Low Foraging Success of Semipalmated Sandpipers at the Edges of Groups

Variation in foraging success in relation to spatial position in a group is little known in species that feed on mobile prey that can hide or flee upon disturbance by foragers. I examined the foraging success of individuals located either at the edge or at the centre of flocks of semipalmated sandpipers (Calidris pusilla) feeding on a burrowing amphipod (Corophium volutator) during migration stopover in the Bay of Fundy, Canada. The rates of pecking, prey capture and success were lower for individuals foraging at the edge than at the centre of flocks. Edge birds spent more time running and more time flying than centre birds. Edge birds moved away from the centre of the flock and made frequent short flights towards the centre. In contrast, centre birds rarely moved in a specific direction and flew mostly to relocate elsewhere with the whole flock. Sandpiper flocks foraged over a large area in a relatively short amount of time. In addition, amphipod density is high in this habitat. It thus appears unlikely that prey depletion or low food availability at the edges of groups could explain the spatial variation in foraging success. Low foraging success at the edges of flocks thus arose mainly because of time costs related to flock expansion and retraction. The effect of mutual interference among foragers and of predation risk by falcons is discussed with respect to flock expansion and retraction.     

Bird mixed‐species flock formation is driven by low temperatures between and within seasons in a Subtropical Andean‐foothill forest

According to both the predation avoidance and foraging efficiency hypotheses, birds within mixed flocks increase their foraging efficiency and/or can spend more time feeding and less time looking out for predators. These hypotheses predict that birds in mixed flocks obtain benefits. Thus, mixed flock formation could serve as a strategy to cope with difficult conditions imposed on birds such as climatic conditions that ultimately result in a change in predation pressure or food resources. We evaluate the hypotheses that forming part of a flock confers benefits to its members and the associated prediction that birds will take advantage of these benefits and flock more often under cold and dry weather conditions between and within seasons to cope with such conditions. We surveyed the presence of mixed flocks, flocking propensity, number of species and individuals in mixed flocks in the Subtropical Yungas foothill of Argentina, to examine seasonality, flocking behavior of birds and their responses to two climatic variables: temperature and humidity. Bird species presented a higher flocking propensity and mixed flocks occurred more frequently during the dry and cold seasons than during the more benign seasons, and lower values of temperature within seasons triggered the flocking behavior. Although effects between seasons were expected, birds also showed a short‐term response to small changes in temperature within seasons. These results strengthen the ideas proposed by the foraging hypothesis. Although benefits derived from flocking have yet to be determined, whatever they are should be understood in the context of seasonal variation in life‐history traits.     

The influence of flocking on the foraging behaviour of the chough (Pyrrhocorax pyrrhocorax) and the Alpine chough (P. graculus) coexisting in the Alps

We studied the flocking and foraging behaviour of the chough Pyrrhocorax pyrrhocorax and the Alpine chough P. gruculus coexisting in the south-western Italian Alps in order to evaluate the costs and benefits of foraging in single- and mixed-species flocks.
In the single-species context, flock size significantly affected the foraging behaviour of the Alpine chough; in larger flocks, the birds stayed for a shorter time in a patch and fed more quickly than in smaller flocks. Flock size did not significantly affect the foraging behaviour of the chough, probably because of the small number of individuals per flock.
The propensity for mixed-species flocking was rather low. The observed frequencies of single-species flocks of choughs and Alpine choughs were significantly higher than those expected on the basis of random flocking, whereas the observed frequencies of mixed-species flocks of the two species were lower than those expected. The stay times became significantly shorter for the chough in the presence of the Alpine chough. Moreover, feeding rates of the Alpine chough were significantly lower in the presence of the dominant chough.
The present study does not confirm the hypothetical foraging advantages of flocking. In single-species flocks, the benefits for the Alpine chough (higher feeding rates in larger flocks) were roughly compensated by the costs (shorter stay times in larger flocks), whereas the chough apparently neither gained benefits nor endured costs.
In mixed-species flocks, the Alpine chough sustained costs due to a reduction of feeding rates and the chough suffered costs due to a reduction of stay times. Hence, on average, single-species flocking gives no evident foraging advantages to either the chough or the Alpine chough, whereas mixed-species flocking provides some disadvantages for both species.     

Reduced flocking by birds on islands with relaxed predation

Adaptive hypotheses for the evolution of flocking in birds have usually focused on predation avoidance or foraging enhancement. It still remains unclear to what extent each factor has contributed to the evolution of flocking. If predation avoidance were the sole factor involved, flocking should not be prevalent when predation is relaxed. I examined flocking tendencies along with mean and maximum flock size in species living on islands where predation risk is either absent or negligible and then compared these results with matched counterparts on the mainland. The dataset consisted of 46 pairs of species from 22 different islands across the world. The tendency to flock was retained on islands in most species, but in pairs with dissimilar flocking tendencies, island species were less likely to flock. Mean and maximum flock size were smaller on islands than on the mainland. Potential confounding factors such as population density, nest predation, habitat type, food type and body mass failed to account for the results. The results suggest that predation is a significant factor in the evolution of flocking in birds. Nevertheless, predation and other factors, such as foraging enhancement, probably act together to maintain the trait in most species.     

Predation by Australian magpies (Gymnorhina tibicen) on pasture invertebrates: Are non-territorial birds less successful?

Diet composition and prey intake rates of Australian magpies in a New Zealand population were investigated to determine whether flocking, non-territorial birds were less successful predators of pasture invertebrates than territorial birds. Time-activity budgets showed that flock magpies foraged throughout the day during February and March, indicative of a prey shortage at that time. Prey abundance was therefore measured in the autumn months but there was no evidence of a large quantitative difference in the food supply of flock and territorial magpies. Significantly fewer invertebrates were collected from pitfall traps in the flock foraging area, but these were counterbalanced by significantly more scarab beetle larvae and more flying insects, obtained by sweep-netting. When compared with territorial magpies, flock birds exhibited similar peck rates, prey intake rates, and earthworm intake rates. While there were qualitative differences in the diets of flock and territorial magpies, determined by faecal analysis, five of seven prey items analysed were captured in the same relative proportions by the flock and territorial birds. Earthworms exceeded their relative availability in the diets of flock birds, and more scarab larvae were found in the diets of territorial magpies. Non-territorial magpies were therefore no less successful than territorial birds during the autumn, and it is suggested that the function of flocking behaviour may eventually be discovered through consideration of why non-territorial birds never occur as solitary individuals, rather than emphasizing comparisons with territorial magpies.     

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.     

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.     

Age-specific differences in the winter foraging strategies of rooks Corvus frugilegus

Summary Foraging efficiency and intraspecific competition were compared between wild adult and immature rooks Corvus frugilegus with respect to flock size. Behavioural time budgets, and observations of prey selection and prey energetic values revealed that adult rooks in large flocks (> 50 individuals) consumed smaller, less profitable prey, but allocated more time to feeding and fed at a faster rate and with greater success than adults in small flocks. By contrast, immature rooks in flocks of more than 30 individuals allocated proportionally less time to feeding, fed at a lower rate and fed with no increase in success rate than when foraging in smaller flocks. Agonistic encounters and the avoidance of adults by immature rooks appeared responsible for such inefficient foraging. Hence immature rooks showed a preference for smaller flocks (< 50 individuals) with low adult: immature ratios while adults preferred larger flocks (> 50 individuals). We discuss the possible influence of competitive disadvantages on immature rook distribution, flock composition and post-natal dispersal.     

Behaviourally mediated indirect effects: interference competition increases predation mortality in foraging redshanks

1. The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an example of this. 2. We show that the foraging success of redshanks Tringa totanus (L.) foraging on active prey was highest in the front of flocks, whereas this was not the case for redshanks foraging on inactive prey. Also, when foraging on active prey, foraging success in a flock decreased as more birds passed through a patch, while overall foraging success was not lower on subsequent visits to the same patch. Thus, redshanks foraging on active prey suffered from interference competition, whereas this was not the case for redshanks foraging on inactive prey. 3. This interference competition led to differences in activity: redshanks attaining a lower foraging success had a higher walking rate. Greater activity was associated with wider flock spacing and shorter distances to cover, which has previously been shown to increase predation risk and mortality from sparrowhawks Accipiter nisus (L.). 4. We conclude that behavioural adaptations of prey species can lead to interference competition in foraging redshanks, and thus can affect their predation risk and mortality through increased activity. This study is one of the first to show how interference competition can be a mechanism for behaviourally mediated indirect effects, and provides further evidence for the suggestion that a single species occupying an intermediate trophic level may be simultaneously top-down controlled by a predator and bottom-up controlled by a behavioural response of its prey.     

The advantages of social foraging in downy woodpeckers

I investigated the advantages gained by downy woodpeckers (Picoides pubescens) which join mixed-species winter flocks. Woodpeckers foraging alone showed high levels of vigilance as measured by head-cocking rates, and low feeding rates. Woodpeckers foraging with one or two flock members showed intermediate rates of head-cocking and feeding, while woodpeckers foraging with flocks of three or more birds showed low head-cocking rates and high feeding rates. Although local enhancement and copying may contribute to the woodpeckers' increased foraging efficiency in a flock, these do not appear to be the main factors. As downy woodpeckers spend less time on vigilance, they devote more time to foraging, thereby increasing their foraging efficiency     

Variations in flocking behaviour from core to peripheral regions of a bird species’ distribution range

Flocking behaviour in birds reflects the outcome of a momentary trade-off between increased foraging efficiency and improved predator avoidance. However, these changing patterns remain poorly known at any spatial or temporal scale. The aim of the present study was first to investigate seasonal fluctuations of flocking behaviour throughout the entire distribution range of a species and secondly to explore behavioural responses to daily temperature variations. From 2000 to 2010, sightings of Peruvian Thick-knees (Burhinus superciliaris) were collected throughout Ecuador, Peru and Chile. There were strong differences in flocking behaviour between Chilean and Peruvian populations. While Thick-knees occurred into few large year-round flocks in Chile, flock occurrence was highly seasonal in Peru, where group size grew gradually from loose flocks at the end of the breeding season to a few large ones by the middle of the year. Time of day seemed not to affect the species detectability but was negatively related to flock size. Variations in the flocking behaviour of Peruvian Thick-knees throughout its distribution range suggest that aggregation patterns might indicate individuals responding to seasonally fluctuating ecological pressures, such as those derived from predation, foraging or climate.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

Aerial hunting behaviour and predation success by peregrine falcons Falco peregrinus on starling flocks Sturnus vulgaris

Predators use diverse hunting strategies to maximize hunting success, while preys adopt anti‐predator strategies to maximize escape chances, among which flocking, communal roosting, and the related collective responses are a common pattern in gregarious species. Prey‐predator interactions involving a single predator and flocks, a common situation in birds, have received little attention. We studied predation behaviour and success of peregrine falcons Falco peregrinus on starlings Sturnus vulgaris, a highly gregarious species, in proximity of two winter roosts. A total of 328 hunting sequences, with an overall success of 23.1% were recorded. They usually consisted of several attacks, predation success being higher when hunting sequences lasted less than 1.5 min, included less than 3 attacks and no other falcons were hunting simultaneously. Predation success was higher when hunts were directed on singletons than on flocks. However, most hunting sequences were directed towards flocks. Nine hunting strategies on flocks were identified. The most frequent was the ‘surprise attack’, which was also the most successful. We suggest that this strategy minimizes the amount of anti‐predator display elicited by flocks and economizes energy spent in hunting. The constant predation pressure did not seem to affect the use of roosts by starlings, consistent with the ‘dilution’ hypothesis, while falcons captured at least one prey item every evening. Communal roosting may benefit predator and prey, as both sides could have reached a mutual local equilibrium.     

Nest and foraging‐site selection in Yellowhammers Emberiza citrinella: implications for chick provisioning

Capsule Vegetation structure and invertebrate abundance interact to influence both foraging sites and nestling provisioning rate; when invertebrate availability is low, adults may take greater risks to provide food for their young.

Aims To investigate nesting and foraging ecology in a declining farmland bird whose fledging success is influenced by the availability of invertebrate prey suitable for feeding to offspring, and where perceived predation risk during foraging can be mediated by vegetation structure.

Methods Provisioning rates of adult Yellowhammers feeding nestlings were measured at nests on arable farmland. Foraging sites were compared with control sites of both the same and different microhabitats; provisioning rate was related to habitat features of foraging‐sites.

Results Foraging sites had low vegetation density, probably enhancing detection of predators, or high invertebrate abundance at high vegetation density. Parental provisioning rate decreased with increasing vegetation cover at foraging sites with high invertebrate abundance; conversely, where invertebrate abundance was low, provisioning rate increased with increasing vegetation cover.

Conclusions Vegetation structure at foraging sites suggests that a trade‐off between predator detection and prey availability influences foraging site selection in Yellowhammers. Associations between parental provisioning rate and vegetation variables suggest that where invertebrate abundance is high birds increase time spent scanning for predators at higher vegetation densities; however, when prey are scarce, adults may take more risks to provide food for their young.     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

Modeling the flocking propensity of passerine birds in two Neotropical habitats

We examined the importance of mixed-species flock abundance, individual bird home range size, foraging height, and foraging patch characteristics in predicting the propensity for five Neotropical passerine bird species (Slaty Antwren, Myrmotherula schisticolor; Golden-crowned Warbler, Basileuterus culicivorus; Slate-throated Redstart, Myioborus miniatus; Wilson’s Warbler, Wilsonia pusilla; and Black-and-white Warbler, Mniotilta varia) to forage within flocks, rather than solitarily. We used study plots in primary mid-elevation forest and in shade coffee fields in western Panama. We expected that all species would spend as much time as possible flocking, but that the social and environmental factors listed above would limit compatibility between flock movements and individual bird movements, explaining variability in flocking propensity both within and among species. Flocking propensity was well predicted by home range size and flock abundance together, for four of the five species. While flock abundance was uniform across plots, home range sizes varied among species and plots, so that home range size appeared to be the principle factor limiting flocking propensity. Estimates of flock abundance were still required, however, for calculating flocking propensity values. Foraging height and patch characteristics slightly improved predictive ability for the remaining species, M. miniatus. In general, individual birds tended to join flocks whenever one was available inside their home range, regardless of a flock’s specific location within the home range. Flocking propensities of individual species were lower in shade coffee fields than in forests, and probably vary across landscapes with variations in habitat. This variability affects the stability and species composition of flocks, and may affect survival rates of individual species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal variations in feeding range and flock structure of the Rook Corvus frugilegus in eastern Ireland

The feeding range and flock structure of Rooks showed temporal variations caused mainly by the dispersion of food and reproductive behaviour.
Feeding range was restricted in spring, autumn and early winter when food availability was high and Rooks were reproductively active. A large feeding range occurred in late summer and late winter, when food availability was either generally low or locally distributed and when Rooks were reproductively inactive. The occurrence of flocks common to several rookeries mirrored variations in feeding range; mixed rookery flocks were more common in late summer and late winter. Similarly, the size of the rookery (as measured by the number of nests) was related to feeding range only when Rooks were reproductively inactive and at such times larger rookeries had greater feeding ranges.
Flock structure showed similar seasonal variations; small widely spaced flocks predominated in summer, autumn, early winter and spring, whereas large dense flocks occurred in late winter. Diurnal variations in flock structure occurred within any one season. Both seasonal and diurnal variations in flock structure may be determined by the dispersion of the prey and the feeding strategy used to obtain it, reproductive behaviour and the risk of predation.     

Semi‐intensive shrimp farms as experimental arenas for the study of predation risk from falcons to shorebirds

Varying environmental conditions and energetic demands can affect habitat use by predators and their prey. Anthropogenic habitats provide an opportunity to document both predation events and foraging activity by prey and therefore enable an empirical evaluation of how prey cope with trade‐offs between starvation and predation risk in environments of variable foraging opportunities and predation danger. Here, we use seven years of observational data of peregrine falcons Falco peregrinus and shorebirds at a semi‐intensive shrimp farm to determine how starvation and predation risk vary for shorebirds under a predictable variation in foraging opportunities. Attack rate (mean 0.1 attacks/hr, equating 1 attack every ten hours) was positively associated with the total foraging area available for shorebirds at the shrimp farm throughout the harvesting period, with tidal amplitude at the adjacent mudflat having a strong nonlinear (quadratic) effect. Hunt success (mean 14%) was higher during low tides and declined as the target flocks became larger. Finally, individual shorebird vigilance behaviors were more frequent when birds foraged in smaller flocks at ponds with poorer conditions. Our results provide empirical evidence of a risk threshold modulated by tidal conditions at the adjacent wetlands, where shorebirds trade‐off risk and rewards to decide to avoid or forage at the shrimp farm (a potentially dangerous habitat) depending on their need to meet daily energy requirements. We propose that semi‐intensive shrimp farms serve as ideal “arenas” for studying predator–prey dynamics of shorebirds and falcons, because harvest operations and regular tidal cycles create a mosaic of foraging patches with predictable food supply. In addition, the relatively low hunt success suggests that indirect effects associated with enhanced starvation risk are important in shorebird life‐history decisions.     

Water depth selection, daily feeding routines and diets of waterbirds in coastal lagoons in Ghana

Water depth requirements, diet, feeding styles and diurnal activity patterns are described for waterbirds using two brackish water lagoon systems in coastal Ghana, the Songor and Keta Lagoons. We project the habitat and activity data on a guild structure defined on the basis of individual feeding style and the sensory mechanism used to detect food. A total of 3199 flocks containing 118,648 individuals of 36 different waterbird species were examined during October-November 1994. Feeding habitats varied from dry mudflats to wet mud and shallow water of not more than 20 cm. The depth of water selected by waterbirds for foraging (but not for roosting) was correlated with tarsus length. Foraging birds exhibited a wide range of feeding styles using visual and/or tactile means for detecting prey: pecking, probing, stabbing, sweeping and ploughing, sometimes feeding singly, communally or socially in loose or dense flocks. Prey items taken ranged from seeds of Widgeongrass Ruppia maritima to invertebrates (mainly polychaetes, molluscs and crabs) and fish, mainly juvenile Tilapia. The daytime was spent on two main activities, feeding and roosting, with a small fraction of the time (average of 10% for 25 species) spent on comfort activities. The waterbirds exhibited either a circadian (most waders, except Common Sandpipers Actitis hypoleucos and Turnstones Arenaria interpres) or a diurnal foraging activity pattern (herons and terns), with no purely nocturnal species. Some species fed throughout the day, others showed peak foraging at various times of the day. The proportion of time spent foraging was related to guild (highest in visual and tactile surfaceforaging waders) and was negatively correlated with the size of the species. We conclude that the observed patterns in the use of the 24-h day by waterbirds for foraging are not species specific but vary depending on conditions on the feeding grounds. Nocturnal foraging is a normal and a regular strategy used by waterbirds to obtain enough food to fulfill their energetic requirements, so that irrespective of the sensory mechanism used to detect prey and the conditions prevailing on the feeding grounds, waterbirds forage day and night as dictated by their energetic needs. Water depth appears to be the key environmental factor controlling the availability of food for the waterbirds in the Ghanaian lagoons.     

Effects of weather and tides on feeding and flock positions of wintering redheads in the Chandeleur Sound,Louisiana

We studied the effects of weather and tides on percent feeding and flock positions of wintering redheads (Aythya americana Eyton) in the Chandeleur Sound, Louisiana, USA. Flock scans (n = 750) were made on 55 flocks from November through March of 1988–1989. The percent of the flock that was feeding was negatively correlated with time of day, temperature, water level, and distance of the flock from shore, and was positively correlated with wind velocity, flock size, fetch, and wave height; birds also fed more in early winter and during northerly winds. Flocks were closer to land earlier in the winter on cloudy, rainy, and windy days when waves were high, and those flocks were smaller than flocks farther from shore. Conditions associated with reduced fetch by flocks included later time of day, smaller waves, warmer air temperatures, northerly wind shifts, stronger winds, increasing cloud cover, and rain. Redheads minimized energy expenditure by foraging during low tides and in shallow water closer to shore; they increased feeding and reduced fetch during times of high thermoregulatory demands.