Effects of size,caudal autotomy,and predator kairomones on the foraging behavior of Allegheny Mountain dusky salamanders (<Emphasis Type="Italic">Desmognathus ochrophaeus</Emphasis>)

Prey must balance the conflicting demands of foraging and defensive behavior. Foraging under the threat of predation may be further complicated among species that engage in caudal autotomy, the loss of a portion of the tail at preformed breakage planes, because the tail may serve as an important energy storage organ and contribute to motility, culminating in a trade-off between foraging and predator avoidance. As a result of the advantages conferred by the presence of a tail, individuals that have recently undergone autotomy may be more motivated to forage despite elevated levels of threat indicated by predator kairomones. We used a full factorial design to evaluate the combined effects of body size, exposure to predator kairomones, and experience with autotomy on the latency to strike at Drosophila prey, number of strikes, and prey captured per strike by Allegheny Mountain dusky salamanders (Desmognathus ochrophaeus). In our study, caudal autotomy was the only significant main effect and influenced both the latency to attack prey and the number of strikes attempted. In terms of latency to attack prey, there was a significant interaction between body size and autotomy such that “small” salamanders (≤3.2 cm SVL) without tails delayed their foraging behavior. In terms of the number of strikes toward prey, there was a significant interaction between autotomy and exposure to predator kairomones such that individuals with intact tails exhibited a greater number of strikes, with the exception of the “large” (>3.2 cm SVL) salamanders, which performed fewer strikes when exposed to the snake kairomones. There was no significant effect on foraging efficiency, although the trend in the data suggests that autotomized individuals forage more efficiently. This study was designed to evaluate the confluence of factors related to size, caudal autotomy, and exposure to stimuli from predators and hints at the magnitude of caudal autotomy on antipredator decision-making. Our data suggest that despite the importance of tail tissue for energy storage, locomotion, and mating, salamanders without tails are cautious when foraging under elevated risk.     

Modeling optimal responses and fitness consequences in a changing Arctic

Animals must balance a series of costs and benefits while trying to maximize their fitness. For example, an individual may need to choose how much energy to allocate to reproduction versus growth, or how much time to spend on vigilance versus foraging. Their decisions depend on complex interactions between environmental conditions, behavioral plasticity, reproductive biology, and energetic demands. As animals respond to novel environmental conditions caused by climate change, the optimal decisions may shift. Stochastic dynamic programming provides a flexible modeling framework with which to explore these trade‐offs, but this method has not yet been used to study possible changes in optimal trade‐offs caused by climate change. We created a stochastic dynamic programming model capturing trade‐off decisions required by an individual adult female polar bear (Ursus maritimus) as well as the fitness consequences of her decisions. We predicted optimal foraging decisions throughout her lifetime as well as the energetic thresholds below which it is optimal for her to abandon a reproductive attempt. To explore the effects of climate change, we shortened the spring feeding period by up to 3 weeks, which led to predictions of riskier foraging behavior and higher reproductive thresholds. The resulting changes in fitness may be interpreted as a best‐case scenario, where bears adapt instantaneously and optimally to new environmental conditions. If the spring feeding period was reduced by 1 week, her expected fitness declined by 15%, and if reduced by 3 weeks, expected fitness declined by 68%. This demonstrates an effective way to explore a species' optimal response to a changing landscape of costs and benefits and highlights the fact that small annual effects can result in large cumulative changes in expected lifetime fitness.     

Linking predator risk and uncertainty to adaptive forgetting: a theoretical framework and empirical test using tadpoles

Hundreds of studies have examined how prey animals assess their risk of predation. These studies work from the basic tennet that prey need to continually balance the conflicting demands of predator avoidance with activities such as foraging and reproduction. The information that animals gain regarding local predation risk is most often learned. Yet, the concept of ‘memory’ in the context of predation remains virtually unexplored. Here, our goal was (i) to determine if the memory window associated with predator recognition is fixed or flexible and, if it is flexible, (ii) to identify which factors affect the length of this window and in which ways. We performed an experiment on larval wood frogs, Rana sylvatica, to test whether the risk posed by, and the uncertainty associated with, the predator would affect the length of the tadpoles'' memory window. We found that as the risk associated with the predator increases, tadpoles retained predator-related information for longer. Moreover, if the uncertainty about predator-related information increases, then prey use this information for a shorter period. We also present a theoretical framework aiming at highlighting both intrinsic and extrinsic factors that could affect the memory window of information use by prey individuals.     

Shifting the balance between foraging and predator avoidance: the importance of food distribution for a schooling pelagic forager

It is widely held that when predator avoidance conflicts with other activities, such as feeding, avoidance of predators often takes precedence. In this study, we examine how predation risk and food distribution interact to influence the schooling behavior and swimming speed of foraging juvenile walleye pollock, Theragra chalcogramma. Fish were acclimated to either spatially and temporally clumped, or spatially and temporally dispersed food for 3 weeks. Fish were then monitored while feeding in the absence and presence of predatory sablefish, Anoplopoma fimbria. Fish foraging for clumped food swam rapidly in a loose school when predators were absent, but swam more slowly and adopted more cohesive schooling in the presence of predators, trading-off foraging opportunity for decreased vulnerability to predators. Fish foraging for dispersed food swam about slowly and did not engage in cohesive schooling in either the absence or presence of predators. These fish accepted greater predation risk in order to continue foraging, suggesting that the cost of schooling, in terms of decreased foraging opportunity, was greater when food was dispersed than when it was clumped. This lower responsiveness to predators among fish receiving dispersed food demonstrates that predator avoidance does not always take precedence over other activities, but rather, that a balance is maintained between predator avoidance and feeding, which shifts as food distribution changes.     

Testing the Threat‐Sensitive Hypothesis with Predator Familiarity and Dietary Specificity

In a system with multiple predators, the threat‐sensitive predator avoidance hypothesis predicts that prey respond differently to predators relative to the risks each poses (e.g., degree of dietary specialization). Aquatic animals often rely heavily on detecting predators via chemical cues (kairomones) and respond with a suite of behaviors including detection and avoidance. However, little is known about how animals respond to kairomones of specialist versus generalist predators. In laboratory experiments, we compared behavioral responses of a poorly studied aquatic salamander, the greater siren (Siren lacertina), to cues from specialist and generalist predator snakes to evaluate threat‐sensitive responses. Sirens exhibited a novel behavior (gill‐flushing) most often in the presence of specialist predator cues. Avoidance behavior (reversing direction following cue detection) was higher in response to specialist predator and novel animal control cues and lowest in response to generalist predator cues. An intermediate response to the animal control, an unfamiliar amphibian predator, indicated that sirens respond cautiously to a novel cue. The gradient of observed responses to different snake cues indicates that sirens may be evaluating predation potential of animals based on their foraging specificity and familiarity.     

The Effect of Feeder Hotspots on the Predictability and Home Range Use of a Small Bird in Winter

Few studies address how resources and predation risk affect movement patterns and the overall spatial use of prey species. Although movement is generally considered to be dangerous, at large scales, movement may be important for predator avoidance and the predictability of such movement may be key. We examine the movement patterns of a small bird (Junco hyemalis) in winter to better understand how these birds might respond to the trade‐off of unpredictable movements for predator avoidance with the foraging benefits of visiting large, predictable food sources. We manipulated resources by adding feeders to junco home ranges and compared the movement patterns of these flocks to those without access to feeders. Juncos with access to feeders were more spatially and temporally predictable, had reduced movement rates and smaller home range sizes. Our results suggest that the influence of resource distribution on junco movements is high. Juncos with highly productive and predictable resource hotspots may place more value on resources than remaining unpredictable. Consequently, they may be employing non‐movement methods of anti‐predator behavior, such as vigilance, at feeders, although this requires further investigation.     

Foraging time allocation in relation to sex by the gulf coast fiddler crab (Uca panacea)

Summary Schoener (1971) proposed that the reproductive demands of animals should be important in shaping their foraging behavior because fitness is affected. He defined two forager types: energy maximizers (reproductive success depends on energetic intake) and time minimizers (reproductive success depends on time spent in activities other than foraging), and suggested that females most often illustrate the former and males the latter. We tested whether mating activities influence the foraging behavior of Uca panacea, and the predictions that females would be energy maximizers because of their reproductive strategy and that males would also be energy maximizers because of their courtship activity. Time allocated to foraging by 800 male and female fiddler crabs (at two sites) was quantified; no significant difference in foraging time was found between the sexes. Both male and female crabs allotted a large portion of their time to foraging because both sexes depend on stored energy during their reproductive bouts. Our results show that the particular forager type can be predicted based on reproductive demands, but a forager type can not always be assigned to a particular sex without consideration of all important ecological and physiological factors determining reproductive success.     

Food for thought: Harbor porpoise foraging behavior and diet inform vulnerability to disturbance

Research has shown that noise disturbance can disrupt the behavior of harbor porpoises. The significance of such disturbance is unclear. However, these animals may be vulnerable to starvation when disturbed due to their high energy requirements. Important parameters determining harbor porpoise energy balance are the size and energy content of prey, their foraging behavior and their energetic requirements for homeostasis, growth, and reproduction. Energy intake can be estimated using published data from tagged animals. Such analysis indicates a broad range of plausible levels of energy intake, in line with those from captive studies. Metabolizable energy intake estimates were most strongly affected by variations in target prey size and to a lesser extent, by the foraging intensity of porpoises. In all but the worst case scenarios, harbor porpoises are well equipped for their ecological niche due to their generalist diet, consisting of a range of moderate to high energy-density prey combined with ultra-high foraging rates and high capture success. If animals can find suitable prey, porpoises may be capable of recovering from some lost foraging opportunities. Minimizing disturbances is, however, important for their health. Further research into prey and the environment are required to fully test the assumption of vulnerability.     

Seeing is believing: information content and behavioural response to visual and chemical cues

Predator avoidance and foraging often pose conflicting demands. Animals can decrease mortality risk searching for predators, but searching decreases foraging time and hence intake. We used this principle to investigate how prey should use information to detect, assess and respond to predation risk from an optimal foraging perspective. A mathematical model showed that solitary bees should increase flower examination time in response to predator cues and that the rate of false alarms should be negatively correlated with the relative value of the flower explored. The predatory ant, Oecophylla smaragdina, and the harmless ant, Polyrhachis dives, differ in the profile of volatiles they emit and in their visual appearance. As predicted, the solitary bee Nomia strigata spent more time examining virgin flowers in presence of predator cues than in their absence. Furthermore, the proportion of flowers rejected decreased from morning to noon, as the relative value of virgin flowers increased. In addition, bees responded differently to visual and chemical cues. While chemical cues induced bees to search around flowers, bees detecting visual cues hovered in front of them. These strategies may allow prey to identify the nature of visual cues and to locate the source of chemical cues.     

Moose movement rates are altered by wolf presence in two ecosystems

Predators directly impact prey populations through lethal encounters, but understanding nonlethal, indirect effects is also critical because foraging animals often face trade‐offs between predator avoidance and energy intake. Quantifying these indirect effects can be difficult even when it is possible to monitor individuals that regularly interact. Our goal was to understand how movement and resource selection of a predator (wolves; Canis lupus) influence the movement behavior of a prey species (moose; Alces alces). We tested whether moose avoided areas with high predicted wolf resource use in two study areas with differing prey compositions, whether avoidance patterns varied seasonally, and whether daily activity budgets of moose and wolves aligned temporally. We deployed GPS collars on both species at two sites in northern Minnesota. We created seasonal resource selection functions (RSF) for wolves and modeled the relationship between moose first‐passage time (FPT), a method that discerns alterations in movement rates, and wolf RSF values. Larger FPT values suggest rest/foraging, whereas shorter FPT values indicate travel/fleeing. We found that the movements of moose and wolves peaked at similar times of day in both study areas. Moose FPTs were 45% lower in areas most selected for by wolves relative to those avoided. The relationship between wolf RSF and moose FPT was nonlinear and varied seasonally. Differences in FPT between low and high RSF values were greatest in winter (?82.1%) and spring (?57.6%) in northeastern Minnesota and similar for all seasons in the Voyageurs National Park ecosystem. In northeastern Minnesota, where moose comprise a larger percentage of wolf diet, the relationship between moose FPT and wolf RSF was more pronounced (ave. across seasons: ?60.1%) than the Voyageurs National Park ecosystem (?30.4%). These findings highlight the role wolves can play in determining moose behavior, whereby moose spend less time in areas with higher predicted likelihood of wolf resource selection.     

Climate change enhances the negative effects of predation risk on an intermediate consumer

Predators are a major source of stress in natural systems because their prey must balance the benefits of feeding with the risk of being eaten. Although this ‘fear’ of being eaten often drives the organization and dynamics of many natural systems, we know little about how such risk effects will be altered by climate change. Here, we examined the interactive consequences of predator avoidance and projected climate warming in a three‐level rocky intertidal food chain. We found that both predation risk and increased air and sea temperatures suppressed the foraging of prey in the middle trophic level, suggesting that warming may further enhance the top‐down control of predators on communities. Prey growth efficiency, which measures the efficiency of energy transfer between trophic levels, became negative when prey were subjected to predation risk and warming. Thus, the combined effects of these stressors may represent an important tipping point for individual fitness and the efficiency of energy transfer in natural food chains. In contrast, we detected no adverse effects of warming on the top predator and the basal resources. Hence, the consequences of projected warming may be particularly challenging for intermediate consumers residing in food chains where risk dominates predator‐prey interactions.     

Thermal and digestive constraints to foraging behaviour in marine mammals

While foraging models of terrestrial mammals are concerned primarily with optimizing time/energy budgets, models of foraging behaviour in marine mammals have been primarily concerned with physiological constraints. This has historically centred on calculations of aerobic dive limits. However, other physiological limits are key to forming foraging behaviour, including digestive limitations to food intake and thermoregulation. The ability of an animal to consume sufficient prey to meet its energy requirements is partly determined by its ability to acquire prey (limited by available foraging time, diving capabilities and thermoregulatory costs) and process that prey (limited by maximum digestion capacity and the time devoted to digestion). Failure to consume sufficient prey will have feedback effects on foraging, thermoregulation and digestive capacity through several interacting avenues. Energy deficits will be met through catabolism of tissues, principally the hypodermal lipid layer. Depletion of this blubber layer can affect both buoyancy and gait, increasing the costs and decreasing the efficiency of subsequent foraging attempts. Depletion of the insulative blubber layer may also increase thermoregulatory costs, which will decrease the foraging abilities through higher metabolic overheads. Thus, an energy deficit may lead to a downward spiral of increased tissue catabolism to pay for increased energy costs. Conversely, the heat generated through digestion and foraging activity may help to offset thermoregulatory costs. Finally, the circulatory demands of diving, thermoregulation and digestion may be mutually incompatible. This may force animals to alter time budgets to balance these exclusive demands. Analysis of these interacting processes will lead to a greater understanding of the physiological constraints within which the foraging behaviour must operate.     

Optimal foraging for specific nutrients in predatory beetles

Evolutionary theory predicts that animals should forage to maximize their fitness, which in predators is traditionally assumed equivalent to maximizing energy intake rather than balancing the intake of specific nutrients. We restricted female predatory ground beetles (Anchomenus dorsalis) to one of a range of diets varying in lipid and protein content, and showed that total egg production peaked at a target intake of both nutrients. Other beetles given a choice to feed from two diets differing only in protein and lipid composition selectively ingested nutrient combinations at this target intake. When restricted to nutritionally imbalanced diets, beetles balanced the over- and under-ingestion of lipid and protein around a nutrient composition that maximized egg production under those constrained circumstances. Selective foraging for specific nutrients in this predator thus maximizes its reproductive performance. Our findings have implications for predator foraging behaviour and in the structuring of ecological communities.     

When to Come Out from a Refuge: Balancing Predation Risk and Foraging Opportunities in an Alpine Lizard

Prey often respond to predator presence by increasing their use of refuges, but because this strategy may be costly, the decision regarding when to come out from a refuge should be optimized. The loss of foraging opportunities may be one of the main costs when safer microhabitats (i.e. refuges) are also the poorest in terms of their foraging profitability. We present the results of an experimental field study to test whether emergence times from a refuge of the Iberian rock lizard, Lacerta monticola, vary as a function of expected foraging opportunities and level of satiation of the lizard. As predicted, short‐term fluctuations in availability of food influenced emergence times; when a lizard had just detected some food in the recent past, emergence times decreased greatly, because the loss of opportunities for foraging increased costs of refuge use. Furthermore, the characteristics and success of the encounter with food, nutritional state of lizards, and the added possibility of capturing new food items influenced the duration of hiding times. Therefore, foraging requirements and avoidance of predators may be conflicting demands that L. monticola lizards balance by modifying the duration of time spent in refuges.     

Does group foraging promote efficient exploitation of resources?

Increased avoidance of food patches previously exploited by other companions has been proposed as one adaptive benefit of group foraging. However, does group foraging really represent the most efficient way to exploit non- or slowly-renewing resources? Here, I used simulations to explore the costs and benefits of exploiting non-renewing resources by foragers searching for food patches independently or in groups in habitats with different types of resource distribution. Group foragers exploited resources in a patch more quickly and therefore spent proportionately more time locating new patches. Reduced avoidance of areas already exploited by others failed to overcome the increased time cost of searching for new food patches and group foragers thus obtained food at a lower rate than solitary foragers. Group foraging provided one advantage in terms of a reduction in the variance of food intake rate. On its own, reduced avoidance of exploitation competition through group foraging appears unlikely to increase mean food intake rate when exploiting non-renewing patches but may provide a way to reduce the risk of an energy shortfall.     

The effects of mandible morphology and photosynthetic pathway on selective herbivory in grasshoppers

Summary In the present study the following hypotheses were tested: 1) Selective herbivory is influenced by the photosynthetic pathway of plants, and 2) selective herbivory is influenced by the mandible morphology of herbivores. To test these hypotheses, grasshoppers with different mandibular patterns were offered C₃ and C₄ plants in controlled feeding experiments.Mandible morphology was found to be significantly associated with energy assimilation and feeding frequency. Grasshoppers with herbivorous mandibles had greater assimilation efficiencies and fed significantly less often than grasshoppers with forbivorous mandibles. These results indicate grasshoppers (with comparable caloric requirements) feed with a frequency determined, in part, by the functional morphology of their mandibles.Photosynthetic pathway also influenced the foraging behavior of grasshoppers tested. As predicted by the C₄ avoidance hypothesis (Caswell et al. 1973), C₄ plants were fed upon significantly less often than C₃ plants. These results suggest grasshoppers may discriminate between C₃ and C₄ plants, but apparently not on the basis of available energy.Recent studies of insect herbivory, previously interpreted within the framework of C₄ plant avoidance, are reviewed. The results of these studies may be critically dependent on incomplete considerations of mandible morphology and photosynthetic pathway.This study is contribution 92 of the Chihuahuan Desert Research Institute     

Evolution of prey behavior in response to changes in predation regime: damselflies in fish and dragonfly lakes

In a large behavioral experiment we reconstructed the evolution of behavioral responses to predators to explore how interactions with predators have shaped the evolution of their prey's behavior. All Enallagma damselfly species reduced both movement and feeding in the presence of coexisting predators. Some Enallagma species inhabit water bodies with both fish and dragonflies, and these species responded to the presence of both predators, whereas other Enallagma species inhabit water bodies that have only large dragonflies as predators, and these species only responded to the presence of dragonflies. Lineages that shifted to live with large dragonflies showed no evolution in behaviors expressed in the presence of dragonflies, but they evolved greater movement in the absence of predators and greater movement and feeding in the presence of fish. These results suggest that Enallagma species have evolutionarily lost the ability to recognize fish as a predator. Because species coexisting with only dragonfly predators have also evolved the ability to escape attacking dragonfly predators by swimming, the decreased predation risk associated with foraging appears to have shifted the balance of the foraging/predation risk trade-off to allow increased activity in the absence of mortality threats to evolve in these lineages. Our results suggest that evolution in response to changes in predation regime may have greater consequences for characters expressed in the absence of mortality threats because of how the balance between the conflicting demands of growth and predation risk are altered.     

Balancing Energy Budget in a Central-Place Forager: Which Habitat to Select in a Heterogeneous Environment?

Foraging animals are influenced by the distribution of food resources and predation risk that both vary in space and time. These constraints likely shape trade-offs involving time, energy, nutrition, and predator avoidance leading to a sequence of locations visited by individuals. According to the marginal-value theorem (MVT), a central-place forager must either increase load size or energy content when foraging farther from their central place. Although such a decision rule has the potential to shape movement and habitat selection patterns, few studies have addressed the mechanisms underlying habitat use at the landscape scale. Our objective was therefore to determine how Ring-billed gulls (Larus delawarensis) select their foraging habitats while nesting in a colony located in a heterogeneous landscape. Based on locations obtained by fine-scale GPS tracking, we used resource selection functions (RSFs) and residence time analyses to identify habitats selected by gulls for foraging during the incubation and brood rearing periods. We then combined this information to gull survey data, feeding rates, stomach contents, and calorimetric analyses to assess potential trade-offs. Throughout the breeding season, gulls selected landfills and transhipment sites that provided higher mean energy intake than agricultural lands or riparian habitats. They used landfills located farther from the colony where no deterrence program had been implemented but avoided those located closer where deterrence measures took place. On the other hand, gulls selected intensively cultured lands located relatively close to the colony during incubation. The number of gulls was then greater in fields covered by bare soil and peaked during soil preparation and seed sowing, which greatly increase food availability. Breeding Ring-billed gulls thus select habitats according to both their foraging profitability and distance from their nest while accounting for predation risk. This supports the predictions of the MVT for central-place foraging over large spatial scales.     

Relation between complexity and stability in food webs with adaptive behavior

We investigate the influence of functional responses (Lotka-Volterra or Holling type), initial topological web structure (randomly connected or niche model), adaptive behavior (adaptive foraging and predator avoidance) and the type of constraints on the adaptive behavior (linear or nonlinear) on the stability and structure of food webs. Two kinds of stability are considered: one is the network robustness (i.e., the proportion of species surviving after population dynamics) and the other is the species deletion stability. When evaluating the network structure, we consider link density as well as the trophic level structure. We show that the types of functional responses and initial web structure do not have a large effect on the stability of food webs, but foraging behavior has a large stabilizing effect. It leads to a positive complexity-stability relationship whenever higher "complexity" implies more potential prey per species. The other type of adaptive behavior, predator avoidance behavior, makes food webs only slightly more stable. The observed link density after population dynamics depends strongly on the presence or absence of adaptive foraging, and on the type of constraints used. We also show that the trophic level structure is preserved under population dynamics with adaptive foraging.