Revisiting GUD: An empirical test of the size-dependency of patch departure behaviour.

Behaviour related to patch resource exploitation is a major determinant of individual fitness. Assuming the size-dependency of patch departure behaviour, model-based approaches have shown size-mediated coexistence in systems of competing species. However, experimental evidence for the influence of body size on patch use behaviour is scarce. In this study, we explore whether allometric principles provide an underlying framework for interspecific patterns of resource use. To this end, we propose a meso-cosm approach using three species of gastropods differing in size as a model system and 32P radio-isotopic techniques as a measure of resource use. Foragers of different size were placed in an artificial patch, provided with a limited amount of labelled resource and let them free to move as resources decrease and scarcity is sensed. We investigated the extent to which individual body size affects the exploitation of resources by examining Giving Up Density (GUD), Giving Up Time (GUT), resource absorption rate and exploitation efficiency as components of individual exploitation behaviour. To compare positive, constant and negative individual size scaling of population energy requirements, experimental trials with an equal numbers and equal biomass of differently sized foragers were carried out, and an experimental trial with equal metabolic requirements was simulated. We observed clear size dependency in the patch departure behaviour of the experimental organisms. Even under conditions of equivalent overall population energy requirements, larger foragers decided to leave the resource patch earlier and at a higher density of resources than smaller ones. Smaller foragers were able to prolong their presence and make more use of the resources, resulting in an inverse body-size scaling of resource exploitation efficiency.