Pathways to social inequality.

Social inequality is ubiquitous in contemporary human societies, and has deleterious social and ecological impacts. However, the factors that shape the emergence and maintenance of inequality remain widely debated. Here we conduct a global analysis of pathways to inequality by comparing 408 non-industrial societies in the anthropological record (described largely between 1860 and 1960) that vary in degree of inequality. We apply structural equation modelling to open-access environmental and ethnographic data and explore two alternative models varying in the links among factors proposed by prior literature, including environmental conditions, resource intensification, wealth transmission, population size and a well-documented form of inequality: social class hierarchies. We found support for a model in which the probability of social class hierarchies is associated directly with increases in population size, the propensity to use intensive agriculture and domesticated large mammals, unigeniture inheritance of real property and hereditary political succession. We suggest that influence of environmental variables on inequality is mediated by measures of resource intensification, which, in turn, may influence inequality directly or indirectly via effects on wealth transmission variables. Overall, we conclude that in our analysis a complex network of effects are associated with social class hierarchies.

Cultural transmission and ecological opportunity jointly shaped global patterns of reliance on agriculture.

The evolution of agriculture improved food security and enabled significant increases in the size and complexity of human groups. Despite these positive effects, some societies never adopted these practices, became only partially reliant on them, or even reverted to foraging after temporarily adopting them. Given the critical importance of climate and biotic interactions for modern agriculture, it seems likely that ecological conditions could have played a major role in determining the degree to which different societies adopted farming. However, this seemingly simple proposition has been surprisingly difficult to prove and is currently controversial. Here, we investigate how recent agricultural practices relate both to contemporary ecological opportunities and the suitability of local environments for the first species domesticated by humans. Leveraging a globally distributed dataset on 1,291 traditional societies, we show that after accounting for the effects of cultural transmission and more current ecological opportunities, levels of reliance on farming continue to be predicted by the opportunities local ecologies provided to the first human domesticates even after centuries of cultural evolution. Based on the details of our models, we conclude that ecology probably helped shape the geography of agriculture by biasing both human movement and the human-assisted dispersal of domesticates.

Hindcasting global population densities reveals forces enabling the origin of agriculture.

The development and spread of agriculture changed fundamental characteristics of human societies1-3. However, the degree to which environmental and social conditions enabled the origins of agriculture remains contested4-6. We test three hypothesized links between the environment, population density and the origins of plant and animal domestication, a prerequisite for agriculture: (1) domestication arose as environmental conditions improved and population densities increased7 (surplus hypothesis); (2) populations needed domestication to overcome deteriorating environmental conditions (necessity hypothesis)8,9; (3) factors promoting domestication were distinct in each location10 (regional uniqueness hypothesis). We overcome previous data limitations with a statistical model, in which environmental, geographic and cultural variables capture 77% of the variation in population density among 220 foraging societies worldwide. We use this model to hindcast potential population densities across the globe from 21,000 to 4,000 years before present. Despite the timing of domestication varying by thousands of years, we show that improving environmental conditions favoured higher local population densities during periods when domestication arose in every known agricultural origin centre. Our results uncover a common, global factor that facilitated one of humanity's most significant innovations and demonstrate that modelling ancestral demographic changes can illuminate major events deep in human history.

Genetic and demographic founder effects have long-term fitness consequences for colonising populations.

Colonisation is a fundamental ecological and evolutionary process that drives the distribution and abundance of organisms. The initial ability of colonists to establish is determined largely by the number of founders and their genetic background. We explore the importance of these demographic and genetic properties for longer term persistence and adaptation of populations colonising a novel habitat using experimental populations of Tribolium castaneum. We introduced individuals from three genetic backgrounds (inbred - outbred) into a novel environment at three founding sizes (2-32), and tracked populations for seven generations. Inbreeding had negative effects, whereas outbreeding generally had positive effects on establishment, population growth and long-term persistence. Severe bottlenecks due to small founding sizes reduced genetic variation and fitness but did not prevent adaptation if the founders originated from genetically diverse populations. Thus, we find important and largely independent roles for both demographic and genetic processes in driving colonisation success.

Three types of rescue can avert extinction in a changing environment.

Setting aside high-quality large areas of habitat to protect threatened populations is becoming increasingly difficult as humans fragment and degrade the environment. Biologists and managers therefore must determine the best way to shepherd small populations through the dual challenges of reductions in both the number of individuals and genetic variability. By bringing in additional individuals, threatened populations can be increased in size (demographic rescue) or provided with variation to facilitate adaptation and reduce inbreeding (genetic rescue). The relative strengths of demographic and genetic rescue for reducing extinction and increasing growth of threatened populations are untested, and which type of rescue is effective may vary with population size. Using the flour beetle (Tribolium castaneum) in a microcosm experiment, we disentangled the genetic and demographic components of rescue, and compared them with adaptation from standing genetic variation (evolutionary rescue in the strictest sense) using 244 experimental populations founded at either a smaller (50 individuals) or larger (150 individuals) size. Both types of rescue reduced extinction, and those effects were additive. Over the course of six generations, genetic rescue increased population sizes and intrinsic fitness substantially. Both large and small populations showed evidence of being able to adapt from standing genetic variation. Our results support the practice of genetic rescue in facilitating adaptation and reducing inbreeding depression, and suggest that demographic rescue alone may suffice in larger populations even if only moderately inbred individuals are available for addition.

The roles of demography and genetics in the early stages of colonization.

Colonization success increases with the size of the founding group. Both demographic and genetic factors underlie this relationship, yet because genetic diversity normally increases with numbers of individuals, their relative importance remains unclear. Furthermore, their influence may depend on the environment and may change as colonization progresses from establishment through population growth and then dispersal. We tested the roles of genetics, demography and environment in the founding of Tribolium castaneum populations. Using three genetic backgrounds (inbred to outbred), we released individuals of four founding sizes (2-32) into two environments (natal and novel), and measured establishment success, initial population growth and dispersal. Establishment increased with founding size, whereas population growth was shaped by founding size, genetic background and environment. Population growth was depressed by inbreeding at small founding sizes, but growth rates were similar across genetic backgrounds at large founding size, an interaction indicating that the magnitude of the genetic effects depends upon founding population size. Dispersal rates increased with genetic diversity. These results suggest that numbers of individuals may drive initial establishment, but that subsequent population growth and spread, even in the first generation of colonization, can be driven by genetic processes, including both reduced growth owing to inbreeding depression, and increased dispersal with increased genetic diversity.