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.     

When to come out from a refuge: risk-sensitive and state-dependent decisions in an alpine lizard

Prey often respond to predator presence by increasing theiruse of refuges. However, unfavorable thermal conditions in refugesmight entail physiological costs for an ectothermic prey. Thus,the decision of when to come out from a refuge should be optimizedby considering the expected fitness effects of diminution ofpredation risk with time, but also by considering the cost of theloss of time spent at optimal body temperature maximizing physiological functions.The model of Ydenberg and Dill describes the trade-off betweenrisk and cost for a prey fleeing to a refuge. We present a specialcase of this model to predict how emergence time from the refugein lizards or other ectotherms should vary as a function ofrisk of predation and thermal costs of refuge use. The analysesof the variation in emergence time from a refuge of Lacertamonticola lizards in the field under two different predation risklevels supported the predictions of the model. As predicted,time spent in the refuge was longer when the threat of the initialattack had been higher, and therefore the subsequent diminutionof risk was slower, but only when lizards emerged at the sameplace where they hid. When initial body temperature was high,some lizards decreased emergence time by emerging from a differentplace. In addition, the effects of thermal costs were more relevant inthe high-risk situation. Time spent in the refuge under highrisk increased when thermal conditions of the refuge were moresimilar to thermal conditions outside (i.e., physiological costsof refuge use were lower). We conclude that optimization ofrefuge-use strategies might help lizards cope with changes in predationrisk without incurring excessive physiological costs.     

Effects of risk, cost, and their interaction on optimal escape by nonrefuging Bonaire whiptail lizards, Cnemidophorus murinus

Optimal escape theory seeks to explain variation in the distanceto an approaching predator at which the prey initiates escape(flight initiation distance). Flight initiation distance increaseswhen predators pose a greater threat and decreases when escapecosts increase. Although optimal escape theory has been highlysuccessful, its predictions have been tested primarily for speciesthat escape to discrete refuges, and most studies have focusedon single risk or cost factors. We present data from two experimentsin which two risks or a risk and a cost varied in Bonaire whiptaillizards (Cnemidophorus murinus) that escaped without enteringrefuges. Our data verify several predictions about optimal escapefor nonrefuging lizard prey. Two risk factors, speed and directnessof approach by the predator, interacted. Directly approachedlizards had greater flight initiation distances than did indirectlyapproached lizards when approached rapidly, but shorter flightinitiation distances when approached slowly. Flight initiationdistance was shorter in the presence of food and during slowversus rapid approaches, but contrary to expectation, food presenceand approach speed did not interact. This would be explainedif cost curves are nonlinear or if they are parallel ratherthan intersecting when the predator reaches the prey. More empiricalwork is needed to determine which risk and cost factors actadditively and which act synergistically. The absence of interactionbetween the risk and cost factors suggests that cost curveswere nonlinear.     

Enhanced growth at low population density in Daphnia: the absence of crowding effects or relief from visual predation?

1. It has been suggested that chemical information from crowded populations of an animal such as Daphnia carries a cue indicating imminent food limitation, and we suggest that in the presence of fish kairomones, it may also convey a hint of the need to enhance antipredation defences. 2. We performed two‐factorial experiments with Daphnia grown in flow‐through plankton chambers in medium containing high levels of Scenedesmus food plus chemical information on either low or high population density levels and in the presence or absence of fish chemical cues (kairomones) and recorded (i) the effects on Daphnia growth rate and reproduction, and (ii) the effects on Daphnia depth selection. Further depth‐selection experiments were performed to test the reaction of Daphnia to crowding information at different Daphnia concentrations and to test its effect on daytime and night‐time depth selection by different Daphnia instars in the presence of kairomones. 3. The effects of crowding information alone (in the absence of kairomones) were weak and were not significantly strengthened by the addition of kairomones. The effects of kairomones alone (in the absence of crowding information) were much stronger and were increased by the presence of crowding chemicals: Daphnia selected greater depths in daylight (the later the instar and the larger its body size, the greater the depth), their body growth was slower and daily reproductive investment reduced, compared with Daphnia grown in the absence of crowding information. This suggested that crowding chemicals carry a cue indicating the need to invest more into antipredation defences. 4. The adaptive significance of these effects was confirmed by the differential vulnerability to predation of the Daphnia when offered as prey to live roach after being grown for 6 days either in the presence (higher vulnerability) or in the absence (lower vulnerability) of information on high density. 5. The strong interaction between crowding information and fish kairomones may be explained either as the reaction to a cue indicating impending food stress or as the reaction to a signal of increased predation risk. While the former scenario is already known from crowding studies, the latter is a novel idea that stems from the old concept of ‘low‐density anti‐predation refuge’. The two scenarios are not mutually exclusive: each stems from the need to invest in survival rather than in growth and reproduction [Corrections were made to this paragraph after first online publication on 4 April 2012].     

Positioning behaviour in fish shoals: a cost–benefit analysis

Shoal position can have a strong influence on individual fitness. Individuals in front positions of shoals were observed to have higher feeding rates than individuals elsewhere. Manipulation of nutritional state showed that hungry individuals had a stronger preference for front positions and that the duration of food-deprivation was positively correlated with the degree of the position preference. On the other hand, front positions (like other peripheral positions) probably incur costs in terms of increased predation risks. Experiments with Schreckstoff showed that frightened individuals seek the central part of the shoal. This suggests that individuals rotate their shoal positions according to the tradeoff between energy intake and predation risk.     

Diel vertical movements by lotic mayfly nymphs under variable predation risk

1. The diel foraging periodicities of two grazing mayfly (Ephemeroptera) nymphs, Heptagenia dalecarlica and Baetis rhodani, under variable fish (European minnow) predation risk were examined in a series of laboratory experiments. 2. Heptagenia dalecarlica were almost exclusively nocturnal in their use of feeding areas on stone tops. There was a sharp increase in the proportion of nymphs out of refuge at nightfall, both in the control and fish-odour treatments. In a treatment with freely moving fish, H. dalecarlica decreased their activity during both the day and night. In feeding trials with three freely foraging minnows, nymphs were completely safe when they had access to refuges beneath stones, whereas almost half the nymphs were consumed when no refuges were available. 3. Baetis rhodani nymphs reduced their use of stone tops when exposed to four caged minnows or a freely moving fish, but this occurred both day and night. In feeding trials, B. rhodani were captured only while in the water column, and their mortality risk was independent of refuge availability. 4. It is suggested that because H. dalecarlica lack efficient post-encounter defences, they must rely on pre-encounter mechanisms to reduce the threat of fish predation. It appears that in order to dwell sympatrically with fish, H. dalecarlica have evolved a coexistence by hiding strategy. In contrast, Baetis are vulnerable to fish attacks only if they enter drift in daylight, and are thus able to remain on stone tops both during the day and at night.     

Mortality risk versus food quality trade-offs in ants: patch use over time

Abstract.

• 1 Foragers of the ant species Lasius pallitarsis (Provancher) and Myrmica incompleta Provancher were given access to two food patches that differed in nutritive quality. In about half of the trials, the higher quality patch also had a mortality risk for the foragers in the form of a large Formica subnuda Emery ant located along the trail. Colonies were given access to the food for 4 or 24 h, every second day.
• 2 Use of the higher quality patch by L.pallitarsis foragers was depressed by the mortality risk. In contrast to the predictions of a food stress hypothesis, foragers used the risky patch more in the longer access trials. This appears due to a decline in the activity level of the F.subnuda, which results in a significantly lower mortality risk per trip.
• 3 M.incompfeta also significantly decreased their use of the higher quality patch when a F.subnuda was present. However, the length of time a colony had access to the patches had no effect on the use of the risky patch. M.incompleta foraging behaviour seems relatively insensitive to short-term food stress or changing levels of mortality risk.
• 4 The differences between species in their exploitation of patches and reactions to F.subnuda are discussed in terms of the species’natural history.

     

Energy,predators and the behavior of feeding hummingbirds

Summary Current evidence suggests that many animals trade off energy gain against the risk of predation while feeding. In contrast, energetic considerations alone have proven successful in explaining and predicting the behaviour of feeding hummingbirds. This success may reflect the relative lack of natural predation on hummingbirds, but this study suggests that it may additionally reflect the lack of studies taking a predation perspective. In particular, when Anna's hummingbirds (Calypte anna) are faced with an obstructed view of their surroundings, they engage in behaviour suggestive of anti-predatory vigilance. In doing so, they voluntarily reduce their rate of energy intake. These birds also forgo better feeding opportunities that occur close to the ground, where observations suggest they are wary of opportunistic predators such as roadrunners (Geococcys californianus). While energy-based concepts will remain useful in the study of hummingbird feeding behaviour, the lack of predation on these birds should not be equated with an insensitivity to the risk of predation. This realization may lead to further insights into hummingbird-plant interactions, and hummingbird biology in general.