Commodity-specific punishment for experimentally induced defection in cooperatively breeding fish

Coercion is an important but underrated component in the evolution of cooperative behaviour. According to the pay-to-stay hypothesis of cooperative breeding, subordinates trade alloparental care for the concession to stay in the group. Punishment of idle subordinates is a key prediction of this hypothesis, which has received some experimental scrutiny. However, previous studies neither allowed separating between punishment and effects of disruption of social dynamics, nor did they differentiate between different helping behaviours that may reflect either mutualistic or reciprocal interaction dynamics. In the cooperative breeder Neolamprologus pulcher, we experimentally engineered the ability of subordinates to contribute to alloparental care by manipulating two different helping behaviours independently from one another in a full factorial design. We recorded the treatment effects on breeder aggression, subordinate helping efforts and submissive displays. We found two divergent regulatory mechanisms of cooperation, dependent on behavioural function. Experimental impediment of territory maintenance of subordinates triggered punishment by dominants, whereas prevented defence against egg predators released a compensatory response of subordinates without any enforcement, suggesting pre-emptive appeasement. These effects occurred independently of one another. Apparently, in the complex negotiation process among members of cooperative groups, behaviours fulfilling different functions may be regulated by divergent interaction mechanisms.