Redshank Tringa totanus flocking behaviour, distance from cover and vulnerability to sparrowhawk Accipiter nisus predation

Over 11 winters I examined the interactions between sparrowhawk Accipiter nisus attack behaviour, the gregariousness of redshanks Tringa totanus and local geography to test hypotheses that suggest birds should flock to reduce their risk of predation and that predation risk should decline with the prey's distance from cover. Sparrowhawk attacks on redshanks feeding on beaches around the high tide mark (the strandline zone) were more frequent and more successful than attacks on redshanks feeding seaward of the strandline zone (in the intertidal zone). The results therefore confirmed hypothetical expectations that predation risk should decline with distance from cover. Flocking only appeared to influence the outcome of hawk attacks at shorter distances from cover on the strandline, with attacks on singletons and small flocks being more successful than attacks on larger flocks. Distance from cover had a stronger influence on the likelihood of attack success than did flock size. Mid-range flock sizes (6–45 birds) were attacked more frequently than expected, but singletons and large flocks were attacked less than expected. Despite these differences an individual redshank's likelihood of predation by a sparrowhawk declined with increasing flock size, thereby confirming the 'dilution effect' and 'vigilance' hypotheses for the evolution of flocking in birds. Food intake rates of redshanks declined with increasing flock size, further indicating that redshanks flocked to avoid predation rather than to increase their food intake rates. The strong interaction between two influences on predation risk revealed by the present study suggests other studies should take great care when considering a single influence on predation risk in isolation from others.     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

Mass-dependence in the predation risk of unequal competitors; some models

Foragers can monitor their survival through the size of body reserves in a starvation/predation risk trade-off. Energy reserves reduce the risk of energetic shortfall, while survival will be maximised at intermediate reserve levels when there is a cost of carrying mass loads. The size of reserves that will maximise survival may not be identical for unequal competitors, when unequal access to resources will affect the costs and benefits of energy reserves. Here, I evaluate the effect of competitive ability (dominance) for the mass-dependence in predation risk and how it is affected by (1) attack rate (attack rate effect), (2) distance to the emergence of an unconcealed predator attack (attack distance effect) and (3) distance to cover (cover distance effect). This general model is illustrated by empirical data for parameters specific for birds. The effect of competitive ability for the mass-dependence in predation risk is ambiguous and depends on how rank is mediated into mass-dependent predation risk. Dominants pay a lower cost in predation risk for mass loads than sub-ordinates when competitive ability entails that they feed closer to cover (cover distance effect) and when the exposure to attacks and attack rate is lower than for sub-ordinates (attack rate effect) . In contrast, a shorter distance to the emergence of an unconcealed attack (attack distance effect) implies a lower increase in predation risk with mass for sub-ordinates. As a consequence of how the cost of mass load varies with conditions there is no unambiguous relationship for how predation risk can be traded off for starvation risk for individuals with different competitive ability.     

Movements and Conflicts in a Flock of Foraging Black‐Tailed Godwits (Limosa limosa): The Influence of Feeding Rates on Behavioural Decisions

We studied movements and conflicts within a small flock of free‐living black‐tailed godwits foraging on benthic invertebrates in a brackish lagoon. To interpret our results in the framework of foraging theory, we studied the influence of individual feeding rate on the decisions to move and to attack flock companions. Birds changed their position within the flock more often when their intake rate was low and sometimes attacked conspecifics to supplant them from their feeding place. Aggressors significantly avoided front attacks and were almost always successful. They attacked individuals having higher feeding rates than themselves and their own feeding rate significantly increased after the attack, although victims were not chased off to particularly poor sites. Our results suggest that aggressors could obtain reliable information about the quality of the foraging site they coveted by observing their victim’s feeding activity before attacking. Although aggression seemed to be caused by a low intake rate, we show that displacing another bird was more time‐consuming than independent foraging. We conclude that it was not the most profitable behaviour in terms of energy intake. Foraging site displacement probably also had social functions, such as reinforcement of social status in a flock of birds preparing for pre‐breeding migration.     

Influence of predation risk on individual competitive ability and growth in Eurasian perch, Perca fluviatilis

We studied the potential influence of predation risk on the competitive ability and habitat use of foraging perch and the effect of these interactions on growth. Groups of four similar-sized young-of-the-year perch were in visual contact with a piscivorous perch during feeding. The fry had the choice of vegetation and open habitat, with food presented in the open habitat. Competitive ability, defined as proportion of prey attacks, varied between perch individuals and was unaffected by predation risk. The variation in proportion of prey attacks was affected by relative size within each replicate group, despite small size differences (±1 mm), with the largest individual being a better competitor than the smallest ones. The degree of boldness, measured as the proportion of time spent in the open habitat, was significantly related to both competitive ability and prey attack order. Observations of aggressive behaviour indicated a possible occurrence of interference competition, which may contribute to the appearance of different competitive abilities between individuals within a group of perch. A significant correlation was found between competitive ability and growth. Growth variation within groups was not affected by predation risk.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

Response to Predation Risk in Urban and Rural House Sparrows

Habitat urbanization may change the density of predators, and it is often assumed that such changes lead to altered predation risk for urban populations of their prey. Although it is difficult to study predation hazard directly, behavior responses of prey species may be informative in inferring such habitat differences. In this study, we compared the risk‐taking behavior of urban and rural house sparrows (Passer domesticus) after simulated attacks by two of their important predators (sparrowhawk Accipiter nisus and domestic cat Felis catus). The birds were startled by moving dummies of these predators and respective control objects, and their risk taking was estimated as their latency to feed after the startle. We found that sparrows responded more strongly (had longer post‐startle feeding latencies) to sparrowhawk attacks than to the control object, and their responses differed between the habitats. First, risk taking of urban birds strongly decreased with age (older birds had longer latencies than young birds), while there was no such age difference in rural birds. Second, young urban birds responded less strongly, while older urban birds responded more strongly to the sparrowhawk than the same age groups of rural birds, respectively. We did not succeed in evoking antipredatory response by simulated cat attacks, because birds responded similarly to the dummy and the control object. Our results support that predation risk, posed at least by avian predators, is different in urban and rural habitats of house sparrows. The increased wariness of older, hence presumably more experienced, urban birds implies that sparrows may be more exposed to predation in cities.     

Intraflock variation in the speed of escape-flight response on attack by an avian predator

The benefits of flocking to prey species, whether through collective vigilance,dilution of risk, or predator confusion, depend on flock members respondingin a coordinated way to attack. We videotaped sparrowhawks attackingredshank flocks to determine if there were differences in thetiming of escape flights between flock members and the factorsthat might affect any differences. Sparrowhawks are surpriseshort-chase predators, so variation in the time taken to takeflight on attack is likely to be a good index of predation risk.Most birds in a flock flew within 0.25 s of the first bird flying,and all birds were flying within 0.7 s. Redshanks that werevigilant, that were closest to the approaching raptor, and thatwere close to their neighbors took flight earliest within aflock. Birds in larger flocks took longer, on average, to takeflight, measured from the time that the first bird in the flockflew. Most birds took flight immediately after near neighbors tookoff, but later flying birds were more likely to fly immediatelyafter more distant neighbors took flight. This result, alongwith the result that increased nearest neighbor distance increasedflight delay, suggests that most redshanks flew in responseto conspecifics flying. The results strongly suggest that thereis significant individual variation in predation risk withinflocks so that individuals within a flock will vary in benefitsthat they gain from flocking.     

Effect of Group Size on Feeding Rate when Patches are Exhaustible

One benefit of group foraging is that individual foragers can join the food discoveries of companions and thus increase encounter rate with food patches. When food patches are exhaustible, however, individual shares of each patch will decrease with group size negating the effect of increased encounter rate. Mean feeding rate may actually decrease with group size as a result of aggression or time wasted joining already depleted patches, or when searching to join the food discoveries of others, which is referred to as scrounging, precludes finding food. I examined the relationship between mean feeding rate and group size in captive flocks of zebra finches (Taenopygia guttata) foraging for small clumps of seeds. Finches in groups of two or four fared better than solitary birds in terms of mean feeding rate despite the fact that birds in groups scrounged a large proportion of their food. Solitary birds initiated feeding activity after a longer delay, which led to their lower success. Early departures by food finders from food patches joined by others may have lessened the impact of scrounging on mean feeding rate. As a result of benefits from the presence of companions, group foraging in zebra finches appears a viable alternative to foraging alone despite the cost of sharing resources.     

Social Learning of a Novel Foraging Skill by White-throated Magpie-jays (Calocitta formosa,Corvidae): a Field Experiment

The cooperatively breeding white-throated magpie-jay (Calocitta formosa) uses a variety of foraging tactics to find, harvest, and process food. Members of territorial groups forage together and may gain information about how to acquire food by observing each other. A field experiment was performed to determine whether a novel skill, door opening to gain access to food, was more rapidly acquired by members of groups in which a trained individual performed the skill. A higher proportion of jays in groups with behavioural ‘models’ (trained birds that opened doors in the presence of group members) acquired the door-opening skill than those groups without models. Young birds acquired the behaviour more frequently than older individuals. Aggressive behaviour at feeders may have affected the spread of the behaviour by reducing the likelihood that individuals performed the behaviour in the presence of other group members but may also have encouraged subordinate individuals to attempt door opening rather than ‘scrounge’.     

The effects of spatial food distribution and group size on foraging behaviour in a benthic fish

Animals foraging in heterogeneous environments benefit from information on local resource density because it allows allocation of foraging effort to rich patches. In foraging groups, this information may be obtained by individuals through sampling or by observing the foraging behaviour of group members. We studied the foraging behaviour of goldfish (Carassius auratus) groups feeding in pools on resources distributed in patches. First, we determined if goldfish use sampling information to distinguish between patches of different qualities, and if this allowed goldfish to benefit from a heterogeneous resource distribution. Then, we tested if group size affected the time dedicated to food searching and ultimately foraging success. The decision of goldfish to leave a patch was affected by whether or not they found food, indicating that goldfish use an assessment rule. Giving-up density was higher when resources were highly heterogeneous, but overall gain was not affected by resource distribution. We did not observe any foraging benefits of larger groups, which indicate that grouping behaviour was driven by risk dilution. In larger groups the proportion searching for food was lower, which suggests interactions among group members. We conclude that competition between group members affects individual investments in food searching by introducing the possibility for alternative strategies, such as scrounging or resource monopolisation.     

How social network structure affects decision-making in Drosophila melanogaster

Animals use a number of different mechanisms to acquire crucial information. During social encounters, animals can pass information from one to another but, ideally, they would only use information that benefits survival and reproduction. Therefore, individuals need to be able to determine the value of the information they receive. One cue can come from the behaviour of other individuals that are already using the information. Using a previous extended dataset, we studied how individual decision-making is influenced by the behaviour of conspecifics in Drosophila melanogaster. We analysed how uninformed flies acquire and later use information about oviposition site choice they learn from informed flies. Our results suggest that uninformed flies adjust their future choices based on how coordinated the behaviours of the informed individuals they encounter are. Following social interaction, uninformed flies tended either to collectively follow the choice of the informed flies or to avoid it. Using social network analysis, we show that this selective information use seems to be based on the level of homogeneity of the social network. In particular, we found that the variance of individual centrality parameters among informed flies was lower in the case of a ‘follow’ outcome compared with the case of an ‘avoid’ outcome.     

Local prey vulnerability increases with multiple attacks by a predator

Theory and empirical evidence suggest that predator activity makes prey more wary and less vulnerable to predation. However if at least some prey in the population are energetically or spatially constrained, then predators may eventually increase local prey vulnerability because of the cumulative costs of anti‐predation behaviour. We tested whether repeated attacks by a predator might increase prey vulnerability in a system where redshanks on a saltmarsh are attacked regularly by sparrowhawks from adjacent woodland. Cumulative attack number led to a reduction in redshank numbers and flock size (but had no effect on how close redshanks fed to predator‐concealing cover) because some redshanks moved to safer but less profitable habitats, leaving smaller flocks on the saltmarsh. This effect held even though numbers of redshank on the saltmarsh increased with time of day. As a result of the change in flock size, predicted attack‐success increased up to 1.6‐fold for the sparrowhawk, while individual risk of capture for the redshank increased up to 4.5‐fold among those individuals remaining on the saltmarsh. The effect did not arise simply because hawks were more likely to attack smaller flocks because attack rate was not dependent on flock size or abundance. Our data demonstrate that when some individual prey are constrained in their ability to feed on alternative, safer foraging sites, their vulnerability to predation increases as predator attacks accumulate, although those, presumably better quality individuals that leave the immediate risky area will have lower vulnerability, so that the mean vulnerability across the entire population may not have changed substantially. This suggests that the selective benefits of multiple low‐cost attacks by predators on prey could potentially lead to 1) locally heightened trait‐mediated interactions, 2) locally reduced interference among competing predators, and 3) the evolution of active prey manipulation by predators.     

Predicting the optimal prey group size from predator hunting behaviour

1.?How group size affects predator attack and success rate, and so prey vulnerability, is important in determining the nonlethal consequences of predation risk on animal populations and communities. Theory predicts that both predator attack success rate and the dilution effect decline exponentially with group size and that selection generates optimal group sizes at a 'risk threshold' above which antipredation benefits are outweighed by costs, such as those owing to higher attack rates. 2.?We examined whether flock size risk thresholds for attack rate, success rate or dilution differed, and therefore whether the strength of selection for group size differed for these three factors, using a system of redshank Tringa totanus flocks being hunted by Eurasian sparrowhawks Accipiter nisus. We also asked which of the three thresholds, on their own or in combination, predicted the most commonly observed group size. 3.?Mean redshank flock size increased with a very gradual quadratic function (i.e. approximately linearly) with population size, although at a rate half that possible; when population size was not limiting, individuals almost always avoided flocks of less than 30 and birds were frequently in flocks up to at least 80. Sparrowhawk attack rate showed a quadratic relationship with flock size and peaked at 55 redshanks. Sparrowhawk attack success rate, however, declined exponentially, becoming less steep at flock sizes of about 40 and remaining uniformly low (a 95% decrease) by 70. Combined with dilution, individual risk of death per attack decreased by 95% when group size reached 30 (20 for the dilution effect alone). 4.?Redshanks most commonly formed group sizes that gained the maximum individual predation risk reduction. They also commonly formed group sizes far above any further substantial advantages from the dilution effect or from reducing attack rate, but that continued to reduce predation risk by lowering attack success rate. Individuals did not always form the largest groups possible which we suggest is because individual variation in risk-taking subdivides the population. This places a constraint on the ability of individuals to compensate for predation risk and will have a variety of important effects on animal populations.     

Is social coordination during escape flights a general phenomenon in birds?

Escape from predatory attack as a socially coordinated group is observed in many social animals, including birds, especially those in more open habitats where the group itself may be the only source of protection from an attacking predator. For many social birds, however, woody vegetative cover is the main refuge from attack, but such birds might nevertheless benefit from social coordination during escape flights to cover. Such benefits could reflect the confusion effect, selfish herd effect, or the simple dilution of risk. We examined this possibility of coordinated escape in mixed flocks of wintering passerellid sparrows (Passerellidae). These free-living birds fed in a patch of food flanked on opposite sides by two refuges composed of woody cover. Under such conditions, coordination in escape behavior should be expressed as a tendency to escape together as a group to the same cover location. Such behavior, however, was not the rule. During spontaneous flushes to cover, a group of escaping birds stayed together only when one cover location was clearly closer than the other. With cover equidistant from the food patch, escaping flocks tended to split about evenly between cover locations. Birds in close proximity prior to an escape flight did not show enhanced escape coordination, nor did those feeding at significant distances from protective cover. Evidence of escape coordination was observed in small groups (two–four birds), but even in such groups, flock splitting during escape was generally the rule. Flock splitting during attacks might reflect some sort of strategic decision-making process that lessens the risk of capture, but the most parsimonious explanation is that (all else equal) birds head for the nearest refuge, largely irrespective of the behavior of their flockmates. Our results thus provide little evidence of flock-wide social coordination during escape flights in cover-dependent birds.     

Predation Risk Perception,Food Density and Conspecific Cues Shape Foraging Decisions in a Tropical Lizard

When foraging, animals can maximize their fitness if they are able to tailor their foraging decisions to current environmental conditions. When making foraging decisions, individuals need to assess the benefits of foraging while accounting for the potential risks of being captured by a predator. However, whether and how different factors interact to shape these decisions is not yet well understood, especially in individual foragers. Here we present a standardized set of manipulative field experiments in the form of foraging assays in the tropical lizard Anolis cristatellus in Puerto Rico. We presented male lizards with foraging opportunities to test how the presence of conspecifics, predation-risk perception, the abundance of food, and interactions among these factors determines the outcome of foraging decisions. In Experiment 1, anoles foraged faster when food was scarce and other conspecifics were present near the feeding tray, while they took longer to feed when food was abundant and when no conspecifics were present. These results suggest that foraging decisions in anoles are the result of a complex process in which individuals assess predation risk by using information from conspecific individuals while taking into account food abundance. In Experiment 2, a simulated increase in predation risk (i.e., distance to the feeding tray) confirmed the relevance of risk perception by showing that the use of available perches is strongly correlated with the latency to feed. We found Puerto Rican crested anoles integrate instantaneous ecological information about food abundance, conspecific activity and predation risk, and adjust their foraging behavior accordingly.     

Predictability of repeated carnivore attacks on livestock favours reactive use of mitigation measures

1. Predation on livestock is one of the main reasons for low tolerance against large carnivores in many parts of the world. Measures to reduce the conflicts have been developed, but resources for using them are often scarce. If wildlife managers as well as farmers learn more about when the risk of predation on livestock is higher, they will be able to make more effective use of resources for reducing predation.
2. In this study, we tested the hypothesis that the risk of predation on livestock immediately after an attack is higher on the affected farm compared with other farms in the same area. Data on sheep predation by brown bear Ursus arctos , lynx Lynx lynx and wolf Canis lupus in Sweden 1998–2006 were used in the analysis.
3. On depredated farms there was approximately a 55 times higher risk for a repeat predation event within 12 months compared to any other farm in the same area. During the first 5 weeks, 63%, 60% and 50% of the repeat attacks had occurred.
4. We suggest that the main mechanism behind repeat attacks on livestock is that carnivores return to the kill site to feed on carrion. Where livestock are still present and unprotected at the kill site when the carnivore returns, the farms will suffer a higher likelihood of a further attack compared to livestock on other farms. This study uses data from Sweden but we argue that the pattern will be the same in any part of the world where the ranges of livestock and large carnivores overlap.
4. Synthesis and applications . As the risk of an attack is higher directly after an initial attack, it will be more cost-effective to implement measures designed to reduce livestock predation by large carnivores at that time, i.e. within the following 5 weeks. Temporary proactive measures are usually simpler and cheaper than permanent deterrents and we recommend their use wherever resources are limited.     

Mechanisms regulating bird predation on a herbivorous Larva guild in boreal coniferous forests

Bird predation was previously found to considerably reduce the abundance of the herbivorous insect larva guild feeding on bilberry Vacanium myrtillus within boreal coniferous forests In this follow-up study, interest was focused on determining whether female pied flycatchers Ficedula hypoleuca tend to use the field layer more frequently than other types of feeding niches In addition, the degree to which the conspicuousness of a larva increases the risk of its being preyed upon by birds was evaluated in order to further understand the mechanisms regulating the bird predation effect Larval predation risks were measured by presenting larvae with contrasting characteristics to seven captured female flycatchers
Female flycatchers hunted more frequently in the tree and field layers than in the air However, the relative degree of utilization of the tree and field layer varied considerably between years, with use of field layer being positively related to the abundance of insect larvae on bilberry
Large larvae did not suffer higher bird predation than small larvae when both size classes were presented to the birds on bilberry Larvae that crawl on leaves and stems during feeding (high degree of exposure) were preyed upon more than larvae that feed between leaves they have spun together (low degree of exposure) No difference in larval predation risk was found between sawflies and geometrids, the two taxa represented in the exposed feeder group However, dark geometrids suffered higher predation than green sawflies and green geometrids
My findings suggest that the herbivorous larva guild constitutes an abundant food resource frequently utilized by small bird predators However, certain members of the guild appeared to be preyed upon more than other members Thus the risk for predation seems to be highest for dark geometnds followed in decreasing order by green geometrids. sawflies pyralids, and tortricids     

Cooperation in defence against a predator

The origin and the evolutionary stability of cooperation between unrelated individuals is one of the key problems of evolutionary biology. In this paper, a cooperative defence game against a predator is introduced which is based on Hamilton's selfish herd theory and Eshel's survival game models. Cooperation is altruistic in the sense that the individual, which is not the target of the predator, helps the members of the group attacked by the predator and during defensive action the helper individual may also die in any attack. In order to decrease the long term predation risk, this individual has to carry out a high risk action. Here I show that this kind of cooperative behaviour can evolve in small groups. The reason for the emergence of cooperation is that if the predator does not kill a mate of a cooperative individual, then the survival probability of the cooperative individual will increase in two cases. If the mate is non-cooperative, then—according to the dilution effect, the predator confusion effect and the higher predator vigilance—the survival probability of the cooperative individual increases. The second case is when the mate is cooperative, because a cooperative individual has a further gain, the active help in defence during further predator attacks. Thus, if an individual can increase the survival rate of its mates (no matter whether the mate is cooperative or not), then its own predation risk will decrease.     

Anti-Predator Strategies and Grouping Patterns in White-Tailed Deer and Mule Deer

White-tailed deer ( Odocoileus virginianus ) and mule deer ( O. hemionus ) are closely related species of similar size that differ in their anti-predator behavior. White-tails flee from coyotes ( Canis latrans ), whereas mule deer typically stand their ground and attack this predator. I used observations of coyotes hunting deer to identify: (i) changes in group structure made in response to coyotes; and (ii) the relationship between group structure and the risk of predation for each species.
In response to coyotes, groups of mule deer merged with other groups and individuals bunched together. Predation attempts were more likely to escalate when groups split and individuals failed to bunch. Coyotes typically attacked mule deer that were in outlying positions, and these deer had to move to central positions to end attacks. Due to the high frequency of attacks on small groups as well as to the level of dilution of risk, individuals in small mule deer groups were at high risk of being attacked compared with those in larger groups. In contrast to mule deer, white-tails made no consistent changes in group size or formation, and coyotes attacked individuals in central as well as in outlying positions. Variation in aspects of group cohesion was not related to the vulnerability of white-tails, and there was no obvious difference in the risk of attack facing individuals in groups of different size. These results suggest that coyote predation selects for relatively large, cohesive groups in mule deer, apparently because this type of group improves their ability to deter coyotes. Coyote predation does not have similar effects on groups formed by white-tails, which use flight rather than deterrence to avoid predation. The benefits of responding cohesively, occupying certain positions within groups, and forming groups of a certain size can vary widely depending on the anti-predator strategies used by an animal.