Defining Higher Levels in the Multilevel Societies of Geladas (Theropithecus gelada)

Multilevel societies, identified by two or more nested levels (or modules) of organization, have been touted as some of the most complex social systems. However, few empirical studies have effectively quantified the association patterns that delineate the various levels in such societies. In particular, the multiple levels of gelada society were first described >3 decades ago, yet no operational definitions exist for the higher levels, i.e., levels above the one-male unit. In geladas, multiple units form aggregations that fission and fuse throughout the day, and throughout the year, blurring the distinctions between previously described higher social levels: teams, bands, and communities. Here we use 5?yr of data on the daily composition of a population of geladas living in the Simien Mountains National Park, Ethiopia to test the hypothesis that higher levels of gelada organization are discrete entities. If gelada aggregations are nothing more than a group of units that share a home range, then we expect a continuous distribution of unit association. If, however, gelada aggregations are indeed discrete organizational levels, then we expect discontinuity in the patterns of association. We found significant discontinuity at the 50% association level, indicating a sharp distinction between members of the same band (>50% association) and members of the same community (<50% association). We also found evidence that recently fissioned units form teams that associate significantly more than other band members. Thus, despite their extremely fluid social organization, gelada social levels are nevertheless clearly identifiable and quantifiable. Based on these results, we suggest that gelada society is an extremely flexible, multilevel society with fission–fusion dynamics, and as such gelada society presents an unusual example for understanding the evolution of modular societies.     

A critique of 'countryside biogeography' as a guide to research in human‐dominated landscapes

Proliferation of redundant terms in ecology and conservation slows progress and creates confusion. ‘Countryside biogeography’ has been promoted as a new framework for conservation in production landscapes, so may offer a replacement for other concepts used by landscape ecologists. We conducted a systematic review to assess whether the 'countryside biogeography' concept provides a distinctive framing for conservation in human‐dominated landscapes relative to existing concepts. We reviewed 147 papers referring to countryside biogeography and 81 papers that did not. These papers were divided into categories representing three levels of use of countryside biogeography concepts (strong, weak, cited only) and two categories that did not use countryside biogeography at all but used similar concepts including fragmentation and matrix. We revealed few distinctions among groups of papers. Countryside biogeography papers made more frequent use of the terms 'ecosystem services', 'intensification' and 'land sparing' compared with non‐countryside biogeography papers. Papers that did not refer to countryside biogeography sampled production areas (e.g. farms) less often, and this related to their focus on habitat specialist species for which patch‐matrix assumptions were reasonable. Countryside biogeography offers a conceptual wrapper rather than a distinctive framework for advancing research in human‐modified landscapes. This and similar wrappers such as ‘conservation biogeography’ and ‘agricultural biogeography’ risk creating confusion among new researchers, and can prevent clear communication about research. To improve communication, we recommend using the suite of well‐established terms already applied to conservation in human‐modified landscapes rather than through an interceding conceptual wrapper.     

Earliest Porotic Hyperostosis on a 1.5-Million-Year-Old Hominin,Olduvai Gorge,Tanzania

Meat-eating was an important factor affecting early hominin brain expansion, social organization and geographic movement. Stone tool butchery marks on ungulate fossils in several African archaeological assemblages demonstrate a significant level of carnivory by Pleistocene hominins, but the discovery at Olduvai Gorge of a child''s pathological cranial fragments indicates that some hominins probably experienced scarcity of animal foods during various stages of their life histories. The child''s parietal fragments, excavated from 1.5-million-year-old sediments, show porotic hyperostosis, a pathology associated with anemia. Nutritional deficiencies, including anemia, are most common at weaning, when children lose passive immunity received through their mothers'' milk. Our results suggest, alternatively, that (1) the developmentally disruptive potential of weaning reached far beyond sedentary Holocene food-producing societies and into the early Pleistocene, or that (2) a hominin mother''s meat-deficient diet negatively altered the nutritional content of her breast milk to the extent that her nursing child ultimately died from malnourishment. Either way, this discovery highlights that by at least 1.5 million years ago early human physiology was already adapted to a diet that included the regular consumption of meat.     

The weird eusociality of polyembryonic parasites

Some parasitoid wasps possess soldier castes during their parasitic larval stage, but are often neglected from our evolutionary theories explaining caste systems in animal societies. This is primarily due to the polyembryonic origin of their societies. However, recent discoveries of polyembryonic trematodes (i.e. flatworms) possessing soldier castes require us to reconsider this reasoning. I argue we can benefit from including these polyembryonic parasites in eusocial discussions, for polyembryony and parasitism are taxonomically vast and influence the evolution of social behaviours and caste systems in various circumstances. Despite their polyembryony, their social evolution can be explained by theories of eusociality designed for parent–offspring groups, which are the subjects of most social evolution research. Including polyembryonic parasites in these theories follows the trend of major evolutionary transitions theory expanding social evolution research into all levels of biological organization. In addition, these continued discoveries of caste systems in parasites suggest social evolution may be more relevant to parasitology than currently acknowledged.     

A social network analysis of primate groups

Primate social systems are difficult to characterize, and existing classification schemes have been criticized for being overly simplifying, formulated only on a verbal level or partly inconsistent. Social network analysis comprises a collection of analytical tools rooted in the framework of graph theory that were developed to study human social interaction patterns. More recently these techniques have been successfully applied to examine animal societies. Primate social systems differ from those of humans in both size and density, requiring an approach that puts more emphasis on the quality of relationships. Here, we discuss a set of network measures that are useful to describe primate social organization and we present the results of a network analysis of 70 groups from 30 different species. For this purpose we concentrated on structural measures on the group level, describing the distribution of interaction patterns, centrality, and group structuring. We found considerable variability in those measures, reflecting the high degree of diversity of primate social organizations. By characterizing primate groups in terms of their network metrics we can draw a much finer picture of their internal structure that might be useful for species comparisons as well as the interpretation of social behavior.     

From social network (centralized vs. decentralized) to collective decision-making (unshared vs. shared consensus)

Relationships we have with our friends, family, or colleagues influence our personal decisions, as well as decisions we make together with others. As in human beings, despotism and egalitarian societies seem to also exist in animals. While studies have shown that social networks constrain many phenomena from amoebae to primates, we still do not know how consensus emerges from the properties of social networks in many biological systems. We created artificial social networks that represent the continuum from centralized to decentralized organization and used an agent-based model to make predictions about the patterns of consensus and collective movements we observed according to the social network. These theoretical results showed that different social networks and especially contrasted ones--star network vs. equal network--led to totally different patterns. Our model showed that, by moving from a centralized network to a decentralized one, the central individual seemed to lose its leadership in the collective movement's decisions. We, therefore, showed a link between the type of social network and the resulting consensus. By comparing our theoretical data with data on five groups of primates, we confirmed that this relationship between social network and consensus also appears to exist in animal societies.     

Information processing in social insect networks

Investigating local-scale interactions within a network makes it possible to test hypotheses about the mechanisms of global network connectivity and to ask whether there are general rules underlying network function across systems. Here we use motif analysis to determine whether the interactions within social insect colonies resemble the patterns exhibited by other animal associations or if they exhibit characteristics of biological regulatory systems. Colonies exhibit a predominance of feed-forward interaction motifs, in contrast to the densely interconnected clique patterns that characterize human interaction and animal social networks. The regulatory motif signature supports the hypothesis that social insect colonies are shaped by selection for network patterns that integrate colony functionality at the group rather than individual level, and demonstrates the utility of this approach for analysis of selection effects on complex systems across biological levels of organization.     

The evolution of altruistic social preferences in human groups

In this paper, we consider three hypotheses to account for the evolution of the extraordinary capacity for large-scale cooperation and altruistic social preferences within human societies. One hypothesis is that human cooperation is built on the same evolutionary foundations as cooperation in other animal societies, and that fundamental elements of the social preferences that shape our species'' cooperative behaviour are also shared with other closely related primates. Another hypothesis is that selective pressures favouring cooperative breeding have shaped the capacity for cooperation and the development of social preferences, and produced a common set of behavioural dispositions and social preferences in cooperatively breeding primates and humans. The third hypothesis is that humans have evolved derived capacities for collaboration, group-level cooperation and altruistic social preferences that are linked to our capacity for culture. We draw on naturalistic data to assess differences in the form, scope and scale of cooperation between humans and other primates, experimental data to evaluate the nature of social preferences across primate species, and comparative analyses to evaluate the evolutionary origins of cooperative breeding and related forms of behaviour.     

Culture as a stressor: A revised model of biocultural interaction

Contemporary urban societies display in high relief the action of social stratification on human biology. Recent studies of biological responses to urban environments and of socioeconomically disadvantaged people indicate that culture allocates risks disproportionately to some individuals and groups within society through its constituent values and related patterns of behavior. Although risk allocation is present in all societies, it is very clear in urban environments within stratified societies where high exposure to harmful materials is many times more likely for some segments of society. In urban environments, culture may be seen as adding stressors to the environment by concentrating naturally occurring materials to levels that are toxic to humans and through the creation of new toxic materials. In stratified societies the risk of exposure to these new stressors is focused on the socioeconomically disadvantaged. This exposure has consequences that increase the likelihood of more exposure and more socioeconomic disadvantage, thereby increasing social stratification. This suggests that models of biocultural interaction include a feedback relationship in which biological factors influence the sociocultural system in addition to the usual action of the sociocultural system on biological features and responses. This model strongly reinforces the view that stressors can originate from cultural arrangements. Am J Phys Anthropol 102:67–77, 1997 © 1997 Wiley-Liss, Inc.     

The sociobiology of molecular systems

It is often assumed that molecular systems are designed to maximize the competitive ability of the organism that carries them. In reality, natural selection acts on both cooperative and competitive phenotypes, across multiple scales of biological organization. Here I ask how the potential for social effects in evolution has influenced molecular systems. I discuss a range of phenotypes, from the selfish genetic elements that disrupt genomes, through metabolism, multicellularity and cancer, to behaviour and the organization of animal societies. I argue that the balance between cooperative and competitive evolution has shaped both form and function at the molecular scale.     

The genetic implications of biological and cultural structuring processes during hominid evolution

Human populations are genetically structured through biological reproduction but reproduction is socially structured through culturally expressed systems of marriage, hence the need to take into account the cultural dimension of human societies when considering the genetic structure of modern human populations. The interplay between these two structuring processes is examined through the implications of a biologically based primate form of social organization versus a culturally based foraging form of social organization for an anomaly between the genetic differentiation implied by micro-evolutionary genetic models versus the species-wide pattern of morphological change implied by hominid fossil evidence. Simulation is used to derive the pattern for genetic differentiation at the level of the living group for both forms of social organization and a competition model is added to the genetic model as a way to resolve the anomaly. The importance of the competition model for the transition from a biologically based to a culturally based form of social organization is discussed.     

Polydomy in ants: what we know, what we think we know, and what remains to be done

The correct identification of colony boundaries is an essential prerequisite for empirical studies of ant behaviour and evolution. Ant colonies function at various organizational levels, and these boundaries may be difficult to assess. Moreover, new complexity can be generated through the presence of spatially discrete subgroups within a more or less genetically homogeneous colony, a situation called polydomy. A colony is polydomous only if individuals (workers and brood) of its constituent nests function as a social and cooperative unit and are regularly interchanged among nests. This condition was previously called polycalic, and the term polydomy was used in a broader sense for a group of daughter nests of the same mother colony (implying limited female dispersal), without regard to whether these different nests continued to exchange individuals. We think that this distinction between ‘polycaly’ and ‘polydomy’ concerns two disparate concepts. We thus prefer the narrower definition of polydomy, which groups individuals that interact socially. Does this new level of organization affect the way in which natural selection acts on social traits? Here, after examining the history of terms, we review all ant species that have been described as expressing polydomous structures. We show that there is no particular syndrome of traits predictably associated with polydomy. We detail the existing theoretical predictions and empirical results on the ecology of polydomy, and the impact of polydomy on social evolution and investment strategies, while carefully distinguishing monogynous from polygynous species. Finally, we propose a methodology for future studies and offer ideas about what remains to be done. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 319–348.     

Power over reproduction in social hymenoptera

Inclusive fitness theory has been very successful in predicting and explaining much of the observed variation in the reproductive characteristics of insect societies. For example, the theory correctly predicts sex-ratio biasing by workers in relation to the queen''s mating frequency. However, within an insect society there are typically multiple reproductive optima, each corresponding to the interest of different individual(s) or parties of interest. When multiple optima occur, which party''s interests prevail? Presumably, the interests of the party with the greatest ''power''; the ability to do or act. This article focuses on factors that influence power over colony reproduction. In particular, we seek to identify the principles that may cause different parties of interest to have greater or lesser power. In doing this, we discuss power from two different angles. On the one hand, we discuss general factors based upon non-idiosyncratic biological features (e.g. information, access to and ability to process food) that are likely to be important to all social Hymenoptera. On the other hand, we discuss idiosyncratic factors that depend upon the biology of a taxon at any hierarchical level. We propose that a better understanding of the diversity of reproductive characteristics of insect societies will come from combining inclusive fitness theory with a wide range of other factors that affect relative power in a conflict situation.     

Inclusive fitness and differential productivity across the life course determine intergenerational transfers in a small-scale human society

Transfers of resources between generations are an essential element in current models of human life-history evolution accounting for prolonged development, extended lifespan and menopause. Integrating these models with Hamilton''s theory of inclusive fitness, we predict that the interaction of biological kinship with the age-schedule of resource production should be a key driver of intergenerational transfers. In the empirical case of Tsimane’ forager–horticulturalists in Bolivian Amazonia, we provide a detailed characterization of net transfers of food according to age, sex, kinship and the net need of donors and recipients. We show that parents, grandparents and siblings provide significant net downward transfers of food across generations. We demonstrate that the extent of provisioning responds facultatively to variation in the productivity and demographic composition of families, as predicted by the theory. We hypothesize that the motivation to provide these critical transfers is a fundamental force that binds together human nuclear and extended families. The ubiquity of three-generational families in human societies may thus be a direct reflection of fundamental evolutionary constraints on an organism''s life-history and social organization.     

社会网络分析法及其在动物行为学研究中的应用

近年来,社会网络分析法被广泛用于动物行为学研究。它通过量化动物社会关系中的特定属性 (如中心度和中心势),可辨识动物群体中的关键个体及其在群体中的作用,揭示动物社会交往的形成机制,深化人们对动物社会性起源、个体行为与种群动态格局间关系的理解。本文简要介绍了社会网络分析法的发展史和基本概念,然后阐述了如何建立网络及选择常用指标,强调了创建原模型的必要性及途径。最后着重评述了社会网络分析法在动物行为学研究中的应用现状,并提出未来应当关注直接身体接触 (如打斗和理毛) 和非肢体接触 (如声音通讯) 等不同交往形式的变动过程,以此构建社会交往的动态网络。同时也需要加强种间社交网络的研究,以便增进人们对社会行为的生态功能以及合作理论等问题的理解。     

Students' cellular and molecular explanations of genetic phenomena

Genetic concepts are assigned to three principal levels of organisation: macroscopic level, microscopic level, and submicroscopic level. In order to probe Israeli students' understanding at each level, and to gain a sense of their ability to connect ideas and concepts across different levels, we asked three different types of question. Two of the question types ask students to make a bridge between levels by asking them to explain a phenomenon at one level using concepts and processes from a different level. For example, to explain the appearance of phenotypic traits (macroscopic level) using concepts like genes or chromosomes (microscopic level). One question dealt with the molecular level only. We investigated three populations: 9^th graders, 12^th graders, and pre-service teachers. Based on our findings we suggest improvements both in terms of teaching methods and curriculum content.     

Social vocalizations can release oxytocin in humans

Vocalizations are important components of social behaviour in many vertebrate species, including our own. Less well-understood are the hormonal mechanisms involved in response to vocal cues, and how these systems may influence the course of behavioural evolution. The neurohormone oxytocin (OT) partly governs a number of biological and social processes critical to fitness, such as attachment between mothers and their young, and suppression of the stress response after contact with trusted conspecfics. Rodent studies suggest that OT''s release is contingent upon direct tactile contact with such individuals, but we hypothesized that vocalizations might be capable of producing the same effect. To test our hypothesis, we chose human mother–daughter dyads and applied a social stressor to the children, following which we randomly assigned participants into complete contact, speech-only or no-contact conditions. Children receiving a full complement of comfort including physical, vocal and non-verbal contact showed the highest levels of OT and the swiftest return to baseline of a biological marker of stress (salivary cortisol), but a strikingly similar hormonal profile emerged in children comforted solely by their mother''s voice. Our results suggest that vocalizations may be as important as touch to the neuroendocrine regulation of social bonding in our species.     

Social network- and community-level influences on contraceptive use: evidence from rural Poland

The diffusion of ‘modern’ contraceptives—as a proxy for the spread of low-fertility norms—has long interested researchers wishing to understand global fertility decline. A fundamental question is how local cultural norms and other people''s behaviour influence the probability of contraceptive use, independent of women''s socioeconomic and life-history characteristics. However, few studies have combined data at individual, social network and community levels to simultaneously capture multiple levels of influence. Fewer still have tested if the same predictors matter for different contraceptive types. Here, we use new data from 22 high-fertility communities in Poland to compare predictors of the use of (i) any contraceptives—a proxy for the decision to control fertility—with those of (ii) ‘artificial’ contraceptives—a subset of more culturally taboo methods. We find that the contraceptive behaviour of friends and family is more influential than are women''s own characteristics and that community level characteristics additionally influence contraceptive use. Highly educated neighbours accelerate women''s contraceptive use overall, but not their artificial method use. Highly religious neighbours slow women''s artificial method use, but not their contraceptive use overall. Our results highlight different dimensions of sociocultural influence on contraceptive diffusion and suggest that these may be more influential than are individual characteristics. A comparative multilevel framework is needed to understand these dynamics.     

Developments in social terminology: semantic battles in a conceptual war

The problems posed by the evolution of the diverse forms of animal sociality are among the most important and fascinating in evolutionary biology. The conceptual and terminological framework guiding studies of social evolution has been based on a particular insect model, namely, that of highly derived family-structured societies. Virtually all other social systems have been categorized as 'less social' relative to these societies. Recently, the ambiguities and constraints inherent in this hierarchical classification have led to numerous proposals to amend social terminology. What is the best framework for studying social evolution? Should the traditional classification be expanded, narrowed or abandoned altogether? In an important respect, most recent proposals present the same wine in a different bottle by retaining and recasting key terms of the traditional social-evolutionary classification.