The evolutionary and ecological roots of human social organization

Social organization among human foragers is characterized by a three-generational system of resource provisioning within families, long-term pair-bonding between men and women, high levels of cooperation between kin and non-kin, and relatively egalitarian social relationships. In this paper, we suggest that these core features of human sociality result from the learning- and skill-intensive human foraging niche, which is distinguished by a late age-peak in caloric production, high complementarity between male and female inputs to offspring viability, high gains to cooperation in production and risk-reduction, and a lack of economically defensible resources. We present an explanatory framework for understanding variation in social organization across human societies, highlighting the interactive effects of four key ecological and economic variables: (i) the role of skill in resource production; (ii) the degree of complementarity in male and female inputs into production; (iii) economies of scale in cooperative production and competition; and (iv) the economic defensibility of physical inputs into production. Finally, we apply this framework to understanding variation in social and political organization across foraging, horticulturalist, pastoralist and agriculturalist societies.     

An individual-orientated model of the emergence of despotic and egalitarian societies

Single behavioural differences between egalitarian and despotic animal societies are often assumed to reflect specific adaptations. However, in the present paper, I will show in an individual-orientated model, how many behavioural traits of egalitarian and despotic virtual societies arise as emergent characteristics. The artificial entities live in a homogeneous world and only aggregate, and upon meeting one another and may perform dominance interactions in which the effects of winning and losing are self-reinforcing. The behaviour of these entities is studied in a similar way to that of real animals. It will be shown that by varying the intensity of aggression only, one may switch from egalitarian to despotic virtual societies. Differences between the two types of society appear to correspond closely to those between despotic and egalitarian macaque species in the real world. In addition, artificial despotic societies show a clearer spatial centrality of dominants and, counter-intuitively, more rank overlap between the sexes than the egalitarian ones. Because of the correspondence with patterns in real animals, the model makes it worthwhile comparing despotic and egalitarian species for socio-spatial structure and rank overlap too. Furthermore, it presents us with parsimonious hypotheses which can be tested in real animals for patterns of aggression, spatial structure and the distribution of social positive and sexual behaviour.     

Behavioural ecology in traditional societies

Can models from behavioural ecology explain cultural diversity in human populations? Studies of variation in reproductive and productive behaviour, both within and between traditional societies, are beginning to show that specific predictions from sexual selection and optimal foraging theory can be developed and tested with human data. Greatest success has been in the study of foraging; whereas attempts to understand patterns of marriage and parental investment have been most convincing in those cases where behaviour is related to specific ecological and social conditions. The aim of human behavioural ecologists in the future will be to determine the constraints that the dual goals of reproduction and production place on individuals.     

The spread of inequality

The causes of socioeconomic inequality have been debated since the time of Plato. Many reasons for the development of stratification have been proposed, from the need for hierarchical control over large-scale irrigation systems to the accumulation of small differences in wealth over time via inheritance processes. However, none of these explains how unequal societies came to completely displace egalitarian cultural norms over time. Our study models demographic consequences associated with the unequal distribution of resources in stratified societies. Agent-based simulation results show that in constant environments, unequal access to resources can be demographically destabilizing, resulting in the outward migration and spread of such societies even when population size is relatively small. In variable environments, stratified societies spread more and are also better able to survive resource shortages by sequestering mortality in the lower classes. The predictions of our simulation are provided modest support by a range of existing empirical studies. In short, the fact that stratified societies today vastly outnumber egalitarian societies may not be due to the transformation of egalitarian norms and structures, but may instead reflect the more rapid migration of stratified societies and consequent conquest or displacement of egalitarian societies over time.     

Mechanisms of sexual egalitarianism in Western Europe

This paper presents historical evidence on marriage patterns in ancient Sparta, Rome, Early Christianity, and the Early Middle Ages. Monogamy occurred in all of these societies but there is a great deal of diversity in origin and function of monogamous mating arrangements. In the case of Sparta, monogamy arose as part of an intensively egalitarian, racially homogenous social structure which fostered intense cooperation and altruism within the group. In the case of Rome monogamy coexisted with pronounced social, political, and economic inequalities, and there was much more ethnic diversity at Rome than at Sparta. The case of early Christianity involved the spread of a more radical ideology of monogamy and sexual restraint among the lower and middle classes of the Roman Empire, but the crucial event in the Christianization of the West was the apparently chance conversion of a single powerful individual, the Emperor Constantine. In the case of the Christianization of barbarian Europe, the movement was spearheaded by a powerful institution and the acceptance among the aristocracy of Christian ideology. The revolution thus proceeded from the top of the society downward. These findings are related to a model of cultural evolution that emphasizes the irreducibility of social controls and ideology in maintaining egalitarian mating practices.     

Is The Premise Of Egalitarianism Inequality?

The article focusses on two questions. One is the conditions under which one might expect to find community level egalitarian ideology in complex societies. The other is a more specific concern with the occurrence of egalitarian ideology among contemporary small scale hunting and gathering societies. A common lower class situation is critical when this translates into community acceptance of the overriding legitimacy of equalisation of status within the community. Also important is the location of the community in opposition to the wider society which it sees as external, threatening, dominating and intrusive.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

Parallel tool industries in New Caledonian crows

Individual specialization in the use of foraging tools occurs in hunter-gatherer societies but is absent in non-human primate tool use. 'Parallel tool industries' in hunter-gatherers are mainly based on strict sexual division of labour that is highly reliant on social conformity. Here, we show that 12 individuals in a population of New Caledonian crows on Maré Island had strong preferences for either stick tools or pandanus tools. Eight of the 12 crows had exclusive preferences. The individual specialization that we found is probably associated with different foraging niches. However, in spite of sexual size dimorphism there was no significant association between the sex of crows and their tool preferences. Our findings demonstrate that highly organized, strict sexual division of labour is not a necessary prerequisite for the evolution of parallel tool industries.     

Sexual dimorphism and sexual segregation in foraging strategies of northern giant petrels, Macronectes halli, during incubation

Giant petrels ( Macronectes spp.) are the most sexually dimorphic of all seabirds. We used satellite-tracking and mass change during incubation to investigate the influence of sexual size dimorphism, in terms of the intersexual food competition hypothesis, on foraging and fasting strategies of northern giant petrels at South Georgia. Females foraged at sea whereas males foraged mainly on the South Georgia coast, scavenging on seal and penguin carcasses. Foraging effort (flight speed, distance covered, duration of foraging trips) was greater for females than for males. In contrast, foraging efficiency (proportionate daily mass gain while foraging) was significantly greater for males than for females. Females were significantly closer to the desertion mass threshold than males and could not compensate for the mass loss during the incubation fast while foraging, suggesting greater incubation costs for females than for males. Both sexes regulated the duration and food intake of foraging trips depending on the depletion of the body reserves. In males the total mass gain was best explained by mass at departure and body size. We suggest that sexual segregation of foraging strategies arose from size-related dominance at carcasses, promoting sexual size dimorphism. Our results indicate that sex-specific differences in fasting endurance, contest competition over food and flight metabolic rates are key elements in maintenance of sexual size dimorphism, segregating foraging strategies and presumably reducing competition between sexes.     

The Fertility of Melanesian Girls, Laboratory Mice, and Prostitutes: A Comment on the "Bruce Effect"

Greenfield has recently proposed that low fertility in both prostitutes and unmarried Trobriand girls may be explained by the "Bruce effect": the inhibition of pregnancy in female mice that have had intercourse with different males within a short period. An examination of Malinowski's data indicates that a Trobriand girl usually had intercourse with only one man over a fairly long period and her sex life was not at all comparable to that of a prostitute or one of Bruce's mice. Low fertility in unmarried girls has been well documented for several Melanesian societies that differ notably in the behavior permitted to girls. The existence of so-called adolescent sterility still seems to be the best explanation to cover all of these cases. [fertility, sexual behavior, Melanesia]     

Variation in foraging behavior facilitates resource partitioning in a polymorphic cichlid, Herichthys minckleyi

We examined foraging behavior (microhabitat use and feeding behavior) in a trophically polymorphic cichlid fish, Herichthys minckleyi, to address several questions regarding resource partitioning in this threatened species. These include: (1) do morphotypes demonstrate different foraging behaviors? (2) do individuals within a morphotype vary in their foraging behavior (e.g. are some individuals specialists, only using a subset of available resources, while other are generalists)? (3) do foraging behaviors vary between isolated pools? (4) do foraging behaviors vary across seasons? We quantified microhabitat use and feeding behavior for over 100 individuals (of two morphotypes) feeding freely in two isolated pools (populations) and across two seasons (winter and summer). We found differences in foraging behavior between morphotypes and individual specializations within morphotypes; i.e. some individuals specialize on certain food resources by using a few feeding behaviors within a subset of microhabitats, whereas others employ a range feeding behaviors across many microhabitats. Foraging behavior also varied between pools and across seasons. This spatial and temporal variation in foraging behavior and resource use may serve to maintain this polymorphism, as the relative fitness of the each morph may vary over space and time.     

The functions of societies and the evolution of group living: spider societies as a test case

Many models have been advanced to suggest how different expressions of sociality have evolved and are maintained. However these models ignore the function of groups for the particular species in question. Here we present a new perspective on sociality where the function of the group takes a central role. We argue that sociality may have primarily a reproductive, protective, or foraging function, depending on whether it enhances the reproductive, protective or foraging aspect of the animal's life (sociality may serve a mixture of these functions). Different functions can potentially cause the development of the same social behaviour. By identifying which function influences a particular social behaviour we can determine how that social behaviour will change with changing conditions, and which models are most pertinent. To test our approach we examined spider sociality, which has often been seen as the poor cousin to insect sociality. By using our approach we found that the group characteristics of eusocial insects is largely governed by the reproductive function of their groups, while the group characteristics of social spiders is largely governed by the foraging function of the group. This means that models relevant to insects may not be relevant to spiders. It also explains why eusocial insects have developed a strict caste system while spider societies are more egalitarian. We also used our approach to explain the differences between different types of spider groups. For example, differences in the characteristics of colonial and kleptoparasitic groups can be explained by differences in foraging methods, while differences between colonial and cooperative spiders can be explained by the role of the reproductive function in the formation of cooperative spider groups. Although the interactions within cooperative spider colonies are largely those of a foraging society, demographic traits and colony dynamics are strongly influenced by the reproductive function. We argue that functional explanations help to understand the social structure of spider groups and therefore the evolutionary potential for speciation in social spiders.     

Consistency pays: sex differences and fitness consequences of behavioural specialization in a wide-ranging seabird

Specialists and generalists often coexist within a single population, but the biological drivers of individual strategies are not fully resolved. When sexes differ in their foraging strategy, this can lead them to different environmental conditions and stability across their habitat range. As such, sexual segregation, combined with dominance, may lead to varying levels of specialization between the sexes. Here, we examine spatial and temporal niche width (intraindividual variability in aspects of foraging behaviour) of male and female black-browed albatrosses (Thalassarche melanophrys), and its consequences for fitness. We show that females, where maximum foraging range is under fluctuating selection, exhibit more variable behaviours and appear more generalist than males, who are under directional selection to forage close to the colony. However within each sex, successful birds had a much narrower niche width across most behaviours, suggesting some specialization is adaptive in both sexes. These results demonstrate that while there are sex differences in niche width, the fitness benefit of specialization in spatial distribution is strong in this wide-ranging seabird.     

Three millennia of human and sea turtle interactions in Remote Oceania

Sea turtles are one of the largest vertebrates in the shallow water ecosystems of Remote Oceania, occurring in both sea grass pastures and on coral reefs. Their functional roles, however, over ecological and evolutionary times scales are not well known, in part because their numbers have been so drastically reduced. Ethnographic and archaeological data is analysed to assess long-term patterns of human–sea turtle interactions (mainly green and hawksbill) prior to western contact and the magnitude of turtle losses in this region. From the ethnographic data two large-scale patterns emerge, societies where turtle capture and consumption was controlled by chiefs and priests versus those where control over turtle was more flexible and consumption more egalitarian. Broadly the distinction is between societies on high (volcanic and raised coral) islands versus atolls, but the critical variables are the ratio of land to shallow marine environments, combined with the availability of refugia. Archaeological evidence further highlights differences in the rate and magnitude of turtle losses across these two island types, with high islands suffering both large and rapid declines while those on atolls are less marked. These long-term historical patterns help explain the ethnographic endpoints, with areas that experienced greater losses apparently developing more restrictive social controls over time. Finally, if current turtle migration patterns held in the past, with annual movements between western foraging grounds and eastern nesting beaches, then intensive harvesting from 2,800 Before Present in West Polynesia probably affected turtle abundance and coral reef ecology in East Polynesia well before the actual arrival of human settlers, the latter a process that most likely began 1,400 years later.     

(Evolutionary) Theories of warfare in preindustrial (foraging) societies

I present an inventory of theories of war causation (and on the origin of war) in preindustrial (traditional, foraging, 'primitive', hunter-gatherer, band- and tribe-level) societies, with emphasis on the roles of natural selection, sexual selection and kin selection. Also the school of sociocultural evolution is briefly discussed.     

Cross-site differences in foraging behavior of white-faced capuchins (Cebus capucinus)

Researchers have identified a variety of cross-site differences in the foraging behavior of free-ranging great apes, most notably among chimpanzees (Pan troglodytes) and more recently orangutans (Pongo pygmaeus), that are not due to obvious genetic or ecological differences. These differences are often referred to as "traditions." What is not known is whether this high level of interpopulation variation in behavior is limited to hominoids. In this study, we use long-term data from three Costa Rican field sites that are geographically close and similar ecologically to identify potential foraging traditions in white-faced capuchins (Cebus capucinus). Foraging traditions are predicted in Cebus because of many behavioral and morphological convergences between this genus and the great apes. The processing techniques used for the same food species were compared across sites, and all differences found were classified as present, habitual, or customary. Proximity data were also analyzed to determine if social learning mechanisms could explain variation in foraging behavior. Of the 61 foods compared, we found that 20 of them are processed differently by capuchins across sites. The differences involve pound, rub, tap, "fulcrum," "leaf-wrap," and "army ant following." For most of the differences with enough data to analyze, the average proximity score of the "matched" dyads (two individuals within a group who shared a "different" processing technique) was statistically higher than the average proximity score of the remaining "unmatched" dyads.     

Farewell to the ‘childhood of man’: ritual,seasonality, and the origins of inequality

Evidence of grand burials and monumental construction is a striking feature in the archaeological record of the Upper Palaeolithic period, between 40 and 10 kya (thousand years ago). Archaeologists often interpret such finds as indicators of rank and hierarchy among Pleistocene hunter‐gatherers. Interpretations of this kind are difficult to reconcile with the view, still common in sociobiology, that pre‐agricultural societies were typically egalitarian in orientation. Here we develop an alternative model of ‘Palaeolithic politics’, which emphasizes the ability of hunter‐gatherers to alternate – consciously and deliberately – between contrasting modes of political organization, including a variety of hierarchical and egalitarian possibilities. We propose that alternations of this sort were an emergent property of human societies in the highly seasonal environments of the last Ice Age. We further consider some implications of the model for received concepts of social evolution, with particular attention to the distinction between ‘simple’ and ‘complex’ hunter‐gatherers.     

Gene flow across species boundaries in sympatric, sexually deceptive Ophrys (Orchidaceae) species

Abstract Orchids of the genus Ophrys (Orchidaceae) are pollinated by male bees and wasps through sexual deception. The Ophrys sphegodes group encompasses several closely related species that differ slightly in floral morphology and are pollinated by different solitary bee species. Populations representing different species of the O. sphegodes group often flower simultaneously in sympatry. To test whether gene flow across the species boundaries occurs in these sympatric populations, or whether they are reproductively isolated, we examined the distribution of genetic variation within and among populations and species of this group. We collected at each of five different localities in southern France and Italy two sympatric, co-flowering Ophrys populations, representing six Ophrys species in total. The six microsatellite loci surveyed were highly variable. Genetic differentiation among geographically distant populations of the same species was lower than differentiation among sympatric populations of different species. However, the strength of genetic differentiation among species was among the lowest reported for orchids. Genotype assignment tests and marker-based estimates of gene flow revealed that gene flow across species boundaries occurred and may account for the low observed differentiation among species. These results suggest that sexual deceit pollination in Ophrys may be less specific than thought, or that rare mistakes occur.     

Family feuds: social competition and sexual conflict in complex societies

Darwin was initially puzzled by the processes that led to ornamentation in males-what he termed sexual selection-and those that led to extreme cooperation and altruism in complex animal societies-what was later termed kin selection. Here, I explore the relationships between sexual and kin selection theory by examining how social competition for reproductive opportunities-particularly in females-and sexual conflict over mating partners are inherent and critical parts of complex altruistic societies. I argue that (i) patterns of reproductive sharing within complex societies can drive levels of social competition and reproductive conflict not only in males but also in females living in social groups, and ultimately the evolution of female traits such as ornaments and armaments; (ii) mating conflict over female choice of sexual partners can influence kin structure within groups and drive the evolution of complex societies; and (iii) patterns of reproductive sharing and conflict among females may also drive the evolution of complex societies by influencing kin structure within groups. Ultimately, complex societies exhibiting altruistic behaviour appear to have only arisen in taxa where social competition over reproductive opportunities and sexual conflict over mating partners were low. Once such societies evolved, there were important selective feedbacks on traits used to regulate and mediate intra-sexual competition over reproductive opportunities, particularly in females.     

Foraging effort, food intake, fat deposition and puberty in female mice

A novel caging system was used to study the interrelationships between foraging effort, food intake, growth and sexual maturation of peripubertal female mice. Females housed in these cages were forced to work (forage) at various intensities in order to obtain food pellets. It is argued that this is a biologically more meaningful approach to understanding the energetics of sexual development than the traditional approach of simple underfeeding. Female mice exhibited a cascade of developmental adjustments and deficits when challenged to forage harder for less food. The functions most sensitive to increased foraging effort were sexual development and growth in body length; growth in body weight was intermediate and fat deposition was least sensitive of all. The relative insensitivity of fat deposition to higher foraging costs suggests a strategy for survival during the postweaning dispersal movements of the wild ancestors of the laboratory mouse. Finally, regression analyses suggested that heavier females who had less than average body fat and higher than average food intake achieved their pubertal ovulation most rapidly.