Early Social Networks Predict Survival in Wild Bottlenose Dolphins

A fundamental question concerning group-living species is what factors influence the evolution of sociality. Although several studies link adult social bonds to fitness, social patterns and relationships are often formed early in life and are also likely to have fitness consequences, particularly in species with lengthy developmental periods, extensive social learning, and early social bond-formation. In a longitudinal study of bottlenose dolphins (Tursiops sp.), calf social network structure, specifically the metric eigenvector centrality, predicted juvenile survival in males. Additionally, male calves that died post-weaning had stronger ties to juvenile males than surviving male calves, suggesting that juvenile males impose fitness costs on their younger counterparts. Our study indicates that selection is acting on social traits early in life and highlights the need to examine the costs and benefits of social bonds during formative life history stages.     

Kin selection does not explain male aggregation at leks of 4 manakin species

In lek-mating systems, males aggregate at display arenas andfemales visit solely for the purpose of mating. This breedingsystem is characterized by high variance in male mating successwith one male often receiving most copulations. High reproductiveskew among males has led to question why males join leks whentheir chances of reproductive success are so low. Kin selectionhas been invoked as a mechanism to explain the evolution oflekking behavior, whereby nonreproducing but genetically relatedmales gain indirect inclusive-fitness benefits. Evidence forkin selection among lek-mating birds is, however, mixed. Here,we show that kin selection is unlikely to be an important explanationfor evolution of lekking behavior in manakins (Aves: Pipridae).We found that for 4 species chosen from several major cladeswithin Pipridae, males within leks were not significantly morerelated than expected from random assortment of males in thepopulation. This means that nonreproducing males do not gainindirect inclusive-fitness benefits by joining leks. This resultsuggests alternative mechanisms must be invoked to explain theevolution of lek-mating systems in manakins.     

Being in the thick of things: context-dependent network centrality in a captive flock of American flamingos

We test the hypothesis that the relationship between networks resulting from unresolved agonistic interactions (URI) and social dominance was context-dependent in a captive group of American Flamingos (Phoenicopterus ruber ruber). URI formed networks that differed substantially from the dominance network and depended on the context in which the interactions occurred. Betweenness centrality in the URI network on the region (“the island”) where nesting took place was strongly correlated with dominance score. This correlation was particularly strong in the case of males, but, in both sexes, the dominant individual in a dyad was likely to have a higher betweenness centrality in the island URI. On the other hand, betweenness centrality in URI networks at the feeder did not show any relationship with dominance, but was associated with observed visits to the feeder. Thus, centrality in both the island URI network and the feeder URI network was associated with access to a resource over which individuals competed. Along with other results suggesting that non-dominance interactions may form social networks distinct from, but related to, dominance networks, our results support the hypothesis that relationships within animal social groups can be modeled as a series of distinct but inter-related networks.     

Formidable females redux:male social integration into female networks and the value of dynamic multilayer networks

The development of multilayer network techniques is a boon for researchers who wish to understand how different interaction layers might influence each other,and how these in turn might influence group dynamics.Here,we investigate how integration between male and female grooming and aggression interaction networks influences male power trajectories in vervet monkeys Chlorocebus pygerythrus.Our previous analyses of this phenomenon used a monolayer approach,and our aim here is to extend these analyses using a dynamic multilayer approach.To do so,we constructed a temporal series of male and female interaction layers.We then used a multivariate multilevel autoregression model to compare cross-lagged associations between a male's centrality in the female grooming layer and changes in male Elo ratings.Our results confirmed our original findings:changes in male centrality within the female grooming network were weakly but positively tied to changes in their Elo ratings.However,the multilayer network approach offered additional insights into this social process,identifying how changes in a male's centrality cascade through the other network layers.This dynamic view indicates that the changes in Elo ratings are likely to be short-lived,but that male centrality within the female network had a much stronger impact throughout the multilayer network as a whole,especially on reducing intermale aggression(i.e.,aggression directed by males toward other males).We suggest that multilayer social network approaches can take advantage of increased amounts of social data that are more commonly collected these days,using a variety of methods.Such data are inherently multilevel and multilayered,and thus offer the ability to quantify more precisely the dynamics of animal social behaviors.     

Bipartite graphs as models of population structures in evolutionary multiplayer games

By combining evolutionary game theory and graph theory, "games on graphs" study the evolutionary dynamics of frequency-dependent selection in population structures modeled as geographical or social networks. Networks are usually represented by means of unipartite graphs, and social interactions by two-person games such as the famous prisoner's dilemma. Unipartite graphs have also been used for modeling interactions going beyond pairwise interactions. In this paper, we argue that bipartite graphs are a better alternative to unipartite graphs for describing population structures in evolutionary multiplayer games. To illustrate this point, we make use of bipartite graphs to investigate, by means of computer simulations, the evolution of cooperation under the conventional and the distributed N-person prisoner's dilemma. We show that several implicit assumptions arising from the standard approach based on unipartite graphs (such as the definition of replacement neighborhoods, the intertwining of individual and group diversity, and the large overlap of interaction neighborhoods) can have a large impact on the resulting evolutionary dynamics. Our work provides a clear example of the importance of construction procedures in games on graphs, of the suitability of bigraphs and hypergraphs for computational modeling, and of the importance of concepts from social network analysis such as centrality, centralization and bipartite clustering for the understanding of dynamical processes occurring on networked population structures.     

Extreme cost of rivalry in a monandrous species: male–male interactions result in failure to acquire mates and reduced longevity

Mating system variation is profound in animals. In insects, female willingness to remate varies from mating with hundreds of males (extreme polyandry) to never remating (monandry). This variation in female behaviour is predicted to affect the pattern of selection on males, with intense pre-copulatory sexual selection under monandry compared to a mix of pre- and post-copulatory forces affecting fitness under polyandry. We tested the hypothesis that differences in female mating biology would be reflected in different costs of pre-copulatory competition between males. We observed that exposure to rival males early in life was highly costly for males of a monandrous species, but had lower costs in the polyandrous species. Males from the monandrous species housed with competitors showed reduced ability to obtain a mate and decreased longevity. These effects were specific to exposure to rivals compared with other types of social interactions (heterospecific male and mated female) and were either absent or weaker in males of the polyandrous species. We conclude that males in monandrous species suffer severe physiological costs from interactions with rivals and note the significance of male–male interactions as a source of stress in laboratory culture.     

Female reproductive strategies, paternity and community structure in wild West African chimpanzees

Although the variability and complexity of chimpanzee behaviour frustrates generalization, it is widely believed that social evolution in this species occurs in the context of the recognizable social group or community. We used a combination of field observations and noninvasive genotyping to study the genetic structure of a habituated community of 55 wild chimpanzees, Pan troglodytes verus, in the Ta? Forest, C?te d'Ivoire. Pedigree relationships in that community show that female mate choice strategies are more variable than previously supposed and that the observed social groups are not the exclusive reproductive units. Genetic evidence based on nuclear microsatellite markers and behavioural obser-vations reveal that females in the Ta? forest actively seek mating partners outside their social unit; noncommunity males accounted for half the paternities over 5 years. This female mating strategy increases male gene flow between communities despite male philopatry, and negates the predicted higher relatedness among community males. Kin selection seems unlikely to explain the frequent cooperation and sharing observed among group males in this population. Similarly, inbreeding avoidance is probably not the sole cause of permanent adolescent female dispersal as a combination of extragroup mating and avoidance of incest with home group males would allow females to avoid inbreeding without the hazards associated with immigration into a new community. Extragroup mating as part of chimpanzee females' reproductive strategy may allow them to choose from a wider variety and number of males, without losing the resources and support provided by their male social group partners. Copyright 1999 The Association for the Study of Animal Behaviour.     

It takes two to tango: reproductive skew and social correlates of male mating success in a lek-breeding bird

Variance in reproductive success among individuals is a defining characteristic of many social vertebrates. Yet, our understanding of which male attributes contribute to reproductive success is still fragmentary in most cases. Male–male reproductive coalitions, where males jointly display to attract females, are of particular interest to evolutionary biologists because one male appears to forego reproduction to assist the social partner. By examining the relationship between social behaviour and reproductive success, we can elucidate the proximate function of coalitions in the context of mate choice. Here, we use data from a 4-year study of wire-tailed manakins (Pipra filicauda) to provide molecular estimates of reproductive skew and to test the hypothesis that male–male social interactions, in the context of coordinated displays, positively influence a male''s reproductive success. More specifically, we quantify male–male social interactions using network metrics and predict that greater connectivity will result in higher relative reproductive success. Our data show that four out of six leks studied had significant reproductive skew, with success apportioned to very few individuals in each lek. Metrics of male social affiliations derived from our network analysis, especially male connectivity, measured as the number of males with whom the focal male has extended interactions, were strong predictors of the number of offspring sired. Thus, network connectivity is associated with male fitness in wire-tailed manakins. This pattern may be the result of shared cues used by both sexes to assess male quality, or the result of strict female choice for coordinated display behaviour.     

Fashion,Cooperation, and Social Interactions

Fashion plays such a crucial rule in the evolution of culture and society that it is regarded as a second nature to the human being. Also, its impact on economy is quite nontrivial. On what is fashionable, interestingly, there are two viewpoints that are both extremely widespread but almost opposite: conformists think that what is popular is fashionable, while rebels believe that being different is the essence. Fashion color is fashionable in the first sense, and Lady Gaga in the second. We investigate a model where the population consists of the afore-mentioned two groups of people that are located on social networks (a spatial cellular automata network and small-world networks). This model captures two fundamental kinds of social interactions (coordination and anti-coordination) simultaneously, and also has its own interest to game theory: it is a hybrid model of pure competition and pure cooperation. This is true because when a conformist meets a rebel, they play the zero sum matching pennies game, which is pure competition. When two conformists (rebels) meet, they play the (anti-) coordination game, which is pure cooperation. Simulation shows that simple social interactions greatly promote cooperation: in most cases people can reach an extraordinarily high level of cooperation, through a selfish, myopic, naive, and local interacting dynamic (the best response dynamic). We find that degree of synchronization also plays a critical role, but mostly on the negative side. Four indices, namely cooperation degree, average satisfaction degree, equilibrium ratio and complete ratio, are defined and applied to measure people’s cooperation levels from various angles. Phase transition, as well as emergence of many interesting geographic patterns in the cellular automata network, is also observed.     

Higher relatedness mitigates mortality in a nematode with lethal male fighting

According to kin selection theory, individuals show less aggression towards their relatives. Limited dispersal promotes interactions among relatives but also increases competition among them. The evolution of cooperation in viscous populations has been subject of mainly theoretical exploration. We investigated the influence of relatedness on aggression in males of entomopathogenic nematode Steinernema longicaudum that engage in lethal fighting. In a series of in vitro experiments, we found that both competitor male group size and relatedness influence male mortality rates. Higher relatedness led to progressively lower rates of male mortality. In experimentally infected insects, wherein large numbers of males and females interact, the proportion of dead and paralysed (= terminally injured) males was higher when infection was established by infective juveniles originating from a mixture of three lines than in those infected by a single line. The results collectively show that Steinernema longicaudum males recognize their kin and consequently male mortality rates are lower in groups consisting of more related males. Furthermore, this monotonic negative relationship between aggression and relatedness suggests that kin selection benefits are still substantial even under extreme competition. Our experiments also suggest that kin recognition in entomopathogenic nematodes has a genetic basis rather than being strictly based on environmental cues. We discuss our findings within the theoretical context of the evolution of altruistic/cooperative behaviour in structured populations.     

Female clustering in cockroach aggregations—A case of social niche construction?

Individuals in groups can suffer costs through interactions with adversarial or unknown conspecifics. Social niche construction allows individuals to buffer such potential costs by only engaging in preferred associations. This may be particularly beneficial in insect aggregations, which are often large and highly fluid. However, little is known regarding the structuring of such aggregations. Here we use social network analyses to test for fine‐scale social structure in resting aggregations of the sub‐social cockroach Diploptera punctata and to explore the social pressures that contribute towards such structure. We showed that females were significantly more gregarious than males and formed the core of the proximity network, thus demonstrating a higher level of social integration. This fine‐scale structure is likely to result from females displacing males; females initiated most displacements whilst males received the majority. We explain this behaviour in terms of social niche construction by showing that females received significantly fewer approaches and investigations at more female‐biased local sex ratios. We therefore suggest that female social clustering occurs in this, and presumably other, species to reduce potential costs associated with male harassment. This demonstrates how social niche construction can lead to higher level social structure; we suggest this approach could be used across a range of species in order to improve our understanding of the evolution of sociality.     

Dynamic social behaviour in a bacterium: Pseudomonas aeruginosa partially compensates for siderophore loss to cheats

Cooperation underlies diverse phenomena including the origins of multicellular life, human behaviour in economic markets and the mechanisms by which pathogenic bacteria cause disease. Experiments with microorganisms have advanced our understanding of how, when and why cooperation evolves, but the extent to which microbial cooperation can recapitulate aspects of animal behaviour is debated. For instance, understanding the evolution of behavioural response rules (how should one individual respond to another's decision to cooperate or defect?) is a key part of social evolution theory, but the possible existence of such rules in social microbes has not been explored. In one specific context (biparental care in animals), cooperation is maintained if individuals respond to a partner's defection by increasing their own investment into cooperation, but not so much that this fully compensates for the defector's lack of investment. This is termed ‘partial compensation’. Here, I show that partial compensation for the presence of noncooperating ‘cheats’ is also observed in a microbial social behaviour: the cooperative production of iron‐scavenging siderophores by the bacterium Pseudomonas aeruginosa. A period of evolution in the presence of cheats maintains this response, whereas evolution in the absence of cheats leads to a loss of compensatory behaviour. These results demonstrate (i) the remarkable flexibility of bacterial social behaviour, (ii) the potential generality of partial compensation as a social response rule and (iii) the need for mathematical models to explore the evolution of response rules in multi‐player social interactions.     

Display behaviour,social organization and vocal repertoire of Blue‐backed Manakin Chiroxiphia pareola napensis in northwest Amazonia

Members of the avian family Pipridae (manakins) have served as a model system for studies aimed at understanding cooperation among individuals, traits involved in sexual selection and the evolution of female preferences. These studies rely on a detailed natural history baseline, yet multiple manakin taxa remain unstudied, precluding comparative analyses and assessment of geographical variation. The Blue‐backed Manakin Chiroxiphia pareola is a widely distributed species with four recognized subspecies; however, most published information on display behaviour, social organization and vocalizations are derived from an island population of one subspecies, Chiroxiphia pareola atlantica. To describe the social organization and patterns of activity of the subspecies C. p. napensis, we studied a population in eastern Ecuador and collected data from seven display areas or leks within the Tiputini Biodiversity Station. We used focal observations, camera traps and automatic song recorders to collect information for the field seasons from January to March 2013 and video cameras from February to March 2014. Males were active most of the day with a peak in activity at noon. Display areas were occupied by one to five males, ranging from > 2 to > 18 years old. The vocal repertoire included duet song displays and a variety of calls. We provide sonograms for all six observed vocalizations, including two previously unrecorded vocalizations for the species, and comment on the possible behavioural context of each vocalization. We describe five distinct display elements and the dance display behaviour, and highlight differences relative to other species of the genus. The first part of the dance display involved cooperative dancing, whereas the second part was a solo display, usually performed by alpha males. Our work reveals some differences between C. p. napensis and C. p. atlantica, such as a possible aggressive interaction between males of different ages and hierarchical status that has not been reported before and the apparent absence of mechanical wing sounds in C. p. napensis. The data reported here help advance our knowledge of the evolution of display behaviour among manakins.     

Kin selection and the evolution of virulence

Social interactions between conspecific parasites are partly dependent on the relatedness of interacting parasites (kin selection), which, in turn, is predicted to affect the extent of damage they cause their hosts (virulence). High relatedness is generally assumed to favour less competitive interactions, but the relationship between relatedness and virulence is crucially dependent on the social behaviour in question. Here, we discuss the rather limited body of experimental work that addresses how kin-selected social behaviours affect virulence. First, if prudent use of host resources (a form of cooperation) maximizes the transmission success of the parasite population, decreased relatedness is predicted to result in increased host exploitation and virulence. Experimental support for this well-established theoretical result is surprisingly limited. Second, if parasite within-host growth rate is a positive function of cooperation (that is, when individuals need to donate public goods, such as extracellular enzymes), virulence is predicted to increase with increasing relatedness. The limited studies testing this hypothesis are broadly consistent with this prediction. Finally, there is some empirical evidence supporting theory that suggests that spiteful behaviours are maximized at intermediate degrees of relatedness, which, in turn, leads to minimal virulence because of the reduced growth rate of the infecting population. We highlight the need for further thorough experimentation on the role of kin selection in the evolution of virulence and identify additional biological complexities to these simple frameworks.     

Dispersal, philopatry and intergroup relatedness: fine-scale genetic structure in the white-breasted thrasher, Ramphocinclus brachyurus

Dispersal is a fundamental process influencing evolution, social behaviour, and the long-term persistence of populations. We use both observational and genetic data to investigate dispersal, kin-clustering and intergroup relatedness in the white-breasted thrasher, Ramphocinclus brachyurus, a cooperatively breeding bird that is globally endangered. Mark-resighting data suggested sex-biased dispersal, with females dispersing over greater distances while males remained philopatric. Accordingly, spatial autocorrelation analysis showed highly significant fine-scale genetic structure among males, but not among females. This fine-scale genetic structuring of the male population resulted in very high levels of relatedness between dominant males at neighbouring nests, similar to that seen within cooperative groups in many species where kin selection is cited as a cause of cooperation. By implication, between-group as well as within-group cooperation may be important, potentially creating a feedback loop in which short-distance dispersal by males leads to the formation of male kin clusters that in turn facilitate nepotistic interactions and favour further local recruitment. The strength of spatial autocorrelation, as measured by the autocorrelation coefficient, r, was approximately two to three times greater than that reported in previous studies of animals. Relatively short dispersal distances by both males and females may have a negative impact on the white-breasted thrasher's ability to colonize new areas, and may influence the long-term persistence of isolated populations. This should be taken into account when designating protected areas or selecting sites for habitat restoration.     

Cooperation prevails when individuals adjust their social ties

Conventional evolutionary game theory predicts that natural selection favours the selfish and strong even though cooperative interactions thrive at all levels of organization in living systems. Recent investigations demonstrated that a limiting factor for the evolution of cooperative interactions is the way in which they are organized, cooperators becoming evolutionarily competitive whenever individuals are constrained to interact with few others along the edges of networks with low average connectivity. Despite this insight, the conundrum of cooperation remains since recent empirical data shows that real networks exhibit typically high average connectivity and associated single-to-broad–scale heterogeneity. Here, a computational model is constructed in which individuals are able to self-organize both their strategy and their social ties throughout evolution, based exclusively on their self-interest. We show that the entangled evolution of individual strategy and network structure constitutes a key mechanism for the sustainability of cooperation in social networks. For a given average connectivity of the population, there is a critical value for the ratio W between the time scales associated with the evolution of strategy and of structure above which cooperators wipe out defectors. Moreover, the emerging social networks exhibit an overall heterogeneity that accounts very well for the diversity of patterns recently found in acquired data on social networks. Finally, heterogeneity is found to become maximal when W reaches its critical value. These results show that simple topological dynamics reflecting the individual capacity for self-organization of social ties can produce realistic networks of high average connectivity with associated single-to-broad–scale heterogeneity. On the other hand, they show that cooperation cannot evolve as a result of “social viscosity” alone in heterogeneous networks with high average connectivity, requiring the additional mechanism of topological co-evolution to ensure the survival of cooperative behaviour.     

Costs for switching partners reduce network dynamics but not cooperative behaviour

Social networks represent the structuring of interactions between group members. Above all, many interactions are profoundly cooperative in humans and other animals. In accordance with this natural observation, theoretical work demonstrates that certain network structures favour the evolution of cooperation. Yet, recent experimental evidence suggests that static networks do not enhance cooperative behaviour in humans. By contrast, dynamic networks do foster cooperation. However, costs associated with dynamism such as time or resource investments in finding and establishing new partnerships have been neglected so far. Here, we show that human participants are much less likely to break links when costs arise for building new links. Especially, when costs were high, the network was nearly static. Surprisingly, cooperation levels in Prisoner''s Dilemma games were not affected by reduced dynamism in social networks. We conclude that the mere potential to quit collaborations is sufficient in humans to reach high levels of cooperative behaviour. Effects of self-structuring processes or assortment on the network played a minor role: participants simply adjusted their cooperative behaviour in response to the threats of losing a partner or of being expelled.     

Male mating behaviour and sperm production characteristics under varying sperm competition risk in guppies

Since natural populations of guppies, Poecilia reticulata, often differ from one another in social structure, the intensity of sperm competition is likely to vary between localities. Guppies are promiscuous, with female choice for colourful males playing a central role in the mating system. In addition, male guppies use forced copulations to circumvent female choice. Both methods of copulation are used interchangeably by individual males, but the degree to which either is used may depend on the social environment into which males are born. Here we show that male mating behaviour varies according to the rearing sex ratio: when reared in male-biased groups, males performed more forced copulations and fewer courtship displays but showed the opposite pattern of behaviour when reared in female-biased groups. Our prediction, based on sperm competition theory, that stripped sperm number would reflect social structure was not supported by our results. Instead, the overall level of sexual activity (gonopodial thrusts+sigmoid displays) was a better predictor of sperm number in the different groups of males. Rearing density, where sex ratio was controlled, did not significantly affect male mating behaviour or sperm traits. Males reared under the different sex ratios continued to show their characteristic behaviour patterns when placed in equal sex ratio tanks. We conclude, therefore, that males adopt mating strategies to suit their social environment, and that these strategies remain fixed, for short periods at least, if population structure changes. Copyright 1999 The Association for the Study of Animal Behaviour.     

Correlated pay-offs are key to cooperation

The general belief that cooperation and altruism in social groups result primarily from kin selection has recently been challenged, not least because results from cooperatively breeding insects and vertebrates have shown that groups may be composed mainly of non-relatives. This allows testing predictions of reciprocity theory without the confounding effect of relatedness. Here, we review complementary and alternative evolutionary mechanisms to kin selection theory and provide empirical examples of cooperative behaviour among unrelated individuals in a wide range of taxa. In particular, we focus on the different forms of reciprocity and on their underlying decision rules, asking about evolutionary stability, the conditions selecting for reciprocity and the factors constraining reciprocal cooperation. We find that neither the cognitive requirements of reciprocal cooperation nor the often sequential nature of interactions are insuperable stumbling blocks for the evolution of reciprocity. We argue that simple decision rules such as ‘help anyone if helped by someone’ should get more attention in future research, because empirical studies show that animals apply such rules, and theoretical models find that they can create stable levels of cooperation under a wide range of conditions. Owing to its simplicity, behaviour based on such a heuristic may in fact be ubiquitous. Finally, we argue that the evolution of exchange and trading of service and commodities among social partners needs greater scientific focus.