A sentinel system in the Florida scrub jay

One postulated advantage of living in groups is increased protection from predators. Living in small, permanent groups of stable composition allows for the formation of a sentinel system, in which individuals coordinate their vigilance. Florida scrub jays, Aphelocoma coerulescens coerulescens, live in family groups, and coordinate their vigilance into a sentinel system. Observed vigilance behaviour was compared with models generating similar behaviour patterns at random. The occurrence of one jay terminating vigilance in the same minute that another initiated vigilance was greater than expected by chance, and the occurrence of more than one vigilant jay at one time was less frequent than expected by chance. The seasonal pattern of scrub jay sentinel behaviour suggests that it functions primarily for predator detection, especially of raptors. Most aerial predator alarm calls were given by sentinels, and a sentinel was most likely to give the alarm call first.     

Information transfer and the facilitation and inhibition of feeding in a schooling fish

Synopsis Recent studies show that fish forage actively when perceived risk is low, but decrease foraging and increase vigilance when perceived risk is high. Isolated juvenile chum salmon,Oncorhynchus keta, were visually exposed to groups of conspecifics engaged in different activities to examine their ability to gain information about foraging opportunity and risk by interpreting conspecific behavior. Isolates ate most when exposed to feeding groups, less when exposed to nonfeedig groups, and least when exposed to alarmed groups. Isolates exposed to alarmed conspecifics also spent more time motionless than did fish exposed to either feeding or nonfeeding conspecifics. These findings indicate that schooling fish gain information by interpreting conspecific behavior, and are consistent with research showing that animals balance the conflicting demands of foraging and vigilance.     

Theory and method in studies of vigilance and aggregation

Predation is considered one of the most important selective pressures on free-ranging animals. Our understanding of it derives mainly from studies of individual vigilance (visual scanning of the surroundings beyond the immediate vicinity) and aggregation in prey. Vigilance bears a direct relationship to aggregation, because animals in groups may rely on associates for early warning of danger. This review addresses the relationship between vigilance and aggregation with particular attention to the prediction that individual vigilance declines with increasing group size. Contrary to most other animals studied, primates do not support the prediction. Exploring this, I examined the assumptions underlying vigilance theory in the light of primate behaviour. First I tested whether manual harvesting and upright processing of food as seen among primates might permit them to feed and scan simultaneously. I found no support for this idea. Next I examined the targets of primate vigilance and found that one component (within-group vigilance) might explain the differences between primates and other animals. Finally, I evaluated whether individual primates in large groups face a lower risk of predation than those in small groups. A conclusion was impossible, but by separating group-level from individual-level risk, I was able to identify several common circumstances in which group size would not predict individual risk or vigilance. These circumstances arose for primates and nonprimates alike. I concluded that the relationship of vigilance to aggregation is not straightforward. The absence of a group-size effect on vigilance among primates is probably due to functional differences in vigilance behaviour or safety in groups, not to methodological differences. Furthermore, future work on animal vigilance and aggregation must fully consider both the targets of glances, and the assumption that larger groups are safer from predators. I predict that animals will not relax vigilance in larger groups if conspecific threat increases with group size. Group size will not predict individual risk of predation nor individual vigilance rates when predators do not rely on surprise, or when predators select a small subset of highly vulnerable group members. Copyright 2000 The Association for the Study of Animal Behaviour.     

Vigilance and fitness in grey partridges Perdix perdix: the effects of group size and foraging-vigilance trade-offs on predation mortality

1. Vigilance increases fitness by improving predator detection but at the expense of increasing starvation risk. We related variation in vigilance among 122 radio-tagged overwintering grey partridges Perdix perdix (L.) across 20 independent farmland sites in England to predation risk (sparrowhawk Accipiter nisus L., kill rate), use of alternative antipredation behaviours (grouping and use of cover) and survival. 2. Vigilance was significantly higher when individuals fed in smaller groups and in taller vegetation. In the covey period (in early winter when partridges are in flocks), vigilance and use of taller vegetation was significantly higher at sites with higher sparrowhawk predation risk, but tall vegetation was used less by larger groups. Individuals were constrained in reducing individual vigilance by group size and habitat choice because maximum group size was determined by overall density in the area during the covey period and by the formation of pairs at the end of the winter (pair period), when there was also a significant twofold increase in the use of tall cover. 3. Over the whole winter individual survival was higher in larger groups and was lower in the pair period. However, when controlling for group size, mean survival decreased as vigilance increased in the covey period. This result, along with vigilance being higher at sites with increasing with raptor risk, suggests individual vigilance increases arose to reduce short-term predation risk from raptors but led to long-term fitness decreases probably because high individual vigilance increased starvation risk or indicated longer exposure to predation. The effect of raptors on survival was less when there were large groups in open habitats, where individual partridges can probably both detect predators and feed efficiently. 4. Our study suggests that increasing partridge density and modifying habitat to remove the need for high individual vigilance may decrease partridge mortality. It demonstrates the general principle that antipredation behaviours may reduce fitness long-term via their effects on the starvation-predation risk trade-off, even though they decrease predation risk short-term, and that it may be ecological constraints, such as poor habitat (that lead to an antipredation behaviour compromising foraging), that cause mortality, rather than the proximate effect of an antipredation behaviour such as vigilance.     

Vigilance and Collective Detection of Predators in Degus (Octodon degus)

Individuals of social and partially social species typically reduce their vigilance activity when foraging in groups. As a result, per capita risk of predation decreases and individuals allocate more time to foraging and other fitness rewarding activities. Reduction of per capita risk is hypothesized to occur because there are more individuals to detect potential predators. If so, collective (i.e. total) vigilance is expected to increase with foraging group size. Increased surveillance during group foraging may occur if group members scan independently of one another, or sequentially to avoid the overlapping of their vigilance bouts. Intriguingly, such coordinated vigilance assumes that individuals monitor not only the presence, but the vigilance behaviour of group mates. We used seasonal records on time budget and grouping patterns of individually marked degus (Octodon degus), a social rodent, to examine if (a) individual vigilance decreases and/or foraging increases with group size, (b) collective vigilance increases with group size and (c) foraging degus coordinate their vigilance. When foraging, degus decreased their individual vigilance and increased their foraging time when in larger groups. Despite this, degus in larger groups increased their collective vigilance, supporting the hypothesis that socially foraging degus decrease predation risk through an improved ability to detect and escape potential predators. Additionally, patterns of collective vigilance suggested that degus scan independently of each other and so, they do not coordinate their vigilance to prevent its temporal overlapping. This finding does not support that foraging degus monitor the vigilance activity of group mates.     

Thirteen-lined ground squirrel (Sciuridae: Spermophilus tridecemlineatus) antipredator vigilance decreases as vigilance cost increases

Vigilance is a commonly studied antipredator phenomenon. However, little research has been conducted on the effects of visual obstruction coupled with a manipulation of the cost of performing vigilance behaviours. We tested the hypothesis that antipredator vigilance decreases as the cost of performing antipredator vigilance increases in free-ranging thirteen-lined ground squirrels. We presented adult ground squirrels with peanut butter within six rectangular Plexiglas boxes (opaque on all sides; clear on top) ranging in length from 15.2 to 91.4 cm. Squirrels repeatedly entered and withdrew from the boxes, to scan visually the surrounding area. As box length increased and, therefore, the cost of the vigilance behaviour increased, the ground squirrels increased their mean time within the box per entry and decreased their alertness per unit of trial time. Squirrels took longer to traverse the longer boxes, but their travel time into the boxes was significantly slower than their travel time out of the boxes. We interpreted this difference in travel time to indicate a behavioural conflict that occurred only when the squirrels entered the box (food versus vigilance) and did not exist once the squirrels had begun to exit from the box. The hypothesis that a decrease in ground squirrel antipredator vigilance would benefit the squirrels by decreasing their time exposed to predators (i.e. decreased trial duration, which was determined by the squirrels' behaviour), was not supported. Copyright 1999 The Association for the Study of Animal Behaviour.     

Different Forms of Vigilance in Response to the Presence of Predators and Conspecifics in a Group-Living Mammal, the European Rabbit

In group‐living mammals, the major functions of vigilance are to detect the presence of predators and to monitor the movements of conspecific competitors, i.e. of potential opponents in agonistic encounters. The minimum distance to such a conspecific competitor that an animal considers safe is usually lower than to a predator, whereas the frequency of encounters with conspecifics is higher. Therefore, the acquisition of information about a predator or about a conspecific could lead to the existence of at least two different modes of vigilance behaviour. The aim of the present study was to describe and compare different forms of vigilance behaviour that European rabbits, Oryctolagus cuniculus, display in anti‐predator and social contexts. We conducted an observational study on individually marked animals from a field enclosure population. We recorded social interactions of the animals, the presence of aerial predators (common buzzard Buteo buteo), and the vigilance behaviour of the rabbits. We distinguished between two forms of vigilance of different intensity: subtle and overt. The frequencies of both forms of vigilance displayed by the rabbits differed significantly in occurrence, duration, and distribution over time. Females and males showed higher frequencies of overt but not subtle vigilance when buzzards were present. In contrast, the presence of conspecifics in close proximity affected the display of subtle but not overt vigilance: males increased the frequency of subtle vigilance when other males were close. Females increased subtle vigilance in proximity of males and females; however, this effect was only apparent in females with a more unstable social situation. In conclusion, European rabbits differentially increased two different forms of vigilance behaviour in social and anti‐predator contexts.     

Impact of increased predation risk on vigilance behaviour in a gregarious waterfowl,the Egyptian goose Alopochen aegyptiaca

Vigilance is amongst the most universal of anti‐predator strategies and commonly declines with increasing group size. We experimentally manipulated predation risk in a system with a known relationship between group size and vigilance levels to explore whether this relationship changes in response to elevated predation risk. We investigated the vigilance levels of Egyptian geese Alopochen aegyptiaca at eight golf courses in the western Cape, South Africa, to assess the perception of and reaction to predation risk. We manipulated predation risk by introducing trained Harris's hawks Parabuteo unicintus where avian predation was otherwise low or absent. The study confirmed the typical reduction in vigilance with group size on control sites, where the risk of predation is low. However, at experimental sites with elevated predation risk, a positive relationship between vigilance and group size was observed. We hypothesize that the mechanism for this relationship might be linked to social information transfer via copying behaviour and manipulation to induce vigilance. Thus, larger groups will have a higher probability of containing individuals with experience of elevated predation risk and their increased vigilance behaviour is copied by naïve individuals. This prediction is based on the intended outcome of introducing avian predation to make the geese feel less safe and to eventually leave the site as a management tool for controlling nuisance geese.     

Prey synchronize their vigilant behaviour with other group members

It is generally assumed that an individual of a prey species can benefit from an increase in the number of its group''s members by reducing its own investment in vigilance. But what behaviour should group members adopt in relation to both the risk of being preyed upon and the individual investment in vigilance? Most models assume that individuals scan independently of one another. It is generally argued that it is more profitable for each group member owing to the cost that coordination of individual scans in non-overlapping bouts of vigilance would require. We studied the relationships between both individual and collective vigilance and group size in Defassa waterbuck, Kobus ellipsiprymnus defassa, in a population living under a predation risk. Our results confirmed that the proportion of time an individual spent in vigilance decreased with group size. However, the time during which at least one individual in the group scanned the environment (collective vigilance) increased. Analyses showed that individuals neither coordinated their scanning in an asynchronous way nor scanned independently of one another. On the contrary, scanning and non-scanning bouts were synchronized between group members, producing waves of collective vigilance. We claim that these waves are triggered by allelomimetic effects i.e. they are a phenomenon produced by an individual copying its neighbour''s behaviour.     

Vigilance and social organization in two species of primates

Animals that live in groups may accrue anti-predator benefits by virtue of their association with other individuals. While a number of factors (e.g. habitat quality, group geometry, dominance rank) have been shown to affect the relationship between group living and individual rates of visual vigilance, sociality and vigilance have not been studied specifically in terms of the two competing demands they impose upon an individual's visual time. Two captive groups each of tamarins, Saguinus labiatus, and squirrel monkeys, Saimiri sciureus, were used in three related experiments to determine whether they looked at the social or non-social environment while they engated in a simulated foraging task. The tamarins, whose social behaviour is quite pacific and cooperative relative to the more hierarchical squirrel monkeys, looked at the non-social environment when they visually interrupted their foraging. Squirrel monkeys, however, looked more often at group mates. Detection of predators is probably more likely when an individual directs its attention not to conspecifics, but to the environment. Hence, social organizations in which individuals must pay social attention in order to monitor threats and avoid aggression may reduce individual rates of vigilance for predators. Failure to consider various targets of attention may well overestimate vigilance for predators by including other sorts of attentional phenomena in the measurement of vigilance.     

Vigilance and grouping in the southern African ground squirrel (Xerus inauris)

Animals may form groups in response to the foraging–vigilance trade‐off, through enhanced predator detection (collective detection hypothesis) or reduced predation risk to the individual (dilution hypothesis), allowing individuals to decrease vigilance levels. Both hypotheses predict decreasing individual vigilance levels with increasing group size; however, the collective detection hypothesis also predicts increasing overall group vigilance with increasing group size. However, in species in which vigilance and foraging are not mutually exclusive, where vigilance may not be as costly, neither of these hypotheses may apply. Here, we examine the relationship between group size and vigilance in the social Cape ground squirrel (Xerus inauris), a species that can combine foraging and vigilance behaviours. Ten groups were observed using scan sampling, measuring both group and individual vigilance and group size. A negative relationship existed between individual vigilance and group size and a positive relationship between group vigilance and group size. Therefore, in Cape ground squirrels, vigilance seems to be costly even though it can be combined with foraging behaviours. Furthermore, group vigilance behaviour gives support to the collective detection hypothesis, whilst individual vigilance gives support to both hypotheses.     

Vigilance Decreases with Time at Loafing Sites in Gulls (Larus spp.)

Understanding how animals partition effort between vigilance for predators and other conflicting activities has been a mainstay of animal behaviour research. Classical theories implicitly assume that animals alternate between high and low vigilance states over short timescales, but that average effort invested in vigilance is constant over an extended bout of such alternations. However, one recent model suggests that vigilance should be adjusted dynamically to short‐term changes in the perception of predation risk and would tend to decrease with time. Indeed, as time passes by without disturbances, perception of the need for vigilance should decrease and prey animals may allocate more time to competing activities. Here, we examined how the proportion of sleeping gulls (Larus spp.) in loafing groups changed over time. Sleeping gulls can only maintain low levels of vigilance against external threats (compared to alert individuals), and we predicted that the proportion of sleeping gulls at loafing sites should increase over time when no disturbances occur. Statistically significant changes in the proportion of sleeping gulls as a function of time occurred in the majority of sequences and an increase was observed significantly more often than predicted by chance alone. This temporal pattern cannot be caused by reduction in hunger levels because gulls are not feeding at loafing sites. The results indicate that vigilance can be adjusted dynamically in response to short‐term temporal changes in the perception of predation risk.     

Negotiation may lead selfish individuals to cooperate: the example of the collective vigilance game

Game-theoretical models have been highly influential in behavioural ecology. However, these models generally assume that animals choose their action before observing the behaviour of their opponents while, in many natural situations, individuals in fact continuously react to the actions of others. A negotiation process then takes place and this may fundamentally influence the individual attitudes and the tendency to cooperate. Here, I use the classical model system of vigilance behaviour to demonstrate the consequences of such behavioural negotiation among selfish individuals, by predicting patterns of vigilance in a pair of animals foraging under threat of predation. I show that the game played by the animals and the resulting vigilance strategies take radically different forms, according to the way predation risk is shared in the pair. In particular, if predators choose their target at random, the prey respond by displaying moderate vigilance and taking turns scanning. By contrast, if the individual that takes flight later in an attack endures a higher risk of being targeted, vigilance increases and there is always at least one sentinel in the pair. Finally, when lagging behind its companion in fleeing from an attacker becomes extremely risky, vigilance decreases again and the animals scan simultaneously.     

The Context and Quality of Social Relationships Affect Vigilance Behaviour in Wild Chimpanzees

Previous studies of vigilance behaviour have focused mainly on the influence of predation threat, whereas the influences of conspecific factors, such as within‐group threats, are relatively unstudied. To elucidate the influences of conspecific factors, this study examined vigilance behaviour in wild chimpanzees (Pan troglodytes schweinfurthii) in Mahale Mountains National Park, Tanzania. Vigilance level was lower during foraging than during resting, which indicated a conflict between vigilance and foraging activity. In addition, vigilance level was higher when chimpanzees were on the ground where an encounter with leopards (Panthera pardus) is likely than when the chimpanzees were in trees. Males, but not females, increased their level of vigilance as the number of individuals within 3 m increased. In both males and females, daily party size – an index of group cohesion – did not affect the vigilance level. The level of maternal vigilance was higher when a dependent infant was separated from its mother than when the offspring was in contact with its mother. Both males and females increased their vigilance when a less‐associated group member was nearby, when compared with when there was no less‐associated group member nearby. This finding suggests that variation in relationship quality influences the vigilance level and that individuals need to increase their level of vigilance when the level of within‐group threats is high. This study indicated that variation in vigilance cannot be understood unless conspecific factors, such as variation in the relationship quality with associates, are considered.     

Evolution of anti-predator traits in response to a flexible targeting strategy by predators

Anti-predator benefits increase with vigilance rate and group size in many species of animal, while simultaneously resource intake rates usually decrease. This implies that there is an optimal group size and vigilance rate that will maximize individual fitness. While this basic theory of vigilance has been modelled and tested extensively, it has often been assumed that the predator represents a 'fixed-risk' such that groups of prey are essentially independent entities that exert little or no effect on one another either directly or indirectly. We argue that this is an over-simplification, and propose that the behaviour of one group of prey will likely affect the fitness of another local group of prey if the predator preferentially attacks the most vulnerable group-rather than attack both with constant rates. Using a numerical simulation model, we make the first examination of this game and allow the prey to dynamically evolve both optimal group size distributions between two habitats and vigilance rates in response to a predator with a preference for whichever group is the more vulnerable. We show that the density of prey in the population and the sensitivity of a predator to differences in prey vulnerability are likely to drive the dynamics of such a game. This novel approach to vigilance theory opens the door to several challenging lines of future research, both experimental and theoretical.     

集群数量和采食距离对储草期布氏田鼠警戒频次的影响

警戒行为是动物对环境中潜在危险做出的反应。为探究影响警戒行为的因素,作者在内蒙古锡林郭勒典型草原区进行了集群数量和采食距离对储草期自由生活状态下布氏田鼠(Lasiopodomys brandtii)警戒频次影响的实验。通过标志重捕法和人工去除法设定3个集群数量梯度(11只、6只和3只),利用人工食物站设置4个采食距离梯度(5 m、10 m、20 m、30 m),共12个梯度组合,每个组合分别观察60次采食过程,共计观察720次,并记录采食过程中的警戒频次。对数据进行双因素方差分析,集群规模和采食距离对布氏田鼠警戒行为频次均有极显著影响(P0.01)。随着集群数量减小,布氏田鼠单次采食的警戒频次显著增加;采食距离越远,单次采食的警戒频次越高;集群数量和采食距离之间还存在极显著的交互作用(P0.01)。集群数量增大意味着采食距离的增加,为保证储草效率,布氏田鼠种群会在增加集群数量和减小采食距离之间权衡,最终集群数量维持在中等水平,支持了最优集群理论。     

The presence of an avian co-forager reduces vigilance in a cooperative mammal

Many animals must trade-off anti-predator vigilance with other behaviours. Some species facilitate predator detection by joining mixed-species foraging parties and ‘eavesdropping’ on the predator warnings given by other taxa. Such use of heterospecific warnings presumably reduces the likelihood of predation, but it is unclear whether it also provides wider benefits, by allowing individuals to reduce their own vigilance. We examine whether the presence of an avian co-forager, the fork-tailed drongo (Dicrurus adsimilis), affects rates of vigilance (including sentinel behaviour) in wild dwarf mongooses (Helogale parvula). We simulate the presence of drongos—using playbacks of their non-alarm vocalizations—to show that dwarf mongooses significantly reduce their rate of vigilance when foraging with this species. This is, to our knowledge, the first study to demonstrate experimentally that a mammal reduces vigilance in the presence of an avian co-forager.     

Stress During Adolescence Shapes Performance in Adulthood: Context‐Dependent Effects on Foraging and Vigilance

Exposure to chronic stress during adolescence can shape behaviour, cognition and physiology in adulthood, but the consequences of these long‐term changes remain unclear. Prior studies reporting altered performance following exposure to stress in adolescence have generally interpreted lasting changes as impairments. However, we have recently shown that exposure to chronic unpredictable stress during adolescence (from post‐natal days 30–70) can enhance performance in a context‐dependent manner during a foraging task. Increases in foraging performance, (previously measured by the number of rewards obtained), are often associated with trade‐offs in other behaviours, such as vigilance. Here, we examined the effect of stress exposure in adolescence on adult foraging in male Sprague Dawley rats to determine (1) whether the increase in foraging performance exhibited by animals exposed to stress in adolescence is balanced by a decrease in vigilance, and (2) whether stress in adolescence alters time allocation between foraging and vigilance behaviours in low‐ and high‐threat conditions. We found no evidence of a trade‐off between foraging and vigilance; under low‐threat conditions, rats exposed to stress in adolescence spent more time being vigilant compared with unstressed rats, suggesting that exposure to stress in adolescence enhances anticipation of threat in adulthood. Under high‐threat conditions, adolescent‐stressed and unstressed rats did not differ in foraging and vigilance behaviours. Given that we have previously found that rats exposed to stress in adolescence nearly double food intake under high‐threat, and we now show that high‐performing rats do not spend more time foraging, it appears that stress exposure in adolescence may enhance foraging efficiency (food consumed/time) under high‐threat conditions rather than time allocation between foraging and competing behaviours. We also examined the relationship, at the level of the individual, between foraging performance and foraging and vigilance behaviours. We found that changes in individual foraging performance between low‐ and high‐threat conditions were independent of behavioural changes (i.e. both highly and poorly performing rats were equally active and contacted a similar number of patches). This suggests that the ability to obtain many rewards under high‐threat conditions may be related to efficiency, rather than the frequency of foraging and effort‐related behaviours.     

Habitat but not group size or recent predator activity affect corvid collective vigilance at carcasses

Vigilance is an important anti-predator behaviour that can be an indicator of the predation risk faced by potential prey animals. Here, we assess the collective vigilance, or the vigilance level of an entire group, of corvids (Family: Corvidae) at experimentally placed carcasses in a desert environment in Australia. Specifically, we explore the relationship between collective vigilance levels and the habitat in which the carcass was placed, the time since a potential predator (dingo Canis dingo, wedge-tailed eagle Aquila audax or red fox Vulpes vulpes) was present at a carcass, and the group size of corvids around the carcass. We found that corvids are more vigilant in open habitat, but that group size and the recent presence of a potential predator does not affect the collective vigilance behaviour of corvids. The results demonstrate the important link between habitat and vigilance, and that animals may adopt anti-predator behaviours irrespective of the size of the group in which they occur or the recent presence of a potential predator.     

Effect of Male Emigration on the Vigilance Behavior of Coresident Males in White-Faced Capuchins (Cebus capucinus)

I examined the vigilance behavior of male Cebus capucinus residing in four groups in Santa Rosa Park, Costa Rica. One male emigrated from each of three study groups, providing ideal experimental conditions for examining the effect of coresident males on male vigilance behavior (social and non-social). Following the predator and conspecific defense theories for the adaptive value of male vigilance behavior and the occurrence of multimale groups, I predicted that male nonsocial vigilance would increase after the emigration of a coresident male. My prediction was supported in only one of the three study groups. Males in two groups decreased their nonsocial vigilance after the emigration of a coresident male, which was probably influenced by seasonal changes and the peripherality of one group's coresident male before his emigration. The social vigilance hypothesis proposes that for species with rigid dominance hierarchies, social vigilance should increase with an increase in same sex competitors. Therefore, I predicted that male social vigilance would decrease after the emigration of a coresident male. The hypothesis is generally supported in my findings, as two of the three groups displayed a decrease in social vigilance after the emigration of a coresident male. Increased social vigilance in the third group is best explained by the change in the alpha male's social behavior after the emigration of his only coresident male.