Context-dependent social benefits drive cooperative predator defense in a bird

Understanding the major evolutionary transition from solitary individuals to complex societies is hampered by incomplete insight into the drivers of living in cooperative groups.^1-3 This may be because the benefits of sociality can derive from group living itself (e.g., dilution of predation risk),^4,5 or depend on social context (e.g., kin or potential mates represent beneficial group members).^6-8 Cooperative breeders, where non-breeding subordinates assist breeders, have provided important insights into the drivers of cooperation, but comprehensive assessment of diverse potential benefits has been hindered by a prevailing focus on benefits deriving from raising offspring.^9-11 We propose a novel paradigm to tease apart different benefits by comparing cooperative responses to predators threatening dependent young and adult group members according to their value for the responding individual. Applying this approach in purple-crowned fairy-wrens, Malurus coronatus, we show that non-breeding subordinates are more responsive to nest predators-a threat to offspring-when their probability of inheriting a breeding position is greater-irrespective of group size, relatedness to offspring, or opportunity to showcase individual quality to potential mates. This suggests that offspring defense is modulated according to the benefits of raising future helpers. Conversely, when predators pose a threat to adults, responsiveness depends on social context: subordinates respond more often when kin or potential mates are under threat, or when group members are associated with mutualistic social bonds, indirect genetic benefits, and future reproductive benefits.^9,12,13 Our results demonstrate that direct and kin-selected benefits of sociality are context dependent, and highlight the importance of predation risk in driving complex sociality.