The energetic equivalence of cover to juvenile coho salmon (Oncorhynchus kisutch): ideal free distribution theroy applied

Cover is often thought to be an important habitat characteristicfor juvenile stream salmonida. In addition to providing protectionfrom predators, cover may be associated with reduced food availability.Thus, an individual's use of cover is likely to reflect a trade-offbetween the conflicting demands of growth and survival. We measuredthe influence of cover on foraging-site selection in groupsof eight juvenile coho salmon (Oncorhynchus kisutch) by examiningtheir distribution across two stream channel patches, one providingaccess to cover but little food (the "poor" patch), the otherproviding more food but no cover (the "good" patch). Becausefish distributions in the absence of cover conformed to an idealfree distribution (IFD) for unequal competitors (i.e., the distributionof competitive abilities matched the distribution of food),we used IFD theory to quantify the energetic equivalence ofcover to the fish. In the presence of cover and a model avianpredator, use of the poor patch increased relative to the predictionsof the IFD model. Using this observed deviation from an IFD,we calculated how much extra food must be added to the goodpatch to return the distribution of fish to the previously observedIFD of unequal competitors. As predicted, adding this amountof food caused the fish to return to their previous distribution,demonstrating that IFD theory can be used to relate energy intakeand risk of predation in a common currency     

Season- and size-dependent risk taking in juvenile coho salmon: experimental evaluation of asset protection

Using juvenile coho salmon, Oncorhynchus kisutch, we tested predictions arising from dynamic optimization models of foraging under predation risk. Coho juveniles from two size groups raised in the laboratory were individually fed varying food rations. Their willingness to risk predation was measured as the time to resume foraging after presentation of a predator model. Small fish (mean weight 1.5 g) resumed feeding earlier than larger fish (3.5 g) as predicted by dynamic models under summer photoperiod but not under autumn photoperiod. Contrary to predictions, larger fish did not increase risk taking and small fish decreased risk taking between summer and autumn treatments. Food ration significantly influenced time to resume feeding only in small coho. A simple mechanistic model we proposed to explain feeding motivation under risk as a function of body size and prior growth rate was not sufficient to explain observed variation in risk taking. This study suggests that coho salmon use photoperiod and their own body size as cues for long-term, state-dependent adjustments of feeding behaviour. The lower risk taking of larger fish is probably an example of asset protection, whereby larger animals accept less predation risk to protect their greater accumulated fitness value. The decrease of risk taking in small fish in the autumn was possibly caused by a switch of life history trajectory towards delayed smolting. Copyright 1999 The Association for the Study of Animal Behaviour.     

Prey selection by experienced and naive juvenile Atlantic salmon

Both in foraging groups and in a sequential prey encounter context, learning had a visible effect on the pattern of selection for three live prey types ( Ecdyonurus larvae, Hydropsyche larvae, and Gammarus ) by juvenile Atlantic salmon Salmo salar . Compared to wild-caught fish, naive, hatchery-reared fish that had not been exposed to natural prey ate Hydropsyche larvae in a remarkably low proportion, and consumed a higher proportion of Gammarus. Ecdyonurus experienced a high and rather steady predation rate across the experience gradient, but after a short period of experience with live prey the consumption rate for Hydropsyche increased drastically, and that of Gammarus decreased, matching the selection pattern exhibited by wild fish. Individual fish offered prey in a sequential encounter context increased consumption rates of all the prey types as they gained experience, but the improvement was higher for the prey that were less consumed initially. Fish became more selective as they approached satiation, conforming to the prediction of optimal foraging theory that higher predator's energy requirements, as well as low food availability, result in reduced selectivity. The results also suggest that fish from distinct populations can differ in the degree of diet selectivity according to their energetic requirements for growth. The fast learning response of Atlantic salmon parr towards novel prey probably allows fish to maintain a high foraging efficiency when faced with frequent changes in the availability of different prey types.     

Structural complexity and fish body size interactively affect habitat optimality

Predator–prey interactions are strongly influenced by habitat structure, particularly in coastal marine habitats such as seagrasses in which structural complexity (SC) may vary over small spatial scales. For seagrass mesopredators such as juvenile fishes, optimality models predict that fitness will be maximized at levels of SC that enhance foraging but minimize predation risk, both of which are functions of body size. We tested the hypothesis that in eelgrass (Zostera marina) habitat, optimal SC for juvenile giant kelpfish (Heterostichus rostratus), an abundant eelgrass mesopredator in southern California, changes through ontogeny. To do this, we quantified eelgrass SC effects on habitat associations, relative predation risk, and foraging efficiency for three size classes of juvenile giant kelpfish. We found that habitat selection differed with fish size: small fish selected dense eelgrass, whereas larger fish selected sparse eelgrass. Small kelpfish experienced the lowest relative predation risk in dense eelgrass but also had higher foraging efficiency in dense eelgrass, suggesting that dense eelgrass is selected by these fish because it minimizes risk and maximizes potential for growth. Surprisingly, larger kelpfish did not experience lower predation risk than small kelpfish. However, larger kelpfish experienced higher foraging efficiency in sparse eelgrass vs. dense eelgrass, suggesting that they select sparse eelgrass to maximize foraging efficiency. Our study highlights that trade-offs between predation risk and foraging can occur within a single habitat type, that studies should consider how habitat value changes through ontogeny, and that seagrass habitat value may be maximal when within-patch variability in SC is high.     

A wide window of migration phenology captures inter‐annual variability of favourable conditions: Results of a whole‐lake experiment with juvenile Pacific salmon

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Age and seasonal effects on predator-sensitive foraging in squirrel monkeys (Saimiri sciureus): a field experiment

A field experiment was conducted to examine the effect of perceived predation risk on the use of foraging areas by juvenile and adult primates under different conditions of local food abundance. Wild squirrel monkeys, Saimiri sciureus, were observed in an experiment conducted during the dry and the wet seasons at a site in Eastern Amazonia, Brazil. Animals were presented with feeding platforms that differed in food quantity and exposure to aerial predators through varying vegetative cover. In the dry season, juveniles and adults chose platforms based solely on food quantity. However, in the wet season, juveniles foraged preferentially on high-reward platforms only if cover level also was high (i.e., potentially offered greater concealment from predators). In contrast, adults showed the same pattern of platform use regardless of season. These results indicate that age and local resource availability based on seasonality affect whether primates forage in a predator-sensitive manner. Juveniles may be more sensitive to predation risk when foraging, and individuals may take fewer risks when resource abundance is high in their environment.     

Effects of ice cover on the diel behaviour and ventilation rate of juvenile brown trout

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Linking recruitment to trophic factors: revisiting the Beverton--Holt recruitment model from a life history and multispecies perspective

The Beverton--Holt recruitment model can be derived from arguments about evolution of life history traits related to foraging and predation risk, along with spatially localized and temporarily competitive relationships in the habitats where juvenile fish forage and face predation risk while foraging. This derivation explicitly represents two key biotic factors, food supply (I) and predator abundance (R), which appear as a risk ratio (R/I) that facilitates modelling of changes in trophic circumstances and analysis of historical data. The same general recruitment relationship is expected whether the juvenile life history is simple or involves a complex sequence of stanzas; in the complex case, the Beverton--Holt parameters represent weighted averages or integrals of risk ratios over the stanzas. The relationship should also apply in settings where there is complex, mesoscale variation in habitat and predation risk, provided that animals sense this variation and move about so as to achieve similar survival at all mesoscale rearing sites. The model predicts that changes in food and predation risk can be amplified violently in settings where juvenile survival rate is low, producing large changes in recruitment rates over time.     

The costs of becoming nocturnal: feeding efficiency in relation to light intensity in juvenile Atlantic Salmon

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging.
2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness.
3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift-feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging.
4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.     

Linking piscivory to spatial–temporal distributions of pelagic prey fishes with a visual foraging model

A visual foraging model (VFM) used light-dependent reaction distance and capture success functions to link observed prey fish abundance and distribution to predation rates and the foraging performance of piscivorous cutthroat trout Oncorhynchus clarki in Lake Washington (WA, U.S.A.). Total prey density did not correlate with predation potential estimated by the foraging model for cutthroat trout because prey were rarely distributed in optically favourable conditions for detection. Predictions of the depth-specific distribution and timing of cutthroat trout foraging were qualitatively similar to diel stomach fullness patterns observed in field samples. Nocturnal foraging accounted for 34–64% of all prey fish consumption in simulations for 2002 and 2003. Urban light contamination increased the access of nocturnally foraging cutthroat trout to vertically migrating prey fishes. These results suggest that VFMs are useful tools for converting observed prey fish density into predictions of predator consumptions and behavioural responses of predators to environmental change.     

Risk‐taking behaviour may explain high predation mortality of GH‐transgenic common carp Cyprinus carpio

The competitive ability and habitat selection of juvenile all‐fish GH‐transgenic common carp Cyprinus carpio and their size‐matched non‐transgenic conspecifics, in the absence and presence of predation risk, under different food distributions, were compared. Unequal‐competitor ideal‐free‐distribution analysis showed that a larger proportion of transgenic C. carpio fed within the system, although they were not overrepresented at a higher‐quantity food source. Moreover, the analysis showed that transgenic C. carpio maintained a faster growth rate, and were more willing to risk exposure to a predator when foraging, thereby supporting the hypothesis that predation selects against maximal growth rates by removing individuals that display increased foraging effort. Without compensatory behaviours that could mitigate the effects of predation risk, the escaped or released transgenic C. carpio with high‐gain and high‐risk performance would grow well but probably suffer high predation mortality in nature.     

Influence of prey foraging posture on flight behavior and predation risk: predators take advantage of unwary prey

Foraging in animals is often associated with characteristicbody postures, such as the head-down posture. When foragingconflicts with the ability to detect predators or to flee, individualsmay incur a greater risk of mortality to predation than otherwise.Here we investigate the influence of different foraging postures(horizontal versus nose-down body posture) on the ability ofindividuals to respond to approaching predators and on the riskof mortality to predation in the guppy (Poecilia reticulata).Individuals engaged in nose-down foraging were assumed to beable to visually scan a smaller area for predators and to escapeless effectively due to their body posture, and thus are morevulnerable to stalking predators than horizontally foragingones. In a first experiment, we separately exposed nonforaging,horizontally foraging, and nose-down foraging guppies to anapproaching cichlid fish predator model. Nonforaging guppiesreacted sooner to and initiated flight further away from theapproaching model than did foraging fish collectively, and horizontallyforaging individuals responded sooner to the model than nose-downforaging ones. Comparing all test guppies, nose-down foragingindividuals were the most likely not to exhibit any responseto the predator model. When presented with a simultaneous choiceof two guppies behind a one-way mirror, individual blue acaracichlid (Aequidens pulcher), a natural predator of the guppy,preferred to attack foraging guppies over nonforaging ones andnose-down foraging guppies over horizontally foraging individuals.In a final experiment with free-swimming cichlids and guppies,we demonstrated that individual risk of predation for guppiesforaging nose down was greater than for guppies foraging horizontally,and both were at greater risk than nonforaging guppies. Thislatter result is consistent with the above differences in theguppy's responsiveness to approaching predators depending ontheir foraging behavior, and with the finding that cichlid predatorspreferred fish that were less likely to show any response tothem. Our results therefore indicate that the ability to respondto approaching predators and the risk of mortality to predationin the guppy is strongly influenced by their foraging activity,and in particular their foraging posture, and that cichlid predatorspreferentially select less wary and more vulnerable guppies.[BehavEcol 7: 264–271 (1996)]     

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.     

Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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The starvation–predation risk trade-off, body mass and population status in the Common Starling Sturnus vulgaris

It is theoretically and empirically well established that body mass variation in small birds reflects a trade-off between starvation risk and predation risk. This occurs because carrying increased fat reserves reduces starvation risk but also results in a higher predation risk due to reduced escape flight performance and/or the increased foraging exposure needed to maintain a higher body mass. In principle, therefore, the theory of mass-dependent predation risk could be used to understand how a bird perceives and responds to the risks in its environment, because its mass will reflect the predictability of foraging opportunities and predation risk. Mass in birds may then provide a relatively straightforward way of assessing the foraging environment of birds and so the potential conservation problems a species faces. This study tests, for the first time for any species, how body mass changes in response to changing starvation risk, changing predation risk and changing population status. Common Starling Sturnus vulgaris mass varies as predicted by starvation–predation risk trade-off theory: mass is lower when foraging conditions are more favourable and when predation risk is increased. The populations that are declining the most strongly have higher mass, which is most likely indicative of a poor foraging environment, leading to lower relative survival. The results suggest that increased mass in Starlings, and possibly in other species, may provide an indication of the poor quality of the foraging environment and/or rapidly declining populations.     

Predation risk influences food‐web structure by constraining species diet choice

The foraging behaviour of species determines their diet and, therefore, also emergent food‐web structure. Optimal foraging theory (OFT) has previously been applied to understand the emergence of food‐web structure through a consumer‐centric consideration of diet choice. However, the resource‐centric viewpoint, where species adjust their behaviour to reduce the risk of predation, has not been considered. We develop a mechanistic model that merges metabolic theory with OFT to incorporate the effect of predation risk on diet choice to assemble food webs. This ‘predation‐risk‐compromise’ (PR) model better captures the nestedness and modularity of empirical food webs relative to the classical optimal foraging model. Specifically, compared with optimal foraging alone, risk‐mitigated foraging leads to more‐nested but less‐modular webs by broadening the diet of consumers at intermediate trophic levels. Thus, predation risk significantly affects food‐web structure by constraining species’ ability to forage optimally, and needs to be considered in future work.     

Avian daily foraging patterns: Effects of digestive constraints and variability

Summary Birds show a typical daily pattern of heavy morning and secondary afternoon feeding. We investigate the pattern of foraging by a bird that results in the lowest long-term rate of mortality. We assume the following: mortality is the sum of starvation and predation. The bird is characterized by two state variables, its energy reserves and the amount of food in its stomach. Starvation occurs during the day if the bird's reserves fall to zero. The bird starves during the night if the total energy stored in reserves and the stomach is less than a critical amount. The probability that the bird is killed by a predator is higher if the bird is foraging than if it is resting. Furthermore, the predation risk while foraging increases with the bird's mass. From these assumptions, we use dynamic programming techniques to find the daily foraging routine that minimizes mortality. The principal results are (1) Variability in food finding leads to routines with feeding concentrated early in the day, (2) digestive constraints cause feeding to be spread more evenly through the day, (3) even under fairly severe digestive constraints, the stomach is generally not full and (4) optimal fat reserve levels are higher in more variable environments and under digestive constraints. This model suggests that the characteristic daily feeding pattern of small birds is not due to digestive constraints but is greatly influenced by environmental variability.     

Revisiting Beverton–Holt recruitment in the presence of variation in food availability

Understanding density-dependent changes in juvenile survival and growth rates is of great importance because these rates determine recovery rates for imperiled populations and/or sustainable harvest rates. Unfortunately, the mechanisms leading to density dependent survival and growth are among the least understood process in biology and fisheries. Previous work has shown that small fish may vary foraging times to achieve a target growth rate, resulting in the well-known Beverton–Holt recruitment function with variation in food availability affected the initial slope of the recruitment curve. We amend their derivation to show that incorporating fish growth under a variety of evolutionary strategies for balancing foraging time and predation risk still leads to recruitment approximately as expected under the Beverton–Holt recruitment model but that changing food availability affects both the initial slope and maximum recruitment level. We demonstrate that when food availability is known to vary over time, these models often result in a more parsimonious alternative than the standard Beverton–Holt function. Further, Beverton–Holt recruitment is expected when foraging times are adjusted to balance fitness gains from growth against mortality risk. Finally, linking recruitment success to food availability warns that species with high scope for density dependent survival (high compensation ratio or steepness) may be extremely sensitive to changes in available food densities. This work emphasizes the sensitivity of stock-recruitment parameters to food availability and strongly suggests a need to carefully monitor lower trophic levels to better understand and predict dramatic changes in juvenile recruitment and carrying capacity.     

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。     

Using niche construction theory to generate testable foraging hypotheses at Liang Bua

Niche construction theory (NCT) has emerged as a promising theoretical tool for interpreting zooarchaeological material. However, its juxtaposition against more established frameworks like optimal foraging theory (OFT) has raised important criticism around the testability of NCT for interpreting hominin foraging behavior. Here, we present an optimization foraging model with NCT features designed to consider the destructive realities of the archaeological record after providing a brief review of OFT and NCT. Our model was designed to consider a foragers decision to exploit an environment given predation risk, mortality, and payoff ratios between different ecologies, like more‐open or more‐forested environments. We then discuss how the model can be used with zooarchaeological data for inferring environmental exploitation by a primitive hominin, Homo floresiensis, from the island of Flores in Southeast Asia. Our example demonstrates that NCT can be used in combination with OFT principles to generate testable foraging hypotheses suitable for zooarchaeological research.