Showing off: Tests of an hypothesis about men's foraging goals

It is widely assumed that among hunter-gatherers, men work to provision their families. However, men may have more to gain by giving food to a wide range of companions who treat them favorably in return. If so, and if some resources better serve this end, men's foraging behavior should vary accordingly. Aspects of this hypothesis are tested on observations of food acquisition and sharing among Ache foragers of Eastern Paraguay. Previous analysis showed that different Ache food types were differently shared. Resources shared most widely were game animals. Further analysis and additional data presented here suggest a causal association between the wide sharing of game and the fact that men hunt and women do not. Data show that men preferentially target resources in both hunting and gathering which are more widely shared, resources more likely to be consumed outside their own nuclear families. These results have implications for 1) the identification of male reproductive trade-offs in human societies, 2) the view that families are units of common interest integrated by the sexual division of labor, 3) current reconstructions of the evolution of foraging and food sharing among early hominids, and 4) assessments of the role of risk and reciprocity in hunter-gatherer foraging strategies.     

Accepting loss: the temporal limits of reciprocity in brown capuchin monkeys

Delayed reciprocity is a potentially important mechanism for cooperation to occur. It is however rarely reported among animals, possibly because it requires special skills like the ability to plan a loss. We tested six brown capuchin monkeys (Cebus apella) in such skills. Subjects were studied in exchange tasks in which they had to retain a food item for a given time lag before returning it to an experimenter and obtaining a more desirable reward. Experiments showed that the subjects could wait for several minutes when allowed to return only part of the initial item. When required to return the full item intact, however, most subjects could not sustain a time lag longer than 10 s. Although the duration of waiting increased with the amount of return expected by subjects, in most cases it did not extend beyond 20 s even when the eperimenter offered a food amount 40 fold the initial item. The failure of capuchin monkeys to sustain long-lasting waiting periods may be explained by limited self-control abilities. This would prevent them achieving reciprocal altruism.     

Distribution Patterns of Tropical Plant Foods as an Evolutionary Stimulus to Primate Mental Development

Primates are noted for their mental abilities but the selective basis for such traits has remained obscure. It is hypothesized that the element of predictability associated with the spatial and temporal distribution patterns of plant foods in tropical forests has served to stimulate mental development in primates taking much of their food from the first trophic level. Primates able to remember the locations and phenological patterns of a wide variety of plant foods could move directly to such foods when and where available without wasting time and energy in random search. This would enhance overall foraging success by lowering procurement costs associated with a varied and patchily distributed plant diet. Membership in a cohesive social unit, that utilized the same supplying area over many consecutive generations, would also enhance foraging success by serving to transmit important information about diet to close kin. Data on the foraging behavior of howler and spider monkeys are presented to test certain implications of this hypothesis. Similar selective pressures, but applied to foods from the second trophic level, may have been of critical importance in the mental development of hominids . [primates, evolution, intelligence, plant foods, Aleles, Alouatta ]     

Crows and common ravens do not reciprocally exchange tokens with a conspecific to gain food rewards

Human economic transactions are based on complex forms of reciprocity, which involve the capacities to share and to keep track of what was given and received over time. Animals too engage in reciprocal interactions, but mechanisms such as calculated reciprocity have only been shown experimentally in few species. Various forms of cooperation, for example food and information sharing, are frequently observed in corvids, and they can engage in exchange interactions with human experimenters and accept delayed rewards. Here, we tested whether crows and common ravens would reciprocally exchange tokens with a conspecific in an exchange task. Birds received a set of three different types of tokens, some valuable for themselves, that is, they could exchange them for a food reward with a human experimenter, some valuable for their partner and some without value. The valuable tokens differed between the birds, which means that each bird could obtain more self-value tokens from their partner's compartment. We did not observe any active transfers, that is one individual giving a token to the experimental partner by placing it in its beak. We only observed 6 indirect transfers, that is one individual transferring a token into the compartment of the partner (3 no-value, 1 partner-value and 2 self-value tokens) and 67 “passive” transfers, that is one subject taking the token lying in reach in the compartment of the partner. Individuals took significantly more self-value tokens compared with no-value and partner-value tokens. This indicates a preference for tokens valuable to focal individuals. Significantly more no-value tokens compared with partner-value tokens were taken, likely to be caused by experimental partners exchanging self-value tokens with the human experimenter, and therefore, more no-value tokens being available in the compartment. Our results presently do not provide empirical support for reciprocity in crows and ravens, most likely caused by them not understanding the potential roles of receiver and donor. We therefore suggest further empirical tests of calculated reciprocity to be necessary in corvids.     

A study of carnivore competition in riparian and open habitats of modern savannas and its implications for hominid behavioral modelling

Some of the models proposed to explain Plio-Pleistocene hominid behavior and the formation of early East African archaeological sites are based on the assumption that the riparian habitats in which most of them occur were places of low interspecific competition. Competition is expressed here in terms of carnivore and hominid interactions. In this paper, a study of carnivore interaction in open and closed habitats is presented. The results indicate that riparian woodland shows the lowest degree of competition in savanna ecosystems. This suggests that if Plio-Pleistocene carnivores were adapted like their modern counterparts, the paleoecological settings of early sites could have provided hominids with enough safety to process carcasses and behave as shown in "central-place", "near-kill location" and "refuge" foraging models.     

Performance of human groups in social foraging: the role of communication in consensus decision making

Early hominids searched for dispersed food sources in a patchy, uncertain environment, and modern humans encounter equivalent spatial-temporal coordination problems on a daily basis. A fundamental, but untested assumption is that our evolved capacity for communication is integral to our success in such tasks, allowing information exchange and consensus decisions based on mutual consideration of pooled information. Here we examine whether communication enhances group performance in humans, and test the prediction that consensus decision-making underlies group success. We used bespoke radio-tagging methodology to monitor the incremental performance of communicating and non-communicating human groups (small group sizes of two to seven individuals), during a social foraging experiment. We found that communicating groups (n = 22) foraged more effectively than non-communicating groups (n = 21) and were able to reach consensus decisions (an 'agreement' on the most profitable foraging resource) significantly more often than non-communicating groups. Our data additionally suggest that gesticulations among group members played a vital role in the achievement of consensus decisions, and therefore highlight the importance of non-verbal signalling of intentions and desires for successful human cooperative behaviour.     

Seasonal selection of soil types and grass swards by roan antelope in a South African savanna

Roan antelope are distributed mainly in regions characterized by infertile soils, offering food of low quality. We hypothesized that roan may select localities with higher soil nutrient levels and/or grass swards with more favourable properties in terms of food abundance or quality than generally available in those regions. Roan antelope were observed in a savanna region in South Africa where soils of widely varying nutrient status occurred. Roan favoured open grassland over wooded savanna areas. During the wet season, roan preferred sites with felsite‐derived soil of intermediate soil nutrient status. Grasslands growing on nutrient enriched alluvial soils were preferred outside of the early wet season, although most of the favourable sward characteristics were present in other landscape units. Food quantity, rather than quality appeared to attract roan to foraging sites in the late wet and early dry seasons. Food quality appeared more important in the early wet and late dry seasons. The higher degree of clustering of leafy material within foraging swards seemed to be an additional discriminating factor. The factors governing the selection of foraging sites by roan did not seem notably different from those influencing other species of grazing ruminant, but roan nevertheless seemed tolerant of stemmy grasslands growing on nutrient richer substrates.     

Estimating hominid life history: the critical interbirth interval

Unlike any great apes, humans have expanded into a wide variety of habitats during the course of evolution, beginning with the transition by australopithecines from forest to savanna habitation. Novel environments are likely to have imposed hominids a demographic challenge due to such factors as higher predation risk and scarcer food resources. In fact, recent studies have found a paucity of older relative to younger adults in hominid fossil remains, indicating considerably high adult mortality in australopithecines, early Homo, and Neanderthals. It is not clear to date why only human ancestors among all hominoid species could survive in these harsh environments. In this paper, we explore the possibility that hominids had shorter interbirth intervals to enhance fertility than the extant apes. To infer interbirth intervals in fossil hominids, we introduce the notion of the critical interbirth interval, or the threshold length of birth spacing above which a population is expected to go to extinction. We develop a new method to obtain the critical interbirth intervals of hominids based on the observed ratios of older adults to all adults in fossil samples. Our analysis suggests that the critical interbirth intervals of australopithecines, early Homo, and Neanderthals are significantly shorter than the observed interbirth intervals of extant great apes. We also discuss possible factors that may have caused the evolutionary divergence of hominid life history traits from those of great apes.     

A marginal model of tolerated theft

Using marginal analysis to represent Blurton Jones's concept of tolerated theft, I show how equilibrium resource transfers among individuals might be affected by foraging behavior, resource qualities, and number of participants. The model applies to hominids and other species that exchange or share food or other resources. Among the results: Tolerated theft enhances the value to be derived from resources, packets intermediate in size are most likely to be subjected to tolerated theft, packet division is more likely to be unequal than equal, division is a function of group size, and tolerated theft is most likely in small groups. The model also suggests that among reciprocators the widest possible exchange or sharing is in the self-interest of the individual procuring the resource. In general, evolutionary cost-benefit accounting should track marginal changes in the value (fitness or utility) of resources. Marginal valuation is conceptually primary and may produce results that differ from direct measures of quantity.     

Risk and the evolution of human exchange

Compared with other species, exchange among non-kin is a hallmark of human sociality in both the breadth of individuals and total resources involved. One hypothesis is that extensive exchange evolved to buffer the risks associated with hominid dietary specialization on calorie dense, large packages, especially from hunting. 'Lucky' individuals share food with 'unlucky' individuals with the expectation of reciprocity when roles are reversed. Cross-cultural data provide prima facie evidence of pair-wise reciprocity and an almost universal association of high-variance (HV) resources with greater exchange. However, such evidence is not definitive; an alternative hypothesis is that food sharing is really 'tolerated theft', in which individuals possessing more food allow others to steal from them, owing to the threat of violence from hungry individuals. Pair-wise correlations may reflect proximity providing greater opportunities for mutual theft of food. We report a laboratory experiment of foraging and food consumption in a virtual world, designed to test the risk-reduction hypothesis by determining whether people form reciprocal relationships in response to variance of resource acquisition, even when there is no external enforcement of any transfer agreements that might emerge. Individuals can forage in a high-mean, HV patch or a low-mean, low-variance (LV) patch. The key feature of the experimental design is that individuals can transfer resources to others. We find that sharing hardly occurs after LV foraging, but among HV foragers sharing increases dramatically over time. The results provide strong support for the hypothesis that people are pre-disposed to evaluate gains from exchange and respond to unsynchronized variance in resource availability through endogenous reciprocal trading relationships.     

Foraging costs drive female resistance to a sensory trap

Male ornaments can evolve through the exploitation of female perceptual biases such as those involved in responding to cues from food. This type of sensory exploitation may lead to confusion between the male signals and the cues that females use to find/recognize food. Such interference would be costly to females and may be one reason why females evolve resistance to the male ornaments. Using a group of species of viviparous fish where resistance to a sensory trap has evolved, we demonstrate that females exposed to an ornament that resembles food have a diminished foraging efficiency, that this effect is apparent when foraging on a food item with which the ornament shares visual attributes, and that not all species are equally affected by such confusion. Our results lend support to the model of ornamental evolution through chase-away sexual conflict.     

Hominid divergence and speech evolution

Hominids evolved from a population which diverged from other hominoids during the Mio-Pliocene. This population was perhaps forced by ecological conditions and competitive exclusion to rely more on tools, gathering, hunting, vocal communication and memory, under whose mutually positively reinforcing effects the hominids diverged. The ape ancestors may have been forced into the forests (or they may have forced hominids onto the savanna), while hominids adapted to a plains, hunting econiche.Speech was selected for because verbal symbols served as retrieval cues for a large number of complex concepts and were transmissible, and thus could be used to influence food-getting and other behavior by the social group.Speech is dependent on three inherited entities: (1) anatomical and neurological adaptations which allow vocalization of a wide range of phonemes in rapid succession and which allow for (2) duality of patterning, thereby promoting a large number of words, and (3) encoding, which greatly increases the rate at which verbal information transfer can occur. Speech may have evolved through small hominid groups using progressively more phonemes in an increasingly blended manner, with encoding subsequently being selected for. Neandertals apparently could not encode speech and could speak only a restricted range of phonemes. Their expanded cranial capacity may have been selected for to store ambiguous and slowly transmitted verbal data.     

Extractive foraging and the evolution of primate intelligence

One of the two major theories regarding the evolution of intelligence in primates is that feeding strategies determine mental development. Evidence for this theory is reviewed and related to extractive foraging, which is the act of locating and/or processing embedded foods such as underground roots and insects or hard-shelled nuts and fruits. It is shown that, although only cebus monkeys and chimpanzees in the wild use tools in extractive foraging, many other species of mammals (including primates) and birds are capable of extracting embedded foods without tools. Extractive foraging by primates is compared to extractive foraging by other mammals and birds to assess whether: 1) extractive foraging involves cognition, and 2) extractive foraging by primates is unique in a way that may mean it played a role in the development of intelligence among primates. This comparison reveals that some acts of extractive foraging by nonprimates are equally sophisticated as those of primates. It is suggested that extractive foraging played no significant role in the evolution of primate intelligence. Hypotheses for testing precise differences in extractive foraging ability across taxa are offered, and the roles of olfactory cues, manual dexterity, and strength in extractive foraging are evaluated. In conclusion, the hominization process is briefly reviewed in relation to foraging behavior. A ?package? of traits that, in combination, is unique to hominids is discussed: tool-aided extractive foraging, division of labor by sex with food exchange, and feeding of juveniles.     

Effects of three food enrichment items on the behavior of black lemurs (Eulemur macaco macaco) and Ringtail Lemurs (Lemur catta) at the Henson Robinson Zoo, Springfield, Illinois

This study tested 3 food enrichment items mentioned in a laboratory primate newsletter with 6 adult Eulemur macaco and 3 adult Lemur catta to examine whether the items would affect the behavior of the lemurs. The results suggest that Food Enrichment Item 3 (a wire box filled with whole grapes, apples, or both hidden in straw hung from a branch within the enclosure) caused a significant decrease in the incidence of resting and a significant increase in the incidences of playing and grooming, with no significant effect on the incidence of feeding or foraging. The lemurs' behavior appeared to be most affected by the food enrichment item that required the most manipulation, closely followed by an enrichment that required a moderate amount of manipulation. The order of the exposure to the food enrichment items and the day of the week appear to have an attenuation effect on these behaviors and did affect the incidence of 3 stereotypic behaviors exhibited by a male L. catta such that 3 behaviors declined in occurrence as the study progressed.     

Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): testing hypotheses on the evolution of bipedalism

A host of ecological, anatomical, and physiological selective pressures are hypothesized to have played a role in the evolution of hominid bipedalism. A referential model, based on the chimpanzee (Pan troglodytes) and bonobo (Pan paniscus), was used to test through experimental manipulation four hypotheses on the evolution of hominid bipedalism. The introduction of food piles (Carry hypothesis) increased locomotor bipedality in both species. Neither the introduction of branches (Display hypothesis) nor the construction of visual barriers (Vigilance hypothesis) altered bipedality in either species. Introduction of raised foraging structures (Forage hypothesis) increased postural bipedality in chimpanzees. These experimental manipulations provided support for carrying of portable objects and foraging on elevated food-items as plausible mechanisms that shaped bipedalism in hominids.     

Why hunt? Why gather? Why share? Hadza assessments of foraging and food-sharing motive

Over the last half century, anthropologists have vigorously debated the adaptive motivations underlying food acquisition choices and food-sharing among hunter-gatherer groups. Numerous explanations have been proposed to account for high-levels of generosity in food-sharing, including self- and family-provisioning, reciprocity, tolerated theft and pro-social- or skill-signaling. However, few studies have asked foragers directly and systematically about the motivations underlying their foraging and sharing decisions. We recruited 110 Hadza participants and employed a combination of free-response, yes/no, ranking and forced-choice questions to do just this. In free-response answers, respondents typically gave outcome-oriented accounts of foraging motive (e.g., to get food) and moralistic accounts of sharing motive (e.g., I have a good heart). In ranking tasks, participants gave precedence to reciprocity as a motive for sharing food beyond the household. We found small but clear gender differences in foraging motive, in line with previous predictions: women were more likely than men to rank family-provisioning highly whereas men were more likely than women to rank skill-signaling highly. However, despite these gender differences, the relative importance of different motivations was similar across genders and skill-signaling, sharing and family-provisioning were the most important motivators of foraging activity for both men and women. Contrary to the expectations of tolerated theft, peer complaints and requests for food ranked very low. There are several compelling reasons that evolutionary thinkers, typically interested in ultimate-level adaptive processes, have traditionally eschewed direct and explicit investigations of motive. However, these data may yet provide important insights.     

Female Bechstein's Bats Share Foraging Sites with Maternal Kin but do not Forage Together with them – Results from a Long‐Term Study

In many social animals, group members exchange information about where to feed. Thereby, they may gain direct benefits, for example, if social hunting enhances individual foraging success. Alternatively, individuals may receive indirect fitness benefits by preferentially sharing information about suitable feeding sites with kin. Indeed, in some species, a positive correlation between the degree of relatedness among individuals and the overlap among their foraging areas was found. However, sharing foraging sites with kin can also have costs if it increases food competition, which is not compensated by direct benefits. The goal of this study was to investigate whether sharing of individual foraging areas in female Bechstein's bats is best explained by kin selection or by direct benefits through social foraging. To assess their individual foraging behaviour, we analysed radio‐tracking data of 22 members of one maternity colony, including nine mother–daughter pairs, seven pairs of less closely related individuals and six pairs of unrelated bats. We examined the bats' fidelity to specific foraging areas during several years and quantified the influence of kinship on the overlap among individual foraging areas. By measuring how close to each other the bats foraged, we assessed whether individuals with overlapping areas are likely to forage together. Our study confirms previous findings that Bechstein's bats show high fidelity to foraging areas across years. Moreover, we found that relatives share foraging areas significantly more often compared with unrelated colony members. Finally, our data reveal for the first time that most colony members that share foraging areas are unlikely to forage together. This suggests that female Bechstein's bats gain no direct benefits from sharing foraging areas with members of the same maternal lineage. Our findings also have implications for conservation as habitat loss within a colony's home range might expose entire matrilines to high risks.     

Frequency-dependent selection on information-transfer strategies at breeding colonies: a simulation study

Through computer simulations, we model three different foodfinding strategies: searcher, no information transfer, watcher,limited information transfer; follower, full information transfer.The aim of this article was to study how frequency-dependentselection affects the proportion of these strategies at a simulatedcolony under different patterns of food distribution. Furthermore,we determined how information transfer in a population witha mixed evolutionarily stable strategy (ESS) modified the averageforaging efficiency of an individual compared to that of anindividual in a population with mutual information exchange.We found that the proportion of information gaining strategiesincreases as the food resources become more clumped. The improvementin foraging efficiency through the operation of an informationcenter need not require mutuality in information exchange. Onthe basis of the presented study, at the ESS only a small percentageof colony members need discover food patches, yet the foragingefficiency may be high because of the operation of an informationcenter.     

Tolerant food sharing and reciprocity is precluded by despotism among bonobos but not chimpanzees

Tolerant food sharing among human foragers can largely be explained by reciprocity. In contrast, food sharing among chimpanzees and bonobos may not always reflect reciprocity, which could be explained by different dominance styles: in egalitarian societies reciprocity is expressed freely, while in more despotic groups dominants may hinder reciprocity. We tested the degree of reciprocity and the influence of dominance on food sharing among chimpanzees and bonobos in two captive groups. First, we found that chimpanzees shared more frequently, more tolerantly, and more actively than bonobos. Second, among chimpanzees, food received was the best predictor of food shared, indicating reciprocal exchange, whereas among bonobos transfers were mostly unidirectional. Third, chimpanzees had a shallower and less linear dominance hierarchy, indicating that they were less despotic than bonobos. This suggests that the tolerant and reciprocal sharing found in chimpanzees, but not bonobos, was made possible by the absence of despotism. To investigate this further, we tested the relationship between despotism and reciprocity in grooming using data from an additional five groups and five different study periods on the main groups. The results showed that i) all chimpanzee groups were less despotic and groomed more reciprocally than bonobo groups, and ii) there was a general negative correlation between despotism and grooming reciprocity across species. This indicates that an egalitarian hierarchy may be more common in chimpanzees, at least in captivity, thus fostering reciprocal exchange. We conclude that a shallow dominance hierarchy was a necessary precondition for the evolution of human‐like reciprocal food sharing. Am J Phys Anthropol 143:41–51, 2010. © 2010 Wiley‐Liss, Inc.     

Out of Africa: origins of the Taenia tapeworms in humans

Phylogenetic and divergence date analyses indicate that the occurrence of Taenia tapeworms in humans pre-dates the development of agriculture, animal husbandry and domestication of cattle (Bos spp.) or swine (Sus scrofa). Taeniid tapeworms in Africa twice independently colonized hominids and the genus Homo prior to the origin of modern humans. Dietary and behavioural shifts, from herbivory to scavenging and carnivory, as early Homo entered the carnivore guild in the Pliocene/Pleistocene, were drivers for host switching by tapeworms to hominids from carnivores including hyaenids and felids. Parasitological data provide a unique means of elucidating the historical ecology, foraging behaviour and food habits of hominids during the diversification of Homo spp.