To the hunter go the spoils? No evidence of nutritional benefit to being or marrying a well-reputed Hadza hunter.

OBJECTIVES: The incentives underlying men's hunting acquisition patterns among foragers are much debated. Some argue that hunters preferentially channel foods to their households, others maintain that foods are widely redistributed. Debates have focused on the redistribution of foods brought to camp, though the proper interpretation of results is contested. Here we instead address this question using two nutritional variables, employed as proxies for longer-term food access. We also report on broader patterns in nutritional status. MATERIALS AND METHODS: We measured male hunting success, hemoglobin concentration and body fatness among bush-living Hadza. Hunting success was measured using an aggregated reputation score. Hemoglobin concentration, a proxy for dietary red meat, was measured from fingerprick capillary blood. Body fatness, a proxy for energy balance, was measured using BMI and bioelectrical impedance. RESULTS: We find no statistically significant relationship between a hunter's success and any measure of his nutritional status or that of his spouse. We further find that: women are, as elsewhere, at greater risk of iron-deficiency anemia than men; men had slightly lower BMIs than women; men but not women had significantly lower hemoglobin levels than in the 1960s. DISCUSSION: The absence of an association between hunting reputation and nutritional status is consistent with generalized food sharing. Null results are difficult to interpret and findings could potentially be a consequence of insufficient signal in the study measures or some confounding effect. In any event, our results add to a substantial corpus of existing research that identifies few nutritional advantages to being or marrying a well-reputed Hadza hunter.

Physical activity patterns and biomarkers of cardiovascular disease risk in hunter-gatherers.

OBJECTIVES: Time spent in moderate-to-vigorous physical activity (MVPA) is a strong predictor of cardiovascular health, yet few humans living in industrialized societies meet current recommendations (150 min/week). Researchers have long suggested that human physiological requirements for aerobic exercise reflect an evolutionary shift to a hunting and gathering foraging strategy, and a recent transition to more sedentary lifestyles likely represents a mismatch with our past in terms of physical activity. The goal of this study is to explore this mismatch by characterizing MVPA and cardiovascular health in the Hadza, a modern hunting and gathering population living in Northern Tanzania. METHODS: We measured MVPA using continuous heart rate monitoring in 46 participants recruited from two Hadza camps. As part of a larger survey of health in the Hadza, we measured blood pressure (n = 198) and biomarkers of cardiovascular health (n = 23) including C-reactive protein, cholesterol (Total, HDL, and LDL), and triglycerides. RESULTS: We show that Hadza participants spend large amounts of time in MVPA (134.92 ± 8.6 min/day), and maintain these activity levels across the lifespan. In fact, the Hadza engage in over 14 times as much MVPA as subjects participating in large epidemiological studies in the United States. We found no evidence of risk factors for cardiovascular disease in this population (low prevalence of hypertension across the lifespan, optimal levels for biomarkers of cardiovascular health). CONCLUSIONS: Our results provide evidence that the hunting and gathering foraging strategy involves high levels of MVPA, supporting the evolutionary medicine model for the relationship between MVPA and cardiovascular health.

Hunter-Gatherers and the Origins of Religion.

Recent studies of the evolution of religion have revealed the cognitive underpinnings of belief in supernatural agents, the role of ritual in promoting cooperation, and the contribution of morally punishing high gods to the growth and stabilization of human society. The universality of religion across human society points to a deep evolutionary past. However, specific traits of nascent religiosity, and the sequence in which they emerged, have remained unknown. Here we reconstruct the evolution of religious beliefs and behaviors in early modern humans using a global sample of hunter-gatherers and seven traits describing hunter-gatherer religiosity: animism, belief in an afterlife, shamanism, ancestor worship, high gods, and worship of ancestors or high gods who are active in human affairs. We reconstruct ancestral character states using a time-calibrated supertree based on published phylogenetic trees and linguistic classification and then test for correlated evolution between the characters and for the direction of cultural change. Results indicate that the oldest trait of religion, present in the most recent common ancestor of present-day hunter-gatherers, was animism, in agreement with long-standing beliefs about the fundamental role of this trait. Belief in an afterlife emerged, followed by shamanism and ancestor worship. Ancestor spirits or high gods who are active in human affairs were absent in early humans, suggesting a deep history for the egalitarian nature of hunter-gatherer societies. There is a significant positive relationship between most characters investigated, but the trait "high gods" stands apart, suggesting that belief in a single creator deity can emerge in a society regardless of other aspects of its religion.

Hadza hunter-gatherer men do not have more masculine digit ratios (2D:4D).

OBJECTIVES: The ratio between the length of the second and the length of the fourth digit (2D:4D) is sexually dimorphic such that males of many species possess a lower ratio than females, particularly in the right hand. Still, men and women often exhibit overlapping 2D:4D ranges and the ratio is highly variable between populations. In order to further explore populational variability, we chose to analyze 2D:4D in the Hadza, a population of hunter-gatherers living in Tanzania. MATERIALS AND METHODS: Data were collected separately by two researchers over the course of three years (1998, 2001, 2006) from 152 adult participants (male: n = 76, female: n = 76). Independent samples t-tests were used to explore sex differences, paired samples t-tests were used to explore directional effects within each sex, and linear regression and one-way ANOVA were used to test possible age effects. RESULTS: In none of the years, or pooled (n = 152), did we find evidence that adult men have a lower 2D:4D than adult women. If anything, the data suggest that women in this population have a significantly lower right hand 2D:4D than men (P < 0.001, d = 0.57). In contrast, left hand 2D:4D did not exhibit a sex difference (P = 0.862, d = 0.03). DISCUSSION: These findings challenge the current view that lower 2D:4D in men is a uniform characteristic of our species. Cross-populational variance in 2D:4D may be related to known patterns of hormonal variation resulting from both genetic and environmental mechanisms, though this relationship merits further investigation.

Assessing digestibility of Hadza tubers using a dynamic in-vitro model.

OBJECTIVES: Bioaccessibility is a useful measure for assessing the biological value of a particular nutrient from food, especially foods such as tubers. The wild tubers exploited by Hadza foragers in Tanzania are of interest because they are nontoxic, consumed raw or briefly roasted, and entail substantial physical barriers to consumers. In this study, we attempted to elucidate the biological value of Hadza tubers by measuring the absorption of glucose through in-vitro digestion. METHODS: We quantified digestibility using data from 24 experimental trials on four species of Hadza tuber using a dynamic in-vitro model that replicates digestion in the stomach and small intestine. Analysis of glucose in the input meal and output dialysate revealed the accessible glucose fraction. We also conducted assays for protein, vitamin, and mineral content on whole tubers and meal fractions. RESULTS: Bioaccessibility of glucose varies depending on tuber species. Holding effects of chewing constant, brief roasting had negligible effects, but high intraspecific variation precludes interpretive power. Overall, Hadza tubers are very resistant to digestion, with between one- and two-thirds of glucose absorbed on average. Glucose absorption negatively correlated with glucose concentration of the tubers. CONCLUSIONS: Roasting may provide other benefits such as ease of peeling and chewing to extract edible parenchymatous tissue. A powerful factor in glucose acquisition is tuber quality, placing emphasis on the skill of the forager. Other nutrient assays yielded unexpectedly high values for protein, iron, and iodine, making tubers potentially valuable resources beyond caloric content.

Energy expenditure and activity among Hadza hunter-gatherers.

OBJECTIVES: Studies of total energy expenditure, (TEE; kcal/day) among traditional populations have challenged current models relating habitual physical activity to daily energy requirements. Here, we examine the relationship between physical activity and TEE among traditional Hadza hunter-gatherers living in northern Tanzania. METHODS: Hadza adults were studied at two camps, with minimal intervention so as to monitor energy expenditure and activity during normal daily life. We measured daily walking distance and walking speed using wearable GPS units for 41 adults. For a subset of 30 adults, we measured TEE using doubly labeled water, three indices of work load (foraging return rate, maternal status, and number of dependent children), and urinary biomarkers of metabolic activity and stress (8-hydroxydeoxyguanosine, cortisol, and testosterone). RESULTS: Fat-free mass was the single strongest predictor of TEE among Hadza adults (r(2) = 0.66, P < 0.001). Hadza men used greater daily walking distances and faster walking speeds compared with that of Hadza women, but neither sex nor any measure of physical activity or work load were correlated with TEE in analyses controlling for fat-free mass. Compared with developed, industrial populations, Hadza adults had similar TEE but elevated levels of metabolic stress as measured by 8-hydroxydeoxyguanosine. CONCLUSION: Our results indicate that daily physical activity may not predict TEE within traditional hunter-gatherer populations like the Hadza. Instead, adults with high levels of habitual physical activity may adapt by reducing energy allocation to other physiological activity.

Toward a reality-based understanding of Hadza men's work: a response to Hawkes et al. (2014).

Observations of Hadza men foraging out of camp and sharing food in camp show that men seeking to maximize the flow of calories to their families should pursue large game, and that hunting large game does not pose a collective action problem. These data also show that Hadza men frequently pursued honey, small game, and fruit, and that by doing so, provided a more regular flow of food to their households than would a putative big game specialist. These data support our earlier studies demonstrating that the goal of family provisioning is a robust predictor of Hadza men's behavior. As before, the show-off and costly signaling hypotheses advanced by Hawkes and colleagues fail as both descriptions of and explanations for Hadza men's work.

Honey, Hadza, hunter-gatherers, and human evolution.

Honey is the most energy dense food in nature. It is therefore not surprising that, where it exists, honey is an important food for almost all hunter-gatherers. Here we describe and analyze widespread honey collecting among foragers and show that where it is absent, in arctic and subarctic habitats, honey bees are also rare to absent. Second, we focus on one hunter-gatherer society, the Hadza of Tanzania. Hadza men and women both rank honey as their favorite food. Hadza acquire seven types of honey. Hadza women usually acquire honey that is close to the ground while men often climb tall baobab trees to raid the largest bee hives with stinging bees. Honey accounts for a substantial proportion of the kilocalories in the Hadza diet, especially that of Hadza men. Cross-cultural forager data reveal that in most hunter-gatherers, men acquire more honey than women but often, as with the Hadza, women do acquire some. Virtually all warm-climate foragers consume honey. Our closest living relatives, the great apes, take honey when they can. We suggest that honey has been part of the diet of our ancestors dating back to at least the earliest hominins. The earliest hominins, however, would have surely been less capable of acquiring as much honey as more recent, fully modern human hunter-gatherers. We discuss reasons for thinking our early ancestors would have acquired less honey than foragers ethnographically described, yet still significantly more than our great ape relatives.

Old stones' song: use-wear experiments and analysis of the Oldowan quartz and quartzite assemblage from Kanjera South (Kenya).

Evidence of Oldowan tools by ∼2.6 million years ago (Ma) may signal a major adaptive shift in hominin evolution. While tool-dependent butchery of large mammals was important by at least 2.0 Ma, the use of artifacts for tasks other than faunal processing has been difficult to diagnose. Here we report on use-wear analysis of ∼2.0 Ma quartz and quartzite artifacts from Kanjera South, Kenya. A use-wear framework that links processing of specific materials and tool motions to their resultant use-wear patterns was developed. A blind test was then carried out to assess and improve the efficacy of this experimental use-wear framework, which was then applied to the analysis of 62 Oldowan artifacts from Kanjera South. Use-wear on a total of 23 artifact edges was attributed to the processing of specific materials. Use-wear on seven edges (30%) was attributed to animal tissue processing, corroborating zooarchaeological evidence for butchery at the site. Use-wear on 16 edges (70%) was attributed to the processing of plant tissues, including wood, grit-covered plant tissues that we interpret as underground storage organs (USOs), and stems of grass or sedges. These results expand our knowledge of the suite of behaviours carried out in the vicinity of Kanjera South to include the processing of materials that would be 'invisible' using standard archaeological methods. Wood cutting and scraping may represent the production and/or maintenance of wooden tools. Use-wear related to USO processing extends the archaeological evidence for hominin acquisition and consumption of this resource by over 1.5 Ma. Cutting of grasses, sedges or reeds may be related to a subsistence task (e.g., grass seed harvesting, cutting out papyrus culm for consumption) and/or a non-subsistence related task (e.g., production of 'twine,' simple carrying devices, or bedding). These results highlight the adaptive significance of lithic technology for hominins at Kanjera.

Hunter-gatherers have less famine than agriculturalists.

The idea that hunter-gatherer societies experience more frequent famine than societies with other modes of subsistence is pervasive in the literature on human evolution. This idea underpins, for example, the 'thrifty genotype hypothesis'. This hypothesis proposes that our hunter-gatherer ancestors were adapted to frequent famines, and that these once adaptive 'thrifty genotypes' are now responsible for the current obesity epidemic. The suggestion that hunter-gatherers are more prone to famine also underlies the widespread assumption that these societies live in marginal habitats. Despite the ubiquity of references to 'feast and famine' in the literature describing our hunter-gatherer ancestors, it has rarely been tested whether hunter-gatherers suffer from more famine than other societies. Here, we analyse famine frequency and severity in a large cross-cultural database, in order to explore relationships between subsistence and famine risk. This is the first study to report that, if we control for habitat quality, hunter-gatherers actually had significantly less--not more--famine than other subsistence modes. This finding challenges some of the assumptions underlying for models of the evolution of the human diet, as well as our understanding of the recent epidemic of obesity and type 2 diabetes mellitus.

Evidence of Levy walk foraging patterns in human hunter-gatherers.

When searching for food, many organisms adopt a superdiffusive, scale-free movement pattern called a Lévy walk, which is considered optimal when foraging for heterogeneously located resources with little prior knowledge of distribution patterns [Viswanathan GM, da Luz MGE, Raposo EP, Stanley HE (2011) The Physics of Foraging: An Introduction to Random Searches and Biological Encounters]. Although memory of food locations and higher cognition may limit the benefits of random walk strategies, no studies to date have fully explored search patterns in human foraging. Here, we show that human hunter-gatherers, the Hadza of northern Tanzania, perform Lévy walks in nearly one-half of all foraging bouts. Lévy walks occur when searching for a wide variety of foods from animal prey to underground tubers, suggesting that, even in the most cognitively complex forager on Earth, such patterns are essential to understanding elementary foraging mechanisms. This movement pattern may be fundamental to how humans experience and interact with the world across a wide range of ecological contexts, and it may be adaptive to food distribution patterns on the landscape, which previous studies suggested for organisms with more limited cognition. Additionally, Lévy walks may have become common early in our genus when hunting and gathering arose as a major foraging strategy, playing an important role in the evolution of human mobility.

Household and kin provisioning by Hadza men.

We use data collected among Hadza hunter-gatherers between 2005 and 2009 to examine hypotheses about the causes and consequences of men's foraging and food sharing. We find that Hadza men foraged for a range of food types, including fruit, honey, small animals, and large game. Large game were shared not like common goods, but in ways that significantly advantaged producers' households. Food sharing and consumption data show that men channeled the foods they produced to their wives, children, and their consanguineal and affinal kin living in other households. On average, single men brought food to camp on 28% of days, married men without children at home on 31% of days, and married men with children at home on 42% of days. Married men brought fruit, the least widely shared resource, to camp significantly more often than single men. A model of the relationship between hunting success and household food consumption indicates that the best hunters provided 3-4 times the amount of food to their families than median or poor hunters. These new data fill important gaps in our knowledge of the subsistence economy of the Hadza and uphold predictions derived from the household and kin provisioning hypotheses. Key evidence and assumptions backing prior claims that Hadza hunting is largely a form of status competition were not replicated in our study. In light of this, family provisioning is a more viable explanation for why good hunters are preferred as husbands and have higher fertility than others.

The human operational sex ratio: effects of marriage, concealed ovulation, and menopause on mate competition.

Among mammals, male-male competition for sexual access to females frequently involves fighting. Larger body size gives males an advantage in fighting, which explains why males tend to be larger than females in many species, including anthropoid primates. Mitani et al. derived a formula to measure the operational sex ratio (OSR) to reflect the degree of male-male competition using the number of reproductively available males to females who are cycling and capable of conceiving. The OSR should predict the degree of sexual dimorphism in body mass-at least if male-male competition involves much fighting or threatening. Here, we use hunter-gatherer demographic data and the Mitani et al. formula to calculate the human OSR. We show that humans have a much lower degree of body mass sexual dimorphism than is predicted by our OSR. We suggest this is because human competition rarely involves fighting. In human hunter-gatherer societies, differences in the ages of marriage have an impact on competition in that the age of males at first marriage is younger when there is a lower percentage of married men with two or more wives, and older when there is a higher percentage of married men with two or more wives. We discuss the implications of this for females, along with the effects of two key life history traits that influence the OSR, concealed ovulation and menopause. While menopause decreases the number of reproductively available females to males and thus increases male-male competition, concealed ovulation decreases male-male competition. Finally, we discuss the importance of mostly monogamous mate bonds in human evolution.

Hunter-gatherer energetics and human obesity.

Western lifestyles differ markedly from those of our hunter-gatherer ancestors, and these differences in diet and activity level are often implicated in the global obesity pandemic. However, few physiological data for hunter-gatherer populations are available to test these models of obesity. In this study, we used the doubly-labeled water method to measure total daily energy expenditure (kCal/day) in Hadza hunter-gatherers to test whether foragers expend more energy each day than their Western counterparts. As expected, physical activity level, PAL, was greater among Hadza foragers than among Westerners. Nonetheless, average daily energy expenditure of traditional Hadza foragers was no different than that of Westerners after controlling for body size. The metabolic cost of walking (kcal kg(-1) m(-1)) and resting (kcal kg(-1) s(-1)) were also similar among Hadza and Western groups. The similarity in metabolic rates across a broad range of cultures challenges current models of obesity suggesting that Western lifestyles lead to decreased energy expenditure. We hypothesize that human daily energy expenditure may be an evolved physiological trait largely independent of cultural differences.

Subsistence and the evolution of religion.

We present a cross-cultural analysis showing that the presence of an active or moral High God in societies varies generally along a continuum from lesser to greater technological complexity and subsistence productivity. Foragers are least likely to have High Gods. Horticulturalists and agriculturalists are more likely. Pastoralists are most likely, though they are less easily positioned along the productivity continuum. We suggest that belief in moral High Gods was fostered by emerging leaders in societies dependent on resources that were difficult to manage and defend without group cooperation. These leaders used the concept of a supernatural moral enforcer to manipulate others into cooperating, which resulted in greater productivity. Reproductive success would accrue most to such leaders, but the average reproductive success of all individuals in the society would also increase with greater productivity. Supernatural enforcement of moral codes maintained social cohesion and allowed for further population growth, giving one society an advantage in competition with others.

Sex differences in Hadza dental wear patterns : a preliminary report.

Among hunter-gatherers, the sharing of male and female foods is often assumed to result in virtually the same diet for males and females. Although food sharing is widespread among the hunting and gathering Hadza of Tanzania, women were observed eating significantly more tubers than men. This study investigates the relationship between patterns of dental wear, diet, and extramasticatory use of teeth among the Hadza. Casts of the upper dentitions were made from molds taken from 126 adults and scored according to the Murphy dental attrition scoring system. Females had significantly greater anterior occlusal wear than males when we controlled for age. Males exhibited greater asymmetry in wear, with greater wear on the left side in canines, first premolars, and first molars. We suggest that these sex differences in wear patterns reflect the differences seen in the diet, as well as in the use of teeth as tools.

Social networks and cooperation in hunter-gatherers.

Social networks show striking structural regularities, and both theory and evidence suggest that networks may have facilitated the development of large-scale cooperation in humans. Here, we characterize the social networks of the Hadza, a population of hunter-gatherers in Tanzania. We show that Hadza networks have important properties also seen in modernized social networks, including a skewed degree distribution, degree assortativity, transitivity, reciprocity, geographic decay and homophily. We demonstrate that Hadza camps exhibit high between-group and low within-group variation in public goods game donations. Network ties are also more likely between people who give the same amount, and the similarity in cooperative behaviour extends up to two degrees of separation. Social distance appears to be as important as genetic relatedness and physical proximity in explaining assortativity in cooperation. Our results suggest that certain elements of social network structure may have been present at an early point in human history. Also, early humans may have formed ties with both kin and non-kin, based in part on their tendency to cooperate. Social networks may thus have contributed to the emergence of cooperation.

Sex differences in Hadza eating frequency by food type.

OBJECTIVES: We investigate sex differences in frequencies of adults eating in a foraging population-the Hadza of Tanzania. METHODS: We use eating frequency data from instantaneous scan observations of the Hadza, to see to how much sharing of foods taken back to camp compensates for the targeting of different foods by each sex while out foraging. RESULTS: Eating in camp differs by sex in terms of overall eating frequency, as well as in terms of diet composition (frequencies of eating each food type). We also control for sex-differences in time spent in camp and still find sex-differences in eating frequencies-women are observed eating significantly more frequently than men. There are also sex-differences in the eating frequencies of particular food types both with and without controlling for presence in camp. Finally, we use data on acquisition of each food type by sex and find that both sexes are more frequently observed eating women's foods in camp than men's foods. CONCLUSIONS: At least in the case of the Hadza, we see pronounced sex differences in the in-camp diet. Hadza men are eating a higher quality diet than are women, but women are able to eat far more frequently, and spend less time foraging than men. It is not yet clear whether a regular caloric intake of lower quality foods would be more beneficial for maintaining fecundity than a more variable diet consisting of higher quality foods.