Hunting income patterns among the Hadza: big game, common goods, foraging goals and the evolution of the human diet.

The assumption that large mammal hunting and scavenging are economically advantageous to hominid foragers is examined in the light of data collected among the Hadza of northern Tanzania. Hadza hunters disregard small prey in favour of larger forms (mean adult mass greater than or equal to 40 kg). Here we report experimental data showing that hunters would reduce their mean rates if they included small animals in the array they target. Still, daily variance in large animal hunting returns is high, and the risk of failure correspondingly great, significantly greater than that associated with small game hunting and trapping. Sharing large kills reduces the risk of meatless days for big game hunters, and obviates the problem of storing large amounts of meat. It may be unavoidable if large carcasses cannot be defended economically against the demands of other consumers. If so, then large prey are common goods. A hunter may gain no consumption advantage from his own big game acquisition efforts. We use Hadza data to model this 'collective action' problem, and find that an exclusive focus on large game with extensive sharing is not the optimal strategy for hunters concerned with maximizing their own chances of eating meat. Other explanations for the emergence and persistence of this practice must be considered.     

Male strategies and Plio-Pleistocene archaeology

Archaeological data are frequently cited in support of the idea that big game hunting drove the evolution of early Homo, mainly through its role in offspring provisioning. This argument has been disputed on two grounds: (1) ethnographic observations on modern foragers show that although hunting may contribute a large fraction of the overall diet, it is an unreliable day-to-day food source, pursued more for status than subsistence; (2) archaeological evidence from the Plio-Pleistocene, coincident with the emergence of Homo can be read to reflect low-yield scavenging, not hunting. Our review of the archaeology yields results consistent with these critiques: (1) early humans acquired large-bodied ungulates primarily by aggressive scavenging, not hunting; (2) meat was consumed at or near the point of acquisition, not at home bases, as the hunting hypothesis requires; (3) carcasses were taken at highly variable rates and in varying degrees of completeness, making meat from big game an even less reliable food source than it is among modern foragers. Collectively, Plio-Pleistocene site location and assemblage composition are consistent with the hypothesis that large carcasses were taken not for purposes of provisioning, but in the context of competitive male displays. Even if meat were acquired more reliably than the archaeology indicates, its consumption cannot account for the significant changes in life history now seen to distinguish early humans from ancestral australopiths. The coincidence between the earliest dates for Homo ergaster and an increase in the archaeological visibility of meat eating that many find so provocative instead reflects: (1) changes in the structure of the environment that concentrated scavenging opportunities in space, making evidence of their pursuit more obvious to archaeologists; (2) H. ergaster's larger body size (itself a consequence of other factors), which improved its ability at interference competition.     

Patch Choice Decisions among Ifaluk Fishers

Studies of patch choice decisions among human foragers have failed to explain why foragers do not exclusively exploit the patch with the highest mean profitability. One possible explanation is that profitability rankings are likely to vary daily; however, this instability is not captured when profitabilities are calculated as a sampled average over a longer time span. Here I present data on the patch choice decisions of Ifaluk fishers to evaluate whether men are responding to daily variation in the profitability of their primary fishing patch. Results show that men choose to fish most frequently in the patch with the highest mean profitability. Men fish in alternative patches (alternative from the most profitable patch) when, on that morning or the previous day, return rates in the most profitable patch are lower than the overall mean per capita return rate of alternative patches. Results also indicate that when fishers pursue alternative patches after fishing in the patch with the highest profitability, their mean per capita return rates are generally higher in the alternative patches exploited. However, variance in the profitability of the most profitable patch cannot explain why men exploit two patches, the Nine-mile reef and the dogtoothed tuna patch, which on average have very low profitability. These results and directions for future research are discussed. [Keywords: human behavioral ecology, patch choice decisions, Micronesia]     

High adult mortality among Hiwi hunter-gatherers: implications for human evolution

Extant apes experience early sexual maturity and short life spans relative to modern humans. Both of these traits and others are linked by life-history theory to mortality rates experienced at different ages by our hominin ancestors. However, currently there is a great deal of debate concerning hominin mortality profiles at different periods of evolutionary history. Observed rates and causes of mortality in modern hunter-gatherers may provide information about Upper Paleolithic mortality that can be compared to indirect evidence from the fossil record, yet little is published about causes and rates of mortality in foraging societies around the world. To our knowledge, interview-based life tables for recent hunter-gatherers are published for only four societies (Ache, Agta, Hadza, and Ju/'hoansi). Here, we present mortality data for a fifth group, the Hiwi hunter-gatherers of Venezuela. The results show comparatively high death rates among the Hiwi and highlight differences in mortality rates among hunter-gatherer societies. The high levels of conspecific violence and adult mortality in the Hiwi may better represent Paleolithic human demographics than do the lower, disease-based death rates reported in the most frequently cited forager studies.     

Macronutrient modifications of optimal foraging theory: An approach using indifference curves applied to some modern foragers

The use of energy (calories) as the currency to be maximized per unit time in Optimal Foraging Models is considered in light of data on several foraging groups. Observations on the Ache, Cuiva, and Yora foragers suggest men do notattempt to maximize energetic return rates, but instead often concentrate on acquiring meat resources which provide lower energetic returns. The possibility that this preference is due to the macronutrient composition of hunted and gathered foods is explored. Indifference curves are introduced as a means of modeling the tradeoff between two desirable commodities, meat (protein-lipid) and carbohydrate, and a specific indifference curve is derived using observed choices in five foraging situatiuons. This curve is used to predict the amount o meat that Mbuti foragers will trade for carbohydrate, in an attempt to test the utility of the approach.     

Partial nectar loads as a cause of multiple nectar transfer in the honey bee (Apis mellifera): a simulation model

Honey bee foragers transfer their nectar loads to receiver bees within the nest. Surprisingly, they often transfer to more than one receiver (published values range from 1.9 to 2.7). Several adaptive hypotheses have been proposed to explain why multiple transfer occurs. One hypothesis, information improvement, states that multiple transfer arises as an adaptive forager-driven process. Foragers use the delay in finding a receiver to assess the relative work capacities of foragers and receivers, performing recruitment dances when appropriate. Multiple transferring improves their delay information. We used a stochastic simulation model to investigate the non-adaptive partial loads hypothesis. We determined the extent to which partial crop loads and receiver filling and emptying rules (i.e. how much nectar to accept before leaving the transfer area) can cause multiple transfer. As many as 1.9 nectar transfers per returning forager were generated within biologically realistic parameter space. We suggest that much multiple transfer arises as a non-adaptive consequence of partitioning nectar foraging between foragers and receivers, but that this will also result in foragers having better information about the relative work capacities of foragers and receivers as a useful consequence. We suggest that the number of transfers caused by partial loads could also be increased by an adaptive forager-driven effort to improve their information concerning the balance of foragers and receivers and we outline a framework wherein the information improvement hypothesis can be directly tested.     

The Organization of Male and Female Labor in Foraging Societies: Implications for Early Paleoindian Archaeology

I use cross-cultural ethnographic data to explore the relationship between male and female subsistence labor among hunter-gatherer populations by examining data regarding resource procurement, time allocation, and task differentiation between the sexes relative to dependence on hunted foods. The findings indicate that female foragers generally perform a variety of nonsubsistence collection activities and preferentially procure high-return resources in hunting-based economies. I develop ideas about predictable relationships concerning the amount of time female foragers expend on subsistence and technological tasks relative to the dietary contribution of meat. I then use ethnographic trends to evaluate archaeological assumptions regarding the sexual division of labor in prehistoric foraging contexts, focusing on the dichotomous views of Clovis labor organization. I argue that archaeological interpretations of prehistoric labor roles in hunting-based foraging societies are commonly polarized between stereotypical views of male and female subsistence behaviors. I develop an interpretation of Early Paleoindian labor organization, emphasizing female labor in the production of material goods and the procurement of low-risk plant and animal resources based on global economic trends among foragers.     

The Mechanisms of Water Exchange: The Regulatory Roles of Multiple Interactions in Social Wasps

Evolutionary benefits of task fidelity and improving information acquisition via multiple transfers of materials between individuals in a task partitioned system have been shown before, but in this paper we provide a mechanistic explanation of these phenomena. Using a simple mathematical model describing the individual interactions of the wasps, we explain the functioning of the common stomach, an information center, which governs construction behavior and task change. Our central hypothesis is a symmetry between foragers who deposit water and foragers who withdraw water into and out of the common stomach. We combine this with a trade-off between acceptance and resistance to water transfer. We ultimately derive a mathematical function that relates the number of interactions that foragers complete with common stomach wasps during a foraging cycle. We use field data and additional model assumptions to calculate values of our model parameters, and we use these to explain why the fullness of the common stomach stabilizes just below 50 percent, why the average number of successful interactions between foragers and the wasps forming the common stomach is between 5 and 7, and why there is a variation in this number of interactions over time. Our explanation is that our proposed water exchange mechanism places natural bounds on the number of successful interactions possible, water exchange is set to optimize mediation of water through the common stomach, and the chance that foragers abort their task prematurely is very low.     

Learning rules for social foragers: implications for the producer-scrounger game and ideal free distribution theory

In population games, the optimal behaviour of a forager depends partly on courses of action selected by other individuals in the population. How individuals learn to allocate effort in foraging games involving frequency-dependent payoffs has been little examined. The performance of three different learning rules was investigated in several types of habitats in each of two population games. Learning rules allow individuals to weigh information about the past and the present and to choose among alternative patterns of behaviour. In the producer-scrounger game, foragers use producer to locate food patches and scrounger to exploit the food discoveries of others. In the ideal free distribution game, foragers that experience feeding interference from companions distribute themselves among heterogeneous food patches. In simulations of each population game, the use of different learning rules induced large variation in foraging behaviour, thus providing a tool to assess the relevance of each learning rule in experimental systems. Rare mutants using alternative learning rules often successfully invaded populations of foragers using other rules indicating that some learning rules are not stable when pitted against each other. Learning rules often closely approximated optimal behaviour in each population game suggesting that stimulus-response learning of contingencies created by foraging companions could be sufficient to perform at near-optimal level in two population games.     

Socially stable territories: the negotiation of space by interacting foragers

This article presents a theory of territoriality that integrates optimal foraging and conflict resolution through negotiation. Using a spatially explicit model of a sit-and-wait forager, we show that when resources are scarce, there is a conflict between foragers: there is not enough space for all individuals to have optimal home ranges. We propose that a division of space that solves this conflict over resources is the outcome of a negotiation between foragers. We name this outcome the socially stable territories (SST). Using game theory we show that in a homogenous patch occupied by two interacting foragers, both individuals receive identical energy yields at the socially stable territories; that is, there is economic equity. Economic inequity can arise in a heterogeneous patch or from asymmetries in fighting abilities between the foragers. Opportunity costs play a role in reducing economic inequity. When the asymmetry in fighting abilities is very large, a negotiated division of space is not possible and the forager with lowest fighting ability may be evicted from the habitat patch. A comparison between territories and overlapping home ranges shows that energy yields from territories are generally higher. We discuss why there are instances in which individuals nevertheless overlap home ranges.     

Increasing the resolution of the Broad Spectrum Revolution in the Southern Levantine Epipaleolithic (19-12 ka)

We analyze terminal Pleistocene archaeofaunal diversity trends in the Southern Levant by examining eight Epipaleolithic (ca. 19-12 ka) assemblages from the Western Galilee/Mt. Carmel, Israel subregion. We test predictions from a Broad Spectrum Revolution model of the population dynamics of human foragers and their prey. The study emphasizes control over geographic variability and archaeological recovery and recording methods, as we analyze a time series that samples the Epipaleolithic more fully than have previous studies. This provides a new opportunity to examine human population and economic change in the long-term transition to sedentism and agriculture.We use the Mantel test to evaluate the significance of temporal trends in body-size-based big game diversity, as well as in diversity of small game prey types. Results demonstrate a highly significant decline through time in the relative abundance of medium and large big game, measured relative to small big game. This suggests that the apparent “gazelle specialization” by Late Epipaleolithic (Natufian) hunters reflects longer-term anthropogenic overexploitation of the largest prey types in the spectrum. While large and medium big game abundance declined, our results show small game increased in economic importance over time.Considered with associated climate change data, the results provide substantial support for the hypothesis that local human populations expanded rapidly in size after the Last Glacial Maximum (LGM). We suggest that following the post-LGM population pulse, human foragers adopted a shifting series of intensification strategies mediated by changes in residential mobility.     

The evolution of multiple mating: Costs and benefits of polyandry to females and of polygyny to males

Polyandry is a paradox: why do females mate multiple times when a single ejaculate often provides enough sperm for lifetime egg production? Gowaty et al. addressed explanations for polyandry in Drosophila pseudoobscura from the perspective of hypotheses based on sex differences in costs of reproduction (CoR). Contrary to CoR, Gowaty et al. showed that (1) a single ejaculate was inadequate for lifetime egg production; (2) polyandry provided fitness benefits to females beyond provision of adequate sperm and (3) fitness benefits of polyandry were not offset by costs. Here, I discuss predictions of the ad hoc hypotheses of CoR and three alternative hypotheses to CoR to facilitate a discussion and further development of a strong inference approach to experiments on the adaptive significance of polyandry for females. Each of the hypotheses makes testable predictions; simultaneous tests of the predictions will provide a strong inference approach to understanding the adaptive significance of multiple mating. I describe a sex-symmetric experiment meant to evaluate variation in fitness among lifelong virgins (V); monogamous females and males with one copulation (M_OC); monogamous females and males with multiple copulations (M_MC); P_AND, polyandrous females; and P_GYN, polygynous males. Last, I recommend the study of many different species, while taking care in choice of study species and attention to the assumptions of specific hypotheses. I particularly urge the study of many more Drosophila species both in laboratory and the wild to understand the “nature of flies in nature,” where opportunities and constraints mold evolutionary responses.     

Are kubo hunters ‘show offs’?

The ‘show off’ hypothesis proposed by K. Hawkes, and tested using data on Ache foragers, makes important connections between food resource choice, reproductive strategies, and food sharing by human foragers. We test predictions derived from that hypothesis concerning contexts of meat acquisition, association between individuals, mobility, and reproductive success among Kubo hunter-horticulturalists of the interior lowlands of Papua New Guinea. Application of the hypothesis to both the Kubo and Ache cases is questioned. Differences between Kubo males in means and variances of returns from hunting arise as a consequence of differential target specialization; they do not map onto variation in reproductive success.     

Eavesdropping foragers use level of collective commotion as public information to target high quality patches

Public information offers a valuable means for social foragers to determine the relative quality of foraging patches. Despite much evidence that foragers use public information based on others’ feeding behavior, no experiments have examined whether foragers might use public information based on others’ competitive behavior, particularly the collective commotion that can be generated by aggregations. Such commotion could potentially provide a rich source of public information: as foragers compete in a patch with an especially high value resource, their heightened competition intensity could enable eavesdropping foragers to target this superior patch, based simply on its higher level of collective commotion. To test the hypothesis that the level of collective commotion is used as public information by eavesdropping foragers I conducted field experiments on terrestrial hermit crabs Coenobita compressus. These animals engage in collective competitive interactions in foraging patches for food and shells, generating variable levels of commotion across different quality patches. By experimentally manipulating the level of collective commotion in sham aggregations in the wild I show that a higher level of commotion is exploited by eavesdropping foragers to differentially target more valuable patches. Broadly, these results highlight an underappreciated significance of competitive by‐products and higher‐ order collective pheno mena as forms of public information for foragers.     

Provisioning of Game Meat to Rural Communities as a Benefit of Sport Hunting in Zambia

Sport hunting has reportedly multiple benefits to economies and local communities; however, few of these benefits have been quantified. As part of their lease agreements with the Zambia Wildlife Authority, sport hunting operators in Zambia are required to provide annually to local communities free of charge i.e., provision a percentage of the meat obtained through sport hunting. We characterized provisioning of game meat to rural communities by the sport hunting industry in Zambia for three game management areas (GMAs) during 2004–2011. Rural communities located within GMAs where sport hunting occurred received on average > 6,000 kgs per GMA of fresh game meat annually from hunting operators. To assess hunting industry compliance, we also compared the amount of meat expected as per the lease agreements versus observed amounts of meat provisioned from three GMAs during 2007–2009. In seven of eight annual comparisons of these GMAs, provisioning of meat exceeded what was required in the lease agreements. Provisioning occurred throughout the hunting season and peaked during the end of the dry season (September–October) coincident with when rural Zambians are most likely to encounter food shortages. We extrapolated our results across all GMAs and estimated 129,771 kgs of fresh game meat provisioned annually by the sport hunting industry to rural communities in Zambia at an approximate value for the meat alone of >US$600,000 exclusive of distribution costs. During the hunting moratorium (2013–2014), this supply of meat has halted, likely adversely affecting rural communities previously reliant on this food source. Proposed alternatives to sport hunting should consider protein provisioning in addition to other benefits (e.g., employment, community pledges, anti-poaching funds) that rural Zambian communities receive from the sport hunting industry.     

Energetic demand of multiple dependents and the evolution of slow human growth

This study investigates the consequences of the human foraging niche and multiple dependent offspring on the optimal growth trajectory of humans. We test the hypothesis that the human pattern of slow human growth between age at weaning and puberty helps defer the compound energetic demand on parents with multiple dependents, by using growth and demographic data from two foraging societies, the Ache of eastern Paraguay and the Dobe Ju/'hoansi of Botswana and Namibia. We run simulations of observed and potential growth trajectories among sub-adults and their consequent energetic demands on parents given profiles of fertility, mortality, consumption and production. We find that either sub-adult production or food subsidies from other people must substantially increase in order to compensate for the dramatic increase in energetic demand on parents if offspring were to grow faster at younger ages. Our conclusion is that slow human growth followed by a rapid adolescent growth spurt may have facilitated rising human fertility rates and greater investments in neural capital.     

The state as a whiteman,the whiteman as a /’hun: personhood,recognition, and the politics of knowability in the Kalahari

The article is dedicated to the loving memory of !A|’xuni.

The Ju|’hoansi of east central Namibia sometimes refer to the state as a whiteman and to the whiteman as a /’hun (steenbok). In this article, I contextualize these naming practices by tracing the history of colonial encounters on the fringes of the Western Kalahari through a small-scale animist perspective. I then discuss what this means for the concept of ‘recognition’, which I treat as a two-way intersubjective process of making oneself un/knowable to others. I argue that the Ju|’hoansi have engaged in parallel processes of mis/recognition vis-à-vis their colonial Others. By failing to enter into reciprocal relations with the Ju|’hoansi, the whiteman and the state have remained outside of the Ju|’hoansi's social universe and have thus compromised their own personhood.     

Climate change,collections and the classroom: using big data to tackle big problems

Preparing students to explore, understand, and resolve societal challenges such as global climate change is an important task for evolutionary and ecological biologists that will require novel and innovative pedagogical approaches. Recent calls to reform undergraduate science education emphasize the importance of engaging students in inquiry-driven, active, and authentic learning experiences. We believe that the vast digital resources (i.e., “big data”) associated with natural history collections provide invaluable but underutilized opportunities to create such experiences for undergraduates in biology. Here, we describe an online, open-access educational module that we have developed that harnesses the power of collections-based information to introduce students to multiple conceptual and analytical elements of climate change, evolutionary, and ecological biology research. The module builds upon natural history specimens and data collected over the span of nearly a century in Yosemite National Park, California, to guide students through a series of exercises aimed at testing hypotheses regarding observed differences in response to climate change by two closely related and partially co-occurring species of chipmunks. The content of the module can readily be modified to meet the pedagogical goals and instructional levels of different courses while the analytical strategies outlined can be adapted to address a wide array of questions in evolutionary and ecological biology. In sum, we believe that specimen-based natural history data represent a powerful platform for reforming undergraduate instruction in biology. Because these efforts will result in citizens who are better prepared to understand complex biological relationships, the benefits of this approach to undergraduate education will have widespread benefits to society.