Foraging decisions: patch choice and exploitation by pigeons

Six pigeons, Columba livia, served as subjects in a foraging analogue testing the predictions, both qualitative and quantitative, of optimal foraging theory as applied to patchy environments. The operant procedure modelled the events of a foraging episode, including depletion of patch resources. The encounter rates with the rich and poor patch-types were varied, either together or separately, in each of four experiments, and the effects on patch choice behaviour and patch exploitation were noted. Most of the qualitative predictions of optimal foraging theory were supported: (1) rich patch-type opportunities were nearly always accepted; (2) as the encounter rate with the rich patch-type decreased, subjects became less selective and began to accept poor patch opportunities; and (3) the acceptance of the poor patch-type closely, though not always, approximated an all-or-none pattern. However, contrary to optimal foraging theory's prediction, variation in the encounter rate with only the poor patch-type did influence patch choice behaviour. As predicted, the extent of patch exploitation increased as environmental quality decreased; however, subjects over-exploited patches relative to the quantitative predictions of optimal foraging theory.     

Foraging decisions in a capital breeder: trade-offs between mass gain and lactation

The high energetic costs of lactation can lead to fundamental trade-offs in life-history traits, particularly in young females that reproduce before completing body growth. We assessed whether lactating female mountain goats (Oreamnos americanus) used behavioural tactics at fine spatio-temporal scales to increase energy intake to compensate for the costs of lactation. Lactating females increased bite rate and chewing rate compared with non-lactating females, but selected similar foraging sites in terms of plant quality and abundance. At peak lactation, forage intake of lactating females was >40% greater than that of non-lactating females. For females that had reached asymptotic body mass (i.e. ≥6 years old), summer mass gain of lactating females was similar to that of non-lactating females. At 4 and 5 years of age, however, daily mass gain of lactating females was about 20% lower than that of non-lactating females. We conclude that increased foraging may allow fully-grown lactating females to compensate for the energetic costs of lactation, but that there is a major trade-off between mass gain and lactation for younger females. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Foraging decisions and behavioural flexibility in trap‐building predators: a review

Foraging theory was first developed to predict the behaviour of widely‐foraging animals that actively search for prey. Although the behaviour of sit‐and‐wait predators often follows predictions derived from foraging theory, the similarity between these two distinct groups of predators is not always obvious. In this review, we compare foraging activities of trap‐building predators (mainly pit‐building antlions and web‐building spiders), a specific group of sit‐and‐wait predators that construct traps as a foraging device, with those of widely‐foraging predators. We refer to modifications of the trap characteristics as analogous to changes in foraging intensity. Our review illustrates that the responses of trap‐building and widely‐foraging predators to different internal and external factors, such as hunger level, conspecific density and predation threat are quite similar, calling for additional studies of foraging theory using trap‐building predators. In each chapter of this review, we summarize the response of trap‐building predators to a different factor, while contrasting it with the equivalent response characterizing widely‐foraging predators. We provide here evidence that the behaviour of trap‐building predators is not stereotypic or fixed as was once commonly accepted, rather it can vary greatly, depending on the individual's internal state and its interactions with external environmental factors.     

Foraging behaviour of predators in heterogeneous landscapes: the role of perceptual ability and diet breadth

Oligophagous and polyphagous predators are confronted with spatially and temporally varying distributions of prey. Their species-specific foraging strategies should be able to cope with this variability. Using an individual based model, we explore how diet breath and the spatial scale at which predators respond to prey affects their capture efficiency in four heterogeneous prey landscapes, and combinations thereof. We interpret the spatial scale of the predator's response as perceptual range, and propose giving-up density as a proxy for diet breadth. Foraging behaviour is evaluated for a total of 121 perceptual range/giving-up density combinations, with four of them reflecting the strategies adopted by real ladybeetle species. Foraging rules of oligophagous ladybeetles were generally not very effective in terms of attained predation rate when foraging in a single prey landscape, but appear to be more effective when foraging in multiple prey landscapes. This finding is compatible with the notion that oligophagous predators do not adopt a foraging strategy that is especially adapted to a specific prey landscape, but to multiple prey landscapes. Simulations further indicated that there was not a 'best' foraging rule that resulted in the highest predation rates for a range of spatial prey distributions and prey densities. The findings thus suggest that strategies of four ladybeetle species are effective in generating sufficient prey capture under a broad range of spatial distributions, rather than maximum capture under a narrower set of distributions.     

Foraging decisions with conservation consequences: Interaction between beavers and invasive tree species

1. Herbivore species can either hinder or accelerate the invasion of woody species through selective utilization. Therefore, an exploration of foraging decisions can contribute to the understanding and forecasting of woody plant invasions. Despite the large distribution range and rapidly growing abundance of beaver species across the Northern Hemisphere, only a few studies focus on the interaction between beavers and invasive woody plants.
2. We collected data on the woody plant supply and utilization at 20 study sites in Hungary, at two fixed distances from the water. The following parameters were registered: taxon, trunk diameter, type of utilization, and carving depth. Altogether 5401 units (trunks and thick branches) were identified individually. We developed a statistical protocol that uses a dual approach, combining whole‐database and transect‐level analyses to examine foraging strategy.
3. Taxon, diameter, and distance from water all had a significant effect on foraging decisions. The order of preference for the four most abundant taxa was Populus spp. (softwood), Salix spp. (softwood), Fraxinus pennsylvanica (invasive hardwood), and Acer negundo (invasive hardwood). The diameter influenced the type of utilization, as units with greater diameter were rather carved or debarked than felled. According to the central‐place foraging strategy, the intensity of the foraging decreased with the distance from the water, while both the taxon and diameter selectivity increased. This suggests stronger modification of the woody vegetation directly along the waterbank, together with a weaker impact further from the water.
4. In contrast to invasive trees, for which utilization occurred almost exclusively in the smallest diameter class, even the largest softwood trees were utilized by means of carving and debarking. This may lead to the gradual loss of softwoods or the transformation of them into shrubby forms. After the return of the beaver, mature stages of softwood stands and thus the structural heterogeneity of floodplain woody vegetation could be supported by the maintenance of sufficiently large active floodplains.
5. The beaver accelerates the shift of the canopy layer''s species composition toward invasive hardwood species, supporting the enemy release hypothesis. However, the long‐term impact will also depend on how plants respond to different types of utilization and on their ability to regenerate, which are still unexplored issues in this environment. Our results should be integrated with knowledge about factors influencing the competitiveness of the studied native and invasive woody species to support floodplain conservation and reconstruction.

     

Foraging trip decisions by the streaked shearwater Calonectris leucomelas depend on both parental and chick state

Parents of albatross and shearwater species employ a dual foraging strategy, feeding their chicks quickly in repeated short trips and then restoring their own fuel reserves during longer trips. A decline in parental body condition is believed to trigger longer trips, but chick body condition and age may also play a role. To investigate these factors in the little-studied streaked shearwater Calonectris leucomelas, we monitored the nest attendance of 17 pairs on Mikura Island in 2005 using an automated identification system. We also monitored body mass changes and meal masses of 5 of the 17 pairs using an automated weighing system. Although the birds did not show a clear dual foraging pattern, trip duration varied widely from 1 to 15 days. On average, the birds fed chicks 67.6 g during nighttime meals at 2.74-day intervals. Since meal mass did not depend on trip duration, feeding efficiency (meal mass delivered per unit trip duration) decreased as trip duration increased. Parents accumulated more energy reserves when they took longer trips. Parents appeared likely to initiate longer trips when their body condition declined or chick body condition recovered.     

Foraging across a variable landscape: behavioral decisions made by woodland caribou at multiple spatial scales

We examined the foraging behavior of woodland caribou (Rangifer tarandus caribou) relative to the spatial and temporal heterogeneity of their environment. We assessed (1) whether caribou altered their behavior over time while making trade-offs between forage abundance and accessibility; and (2) whether foraging decisions were consistent across spatial scales (i.e., as scale increased, similar decision criteria were used at each scale). We discuss whether caribou adjusted their behavior to take advantage of changing forage availability through time and space. At the scale of the feeding site (as revealed by discriminant function analyses), caribou in both forested and alpine (above tree-line) environments selected sites where the biomass of particular lichen species was greatest and snow the least deep. Caribou did not select those species with the highest nutritional value (i.e., digestible protein and energy) in either area. Where snow depth, density, and hardness limited access to terrestrial lichens in the forest, caribou foraged instead at those trees with the greatest amount of arboreal lichen. Selection of lichen species and the influence of snow differed across time, indicating that in this system the abundance or accessibility of forage temporally influenced foraging behavior. A path analysis of forest data and multiple regression analysis of alpine data were used to test the hypothesis that variables important at the scale of the feeding site explained foraging effort at the scale of the patch. For forest patches, our hypothesized model reliably explained foraging effort, but not all variables that were statistically important at the scale of the feeding site were significant predictors at the scale of the patch. For alpine patches, our hypothesized model did not explain a statistically significant portion of the variation in the number of feeding sites within the patch, and none of the individual variables from the feeding site remained statistically significant at the patch scale. The incongruity between those variables important at the scale of the feeding site and those important at the patch showed that spatial scale affects the foraging decisions of woodland caribou. At the scale of the landscape, there was a trade-off between forage abundance and accessibility. Relative to the alpine environment, caribou in the forest foraged at feeding sites and patches with greater amounts of less variably distributed lichen, but deeper less variable snow depths. Considering the behavioral plasticity of woodland caribou, there may be no distinct advantage to foraging in one landscape over the other.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Foraging decisions underlying restricted space use: effects of fire and forage maturation on large herbivore nutrient uptake

Recent models suggest that herbivores optimize nutrient intake by selecting patches of low to intermediate vegetation biomass. We assessed the application of this hypothesis to plains bison (Bison bison) in an experimental grassland managed with fire by estimating daily rates of nutrient intake in relation to grass biomass and by measuring patch selection in experimental watersheds in which grass biomass was manipulated by prescribed burning. Digestible crude protein content of grass declined linearly with increasing biomass, and the mean digestible protein content relative to grass biomass was greater in burned watersheds than watersheds not burned that spring (intercept; F_1,251 = 50.57, P < 0.0001). Linking these values to published functional response parameters, ad libitum protein intake, and protein expenditure parameters, Fryxell's (Am. Nat., 1991, 138 , 478) model predicted that the daily rate of protein intake should be highest when bison feed in grasslands with 400–600 kg/ha. In burned grassland sites, where bison spend most of their time, availability of grass biomass ranged between 40 and 3650 kg/ha, bison selected foraging areas of roughly 690 kg/ha, close to the value for protein intake maximization predicted by the model. The seasonal net protein intake predicted for large grazers in this study suggest feeding in burned grassland can be more beneficial for nutrient uptake relative to unburned grassland as long as grass regrowth is possible. Foraging site selection for grass patches of low to intermediate biomass help explain patterns of uniform space use reported previously for large grazers in fire‐prone systems.     

Quality versus quantity: Foraging decisions in the honeybee (Apis mellifera scutellata) feeding on wildflower nectar and fruit juice

Foraging animals must often decide among resources which vary in quality and quantity. Nectar is a resource that exists along a continuum of quality in terms of sugar concentration and is the primary energy source for bees. Alternative sugar sources exist, including fruit juice, which generally has lower energetic value than nectar. We observed many honeybees (Apis mellifera scutellata) foraging on juice from fallen guava (Psidium guajava) fruit near others foraging on nectar. To investigate whether fruit and nectar offered contrasting benefits of quality and quantity, we compared honeybee foraging performance on P. guajava fruit versus two wildflowers growing within 50 m, Richardia brasiliensis and Tridax procumbens. Bees gained weight significantly faster on fruit, 2.72 mg/min, than on either flower (0.17 and 0.12 mg/min, respectively). However, the crop sugar concentration of fruit foragers was significantly lower than for either flower (12.4% vs. 37.0% and 22.7%, respectively). Fruit foragers also spent the most time handling and the least time flying, suggesting that fruit juice was energetically inexpensive to collect. We interpret honeybee foraging decisions in the context of existing foraging models and consider how nest‐patch distance may be a key factor for central place foragers choosing between resources of contrasting quality and quantity. We also discuss how dilute solutions, such as fruit juice, can help maintain colony sugar–water balance. These results show the benefits of feeding on resources with contrasting quality and quantity and that even low‐quality resources have value.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.     

Foraging of multimammate mice, Mastomys natalensis, under different predation pressure: cover, patch-dependent decisions and density-dependent GUDs

Patch use under predation risk often results in a change of feeding behaviour in the prey animals. However, such changes only appear if the animals are able to assess under which predation pressure they live. We investigated patch use of Mastomys natalensis under different conditions of avian predation pressure.
In replicated maize field plots in Morogoro, Tanzania, avian predators were allowed under natural conditions (control), attracted with perches and nest boxes or kept out with nets. During four one‐week periods in late 1999, we measured rodent feeding decisions with the giving‐up density (GUD) method. Trays with known amounts of millet seeds in sand were placed in pairs, one of them under a cover, the other one in the open. M. natalensis mice were expected to give up sooner in the open trays than in those with cover. We hypothesised that M. natalensis mice could assess the ambient predation pressure leading to larger difference in GUD between covered and non‐covered trays in the plots where predators were attracted. We also made video recordings of the rodent activity at a pair of trays in each treatment. The GUD‐values were significantly lower for the covered trays but predation pressure did not affect this difference. The video observations showed that in the control and netted plots the animals visited trays equally frequently regardless of the cover, while the visits in the predator‐attracted plots occurred significantly more often in the covered trays. We conclude that M. natalensis can assess the ambient predation pressure and adapt its behaviour at a feeding patch. However, the variation in predation pressure in our experiment was not obvious from the GUD. Moreover, we found a strong relation between rodent density and GUD, which may mask variations in perceived predation pressure. Similar GUD values may be reached in different ways and we present models to investigate whether animals’ decision to forage at a food patch is only affected by the seed density at that patch, not by that at a neighbour patch.     

Context-Sensitivity in Crab Foraging Studies

Initial work examining crab foraging from optimality premisesexplored fundamental foraging scope (the capacity of animalsto graze), often within an abstract experimental context. Anemergent theme involves explicit consideration of biologicalconstraints (e.g., predation risk) and environmental factors(e.g., substrate type) which, by modulating grazing capacity,determine the realized foraging pattern seen in nature. We briefly review two studies that illustrate the contemporaryfocus on realized foraging pattern. One defines field grow-outtechniques for bivalves (raised in mariculture and resourceenhancement programs) that minimize losses from predators suchas the portunid crab, Ovalipes ocellatus. By focusing on marginalregions of predator: prey interaction (in this case, foragingon low densities of clams planted in heterogenous substrates),the study yielded novel insight into limits on portunid crabforaging on infaunal clams. The second study analyses the foragingperformance of a deposit feeding ocypodid crab, Scopimera inflata,over different temporal and spatial scales. We demonstrate thatwhereas S. inflata performs sub-optimally at micro-scales (secondsto minutes; mm to cm), the crabs nearly optimize performanceover macro-scales (days to years; cm to m). Continued research on the fundamental foraging scope of crabsis warranted, but should be explicitly referenced to naturalhistorical context and, in particular, to the forager's ontogeneticstage. We also perceive a need for collaborative research incorporatingbehavioral, physiological, and biochemical facets in an integratedexperimental setting. This would ensure that context does notbias information, as can occur in studies that emphasise a particularresearch perspective, methodological approach, or scale at whichforaging is analysed.     

Foraging behavior in stochastic environments

How do temporally stochastic environments affect risk sensitivity in foraging behavior? We build a simple model of foraging under predation risks in stochastic environments, where the environments change over generations. We analyze the effects of stochastic environments on risk sensitivity of foraging animals by means of the difference between the geometric mean fitness and the arithmetic mean fitness. We assume that foraging is associated with predation risks whereas resting in the nest is safe because it is free of predators. In each generation, two different environments with given food amounts and predation risks occur with a certain probability. The geometric mean optimum is independent of food amounts. In most cases of stochastic environments, risk-averse tendency is increased, but in some limited conditions, more risk-prone behavior is favored. Specifically, risk-prone tendency is increased when the variation in food amount increases. Our results imply that the optimal behavior depends on the probability distribution of environmental effects under all selection regimes.     

Foraging innovation in the guppy

When novel behaviour patterns spread through animal populations, typically one animal will initiate the diffusion. It is not known whether such 'innovators' are particularly creative individuals, individuals exposed to the appropriate environmental contingencies, or individuals in a particular motivational state. We describe three experiments that investigated the factors influencing foraging innovation in the guppy Poecilia reticulata. We exposed small laboratory populations of fish to novel foraging tasks, which involved exploration and problem solving to locate a novel food source. Experiments 1 and 2 found that (1) females were more likely to innovate than males, (2) food-deprived fish were more likely to innovate than nonfood-deprived subjects, and (3) smaller fish were more likely to innovate than larger fish. We suggest that the sex difference may reflect parental investment asymmetries in males and females. Experiment 3 found that past innovators were more likely to innovate than past noninnovators. Collectively, the results suggest that differences in foraging innovation in guppies are best accounted for by differences in motivational state, but, in addition, guppies may vary in their predisposition to innovate. Copyright 1999 The Association for the Study of Animal Behaviour.     

Central Place Foraging in Nonpatchy Habitats

This paper deal with a model of optimal foraging in a habitat with arbitrary food distribution. It takes into account an arbitrary risk cost related to the distance to the animal's nest. Food acquisition and risk cost are accounted for in common units of fitness. The resulting problem is solved in the context of Calculus of Variations. The optimal duration of absence from the nest and the optimal spatial allocation of foraging time are obtained: the optimal strategy leads to separate the habitat into a region to exploit and a region to ignore. The definition of these two distinct regions depends on the relative importance of risk and food availability. With realistic risk costs, the resulting strategy indicates a highly selective behaviour when far from the nest, as observed in field studies. The model is also extended to take account of the need of returning to the nest to guard it or to feed the young.     

At-Sea Associations in Foraging Little Penguins

Prey distribution, patch size, and the presence of conspecifics are important factors influencing a predator’s feeding tactics, including the decision to feed individually or socially. Little is known about group behaviour in seabirds as they spend most of their lives in the marine environment where it is difficult to observe their foraging activities. In this study, we report on at-sea foraging associations of little penguins (Eudyptula minor) during the breeding season. Individuals could be categorised as (1) not associating; (2) associating when departing from and/or returning to the colony; or (3) at sea when travelling, diving or performing synchronised dives. Out of 84 separate foraging tracks, 58 (69.0%) involved associations with conspecifics. Furthermore, in a total of 39 (46.4%), individuals were found to dive during association and in 32 (38.1%), individuals were found to exhibit synchronous diving. These behaviours suggest little penguins forage in groups, could synchronise their underwater movements and potentially cooperate to concentrate their small schooling prey.     

Foraging strategies of caterpillars

Summary An analysis of the foraging behaviors of several species of palatable and unpalatable lepidopterous larvae indicates that palatable caterpillars partition their time between feeding and behaviors that could be related to escape visually oriented predators. Depending on the species, palatable caterpillars do all or several of the following: 1) restrict themselves to the underside of leaves at all times, 2) restrict foraging to night-time, 3) commute to and from their feeding area on leaves, 4) move from the unfinished leaf to a distant leaf after a feeding bout, thus removing themselves from the evidence of their eating, 5) snip off partially-eaten leaves after feeding on them. The less palatable, or unpalatable, caterpillars do not snip off partially-eaten leaves, feed from leaves leaving tattered edges, and are often exposed resting and feeding on the leaf surfaces in direct sunshine. I conclude that some caterpillar foraging behaviors may have evolved under the selective pressure of visually-oriented predators that use leaf-damage as a cue in their searching behavior.