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.     

Is Foraging Time Limited During Winter? – A Feeding Experiment with Tree Sparrows Under Different Predation Risk

Small passerines are faced with a trade‐off when foraging during winter. Increasing energy reserves makes them more vulnerable to predators, while a low level of reserves exposes them to a high risk of starvation. Whether small birds under these circumstances are allowed to reduce their foraging activity under increased predation risk, for example in feeding sites more exposed to predators, remains controversial in former behavioural and ecological researches. In this study, we investigated the foraging activity of free‐living Tree Sparrow Passer montanus flocks feeding on an artificial feeding platform. The predation risk perceived by the sparrows was manipulated by placing the platform either close to or far from a bushy shelter. Foraging activity, assessed as cumulative activity of sparrows per unit time on the platform, did not differ between the low‐risk and the high‐risk conditions and did not significantly change during the day. Feeding efficiency, assessed as pecking rate, was not either reduced under the high‐risk condition. Our results suggest that sparrows were forced to feed almost continuously during the day in order to maintain their preferred level of energy reserves. However, several behavioural changes helped sparrows to adopt a safer foraging policy when feeding far from cover, as we found in another study. Altogether, sparrow flocks feeding far from cover decreased the overall foraging time (the time when any sparrow stayed on the platform) by approximately 20% as compared to the near cover condition. A possible way to maintain the same level of foraging activity despite of the reduction in overall foraging time is discussed.     

Mass regulation in response to predation risk can indicate population declines

In theory, survival rates and consequent population status might be predictable from instantaneous behavioural measures of how animals prioritize foraging vs. avoiding predation. We show, for the 30 most common small bird species ringed in the UK, that one quarter respond to higher predation risk as if it is mass-dependent and lose mass. Half respond to predation risk as if it only interrupts their foraging and gain mass thus avoiding consequent increased starvation risk from reduced foraging time. These mass responses to higher predation risk are correlated with population and conservation status both within and between species (and independently of foraging habitat, foraging guild, sociality index and size) over the last 30 years in Britain, with mass loss being associated with declining populations and mass gain with increasing populations. If individuals show an interrupted foraging response to higher predation risk, they are likely to be experiencing a high quality foraging environment that should lead to higher survival. Whereas individuals that show a mass-dependent foraging response are likely to be in lower quality foraging environments, leading to relatively lower survival.     

Asset protection in juvenile salmon: how adding biological realism changes a dynamic foraging model

The "asset-protection principle" created by Clark is based ona dynamic programming model and states that individuals should(1) become more averse to predation risk as they accumulatefitness assets but (2) generally be more willing to acceptpredation risk later in the foraging season. To test whetherthese predictions hold under biologically meaningful foraging parameters, I constructed a dynamic model of the optimal trade-offbetween foraging and predator avoidance in juvenile salmon.The model incorporates temperature and body-size dependentbio-energetic constraints typical for juvenile fish, whichgrow by orders of magnitude over a season. In its simplestform using seasonally constant growth potential and a linear over-winter survival function, my results equal those of Clark'smodel. Adding a fitness function and environmental data fromfield studies accentuates the asset-protection effect and fundamentallychanges the seasonal pattern of optimal effort. Simulationof typical poor feeding conditions in mid-summer yields theprediction of increased foraging in the spring in anticipationof worsening conditions. Increasing overall predation riskresults in smaller fish at the end of the season with a trade-offbetween summer and winter survival. The model generates testablepredictions for juvenile salmon and provides insights for otherorganisms (particularly poikilotherms) that are subject tosize-dependent or seasonally changing foraging dynamics.     

Vigilance and food intake rate in paired and solitary Zenaida Doves Zenaida aurita

We quantified vigilance during feeding in the Zenaida Dove Zenaida aurita, a tropical species with stable pair‐bonds and year‐round territoriality. Both males and females decreased the proportion of time spent vigilant by 30% when feeding with their partner compared with when feeding alone. This reduction was achieved through increasing the length of inter‐scan duration, while scan duration remained constant. No evidence was found for coordination of vigilance between pair members. The equal investment in vigilance by male and female Zenaida Doves might be related to the mutual benefits of long‐term pair‐bonding.     

Why don't predators have positive effects on prey populations?

Summary Three mechanisms by which increasing predation can increase prey population density are discussed: (1) Additional predation on species which have negative effects on the prey; (2) Predation on consumer species whose relationship with their own prey is characterized by a unimodal prey isocline; (3) Predation on species which adaptively balance predation risk and food intake while foraging. Possible reasons are discussed for the rarity of positive effects in previous predator-manipulation studies; these include the short-term nature of experiments, the large magnitudes of predator density manipulation, and various sources of bias in choice of system and interpretation of results.     

Mean-variance sets for dietary choice models: simplicity in a complex world

Summary This paper presents a series of simulations designed to determine optimal diet breadth under shortfall avoidance models. Profitability and encounter rate functions were varied, and means and variances of energy intake rate were generated using a simple simulation procedure. The resulting mean-variance sets assumed three distinct shapes: u-shaped, arched, and looped. These simulations show that certain mean-variance sets allow the forager to employ simple behavioural rules to determine the optimal diet breadth. This situation occurs when low ranking diet items have small handling times, and these conditions may be quite common. In other cases, mean-variance sets may be too complicated to allow for easy behavioural rules designed to minimize starvation probability. The ability to characterize foraging problems into a limited series of mean-variance set types benefits workers examining the evolution and maintenance of foraging strategies, since these sets have clear implications for the ability of animals to develop simple behavioural rules. Unfortunately data are lacking on the profitability and encounter rate distributions animals face in nature.     

Alternative forms of competition and predation dramatically affect habitat selection under foraging--predation-risk trade-offs

Habitat selection under foraging—predation-risk trade-offshas been a frequent topic of interest to theoretical behavioraland evolutionary ecologists. However, most habitat selectionmodels assume that individuals compete exploitatively for resourcesand that predation is either density independent or dilutedcompletely by competitor number, despite empirical evidencethat other forms of competition and predation also occur innature. I developed an individual-based model for studyingthe effects of alternative forms of competition and predationon the process of habitat selection under foraging—predation-risktrade-offs. To make the model more relevant to natural populations,I assumed that individuals vary continuously in traits relatedto competitive ability and vulnerability to predation and allowedresources and predators to be distributed across more than twohabitats. The results of my investigation demonstrate thatthe predicted pattern of habitat selection can be affecteddramatically by the form predation is assumed to take. Whenpredation is density dependent or frequency dependent, individualswill tend to be distributed across habitats according to theirabsolute vulnerability to predation. In contrast, when predationis density dependent or vulnerability dependent, individualswill tend to segregate by competitive ability. Whether oneassumes that individuals compete for resources via exploitationor interference also influences the predicted pattern of habitatselection. In general, interference competition results in amore even distribution of competitors across habitats.     

Seeing a Ghost? Vigilance and Its Drivers in a Predator‐free World

Vigilance is a key to the early detection of predators, but may be costly if it impairs foraging efficiency. Hence, we would expect vigilance to be suppressed and/or counter‐selected in predator‐free environments, although this might depend on the environmental drivers influencing perceived predation risk. We studied vigilance in two populations of Sitka black‐tailed deer (Odocoileus hemionus sitkensis) on Haida Gwaii (Canada) which have not been exposed to predators since they colonized the study islands approx. 60 yr ago. In this context, anti‐predator behavior should not have any obvious current benefit. Moreover, its maintenance should be particularly costly in our study populations because these deer have depleted their food resources and, thus, anti‐predator behaviors should interfere with time spent searching for scarce resources. We used bait stations equipped with camera traps to assess vigilance under standardized feeding conditions. We expected to observe lower vigilance levels than those observed elsewhere in locations with predators. We investigated how vigilance varied in relation to the amount of bait, the level of visibility, and between day and night. During the day, deer spent, on average, 14% of their time in overt vigilance during foraging bouts, a level similar to, although in the lower range of, values reported at sites where predators are present. Levels of vigilance were lower at night, and decreased with increasing visibility, but not during the day. Deer were less vigilant when bait availability was high, but only when visibility was also high. We discuss why the maintenance of vigilance is here best explained by the ghosts of predators past, and how, at the temporal scale of a few generations, the ecological factors driving vigilance levels might override the absence of significant risk from large predators.     

Barí loricarid collection and the value of information: An application of optimal foraging theory

Analysis of Barí collection of loricarid species suggests that the length of time that has elapsed since each collecting site has last been exploited significantly guides site selection. Information on patch recovery time, gathered through intra-village monitoring of independent foraging groups, allows foragers to choose those sites with a high probability of generating good returns. Comparison of actual returns with those predicted by a model of random site selection indicates that the observed pattern of patch exploitation increases the return of kilograms of loricarids for time invested in foraging substantially above that predicted by random returns to sites. The saving of time as well as the increase in food afforded by this system represent currencies for evaluating the value of information on patch recovery time.     

捕食风险及其对动物觅食行为的影响

对捕食风险的涵义及其对猎物动物觅食行为的影响、猎物动物面对捕食风险时的反应进行了论述。捕食风险可以简单地理解为一定时间内猎物动物被杀死的概率。当捕食风险存在时 ,动物会选择相对安全但觅食效益较低的地点觅食 ;由于死亡率和消化方面的限制 ,一般都会产生食谱收缩 ;觅食活动方式的时间格局也会因捕食风险而发生改变 ,如水生动物的昼夜垂直迁移、某些陆生动物昼行性与夜行性活动的转换、月光回避等。在与捕食者发生遭遇时 ,猎物动物的主要反应是 :①发出某些信号以阻止捕食者的追捕 ;②靠近并注视捕食者 ;③逃逸 ;④在一定的时间恢复觅食活动。在以往的研究中 ,对捕食者种类已经有了较多的了解 ,而对猎物如何判断捕食者丰富度信息、估计风险程度等方面则知之甚少 ;同时 ,对捕食风险水平的调控、对多种因素的综合分析也较少涉及。在今后的研究中 ,还应该考虑研究的尺度问题 ,因为在不同尺度的环境条件下 ,猎物动物对于捕食风险的反应可能大相径庭。