Olivier Pays

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 groups 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 neighbours behaviour.

Interactions among social monitoring, anti-predator vigilance and group size in eastern grey kangaroos

Group size is known to affect both the amount of time that prey animals spend in vigilance and the degree to which the vigilance of group members is synchronized. However, the variation in group-size effects reported in the literature is not yet understood. Prey animals exhibit vigilance both to protect themselves against predators and to monitor other group members, and both forms of vigilance presumably influence group-size effects on vigilance. However, our understanding of the patterns of individual investment underlying the time sharing between anti-predator and social vigilance is still limited. We studied patterns of variation in individual vigilance and the synchronization of vigilance with group size in a wild population of eastern grey kangaroos (Macropus giganteus) subject to predation, in particular focusing on peripheral females because we expected that they would exhibit both social and anti-predator vigilance. There was no global effect of group size on individual vigilance. The lack of group-size effect was the result of two compensating effects. The proportion of time individuals spent looking at other group members increased, whereas the proportion of time they spent scanning the environment decreased with group size; as a result, overall vigilance levels did not change with group size. Moreover, a degree of synchrony of vigilance occurred within groups and that degree increased with the proportion of vigilance time peripheral females spent in anti-predator vigilance. Our results highlight the crucial roles of both social and anti-predator components of vigilance in the understanding of the relationship between group size and vigilance, as well as in the synchronization of vigilance among group members.

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.

Individual variation in the relationship between vigilance and group size in eastern grey kangaroos

The mean vigilance of animals in a group often decreases as their group size increases, yet nothing is known about whether there is individual variability in this relationship in species that change group sizes frequently, such as those that exhibit fission–fusion social systems. We investigated variability in the relationship between group size and vigilance in the eastern grey kangaroo (Macropus giganteus) by testing whether all individuals showed decreased vigilance with increased group size, as has been commonly assumed. We carried out both behavioural observations of entire groups of kangaroos and focal observations of individually recognised wild female kangaroos. As in other studies, we found a collective group-size effect on vigilance; however, individuals varied in their vigilance patterns. The majority (57%) of the identified individual kangaroos did not show significant group-size effects for any of the recorded measures of vigilance. The females that did not show a negative group-size effect were, on average, more vigilant than those females that did show a group-size effect, but this difference was not significant. We propose that some females exhibit higher levels of social vigilance than others, and that this social vigilance increases with group size, cancelling out any group-size effect on anti-predator vigilance for those females. Our results therefore suggest that only some prey individuals may gain anti-predator benefits by reducing their time spent scanning when in larger groups. The large amount of variation that we found in the vigilance behaviour of individual kangaroos highlights the importance of collecting and analysing vigilance data at the individual level, which requires individual recognition.

Does sex affect both individual and collective vigilance in social mammalian herbivores: the case of the eastern grey kangaroo?

In several vertebrate taxa, males and females differ in the proportions of time they individually devote to vigilance, commonly attributed to sex differences in intra-specific competition or in absolute energy requirements. However, an effect of sex on collective vigilance is less often studied (and therefore rarely predicted), despite being relevant to any consideration of the adaptiveness of mixed- vs single-sex grouping. Controlling for group size, we studied the effect of sex on vigilance in the sexually dimorphic eastern grey kangaroo Macropus giganteus, analysing vigilance at two structural levels: individual vigilance and the group’s collective vigilance. Knowing that group members in this species tend to synchronise their bouts of vigilance, we tested (for the first time) whether sex affects the degree of synchrony between group members. We found that females were individually more vigilant than males and that their vigilance rate was unaffected by the presence of males. Collective vigilance did not differ between female-only and mixed-sex groups of the same size. Vigilance in mixed-sex groups was neither more nor less synchronous than in single-sex groups of females, and the presence of males seemed not to affect the degree of synchrony between females. Sixty-six percent of vigilant acts were unique (performed when no other kangaroo was alert), and only about one unique vigilant act in every three induced a collective wave of vigilance. The proportions of vigilant acts that were unique were 60% for females but only 46% for males. However, the sexes differed little in the rates at which their unique vigilant acts were copied. This limited study shows that the differences in vigilance between male and female kangaroos had no discernible effect upon collective vigilance.

Within-population differences in personality and plasticity in the trade-off between vigilance and foraging in kangaroos

Behavioural traits can vary between individuals from the same population. These differences can involve consistent variation in the level of a particular behaviour (personality) or differences in the way individuals adjust their behaviour to environmental gradients (plasticity). In prey species, feeding rates and vigilance vary with environmental, social and individual factors and the feeding rate/vigilance relationship reflects the trade-off between food acquisition and safety. While feeding rates and vigilance have been shown to vary between individuals in relation to group size and predation risk, how they relate to other factors has not yet been investigated, nor has between-individual variation in this tradeoff. We studied between-individual variation in vigilance, feeding rates and their trade-off in female eastern grey kangaroos, Macropus giganteus, to see whether females showed consistent behavioural differences and different plasticity in relation to ecological (food patch richness), social (group sizes) and physiological (reproductive states) conditions. We addressed two contrasting hypotheses: an ‘ecological’ hypothesis under which individuals facing the same conditions should behave similarly, and a ‘behavioural’ hypothesis under which they should behave differently because of their own personality or plasticity. Female kangaroos tended to adjust their behaviours similarly in relation to ecological and social conditions, supporting the ecological hypothesis. However, they also showed differences in personality and plasticity in relation to their reproductive states that could not be explained by energetic demand alone; this was suggestive of different maternal strategies, thus supporting the behavioural hypothesis. Altogether these results suggest that consistent differences in animals’ personality and behavioural plasticity can be promoted by physiological conditions and are not necessarily repeatable across different ecological contexts.

Group Dynamics and Landscape Features Constrain the Exploration of Herds in Fusion-Fission Societies: The Case of European Roe Deer

Despite the large number of movement studies, the constraints that grouping imposes on movement decisions remain essentially unexplored, even for highly social species. Such constraints could be key, however, to understanding the dynamics and spatial organisation of species living in group fusion-fission systems. We investigated the winter movements (speed and diffusion coefficient) of groups of free-ranging roe deer (Capreolus capreolus), in an agricultural landscape characterised by a mosaic of food and foodless patches. Most groups were short-lived units that merged and split up frequently during the course of a day. Deer groups decreased their speed and diffusion rate in areas where food patches were abundant, as well as when travelling close to main roads and crest lines and far from forests. While accounting for these behavioural adjustments to habitat features, our study revealed some constraints imposed by group foraging: large groups reached the limit of their diffusion rate faster than small groups. The ability of individuals to move rapidly to new foraging locations following patch depression thus decreases with group size. Our results highlight the importance of considering both habitat heterogeneity and group dynamics when predicting the movements of individuals in group fusion-fission societies. Further, we provide empirical evidence that group cohesion can restrain movement and, therefore, the speed at which group members can explore their environment. When maintaining cohesion reduces foraging gains because of movement constraints, leaving the group may become a fitness-rewarding decision, especially when individuals can join other groups located nearby, which would tend to maintain highly dynamical group fusion-fission systems. Our findings also provide the basis for new hypotheses explaining a broad range of ecological patterns, such as the broader diet and longer residency time reported for larger herbivore groups.

Investigating Differences in Vigilance Tactic Use within and between the Sexes in Eastern Grey Kangaroos

Aggregation is thought to enhance an animal’s security through effective predator detection and the dilution of risk. A decline in individual vigilance as group size increases is commonly reported in the literature and called the group size effect. However, to date, most of the research has only been directed toward examining whether this effect occurs at the population level. Few studies have explored the specific contributions of predator detection and risk dilution and the basis of individual differences in the use of vigilance tactics. We tested whether male and female (non-reproductive or with young) eastern grey kangaroos (Macropus giganteus) adopted different vigilance tactics when in mixed-sex groups and varied in their reliance on predator detection and/or risk dilution as group size changed. This species exhibits pronounced sexual dimorphism with females being much smaller than males, making them differentially vulnerable toward predators. We combined field observations with vigilance models describing the effects of detection and dilution on scanning rates as group size increased. We found that females with and without juveniles relied on predator detection and risk dilution, but the latter adjusted their vigilance to the proportion of females with juveniles within their group. Two models appeared to equally support the data for males suggesting that males, similarly to females, relied on predator detection and risk dilution but may also have adjusted their vigilance according to the proportion of mothers within their group. Differential vulnerability may cause sex differences in vigilance tactic use in this species. The presence of males within a group that do not, or only partially, contribute to predator detection and are less at risk may cause additional security costs to females. Our results call for reexamination of the classical view of the safety advantages of grouping to provide a more detailed functional interpretation of gregariousness.

On the dynamics of predation risk perception for a vigilant forager.

Foraging animals often raise their head to scan for predators. Scanning intervals have variable durations, and occur more or less frequently, depending on ecological conditions. Our study relies on the assumption that temporal patterns of vigilance depend on the speed with which information concerning the likelihood of a predators presence in the neighbourhood is gathered when an animal is vigilant, and lost when it is not. Using an analytical model, we study how the perceived level of risk progressively decreases, when the individual is vigilant and detects no predator, then increases again, when it lowers its head to feed, thereby losing most of its detection abilities. The speed of these variations is affected by the likelihood of the presence of a predator in the whole environment, by the mobility of this predator, and by the detection capacities of the prey. We show how, combined with the range of risk levels tolerated by this animal, this dynamics determines the frequency and the duration of its scanning intervals. The dynamics of risk perception can also explain particular behavioural patterns, such as the progressive decrease of vigilance that may occur after the arrival into a novel environment, and the central tendency in the distribution of interscan durations reported by many studies. Next, we use the model to compute optimal vigilance strategies, taking into account the trade-off between feeding and limiting exposure to predators. The model predicts that a forager will scan more often, and for longer periods, when the likelihood of a predators presence in the surrounding environment is increased. A similar response is expected when the mobility of the predator is increased. By contrast, when the detection capacities of the prey are reduced, it will increase its vigilance by scanning for longer periods, but scanning intervals will be separated by longer interscans.

How unpredictable is the individual scanning process in socially foraging mammals

In group-forming prey species, theory assumes that individuals within groups should scan independently of one another, with vigilance sequences being relatively unpredictable, making interscan durations highly variable. We attempted to detect any divergence from randomness in the scanning process in three mammalian prey species phylogenetically and geographically separated and exposed to different levels of predation: waterbuck, Kobus ellipsiprymnus defassa, under a high observed predation risk, eastern grey kangaroo, Macropus giganteus, still experiencing occasional predation and European roe deer, Capreolus capreolus, under a very low natural predation risk. Our results revealed that the focal interscan duration increased when the duration of the preceding interscan increased, whatever the studied species and the predation risk that its individuals experienced, and decreased with the preceding scan duration in two species under, respectively, occasional and low predation risks. The exponential distribution was the tested model that fitted the observed distributions of interscan durations least well. We discuss what can trigger non-randomness in scanning, through a non-homogenous Poisson process, at both intra-individual and inter-individual levels, particularly with regard to previous studies that have demonstrated synchronisation of vigilance in such mammals. Our results suggest the need to reconsider any assumption of randomness in scanning in the basic model predicting form and frequency of scanning behaviour by prey species.