Predator detection and dilution as benefits of associations between yellow mongooses and Cape ground squirrels

Associations among organisms are thought to form because the benefits, such as increased foraging efficiency or decreased risk of predation, outweigh any costs, such as resource competition. Though many interspecific associations have been described for closely related mammals, few studies have examined the associations between mammals in different orders. The yellow mongoose (Cynictus pencillata), a carnivore, and the Cape ground squirrel (Xerus inauris), a rodent, co-occur in arid and semi-arid South Africa where they share sleeping burrows, predators, a similar body size, and the capability to emit alarm calls in response to predators. To investigate enhanced predator avoidance as a potential benefit explaining the persistence of this association, we assessed individual mongoose vigilance alone and with squirrels or other mongooses, and with varying interspecific group size, using field observations. We also tested for responses to conspecific and heterospecific alarm calls in both study species using playback experiments. The proportion of time mongoose individuals spent vigilant decreased in the presence of squirrels or other mongooses and was negatively correlated with interspecific group size; a similar pattern was previously shown for conspecific groups of Cape ground squirrels. These results are predicted by both the dilution and collective detection hypotheses. In addition, hetero- and conspecific alarm calls elicited vigilance responses in both species. These results suggest that both species can benefit from the collective detection and dilution arising from their interspecific association and that this interspecific association could be mutualistic.     

Vigilance for predators: detection and dilution effects

Summary Grouped individuals are less vigilant for predators than solitary conspecifics because (1) grouping increases the likelihood of predator detection (detection effect) and (2) grouping makes it less likely that any given individual will be preyed upon (dilution effect). However, many models of vigilance behaviour consider only the detection effect, and the interaction of the two effects has been insufficiently considered. I present two models of vigilance behaviour and test them using data on the vigilance of elk, Cervus elaphus. The first model, based on the detection effect alone, is implicit in many published formulations of the relationship between vigilance and group size. Although it predicts the direction of the relationship between vigilance and group size, it provides a poor explanation of the form of the relationship. The second model incorporates both detection and dilution effects. Regression analysis on this security model indicates that the model provides a good prediction of both the direction and the form of the relationship between vigilance and group size, explaining 69% of the variance in vigilance frequency. The security model demonstrates that both detection and dilution are important in determining the frequency of vigilance behaviour but that the relative importance of these two effects changes across group size, with detection providing relatively less benefit as group size increases. However, even when groups are large, individuals should exhibit at least some vigilance because although dilution alone provides much protection from predation, even a low level of vigilance greatly increases the likelihood that an individual will survive repeated predation attempts.     

Cost-free vigilance during feeding in folivorous primates? Examining the effect of predation risk,scramble competition,and infanticide threat on vigilance in ursine colobus monkeys (<Emphasis Type="Italic">Colobus vellerosus</Emphasis>)

Vigilance often decreases with increasing group size, due to lower predation risk or greater scramble competition for food. A group size effect on vigilance is seldom seen in primates, perhaps because scanning and feeding often occur simultaneously or because the distinction between routine and induced vigilance has not been investigated. We analyzed feeding and resting observations separately while distinguishing between routine and induced scans in four groups of wild ursine colobus monkeys (Colobus vellerosus) experiencing scramble competition for food and infanticide risk. We used linear mixed-effect models to test the effect of group size, age–sex class, number of neighbors, number of adult male neighbors, and height in the canopy on scanning rates (vigilance) with and without evident conspecific threat. Food type was also examined in the feeding models. Perceived predation risk affected vigilance more than scramble competition for food and infanticide risk. Routine and induced vigilance were greatest at lower canopy heights during feeding and resting and increased when individuals had fewer neighbors while resting. A group size effect was found on induced vigilance while resting, but scanning increased with group size, which probably indicates visual monitoring of conspecifics. Scanning rates decreased while feeding on foods that required extensive manipulation. This supports the idea that vigilance is relatively cost free for upright feeders when eating food that requires little manipulation, a common feature of folivore diets. In the presence of threatening conspecific males, close proximity to resident males decreased individual vigilance, demonstrating the defensive role of these males in the group.     

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.     

Predation, scramble competition, and the vigilance group size effect in dark-eyed juncos (Junco hyemalis)

In socially feeding birds and mammals, as group size increases, individuals devote less time to scanning their environment and more time to feeding. This vigilance “group size effect” has long been attributed to the anti-predatory benefits of group living, but many investigators have suggested that this effect may be driven by scramble competition for limited food. We addressed this issue of causation by focusing on the way in which the scan durations of free-living dark-eyed juncos (Junco hyemalis) decrease with group size. We were particularly interested in vigilance scanning concomitant with the handling of food items, since a decrease in food handling times (i.e. scan durations) with increasing group size could theoretically be driven by scramble competition for limited food resources. However, we showed that food-handling scan durations decrease with group size in an environment with an effectively unlimited food supply. Furthermore, this food-handling effect was qualitatively similar to that observed in the duration of standard vigilance scans (scanning exclusive of food ingestion), and both responded to changes in the risk of predation (proximity of a refuge) as one might expect based upon anti-predator considerations. The group size effects in both food-handling and standard scan durations may reflect a lesser need for personal information about risk as group size increases. Scramble competition may influence vigilance in some circumstances, but demonstrating an effect of competition beyond that of predation may prove challenging. Received: 22 September 1998 / Received in revised form: 1 February 1999 / Accepted: 14 February 1999     

Ideal free distributions under predation risk

 We examine the trade-off between gathering food and avoiding predation in the context of patch use by a group of animals. Often a forager will have to choose between feeding sites that differ in both energetic gain rate and predation risk. The ideal site will have a high gain rate and low risk of predation. However, intake rate will often decrease when the patch is shared with other foragers and it may be optimal for some individuals to feed elsewhere. Within the framework of ideal free theory, we investigate the distribution of foragers that will equalise individual fitness gains. We focus on a two-patch environment with continuous inputs of food. With reference to existing experimental studies, we examine the effects of risk dilution, food input rates and an animal’s expectations of the future. We identify the effect of total animal numbers when one patch is subject to predation risk and the other is safe. Conditions under which the difference in intake rate in the two patches is constant are identified, as are conditions in which the ratio of animals in the two patches is constant. If current conditions do not alter future expectations an increase in input rates to the patches promotes increased use of the risky patch. Yet, if conditions are assumed to persist indefinitely the opposite effect is seen. When both patches are subject to predation risk, dilution of risk favours more extreme distributions, and may lead to more than one stable distribution. The results of these models are used to critically analyse previous work on the energetic equivalence of risk. This paper is intended to help guide the development of new experimental studies into the energy-risk trade-off. Received: 10 February 1995/Accepted after revision: 1 October 1995     

Vigilance and aggregation in black howler monkeys (Alouatta pigra)

Animals are expected to reduce per capita vigilance with increasing group size. Widespread support for this expectation has been found in studies of birds and ungulates. Primate vigilance patterns appear to differ, but studies of primates may have sampled groups with too many members to detect intergroup differences in vigilance. We tested this idea by measuring individual vigilance rates in wild black howler monkeys (Alouatta pigra) living in stable groups of two to ten individuals. No group size effect on vigilance emerged. As in previous studies, inter-individual proximity explained variation in individual vigilance rates. Focal animals with associates within 2 m and with multiple associates in the same tree were less vigilant than isolated animals. Activity and class of focal animals as well as the presence of neonates influenced the average vigilance of adult and subadult black howler monkeys. We conclude that functional explanations must be considered for the general lack of a group size effect on primate vigilance.     

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.     

Balancing the response to predation—the effects of shoal size,predation risk and habituation on behaviour of juvenile perch

Group size, predation risk and habituation are key drivers of behaviour and evolution in gregarious prey animals. However, the extent to which they interact in shaping behaviour is only partially understood. We analyzed their combined effects on boldness and vigilance behaviour in juvenile perch (Perca fluviatilis) by observing individuals in groups of one, two, three and five faced with four different levels of predation risk in a repeated measures design. The perch showed an asymptotic increase in boldness with increasing group size and the highest per capita vigilance in groups of two. With increasing predation risk, individuals reduced boldness and intensified vigilance. The interaction between group size and predation risk influenced vigilance but not boldness. In this context, individuals in groups of two elevated their vigilance compared to individuals in larger groups only when at higher risk of predation. Further, as only group size, they significantly reduced vigilance at the highest level of risk. With increasing habituation, solitary individuals became considerably bolder. Also, predation risk affected boldness only in the more habituated situation. Hence, repeated measures may be essential to correctly interpret certain relationships in behaviour. Our results suggest that perch may adjust boldness behaviour to group size and predation risk independently. This is rather unexpected since in theory, natural selection would strongly favour an interactive adjustment. Finally, vigilance might be particularly effective in groups of two due to the intense monitoring and detailed response to changing levels of risk.     

Vigilance behaviour and fitness consequences: comparing a solitary foraging and an obligate group-foraging mammal

Vigilance behaviour in gregarious species has been studied extensively, especially the relationship between individual vigilance and group size, which is often negative. Relatively little is known about the effect of conspecifics on vigilance in non-obligate social species or the influence of sociality itself on antipredator tactics. We investigated predator avoidance behaviour in the yellow mongoose, Cynictis penicillata, a group-living solitary forager, and compared it with a sympatric group-living, group-foraging herpestid, the meerkat, Suricata suricatta. In yellow mongooses, the presence of conspecifics during foraging—an infrequent occurrence—reduced their foraging time and success and increased individual vigilance, contrary to the classical group-size effect. Comparing the two herpestids, sociality did not appear to affect overt vigilance or survival rates but influenced general patterns of predator avoidance. Whereas meerkats relied on communal vigilance, costly vigilance postures, and auditory warnings against danger, yellow mongooses avoided predator detection by remaining close to safe refuges and increasing “low-cost” vigilance, which did not interfere with foraging. We suggest that foraging group size in herpestids is constrained by species-distinct vigilance patterns, in addition to habitat and prey preference.     

Determinants of vigilance behavior in the ring-tailed coati (Nasua nasua): the importance of within-group spatial position

Individuals living in social groups are predicted to live under unequal predation risk due to their spatial location within the group. Previous work has indicated that individuals located at the edge of groups have higher “domains of danger”, thus are more likely to engage in vigilance or antipredator behavior. We studied the determinants of vigilance behavior in two groups of ring-tailed coatis in Iguazu National Park, Argentina. In addition to the expected pattern that coatis were more vigilant at the edge of the group, we found that individuals were particularly vigilant at the front edge of the group. This pattern conforms to predictions of differing predation risk caused by sit-and-wait predators with respect to mobile animal groups. In addition, coatis exhibited less vigilance when the number of neighbors within 5 m and group size increased. Of the three spatial variables tested, within-group spatial position was the most important predictor variable determining vigilance levels. These results confirm that spatial position has major effects on vigilance behavior, and that group directionality is an important factor which should be taken into account when measuring vigilance behavior. Coatis were more vigilant when juveniles less than 6 months old were in the groups. The presence of these young juveniles also affected the relationship between alarm response and vigilance levels. Coatis were more vigilant after strong alarm reactions, but only when young juveniles were not present in the groups. This may indicate that coatis give differential responses to alarm calls depending on the age of the caller. A comparison of antipredator vigilance between coatis and sympatric capuchin monkeys is consistent with the hypothesis that terrestriality leads to higher perceive predation risk for coatis.     

Within-group spatial position and vigilance: a role also for competition? The case of impalas (<Emphasis Type="Italic">Aepyceros melampus</Emphasis>) with a controlled food supply

Theory predicts that individuals at the periphery of a group should be at higher risk than their more central conspecifics since they would be the first to be encountered by an approaching terrestrial predator. As a result, it is expected that peripheral individuals display higher vigilance levels. However, the role of conspecifics in this “edge effect” may have been previously overlooked, and taking into account the possible role of within-group competition is needed. Vigilance behavior in relation to within-group spatial position was studied in impalas (Aepyceros melampus) feeding on standardized patches. We also controlled for food distribution in order to accurately define a “central” as opposed to a “peripheral” position. Our data clearly supported an edge effect, with peripheral individuals spending more time vigilant than their central conspecifics. Data on social interactions suggest that it was easier for a foraging individual to defend its feeding patch with its head lowered, and that more interactions occurred at the center of the group. Together, these results indicate that central foragers may reduce their vigilance rates in response to increased competition. Disentangling how the effects of competition and predation risk contribute to the edge effect requires further investigations.     

Self-organization in collective honeybee foraging: emergence of symmetry breaking,cross inhibition and equal harvest-rate distribution

De Vries and Biesmeijer described in 1998 an individual-oriented model that simulates the collective foraging behaviour of a colony of honeybees. Here we report how this model has been expanded and show how, through self-organization, three colony-level phenomena can emerge: symmetry breaking, cross inhibition and the equal harvest-rate distribution. Symmetry breaking is the phenomenon that the numbers of foragers visiting two equally profitable food sources will diverge after some time. Cross inhibition is the phenomenon that, by increasing the profitability of one of two equal food sources, the number of foragers visiting the other source will decrease. In some circumstances, the bees foraging on two sources of different profitabilities will be distributed between these sources such that the two average energy harvest rates are equal. We will refer to this phenomenon as the equal harvest-rate distribution. For each of these three phenomena, we show what the necessary behavioural rules to be followed by the individual forager bees are, and what the necessary circumstances are (that is, what values the model parameters should take) in order for these phenomena to arise. It seems that patch size and forager group size largely determine when each of these phenomena will arise. Experimenting with two types of currency, net gain rate and net gain efficiency, revealed that only gain rate may result in an equal harvest-rate distribution of foragers visiting different food sources.     

Division of foraging labor in ants can mediate demands for food and safety

Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström     

Look before you leap - individual variation in social vigilance shapes socio-spatial group properties in an agent-based model

Next to predator detection, primate vigilance also serves to keep track of relevant conspecifics. The degree of vigilance towards group members often reflects the dominance rank of an individual: subordinates pay attention to dominants. Although it has been suggested that subordinates' vigilance may result in spatial centrality of dominants, this has not been addressed in either empirical or modeling studies. Using agent-based models, we determined how social vigilance affects socio-spatial properties of primate groups. A basic model without social vigilance, where individuals avoid potential aggressors (avoidance model), was contrasted with two models that each additionally included a different type of social vigilance: a) monitoring a specific potential aggressor to remain informed on its whereabouts (monitoring model) or b) scanning the whole group to detect potential aggressors (scanning model). Adding monitoring or scanning behavior to the avoidance model reinforced spatial centrality of dominants, a pattern often observed in primates, and resulted in more spread out groups. Moreover, variation in scanning tendency alone was already sufficient to generate spatial centrality of dominants: frequently scanning subordinates could move further away from the group center than dominants, before losing sight of group members. In the monitoring model, two mechanisms caused decreased encounter frequencies among subordinates: a) increased inter-individual distances, and b) frequent monitoring of central dominants. In the scanning model, encounters among subordinates decreased due to increased inter-individual distances. This agent-based model study provides a clear indication that individual variation in social vigilance may be an important structuring feature of primate social groups.     

Should vigilance always decrease with group size?

Levels of anti-predator vigilance often decrease with group size in birds and mammals. Vigilance may also serve other purposes such as scrounging. While scrounging, individual adopt the vigilant posture to locate and eventually exploit the food discoveries of their companions. Models that combine anti-predator alertness and scrounging were developed to examine how changes in scrounger use with group size could alter the shape of the function relating vigilance to group size and whether the additional investment in vigilance while searching could be used to reduce the allocation of time to vigilance while in the food patch. Results indicate that increased allocation of time to scrounging during search can add to the overall level of vigilance and even counteract the expected decrease in vigilance with group size. However, the addition of scrounging was found to have little impact on vigilance while in the food patch. Behavioral effects that are dependent on group size, such as scrounging, may lead to changes in the shape of the vigilance function and may mitigate against any gain garnered from a reduction in individual levels of vigilance in larger groups.     

All clear? Meerkats attend to contextual information in close calls to coordinate vigilance

Socio-demographic factors, such as group size and their effect on predation vulnerability, have, in addition to intrinsic factors, dominated as explanations when attempting to understand animal vigilance behaviour. It is generally assumed that animals evaluate these external factors visually; however, many socially foraging species adopt a foraging technique that directly compromises the visual system. In these instances, such species may instead rely more on the acoustical medium to assess their relative risk and guide their subsequent anti-predator behaviour. We addressed this question in the socially foraging meerkat (Suricata suricatta). Meerkats forage with their head down, but at the same time frequently produce close calls (‘Foraging’ close calls). Close calls are also produced just after an individual has briefly scanned the surrounding environment for predators (‘Guarding’ close calls). Here, we firstly show that these Guarding and Foraging close call variants are in fact acoustically distinct and secondly subjects are less vigilant (in terms of frequency and time) when exposed to Guarding close call playbacks than when they hear Foraging close calls. We argue that this is the first evidence for socially foraging animals using the information encoded within calls, the main adaptive function of which is unrelated to immediate predator encounters, to coordinate their vigilance behaviour. In addition, these results provide new insights into the potential cognitive mechanisms underlying anti-predator behaviour and suggest meerkats may be capable of signalling to group members the ‘absence’ of predatory threat. If we are to fully understand the complexities underlying the coordination of animal anti-predator behaviour, we encourage future studies to take these additional auditory and cognitive dimensions into account.     

Vigilance of mustached tamarins in single-species and mixed-species groups—the influence of group composition

Species that participate in mixed-species groups (MSG) may have complementary roles in antipredator strategies. We studied vigilance in mustached tamarins (Saguinus mystax), small arboreal primates that form stable mixed-species groups with saddleback tamarins (Saguinus fuscicollis), in order to examine how the direction of vigilance changes with different species group compositions and whether the division of labor between the two species can be confirmed. We did so by comparing quantitative and qualitative differences in vigilance behavior between same individuals in and out of association (case A); MSG and single-species groups of the same total group size from two different populations (case B); and MSG of the same group size but with a different ratio of conspecifics to heterospecifics (case C). We predicted that individuals would increase downward scanning when heterospecifics are absent or their percentage is low, but total vigilance would increase only in case A due to the group size effect. However, mustached tamarins increased total vigilance due to horizontal scanning in cases A and C, and the predictions were confirmed only in small-sized groups in case B. Thus, we found indications that associating tamarin species in MSG might complement each other in the direction of vigilance, but the division of labor alone does not satisfactorily explain all the findings. There appear to be other mechanisms at work that define how direction of vigilance changes with group size and species composition. Complementarity of species probably occurs due to species vertical stratification rather than differences in the direction of vigilance.     

The influence of pollen quality on foraging behavior in honeybees (Apis mellifera L.)

Individual and colony-level foraging behaviors were evaluated in response to changes in the quantity or nutritional quality of pollen stored within honeybee (Apis mellifera L.) colonies. Colonies were housed in vertical, three-frame observation hives situated inside a building, with entrances leading to the exterior. Before receiving treatments, all colonies were deprived of pollen for 5 days and pollen foragers were marked. In one treatment group, colony pollen reserves were quantitatively manipulated to a low or high level, either by starving colonies of pollen or by providing them with a fully provisioned frame of pollen composed of mixed species. In another treatment group, pollen reserves were qualitatively manipulated by removing pollen stores from colonies and replacing them with low- or high-protein pollen supplements. After applying treatments, foraging rates were measured four times per day and pollen pellets were collected from experienced and inexperienced foragers to determine their weight, species composition, and protein content. Honeybee colonies responded to decreases in the quantity or quality of pollen reserves by increasing the proportion of pollen foragers in their foraging populations, without increasing the overall foraging rate. Manipulation of pollen stores had no effect on the breadth of floral species collected by colonies, or their preferences for the size or protein content of pollen grains. In addition, treatments had no effect on the weight of pollen loads collected by individual foragers or the number of floral species collected per foraging trip. However, significant changes in foraging behavior were detected in relation to the experience level of foragers. Irrespective of treatment group, inexperienced foragers exerted greater effort by collecting heavier pollen loads and also sampled their floral environment more extensively than experienced foragers. Overall, our results indicate that honeybees respond to deficiencies in the quantity or quality of their pollen reserves by increasing the gross amount of pollen returned to the colony, rather than by specializing in collecting pollen with a greater protein content. Individual pollen foragers appear to be insensitive to the quality of pollen they collect, indicating that colony-level feedback is necessary to regulate the flow of protein to and within the colony. Colonies may respond to changes in the quality of their pollen stores by adjusting the numbers of inexperienced to experienced foragers within their foraging populations.     

Spatio-temporal vigilance architecture of an Australian flying-fox colony

The social structure of animal aggregations may vary considerably in both space and time, yet little is known about how this affects vigilance. Here, we investigate the vigilance architecture of a colony of wild-living grey-headed flying-foxes (Pteropus poliocephalus) in Australia and examine how spatial as well as temporal variation in social organization influences social and environmental vigilance. We sampled color-marked individuals at different stages of the reproductive cycle and the year and at different locations in the colony to examine the effects of temporal and spatial factors on social and environmental vigilance. We found that vigilance architecture reflected the social structure of the colony, with the highest environmental vigilance being displayed by bats at the periphery of the colony, and the highest social vigilance by bats that roosted at intermediate distances from the colony’s edge. Furthermore, we found that vigilance levels reflected changes in reproductive state, with social vigilance increasing toward the mating season, particularly in males. Our findings show that spatial and temporal variation in social structure can have differential effects on social and environmental vigilance. This highlights the necessity to differentiate between functions of vigilance to understand fully vigilance architecture in aggregations of social animals.