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Models relating intake rate to food abundance and competitor density (generalized functional response models) can predict forager distributions and movements between patches, but we lack understanding of how distributions and small-scale movements by the foragers themselves affect intake rates., Using a state-of-the-art approach based on continuous-time Markov chain dynamics, we add realism to classic functional response models by acknowledging that the chances to encounter food and competitors are influenced by movement decisions, and, vice versa, that movement decisions are influenced by these encounters., We used a multi-state modelling framework to construct a stochastic functional response model in which foragers alternate between three behavioural states: searching, handling and moving., Using behavioural observations on a molluscivore migrant shorebird (red knot, Calidris canutus canutus), at its main wintering area (Banc d'Arguin, Mauritania), we estimated transition rates between foraging states as a function of conspecific densities and densities of the two main bivalve prey., Intake rate decreased with conspecific density. This interference effect was not due to decreased searching efficiency, but resulted from time lost to avoidance movements., Red knots showed a strong functional response to one prey ( Dosinia isocardia), but a weak response to the other prey ( Loripes lucinalis). This corroborates predictions from a recently developed optimal diet model that accounts for the mildly toxic effects due to consuming Loripes., Using model averaging across the most plausible multi-state models, the fully parameterized functional response model was then used to predict intake rate for an independent data set on habitat choice by red knot., Comparison of the sites selected by red knots with random sampling sites showed that the birds fed at sites with higher than average Loripes and Dosinia densities, that is sites for which we predicted higher than average intake rates., We discuss the limitations of Holling's classic functional response model which ignores movement and the limitations of contemporary movement ecological theory that ignores consumer-resource interactions. With the rapid advancement of technologies to track movements of individual foragers at fine spatial scales, the time is ripe to integrate descriptive tracking studies with stochastic movement-based functional response models. [ABSTRACT FROM AUTHOR]

Mechanistic insights and predictive understanding of the spatial distributions of foragers are typically derived by fitting either field measurements on intake rates and food abundance, or observations from controlled experiments, to functional response models. It has remained unclear, however, whether and why one approach should be favoured above the other, as direct comparative studies are rare., The field measurements required to parameterize either single or multi-species functional response models are relatively easy to obtain, except at sites with low food densities and at places with high food densities, as the former will be avoided and the second will be rare. Also, in foragers facing a digestive bottleneck, intake rates (calculated over total time) will be constant over a wide range of food densities. In addition, interference effects may depress intake rates further. All of this hinders the appropriate estimation of parameters such as the 'instantaneous area of discovery' and the handling time, using a type II functional response model also known as 'Holling's disc equation'., Here we compare field- and controlled experimental measurements of intake rate as a function of food abundance in female bar-tailed godwits Limosa lapponica feeding on lugworms Arenicola marina., We show that a fit of the type II functional response model to field measurements predicts lower intake rates (about 2·5 times), longer handling times (about 4 times) and lower 'instantaneous areas of discovery' (about 30-70 times), compared with measurements from controlled experimental conditions., In agreement with the assumptions of Holling's disc equation, under controlled experimental settings both the instantaneous area of discovery and the handling time remained constant with an increase in food density. The field data, however, would lead us to conclude that although handling time remains constant, the instantaneous area of discovery decreased with increasing prey densities. This will result into highly underestimated sensory capacities when using field data., Our results demonstrate that the elucidation of the fundamental mechanisms behind prey detection and prey processing capacities of a species necessitates measurements of functional response functions under the whole range of prey densities on solitary feeding individuals, which is only possible under controlled conditions. Field measurements yield 'consistency tests' of the distributional patterns in a specific ecological context. [ABSTRACT FROM AUTHOR]

1. During migratory stopovers, animals are under strong time stress and need to maximize intake rates. We examine how foragers react to resource depletion by studying the foraging ecology and foraging site selection of black-tailed godwits Limosa l. limosa staging in rice fields during their northward migration stopover (January–March 2007). 2. We analysed godwit abundance and foraging behaviour, sampled the availability of rice in the fields and used the functional response model to predict the giving-up density (GUD) of rice kernels when godwits should give up a rice field. Sightings of individually colour-marked birds were used to verify whether individuals moving between rice fields confirmed the predicted GUD. 3. Black-tailed godwit intake rates at different rice densities fitted Holling’s functional response curve. The predicted GUD of rice necessary to balance allometric estimates of daily energy expenditure (DEE) and measured time budgets were confirmed by GUD measured in the field. 4. Individually marked birds moved towards rice fields with higher rather than lower rice densities more often than randomly expected. These birds increased the measured intake rates after this move. 5. Godwit foraging caused a decrease in the rice density of individual fields during the stopover period. Despite this, overall intake rates remained constant as godwits reacted to resource depletion by moving to a new foraging site as soon as their intake rate falls below the required levels to achieve DEE. [ABSTRACT FROM AUTHOR]

1. Within the broad field of optimal foraging, it is increasingly acknowledged that animals often face digestive constraints rather than constraints on rates of food collection. This therefore calls for a formalization of how animals could optimize food absorption rates. 2. Here we generate predictions from a simple graphical optimal digestion model for foragers that aim to maximize their (true) metabolizable food intake over total time (i.e. including nonforaging bouts) under a digestive constraint. 3. The model predicts that such foragers should maintain a constant food retention time, even if gut length or food quality changes. For phenotypically flexible foragers, which are able to change the size of their digestive machinery, this means that an increase in gut length should go hand in hand with an increase in gross intake rate. It also means that better quality food should be digested more efficiently. 4. These latter two predictions are tested in a large avian long-distance migrant, the Bewick's swan ( Cygnus columbianus bewickii), feeding on grasslands in its Dutch wintering quarters. 5. Throughout winter, free-ranging Bewick's swans, growing a longer gut and experiencing improved food quality, increased their gross intake rate (i.e. bite rate) and showed a higher digestive efficiency. These responses were in accordance with the model and suggest maintenance of a constant food retention time. 6. These changes doubled the birds’ absorption rate. Had only food quality changed (and not gut length), then absorption rate would have increased by only 67%; absorption rate would have increased by only 17% had only gut length changed (and not food quality). 7. The prediction that gross intake rate should go up with gut length parallels the mechanism included in some proximate models of foraging that feeding motivation scales inversely to gut fullness. We plea for a tighter integration between ultimate and proximate foraging models. [ABSTRACT FROM AUTHOR]

1. Rate-maximizing foragers that only divide their time between searching and handling prey should, according to the classical contingency model (CM), only select those prey whose energy content per unit handling time (i.e. profitability) exceeds or equals long-term average energy intake rate.2. However, if digestively constrained foragers were to follow this so-called‘zero-one rule’, they would need to take digestive breaks and their energy intake over total time would not be maximized. They should, according to the digestive rate model (DRM), also consider the rate at which a prey type is digested (i.e. digestive quality), such that time lost to digestive breaks is minimized.3. In three different contexts, we tested these competing models in a mollusc-eating shorebird, the red knot (Calidris canutus), that is often digestively constrained due to its habit of ingesting its bulky prey whole. Measurements on gizzard size (using ultrasonography) and prey-characteristics confirmed that in each test the birds were digestively bottlenecked and should thus follow the DRM in order to maximize long-term energy intake.4. In the first experiment, knots were offered a choice between two fully exposed prey, and tended to select prey by the criterion of digestive quality rather than profitability.5. In the second experiment, knots were offered two buried prey types and preferred the highest quality prey to the most profitable prey.6. In the wild, knots mainly fed on high qualityMyaand largely ignored poor quality, but equally profitable,Cerastoderma.7. Thus, each test verified the predictions of the DRM and rejected those of the CM. Given that many species face digestion constraints, we expect that the DRM is likely to explain diet composition in many more studies.Journal of Animal Ecology(2005) doi: 10.1111/j.1365-2656.2004.00903.x [ABSTRACT FROM AUTHOR]

1. Theoretical studies of large herbivore foraging assume that total daily grazing time is a key constraint on daily intake and diet choice. We experimentally tested this assumption and investigated the effects of food availability on the ability of grazing sheep to compensate for restriction of available daily grazing time. 2. Foraging behaviour, intake and diet digestibility by sheep, were measured on grass pastures in a replicated 2 × 2 factorial experiment, in which overnight access to pasture was varied (restricted overnight and continuous access) on two sward heights (5·5 and 3·0 cm), representing high and low food availability. 3. Regardless of food availability, the overnight-restricted sheep fed for almost all of the available grazing time by grazing for fewer, longer foraging bouts, but still had much shorter total daily grazing time than the continuous access sheep. 4. In response to overnight penning, the sheep had a significantly higher instantaneous rate of intake achieved mainly via larger bites. The continuous access sheep were hence not maximizing their short-term rate of intake, whilst grazing according to the daily schedule considered normal for sheep. 5. The behavioural responses to overnight food restriction were able to counteract the reduction in daily grazing time only where food availability was high. In contrast on short swards overnight grazing restriction led to a reduction in total daily intake. We suggest that the interactions between the factors considered as constraints on foraging behaviour of herbivores are, as yet, only poorly quantified. [ABSTRACT FROM AUTHOR]

Models relating intake rate to food abundance and competitor density (generalized functional response models) can predict forager distributions and movements between patches, but we lack understanding of how distributions and small-scale movements by the foragers themselves affect intake rates. Using a state-of-the-art approach based on continuous-time Markov chain dynamics, we add realism to classic functional response models by acknowledging that the chances to encounter food and competitors are influenced by movement decisions, and, vice versa, that movement decisions are influenced by these encounters. We used a multi-state modelling framework to construct a stochastic functional response model in which foragers alternate between three behavioural states: searching, handling and moving. Using behavioural observations on a molluscivore migrant shorebird (red knot, Calidris canutus canutus), at its main wintering area (Banc d'Arguin, Mauritania), we estimated transition rates between foraging states as a function of conspecific densities and densities of the two main bivalve prey. Intake rate decreased with conspecific density. This interference effect was not due to decreased searching efficiency, but resulted from time lost to avoidance movements. Red knots showed a strong functional response to one prey (Dosinia isocardia), but a weak response to the other prey (Loripes lucinalis). This corroborates predictions from a recently developed optimal diet model that accounts for the mildly toxic effects due to consuming Loripes. Using model averaging across the most plausible multi-state models, the fully parameterized functional response model was then used to predict intake rate for an independent data set on habitat choice by red knot. Comparison of the sites selected by red knots with random sampling sites showed that the birds fed at sites with higher than average Loripes and Dosinia densities, that is sites for which we predicted higher than average intake rates. We discuss the limitations of Holling's classic functional response model which ignores movement and the limitations of contemporary movement ecological theory that ignores consumer-resource interactions. With the rapid advancement of technologies to track movements of individual foragers at fine spatial scales, the time is ripe to integrate descriptive tracking studies with stochastic movement-based functional response models., (© 2014 The Authors. Journal of Animal Ecology © 2014 British Ecological Society.)