Women's subsistence strategies predict fertility across cultures, but context matters.

While it is commonly assumed that farmers have higher, and foragers lower, fertility compared to populations practicing other forms of subsistence, robust supportive evidence is lacking. We tested whether subsistence activities-incorporating market integration-are associated with fertility in 10,250 women from 27 small-scale societies and found considerable variation in fertility. This variation did not align with group-level subsistence typologies. Societies labeled as "farmers" did not have higher fertility than others, while "foragers" did not have lower fertility. However, at the individual level, we found strong evidence that fertility was positively associated with farming and moderate evidence of a negative relationship between foraging and fertility. Markers of market integration were strongly negatively correlated with fertility. Despite strong cross-cultural evidence, these relationships were not consistent in all populations, highlighting the importance of the socioecological context, which likely influences the diverse mechanisms driving the relationship between fertility and subsistence.

Reproductive inequality in humans and other mammals.

To address claims of human exceptionalism, we determine where humans fit within the greater mammalian distribution of reproductive inequality. We show that humans exhibit lower reproductive skew (i.e., inequality in the number of surviving offspring) among males and smaller sex differences in reproductive skew than most other mammals, while nevertheless falling within the mammalian range. Additionally, female reproductive skew is higher in polygynous human populations than in polygynous nonhumans mammals on average. This patterning of skew can be attributed in part to the prevalence of monogamy in humans compared to the predominance of polygyny in nonhuman mammals, to the limited degree of polygyny in the human societies that practice it, and to the importance of unequally held rival resources to women's fitness. The muted reproductive inequality observed in humans appears to be linked to several unusual characteristics of our species-including high levels of cooperation among males, high dependence on unequally held rival resources, complementarities between maternal and paternal investment, as well as social and legal institutions that enforce monogamous norms.

Nahua mushroom gatherers use area-restricted search strategies that conform to marginal value theorem predictions.

We develop a method of analysis for testing the marginal value theorem (MVT) in natural settings that does not require an independent definition or mapping of patches. We draw on recent theoretical work on area-restricted search (ARS) that links turning-angle and step-size changes to geographically localized encounter-rates. These models allow us to estimate "giving-up times" using encounter-annotated GPS tracking data. Applied to a case study of Nahua mushroom foragers, these models identify distinct forms of intrapatch and interpatch search behavior, with intrapatch search transitioning to interpatch search after a predictable interval of time since the last encounter with a harvested mushroom. Our empirical estimate of giving-up time coincides with the theoretically optimal giving-up time derived under the MVT in the same environment. The MVT is currently underused in studies of human foraging and settlement patterns, due in large part to the difficulty of identifying discrete resource patches and quantifying their characteristics. Our methods mitigate the need to make such discrete maps of patches and thus have the potential to broaden the scope for empirical evaluations of the MVT and related theory in humans. Beyond studies of naturalistic foraging in humans and other animals, our approach has implications for optimization of search behavior in a range of applied fields where search dynamics must be adapted to shifting patterns of environmental heterogeneity affecting prey density and patchiness.

A general model of forager search: Adaptive encounter-conditional heuristics outperform Lévy flights in the search for patchily distributed prey.

A theoretical and applied literature has suggested that foragers search using Lévy flights, since Lévy flights can maximize the efficiency of search in the absence of information on the location of randomly distributed prey. Foragers, however, often have available to them at least some information about the distribution of prey, gained either through evolved mechanisms, experience and memory, or social transmission of information. As such, we might expect selection for heuristics that make use of such information to further improve the efficiency of random search. Here we present a general model of random search behavior that includes as special cases: area-restricted search, correlated random walks, Brownian search, and Lévy flights. This generative model allows foragers to adjust search parameters based on encounter-conditional and other heuristics. Using a simulation model, we demonstrate the efficiency gains of these search heuristics, and illustrate the resulting differences in the distributions of step-size and heading angle change they imply, relative to Lévy flights. We conclude by presenting a statistical model that can be fit to empirical data and a set of testable, quantitative predictions that contrast our model of adaptive search with the Lévy flight foraging hypothesis.

A hierarchical bayesian analysis of parasite prevalence and sociocultural outcomes: The role of structural racism and sanitation infrastructure.

OBJECTIVES: We conduct a revaluation of the Thornhill and Fincher research project on parasites using finely-resolved geographic data on parasite prevalence, individual-level sociocultural data, and multilevel Bayesian modeling. In contrast to the evolutionary psychological mechanisms linking parasites to human behavior and cultural characteristics proposed by Thornhill and Fincher, we offer an alternative hypothesis that structural racism and differential access to sanitation systems drive both variation in parasite prevalence and differential behaviors and cultural characteristics. METHODS: We adopt a Bayesian framework to estimate parasite prevalence rates in 51 districts in eight Latin American countries using the disease status of 170,220 individuals tested for infection with the intestinal roundworm Ascaris lumbricoides (Hürlimann et al., []: PLoS Negl Trop Dis 5:e1404). We then use district-level estimates of parasite prevalence and individual-level social data from 5,558 individuals in the same 51 districts (Latinobarómetro, 2008) to assess claims of causal associations between parasite prevalence and sociocultural characteristics. RESULTS: We find, contrary to Thornhill and Fincher, that parasite prevalence is positively associated with preferences for democracy, negatively associated with preferences for collectivism, and not associated with violent crime rates or gender inequality. A positive association between parasite prevalence and religiosity, as in Fincher and Thornhill (: Behav Brain Sci 35:61-79), and a negative association between parasite prevalence and achieved education, as predicted by Eppig et al. (: Proc R S B: Biol Sci 277:3801-3808), become negative and unreliable when reasonable controls are included in the model. We find support for all predictions derived from our hypothesis linking structural racism to both parasite prevalence and cultural outcomes. CONCLUSIONS: We conclude that best practices in biocultural modeling require examining more than one hypothesis, retaining individual-level data and its associated variance whenever possible, and adopting multilevel techniques suited to the structuring of the data.

Sit-and-wait versus active-search hunting: A behavioral ecological model of optimal search mode.

Drawing on Skellam׳s (1958) work on sampling animal populations using transects, we derive a behavioral ecological model of the choice between sit-and-wait and active-search hunting. Using simple, biologically based assumptions about the characteristics of predator and prey, we show how an empirically definable parameter space favoring active-search hunting expands as: (1) the average rate of movement of prey decreases, or (2) the energetic costs of hunter locomotion decline. The same parameter space narrows as: (3) prey skittishness increases as a function of a hunter׳s velocity, or (4) prey become less detectable as a function of a hunter׳s velocity. Under either search tactic, encounter rate increases as a function of increasing prey velocity and increasing detection zone radius. Additionally, we investigate the roles of habitat heterogeneity and spatial auto-correlation or grouping of prey on the optimal search mode of a hunter, finding that habitat heterogeneity has the potential to complicate application of the model to some empirical examples, while the effects of prey grouping lead to relatively similar model outcomes. As predicted by the model, the introduction of the horse to the Great Plains and the introduction of the snowmobile to Arctic foraging communities decreased the metabolic costs of active-search and led to a change in normative hunting strategies that favored active-search in place of sit-and-wait hunting.

Ethnography and ethnohistory support the efficiency of hunting through endurance running in humans.

Humans have two features rare in mammals: our locomotor muscles are dominated by fatigue-resistant fibres and we effectively dissipate through sweating the metabolic heat generated through prolonged, elevated activity. A promising evolutionary explanation of these features is the endurance pursuit (EP) hypothesis, which argues that both traits evolved to facilitate running down game by persistence. However, this hypothesis has faced two challenges: running is energetically costly and accounts of EPs among late twentieth century foragers are rare. While both observations appear to suggest that EPs would be ineffective, we use foraging theory to demonstrate that EPs can be quite efficient. We likewise analyse an ethnohistoric and ethnographic database of nearly 400 EP cases representing 272 globally distributed locations. We provide estimates for return rates of EPs and argue that these are comparable to other pre-modern hunting methods in specified contexts. EP hunting as a method of food procurement would have probably been available and attractive to Plio/Pleistocene hominins.

The life history of human foraging: Cross-cultural and individual variation.

Human adaptation depends on the integration of slow life history, complex production skills, and extensive sociality. Refining and testing models of the evolution of human life history and cultural learning benefit from increasingly accurate measurement of knowledge, skills, and rates of production with age. We pursue this goal by inferring hunters' increases and declines of skill from approximately 23,000 hunting records generated by more than 1800 individuals at 40 locations. The data reveal an average age of peak productivity between 30 and 35 years of age, although high skill is maintained throughout much of adulthood. In addition, there is substantial variation both among individuals and sites. Within study sites, variation among individuals depends more on heterogeneity in rates of decline than in rates of increase. This analysis sharpens questions about the coevolution of human life history and cultural adaptation.

Evidence for encounter-conditional, area-restricted search in a preliminary study of Colombian blowgun hunters.

Active search for prey is energetically costly, so understanding how foragers optimize search has been central to foraging theory. Some theoretical work has suggested that foragers of randomly distributed prey should search using Lévy flights, while work on area-restricted and intermittent search strategies has demonstrated that foragers can use the information provided by prey encounters to more effectively adapt search direction and velocity. Previous empirical comparisons of these search modes have tended to rely on distribution-level analyses, due to the difficulty of collecting event-level data on encounters linked to the GPS tracks of foragers. Here we use a preliminary event-level data-set (18.7 hours of encounter-annotated focal follows over 6 trips) to show that two Colombian blowgun hunters use adaptive encounter-conditional heuristics, not non-conditional Lévy flights, when searching for prey. Using a theoretically derived Bayesian model, we estimate changes in turning-angle and search velocity as a function of encounters with prey at lagged time-steps, and find that: 1) hunters increase average turning-angle in response to encounters, producing a more tortuous search of patches of higher prey density, but adopt more efficient uni-directional, inter-patch movement after failing to encounter prey over a sufficient period of time; and, 2) hunters reduce search velocity in response to encounters, causing them to spend more of their search time in patches with demonstrably higher prey density. These results illustrate the importance of using event-level data to contrast encounter-conditional, area-restricted search and Lévy flights in explaining the search behavior of humans and other organisms.

Greater wealth inequality, less polygyny: rethinking the polygyny threshold model.

Monogamy appears to have become the predominant human mating system with the emergence of highly unequal agricultural populations that replaced relatively egalitarian horticultural populations, challenging the conventional idea-based on the polygyny threshold model-that polygyny should be positively associated with wealth inequality. To address this polygyny paradox, we generalize the standard polygyny threshold model to a mutual mate choice model predicting the fraction of women married polygynously. We then demonstrate two conditions that are jointly sufficient to make monogamy the predominant marriage form, even in highly unequal societies. We assess if these conditions are satisfied using individual-level data from 29 human populations. Our analysis shows that with the shift to stratified agricultural economies: (i) the population frequency of relatively poor individuals increased, increasing wealth inequality, but decreasing the frequency of individuals with sufficient wealth to secure polygynous marriage, and (ii) diminishing marginal fitness returns to additional wives prevent extremely wealthy men from obtaining as many wives as their relative wealth would otherwise predict. These conditions jointly lead to a high population-level frequency of monogamy.

Frequency-Dependent Social Transmission and the Interethnic Transfer of Female Genital Modification in the African Diaspora and Indigenous Populations of Colombia.

We present a quantitative account based on ethnographic and documentary research of the prevalence of female genital modification (FGMo) in the African diaspora and indigenous populations of Colombia. We use these data to test hypotheses concerning the cultural evolutionary drivers of costly trait persistence, attenuation, and intergroup transmission. The uptake of FGMo by indigenous populations in Colombia is consistent with frequency-dependent hypotheses for the social transmission of the FGMo trait from the African diaspora population in the period following the era of slavery in Colombia. The prevalence and severity of practices related to FGMo decline with level of sociocultural integration into mainstream Colombian culture. Our results provide empirical support for the cultural evolutionary models proposed by Ross et al. (2015) to describe the transmission dynamics of FGMo and other costly traits. Analysis of costly trait dynamics contributes knowledge useful to applied anthropology and may be of interest in policy design and human rights monitoring in Colombia and elsewhere.

Epidemics in Ming and Qing China: Impacts of changes of climate and economic well-being.

We investigated the mechanism of epidemics with the impacts of climate change and socio-economic fluctuations in the Ming and Qing Dynasties in China (AD 1368-1901). Using long-term and high-quality datasets, this study is the first quantitative research that verifies the 'climate change â†’ economy â†’ epidemics' mechanism in historical China by statistical methods that include correlation analysis, Granger causality analysis, ARX, and Poisson-ARX modeling. The analysis provides the evidences that climate change could only fundamentally lead to the epidemics spread and occurrence, but the depressed economic well-being is the direct trigger of epidemics spread and occurrence at the national and long term scale in historical China. Moreover, statistical modeling shows that economic well-being is more important than population pressure in the mechanism of epidemics. However, population pressure remains a key element in determining the social vulnerability of the epidemics occurrence under climate change. Notably, the findings not only support adaptation theories but also enhance our confidence to address climatic shocks if economic buffering capacity can be promoted steadily. The findings can be a basis for scientists and policymakers in addressing global and regional environmental changes.

The population ecology of despotism. Concessions and migration between central and peripheral habitats.

Since despotism is a common evolutionary development in human history, we seek to understand the conditions under which it can originate, persist, and affect population trajectories. We describe a general system of population ecology equations representing the Ideal Free and Despotic Distributions for one and two habitats, one of which contains a despotic class that controls the distribution of resources. Using analytical and numerical solutions we derive the optimal concession strategy by despots with and without subordinate migration to an alternative habitat. We show that low concessions exponentially increase the time it takes for the despotic habitat to fill, and we discuss the trade-offs despots and subordinates confront at various levels of exploitation. Contrary to previous hypotheses, higher levels of despotism do not necessarily cause faster migration to alternative habitats. We further show how, during colonization, divergent population trajectories may arise if despotic systems experience Allee-type economies of scale.

The invisible cliff: abrupt imposition of Malthusian equilibrium in a natural-fertility, agrarian society.

Analysis of a natural fertility agrarian society with a multi-variate model of population ecology isolates three distinct phases of population growth following settlement of a new habitat: (1) a sometimes lengthy copial phase of surplus food production and constant vital rates; (2) a brief transition phase in which food shortages rapidly cause increased mortality and lessened fertility; and (3) a Malthusian phase of indefinite length in which vital rates and quality of life are depressed, sometimes strikingly so. Copial phase duration declines with increases in the size of the founding group, maximum life expectancy and fertility; it increases with habitat area and yield per hectare; and, it is unaffected by the sensitivity of vital rates to hunger. Transition phase duration is unaffected by size of founding population and area of settlement; it declines with yield, life expectancy, fertility and the sensitivity of vital rates to hunger. We characterize the transition phase as the Malthusian transition interval (MTI), in order to highlight how little time populations generally have to adjust. Under food-limited density dependence, the copial phase passes quickly to an equilibrium of grim Malthusian constraints, in the manner of a runner dashing over an invisible cliff. The three-phase pattern diverges from widely held intuitions based on standard Lotka-Verhulst approaches to population regulation, with implications for the analysis of socio-cultural evolution, agricultural intensification, bioarchaeological interpretation of food stress in prehistoric societies, and state-level collapse.

Development and disintegration of Maya political systems in response to climate change.

The role of climate change in the development and demise of Classic Maya civilization (300 to 1000 C.E.) remains controversial because of the absence of well-dated climate and archaeological sequences. We present a precisely dated subannual climate record for the past 2000 years from Yok Balum Cave, Belize. From comparison of this record with historical events compiled from well-dated stone monuments, we propose that anomalously high rainfall favored unprecedented population expansion and the proliferation of political centers between 440 and 660 C.E. This was followed by a drying trend between 660 and 1000 C.E. that triggered the balkanization of polities, increased warfare, and the asynchronous disintegration of polities, followed by population collapse in the context of an extended drought between 1020 and 1100 C.E.

Four neglected concepts with a role to play in explaining the origins of agriculture.

Explanations of the socioeconomic changes accompanying the transition from foraging through mixed economies to societies that take up full-time agriculture will entail concepts of risk, discounting, economies of scale, and transaction costs. The spatial form and temporal scale of agricultural production fundamentally change the parameters of risk management. Delays between investment decisions and consumption of yields grow in duration and significance, elevating the salience of discounting. Localization and control over property and productivity generate opportunities for specialization and economies of scale, at the same time setting up the possibility of exchange among specialists and between locales with differential production advantages and consumption needs. Full evolutionary analysis of the origins of agriculture entails these ideas; their use in anthropology will require that we temper our history of economic substantivism with recognition that certain concepts forged to understand market economies are applicable much more broadly.

Demographic consequences of unpredictability in fertility outcomes.

Child survival is probabilistic, but the unpredictability in family formation and completed family size has been neglected in the fertility literature. In many societies, ending the family cycle with too few or too many surviving offspring entails serious social, economic, or fitness consequences. A model of risk- (or variance-) sensitive adaptive behavior that addresses long-term fertility outcomes is presented. The model shows that under conditions likely to be common, optimal, risk-sensitive reproductive strategies deviate systematically from the completed family size that would be expected if reproductive outcome is were predictable. This is termed the "variance compensation hypothesis." Variance compensation may be either positive or negative, resulting in augmented or diminished fertility. Which outcome obtained is a function of identifiable social, economic, and environmental factors. Through its effect on fertility behavior, variance compensation has a direct bearing on birth spacing and completed fertility, and thereby on problems in demography and human population biology ranging from demographic transitions to maternal depletion and child health. Risk-sensitive models will be a necessary component of a general theory of fertility.