Diet quality in a wild grazer declines under the threat of an ambush predator

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator–prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.     

Effects of lions on behaviour and endocrine stress in plains zebras

Living under predation risk may alter both behaviour and physiology of potential prey. In extreme cases, such alterations may have serious demographic consequences, and recent studies support that non‐lethal effects of predation may have broad ecological consequences. However, behavioural and physiological responses to predation risk may be related to trade‐offs associated with resource acquisition and direct predation risk. We validated an enzyme‐linked immunoassay (EIA) for non‐invasive monitoring of stress in plains zebras (Equus quagga) from faecal material. We used this assay in combination with behavioural data to assess if plains zebras living with and without lions (Panthera leo) in a mountain savannah in southern Africa differed in behaviour and physiology, and if such differences were influenced by seasons with contrasting resource availability. Zebra group sizes did not differ between areas with and without lions, but zebra groups had more juveniles in an area with lions than groups in an area without lions, but only during the wet season. Similarly, we observed differences in individual vigilance, foraging behaviour and stress hormone concentrations, but all these differences were influenced by seasons. Despite these seasonal influences, our study did not suggest that zebras in an area with lions spent a higher proportion of time being vigilant, a lower proportion of time foraging, or had higher stress hormone levels. Our results instead suggest that zebras' responses to lion presence were highly context dependent and the result of complex interactions between resource abundance and cues about predation risk. Because of the obvious ecological and evolutionary ramifications of such findings, we argue that further research is needed to define the spatial and temporal scales over which predators impose indirect effects on their prey.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes

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Bottom-up and top-down processes in African ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids

African ungulate populations appear to be limited principally by their food resources. Within ungulate communities, plains zebras coexist with grazing bovids of similar body size, but rarely are the dominant species. Given the highly effective nutritional strategy of the equids and the resistance of zebras to drought, this is unexpected and suggests that zebra populations may commonly be limited by other mechanisms. Long-term research in the Serengeti ecosystem and in the Kruger National Park suggests that zebra could be less sensitive to food shortage, and more sensitive to predation, than grazing bovids: if this is a general principle, then, at a larger scale, resource availability should have a weaker effect on the abundance of zebra than on grazing ruminants of similar body size (wildebeest and buffalo), and zebras should be relatively more abundant in ecosystems where predators are rare or absent. We test these expectations using data on 23 near-natural ecosystems in east and southern Africa. The abundance of wildebeest is more closely related to resources than is that of zebra; buffalo are intermediate. We show that hyena densities are closely correlated with those of lions, and use the abundance of lions as an index of predation by large predators. The numerical response of lions to increases in the abundance of their prey was linear for mesoherbivores, and apparently so for the three species alone. Finally, the abundance of zebra relative to grazing bovids is lower in ecosystems with high biomasses of lions. These results indicate that zebras may commonly be more sensitive to top-down processes than grazing bovids: the mechanism(s) have not been demonstrated, but predation could play a role. If it is true, then when numbers of the large mammalian predators decline, zebra populations should increase faster than buffalo and wildebeest.     

The Effect of Simulated African Wild Dog Presence on Anti‐predator Behaviour of Kudu and Impala

In this study, we examined the behavioural, temporal and spatial effects of simulated African wild dog (Lycaon pictus) presence on its two main prey species: kudu (Tragelaphus strepsiceros) and impala (Aepyceros melampus). We spread African wild dog faeces around waterholes and played African wild dog sounds at different intervals to mimic immediate and non‐immediate predation pressure. We looked at anti‐predator behaviour at both a herd and individual level and distinguished between high‐quality (detracts from all other activities), high‐cost vigilance and low‐quality (used to monitor the surrounding in spare time), low‐cost vigilance to determine costs involved. We found that simulated African wild dog presence had little effect on anti‐predator behaviour of their free‐ranging prey. Only when immediate predation risk was mimicked did kudu invest in (additional) high‐quality vigilance, whereas impala showed no response. Regardless of direct cues of African wild dog presence, behavioural adjustments to reduce predation risk were primarily based on environmental factors such as time of the day and broad‐scale habitat structure. Predators have been shown to utilize waterholes to hunt, and prey species are therefore likely to maximize anti‐predator behaviour in this high‐risk environment based on environmental variables affecting predation risk, the main predator within the system, and water requirements, leaving little flexibility to respond to (simulated) African wild dog presence.     

Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

Dynamic Risk Assessment: Prey Rapidly Adjust Flight Initiation Distance to Changes in Predator Approach Speed

The pre‐eminent model of flight initiation distance assumes that the function relating predation risk to distance between predator and prey is constant. However, the risk–distance function can change dramatically during approaches by predators. Changes in predator behavior during approach and in availability of benefits (e.g. food or potential mates) may alter risks and/or costs during encounters. Thus, prey should be able to respond appropriately to changes in cues to risk, such as predator approach speed. Under the assumption that prey assess risk in real time, it was predicted that flight initiation distance (distance between predator and prey when escape begins) decreases when approach speed increases and increases when approach speed decreases during an encounter. Effects of single, abrupt changes from slower to faster approach or the reverse were studied in a lizard, Anolis lineatopus. Flight initiation distances were determined solely by final approach speed, being nearly identical for: (1) continuously fast approaches and approaches initially at the slower and finally at the faster speed and (2) for continuously slower approaches and approaches initially at faster and finally at slower speed. Escape should be adjusted to match changes in risk and cost caused by changes in predator behavior, ability to escape, and costs of escape as attacks unfold. A recent model by Broom and Ruxton [Behavioural Ecology (2004) vol. 16, pp. 534—540] predicts that cryptic prey should stay motionless until detected, then flee immediately. Our results suggest that current escape models can be applied to prey escape strategies when cues to risk change, by assuming that prey base decisions on the current relationship between risk and distance. Empirical studies are needed to test predictions concerning continuous risk assessment.     

Temperature effects on ballistic prey capture by a dragonfly larva

Understanding the effects of temperature on prey–predator interactions is a key issue to predict the response of natural communities to climate change. Higher temperatures are expected to induce an increase in predation rates. However, little is known on how temperature influences close‐range encounter of prey–predator interactions, such as predator's attack velocities. Based on the speed–accuracy trade‐off concept, we hypothesized that the increase in predator attack velocity by increasing temperature reduces the accuracy of the attack, leading to a lower probability of capture. We tested this hypothesis on the dragonfly larvae Anax imperator and the zooplankton prey Daphnia magna. The prey–predator encounters were video‐recorded at high speed, and at three different temperatures. Overall, we found that (1) temperature had a strong effect on predator's attack velocities, (2) prey did not have the opportunity to move and/or escape due to the high velocity of the predator during the attack, and (3) neither velocity nor temperature had significant effects on the capture success. By contrast, the capture success mainly depended on the accuracy of the predator in capturing the prey. We found that (4) some 40% of mistakes were undershooting and some 60% aimed below or above the target. No lateral mistake was observed. These results did not support the speed–accuracy trade‐off hypothesis. Further studies on dragonfly larvae with different morphological labial masks and speeds of attacks, as well as on prey with different escape strategies, would provide new insights into the response to environmental changes in prey–predator interactions.     

Do Lions Panthera leo Actively Select Prey or Do Prey Preferences Simply Reflect Chance Responses via Evolutionary Adaptations to Optimal Foraging?

Research on coursing predators has revealed that actions throughout the predatory behavioral sequence (using encounter rate, hunting rate, and kill rate as proxy measures of decisions) drive observed prey preferences. We tested whether similar actions drive the observed prey preferences of a stalking predator, the African lion Panthera leo. We conducted two 96 hour, continuous follows of lions in Addo Elephant National Park seasonally from December 2003 until November 2005 (16 follows), and compared prey encounter rate with prey abundance, hunt rate with prey encounter rate, and kill rate with prey hunt rate for the major prey species in Addo using Jacobs' electivity index. We found that lions encountered preferred prey species far more frequently than expected based on their abundance, and they hunted these species more frequently than expected based on this higher encounter rate. Lions responded variably to non-preferred and avoided prey species throughout the predatory sequence, although they hunted avoided prey far less frequently than expected based on the number of encounters of them. We conclude that actions of lions throughout the predatory behavioural sequence, but particularly early on, drive the prey preferences that have been documented for this species. Once a hunt is initiated, evolutionary adaptations to the predator-prey interactions drive hunting success.     

Escape and alerting responses by Balearic lizards (Podarcis lilfordi) to movement and turning direction by nearby predators

Escape theory predicts that prey monitoring an approaching predator delay escape until predation risk outweighs costs of fleeing. However, if a predator is not detected until it is closer than the optimal flight initiation distance (FID = distance between predator and prey when escape begins), escape should begin immediately. Similarly, if a change in a nearby predator’s behavior indicates increased risk, the optimal FID increases, sometimes inducing immediate escape. If a predator that has been standing immobile near a prey suddenly turns toward the prey, greater risk is implied than if the predator turns away. If the immobile predator suddenly moves its foot without turning, it might be launching an attack. Therefore, we predicted that frequency of fleeing and preparation to flee are greater when a predator turns toward than away from prey and that frequency of fleeing when a predator suddenly moves decreases as distance between predator and prey increases. We verified these predictions in the Balearic lizard Podarcis lilfordi in field experiments in which an investigator simulated the predator. Lizards fled and performed alerting responses indicating readiness to flee more frequently when the predator turned toward than away from them, and fled more frequently the nearer the predator.     

Effects of predator behavior and proximity on risk assessment by Columbian black-tailed deer

In predator-prey encounters, many factors influence risk perceptionby prey and their decision to flee. Previous studies indicatethat prey take flight at longer distances when they detect predatorsat longer distances and when the predator's behavior indicatesthe increased likelihood of attack. We examined the flight decisionsof Columbian black-tailed deer (Odocoileus hemionus columbianus)using an approaching human whose speed, directness of approach,directness of gaze, and simulated gun carrying varied. Deerfled at greater distances when approached more quickly and directly,and there was a concave-down quadratic trend in the relationshipbetween the distances at which the predator began its approachand at which the deer became alert (alert distance [AD]), indicatingthat deer have a zone of awareness beyond which there is a delayin detecting an approaching predator. Time spent assessing theapproacher (assessment time) was shorter during faster approachesand was positively related with AD. Deer fled at longer distancesand had shorter assessment times when they were already alertto the predator at the initiation of approach. Males fled atshorter distances than females when approached during the gun-holdingcondition, and males had shorter assessment times than femaleswhen the approacher averted his gaze. Such sex differences inrisk assessment might reflect male motivation during the matingseason as well as exposure to human hunting. We suggest thatrisk assessment is affected the by the predator's behavior,the state of awareness of the prey, and the distance at whichthey detect the predator.     

The relationship between vegetation density and its protective value depends on the densities and traits of prey and predators

Densities of submerged vegetation and those of associated animals tend to co‐vary. This relationship is often attributed to the positive correlation between the density of vegetation and its protective value against predators. However, two counteracting basic elements underlying this paradigm limit its generality. That is, increasing vegetation density should result in decreased predator–prey encounters, whilst at the same time predator–prey encounters should increase as animal densities increase. These two mechanisms should thus counteract each other when the densities of vegetation and associated animals, including both prey and predators, co‐vary. Experimental designs that expose fixed densities of prey and predators to varying densities of vegetation assess only the former mechanism and may thus not properly evaluate the protective value of vegetation in such conditions. By contrast, designs that mimic the naturally co‐varying organism densities test both mechanisms and thus their counteractive impacts on predator–prey encounters. We compared the outcomes of the two alternative designs and carried out additional experiments to explain the putative discrepancy. Increasing vegetation density (mimics of Potamogeton pectinatus) enhanced prey (Daphnia magna) survival only when fixed densities of prey and predators (Perca fluviatilis or Rutilus rutilus) were used. When the animal densities were allowed to co‐vary with vegetation density, vegetation had no impact on prey survival. Instead, prey survival was determined by the aggregate density of prey and predators, shaped by the species‐specific traits of the latter. Thus, the impact of the increased animal densities overrode the impact of the increased vegetation density on predator–prey encounters. It may be insufficient to attribute the co‐variation of vegetation, prey and predator densities simply to the association between vegetation density and its protective value. Increased food resources and reduced competition within vegetation may promote prey and thereby also predator abundance to a greater extent than previously thought.     

The effectiveness of short‐term fox control in protecting a seasonally vulnerable species,the Eastern Long‐necked Turtle (Chelodina longicollis)

Reducing predation by introduced predators on seasonally vulnerable prey is of interest to biodiversity and game managers around the world. In Australia, the Red Fox (Vulpes vulpes) is a significant predator of freshwater turtle nests, destroying up to 93% of nests. We used a nonrandomized intervention study to assess the effectiveness of a short‐term (3‐week) but broad‐scale baiting operation in reducing the level of nest predation on artificial turtle nests around a complex lake system during a major flooding event in north‐western Victoria. Estimates of fox occupancy declined from 0.58 (0.44–0.70 95% CI) to 0.34 (0.21–0.46 95% CI) following fox control. Modelling of nest‐survival rates indicated there was no significant change in survival rates. Effective short‐term predator control to protect seasonally vulnerable prey is desirable and achievable. Knowledge of underlying predator density, predator–bait encounter and consumption rates, and the optimal duration of short‐term control is needed to reduce the risk to prey.     

The role of size and colour pattern in protection of developmental stages of the red firebug (Pyrrhocoris apterus) against avian predators

We investigated how predator/prey body‐size ratio and prey colour pattern affected efficacy of prey warning signals. We used great and blue tits (Parus major and Cyanistes caeruleus), comprising closely related and ecologically similar bird species differing in body size, as experimental predators. Two larval instars and adults of the unpalatable red firebug (Pyrrhocoris apterus), differing in body size and/or coloration, were used as prey. We showed that prey body size did not influence whether a predator attacked the prey or not during the first encounter. However, smaller prey were attacked, killed, and eaten more frequently in repetitive encounters. We assumed that body size influences the predator through the amount of repellent chemicals better than through the amount of optical warning signal. The larger predator attacked, killed and ate all forms of firebug more often than the smaller one. The difference between both predators was more pronounced in less protected forms of firebug (chemically as well as optically). Colour pattern also substantially affected the willingness of predators to attack the prey. Larval red–black coloration did not provide a full‐value warning signal, although a similarly conspicuous red‐black coloration of the adults reliably protected them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 890–898.     

Increased prey vulnerability as a result of prey-prey interactions

This study examines indirect effects in a trophic system with three levels, consisting of two prey species, a top predator, and an intermediate predator. Qualitative data showed that the activity of both the top predator Aeshna juncea (Odonata) and the active prey Heterocope saliens, (Copepoda) caused bouts of swimming in the sedentary prey Sida crystallina (Cladocera). These swimming bouts caused encounters, reactions, attacks and captures of S. crystallina by the intermediate predator Coenagrion hastulatum (Odonata). Quantitative data showed that C. hastulatum had a higher encounter frequency and a higher attack frequency on the sedentary prey when the active prey was present. This result was an effect of encounters between the two prey which increased swimming activity of the sedentary S. crystallina. The results suggest that interactions between prey 1 and prey 2, and between prey and predators, could influence the structure of natural communities.     

Responses of Spotted Hyenas to Lions Reflect Individual Differences in Behavior

Competition among mammalian carnivores can be particularly intense and can influence population dynamics at lower trophic levels. One strategy employed by carnivores to minimize potentially costly interspecific competition is avoidance of dominant species. Recent research has highlighted the importance of consistent individual differences in behavior (i.e. temperament traits) in understanding behavioral variation during predator–prey interactions and intraspecific interactions. However, the importance of such individual differences to interspecific competition has received little attention. Here, we examined the responses of spotted hyenas (Crocuta crocuta) to their primary competitors, African lions (Panthera leo), to (1) determine whether hyenas avoid lions and (2) evaluate the potential importance of individual differences in behavior during interspecific competition. Spotted hyenas and lions co‐occur throughout much of Africa and are vigorous competitors. Whereas lions sometimes kill hyenas and steal their food, lions also represent a source of food for hyenas via scavenging. Using audio playback experiments, we found that hyenas do not uniformly avoid potential encounters with lions. Indeed, we noted considerable variation among individuals in their responses to lion roars, and this variation reflected consistent individual differences in risk‐taking and vigilance tendencies. Individual differences in vigilance behavior were specific to interactions with lions. We conclude that individual differences in behavior have the potential to play an important role in determining the nature and outcome of interspecific competition.     

Plesiomorphic Escape Decisions in Cryptic Horned Lizards (Phrynosoma) Having Highly Derived Antipredatory Defenses

Escape theory predicts that the probability of fleeing and flight initiation distance (predator–prey distance when escape begins) increase as predation risk increases and decrease as escape cost increases. These factors may apply even to highly cryptic species that sometimes must flee. Horned lizards (Phrynosoma) rely on crypsis because of coloration, flattened body form, and lateral fringe scales that reduce detectability. At close range they sometimes squirt blood‐containing noxious substances and defend themselves with cranial spines. These antipredatory traits are highly derived, but little is known about the escape behavior of horned lizards. Of particular interest is whether their escape decisions bear the same relationships to predation risk and opportunity costs of escaping as in typical prey lacking such derived defenses. We investigated the effects of repeated attack and direction of predator turning on P. cornutum and of opportunity cost of fleeing during a social encounter in P. modestum. Flight initiation distance was greater for the second of two successive approaches and probability of fleeing decreased as distance between the turning predator and prey increased, but was greater when the predator turned toward than away from a lizard. Flight initiation distance was shorter during social encounters than when lizards were solitary. For all variables studied, risk assessment by horned lizards conforms to the predictions of escape theory and is similar to that in other prey despite their specialized defenses. Our findings show that these specialized, derived defenses coexist with a taxonomically widespread, plesiomorphic method of making escape decisions. They suggest that escape theory based on costs and benefits, as intended, applies very generally, even to highly cryptic prey that have specialized defense mechanisms.     

Prey Responses to Predator's Sounds: A Review and Empirical Study

Many animals assess their risk of predation by listening to and evaluating predators' vocalizations. We reviewed the literature to draw generalizations about predator discrimination abilities, the retention of these abilities over evolutionary time, and the potential underlying proximate mechanisms responsible for discrimination. Broadly, we found that some prey possess an ability to respond to a predator after having been evolutionarily isolated from a specific predator (i.e., predators are allopatric) and that some prey are predisposed to respond to certain types of predators that they coevolved with but without having ecological experience. However, these types of studies are lacking, and relatively, few studies have examined predator discrimination abilities in ungulates. To begin addressing these knowledge gaps, we performed field experiments on Mule deer (Odocoileus hemionus) in which we investigated the ability of deer to discriminate among familiar predators [coyotes (Canis latrans) and mountain lions (Puma concolor)] and an evolutionary relevant predator with which deer have had no recent exposure [locally extinct wolves (Canis lupus)]. We found that Mule deer respond to and discriminate among predators based on predator vocalizations and have retained an ability to respond to wolves that have been extinct from the study area since the early 20th century. Previous playback studies have shown that responses vary among human‐habituated and non‐habituated populations and differ according to human proximity. Deer greater than 0.5 km from human residences allocated more time to heightened responses both before and after stimulus playback. Our findings may help predict how prey–predator interactions may change as a result of the recovering wolf population with a basis in ecological and evolutionary experience in predator discrimination and desensitization.     

Multi-behavioral strategies in a predator–prey game: an evolutionary algorithm analysis

Behavioral games between predators and prey often involve two sub-games: 'pre-encounter' games affecting the rate of encounter between predators and prey (e.g. predator–prey space games, Sih 2005 ), and 'post-encounter' games that influence the outcome of encounters (e.g. waiting games at prey refugia, Hugie 2003 , and games of vigilance, Brown et al. 1999 ). Most models, however, focus on only one or the other of these two sub-games.
I investigated a multi-behavioral game between predators and prey that integrated both pre-encounter and post-encounter behaviors. These behaviors included landscape-scale movements by predators and prey, a type of prey vigilance that increases immediately after an encounter and then decays over time ('ratcheting vigilance'), and predator management of prey vigilance. I analyzed the game using a computer-based evolutionary algorithm. This algorithm embedded an individual-based model of ecological interactions within a dynamic adaptive process of mutation and selection. I investigated how evolutionarily stable strategies (ESS) varied with the predators' learning ability, killing efficiency, density and rate of movement. I found that when predators learn prey location, random prey movement can be an ESS. Increased predator killing efficiency reduced prey movement, but only if the rate of predator movement was low. Predators countered ratcheting vigilance by delaying their follow-up attacks; however, this delay was reduced in the presence of additional predators. The interdependence of pre-and post-encounter behaviors revealed by the evolutionary algorithm suggests an intricate co-evolution of multi-behavioral predator–prey behavioral strategies.