Emergence of size-structured dominance hierarchies through size-dependent feedback

Size-based dominance hierarchies influence fitness, group size and population dynamics and link dominance structure to evolutionary and ecological outcomes. While larger individuals often gain dominance, social status may influence growth and size in return, resulting in feedbacks among status, growth and size. Here, we present two models evaluating how these feedbacks influence the emergence of size structure in a dominance hierarchy. In the first, size influences competition for food and investment in suppressing growth of groupmates. Stable size differences emerged when suppression was greatest for similarly sized individuals and size had little effect on competition for food. The model predicted size divergence when size strongly affected competition for food. In the second model, we used a dynamic game to solve for optimal investment in growth suppression as a function of size structure. Investment in growth suppression was favoured only when dominants and subordinates were similar in size, generating size ratios different than those expected by chance. Variation in the feedbacks among growth, size and status can explain variation in emergent size structure of dominance hierarchies and its consequences for conflict within groups. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

The establishment and maintenance of dominance hierarchies

Animal groups are often organized hierarchically, with dominant individuals gaining priority access to resources and reproduction over subordinate individuals. Initial dominance hierarchy formation may be influenced by multiple interacting factors, including an animal's individual attributes, conventions and self-organizing social dynamics. After establishment, hierarchies are typically maintained over the long-term because individuals save time, energy and reduce the risk of injury by recognizing and abiding by established dominance relationships. A separate set of behaviours are used to maintain dominance relationships within groups, including behaviours that stabilize ranks (punishment, threats, behavioural asymmetry), as well as signals that provide information about dominance rank (individual identity signals, signals of dominance). In this review, we describe the behaviours used to establish and maintain dominance hierarchies across different taxa and types of societies. We also review opportunities for future research including: testing how self-organizing behavioural dynamics interact with other factors to mediate dominance hierarchy formation, measuring the long-term stability of social hierarchies and the factors that disrupt hierarchy stability, incorporating phenotypic plasticity into our understanding of the behavioural dynamics of hierarchies and considering how cognition coevolves with the behaviours used to establish and maintain hierarchies. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Dominance in a socially dynamic setting: hierarchical structure and conflict dynamics in ravens' foraging groups

Dominance hierarchies typically emerge in systems where group members regularly encounter and compete for resources. In birds, the ‘open’ and dynamic structure of foraging groups may prevent the emergence of structured hierarchies, although this assumption have hardly been tested. We report on agonistic data for ravens Corvus corax , collected over two 18-month periods for 183 marked individuals of a wild (fluid) population and 51 birds from six captive (stable) groups. We show that the dominance structure (steep and transitive) in wild foraging groups is strikingly similar to that found in captivity. In the wild, we found that higher ranks are mainly occupied by males, older and more aggressive individuals that also tend to receive fewer aggressions. Exploring the mechanisms sustaining the wild dominance structure, we confirmed that males are more aggressive than females and, with age, tend to receive fewer aggressions than females. Males that are about to leave the foraging groups for some months are less aggressive than newcomers or locals, while newcomers are specifically targeted by aggressions in their first year (as juveniles). Taken together, our results indicate that the socially dynamic conditions ravens face during foraging do not hinder, but provide opportunities for, using (advanced) social cognition. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Distinct gene regulatory signatures of dominance rank and social bond strength in wild baboons

The social environment is a major determinant of morbidity, mortality and Darwinian fitness in social animals. Recent studies have begun to uncover the molecular processes associated with these relationships, but the degree to which they vary across different dimensions of the social environment remains unclear. Here, we draw on a long-term field study of wild baboons to compare the signatures of affiliative and competitive aspects of the social environment in white blood cell gene regulation, under both immune-stimulated and non-stimulated conditions. We find that the effects of dominance rank on gene expression are directionally opposite in males versus females, such that high-ranking males resemble low-ranking females, and vice versa. Among females, rank and social bond strength are both reflected in the activity of cellular metabolism and proliferation genes. However, while we observe pronounced rank-related differences in baseline immune gene activity, only bond strength predicts the fold-change response to immune (lipopolysaccharide) stimulation. Together, our results indicate that the directionality and magnitude of social effects on gene regulation depend on the aspect of the social environment under study. This heterogeneity may help explain why social environmental effects on health and longevity can also vary between measures. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Quantifying the dynamics of nearly 100 years of dominance hierarchy research

Dominance hierarchies have been studied for almost 100 years. The science of science approach used here provides high-level insight into how the dynamics of dominance hierarchy research have shifted over this long timescale. To summarize these patterns, I extracted publication metadata using a Google Scholar search for the phrase ‘dominance hierarchy’, resulting in over 26 000 publications. I used text mining approaches to assess patterns in three areas: (1) general patterns in publication frequency and rate, (2) dynamics of term usage and (3) term co-occurrence in publications across the history of the field. While the overall number of publications per decade continues to rise, the percent growth rate has fallen in recent years, demonstrating that although there is sustained interest in dominance hierarchies, the field is no longer experiencing the explosive growth it showed in earlier decades. Results from title term co-occurrence networks and community structure show that the different subfields of dominance hierarchy research were most strongly separated early in the field’s history while modern research shows more evidence for cohesion and a lack of distinct term community boundaries. These methods provide a general view of the history of research on dominance hierarchies and can be applied to other fields or search terms to gain broad synthetic insight into patterns of interest, especially in fields with large bodies of literature. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Dominance in humans

Dominance captures behavioural patterns found in social hierarchies that arise from agonistic interactions in which some individuals coercively exploit their control over costs and benefits to extract deference from others, often through aggression, threats and/or intimidation. Accumulating evidence points to its importance in humans and its separation from prestige—an alternate avenue to high status in which status arises from information (e.g. knowledge, skill, etc.) or other non-rival goods. In this review, we provide an overview of the theoretical underpinnings of dominance as a concept within evolutionary biology, discuss the challenges of applying it to humans and consider alternative theoretical accounts which assert that dominance is relevant to understanding status in humans. We then review empirical evidence for its continued importance in human groups, including the effects of dominance—independently of prestige—on measurable outcomes such as social influence and reproductive fitness, evidence for specialized dominance psychology, and evidence for gender-specific effects. Finally, because human-specific factors such as norms and coalitions may place bounds on purely coercive status-attainment strategies, we end by considering key situations and contexts that increase the likelihood for dominance status to coexist alongside prestige status within the same individual, including how: (i) institutional power and authority tend to elicit dominance; (ii) dominance-enhancing traits can at times generate benefits for others (prestige); and (iii) certain dominance cues and ethology may lead to mis-attributions of prestige. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

The build-up of dominance hierarchies in eusocial insects

Reproductive division of labour is a hallmark of eusocial insects. However, its stability can often be hampered by the potential for reproduction by otherwise sterile nest-mates. Dominance hierarchy has a crucial role in some species in regulating which individuals reproduce. Compared with those in vertebrates, the dominance hierarchies in eusocial insects tend to involve many more individuals, and should require additional selective forces unique to them. Here, we provide an overview of a series of studies on dominance hierarchies in eusocial insects. Although reported from diverse eusocial taxa, dominance hierarchies have been extensively studied in paper wasps and ponerine ants. Starting from molecular physiological attributes of individuals, we describe how the emergence of dominance hierarchies can be understood as a kind of self-organizing process through individual memory and local behavioural interactions. The resulting global structures can be captured by using network analyses. Lastly, we argue the adaptive significance of dominance hierarchies from the standpoint of sterile subordinates. Kin selection, underpinned by relatedness between nest-mates, is key to the subordinates' acceptance of their positions in the hierarchies. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Social hierarchies and social networks in humans

Across species, social hierarchies are often governed by dominance relations. In humans, where there are multiple culturally valued axes of distinction, social hierarchies can take a variety of forms and need not rest on dominance relations. Consequently, humans navigate multiple domains of status, i.e. relative standing. Importantly, while these hierarchies may be constructed from dyadic interactions, they are often more fundamentally guided by subjective peer evaluations and group perceptions. Researchers have typically focused on the distinct elements that shape individuals’ relative standing, with some emphasizing individual-level attributes and others outlining emergent macro-level structural outcomes. Here, we synthesize work across the social sciences to suggest that the dynamic interplay between individual-level and meso-level properties of the social networks in which individuals are embedded are crucial for understanding the diverse processes of status differentiation across groups. More specifically, we observe that humans not only navigate multiple social hierarchies at any given time but also simultaneously operate within multiple, overlapping social networks. There are important dynamic feedbacks between social hierarchies and the characteristics of social networks, as the types of social relationships, their structural properties, and the relative position of individuals within them both influence and are influenced by status differentiation. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Hot-headed peckers: thermographic changes during aggression among juvenile pheasants ( Phasianus colchicus )

In group-living vertebrates, dominance status often covaries with physiological measurements (e.g. glucocorticoid levels), but it is unclear how dominance is linked to dynamic changes in physiological state over a shorter, behavioural timescale. In this observational study, we recorded spontaneous aggression among captive juvenile pheasants ( Phasianus colchicus ) alongside infrared thermographic measurements of their external temperature, a non-invasive technique previously used to examine stress responses in non-social contexts, where peripheral blood is redirected towards the body core. We found low but highly significant repeatability in maximum head temperature, suggesting individually consistent thermal profiles, and some indication of lower head temperatures in more active behavioural states (e.g. walking compared to resting). These individual differences were partly associated with sex, females being cooler on average than males, but unrelated to body size. During pairwise aggressive encounters, we observed a non-monotonic temperature change, with head temperature dropping rapidly immediately prior to an attack and increasing rapidly afterwards, before returning to baseline levels. This nonlinear pattern was similar for birds in aggressor and recipient roles, but aggressors were slightly hotter on average. Our findings show that aggressive interactions induce rapid temperature changes in dominants and subordinates alike, and highlight infrared thermography as a promising tool for investigating the physiological basis of pecking orders in galliforms. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

Neural systems that facilitate the representation of social rank

Across species, animals organize into social dominance hierarchies that serve to decrease aggression and facilitate survival of the group. Neuroscientists have adopted several model organisms to study dominance hierarchies in the laboratory setting, including fish, reptiles, rodents and primates. We review recent literature across species that sheds light onto how the brain represents social rank to guide socially appropriate behaviour within a dominance hierarchy. First, we discuss how the brain responds to social status signals. Then, we discuss social approach and avoidance learning mechanisms that we propose could drive rank-appropriate behaviour. Lastly, we discuss how the brain represents memories of individuals (social memory) and how this may support the maintenance of unique individual relationships within a social group. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.