Leadership solves collective action problems in small-scale societies

Observation of leadership in small-scale societies offers unique insights into the evolution of human collective action and the origins of sociopolitical complexity. Using behavioural data from the Tsimane forager-horticulturalists of Bolivia and Nyangatom nomadic pastoralists of Ethiopia, we evaluate the traits of leaders and the contexts in which leadership becomes more institutional. We find that leaders tend to have more capital, in the form of age-related knowledge, body size or social connections. These attributes can reduce the costs leaders incur and increase the efficacy of leadership. Leadership becomes more institutional in domains of collective action, such as resolution of intragroup conflict, where collective action failure threatens group integrity. Together these data support the hypothesis that leadership is an important means by which collective action problems are overcome in small-scale societies.     

Variation in Swidden practices in four central Brazilian Indian societies

It has been suggested that indigenous tropical forest agriculture, in contrast with other agricultural systems, is characterized by a high degree of polyculture, and, being similar in diversity to the forest itself, has little disturbing effect on the generalized ecosystem that surrounds native gardens. Our comparative study of four Central Brazilian Indian groups shows, however, that while each of these groups practices polyculture to some extent, the crop mix found in their swidden plots is highly patterned, and includes single crop stands at certain stages of garden life. Different crops are planted in the same swidden plot from year to year, in accordance with variation in soil fertility within the swidden cycle. None of the swiddens we observed compares in complexity to the surrounding forest. We suggest that polyculture, rather than being regarded as the distinguishing characteristic of swidden cultivation, should be considered as a varying dimension—along with other variables such as use of wild plants, soil fertility and exhaustion, tillage practices, the introduction of novel crops, and production for the market—within the overall subsistence strategy of a group.     

The use of AFLPs to examine genetic relatedness in barley

The generation of AFLPs in spring barley cultivars provided genetic information relating to the development of the crop in the UK since 1953. Principal co-ordinate (PCO) analysis of genetic similarities (gs) confirmed the marked contrast in the cultivars used in the 1970s and 1980s. The earliest cultivars, many derived from Proctor, were succeeded by tall-strawed, disease-resistant types with high yield but poor malting potential. In the 1980s they were in turn replaced by short-strawed cultivars with excellent yield and good malting quality, which originated from Triumph. A PCO plot of gs provided insight into the effects of selection for disease resistance and the antagonism between malting quality and particular resistance genes. The analysis of gs was more useful than pedigrees and estimates of kinship in revealing the genetic relationship between cultivars. Theoretical considerations for maximising the efficiency of an AFLP genotyping programme are discussed in the context of the number of primer pairs required to distinguish genotypes at varying levels of similarity.     

Nutritional consequences of the transformation from subsistence to commercial agriculture in Tabasco,Mexico

Recent changes in the area of the Plan Chontalpa in Tabasco have greatly reduced the production of subsistence crops by rural families, resulting in decreased crop diversity and a concomitant increase in the degree of dependence on outside sources of food. Results from a nutrition survey of 149 families demonstrate that dietary diversity, dietary quality, and nutritional status of preschool children are negatively associated with lower crop diversity and increased dependence on purchased foods. Dietary deterioration is illustrated by the negative relationship found between nutritional status and increased sugar consumption. The assumption that a rise in income accompanying the adoption of commercial production will automatically lead to improved nutrition is challenged: income levels were not found to be consistently related to nutritional status. Children of families that have converted to cattle production, despite greater land availability and family incomes, do not have improved nutritional status. In the study area, where wages are low and food prices are very high, the value of a higher degree of self-sufficiency in food is recognized, yet families continue to switch to cash crops due to the environmental, economic, and time constraints imposed by the system of commercial agriculture in which they participate. The solution is not to return to traditional subsistence farming, however, but to determine under what conditions a more progressive form of agricultural change can occur.     

Genetic relatedness in wintering groups of house sparrows (Passer domesticus)

Social behaviour of group-living animals is often influenced by the relatedness of individuals, thus understanding the genetic structure of groups is important for the interpretation of costs and benefits of social interactions. In this study, we investigated genetic relatedness in feeding aggregations of free-living house sparrows ( Passer domesticus ) during the nonbreeding season. This species is a frequent model system for studies of social behaviour (e.g. aggression, social foraging), but we lack adequate information on the kin structure of sparrow flocks. During two winters, we ringed and observed sparrows at feeding stations, and used resightings to identify stable flock-members and to calculate association indices between birds. We genotyped the birds using seven highly polymorphic microsatellite loci, and estimated pairwise relatedness coefficients and relatedness categories (close kin vs. unrelated) by maximum likelihood method. We found that most birds were unrelated to each other in the flocks (mean ± SE relatedness coefficient: 0.06 ± 0.002), although most individuals had at least a few close relatives in their home flock (14.3 ± 0.6% of flock-mates). Pairwise association between individuals was not significantly related to their genetic relatedness. Furthermore, there was no difference between within-flock vs. between-flock relatedness, and birds had similar proportions of close kin within and outside their home flock. Finally, relatedness among members of different flocks was unrelated to the distance between their flocks. Thus, sparrow flocks were not characterized by association of relatives, nevertheless the presence of some close kin may provide opportunity for kin-biased behaviours to evolve.     

Behavioural variation in 172 small-scale societies indicates that social learning is the main mode of human adaptation

The behavioural variation among human societies is vast and unmatched in the animal world. It is unclear whether this variation is due to variation in the ecological environment or to differences in cultural traditions. Underlying this debate is a more fundamental question: is the richness of humans’ behavioural repertoire due to non-cultural mechanisms, such as causal reasoning, inventiveness, reaction norms, trial-and-error learning and evoked culture, or is it due to the population-level dynamics of cultural transmission? Here, we measure the relative contribution of environment and cultural history in explaining the behavioural variation of 172 Native American tribes at the time of European contact. We find that the effect of cultural history is typically larger than that of environment. Behaviours also persist over millennia within cultural lineages. This indicates that human behaviour is not predominantly determined by single-generation adaptive responses, contra theories that emphasize non-cultural mechanisms as determinants of human behaviour. Rather, the main mode of human adaptation is social learning mechanisms that operate over multiple generations.     

The nutritional consequences of rejecting food in the Ituri Forest of Zaire

Based on a representative sample of 449 different individuals, food avoidances are shown to be nutritionally insignificant on average for a population of horticulturalists and foragers living in the Ituri Forest of northeastern Zaire. However, due to variability in food-related beliefs, nutritional costs may be biologically significant for a few individuals. Less than 2% of potential dietary calories are restricted to horticulturalists on average due to food avoidances, while less than 1% is the average cost of these beliefs among foragers. When nutritional costs are calculated from reported avoidance rules rather than consumption practice, estimated costs are significantly inflated. These lower actual costs arise because individuals make use of rules that permit them to eat otherwise restricted foods. For example, eating specific plants is believed to prevent the illness that would otherwise follow the consumption of a tabooed food. Use of such exception rules significantly reduces the nutritional cost of food avoidances for many individuals in this population.     

General patterns of niche construction and the management of 'wild' plant and animal resources by small-scale pre-industrial societies

Niche construction efforts by small-scale human societies that involve 'wild' species of plants and animals are organized into a set of six general categories based on the shared characteristics of the target species and similar patterns of human management and manipulation: (i) general modification of vegetation communities, (ii) broadcast sowing of wild annuals, (iii) transplantation of perennial fruit-bearing species, (iv) in-place encouragement of economically important perennials, (v) transplantation and in-place encouragement of perennial root crops, and (vi) landscape modification to increase prey abundance in specific locations. Case study examples, mostly drawn from North America, are presented for each of the six general categories of human niche construction. These empirically documented categories of ecosystem engineering form the basis for a predictive model that outlines potential general principles and commonalities in how small-scale human societies worldwide have modified and manipulated their 'natural' landscapes throughout the Holocene.     

Mating frequency and genetic relatedness of workers in the hornet Vespa analis (Hymenoptera: Vespidae)

Mating frequency of Vespa analis queens and the genetic relatedness of their workers was analyzed by DNA microsatellite genotyping. Of 20 colonies studied, 18 had a queen inseminated by a single male and two had queens each inseminated by two males. The estimated effective number of matings was 1.05 ± 0.037 (mean ± SE), with 75–85% of the offspring of the two multiply mated queens sired by a single male. The pedigree relatedness between nestmate workers averaged over the 20 colonies was estimated to be 0.74 ± 0.008, almost identical to the predicted value of 0.75 for colonies headed by a singly mated queen. Multiple matrilines; that is, the presence of workers not related to the current queens, were detected in six colonies, suggesting that queen replacement occurred via usurpation of the founding queens in these six colonies. These results demonstrate that the kin structure of V. analis is similar to that reported in other vespid species.     

Genetic relatedness in early associations of Polistes dominulus: from related to unrelated helpers

Indirect benefits obtained through the reproduction of relatives are fundamental in the formation and maintenance of groups. Here, we examine the hypothesis that females of the temperate paper wasp Polistes dominulus preferentially form groups with close relatives. Genetic relatedness data were obtained for 180 groups of females collected at the early stages of the nesting cycle of a large population of P. dominulus in two sites in southwestern Spain. Average within-group relatedness values ranged from 0.189 to 0.491. Foundresses on early nests were significantly more closely related than females in winter aggregations or in stable groups (just before workers emerged). Within-group relatedness values were independent of group size. The vast majority of worker-producing nests ( c . 85%) had one or more females that were unrelated (or distantly related) to the remaining members of the group. These results provide further support to the hypothesis that indirect fitness benefits alone are unlikely to explain why P. dominulus foundresses form cooperative associations.     

Implications of multiple mating for offspring relatedness and shoaling behaviour in juvenile guppies

Polyandry (female multiple mating) can confer important benefits to females, but few studies have considered its potential costs. One such cost may arise through differences in the relatedness of offspring born to females with different mating histories; offspring born to monandrous females are always full siblings, while those produced by polyandrous females may be full or half siblings. These differences may have important consequences for social interactions among offspring. We used artificial insemination in the guppy (Poecilia reticulata), a promiscuous live-bearing fish, to evaluate shoaling behaviour in polyandrous and monandrous broods. We combined this information with known parentage data for the polyandrous broods to determine whether sibling relatedness influenced offspring shoaling behaviour. While we detected no effect of mating treatment (polyandry/monandry) on shoaling behaviour, we found that pairs of full siblings spent significantly more time shoaling (and in close proximity) than pairs of half siblings. This latter finding confirms the ability of newborn guppies to distinguish brood mates on the basis of kinship, but also suggests an important and hitherto unrealized potential cost of polyandry: a reduction in within-brood relatedness with potentially important implications for offspring social behaviour.     

Fitness consequences of ovicide in a parasitoid wasp

Ovicide, superparasitism and host rejection are alternative reproductive tactics facing female parasitoid wasps encountering an already-parasitized host. Superparasitism is simply the addition of an egg or a clutch of eggs by the secondary parasitoid, but under ovicide the primary clutch is removed or destroyed. Host rejection occurs if the wasp leaves without laying a clutch. The ectoparasitoid Laelius pedatus (Say) (Hymenoptera: Bethylidae) performs ovicide in this situation. Clutch manipulation experiments show that secondary clutches suffer high mortality in competition with primary clutches, which increases with increasing time delay between clutches. Primary clutches however suffer little in competition with secondary clutches, even if there is minimal time delay between clutches. These data suggest that the offspring of ovicidal females are substantially fitter than the offspring of superparasitizing females. Handling time and clutch size do not differ significantly between first (sole) parasitoids and second (ovicidal) parasitoids. The same is true for offspring survival and development time. However, offspring of second females are slightly smaller. This suggests that parasitized and unparasitized hosts are resources of similar quality when ovicide is performed. These data strongly support the predictions of evolutionary models of ovicide. They may also give some insight into the taxonomic distribution of ovicide in parasitoids.     

Fitness consequences of centrality in mutualistic individual-based networks

The relationships among the members of a population can be visualized using individual networks, where each individual is a node connected to each other by means of links describing the interactions. The centrality of a given node captures its importance within the network. We hypothesize that in mutualistic networks, the centrality of a node should benefit its fitness. We test this idea studying eight individual-based networks originated from the interaction between Erysimum mediohispanicum and its flower visitors. In these networks, each plant was considered a node and was connected to conspecifics sharing flower visitors. Centrality indicates how well connected is a given E. mediohispanicum individual with the rest of the co-occurring conspecifics because of sharing flower visitors. The centrality was estimated by three network metrics: betweenness, closeness and degree. The complex relationship between centrality, phenotype and fitness was explored by structural equation modelling. We found that the centrality of a plant was related to its fitness, with plants occupying central positions having higher fitness than those occupying peripheral positions. The structural equation models (SEMs) indicated that the centrality effect on fitness was not merely an effect of the abundance of visits and the species richness of visitors. Centrality has an effect even when simultaneously accounting for these predictors. The SEMs also indicated that the centrality effect on fitness was because of the specific phenotype of each plant, with attractive plants occupying central positions in networks, in relation to the distribution of conspecific phenotypes. This finding suggests that centrality, owing to its dependence on social interactions, may be an appropriate surrogate for the interacting phenotype of individuals.     

Families in space: relatedness in the Barents Sea population of polar bears (Ursus maritimus)

The kin structure and dispersal pattern of polar bears ( Ursus maritimus ) of the Barents Sea was investigated during the spring mating season using two complementary approaches. First, individual genotypes based on the analyses of 27 microsatellite loci of 583 polar bears were related to field information gathered from 1146 bears in order to reconstruct the animals' pedigrees and to infer geographical distances between adult bears of different relatedness categories. According to the data, the median natal dispersal distance of the male animals was 52 km while that of the females was 93 km. Second, the relatedness of pairs of adult bears was estimated and correlated to the geographical distance between them. The female dyads had a much stronger kin structure than the male dyads. The 'pedigree approach' revealed a male kin structure which could not be detected using the 'relatedness approach'. This suggests that, on a broader scale, effective dispersal is slightly male biased. Despite fidelity to natal areas, male-mediated gene flow may nevertheless prevent genetic differentiation. Males might occasionally shift their home range which could therefore lead to a male-biased breeding dispersal. Our results showed that a nonterritorial species such as the polar bear that has a high dispersal potential, lives in a highly unstable environment and migrates seasonally is still able to exhibit a distinct kin structure during the mating season.     

Children's Help and the Pace of Reproduction: Cooperative Breeding in Humans

Because children's ability to support themselves falls below their consumption, human young are subsidized by others throughout much of their growth and development. Mothers, however, who often have multiple dependents of different ages, are faced with an allocation problem (Fig. 1). This has led to important debate about the evolution of a long period dependence and the development of nonmaternal strategies to provision young. This article focuses on the critical role that children themselves play. Because the human subsistence niche incorporates a broad diversity of resources that require variable procurement and processing costs, dependent children can also be important producers, furthering both a need and an opportunity for cooperative breeding.     

Reproductive skew and relatedness in social groups of European badgers, Meles meles

Reproductive skew is a measure of the proportion of individuals of each sex that breed in a group and is a valuable measure for understanding the evolution and maintenance of sociality. Here, we provide the first quantification of reproductive skew within social groups of European badgers Meles meles , throughout an 18-year study in a high-density population. We used 22 microsatellite loci to analyse within-group relatedness and demonstrated that badger groups contained relatives. The average within-group relatedness was high ( R =  0.20) and approximately one-third of within-group dyads were more likely to represent first-order kin than unrelated pairs. Adult females within groups had higher pairwise relatedness than adult males, due to the high frequency of extra-group paternities, rather than permanent physical dispersal. Spatial clustering of relatives occurred among neighbouring groups, which we suggest was due to the majority of extra-group paternities being attributable to neighbouring males. Reproductive skew was found among within-group candidate fathers ( B  = 0.26) and candidate mothers ( B  = 0.07), but not among breeding individuals; our power to detect skew in the latter was low. We use these results to evaluate reproductive skew models. Although badger society best fits the assumptions of the incomplete-control models, our results were not consistent with their predictions. We suggest that this may be due to female control of paternity, female–female reproductive suppression occurring only in years with high food availability resulting in competition over access to breeding sites, extra-group paternity masking the benefits of natal philopatry, and/or the inconsistent occurrence of hierarchies that are linear when established.     

Fitness consequences of male parental care in Savannah sparrows

The fitness consequences of male parental care are central tolife-history models of reproductive effort and the evolutionof social and genetic mating system However in a 4-year study(1992–1995) of Savannah sparrows (Parsserculs sandwichensis)nesting on Kent Island, New Brunswick, Canada, male parentalcare during the nestling stage was only weakly related to offspringquality and survivorship. Females compensated for poor paternalperformance by elevating their feeding rates, and they thusameliorated selection on male parental care. The likelihoodand size of subsequent broods was also unrelated to male feedingeffort, suggesting the absence of some indirect (or ‘female-mediated’)benefits. These results were confirmed by an experiment in whichmale removal resulted in increased female feeding rates buthad little effect on the number or quality of young. In contrast,previous research with multilocus DNA fingerpnnting revealeda strong, positive relationship between a male's parental effortand his subsequent fertilization success. Together, this researchsuggests that the proximate and ultimate targets of male feedingeffort can be disiinct traditional assumptions regarding thefunction of male parental care in birds may therefore be incomplete.     

A social learning model of conflict and cooperation in human societies

Patterns of conflict and cooperation both within and between societies may be related to the degree of cultural similarity within and between the same societies. A simple model of social learning is used to predict patterns of conflict and cooperation in hypothetical societies that differ in the roles of relatives and nonrelatives in the enculturation of children. The model is illustrated by comparing its predictions to known differences in the patterns of conflict between males inpatrilocal and matrilocal societies.