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

Date Published: September 27, 2018

Publisher: Public Library of Science

Author(s): Francesco Cozzoli, Giovanna Ligetta, Fabio Vignes, Alberto Basset, Samantha Munroe.


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.

Partial Text

The acquisition of resources is a major determinant of vagile organisms’ ecology and behaviour. It involves decisions regarding where to search, when to feed, which food types to consume and when to terminate feeding [1,2,3,4,5]. An optimal foraging strategy serves to maximize the resource acquisition under the constraints of the environment [6]. Short-term feeding goals [7] can be achieved by following criteria such as maximising the rate of net energy intake [8,9], minimising the variance in the rate of net energy intake [10,11,2,7] and minimising the time spent foraging [3].

On average, the E. ventrosa individuals used in this experiment weighed 0.37 mg AFDW [± 0.05 95% Confidence Interval]. The average weight of B. tentaculata was 7.77 mg AFDW [± 2.99] and that of G. truncatula was 12.88 mg AFDW [± 2.54] (Table 1). Average individual metabolic rates estimated from Brey’s empirical model (Brey, An empirical model for estimating aquatic invertebrate respiration, 2010) were 0.31 mJ day-1 [± 0.11] for E. ventrosa, 3.49 mJ day-1 [± 1.42] for B. tentaculata and 5.25 mJ day-1 [± 2.16] for G. truncatula (Table 1).

Overall, we observed clear size dependencies in the patch departure behaviour of the studied model organisms. Larger foragers tended to leave the resource patch earlier and at a higher density of resources than smaller ones. Smaller foragers, able to sustain themselves with lower ingestion rates, had a longer Giving Up Times and a lower Giving Up Densities, and made a more efficient use of resources at the patch level. The observed negative size-scaling of GUT and GUD and positive size-scaling of patch exploitation efficiency are in agreement with both theoretical expectations [27,30,24] and previous empirical evidence [31,33,34].

While much effort has been devoted to investigating the influence of extrinsic factors (e.g. habitat structure and ecological interactions such as predation) on resource exploitation behaviour, the intrinsic influence of forager energetics has yet to be fully addressed in GUD theory [17]. Previous research in this field has mainly focused on contextual and idiosyncratic issues such as satiation and forager development, physiology, health and reproductive state (Bedoya-Peréz et al. 2013 [17] and references within). In contrast, we investigated the systematic variation of patch departure behaviour using body size as a proxy for forager energy requirements. This study therefore frames GUD theory in the broader context of energy scaling research (e.g. the Metabolic Theory of Ecology, [22]). This topic is of particular interest in ecology, because size-dependencies in patch departure behaviour are believed to affect space use, home range size [20,95] species coexistence and resource partitioning [27,30,96]. The results of our experiments can be used to parametrise theoretical models of energetic carrying capacity [97,14], size-related species coexistence [30] and ecological community responses to climate change [93].




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