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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]

The restricted area of space used by most mobile animals is thought to result from fitness-rewarding decisions derived from gaining information about the environment. Yet, assessments of how animals deal with uncertainty using memory have been largely theoretical, and an empirically derived mechanism explaining restricted space use in animals is still lacking. Using a patch-to-patch movement analysis, we investigated predictions of how free-ranging bison ( Bison bison) living in a meadow-forest matrix use memory to reduce uncertainty in energy intake rate. Results indicate that bison remembered pertinent information about location and quality of meadows, and they used this information to selectively move to meadows of higher profitability. Moreover, bison chose profitable meadows they had previously visited, and this choice was stronger after visiting a relatively poor quality meadow. Our work demonstrates a link between memory, energy gains and restricted space use while establishing a fitness-based integration of movement, cognitive and spatial ecology. [ABSTRACT FROM AUTHOR]

Understanding the role that consumers play in an ecosystem requires knowledge about food selection and intake rates. However, such basic data are often difficult to obtain, particularly for marine animals that are not easy to observe. To overcome this problem, a beak movement sensor was employed on a free-ranging loggerhead turtle in a neritic foraging habitat at the Domitian littoral ( SW Italy). In combination with gastrointestinal content analysis from six turtles found dead in the same area we sought to identify which beak movement patterns were associated with which prey type, and to quantify the ingestion of the various prey types. Brachyuran crabs (100% occurrence), in particular Liocarcinus vernalis, and small molluscs (66% occurrence) were found most frequently in the stomach and intestine of the turtles. Beak movements revealed average ingestion rates of (mean ± SE) 0.27 ± 0.13 food items per minute and that feeding occurred predominantly during dives >4 m and during early morning and evening. Interestingly, the time spent feeding amounted to only 2.2% of the total observation time, whereas feeding-associated dives added up to just above 10% of the total time. We thus established that loggerhead turtles in this area are specialised on brachyuran crabs, on which they prey with high success during the short time dedicated to foraging. This information strengthens our knowledge about turtle-prey and turtle-habitat interactions, which are essential data to delineate the role that turtles play in this and similar marine ecosystems. Moreover, since the same area is also intensively used by the regional bottom trawl fishery, our results have important conservation implications, because they clearly show the time of day and water depths for which fishing activity should be regulated to reduce the number of turtles that are currently being incidentally caught in this area. [ABSTRACT FROM AUTHOR]

Cadmium and zinc accumulation and toxicity were assessed in whole plants and callus culture of two Zygophyllum fabago populations originating from two metallicolous sites in Murcia (southeast Spain), La Peña and Mazarrón, the first containing 2.8-times more Cd and five-times more Zn than the second. Seedlings from both ecotypes were exposed for 3 weeks to 1 or 10 μm Cd, and to 10 or 100 μm Zn in nutrient solution in a controlled environment. Calli from both ecotypes were exposed to 0.01, 0.1 or 1 mm Cd, and to 0.1, 1 or 5 mm Zn. Plants from both populations exhibited similar tolerance to Zn, while tolerance to Cd appeared more important in plants from La Peña than those from Mazarrón. Only minor differences were recorded in final Cd accumulation, with higher Cd retention in roots and stems of plants from La Peña. In both populations, transient decreases in the rate of Zn intake and translocation from root to shoot were recorded. This reduction in ion uptake was not more efficient for the population from the most contaminated area compared to the less contaminated area. Similar concentrations of Cd were found in cotyledon-derived calli from the two populations, but absorbed Cd induced conspicuous water stress in calli issues from Mazarrón but not in those from La Peña. It is concluded that, beside comparable levels of heavy metal concentration in tissues, the physiological strategy of tolerance may differ according to the metal and according to the considered population. [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]

Mechanistic studies on herbivore functional responses have largely taken place in mammals; very little has been done in herbivorous birds so far. Here we aim to fill that gap by experimentally quantifying the (short-term) functional response of a large avian herbivore, the Bewick's Swan (Cygnus columbianus bewickii). We explicitly distinguish between encounter-limited and handling-limited foraging by analyzing the results in the framework of the models of D. E. Spalinger and N. T. Hobbs, originally developed for mammalian herbivory. Bite size in captive swans was experimentally manipulated by varying sward height. The time interval between two bites increased with bite size, which supports the handling-limited model (process 3) and rejects the encounter-limited models (processes 1 and 2). Subsequently, we took the obtained functional response parameters into the field in order to predict, from measurements of sward height, (1) bite sizes, (2) handling times, and (3) short-term intake rates in free-ranging swans. Indeed, for all three variables, the observed values closely matched the experimentally based predictions. Finally, we review functional response parameters available in the literature on avian herbivores and scale them allometrically in relation to mammals. This analysis revealed that maximum bite sizes, and therefore maximum intake rates, in herbivorous birds are smaller than in herbivorous mammals. We hypothesize and provide evidence that birds compensate by longer daily foraging times. [ABSTRACT FROM AUTHOR]

We present a novel technique using a video serial interface to link a motion-activated, infrared video camera and a weigh-scale to measure supplemental feed consumption by wildlife. This "video-scale" allows us to quantify consumption of various feeds by individual animals to an accuracy of 5 g and to record feeding duration to the second. Using intake rate and total time spent feeding, we calculated comparative use of supplemental feed by animals categorized by species, age, and sex. We present preliminary data on feed consumption and intake rates of wild and tame white-tailed deer (Odocoileus virginianus) and raccoons (Procyon lotor) eating shelled corn, pelleted supplement, and whole cottonseed. Other uses and limitations of the video-scale are discussed. [ABSTRACT FROM AUTHOR]

The distance that mammalian herbivores can travel without interrupting food processing corresponds to a distance threshold (d*) in plant spacing where change occurs in the mechanisms regulating the functional response. The instantaneous rate of food consumption is controlled by food encounter rate when plant spacing exceeds d°. Below this threshold, food processing in the mouth controls instantaneous intake rate. The distance threshold provides a mechanistic definition of the scale of heterogeneity of the food resource. Recent work indicates that d* should scale positively with the body mass of mammalian herbivores. Here I evaluated the empirical evidence for this positive scaling by investigating (I) herbivores consuming only alfalfa (Medicago sativa), (2) gazers, and (3) herbivores consuming any kind of vegetation. Overall, I found greater evidence for a negative than for a positive scaling of as. Out of the three groups, only herbivores consuming alfalfa could yield a positive covariation between d* and body mass. However, even this positive scaling became negative when herbivores consumed bites of alfalfa that were only a fraction of their maximum size. Finally, d* also scaled negatively among herbivores foraging in similar food patches. Overall, differences in foraging decisions among mammalian herbivores seem more likely to have been shaped by a negative than a positive scaling of d*. [ABSTRACT FROM AUTHOR]

Intake rate and prey size selection of 0+ perch, Perca fluviatilis, from Lake Wallersee (Austria) was studied at different illumination (day light: 400 lx, twilight: 2 lx) during 5 days of habituation to novel surroundings in aquaria. The hypothesis was tested that high illumination and novelty of surroundings (transfer from holding to test aquaria) as indirect cues of predation risk influences 0+ perch foraging behaviour. Significantly lower total intake and lower proportion of large Daphnia magna (two prey size groups, 2.9 and 1.2 mm, were used) were observed at higher illumination and under novel surroundings. Habituation to novelty caused an increase in consumption of large prey and decrease in consumption of small prey. During the whole period of habituation, more large prey was ingested at twilight than at daylight; no light-induced difference in ingestion of small prey was found. Foraging 0+ perch responded to indirect nonspecific cues of predation risk by reduction of intake of large prey, which are costly in terms of handling time. This allows fish to be more vigilant without ceasing their foraging activity even in potentially dangerous situations. In the lake, young perch are most vulnerable to abundant piscivorous fish and birds during the day; in twilight perch can afford a more time-consuming foraging activity. [ABSTRACT FROM AUTHOR]

Four intermediate-heading perennial ryegrass ( Lolium perenne L.) varieties, which in previous studies had been associated with high- or low-intake characteristics when swards containing them had been continuously stocked with sheep, were sown as monocultures. They were rotationally grazed, using 1-d paddocks, with core groups of four yearling Simmental × Holstein beef heifers in 2002 and 2003 and ingestive and ruminative behaviour, and sward factors, were measured. There were two diploid (Belramo and Glen) and one tetraploid (Rosalin) perennial ryegrass varieties and one tetraploid hybrid ( Lolium × boucheanum Kunth) (AberExcel) variety. Intake rate (IR) was significantly higher in August 2003 for heifers grazing Glen than those grazing Belramo [27·5 vs. 20·6 g dry matter (DM) min−1; P = 0·019], but there were no significant differences between varieties in two other measurement periods. This is in contrast to previous results with sheep when IR were significantly higher for Glen than Belramo and for AberExcel than Rosalin. Total jaw movement rates during grazing were significantly higher for heifers on the tetraploid swards than those on the diploid swards (87·7 vs. 83·6 jaw movements min−1; P = 0·023) in September 2002. Ruminating time was significantly lower for heifers on the tetraploid swards than those on the diploid swards (453 vs. 519 min 24 h−1; P = 0·012) in July 2002. Digestibility of grass snips was significantly higher on the tetraploid than the diploid swards [697 vs. 680 g digestible organic matter (DOM) kg−1 DM; P = 0·042] in September 2003 and, within diploids, was significantly higher for Glen than Belramo (696 vs. 663 g DOM kg−1 DM; P = 0·014). There were significant differences in sheath tube and leaf lengths and in the population density of tillers between and within ploidies, which might have been expected to have influenced intake characteristics, but this was not generally found under rotational grazing with cattle. In order to separate the effects of defoliation interval from those of grazing style of the different ruminant species, it is suggested that grass variety evaluations using continuously stocked cattle swards are required. [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]

Mule (Odocoileus hemionus) and white‐tailed deer (Odocoileus virginianus) are congeneric and share similar life histories, yet their distribution is segregated across much of North America. Extensive research on both species within and outside their zone of co‐occurrence has not fully explained these distribution patterns, especially the potential role of diet and foraging behavior. Therefore, we used a common garden experiment to compare diet composition, diet quality, foraging behavior, and intake of tractable mule and white‐tailed deer foraging together within the dry Douglas‐fir (Pseudotsuga menziesii)/ponderosa pine (Pinus ponderosa) forests of northeastern Washington. We sampled at 21 0.5‐ha sites from June to August 2016. We used standard bite count techniques coupled with forage biomass sampling, behavioral observations, and nutritional analyses to compare the foraging ecology of the two species. Mule and white‐tailed deer had similar activity patterns. However, mule deer took larger bites and harvested food faster than white‐tailed deer, and white‐tailed deer consumed more diverse but higher‐quality diets than mule deer. These differences resulted in mule deer acquiring ~25% more dry matter and digestible energy per day. About 90% of the diets consumed by both deer species consisted of deciduous shrubs and forbs, and they selected many of the same plant species. However, overall diet composition was 38% dissimilar, with mule deer consuming diets that were more likely to contain shrubs with higher levels of tannins and lower levels of dry matter digestibility than diets eaten by white‐tailed deer. Dietary overlap was greatest at both very low and very high forage biomass, indicating potential for modest resource competition or partitioning. Our research provides evidence that differences in diet composition of mule and white‐tailed deer do not merely reflect differences in habitat selection, but also suggest the species differ in their fundamental nutritional niches. [ABSTRACT FROM AUTHOR]