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Micro-society experimental setups are increasingly used to infer aspects of human behavioural evolution. A key part of human society today is our dependence on, and use of, technology-whether simple (such as a knife) or complex (such as the technology that underpins AI). Previously, two groups of researchers used an abstracted-wheel experiment to explore the evolution of human technical behaviour, reaching fundamentally different outcomes. Whereas one group saw their results as indicating social learning only (void of causal understanding), the other inferred non-social technical reasoning as part of human technical behaviour. Here we report on the third generation of the micro-society abstracted-wheel experiment. We argue that causal reasoning is inseparable from both social learning and technical reasoning, and that these traits probably co-evolved into the current human socio-technical niche. Based on our outcomes, we present a critical assessment of what this experiment may (or may not) reveal about the evolution of human technical behaviour. We show that the abstracted-wheel experiment reflects behavioural output only, instead of testing for cognition. It is therefore limited in its ability to inform on aspects of human cognitive evolution, but it can provide useful insights into the interrelatedness of social learning, technical reasoning, and causal reasoning. Such a co-evolutionary insight has the potential to inform on aspects of human socio-technical evolution throughout the Pleistocene., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Högberg et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Behavioural variation among individuals is a hallmark of cooperative societies, which commonly contain breeders and non-breeders, helpers and non-helpers. In some cases, labour is divided with non-breeders "helping." Conversely, in some societies, subordinate non-breeders may not help. These individuals may be (i) an insurance workforce ensuring continuity of help for breeders when other helpers are lost, (ii) conserving energy while waiting to breed themselves or (iii) simply of too poor physiological quality either to help or breed. In the Australian Outback, Acacia thrips Dunatothrips aneurae (Thysanoptera) glue Acacia phyllodes into "domiciles" using silk-like secretions, either alone or cooperatively. Domicile maintenance is important for humidity, so repair can be interpreted as helping. I found that not all females helped to repair damage; some repaired partially or not at all ("non-helpers"). At the same time, some co-foundresses are non- or partially reproductive ("non-breeders"), and their role is currently unknown. I first tested the possibility that helping and breeding are divided, with non-helping females breeding, and non-breeders helping. In a lab experiment, I rejected this idea. Experimentally damaged domiciles were typically repaired by reproductive females, and less so by non- or partially reproductive individuals. To test whether non-helpers are an insurance workforce, I successively removed repairing females and found that non-helping females from the same domicile did not increase effort as a result, rejecting this hypothesis. Then I tested whether non-helpers were conserving energy waiting to breed. In a field experiment, I removed all other females, allowing either a helpful female or a non-helper to "inherit" her domicile. Isolated like this, non-helpers laid very few eggs compared to helpers or naturally occurring single foundresses, despite similar ovarian development. My findings show that labour was not divided: reproduction and helping covaried positively, probably depending on individual variation in female quality and intra-domicile competition. Non-helping females were neither an insurance workforce nor conserving energy waiting to breed. They are likely simply of poor quality, freeloading by benefiting from domicile maintenance by others. I hypothesize they are tolerated because of selection for indiscriminate communal brood care in the form of domicile repair., (© 2024 The Author(s). Journal of Animal Ecology © 2024 British Ecological Society.)

Cooperatively breeding societies show distinct interspecific variations in social and genetic organization. Long-term studies provide invaluable data to further our understanding of the evolution and maintenance of cooperative breeding but have also demonstrated how variation exists within species. Here we integrate life-history, behavioural and genetic data from a long-term study of dwarf mongooses Helogale parvula in South Africa to document mating, breeding, dispersal and relatedness patterns in this population and compare them to those found in a Tanzanian population at the other extreme of the species' range. Our genetic data reveal high levels of reproductive skew, above that expected through observational data. Dispersal was male-biased and was seen more frequently towards the onset of the breeding season, but females also regularly switched between groups. These patterns of breeding and dispersal resulted in a genetically structured population: individuals were more related to groupmates than outsiders, apart from the unrelated dominant pair, ultimately resulting in reduced inbreeding risk. Our results also demonstrate that dwarf mongooses are largely consistent in their social structure across their sub-Saharan distribution. This work demonstrates the direct and indirect pathways to reproductive success for dwarf mongooses and helps to explain the maintenance of cooperative breeding in the species., Competing Interests: We declare we have no competing interests., (© 2024 The Author(s).)

Hamilton's rule provides the cornerstone for our understanding of the evolution of all forms of social behavior, from altruism to spite, across all organisms, from viruses to humans. In contrast to the standard prediction from Hamilton's rule, recent studies have suggested that altruistic helping can be favored even if it does not benefit relatives, as long as it decreases the environmentally induced variance of their reproductive success ("altruistic bet-hedging"). However, previous predictions both rely on an approximation and focus on variance-reducing helping behaviors. We derived a version of Hamilton's rule that fully captures environmental variability. This shows that decreasing (or increasing) the variance in the absolute reproductive success of relatives does not have a consistent effect-it can either favor or disfavor the evolution of helping. We then empirically quantified the effect of helping on the variance in reproductive success across 15 species of cooperatively breeding birds. We found that a) helping did not consistently decrease the variance of reproductive success and often increased it, and b) the mean benefits of helping across environments consistently outweighed other variability components of reproductive success. Altogether, our theoretical and empirical results suggest that the effects of helping on the variability components of reproductive success have not played a consistent or strong role in favoring helping., Competing Interests: Competing interests statement:The authors declare no competing interest.

Competing Interests: Competing interests statement:The author declares no competing interest.

The rapid increase of antibiotic-resistant pathogens is severely limiting our current treatment possibilities. An important subset of the resistance mechanisms conferring antibiotic resistance have public effects, allowing otherwise susceptible bacteria to also survive antibiotic treatment. As susceptible bacteria can survive treatment without bearing the metabolic cost of producing the resistance mechanism, there is potential to increase their relative frequency in the population and, as such, select against resistant bacteria. Multiple studies showed that this altered selection for resistance is dependent on various environmental and treatment parameters. In this review, we provide a comprehensive overview of their most important findings and describe the main factors impacting the selection for resistance. In-depth understanding of the driving forces behind selection can aid in the design and implementation of alternative treatments which limit the risk of resistance development.

Models of indirect reciprocity study how social norms promote cooperation. In these models, cooperative individuals build up a positive reputation, which in turn helps them in their future interactions. The exact reputational benefits of cooperation depend on the norm in place, which may change over time. Previous research focused on the stability of social norms. Much less is known about how social norms initially evolve when competing with many others. A comprehensive evolutionary analysis, however, has been difficult. Even among the comparably simple space of so-called third-order norms, there are thousands of possibilities, each one inducing its own reputation dynamics. To address this challenge, we use large-scale computer simulations. We study the reputation dynamics of each third-order norm and all evolutionary transitions between them. In contrast to established work with only a handful of norms, we find that cooperation is hard to maintain in well-mixed populations. However, within group-structured populations, cooperation can emerge. The most successful norm in our simulations is particularly simple. It regards cooperation as universally positive, and defection as usually negative-unless defection takes the form of justified punishment. This research sheds light on the complex interplay of social norms, their induced reputation dynamics, and population structure., Competing Interests: Competing interests statement:The authors declare no competing interest.

We proposed a neighbor selection mechanism based on memory and target payoff, where the target payoff is the maximum value of the group's average expected payoff. According to this mechanism, individuals prioritize selecting neighbors whose average payoffs in the last M rounds are close to the target payoff for strategy learning, aiming to maximize the group's expected payoff. Simulation results on the grid-based Prisoner's Dilemma and Snowdrift games demonstrate that this mechanism can significantly improve the group's payoff and cooperation level. Furthermore, the longer the memory length, the higher the group's payoff and cooperation level. Overall, the combination of memory and target payoff can lead to the emergence and persistence of cooperation in social dilemmas as individuals are motivated to cooperate based on both their past experiences and future goals. This interplay highlights the significance of taking into account numerous variables in comprehending and promoting cooperation within evolutionary frameworks., (© 2024 Author(s). Published under an exclusive license by AIP Publishing.)

Resistance to and avoidance of stress slow aging and confer increased longevity in numerous organisms. Honey bees and other superorganismal social insects have two main advantages over solitary species to avoid or resist stress: individuals can directly help each other by resource or information transfer, and they can cooperatively control their environment. These benefits have been recognised in the context of pathogen and parasite stress as the concept of social immunity, which has been extensively studied. However, we argue that social immunity is only a special case of a general concept that we define here as social stress protection to include group-level defences against all biotic and abiotic stressors. We reason that social stress protection may have allowed the evolution of reduced individual-level defences and individual life-history optimization, including the exceptional aging plasticity of many social insects. We describe major categories of stress and how a colonial lifestyle may protect social insects, particularly against temporary peaks of extreme stress. We use the honey bee (Apis mellifera L.) to illustrate how patterns of life expectancy may be explained by social stress protection and how modern beekeeping practices can disrupt social stress protection. We conclude that the broad concept of social stress protection requires rigorous empirical testing because it may have implications for our general understanding of social evolution and specifically for improving honey bee health., (© 2024 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Social insect workers, renowned for their altruism, are frequently perceived as 'disposable'. A new study finds that ants amputate the limbs of nestmates, which saves them from infection, and indicates that worker care is as critical to colony success as sacrifice., Competing Interests: Declaration of interests The author declares no competing interests., (Crown Copyright © 2024. Published by Elsevier Inc. All rights reserved.)

Understanding the influence of social interactions on individual fitness is key to improving our predictions of phenotypic evolution. However, we often overlook the different components of selection regimes arising from interactions among organisms, including social, correlational, and indirect selection. This is due to the challenging sampling efforts required in natural populations to measure phenotypes expressed during interactions and individual fitness. Furthermore, behaviours are crucial in mediating social interactions, yet few studies have explicitly quantified these selection components on behavioural traits. In this study, we capitalize on an online multiplayer video game as a source of extensive data recording direct social interactions among prey, where prey collaborate to escape a predator in realistic ecological settings. We estimate natural and social selection and their contribution to total selection on behavioural traits mediating competition, cooperation, and predator-prey interactions. Behaviours of other prey in a group impact an individual's survival, and thus are under social selection. Depending on whether selection pressures on behaviours are synergistic or conflicting, social interactions enhance or mitigate the strength of natural selection, although natural selection remains the main driving force. Indirect selection through correlations among traits also contributed to the total selection. Thus, failing to account for the effects of social interactions and indirect selection would lead to a misestimation of the total selection acting on traits. Dissecting the contribution of each component to the total selection differential allowed us to investigate the causal mechanisms relating behaviour to fitness and quantify the importance of the behaviours of conspecifics as agents of selection. Our study emphasizes that social interactions generate complex selective regimes even in a relatively simple ecological environment., (© The Author(s) 2024. Published by Oxford University Press on behalf of the European Society of Evolutionary Biology.)

Individuals delay natal dispersal for many reasons. There may be no place to disperse to; immediate dispersal or reproduction may be too costly; immediate dispersal may mean that the individual and their relatives miss the benefits of group living. Understanding the factors that lead to the evolution of delayed dispersal is important because delayed dispersal sets the stage for complex social groups and social behavior. Here, we study the evolution of delayed dispersal when the quality of the local environment is improved by greater numbers of individuals (e.g., safety in numbers). We assume that individuals who delay natal dispersal also expect to delay personal reproduction. In addition, we assume that improved environmental quality benefits manifest as changes to fecundity and survival. We are interested in how do the changes in these life-history features affect delayed dispersal. We use a model that ties evolution to population dynamics. We also aim to understand the relationship between levels of delayed dispersal and the probability of establishing as an independent breeder (a population-level feature) in response to changes in life-history details. Our model emphasizes kin selection and considers a sexual organism, which allows us to study parent-offspring conflict over delayed dispersal. At evolutionary equilibrium, fecundity and survival benefits of group size or quality promote higher levels of delayed dispersal over a larger set of life histories with one exception. The exception is for benefits of increased group size or quality reaped by the individuals who delay dispersal. There, the increased benefit does not change the life histories supporting delay dispersal. Next, in contrast to previous predictions, we find that a low probability of establishing in a new location is not always associated with a higher incidence of delayed dispersal. Finally, we find that increased personal benefits of delayed dispersal exacerbate the conflict between parents and their offspring. We discuss our findings in relation to previous theoretical and empirical work, especially work related to cooperative breeding., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Sociality underpins major evolutionary transitions and significantly influences the structure and function of complex ecosystems. Social insects, seen as the pinnacle of sociality, have traits like obligate sterility that are considered 'master traits', used as single phenotypic measures of this complexity. However, evidence is mounting that completely aligning both phenotypic and evolutionary social complexity, and having obligate sterility central to both, is erroneous. We hypothesize that obligate and functional sterility are insufficient in explaining the diversity of phenotypic social complexity in social insects. To test this, we explore the relative importance of these sterility traits in an understudied but diverse taxon: the termites. We compile the largest termite social complexity dataset to date, using specimen and literature data. We find that although functional and obligate sterility explain a significant proportion of variance, neither trait is an adequate singular proxy for the phenotypic social complexity of termites. Further, we show both traits have only a weak association with the other social complexity traits within termites. These findings have ramifications for our general comprehension of the frameworks of phenotypic and evolutionary social complexity, and their relationship with sterility.

A long-standing problem in evolutionary theory is to clarify in what sense (if any) natural selection cumulatively improves the design of organisms. Various concepts, such as fitness and inclusive fitness, have been proposed to resolve this problem. In addition, there have been attempts to replace the original problem with more tractable questions, such as whether a given gene or trait is favored by selection. Here, we ask what theoretical properties the concept fitness should possess to encapsulate the improvement criterion required to talk meaningfully about adaptive evolution. We argue that natural selection tends to shape phenotypes based on the causal properties of individuals and that this tendency is, therefore, best captured by a fitness concept that focuses on these properties. We highlight a fitness concept that meets this role under broad conditions but requires adjustments in our conceptual understanding of adaptive evolution. These adjustments combine elements of Dawkinsian gene selectionism and Egbert Leigh's "parliament of genes.", (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE).)

Social reputations facilitate cooperation: those who help others gain a good reputation, making them more likely to receive help themselves. But when people hold private views of one another, this cycle of indirect reciprocity breaks down, as disagreements lead to the perception of unjustified behavior that ultimately undermines cooperation. Theoretical studies often assume population-wide agreement about reputations, invoking rapid gossip as an endogenous mechanism for reaching consensus. However, the theory of indirect reciprocity lacks a mechanistic description of how gossip actually generates consensus. Here, we develop a mechanistic model of gossip-based indirect reciprocity that incorporates two alternative forms of gossip: exchanging information with randomly selected peers or consulting a single gossip source. We show that these two forms of gossip are mathematically equivalent under an appropriate transformation of parameters. We derive an analytical expression for the minimum amount of gossip required to reach sufficient consensus and stabilize cooperation. We analyze how the amount of gossip necessary for cooperation depends on the benefits and costs of cooperation, the assessment rule (social norm), and errors in reputation assessment, strategy execution, and gossip transmission. Finally, we show that biased gossip can either facilitate or hinder cooperation, depending on the direction and magnitude of the bias. Our results contribute to the growing literature on cooperation facilitated by communication, and they highlight the need to study strategic interactions coupled with the spread of social information., Competing Interests: Competing interests statement:The authors declare no competing interest.

Greenbeards are selfish genetic elements that make their bearers behave either altruistically towards individuals bearing similar greenbeard copies or harmfully towards individuals bearing different copies. They were first proposed by W. D. Hamilton over 50 yr ago, to illustrate that kin selection may operate at the level of single genes. Examples of greenbeards have now been reported in a wide range of taxa, but they remain undocumented in plants. In this paper, we discuss the theoretical likelihood of greenbeard existence in plants. We then question why the greenbeard concept has never been applied to plants and speculate on how hypothetical greenbeards could affect plant-plant interactions. Finally, we point to different research directions to improve our knowledge of greenbeards in plants., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Across evolutionary lineages, insects vary in social complexity, from those that exhibit extended parental care to those with elaborate divisions of labor. Here, we synthesize the sociogenomic resources from hundreds of species to describe common gene regulatory mechanisms in insects that regulate social organization across phylogeny and levels of social complexity. Different social phenotypes expressed by insects can be linked to the organization of co-expressing gene networks and features of the epigenetic landscape. Insect sociality also stems from processes like the emergence of parental care and the decoupling of ancestral genetic programs. One underexplored avenue is how variation in a group's social environment affects the gene expression of individuals. Additionally, an experimental reduction of gene expression would demonstrate how the activity of specific genes contributes to insect social phenotypes. While tissue specificity provides greater localization of the gene expression underlying social complexity, emerging transcriptomic analysis of insect brains at the cellular level provides even greater resolution to understand the molecular basis of social insect evolution., Competing Interests: The authors declare there are no competing interests.

Organisms from microbes to humans engage in a variety of social behaviors, which affect fitness in complex, often nonlinear ways. The question of how these behaviors evolve has consequences ranging from antibiotic resistance to human origins. However, evolution with nonlinear social interactions is challenging to model mathematically, especially in combination with spatial, group, and/or kin assortment. We derive a mathematical condition for natural selection with synergistic interactions among any number of individuals. This result applies to populations with arbitrary (but fixed) spatial or network structure, group subdivision, and/or mating patterns. In this condition, nonlinear fitness effects are ascribed to collectives, and weighted by a new measure of collective relatedness. For weak selection, this condition can be systematically evaluated by computing branch lengths of ancestral trees. We apply this condition to pairwise games between diploid relatives, and to dilemmas of collective help or harm among siblings and on spatial networks. Our work provides a rigorous basis for extending the notion of "actor", in the study of social evolution, from individuals to collectives., (© The Author(s) 2024. Published by Oxford University Press on behalf of National Academy of Sciences.)

The neurogenomic mechanisms mediating male-male reproductive cooperative behaviours remain unknown. We leveraged extensive transcriptomic and behavioural data on a neotropical bird species (Pipra filicauda) that performs cooperative courtship displays to understand these mechanisms. In this species, the cooperative display is modulated by testosterone, which promotes cooperation in non-territorial birds, but suppresses cooperation in territory holders. We sought to understand the neurogenomic underpinnings of three related traits: social status, cooperative display behaviour and testosterone phenotype. To do this, we profiled gene expression in 10 brain nuclei spanning the social decision-making network (SDMN), and two key endocrine tissues that regulate social behaviour. We associated gene expression with each bird's behavioural and endocrine profile derived from 3 years of repeated measures taken from free-living birds in the Ecuadorian Amazon. We found distinct landscapes of constitutive gene expression were associated with social status, testosterone phenotype and cooperation, reflecting the modular organization and engagement of neuroendocrine tissues. Sex-steroid and neuropeptide signalling appeared to be important in mediating status-specific relationships between testosterone and cooperation, suggesting shared regulatory mechanisms with male aggressive and sexual behaviours. We also identified differentially regulated genes involved in cellular activity and synaptic potentiation, suggesting multiple mechanisms underpin these genomic states. Finally, we identified SDMN-wide gene expression differences between territorial and floater males that could form the basis of 'status-specific' neurophysiological phenotypes, potentially mediated by testosterone and growth hormone. Overall, our findings provide new, systems-level insights into the mechanisms of cooperative behaviour and suggest that differences in neurogenomic state are the basis for individual differences in social behaviour., (© 2024 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Crozier's paradox suggests that genetic kin recognition will not be evolutionarily stable. The problem is that more common tags (markers) are more likely to be recognised and helped. This causes common tags to increase in frequency, eliminating the genetic variability that is required for genetic kin recognition. In recent years, theoretical models have resolved Crozier's paradox in different ways, but they are based on very complicated multi-locus population genetics. Consequently, it is hard to see exactly what is going on, and whether different theoretical resolutions of Crozier's paradox lead to different types of kin discrimination. I address this by making unrealistic simplifying assumptions to produce a more tractable and understandable model of Crozier's paradox. I use this to interpret a more complex multi-locus population genetic model where I have not made the same simplifying assumptions. I explain how Crozier's paradox can be resolved, and show that only one known theoretical resolution of Crozier's paradox - multiple social encounters - leads without restrictive assumptions to the type of highly cooperative and reliable form of kin discrimination that we observe in nature. More generally, I show how adopting a methodological approach where complex models are compared with simplified ones can lead to greater understanding and accessibility., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

We review theoretical approaches for modelling the origin, persistence and change of social norms. The most comprehensive models describe the coevolution of behaviours, personal, descriptive and injunctive norms while considering influences of various authorities and accounting for cognitive processes and between-individual differences. Models show that social norms can improve individual and group well-being. Under some conditions though, deleterious norms can persist in the population through conformity, preference falsification and pluralistic ignorance. Polarization in behaviour and beliefs can be maintained, even when societal advantages of particular behaviours or belief systems over alternatives are clear. Attempts to change social norms can backfire through cognitive processes including cognitive dissonance and psychological reactance. Under some conditions social norms can change rapidly via tipping point dynamics. Norms can be highly susceptible to manipulation, and network structure influences their propagation. Future models should incorporate network structure more thoroughly, explicitly study online norms, consider cultural variations and be applied to real-world processes. This article is part of the theme issue 'Social norm change: drivers and consequences'.

Humans operate in groups that are oftentimes nested in multilayered collectives such as work units within departments and companies, neighborhoods within cities, and regions within nation states. With psychological science mostly focusing on proximate reasons for individuals to join existing groups and how existing groups function, we still poorly understand why groups form ex nihilo, how groups evolve into complex multilayered social structures, and what explains fission-fusion dynamics. Here we address group formation and the evolution of social organization at both the proximate and ultimate level of analysis. Building on models of fitness interdependence and cooperation, we propose that socioecologies can create positive interdependencies among strangers and pave the way for the formation of stable coalitions and groups through reciprocity and reputation-based partner selection. Such groups are marked by in-group bounded, parochial cooperation together with an array of social institutions for managing the commons, allowing groups to scale in size and complexity while avoiding the breakdown of cooperation. Our analysis reveals how distinct group cultures can endogenously emerge from reciprocal cooperation, shows that social identification and group commitment are likely consequences rather than causes of group cooperation, and explains when intergroup relations gravitate toward peaceful coexistence, integration, or conflict.

Cooperation is prevalent across bacteria, but risks being exploited by non-cooperative cheats. Horizontal gene transfer, particularly via plasmids, has been suggested as a mechanism to stabilize cooperation. A key prediction of this hypothesis is that genes which are more likely to be transferred, such as those on plasmids, should be more likely to code for cooperative traits. Testing this prediction requires identifying all genes for cooperation in bacterial genomes. However, previous studies used a method which likely misses some of these genes for cooperation. To solve this, we used a new genomics tool, SOCfinder, which uses three distinct modules to identify all kinds of genes for cooperation. We compared where these genes were located across 4648 genomes from 146 bacterial species. In contrast to the prediction of the hypothesis, we found no evidence that plasmid genes are more likely to code for cooperative traits. Instead, we found the opposite-that genes for cooperation were more likely to be carried on chromosomes. Overall, the vast majority of genes for cooperation are not located on plasmids, suggesting that the more general mechanism of kin selection is sufficient to explain the prevalence of cooperation across bacteria.

Gene transfer agents (GTAs) are enigmatic elements that resemble small viruses and are known to be produced during nutritional stress by some bacteria and archaea. The production of GTAs is regulated by quorum sensing, under which a small fraction of the population acts as GTA producers, while the rest becomes GTA recipients. In contrast to canonical viruses, GTAs cannot propagate themselves because they package pieces of the producing cell's genome. In alphaproteobacteria, GTAs are mostly vertically inherited and reside in their hosts' genomes for hundreds of millions of years. While GTAs' ability to transfer genetic material within a population and their long-term preservation suggest an increased fitness of GTA-producing microbes, the associated benefits and type of selection that maintains GTAs are poorly understood. By comparing rates of evolutionary change in GTA genes to the rates in gene families abundantly present across 293 alphaproteobacterial genomes, we detected 59 gene families that likely co-evolve with GTA genes. These gene families are predominantly involved in stress response, DNA repair, and biofilm formation. We hypothesize that biofilm formation enables the physical proximity of GTA-producing cells, limiting GTA-derived benefits only to a group of closely related cells. We further conjecture that the population structure of biofilm-forming sub-populations ensures that the trait of GTA production is maintained despite the inevitable rise of "cheating" genotypes. Because release of GTA particles kills the producing cell, maintenance of GTAs is an exciting example of social evolution in a microbial population.IMPORTANCEGene transfer agents (GTAs) are viruses domesticated by some archaea and bacteria as vehicles for carrying pieces of the host genome. Produced under certain environmental conditions, GTA particles can deliver DNA to neighboring, closely related cells. The function of GTAs remains uncertain. While making GTAs is suicidal for a cell, GTA-encoding genes are widespread in genomes of alphaproteobacteria. Such GTA persistence implies functional benefits but raises questions about how selection maintains this lethal trait. By showing that GTA genes co-evolve with genes involved in stress response, DNA repair, and biofilm formation, we provide support for the hypothesis that GTAs facilitate DNA exchange during the stress conditions and present a model for how GTAs persist in biofilm-forming bacterial populations despite being lethal., Competing Interests: The authors declare no conflict of interest.

Much research on the evolution of altruism via kin selection, group selection, and reciprocity focuses on the role of a single locus or quantitative trait. Very few studies have explored how linked selection, or selection at loci neighboring an altruism locus, impacts the evolution of altruism. While linked selection can decrease the efficacy of selection at neighboring loci, it might have other effects including promoting selection for altruism by increasing relatedness in regions of low recombination. Here, we used population genetic simulations to study how negative selection at linked loci, or background selection, affects the evolution of altruism. When altruism occurs between full siblings, we found that background selection interfered with selection on the altruistic allele, increasing its fixation probability when the altruistic allele was disfavored and reducing its fixation when the allele was favored. In other words, background selection has the same effect on altruistic genes in family-structured populations as it does on other, nonsocial, genes. This contrasts with prior research showing that linked selective sweeps can favor the evolution of cooperation, and we discuss possibilities for resolving these contrasting results., Competing Interests: The authors declare no conflicts of interest., (© 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Many species of microbes cooperate by producing public goods from which they collectively benefit. However, these populations are under the risk of being taken over by cheating mutants that do not contribute to the pool of public goods. Here we present theoretical findings that address how the social evolution of microbes can be manipulated by external perturbations to inhibit or promote the fixation of cheaters. To control social evolution, we determine the effects of fluid-dynamical properties such as flow rate or domain geometry. We also study the social evolutionary consequences of introducing beneficial or harmful chemicals at steady state and in a time-dependent fashion. We show that by modulating the flow rate and by applying pulsed chemical signals, we can modulate the spatial structure and dynamics of the population in a way that can select for more or less cooperative microbial populations., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Biophysical Society. Published by Elsevier Inc. All rights reserved.)

The dynamics of animal social structures are heavily influenced by environmental patterns of competition and cooperation. In folivorous colobine primates, prevailing theories suggest that larger group sizes should be favored in rainforests with a year-round abundance of food, thereby reducing feeding competition. Yet, paradoxically, larger groups are frequently found in high-altitude or high-latitude montane ecosystems characterized by a seasonal scarcity of leaves. This contradiction is posited to arise from cooperative benefits in heterogeneous environments. To investigate this hypothesis, we carried out a six-year field study on two neighboring groups of golden snub-nosed monkey ( Rhinopithecus roxellana ), a species representing the northernmost distribution of colobine primates. Results showed that the groups adjusted their movement and habitat selection in response to fluctuating climates and spatiotemporal variability of resources, indicative of a dynamic foraging strategy. Notably, during the cold, resource-scarce conditions in winter, the large group occupied food-rich habitats but did not exhibit significantly longer daily travel distances than the smaller neighboring group. Subsequently, we compiled an eco-behavioral dataset of 52 colobine species to explore their evolutionary trajectories. Analysis of this dataset suggested that the increase in group size may have evolved via home range expansion in response to the cold and heterogeneous climates found at higher altitudes or latitudes. Hence, we developed a multi-benefits framework to interpret the formation of larger groups by integrating environmental heterogeneity. In cold and diverse environments, even smaller groups require larger home ranges to meet their dynamic survival needs. The spatiotemporal distribution of high-quality resources within these expanded home ranges facilitates more frequent interactions between groups, thereby encouraging social aggregation into larger groups. This process enhances the benefits of collaborative actions and reproductive opportunities, while simultaneously optimizing travel costs through a dynamic foraging strategy.

In the field of social evolution, inclusive fitness theory has been successful in making a wide range of qualitative predictions on expected patterns of cooperation and conflict. Nevertheless, outside of sex ratio theory, inclusive fitness models that make accurate quantitative predictions remain relatively rare. Past models dealing with caste fate conflict in insect societies, for example, successfully predicted that if female larvae can control their own caste fate, an excess should opt to selfishly develop as queens. Available models, however, were unable to accurately predict levels of queen production observed in Melipona bees-a genus of stingless bees where caste is self-determined-as empirically observed levels of queen production are approximately two times lower than the theoretically predicted ones. Here, we show that this discrepancy can be resolved by explicitly deriving the colony-level cost of queen overproduction from a dynamic model of colony growth, requiring the incorporation of parameters of colony growth and demography, such as the per-capita rate at which new brood cells are built and provisioned, the percentage of the queen's eggs that are female, costs linked with worker reproduction and worker mortality. Our revised model predicts queen overproduction to more severely impact colony productivity, resulting in an evolutionarily stable strategy that is approximately half that of the original model, and is shown to accurately predict actual levels of queen overproduction observed in different Melipona species. Altogether, this shows how inclusive fitness models can provide accurate quantitative predictions, provided that costs and benefits are modeled in sufficient detail and are measured precisely., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE) and European Society for Evolutionary Biology (ESEN).)

The size-complexity rule posits that the evolution of larger cooperative groups should favour more division of labour. Examples include more cell types in larger multicellular organisms, and more polymorphic castes in larger eusocial colonies. However, a correlation between division of labour and group size may reflect a shared response of both traits to resource availability and/or profitability. Here, this possibility was addressed by investigating the evolution of sterile caste number (worker and soldier morphotypes) in termites, a major clade of eusocial insects in which the drivers of caste polymorphism are poorly understood. A novel dataset on 90 termite species was compiled from the published literature. The analysis showed that sterile caste number did increase markedly with colony size. However, after controlling for resource adaptations and phylogeny, there was no evidence for this relationship. Rather, sterile caste number increased with increasing nest-food separation and decreased with soil-feeding, through changes in worker (but not soldier) morphotype number. Further, colony size increased with nest-food separation, thus driving the false correlation between sterile caste number and colony size. These findings support adaptation to higher energy acquisition as key to the rise of complex insect societies, with larger size being a by-product.

Fisher's geometric model provides a powerful tool for making predictions about key properties of Darwinian adaptation. Here, I apply the geometric model to predict differences between the evolution of altruistic versus nonsocial phenotypes. I recover Kimura's prediction that probability of fixation is greater for mutations of intermediate size, but I find that the effect size that maximises probability of fixation is relatively small in the context of altruism and relatively large in the context of nonsocial phenotypes, and that the overall probability of fixation is lower for altruism and is higher for nonsocial phenotypes. Accordingly, the first selective substitution is expected to be smaller, and to take longer, in the context of the evolution of altruism. These results strengthen the justification for employing streamlined social evolutionary methodologies that assume adaptations are underpinned by many genes of small effect., Competing Interests: Declaration of Competing Interest The author declares that he has no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Although social interactions are known to drive pathogen transmission, the contributions of socially transmissible host-associated mutualists and commensals to host health and disease remain poorly explored. We use the concept of the social microbiome-the microbial metacommunity of a social network of hosts-to analyze the implications of social microbial transmission for host health and disease. We investigate the contributions of socially transmissible microbes to both eco-evolutionary microbiome community processes (colonization resistance, the evolution of virulence, and reactions to ecological disturbance) and microbial transmission-based processes (transmission of microbes with metabolic and immune effects, inter-specific transmission, transmission of antibiotic-resistant microbes, and transmission of viruses). We consider the implications of social microbial transmission for communicable and non-communicable diseases and evaluate the importance of a socially transmissible component underlying canonically non-communicable diseases. The social transmission of mutualists and commensals may play a significant, under-appreciated role in the social determinants of health and may act as a hidden force in social evolution., Competing Interests: Declaration of interests A.S., C.J.A.M., A.R., N.G.O.I., K.V.-A.J., M.V.-C., I.L.B., J.H., E.A.A., L.B.B., B.B.F., R.N.C., and A.H.M. declare no competing interests. S.H. is a current employee of SilverCloud Health and SilverCloud Health is a subsidiary of Amwell. S.H. holds shares in Amwell. F.S.G. is listed as an inventor on patent applications pertaining to the treatment of cancer or the study of the human microbiome. E.V.K. is listed as an inventor on patents and patent applications pertaining to CRISPR enzymes and systems or protein regulation., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Many microbes interact with one another, but the difficulty of directly observing these interactions in nature makes interpreting their adaptive value complicated. The social amoeba Dictyostelium discoideum forms aggregates wherein some cells are sacrificed for the benefit of others. Within chimaeric aggregates containing multiple unrelated lineages, cheaters can gain an advantage by undercontributing, but the extent to which wild D. discoideum has adapted to cheat is not fully clear. In this study, we experimentally evolved D. discoideum in an environment where there were no selective pressures to cheat or resist cheating in chimaeras. Dictyostelium discoideum lines grown in this environment evolved reduced competitiveness within chimaeric aggregates and reduced ability to migrate during the slug stage. By contrast, we did not observe a reduction in cell number, a trait for which selection was not relaxed. The observed loss of traits that our laboratory conditions had made irrelevant suggests that these traits were adaptations driven and maintained by selective pressures D. discoideum faces in its natural environment. Our results suggest that D. discoideum faces social conflict in nature, and illustrate a general approach that could be applied to searching for social or non-social adaptations in other microbes.

The ability to recognize others is a frequent assumption of models of the evolution of cooperation. At the same time, cooperative behavior has been proposed as a selective agent favoring the evolution of individual recognition abilities. Although theory predicts that recognition and cooperation may co-evolve, data linking recognition abilities and cooperative behavior with evidence of selection are elusive. Here, we provide evidence of a selective link between individual recognition and cooperation in the paper wasp Polistes fuscatus through a combination of clinal, common garden, and population genomics analyses. We identified latitudinal clines in both rates of cooperative nesting and color pattern diversity, consistent with a selective link between recognition and cooperation. In behavioral experiments, we replicated previous results demonstrating individual recognition in cooperative and phenotypically diverse P. fuscatus from New York. In contrast, wasps from a less cooperative and phenotypically uniform Louisiana population showed no evidence of individual recognition. In a common garden experiment, groups of wasps from northern populations formed more stable and individually biased associations, indicating that recognition facilitates group stability. The strength of recent positive selection on cognition-associated loci likely to mediate individual recognition is substantially greater in northern compared with southern P. fuscatus populations. Collectively, these data suggest that individual recognition and cooperative nesting behavior have co-evolved in P. fuscatus because recognition helps stabilize social groups. This work provides evidence of a specific cognitive phenotype under selection because of social interactions, supporting the idea that social behavior can be a key driver of cognitive evolution., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Raymond Kelly's widely cited Warless Societies and the Origin of War (University of Michigan Press, 2000) seeks to explain the origins of two central signatures of human society: war and segmented-i.e., multilevel-societies. Both, he argues, arose with the emergence of a social-substitutability principle, a rule that establishes a collective identity among a set of individuals such that any one member becomes equivalent to, and responsible for the actions of, the others. This principle emerged during the Holocene, when population increase gave rise to the first lethal ambushes. By its nature, ambush obscures attackers' identities. Those attempting to retaliate for the ambush were therefore obliged to target members of the ambushers' group indiscriminately-i.e., based on a social-substitutability principle. Kelly's proposals draw welcome attention to a widespread, deeply influential, and unsettling human behavior, the disposition to hold everyone in a group culpable for the actions of a few, a proclivity that all too often results in mass slaughter. His general argument, however, is logically and empirically deficient, and cross-cultural evidence on ambush in contact-era New Guinea undermines his anonymity-of-ambush hypothesis. What then accounts for war and multilevel society? The New Guinea evidence strongly supports a contention that social-substitutability behavior arose not from offensive military action (i.e., ambush) but from the defensive military response to ambush. These findings render the social-substitutability argument's unconventional definition of war superfluous, undermine its chronology for the emergence of war, and underwrite an alternative scenario for the origins of multilevel, segmented society., (© 2023. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Ever since, people live in contact with nature and animals, even in relatively non-utilitarian ways. Erich Fromm and Edward Wilson termed this human universal "Biophilia". But why different species can live together in a social way, is explained by a "common social toolbox" of neural, psychological and physiologicalmechanisms, which evolved over phylogeny.Major components of this toolbox are found in the vertebrate brain, which evolved over the past 600 million years in a succession of key innovations and conservative preservation.The tegmental and diencephalic brain hosts a 450 million year old, structurally and functionally virtually unchanged "social network" which, in crosstalk with the mammalian prefrontal cortex or the analogous bird forebrain, enables complex social behaviour - within as well as between species. In addition, this toolbox features common principles of behavioural organization, including the expression and reading of emotions, as well as shared emotional, stress and calming systems. Such a common ground for social behaviour also explains the potential effectiveness of animal-assisted interventions in a wide range of pedagogic and therapeutic settings. However, positive effects aremostly revealed by experience and plausibility, whereas studies on animal- assisted activities and interventions according to biomedical scientific standards are still rare.

Dominance and leverage are both possible causes of social inequality. If sexual dimorphism influences patterns of intersexual dominance, we predicted that highly dimorphic species are constrained to exhibit male-biased power (MP), but species with low sexual dimorphism are free to demonstrate a broader range of intersexual power relationships. If market effects influence intersexual leverage, we predicted that females have more power when group composition is more male-biased and estrus is asynchronous. We analyzed data on intersexual power, sexual dimorphism, expected estrous overlap, and sex ratio for 79 extant primate species using phylogenetic logistic regression and ancestral state reconstructions. Although MP is more common, every major primate clade includes non-MP species. MP was associated with greater body mass and canine length dimorphism and with female-biased sex-ratios. Low estrous overlap was associated with non-MP. Although MP was reconstructed as likely ancestral for anthropoids, the last common ancestor of this clade probably did not exhibit high sexual dimorphism. The last common ancestor of catarrhines was probably highly dimorphic, potentially constraining intersexual power relationships. Non-MP probably evolved multiple times in primates and may be less common because multiple traits are linked to MP while fewer traits are associated with female-biased power or equality.

Social organization is commonly dynamic, with extreme examples in annual social insects, but little is known about the underlying signals and mechanisms. Bumble bee larvae with close contact to a queen do not differentiate into gynes, pupate at an earlier age, and are commonly smaller than siblings that do not contact a queen. We combined detailed observations, proteomics, microRNA transcriptomics, and gland removal surgery to study the regulation of brood development and division of labor in the annual social bumble bee Bombus terrestris . We found that regurgitates fed to larvae by queens and workers differ in their protein and microRNA composition. The proteome of the regurgitate overlaps significantly with that of the mandibular (MG) and hypopharyngeal glands (HPG), suggesting that these exocrine glands are sources of regurgitate proteins. The proteome of the MG and HPG, but not the salivary glands, differs between queens and workers, with caste-specificity preserved for the MG and regurgitate proteomes. Queens subjected to surgical removal of the MG showed normal behavior, brood care, and weight gain, but failed to shorten larval development. These findings suggest that substances in the queen MG are fed to larvae and influence their developmental program. We suggest that when workers emerge and contribute to larval feeding, they dilute the effects of the queen substances, until she can no longer manipulate the development of all larvae. Longer developmental duration may allow female larvae to differentiate into gynes rather than to workers, mediating the colony transition from the ergonomic to the reproductive phase.

Host-parasite dynamics involve coevolutionary arms races, which may lead to host specialization and ensuing diversification. Our general understanding of the evolution of host specialization in brood parasites is compromised by a restricted focus on bird and insect lineages. The cuckoo catfish (Synodontis multipunctatus) is an obligate parasite of parental care of mouthbrooding cichlids in Lake Tanganyika. Given the ecological and taxonomic diversity of mouthbrooding cichlids in the lake, we hypothesized the existence of sympatric host-specific lineages in the cuckoo catfish. In a sample of 779 broods from 20 cichlid species, we found four species parasitized by cuckoo catfish (with prevalence of parasitism of 2%-18%). All parasitized cichlids were from the tribe Tropheini, maternal mouthbrooders that spawn over a substrate (rather than in open water). Phylogenetic analysis based on genomic (ddRAD sequencing) and mitochondrial (Dloop) data from cuckoo catfish embryos showed an absence of host-specific lineages. This was corroborated by analyses of genetic structure and co-ancestry matrix. Within host species, parasitism was not associated with any individual characteristic we recorded (parent size, water depth), but was costly as parasitized parents carried smaller clutches of their own offspring. We conclude that the cuckoo catfish is an intermediate generalist and discuss costs, benefits and constraints of host specialization in this species and brood parasites in general., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Social behaviours are typically modelled using neighbour-modulated fitness, which focuses on individuals having their fitness altered by neighbours. However, these models are either interpreted using inclusive fitness, which focuses on individuals altering the fitness of neighbours, or not interpreted at all. This disconnect leads to interpretational mistakes and obscures the adaptive significance of behaviour. We bridge this gap by presenting a systematic methodology for constructing inclusive-fitness models. We find a behaviour's 'inclusive-fitness effect' by summing primary and secondary deviations in reproductive value. Primary deviations are the immediate result of a social interaction; for example, the cost and benefit of an altruistic act. Secondary deviations are compensatory effects that arise because the total reproductive value of the population is fixed; for example, the increased competition that follows an altruistic act. Compared to neighbour-modulated fitness methodologies, our approach is often simpler and reveals the model's inclusive-fitness narrative clearly. We implement our methodology first in a homogeneous population, with supplementary examples of help under synergy, help in a viscous population and Creel's paradox. We then implement our methodology in a class-structured population, where the advantages of our approach are most evident, with supplementary examples of altruism between age classes, and sex-ratio evolution.

A basic mechanism of kin selection is limited dispersal, whereby individuals remain close to their place of origin such that even indiscriminate social interaction tends to modify the fitness of genealogical kin. Accordingly, the causes and consequences of dispersal have received an enormous amount of attention in the social evolution literature. This work has focused on dispersal of individuals in space, yet similar logic should apply to dispersal of individuals in time (e.g. dormancy). We investigate how kin selection drives the evolution of dormancy and how dormancy modulates the evolution of altruism. We recover dormancy analogues of key results that have previously been given for dispersal, showing that: (1) kin selection favours dormancy as a means of relaxing competition between relatives; (2) when individuals may adjust their dormancy behaviour to local density, they are favoured to do so, resulting in greater dormancy in high-density neighbourhoods and a concomitant 'constant non-dormant principle'; (3) when dormancy is constrained to be independent of density, there is no relationship between the rate of dormancy and the evolutionary potential for altruism; and (4) when dormancy is able to evolve in a density-dependent manner, a greater potential for altruism is expected in populations with lower dormancy.

Cooperative social behaviors, such as parental care, have long been hypothesized to relax selection leading to the accumulation of genetic variation in populations. Although the idea has been discussed for decades, there has been relatively little experimental work to investigate how social behavior contributes to genetic variation in populations. Here, we investigate how parental care can shape molecular genetic variation in the subsocial insect, Nicrophorus vespilloides . Using whole-genome sequencing of populations that had evolved in the presence or absence of parental care for 30 generations, we show that parental care maintains levels of standing genetic variation. In contrast, under a harsh environment without care, strong directional selection caused a reduction in genetic variation. Furthermore, we show that adaptation to the loss of care is associated with genetic divergence between populations at loci related to stress, morphological development, and transcriptional regulation. These data reveal how social behavior is linked to the genetic processes that shape and maintain genetic diversity within populations, and provides rare empirical evidence for an old hypothesis., (© The Author(s) 2023. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE) and European Society for Evolutionary Biology (ESEN).)

Research in medicine and evolutionary biology suggests that the sequencing of parental investment has a crucial impact on offspring life history and health. Here, we take advantage of the synchronous birth system of wild banded mongooses to test experimentally the lifetime consequences to offspring of receiving extra investment prenatally versus postnatally. We provided extra food to half of the breeding females in each group during pregnancy, leaving the other half as matched controls. This manipulation resulted in two categories of experimental offspring in synchronously born litters: (i) 'prenatal boost' offspring whose mothers had been fed during pregnancy, and (ii) 'postnatal boost' offspring whose mothers were not fed during pregnancy but who received extra alloparental care in the postnatal period. Prenatal boost offspring lived substantially longer as adults, but postnatal boost offspring had higher lifetime reproductive success (LRS) and higher glucocorticoid levels across the lifespan. Both types of experimental offspring had higher LRS than offspring from unmanipulated litters. We found no difference between the two experimental categories of offspring in adult weight, age at first reproduction, oxidative stress or telomere lengths. These findings are rare experimental evidence that prenatal and postnatal investments have distinct effects in moulding individual life history and fitness in wild mammals. This article is part of the theme issue 'Evolutionary ecology of inequality'.

Many groups of animals have evolved social behaviours in different forms, from intimate familial associations to the complex eusocial colonies of some insects. The subfamily Xylocopinae, including carpenter bees and their relatives, is a diverse clade exhibiting a wide range of social behaviours, from solitary to obligate eusociality with distinct morphological castes, making them ideal focal taxa in studying the evolution of sociality. We used ultraconserved element data to generate a broadly sampled phylogeny of the Xylocopinae, including several newly sequenced species. We then conducted ancestral state reconstructions on the evolutionary history of sociality in this group under multiple coding models. Our results indicate solitary origins for the Xylocopinae with multiple transitions to sociality across the tree and subsequent reversals to solitary life, demonstrating the lability and dynamic nature of social evolution in carpenter bees. Ultimately, this work clarifies the evolutionary history of the Xylocopinae, and expands our understanding of independent origins and gains and losses of social complexity.

Changes in avian breeding phenology are among the most apparent responses to climate change in free-ranging populations. A key question is whether populations will be able to keep up with the expected rates of environmental change. There is a large body of research on the mechanisms by which avian lay-dates track temperature change and the consequences of (mal)adaptation on population persistence. Often overlooked is the role of males, which can influence the lay-date of their mate through their effect on the prelaying environment. We explore how social plasticity causing male indirect genetic effects can help or hinder population persistence when female genes underpinning lay-date and male genes influencing female's timing of reproduction both respond to climate-mediated selection. We extend quantitative genetic moving optimum models to predict the consequences of social plasticity on the maximum sustainable rate of temperature change, and evaluate our model using a combination of simulated data and empirical estimates from the literature. Our results suggest that predictions for population persistence may be biased if indirect genetic effects and cross-sex genetic correlations are not considered and that the extent of this bias depends on sex differences in how environmental change affects the optimal timing of reproduction. Our model highlights that more empirical work is needed to understand sex-specific effects of environmental change on phenology and the fitness consequences for population dynamics. While we discuss our results exclusively in the context of avian breeding phenology, the approach we take here can be generalized to many different contexts and types of social interaction., (© The Author(s) 2023. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE) and European Society for Evolutionary Biology (ESEN).)

Rudimentary social systems have the potential to both advance our understanding of how complex sociality may have evolved and our understanding of how changes in social environment may influence gene expression and cooperation. Recently, studies of primitively social Hymenoptera have greatly expanded empirical evidence for the role of social environment in shaping behavior and gene expression. Here, we compare brain gene expression profiles of foragers across social contexts in the small carpenter bee, Ceratina calcarata. We conducted experimental manipulations of field colonies to examine gene expression profiles among social contexts including foraging mothers, regular daughters, and worker-like dwarf eldest daughters in the presence and absence of mother. Our analysis found significant differences in gene expression associated with female age, reproductive status, and social environment, including circadian clock gene dyw, hexamerin, and genes involved in the regulation of juvenile hormone and chemical communication. We also found that candidate genes differentially expressed in our study were also associated with division of labor, including foraging, in other primitively and advanced eusocial insects. Our results offer evidence for the role of the regulation of key developmental hormones and circadian rhythms in producing cooperative behavior in rudimentary insect societies., (© The Author(s) 2023. Published by Oxford University Press on behalf of Society for Molecular Biology and Evolution.)

In social hymenopterans, workers specialize in different tasks. Whether a worker nurses the brood or forages is influenced by the responsiveness for task-related cues which in turn is determined by gene expression. Task choice is dynamic and changes throughout a worker's life, e.g. with age or in response to increased demands for certain tasks. Behavioural switches require the ability to adjust gene expression but the mechanisms regulating such transcriptional adaptations remain elusive. We investigated the role of histone acetylation in task specialization and behavioural flexibility in Temnothorax longispinosus ants. By inhibiting p300/CBP histone acetyltransferases (HAT) and manipulating colony composition, we found that HAT inhibition impairs the ability of older workers to switch to brood care. Yet, HAT inhibition increased the ability of young workers to accelerate their behavioural development and switch to foraging. Our data suggest that HAT in combination with social signals indicating task demands play an important role in modulating behaviour. Elevated HAT activity may contribute to keeping young brood carers from leaving the nest, where they would be exposed to high mortality. These findings shed light on the epigenetic processes underlying behavioural flexibility in animals and provide insight into the mechanisms of task specialization in social insects.

Reputations provide a powerful mechanism to sustain cooperation, as individuals cooperate with those of good social standing. But how should someone's reputation be updated as we observe their social behavior, and when will a population converge on a shared norm for judging behavior? Here, we develop a mathematical model of cooperation conditioned on reputations, for a population that is stratified into groups. Each group may subscribe to a different social norm for assessing reputations and so norms compete as individuals choose to move from one group to another. We show that a group initially comprising a minority of the population may nonetheless overtake the entire population-especially if it adopts the Stern Judging norm, which assigns a bad reputation to individuals who cooperate with those of bad standing. When individuals do not change group membership, stratifying reputation information into groups tends to destabilize cooperation, unless individuals are strongly insular and favor in-group social interactions. We discuss the implications of our results for the structure of information flow in a population and for the evolution of social norms of judgment.

Animals adjust their use of alarm calls depending on social environments. We tested whether dominant (adult) and subordinate (juvenile non-kin) male crested tits ( Lophophanes cristatus ) warn each other and heterospecific willow tits ( Poecile montanus ) across the wintering season. Birds rarely alarm called when feeding alone. Both adult and juvenile crested tits warned each other in early winter, and adults did so in the middle of wintering season. However, juvenile males rarely warned conspecific adult males in the middle of the winter. Both adult and juvenile males stopped giving alarm calls when feeding together at the end of wintering season. The results suggest that the mid-winter reduction of juvenile alarms could increase the likelihood of successful predator attacks on adults, increasing the chances for juveniles to replace adults and acquire their territories. By contrast, both adult and juvenile males produced alarm calls throughout the season when foraging together with willow tits. Whether juvenile male crested tits could be selectively altering alarm call propensity to endanger adult males, thereby selfishly enhancing their own succession to territory ownership, is discussed. The results add to the understanding of the origin of mixed-species groups and explain the dynamics of social communication. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.