The effect of crowding and density on male mating behaviour in the rose bitterling (Rhodeus ocellatus)

Female density and resource availability are two key variables that shape mating systems. Theory predicts that reproductive skew will amplify with increased male density and decreasing availability of resources, though limited empirical evidence suggests that this may not always be the case. Here we tested mean crowding, defined as the number of males per unit of resource, and density per se, defined as the number of individuals present per unit area, to investigate their effect on the mating system of Rhodeus ocellatus, a fish with a promiscuous, resource-based mating system. Males were exposed to combinations of high and low levels of crowding and density, while the operational sex ratio was held constant. High levels of crowding significantly affected the proportion of mussel spawning sites defended by males and the proportion of mussels into which sperm was released. In contrast to theoretical predictions, neither density nor crowding influenced overall male aggressive behaviours. Density, but not crowding, had a significant effect on male courtship rate, which arose as a possible trade-off between intra-sexual competition and inter-sexual behaviour. We discuss the results in the context of mating system evolution.