Cooperating to show that you care: costly helping as an honest signal of fitness interdependence.

Social organisms often need to know how much to trust others to cooperate. Organisms can expect cooperation from another organism that depends on them (i.e. stake or fitness interdependence), but how do individuals assess fitness interdependence? Here, we extend fitness interdependence into a signalling context: costly helping behaviour can honestly signal one's stake in others, such that those who help are trusted more. We present a mathematical model in which agents help others based on their stake in the recipient's welfare, and recipients use that information to assess whom to trust. At equilibrium, helping is a costly signal of stake: helping is worthwhile for those who value the recipient (and thus will repay any trust), but is not worthwhile for those who do not value the recipient (and thus will betray the trust). Recipients demand signals when they value the signallers less and when the cost of betrayed trust is higher; signal costs are higher when signallers have more incentive to defect. Signalling systems are more likely when the trust games resemble Prisoner's Dilemmas, Stag Hunts or Harmony Games, and are less likely in Snowdrift Games. Furthermore, we find that honest signals need not benefit recipients and can even occur between hostile parties. By signalling their interdependence, organisms benefit from increased trust, even when no future interactions will occur. This article is part of the theme issue 'The language of cooperation: reputation and honest signalling'.

People have shaped most of terrestrial nature for at least 12,000 years.

Archaeological and paleoecological evidence shows that by 10,000 BCE, all human societies employed varying degrees of ecologically transformative land use practices, including burning, hunting, species propagation, domestication, cultivation, and others that have left long-term legacies across the terrestrial biosphere. Yet, a lingering paradigm among natural scientists, conservationists, and policymakers is that human transformation of terrestrial nature is mostly recent and inherently destructive. Here, we use the most up-to-date, spatially explicit global reconstruction of historical human populations and land use to show that this paradigm is likely wrong. Even 12,000 y ago, nearly three quarters of Earth's land was inhabited and therefore shaped by human societies, including more than 95% of temperate and 90% of tropical woodlands. Lands now characterized as "natural," "intact," and "wild" generally exhibit long histories of use, as do protected areas and Indigenous lands, and current global patterns of vertebrate species richness and key biodiversity areas are more strongly associated with past patterns of land use than with present ones in regional landscapes now characterized as natural. The current biodiversity crisis can seldom be explained by the loss of uninhabited wildlands, resulting instead from the appropriation, colonization, and intensifying use of the biodiverse cultural landscapes long shaped and sustained by prior societies. Recognizing this deep cultural connection with biodiversity will therefore be essential to resolve the crisis.

Climate, landscape diversity, and food sovereignty in arid Australia: The firestick farming hypothesis.

OBJECTIVE: Climate change has long been recognized as a significant driver of dietary diversity and dietary quality. An often overlooked aspect of climate change are shifts in fire regimes, which have the potential to drastically affect landscape diversity, species distributions, and ultimately, human diets. Here, we investigate whether the fire regimes shaped by Indigenous Australians change landscape diversity in ways that improve dietary quality, considering both the diversity and the quantity of traditional foods in the diet. METHODS: We use structural equation modeling to explore two causal models of dietary quality, one focused on the direct effects of climate change and resource depression, the other incorporating the dietary effects of landscape diversity, itself a product of fire-created patchiness. We draw on a focal camp dataset covering 10 years of observations of Martu foraging income in the Western Desert of Australia. RESULTS: We find strong support for the hypothesis that fire-created patchiness improves diet quality. Climate change (cumulative 2-year rainfall) has only an indirect effect on dietary quality; the availability of traditional foods is mediated primarily through the landscape diversity shaped by fire. CONCLUSIONS: Our model suggests that the loss of the indigenous fire mosaic may lead to worsening availability of traditional foods, measured as both caloric intake and diet diversity. Because the effects of rainfall are mediated through landscape diversity, increased rainfall may not compensate for the recent changes in fire regimes resulting from the loss of Aboriginal fire from the landscape.

Fire mosaics and habitat choice in nomadic foragers.

In the mid-1950s Western Desert of Australia, Aboriginal populations were in decline as families left for ration depots, cattle stations, and mission settlements. In the context of reduced population density, an ideal free-distribution model predicts landscape use should contract to the most productive habitats, and people should avoid areas that show more signs of extensive prior use. However, ecological or social facilitation due to Allee effects (positive density dependence) would predict that the intensity of past habitat use should correlate positively with habitat use. We analyzed fire footprints and fire mosaics from the accumulation of several years of landscape use visible on a 35,300-km2 mosaic of aerial photographs covering much of contemporary Indigenous Martu Native Title Lands imaged between May and August 1953. Structural equation modeling revealed that, consistent with an Allee ideal free distribution, there was a positive relationship between the extent of fire mosaics and the intensity of recent use, and this was consistent across habitats regardless of their quality. Fire mosaics build up in regions with low cost of access to water, high intrinsic food availability, and good access to trade opportunities; these mosaics (constrained by water access during the winter) then draw people back in subsequent years or seasons, largely independent of intrinsic habitat quality. Our results suggest that the positive feedback effects of landscape burning can substantially change the way people value landscapes, affecting mobility and settlement by increasing sedentism and local population density.

Animal movements in fire-prone landscapes.

Movement is a trait of fundamental importance in ecosystems subject to frequent disturbances, such as fire-prone ecosystems. Despite this, the role of movement in facilitating responses to fire has received little attention. Herein, we consider how animal movement interacts with fire history to shape species distributions. We consider how fire affects movement between habitat patches of differing fire histories that occur across a range of spatial and temporal scales, from daily foraging bouts to infrequent dispersal events, and annual migrations. We review animal movements in response to the immediate and abrupt impacts of fire, and the longer-term successional changes that fires set in train. We discuss how the novel threats of altered fire regimes, landscape fragmentation, and invasive species result in suboptimal movements that drive populations downwards. We then outline the types of data needed to study animal movements in relation to fire and novel threats, to hasten the integration of movement ecology and fire ecology. We conclude by outlining a research agenda for the integration of movement ecology and fire ecology by identifying key research questions that emerge from our synthesis of animal movements in fire-prone ecosystems.

Emergent sustainability in open property regimes.

Current theoretical models of the commons assert that common-pool resources can only be managed sustainably with clearly defined boundaries around both communities and the resources that they use. In these theoretical models, open access inevitably leads to a tragedy of the commons. However, in many open-access systems, use of common-pool resources seems to be sustainable over the long term (i.e., current resource use does not threaten use of common-pool resources for future generations). Here, we outline the conditions that support sustainable resource use in open property regimes. We use the conceptual framework of complex adaptive systems to explain how processes within and couplings between human and natural systems can lead to the emergence of efficient, equitable, and sustainable resource use. We illustrate these dynamics in eight case studies of different social-ecological systems, including mobile pastoralism, marine and freshwater fisheries, swidden agriculture, and desert foraging. Our theoretical framework identifies eight conditions that are critical for the emergence of sustainable use of common-pool resources in open property regimes. In addition, we explain how changes in boundary conditions may push open property regimes to either common property regimes or a tragedy of the commons. Our theoretical model of emergent sustainability helps us to understand the diversity and dynamics of property regimes across a wide range of social-ecological systems and explains the enigma of open access without a tragedy. We recommend that policy interventions in such self-organizing systems should focus on managing the conditions that are critical for the emergence and persistence of sustainability.

The social significance of subtle signals.

Acts of prosociality, such as donating to charity, are often analysed in a similar way to acts of conspicuous advertising; both involve costly signals revealing hidden qualities that increase the signaller's prestige. However, experimental work suggests that grand gestures, even if prosocial, may damage one's reputation for trustworthiness and cooperativeness if they are perceived as prestige enhancing: individuals may gain some types of cooperative benefits only when they perform prosocial acts in particular ways. Here, we contrast subtle, less obviously costly, interpersonal forms of prosocial behaviour with high-cost displays to a large audience, drawing on the example of food sharing in subsistence economies. This contrast highlights how highly visible prosocial displays may be effective for attracting new partners, while subtle signals may be crucial for ensuring trust and commitment with long-term partners. Subtle dyadic signals may be key to understanding the long-term maintenance of interpersonal networks that function to reduce unanticipated risks.

Pyrodiversity and the anthropocene: the role of fire in the broad spectrum revolution.

The Anthropocene colloquially refers to a global regime of human-caused environmental modification of earth systems associated with profound changes in patterns of human mobility, as well as settlement and resource use compared with prior eras. Some have argued that the processes generating the Anthropocene are mainly associated with population growth and technological innovation, and thus began only in the late Holocene under conditions of dense sedentism and industrial agriculture.(1) However, it now seems clear that the roots of the Anthropocene lie in complex processes of intensification that significantly predate transitions to agriculture.(2,3) What intensification is remains less clear. For some it is increasing economic productivity that increases carrying capacity, the drivers of which may be too diverse and too local to generalize.(4,5) For others using Boserup's ideas about agrarian intensification, increasing density in hunter-gatherer populations can produce declines in subsistence efficiency that increase incentives for investing labor to boost yield per unit area, which then elevates Malthusian limits on carrying capacity.(6-8) As Morgan(9) demonstrates in a comprehensive review, the legacy of such Boserupian intensification is alive, well, and controversial in hunter-gatherer archeology. This is a result of its potential for illuminating processes involved in transformations of forager socio-political and economic systems, including those dominated by harvesting more immediate-return resources and high residential mobility as well as those characterized by more delayed-return material economies with reduced residential mobility, a broader spectrum of resources, degrees of storage, and greater social stratification. Here we detail hypotheses about the processes involved in such transitions and explore the way that anthropogenic disturbance of ecosystems, especially the use of landscape fire, could be fundamentally entangled with many broad-spectrum revolutions associated with intensified foraging systems.

People, El Niño southern oscillation and fire in Australia: fire regimes and climate controls in hummock grasslands.

While evidence mounts that indigenous burning has a significant role in shaping pyrodiversity, the processes explaining its variation across local and external biophysical systems remain limited. This is especially the case with studies of climate-fire interactions, which only recognize an effect of humans on the fire regime when they act independently of climate. In this paper, we test the hypothesis that an anthropogenic fire regime (fire incidence, size and extent) does not covary with climate. In the lightning regime, positive El Niño southern oscillation (ENSO) values increase lightning fire incidence, whereas La Niña (and associated increases in prior rainfall) increase fire size. ENSO has the opposite effect in the Martu regime, decreasing ignitions in El Niño conditions without affecting fire size. Anthropogenic ignition rates covary positively with high antecedent rainfall, whereas fire size varies only with high temperatures and unpredictable winds, which may reduce control over fire spread. However, total area burned is similarly predicted by antecedent rainfall in both regimes, but is driven by increases in fire size in the lightning regime, and fire number in the anthropogenic regime. We conclude that anthropogenic regimes covary with climatic variation, but detecting the human-climate-fire interaction requires multiple measures of both fire regime and climate.This article is part of the themed issue 'The interaction of fire and mankind'.

Niche construction and Dreaming logic: aboriginal patch mosaic burning and varanid lizards (Varanus gouldii) in Australia.

Anthropogenic fire is a form of ecosystem engineering that creates greater landscape patchiness at small spatial scales: such rescaling of patch diversity through mosaic burning has been argued to be a form of niche construction, the loss of which may have precipitated the decline and extinction of many endemic species in the Western Desert of Australia. We find evidence to support this hypothesis relative to one keystone species, the sand monitor lizard (Varanus gouldii). Paradoxically, V. gouldii populations are higher where Aboriginal hunting is most intense. This effect is driven by an increase in V. gouldii densities near successional edges, which is higher in landscapes that experience extensive human burning. Over time, the positive effects of patch mosaic burning while hunting overwhelm the negative effects of predation in recently burned areas to produce overall positive impacts on lizard populations. These results offer critical insights into the maintenance of animal communities in the desert, supporting the hypothesis that the current high rate of endemic species decline among small animals may be linked to the interaction between invasive species and mid-century removal of Aboriginal niche construction through hunting and patch mosaic burning.

Aboriginal hunting buffers climate-driven fire-size variability in Australia's spinifex grasslands.

Across diverse ecosystems, greater climatic variability tends to increase wildfire size, particularly in Australia, where alternating wet-dry cycles increase vegetation growth, only to leave a dry overgrown landscape highly susceptible to fire spread. Aboriginal Australian hunting fires have been hypothesized to buffer such variability, mitigating mortality on small-mammal populations, which have suffered declines and extinctions in the arid zone coincident with Aboriginal depopulation. We test the hypothesis that the relationship between climate and fire size is buffered through the maintenance of an anthropogenic, fine-grained fire regime by comparing the effect of climatic variability on landscapes dominated by Martu Aboriginal hunting fires with those dominated by lightning fires. We show that Aboriginal fires are smaller, more tightly clustered, and remain small even when climate variation causes huge fires in the lightning region. As these effects likely benefit threatened small-mammal species, Aboriginal hunters should be considered trophic facilitators, and policies aimed at reducing the risk of large fires should promote land-management strategies consistent with Aboriginal burning regimes.

What Explains Differences in Men's and Women's Production? : Determinants of Gendered Foraging Inequalities among Martu.

Researchers commonly use long-term average production inequalities to characterize cross-cultural patterns in foraging divisions of labor, but little is known about how the strategies of individuals shape such inequalities. Here, we explore the factors that lead to daily variation in how much men produce relative to women among Martu, contemporary foragers of the Western Desert of Australia. We analyze variation in foraging decisions on temporary foraging camps and find that the percentage of total camp production provided by each gender varies primarily as a function of men's average bout successes with large, mobile prey. When men target large prey, either their success leads to a large proportional contribution to the daily harvest, or their failure results in no contribution. When both men and women target small reliable prey, production inequalities by gender are minimized. These results suggest that production inequalities among Martu emerge from stochastic variation in men's foraging success on large prey measured against the backdrop of women's consistent production of small, low-variance resources.

Group Structure and Female Cooperative Networks in Australia's Western Desert.

The division of labor has typically been portrayed as a complementary strategy in which men and women work on separate tasks to achieve a common goal of provisioning the family. In this paper, we propose that task specialization between female kin might also play an important role in women's social and economic strategies. We use historic group composition data from a population of Western Desert Martu Aborigines to show how women maintained access to same-sex kin over the lifespan. Our results show that adult women had more same-sex kin and more closely related kin present than adult men, and they retained these links after marriage. Maternal co-residence was more prevalent for married women than for married men, and there is evidence that mothers may be strategizing to live with daughters at critical intervals-early in their reproductive careers and when they do not have other close female kin in the group. The maintenance of female kin networks across the lifespan allows for the possibility of cooperative breeding as well as an all-female division of labor.

Children on the reef : Slow learning or strategic foraging?

Meriam children are active reef-flat collectors. We demonstrate that while foraging on the reef, children are significantly less selective than adults. This difference and the precise nature of children's selectivity while reef-flat collecting are consistent with a hypothesis that both children and adults attempt to maximize their rate of return while foraging, but in so doing they face different constraints relative to differences in walking speeds while searching. Implications of these results for general arguments about factors that shape differences between child and adult behavior and human life-histories are explored.