The experimental emergence of convention in a non-human primate

Conventions form an essential part of human social and cultural behaviour and may also be important to other animal societies. Yet, despite the wealth of evidence that has accumulated for culture in non-human animals, we know surprisingly little about non-human conventions beyond a few rare examples. We follow the literature in behavioural ecology and evolution and define conventions as systematic behaviours that solve a coordination problem in which two or more individuals need to display complementary behaviour to obtain a mutually beneficial outcome. We start by discussing the literature on conventions in non-human primates from this perspective and conclude that all the ingredients for conventions to emerge are present and therefore that they ought to be more frequently observed. We then probe the emergence of conventions by using a unique novel experimental system in which pairs of Guinea baboons ( Papio papio ) can voluntarily participate together in touchscreen-based cognitive testing and we show that conventions readily emerge in our experimental set-up and that they share three fundamental properties of human conventions (arbitrariness, stability and efficiency). These results question the idea that observational learning, and imitation in particular, is necessary to establish conventions; they suggest that positive reinforcement is enough. This article is part of a discussion meeting issue ‘The emergence of collective knowledge and cumulative culture in animals, humans and machines’.

Cooperative breeding and the emergence of multilevel societies in birds

Multilevel societies (MLSs), where social levels are hierarchically nested within each other, are considered one of the most complex forms of animal societies. Although thought to mainly occur in mammals, it is suggested that MLSs could be under-detected in birds. Here we propose that the emergence of MLSs could be common in cooperatively breeding birds, as both systems are favoured by similar ecological and social drivers. We first investigate this proposition by systematically comparing evidence for multilevel social structure in cooperative and non-cooperative birds in Australia and New Zealand, global hotspots for cooperative breeding. We then analyse non-breeding social networks of cooperatively breeding superb fairy-wrens (Malurus cyaneus) to reveal their structured multilevel society, with three hierarchical social levels that are stable across years. Our results confirm recent predictions that MLSs are likely to be widespread in birds and suggest that these societies could be particularly common in cooperatively breeding birds.

Submission signals in animal groups

Aggression is costly, and animals have evolved tactics to mitigate these costs. Submission signals are an underappreciated example of such adaptations. Here we review submissive behaviour, with an emphasis on non-primates. We highlight the design of submission signals and how such signals can reduce costs. Animal societies necessitate frequent social interactions, which can increase the probability of conflict. Where maintaining group proximity is essential, animals cannot avoid aggression by fleeing. Mutual interest between group members may also select for efficient conflict avoidance and resolution mechanisms. As a result, submission signals may be especially well developed among group living species, helping social animals to overcome potential costs of recurring conflict that could otherwise counter the benefits of group living. Therefore, submission signalling can be a crucial aspect of social living and is deserving of specific attention within the broader context of social evolution and communication.

The ecology of inequality in animal societies

Individuals vary in their access to resources, social connections, and phenotypic traits, and a central goal of behavioral ecology is to understand how this variation influences reproductive success and longevity. Parallel research on human societies has focused on the causes and consequences of variation in material possessions, opportunity, and health among individuals. At the core in both fields of study is that unequal distribution of benefits is an important component of social structure, but an explicit study of inequality is largely missing from evolutionary biology and ecology. Here we advance a research framework and agenda for studying inequality within an ecological and evolutionary context, drawing upon work in the human-oriented literature where applicable. We present four broad arguments for the ecological study of inequality: (1) wealth and inequality are taxonomically broad features of societies, (2) feedback loops link inequality to individual and societal outcomes, (3) very little is known about what makes some societies more unequal than others, and (4) inequality is dynamic, and these dynamics are relevant for social evolution. We hope that this framework will motivate a cohesive interdisciplinary approach to understanding inequality as a widespread and diverse biological phenomenon.

On the Contribution of Reproductive and Offspring Investment on Fertility: Human and Animal Societies

Differences in investment into reproduction or offspring rearing are plentiful throughout the world, from the cells inside our bodies to complex sociological interactions among humans. Such differences can lead to profound impacts on species' fitness, fertility, and reproductive rates, sometimes in startling ways. In this paper, we create a simple game-theoretical model to qualitatively investigate the effects of such differential investment. We focus on fertility in human societies and show that more wealthy individuals produce more offspring within a a mating group. However, when assortative mating mechanisms are introduced, this effectively leads to a speciation event, and a higher reproduction rate for poorer individuals is noticed, capturing what we call the "wealthy-to-poor switch". We discuss extensions and implications of this work to nupital gifts in ecology and to clonal competition in cancer cell lines under the influence of treatment.

Group size and modularity interact to shape the spread of infection and information through animal societies

Social interactions between animals can provide many benefits, including the ability to gain useful environmental information through social learning. However, these social contacts can also facilitate the transmission of infectious diseases through a population. Animals engaging in social interactions must therefore face a trade-off between the potential informational benefits and the risk of acquiring disease. In order to understand how this trade-off can influence animal sociality, it is necessary to quantify the effects of different social structures on individuals' likelihood of acquiring information versus infection Theoretical models have suggested that modular social networks, associated with the formation of groups or sub-groups, can slow spread of infection by trapping it within particular groups. However these social structures will not necessarily impact the spread of information in the same way if its transmission is considered as a "complex contagion", e.g. through individuals copying the majority (conformist learning). Here we use simulation models to demonstrate that modular networks can promote the spread of information relative to the spread of infection, but only when the network is fragmented and group sizes are small. We show that the difference in transmission between information and disease is maximised for more well-connected social networks when the likelihood of transmission is intermediate. Our results have important implications for understanding the selective pressures operating on the social structure of animal societies, revealing that highly fragmented networks such as those formed in fission-fusion social groups and multilevel societies can be effective in modulating the infection-information trade-off for individuals within them.

Cooperation-based concept formation in male bottlenose dolphins

In Shark Bay, Western Australia, male bottlenose dolphins form a complex nested alliance hierarchy. At the first level, pairs or trios of unrelated males cooperate to herd individual females. Multiple first-order alliances cooperate in teams (second-order alliances) in the pursuit and defence of females, and multiple teams also work together (third-order alliances). Yet it remains unknown how dolphins classify these nested alliance relationships. We use 30 years of behavioural data combined with 40 contemporary sound playback experiments to 14 allied males, recording responses with drone-mounted video and a hydrophone array. We show that males form a first-person social concept of cooperative team membership at the second-order alliance level, independently of first-order alliance history and current relationship strength across all three alliance levels. Such associative concepts develop through experience and likely played an important role in the cooperative behaviour of early humans. These results provide evidence that cooperation-based concepts are not unique to humans, occurring in other animal societies with extensive cooperation between non-kin.