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.     

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.     

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.     

Modelling antipredator vigilance and flight response in group foragers when warning signals are ambiguous

The trade-off between feeding and vigilance in flocks of birds has been extensively studied and modelled. An assumption of many models is that if one bird spots the predator, it gives a signal and the rest of the flock takes flight. However, it has been observed that birds do not always respond to signals and in fact many signals turn out to be false alarms. Since taking flight is both costly in time and energy, it may be advantageous for birds not to respond to all alarm calls. A model is developed to show under what circumstances birds should respond to a signal. The model predicts that under most, but not all, circumstances, birds should respond to multiple detections but not to single detections. The model also predicts that if birds respond to all flights, they will have to compensate for the time lost to feeding and the greater energy requirement of spending more time in flight, by being less vigilant, and they have a lower probability of survival than birds which only respond to multiple detections.     

Temporal and Contextual Consistency of Leadership in Homing Pigeon Flocks

Organized flight of homing pigeons (Columba livia) was previously shown to rely on simple leadership rules between flock mates, yet the stability of this social structuring over time and across different contexts remains unclear. We quantified the repeatability of leadership-based flock structures within a flight and across multiple flights conducted with the same animals. We compared two contexts of flock composition: flocks of birds of the same age and flight experience; and, flocks of birds of different ages and flight experience. All flocks displayed consistent leadership-based structures over time, showing that individuals have stable roles in the navigational decisions of the flock. However, flocks of balanced age and flight experience exhibited reduced leadership stability, indicating that these factors promote flock structuring. Our study empirically demonstrates that leadership and followership are consistent behaviours in homing pigeon flocks, but such consistency is affected by the heterogeneity of individual flight experiences and/or age. Similar evidence from other species suggests leadership as an important mechanism for coordinated motion in small groups of animals with strong social bonds.     

Predator–prey interactions,flight initiation distance and brain size

Prey avoid being eaten by assessing the risk posed by approaching predators and responding accordingly. Such an assessment may result in prey–predator communication and signalling, which entail further monitoring of the predator by prey. An early antipredator response may provide potential prey with a selective advantage, although this benefit comes at the cost of disturbance in terms of lost foraging opportunities and increased energy expenditure. Therefore, it may pay prey to assess approaching predators and determine the likelihood of attack before fleeing. Given that many approaching potential predators are detected visually, we hypothesized that species with relatively large eyes would be able to detect an approaching predator from afar. Furthermore, we hypothesized that monitoring of predators by potential prey relies on evaluation through information processing by the brain. Therefore, species with relatively larger brains for their body size should be better able to monitor the intentions of a predator, delay flight for longer and hence have shorter flight initiation distances than species with smaller brains. Indeed, flight initiation distances increased with relative eye size and decreased with relative brain size in a comparative study of 107 species of birds. In addition, flight initiation distance increased independently with size of the cerebellum, which plays a key role in motor control. These results are consistent with cognitive monitoring as an antipredator behaviour that does not result in the fastest possible, but rather the least expensive escape flights. Therefore, antipredator behaviour may have coevolved with the size of sense organs, brains and compartments of the brain involved in responses to risk of predation.     

Contagious fear: Escape behavior increases with flock size in European gregarious birds

Flight initiation distance (FID), the distance at which individuals take flight when approached by a potential (human) predator, is a tool for understanding predator–prey interactions. Among the factors affecting FID, tests of effects of group size (i.e., number of potential prey) on FID have yielded contrasting results. Group size or flock size could either affect FID negatively (i.e., the dilution effect caused by the presence of many individuals) or positively (i.e., increased vigilance due to more eyes scanning for predators). These effects may be associated with gregarious species, because such species should be better adapted to exploiting information from other individuals in the group than nongregarious species. Sociality may explain why earlier findings on group size versus FID have yielded different conclusions. Here, we analyzed how flock size affected bird FID in eight European countries. A phylogenetic generalized least square regression model was used to investigate changes in escape behavior of bird species in relation to number of individuals in the flock, starting distance, diet, latitude, and type of habitat. Flock size of different bird species influenced how species responded to perceived threats. We found that gregarious birds reacted to a potential predator earlier (longer FID) when aggregated in large flocks. These results support a higher vigilance arising from many eyes scanning in birds, suggesting that sociality may be a key factor in the evolution of antipredator behavior both in urban and rural areas. Finally, future studies comparing FID must pay explicit attention to the number of individuals in flocks of gregarious species.     

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.     

Flock feeding and time budgets in the house sparrow (Passer domesticus L.)

A winter population of house sparrows at a farm fed on barley seed in two distinct types of habitat: cattlesheds and open fields. The risk of predation was apparently higher in the fields where birds scanned more frequently than in the cattlesheds and where scanning was negatively influenced by flock size but positively influenced by distance from cover. Individual time budgets were more influenced by flock size than by seed density in the fields but more influenced by seed density than by flock size in the cattlesheds. Higher rates of scanning resulted in greater flock vigilance and longer flight distances in the fields but flight distance was negatively influenced by the density of seeds on which birds were feeding.     

ROOSTING MOVEMENTS OF BIRDS AND MIGRATION DEPARTURES FROM ROOSTS AS SEEN BY RADAR

Unusual radar echoes attributable to the roosting movements of large flocks of birds, and to departures from roosts on migration flight are illustrated. Probably the most striking displays are received from Starlings Sturnus vulgaris. Radar evidence is shown of a much more rapid break-up of flocks on migration at night than by day. Weather at departure points is studied.     

Patterns and correlates of songbird movements at an ecological barrier during autumn migration assessed using landscape‐ and regional‐scale automated radiotelemetry

Departure decisions of songbirds at ecological barriers they encounter en route can strongly influence time, energy and survival costs of migration. To date, most field studies of departure decisions and their correlates have used indirect methods and followed migrants at a single stopover site, with limited information on what happens to individuals after they depart from the site. We used an automated radiotelemetry array extending 350 km from southwest Nova Scotia to southern Maine to study the migratory and stopover movements of Northern Waterthrushes Parkesia noveboracensis, Red‐eyed Vireos Vireo olivaceus and Yellow‐rumped Warblers Setophaga coronata in relation to fuel load and weather at the northeastern edge of the Gulf of Maine. From the 105 radio‐transmitters we deployed in southwest Nova Scotia, we recorded 42 landscape‐scale stopover flights and 47 migratory flights by 75 individuals. Of the migratory flights, 57% were orientated southwest, a trajectory that, if held, would require individuals to complete a 350–440 km overwater flight. The remaining 43% of migratory flights were orientated northwest, away from the Gulf of Maine, and 15 individuals were confirmed to have detoured around all or a portion of the barrier, as evidenced by their being re‐detected over the Bay of Fundy and/or along the coast of Maine between 4 h and 15 days later. Across all individuals, initial fat score had a positive effect on departure probability, especially for individuals that made stopover flights. Among weather variables, tailwind assistance was the best predictor of migratory departures but did not appear to be the main factor determining whether individuals orientated towards or away from the Gulf of Maine. Weather had little effect on departure decisions of individuals that made stopover flights. These differences in the correlates of migratory departures and stopover flights would probably not have been distinguishable had our study been restricted to a local scale. Therefore, our findings highlight the importance of expanding the scale at which departure decisions and the ecology of stopover in general are studied.     

Wingbeat frequency and flap-pause ratio during natural migratory flight in thrushes

Powered flapping flight has evolved independently in many differenttaxa. For flapping fliers, wingbeat parameters such as frequencyand amplitude are the primary determinants of these animals’energetic expenditure during flight. Here we present data onwingbeat frequency and amplitude for three New World thrushspecies during 15 entire nocturnal migratory flights over theMidwestern United States. Using continuous (non-pulsing) radiotransmitters, we were able to measure wingbeat frequency andrelative amplitude of wingbeats as well as the characteristicsof flap-pauses. Contrary to previous telemetric findings, allof the individuals we followed used both flapping-only and flap-pauseflight. During migratory flights, wingbeat frequency, effectivewingbeat frequency, and amplitude were highest during initialascent. Effective wingbeat frequency and amplitude were lowestduring final descent. We show that identification of speciesbased solely on characteristics of the wingbeat e.g., duringradar studies, can be difficult because variables such as wingbeatfrequency and amplitude, wingbeat pausing, and pattern of beatsand pauses vary between individuals of the same species andeven within individual flights. We also show that observed wingbeatfrequencies were lower than those predicted by theoretical models.We speculate that this may be because theoretical predictionsare generally based on (1) data from larger birds and (2) datafrom diurnal flights. We found that diurnal wingbeat frequenciesof thrushes were generally higher than were those during nocturnalmigratory flight. Finally, we suggest that rather than remainingat a single altitude during flight or climbing slightly as theoreticalmodels predict, thrushes often moved up and down in the aircolumn, perhaps searching for favorable atmospheric conditions.     

Great flights by great snipes: long and fast non-stop migration over benign habitats

Migratory land birds perform extreme endurance flights when crossing ecological barriers, such as deserts, oceans and ice-caps. When travelling over benign areas, birds are expected to migrate by shorter flight steps, since carrying the heavy fuel loads needed for long non-stop flights comes at considerable cost. Here, we show that great snipes Gallinago media made long and fast non-stop flights (4300-6800 km in 48-96 h), not only over deserts and seas but also over wide areas of suitable habitats, which represents a previously unknown migration strategy among land birds. Furthermore, the great snipes achieved very high ground speeds (15-27 m s(-1)), which was not an effect of strong tailwind support, and we know of no other animal that travels this rapidly over such a long distance. Our results demonstrate that some migratory birds are prepared to accept extreme costs of strenuous exercise and large fuel loads, even when stopover sites are available along the route and there is little tailwind assistance. A strategy of storing a lot of energy before departure, even if migration is over benign habitats, may be advantageous owing to differential conditions of fuel deposition, predation or infection risk along the migration route.     

Arrivals of Hitchhiking Insect Pests on International Cargo Aircraft at Miami International Airport

In a study of hitchhiking or contaminating insect pests on international cargo aircraft at Miami International Airport from 1998 to 1999, it was found that contamination rates were greatest, 23%, on cargo flights from Central America and much lower, near 5%, on flights from all other regions. We reanalyzed the study data to test for associations between contaminated flights and factors such as season, cargo type, and time of departure (night or day), and developed probabilistic models for predicting insect pest arrivals by region and pest risk levels. Significant (P < 0.05) associations were detected between contaminated flights and (1) wet season flights from Central America, (2) flights carrying plant products and clothing or fabrics, and (3) flights departing at night from the country of origin. In Monte Carlo simulations, numbers of arriving mated insect pests were greatest for cargo flights from Central America, because of great contamination rates, and South America, because of the large volume of flights from there. Few insects arrived on flights from the Caribbean, and few high-risk insects arrived from anywhere. Although the likelihood of establishment in South Florida via this pathway could not be estimated, based upon arrivals the greatest threats were posed by moderate-risk insect pests on flights from Central and South America. Simulations indicated that switching to daytime departures only reduced pest arrivals by one-third. The simplest mechanism for pathway entry that explains the associations found is that insects entered aircraft randomly but sometimes remained because of the presence of certain cargo types. Hence, contamination rates were greater during the wet season because of greater abundance locally, and on nighttime flights because of greater abundance around lighted loading operations. Empty planes probably had no pests because pests had no access to holds. Thus, the best mitigation strategies for this pathway will likely be those that exclude insects from holds or reduce the attractiveness of night loading operations. Optimizing inspections based on associations is also possible but will be less effective for regions such as South America, with high flight volumes and low contamination rates. Comparisons to other pathways indicates the potential importance of hitchhikers on cargo aircraft at MIA.     

Spring nocturnal migration of Reed Warblers Acrocephalus scirpaceus: departure, landing and body condition

Nocturnal migration of Reed Warblers Acrocephalus scirpaceus was studied by trapping with 'high nets' on the Courish Spit (Eastern Baltic) during spring 1998–2000. In spring, Reed Warblers left the stopover site between 45 and 240 min after sunset (median 84 min), although 85% of birds took off between 45 and 120 min after sunset. Birds did not arrive until the fifth hour after sunset; 67% of birds ended their nocturnal flights in the penultimate hour before sunrise, i.e. at dawn. At the moment of migratory departure, the average Reed Warbler body mass was 12.79 ± 0.66 g ( n  = 60). Average body mass of birds ending migratory flight was 11.69 ± 0.67 g ( n  = 18). The difference was highly significant. However, more than half of the birds completed migratory flights with a considerable fuel load, and some even had energy stores sufficient for a migratory flight on the next night. The spring migratory strategy of Reed Warblers over Central and Northern Europe probably includes a succession of short migratory flights (4–6 h) during several subsequent nights with 1-day stopovers.     

Barometer logging reveals new dimensions of individual songbird migration

Recent advances in tracking technology are based on the use of miniature sensors for recording new aspects of individual migratory behaviour. In this study, we have used activity data loggers with barometric and temperature sensors to record the flight altitudes as well as ground elevations during stationary periods of migratory songbirds. We tracked one individual of red‐backed shrike and one great reed warbler along their autumn migration from Europe to Africa. Both individuals performed their migration stepwise in travel segments and climbed most metres during the passage across the Mediterranean Sea and the Sahara Desert and least metres during the first flight segment in Europe. The great reed warbler reached its highest flight altitude of 3950 m a.s.l. during the travel segment from Europe to west Africa, while the red‐backed shrike reached 3650 m a.s.l as maximum flight altitude during its travel segment from Sahel to southern Africa. Both individuals used both lowlands and highlands for resting periods along their migrations. Furthermore, temperature decreased with increasing altitude during migratory flights for both individuals, highlighting the potential to determine flight duration from temperature measurements. Finally, we discuss how barometric data could be used to investigate birds’ responses to changes in air pressure as a cue for departures on migratory flights. This new technique, i.e. using a miniature data logger with barometric pressure sensor to estimate flight altitudes and ground elevations, will open up new avenues for research and importantly advance our understanding on how small birds behave during migratory flights.     

Vigilance in Greater Flamingos Wintering in Southern Tunisia: Age-Dependent Flock Size Effect

Decrease in individual vigilance with flock size is a widely recognized pattern in group‐living species. However such a relationship may be affected by other factors, such as age and flock composition. For instance, because young animals generally lack experience and have higher nutritional needs than adults, they can be expected not only to be less vigilant than adults but also to decrease their vigilance level by a greater extent when flock size increases than adults do. We investigated this issue using data on greater flamingos wintering in the gulf of Gabès, in southern Tunisia. Flamingos tended to congregate in small single‐age flocks for feeding, but as flock size increased, flocks became mixed. We found that when flock size increased, young flamingos significantly decreased their vigilance time, while adult did not, suggesting an age‐dependent flock size effect on vigilance. However, when flock composition (single‐age vs. mixed) was taken into account, a more complex pattern was found. Within single‐age and small flocks, no difference was found between young flamingos and adult ones regarding their vigilance level and their response to increasing flock size. However, within mixed and large flocks, adult flamingos were more vigilant than young ones, while variation in flock size did not result in a significant change in vigilance. These results suggest that young birds relied on the presence of adults, and hence more experienced individuals in detecting dangers, to reduce their vigilance and to increase their foraging time in order to satisfy their higher nutritional requirements. They could also be interpreted as a possible consequence of increasing competition with flock size which constrained more nutritionally stressed young flamingos to increase their foraging time to the detriment of vigilance.     

Multicontinental community phylogenetics of avian mixed‐species flocks reveal the role of the stability of associations and of kleptoparasitism

If understood as a way to forage socially without incurring intra‐specific competition for mates or other resources, mixed‐species foraging flocks are predicted to be composed of functionally similar species. In the most intensively studied mixed‐species foraging system, understory forest birds, relevant functional traits are however extremely difficult to measure and best replaced by phylogenetic relatedness. A multicontinental analysis of flock phylogenetic structure revealed departures from the expected phylogenetic clustering. Long‐lasting associations (> one day) were phylogenetically overdispersed, indicating that these associations are affected by competitive exclusion or by mutualistic interactions. However, where kleptoparasites occurred, this effect disappeared completely, as expected if the dilution of kleptoparasitism risk compensated competition between related species. Mixed‐species flocks should not be analyzed as a homogeneous phenomenon.     

The ecological significance of extremely large flocks of birds

Population size is generally limited by resource availability during and outside the breeding season. Therefore, maximum size of flocks may provide important information on population regulation and the influence of diet and trophic level on maximal degree of sociality. We hypothesized that (a) flock size should increase with nutrient availability; (b) flock size should decrease with latitude because productivity is higher at lower latitude; (c) aquatic habitats should have larger flocks than terrestrial habitats because the former are less accessible; (d) smaller species should have larger flocks because they require overall less food; (e) human‐impacted species that live in perturbed habitats should have smaller flocks than other species; (f) flock size should decrease with increasing trophic level because there is a reduction in biomass due to conversion at each trophic level; and (g) flocks of species depending on ancestral landscapes should have decreased in size in recent years due to human impact (e.g., land‐use). We obtained 1564 observations of flocks that exceeded 100,000 individuals in order to test the predictions listed above. Most effect sizes were small to medium accounting for 1%–9% of the variance, while large effects accounting for 25% or more were only found for total nitrogen used per km² and area used for agriculture. Changes in large bird flocks were caused by habitat degradation and persecution, and temporal decline in size of large flocks revealed changes in nutrient use, reductions in nutrient cycling, and changes in flock size linked to trophic level.