equivalent. The regional simulation was constructed by linking four of the local equilibrium models with a set of explicit migration
provisions. Each local unit was subject to immigration and emigration rules developed
from Lowman's hypothesis, i.e., when forest stocks were large and productivity high in a
given local unit, that unit would attract immigrants from the pool or potential migrants
from other groups. The immigration rule draws from the pool, and all units with net
outmigrationpotential contribute equally to that pool. Conversely, when population density is high, productivity is down, and forest stocks are limited, a local group shifts to net
emigration, corresponding to the hypothesis that it loses its attractiveness both to outsiders and local residents. In essence, local groups are attractive when forest stocks are
good and diets are adequate, and unattractive when forests decrease and dietary quality
declines.
There are three equations governing migration behavior:
(3)
PMS(i,t)= MF(i,t)/MF(t)
when PMS is the propensity to migrate for an individual in the ith group at time t, MF(i,t)
is the proportion of standing mature forest in the ith group at t, and MF(t) is the total
mature forest in all groups at t;
(4)
CMP(i,t)= DT(i,t)*HP(i,t)
where CMP is the contribution of the ith group to the migrant pool at t, HP is the population size of group i, and DT is the proportion of the group that will migrate. DT is
specified as a TABLE function which outputs the proportion of the population that joins
the pool as a function of diet (range of output: 23% to 98%); and
(5)
M(i,t) = PMS(i,t)*MP(t)
where M is the actual number of migrants and MP is the size of the migrant pool at t.
The migrant pool is emptied at each step of the simulation by allocating all individuals
in proportion to the forest stock available locally. Simulation control parameters for this
model were the same as for the local stability model above.
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Complete listings of both simulations, including lists of the variable names, are available on request from the senior author. The DYNAMO language and simulations are
implemented on the DEC 1173 system.
Results
Behaviorof theBaselineSimulation
Inspection of the outputs of the baseline simulation (Figs. 2-6) shows that the system
is inherently stable with the parameters chosen, in the sense that the derivatives of important variables go to zero, even though the equilibrium values of these variables are
not the same as the initial ones. The Maring population initially grows rapidly, approaching an asymptotic equilibrium at 295 individuals approximately 170 years into the simulation (Fig. 2). Simultaneously mature secondary forest declines from -700 acres to
200 acres, to be replaced by successional forest (coded as immature secondary forest, IMF
in Fig. 3). All vegetation types reach steady state in less than 50 years. The baseline simulation predicts very little grassland invasion into old swiddens because plots are abandoned quickly enough to permit the forest to regenerate normally. Average utilization
times (UT, Fig. 4) fluctuate around 2 years, which is insufficient to triggermuch reversion
to grasslands. Return times (RT), defined as the time between use of a particular piece
of land, fluctuate more widely but seldom fall below 15 years. Both estimates compare
favorably to the literature, although RT may be slightly too high.
The derivatives associated with population pressure on the land (Fig. 5) all fall to zero
asymptotically or go to a limit cycle as a consequence of the diet-disease loop. Disease
incidence rises as population pressure on the forest reduces productivity, and ultimately
forces the population growth rate to zero. In turn, reduction of population growth derivatives permits the derivatives associated with the state of the forest (Fig. 6) to go to zero
as well. The process of stabilization by increased impact of disease occurs only after di400
0
P
U
L
A
T
300HP
I
o
N
S
I
200
100
0
100
200
300
TIME IN YEARS
Figure 2
The population growth curve for the baseline simulation.HP = human population.
400
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800
F
0
R
E
-~
. ..---
"
-.-
600
I
v
E
R
400
I
N
A
C
200
R
Sxxxx
S
EXXX
_X._X
X_
XXX
XX
xx
xx
xx X
xx
cp
$
00
100
200
300
400
TIME IN YEARS
Figure 3
Growthcurves for four vegetation types in the baseline simulation. P = mature primary
forest; S = mature secondary forest; I = immature forest; and G = anthropogenic grasslands.
etary quality begins to fall, which means that the forest resources are under increased
pressure when disease increases, in accord with the field observations made by both
Buchbinder and Lowman. Consequently, most of the response measures show limited
fluctuations as the system moves toward equilibrium (e.g., food per capita, return time,
utilization time, and ephemeral appearance of grass invasions in a small proportion of
the swiddens). The exceptions are the behavior of the pig population, which is expected
to show continuing variation due to harvesting for the ritual festival, and the group of
variables associated with quality of the diet. Since the diet variables control system regulation, this variation is to be expected. Extensive simulation with differentvalues of the
malarial mortality vector showed that the critical rates are those operating when diet is
80% to 100% of the desired value. This is due to the low potential rate of population
growth; not many deaths are required to limit population increase, so even low mortality
rates are sufficient. Thus, the amelioration of malaria by malnutrition may be a real phenomenon, but should have only limited effects on the stability of the system.
These results suggest that local stabilization is a reasonable model for the New Guinea
Highlands, but it is important to remember that the model has very limited ability to
choose from the number of competing mechanisms for regulation. Nevertheless, the simulation model supports the feasibility of malarial mortality as a control agent in the Maring ecosystem, and it rejects the Rappaport model. The baseline simulation, as well as a
number of variants, all show that disease mortality is about ten times greater than war
mortality in the system. War mortality becomes important only under extreme conditions. In the simulation, at least, it is malarial mortality stimulated by malnutrition which
imposes control on population growth.
Behaviorof theRegionalStabilitySimulation
Linking four identical groups having equal territorysizes using simple migration rules,
in accord with Lowman's hypothesis, fails to produce local disequilibrium (Fig. 7). In-
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30
R T
E220
I
I
IY
IS
TII
10
YE
200
100
TIME
IN
300
400
YEARS
Figure 4
Changes in return (RT) and utilization (UT) times indicate changes in the stability of the
agroecosystem. Neither return nor utilization time exhibits any significant trend.
stead, all four groups converge to equilibrium values (2-375 persons) which are held
thereafter.Differences in initial conditions do not affect the final outcome.
The failure of the model to reproduce the hypothesized developmental cycle with the
simple migration rules used can be explained as follows. For a group early in the development, with low population densities and abundant standing forest, early population
growth will be rapid due to the influx of immigrants from other groups with limited forest
stocks. However, growth rates will slow as the surplus of forest is used. Groups that have
reached peak densities and may have begun to decline will experience net emigration,
which initially will accelerate their decline; but as densities fall and forest regeneration
begins, the probabilities of emigration and immigration will converge and densities will
stabilize. Thus, it is obvious that each group can be expected to reach an equilibrium
density.
One way to destabilize each local group would be to make migration behavior more
complex. For example, it is possible that at the early phase immigration should be strong,
but that at or after the peak population is reached, that emigration should be limited by
some combination of social and political factors. This supposition has a problem, however; if emigration is limited, then that group will remain large enough to survive chronic
warfare and will dominate neighboring groups indefinitely. We require emigration to
weaken that group relative to others, but emigration must cease before that group simply
disperses. In any case, simulation of these rules produces no change in outcome; each
group still reaches a stable equilibrium. We were unable to find any combination of migration rules that produces the desired cyclic behavior.
Lowman's model embodies the implicit assumption that the resources controlled by
each group are approximately equal. Groups that are founded in small territories (limited, for example, by steep slopes, small size, or low soil fertility) should not be as successful as groups with greater resources, if only because their maximum population size
will be limited. If the population is small enough and not successful in gaining depend-
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[89, 1987
15
E
R
I
V
A
T
V
10
\
\
-\R
5-
A
L
I
IFP;
100
200
300
400
TIME IN YEARS
Figure 5
Three system derivatives: human population growth rate (R), change in population size
(HP), and caloric intake per capita (FPC).
able allies, it will be defeated in war sooner or later, never having attained enough population and power to reach the peak of the developmental cycle. Simulation of this alternative was easily achieved by simply changing the parameters controlling territory size
for each of the four groups. The results of this change (Fig. 8) show that the dynamics of
the four-group system are not fundamentally altered, except to change the equilibrium
level of each group to reflect the resource base (measured by territorysize) of each group.
We also examined the two criteria for a rout to occur. The principal criterion is that
there be a numerical disadvantage of about 2:1 for the group in degraded forest (following
Rappaport 1968). Although it was easy to simulate small groups they would not be defeated routinely unless their territorieswere so small that a neighbor in a larger territory
could fulfill the 2:1 rout criterion most of the time, or nutritional state so poor that net
emigration reduced the population size to the critical proportion compared to hostile
neighbors. The original criterion for emigration was that the average diet was only 60%
of normal, but it did not create sufficient outmigration to ensure a defeat. When the simulation model was changed to permit outmigration at 90% of normal caloric intake, routs
did occur, but far too frequently to permit a newly founded, small group to undergo the
hypothesized developmental cycle (Fig. 9). The village with the smallest territorycannot
grow large enough to escape frequent routs by its larger neighbors, while the other three
groups are unaffected. This points out how sensitive the dynamics of the "cycle" are to
small differences in parameters.
This analysis casts doubt upon the veracity of Lowman's hypothesis as a model for
Maring population dynamics. There must exist conditions under which a 200-year cycle
can occur, but they are empirically unrealistic. Our simulations show that, as intuitively
appealing as Lowman's model is, it is not very likely to exist in nature.
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20
D
E
R
I
V
A
10
-
UT
"
*
I
E
V
A
L
U
E
--_-- - - - - -
PSL
-10
- - - . -- .
.
-.
.. . -.--
-M.-
"
PSF
-200
100
200
300
400
TIME IN YEARS
Figure 6
System derivatives for UT, RT, and PSF (% secondary forest).
500
400
L-D
T
-
0
N
200
C/
0-
D
0
100
200
300
400
TIME IN YEARS
Figure 7
Population growth curves for four villages with initial values of A = 450, B = 300, C =
150, D = 20.
ANTHROPOLOGIST
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[89, 1987
D
P
0
P
4 00
z
A
A
II
N
E
s
I
200
100
200
300
400
TIME IN YEARS
Figure 8
Behavior of the regional equilibrium simulation model with the same initial populations,
but also with differences in territory sizes. Territory size of A = 400 acres, B and C = 1,000,
and D = 1,600.
EstimatesofReturnTimesin theLocalSimulationModel
Elasticity, or return time, may be defined as the time required for selected response
variables to return to their equilibrium values after a perturbation of known timing and
magnitude. Estimating return times is important in differentiating equilibrium and disequilibrium models. If return times are long, the possibility that these New Guinea Highland systems are in disequilibrium much of the time is enhanced, while if return times
are short, disequilibrium models are probably irrelevant.
One important aspect of return time analysis is the selection of response variables.
Those selected in the Maring model were human population size and standing mature
secondary forest. These two were chosen because they are the principal state variables in
the model, one representing the population requiring some form of regulation, the other
representing the principal environmental resource. Three variables were chosen for perturbation: population size, forest stock, and secondary forest fertility. The first two of
these variables were subjected to a one-time, specified percentage change imposed at year
200 in the simulation (year 200 was chosen because the system had reached equilibrium
by that point). Site fertility was changed at year 200 but maintained at the new level
throughout the remainder of the simulation. Return and relaxation times are estimated
as the time, to the nearest ten years, to return to the original value at year 200. Where
these times exceeded the remaining time in the simulation, mean values of the response
variables for years 300 to 400 inclusive were calculated and normalized on the corresponding values for the baseline unmodified simulation results.
The results of this analysis are shown in Table 1. The effects of population reduction
<20% are temporary but persist for many years, even for reductions as small as 5%. A
20% reduction has a recovery time >200 years and keeps the forest above baseline values
for an equal time. This emphasizes that an equilibrium-seekingmodel can have such long
return times that it could not be distinguished from a disequilibrium one in the field. For
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25
500
R,
--
)
...
p/
00
4 00
A
T
L
U-300
~r
0S"I
/7
B
B
N
S
200
zZ
E
100-
0-
100
200
300
400
TIME IN YEARS
Figure 9
Behavior of the regional simulation with the same conditions as Fig. 8, but with the battle
defeat criterion changed from actual diet = 60% to 90% of the lineal value.
26
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Table 1
Estimatedrecovery times (or relaxation times, for positive changes in parametervalues)
for some variablesin the local simulation model.
Response variables
Human population
Mature secondary forest
Recovery
time
Mean
%
Input variable
A. Population reduction
- 2%
30
- 5%
80
-10%
110
-15%
140
- 20%
> 200
- 4.8
281.4
B. Population supplementation
- 0.3
+ 2%
>200
294.5
+ 5%
1.4
>200
299.8
+ 10%
0.9
>200
298.3
+ 15%
1.0
>200
298.7
+ 20%
1.4
>200
296.8
C. Reduction in forest stock
-5%
10
10
-10%
-15%
30
- 20%
- 4.9
>200
281.1
D. Addition of new secondary forest
> 200
+ 5%
0.7
297.7
+ 10%
1.9
301.1
>200
+ 15%
305.9
3.5
>200
+ 20%
4.6
>200
309.2
E. Lower swidden productivity from secondary forest
- 10%
- 10.4
NE
264.8
- 50%
NE
114.4
-61.3
F. Enhanced fertility of secondary forest
+ 10%
NE
11.6
329.8
+ 50%
NE
469.6
58.8
+ 100%
NE
110.3
621.8
Recovery
time
Mean
168.8
3.7
NE
NE
183.4
392.9
12.6
141.3
NE
NE
NE
145.1
97.8
70.5
- 10.9
- 65.0
- 92.9
80
110
160
160
>200
10
110
150
150
180
10
40
40
90
80
80
80
120
Note: Where recovery or relaxation times exceed 200 years, the mean of the response variable
averaged over the last century of the simulation, and the % change from the baseline simulation
are given. NE = new equilibrium value. Baseline mean population: 295.6. Mean secondary forest:
162.8 acres.
times. If such disturbances are realistic for New Guinea populations, the disequilibrium
model is the one which is the best description of reality.
Discussion
TheValidityof theRegionalEquilibriumModel
Of the three Highlands models considered in this paper, the regional equilibrium
model is most easily rejected. Recall that we were unable to find any set of simple rules
that produced local instability. Indeed, local stabilization appears to be a robust feature
of this model, even when more elaborate migration rules are used. The only migration
rules that would work are those stipulated by the hypothesis, requiring potential emigrants to respond to different causation depending on the size of the population. It is
FoinandDavis]
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27
difficult to imagine how these might arise, suggesting that the regional equilibrium model
should be rejected.
There are also logical difficulties associated with this model, some of which are raised
by our simulation analysis. Forest recovery rates are relatively rapid, on the order of 12
years (Sanchez 1976). This is very short relative to the estimated length of the cycle,
which argues for rapid stabilization of population by changes in migration, and thus adds
to the credibility of our analysis. Although delayed forest recovery could prevent stabilizing immigration into groups late in the developmental cycle, empirical information
indicates that this possibility is unlikely.
The second problem is that differences in available resources stemming from variable
territorysize, rather than a cycle of resource depletion, can account for some local disequilibrium. Even then, the pattern of disequilibrium is not identical to the long-term
growth and depletion cycle proposed by Lowman. Third, it is not clear why a population
would not, while at the optimum point of its ecological and political success, simply use
its warriorsand status to annex quality territoriesoccupied by neighbors both earlier and
later in their own cycles of development, rather than allow itself to degenerate.
Collectively, the evidence available does not support the local disequilibrium/regional
equilibrium model. Furthermore, we have raised a number of logical difficulties which
cast doubt upon the probability of ever demonstrating its application to real populations.
It is difficult to see how sufficient empirical research could be marshalled to test this
model; moreover, our results suggest that such an empirical study would not be worthwhile.
EquilibriumVersusDisequilibrium
The local equilibrium model, especially Rappaport's version, has been criticized on
several points. Both MacArthur (1974) and Buchbinder (1977) have presented evidence
that the Tsembaga Maring are not very well nourished, a condition that is inconsistent
with the hypothesis that the population is regulated below the critical point of sustainable
yield. Bates and Lees (1979), Samuels (1982), and Peoples (1982) all criticize the group
selection aspects of Rappaport's model, on the grounds that it is not clear how the equilibrium-seeking requirements of collective action could be translated into incentives for
individual behavior-how, in short, such a system might have evolved. Salisbury (1975)
points out that the Tsembaga have been in place far too briefly (perhaps only 200 years)
to have had the opportunity to develop such a finely tuned adaptation as Rappaport postulates. Foin and Davis (1984) reconstructed the Shantzis-Behrens simulation model, but
used parameter values more consistent with the data provided by Maring scholars. As
the model is highly sensitive to any change in key parameters, the system destabilizes
very rapidly when war death rates are reduced to realistic values.
These lines of evidence argue against a highly adaptive and efficient scheme such as
Rappaport proposed for the Tsembaga. However, our simulations support less finely
tuned models, such as the one proposed by Buchbinder. The Maring models have strong
equilibrium-seekingbehavior within each group which are inconsistent with the Lowman
model.
The long return times, however, are more consistent with the disequilibrium model,
since it is likely that population disturbance and technological change are frequent
enough to prevent the population from maintaining an equilibrium very much of the
time. Return time analysis using the simulation model indicates that one significant displacement every few decades would be sufficient to keep the system in disequilibrium.
Those who have argued for the importance of nonequilibrium models for New Guinea
ecology (Strathern 1971; Salisbury 1975; Golson 1982) are correct insofar as they refer to
disequilibrium. There is an important distinction to be made, however, between disequilibrium and nonequilibrium (the term used by Salisbury 1975), since our research
supports the former strongly but not the latter, at least for shifting agriculturists in Highland New Guinea.
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The most important conclusions to be drawn here include (1) the Maring populations
are equilibrium seeking; (2) with limited rates of change and sensitivity to disturbance,
the Maring are most often moving transitionally from some displacement toward some
equilibrium; and (3) it is extremely difficult empirically to detect equilibria under these
conditions. Even in a system seeking equilibrium, long recovery times render it likely that
significant perturbations will occur in the interval.
The possibility that Maring populations are equilibrium seeking but in disequilibrium
much of the time has several important implications. First, there may be a Heisenberg
Uncertainty Principle in operation, especially if exogenous variables are constantly redefining the equilibrium population size. In that case, it may be impossible to conduct a
meaningful empirical study of population regulation, since the system itself may well
have changed by the time any one study has been completed. If this is true, empirical
studies of population regulation should attempt to define the regulating mechanisms in
force at the time, with less emphasis on the equilibrium-disequilibrium issue. Alternatively, one could identify the most important disturbances, measure their frequency, and
estimate population response rates. At present, it could be claimed that Maring agroecosystems are stable, since they will return to equilibrium in the absence of disturbance,
but it could also be claimed that they are unstable, since they are seldom at equilibrium.
The probability that episodic mortality and technological or environmental changes occur argues that the disequilibrium model is the best descriptor of Maring dynamics.
It is important to emphasize that the key issue is the quality of information pertaining
to mechanisms governing equilibrium-seeking behavior and the nature and frequency of
factors that disrupt system equilibria. This restates Harris's (1968) comment. Such information is relevant to human social and ecological systems in general, not merely the
Maring or shifting agroecosystems. Whether the system is in equilibrium or exists in a
more or less permanent state of disequilibrium is less relevant. In any case, it may be
impossible to determine.
We acknowledgethe supportof the UC MEXUS Program,USAID StrengthAcknowledgments.
eningGrantProgram,andThe Instituteof Ecology,Universityof California,forfinancialsupport
of theresearchreportedhere.DavidBoydand StephenBrushcarefullyreada muchlongerversion
of the manuscript, to our great benefit. We thank four anonymous reviewers for their extensive,
constructive comments on the manuscript.
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