Turn-taking in cooperative offspring care: by-product of individual provisioning behavior or active response rule?

ABSTRACT: For individuals collaborating to rear offspring, effective organization of resource delivery is difficult because each carer benefits when the others provide a greater share of the total investment required. When investment is provided in discrete events, one possible solution is to adopt a turn-taking strategy whereby each individual reduces its contribution rate after investing, only increasing its rate again once another carer contributes. To test whether turn-taking occurs in a natural cooperative care system, here we use a continuous time Markov model to deduce the provisioning behavior of the chestnut-crowned babbler (Pomatostomus ruficeps), a cooperatively breeding Australian bird with variable number of carers. Our analysis suggests that turn-taking occurs across a range of group sizes (2-6), with individual birds being more likely to visit following other individuals than to make repeat visits. We show using a randomization test that some of this apparent turn-taking arises as a by-product of the distribution of individual inter-visit intervals ("passive" turn-taking) but that individuals also respond actively to the investment of others over and above this effect ("active" turn-taking). We conclude that turn-taking in babblers is a consequence of both their individual provisioning behavior and deliberate response rules, with the former effect arising through a minimum interval required to forage and travel to and from the nest. Our results reinforce the importance of considering fine-scale investment dynamics when studying parental care and suggest that behavioral rules such as turn-taking may be more common than previously thought. SIGNIFICANCE STATEMENT: Caring for offspring is a crucial stage in the life histories of many animals and often involves conflict as each carer typically benefits when others contribute a greater share of the work required. One way to resolve this conflict is to monitor when other carers contribute and adopt a simple "turn-taking" rule to ensure fairness, but natural parental care has rarely been studied in sufficient detail to identify such rules. Our study investigates whether cooperatively breeding chestnut-crowned babblers "take turns" delivering food to offspring, and (if so) whether this a deliberate strategy or simply a by-product of independent care behavior. We find that babblers indeed take turns and conclude that part of the observed turn-taking is due to deliberate responsiveness, with the rest arising from the species' breeding ecology