Prey selection by plovers: Optimal foraging in mixed-species groups

Prey (earthworm) size selection was investigated in lapwings (Vanellus vanellus) and golden plovers (Pluvialis apricaria) feeding in mixed species flocks and compared with that predicted by an optimal foraging model based on energy intake. As well as the usual constraints of searching and handling time, our model incorporated the difficulty of capturing concealed prey, the orientation time needed to locate prey and the risk of theft by gulls (Larus ridibundus). When costs were taken into account, small worms turned out to be the most profitable. The relative profitability of size classes changed when gulls were present and birds shifted their intake accordingly so that they always took mainly the most profitable worms. Birds were expected to do best by taking the three most profitable size classes and the size range taken was consistent with this. In addition there was an inverse relationship between the probabilities of taking profitable and unprofitable worm sizes. Observations of birds were supported by field enclosure experiments which prevented birds feeding in certain areas. Departures from predictions of the model are interpreted as sampling errors due to birds using depth as an approximate indicator of worm size.     

Foraging behaviour of individual coal tits,Parus ater,in relation to their age,sex and morphology

The aim of this study was to reveal the causes and consequences of individual differences in foraging behaviour of coal tits, Parus ater, on the island of Gotland in the Baltic. On Gotland the willow tit, P. montanus, and the crested tit, P. cristatus, are absent, while these larger species are abundant competitors on the mainland. The Gotland population of coal tits exploited parts of the foraging niches of their absent competitors. Gotland coal tits are bigger than those on the mainland; they have thus evolved towards the size of the missing competitors. Individual differences in niche use were related to their morphology. Large birds more often used the inner parts of trees whereas small individuals foraged more on needles in the outer parts of the trees. Also, individuals with wing shapes more suitable for hovering and slow flight foraged more on needles. It is argued that morphology affects the profitability of different foraging sites. However, for foraging site selection, age was more important than morphology. Older, and probably more dominant, individuals occupied the most profitable foraging sites.     

When Parasitoid Males Make Decisions: Information Used when Foraging for Females

Optimal foraging models predict how an organism allocates its time and energy while foraging for aggregated resources. These models have been successfully applied to organisms such as predators looking for prey, female parasitoids looking for hosts, or herbivorous searching for food. In this study, information use and patch time allocation were investigated using male parasitoids looking for mates. The influence of the former presence of females in absence of mates and the occurrence of mating and other reproductive behaviours on the patch leaving tendency was investigated for the larval parasitoid Asobara tabida. Although males do not modify their patch residence time based on the number of females that visited the patch, they do show an increase in the patch residence time after mating a virgin female and performing courtship behaviour such as opening their wings. These results are in concordance with an incremental mechanism, as it has been described for females of the same species while foraging for hosts. The similarities between males and females of the same species, and the conditions under which such a patch-leaving decision rule is fitted are discussed. This is the first study describing an incremental effect of mating on patch residence time in males, thus suggesting that similar information use are probably driving different organisms foraging for resource, regardless of its nature.     

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.     

Heritability of selectively advantageous foraging behaviour in a small passerine

Summary I measured the heritability of foraging patch choice in a laboratory population of zebra finches (Taeniopygia guttata). Mothers and offspring were tested for their ability to discriminate between four foraging patches which provided four different rates of energy gain. Use of a foraging patch with a high rate of energy gain has been shown to confer a selective advantage on zebra finches in a similar experimental system. In this population of zebra finches there was a large amount of variation in foraging patch choice behaviour both within and among individuals. I determined that foraging patch choice was a phenotypically labile trait with a degree of stereotypy or repeatability, much lower than those typically recorded for morphological traits. The mating behaviour of zebra finches required that heritability be determined from a mother—offspring regression, which showed that narrow sense heritability of foraging patch choice was approximately 0.346. This heritability was significantly different than zero, as was heritability when it was limited by repeatability to 0.246. Foraging patch choice, a behaviour that has a demonstrated fitness consequence, had a heritable component in this laboratory population of zebra finches.     

Context dependence in foraging behaviour of Achillea millefolium

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.     

Optimal foraging and risk of claw damage: How flexible are shore crabs in their prey size selectivity?

Understanding the behavioural mechanisms that underlie prey size preference of predators is an essential component of unravelling the processes that govern predator-prey dynamics. In marine systems, despite being able to consume larger and more profitable prey, many molluscivorous predators show a preference for smaller, less profitable prey, most likely to minimize the risk of damaging feeding extremities. Here we assessed the flexibility of this prey size preference. We observed that shore crabs (Carcinus maenas) that were food deprived, and which were offered mussels (Mytilus edulis) of different sizes in dichotomous preference tests, preferred smaller, less profitable mussels. The same result was observed for crabs foraging with a conspecific competitor. Only crabs that were conditioned to feed on the larger, most profitable mussels shifted their prey size preference and ranked the most profitable mussels as highest. Although shore crabs showed flexibility in prey size preference, through which they would be able to cope with environmental variability, our results in general emphasize preference for smaller prey. We discuss the possibility that crabs maximize their long-term feeding rate, in which case it can be optimal to select these smaller mussels.     

Frequency-dependent herbivory by a leaf beetle,Phaedon brassicae,on hairy and glabrous plants of Arabidopsis halleri subsp. gemmifera

Frequency-dependent prey choice by natural enemies may influence the coexistence of multiple prey types, but little is known about whether frequency-dependent foraging choice occurs in herbivory on plants showing resistance polymorphism within a single population. Here we examined frequency-dependent foraging by a crucifer-feeding leaf beetle, Phaedon brassicae, on trichome-producing (hairy) and trichomeless (glabrous) plants coexisting within a natural population of the perennial herb Arabidopsis halleri subsp. gemmifera. Larvae of P. brassicae fed on hairy leaves showed slower growth than those fed on glabrous leaves. Although adult beetles consumed similar amounts of leaves when they were fed either hairy or glabrous leaves in no-choice conditions, our choice experiment showed that adult beetles fed at less than the proportionally expected level on hairy leaves compared to glabrous leaves when the hairy leaves were less or equally abundant. Both types of leaves were consumed at the proportionally expected levels when the hairy leaves were more abundant than the glabrous leaves. In a natural population, the leaf damage on the hairy plants was negatively correlated with the local proportion of the glabrous plants in a 1-m diameter patch across 2 years, while correlations between the leaf damage on the glabrous plants and their proportion differed between the 2 years. Additionally, we found five glucosinolates in leaves of A. halleri, but their accumulation did not differ between hairy and glabrous plants. Our experimental results indicate that hairy plants incur less herbivory by P. brassicae when glabrous plants are abundant. The field pattern provides evidence suggestive of frequency-dependent herbivory acting on hairy plants. The present study highlights one of the putative mechanisms of maintaining plant resistance polymorphism.     

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.     

Nectar provisioning close to host patches increases parasitoid recruitment,retention and host parasitism

In the adult stage, many parasitoids require hosts for their offspring growth and plant-derived food for their survival and metabolic needs. In agricultural fields, nectar provisioning can enhance biological control by increasing the longevity and fecundity of many species of parasitoids. Provided in a host patch, nectar can also increase patch quality for parasitoids and affect their foraging decisions, patch time residence, patch preference or offspring allocation. The aim of this study was to investigate the impact of extrafloral nectar (EFN) provisioning close to hosts on parasitoid aggregation in patches. The aphid parasitoid Diaeretiella rapae (M’Intosh) was released inside or outside patches containing Brassica napus L. infested by Brevicoryne brassicae L. aphids and Vicia faba L. with or without EFN. When parasitoids were released outside patches, more parasitoids were observed in patches with EFN than in patches deprived of EFN. This higher recruitment could be linked to a higher attraction of a combination of host and food stimuli or a learning process. A release–recapture experiment of labeled parasitoids released within patches showed the higher retention of parasitoids in patches providing EFN and hosts, suggesting that food close to the host patch affects patch residence time. Both attractiveness and patch retention could be involved in the higher number of parasitoids foraging in host patches surrounded by nectar and for the higher parasitism recorded. Nectar provisioning in host patches also affected female offspring allocation inside the patch.     

Foraging on patchily distributed prey by a cichlid fish (Teleostei,Cichlidae): A test of the ideal free distribution theory

Cichlid fish (Aequidens curviceps) distributed themselves and allocated their foraging time between two drift food patches in close approximation to the patch profitability ratio, as predicted by the ideal free distribution theory. The fish thereby achieved similar average feeding rates in the two patches, in two of three patch profitability ratio experiments. However, one major assumption of the ideal free model was violated, since individual fish differed in their competitive abilities for limited food resources, which resulted in unequal payoffs among individuals within each patch. Individual variation in feeding rates, and thus in competitive ability, was not related to despotism, but perhaps rather to individual differences in perceptual ability and in the ability to learn which patch was currently the more profitable. The strategy used by the fish to assess patch profitability included sampling available patches. However, individual fish switched (sampled) patches with varying frequency. Sampling had an associated cost, since high-frequency switchers had lower feeding rates on average than low-frequency switchers. Differences in foraging strategy among the fish therefore contributed to the observed in-equality in individual payoffs within patches.     

Nutrient balance affects foraging behaviour of a trap-building predator

Predator foraging may be affected by previous prey capture, but it is unknown how nutrient balance affects foraging behaviour. Here, we use a trap-building predator to test whether nutrients from previous prey captures affect foraging behaviour. We fed orb-weaving spiders (Zygiella x-notata) prey flies of different nutrient composition and in different amounts during their first instar and measured the subsequent frequency of web building and aspects of web architecture. We found that both the likelihood of web building and the number of radii in the web were affected by prey nutrient composition while prey availability affected capture area and mesh height. Our results show that both the balance of nutrients in captured prey and the previous capture rate may affect future foraging behaviour of predators.     

Local floral composition and the behaviour of pollinators: attraction to and foraging within experimental patches

1. Understanding how foraging decisions take place at the local scale is relevant because they may directly affect the fitness of individual plants. However, little is known about how local diversity and density affect the foraging behaviour of most pollinator groups. 2. By introducing two potted plant species (Salvia farinacae and Tagetes bonanza) into two populations of Taraxacum officinale, we investigated how plant identity, the mixtures of these plant species, and total plant density affected the attraction to and the foraging within a patch for six pollinator groups. 3. The foraging behaviour was mainly driven by the availability of the preferred plant species, and secondly by patch diversity and density. In general, dense patches and those containing the three‐species mixture were preferred by all insect groups for arrival, although muscoid and hover flies responded less to local floral composition than bees. Local diversity and density had, however, a weaker effect on foraging behaviour within patches. Site dependence in response to floral treatments could be attributable to differences between sites in pollinator assemblage and Taraxacum density. 4. Studies like ours will help to understand how foraging decisions occur at the local scale and how foraging patterns may differ between pollinators and sites.     

Experimental analyses of patch selection by foraging black surfperch (Embiotoca jacksoni Agazzi)

The dynamics of microhabitat use by foraging adult and juvenile black surfperch (Embiotocajacksoni Agazzi) were explored. Detailed observations of black surfperch feeding at Santa Catalina Island, California, revealed that adults and young-of-year juveniles co-occurred in the same habitat but used different algal substrata as foraging sites. Juveniles selected invertebrate prey almost exclusively from the surface of foliose algae. The occurrence of young E. jacksoni was highly correlated with that of foliose algae. Adults tended to bite most frequently from turf, a low-growing matrix of plants, colonial animals, and debris covering the rocky substratum. The abundance of adults was negatively correlated with the occurrence of foliose algae. Adults and juveniles showed marked, but different, preferences in their utilization of taxa of algae as foraging substrata. Certain algae (e.g., Zonaria farlowii Setchell & Gardner) were preferred while other taxa (e.g., Sargassum palmeri Grun) were avoided by both age groups. However, most types of algae were preferred by one group but not the other. To test the hypothesis that knowledge of algal substratum composition allows prediction of fish occurrence and foraging behavior in a patch, algal cover on 2 × 2 m² areas of bottom was manipulated creating plots dominated by turf, Zonaria farlowii, or Sargassum palmeri. Fish occurrence could be accurately predicted on the basis of abundance of foliose algae, but foraging activity of fish was highly dependent on the algal taxon that dominated the patch. Differential prey availabilities among foraging substrata provided some insight into the patterns of foraging patch preferences displayed by adult and juvenile Embiotoca jacksoni.     

Foraging Patch Selection and Departure by Non-Omniscient Foragers: A Field Example in White-Fronted Geese

Animals often face great uncertainty as to the quality of foraging patches. There have been a number of theoretical studies investigating how non‐omniscient predators, i.e. predators that are unable to assess foraging patch quality prior to patch exploitation, should forage in a heterogeneous environment, but empirical studies, especially in the field, are scarce. This paper describes the way in which white‐fronted geese Anser albifrons forage on harvest remains of rice, focusing on the processes of patch selection and departure. Not only in autumn, but also in spring when rice depletion has progressed, patch rice density showed no positive effect on patch selection by geese, indicating the incapability of geese to select the most profitable patch. Instead, the geese tended to select patches with a large proportion of rice fields that were near the roost and a previously visited patch and bordered by a small number of windbreaks only. The rice consumption volume by geese increased with increasing initial rice density, while giving‐up density was independent of initial rice density and was positively correlated to the mean rice density of the habitat. This suggests that the geese could compensate their lack of information on patch quality at the moment of patch selection by leaving less profitable patches earlier. We discuss the necessity of predictive models based on random patch selection and an appropriate departure rule to explain the distribution of individuals of species with limited information on patch quality.     

Impact of landscape fragmentation on a specialised woodland bat,Rhinolophus hipposideros

For habitat specialists, fragmentation has major consequences as it means less suitable habitat for the species to live in. In a fragmented landscape, we would expect larger, but spatially more clustered, foraging ranges. We studied the impact of landscape fragmentation on the foraging range and habitat exploitation of a specialised forest bat by radiotracking 16 female lesser horseshoe bats Rhinolophus hipposideros in a landscape with connected woodland structures and in a highly fragmented landscape in Carinthia, Austria.Contrary to our expectations, spatial foraging behaviour was not influenced by fragmentation. No differences in the behaviour of the bats between the sites were evident for the foraging ranges (minimum convex polygon, MCP), the core foraging areas (50% kernel), nor the mean or the maximum distances from the roost. However, in the highly fragmented landscape, the foraging activity of individuals was spatially more clustered and the overall MCP of all bats of a colony was greater compared to the less fragmented landscape.Woodland was the most important foraging habitat for the lesser horseshoe bats at both study sites. Habitat selection at the individual MCPs was evident only at the site with low fragmentation. However, in the core foraging areas, woodland was significantly selected over all other habitat types at both study sites.We conclude that (1) conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost, (2) woodland is the key foraging habitat particularly in the vicinity of the roost, and (3) any loss of woodland near the colonial roosts are likely to negatively influence the colony, since these bats do not seem to be able to adapt their spatial foraging behaviour in a degraded landscape. The inflexible spatial behaviour of this specialised bat highlights the need to compensate for any habitat loss within the foraging range of a bat colony.     

Predator interference alters foraging behavior of a generalist predatory arthropod

Interactions between predators foraging in the same patch may strongly influence patch use and functional response. In particular, there is continued interest in how the magnitude of mutual interference shapes predator–prey interactions. Studies commonly focus on either patch use or the functional response without attempting to link these important components of the foraging puzzle. Predictions from both theoretical frameworks suggest that predators should modify foraging efforts in response to changes in feeding rate, but this prediction has received little empirical attention. We study the linkage between patch departure rates and food consumption by the hunting spider, Pardosa milvina, using field enclosures in which prey and predator densities were manipulated. Additionally, the most appropriate functional response model was identified by fitting alternative functional response models to laboratory foraging data. Our results show that although prey availability was the most important determinant of patch departure rates, a greater proportion of predators left enclosures containing elevated predator abundance. Functional response parameter estimation revealed significant levels of interference among predators leading to lower feeding rates even when the area allocated for each predator was kept constant. These results suggest that feeding rates determine patch movement dynamics, where interference induces predators to search for foraging sites that balance the frequency of agonistic interactions with prey encounter rates.     

Feeding rate and attack specialization: the roles of predator experience and energetic tradeoffs

Synopsis Individual mosquitofish, Gambusia affinis, can adopt a broad range of attack selectivities. In part, this variation can be explained by the past experiences of a fish. Individuals selected the more profitable Ceriodaphnia dubia (Cladocera) over less profitable cyclopoid copepods to a greater degree after being exposed to both prey types than did individuals experienced with only one of the prey types. Feeding rate (biomass ingested per unit time) declined with increased attack specialization on the profitable prey (Ceriodaphnia) when such prey were scarce, a result in agreement with assumptions of optimal diet theory. When profitable prey were abundant feeding rate was a bimodal function of the intensity of specialization on profitable prey; fish that specialized on cyclopoid copepods (the less profitable prey type) fed at higher rates than did generalists. This may be the result of antagonistic learning that precluded feeding efficiently on more than one type of prey at a time. The data are consistent with the hypothesis that rejection of unsuitable prey involves a time cost. The two preceeding aspects of foraging behavior, which are absent from most optimal diet models, could lead to failure in predicting the attack specialization of some predators, An additional aspect of the results was the generally weak relationship between feeding efficiency and specialization behavior. This suggests that feeding rate may not have been as tightly linked to the specialization behavior a predator adopts as is assumed by current foraging theory.     

Optimal patch use in a stochastic environment

This paper considers an animal foraging on prey which are distributed in well-defined patches. It is assumed that the environment may be stochastic and that the animal can gain information on patch type as it forages. The foraging policy which maximises mean reward rate for the environment is characterised in terms of a function of state called the potential function. This policy is shown to be given by the rule: continue foraging on the present patch while the potential is positive, when the potential falls to zero move on to the next patch. Let r denote the current reward rate on a patch and let γ denote the maximum mean reward rate for the environment. It is shown that r ? γ if it is optimal to leave. Conditions which ensure r < γ are also given. For a large class of environments the optimal policy is stated in terms of a revised reward rate r?, and is given by the rule: continue on the present patch while $\text{r}\text{?}\text{> γ}$, when r? falls to γ move on to the next patch. Finally, it is shown that the stay time on a patch is a decreasing function of γ.     

Aspects of the biology of Pantinonemertes californiensis,a high intertidal nemertean

I studied the distribution, feeding biology, and reproductive biology of Pantinonemertes californiensis, described as a semi-terrestrial nemertean, along the central California coast. At the sites used in this study, maximal tidal height is about 2 m, and P. californiensis typically occurred under boulders between 1.3 and 1.7 m tidal height. Worms fed primarily on the semi-terrestrial amphipod Traskorchestia traskiana. Distribution of nemerteans was similar to that of the prey, although prey extended higher on the beach than did the worms. Nemerteans were largest and most abundant at the site with highest abundance of T. traskiana and smallest and least abundant at the lowest prey abundance site. In laboratory feeding trials, nemerteans from the site with lowest prey abundance fed most readily. Non-reproductive nemerteans lived for at least a week when submerged in sea water; some prey died within a week of being submerged. Nemerteans only lived minutes when submerged in fresh water; 50% of prey lived 4.5 h. Eggs are approximately 90–100 μm in diameter and hundreds to thousands are shed per female. Larvae are planktonic and apparently planktotrophic, and are morphologically similar to other marine hoplonemertean larvae. At the sites studied life history characteristics of P. californiensis provided little evidence of adaptations to terrestrial life in these worms and were not helpful in elucidating the role of semi-terrestrial nemerteans in the evolution of terrestrial nemerteans.