Using Past Experience in Web Relocation Decisions Enhances the Foraging Efficiency of the Spider Cyclosa argenteoalba

The purpose of this study is to test whether the integration of past and present foraging experience in web relocation decision enhances foraging efficiency of the spider Cyclosa argenteoalba in its natural environment. We measured daily changes in the prey availability at several fixed sites in a natural environment and constructed a model environment based on these observational data. In the model environment, we simulated the behavior of spiders that foraged and relocated their webs according to several decision rules, which differed in terms of how a spider used its past experience. Results of the simulation revealed that the less past experience is discounted in making web relocation decisions, the more prey the spider is expected to capture. The expected number of web relocations decreased as spiders kept past foraging experience longer. These results suggest that C. argenteoalba enhances foraging efficiency by using past foraging experience for long times in the decision of web relocation in its natural environment.     

Foraging strategy predicts foraging economy in a facultative secondary nectar robber

In mutualistic interactions, the decision whether to cooperate or cheat depends on the relative costs and benefits of each strategy. In pollination mutualisms, secondary nectar robbing is a facultative behavior employed by a diverse array of nectar‐feeding organisms, and is thought to be a form of cheating. Primary robbers create holes in floral tissue through which they feed on nectar, whereas secondary robbers, which often lack chewing mouthparts, feed on nectar through existing holes. Because primary robbers make nectar more readily available to secondary robbers, primary robbers facilitate the behaviors of secondary robbers. However, the net effect of facilitation on secondary robber fitness has not been empirically tested: it is unknown whether the benefit secondary robbers receive is strong enough to overcome the cost of competing with primary robbers for a shared resource. We conducted foraging experiments using the bumble bee Bombus bifarius, which can alternatively forage ‘legitimately’ (from the floral opening) or secondary‐rob. We measured the relative foraging efficiencies (handling time per flower, flowers visited per minute, proportion of foraging bout spent consuming nectar) of these alternative behaviors, and tested whether the frequency of primary robbing and nectar standing crop in primary‐robbed flowers of Linaria vulgaris (Plantaginaceae) affected foraging efficiency. Surprisingly, there was no effect of primary robbing frequency on the foraging efficiency of secondary‐robbing B. bifarius. Instead, foraging strategy was a major predictor of foraging efficiency, with legitimate foraging being significantly more efficient than secondary robbing. Legitimate foraging was the more common strategy used by B. bifarius in our study; however, it is rarely used by B. bifarius foraging on L. vulgaris in nature, despite indications that it is more efficient. Our results suggest the need for deeper investigations into why bees adopt secondary robbing as a foraging strategy, specifically, the environmental contexts that promote the behavior.     

Exploring foraging decisions in a social primate using discrete-choice models

Abstract There is a growing appreciation of the multiple social and nonsocial factors influencing the foraging behavior of social animals but little understanding of how these factors depend on habitat characteristics or individual traits. This partly reflects the difficulties inherent in using conventional statistical techniques to analyze multifactor, multicontext foraging decisions. Discrete-choice models provide a way to do so, and we demonstrate this by using them to investigate patch preference in a wild population of social foragers (chacma baboons Papio ursinus). Data were collected from 29 adults across two social groups, encompassing 683 foraging decisions over a 6-month period and the results interpreted using an information-theoretic approach. Baboon foraging decisions were influenced by multiple nonsocial and social factors and were often contingent on the characteristics of the habitat or individual. Differences in decision making between habitats were consistent with changes in interference-competition costs but not with changes in social-foraging benefits. Individual differences in decision making were suggestive of a trade-off between dominance rank and social capital. Our findings emphasize that taking a multifactor, multicontext approach is important to fully understand animal decision making. We also demonstrate how discrete-choice models can be used to achieve this.     

Influence of environmental and colony factors on the initial commodity choice of foragers of the stingless bee Plebeia tobagoensis (Hymenoptera, Meliponini)

Novice foragers of social bees have to decide what food commodity to collect when they start foraging for the first time. In this decision making process two types of factors are involved: internal factors (the response threshold) and external factors (environmental and colony conditions). In this study we will focus on the importance of two external factors, pollen storage level and information from experienced foragers about food availability in the field, on the initial commodity choice of foragers of the stingless bee species Plebeia tobagoensis. We also studied the effect of the initial choice of individuals on their subsequent foraging career. This study was performed in a closed greenhouse compartment, where food availability and colony condition could be controlled. Information on food availability in the field from experienced foragers and pollen storage level both greatly influenced the initial commodity choice of individuals, with more choices for the commodity communicated by experienced foragers or lacking in storage. The initial choice of foragers is of importance for their future foraging career, although a substantial proportion of foragers did switch between food commodities. Because of the ability of novice foragers to become flexibly distributed over foraging tasks, social bees are able to react to changes in their environment without directly having to decrease foraging effort devoted to other foraging tasks. This, in combination with individual flexibility during foraging careers makes it possible for colonies of P. tobagoensis to forage efficiently in an ever-changing environment. Received 7 November 2005; revised 12 January 2006; accepted 16 February 2006.     

Central place foraging in larvae of the charaxine butterfly,Polyura pyrrhus (L.): A case study in a herbivore

Central place foraging by larvae of the charaxine butterfly,Polyura pyrrhus, was studied. Larvae made foraging trips from the silken pads they constructed on leaflets of their foodplant,Acacia sp. A foraging trip sometimes involved complete depletion of a single patch of foodplant pinnules. Larvae which did not deplete a patch appeared to eat until they were satiated, whereas larvae which depleted a patch either visited another patch (multiple-patch foraging) or returned directly to the pad (single-patch foraging). If the food intake at the first patch was small a larva tended to make a “multiple-patch” decision, especially when the pinnule-patch was distant from the resting pad. The duration between successive foraging trips (resting time on the pad) was much longer than the round trip duration: on average about 3 h and 15 min, respectively. The resting time is suggested to be a handling time (i.e., digesting food in the gut) and was disproportional to the amount of food consumed, i.e., the handling efficiency was higher when the larva consumed a larger amount of food. This may be the reason why larvae usually ate until they were satiated. A food-intake-rate maximizing model was constructed to describe the decision rule as to whether a larva should make a single-patch or a multiple-patch foraging trip. One of the model's predictions (i.e., larvae should engage in multiple-patch foraging when the food intake at the first patch is small) qualitatively corresponds with data, however, the model does not explain the effect of travelling time on decision making in larvae. Several other factors which may influence the decision making of larvae are discussed.     

To be or not to be faithful: flexible fidelity to foraging trails in the leaf‐cutting ant Acromyrmex lobicornis

1. Ants using trails to forage have to select between two alternative routes at bifurcations, using two, potentially conflicting, sources of information to make their decision: individual experience to return to a previous successful foraging site (i.e. fidelity) and ant traffic. In the field, we investigated which of these two types of information individuals of the leaf‐cutting ant Acromyrmex lobicornis Emery use to decide which foraging route to take. 2. We measured the proportion of foraging ants returning to each trail of bifurcations the following day, and for 4–7 consecutive days. We then experimentally increased ant traffic on one trail of the bifurcation by adding additional food sources to examine the effect of increased ant traffic on the decision that ants make. 3. Binomial tests showed that for 62% of the trails, ant fidelity was relatively more important than ant traffic in deciding which bifurcation to follow, suggesting the importance of previous experience. 4. When information conflict was generated by experimentally increasing ant traffic along the trail with less foraging activity, most ants relied on ant traffic to decide. However, in 33% of these bifurcations, ants were still faithful to their trail. Thus, there is some degree of flexibility in the decisions that A. lobicornis make to access food resources. 5. This flexible fidelity results in individual variation in the response of workers to different levels of ant traffic, and allows the colony to simultaneously exploit both established and recently discovered food patches, aiding efficient food gathering.     

Does Wood Fuel Gathering for Household Use Follow an Optimality Model? A Study from Kakamega Forest,Western Kenya

Successful management of forests, especially in the context of subsistence wood fuel use, can be improved by the application of theoretic models that predict patterns of use. A first step in this approach is understanding the decision rules of people using forests. While optimal foraging theory is normally applied to foraging for food, it also makes sense to apply it in the context of “foraging” for wood fuel. In this study, we applied a time allocation model of optimal foraging theory to wood gathering decisions in communities around Kakamega Forest, a mid-altitude seasonal tropical rain forest. The model predicts that the amount of wood gathered should increase with the distance to the wood source. We found that the predictions of OFT are supported, but only for adults and more strongly on a weekly scale. Based upon the results, we then discuss future improvements of the model to better understand and predict human use.     

State dependence,personality, and plants: light‐foraging decisions in Mimosa pudica (L.)

Plants make foraging decisions that are dependent on ecological conditions, such as resource availability and distribution. Despite the field of plant behavioral ecology gaining momentum, ecologists still know little about what factors impact plant behavior, especially light‐foraging behavior. We made use of the behavioral reaction norm approach to investigate light foraging in a plant species that exhibits rapid movement: Mimosa pudica. We explored how herbivore avoidance behavior in M. pudica (which closes its leaflets temporarily when disturbed) is affected by an individual's energy state and the quality of the current environment and also repeatedly tested the behavior of individuals from two seed sources to determine whether individuals exhibit a “personality” (i.e., behavioral syndrome). We found that when individuals are in a low‐energy state, they adopt a riskier light‐foraging strategy, opening leaflets faster, and not closing leaflets as often in response to a disturbance. However, when plants are in a high‐energy state, they exhibit a plastic light‐foraging strategy dependent on environment quality. Although we found no evidence that individuals exhibit behavioral syndromes, we found that individuals from different seed sources consistently behave differently from each other. Our results suggest that plants are capable of making state‐dependent decisions and that plant decision making is complex, depending on the interplay between internal and external factors.     

Molecular and spatial analyses reveal links between colony‐specific foraging distance and landscape‐level resource availability in two bumblebee species

Foraging distance is a key determinant of colony survival and pollination potential in bumblebees Bombus spp. However this aspect of bumblebee ecology is poorly understood because of the difficulty in locating colonies of these central place foragers. Here, we used a combination of molecular microsatellite analyses, remote sensing and spatial analyses using kernel density estimates to estimate nest location and foraging distances for a large number of wild colonies of two species, and related these to the distribution of foraging habitats across an experimentally manipulated landscape. Mean foraging distances were 755 m for Bombus lapidarius and 775 m for B. pascuorum (using our most conservative estimation method). Colony‐specific foraging distances of both species varied with landscape structure, decreasing as the proportion of foraging habitats increased. This is the first time that foraging distance in wild bumblebees has been shown to vary with resource availability. Our method offers a means of estimating foraging distances in social insects, and informs the scale of management required to conserve bumblebee populations and enhance their pollination services across different landscapes.     

Allocation of Colony‐Level Foraging Effort in Vespula germanica in Response to Food Resource Quantity,Quality, and Associated Olfactory Cues

In social insects, selection takes place primarily at the level of the colony. Therefore, unlike solitary insects, social species are expected to forage at rates that maximize colony fitness rather than individual fitness. Workers can increase the net benefit of foraging by responding to increased resource availability, by responding more strongly to higher‐quality resources, and by decreasing the uncertainty with which nestmates find resources. Unlike many ants and social bees, no social wasp is known to utilize a nest‐based recruitment signal to inform nestmates of food location. On the other hand, wasps do learn the odor of food brought to the nest and use this cue to locate the food source outside the nest. Here, we quantify the effects of three food‐associated variables on the allocation of foraging effort in the yellowjacket Vespula germanica. We used an experimental approach to assess whether resource quantity, quality, or associated olfactory information affect the probability that a forager will leave the nest on a foraging trip. We addressed these questions by inserting a known amount of sucrose solution directly into nests and recording foraging effort (departure rate) over the subsequent hour‐long observation period. No differences were found in foraging effort because of the presence/absence of olfactory cues, but there was strong evidence that foraging effort increased in response to resource influx and resource quality. Thus, while olfactory cues are learned in the nest, only resource quality and the cue of increased amount of food in the nest factor into a forager's decision of whether or not to depart on a foraging trip. However, as prior work has shown, once a wasp forager leaves the nest, it uses the learned olfactory cues to aid in finding resources.     

Assessing the effectiveness of foraging radius models for seabird distributions using biotelemetry and survey data

Relatively simple foraging radius models have the potential to generate predictive distributions for a large number of species rapidly, thus providing a cost-effective alternative to large-scale surveys or complex modelling approaches. Their effectiveness, however, remains largely untested. Here we compare foraging radius distribution models for all breeding seabirds in Ireland, to distributions of empirical data collected from tracking studies and aerial surveys. At the local/colony level, we compared foraging radius distributions to GPS tracking data from seabirds with short (Atlantic puffin Fratercula arctica, and razorbill Alca torda) and long (Manx shearwater Puffinus puffinus, and European storm-petrel Hydrobates pelagicus) foraging ranges. At the regional/national level, we compared foraging radius distributions to extensive aerial surveys conducted over a two-year period. Foraging radius distributions were significantly positively correlated with tracking data for all species except Manx shearwater. Correlations between foraging radius distributions and aerial survey data were also significant, but generally weaker than those for tracking data. Correlations between foraging radius distributions and aerial survey data were benchmarked against generalised additive models (GAMs) of the aerial survey data that included a range of environmental covariates. While GAM distributions had slightly higher correlations with aerial survey data, the results highlight that the foraging radius approach can be a useful and pragmatic approach for assessing breeding distributions for many seabird species. The approach is likely to have acceptable utility in complex, temporally variable ecosystems and when logistic and financial resources are limited.     

Difference in Web Construction Behavior at Newly Occupied Web Sites Between Two Cyclosa Species

Animals make decisions based on subjective assessments of their environment. To determine their future foraging activities, animals probably assess food availability from past foraging experiences. Thus, foraging also functions as a way for animals to collect information, with the uncertainty of an assessment decreasing as foraging activity increases. This suggests that different needs for a correct assessment may affect the investment made in foraging activities. Orb‐web spiders sometimes relocate their webs and relocation rate differs among species. After web relocation, several spider species have been reported to construct the first webs at newly occupied web sites using less silk than usual, possibly to avoid the risk of an overinvestment at sites where food availability has not been determined. Nevertheless, they may pay a cost, because of inadequate decision‐making, if webs constructed with less silk convey less information and increase the uncertainty of an assessment. We expect that stronger site tenacity necessitates a greater requirement for correct assessment of web site and the degree to which spiders reduce the amount of web silk in the first web after web relocation is smaller in species that use the same site longer. To test this hypothesis, we examined web construction in two orb‐web spiders, Cyclosa octotuberculata and C. argenteoalba. At the same time we found that these two species exhibit different web‐site tenacity, as C. octotuberculata does not relocate its webs as frequently as does C. argenteoalba. After artificially induced web relocation, C. argenteoalba constructed webs that were initially smaller and contained only about 2/3 of the silk in control webs that were constructed at the original site. In contrast, C. octotuberculata did not exhibit such decreases in web size or in the amount of web silk used. This result is consistent with our hypothesis.     

东方田鼠社群气味对家族群成员个体觅食行为的影响

东方田鼠家族群成员个体的觅食行为是否因食物斑块存有家族群自身及非亲缘家族群气味而发生变异,进而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,分别配置家族群自身巢垫物及非亲缘家族群巢垫物作为社群气味,测定东方田鼠家族群在食物斑块觅食时,其成员个体觅食行为的序列过程及参数,检验家族群自身气味及非亲缘家族群气味对成员个体觅食行为的影响。结果表明,家族群自身气味能显著地缩短本群成员个体的觅食决定时间,通过减少成员个体的嗅闻及直立扫视动作时间比例、增大一般扫视、盯视及静听动作时间比例,降低觅食中断时间比例,提高其摄入率;而非亲缘家族群气味则能显著地延长家族群成员个体的觅食决定时间,通过增大家族群成员个体的嗅闻和一般扫视动作时间比例、减小直立扫视、盯视及静听动作时间比例,增大觅食中断时间比例,降低其摄入率。结果揭示,熟悉的社群气味会促使觅食活动中的家族群成员个体,在监测环境风险时,将精力更多地用于观察和监听群内其他成员个体的行为及其发出的警报信息,以便在有效规避环境风险的同时减缓个体间因干扰性竞争对觅食活动所造成的不利影响;而陌生的社群气味会迫使成员个体,将精力由依赖群内其他成员个体的行为转向凭借自身直接警觉周围环境。     

Sexual differences in the foraging ecology of 19th century beluga whales (Delphinapterus leucas) from the Canadian High Arctic

Marine mammals often exhibit significant sexual segregation in their diet and habitat use but these differences have not been studied systematically in historic or ancient populations due to the difficulties associated with determining the sex of skeletal elements based on gross morphology. Using a combined ancient DNA and stable isotope approach, we document a sexual difference in the foraging ecology of late 19th century beluga whales (Delphinapterus leucas) from the Canadian High Arctic. Using two PCR assays that coamplify fragments of the Y-linked SRY and X-linked ZFX genes, we assigned reproducible sex identities to 35 beluga specimens. This provided a basis for investigating sex-specific differences in foraging ecology using stable carbon and nitrogen isotope analyses of bone collagen. These isotopic data demonstrate that although both males and females primarily consumed Arctic cod, males utilized a wider range of prey than females, feeding on high trophic level benthic prey (sculpins) to a greater extent. Because bone collagen integrates prey isotopic compositions over the course of several years these sex-based differences in beluga bone collagen isotopic compositions reflect long-term and sustained sexual differences in foraging.     

The behavioral context of common dolphin (Delphinus sp.) vocalizations

Correlations between surface behavior and concurrent underwater vocalizations were modeled for common dolphins (Delphinus spp.) in the Southern California Bight (SCB) over multiple field seasons. Clicks, pulsed calls, and whistles were examined, with a total of 50 call features identified. Call features were used to classify behavior using random forest decision trees, with rates of correct classification reaching 80.6% for fast travel, 84.6% for moderate travel, 59.8% for slow travel, and 58% for foraging behavior. Common dolphins spent most of their time traveling. The highest number of clicks, pulsed calls, and complex whistles were produced during fast travel. In contrast, during foraging there were few pulsed calls and whistles produced, and the whistles were simple with narrow bandwidths and few harmonics. Behavior and vocalization patterns suggest nocturnal foraging in offshore waters as the primary feeding strategy. Group size and spacing were strongly correlated with behavior and rates of calling, with higher call rates in dispersed traveling groups and lower call rates in loosely aggregated foraging groups. These results demonstrate that surface behavior can be classified using vocalization data, which builds the framework for behavioral studies of common dolphins using passive acoustic monitoring techniques.     

Why do honeybees reject certain flowers?

Summary Honeybees often approach flowers of Lotus corniculatus and then fly away without attempting to extract nectar. These rejected flowers contained 41% less nectar than my random sample. The accepted flowers contained 24% more nectar than my random sample. The differences among these three flower-groups were due to differences in the percent of empty flowers in each group rather than the differences in the absolute amount of nectar. Honeybees increased their foraging efficiency by accepting less empty flowers and rejecting more empty flowers than would be expected if they foraged randomly. There are two possible mechanisms for this discrimination-behavior: either the bees are smelling nectar odor or they are smelling bee scent left by previous visitors to the flower. My results are inconsistent with the hypothesis that bees are basing their decision on nectar smell and suggest that they are using bee scent as a means of identifying empty flowers.     

Density-dependent prey behaviours and mutable predator foraging modes induce Allee effects and over-prediction of prey mortality rates

1. Predator–prey models are often used to represent consumptive interactions between species but, typically, are derived using simple experimental systems with little plasticity in prey or predator behaviours. However, many prey and predators exhibit a broad suite of behaviours. Here, we experimentally tested the effect of density-dependent prey and predator behaviours on per capita relative mortality rates using Florida bass (Micropterus floridanus) consuming juvenile Bluegill (Lepomis macrochirus).
2. Experimental ponds were stocked with a factorial design of low, medium, and high prey and predator densities. Prey mortality, prey–predator behaviours, and predator stomach contents were recorded over or after 7 days. We assumed the mortality dynamics followed foraging arena theory. This pathologically flexible predator–prey model separates prey into invulnerable and vulnerable pools where predators can consume prey in the latter. As this approach can represent classic Lotka–Volterra and ratio-dependent dynamics, we fit a foraging arena predator–prey model to the number of surviving prey.
3. We found that prey exhibited density-dependent prey behaviours, hiding at low densities, shoaling at medium densities, and using a provided refuge at high densities. Predators exhibited ratio-dependent behaviours, using an ambush foraging mode when one predator was present, hiding in the shadows at low prey–high predator densities, and shoaling at medium and high prey–high predator densities. The foraging arena model predicted the mortality rates well until the high prey–high predator treatment where group vigilance prey behaviours occurred and predators probably interfered with one another resulting in the model predicting higher mortality than observed.
4. This is concerning given the ubiquity of predator–prey models in ecology and natural resource management. Furthermore, as Allee effects engender instability in population regulation, it could lead to inaccurate predictions of conservation status, population rebuilding or harvest rates.

     

Optimal foraging: A case for random movement

Summary The bumblebee, Bombus flavifrons, forages randomly with respect to direction on Polemonium foliosissimum. This foraging pattern is as predicted for a system where there is a low probability of revisiting any given flower upon returning to a patch. This low revisitation probability is a function of the floral resource arrangement. It is further shown that B. flavifrons is using the resource distribution to direct its movements. A large percentage of all movements are to nearest neighbors with maximal foraging efficiency gained through minimization of flight distances.     

东方田鼠家族群对成员个体觅食行为的影响

植食性哺乳动物在分享社群觅食带来好处的同时,是否因个体间的相互干扰而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,测定东方田鼠家族群成员个体在食物斑块上的觅食行为序列过程及行为参数,检验家族群存在对成员个体觅食行为的影响。结果发现,东方田鼠家族群雌、雄成员个体的觅食行为参数均无显著差异。然而,与单只个体相比,家族群觅食尽管能显著地缩短成员个体的觅食决定时间,但却显著地降低了成员个体的摄入率。分析觅食行为参数觅食中断时间发现,相较于单只个体,家族群成员个体间因相互干扰而引起的觅食中断时间的增加,不但增大了收获每口食物的时间,而且导致其摄入率下降。检测家族群成员个体各警觉行为动作参数,发现,成员个体间的相互干扰能引致个体的一般扫视、盯视及嗅闻动作时间比例显著增大,尽管直立扫视和静听动作时间比例减少显著,但并未使个体的觅食中断时间减小。结果充分说明,东方田鼠家族群成员个体间的相互干扰能使个体觅食行为参数发生变异,导致觅食中断时间增加,摄入率降低。     

Individual differences in foraging site fidelity are not related to time-activity budgets in Herring Gulls

Many populations consist of individuals that differ consistently in their foraging behaviour through resource or foraging site selection. Foraging site fidelity has been reported in several seabird species as a common phenomenon. It is considered especially beneficial in spatially and/or temporally predictable environments in which fidelity is thought to increase energy intake, thereby affecting time-energy budgets. However, the consequences for activity and energy budget have not been adequately tested. In this paper, we studied the consequences of fine-scale foraging site fidelity in adult Herring Gulls Larus argentatus in a highly predictable foraging environment with distinct foraging patches. We measured their time-activity budgets using GPS tracking and tri-axial acceleration measurements, which also made it possible to estimate energy expenditure. Individual variation in foraging site fidelity was high, some individuals spending most of their time on a single foraging patch and others spending the same amount of time in up to 21 patches. While time and activity budgets differed between individuals, we found no clear relationship with foraging site fidelity. We did find a relationship between the size of the birds and the level of site fidelity; faithful birds tend to have a larger body size. Although differences in foraging time and habitat use between individuals could play a role in the results of the current study, short-term consequences of variation in foraging site fidelity within a population remain elusive, even when focusing on individuals with a similar foraging specialization (Blue Mussels Mytilus edulis). Studying individuals over multiple years and under varying environmental conditions may provide better insight into the consequences and plasticity of foraging site fidelity.