Risk-sensitive foraging in common shrews (Sorex araneus L.)

Summary Risk-sensitive foraging theory predicts that predators which face starvation if there is a temporary shortfall in their food supply should choose feeding sites on the basis of variation in as well as mean expected reward rate. For a given mean reward rate they should choose high variance feeding sites (be risk-prone) if they are running below energy requirement, but low variance sites (be risk-averse) if they are running above.Common shrews presented with a choice between constant and variable feeding stations were more likely to visit the variable station when they were running below energy requirement and more likely to visit the constant station when they were running above. However, the tendency towards risk-aversion above requirement was greater than that towards risk-proneness below.When all shrews were considered together, the probability of visiting the variable station correlated negatively and continuously with intake relative to requirement.     

Nectar-receiver behavior in relation to the reward rate experienced by foraging honeybees

Since forager honeybees change their food-unloading behavior according to nectar-source profitability, an experiment was performed in order to analyze whether food-receivers modify their within-hive tasks related to different reward conditions. We offered individual foragers two reward conditions at a rate feeder while an additional feeder offered a constant reward and was of free access to the rest of the hive. Both feeders were the only food sources exploited by the colony during the assays since a flight chamber was used. After receiving nectar, hive bees performed processing cycles that involved several behaviors and concluded when they returned to the delivery area to receive a new food sample. During these cycles, receivers mainly performed oral contacts offering food, or inspected cells, and often both. In the latter case, both behaviors occurred simultaneously and at the same distance from the hive entrance. When they performed a single task, either the occurrence of cell inspections increased or contact offerings decreased for the highest reward rate offered to the donor-forager. Receivers also begged for food more often after interacting with low-profit foragers. Thus, the profitability of the food source exploited by nectar-forager honeybees could affect receiver behaviors within the hives based on individual-to-individual interactions.Communicated by R.F.A. Moritz     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Social foraging and dominance relationships: the effects of socially mediated interference

In socially foraging animals, it is widely acknowledged that the position of an individual within the dominance hierarchy of the group has a large effect upon its foraging behaviour and energetic intake, where the intake of subordinates can be reduced through socially mediated interference. In this paper, we explore the effects of interference upon group dynamics and individual behaviour, using a spatially explicit individual-based model. Each individual follows a simple behavioural rule based upon its energetic reserves and the actions of its neighbours (where the rule is derived from game theory models). We show that dominant individuals should have larger energetic reserves than their subordinates, and the size of this difference increases when either food is scarce, the intensity of interference suffered by the subordinates increases, or the distance over which dominant individuals affect subordinates increases. Unlike previous models, the results presented in this paper about differences in reserves are not based upon prior assumptions of the effects of social hierarchy and energetic reserves upon predation risk, and emerge through nothing more than a reduction in energetic intake by the subordinates when dominants are present. Furthermore, we show that increasing interference intensity, food availability or the distance over which dominants have an effect also causes the difference in movement between ranks to increase (where subordinates move more than dominants), and the distance over which dominants have an effect changes the size of the groups that the different ranks are found in. These results are discussed in relation to previous studies of intra- and interspecific dominance hierarchies.     

Temporal variance reverses the impact of high mean intensity of stress in climate change experiments

Extreme climate events produce simultaneous changes to the mean and to the variance of climatic variables over ecological time scales. While several studies have investigated how ecological systems respond to changes in mean values of climate variables, the combined effects of mean and variance are poorly understood. We examined the response of low-shore assemblages of algae and invertebrates of rocky seashores in the northwest Mediterranean to factorial manipulations of mean intensity and temporal variance of aerial exposure, a type of disturbance whose intensity and temporal patterning of occurrence are predicted to change with changing climate conditions. Effects of variance were often in the opposite direction of those elicited by changes in the mean. Increasing aerial exposure at regular intervals had negative effects both on diversity of assemblages and on percent cover of filamentous and coarsely branched algae, but greater temporal variance drastically reduced these effects. The opposite was observed for the abundance of barnacles and encrusting coralline algae, where high temporal variance of aerial exposure either reversed a positive effect of mean intensity (barnacles) or caused a negative effect that did not occur under low temporal variance (encrusting algae). These results provide the first experimental evidence that changes in mean intensity and temporal variance of climatic variables affect natural assemblages of species interactively, suggesting that high temporal variance may mitigate the ecological impacts of ongoing and predicted climate changes.     

Nest defense and central place foraging: A model and experiment

Summary A graphical model presented here indicates that a nest-defending forager should stay closer to its nest, forage for shorter times per patch, and deliver smaller loads than predicated for delivery rate maximization. The effect is more pronounced farther from the nest, so that if nest defense is especially important, the predator should leave far patches sooner than near ones, and deliver smaller loads from farther away. Moreover, if the attack rate at the nest is increased, the defending forager should move closer and deliver smaller prey.Experimental attacks with stuffed specimens at Gila woodpecker (Melanerpes uropygialis) nests produced the predicted changes in the foraging behavior of males, but not of females.Mated pairs may work as a team to pursue simultaneously two conflicting goals—food delivery and nest protection—both of which affect the survivorship of the young. Sexual dimorphism in monogamous species may result in part from specialization in these roles.     

Exploring the effects of reductions in krill biomass in the Southern Ocean on blue whales using a state-dependent foraging model

Many species of baleen whales were hunted to near extinction in the Southern Hemisphere. The recovery of these populations will be affected by the availability of krill, a major dietary component, in the Southern Ocean. We combine a novel energetics model for baleen whales with a state dependent foraging model to explore the impacts of an expanding krill fishery on baleen whales. We parameterize the model for blue whales, but with simple modifications it could be applied to most baleen whales. We predict that an expanding fishery will have a small but significant impact on the blue whale population through decreased birth rates. However, spreading the catch limit throughout the range of krill can reduce these effects. In addition, whales may be able to reduce these impacts through adaptive changes in foraging behavior. The relationship between krill abundance and blue whale foraging and reproductive success is nonlinear, such that larger reductions in krill biomass, potentially following a loss of sea ice due to climate change, could have a much larger negative impact on the recovery of blue whales.     

Difficulties with the lognormal model in mean estimation and testing

A frequent assumption in environmental risk assessment is that the underlying distribution of an analyte concentration is lognormal. However, the distribution of a random variable whose log has a t-distribution has infinite mean. Because of the proximity of the standard normal and t-distribution, this suggests that a distribution such as the gamma or truncated normal, with smaller right tail probabilities, might make a better statistical model for mean estimation than the lognormal. In order to assess the effect of departures from lognormality on lognormal-based statistics, we simulated complete lognormal, truncated normal, and gamma data for various sample sizes and coefficients of variation. In these cases, departures from lognormality were not easily detected with the Shapiro-Wilk test. Various lognormal-based estimates and tests were compared with alternate methods based on the ordinary sample mean and standard error. The examples were also considered in the presence of random left censoring with the mean and standard error of the product limit estimate replacing the ordinary sample mean and standard error. The results suggest that in the estimation of or tests about a mean, if the assumption of lognormality is at all suspect, then lognormal-based approaches may not be as good as the alternative methods.     

Body size,demography and foraging in a socially plastic sweat bee: a common garden experiment

Phenotypic plasticity may evolve when conditions vary temporally or spatially on a small enough scale. Plasticity is thought to play a central role in the early stages of evolutionary transitions, including major transitions such as those between non-sociality and sociality. The sweat bee Halictus rubicundus is of special interest in this respect, because it is socially plastic in the British Isles: Nests are social or non-social depending on the environment. However, sociality comprises a complex suite of inter-related traits. To further investigate social plasticity in H. rubicundus, we measured traits that are potentially integral to social phenotype at a northern site, where nests are non-social, and a southern site where nests can be social. We found that foundresses at non-social sites were smaller, produced offspring of a size more similar to themselves, initiated nesting later, and took longer to produce their first female offspring. They began provisioning earlier in the day, finished earlier, and collected more pollen loads. Common garden experiments suggested that these differences represent mainly plasticity, as expected for traits involved in the overall plastic social phenotype, with only limited evidence for fixed genetic differences in foraging. Conditions during overwintering did not have major effects on a foundress' subsequent behaviour.     

Effects of parasitic mites and protozoa on the flower constancy and foraging rate of bumble bees

Parasites can affect host behavior in subtle but ecologically important ways. In the laboratory, we conducted experiments to determine whether parasitic infection by the intestinal protozoan Crithidia bombi or the tracheal mite Locustacarus buchneri alters the foraging behavior of the bumble bee Bombus impatiens. Using an array of equally rewarding yellow and blue artificial flowers, we measured the foraging rate (flowers visited per minute, flower handling time, and flight time between flowers) and flower constancy (tendency to sequentially visit flowers of the same type) of bees with varying intensities of infection. Bumble bee workers infected with tracheal mites foraged as rapidly as uninfected workers, but were considerably more constant to a single flower type (yellow or blue). In contrast, workers infected with intestinal protozoa showed similar levels of flower constancy, but visited 12% fewer flowers per minute on average than uninfected bees. By altering the foraging behavior of bees, such parasites may influence interactions between plants and pollinators, as well as the reproductive output of bumble bee colonies. Our study is the first to investigate the effects of parasitic protozoa and tracheal mites on the foraging behavior of bumble bees, and provides the first report of Crithidia bombi in commercial bumble bees in North America.     

Calculation of search volume on cruise-searching planktivorous fish in foraging model

Search volume of cruising planktivorous fish was calculated based on its detailed behavior Th examine the factors influencing search volume, a series of experiments were conducted by varying ambient conditions, such as structural complexity light intensity and turbidity Pseudorasbora parva were used in experiment as predator and Daphnia pulex was selected as prey The shape of scanning area of P parva showed elliptic and the search volume changed drastically depending on ambient conditions. Compared with the results of previous foraging model, the search volumes of the fish under previous study were larger (1.2 to 2.4 times) than those from our study These results on the changes in feeding rate can be useful in determining microhabitat requirement of P parva and othercyprinids with a similar foraging behavior The calculated search volume is compared with other foraging model andthe effect of zooplankton-planktivore interactions on aquatic ecosystem is discussed.     

Evaluating the effects of foraging habitat restoration on shorebird reproduction: the importance of performance criteria and comparative design

Coastal development and engineering projects preclude ecosystem processes that provide habitat for beach nesting birds. Management for coastal species may depend on actions that attempt to restore important habitat features and mitigate disturbance effects. However, species response to restoration or other management actions may be difficult to predict or measure. At Jones Beach State Park, on Long Island, New York, a 0.49 ha restoration project provided moist substrate foraging habitat for breeding Piping Plovers (Charadrius melodus) from 2002 to 2005. We examined whether foraging habitat restoration affected Piping Plover breeding population size, productivity, and fledgling production within 300 m of the restoration site. We found a positive relationship between habitat restoration and the number of fledglings produced per year. However, foraging habitat restoration did not significantly increase the number of Plover pairs breeding at Jones Beach. Our ability to evaluate restoration effects on Plovers depended on: 1) use of multiple performance criteria; 2) a design that allowed comparison of pre- and post-restoration data; and 3) a spatial control that allowed comparison of similar areas that were near and far from the restoration site. Despite the small size of the restoration project, there were measurable benefits to Plovers, indicating that foraging habitat restoration may be an effective tool for species recovery.     

Effects of experience and weather on foraging rate and pollen versus nectar collection in the bumblebee, Bombus terrestris

This study examines factors that affect foraging rate of free-flying bumblebees, Bombus terrestris, when collecting nectar, and also what factors determine whether they collect pollen or nectar. We show that nectar foraging rate (mass gathered per unit time) is positively correlated with worker size, in accordance with previous studies. It has been suggested that the greater foraging rate of large bees is due to their higher thermoregulatory capacity in cool conditions, but our data suggest that this is not so. Workers differing in size were not differentially affected by the weather. Regardless of size, naïve bees were poor foragers, often using more resources than they gathered. Foraging rate was not maximised until at least 30 trips had been made from the nest. Foraging rates were positively correlated with humidity, perhaps because nectar secretion rates were higher or evaporation of nectar lower at high humidity. Temperature, wind speed and cloud cover did not significantly influence foraging rate, within the summertime range that occurred during the study. Weather greatly influenced whether bees collected pollen or nectar. Pollen was preferably collected when it was warm, windy, and particularly when humidity was low; and preferably during the middle of the day. We suggest that bees collect pollen in dry conditions, and avoid collecting pollen when there is dew or rain-water droplets on the vegetation, which would make grooming pollen into the corbiculae difficult. Availability of sufficient dry days for pollen collection may be an important factor determining the success of bumblebee colonies.Communicated by M. Giurfa     

Alternative reproductive tactics in male common shrews: relationships between mate-searching behaviour,sperm production,and reproductive success as revealed by DNA fingerprinting

The common shrew (Sorex araneus) is a solitary small mammal with a promiscuous mating system. Previous studies of this species suggest that females typically mate multiply, and that males may adopt alternative mate-searching tactics. We studied two generations of common shrews in a population near Oxford, England. Males were found to adopt two different mate-searching tactics. Those classed as type A occupied relatively small exclusive ranges during March, and made repeated long-distance movements to visit female ranges around the time of first oestrus in April. Males classed as type B established large overlapping ranges in areas of relatively high female density during March, and maintained these ranges throughout April. Type B males were larger than type A males at an early stage of sexual maturation, but there was no difference in the adult body size of the two types of male. Type A males had significantly higher epididymal sperm counts than type B males. Paternity analyses of litters born during the first year of the study reveal that the mean number of offspring fathered by type B males was greater than the mean number fathered by type A males. It is concluded that different mate-searching tactics may be conditional upon the timing of sexual maturation. Differences in sperm production are discussed in relation to sperm competition theory.     

Robust analysis of variance with imprecise data: an ad hoc algorithm

We present an easy to implement algorithm, which is valid to analyse the variance of data under several robust conditions. Firstly, the observations may be precise or imprecise. Secondly, the error distributions may vary within the wide class of the strongly unimodal distributions, symmetrical or not. Thirdly, the variance of the errors is unknown. The algorithm starts by estimating the parameters of the ANOVA linear model. Then, the asymptotic covariance matrix of the effects is estimated. Finally, the algorithm uses this matrix estimate to test ANOVA hypotheses posed in terms of linear combinations of the effects.     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

Regulation of pollen foraging in honeybee colonies: effects of young brood, stored pollen, and empty space

Pollen storage in a colony of Apis mellifera is actively regulated by increasing and decreasing pollen foraging according to the “colony's needs.” It has been shown that nectar foragers indirectly gather information about the nectar supply of the colony from nestmates without estimating the amount of honey actually stored in the combs. Very little is known about how the actual colony need is perceived with respect to pollen foraging. Two factors influence the need for pollen: the quantity of pollen stored in cells and the amount of brood. To elucidate the mechanisms of perception, we changed the environment within normal-sized colonies by adding pollen or young brood and measured the pollen-foraging activity, while foragers had either direct access to them or not. Our results show that the amount of stored pollen, young brood, and empty space directly provide important stimuli that affect foraging behavior. Different mechanisms for forager perception of the change in the environment are discussed. Received: 13 June 1998 / Accepted after revision: 25 October 1998     

The effects of variable predation risk on foraging and growth: less risk is not necessarily better

There is strong evidence that the way prey respond to predation risk can be fundamentally important to the structuring and functioning of natural ecosystems. The majority of work on such nonconsumptive predator effects (NCEs) has examined prey responses under constant risk or constant safety. Hence, the importance of temporal variation in predation risk, which is ubiquitous in natural systems, has received limited empirical attention. In addition, tests of theory (e.g., the risk allocation hypothesis) on how prey allocate risk have relied almost exclusively on the behavioral responses of prey to variation in risk. In this study, we examined how temporal variation in predation risk affected NCEs on prey foraging and growth. We found that high risk, when predictable, was just as energetically favorable to prey as safe environments that are occasionally pulsed by risk. This pattern emerged because even episodic pulses of risk in otherwise safe environments led to strong NCEs on both foraging and growth. However, NCEs more strongly affected growth than foraging, and we suggest that such effects on growth are most important to how prey ultimately allocate risk. Hence, exclusive focus on behavioral responses to risk will likely provide an incomplete understanding of how NCEs shape individual fitness and the dynamics of ecological communities.     

Using an individual-based model to quantify scale transition in demographic rate functions: Deaths in a coral reef fish

Scientifically informed population management requires quantitatively accurate demographic rate functions that apply at the spatial scale at which populations are actually managed, but practical constraints confine most field measurements of such functions to small study plots. This paper employs an individual-based population growth model to extrapolate the death rate function in a well-studied coral reef fish, the bridled goby Coryphopterus glaucofraenum, from the scale of coral reef “cells” at which it was measured to the larger scale of an entire coral reef. Density dependence in the whole-reef function actually proves stronger than in the local function because high goby density occasionally arises in local patches with few refuges from predators, producing very high mortality there. This IBM-based approach extends the reach of scale transition theory by examining considerably more realistic models than standard analytical methods can presently handle.