Predation by Kelp Gulls Larus dominicanus at a mixed-species colony of Royal Terns Sterna maxima and Cayenne Terns Sterna eurygnatha in Patagonia

We studied the predator-prey interaction between Kelp Gulls Larus dominicanus and Royal Terns Sterna maxima and Cayenne Terns Sterna eurygnatha at Punta León, Argentina, from 1990 to 1992. Kelp Gulls were the main predator of tern eggs in all 3 years, accounting for 99% of observed predations (n = 454). A total of 99% of predations occurred on peripheral tern nests. Total predation rate on both tern species was highest in 1990 and varied significantly between years, with between 0.01 and 0.03 cases of predation per hour. Predation rates in all years were significantly higher on Cayenne than on Royal Terns probably because the latter were significantly more aggressive towards gulls than were Cayenne Terns. Royal and Cayenne Terns displayed aggressively towards gulls at least once in 64% and 10% of the observation periods, respectively. Defence behaviour, such as attacking and chasing the predator, was used more frequently by Royal than Cayenne Terns (13.2% v 3.6%). Predation on tern eggs was restricted to only a few Kelp Gulls which nested close to the tern colony periphery, and 50% of the eggs that were stolen were taken by only two pairs during 1991 and 47% by three pairs in 1992. This suggests that the current expansion of the Kelp Gull numbers at Punta León might not necessarily result in a proportional increase in predation rate on both tern species. Stealing eggs from the ground was more successful than from the air (77% v 17%). The nesting strategy shown by Royal and Cayenne Terns at Punta León appears to be an important factor which reduced the impact on their breeding success of high potential predation pressure by gulls.     

Behavioural ecology of lonnbergi skuas in relation to environment on the Chatham Islands,New Zealand

Abstract

A population of lonnbergi skuas (family Stercorariidae) was studied over December and January 1974–75 on Rangatira Island, Chatham group (44°22′S,176°11′W). The main aim was to compare the behavioural ecology of these birds with that of maccormicki skuas of Antarctica, and to assess the moderating effect of environment on habits. The skuas held extensive territories along the coast and in inland areas clear of forest and bracken. Five of the 11 study territories were occupied by trios of adult birds. A major effect of the vegetation was to reduce contact between neighbours on the ground at territorial boundaries, and most territorial defence was by aerial display and attack. Eggs were laid from late September, and chicks began flying in late December and January. This breeding schedule is about 2 months earlier than for lonnbergi at Signy Island or for skuas in Antarctica, but falls on a trend line of latitude against egg-laying when all populations are considered. Seventeen chicks were reared on the 11 territories, a breeding success of 1.55 chicks per nest. This high success rate was attributable largely to success in rearing both chicks when two were hatched. This is seldom achieved in more extreme climates, and is apparently determined by the intensity of sibling competitiveness. It is hypothesised that this intensity is graded, and aggression by the older chick of the pair is provoked more easily by food shortage as environmental conditions become increasingly severe. The most important prey items recorded were adult petrels, especially broad-billed prion (Pachyptila vittata vittata) and white-faced storm petrel (Pelagodroma marina maoriana) captured on the ground at night. The skuas were mostly inactive during the day, and were not seen to fish or to chase or harass any other bird on or about the island. Southern blue penguins (Eudyptula minor minor), though very common on the island, were not attacked by the skuas. Skuas were not seen to attempt to dig out petrel or penguin nesting burrows. Of the differences in ecology recorded, only the presence of three adults on territories appears to be characteristic of lonnbergi; it has not been recorded for maccormicki or hamiltoni, and is either unknown or very rare in skua.     

THE SIGNIFICANCE OF KLEPTOPARASITISM DURING THE BREEDING SEASON IN A COLONY OF ARCTIC SKUAS STERCORARIUS PARASITICUS IN ICELAND

Breeding and kleptoparasitism of Arctic Skuas was studied at Vik, Iceland, in 1973. Hatching success was 88.9%; fledging success was 0.27 fledglings per pair. The heavy chick mortality occurred mostly in the first week after hatching. In the early part of the breeding season skuas fed by robbing kittiwakes of their food at sea and by eating arthropods at the breeding grounds. At the time of hatching of skua eggs, which coincided with the hatching of Puffin eggs, skuas switched their feeding activities to kleptoparasitism of Puffins and fish so obtained was the principal item of most skuas' diet thereafter. In the first week post-hatching, the energy balance of an average adult skua pirating Puffins was estimated to be negative, but changed to positive a week later, although later, coinciding with an influx of non-breeders it turned back to negative. Arctic Terns which normally breed at Vik, and are exploited by skuas, failed to breed in 1973, and the abnormally heavy chick loss is therefore attributed to this failure of the terns. The success of the skuas kleptoparasitizing Puffins depends on the proximity of the interactions to the cliff or to the ocean, where Puffins seck refuge. Skuas catch fish mostly in the air, especially if it is dropped from high. Fish dropped from low is mostly taken by competing gulls and Ravens, which mostly control the ground and lower airspace. Skuas chasing in groups enhance their success, but the average success per member decreased with group increase. However, the success of the group-member in the ‘best’ position was equal to that of a single skua. Puffins carry 1 (large)-6 (smaller) fish, an approximately equal load irrespective of number of fish. Skuas preferentially chase Puffins carrying ‘large’ fish, thus maximizing their yield per effort. Arctic Skuas responded to changes in the numbers of arriving Puffins with a functional response, but their monitoring of the food supply was far from perfect.     

Effects of opportunistic predation on anti-predator behavioural responses in a guild of ground foragers

We studied factors that affect prey selection by a generalist predator that opportunistically attacks prey species, and the associated inter- and intra-specific responses of prey to this type of predation. Our model system was a guild of ground-foraging birds that are preyed upon by magpies (Pica pica) during the breeding season. We found that magpies attacked up to 12 species during three consecutive breeding seasons. The overall capture success was estimated to be 4.9%. Magpies tended to attack from the air, targeting solitary prey, either on the ground or flying. Inter-specific prey responses to the risk of magpie predation included a reduction in the mean number of species occupying a foraging patch when magpies were present and a decrease in the distance between heterospecific neighbours. Intra-specific responses to magpie predation varied between species that were subject to different attack rates. Preferentially attacked prey enhanced their risk responses (increase in scanning time and scanning rate in the presence of magpies) relative to those species attacked in proportion to their abundance (increase only in scanning rate with magpies). Species attacked infrequently, relative to their abundance, showed no antipredator response. The probability of being attacked, rather than mortality rate, appears to be the factor to which prey species respond.     

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.     

Prey susceptibilities,prey utilization and variable attack efficiencies of Ural owls

Summary To investigate the factors that influence prey utilization among predators with active prey, three series of experiments were performed in which Ural owls (Strix uralensis) searched for and attacked three prey species of wild mice, Microtus montebelli, Apodemus speciosus, and A. argenteus, in a large flight cage. Over the whole study, owls attacked mice about ten times a night. The number of attacks on each prey species did not differ from that predicted by a random attack model. M. montebelli was taken more than either Apodemus species. Prey utilization appears to be influenced by prey susceptibilities only and it is unlikely that prey selection by the owls affected prey utilization patterns. Under the experimental conditions, random attack is predicted by optimal foraging theory. However, random attack may be explained just as well by the inability of the owl to discriminate prey type. The owls, energy gain was adjusted not by alteration in the number of attacks on a prey species but rather by alteration in the capture success between experiments. Capture success increased in poor food conditions for the same prey species. This flexibility in capture success has not been considered in the assumptions of optimal foraging theory. In conventional optimal foraging theory, the probability of capture success is implicitly assumed as constant and unity. We suggest that this assumption is inadequate to understand the foraging behavior of owls.     

Feeding ecology of wintering terns in Guinea-Bissau

We studied the feeding ecology of Little Terns Sterna albifrons , Sandwich Terns S. sandvicensis and Royal Terns S. maxima in the Archipélago dos Bijagós (11°40'N, 15°45'W) in Guinea-Bissau (West Africa) during the winter of 1992/1993. More than 95% of all prey taken by these terns were roundfish, ranging in weight from 0.3 to 40 g. Birds usually fed alone, but sometimes they were observed feeding in mixed-species flocks consisting of 15–200 individuals. Capture rate ( n fish per hour foraging) in these flocks was higher than that of solitary birds. However, smaller fish were caught by birds foraging in flocks, so food intake rate (g/h) did not differ between solitary and flock-feeding birds. The relationships between foraging behaviour of the three tern species and abiotic factors, such as time, tide and water clarity, have been investigated. Capture rate of Royal Terns increased with water clarity. For Little Terns and Sandwich Terns, food intake rate was lower in the most turbid waters compared to clearer waters. There was very little foraging activity during high tide. For Little Terns and Royal Terns, food intake rate was about twice as high during receding and low tides as during an incoming tide. Food intake rate averaged 8 g/h in Little Terns, 60 g/h in Sandwich Terns and 45 g/h in Royal Terns. With a rough model, we estimate the maximum rate of daily energy expenditure of terns wintering in the tropics at 3 × BMR (defined as energy expenditure of inactive bird at thermoneutrality in a post-absorptive state during the resting phase of the daily cycle). From an energetic viewpoint, wintering Sandwich Terns in Guinea-Bissau seem to have an easy living.     

Surprise as a winter hunting strategy in Sparrowhawks Accipiter nisus,Peregrines Falco peregrinus and Merlins F. columbarius

Sparrowhawks Accipiter nisus, Peregrines Falco peregrinus and Merlins Falco columbarius were studied hunting Redshanks Tringa totanus, Dunlins Calidris alpina and Skylarks Alauda arvensis over three winters on a small Scottish estuary. Most Sparrowhawk and Merlin hunts consisted of a single attack (mean = 1.0 and 1.1, respectively), whereas Peregrine hunts often consisted of several attacks (mean = 1.8). Most hunts were short (<1 min), but Peregrine and Merlin hunts occasionally lasted over 5 min. In general, all three raptor species attacked by surprise, although Peregrines were more likely to make nonsurprise attacks. Prey attacked were usually initially very close to the raptor (<100 m); Peregrines attacked prey most often at long distances (>500 m). Chase lengths were mostly <5 second in length, although Peregrines, and particularly Merlins hunting Skylarks, often chased for several minutes. Peregrines attacked most prey in flight from flight, while Merlins and Sparrowhawks attacked birds on the ground with a flight from a perch. All three raptor species preferentially attacked larger Dunlin flocks, but Peregrines also favoured single birds. Capture rates of Redshanks and Dunlins were similar for the three raptor species (C. 10%), but for Skylarks, capture rate by Merlins was much higher (12%) than by Sparrowhawks (3%) or Peregrines (0%). Capture rates were highest when raptors attacked by surprise, particularly for a Peregrine hunting in the first minute of arrival on the study site if no Peregrines had been hunting there for the previous hour (16% success for the first minute compared with 2% in subsequent minutes). Sparrowhawks were more successful when attacking small rather than large Dunlin flocks. The use of short surprise attacks interspersed with long periods of inactivity was common to the three raptor species and was interpreted as a strategy to minimize the amount of energy and risk involved in hunting during the nonbreeding season.     

Effect of predation by skuas on breeding success of the Cape petrel Daption capense at Nelson Island, Antarctica

Breeding success of Cape petrels at Nelson Island (South Shetland Islands) in 1991/1992 averaged 29%. Predation by skuas accounted for 76% of all nesting failures. Direct attacks prevailed over opportunistic predation. Intensity of observer disturbance alone could not account for the inter-colony variation in predation rate, which tended to increase with colony size, presence of nearby breeding skuas and local availability of other prey species (penguins). Predation was significantly lower on nests that could be attacked only, or more easily, from the air and nests with close neighbours. Selective predation on nests in which relatively narrower eggs were laid was independent of nest-site characteristics and laying date, suggesting an effect of parental age/experience on breeding success through antipredator behaviour. Development of oil-spitting behaviour in growing chicks balanced the parental attendance and effort in nest defence, which declined as chick age increased. Increased predation on lighter but not smaller chicks may reflect the lower ability of such chicks to defend themselves by spitting stomach oil. Seasonal variations of predation rate on Cape petrel nests did not result from predator swamping by numbers of simultaneously active nests, but rather reflected temporal availability of alternative prey. Success of all phases of nesting declined with later laying date. Received: 15 December 1997 / Accepted: 11 April 1998     

EFFECT OF AGE ON THE FISHING ABILITY OF SANDWICH TERNS STERNA SANDVICENSIS

The fishing ability of first-winter and older Sandwich Terns was compared at foraging sites used by both age-groups on the coast of Sierra Leone between January and March 1970 when first-winter birds were about 7–9 months old. The methods used for measuring various parameters of fishing ability are described. It was found that a significantly larger proportion of plunge dives by the older age-group yielded prey although the difference between the two age-groups was only slight.
There were no major differences between the age-groups in diving rates or in size of fish captured. Older birds tended to dive from higher than first-winter birds.
By driving shoals of small fish to the surface, predatory fish enhanced the fishing success of both age-groups on certain days. Where man's beach seining operations yielded a superabundance of small fish near the tideline, these were captured equally well by both age-groups.
First-winter birds were not seen being fed by older birds, and although food-begging behaviour was observed once in late January it is clear that the first-winter birds were self-sufficient for food by the time this study was made.
Some implications of the observed differences in fishing success are discussed. It is suggested that, after fledging, Sandwich Terns gradually gain the judgment and skill necessary to dive profitably from increasing heights and so learn to exploit prey in a greater range of water depth as they get older.     

Vigilance during food handling by Oystercatchers Haematopus ostralegus reduces the chances of losing prey to kleptoparasites

Oystercatchers Haematopus ostralegus periodically pause while handling mussels Mytilus edulis to make visual scans. This paper presents evidence that scanning is associated with the high incidence of intra-specific food stealing among mussel-eating Oystercatchers. Scanning increased in frequency as bird density - and the risk of being attacked for mussels increased and the duration of attacks decreased. Additionally, among a sample of individually marked adults, the aggressive dominant birds spent half as much time scanning as the less aggressive subdominants and were also less likely to be attacked. Whereas detecting an attack made no difference to the success with which the dominants defended their mussels, subdominants increased their chances of retaining the mussel if they detected and carried the mussel away from the approaching attacker. The extra time which the less aggressive birds spent in vigilance seems best understood as a tactic for reducing food loss to kleptoparasites.     

Aggression by adult South American Terns toward conspecific chicks

ABSTRACT Infanticide and nonfatal aggression by adults toward unfamiliar chicks have been widely reported in colonial birds, and can be an important cause of chick mortality. We studied intraspecific aggression by adults toward chicks at a South American Tern (Sterna hirundinacea) colony in Patagonia, Argentina, during 2005 to characterize this behavior, evaluate its relationship with nesting density, chick age and microhabitat characteristics, and assess its effect on breeding success. Of 111 chicks in the study area, 45% were attacked at least once. Chicks older than 9 d posthatching were more likely to be attacked than younger chicks, and unattended chicks were more likely to be attacked than guarded chicks (88 vs. 12%). Chicks were also attacked more often when in their own territories (76% of cases), but were less likely to be attacked in territories with more vegetation cover and high‐quality shelters (i.e., vegetation with characteristics that prevented adults from reaching chicks). The number of aggression events was not related to nest density. At least 8% of the chicks in our study area died as a result of adult intraspecific aggression. Our results indicate that intraspecific aggression by adult South American Terns toward chicks is relatively common in the Punta Loma colony and should not be underestimated as a factor affecting their breeding success.     

Brood defence and parental roles in a biparental cichlid fishLamprologus toae in Lake Tanganyika

Brood defence of a cichlid fish,Lamprologus toae, was investigated in its natural habitat in Lake Tanganyika. Both parents guarding a brood attacked both conspecific and heterospecific intruder fishes at different locations. The heterospecific intruder fishes could be classified into three groups on the basis of the locations at which the attacks against each species took place. The distinction of groups by the parents seemed to be primarily based on food habits and feeding behaviour of the intruder fishes. The piscivorous species which were more dangerous for the brood were attacked by both parents at more distant locations from the brood. Parental defence of breeding territory changed with the development of the young. The frequency of attacks against each group decreased after the young reached the size too large for the fishes of the group to prey on. Division of labour in the territorial defence was recognized between male and female. The male parent mainly defended the peripheral region of the territory and the female parent defended the inner region. Significance of the selective attack against intruders and the division of labour between the two sexes in brood defence is discussed.     

The Role of Body Size in Mating Interactions of the Sexually Cannibalistic Fishing Spider Dolomedes triton

Some arachnids display extreme sexual size dimorphism (SSD) with adult females being several times larger than adult males. One explanation for SSD in species that exhibit pre‐copulatory sexual cannibalism (female attack, kill and consumption of the male prior to mating) is that smaller males may be less likely victims of predatory attacks by females. However, in some sexually cannibalistic species SSD is relatively moderate (i.e. males are similar in size to females) suggesting benefits of large male body size. Here, I report the results of an experiment designed to explore the ramifications of body size in mating interactions of the sexually cannibalistic, North American fishing spider (Dolomedes triton). Results suggest that male size does not influence courtship behavior, the likelihood of being attacked, or the male's ability to secure a mounting. However, large males were superior at gaining copulations once mounted. Sexual cannibalism may also be predicated on female size. Female condition (mass/cephalothorax area) did not explain any of these behaviors from the copulatory sequence, however, females with a smaller cephalothorax area were more likely to attack courting males. Finally, analysis of the ratio of female size to male size showed that when SSD is weak males are more likely to escape attacks and mate successfully. Results are discussed in light of several hypotheses for sexual cannibalism, and the benefits of large male body size illustrated here are put forth as potential explanations for the relatively moderate extent of SSD found in this sexually cannibalistic species.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

Island shape,size, and isolation affect nest‐site selection by Little Terns

ABSTRACT Populations of many seabirds and other species that nest along coasts are declining due to habitat degradation and loss. An improved understanding of the species‐specific factors that determine nest density across a landscape is therefore critical for conservation efforts. We examined factors that affected the density (number per hectare) and abundance (number at a sampling site) of nests of Little Terns (Sternula albifrons) on the Sinai Peninsula, Egypt. Terns preferred to nest on islands rather than the mainland, with islands constituting 64% of the area surveyed, but containing 99% of the 439 tern nests we found. Nest densities were highest on islands that were small, located at moderate distances from the mainland, and irregularly shaped or elongated. Most nests (69%) were on islands with areas < 3 ha, although these islands represented < 5% of total island area, and islands with the highest nest densities were 80–300 m from the mainland. Terrestrial predators were more likely to occur on larger islands, visiting three of the largest four islands. Most tern nests were within 1 m of shorelines, causing island perimeter to be a strong influence on nest density. Island shape was the only factor that significantly affected nest abundance, with more nests on islands with relatively long perimeters for their size. Our results suggest that protection or creation of relatively small, slender islands at moderate distances from shore may be an effective means of increasing the number of breeding sites for Little Terns. Although not generally considered a potential determinant of nest site preferences for seabirds, island shape is likely to be important for species that prefer sites adjacent to water, including species that nest on beaches and seaside cliffs.     

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.     

Is there always an influence of shoal size on predator hunting success?

Theoretical and empirical studies predict that there should be a decrease in hunting success of predators with increasing prey group size. Most of these studies investigated situations in which predator and prey were in full view of each other before, during and after an attack. In this study, single rock bass Ambloplites rupestris were given an opportunity to launch surprise attacks at shoals of creek chub Semotilus atromaculatus that ranged in size from two to 13 fish. There was no significant influence of either shoal size or attack distance on predator success rate and no significant relationship between attack distance and shoal size. Furthermore, it was found that the leading fish of a shoal was attacked significantly more often than fish in other shoal positions, indicating that predation risk was not shared equally among shoal members. Also, leading fish in larger shoals (eight to 13 fish) were not more likely to survive a predator attack than ones in small shoals (two to seven fish).The consequences of these results are discussed in the general context of antipredator benefits of grouping.     

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.     

Bird attacks on a butterfly with marginal eyespots and the role of prey concealment against the background

Small eyespots on butterflies have long been thought to deflect attacks, and birds are the presumptive drivers selecting for these patterns; however, evidence of this function is still ambiguous. Marginal eyespots typically consist of a UV‐reflective white pupil, surrounded by one black and one yellowish ring. We have recently shown that Cyanistes caeruleus (blue tits) attack such eyespots, but only under low light intensities with accentuated UV levels: the increased salience of the eyespots relative to the rest of the butterfly probably explains this result. Possibly the background against which the butterfly is concealed may deceive birds to make similar errors. We therefore presented speckled wood butterflies decorated with eyespots (or controls without eyespots) to C. caeruleus against two backgrounds: oak and birch bark. Our results show that: (1) eyespots, independent of background, were effective in deflecting attacks; (2) the time elapsed between a bird landing and the attack was interactively dependent on the background and whether the butterfly bore an eyespot; and (3) the speed at which a butterfly was attacked predicted the outcome, with faster birds being more prone to errors than slower birds. This underscores a speed–accuracy trade‐off in the predators, and that background plays a role in the defensive qualities of marginal eyespots. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 290–297.