Cheats in a cooperative behaviour? Behavioural differences and breakdown of cooperative behaviour in aggregating, intertidal littorinids (Mollusca)

Cooperative behaviours result in the evolution of cheats: individuals that benefit from the behaviour without sharing the costs required to generate the behaviour. Normally the proportion of cheats is small, as large numbers of cheats will result in the breakdown of the behaviour. Using empirical and simulation-derived results we demonstrate a cooperative behaviour (aggregation between two species of intertidal snails that provides a benefit by reducing desiccation stress) that shows many characteristics similar to those of a cooperative system with cheats present. In this system, the high rocky shore littorinid Echinolittorina malaccana forages for longer after high water than Echinolittorina radiata, which stops foraging and begins to form aggregations earlier. Nevertheless, E. malaccana, the 'cheat' in this system, still occupies an equal proportion of the most beneficial places in aggregations. Computer simulations demonstrate that up to 65% of individual snails can show the behaviour of E. malaccana before the breakdown of this aggregation behaviour begins to occur through aggregations becoming smaller, and hence less effective against desiccation. The high proportion of 'cheats' possible in this cooperative behaviour implies that different selective pressures may act on individuals of different species in multi-species cooperative behaviours to those acting on individuals engaging in single species cooperative behaviours. Social symbiosis appears to be occurring between the species, but it appears that both mutualistic and parasitic symbioses are occurring.