Predicted fitness consequences of threat-sensitive hiding behavior

In studies of refuge use as a form of antipredator behavior,where prey hide in response to a predator's approach, factorssuch as foraging costs and the perceived risk in a predator'sapproach have been shown to influence the hiding behavior ofprey. Because few studies of waiting games have focused on mammals,we studied the hiding behavior of the yellow-bellied marmot(Marmota flaviventris), a ground-dwelling rodent. We testedthe prediction that marmots vary hiding time as a function ofpredator approach speed and presence and absence of food outsidetheir refuge and that marmots hide differently depending ontheir relative condition. We conducted "fast approaches" and"slow approaches" in the presence and absence of extra foodand evaluated hiding times. Multiple regression analyses demonstratedthat the interaction between the approach speed and the presenceand absence of food influenced hiding behavior; body conditionhad a smaller, but nonsignificant effect. We then developeda state-dependent dynamic model to explore potential fitnessconsequences of these decisions. The model suggested that theoverall survival of a population is substantially reduced whenindividuals make suboptimal decisions. Our research builds onprevious studies, indicating that animals integrate both costsand benefits of hiding when determining their hiding times.     

To feed or not to feed? Bioenergetic impacts of fear‐driven behaviors in lactating dolphins

In mammals, lactation can be the most energetically expensive part of the reproductive cycle. Thus, when energy needs are compromised due to predation risk, environmental disturbance, or resource scarcity, future reproductive success can be impacted. In marine and terrestrial environments, foraging behavior is inextricably linked to predation risk. But quantification of foraging energetics for lactating animals under predation risk is less understood. In this study, we used a spatially explicit individual‐based model to study how changes in physiology (lactating or not) and the environment (predation risk) affect optimal behavior in dolphins. Specifically, we predicted that an adult dolphin without calf would incur lower relative energetic costs compared to a lactating dolphin with calf regardless of predation risk severity, antipredator behavior, or prey quality consumed. Under this state‐dependent analysis of risk approach, we found predation risk to be a stronger driver in affecting total energetic costs (foraging plus locomotor costs) than food quality for both dolphin types. Further, contrary to our hypothesis, after accounting for raised energy demands, a lactating dolphin with calf does not necessarily have higher relative‐to‐baseline costs than a dolphin without calf. Our results indicate that both a lactating (with calf) and non‐lactating dolphin incur lowered energetic costs under a risk‐averse behavioral scheme, but consequently suffer from lost foraging calories. A lactating dolphin with calf could be particularly worse off in lost foraging calories under elevated predation risk, heightened vigilance, and increased hiding time relative to an adult dolphin without calf. Further, hiding time in refuge could be more consequential than detection distance for both dolphin types in estimated costs and losses incurred. In conclusion, our study found that reproductive status is an important consideration in analyzing risk effects in mammals, especially in animals with lengthy lactation periods and those exposed to both biological and nonbiological stressors.     

Spine and dine: A key defensive trait promotes ecological success in spiny ants

A key focus of ecologists is explaining the origin and maintenance of morphological diversity and its association with ecological success. We investigate potential benefits and costs of a common and varied morphological trait, cuticular spines, for foraging behavior, interspecific competition, and predator–prey interactions in naturally co‐occurring spiny ants (Hymenoptera: Formicidae: Polyrhachis) in an experimental setting. We expect that a defensive trait like spines might be associated with more conspicuous foraging, a greater number of workers sent out to forage, and potentially increased competitive ability. Alternatively, consistent with the ecological trade‐off hypothesis, we expect that investment in spines for antipredator defense might be negatively correlated with these other ecological traits. We find little evidence for any costs to ecological traits, instead finding that species with longer spines either outperform or do not differ from species with shorter spines for all tested metrics, including resource discovery rate and foraging effort as well as competitive ability and antipredator defense. Spines appear to confer broad antipredator benefits and serve as a form of defense with undetectable costs to key ecological abilities like resource foraging and competitive ability, providing an explanation for both the ecological success of the study genus and the large number of evolutionary origins of this trait across all ants. This study also provides a rare quantitative empirical test of ecological effects related to a morphological trait in ants.     

Population dynamics of prey exhibiting inducible defenses: the role of associated costs and density-dependence

The effect of antipredator behavior on the dynamics of a resource-consumer model was analyzed in relation to the magnitude of associated costs, and the strength of density-dependence. For this purpose, I present a deterministic continuous resource-consumer model that exhibits biomass conversion, structural homogeneity, and competition for renewable and fixed resources as separate processes. Antipredator behavior is incorporated as an inducible response to consumer density, and has metabolic and feeding costs. By means of numerical methods, I show: (1) that antipredator behavior is stabilizing for certain parameter ranges, where other stabilizing forces do not dominate the dynamics; (2) intraspecific competition for both fixed and renewable resources have a stabilizing role; (3) metabolic cost is always stabilizing, and feeding cost can be stabilizing or destabilizing, depending on the relative strength of the two competition forces.     

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.     

Optimal traits when there are several costs: the interaction of mortality and energy costs in determining foraging behavior

I analyzed the interaction of different types of costs in determiningoptimal behavior using mathematical models. The analysis concentrateson foraging behavior and asks (1) whether the cost factor thathas the greatest effect on fitness generally has the greatesteffect on optimal trait values and (2) whether increasing thesize of one type of cost makes the optimal behavior absolutelyor relatively more sensitive to that cost. The foraging costsconsidered are energy expenditure, predation risk, and othermortality factors. It is shown that increasing the magnitudeof one cost often decreases the relative and absolute sensitivityof the optimal foraging strategy to that cost. The relativefitness effects of different costs generally differ from therelative sensitivities of the optimal strategies to the costfactors. Researchers should therefore measure the shapes ofcost curves rather than their average magnitudes to determinewhich of several costs can be ignored in cost-benefit analyses.     

Numerical bifurcation analysis of distance-dependent on-center off-surround shunting neural networks

 On-center off-surround shunting neural networks are often applied as models for content-addressable memory (CAM), the equilibria being the stored memories. One important demand of biological plausible CAMs is that they function under a broad range of parameters, since several parameters vary due to postnatal maturation or learning. Ellias, Cohen and Grossberg have put much effort into showing the stability properties of several configurations of on-center off-surround shunting neural networks. In this article we present numerical bifurcation analysis of distance-dependent on-center off-surround shunting neural networks with fixed external input. We varied four parameters that may be subject to postnatal maturation: the range of both excitatory and inhibitory connections and the strength of both inhibitory and excitatory connections. These analyses show that fold bifurcations occur in the equilibrium behavior of the network by variation of all four parameters. The most important result is that the number of activation peaks in the equilibrium behavior varies from one to many if the range of inhibitory connections is decreased. Moreover, under a broad range of the parameters the stability of the network is maintained. The examined network is implemented in an ART network, Exact ART, where it functions as the classification layer F2. The stability of the ART network with the F2-field in different dynamic regimes is maintained and the behavior is functional in Exact ART. Through a bifurcation the learning behavior of Exact ART may even change from forming local representations to forming distributed representations. Received: 23 January 1996 / Accepted in revised form: 1 July 1996     

Combined effects of chemical and visual information in eliciting antipredator behaviour in juvenile Atlantic salmon Salmo salar

Under natural conditions, both young-of-the-year (YOY; 0+ year) and parr (1+ year) Atlantic salmon Salmo salar exhibited strong antipredator behaviour ( e.g. increase in latency to resume foraging) following the exposure to damage-released chemical alarm cues relative to a stream water control. Subsequent exposure to a novel visual stimulus had contrasting results. Parr increased their reactive distance to the visual stimulus if they had been previously exposed to a chemical alarm cue, whereas YOY did not. On the other hand, both YOY and parr took significantly longer to resume foraging when exposed to a visual stimulus if they had been previously exposed to a chemical alarm cue than control groups. While YOY and parr differed in the type and intensity of antipredator responses to both chemical and visual stimuli, perhaps due to differential costs and benefits associated with age, both used the chemical and the visual information in a combined manner.     

Antipredator responses of aphids to parasitoids change as a function of aphid physiological state

Under predation risk, prey may prioritize antipredator behaviours and sacrifice feeding. However, energetically constrained animals may choose to sacrifice or change antipredator responses and accept relatively greater risk in order to secure food. In this last case, the antipredator tactics chosen must balance safety and feeding in such a way that costs are minimized and benefits maximized. We studied the antipredator behaviour of pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae) subjected to different periods of food deprivation, against the parasitoid Aphidius ervi (Hymenoptera: Braconidae). As the energetic internal stress of aphids increased, the predominant antipredator response changed from walking away and dropping to kicking behaviour, and parasitization avoidance decreased. Parasitoids did not show preference between food-deprived and nonfood-deprived aphids. Dropping and walking away reduced parasitization from 50 to 33%. These results support the hypothesis that the antipredator behaviour of an aphid changes as a function of internal stress. By performing less costly behaviour such as kicking under energetically constrained conditions, aphids seem to minimize their probability of energy shortfall. Given that aphid antipredator behaviour is a function of nutritional state, its occurrence under natural conditions may match host quality spatial distribution. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Ecology shapes the evolutionary trade‐off between predator avoidance and defence in coral reef butterflyfishes

Antipredator defensive traits are thought to trade‐off evolutionarily with traits that facilitate predator avoidance. However, complexity and scale have precluded tests of this prediction in many groups, including fishes. Using a macroevolutionary approach, we test this prediction in butterflyfishes, an iconic group of coral reef inhabitants with diverse social behaviours, foraging strategies and antipredator adaptations. We find that several antipredator traits have evolved adaptively, dependent primarily on foraging strategy. We identify a previously unrecognised axis of diversity in butterflyfishes where species with robust morphological defences have riskier foraging strategies and lack sociality, while species with reduced morphological defences feed in familiar territories, have adaptations for quick escapes and benefit from the vigilance provided by sociality. Furthermore, we find evidence for the constrained evolution of fin spines among species that graze solely on corals, highlighting the importance of corals, as both prey and structural refuge, in shaping fish morphology.     

Tadpoles of the bronze frog (Rana temporalis) assess predation risk before evoking antipredator defense behavior

Predation threat-associated behavioral response was studied in Rana temporalis tadpoles to discover the importance of predators’ visual and chemical cues (kairomones and diet-derived metabolites of consumed prey) in evoking antipredator behavior. The caged predators (dragonfly larvae) fed on prey tadpoles or insects (Notonecta spp.) and water conditioned with the predators provided the threat stimuli to the tadpole prey. The predators’ visual cues were ineffective in evoking antipredator behaviors in the tadpole prey. However, exposure to caged tadpole-fed predators or water conditioned with tadpole-fed predators elicited predator avoidance behavior in the tadpoles; they stayed away from the predators, significantly reduced swimming activity (swimming time and distance traveled), and increased burst speed. Interestingly, exposure to water conditioned with starved predators did not elicit any antipredator behavior in the prey. Further, the antipredator responses of predator-experienced tadpoles were significantly greater than those exhibited by predator-na?ve tadpoles. The study shows that R. temporalis tadpoles assess predation threat based exclusively on chemical cues emanating from the predators’ dietary metabolites and that the inclusion of conspecific prey items in the diet of the predators is perceived as a threat. The study also shows that antipredator behavior in these tadpoles is innate and is enhanced during subsequent encounters with the predators.     

Ontogenetic colour change and the evolution of aposematism: a case study in panic moth caterpillars

1. Aposematism is a widely used antipredator strategy in which an organism possesses both warning coloration and unprofitable characters. Theoretical evidence suggests that aposematic colour should develop when high opportunity costs imposed by crypsis force an organism to engage in conspicuous behaviours. Hence, it is expected that ontogenetic colour change (OCC) in larval insects should include aposematism when foraging needs compel behavioural modifications that preclude a continued state of crypsis. 2. To test this idea, I first investigated whether OCC in caterpillars of the panic moth Saucrobotys futilalis was indicative of a switch from cryptic to aposematic coloration. I then examined the context of panic moth OCC as it related to foraging patterns and behavioural conspicuousness. 3. Early Saucrobotys instars are a cryptic green, but later instars become progressively more orange and develop black spots. Early instar larvae forage cryptically on the inner parenchyma of silked-together host plant leaves to avoid predation, but are rapidly forced to engage in conspicuous foraging behaviours as they outgrow the resources afforded by their shelters. Both coloration and behaviour reach maximal conspicuousness in final instar larvae. 4. As predicted, OCC encompassed a change from crypsis to aposematism in Saucrobotys. Aposematic function was demonstrated by changes in both antipredator behaviour patterns and effectiveness of predator deterrence in early and late instars. Moreover, increased opportunity costs of crypsis and behavioural conspicuousness coincided with the onset of aposematic coloration. 5. This pattern of OCC suggests that aposematic coloration in Saucrobotys develops as a response to constraints imposed by crypsis. Moreover, my study illustrates the importance of the study of ontogenetic patterns in determining how behaviour, morphology, and predator responses interact to influence the initial evolution of phenomena such as aposematism.     

Computational studies on carbohydrates: solvation studies on maltose and cyclomaltooligosaccharides (cyclodextrins) using a DFT/ab initio-derived empirical force field, AMB99C

An empirical force field, denoted AMB99C, has been used to study molecular properties of alpha-(1-->4)-linked carbohydrates in solution. AMB99C was parameterized using structural and energetic parameters from density functional ab initio methodology. In this work we examine the solution behavior of the beta anomer of maltose and cyclohexa-, cyclohepta-, and cyclooctaamyloses (alpha-, beta-, and gamma-cyclodextrins or alpha-, beta-, and gamma-CDs, respectively), as well as of two larger (DP 10, epsilon-CD; DP 21) cyclomaltooligosaccharides, CA10 and CA21. Experimental data used for comparison purposes include X-ray structures, small-angle scattering radius of gyration values, NMR nuclear Overhauser enhancements (NOEs), and proton coupling constants. Molecular dynamics simulations were carried out using explicit water molecules (TIP3P) to establish equilibrium populations of conformations in solution, and these results are compared with other calculated values and a variety of experimental parameters, such as average H-1-H-4' distances between the rings in beta-maltose, and the primary hydroxyl groups' conformational populations. Medium-to-large cyclomaltooligosaccharide molecules were studied to test for glucose ring puckering and stability of kinked and 'flipped' conformations. The results of the solvation studies are in excellent agreement with experimental structural parameters.     

Teasing Apart the Effects of Seed Size and Energy Content on Rodent Scatter-Hoarding Behavior

Scatter-hoarding rodents are known to play a crucial role in the seed dispersal of many plant species. Numerous studies have indicated that both seed size and the energy content of seeds can affect rodent foraging behavior. However, seed size is usually associated with energy content per seed, making it difficult to isolate how seed size and energy affect rodent foraging preferences. This study used 99 treatments of artificial seeds (11 seed sizes×9 levels of energy content) to tease apart the effect of seed size and energy content on rodent seed-caching behavior. Both seed traits showed significant effects, but their details depended on the stage of the rodent foraging process. Seeds with higher energy content were harvested more rapidly while seed size only had a modest effect on harvest rate. However, after harvesting, seed size showed a much stronger effect on rodent foraging behavior. Rodents’ choice of which seeds to remove and cache, as well as seed dispersal distance, seemed to reflect an optimal seed size. Our findings could be adapted in future studies to gain a better understanding of scatter-hoarding rodent foraging behavior, and the co-evolutionary dynamics between plant seed production and seed dispersers.     

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.     

Scrounging behavior regulates population dynamics

The integration of behavioral and population ecology is necessary when behavior both feeds into demographic parameters and depends on population parameters. We show that scrounging behavior, the exploitation of others' resources, can affect both demographic parameters and population dynamics, including the stability of interactions with prey. Scrounging is a common tactic and its pay-offs exhibit both density- and frequency-dependence. We demonstrate that scrounging can act as a population regulator through its effects on individuals' reproductive rate and mortality. We also explore its effects on predator-prey population dynamics and show that the presence of scrounging predators allows an increased predator population size and contributes to the regulation of both predator and prey populations. Behavioral ecologists will appreciate that although scrounging is often pictured as imposing a social foraging cost to group membership, at the population level it also allows higher numbers of both prey and predators to coexist at equilibrium.     

Reduction of predation risk under the cover of darkness: Avoidance responses of mayfly larvae to a benthic fish

Summary Mayfly larvae of Paraleptophlebia heteronea (McDunnough) had two antipredator responses to a nocturnal fish predator (Rhinichthys cataractae (Valenciennes)): flight into the drift and retreat into interstitial crevices. Drift rates of Paraleptophlebia abruptly increased by 30 fold when fish were actively foraging in the laboratory streams but, even before fish were removed, drift began returning to control levels because larvae settled to the substrate and moved to areas of low risk beneath stones. This drifting response was used as an immediate escape behavior which likely decreases risk of capture from predators which forage actively at night. Surprisingly, drift most often occurred before contact between predator and prey, and we suggest that in darkness this mayfly may use hydrodynamic pressure waves for predator detection, rather than chemical cues, since fish forage in an upstream direction. Although drifting may represent a cost to mayfly larvae in terms of relocation to a new foraging area with unknown food resources, the immediate mortality risk probably out-weighs the importance of staying within a profitable food patch because larvae can survive starvation for at least 2 d. In addition to drifting, mayflies retreated from upper, exposed substrate surfaces to concealed interstitial crevices immediately after a predator encounter, or subsequent to resettlement on the substrate after predator-induced drift. A latency period was associated with this response and mayflies remained in these concealed locations for at least 3 h after dace foraging ceased. Because this mayfly feeds at night and food levels are significantly lower in field refugia under stones, relative to exposed stone surfaces, predator avoidance activity may limit foraging time and, ultimately, reduce the food intake of this stream mayfly.     

Temperature and Cloud Cover, but Not Predator Urine, Affect Winter Foraging of Mice

Although homeotherms likely experience costs of both predation risk and thermoregulation while foraging, it is unclear how foragers contend with these costs. We used foraging trays placed in sheltered microsites to determine whether temperature, a direct cue of predator presence (predator urine) and an indirect cue of predation risk (cloudy nights) affect foraging of white-footed mice, Peromyscus leucopus , in winter. Mice were presented with urine from bobcats, Lynx rufus , red foxes, V ulpes vulpes , and coyotes, Canis latrans , an herbivore (whitetailed deer, Odocoileus virginianus ), and a water control. To measure rodent foraging, we used seeds of millet mixed with sand to quantify giving-up densities (the number of seeds left in each foraging tray). Giving-up density was not affected by predator urine. Rather, rodent foraging was affected by an interaction of temperature and weather. On overcast nights, when predation risk was likely lower, mice foraged more when soil temperature was higher, presumably reducing thermoregulatory costs. On clear nights, foraging was low regardless of soil temperature, presumably because foraging was more risky. These results suggest that mice consider thermoregulatory costs and predation risk when making foraging decisions, and that the indirect cue afforded by weather, rather than the direct cue of predator urine, is among the cues used to make foraging decisions. Moreover, these results suggest that sensitivity to a particular cue is likely to be context-dependent.     

Effects of fat and protein levels on foraging preferences of tannin in scatter-hoarding rodents

Both as consumers and dispersers of seeds, scatter-hoarding rodents often play an important role in the reproductive ecology of many plant species. However, the seeds of many plant species contain tannins, which are a diverse group of water-soluble phenolic compounds that have a high affinity for proteins. The amount of tannins in seeds is expected to affect rodent foraging preferences because of their major impact on rodent physiology and survival. However, variable results have been obtained in studies that evaluated the effects of tannin on rodent foraging behavior. Hence, in this study, we aimed to explain these inconsistent results and proposed that a combination of seed traits might be important in rodent foraging behavior, because it is difficult to distinguish between the effects of individual traits on rodent foraging behavior and the interactions among them. By using a novel artificial seed system, we manipulated seed tannin and fat/protein levels to examine directly the univariate effects of each component on the seed preferences of free-ranging forest rats (Apodemus latronum and Apodemus chevrieri) during the behavioral process of scatter hoarding. Our results showed that both tannin and fat/protein had significant effects on rodent foraging behavior. Although only a few interactive effects of tannin and fat/protein were recorded, higher concentrations of both fat and protein could attenuate the exclusion of seeds with higher tannin concentrations by rodents, thus influencing seed fate. Furthermore, aside from the concentrations of tannin, fat, and protein, numerous other traits of plant seeds may also influence rodent foraging behavior. We suggest that by clarifying rodent foraging preferences, a better understanding of the evolution of plant seed traits may be obtained because of their strong potential for selective pressure.     

视野受阻对东方田鼠觅食行为的影响

植食性小型哺乳动物的觅食行为是对特定生境的适应性产物。食物斑块周边植被对动物视野遮挡是否通过作用于其觅食活动中的警觉而影响摄入率。采用新鲜白三叶叶片构建东方田鼠密集均质食物斑块,以牛皮纸模拟食物斑块周边植被遮挡田鼠视野,测定其在食物斑块上的觅食行为序列过程及行为参数,检验食物斑块周边植被高度对东方田鼠摄入率的影响。结果发现,个体在不同程度视野受阻条件下食物摄入率无显著差异。分析觅食行为参数动态发现,在不同视野受阻条件下,个体能通过调整各采食回合内警觉行为动作的发生频次和持续时间,维持觅食回合内总的觅食中断时间的稳定,进而保证进食时间的稳定。东方田鼠在不同程度的视野遮挡条件下均能通过行为变异和优化使摄入率保持稳定。结果亦充分说明,东方田鼠在警觉强度上的变化不能反映觅食中断所带来的食物收益减损的代价,但觅食活动中各警觉动作的持续时间的变异却能够明确指示个体摄入率的动态变化,因此,以觅食活动中警觉引起的觅食中断时间代价为线索,检验其对摄入率的影响,是评价植食性小型哺乳动物在不同生境觅食适应性策略的有效的方法。