An Experimental Test of Predator Detection Rates Using Groups of Free-living Emus

Abstract Improved predator detection is often stated to be one of the principal benefits of social foraging. However, actual field evidence supporting this assumption is scarce. This may be the result of the fact that most observations are conducted on social animals acting in the absence of an acute predation threat, yet the benefits of grouping come to the fore in that brief moment when an individual's life is at risk. As predation attempts are typically rare in nature, experimental manipulations are necessary to further explore the costs and benefits of social foraging. This study utilizes simple predator simulations (by humans) to experimentally test the predator‐detecting abilities and escape strategies of groups of free‐living emus Dromaius novaehollandiae. Emus in larger groups spent less time in vigilance and more time foraging. Nonetheless, the combined vigilance of group members ensured that emus detected the ‘simulated predator’ sooner as group size increased. After detecting the ‘predator’, larger groups waited longer until opting to flee, and then spent less time and energy doing so. Thus, the results of this study provide experimental evidence that emus benefit from grouping in terms of both the ‘many‐eyes effect’ and the ‘dilution effect’.     

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.     

You watch my back,I'll watch yours: Emergence of collective risk monitoring through tacit coordination in human social foraging

Predation risk is a significant concern when social animals including humans engage in foraging. When people search for resources together, individuals often find themselves in a producer–scrounger game, in which some individuals bear the cost of risk monitoring while others can free ride on those efforts. A theoretically rational strategy is to mix foraging and risk monitoring randomly with the same probability across all members, but such uncoordinated action often yields inefficiencies of under- or over-supply of risk monitoring in a group. Here, we examined whether people could spontaneously develop a coordinated risk-monitoring system, alternating vigilance and foraging in a pair. Given that human cooperation is vulnerable to fear of exploitation and emotional arousal under risk, we hypothesized that such sources of anxiety would be potential disruptors to coordination. In a laboratory experiment, two participants worked on a “treasure hunt” task simultaneously, in which they chose between low or high vigilance against predators during foraging without verbal communication. If one chose high vigilance with personal cost, it yielded a spillover benefit to the other. Besides behavioral choices, each participant's physiological arousal (skin conductance response) and cognitive effort (tonic pupil dilation) were measured during the task. Results showed that some pairs were actually able to develop a role-alternating system over time through tacit coordination, but coordinated action was also vulnerable to anxiety and mistrust among participants. Overall, these results imply that, besides the mutual behavioral control that often characterizes repeated interaction, cognitive control of emotional arousal may be a critical psychological factor for the emergence of coordinated cooperation.     

夏季雌性藏羚昼间行为时间分配及活动节律

2003 至2005 年的6 ～ 9,采用目标动物观察法在可可西里观察夏季雌性藏羚的昼间行为。将雌性藏羚的行为分为觅食、警戒、卧息、移动和“其他”5 种类型,各类行为所占时间比例分别为59.1% 、7.3% 、19.4% 、13.0% 和1.3% 。觅食消耗的时间最多,是藏羚的常见行为,卧息、移动和警戒次之,而“其他”行为消耗时间最短。觅食、警戒、卧息和移动的累计时间在不同时段之间均存在显著(P<0.05) 或极显著(P< 0.01)差异。觅食行为存在3 个高峰,分别在10:00 ～11:00,13: 00 ～14: 00 和18: 00 ～19: 00;警戒的高峰期出现在早上08: 00 ～09 00,在15: 00 ～16:00 以及17: 00 ～18: 00 也分别出现小的高峰,但不明显;卧息行为表现为双峰形,分别为11: 00 ～12: 00 和16: 00 ～17: 00。移动行为也表现为双峰,出现在08: 00 ～09: 00 和15:00 ～16:00; “其他”行为在各时段间没有明显的变化趋势。     

Coordination and synchronisation of anti-predation vigilance in two crane species

Much of the previous research on anti-predation vigilance in groups has assumed independent scanning for threats among group members. Alternative patterns that are based on monitoring the vigilance levels of companions can also be adaptive. Coordination of vigilance, in which foragers avoid scanning at the same time as others, should decrease the odds that no group member is alert. Synchronisation of vigilance implies that individuals are more likely to be vigilant when companions are already vigilant. While synchronisation will increase the odds that no one is vigilant, it may allow a better assessment of potential threats. We investigated temporal sequences of vigilance in family flocks consisting of two parents and at most two juveniles in two species of cranes in coastal China. We established whether the observed probability that at least one parent is alert was greater (coordination) or lower (synchronisation) than that predicted under the null hypothesis of independent vigilance. We documented coordination of vigilance in common cranes (Grus grus) foraging in an area with high potential for disturbance by people. We documented synchronisation of vigilance in red-crowned cranes (Grus japonensis) in the less but not in the more disturbed area. Coordination in small flocks leads to high collective vigilance but low foraging rates that may not be suitable in areas with low disturbance. We also argue that synchronisation should break down in areas with high disturbance because periods with low vigilance are riskier. Results highlight the view that temporal patterns of vigilance can take many forms depending on ecological factors.     

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.     

Foraging for animal prey by outdoor groups of Geoffroy’s marmosets (Callithrix geoffroyi)

I studied animal prey foraging by three outdoor groups of Geoffroy’s marmosets, Callithrix geoffroyi, over the course of two summers (June–August 1994; August–September, 1995. Marmosets are highly motivated to forage for animal prey, as demonstrated by the amount of time spent foraging for insects and small vertebrates even in the presence of provisioned, high-quality food. Like their wild congeners, the marmosets engaged in prolonged visual searches of selected areas while slowly locomoting. They seldom used their hands to manipulate substrates. The marmosets were startled proportionately more often while foraging for animal prey than would be predicted by their overall time budgets, and they regularly, if briefly, interrupted their search/locomote pattern with sweeping scans of the surroundings that may contribute to a vigilance function. While carrying infants, they almost never foraged. These results are consistent with the notion that marmosets may be particularly vulnerable to predation while searching for animal prey. Most of the data on marmoset and tamarin feeding ecology have focused on fruits and gums. My data underscore the need to understand better the role of animal prey in the behavioral ecology of callitrichids.     

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 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.     

Vigilance and its complex synchrony in the red-necked pademelon, Thylogale thetis

Several adaptive functions, including gaining information fromother group members and detecting predators, are generally ascribedto vigilance in groups of animals subject to predation. Moststudies of the effects of neighbors on vigilance have focusedon individual vigilance. We investigated the effects of neighborson vigilance in wild red-necked pademelons Thylogale thetisforaging at night in nonpersistent aggregations in a clearingin rain forest. Neither the total number of pademelons in theclearing nor the numbers at various distances around focal individualsaffected the individual vigilance of focal animals. However,focal animals’ individual vigilance did change with thedistance to their nearest neighbor and also with distance tocover. Pairs of individuals closer than 10 m apart tended tosynchronize their bouts of individual vigilance and foraging.The degree of synchrony within pairs increased with both distanceto cover and the total number of pademelons foraging in thearea and decreased with increasing distance to the pair's nearestneighbor but did not vary with the distance separating the membersof the pair. Thus, despite their individual vigilance beingunaffected by the number of other pademelons in the feedingaggregation, pademelons were nonetheless sensitive to the presenceof conspecifics and adjusted their behavior in relation to theirseparation from neighbors. Thus, some vigilance benefits maybe obtained from the presence of conspecifics even in speciesthat aggregate only temporarily on food patches without formingmore permanent social groups.     

The value of constant surveillance in a risky environment

In risky environments, where threats are unpredictable and the quality of information about threats is variable, all individuals face two fundamental challenges: balancing vigilance against other activities, and determining when to respond to warning signals. The solution to both is to obtain continuous estimates of background risk, enabling vigilance to be concentrated during the riskiest periods and informing about the likely cost of ignoring warnings. Human surveillance organizations routinely produce such estimates, frequently derived from indirect cues. Here we show that vigilant individuals in an animal society (the pied babbler, Turdoides bicolor) perform a similar role. We ask (i) whether, in the absence of direct predator threats, pied babbler sentinels react to indirect information associated with increased risk and whether they communicate this information to group mates; (ii) whether group mates use this information to adjust their own vigilance, and whether this influences foraging success; and (iii) whether information provided by sentinels reduces the likelihood of inappropriate responses to alarm calls. Using playback experiments, we show that: (i) sentinels reacted to indirect predator cues (in the form of heterospecific alarm calls) by giving graded surveillance calls; (ii) foragers adjusted their vigilance in reaction to changes in surveillance calls, with substantial effects on foraging success; and (iii) foragers reduced their probability of responding to alarm calls when surveillance calls indicated lowered risk. These results demonstrate that identifying attacks as they occur is only part of vigilance: equally important is continuous surveillance providing information necessary for individuals to make decisions about their own vigilance and evasive action. Moreover, they suggest that a major benefit of group living is not only the increased likelihood of detecting threats, but a marked improvement in the quality of information available to each individual.     

The group size effect and synchronization of vigilance in the Tibetan wild ass

Vigilance behavior is considered as an effective strategy for prey species to detect predators.An individual benefits from living in a group by reducing the time spent being vigilant without affecting the probability of detecting a predator.However,the mechanism producing a decrease in vigilance with increasing group size is unclear.Many models of vigilance assume that group members scan independently of one another.Yet in recent studies,the other 2 patterns of vigilance,coordination and synchronization,were reported in some species.In 2 summers(2018 and 2019),we studied the group-size effect on vigilance and foraging of Tibetan wild ass in Chang Tang Nature Reserve of Tibet.We also tested whether individuals scan the environment independently,tend to coordinate their scans,or tend to synchronize their vigilance.The results showed that individuals decreased the time spent on vigilance with increasing group size,while increased the time spent foraging.Group members scanned the environment at the same time more frequently and there was a positive correlation between group members'behaviors,indicating that Tibetan wild asses tend to synchronize their vigilance.     

Relatedness does not predict vigilance in a population of the social rodent Octodon degus

The possibility that social foragers adjust and coordinate their scanning activity when in the presence of close relatives to attain inclusive fitness benefits remains controversial and scarcely examined. To this aim, we first tested the null hypothesis of no association between foraging individuals of the diurnal rodent, Octodon degus and their pairwise relatedness (six microsatellite loci), under natural conditions. Secondly, we examined the influence of relatedness on scan effort (percent overlapping) and temporal distribution of scanning using linear regression. Finally, we evaluated whether temporal distributions of scanning were significantly lower (coordination) or higher (synchrony) than random expectations using bootstrapping. We found that pairwise relatedness between focal degus and their foraging partner did not influence the scan effort or the temporal distribution of scanning. These original, field-based findings imply that vigilance behavior in socially foraging degus is unlikely to be kin-selected and adds to results from previous lab studies in that kinship remains a poor predictor of social behavior in these animals. Overall, our study adds to others revealing that kin selection may not have had an impact on aspects of social behavior such as vigilance during social foraging.     

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.     

Self-Improvement for Team-Players: The Effects of Individual Effort on Aggregated Group Information

By putting effort into behaviours like foraging or scanning for predators, an animal can improve the correctness of its personal information about the environment. For animals living in groups, the individual can gain further information if it is able to assess public information about the environment from other group members. Earlier work has shown that consensus group decisions based upon the public information available within the group are more likely to be correct than decisions based upon personal information alone, given that each individual in a group has a fixed probability of being correct. This study develops a model where group members are able to improve their personal likelihood of making a correct decision by conducting some level of (costly) effort. I demonstrate that there is an evolutionarily stable level of effort for all the individuals within the group, and the effort made by an individual should decrease with increasing group size. The relevance of these results to social decision making is discussed: in particular, these results are similar to standard theoretical predictions about the amount of vigilance shown by individuals decreasing with increasing group size. However, this model suggests that these results could come about where individuals are coordinating their effort within the group (unlike standard models, which assume that all individual effort is independent of the actions of others). This ties in with experimental findings where individuals have been shown to monitor the efforts of others.     

Testing the Threat‐Sensitive Hypothesis with Predator Familiarity and Dietary Specificity

In a system with multiple predators, the threat‐sensitive predator avoidance hypothesis predicts that prey respond differently to predators relative to the risks each poses (e.g., degree of dietary specialization). Aquatic animals often rely heavily on detecting predators via chemical cues (kairomones) and respond with a suite of behaviors including detection and avoidance. However, little is known about how animals respond to kairomones of specialist versus generalist predators. In laboratory experiments, we compared behavioral responses of a poorly studied aquatic salamander, the greater siren (Siren lacertina), to cues from specialist and generalist predator snakes to evaluate threat‐sensitive responses. Sirens exhibited a novel behavior (gill‐flushing) most often in the presence of specialist predator cues. Avoidance behavior (reversing direction following cue detection) was higher in response to specialist predator and novel animal control cues and lowest in response to generalist predator cues. An intermediate response to the animal control, an unfamiliar amphibian predator, indicated that sirens respond cautiously to a novel cue. The gradient of observed responses to different snake cues indicates that sirens may be evaluating predation potential of animals based on their foraging specificity and familiarity.     

Risk management in optimal foragers: the effect of sightlines and predator type on patch use,time allocation,and vigilance in gerbils

In the foraging game between gerbils and their predators, gerbils manage risk of predation using the tools of time allocation (where, when and for how long to forage) and vigilance. The optimal level of a forager's vigilance should be affected by its encounter rate with predators and the effectiveness of its vigilance in reducing mortality risk. The physical structure of the environment can alter the effectiveness of its vigilance and therefore alter its foraging behaviour. We tested this for gerbils at risk of predation from barn owls or foxes in a large vivarium. In particular, we reduced the effectiveness of vigilance by placing obstructions around feeding trays that blocked sight lines along either the vertical (vigilance directed against owls) or horizontal axis (vigilance directed against foxes), thereby changing the physical structure of the environment. In addition, we manipulated the presence of foxes and owls. In general, gerbils harvested fewer seeds, allocated less time to foraging in dangerous patches, and used more vigilance while foraging where and when risks were higher (i.e. in the presence of predators and in bright moonlight). Vertical and horizontal sightline treatments interacted synergistically to further raise perceived risk. These results imply that blocking sight lines reduces the effectiveness of vigilance, causing gerbils to use it less. Moreover, in the presence of a predator, the gerbils’ response to the blocked sightlines was more severe – harvesting less food and spending less time and vigilance – in the patches with the increased risk. This was especially so in the presence of the predator that was expected to most benefit from blocking that particular type of sight line: cover that blocked vertical sight lines was riskiest in the presence of owls, and cover that blocked horizontal sight lines was riskiest in the presence of foxes. These results strongly indicate the importance of sightlines and landscape features such as bushes in the risk management and forging decisions of gerbils, demonstrating that bush cover provides mixed blessing to gerbils by providing cover, but making vigilance ineffective.     

Group‐Size Effect on Vigilance and Foraging in a Predator‐Free Population of Feral Goats (Capra hircus) on the Isle of Rum,NW Scotland

Many previous studies have found that as group size increases, individual vigilance levels decrease and forage intake increases (group‐size effect), but few such studies have considered the impact of within‐group interactions and other confounding factors on the direction of group‐size effects. A free‐ranging population of feral goats (Capra hircus), with little predation threat, was studied on the Isle of Rum (northwest Scotland), from Jun. to Nov. 2000, to investigate the effects of group size on individual vigilance levels and foraging efficiency after taking into account the effect of confounding factors (e.g. sex, season, time of day, habitat, predation risk) and within‐group interactions (via changes in movement rates while feeding). Our results show that, while group size exerted a negative influence on individual vigilance levels and a positive effect on movement rate, foraging efficiency never increased with group size and even decreased at certain times of day. There was no sex difference in individual vigilance in feral goats, but foraging efficiency was higher in females than in males. Goats were more vigilant in fall than in summer. The results imply that the benefits for foraging obtained from the reduced vigilance level in larger groups may be constrained or offset by increased interaction (or competition) within larger groups even in a population that faces negligible predation risk.     

Vigilance in Greater Flamingos Wintering in Southern Tunisia: Age-Dependent Flock Size Effect

Decrease in individual vigilance with flock size is a widely recognized pattern in group‐living species. However such a relationship may be affected by other factors, such as age and flock composition. For instance, because young animals generally lack experience and have higher nutritional needs than adults, they can be expected not only to be less vigilant than adults but also to decrease their vigilance level by a greater extent when flock size increases than adults do. We investigated this issue using data on greater flamingos wintering in the gulf of Gabès, in southern Tunisia. Flamingos tended to congregate in small single‐age flocks for feeding, but as flock size increased, flocks became mixed. We found that when flock size increased, young flamingos significantly decreased their vigilance time, while adult did not, suggesting an age‐dependent flock size effect on vigilance. However, when flock composition (single‐age vs. mixed) was taken into account, a more complex pattern was found. Within single‐age and small flocks, no difference was found between young flamingos and adult ones regarding their vigilance level and their response to increasing flock size. However, within mixed and large flocks, adult flamingos were more vigilant than young ones, while variation in flock size did not result in a significant change in vigilance. These results suggest that young birds relied on the presence of adults, and hence more experienced individuals in detecting dangers, to reduce their vigilance and to increase their foraging time in order to satisfy their higher nutritional requirements. They could also be interpreted as a possible consequence of increasing competition with flock size which constrained more nutritionally stressed young flamingos to increase their foraging time to the detriment of vigilance.     

青藏铁路对雄性藏羚行为时间分配的影响

行为时间分配是动物对资源获取和风险防御的权衡,受到诸多因素影响。本研究采用目标动物观察法记录了可可西里冬季（交配季）雄性藏羚的行为表现,并将行为分为觅食、警戒、卧息、移动和“其他”5种类型。首先,本文通过对比青藏铁路运营前后两个交配季（2003-2004年建设期和2017-2018年运行期）之间雄性藏羚的行为时间分配差异,探讨青藏铁路的出现对雄性藏羚行为的影响。随后,基于2017-2018年交配季雄性藏羚的行为数据,分析了年龄对其行为时间分配的影响。结果表明：与建设期相比,雄性藏羚的觅食和“其他”行为时间比例明显增加,而警戒和卧息的行为时间比例显著降低,这说明铁路稳定运营后对雄性藏羚的影响降低。交配群中成年雄性藏羚的警戒、移动,以及“其他”行为中的繁殖行为的时间比例显著高于亚成体,而觅食和卧息行为时间比例显著低于亚成体, 这与交配群中不同年龄雄性个体所处的地位等级有关。成年雄性藏羚在交配群中占据主导地位,拥有更多配偶资源,增加警戒和移动,减少觅食和卧息有助于其维持交配群的稳定。