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The social structure of populations plays a key role in shaping variation in sexual selection. In nature, sexual selection occurs in communities of interacting species; however, heterospecifics are rarely included in characterizations of social structure. Heterospecifics can influence the reproductive outcomes of intrasexual competition by interfering with intraspecific sexual interactions (interspecific reproductive interference [IRI]). We outline the need for studies of sexual selection to incorporate heterospecifics as part of the social environment. We use simulations to show that classic predictions for the effect of social structure on sexual selection are altered by an interaction between social structure and IRI. This interaction has wide‐ranging implications for patterns of sexual conflict and kin‐selected reproductive strategies in socially structured populations. Our work bridges the gap between sexual selection research on social structure and IRI, and highlights future directions to study sexual selection in interacting communities. [ABSTRACT FROM AUTHOR]

The distribution of group sizes in woodland baboons forms a pair of demographic oscillators that trade fertility off against predation risk. Fertility rates, however, set an upper limit on group size of around 90–95 animals. Despite this, two species of baboons (hamadryas and gelada) have groups that significantly exceed this limit, suggesting that these two species have been able to break through this fertility constraint. We suggest that they have done so by adopting a form of social substructuring that uses males as 'hired guns' to minimize the stresses of living in the unusually large groups required by high predation risk habitats. [ABSTRACT FROM AUTHOR]

Obligate brood-parasitic cheats have fascinated natural historians since ancient times. Passing on the costs of parental care to others occurs widely in birds, insects and fish, and often exerts selection pressure on hosts that in turn evolve defences. Brood parasites have therefore provided an illuminating system for researching coevolution. Nevertheless, much remains unknown about howecology and evolutionary history constrain or facilitate brood parasitism, or the mechanisms that shape or respond to selection. In this special issue, we bring together examples fromacross the animal kingdomto illustrate the diverseways in which recent research is addressing these gaps. This special issue also considers how research on brood parasitism may benefit from, and in turn inform, related fields such as social evolution and immunity. Here, we argue that progress in our understanding of coevolution would benefit from the increased integration of ideas across taxonomic boundaries and across Tinbergen's Four Questions: mechanism, ontogeny, function and phylogeny of brood parasitism. We also encourage renewed vigour in uncovering the natural history of the majority of the world's brood parasites that remain little-known. Indeed, it seems very likely that some of nature's brood parasites remain entirely unknown, because otherwise we are left with a puzzle: if parental care is so costly, why is brood parasitism not more common?. [ABSTRACT FROM AUTHOR]

Animal gut microbiota affect host physiology and behaviour. In social insects, where colony level integrity is preserved via a nestmate discrimination system based on cuticular hydrocarbon mixtures, microorganismal effects may therefore influence social dynamics. Although nestmate recognition has undergone a thorough exploration during the last four decades, few studies have investigated the putative role of gut microbes. Here, we integrated 16S rRNA-based microbial community profiling, chemical and behavioural approaches to test whether gut microbes affect nestmate recognition in Acromyrmex echinatior leaf-cutting ants. Treating workers with a sterile diet or with antibiotics resulted in a substantial alteration of their gut microbial communities. In pairwise social interactions, untreated versus antibiotic-treated nestmates behaved more aggressively than other nestmate and non-nestmate pairs, suggesting that bacterial suppression may alter chemical social cues and trigger aggressive behaviour. Chemical analyses of treated individuals revealed a decrease in the abundance of two metapleural gland antifungal compounds, and confirmed the correspondence between aggression levels and chemical profile differences. Feeding microbiota-remodelled ants with conspecific faecal droplets partially restored the original bacterial communities. Ants fed with faecal droplets from different colonies were unusually aggressive compared to pairs fed with faecal droplets from the same colony. We cannot exclude confounding effects resulting from the potentially harmful action of antibiotics on ant hosts. However, our results suggest a correlation between chemical profiles and the presence of certain microbial species in the gut, which may affect nestmate recognition and division of labour. This opens novel questions about the role of symbiotic microorganisms in the evolution of social insect behaviour. Highlights • We tested whether gut microbes influence nestmate recognition in leaf-cutting ants. • Gut microbiota suppression appears to affect aggression and cuticular hydrocarbons. • Partial microbiota restoration partly reverses induced changes in aggression. • 3-way interactions of gut microbes, cuticular hydrocarbons and aggression may exist. [ABSTRACT FROM AUTHOR]

This article situates actor-network theory in the history of evolutionary anthropology. In the 1980s, this attempt at explaining the social through the mediation of nonhumans received important impulses from Bruno Latour's conversations with primatologist Shirley Strum. In a re-articulation of social evolutionism, they proposed that the utilization of objects distinguished humans from baboons and that the use of a growing number of objects set industrialized human populations apart from hunter-gatherers, enabling the formation of larger collectives. While Strum's and Latour's early work presented baboons as almost human and suggested that we moderns had never been modern, the Anthropocene has reawakened curiosity about the original question of anthropology: how do modern humans, including modern scientists, differ from premoderns and animals? This 18th-century question is gaining new significance and urgency as we recognize our transmutation into a super-dominant species. But the answer might not lie solely in the use of more objects. [ABSTRACT FROM AUTHOR]

Group-living behavior is taxonomically widespread but rare in spiders. The conventional view is that the main pathways to group-living in spiders are either sub-social, where extended maternal care leads to prolonged sibling association; or communal living, where individuals aggregate to exploit a common resource. Female egg-sac guarding behavior occurs throughout kleptoparasitic spiders in the subfamily Argyrodinae (Theridiidae), while individuals in group-living species cohabit in the resource rich webs of their host spiders. These attributes fit both sub-social and communal routes to group-living, which offers new insights to study the early stages of social evolution. We investigated whether members of kleptoparasitic groups in natural populations comprise related individuals by comparing the population structure of two group-living species, Argyrodes miniaceus and A. cf. fissifrons, and two solitary species, A. fasciatus and Neospintharus trigonum. We found that: (1) genetic-spatial autocorrelation in group-living species was highest among spiders sharing the same host web and declined steeply with increasing distance, but no significant autocorrelation at any scale for solitary species; (2) there was high relatedness among group members in two cases of group-living species, which indicated relatedness was not an adhesive agent in most of the groups, but no high relatedness in solitary species; and (3) the host web boundary was not the sole predictor of genetic structures in group-living species. These results suggest that population genetic structure in the group-living species is caused by limited dispersal of group members that is favored by ecological conditions, including the nature and size of resources. In contrast, the absence of genetic structuring in populations of solitary species indicates a high level of dispersal with individual interactions unlikely to have fitness benefits. [ABSTRACT FROM AUTHOR]

In recent years, the phenomenon of cumulative cultural evolution (CCE) has become the focus of major research interest in biology, psychology and anthropology. Some researchers argue that CCE is unique to humans and underlies our extraordinary evolutionary success as a species. Others claim to have found CCE in non-human species. Yet others remain sceptical that CCE is even important for explaining human behavioural diversity and complexity. These debates are hampered by multiple and often ambiguous definitions of CCE. Here, we review how researchers define, use and test CCE. We identify a core set of criteria for CCE which are both necessary and sufficient, and may be found in non-human species. We also identify a set of extended criteria that are observed in human CCE but not, to date, in other species. Different socio-cognitive mechanisms may underlie these different criteria. We reinterpret previous theoretical models and observational and experimental studies of both human and non-human species in light of these more fine-grained criteria. Finally, we discuss key issues surrounding information, fitness and cognition. We recommend that researchers are more explicit about what components of CCE they are testing and claiming to demonstrate. [ABSTRACT FROM AUTHOR]

Understanding the behavioral and psychological mechanisms underlying social behaviors is one of the major goals of social evolutionary theory. In particular, a persistent question about animal cooperation is to what extent it is supported by other-regarding preferences--the motivation to increase the welfare of others. In many situations, animals adjust their behaviors through learning by responding to the rewards they experience as a consequence of their actions. Therefore, we may ask whether learning in social situations can be driven by evolved other-regarding rewards. Here we develop a mathematical model in order to ask whether the mere act of cooperating with a social partner will evolve to be inherently rewarding. Individuals interact repeatedly in pairs and adjust their behaviors through reinforcement learning. We assume that individuals associate with each game outcome an internal reward value. These perceived rewards are genetically evolving traits. We find that conditionally cooperative rewards that value mutual cooperation positively but the sucker's outcome negatively tend to be evolutionarily stable. Purely other-regarding rewards can evolve only under special parameter combinations. On the other hand, selfish rewards that always lead to pure defection are also evolutionarily successful. These findings are consistent with empirical observations showing that humans tend to display conditionally cooperative behavior and also exhibit a diversity of preferences. Our model also demonstrates the need to further integrate multiple levels of biological causation of behavior. [ABSTRACT FROM AUTHOR]

Facultatively social species exhibit varying degrees of reproductive skew that provide valuable insights into the possible evolutionary forces shaping the origins of obligate eusocial colony organisation, wherein the majority of individuals (workers) forgo direct reproduction. Here we report aspects of the nesting biology of a semi-arid population of the allodapine beeExoneurella setosa, which forms social colonies that lack reproductive hierarchies and are therefore ‘casteless’; an intriguing discovery given that a congeneric eusocial species exhibits the greatest morphological distinction between queen and worker castes in the entire subfamily Xylocopinae (Apidae).Exoneurella setosaexhibited a modal colony size of two females per nest and we analysed nest-mate differentiation in ovarian development, body size and wing wear (a proxy for foraging activity). We then contrasted empirical results with Monte Carlo-simulated colonies to ascertain that multifemale nests lack evidence for reproductive skew. Our results suggest that nest-site limitation is a key driver for eusocial organisation within the Xylocopinae and that the absence of such environmental limitations, combined with minor benefits for group nesting, can select for casteless social organisation. [ABSTRACT FROM AUTHOR]

Background: Social learning allows animals to eavesdrop on ecologically relevant knowledge of competitors in their environment. This is especially important when selecting a habitat if individuals have relatively little personal information on habitat quality. It is known that birds can use both conspecific and heterospecific information for social learning, but little is known about the relative importance of each information type. If provided with the choice between them, we expected that animals should copy the behaviour of conspecifics, as these confer the best information for that species. We tested this hypothesis in the field for Pied Flycatchers Ficedula hypoleuca arriving at their breeding grounds to select a nest box for breeding. We assigned arbitrary symbols to nest boxes of breeding pied flycatchers (conspecifics) and blue and great tits, Cyanistes caeruleus and Parus major (heterospecifics), in 2014 and 2016 in two areas with different densities of tits and flycatchers. After ca 50% of flycatchers had returned and a flycatcher symbol was assigned to their nest box, we gave the later arriving flycatchers the choice between empty nest boxes with either a conspecific (flycatcher) or a heterospecific (tit) symbol. Results: As expected, Pied Flycatchers copied the perceived nest box choice of conspecifics, but only in areas that were dominated by flycatchers. Against our initial expectation, flycatchers copied the perceived choice of heterospecifics in the area heavily dominated by tits, even though conspecific minority information was present. Conclusions: Our results confirm that the relative density of conspecifics and heterospecifics modulates the propensity to copy or reject novel behavioural traits. By contrasting conspecific and heterospecific ecology in the same study design we were able to draw more general conclusions about the role of fluctuating densities on social information use. [ABSTRACT FROM AUTHOR]

Among social insects, colony-level variation is likely to be widespread and has significant ecological consequences. Very few studies, however, have documented how genetic factors relate to behaviour at the colony level. Differences in expression of the foraging gene have been associated with differences in foraging and activity of a wide variety of organisms. We quantified expression of the red imported fire ant foraging gene ( sifor) in workers from 21 colonies collected across the natural range of Texas fire ant populations, but maintained under standardized, environmentally controlled conditions. Colonies varied significantly in their behaviour. The most active colonies had up to 10 times more active foragers than the least active colony and more than 16 times as many workers outside the nest. Expression differences among colonies correlated with this colony-level behavioural variation. Colonies with higher sifor expression in foragers had, on average, significantly higher foraging activity, exploratory activity and recruitment to nectar than colonies with lower expression. Expression of sifor was also strongly correlated with worker task (foraging vs. working in the interior of the nest). These results provide insight into the genetic and physiological processes underlying collective differences in social behaviour. Quantifying variation in expression of the foraging gene may provide an important tool for understanding and predicting the ecological consequences of colony-level behavioural variation. [ABSTRACT FROM AUTHOR]

The genetic mechanisms behind aggressive behaviors are important for understanding the formation of dominance hierarchies, and thus social systems in general. Studies into the effects of social experience and agonistic contest outcomes have shown significant changes in brain gene expression resulting from repeated winning and losing, as well as changing dominance rank, primarily in obligately social species. However, our knowledge of the genetic underpinnings of behavior in subsocial organisms is relatively poor, yet understanding the behavioral genetics of this simplest form of sociality provides the basis for understanding all other forms of social living. Here, we measured the effects of aggression on brain gene expression in the subsocial bee, Ceratina calcarata, in individuals that had experienced repeated winning, repeated losing, or a change in rank during repeated encounters. Consistent winning accounted for the majority of variation in brain gene expression, followed by changing rank over maintaining rank. Candidate genes for aggression are identified through comparative transcriptomics across 21 invertebrate and 6 vertebrate taxa. Lastly, we identified significantly over-represented cis-regulatory elements, namely transcription factor binding motifs deeply conserved across a wide range of taxa and broadly implicated in differential regulation of genes related to aggressive/dominant behavior. We present evidence for both genetic and cis-regulatory mechanisms for aggression that may have broad importance to social evolution. [ABSTRACT FROM AUTHOR]

Switching rate between cooperating and non-cooperating genotypes is a crucial social evolution factor, often neglected by game theory-inspired theoretical and experimental frameworks. We show that the evolution of alleles increasing the mutation or phenotypic switching rates toward cooperation is in itself a social dilemma. Although cooperative offspring are often unlikely to reproduce, due to high cost of cooperation, they can be seen both as a living public good and a part of the extended parental phenotype. The competition between individuals that generate cooperators and ones that do not is often more relevant than the competition between cooperators and non-cooperators. The dilemma of second-order cooperation we describe relates directly to eusociality, but can be also interpreted as a division of labor or a soma-germline distinction. The results of our simulations shine a new light on what Darwin had already termed a 'special difficulty' of evolutionary theory and describe a novel type of cooperation dynamics. [ABSTRACT FROM AUTHOR]

Identifying and quantifying variation in vocalizations is fundamental to advancing our understanding of processes such as speciation, sexual selection, and cultural evolution. The song of the humpback whale (Megaptera novaeangliae) presents an extreme example of complexity and cultural evolution. It is a long, hierarchically structured vocal display that undergoes constant evolutionary change. Obtaining robust metrics to quantify song variation at multiple scales (from a sound through to population variation across the seascape) is a substantial challenge. Here, the authors present a method to quantify song similarity at multiple levels within the hierarchy. To incorporate the complexity of these multiple levels, the calculation of similarity is weighted by measurements of sound units (lower levels within the display) to bridge the gap in information between upper and lower levels. Results demonstrate that the inclusion of weighting provides a more realistic and robust representation of song similarity at multiple levels within the display. This method permits robust quantification of cultural patterns and processes that will also contribute to the conservation management of endangered humpback whale populations, and is applicable to any hierarchically structured signal sequence. [ABSTRACT FROM AUTHOR]

Conflict between groups is a notable feature of many animal societies. Recent theoretical models suggest that violent intergroup conflict can shape patterns of within-group cooperation. However, despite its prevalence in social species, the adaptive significance of violent intergroup conflict has been little explored outside of humans and chimpanzees, Pan troglodytes . A barrier to current understanding of the role of intergroup conflict in the evolution of social behaviour is a lack of information on the causes and consequences of aggression between groups. Here, we examined the causes and fitness consequences of intergroup conflict in the banded mongoose, Mungos mungo , using a 16-year data set of observed intergroup interactions, life history and behaviour. Banded mongooses are cooperative breeders that live in highly territorial groups and engage in frequent, aggressive and violent intergroup interactions. We found that intensified population-wide competition for food and mates increased the probability of intergroup interactions, and that increased intergroup conflict was associated with periods in which groups were growing in size. Intergroup conflict had fitness costs in terms of reduced litter and adult survival but no cost to pregnant females: in fact, females were less likely to abort following an intergroup interaction than when there had been no recent intergroup conflict. Our results suggest that intergroup conflict has measurable costs to both individuals and groups in the long and short term, and that levels of conflict among groups could be high enough to affect patterns of within-group cooperative behaviour. Establishing the consequences of intergroup conflict in cooperative species can shed light on patterns of conflict and cooperation within groups and, in turn, facilitate our understanding of social evolution. [ABSTRACT FROM AUTHOR]

Visual signals, including changes in coloration and color patterns, are frequently used by animals to convey information. During contests, body coloration and its changes can be used to assess an opponent's state or motivation. Communication of aggressive propensity is particularly important in group-living animals with a stable dominance hierarchy, as the outcome of aggressive interactions determines the social rank of group members. Neolamprologus pulcher is a cooperatively breeding cichlid showing frequent within-group aggression. Both sexes exhibit two vertical black stripes on the operculum that vary naturally in shape and darkness. During frontal threat displays these patterns are actively exposed to the opponent, suggesting a signaling function. To investigate the role of operculum stripes during contests we manipulated their darkness in computer animated pictures of the fish. We recorded the responses in behavior and stripe darkness of test subjects to which these animated pictures were presented. Individuals with initially darker stripes were more aggressive against the animations and showed more operculum threat displays. Operculum stripes of test subjects became darker after exposure to an animation exhibiting a pale operculum than after exposure to a dark operculum animation, highlighting the role of the darkness of this color pattern in opponent assessment. We conclude that (i) the black stripes on the operculum of N. pulcher are a reliable signal of aggression and dominance, (ii) these markings play an important role in opponent assessment, and (iii) 2D computer animations are well suited to elicit biologically meaningful short-term aggressive responses in this widely used model system of social evolution. [ABSTRACT FROM AUTHOR]

Reproductive skew in multimale groups may be determined by the need for alpha males to offer reproductive opportunities as staying incentives to subordinate males (concessions), by the relative fighting ability of the alpha male (tug-of-war) or by how easily females can be monopolized (priority-of-access). These models have rarely been investigated in species with exceptionally long male tenures, such as white-faced capuchins, where female mate choice for novel unrelated males may be important in shaping reproductive skew. We investigated reproductive skew in white-faced capuchins at Sector Santa Rosa, Costa Rica, using 20 years of demographic, behavioural and genetic data. Infant survival and alpha male reproductive success were highest in small multimale groups, which suggests that the presence of subordinate males can be beneficial to the alpha male, in line with the concession model's assumptions. None of the skew models predicted the observed degree of reproductive sharing, and the probability of an alpha male producing offspring was not affected by his relatedness to subordinate males, whether he resided with older subordinate males, whether he was prime aged, the number of males or females in the group or the number of infants conceived within the same month. Instead, the alpha male's probability of producing offspring decreased when he was the sire of the mother, was weak and lacked a well-established position and had a longer tenure. Because our data best supported the inbreeding avoidance hypothesis and female choice for strong novel mates, these hypotheses should be taken into account in future skew models. [ABSTRACT FROM AUTHOR]

How do animal social skills influence evolution? Complex animal social behaviors require many cognitive skills including individual recognition and observational learning. For social systems to evolve, these abilities need to be transmitted genetically or culturally and supported by the evolution of underlying neural systems. Because animal skill sets are so varied, it seems best to describe animal cognitive behaviors as being a social calculus that can change with experience, which has evolved to match and facilitate the complexity of the social system where it arose. That is, acquiring and using social information in response to a rapidly changing complex world leads to social competence enabling success in essential behavioral interactions. Here, we describe the remarkable suite of social skills discovered in the African cichlid fish Astatotilapia burtoni, including an attention hierarchy, male deception, transitive inference, the mechanistic bases of social dominance, female mate choice and the neural control of female reproductive behavior. The social calculus of this species is presented as an example of a potential causal factor in the evolution of sophisticated social behavior necessary for the evolutionary success of their social system. [ABSTRACT FROM AUTHOR]

Animal personality studies rely on collecting repeated behavioural data either in the field or under laboratory conditions. Conditions in the field should be far less stable than controlled laboratory conditions, and hence represent a potential source of variation in behaviour. Here we report on the first experiment to our knowledge that formally compares the repeatability of identical behaviours in the laboratory and the field, and across the transition from laboratory to field. Using a design that controls for observation number we compared two groups of sea anemones, observed across two experimental phases, either (1) in the field followed by the laboratory or (2) in the laboratory only. We analysed differences in behaviour across a range of levels including repeatability and its between- and within-individual variance components. Although mean startle response durations varied between the laboratory and field, there was no significant difference in repeatability across situations. Within-individual variance differed between the two periods of the experiment for animals observed only in the laboratory but this effect was not present for those that transitioned from field to laboratory. Furthermore, the rank order of individual responses was stable for animals observed only in the laboratory but changed for those that transitioned from field to laboratory. These results show that although repeatability estimates in the laboratory can yield results like those obtained in the field, the underlying components of consistent variation in behaviour might be influenced by an interaction between prior experiences and the current situation in which the animals are observed. [ABSTRACT FROM AUTHOR]

Indirect genetic effects (IGEs) are a major driver of social evolution, but much of the experimental work pertaining to IGEs on social behaviour has focused on the effect of stimulus individuals on single focal individuals. We extended IGE research to examine how stimulus individuals influence social interactions among several focal individuals. Specifically, we relied on recent work on social behaviour in fruit flies to examine whether IGEs cause 12 stimulus flies of distinct genotypes to alter social interactions within groups of six focal flies. The social behaviour of focals was significantly affected by the genotype of the stimulus flies. Focals were closer together when grouped with stimulus flies from genotypes that were close together than when grouped with stimulus flies from genotypes that were farther apart. A mechanism mediating this effect was the encounter rate between focal flies, which was lowest when the focal flies were grouped with stimulus flies of the more cohesive genotypes. [ABSTRACT FROM AUTHOR]

Odontocete sounds are produced by two pairs of phonic lips situated in soft nares below the blowhole; the right pair is larger and is more likely to produce clicks, while the left pair is more likely to produce whistles. This has important implications for the cultural evolution of delphinid sounds: the greater the physical constraints, the greater the probability of random convergence. In this paper the authors examine the call structure of eight killer whale populations to identify structural constraints and to determine if they are consistent among all populations. Constraints were especially pronounced in two-voiced calls. In the calls of all eight populations, the lower component of two-voiced (biphonic) calls was typically centered below 4 kHz, while the upper component was typically above that value. The lower component of two-voiced calls had a narrower frequency range than single-voiced calls in all populations. This may be because some single-voiced calls are homologous to the lower component, while others are homologous to the higher component of two-voiced calls. Physical constraints on the call structure reduce the possible variation and increase the probability of random convergence, producing similar calls in different populations. [ABSTRACT FROM AUTHOR]

An important issue in the evolution of group living is the risk of pathogen and predator exposure entailed by the inherent accumulation of feces within a nesting site. While many group living species limit this risk by cleaning the nest, others do not, raising questions about the benefits of maintaining feces in the nest and their importance in social evolution. Here, we investigated whether one of these benefits could be mediated by coprophagy in families of the European earwig, Forficula auricularia. In this insect species, mothers and mobile juveniles (nymphs) line their nests with feces and consume them. In a first experiment, we tested whether access to feces produced by either nymphs or mothers affects nymph survival in both presence and absence of food. The results showed that access to sibling feces, but not mother feces, enhanced offspring survival under food deprivation. Such an effect did not occur when regular food was available. We then conducted a food-choice experiment to reveal whether nymphs prefer food to feces, and if they discriminate between feces from their mother, unrelated adult females, unrelated nymphs, or their siblings. We found that offspring generally preferred regular food to feces, but nevertheless always consumed some feces. By contrast, nymphs showed no preference between related sibling or mother feces and did not discriminate between feces from related and unrelated individuals. Overall, our results suggest that the benefits of coprophagy could favor the maintenance of feces within the nest and promote the evolution of social life. [ABSTRACT FROM AUTHOR]

The evolutionary stability of quantitative traits depends on whether a population can resist invasion by any mutant. While uninvadability is well understood in well-mixed populations, it is much less so in subdivided populations when multiple traits evolve jointly. Here, we investigate whether a spatially subdivided population at a monomorphic equilibrium for multiple traits can withstand invasion by any mutant or is subject to diversifying selection. Our model also explores the correlations among traits arising from diversifying selection and how they depend on relatedness due to limited dispersal. We find that selection tends to favor a positive (negative) correlation between two traits when the selective effects of one trait on relatedness is positively (negatively) correlated to the indirect fitness effects of the other trait. We study the evolution of traits for which this matters: dispersal that decreases relatedness and helping that has positive indirect fitness effects. We find that when dispersal cost is low and the benefits of helping accelerate faster than its costs, selection leads to the coexistence of mobile defectors and sessile helpers. Otherwise, the population evolves to a monomorphic state with intermediate helping and dispersal. Overall, our results highlight the effects of population subdivision for evolutionary stability and correlations among traits. [ABSTRACT FROM AUTHOR]

Cultural transmission in nonhuman animals is often sex biased, with females more frequently or efficiently learning cultural behaviors than males. The evolutionary origins of sex-biased cultural transmission have been a mystery, though it has been proposed that female offspring may gain greater reproductive benefit from cultural traits than sons—the "disparate benefits" hypothesis. I propose a different, "uniparental teaching," hypothesis where sex-biased transmission evolves in uniparental species if mothers teach, that is, invest in their offsprings' learning. I show, with theoretical models, that mothers evolve to invest more in teaching daughters than sons because teaching daughters results in greater inclusive fitness benefits. Teaching a son gives him a reproductive benefit for one generation. However, I show that because daughters may teach future generations, teaching a daughter can be a better long-term investment. I also model the disparate benefits hypothesis and show that the uniparental teaching hypothesis better fits the empirical patterns of sex-biased transmission in the well-studied example of "sponging" in bottlenose dolphins. Uniparental teaching may also explain sex-biased transmission in other species, including chimpanzees. My findings suggest that controversial mechanisms of cultural transmission in nonhumans, such as teaching, may be inferred from population-level patterns of transmission even when it is difficult to observe transmission directly in the field or laboratory. [ABSTRACT FROM AUTHOR]

The so-called "strength of weak ties" is a central concept in social network theory, especially for understanding how information and diseases are transmitted through socially structured populations. In general, weak ties occur in networks where relatively few individuals are responsible for maintaining linkages between groups of individuals that would otherwise be poorly connected. This common structural motif can be seen in the social networks of species with fission–fusion social organization, such as giraffe (Giraffa camelopardalis). Giraffe social networks are characterized by social cliques in which individuals associate more with members of their own social clique than with those outside their clique. Individuals involved in weak, between-clique social interactions are hypothesized to serve as bridges by which an infection may enter a clique and, hence, may experience higher infection risk. Here, we address this and other hypotheses explaining helminth infection patterns in wild giraffe, exploring the relative roles of the social network and ranging behavior in determining infection risk. We show that infection risk is more influenced by weak ties with individuals outside one's clique than by repeated contact with a core set of associates. Even when controlling for age and home range size, individuals who engaged in more between-clique associations, that is, those with multiple weak ties, were more likely to be infected with gastrointestinal helminth parasites. Our results suggest that diverse social interactions with giraffe from multiple cliques may increase exposure to pathogens. The importance of weak ties in pathogen transmission has only rarely been empirically demonstrated in wildlife. [ABSTRACT FROM AUTHOR]

One key question in social evolution is the identification of factors that promote the formation and maintenance of stable bonds between females and males beyond the mating context. Baboons lend themselves to examine this question, as they vary in social organisation and male-female association patterns. We report the results from the first systematic observations of individually identified wild female Guinea baboons. Guinea baboons live in a multilevel society with female-biased dispersal. Although several males could be found within 5 m of females, each female chiefly associated with one 'primary' male at the 2 m distance. Social interactions occurred predominantly with the primary male, and female reproductive state had little influence on interaction patterns. The number of females per primary male varied from 1 to 4. During the 17-month study period, half of the females transferred between different males one or multiple times. A subset of females maintained weaker affiliative nonsexual relationships with other 'secondary' males. Units composed of primary males with females, and occasional secondary males, apparently form the core of the Guinea baboon society. The social organisation and mating patterns of Guinea and hamadryas baboons may have a common evolutionary origin, despite notable differences in relationship quality. Specifically, Guinea baboon females appear to have greater leverage in their association patterns than hamadryas baboon females. Although we cannot yet explain the lack of overt male control over females, results generally support the notion that phylogenetic descent may play an important role in shaping social systems. [ABSTRACT FROM AUTHOR]

Social immunity refers to any immune defence that benefits others, besides the individual that mounts the response. Since contributions to social immunity are known to be personally costly, they are contributions to a public good. However, individuals vary in their contributions to this public good and it is unclear why. Here we investigate whether they are responding to contributions made by others with experiments on burying beetle ( Nicrophorus vespilloides) families. In this species, females, males and larvae each contribute to social immunity through the application of antimicrobial exudates upon the carrion breeding resource. We show experimentally that mothers reduce their contributions to social immunity when raising large broods, and test two contrasting hypotheses to explain why. Either mothers are treating social immunity as a public good, investing less in social immunity when their offspring collectively contribute more, or mothers are trading off investment in social immunity with investment in parental care. Overall, our experiments yield no evidence to support the existence of a trade-off between social immunity and other parental care traits: we found no evidence of a trade-off in terms of time allocated to each activity, nor did the relationship between social immunity and brood size change with female condition. Instead, and consistent with predictions from models of public goods games, we found that higher quality mothers contributed more to social immunity. Therefore our results suggest that mothers are playing a public goods game with their offspring to determine their personal contribution to the defence of the carrion breeding resource. [ABSTRACT FROM AUTHOR]

Hamilton's theory of kin selection has revolutionized and inspired fifty years of additional theories and experiments on social evolution. Whereas Hamilton's broader intent was to explain the evolutionary stability of cooperation, his focus on shared genetic history appears to have limited the application of his theory to populations within a single species rather than across interacting species. The evolutionary mechanisms for cooperation between species require both spatial and temporal correlations among interacting partners for the benefits to be not only predictable but of sufficient duration to be reliably delivered. As a consequence when the benefits returned by mutualistic partners are redirected to individuals other than the original donor, cooperation usually becomes unstable and parasitism may evolve. However, theoretically, such redirection of mutualistic benefits may actually reinforce, rather than undermine, mutualisms between species when the recipients of these redirected benefits are genetically related to the original donor. Here, I review the few mathematical models that have used Hamilton's theory of kin selection to predict the evolution of mutualisms between species. I go on to examine the applicability of these models to the most well-studied case of mutualism, pollinating seed predators, where the role of kin selection may have been previously overlooked. Future detailed studies of the direct, and indirect, benefits of mutualism are likely to reveal additional possibilities for applying Hamilton's theory of kin selection to mutualisms between species. [ABSTRACT FROM AUTHOR]

The importance of between-group competition in the social evolution of animal societies is controversial, particularly with respect to understanding the origins and maintenance of cooperation in our own species. Among primates, aggressive between-group encounters are often rare or strikingly absent, a phenomenon that in some species has been ascribed to the presence of collective action problems. Here, we report on a series of comparative tests that show that the intensity of between-group contest competition is indeed lower in species that experience a collective action problem while controlling for predictions from an "ideal gas" model of animal encounters and general species' ecology. Species that do not succumb to the collective action problem are either cooperative breeders, are characterized by philopatry of the dominant sex, or live in relatively small groups with few individuals of this dominant sex. This implies that collective action problems are averted either through shared genes and benefits or a by-product mutualism in which the territorial behavior of some privileged individuals is not affected by the behavior of others. We conclude that across the primate taxon, the intensity of between-group competition is predominantly constrained by a social dilemma among group members, rather than ecological conditions, and that the collective action problem is thus an important selective pressure in the evolution of primate (including human) cooperation and sociality. [ABSTRACT FROM AUTHOR]

Studies of annual and geographic variation in eusocial bee populations suggest that more stringent environmental conditions result in stronger reproductive skew favouring queens, while moderate conditions favour increasing worker reproduction. To test these predictions, we compared the phenology and colony development of H. ligatus nesting in St. Catharines, Ontario, Canada to a previously studied aggregation 90 km north of St. Catharines, in Victoria, Ontario. Despite the close proximity of these two locations, St. Catharines has markedly shorter winters and longer summers. Comparisons between St. Catharines in 2006 and Victoria in the 1980s and 1990s incorporate both geographic differences in climate and temporal differences due to climate change. We predicted that St. Catharines foundress queens should emerge from hibernation and initiate nests earlier in spring, giving them time to produce more workers. Since earlier studies indicated that queens have difficulty suppressing worker reproduction in larger colonies, we also predicted higher rates of worker ovarian development in St. Catharines. In spring and summer 2006, we excavated 65 H. ligatus nests, comparing their contents to 713 specimens collected in pan traps. As predicted, nests were initiated about a month earlier in St. Catharines than in Victoria, but contrary to prediction, fewer workers were produced in St. Catharines. St. Catharines workers were just as likely to have developed ovaries as Victoria workers. About 40% of St. Catharines workers were classified as reproductive, and larger reproductive workers tended to have higher ovarian scores. Early queen mortality in the longer nest cycle of St. Catharines bees may have enhanced opportunities for worker reproduction despite their smaller numbers. Novel features of H. ligatus sociobiology in St. Catharines included evidence that queens can initiate new nests following the loss of their first brood, overlap between worker and gyne production within some nests, and high rates of independent nest founding by worker-sized females, suggesting that many worker-brood females overwinter. Overall, the distinctly warmer climate of St. Catharines compared to Victoria led to earlier nest initiation and lengthening of the flight season, but not to the predicted differences in colony social organisation or queen-worker reproductive skew. A second objective of our study was to assess how well pan trap collections capture important information about demographic and social parameters important in assessing social variability in sweat bees. Nest excavations and pan traps produced similar results, suggesting that pan traps are a good alternative when nest excavations are impossible. [ABSTRACT FROM AUTHOR]

Gregarious animals need to make collective decisions in order to keep their cohesiveness. Several species of them live in multilevel societies, and form herds composed of smaller communities. We present a model for the development of a leadership hierarchy in a herd consisting of loosely connected sub-groups (e.g. harems) by combining self organization and social dynamics. It starts from unfamiliar individuals without relationships and reproduces the emergence of a hierarchical and modular leadership network that promotes an effective spreading of the decisions from more capable individuals to the others, and thus gives rise to a beneficial collective decision. Our results stemming from the model are in a good agreement with our observations of a Przewalski horse herd (Hortobágy, Hungary). We find that the harem-leader to harem-member ratio observed in Przewalski horses corresponds to an optimal network in this approach regarding common success, and that the observed and modeled harem size distributions are close to a lognormal. [ABSTRACT FROM AUTHOR]