Client fish ectoparasite loads and cleaner shrimp Urocaridella sp. c hunger levels affect cleaning behaviour

Cleaning is a classic example of mutualism and determining the factors that maintain the balance between the costs and benefits for mutualist partners can assist our understanding of how cleaning relationships are maintained. Optimal foraging theory suggests two factors that might help to maintain the relationship between cleaners and their clients: client ectoparasite load and cleaner hunger levels. The ecological relevance and importance of foraging by cleaner fish in marine systems has been demonstrated repeatedly, yet there is little information available on this behaviour in cleaner shrimp. To determine whether cleaner shrimp base their choice of client fish on food patch quality (i.e. client fish ectoparasite load) we offered the yellow-beaked cleaner shrimp Urocaridella sp. c a choice of parasitized and unparasitized rock cods, Cephalopholis cyanostigma. To determine whether cleaner shrimp hunger levels influence cleaning time, we manipulated hunger levels in Urocaridella sp. c and examined their behaviour towards parasitized client fish. Cleaner shrimp preferred parasitized to unparasitized client fish and food-deprived cleaner shrimp cleaned parasitized rock cods more frequently than satiated cleaner shrimp did. Therefore, variations in client fish ectoparasite load and cleaner shrimp hunger level are two factors that affect the balance in this mutualism. Finally, our results meet some of the assumptions of biological market theory, a framework used to understand cooperative interactions, and thus this framework is suggested for future studies on this cleaning system.