Is peace a human phenomenon?

Peace is a hallmark of human societies. However, certain ant species engage in long-term intergroup resource sharing, which is remarkably similar to peace among human groups. We discuss how individual and group payoff distributions are affected by kinship, dispersal, and age structure; the challenges of diagnosing peace; and the benefits of comparing convergent complex behaviours in disparate taxa.

Competition and habitat availability interact to structure arboreal ant communities across scales of ecological organization.

Understanding how resource limitation and biotic interactions interact across spatial scales is fundamental to explaining the structure of ecological communities. However, empirical studies addressing this issue are often hindered by logistical constraints, especially at local scales. Here, we use a highly tractable arboreal ant study system to explore the interactive effects of resource availability and competition on community structure across three local scales: an individual tree, the nest network created by each colony and the individual ant nest. On individual trees, the ant assemblages are primarily shaped by availability of dead wood, a critical nesting resource. The nest networks within a tree are constrained by the availability of nesting resources but also influenced by the co-occurring species. Within individual nests, the distribution of adult ants is only affected by distance to interspecific competitors. These findings demonstrate that resource limitation exerts the strongest effects on diversity at higher levels of local ecological organization, transitioning to a stronger effect of species interactions at finer scales. Collectively, these results highlight that the process exerting the strongest influence on community structure is highly dependent on the scale at which we examine the community, with shifts occurring even across fine-grained local scales.

Cost, risk, and avoidance of inbreeding in a cooperatively breeding bird.

Inbreeding is often avoided in natural populations by passive processes such as sex-biased dispersal. But, in many social animals, opposite-sexed adult relatives are spatially clustered, generating a risk of incest and hence selection for active inbreeding avoidance. Here we show that, in long-tailed tits (Aegithalos caudatus), a cooperative breeder that risks inbreeding by living alongside opposite-sex relatives, inbreeding carries fitness costs and is avoided by active kin discrimination during mate choice. First, we identified a positive association between heterozygosity and fitness, indicating that inbreeding is costly. We then compared relatedness within breeding pairs to that expected under multiple mate-choice models, finding that pair relatedness is consistent with avoidance of first-order kin as partners. Finally, we show that the similarity of vocal cues offers a plausible mechanism for discrimination against first-order kin during mate choice. Long-tailed tits are known to discriminate between the calls of close kin and nonkin, and they favor first-order kin in cooperative contexts, so we conclude that long-tailed tits use the same kin discrimination rule to avoid inbreeding as they do to direct help toward kin.

From foraging trails to transport networks: how the quality-distance trade-off shapes network structure.

Biological systems are typically dependent on transportation networks for the efficient distribution of resources and information. Revealing the decentralized mechanisms underlying the generative process of these networks is key in our global understanding of their functions and is of interest to design, manage and improve human transport systems. Ants are a particularly interesting taxon to address these issues because some species build multi-sink multi-source transport networks analogous to human ones. Here, by combining empirical field data and modelling at several scales of description, we show that pre-existing mechanisms of recruitment with positive feedback involved in foraging can account for the structure of complex ant transport networks. Specifically, we find that emergent group-level properties of these empirical networks, such as robustness, efficiency and cost, can arise from models built on simple individual-level behaviour addressing a quality-distance trade-off by the means of pheromone trails. Our work represents a first step in developing a theory for the generation of effective multi-source multi-sink transport networks based on combining exploration and positive reinforcement of best sources.

Ant colony nest networks adapt to resource disruption.

Animal social structure is shaped by environmental conditions, such as food availability. This is important as conditions are likely to change in the future and changes to social structure can have cascading ecological effects. Wood ants are a useful taxon for the study of the relationship between social structure and environmental conditions, as some populations form large nest networks and they are ecologically dominant in many northern hemisphere woodlands. Nest networks are formed when a colony inhabits more than one nest, known as polydomy. Polydomous colonies are composed of distinct sub-colonies that inhabit spatially distinct nests and that share resources with each other. In this study, we performed a controlled experiment on 10 polydomous wood ant (Formica lugubris) colonies to test how changing the resource environment affects the social structure of a polydomous colony. We took network maps of all colonies for 5 years before the experiment to assess how the networks changes under natural conditions. After this period, we prevented ants from accessing an important food source for a year in five colonies and left the other five colonies undisturbed. We found that preventing access to an important food source causes polydomous wood ant colony networks to fragment into smaller components and begin foraging on previously unused food sources. These changes were not associated with a reduction in the growth of populations inhabiting individual nests (sub-colonies), foundation of new nests or survival, when compared with control colonies. Colony splitting likely occurred as the availability of food in each nest changed causing sub-colonies to change their inter-nest connections. Consequently, our results demonstrate that polydomous colonies can adjust to environmental changes by altering their social network.

Social tipping points in animal societies.

Animal social groups are complex systems that are likely to exhibit tipping points-which are defined as drastic shifts in the dynamics of systems that arise from small changes in environmental conditions-yet this concept has not been carefully applied to these systems. Here, we summarize the concepts behind tipping points and describe instances in which they are likely to occur in animal societies. We also offer ways in which the study of social tipping points can open up new lines of inquiry in behavioural ecology and generate novel questions, methods, and approaches in animal behaviour and other fields, including community and ecosystem ecology. While some behaviours of living systems are hard to predict, we argue that probing tipping points across animal societies and across tiers of biological organization-populations, communities, ecosystems-may help to reveal principles that transcend traditional disciplinary boundaries.

Fine-scale genetic structure and helping decisions in a cooperatively breeding bird.

In animal societies, characteristic demographic and dispersal patterns may lead to genetic structuring of populations, generating the potential for kin selection to operate. However, even in genetically structured populations, social interactions may still require kin discrimination for cooperative behaviour to be directed towards relatives. Here, we use molecular genetics and long-term field data to investigate genetic structure in an adult population of long-tailed tits Aegithalos caudatus, a cooperative breeder in which helping occurs within extended kin networks, and relate this to patterns of helping with respect to kinship. Spatial autocorrelation analyses reveal fine-scale genetic structure within our population, such that related adults of either sex are spatially clustered following natal dispersal, with relatedness among nearby males higher than that among nearby females, as predicted by observations of male-biased philopatry. This kin structure creates opportunities for failed breeders to gain indirect fitness benefits via redirected helping, but crucially, most close neighbours of failed breeders are unrelated and help is directed towards relatives more often than expected by indiscriminate helping. These findings are consistent with the effective kin discrimination mechanism known to exist in long-tailed tits and support models identifying kin selection as the driver of cooperation.

Inter-group cooperation in humans and other animals.

Social interactions are often characterized by cooperation within groups and conflict or competition between groups. In certain circumstances, however, cooperation can arise between social groups. Here, we examine the circumstances under which inter-group cooperation is expected to emerge and present examples with particular focus on groups in two well-studied but dissimilar taxa: humans and ants. Drivers for the evolution of inter-group cooperation include overarching threats from predators, competitors or adverse conditions, and group-level resource asymmetries. Resources can differ between groups in both quantity and type. Where the difference is in type, inequalities can lead to specialization and division of labour between groups, a phenomenon characteristic of human societies, but rarely seen in other animals. The ability to identify members of one's own group is essential for social coherence; we consider the proximate roles of identity effects in shaping inter-group cooperation and allowing membership of multiple groups. Finally, we identify numerous valuable avenues for future research that will improve our understanding of the processes shaping inter-group cooperation.

How collective comparisons emerge without individual comparisons of the options.

Collective decisions in animal groups emerge from the actions of individuals who are unlikely to have global information. Comparative assessment of options can be valuable in decision-making. Ant colonies are excellent collective decision-makers, for example when selecting a new nest-site. Here, we test the dependency of this cooperative process on comparisons conducted by individual ants. We presented ant colonies with a choice between new nests: one good and one poor. Using individually radio-tagged ants and an automated system of doors, we manipulated individual-level access to information: ants visiting the good nest were barred from visiting the poor one and vice versa. Thus, no ant could individually compare the available options. Despite this, colonies still emigrated quickly and accurately when comparisons were prevented. Individual-level rules facilitated this behavioural robustness: ants allowed to experience only the poor nest subsequently searched more. Intriguingly, some ants appeared particularly discriminating across emigrations under both treatments, suggesting they had stable, high nest acceptance thresholds. Overall, our results show how a colony of ants, as a cognitive entity, can compare two options that are not both accessible by any individual ant. Our findings illustrate a collective decision process that is robust to differences in individual access to information.

Bourgeois behavior and freeloading in the colonial orb web spider Parawixia bistriata (Araneae, Araneidae).

Spiders of the tropical American colonial orb weaver Parawixia bistriata form a communal bivouac in daytime. At sunset, they leave the bivouac and construct individual, defended webs within a large, communally built scaffolding of permanent, thick silk lines between trees and bushes. Once spiders started building a web, they repelled other spiders walking on nearby scaffolding with a "bounce" behavior. In nearly all cases (93%), this resulted in the intruder leaving without a fight, akin to the "bourgeois strategy," in which residents win and intruders retreat without escalated contests. However, a few spiders (6.5%) did not build a web due to lack of available space. Webless spiders were less likely to leave when bounced (only 42% left) and instead attempted to "freeload," awaiting the capture of prey items in nearby webs. Our simple model shows that webless spiders should change their strategy from bourgeois to freeloading satellite as potential web sites become increasingly occupied.

Exploration versus exploitation in polydomous ant colonies.

In socially foraging species resource information can be shared between individuals, increasing foraging success. In ant colonies, nestmate recruitment allows high exploitation rates at known resources however, to maximise foraging efficiency this must be balanced with searching for new resources. Many ant species form colonies inhabiting two or more spatially separated but socially connected nests: this type of organisation is known as polydomy. Polydomous colonies may benefit from increased foraging efficiency by carrying out dispersed-central place foraging. However, decentralisation of the colony may affect recruitment success by limiting interaction between ants based in separate nests. We use an agent-based model which compares the foraging success of monodomous and polydomous colonies in different food environments, incorporating recruitment through pheromone trails and group foraging. In contrast to previous results we show that polydomy is beneficial in some but not all cases. Polydomous colonies discover resources at a higher rate, making them more successful when food is highly dispersed, but their relative success can be lowered by limitations on recruitment success. Monodomous colonies can have higher foraging efficiency than polydomous colonies by exploiting food more rapidly. The results show the importance of interactions between recruitment strategy, colony size, and colony organisation.

The evolution of intergroup cooperation.

Sociality is widespread among animals, and involves complex relationships within and between social groups. While intragroup interactions are often cooperative, intergroup interactions typically involve conflict, or at best tolerance. Active cooperation between members of distinct, separate groups occurs very rarely, predominantly in some primate and ant species. Here, we ask why intergroup cooperation is so rare, and what conditions favour its evolution. We present a model incorporating intra- and intergroup relationships and local and long-distance dispersal. We show that dispersal modes play a pivotal role in the evolution of intergroup interactions. Both long-distance and local dispersal processes drive population social structure, and the costs and benefits of intergroup conflict, tolerance and cooperation. Overall, the evolution of multi-group interaction patterns, including both intergroup aggression and intergroup tolerance, or even altruism, is more likely with mostly localized dispersal. However, the evolution of these intergroup relationships may have significant ecological impacts, and this feedback may alter the ecological conditions that favour its own evolution. These results show that the evolution of intergroup cooperation is favoured by a specific set of conditions, and may not be evolutionarily stable. We discuss how our results relate to empirical evidence of intergroup cooperation in ants and primates. This article is part of a discussion meeting issue 'Collective behaviour through time'.

Experience, corpulence and decision making in ant foraging.

Social groups are structured by the decisions of their members. Social insects typically divide labour: some decide to stay in the nest while others forage for the colony. Two sources of information individuals may use when deciding whether to forage are their own experience of recent task performance and their own physiology, e.g. fat reserves (corpulence). The former is primarily personal information; the latter may give an indication of the food reserves of the whole colony. These factors are hard to separate because typically leaner individuals are also more experienced foragers. We designed an experiment to determine whether foraging specialisation is physiological or experience based (or both). We invented a system of automatic doors controlled by radio-tag information to manipulate task access and decouple these two sources of information. Our results show that when information from corpulence and recent experience conflict, ants behave only in accordance with their corpulence. However, among ants physiologically inclined to forage (less corpulent ants), recent experience of success positively influenced their propensity to forage again. Hence, foraging is organised via long-term physiological differences among individuals resulting in a relatively stable response threshold distribution, with fine-tuning provided by short-term learning processes. Through these simple rules, colonies can organise their foraging effort both robustly and flexibly.

Do ants need to be old and experienced to teach?

Learning is widespread in invertebrates. However, whether social insects improve their recruitment skills with experience is only beginning to be investigated. Tandem running is a one-to-one form of recruitment used by certain species of ant. It is a remarkable communication system that meets widely accepted criteria for teaching in non-human animals. Here, we determined experimentally to what extent participation in, and efficient execution of, tandem running depends on either the age or the experience of worker ants. To investigate these issues, we constructed colonies of the ant Temnothorax albipennis with different compositions of inexperienced and experienced workers from different age cohorts and then examined which ants participated in tandem runs when they emigrated. Our results show that the ability to participate actively in recruitment by tandem running is present in all worker age groups but the propensity to participate varies with experience rather than age per se. Experienced individuals were more likely to engage in tandem runs, either as leaders or as followers, than young inexperienced individuals, and older experienced ants were more likely to lead tandems than older inexperienced ants. Young inexperienced ants led faster, more rapidly dispersing and less accurately orientated tandem runs than the older experienced ants. Our study suggests that experience (rather than age per se) coupled to stimulus threshold responses might interact to promote a division of labour so that a suitable number of workers actively participate in tandem runs.

From inter-group conflict to inter-group cooperation: insights from social insects.

The conflict between social groups is widespread, often imposing significant costs across multiple groups. The social insects make an ideal system for investigating inter-group relationships, because their interaction types span the full harming-helping continuum, from aggressive conflict, to mutual tolerance, to cooperation between spatially separate groups. Here we review inter-group conflict in the social insects and the various means by which they reduce the costs of conflict, including individual or colony-level avoidance, ritualistic behaviours and even group fusion. At the opposite extreme of the harming-helping continuum, social insect groups may peacefully exchange resources and thus cooperate between groups in a manner rare outside human societies. We discuss the role of population viscosity in favouring inter-group cooperation. We present a model encompassing intra- and inter-group interactions, and local and long-distance dispersal. We show that in this multi-level population structure, the increased likelihood of cooperative partners being kin is balanced by increased kin competition, such that neither cooperation (helping) nor conflict (harming) is favoured. This model provides a baseline context in which other intra- and inter-group processes act, tipping the balance toward or away from conflict. We discuss future directions for research into the ecological factors shaping the evolution of inter-group interactions. This article is part of the theme issue 'Intergroup conflict across taxa'.

Insect communication: 'no entry' signal in ant foraging.

Forager ants lay attractive trail pheromones to guide nestmates to food, but the effectiveness of foraging networks might be improved if pheromones could also be used to repel foragers from unrewarding routes. Here we present empirical evidence for such a negative trail pheromone, deployed by Pharaoh's ants (Monomorium pharaonis) as a 'no entry' signal to mark an unrewarding foraging path. This finding constitutes another example of the sophisticated control mechanisms used in self-organized ant colonies.

Helping decisions and kin recognition in long-tailed tits: is call similarity used to direct help towards kin?

Most cooperative breeders live in discrete family groups, but in a minority, breeding populations comprise extended social networks of conspecifics that vary in relatedness. Selection for effective kin recognition may be expected for more related individuals in such kin neighbourhoods to maximize indirect fitness. Using a long-term social pedigree, molecular genetics, field observations and acoustic analyses, we examine how vocal similarity affects helping decisions in the long-tailed tit Aegithalos caudatus. Long-tailed tits are cooperative breeders in which help is typically redirected by males that have failed in their own breeding attempts towards the offspring of male relatives living within kin neighbourhoods. We identify a positive correlation between call similarity and kinship, suggesting that vocal cues offer a plausible mechanism for kin discrimination. Furthermore, we show that failed breeders choose to help males with calls more similar to their own. However, although helpers fine-tune their provisioning rates according to how closely related they are to recipients, their effort was not correlated with their vocal similarity to helped breeders. We conclude that although vocalizations are an important part of the recognition system of long-tailed tits, discrimination is likely to be based on prior association and may involve a combination of vocal and non-vocal cues. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

Flexible task allocation and the organization of work in ants.

Flexibility in task performance is essential for a robust system of division of labour. We investigated what factors determine which social insect workers respond to colony-level changes in task demand. We used radio-frequency identification technology to compare the roles of corpulence, age, spatial location and previous activity (intra-nest/extra-nest) in determining whether worker ants (Temnothorax albipennis) respond to an increase in demand for foraging or brood care. The less corpulent ants took on the extra foraging, irrespective of their age, previous activity or location in the nest, supporting a physiological threshold model. We found no relationship between ants that tended the extra brood and corpulence, age, spatial location or previous activity, but ants that transported the extra brood to the main brood pile were less corpulent and had high previous intra-nest activity. This supports spatial task-encounter and physiological threshold models for brood transport. Our data suggest a flexible task-allocation system allowing the colony to respond rapidly to changing needs, using a simple task-encounter system for generalized tasks, combined with physiologically based response thresholds for more specialized tasks. This could provide a social insect colony with a robust division of labour, flexibly allocating the workforce in response to current needs.

Do ants make direct comparisons?

Many individual decisions are informed by direct comparison of the alternatives. In collective decisions, however, only certain group members may have the opportunity to compare options. Emigrating ant colonies (Temnothorax albipennis) show sophisticated nest-site choice, selecting superior sites even when they are nine times further away than the alternative. How do they do this? We used radio-frequency identification-tagged ants to monitor individual behaviour. Here we show for the first time that switching between nests during the decision process can influence nest choice without requiring direct comparison of nests. Ants finding the poor nest were likely to switch and find the good nest, whereas ants finding the good nest were more likely to stay committed to that nest. When ants switched quickly between the two nests, colonies chose the good nest. Switching by ants that had the opportunity to compare nests had little effect on nest choice. We suggest a new mechanism of collective nest choice: individuals respond to nest quality by the decision either to commit or to seek alternatives. Previously proposed mechanisms, recruitment latency and nest comparison, can be explained as side effects of this simple rule. Colony-level comparison and choice can emerge, without direct comparison by individuals.

The costs and benefits of decentralization and centralization of ant colonies.

A challenge faced by individuals and groups of many species is determining how resources and activities should be spatially distributed: centralized or decentralized. This distribution problem is hard to understand due to the many costs and benefits of each strategy in different settings. Ant colonies are faced by this problem and demonstrate two solutions: 1) centralizing resources in a single nest (monodomy) and 2) decentralizing by spreading resources across many nests (polydomy). Despite the possibilities for using this system to study the centralization/decentralization problem, the trade-offs associated with using either polydomy or monodomy are poorly understood due to a lack of empirical data and cohesive theory. Here, we present a dynamic network model of a population of ant nests which is based on observations of a facultatively polydomous ant species (Formica lugubris). We use the model to test several key hypotheses for costs and benefits of polydomy and monodomy and show that decentralization is advantageous when resource acquisition costs are high, nest size is limited, resources are clustered, and there is a risk of nest destruction, but centralization prevails when resource availability fluctuates and nest size is limited. Our model explains the phylogenetic and ecological diversity of polydomous ants, demonstrates several trade-offs of decentralization and centralization, and provides testable predictions for empirical work on ants and in other systems.