Direct benefits explain interspecific variation in helping behaviour among cooperatively breeding birds

Kin selection theory provides one important explanation for seemingly altruistic helping behaviour by non-breeding subordinates in cooperative breeding animals. However, it cannot explain why helpers in many species provide energetically costly care to unrelated offspring. Here, I use comparative analyses to show that direct fitness benefits of helping others, associated with future opportunities to breed in the resident territory, are responsible for the widespread variation in helping effort (offspring food provisioning) and kin discrimination across cooperatively breeding birds. In species where prospects of territory inheritance are larger, subordinates provide more help, and, unlike subordinates that cannot inherit a territory, do not preferentially direct care towards related offspring. Thus, while kin selection can underlie helping behaviour in some species, direct benefits are much more important than currently recognised and explain why unrelated individuals provide substantial help in many bird species

The cost of prospecting for dispersal opportunities in a social bird

Understanding why individuals delay dispersal and become subordinates within a group is central to studying the evolution of sociality. Hypotheses predict that dispersal decisions are influenced by costs of extra-territorial prospecting that are often required to find a breeding vacancy. Little is known about such costs, partly because it is complicated to demonstrate them empirically. For example, prospecting individuals may be of inferior quality already before prospecting and/or have been evicted. Moreover, costs of prospecting are mainly studied in species where prospectors suffer from predation risk, so how costly prospecting is when predators are absent remains unclear. Here, we determine a cost of prospecting for subordinate Seychelles warblers, Acrocephalus sechellensis, in a population where predators are absent and individuals return to their resident territory after prospecting. Prospecting individuals had 5.2% lower body mass than non-prospecting individuals. Our evidence suggests this may be owing to frequent attacks by resident conspecifics, likely leading to reduced food intake by prospectors. These results support the hypothesis that energetic costs associated with dispersal opportunities are one factor influencing dispersal decisions and shaping the evolution of delayed dispersal in social animals

The evolution of delayed dispersal and different routes to breeding in social birds

Many animals live in stable groups, where sexually mature individuals delay dispersal and stay as nonbreeding subordinates, seemingly counter to their own evolutionary interests. Revealing what circumstances drive the evolution of delayed dispersal is central to understanding sociality, family living and cooperative breeding across the animal kingdom, but there is as yet no general consensus about the relative importance of the various ecological and social conditions and the reproductive benefits proposed to drive delayed dispersal. We argue that two components may facilitate further progress in this respect: firstly, full consideration of the various routes that individuals can follow to obtain an independent breeding position. Here, we provide a comprehensive review of these routes: inheritance of a natal territory, budding off part of the natal territory, shifting to a neighboring vacancy, making temporary prospecting trips throughout the population; or permanently leaving to float in search of a breeding position or to stage as subordinate in a non-natal territory. Second, we illustrate that in order to understand delayed dispersal, we need to consider that the fitness consequences of these different routes apply across the lifetime: as subordinate (e.g., benefits of philopatry and indirect fitness); while waiting or searching for a position; and after obtaining a breeding position. Overall, we conclude that by which route and under what circumstances individuals can obtain a breeding position must be considered in order to make more comprehensive inferences about the evolution of delayed dispersal, cooperative breeding and animal sociality as a whole.</p

Experimentally induced antipredator responses are mediated by social and environmental factors

Nest predation is a common cause of reproductive failure for many bird species, and various antipredator defense behaviors have evolved to reduce the risk of nest predation. However, trade-offs between current reproductive duties and future reproduction often limit the parent’s ability to respond to nest predation risk. Individual responses to experimentally increased nest predation risk can give insights into these trade-offs. Here, we investigate whether social and ecological factors affect individual responses to predation risk by experimentally manipulating the risk of nest predation using taxidermic mounts in the cooperative breeding Seychelles warbler (Acrocephalus sechellensis). Our results show that dominant females, but not males, alarm called more often when they confront a nest predator model alone than when they do so with a partner, and that individuals that confront a predator together attacked more than those that did so alone. Dominant males increased their antipredator defense by spending more time nest guarding after a presentation with a nest predator, compared with a nonpredator control, but no such effect was found for females, who did not increase the time spent incubating. In contrast to incubation by females, nest guarding responses by dominant males depended on the presence of other group members and food availability. These results suggest that while female investment in incubation is always high and not dependent on social and ecological conditions, males have a lower initial investment, which allows them to respond to sudden changes in nest predation risk

Delayed dispersal and the cost and benefits of different routes to independent breeding in a cooperative breeding bird

Why sexually mature individuals stay in groups as non-reproductive subordinates is central to the evolution of sociality and cooperative breeding. To understand such delayed dispersal, its costs and benefits need to be compared with those of permanently leaving to float through the population. However, comprehensive comparisons, especially regarding differences in future breeding opportunities, are rare. Moreover, extra-territorial prospecting by philopatric individuals has generally been ignored, even though the factors underlying this route to independent breeding may differ from those of strict philopatry or floating. We use a comprehensive predictive framework to explore how various costs, benefits and intrinsic, environmental and social factors explain philopatry, prospecting and floating in Seychelles warblers (Acrocephalus sechellensis). Floaters more likely obtained an independent breeding position before the next season than strictly philopatric individuals, but also suffered higher mortality. Prospecting yielded similar benefits to floating but lower mortality costs, suggesting that it is overall more beneficial than floating and strict philopatry. Whereas prospecting is probably individual-driven, though limited by resource availability, floating likely results from eviction by unrelated breeders. Such differences in proximate and ultimate factors underlying each route to independent breeding highlight the need for simultaneous consideration when studying the evolution of delayed dispersal

More than kin:Subordinates foster strong bonds with relatives and potential mates in a social bird

Social interactions shape relationships between individuals in complex societies. Affiliative interactions are associated with benefits and strengthen social bonds, while aggressive interactions are costly and negatively affect social bonds. Individuals may attempt to reduce aggressive encounters through submissive displays directed at higher-ranking individuals. Thus, fine-scale patterns of affilia-tive, aggressive, and submissive interactions may reflect costly and beneficial social relationships within groups, providing insight into the benefits of group living and the mechanisms of conflict resolution. So far, however, most studies have looked at social interactions and benefits of group living in isolation. We investigated how the strength of social bonds (affiliative vs. aggressive interactions) and submissive displays varied with kin-selected and potential mating benefits, and with reproductive conflict in the cooperatively breeding purple-crowned fairy-wren, Malurus coronatus. Our results revealed that subordinates formed equally strong social bonds with kin and potential mates (unrelated opposite-sex individuals) while they formed antagonistic relationships with reproductive competitors that offered no kin-selected or mating benefits (unrelated same-sex individuals). Submissive displays were directed exclusively at same-sex breeders, regardless of relatedness. Affiliation and submission were associated with reduced foraging time when food was limited, indicating a cost to maintaining positive relationships. Together, our results suggest that the strength of social bonds is determined by (potential) benefits obtained from group members, while submission likely serves to reduce conflict. Our findings highlight the importance of time-costly social interactions for maintaining relationships with group members, providing insight into how social groups of individuals with (partly) divergent interests can remain stable.</p