Foraging-area size and food density: Some predictive models

It has been observed that when food increases in density along an environmental gradient, the size of foraging areas used by desert ant colonies decreases. Factors that could cause this inverse relationship are explored. Four models of ant foraging are developed and presented in the form of equations for calculating net foraging energy as a function of size of the foraging area. These are used to predict the optimal sizes of foraging areas under conditions of different food densities. Only one of the models predicts an inverse relationship between density of food and size of foraging area when food is the limiting factor in colony reproduction. In this model, the foraging areas of adjacent colonies are assumed to be overlapping and the number of foragers assigned to each square meter of the foraging area is constant, regardless of food density or distance from the nest entrance. Tests for distinguishing among the four models, as well as for determining whether colonies are or are not food limited, are discussed.     

Experimental functional response and inter-individual variation in foraging rate of teal (Anas crecca)

The functional response, i.e. the change in per capita food intake rate per time unit with changed food availability, is a widely used tool for understanding the ecology and behaviour of animals. However, waterfowl remain poorly explored in this context. In an aviary experiment we derived a functional response curve for teal (Anas crecca) foraging on rice (Oryza sativa) seeds. We found a linear relationship between intake rate and seed density, as expected for a filter-feeder. At high seed densities we found a threshold, above which intake rate still increased linearly but with a lower slope, possibly reflecting a switch from filter-feeding to a scooping foraging mode. The present study shows that food intake rate in teal is linearly related to food availability within the range of naturally occurring seed densities, a finding with major implications for management and conservation of wetland habitats.     

Opportunity costs influence food selection and giving‐up density of dabbling ducks

The interaction of animals with their food can yield insights into habitat characteristics, such as perceived predation risk and relative quality. We deployed experimental foraging patches in wetlands used by migrating dabbling ducks Anas spp. in the central Illinois River Valley to estimate variation in seed removal and giving‐up density (GUD; i.e. density of food remaining in patches following abandonment) with respect to seed density, seed size, seed depth in the substrate, substrate firmness, perceived predation risk, and an energetic profitability threshold (i.e. critical food density). Seed depth and the density of naturally‐occurring seeds outside of experimental plots affected seed removal and GUD in experimental patches more than perceived predation risk, seed density, seed size or substrate firmness. The greatest seed removal and lowest GUDs in experimental patches occurred when food resources in alternative foraging locations outside of plots (i.e. opportunity costs) appeared to be near or below a critical food density (i.e. 119–181 kg ha^–1). Giving‐up densities varied substantially from a critical food density across a range of food densities in alternative foraging locations suggesting that fixed GUDs should not be used as surrogates for critical food densities in energetic carrying capacity models. Foraging and resting rates in and near experimental foraging patches did not reflect patterns of seed removal and were poor predictors of GUD and foraging habitat quality. Our results demonstrated the usefulness of GUDs as indicators of habitat quality for subsurface, benthic foragers relative to other available foraging patches and suggested that food may be limited for dabbling ducks during spring migration in some years in the midwestern USA.     

Sex-Related Differences in the Trade-Off between Foraging and Vigilance in a Granivorous Forager

The relationship between intake rate and food density can provide the foundation for models that predict the spatiotemporal distribution of organisms across a range of resource densities. The functional response, describing the relationship between resource density and intake rate is often interpreted mechanistically as the relationships between times spend searching and handling. While several functional response models incorporate anti-predator vigilance (defined here as an interruption of feeding or some other activity to visually scan the environment, directed mainly towards detecting potential predators), the impacts of environmental factors influencing directly anti-predator vigilance remains unclear. We examined the combined effects of different scenarios of predation risk and food density on time allocation between foraging and anti-predator vigilance in a granivorous species. We experimentally exposed Skylarks to various cover heights and seed densities, and measured individual time budget and pecking and intake rates. Our results indicated that time devoted to different activities varied as a function of both seed density and cover height. Foraging time increased with seed density for all cover heights. Conversely, an increased cover height resulted in a decreased foraging time. Contrary to males, the decreased proportion of time spent foraging did not translate into a foraging disadvantage for females. When vegetation height was higher, females maintained similar pecking and intake rates compared to intermediate levels, while males consistently decreased their energy gain. This difference in anti-predator responses suggests a sexually mediated strategy in the food-safety trade-off: when resource density is high a females would adopt a camouflage strategy while an escape strategy would be adopted by males. In other words, males would leave risky-areas, whereas females would stay when resource density is high. Our results suggest that increased predation risk might generate sexually mediated behavioural responses that functional response models should perhaps better consider in the future.     

Experimental studies on the foraging behavior of the sand fiddler crab Uca pugilator (Bosc, 1802)

The purpose of this study was to test the hypothesis that foraging sand fiddler crabs. Uca pugilator (Bosc), move through the habitat in response to low substratum food levels even though these movements may take the crabs considerable distances from the safety of the burrow area. Chl a and ATP concentrations were used as measures of food density in foraged and unforaged substratum. Field and laboratory feeding experiments showed that crab foraging intensity in a habitat patch was directly correlated with food density in the patch either in the presence or absence of alternative food patches. Other experiments showed that sand fiddlers can respond to differences in food level on a scale of millimeters and do this by probing the substratum with minor chelae. Food levels in aggregations of non-ingested particles harvested by sand fiddlers, feeding pellets, correspond to low foraging intensities predicted from foraging experiments and crabs exhibit low foraging intensities on substratum patches derived from feeding pellets. Substratum food levels in two distinct areas corresponded to high predicted foraging intensities and there was no consistent trend in the level of food in the burrow vs. the nonburrow microhabitats. These results suggest that the movements of foraging sand fiddlers are to some extent controlled by the reduction in substratum food levels due to feeding during a single foraging episode. Sand fiddlers can extract over 70% of the food from harvested substratum over a broad range of substratum food densities but harvest only 42% of the available substratum.     

Anthropogenically-Mediated Density Dependence in a Declining Farmland Bird

Land management intrinsically influences the distribution of animals and can consequently alter the potential for density-dependent processes to act within populations. For declining species, high densities of breeding territories are typically considered to represent productive populations. However, as density-dependent effects of food limitation or predator pressure may occur (especially when species are dependent upon separate nesting and foraging habitats), high territory density may limit per-capita productivity. Here, we use a declining but widespread European farmland bird, the yellowhammer Emberiza citrinella L., as a model system to test whether higher territory densities result in lower fledging success, parental provisioning rates or nestling growth rates compared to lower densities. Organic landscapes held higher territory densities, but nests on organic farms fledged fewer nestlings, translating to a 5 times higher rate of population shrinkage on organic farms compared to conventional. In addition, when parental provisioning behaviour was not restricted by predation risk (i.e., at times of low corvid activity), nestling provisioning rates were higher at lower territory densities, resulting in a much greater increase in nestling mass in low density areas, suggesting that food limitation occurred at high densities. These findings in turn suggest an ecological trap, whereby preferred nesting habitat does not provide sufficient food for rearing nestlings at high population density, creating a population sink. Habitat management for farmland birds should focus not simply on creating a high nesting density, but also on ensuring heterogeneous habitats to provide food resources in close proximity to nesting birds, even if this occurs through potentially restricting overall nest density but increasing population-level breeding success.     

Population density affects foraging behavior of male Black-throated Blue Warblers during the breeding season

ABSTRACT.   Foraging behavior often reflects food availability, a resource that may increasingly limit breeding birds as intraspecific crowding increases. Measuring foraging behavior, therefore, provides a way to investigate effects of population density on food limitation, an important link in understanding how crowding functions to regulate populations. We quantified three components of foraging behavior (prey attack rate, foraging speed, and relative use of morphologically constrained attack maneuvers) for male Black-throated Blue Warblers ( Dendroica caerulescens ) breeding under experimentally manipulated density conditions. Building on the previous work showing the density of conspecific neighbors affects territory size, reproductive success, and the time budgets of males ( Sillett et al. 2004 , Ecology 85: 2467–2477), we further show that density affects male foraging strategies. Although not differing in attack rate or foraging speed, male Black-throated Blue Warblers on territories with reduced neighbor densities used energetically expensive aerial attack maneuvers significantly less frequently than males in control (high-density) territories during both the incubation period and when provisioning nestlings and fledglings. We conclude that males altered their foraging behavior to compensate for density-related reductions in time available for foraging and that population density may constrain the time available for foraging.     

Taxonomic status of the Forest Buzzard Buteo oreophilus trizonatus

The foraging efficiencies of four sympatric southern African seed-eating birds, namely Bronze Mannikin Spermestes cucullatus, Cape Sparrow Passer melanurus, Southern Red Bishop Euplectes orix and Thick-billed Weaver Amblyospiza albifrons, and a domesticated species, the Bengalese Finch Lonchura domestica, were measured and compared using giving-up densities (the amount of food remaining following patch exploitation) in experimental food patches. Foraging efficiency was quantified using giving-up densities by offering individual birds equal foraging opportunities. A low giving-up density displays the ability of a forager to profitably harvest food at low resource densities and to gainfully exploit the foraging opportunities overlooked by a less efficient forager. Ten individuals of each of the five species were allowed to forage on six different seed types. Thick-billed Weavers had significantly lower giving-up densities for all seed types except the smallest, namely red manna. Bronze Mannikins showed the converse trend, foraging most efficiently on the smallest seeds. The results of the present study revealed that Thick-billed Weavers were the most efficient foragers (i.e. had the lowest giving-up densities on seeds in feeding trays).     

Optimal foraging: The responses of Peromyscus leucopus to experimental changes in processing time and hunger

Summary Food density, degree of hunger prior to foraging, and food processing difficulty were varied in order to determine their effects on the diet diversity of captive Peromyscus leucopus foraging for buried seeds. No relationship was found between diet diversity and food density. However, diet diversity exhibited a significantly positive relationship with hunger, and a significantly negative relationship with seed processing time. These results strongly support optimal foraging theory.     

Modelling the energy budget of a colonial bird of prey, the Ruppell's griffon vulture, and consequences for its breeding ecology

Colonial nesting is rare in birds of prey. In this study we develop further Pennycuick's (1979 ) model of energy balance to consider the implications of colonial nesting for the breeding ecology of Ruppell's griffon vultures. To achieve a realistic foraging range, and remain in energy balance, the birds need to do more than fill their crop once on each foraging trip. They must remain in the feeding area and digest some of this food and refill the crop to obtain sufficient energy to pay for the flight costs and have sufficient energy to satisfy their own requirements and that of the chick. Given the known distances that the birds have to travel to forage, it would be impossible for them to rear more than one chick. The low growth rate of griffon vulture chicks may be an adaptation to the low rate at which energy can be delivered by the parents. The optimal time for a bird to be away from the nest changes with the distance they have to travel. Assuming that one parent remains on the nest at all times to guard the chick, it is optimal for both parents to take turns to forage on the same day if the distance to a feeding area is under 150 km, but to switch to each parent being away for a whole day when the distance is greater than this. Soaring flight is essential for such a scavenger, because of the low energy expenditure. If a vulture relied on the more energetically demanding flapping flight its maximum foraging range would be under 40 km. Griffon vultures are known to be able to depress their basal metabolic rate, and this has major implications for their foraging range, which then becomes constrained by the flight speed rather than by the amount of food they need to obtain. Griffon vultures minimize energy expenditure on all activities, because even small increases in their energy demands have a large impact on the foraging range that the bird can use.     

Aggressive and Foraging Behavioral Interactions Among Ruffe

The ruffe, Gymnocephalus cernuus, is a nonindigenous percid in the Great Lakes. Ruffe are aggressive benthivores and forage over soft substrates. Laboratory studies in pools (100cm diameter, 15cm water depth) were conducted to determine whether fish density (low=2, medium=4, high=6 ruffe per pool) changed foraging and aggressive behaviors with a limited food supply of chironomid larvae. All fish densities demonstrated a hierarchy based on aggressive interactions, but ruffe were most aggressive at low and high fish densities. Time spent in foraging was lowest at the low fish density. The best forager at the low fish density was the most aggressive individual, but the second most aggressive fish at the medium and high fish density was the best forager and also the one chased most frequently. A medium fish density offered the best energetic benefits to ruffe by providing the lowest ratio of time spent in aggression to that spent foraging. Based on our results, ruffe should grow best at an intermediate density. With high ruffe densities, we would also expect disparity in size as the more aggressive fish are able to garner a disproportionate amount of the resources. Alternatively, as the Great Lakes are a fairly open system, ruffe could migrate out of one area to colonize another as populations exceed optimal densities.     

Determination of Foraging Thresholds and Effects of Application on Energetic Carrying Capacity for Waterfowl

Energetic carrying capacity of habitats for wildlife is a fundamental concept used to better understand population ecology and prioritize conservation efforts. However, carrying capacity can be difficult to estimate accurately and simplified models often depend on many assumptions and few estimated parameters. We demonstrate the complex nature of parameterizing energetic carrying capacity models and use an experimental approach to describe a necessary parameter, a foraging threshold (i.e., density of food at which animals no longer can efficiently forage and acquire energy), for a guild of migratory birds. We created foraging patches with different fixed prey densities and monitored the numerical and behavioral responses of waterfowl (Anatidae) and depletion of foods during winter. Dabbling ducks (Anatini) fed extensively in plots and all initial densities of supplemented seed were rapidly reduced to 10 kg/ha and other natural seeds and tubers combined to 170 kg/ha, despite different starting densities. However, ducks did not abandon or stop foraging in wetlands when seed reduction ceased approximately two weeks into the winter-long experiment nor did they consistently distribute according to ideal-free predictions during this period. Dabbling duck use of experimental plots was not related to initial seed density, and residual seed and tuber densities varied among plant taxa and wetlands but not plots. Herein, we reached several conclusions: 1) foraging effort and numerical responses of dabbling ducks in winter were likely influenced by factors other than total food densities (e.g., predation risk, opportunity costs, forager condition), 2) foraging thresholds may vary among foraging locations, and 3) the numerical response of dabbling ducks may be an inconsistent predictor of habitat quality relative to seed and tuber density. We describe implications on habitat conservation objectives of using different foraging thresholds in energetic carrying capacity models and suggest scientists reevaluate assumptions of these models used to guide habitat conservation.     

Foraging arena size and structural complexity affect the dynamics of food distribution in ant colonies

Food acquisition by ant colonies is a complex process that starts with acquiring food at the source (i.e., foraging) and culminates with food exchange in or around the nest (i.e., feeding). While ant foraging behavior is relatively well understood, the process of food distribution has received little attention, largely because of the lack of methodology that allows for accurate monitoring of food flow. In this study, we used the odorous house ant, Tapinoma sessile (Say) to investigate the effect of foraging arena size and structural complexity on the rate and the extent of spread of liquid carbohydrate food (sucrose solution) throughout a colony. To track the movement of food, we used protein marking and double-antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Variation in arena size, in conjunction with different colony sizes, allowed us to test the effect of different worker densities on food distribution. Results demonstrate that both arena size and colony size have a significant effect on the spread of the food and the number of workers receiving food decreased as arena size and colony size increased. When colony size was kept constant and arena size increased, the percentage of workers testing positive for the marker decreased, most likely because of fewer trophallactic interactions resulting from lower worker density. When arena size was kept constant and colony size increased, the percentage of workers testing positive decreased. Nonrandom (clustered) worker dispersion and a limited supply of food may have contributed to this result. Overall, results suggest that food distribution is more complete is smaller colonies regardless of the size of the foraging arena and that colony size, rather than worker density, is the primary factor affecting food distribution. The structural complexity of foraging arenas ranged from simple, two-dimensional space (empty arenas) to complex, three-dimensional space (arenas filled with mulch). The structural complexity of foraging arenas had a significant effect on food distribution and the presence of substrate significantly inhibited the spread of food. Structural complexity of foraging arenas and the resulting worker activity patterns might exert considerable influence on socioecological processes in ants and should be considered in laboratory assays.     

Biomechanics and foraging profitability: an approach to assessing trophic needs and impacts of diving ducks

A biomechanical model of underwater locomotion is described, and data required by the model presented for 3 species of diving duck (Aythya spp.). Based on field observations of behavior and foods consumed, the model is used to estimate energy costs of foraging and minimum food intake rates of canvasbacks (Aythya valisineria) in two habitats in North Carolina. Increased water depth from 0.5 m in Lake Mattamuskeet to 1.5 m in Pamlico Sound increased the net cost of time spent foraging at the bottom by 43%. Biomechanical calculations are combined with data on intake rates at different food densities (Takekawa, 1987) to determine minimum food densities for profitable foraging in Lake Mattamuskeet. Field observations of behavior are used to adjust minimum intake per dive for unsuccessful dives spent locating food patches. Density and dispersion of plant tuber foods in Lake Mattamuskeet, before and after the fall staging period, suggest that the fraction of habitat with tuber densities above a profitability threshold is more critical to canvasbacks than average tuber density. Such factors are important in relating bird energy requirements and benthic sampling data to carrying capacity and total area of usable habitat. The proportion of foods that can be fed upon profitably also determines the fraction of food organisms subject to depletion as components of trophic pathways.     

Competition between birds and mammals: A comparison of giving-up densities between crested larks and gerbils

We combined the concept of mechanisms of co-existence with the approach of giving-up densities to study inter-taxon competition between seed-eating birds and mammals. We measured feeding behaviour in food patches to define and study the guild of seed-eating vertebrates occupying sandy habitats at Bir Asluj, Negev Desert, Israel. Despite a large number of putatively granivorous rodents and birds at the site, two gerbil species (Allenbys gerbil, Gerbillus allenbyi, and the greater Egyptian gerbil, G. pyramidum) dominated nocturnal foraging, and a single bird species (crested lark, Galerida cristata) contributed all of the daytime foraging. We used giving-up densities to quantify foraging behaviour and foraging efficiencies. A low giving-up density demonstrates the ability of a forager to profitably harvest food at low abundances and to profitably utilize the foraging opportunities left behind by the less efficient forager. Gerbils had lower giving-up densities in the bush than open microhabitat, and lower giving-up densities in the semi-stabilized than stabilized sand habitats. Crested larks showed the opposite: lower giving-up densities in the open than bush, and on the stabilized than semi-stabilized sand habitats. Despite these patterns, gerbils had substantially lower giving-up densities than crested larks in both microhabitats, all sand habitats, and during each month. Several mechanisms may permit the crested lark to co-exist with the gerbils. Larks may be cream skimmers on the high spatial and temporal variability in seed abundances. Larks may rely on insects, fruit or smaller seeds. Or, larks may rely on adjacent rocky habitats.     

Predator foraging success and habitat complexity: quantitative test of the threshold hypothesis

Summary Numerous studies have demonstrated a negative relationship between increasing habitat complexity and predator foraging success. Results from many of these studies suggest a non-linear relationship, and it has been hypothesised that some threshold level of complexity is required before foraging success is reduced significantly. We examined this hypothesis using largemouth bass (Micropterus salmoides) foraging on juvenile bluegill sunfish (Lepomis macrochirus) in various densities of artificial vegetation. Largemouth foraging success differed significantly among the densities of vegetation tested. Regression analysis revealed a non-linear relationship between increasing plant stem density and predator foraging success. Logistic analysis demonstrated a significant fit of our data to a logistic model, from which was calculated the threshold level of plant stem desity necessary to reduce predator foraging success. Studies with various prey species have shown selection by prey for more complex habitats as a refuge from predation. In this stydy, we also examined the effects of increasing habitat complexity (i.e. plant stem density) on choice of habitat by juvenile bluegills while avoiding predation. Plant stem density significantly effected choice of habitat as a refuge. The relationship between increasing habitat complexity and prey choice of habitat was found to be positive and non-linear. As with predator foraging success, logistic analysis demonstrated a significant fit of our data to a logistic model. Using this model we calculated the threshold level of habitat complexity required before prey select a habitat as a refuge. This density of vegetation proved to be considerably higher than that necessary to significantly reduce predator foraging success, indicating that bluegill select habitats safe from predation.Implications of these results and various factors which may affect the relationships described are discussed.     

Density-dependent and frequency-dependent selection by bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

The behaviour of bumblebee workers foraging on arrays of artificial flowers of two colour morphs was observed. Experiments were conducted on arrays of varying morph frequencies and at three different total flower densities. Bumblebees consistently showed a preference for the commonest colour morph, and this behaviour was not significantly affected by changing density. In contrast, frequency-independent preferences changed significantly with density. At low densities, there was a strong bias towards the more conspicuous colour, whereas at higher densities there was no overall colour bias. Flight distances between flowers decreased significantly at high density. Bumblebees also visited flowers of similar colours sequentially, but this behaviour was not density-dependent. It is suggested that as densities increase, there is an increased probability that bumblebees detect yellow flowers, which were probably less conspicuous compared with blue flowers, and that this might be caused by changes in flight speed with flight distance. Where there is a positive relationship between pollinator visitation and the relative fitness of a floral morph, the observed behaviour would induce positive frequency-dependent selection on a plant population with two corolla colour morphs on which the bumblebees were foraging, which would result in stabilizing selection for a single corolla colour, irrespective of density. There was no indication that rare colour morphs would be preferred at high density. The probability of different corolla colour morphs going to fixation would, however, be affected by density.     

Investigations of density interactions among breeding birds in ponderosa pine forests: correlative and experimental evidence

Summary We assessed the importance of interspecific competition among insectivorous birds breeding in northcentral Arizona's ponderosa pine forests. We examined density interactions among species using two analytic approaches; correlative and experimental. The correlative approach examined patterns of change in breeding densities over four years at the community level and within two foraging guilds; picker-gleaners and aerial feeders. The relationships between morphological and behavioral similarity with pairwise density interactions were also assessed. Our experimental approach involved placement of nest boxes on two treatment plots to increase breeding densities of secondary cavity nesting birds that were in foraging guilds with open nesting insectivores.We found little evidence of interspecific competition. Patterns of density fluctuations indicated large positive covariances among species at both the community level and within guilds. Pairwise density interactions were independent of morphological or behavioral similarity. Nest boxes significantly increased breeding densities of the secondary cavity nesters. However, these increases did not induce reprocal density changes in the open nesting species. Interspecific competition for food during the breeding season appears to be unimportant in ponderosa pine bird communities.     

Testing mechanisms of coexistence among two species of frugivorous primates

1. We examined mechanisms of coexistence between two congeneric species of frugivorous primates, the blue monkey (Cercopithecus mitis) and the red-tailed monkey (C. ascanius). 2. We used giving-up densities (the amount of food which animals leave in a patch) in fruit trees to measure foraging efficiency and to evaluate possible mechanisms of coexistence. Animals with higher giving-up densities are less likely to persist in the company of those with lower giving-up densities because the former are not able to exploit food patches used previously by the latter. We climbed trees to estimate giving-up densities by counting the fruit which primates left behind. 3. We tested five possible mechanisms of coexistence. Three mechanisms proposed that each frugivorous species has a lower giving-up density than the other in at least one of the following: (1) different tree species, (2) within-tree foraging zones or (3) seasons. The fourth mechanism predicted that the socially dominant species exploits resources first and that the subordinate species has lower giving-up densities. The final mechanism predicted that one species would find resources more quickly than the other, which would in turn have a lower giving-up density. 4. Four of the five mechanisms received no support from our data. Only a trade-off between interspecific dominance and giving-up densities was supported. 5. We discuss the generality of our results and possible interactions with other factors.     

Combined effect of hemipteran control and liquid bait on Argentine ant populations

The invasive Argentine ant, Linepithema humile (Mayr), has become a worldwide problem capable of inflicting significant ecological and economic injury on urban, agricultural, and natural environments. The mobility of this pest ant has long been noted, rapidly moving nests to new food resources and then away as resources are depleted. This ant, like many pest ant species, has a special affinity for honeydew excreted by phloem-feeding Hemiptera. We investigated the effect of various hemipteran control strategies on terrapin scale densities and measured their indirect effect on local Argentine ant densities and foraging effort. We then determined whether this indirect treatment strategy improved the performance of an ant bait. We predicted that Argentine ants would move nests away from trees treated for Hemiptera and then move nests back when a liquid bait was offered, followed by a decline in ant numbers due to intake of the toxicant. A horticultural oil spray and soil application of the systemic insecticide, imidacloprid, had no effect on terrapin scale numbers. However, trunk-injected dicrotophos caused a reduction in scale and a decline in local Argentine ant nest density and canopy foraging effort. We also recorded a reduction in local Argentine ant ground foraging when large amounts of liquid bait were applied, and we found no evidence that combining dicrotophos with liquid ant bait performed better than each treatment alone. We suggest that a strategy of combined hemipteran control plus application of liquid ant bait can reduce local Argentine ant densities, when both components of this system are highly efficacious.