Subordinate removal affects parental investment, but not offspring survival in a cooperative cichlid

Summary Subordinates in cooperative breeding systems may provide help to dominant pairs, who can benefit by either an increased total investment in their current brood or a reduced personal contribution to this investment. In the social cichlid Julidochromis ornatus, one large male subordinate generally spends 90% of his time in close proximity to the breeding shelter, whereas the dominants only spend 50% of their time close to the shelter. We experimentally removed the large subordinate for 30 days (approximating one breeding cycle) to study the investment strategies of dominants and the effects on offspring survival, while accounting for subordinate immigration. Experimental groups were compared with control groups, from which subordinates were also caught but not removed. On day one following removal, we tested whether dominants overcompensated, fully compensated or undercompensated for absence of the subordinate on several parental behaviours. Moreover, we tested whether the pairs' potential compensatory behaviour remained high seven days following large subordinate removal. One day following removal, dominants increased their time spent in the territory and their frequency of breeding shelter visits and defence, compared with the pre-removal phase and control groups. The dominant pair overcompensated for the loss of subordinate help in their breeding shelter visits, fully compensated in defence and undercompensated their time spent in the territory. Seven days after large subordinate removal, behavioural differences between treatments had disappeared. However, when distinguishing between groups with or without a new immigrant subordinate, dominant pairs only diminished investment in the presence of an immigrant, suggesting a compensatory role of the large subordinate. Finally, survival of juvenile group members was not affected by the treatment. Our experiments indicate that the presence of a large subordinate does not increase the dominant pairs' current reproductive success, but instead allows them to reduce their personal contribution to investment in the current brood. In addition, we illustrate that dominants may show strikingly different compensatory responses depending on the type of behaviour and emphasize the importance of immigrant subordinates to relieve dominants from costly compensatory responses in cooperative breeding systems.