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still occur before final publication
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online and in print.)
C E
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D V A
Disturbance, Complexity, Scale:

New Approaches to the Study
of Human–Environment
Interactions∗
Access provided by Florida Atlantic University on 09/24/15. For personal use only.
Annu. Rev. Anthropol. 2015.44. Downloaded from www.annualreviews.org
Rebecca Bliege Bird

Department of Anthropology, Pennsylvania State University, University Park,
Pennsylvania, 16802; email: [email protected]
Annu. Rev. Anthropol. 2015. 44:241–57 Keywords

The Annual Review of Anthropology is online at human behavioral ecology, sustainability, ecosystem engineering, niche
anthro.annualreviews.org
construction, complex adaptive systems
This article’s doi:
10.1146/annurev-anthro-102214-013946 Abstract
Copyright c 2015 by Annual Reviews. New approaches to human–environment interactions are beginning to move
All rights reserved
beyond a narrow focus on individuals and simple (patch-level) predatory or
∗
This article is part of a special theme on competitive interactions. These approaches link nonequilibrium theory from
Resources. For a list of other articles in this theme,
see http://www.annualreviews.org/doi/full/ community and landscape ecology with theories of individual decision mak-
10.1146/annurev-an-44-themes ing from behavioral ecology to explore new ways of approaching complex
issues of diachronic change in behavior, subsistence, and social institutions.
I provide an overview of two such approaches, one to understand long-term
hunting sustainability among mixed forager-horticulturalists in the wet trop-
ics and the other to understand how foragers act as ecosystem engineers in
a dry perennial grassland in Australia. I conclude by describing the impli-
cations of new approaches that incorporate anthropogenic “intermediate”
disturbance (an emergent property of human–environment interaction) as
a force shaping environments through time and space, and in so doing pat-
terning the sustainability of subsistence, ways of sharing, ownership norms,
and even structures of gendered production.
241
Changes may still occur before final publication online and in print
INTRODUCTION
Models of individual decision making developed in behavioral ecology have proven remarkably
suitable for explaining the adaptive nature of human–environment interaction (Smith 2013, Smith
& Wishnie 2000), especially when considering simple interactions: the decision of whether and
when to take certain shellfish or pass them over (Bird & Bliege Bird 1997), which habitats to
colonize first and whether to defend them (Winterhalder et al. 2010), or how far to transport
items with low utility (Bettinger et al. 1997). Likewise, simple ecological models of predation and
competition are utilized to explain how the environment is affected by human decision-making
processes (Charnov et al. 1976, Nagaoka 2002). Such simple models have also proven useful in
unveiling many of the dynamics that accompany more complex phenomena, such as the interac-
tive effects of the environment on the development of social hierarchy and the colonization of
continents (Kennett et al. 2006, 2009). However, most larger-scale, dynamic social and ecological
processes have proven more difficult to explain, as evidenced by the substantial debate over their
applicability to the understanding of subsistence intensification and the transition from hunting
and gathering to agriculture (Gremillion et al. 2014; Smith 2007, 2011a,b; Zeder 2012).
The most common ecological approach to questions of human–environment interaction, such
as the processes driving subsistence intensification, combines models of human response to differ-
ent environmental conditions (i.e., central place foraging models; prey, patch, and habitat choice
models; and territorial defense/marginal valuation models) with models of human impact on the
environment, primarily predator-prey models. Such models are used to generate predictions about
how human populations cause resource depression, how this influences patterns of mobility and
residence, and how population density and the colonization of less suitable habitat may result in
declining residential mobility and increased sociopolitical stratification (Bird & O’Connell 2006,
Kennett & Winterhalder 2006, Lupo 2007, Morgan 2014). The models predict that the costs
of traveling elsewhere increase as more human populations occupy neighboring habitats, caus-
ing foragers to spend proportionally more time handling lower-ranked resources, and reducing
the opportunity costs of investing in technology to reduce handling costs (Bettinger et al. 2006,
Hawkes & O’Connell 1992). This leads eventually to plant domestication and greater investments
in agricultural production.
Debates over the limitations of these models have highlighted two main problems. First, these
models do not explain the origins of or variations in emergent social phenomena such as the
institutions that accompany complex events such as the origins of agriculture. Coevolutionary
approaches, such as that of Bowles & Choi (2013), address this limitation by proposing a sort
of ratcheting-up link between economic and social institutions. However, in these models the
absence of ecological dynamics and of attention to individual decision making does not allow for
the incorporation of adaptive processes other than group selection transforming social institutions.
Second, these models are designed to explain ecological interactions at very local spatial and
temporal scales, and not population- or community-level dynamic interactions. They focus almost
exclusively on dyadic relationships between species and on negative interactions such as competi-
tion and predation, and they do not incorporate nonequilibrium scale-sensitive approaches, which
would allow one to capture how negative effects at one spatial scale can result in positive outcomes
at the level of the landscape or the community as a whole (Huston 1979). Niche construction
or ecosystems engineering approaches such as those of Zeder (2012) and Smith (2007, 2011a,b)
propose that such complex phenomena as the origins of agriculture are best explained by the
anthropogenic construction and management of ecosystems to improve subsistence productivity.
However, as Smith (2013) points out, although such models succeed in calling our attention to
the dynamic and interactive effects of ecology and social structure, they provide no framework
242 Bliege Bird
for understanding individual decision making with regard to resource use beyond a vague notion
of intentionality in innovation, according to which actors intentionally modify habitats to gain
future, delayed payoffs. The solution to these issues is not to discard models of agency, nor to
model social evolution in an ecological vacuum, but rather to generate new integrative models
that can scale up from individual decision making to its social and ecological causes and conse-
quences at the level of populations and ecological communities. As Layton put it, “our best hope
for reconciling Social and Evolutionary Theory lies in an ecological approach to social evolution
that acknowledges the emergent properties of social interaction and their capacity to modify the
environment to which individuals are adapting” (Layton 2010, p. 149).
I suggest that if behavioral ecology is to have any hope of generating better models of complex,
dynamic social-environmental interactions, it must find ways of integrating individual decision
making within ecological and evolutionary models that operate at broader spatial and temporal
scales. This new approach should seamlessly integrate what are often perceived as theoretical di-
vides between nature and culture, agency and adaptation, behavior and institutions, humans and
other species, and individuals and communities. It should include humans as part of the funda-
mental structure and function of ecosystems; pay attention to the dynamic interactions among
subsistence, social structure, social institutions, and the environment; and include considerations
of different temporal and spatial scales, of variation, and of the complex ecological interactions
between humans and other species. Finally, it should take a coevolutionary perspective to account
for the interactive effects that take place over time between individuals (cooperation, conflict,
cultural transmission), between species (mutualism, predation, competition, facilitation, niche
construction/ecosystem engineering), and between scales of organization, from the individual to
the population to the community.
Here I provide an example of such an approach, which reconsiders existing explanations of
subsistence intensification by linking insights from research on community and landscape ecology
with theories of individual decision making from behavioral ecology.
A THEORY OF ECOLOGICAL INTERACTIONS

Typical models of subsistence intensification usually assume that predation reduces populations
of economically profitable species, causing shifts to lower-profitability resources. Although there
is ample evidence of local population depression, evidence for the hypothesis that indigenous
hunters completely exterminated species is quite limited and contentious (Meltzer 2015). There
is actually a long history of sustainable exploitation of most human prey: Even the flightless
duck in California, an easy mark for native hunters, coexisted with California coastal foragers for
8,000 years ( Jones et al. 2008). A narrow focus on the local-scale, dyadic interactions between
humans and their prey overlooks the more complex effects of predation at larger scales and its
indirect effects on other species that might be facilitating these sustainable interactions.
Predation is generally a source of disturbance to populations, and such disturbance, although
negative at some temporal and spatial scales, can produce positive effects at others. Positive ecolog-
ical interactions, often referred to as facilitation, include any direct or indirect force that positively
affects the growth or reproduction of one or more organisms without negatively affecting other
species (Bertness & Leonard 1997). At the level of the community as a whole, positive effects
are modeled as a function of the scale at which disturbance (mortality) occurs; intermediate dis-
turbances are hypothesized to produce heterogeneous landscapes and source-sink dynamics that
maximize the persistence of a wider range of coexisting species (Connell 1978, Huston 1979).
Positive effects have been documented for a wide range of predatory interactions; in some
interactions, predation actually increases prey populations. Agouti (large rodents of the genus
www.annualreviews.org • The Study of Human–Environment Interactions 243
Dasyprocta) prey extensively on Brazil nuts, but they habitually bury seeds intact for later con-
sumption, which increases seedling survival over unburied nuts and results in a higher density of
Brazil nut trees where agouti are present (Asquith et al. 1999). Other interactions have negative di-
rect effects on the species that are preyed on but positive effects on other species or the community
as a whole. For example, in the absence of top carnivores, white-tailed deer suppress tree seedling
recruitment and deplete many herbaceous plants, which reduces habitat for ground-nesting birds;
with the introduction of top carnivores, both plant populations and the small animals that rely on
them rebound (Côté et al. 2004). Removing otters from nearshore environments causes a collapse
of macroalgae and sea grasses as herbivores such as urchins and grazing snails increase (Estes &
Palmisano 1974). Kelp forests in turn affect other species by increasing food availability for graz-
ers and reducing wave height and current velocity. In otter-dominated kelp forests, barnacles and
mussels grow three to four times faster (Duggins et al. 1989); rock greenling are roughly ten times
more abundant (Reisewitz et al. 2006); glaucous winged gulls eat more fish (Irons et al. 1986); and
bald eagles have a more diverse diet of fish, marine mammals, and seabirds (Anthony et al. 2008).
In the absence of a disturbance such as a storm or fire, pines or fir trees will gradually take over
an oak woodland (Cocking et al. 2012), reducing overall plant diversity and abundance, especially
on the forest floor, and increasing the risk of pathogen spread (Real & Biek 2007). The decline
of a top predator, such as the coyote, releases smaller predators from competition, allowing their
populations to surge and deplete prey populations. When coyotes are rare, smaller predators such
as foxes and cats are more common, and populations of scrub-breeding birds decline (Crooks &
Soulé 1999). Positive effects can also come about as a result of the engineering activities that are
a side effect of some organism’s foraging, housing, or predator evasion strategies. When such
processes construct new niches or enhance existing ones, they are often referred to as ecosystem
engineering ( Jones et al. 1994). The classic example is beaver dam construction, which increases
wetland areas and produces more environmental heterogeneity, supporting larger populations of
a wider range of species at the landscape scale (Wright et al. 2002).
The sum total of these facilitative ecological interactions affects the assembly of entire ecological
communities. When ecosystem engineers affect landscape heterogeneity, this may stabilize species
interactions (Holt 1984, Roff 1974, Roxburgh et al. 2004) and produce rescaling and habitat
protection effects on habitat generalists, which are species that require a variety of habitats for
food and shelter (Futuyma 1988, Marvier et al. 2004, Wiggins et al. 2006). When predators hunt
many different prey species at multiple trophic levels—that is, when they have wide diet breadths
(Beckerman et al. 2006, Redford & Robinson 1987)—food web stability might increase, allowing
more species to persist with more stable populations (Gross et al. 2009). Whereas individual
populations may decline and become locally extinct, the larger meta-population may be more
likely to persist, particularly if habitats are patchy and heterogeneous and predators face lower
costs of prey switching (Holt 1984, McCann et al. 2005, Rooney & McCann 2012). The species
providing more positive ecosystem effects may contribute to more stable ecological communities,
minimizing destabilizing population fluctuations and allowing many more species to coexist over
the long term. Organisms playing a key role in holding communities together are termed keystone
species. When these keystones are removed from the food webs they support, catastrophic species
loss often follows that ripples down through the trophic levels of the web, from top consumers to
herbivores, and even to the plants they consume.
Although charismatic nonhuman species take front and center in most ecological studies of
facilitation, evidence of more complex and/or positive human–environment interactions is more
slowly accumulating. One of the first ecological studies to suggest this was conducted by Castilla
and colleagues (Castilla 1999, Castilla & Bustamante 1989). Through exclusion experiments
244 Bliege Bird
on the Chilean coast, they demonstrated that moderate levels of human predation (subsistence
foraging) on a wide range of shellfish—including mussels (detritivores), limpets (herbivores), and
gastropods (predators)—increased species diversity and shifted the intertidal from one dominated
by barnacles to one evenly composed of mussels, algae, and barnacles. When humans were ex-
cluded, the predatory gastropods rapidly decimated the mussel beds, allowing barnacles to invade
and therefore preventing mussels from reestablishing. Human exclusion also increased the size of
keyhole limpets (as commonly observed archaeologically, human foragers preferentially choose
larger individuals), but this had a devastating effect on algae because big keyhole limpets graze
them down more effectively; the loss of algae also left more room for barnacles to invade. Whereas
foraging strategies with wide diet breadths seem to actually improve coastal resources for human
exploitation, narrow strategies focused on a single intertidal keystone species, such as the sea
otter, cause trophic collapse, as commercial hunting did during the nineteenth century. Castilla
(1999, p. 282) concludes that “humans affect the functioning of food webs by acting as an effi-
cient and selective keystone predator. Humans can regulate predatory efficiency and/or selectivity,
and thereby manage, enhance, reduce, conserve or reserve species or communities.” Castilla also
implies that they may do so simply by virtue of their immediate subsistence strategies, and not
necessarily through deliberate environmental modification designed to produce these community-
level effects. In the following section, I pursue this hypothesis further with two case studies of very
different ecosystems: the neotropical rain forest and the desert grasslands of Australia.
CASE 1: GARDEN GAME IN THE NEOTROPICAL RAIN FOREST

The phenomenon of garden game in the neotropical rain forest is receiving increasing attention
as researchers critically evaluate the differential effects of anthropogenic disturbance on animal
communities. Disturbed forest tends to support a broader range of species than primary forest
(Connell 1978); following this logic, it has been suggested that extensive forest-fallow horticultural
systems would enhance biodiversity and augment populations of some species by improving access
to food or new habitat niches (Fa et al. 2005, Linares 1976, Posey 1985, Smith 2005). Whereas
some species of large game become depleted around indigenous settlements in the Amazon basin
because of hunting (Alvard 1993), other species, particularly agouti, collared peccary, and deer,
seem to be scarcely affected by it (Chacon 2012, Koster 2008, Smith 2005). Likewise, whereas
some communities experience a decline in populations of preferred species over time, others,
such as the Ache, show no effects (Hill & Padwe 2000, Hill et al. 2003). Some suggest that this
is a function of population density and mobility, with seemingly sustainable offtakes expected
where hunters are at low density or are highly mobile (e.g., Alvard 1998); another hypothesis is
that significant source-sink dynamics result in population influx from more remote regions (Hill
& Padwe 2000). But there is also evidence that the increased food availability in anthropogenic
environments may actually enhance populations of these species (Smith 2005). In one community
in western Panama, a majority of agoutis, armadillos, and collared peccaries were found in the
anthropogenically modified forests and gardens surrounding the village (Smith 2005). Because
hunting may be more predictable in anthropogenic habitats, returns may actually be higher: In at
least two studies, hunters had higher returns in gardens and secondary forests, measured as catch
per unit effort and catch per unit area (Dunn & Smith 2011, Parry et al. 2009).
But the reason for these effects may be not the presence of anthropogenic habitat per se,
but rather the way that anthropogenic habitats are spatially patterned across a larger landscape.
Dunn and colleagues (Dunn et al. 2012) suggested that sustainable hunting of Baird’s tapir by
Miskitu was possible because the landscape surrounding the Miskitu community was a diverse one,
including extensive areas of primary forest experiencing low hunting pressure as well as agricultural
landscapes with a diverse set of crops offering feeding opportunities for tapirs and other species.
However, the positive effects of anthropogenic forest modification may peak at intermediate
spatial extents: As agricultural production intensifies, landscape diversity decreases, and refuge
habitat becomes more fragmented and dispersed, the potential for anthropogenic landscapes to
support a wide range of animal species may decline. In southern Ghana, cocoa monocultures,
which dominate the surrounding fragmented forest reserves, are highly depauperate and mainly
support very small animals (Schulte-Herbrüggen et al. 2013).
Although anthropogenic forest modification has been characterized as an intentional forest
management strategy (Posey 2003), the large-scale landscape level effects of human disturbance
do not need to be intentional. Indeed, even processes as complex as plant domestication (and
the patchiness and distribution of plants across a landscape) may be the emergent outcome of
the way people forage for wild plants, as Rindos (1984) first hypothesized. In Central Africa,
the fact that Baka harvest and consume wild yam at a central place facilitates the dispersal of
yams, which do not colonize new regions very efficiently. Dense patches of wild yams and other
secondary forest species colonize old habitation sites, and the activity of foraging alone is sufficient
to generate enough patches of wild yams to support populations at their current densities (Yasuoka
2013).
CASE 2: LIZARDS AND KANGAROOS IN THE AUSTRALIAN

WESTERN DESERT
Morton et al. (2011) note that among the major drivers of plant and animal adaptation in Australia
are long-standing feedback relationships between humans and environmental structure and func-
tion, mediated primarily via fire. Fire plays an especially prominent role in the summer rainfall–
dominated grasslands, savannahs, and woodlands north of the Tropic of Capricorn. In the spinifex
grasslands in this arid and semiarid zone, anthropogenic fires are mainly lit during the early dry
season, in the winter cold months, primarily to hunt burrowed sand monitor lizards (Varanus
gouldii ). Martu hunters in this region are significant predators of sand monitors, which make up
more than 70% of all foraged foods acquired per foraging day. Sand monitor and hill kangaroo
are the two major subsistence prey: Hill kangaroos are limited to rocky hills and ranges, whereas
sand monitors are most abundant in sandy substrates with extensive spinifex cover. During the
colder winter months, sand monitors retreat to shallow burrows to live off extensive stores of
abdominal fat, and at this time hunters burn regions of mature spinifex hummocks to expose the
burrows. During the summer months, monitors are tracked on the surface, and so hunts target
recently burned patches where tracks can be followed more easily. Returns are highest within
6 months following fire and decline as the patch revegetates, becoming almost negligible in grass-
lands more than 5–7 years after the fire. Due to high encounter rates and the high chances of
pursuit success, sand monitors are a staple food resource, the harvest of which can be adjusted to
need on a daily basis (Codding et al. 2010). Variance discounting models show that although sand
monitor hunting has lower mean returns than kangaroo hunting, its lower variance and higher
chances of success make it more useful for consumption than kangaroo hunting ( Jones et al.
2013). In the context of a hybrid economy that includes some reliance on purchased goods, sand
monitor hunting, unlike kangaroo hunting, responds to economic scarcity: It has been shown
that more women hunt significantly more often when money is short (Scelza et al. 2014). As
such, hunting sand monitor lizards is an important way that women, especially postmenopausal
ones, provide for their grandchildren and other dependents (Scelza 2009, Scelza & Bliege Bird
2008).
246 Bliege Bird
The high returns from sand monitor hunting depend not only on the immediate use of fire,
but also on the history of cumulative effects of Martu subsistence activities on the structure of
environmental variation. Sand monitor hunting is dependent on an anthropogenic fire mosaic
built through the accumulation of approximately 10 years’ worth of hunting fires. In two Martu
communities, Parnngurr and Punmu, between 60 and 240 individuals set broadcast fires every
3–4 days, resulting in 360 hunting fires per year in an area of approximately 500,000 ha (Bliege
Bird et al. 2012). Our analysis of 10 years of satellite images covering 46,000 km2 shows that the
landscapes emerging from hunting fires are very different from those dominated by lightning fires:
Hunting fires are much smaller and closer together than lightning fires (Bliege Bird et al. 2008).
Martu hunting fires are about 1 km apart, whereas lightning fires are nearly 9 km apart (Bliege
Bird et al. 2012). The zone of Martu influence attenuates rapidly outside of hunting regions:
The anthropogenic mosaic is tightly linked to the cost of travel from centralized places, and
regions closer to roads, settlements, and frequent camping places are more strongly affected by
anthropogenic fire than regions farther away (Bliege Bird et al. 2008).
Cumulatively, Martu burning radically rescales the temporal and spatial impact of fire distur-
bance across the region. Patches of regrowing vegetation are smaller and closer together, and the
diversity of the types of patches regenerating after fire (successional or seral patch diversity) is
greater at spatial scales that are typical of a human foraging range (Bliege Bird et al. 2008). Martu
fires also prevent more habitat from burning, protecting critical refuge for many small animals
and increasing the number and evenness of such patches throughout the landscape (Bliege Bird
et al. 2012). The effects of anthropogenic fire restructure the distribution not only of successional
mosaics and the attendant vegetation, but also of some animals. Sand monitor density is increased
in regions where there is greater environmental heterogeneity: the higher the density of habitat
edges—contrasts between new burns, regrowing vegetation, and old growth—the higher the den-
sity of sand monitor lizards (Bliege Bird et al. 2013). Mean returns in hunting sand monitor are
1.6 times higher in more heavily hunted regions than in regions that are rarely visited by Martu
hunters, and success rates are 6 times higher. Martu hunting fires also shape population distribu-
tions of hill kangaroo (Codding et al. 2014). Hill kangaroo scat density is linked significantly to
successional-stage heterogeneity: Scat counts increase with the diversity of habitat edges created
through sand monitor hunting fires. Fires also increase encounter rates with patches of seed grass,
which grow mainly in early to mid-successional patches, suggesting that fire played a fundamental
role in the late Holocene proliferation of seed-grinding technologies in the arid zone (Zeanah
et al. 2015).
The characteristics of many of the animal species that have recently disappeared or are in
decline also suggest that they too may have been advantaged by Martu fire mosaics. In the mid-
1960s, most of the last groups of desert nomads were cleared from or migrated out of the heart
of the Western Desert, returning in the mid-1980s (Davenport et al. 2005). Their departure
coincided with the local extinction of 21 species of native marsupial and the decline of 43 more
(Burbidge et al. 1988, Burrows et al. 2006, Finlayson 1961), including several common prey such
as the rufous hare-wallaby and the brush-tail possum. In their place were feral housecats, camels,
donkeys, and foxes. Recent studies have suggested that the mammal decline in this area during
the past 60 years may be fundamentally linked to the disruption of traditional hunting and land
use practices.
Anthropogenic disturbance causes overall foraging returns (including searches for large ani-
mals, small animals, and plant resources) to peak in regions of intermediate anthropogenic activity,
to decline in the most heavily used regions, and to be lowest in remote regions barely touched
by human activity (Bird et al. 2015). Small animals provide a greater proportion of the diet in
anthropogenic regions. Daily variance between individuals foraging from the same central place
(dinnertime camp, in Martu parlance) is lower in anthropogenic environments, mainly because

more individuals are successful at bringing in small game. Everyone in the camp does equally well
in an anthropogenic environment, whereas in a nonanthropogenic environment there is more
variation in returns across individuals: Some do well, but most do not. Landscape modification,
thus, at least in the Martu case, seems to reduce the temporal and spatial variance in resource
access across a camp, mainly by increasing access to high-ranked small game and plant foods (by
reducing foraging cost or increasing its productivity).
These processes have long been referred to as “fire-stick farming” (cf. Jones 1969), which
unfortunately implies that burning by individuals is designed to produce optimally diverse land-
scapes and that the adaptive maintenance of burning is a consequence of the long-term benefits
derived from the increased foraging returns that are possible in such landscapes. To the con-
trary, what seems clear is that burning is only relative to immediate hunting returns and that the
long-term benefits are emergent properties of a complex adaptive system in which Martu play an
integral role in sustaining a diverse and richly structured food web in an arid and hypervariable
environment.
FEEDBACKS INTO SOCIAL INSTITUTIONS

AND SOCIAL ORGANIZATION
Disturbances of both plants and animals, when applied over certain temporal and spatial di-
mensions can have positive effects on the stability and diversity of species interactions and may
actually favor populations of some species. These positive effects are not accomplished through
conservation, management, or foraging restraint, but through the emergent properties of foraging
decisions. The next step in this theory of human–environment interaction is to consider not only
the impact of subsistence on ecological structure and function, but also the structure and function
of both individual decision making and the properties of social groups living on these landscapes.
The impact of human subsistence on populations, communities, and landscapes should feed back
into human social organization and social institutions via foraging decisions, mobility, sharing,
gender-based division of labor, and resource defense. In this section, I explore one method of
accomplishing this by considering how the emergent properties of disturbance caused by human
subsistence activities affect social phenomena.
In both case studies presented here, disturbance seems to have similar effects: increased
landscape-level heterogeneity and a reduction in the variance associated with hunting. Central
place subsistence activities produce spatial variation in human disturbance (predation and ecosys-
tem engineering) across the landscape, focusing the strongest effects on the core regions close
to communities and diminishing outward as travel cost increases. Spatially varying engineering
activities create greater patch diversity at more intermediate spatial scales in regions close to
communities. One common effect of this seems to be a shift in the structure of animal commu-
nities: Different types of disturbance seem to favor different types of animal communities, with
smaller animals possibly favored over larger ones in more heterogeneous landscapes. Smaller ani-
mals with generalized foraging strategies seem to do very well in anthropogenic environments, so
that in some cases the benefits of habitat modification may even outweigh the costs of increased
predation by human hunters. Theoretically, a patchy, heterogeneous environment would reduce
the cost of prey switching, allowing a generalist forager to encounter more high-ranked prey
animals and to sustain lower travel costs between encounters (MacArthur & Pianka 1966). Al-
though very little is known about how human landscapes affect animal communities, some species
seem to thrive in more heterogeneous environments. Deer, which are habitat generalists, have
248 Bliege Bird
smaller home ranges where disturbance creates greater landscape heterogeneity at smaller spatial
scales (Saı̈d & Servanty 2005), and a number of species reproduce at higher rates in more het-
erogeneous landscapes, including meadow voles (Bowers & Dooley 1999) and raccoons (Dijak &
Thompson III 2000). At the same time, however, many birds experience higher rates of predation
due to increased mesopredator activity (Kurki et al. 2000), as diverse landscapes tend to increase
the proportion of a community that is composed of generalists, especially generalist predators
(Marvier et al. 2004). Increased levels of disturbance and fragmentation may act as an evolution-
ary force to favor medium-sized generalists over very small or very large animals: Over 175 years of
increasing habitat fragmentation, very small Danish animals got larger, large animals got smaller,
and medium-sized ones did not change (Schmidt & Jensen 2003, 2005).
Archaeologists are beginning to recognize the long history of human and medium-sized an-
imal coexistence, providing some support for the notion that species more likely to thrive in
anthropogenic environments tend to flourish over time, coming to dominate species assemblages
(Schollmeyer & Driver 2012). These authors review 159 assemblages from 129 sites across west-
ern North America and 57 assemblages from 23 sites on the Iberian peninsula and conclude
that most assemblages show an increased emphasis on smaller terrestrial animals over large ones.
They argue that this is not just a shift in hunting focus, but an actual increase in populations of
smaller animals relative to larger ones as foragers modify environments through their activities.
At some sites, source-sink dynamics in heterogeneous landscapes keep large game populations
high relative to small game (or some large species are advantaged by the kinds of environmental
modification humans engage in, such as tapirs or kangaroos); at other sites, smaller animals may ac-
tually increase due to the habitat improvements offered by human activity (Schollmeyer & Driver
2013).
If human disturbance tends to increase patchiness and the predictability of prey animals, as
well as encounter rates with medium-sized prey species, it is likely to decrease the costs of travel
and reduce stochasticity in returns for human foragers. Larger animals (cf. Morin 2012) tend
to be encountered less frequently than smaller animals, and they run faster. Therefore, both
on-encounter returns and daily trip returns (including search and travel time to and from the
foraging locale) tend to be more variable: There is a higher risk of complete harvest failure and
a higher chance of coming home empty-handed, but also a higher chance of a harvest bonanza
(see Bliege Bird & Bird 2008). Whereas larger animals may provide big bonanzas on average,
when the variance in reward is taken into account, such bonanzas may actually provide less utility
than a lower mean return on less variable prey ( Jones et al. 2013). In general, foraging portfolios
dominated by smaller or less mobile types of prey tend to be associated with lower intra- and
interindividual variance than those dominated by larger prey.
Sharing and Social Structure

Patchiness and predictability in turn have a suite of ramifying effects on social structure via their
impacts on mobility and on the costs and benefits of ownership and sharing. The marginal valuation
of resource defense both at the scale of individual harvest (Blurton Jones 1987, Winterhalder 1996)
and at the scale of the patch or landscape (Dyson-Hudson & Smith 1978) is predicted to affect the
benefits of defending claims to exclusive use rights over a resource. Individuals tend to share more
and defend less when there is high intertemporal variation in foraging success (stochasticity), when
package size and perishability is high, and when the degree of interforager correlation in harvest
success is low (Winterhalder 1986). Empirical observations of sharing among hunter-gatherers and
experimental work on the links between sharing norms and unpredictable rewards among Japanese
and American participants tend to support these predictions about the way people share (what
benefits they receive from sharing is another matter). Resources acquired more synchronously
and predictably, like small game, tend to be kept in larger amounts for the consumption of the
acquirer and his/her family (Gurven 2004b), whereas high-variance foods are retained less by
the producer (Hiwi: Gurven et al. 2000, Ache: Gurven et al. 2002). Independently of variance,
foods that come in large packages are shared with more people or in larger amounts than food
in small packages (Gurven et al. 2001, 2002; Kaplan & Hill 1985). Low-variance resources (e.g.,
cultivated or collected foods) may be shared more contingently, whereas high-variance resources
(e.g., stochastic income such as large animals) show evidence of long-term one-way flows (free
riding) and lack of discrimination between recipients. Among the Ache in the forest (see Gurven
et al. 2001, 2002), women worked hard to acquire large harvests of low-variance foods such
as fruit, palm heart, and palm starch, keeping a consistent portion for themselves and giving
away the remainder, usually in the form of cooked food. But they seemed to be choosy about
recipients: They gave shares to fewer other families, and such giving was more contingent (more
was given to those from whom one had received more) compared to the sharing of higher-variance
resources such as meat, which were given away without contingency to twice as many recipients,
with hunters often relinquishing control over distribution (Gurven 2004a). Similarly, Altman &
Peterson (1988) noted that among the Kuninjku (Gunwinggu), individual producers were expected
to exert stronger claims to ownership (i.e., to keep more within their own families) for small game
than for larger game. The key feature, they suggest, is synchronicity: Even cash, when acquired
asynchronously and unequally across the community, is widely shared, whereas it is more likely
to be kept for personal use when it is simultaneously and equitably provided. In industrialized
societies as well (e.g., Japan and the United States), the expectations about sharing versus keeping
can be manipulated by providing resources associated with an unpredictable link between labor
and reward: Windfall (stochastic) resources (such as high-variance animals) are commonly shared
more widely than resources with strong links between labor and production (Kameda et al. 2002),
which suggests a common mechanism behind such norm development.
Shifts in lower-variance resources, thus, are likely to bring about shifts in community-level
patterns of ownership and sharing. Those who have more low-variance resources have a surplus
because they worked longer hours to get them; this sets up an association between resources and
work effort by which things tend to be recognized as belonging to those who worked to produce
them. Thus, ownership may be likely to emerge and spread in a population when everyone has
access to economically defendable resources, when these resources are predictable in space and
time and densely distributed (so that the benefits of sole consumption outweigh the costs of
defense), and where the amount of time invested predicts harvest size. Norms of ownership and
sharing can thus emerge from the spatially explicit disturbance effects of human subsistence on
the landscape.
Social Inequality and Gender-Based Divisions of Labor

With ownership comes the potential for despotism and increasing social hierarchy and inequal-
ity, and this may be more likely where there is spatial variation in how ecosystems respond
to anthropogenic influence. If anthropogenic activity has little positive effect on resource
distribution, despotism and social inequalities may develop over time as only those who gain
access to “naturally” rich patches enjoy the benefits of owning them. Kennett et al. (2009) and
Winterhalder et al. (2010) use an ideal free distribution model to document the increase in
despotism over time on the Channel Islands off the California coast. Initial occupation was
concentrated in regions of dense, predictable resources. Subsequent occupation of habitats with
250 Bliege Bird
dispersed or more unpredictable resource bases resulted in increasing differentiation in the

population between the haves and the have-nots, as the development of dominance relations
excluded later arrivals from resource ownership (Winterhalder et al. 2010).
Environmental shifts in resource variance caused by increasing landscape heterogeneity can
also be expected to influence the gender division of labor by affecting resource acquisition variance.
Here variance is defined as the unpredictable risk of harvest failure and is observed in the unpre-
dictable range of harvest values. Men and women tend to respond to resource variance differently:
Both men and women include in their subsistence repertoires a suite of activities or resource
types that show a variety of variances, but they differ in the relative proportions of time spent on
high- and low-variance activities. Whereas men do forage for some low-variance resources, they
include many more high-variance resources than do women. The extent to which men include
such high-variance resources predicts the degree to which women are significant producers in the
subsistence economy (Bliege Bird et al. 2009, Codding et al. 2011): Where women produce more,
they may have more autonomy and engage more in cooperative intergenerational partnerships
with other women. If anthropogenic landscapes reduce interforager variance, increase encounter
rates with some prey and thus augment success predictability, and foster a greater reliance on low-
variance prey relative to high-variance prey, women’s economic production is likely to become
equal to men’s, if not greater.
CONCLUSIONS
Despite calls to incorporate niche construction as an alternative to models of human–environment
interaction that typically consider only the way humans respond to environmental conditions (be-
havioral ecology), the two approaches can be integrated by drawing on complex adaptive systems
theory where human–environment interactions at larger spatial and temporal scales are an emer-
gent property of a coevolved social-ecological system maintained by short-term benefits to indi-
vidual agents (Lansing 2003, Lansing & Fox 2011, Lansing et al. 1998). The putative weaknesses
of behavioral ecology in explaining complex phenomena such as subsistence intensification can be
overcome by an approach that integrates across spatial scales and considers the dynamic feedbacks
among subsistence, ecological structure, and social organization. Rather than seeing environmen-
tal modification as the result of an intentional attempt to increase environmental productivity, as
niche construction models suggest, this approach views the emergent properties of disturbance
as a fundamental component of human subsistence, whose effects shape social norms of sharing
through the way disturbance changes the nature of resource patchiness, defensibility, and inter-
and intraindividual variance in returns (see Figure 1).
The approach I have outlined here shows how to incorporate issues of spatial and temporal
scale and links patterns of individual behavior, explained through decision theory, with large-
scale landscape and group outcomes that, in turn, feed back over time to influence individual
behavior. Integrating human behavioral ecology with landscape and community ecology demon-
strates how social norms may emerge from the fundamental properties of human disturbance.
This approach considers humans as integral components of the ecosystem upon which depend a
host of other networked species supported by the emergent properties of human–environment
interaction. This perspective does not require that ecosystem engineering be intentional and thus
dissolves the perennial debate over whether foragers are intentional conservationists or destruc-
tive predators. As Bird et al. (2015) conclude, disturbance processes may lie at the heart of many
complex social-environmental phenomena, such as subsistence intensifications and diet breadth
shifts in prehistory.
Anthropogenic
disturbance to
landscapes
Increased fine-scale
heterogeneity
Increased patchiness,
predictability, and density
of generalist prey and plant
resources near central places
Reduced Higher returns Social

travel on predictable intensification,
cost resources ownership norms
Reduced Increased
residential women’s
mobility production?
Higher
reproductive
rates
Figure 1
Conceptual model for the positive human–environment feedbacks described in the text.
DISCLOSURE STATEMENT
The author is not aware of any affiliations, memberships, funding, or financial holdings that might
be perceived as affecting the objectivity of this review.
ACKNOWLEDGMENTS
I am grateful for the support I received from friends and family in the Parnngurr community
while conducting the research described here. Douglas Bird and Brian Codding contributed to
the development of many of the ideas in this article. The Martu research project has been funded
by grants from the National Science Foundation (BCS-0850664, BCS-0314406, SBR-0211265,
BCS-0127681, BCS-0075289), The Leakey Foundation, The Christensen Fund, The University
of Maine, the Stanford University Office of Teaching and Learning, the Lang Fund for
252 Bliege Bird
Environmental Anthropology, the Stanford Institute for Research in the Social Sciences, and the
Woods Institute Environmental Venture Program.
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AN44CH15-BliegeBird ARI 31 July 2015 8:20

Review in Advance first posted online

still occur before final publication

Disturbance, Complexity, Scale:

Rebecca Bliege Bird

Annu. Rev. Anthropol. 2015. 44:241–57 Keywords

242 Bliege Bird

A THEORY OF ECOLOGICAL INTERACTIONS

www.annualreviews.org • The Study of Human–Environment Interactions 243

244 Bliege Bird

CASE 1: GARDEN GAME IN THE NEOTROPICAL RAIN FOREST

www.annualreviews.org • The Study of Human–Environment Interactions 245

CASE 2: LIZARDS AND KANGAROOS IN THE AUSTRALIAN

246 Bliege Bird

www.annualreviews.org • The Study of Human–Environment Interactions 247

(dinnertime camp, in Martu parlance) is lower in anthropogenic environments, mainly because

FEEDBACKS INTO SOCIAL INSTITUTIONS

248 Bliege Bird

Sharing and Social Structure

www.annualreviews.org • The Study of Human–Environment Interactions 249

Social Inequality and Gender-Based Divisions of Labor

250 Bliege Bird

dispersed or more unpredictable resource bases resulted in increasing differentiation in the

www.annualreviews.org • The Study of Human–Environment Interactions 251

Reduced Higher returns Social

252 Bliege Bird

fate when its disperser is absent. J. Trop. Ecol. 15(2):229–35

www.annualreviews.org • The Study of Human–Environment Interactions 253

254 Bliege Bird

gain resources. Proc. R. Soc. B 280:20131210

www.annualreviews.org • The Study of Human–Environment Interactions 255

J. R. Soc. Interface 4(16):935–48

ics. Am. Anthropol. 89(3):650–67

256 Bliege Bird

tion, toward the historical ecology of rainforests. Hum. Ecol. 41(3):465–75

www.annualreviews.org • The Study of Human–Environment Interactions 257

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