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.     

Modeling neolithic dispersal in central Europe: demographic implications

On the basis of new examination of ancient DNA and craniometric analyses, Neolithic dispersal in Central Europe has been recently explained as reflecting colonization or at least a major influx of near eastern farmers. Given the fact that Neolithic dispersal in Central Europe was very rapid and extended into a large area, colonization would have to be associated with high population growth and fertility rates of an expanding Neolithic population. We built three demographic models to test whether the growth and fertility rates of Neolithic farmers were high enough to allow them to colonize Central Europe without admixture with foragers. The principle of the models is based on stochastic population projections. Our results demonstrate that colonization is an unlikely explanation for the Neolithic dispersal in Central Europe, as the majority of fertility and growth rate estimates obtained in all three models are higher than levels expected in the early Neolithic population. On the basis of our models, we derived that colonization would be possible only if (1) more than 37% of women survived to mean age at childbearing, (2) Neolithic expansion in Central Europe lasted more than 150 years, and (3) the population of farmers grew in the entire settled area. These settings, however, represent very favorable demographic conditions that seem unlikely given current archaeological and demographic evidence. Therefore, our results support the view that Neolithic dispersal in Central Europe involved admixture of expanding farmers with local foragers. We estimate that the admixture contribution from foragers may have been between 55% and 72%.     

The effect of tied rank numbers on the linearity of dominance hierarchies

The occurence of tied rank numbers in dominance hierarchies is discussed, especially its effect on the linearity of the hierarchy. This linearity is measured with Landau's index, that is calculated for several hierarchies with tied ranks on one, two of three levels. Linearity is mostly affected by ties in small groups with many ties. A distinction is made between a hierarchy of individuals and hierarchical levels. The phenomenon of despotism is called an extreme case of tied ranks. It is proposed to regard hierarchies with a linearity in a continuous scale.     

Host–parasitoid spatial models: the interplay of demographic stochasticity and dynamics

Host–parasitoid metapopulation models have typically been deterministic models formulated with population numbers as a continuous variable. Spatial heterogeneity in local population abundance is a typical (and often essential) feature of these models and means that, even when average population density is high, some patches have small population sizes. In addition, large temporal population fluctuations are characteristic of many of these models, and this also results in periodically small local population sizes. Whenever population abundances are small, demographic stochasticity can become important in several ways. To investigate this problem, we have reformulated a deterministic, host–parasitoid metapopulation as an integer-based model in which encounters between hosts and parasitoids, and the fecundity of individuals are modelled as stochastic processes. This has a number of important consequences: (1) stochastic fluctuations at small population sizes tend to be amplified by the dynamics to cause massive population variability, i.e. the demographic stochasticity has a destabilizing effect; (2) the spatial patterns of local abundance observed in the deterministic counterpart are largely maintained (although the area of ''spatial chaos'' is extended); (3) at small population sizes, dispersal by discrete individuals leads to a smaller fraction of new patches being colonized, so that parasitoids with small dispersal rates have a greater tendency for extinction and higher dispersal rates have a larger competitive advantage; and (4) competing parasitoids that could coexist in the deterministic model due to spatial segregation cannot now coexist for any combination of parameters.     

Ancient DNA from an Early Neolithic Iberian population supports a pioneer colonization by first farmers

The Neolithic transition has been widely debated particularly regarding the extent to which this revolution implied a demographic expansion from the Near East. We attempted to shed some light on this process in northeastern Iberia by combining ancient DNA (aDNA) data from Early Neolithic settlers and published DNA data from Middle Neolithic and modern samples from the same region. We successfully extracted and amplified mitochondrial DNA from 13 human specimens, found at three archaeological sites dated back to the Cardial culture in the Early Neolithic (Can Sadurní and Chaves) and to the Late Early Neolithic (Sant Pau del Camp). We found that haplogroups with a low frequency in modern populations-N* and X1-are found at higher frequencies in our Early Neolithic population (～31%). Genetic differentiation between Early and Middle Neolithic populations was significant (F(ST) ～0.13, P<10(-5)), suggesting that genetic drift played an important role at this time. To improve our understanding of the Neolithic demographic processes, we used a Bayesian coalescence-based simulation approach to identify the most likely of three demographic scenarios that might explain the genetic data. The three scenarios were chosen to reflect archaeological knowledge and previous genetic studies using similar inferential approaches. We found that models that ignore population structure, as previously used in aDNA studies, are unlikely to explain the data. Our results are compatible with a pioneer colonization of northeastern Iberia at the Early Neolithic characterized by the arrival of small genetically distinctive groups, showing cultural and genetic connections with the Near East.     

Skull and limb morphology differentially track population history and environmental factors in the transition to agriculture in Europe

The Neolithic transition in Europe was a complex mosaic spatio-temporal process, involving both demic diffusion from the Near East and the cultural adoption of farming practices by indigenous hunter–gatherers. Previous analyses of Mesolithic hunter–gatherers and Early Neolithic farmers suggest that cranial shape variation preserves the population history signature of the Neolithic transition. However, the extent to which these same demographic processes are discernible in the postcranium is poorly understood. Here, for the first time, crania and postcranial elements from the same 11 prehistoric populations are analysed together in an internally consistent theoretical and methodological framework. Results show that while cranial shape reflects the population history differences between Mesolithic and Neolithic lineages, relative limb dimensions exhibit significant congruence with environmental variables such as latitude and temperature, even after controlling for geography and time. Also, overall limb size is found to be consistently larger in hunter–gatherers than farmers, suggesting a reduction in size related to factors other than thermoregulatory adaptation. Therefore, our results suggest that relative limb dimensions are not tracking the same demographic population history as the cranium, and point to the strong influence of climatic, dietary and behavioural factors in determining limb morphology, irrespective of underlying neutral demographic processes.     

The evolution of social philopatry in female primates

The transition from solitary life to sociality is considered one of the major transitions in evolution. In primates, this transition is currently not well understood. Traditional verbal models appear insufficient to unravel the complex interplay of environmental and demographic factors involved in the evolution of primate sociality, and recent phylogenetic reconstructions have produced conflicting results. We therefore analyze a theoretical model for the evolution of female social philopatry that sheds new light on the question why most primates live in groups. In individual-based simulations, we study the evolution of dispersal strategies of both resident females and their offspring. The model reveals that social philopatry can evolve through kin selection, even if retention of offspring is costly in terms of within-group resource competition and provides no direct benefits. Our model supports the role of predator avoidance as a selective pressure for group-living in primates, but it also suggests that a second benefit of group-living, communal resource defense, might be required to trigger the evolution of sizable groups. Lastly, our model reveals that seemingly small differences in demographic parameters can have profound effects on primate social evolution.     

The co‐evolution of social institutions,demography, and large‐scale human cooperation

Human cooperation is typically coordinated by institutions, which determine the outcome structure of the social interactions individuals engage in. Explaining the Neolithic transition from small‐ to large‐scale societies involves understanding how these institutions co‐evolve with demography. We study this using a demographically explicit model of institution formation in a patch‐structured population. Each patch supports both social and asocial niches. Social individuals create an institution, at a cost to themselves, by negotiating how much of the costly public good provided by cooperators is invested into sanctioning defectors. The remainder of their public good is invested in technology that increases carrying capacity, such as irrigation systems. We show that social individuals can invade a population of asocials, and form institutions that support high levels of cooperation. We then demonstrate conditions where the co‐evolution of cooperation, institutions, and demographic carrying capacity creates a transition from small‐ to large‐scale social groups.     

A craniometric perspective on the transition to agriculture in Europe

Debates surrounding the nature of the Neolithic demographic transition in Europe have historically centered on two opposing models: a "demic" diffusion model whereby incoming farmers from the Near East and Anatolia effectively replaced or completely assimilated indigenous Mesolithic foraging communities, and an "indigenist" model resting on the assumption that ideas relating to agriculture and animal domestication diffused from the Near East but with little or no gene flow. The extreme versions of these dichotomous models were heavily contested primarily on the basis of archeological and modern genetic data. However, in recent years a growing acceptance has arisen of the likelihood that both processes were ongoing throughout the Neolithic transition and that a more complex, regional approach is required to fully understand the change from a foraging to a primarily agricultural mode of subsistence in Europe. Craniometric data were particularly useful for testing these more complex scenarios, as they can reliably be employed as a proxy for the genetic relationships among Mesolithic and Neolithic populations. In contrast, modern genetic data assume that modern European populations accurately reflect the genetic structure of Europe at the time of the Neolithic transition, while ancient DNA data are still not geographically or temporally detailed enough to test continent-wide processes. Here, with particular emphasis on the role of craniometric analyses, we review the current state of knowledge regarding the cultural and biological nature of the Neolithic transition in Europe.     

Indo-European origins: A computer-simulation test of five hypotheses

Allele frequency distributions were generated by computer simulation of five models of microevolution in European populations. Genetic distances calculated from these distributions were compared with observed genetic distances among Indo-European speakers. The simulated models differ in complexity, but all incorporate random genetic drift and short-range gene flow (isolation by distance). The best correlations between observed and simulated data were obtained for two models where dispersal of Neolithic farmers from the Near East depends only on population growth. More complex models, where the timing of the farmers' expansion is constrained by archaeological time data, fail to account for a larger fraction of the observed genetic variation; this is also the case for a model including late Neolithic migrations from the Pontic steppes. The genetic structure of current populations speaking Indo-European languages seems therefore to largely reflect a Neolithic expansion. This is consistent with the hypothesis of a parallel spread of farming technologies and a proto-Indo-European language in the Neolithic. Allele-frequency gradients among Indo-European speakers may be due either to incomplete admixture between dispersing farmers, who presumably spoke proto-Indo-European, and pre-existing hunters and gatherers (as in the traditional demic diffusion hypothesis), or to founder effects during the farmers' dispersal. By contrast, successive migrational waves from the East, if any, do not seem to have had genetic consequences detectable by the present comparison of observed and simulated allele frequencies. © 1995 Wiley-Liss, Inc.     

Space Competition and Time Delays in Human Range Expansions. Application to the Neolithic Transition

Space competition effects are well-known in many microbiological and ecological systems. Here we analyze such an effect in human populations. The Neolithic transition (change from foraging to farming) was mainly the outcome of a demographic process that spread gradually throughout Europe from the Near East. In Northern Europe, archaeological data show a slowdown on the Neolithic rate of spread that can be related to a high indigenous (Mesolithic) population density hindering the advance as a result of the space competition between the two populations. We measure this slowdown from a database of 902 Early Neolithic sites and develop a time-delayed reaction-diffusion model with space competition between Neolithic and Mesolithic populations, to predict the observed speeds. The comparison of the predicted speed with the observations and with a previous non-delayed model show that both effects, the time delay effect due to the generation lag and the space competition between populations, are crucial in order to understand the observations.     

Phylogeography,genetic diversity and demographic history of the Iranian Kurdish groups based on mtDNA sequences

Throughout the history of modern humans, the current Kurdish-inhabited area has served as part of a tricontinental crossroad for major human migrations. Also, a significant body of archaeological evidence points to this area as the site of Neolithic transition. To investigate the phylogeography, origins and demographic history, mtDNA D-loop region of individuals representing four Kurdish groups from Iran were analysed. Our data indicated that most of the Kurds mtDNA lineages belong to branches of the haplogroups with the Western Eurasian origin; with small fractions of the Eastern Eurasian and sub-Saharan African lineages. The low level of mtDNA diversity observed in the Havrami group presented a bias towards isolation or increased drift due to small population size; while in the Kurmanji group it indicated a bias towards drift or mass migration events during the 5–18th century AD. The Mantel test showed strong isolation by distance, and AMOVA results for global and regional scales confirmed that the geography had acted as the main driving force in shaping the current pattern of mtDNA diversity, rather than linguistic similarity. The results of demographic analyses, in agreement with archaeological data, revealed a recent expansion of the Kurds (～9,500 years before present) related to the Neolithic transition from hunting and gathering, to farming and cattle breeding in the Near East. Further, the high frequencies of typical haplogroups for early farmers (H; 37.1%) and hunter-gatherers (U; 13.8%) in the Kurds may testify the earlier hunter-gatherers in the Kurdish-inhabited area that adopted and admixed the Kurds ancestors following the Neolithic transition.     

Self-organization and natural selection in the evolution of complex despotic societies

Differences between related species are usually explained as separate adaptations produced by individual selection. I discuss in this paper how related species, which differ in many respects, may evolve by a combination of individual selection, self-organization, and group-selection, requiring an evolutionary adaptation of only a single trait. In line with the supposed evolution of despotic species of macaques, we take as a starting point an ancestral species that is egalitarian and mildly aggressive. We suppose it to live in an environment with abundant food and we put the case that, if food becomes scarce and more clumped, natural selection at the level of the individual will favor individuals with a more intense aggression (implying, for instance, biting and fierce fighting). Using an individual-centered model, called DomWorld, I show what happens when the intensity of aggression increases. In DomWorld, group life is represented by artificial individuals that live in a homogeneous world. Individuals are extremely simple: all they do is flock together and, upon meeting one another, they may perform dominance interactions in which the effects of winning and losing are self-reinforcing. When the intensity of aggression in the model is increased, a complex feedback between the hierarchy and spatial structure results; via self-organization, this feedback causes the egalitarian society to change into a despotic one. The many differences between the two types of artificial society closely correspond to those between despotic and egalitarian macaques in the real world. Given that, in the model, the organization changes as a side effect of the change of one single trait proper to an egalitarian society, in the real world a despotic society may also have arisen as a side effect of the mutation of a single trait of an egalitarian species. If groups with different intensities of aggression evolve in this way, they will also have different gradients of hierarchy. When food is scarce, groups with the steepest hierarchy may have the best chance to survive, because at least a small number of individuals in such a group may succeed in producing offspring, whereas in egalitarian societies every individual is at risk of being insufficiently fed to reproduce. Therefore, intrademic group selection (selection within an interbreeding group) may have contributed to the evolution of despotic societies.     

The Neolithic Demographic Transition in Europe: Correlation with Juvenility Index Supports Interpretation of the Summed Calibrated Radiocarbon Date Probability Distribution (SCDPD) as a Valid Demographic Proxy

Analysis of the proportion of immature skeletons recovered from European prehistoric cemeteries has shown that the transition to agriculture after 9000 BP triggered a long-term increase in human fertility. Here we compare the largest analysis of European cemeteries to date with an independent line of evidence, the summed calibrated date probability distribution of radiocarbon dates (SCDPD) from archaeological sites. Our cemetery reanalysis confirms increased growth rates after the introduction of agriculture; the radiocarbon analysis also shows this pattern, and a significant correlation between both lines of evidence confirms the demographic validity of SCDPDs. We analyze the areal extent of Neolithic enclosures and demographic data from ethnographically known farming and foraging societies and we estimate differences in population levels at individual sites. We find little effect on the overall shape and precision of the SCDPD and we observe a small increase in the correlation with the cemetery trends. The SCDPD analysis supports the hypothesis that the transition to agriculture dramatically increased demographic growth, but it was followed within centuries by a general pattern of collapse even after accounting for higher settlement densities during the Neolithic. The study supports the unique contribution of SCDPDs as a valid demographic proxy for the demographic patterns associated with early agriculture.     

Female and Male Perspectives on the Neolithic Transition in Europe: Clues from Ancient and Modern Genetic Data

The arrival of agriculture into Europe during the Neolithic transition brought a significant shift in human lifestyle and subsistence. However, the conditions under which the spread of the new culture and technologies occurred are still debated. Similarly, the roles played by women and men during the Neolithic transition are not well understood, probably due to the fact that mitochondrial DNA (mtDNA) and Y chromosome (NRY) data are usually studied independently rather than within the same statistical framework. Here, we applied an integrative approach, using different model-based inferential techniques, to analyse published datasets from contemporary and ancient European populations. By integrating mtDNA and NRY data into the same admixture approach, we show that both males and females underwent the same admixture history and both support the demic diffusion model of Ammerman and Cavalli-Sforza. Similarly, the patterns of genetic diversity found in extant and ancient populations demonstrate that both modern and ancient mtDNA support the demic diffusion model. They also show that population structure and differential growth between farmers and hunter-gatherers are necessary to explain both types of data. However, we also found some differences between male and female markers, suggesting that the female effective population size was larger than that of the males, probably due to different demographic histories. We argue that these differences are most probably related to the various shifts in cultural practices and lifestyles that followed the Neolithic Transition, such as sedentism, the shift from polygyny to monogamy or the increase of patrilocality.     

Scaling population responses to spatial environmental variability in advection-dominated systems

We model the spatial dynamics of an open population of organisms that disperse solely through advection in order to understand responses to multiscale environmental variability. We show that the distance over which a population responds to a localized perturbation, called the response length, can be characterized as an organisms average lifetime dispersal distance, unless there is strong density‐dependence in demographic or dispersal rates. Continuous spatial fluctuations in demographic rates at scales smaller than the response length will be largely averaged in the population distribution, whereas those in per capita emigration rates will be strongly tracked. We illustrate these results using a parameterized example to show how responses to environmental variability may differ in streams with different average current velocities. Our model suggests an approach to linking local dynamics dominated by dispersal processes to larger‐scale dynamics dominated by births and deaths.     

Genetic structuring in fluctuating populations

The effect of genetic drift in spatially distributed dispersal-linked and density-regulated populations is studied in a classical one-locus two-allele system. We analyse emergence of genetic differentiation assuming random drift only, where the noise-like variability is due to demographic stochasticity. We find emergence of clusters of sub-units with local allele fixation and persistence of both alleles in lengthy simulations. We demonstrate that local allele fixation (extending over a number of adjoining spatial sub-units) – without global loss of alleles – may occur when the carrying capacities of local patches are small, under a full range population dynamic regimes, when dispersal rate is small, and when redistribution (through dispersal) does not act as global mixer. These results are novel. The key to the observations is that drift is simultaneously influenced by distance-dependent dispersal, demographic stochasticity and autocorrelated population fluctuations due to delayed-density dependence. These are standard elements of contemporary population models in spatially structured context. With stable large populations, no stochasticity and dispersal limited to neighbours only, our model collapses to the stepping-stone model, while with dispersal being random and global, the model collapses to Wright's island model.     

Differences in Nutrient Requirements Imply a Non-Linear Emergence of Leaders in Animal Groups

Collective decision making and especially leadership in groups are among the most studied topics in natural, social, and political sciences. Previous studies have shown that some individuals are more likely to be leaders because of their social power or the pertinent information they possess. One challenge for all group members, however, is to satisfy their needs. In many situations, we do not yet know how individuals within groups distribute leadership decisions between themselves in order to satisfy time-varying individual requirements. To gain insight into this problem, we build a dynamic model where group members have to satisfy different needs but are not aware of each other''s needs. Data about needs of animals come from real data observed in macaques. Several studies showed that a collective movement may be initiated by a single individual. This individual may be the dominant one, the oldest one, but also the one having the highest physiological needs. In our model, the individual with the lowest reserve initiates movements and decides for all its conspecifics. This simple rule leads to a viable decision-making system where all individuals may lead the group at one moment and thus suit their requirements. However, a single individual becomes the leader in 38% to 95% of cases and the leadership is unequally (according to an exponential law) distributed according to the heterogeneity of needs in the group. The results showed that this non-linearity emerges when one group member reaches physiological requirements, mainly the nutrient ones – protein, energy and water depending on weight - superior to those of its conspecifics. This amplification may explain why some leaders could appear in animal groups without any despotism, complex signalling, or developed cognitive ability.     

Population continuity, demic diffusion and Neolithic origins in central-southern Germany: The evidence from body proportions

The transition to agro-pastoralism in central Europe has been framed within a dichotomy of “regional continuity” versus exogenous “demic diffusion”. While substantial genetic support exists for a model of demographic diffusion from an ancestral source in the Near East, archaeological data furnish weak support for the “wave of advance” model. Nevertheless, archaeological evidence attests the widespread introduction of an exogenous “package” comprising ceramics, cereals, pulses and domesticated animals to central Europe at 5600 cal BCE.Body proportions are under strong climatic selection and evince remarkable stability within regional lineages. As such, they offer a viable and robust alternative to cranio-facial data in assessing hypothesised continuity and replacement with the transition to agro-pastoralism in central Europe. Humero-clavicular, brachial and crural indices in a large sample (n=75) of Linienbandkeramik (LBK), Late Neolithic and Early Bronze Age specimens from the middle Elbe-Saale-Werra valley (MESV) were compared with Eurasian and African terminal Pleistocene, European Mesolithic and geographically disparate recent human specimens.Mesolithic Europeans display considerable variation in humero-clavicular and brachial indices yet none approach the extreme “hyper-polar” morphology of LBK humans from the MESV. In contrast, Late Neolithic and Early Bronze Age peoples display elongated brachial and crural indices reminiscent of terminal Pleistocene and “tropically adapted” recent humans. These marked morphological changes likely reflect exogenous immigration during the terminal Fourth millennium cal BC. Population expansion and diffusion is a function of increased mobility and settlement dispersal concomitant with significant technological and subsistence changes in later Neolithic societies during the late fourth millennium cal BCE.     

Evolution of Complex Density-Dependent Dispersal Strategies

The question of how dispersal behavior is adaptive and how it responds to changes in selection pressure is more relevant than ever, as anthropogenic habitat alteration and climate change accelerate around the world. In metapopulation models where local populations are large, and thus local population size is measured in densities, density-dependent dispersal is expected to evolve to a single-threshold strategy, in which individuals stay in patches with local population density smaller than a threshold value and move immediately away from patches with local population density larger than the threshold. Fragmentation tends to convert continuous populations into metapopulations and also to decrease local population sizes. Therefore we analyze a metapopulation model, where each patch can support only a relatively small local population and thus experience demographic stochasticity. We investigated the evolution of density-dependent dispersal, emigration and immigration, in two scenarios: adult and natal dispersal. We show that density-dependent emigration can also evolve to a nonmonotone, “triple-threshold” strategy. This interesting phenomenon results from an interplay between the direct and indirect benefits of dispersal and the costs of dispersal. We also found that, compared to juveniles, dispersing adults may benefit more from density-dependent vs. density-independent dispersal strategies.