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.     

Spread of false alarms in foraging flocks of house sparrows

In group‐foraging species with no alarm signals, the sudden departure of neighbours can be used to inform the rest of the group about the detection of a threat. However, sudden departures are ambiguous because they can be triggered by factors unrelated to predator detection. We evaluated how animals react to the sudden departure of neighbours in groups of foraging house sparrows (Passer domesticus). We focussed on false alarms that occurred for no apparent reasons to us because predation attempts were not frequent. Three factors can explain how the sudden departure of a neighbour can influence reaction times, namely group size, the distance between neighbours, and predation risk. We predicted reaction times to be longer in larger groups where individual vigilance levels are low, and when group members are further apart and cannot easily monitor each other. In addition, we expected reaction times to be longer when predation risk is lower. Departures that are more temporally clumped are also expected to be less ambiguous and should trigger faster reaction times. Our results show that sparrows reacted faster, not more slowly, to the sudden departures of neighbours in larger groups, and, as predicted, more slowly when neighbours were more distant from each other. Reaction times were longer in one of the two study years in which predation risk was deemed lower. Sparrows reacted more quickly when earlier departures were more temporally clumped. The results thus provided partial support for the predictions, and future work is needed to assess how individuals react to fleeing responses by their neighbours in species with no alarm signals.     

Social context affects risk taking by a satellite species in a mixed-species foraging group

Mixed-species flocks of birds form during winter in the easterndeciduous forests of North America. These flocks consist oftwo flock-leading nuclear species, tufted titmouse (Baeolophusbicolor) and Carolina chickadee (Poecile carolinensis), andseveral follower, or satellite, species, including downy woodpecker(Picoides pubescens) and white-breasted nuthatch (Sitta carolinensis).Hypotheses explaining the adaptiveness of participation in suchmixed-species foraging groups have focused on increased foragingsuccess and/or decreased predation risk. We tested the predictionthat if nuthatches join nuclear species to reduce predationrisk, they should be more reluctant to visit an exposed feederin the absence of titmice than in their presence. When the feederwas positioned 16 m from forest cover, latency to visit thefeeder was greater for both male and female nuthatches whentitmice were absent. Removal of titmice had no effect on latencyat 8 m. In the absence of titmice, nuthatches visited the feederless frequently at both distances. These results indicate thatreduced predation risk is a benefit that satellite species gainby flocking with nuclear species.     

On the Advantages of Mixed‐Species Groups: Impalas Adjust Their Vigilance When Associated With Larger Prey Herbivores

Prey can obtain valuable benefits from associating with other species if heterospecifics help to detect predators or locate good food patches. In mixed‐species groups, how species respond to the presence of other species remains a poorly explored question although it might give crucial insights into mechanisms underlying the interspecific coexistence. We studied temporary mixed‐species groups of large herbivores in Hwange National Park (Zimbabwe) between the common impala (Aepyceros melampus), the focal species here, and bigger species including the plains zebra (Equus quagga), the greater kudu (Tragelaphus strepsiceros) or the blue wildebeest (Connochaetes taurinus). In the Hwange savanna, the focal and smaller species are exposed to a larger range of predators than the associated species. In this context, we investigated how impalas adjusted their vigilance with group size comparing impala‐only and mixed‐species groups and whether the identity of heterospecifics affected vigilance of impalas. Our study showed that the time impalas spent in vigilance significantly decreased with group size when they formed impala‐only groups, whereas it did not significantly vary with group size in mixed‐species groups. Moreover, in mixed‐species groups, impalas did not adjust their time spent in vigilance with the proportion of conspecifics and the identity of the associated species. Thus, the mechanism underlying the difference of impalas' behavioural adjustment of vigilance with group size between single‐ and mixed‐species groups seemed to be related to the presence but not to the number and the identity of heteropecifics. Finally, we discuss the concept that larger and dominant heterospecifics were likely to increase competition for food access, thereby forcing higher vigilance of impalas, outweighing any reduction from collective vigilance.     

Dilution games: use of protective cover can cause a reduction in vigilance for prey in groups

Antipredatory vigilance usually decreases in groups. The generallyaccepted "collective detection" explanation implies that becausethere are more eyes to scan the surroundings for predators,individuals in a group can lower their personal investment invigilance without increasing their predation risk. The roleof other factors, such as numerical risk dilution caused bythe mere presence of companions, has been neglected. In a model,we explore a dilution game when foragers in groups have accessto protective cover. We show that foragers can take advantageof risk dilution and that this leads to changes in vigilancewith group size without the need to invoke collective detection.We identify a cost to maintaining high levels of vigilance asless vigilant foragers gather food faster and so depart thegroup sooner (to reach cover) leaving more vulnerable stragglersbehind. In groups, there is a scramble to reach safe sites thatcan induce a reduction in vigilance levels. Such a mechanismoperates less forcefully in large groups because individualsin these groups are less vulnerable to the departure of an individual.We also demonstrate that individuals should adopt lower levelsof vigilance, to reach safe sites sooner, when predator evasionis compromised or when the rate of food intake is high. Themodel provides new insights into the mechanisms underlying changesin vigilance with group size in animals.     

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.     

Optimization of foraging and diet by the piscivorous Othos dentex (Serranidae)

The aim of this study was to determine the dietary characteristics and mouth morphology of Othos dentex and to use these data, together with in situ observations of feeding behaviour, to elucidate how foraging and diet are optimized by this piscivorous serranid. Seasonal spear and line fishing over reefs in south‐western Australia yielded 426 O. dentex (total length, L_T, 183–605 mm), among which the stomachs of 95 contained food. The food in the stomachs of 76 fish was sufficiently undigested to be seen to contain, almost invariably, a single fish prey, which was typically identifiable to family and often to species. The prey of O. dentex, which were measured (L_T), represented 10 families, of which the Labridae and Pempheridae constituted nearly two‐thirds of the prey volume. Two‐way crossed analysis of similarities of volumetric data for stomach contents showed that the dietary compositions of the different length classes of O. dentex in the various seasons were significantly related to length class of prey, but not to prey family, length class within the various prey families or season. Furthermore, an inverse (Q‐mode) analysis, including one‐way analysis of similarities, showed that the patterns in the prey consumed by the different length classes of O. dentex in the various seasons were related more strongly to length class than prey family. The former trend is exemplified in a shade plot, by a marked diagonality of the length classes of prey with increasing predator size. The ingestion of typically a single teleost prey, whose body size increases as that of O. dentex increases, reduces the frequency required for seeking prey, thus saving energy and reducing the potential for intraspecific competition for food. The ability of O. dentex to ingest large prey is facilitated by its possession of a very large gape, prominent recurved teeth, dorsal and independently‐moveable eyes, cryptic colouration and effective ambush behaviour. Othos dentex has thus evolved very cost‐effective mechanisms for optimizing its foraging and diet.     

Traffic noise and responses to a simulated approaching avian predator in mixed-species flocks of chickadees,titmice, and nuthatches

Traffic noise likely reaches a wide range of species and populations throughout the world, but we still know relatively little about how it affects anti-predator behavior of populations. We tested for possible effects of traffic noise on responses to predator acoustic cues in Carolina chickadees (Poecile carolinensis), tufted titmice (Baeolophus bicolor), and white-breasted nuthatches (Sitta carolinensis) near 14 independent feeding stations in eastern Tennessee. We compared anti-predator calling and seed-taking behavior in response to playbacks of predator stimuli (screech owl calls) at sites naturally exposed to traffic noise and at sites that faced relatively little traffic noise. The screech owl call playback was designed to simulate the approach of this dangerous predator to a feeder being used by these small songbirds. We found that chickadees responded consistently to the owl stimuli across different levels of traffic noise. However, titmice, and nuthatches exhibited different behavioral responses to the predator stimulus, suggesting that traffic noise masked these low-frequency predator calls. Overall, chickadees and nuthatches showed the broadest anti-predator behavioral responses in comparison to titmice, corroborating earlier published work with an Indiana population. Finally, populations exposed to traffic noise overall seemed less able to detect predator cues potentially masked by that noise, and future work will need to assess likely seasonal variation in these responses as well as species-level variation in anti-predator responses in mixed-species groups.     

Conspecific Influences on Vigilance Behavior in Wild Chimpanzees

Diurnal primates rely on visual monitoring behavior to collect various kinds of ecological and social information. Vigilance behavior is monitoring specifically to detect external threats. Previous studies of vigilance behavior were focused mainly on the influence of predation threats, whereas the influences of conspecific factors, such as intragroup threats, have been relatively unstudied. Individual vigilance is predicted to be inversely related to the group size or the number of individuals nearby if the main target of the vigilance is a predation threat and positively related if the main target of the vigilance is a conspecific threat. I studied wild chimpanzees (Pan troglodytes schweinfurthii) in Mahale Mountains National Park, Tanzania, and measured the vigilance duration when they are resting on the ground via 2-min focal observation. In both males and females, vigilance duration increased as the number of individuals nearby increased. This result agrees with the idea that the chimpanzees are vigilant toward other group members. In addition, maternal vigilance monitors and protects the safety of dependent offspring as the duration of maternal vigilance was longer when a dependent infant was separated from its mother than when the offspring was in contact with its mother. The results indicate that the vigilance behavior in wild chimpanzees was affected by conspecific factors.     

Overriding the oddity effect in mixed-species aggregations: group choice by armored and nonarmored prey

Because "odd" individuals often suffer disproportionately highrates of predation, solitary individuals should join groupswhose members are most similar to themselves in appearance.We examined group-choice decisions by individuals in armoredand nonarmored species and predicted that either (1) the oddityeffect would result in preference for conspecific groups forsolitary individuals of both species, or (2) individuals inthe armored species would prefer to associate with groups containingindividuals of the more vulnerable species. Armored brook sticklebacks(Culaea inconstans) and nonarmored fathead minnows (Pimephalespromelas) have the same predators and often occur together instreams. In mixed-species shoals, yellow perch (Perca flavescens)attacked minnows earlier and more often than sticklebacks. Wetested whether solitary minnows and sticklebacks preferred toassociate with conspecific or heterospecific shoals under conditionsof both low and high predation risk. When predation risk washigh, minnows preferred to associate with conspecifics overheterospecifics, as predicted by the oddity effect. In contrast,sticklebacks preferentially associated with groups of minnowsover groups of conspecifics when predation risk was high. Whenpredation risk was low, solitary individuals of both speciespreferentially associated with conspecific over heterospecificshoals. Stickleback shoal choices under low-risk conditionsmay have been influenced by interspecific competition for food.In feeding experiments, minnows were more efficient foragersthan sticklebacks, so it should benefit sticklebacks to avoidminnows unless predation risk is high. Therefore, for armoredprey, the benefits of associating with more vulnerable preyappear to override the costs of both the oddity effect and foodcompetition when predation risk is high.     

Mixed-species primate groups in the kibale forest: Ecological constraints on association

Five species of diurnal primates in the Kibale Forest of western Uganda— red colobus (Colobus badius),black- and- white colobus (Colobus guereza),redtail monkeys (Cercopithecus ascanius),blue monkeys (Cercopithecus mitis),mangabeys (Cercocebus albigena)-often associate in mixed- species groups that vary in size and composition from day to day. Across this range of species, we found no consistent effect of association on feeding rate. In addition, there is no systematic difference between the species- specific individual feeding rates when animals were in mixed- species groups feeding in a specific tree on one day and when individuals of one of the same species were feeding in the same individual tree on a subsequent day. If associating in a mixed- species group lowers the risk of predation, one might expect that the number of vigilant events would decrease in mixed- species groups. However, the only species to exhibit a consistent decrease in vigilant behavior when in association was the red colobus. Redtail monkeys were more vigilant when in association. We predicted that the density and distribution of food resources would both constrain the frequency of association and the size of mixed- species groups. Based on 22 months of data on food resources and bimonthly censuses, we found no relationship between the frequency of association (except mangabeys) or mean mixed- species group size and the density and distribution of food resources for all species. Finally, we examined the behavior of the monkeys in and out of association before and after the playback of a crowned hawk eagle call (Spizaetus coronatus),a known predator. When more species were in association, the amount of time they spent being vigilant following the playback was greater and the response more intense than when fewer species were in association or when the group was alone. The results of this study illustrate that the nature of the costs and benefits of polyspecific associations for these different monkey species are complex and vary from species to species.     

A context analysis of bobbing and fin‐flicking in a small marine benthic fish

Most antipredator strategies increase survival of individuals by signaling to predators, by reducing the chances of being recognized as prey, or by bewildering a predator''s perception. In fish, bobbing and fin‐flicking are commonly considered as pursuit‐deterrent behaviors that signal a predator that it has been detected and thus lost its surprise‐attack advantage. Yet, very few studies assessed whether such behavioral traits are restricted to the visual presence of a predator. In this study, we used the yellow black‐headed triplefin Tripterygion delaisi to investigate the association between these behaviors and the visual exposure to (a) a black scorpionfish predator (Scorpaena porcus), (b) a stone of a size similar to that of S. porcus, (c) a conspecific, and (d) a harmless heterospecific combtooth blenny (Parablennius sanguinolentus). We used a laboratory‐controlled experiment with freshly caught fish designed to test for differences in visual cues only. Distance kept by the focal fish to each stimulus and frequency of bobbing and fin‐flicking were recorded. Triplefins kept greater distance from the stimulus compartment when a scorpionfish predator was visible. Bobbing was more frequent in the visual presence of a scorpionfish, but also shown toward the other stimuli. However, fin flicks were equally abundant across all stimuli. Both behaviors decreased in frequency over time suggesting that triplefin become gradually comfortable in a nonchanging new environment. We discuss why bobbing and fin‐flicking are not exclusive pursuit‐deterrent behaviors in this species, and propose additional nonexclusive functions such as enhancing depth perception by parallax motion (bobbing) or signaling vigilance (fin‐flicking).     

Vigilance in Single- and Mixed-Species Groups of Tamarins (Saguinus labiatus and Saguinus fuscicollis)

By forming larger sizes of groups, individuals benefit from a decrease in vigilance, but the collective vigilance of the group as a whole is not compromised. We examined whether this group size effect is apparent in mixed-species groups of red-bellied tamarins (S. labiatus) and saddleback tamarins (S. fuscicollis) which form stable and permanent associations in the wild. We studied general vigilance and responses to hidden threatening stimuli in five captive groups of each species, while they were housed in single- and mixed-species groups. For vigilance, the individual rate was lower in the larger mixed-species groups than in the smaller single-species groups. In addition, the amount of time when at least one individual was vigilant was higher in mixed-species groups. This suggests that the tamarins alter their vigilance behavior in the presence of the other species. In response to hidden threats, both species performed brief vigilance checks and frequencies of checking did not differ in single- and mixed-species groups. However, both species had a significant reduction in the mean duration per check, and there was a reduced total amount of time spent vigilance checking in the mixed-species groups compared to the single-species groups, demonstrating the group size effect. Overall the mixed-species groups had a higher number and mean duration of checking than the smaller single-species groups. Given that the two species share a common set of predators, and respond to each other's alarm calls, these findings provide strong evidence that individuals of both tamarin species may be able to benefit from forming mixed-species groups via improved vigilance and monitoring of threats.     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

Costs and benefits of flocking in foraging white-fronted geese (Anser albifrons): effects of resource depletion

This study investigated the costs and benefits of flocking in white-fronted geese Anser albifrons foraging on rice grains in Japan. The time budgets of focal geese were recorded, and the effects of flock size on the proportions of time spent in vigilant and agonistic behaviour were tested. The results showed that the decline in vigilance level and consequent increase in foraging time were beneficial results of flocking whereas agonistic interactions, a potential cost of flocking, did not increase with increasing flock size. However, seasonal variation in flock size suggested that exploitative competition could be a cost of flocking; the sizes of flocks in spring, when resource depletion had progressed, were significantly reduced compared with those in autumn. An experimental increase in rice density resulted in a significant increase in flock size. We conclude that the flock size of foraging white-fronted geese is a result of compromise between a constant benefit of flocking (i.e. decline in vigilance level) and a cost of flocking varying with food abundance (i.e. exploitative competition).